JOURNAL OF THE BOTANICAL RESEARCH INSTITUTE OF TEXAS

HISTORY AND DEDICATION

1962— Lloyd H. Shinners

(left), a member of the

Southern Methodist University

(SMU) faculty and a prolific

researcher and writer, published the first issues of Sida,

Contributions to Botany (now J. Bot. Res. Inst. Texas)

1971—William F. Mahler (right), professor of

botany at SMU and director emeritus of BRIT,

inherited editorshi

nvriaht

PES: z

1993— BRIT becomes publisher/copyright holder.

2007—First issue of J. Bot. Res. Inst. Texas.

MISSION AND GOALS

The BRIT Press seeks innovation and excellence in

preparation, manufacture, and distribution

of botanical research and scientific discoveries

for the twenty-first century.

The BRIT Press—bringing out the best in botanical

science for plant conservation and education.

DIRECTION AND COVERAGE

The BRIT Press considers original research papers

concerned with classical and modern systematic

botany, sensu lato, for publication in J. Bot. Res. Inst. Texas.

All submissions are peer-reviewed.

Guidelines for submissions are available

from the BRIT Press website, http://www.britpress.org.

BIBLIOGRAPHICAL

Citation abbreviation for the

Journal of the Botanical Research Institute of Texas is

J. Bot. Res. Inst. Texas following the principles

of B.P.H. (informally JBRIT).

International Standard Serial No. (ISSN 1934-5259)

FREQUENCY OF PUBLICATION

J. Bot. Res. Inst. Texas is published semiannually

(summer/fall) ne volume

by the Botanical Research Institute of Texas.

J. Bot. Res. Inst. Texas ISSN 1934-5259

VOLUME 1 NUMBER 1 10 AUGUST 2007

Botanical Research Institute of Texas (BRIT)

509 Pecan Street

Fort Worth, Texas 76102-4060, USA

EDITOR

Barney L Lipscomb

Botanical Research Institute of Texas

509 Pecan Street

Fort Worth, Texas 76102-4060, USA

817-332-7432; 817-332-4112 fax

Electronic il: I brit.org; jbritebrit.org

CONTRIBUTING SPANISH EDITOR

Félix Llamas

Dpto. de Botánica, Facultad de Biologia

Universidad de Léon

E2471 Léon, SPAIN

SUBSCRIPTION PRICES (2007)

$41. Personal (Individual/Family)

$85. USA (Institutional)

$95. Outside USA (Institutional)

SUBSCRIPTIONS/BACK VOLUMES

J. Bot. Res. Inst. Texas and Sida, Contr. Bot.

Judy MacKenzie, Publications Assistant

Botanical Research Institute of Texas

509 Pecan Street

Fort Worth, Texas 76102-4060, USA

Electronic mail: jmackenzie@brit.org

COMPOSITION

rhorngraphics, Plano, Texas; rlrc! izon.net

PRINTING

Prepress production and printing in the United States of

America by Millet the Printer, Dallas, Texas

www.millettheprinter.com

COVER ILLUSTRATION

Electronically tinted botanical illustration

of Liatris aestivalis originally used on

BRIT's anniversary poster 2001.

Summer gayfeather flowers mid

Jul-Aug(-Sep) and is endemic to

Oklahoma and Texas.

Sida 19:768. 2001.

Botanical illustration by

Linny Heagy €2001.

TABLE OF CONTENTS

SYSTEMATICS

The world's smallest bamboo: Raddiella vanessiae (Poaceae: Bambusoideae: Olyreae),

a new species from French Guiana

EMMET J. JUDZIEWICZ AND SOL SEPSENWOL

Kalmiopsis fragrans (Ericaceae), a new distylous species from the southern Cascade

Mountains of Oregon

ROBERT J. MENKE AND THOMAS N. KAYE

Taxonomic overview of the Heterotheca fulcrata complex (Asteraceae: Astereae)

Guy L. Nesom

Generic realignments in tribe Potentilleae and revision of Drymocallis (Rosoideae:

Rosaceae) in North America

BARBARA ERTTER

Lectotypifications and new taxa in Potentilla sect. Subviscosae (Rosaceae) in Arizona

BARBARA ERTTER

A new variety of Festuca roemeri (Poaceae) from the California Floristic Province of

North America

BARBARA L. WILSON

Description of Carex klamathensis (Cyperaceae), a rare sedge of the Klamath Region

of Oregon and California, U.S.A.

BARBARA L. WILSON, RICHARD E. BRAINERD, LAWRENCE P. JANEWAY, KELI KUYKENDALL,

DANNA LYTJEN, BRUCE NEWHOUSE, NICK OTTING, STEPHEN MEYERS, AND PETER F. ZIKA

Una nueva especie de Agave, subgenero Littaea (Agavaceae) de Tamaulipas, México

ABISAÍ GARCÍA-MENDOZA, CUAUHTÉMOC JACQUES-HERNÁNDEZ, Y ANGEL SALAZAR BRAVO

Re-examination of Muhlenbergia capillaris, M. expansa, and M. sericea (Poaceae:

Muhlenbergiinae)

Danny J. GUSTAFSON AND PAUL M. PETERSON

Eoépigynia burmensis gen. and sp. nov., an Early Cretaceous eudicot flower

(Angiospermae) in Burmese amber

GEORGE POINAR JR., KENTON L. CHAMBERS, AND RON BUCKLEY

Lectotypification of Gaultheria pyrolifolia and G. pyroloides (Ericaceae)

PETER W. FRITSCH AND DEBRA K. TROCK

A new name for the well-known Asplenium (Aspleniaceae) from Hale County, Alabama

BRIAN R. KEENER AND L.J. DAVENPORT

Relationships of Houstonia prostrata (Rubiaceae) of Mexico and Arizona and a review

of Houstonia subgenera and sections

EDwARD E. TERRELL

A new variety of Humboldtia (Fabaceae: Caesalpinioideae) from the Western Ghats of India

P.S. UDAYAN, K.V. TUSHAR, AND SATHEESH GEORGE

Agasthiyamalaia (Clusiaceae), a new genus for Poeciloneuron pauciflorum, an endemic

and endangered tree of Western Ghats, India

S. RAJKUMAR AND M.K. JANARTHANAM

New names for bamboos of Nepal (Poaceae: Bambusoideae)

C.M.A. STAPLETON

129

135

Folia taxonomica 1. Validation of two taxa from northern South America

CHRISTIAN FEUILLET

Notes on the disarticulation of Xylothamia (Asteraceae: Astereae)

Guy L. Nesom

Two new species of Gratiola (Plantaginaceae) from eastern North America and

an updated circumscription for Gratiola neglecta

Dwayne ESTES AND RANDALL L. SMALL

Review of Crataegus series Apricae, ser. nov., and C. flava (Rosaceae)

J.B. Puiprs AND K.A. Dvorsky

Taxonomy and nomenclature of Taxus (Taxaceae)

RICHARD W. SpjuT

A phytogeographical analysis of Taxus (Taxaceae) based on leaf anatomical characters

RICHARD W. SPJUT

Notes on the Gaylussacia dumosa complex (Ericaceae)

BRUCE A. SORRIE AND ALAN S. WEAKLEY

Reinstatement of Sagittaria macrocarpa (Alismataceae)

BRUCE A. SORRIE, BRIAN R. KEENER, AND ADRIENNE L. EDWARDS

Variation in Petradoria pumila (Asteraceae: Astereae)

Guy L. NESOM AND CALEB A. MORSE

The identity of cultivated Phellodendron (Rutaceae) in North America

JINSHUANG MA AND ANTHONY R. BRACH

Notes on Lechea maritima var. virginica (Cistaceae)

Bruce A. SORRIE AND ALAN S. WEAKLEY

Recognition of Lechea pulchella var. ramosissima (Cistaceae)

Bruce A. SORRIE AND ALAN S. WEAKLEY

Seed and capsule morphology in six genera of Hedyotideae (Rubiaceae):

Thecagonum, Neanotis, Dentella, Kohautia, Pentodon, and Oldenlandiopsis

EDWARD E. TERRELL AND HAROLD ROBINSON

Notes on Phragmites australis (Poaceae: Arundinoideae) in North America

KRISTIN SALTONSTALL AND DONALD HAUBER

The taxonomy of Carex trisperma (Cyperaceae)

CHAD D. KIRSCHBAUM

Thomas Walter Typification Project, II: the known Walter types

DANIEL B. WARD

Thomas Walter Typification Project, III: lectotypes and neotypes for 20 Walter names,

as recognized in the Fraser/Walter herbarium

DANIEL B. WARD

CHROMOSOME NUMBERS

Chromosome number of Laubertia contorta (Apocynaceae: Apocynoideae) and its

phylogenetic importance

Justin K. WILLIAMS AND DAWN P. DERR

143

145

149

171

HERBARIA AND BOTANICAL HISTORY

Vascular plant type specimens in the University of British Columbia Herbarium (UBC)

JEFFERY M. SAARELA, LINDA LiPSEN, CINDY M. SAYRE, AND JEANNETTE WHITTON

ANATOMY AND MORPHOLOGY

Araucarian source of fossiliferous Burmese amber: spectroscopic and anatomical evidence

GEORGE POINAR JR., JOSEPH B. LAMBERT, AND YUYANG Wu

FLORISTICS, ECOLOGY, AND CONSERVATION

Status of Schoenoplectus hallii (Hall’s bulrush) (Cyperaceae) in the United States

PauL M. McKENZIE, S. GALEN SMITH, AND MARIAN SMITH

A note on the type locality of Oenothera arizonica (Onagraceae)

KATHRYN MAUZ

Notes on Conocarpus erectus (Combretaceae) in the Baja California Peninsula, Mexico

José Luis LEON-DE LA LUZ AND RAYMUNDO DOMÍNGUEZ-CADENA

Redescubrimiento de Axiniphyllum sagittalobum (Asteraceae) en la Sierra Madre del

sur y notas de las especies de este género que habitan en el estado de Guerrero, México

ALBERTO GONZALEZ-ZAMORA, ISOLDA LUNA-VEGA, Y JOSE LUIS VILLASENOR

Inventory and distribution of Agave (Agavaceae) species in Jalisco, Mexico

GERARDO HERNANDEZ-VERA, MIGUEL CHÁZARO BASANEZ, AND ERICKA FLORES-BERRIOS

Diversidad y distribucion de la flora vascular acuatica de Tamaulipas, México

ARTURO Mora-OLivo v José Luis VILLASEÑOR

The Cactaceae of the Natural Municipal Park of Prainha, Rio de Janeiro, Brazil: taxonomy

and conservation

ALICE DE MORAES CALVENTE AND REGINA HELENA POTSCH ANDREATA

Assessment of plant biodiversity in Wechiau Community Hippopotamus Sanctuary

in Ghana

ALEX ASASE AND ALFRED A. OTENG-YEBOAH

Botanical composition and multivariate analysis of vegetation on the Pothowar Plateau,

Pakistan

ALTAF A. DASTI, SHEHZADI SAIMA, MOHAMMAD ATHAR, ATTIQ-UR-RAHMAN, AND SAEED A. MALIK

Seed dispersal and soil seed bank of Seriphidium quettense (Asteraceae) in Highland

Balochistan, Pakistan

SARFRAZ AHMAD, SHAMIM GUL, MUHAMMAD ISLAM, AND MOHAMMAD ATHAR

Vascular plant species/area relationships (species richness) in the West Gulf Coastal Plain:

a first approximation

MicHAEL H. MAcRoserts, BARBARA R. MACROBERTS, AND ROBERT G. KALINSKY

Distribution of hanging garden vegetation associations on the Colorado Plateau, USA

JAMES F. Fow er, N.L. STANTON, AND RONALD L. HARTMAN

The vascular flora of the Hancock Biological Station, Murray State University, Calloway

County, Kentucky

RALPH L. THOMPSON

Checklist of the vascular plants of Crawford County, Pennsylvania

CYNTHIA M. MORTON, LOREE SPEEDY, AND JAMES K. BISSELL

529

549

569

Vascular flora of the Four Canyons Preserve, Ellis County, Oklahoma

Bruce W. HOAGLAND AND Amy K. BUTHOD

The vascular flora of three abandoned rice fields, Georgetown, South Carolina:

a 39 year comparison

RICHARD STALTER, JOHN BADEN, AND DWIGHT KINCAID

The vascular flora of Nash Prairie: a Coastal Prairie remnant in Brazoria County, Texas

Davip J. ROSEN

The vascular flora of a woodland park site in east Harris County, Texas

P.A. HARCOMBE, l.S. ELsik, W.W. PRUESS, AND L.E. BROWN

Hypericum adpressum (Clusiaceae) new to Arkansas and the Ouachita Mountains, U.S.A

C. THEO WITSELL

Monarda lindheimeri (Lamiaceae): new to Arkansas

WALTER C. HOLMES AND JASON R. SINGHURST

New records of wetland and riparian plants in southern California, with recommendations

and additions to the National List of Plant Species that Occur in Wetlands

RICHARD E. RIEFNER, JR. AND STEVE BOYD

Noteworthy plants from north Florida. VIII

LORAN C. ANDERSON

New, corrected, and interesting records for the Kansas vascular flora

CALEB A. MORSE, CRAIG C. FREEMAN, AND RONALD L. MCGREGOR

Minuartia drummondii (Caryophyllaceae) and Gratiola flava (Plantaginaceae)

rediscovered in Louisiana and Gratiola flava historically in Arkansas

MicHAEL H. MACROBERTS, BARBARA R. MACROBERTS, CHRISTOPHER S. REID, PATRICIA L. FAULKNER,

AND DWAYNE ESTES

Noteworthy collections from the Yazoo-Mississippi Delta Region of Mississippi

DANIEL A. SKOJAC, JR., CHARLES T. BRYSON, AND CHARLES H. WALKER II

Talinum rugospermum (Portulacaceae) new to Oklahoma

CHRISTOPHER S. REID, PATRICIA L. FAULKNER, BARBARA R. MACROBERTS, AND MICHAEL H. MACROBERTS

Additional noteworthy collections of Cyperus drummondii (Cyperaceae) from

Texas and first report from Mexico

Davip J. ROSEN AND RICHARD CARTER

IN MEMORIAM

Lawrence K. Magrath (1943-2007)

BARNEY LIPSCOMB

Book Reviews AND Notices 8, 20, 58, 68, 78, 90, 120, 128, 134, 290, 366, 372, 406, 424, 436, 456, 482,

486, 510, 528, 576, 584, 005, 654, 678, 712, 752, 762, 768, 785

Announcements 776

713

717

719

741

781

INDEX to new names and new combinations in J. Bot. Res. Inst. Texas 1(1), 2007

Agasthiyamalaia 5. Rajkumar & Janarth., gen. nov.—130

Agasthiyamalaia pauciflora (Bedd.) S. Rajkumar & Janarth., comb. nov.—131

Agave montium-sancticaroli García-Mend., sp. nov.—79

Aristolochia kanukuensis Feuillet, sp. nov.—143

Asplenium tutwilerae B.R. Keener & L.J. Davenport, sp. nov.—104

Carex billingsii (O.W. Knight) C.D. Kirschbaum, comb. et stat. nov.—401

Carex klamathensis B.L. Wilson & L.P Janeway, sp. nov.—71

Crataegus series Apricae J.B. Phipps, ser. nov.—173

Drymocallis campanulata (C.L. Hitchc.) Ertter, stat. et comb. nov.—43

Drymocallis cuneifolia var. ewanii (D.D. Keck) Ertter, stat. et comb. nov.—44

Drymocallis deseretica Ertter, sp. nov. —41

Drymocallis glandulosa var. reflexa (Greene) Ertter, comb. nov.—43

Drymocallis glandulosa var. viscida (Parish) Ertter, stat. et comb. nov.—43

Drymocallis glandulosa var. wrangelliana (Fisch. & Avé-Lall.) Ertter, comb. nov.—43

Drymocallis lactea var. austiniae (Jeps.) Ertter, comb. nov.—36

Drymocallis pseudorupestris var. crumiana D.D. Keck ex Ertter, var. nov.—39

Drymocallis pseudorupestris var. saxicola Ertter, var. nov.—37

Eoépigynia Poinar, Chambers & Buckley, gen. nov.—92

Eoépigynia burmensis Poinar, Chambers & Buckley, sp. nov.—92

Festuca roemeri var. klamathensis B.L. Wilson, var. nov.—59

Gaylussacia bigeloviana (Fernald) Sorrie & Weakley, comb. nov.—336

Gratiola graniticola D. Estes, sp. nov.—166

Gratiola quartermaniae D. Estes, sp. nov.—163

Heterotheca arizonica (Semple) Nesom, comb. et stat. nov.—27

Heterotheca nitidula (Woot. & Standl.) Nesom, comb. nov. —24

Himalayacalamus planatus Stapleton, sp. nov.—137

Houstonia subg. Porotis Terrell, subg. nov.—117

Humboldtia brunonis var. raktapushpa PS. Udayan, K.V. Tushar & Satheesh George, var. nov.—121

Kalmiopsis fragrans Meinke & Kaye, sp. nov.—10

Lechea pulchella var. ramosissima (Hodgdon) Sorrie & Weakley, comb. nov.—370

Medranoa johnstonii (G.L. Nesom) G.L. Nesom, comb. nov.—148

Medranoa palmeri (A. Gray) G.L. Nesom, comb. nov.—148

Medranoa pseudobaccharis (S.F Blake) G.L. Nesom, comb. nov.—148

Medranoa purpusii (Brandeg.) G.L. Nesom, comb. nov.—148

Passiflora foetida var. orinocensis (Killip) Feuillet, stat. nov.—144

Phragmites australis subsp. berlandieri (E. Fourn.) Saltonstall & Hauber, comb. nov.—387

Potentilla demotica Ertter, sp. nov.—53

Potentilla rhyolitica Ertter, sp. nov.—50

Potentilla rhyolitica var. chiricahuensis Ertter, var. nov.—52

Raddiella vanessiae Judz., sp. nov.—1

Taxus biternata Spjut, sp. nov.—266

Taxus brevifolia var. polychaeta Spjut, var. nov.—217

Taxus brevifolia var. reptaneta Spjut, var. nov.—219

Taxus caespitosa var. angustifolia Spjut, var. nov.—268

Taxus caespitosa var. latifolia (Pilger) Spjut, comb. nov.—269

Taxus canadensis var. adpressa (Hort. ex Carriére) Spjut, comb. nov.—274

Taxus canadensis var. minor (Michx.) Spjut, comb. nov.—274

Taxus contorta var. mucronata Spjut, var. nov.—260

Taxus florinii Spjut, sp. nov.—222

Taxus globosa var. floridana (Nutt. ex Chapm.) Spjut, comb. nov.—224

Taxus kingstonii Spjut, sp. nov.—240

Taxus mairei var. speciosa (Florin) Spjut, comb. et stat. nov.—246

Taxus obscura Spjut, sp. nov.—235

Taxus phytonii Spjut, sp. nov.—237

Taxus suffnessii Spjut, sp. nov.—226

Taxus umbraculifera var. hicksii (Hort. ex Rehder) Spjut, comb. nov.—279

Taxus umbraculifera var. microcarpa (Trautv.) Spjut, comb. nov.—279

Taxus umbraculifera var. nana (Rehder) Spjut, comb. nov.—281

Thamnocalamus chigar (Stapleton) Stapleton, comb. nov.—140

Thamnocalamus nepalensis (Stapleton) Stapleton, stat. nov.—140

Thamnocalamus occidentalis (Stapleton) Stapleton, stat. nov.—140

Thecagonum strigulosum (DC.) Terrell & H. Rob., comb. nov.—377

THE WORLDS SMALLEST BAMBOO:

RADDIELLA VANESSIAE (POACEAE: BAMBUSOIDEAE: OLYREAB),

A NEW SPECIES FROM FRENCH GUIANA

Emmet J. Judziewicz and Sol Sepsenwol

PUI UIC OL Biology

University of Wisconsin-Stevens Point

Stevens Point, Wisconsin 54481, U.S.A.

emmet.judziewiczauwsp.edu

ABSTRACT

Raddiella iae (P : Bambusoideae: Olyreae), a new species of herbaceous bamboo, is described and illustrated. It is prob-

ably the smallest bamboo in the GE Known o from savannas in French Guiana, it appears to be related to R. esenbeckii and R.

minima, differing from botl d leaf blades

Key wonps: Bamboo, Bambusoideae, French Guiana, Olyreae, Poaceae, Raddiella

RESUMEN

Raddiella vanessiae (Poaceae: Bambusoideae: Olyreae), una nueva especie de bambú herbáceo probablemente el más pequeño del

mundo, es descrita e ilustrada. Es conocido solamente de las sae us la cen Francesa 3 está relacionado con R. esenbechii y R.

minima, difi d b p i D su menor tamano yl j peq

While some of the approximately 1,000 members of the Bambusoideae may be over 30 m tall, those of

the herbaceous tribe Olyreae can be only a few tens of centimeters tall (Judziewicz et al. 1999). I herein

describe a new species of Raddiella Swallen from French Guiana that, at maturity, is only 2 cm tall and is

thus world’s smallest bamboo:

aag vanessiae Judz., sp. nov. (Figs. 1-2). Tver: FRENCH GUIANA: Savane Lambert 1 (near ip 4°53'N,

31'W, elev. 10 m, savanes herbacées, 15 May 2001, flowering, Vanessa Hequet 1281 (HoLoTYPE: US!; isotypes: CAY, K

A Raddiella minima Ea minore (10-20 mm), lamina minore (2.7-3.3 x 1.8-2.1 mm), flosculis femina minoribus (0.7-0.9 mm),

decl

et carv Opsi ili neare

Mat-forming annual grass with freely-branching culms 10-20 mm tall; culm terete, glabrous, purplish,

shining, the nodes retrorsely ciliolate. Leaves in loose complements of 3-5, with sheaths 1.8-3.2 mm long,

inflated, striate, 7-nerved, retrorsely pubescent with hairs ca. 0.2 mm long; outer ligule absent; inner ligule

a line of erect cilia 0.2-0.3 mm long; pseudopetioles 0.2-0.3 mm long, puberulent; blades 2.7-3.3 mm long,

1.8-2.1 mm wide (area ca. 3.8—5.5 sq. mm), ovate, acute to slightly apiculate at apex, truncate and slightly

asymmetrical at base, apparently folding upwards (involute) under drought stress or at night, with a central

midvein and 6-7 pairs of lateral veins, the upper (adaxial) surface with veins with scattered appressed macro-

hairs 0.05-0.15 mm long, especially near the blade base and margins, the abaxial (lower) surface purple,

glabrous, the blade margins antrorsely scaberulous. Female inflorescences barely protruding from middle and

upper leaf sheaths, a reduced, contracted panicle bearing 2-5 spikelets, the branches and pedicels 0.3-1 mm

long, filiform, glabrous, slightly cupulate at the apex; female spikelets 1-1.4 mm long, ultimately deciduous

but the floret falling first; rachilla internode between glumes somewhat swollen and adherent to the base of

the lower glume; glumes as long as spikelet, subequal, ovate-lanceolate, acute, green, membranous, somewhat

laterally compressed, gaping 20—30? at maturity, 1-3-nerved, retrorsely pubescent with macrohairs ca. 0.2

mm long, the margins cartilaginous; floret 0.7-0.9 mm long, 0.35-0.45 mm wide, lanceolate-ellipsoid to

ovoid, 3-nerved, white, shining, cartilaginous at maturity, deciduous, glabrous, the apex of the lemma acute

and slightly cucullate; lemma much-enfolding the rounded, bicarinate palea, the palea of the same texture as

J. Bot. Res. Inst. Texas 1(1): 1 — 7. 2007

2 Journal of the Botanical R h Institute of Texas 1(1)

Fic. 1. Phot L T | + anaf DAAA all o L . I £hi=A harris || o IRE sehr

le I J LY LAPpvIC VIEH LN

(US).

the lemma; female flower with two subplumose stigmas. Caryopsis 0.65-0.95 mm long, 0.35-0.4 mm wide,

ovoid-ellipsoid, tan to brown, glabrous, slightly dorsally compressed, the embryo basal, 0.1 mm long and

0.2 mm wide, the hilum short-linear, 0.15 mm long, dark brown, located about 0.1 mm above the base of the

caryopsis. Male inflorescence a reduced panicle included in or barely protruding from terminal leaf sheath,

consisting of just one or two spikelets on filiform, glabrous, slightly cupulate pedicels; male spikelets 1.1-1.2

mm long, narrowly lanceolate, hyaline, glabrous, soon deciduous; glumes absent, the lemma 3-nerved, the

palea ca. 1 mm long, bicarinate; stamens 3, the mature anthers brown, 0.4—0.5 mm long.

Judziewicz and Sepsenwol, Raddiella vanessiae, the world's smallest bamboo 3

0.2 mm

Fic 2 R Lae Add P AF I *LaAlat I | ER Ļ J £4 t R Famaloe florat Lahel I

T Li J It J J F F 1 y VMI ventral

view. C. Caryopsis, ventral view showing short-linear hilum. D. Male spikelet, lateral view. Based on Hequet 1281 (US). Illustration by E.J. Judziewicz.

Additional collection examined (paratype): FRENCH GUIANA: Roura, savane marécageuse incluse dans la forêt á 8 km ESE du de-

grad de Roura [approximately 4°40'N, 52°20'W], bord de mare, trés petite herbs en touffes, sur la berge nue, au ras de leau; feuilles

rougeâtres dessous; epillets axillaries, sessiles, vert clair, en partie caches dans les gaines foliares, 21 Apr 1979, A. Raynal-Roques & J.

Jérémie 21288 (CAY).

LEAF AND SPIKELET ANATOMY

A Hitachi S3400 scanning electron microscope was used in the environmental mode to record features of

the leaves and spikelets of Raddiella vanessiae; uncoated, air-dried material was used. Hand cross-sections

of the leaf blades were also made after softening in Pohlstoffe wetting agent. The descriptions below follow

the format of Watson and Dallwitz (1992 onwards):

Abaxial leaf blade epidermis (Fig. 3). Papillae present and abundant in both costal and intercostal zones,

simple, 4—5 um in diameter, the papillae on intercostals long cells generally in two rows. Intercostal long cells

with very sinuous walls. Bicellular microhairs present, elongated, clearly two-celled, ca. 40 um long, the distal

cell ca. 23 um long, ca. 8 um in diameter and remaining inflated in the SEM, the apical cell ca. 17 um long

and deflated by the SEM. Stomata common, 18-23 um long, the subsidiary cells papillate (two on each but

fairly inconspicuous). Intercostal silica bodies vertically elongated, ca. 20 um tall, nodular.

Leaf blade transverse section. Blade ca. 40 um thick at midnerve. Mesophyll consisting or a single abaxial

palisade of arm cells. Fusoid cells absent. Bulliform cells in discrete, regular, intercostals, adaxial, fan-shaped

groups of 3-6 cells, the largest cells 22-25 um tall and 18-22 um wide. Marginal scabrae 15-20 um long.

Vascular bundles all accompanied by sclerenchyma.

Female floret (Fig. 4). Floret smooth except for a few short vertical files of 6-12 um long circular excava-

tions or pits near the summit of the palea, or occasionally one or two ca. 40 um long bicellular microhairs

present on the uppermost margins of the lemma.

fal, Dat o ID L

4 Journal of titute of Texas 1(1)

TÍ) 419 n

Fic. 3. Scanning electron micrograph of lower (abaxial) leaf blade surface of Raddiella vanessiae showing papillate long cells, stomates, bicellular

F 7 Y 4

F YA

DISCUSSION

Raddiella vanessiae is the smallest bambusoid grass yet known, even smaller than Raddiella minima Judz.

& Zuloaga (Zuloaga & Judziewicz 1991; Zuloaga et al. 1993; Judziewicz et al. 1999). Table 1 shows that

both R. minima and R. vanessiae have smaller leaves than the endemic Cuban olyroid Mniochloa pulchella

(Griseb.) Chase (Zuloaga et al. 1993). The new species appears to be related to both R. minima (known only

from the Brazilian type collection made about 1600 km to the south of French Guiana) and the widespread

Neotropical species R. esenbeckii (Steud.) C.E. Calderón & Soderstr. but is much smaller than either one.

The new species is also distinguished from these related species by its smaller female florets and female

lemmas that are slightly cucullate at the apex. The follow key distinguishes the three taxa in the Raddiella

esenbeckii species complex:

1. Plants 8-40 cm tall, perennial; leaf blades 9-22 mm long, 4-11 mm wide, with fusoid cells present;

female spikelets 1.9-2.7 mm long, the glumes firmly membranous, the floret 1.6-2 mm long; anthers

1.3-3 mm long; widespread in tropical South America, Trinidad, and Panama Raddiella esenbeckii

1. Plants 1-6 cm tall, annual; leaf blades 2.7-6 mm long, 1.8-3.3 mm wide, with fusoid cells absent, at least

in R. vanessiae; female spikelets 1-1.4 mm long, the glumes just membranous, the floret 0.7-1.2 mm

long; anthers 0.4-0.6 mm long; rare endemics of tropical South America (French Guiana and Pará, Brazil)

2. Plants 3-6 cm tall; leaf blades 4-6 mm long, 2.7-3.3 mm wide (area ca. 8.5-15.5 sq. mm); female glumes

3-nerved; female floret 0.9-1.2 mm long; hilum punctiform; southern Pará, Brazil Raddiella minima

2. Plants 1-2 cm tall; leaf blades 2.7-3.3 mm long, 1.8-2.1 wide (area ca. 3.8-5.5 sq. mm); female glumes

1-3-nerved; female floret 0.7-0.9 mm long; hilum short-linear; French Guiana Raddiella vanessiae

Judziewicz and Sepsenwol, Raddiella vanessiae, the world's smallest bamboo 5

— -— Miu Tum

UWSP-0909 5.00kV 12.0mm x500 ESED 60Pa 9/15/2006

ln £1 af p PE

CALGVGLUVUITS Yi

J J F

ligi tly TEM | £ +1 £alAs gl ) Racad an LH, ] f 1281 (US).

Cc | I " ° I £ A a PERIE PRR | ££

Fic. 4 I velit al JUE vi

I f I

All three species in the putative Raddiella esenbeckii complex share 1) asymmetrically-based leaf blades that

are often anthocyanic (purple) beneath; 2) leaves that exhibit sleep movements (the blades folding upwards);

3) female spikelets with essentially glabrous, smooth, shining florets that fall before the tardily deciduous

glumes; and 4) an open savanna rather than waterfall-base habitat. The remaining species of Raddiella gener-

ally have elliptical leaf blades lacking anthocyanic pigment and apparently not exhibiting sleep movements;

female florets that are variously papillate throughout; and a waterfall-base (phreatophyte) habitat.

Raddiella vanessiae may be one of the few, or perhaps the only, member of the Bambusoideae with a truly

annual habit. The Cuban herbaceous bamboos, although small, are all cormose perennials (Zuloaga et al.

1993); congeneric Raddiella species are apparently all perennial except possibly for R. minima; and the only

other putative annual is Olyra filiformis Trin. from Bahia, Brazil, which is reported (Soderstrom & Zuloaga

1989) to be either a cespitose perennial or perhaps an annual. In any case it is a much larger plant 40-125

cm tall.

HABITAT

The type locality of Raddiella vanessiae is a lightly-vegetated savanna dominated by grasses (Poaceae), espe-

cially Andropogon bicornis L. and Panicum cyanescens Nees, and sedges (Cyperaceae) including Rhynchospora

holoschoenoides (Rich.) Herter and Scleria cyperina Kunth. The soil is sandy and the lower parts of the savanna

seasonally flood, but Raddiella vanessiae grows in the driest (highest) part, where it is uncommon (Chaix et

al. 2002). See http://www.cayenne.ird.fr/aublet2/Selection_Collecteur.php3 for a complete list of Vanessa

f4L,D o ID L

6 Journal of t titute of Texas 1(1)

TABLE 1. Comparison of Raddiella vanessiae with related species and Mniochloa pulchella. Approximate leaf blade area was

calculated from the formula a = rrlw/4 (the formula for the area of an ellipse), where | is leaf blade length and w is the blade

widt

Character Raddiella esenbeckii Raddiella minima Raddiella vanessiae Mniochloa pulchella

Habit Cespitose perennial Annual? Mat-forming annual Cormose perennial

Plant height (cm) 8-40 3-6 1-2 3-12

Leaf blade length (mm) 9-22 4-6 2.7-3.3 7-15

Leaf blade width (mm) 4-11 2./-33 1.8-2.1 2-4

Leaf blade area (square mm) 28-104 8.5-15.5 3.8-5.5 11-47

Fusoid cells present or absent ? absent absent

Leaf blade sleep movements? yes yes yes no?

Female spikelet length (mm) 1.9-2.7 1-14 1-14 2.2-2.8

Female spikelet glume texture firmly membranous; membranous; membranous; delicately

becoming blackish remaining green remaining green membranous,

at maturity at maturity at maturity green at maturity

Female spikelet glume 3 9 les 3

nerve number

Female floret length (mm) 1.6-2 0.9-1.2 0.7-0.9 2.2-2.8

Male spikelets/inflorescence (1-)2-4 1 Ora (3-)7-12

Male spikelet length (mm) 3- Cale 1,2 1.3-1.7

Anther length (mm) 1.3-3 0.6 0.4-0.5 0.8-1

Caryopsis length x width (mm) 1-1.2 x 0.7-0.8 0.7 X 0.6 0.65-0.95 x0.35-0.44 1.5-2 x 0.5-0.6

Hilum morphology short-linear punctiform short-linear linear

Distribution Panama, tropical South Southern Pará, French Guiana Cuba

America, Trinidad Brazil

Hequet's specimens (1238-1245, 1250-1263, 1268-1284) collected on 15 May 2001 in the Savane Lambert.

Google Earth (http://earth.google.com/) shows “Savane Lambert” to be several square kilometers in area and

situated about 20 km WSW of the city of Cayenne. The Raynal-Roques & J. Jérémie specimen of Raddiella

vanessiae is from a swamp savanna on the Roura-Kaw road, about 27 km by air from the type locality.

Although widespread in South America, there are only two records of the related species Raddiella

esenbeckii from French Guiana, from Passoura and the Savane des Singes (Judziewicz 1991), about 25 km

NW of the type locality of R. vanessiae.

LEAF AND SPIKELET ANATOMY

The leaf anatomy of Raddiella vanessiae is more or less consistent with descriptions of the anatomy of R.

esenbeckii (Watson & Dallwitz 1992 onwards) with one possible exception. Fusoid cells are absent in Rad-

diella vanessiae, but apparently can be either absent (Calderón & Soderstrom 1967; Renvoize 1985; this study,

based on hand sections of R. potaroensis Soderstr. (Redden et al. 1465, Guyana, UWSP)), or present (Watson

& Dallwitz 1992 onwards, based on R. esenbeckii; this study, based on hand sections of R. esenbeckii (Schwab

491, Aripo Savanna, Trinidad, UWSP) in species of Raddiella. While fusoid cells are a characteristic feature

of the leaf blades of most species of the Bambusoideae, they are absent in several distantly related taxa in the

herbaceous Olyreae (Judziewicz et al. 1999: 33), and this absence is probably a derived condition: the small

Cuban genera Ekmanochloa Hitchc. and Mniochloa Chase (Zuloaga et al. 1993) and the Brazilian Parodiolyra

ramosissima (Trin.) Soderstr. & Zuloaga (Renvoize 1985; Soderstrom & Ellis 1987) all lack fusoid cells.

In leaf blade transverse section, Watson and Dallwitz (1992 onwards) could not discern arm cells in

Raddiella esenbeckii; however, these were easily visible in hand-cut sections of R. vanessiae blades and in

blades of R. esenbeckii (based on Schwab 491). The micromorphology of the female floret of Raddiella vanessiae

is similar to that of R. esenbeckii (Zuloaga & Judziewicz 1991): its surface is essentially smooth except for

small longitudinal files of curious tiny “excavations” or “pits” at the summit of the palea that may represent

Judziewicz and Sepsenwol, Raddiella vanessiae, the world's smallest bamboo 7

deflated cells. Seemingly identical *pits" are present on the female paleas of other olyroid bamboos such as

Parodiolyra lateralis (Nees) Soderstr. & Zuloaga and various species of Olyra such as the widespread common

species O. latifolia L. (Soderstrom & Zuloaga 1989).

ETYMOLOGY

At the suggestion of my colleague Isabelle Girard, we had considered naming this new species for the mythical

country of Lilliput (from Jonathan Swift's novel Gulliver's Travels) where it would be an appropriately-sized

bamboo for the use of the Lilliputians. Instead, we have chosen to recognize Vanessa Hequet, whose type

collection and careful and detailed characterization of the species’ savanna community habitat was so useful

in preparing this paper. The epithet seems doubly appropriate: Vanessa was Swift's nickname for his close

friend Esther Van Homrigh.

ACKNOWLEDGMENTS

We thank Fernando O. Zuloaga and Reyjane Patricia de Oliveira for helpful reviews, Lynn G. Clark and

Carol Annable for useful advice, Paul M. Peterson and Steve Smith (both US) for the loan, Jean-Jacques

de Granville (CAY) for type specimen information and alerting me to the presence of a second collection,

type specimen collector Vanessa Hequet for extensive habitat information, to Virginia Freire for help with

the Spanish translation, and Isabelle Girard for suggesting species epithets. Funding for this research was

provided by a University of Wisconsin-Stevens Point Faculty Development Grant for travel to Washington,

D.C. (Judziewicz) and National Science Foundation CCLI Program grant 40511131 for the purchase of the

Hitachi $3400 SEM (Sepsenwol).

REFERENCES

CALDERÓN, C.E. and T.R. Soperstrom. 1967. Las gramíneas tropicales afines a Olyra L. Atas do Simpósio sôbre a Biota

Amazónica (Conselho de Pesquisas, Rio de Janeiro) 4 (Botanica):67—76.

Chaix, M. V. HEQuET, M. BLANC, O. Tostar, T. DeviLLe, and P. GomBAULD. 2002. Connaissance et conservation des savanes

de Guyane. Rap. IFRD-WWF Guyane.

Jupziewicz, E.J. Family 187. Poaceae. 1991 [as 1990]. In A.R.A. Górts-Van Rijn, ed., Flora of the Guianas, Series A:

Phanerogams. Koeltz Scientific Publications, Kónigstein, Germany.

JupziEWICZ, E.J., L.G. CLARK, X. LonboÑo, and MJ. Stern. American bamboos. 1999. Smithsonian Institution Press,

Washington, DC.

Renvoize, S.A. 1985. A survey of the leaf-blade anatomy in grasses. V. The bamboo allies. Kew Bull. 40:509-535.

SODERSTROM, T.R. and F.O. ZULOAGA. 1989. A revision of the genus Olyra and the new segregate genus Parodiolyra

(Poaceae: Bambusoideae: Olyreae). Smithsonian Contr. Bot. 69:1—79.

Soperstrom, T.R. and R.P. Elus. 1987. The position of bamboo genera and allies in a system of grass classification.

In: Soderstrom, T.R. et al., eds., Grass Systematics and Evolution. Smithsonian Institution Press, Washington,

D.C. Pp. 225-238.

Watson, L. and MJ. Dattwitz. 1992 onwards. The grass genera of the world: descriptions, illustrations, identifica-

tion, and information retrieval; including synonyms, morphology, anatomy, physiology, phytochemistry,

cytology, classification, pathogens, world and local distribution, and references. Version: 28th November

2005. http://delta-intkey.com.

ZULOAGA, F.O. and E.J. Jubziewicz. 1991. A revision of Raadiella (Poaceae: Bambusoideae: Olyreae). Ann. Missouri

Bot. Gard. 78:928-941.

ZULOAGA, F.O., O. Morrone, and EJ. Jupziewicz. 1993. Endemic herbaceous bamboo genera of Cuba. Ann. Missouri

Bot. Gard. 80:846-861.

8 Journal of the Botanical R h Institute of Texas 1(1)

BOOK REVIEWS

PIERRE DELFORGE. 2006. Orchids of Europe, North Africa and the Middle East. (ISBN-13: 9780881927542;

N-10: 0881927546, hbk.). Timber Press Inc, 133 S.W. Second Avenue, Suite 450, Portland, OR

97204-3527, U.S.A. (Orders: www.timberpress.com, mail@timberpress.com, 503-227-2878, 1-800-

327-5680, 503-227-3070 fax). $39.95, 640 pp., 1270 color photos plus several line drawings and

watercolors, 5" x 715" (190 x 125 mm).

MATE : ee} 1 1 1] a ] m

ld be. His work is meticulous and although limited to brief entries

for Es species, as complete as space permits. The gallery of photographs is mind-boggling. This current book is no exception. It com-

bines the in a: all of his ome books and 2o a es volume that is both an e of previous works D us new material

Technically a 3rd edi

ae more than 200 additional pages than previous French editions

oe chapters on basic orchid information anatomy i cycles, reproduction and orchid identification are detailed

and easy to understand with many excellent graphics. The f ters of MUN accounts are broken into four chapters and

d by subtribes and the genera arranged systematically. i n de t for the 2d it does make it

difficult to find a specific aed or d quickly without consulting the index. HK of the large geograp covered by the

book, one must read throug very carefully to find tl Į igl 1 1 ler is exploring. This

gati pect f the book but d f ] g y more species.

A few i itl l i and fortunately they in the ind ell. Several taxonomical and nomen-

clatural points should be em

The gens Listera is pu. into Neottia, Pseudorchis into lea: OS is maintained in its own genus rather than in

] white-flowered forms and other various

Dactylorhiza. t

color variants.

The largest genera treated—Epipactis, Dactylorhiza, Orchis, and Ophrys are as complete as they can possibly be with extensive

illustrations to isis sort out p bus pas. odd and pu Ophrys alone has over 250 dn pas to the genus!

habitat

35 years in Europe,

North Africa, and 7 Middle East and is an expert on C Mis for the IUCN—the Wont Conservation Union.

ilahlle! Paul

Martin Brown, Author, Wild 9 Ui orida, The Wild Orchids of North America, Wild Orchids of the Southeastern United States, 10896 SW

90" Terrace, Ocala, FL 34481, U.S.A., naorchid@aol.com

Note.—This title has been D in Great Britain by A & C Black (ISBN 9780713675252) and cooperatively with Timber Press in

the North America. They h images but the body of the books is the same

Mary Toomey with Everett Leeos and CHARLES CHESSHIRE. 2006. Timber Press Pocket Guide to Clematis.

(ISBN 0-88192-814-3, pbk.). Timber Press Inc, 133 S.W. Second Avenue, Suite 450, Portland, OR

97204-3527, U.S.A. (Orders: www.timberpress.com, mail@timberpress.com, 503-227-2878, 1-800-

327-5680, 503-227-3070 fax). $19.95, 232 pp., 319 color photographs, 1 b/w map of USDA Hardiness

Zone, 1 full color map of European Hardiness Zones, 5%" x 834".

The famous Timber Press Pocket Guides are always first-rate and this particular one ins not nu From its soft, pliable cover

ith ded t into a large pocket, purse, or garden trug I 1 desi ]

pl lotograpiy and design; what's not

to like. It must have been a daunting task, indeed, to condense the selection a Clematis from 550 in the original beautifully published

[2]

version, An Illustrated Encyclopedia of Clematis (2001) to the 300 in this ‘pocket’ version, but what a success! Diminutive in size but

packed with brilliant full-color photography of each of the 300 stunning Clematis cultivars that any gardener would be proud to have

in their garden. Though Dos e brilliance, it is not busy looking. Just the right amount of margin and leading between the lines

of san serif type to give ea ability and space around the photos. Complete with Nursery Sources for 16 countries, a Glossary,

Further Reading resources, and ben

The authors' ] full ] I iel lematis for tl ] i i } d d Clematis

for specific purposes. “Careful selection, mM judicious pruning—never let that tax your dedu with a modicum of patience and

diligence will pay handsome dividends."—Linny Heagy, Linny/Designer, Illustrator, email: a0005835@airmail.net.

J. Bot. Res. Inst. Texas 1(1): 8. 2007

KALMIOPSIS FRAGRANS (ERICACEAE), A NEW DISTYLOUS SPECIES FROM THE

SOUTHERN CASCADE IMOUN TAINS OP OREGON

Robert J. Meinke and Thomas N. Kaye

Department of Botal y dal 1d Plant t Pathology

2082 Cordley Hall

Oregon State University

Corvallis, Oregon 97331-2902, U.S.A.

ABSTRACT

The azalea-like genus Kalmiopsis Rehder (Ericaceae) is endemic to southwestern Oregon, U.S.A., and has two disjunct centers of dis-

tribution. One is found in the Klamath (Siskiyou) Mountains of pd and Josephine cos. Mm the U.S. Forest Service administered

Kalmiopsis Wilderness Area. The other is located 150 km northeast in the southern Cascade Mountains of Douglas Co.

Kalmiopsis is horticulturally significant, and plants from both localities have been established in the garden trade since shortly after

their initial discoveries in 1930 and 1954, respectively. The genus has traditionally been considered monotypic, consisting only of

Kalmiopsis leachiana (Hend.) Rehder. However, comparative studies of the morphology, floral biology, and ecology of the northern and

southern population groups have determined they are best treated as distinct taxa. The northern populations are described here as the

new species Kalmiopsis fragrans, a rare endemi urring on silicified tuffaceou ps within a narrow segment of the Umpqua

National Forest.

RESUMEN

El género de azaleas Kalmiopsis Rehder (Ericaceae) es endémico del suroeste de Oregon, U.S.A., y tiene dos centros disyuntos de dis-

tribución. Uno se encuentra en las montafias Klamath (Siskiyou) de los condados de Curry y Josephine, en la Kalmiopsis Wilderness

Area administrada por el U.S. Forest Service. La otra esta localizada a unos 150 km al noreste en el sur de las Cascade Mountains de

Douglas Co. Kalmiopsis es importante desde el punto de vista de la horticultura, y ur pianti po aces de ee localidades

desde poco después de su descubrimiento inicial en 1930 y 1954, respectivamente. El g

monotípico, ánicamente con Kalmiopsis leachiana (Hend.) Rehder. Sin embargo, estudi mparativos de la morfología, biología floral,

y ecología de los grupos de poblaciones del norte y del sur han determinado que es mejor como taxa distintos. Las poblaciones

del norte se describen aquí como la especie nueva Kalmiopsis fragrans, una especie endémica que aparece en afloramientos silíceos

en una pequefia parte del Umpqua National Forest.

INTRODUCTION

Kalmiopsis Rehder is the only vascular plant genus endemic to the state of Oregon. The interesting cir-

cumstances of its discovery are chronicled in Kirkpatrick et al. (1994). Populations were first observed by

botanists in 1930 in the mountains of southwestern Oregon (Henderson 1931; Rehder 1932), a rugged area

with few trails and limited access. Although this craggy and remote region was already gaining signifi-

cance as a haven for unusual and relict species (Whittaker 1960), the report of this beautiful, low-growing

shrub, reminiscent of Kalmia and various cultivated azaleas (Rhododendron spp.), attracted considerable

taxonomic and horticultural interest (Love 1991). Within months of its discovery, efforts were underway to

introduce Kalmiopsis to the nursery industry. Although early tradesmen found Kalmiopsis a difficult subject

for propagation (Love 1991; Kirkpatrick et al. 1994), plants may be grown with perseverance (Mulligan

1973; Kruckeberg 1982) and are today occasionally established in private and public gardens in the Pacific

Northwest, Europe, and elsewhere.

Kalmiopsis has a bimodal distribution pattern, occurring most abundantly in the Klamath (Siskiyou)

Mountains of Curry and Josephine cos., Oregon, just inland from the Pacific Ocean near the California border.

A second, more restricted series of populations is clustered approximately 150 km to the northeast within

the Umpqua River watershed of the southern Cascade Mountains, an area with a very different geologic and

vegetative history (Whittaker 1960; Marquis 1977). The older Klamath Mountains connect the Coast Ranges

J. Bot. Res. Inst. Texas 1(1): 9 — 19. 2007

£s+haD o ID L

10 Journal of t titute of Texas 1(1)

of Oregon and California, and encompass a complex series of Mesozoic (Jurassic) formations including brec-

cias, tuffs, sandstones, cherts, and conglomerates, much of which has been altered to metavolcanics (Baldwin

1974; Ramp 1975; Marquis 1977). Ultramafic substrates are widespread in the Klamath Mountains, where

Kalmiopsis populations routinely occur on harsh, open serpentine habitats, though they are not necessarily

restricted to them. The geology of the North Umpqua River drainage in the southern Cascades is mostly of

Cenozoic origin (Peck et al. 1964, with the handful of Kalmiopsis populations here endemic to localized pin-

nacles of siliceous tuff, mostly in deep coniferous forests.

Henderson (1931) described the newly discovered species as Rhododendron leachianum, initially align-

ing it with the arctic-alpine R. lapponicum. Shortly thereafter, Rehder (1932) evaluated the new species and

considered it to be closely related to the montane Eurasian genus Rhodothamnus, with possible affinities to

the circumboreal genera Kalmia, Rhododendron, and Phyllodoce. Rehder ultimately concluded that the Oregon

plants were unique enough to merit establishing the monotypic genus Kalmiopsis. Copeland (1943, 1954

subsequently placed Kalmiopsis leachiana (Hend.) Rehder within Rhodothamnus, proposing to drop the new

genus based primarily on anatomical similarities between the taxa. However, Davis (1962), in his more recent

study of Rhodothamnus, disagreed with the assessments by Copeland, and Kalmiopsis leachiana remains the

currently accepted epithet (Stevens 1971; Harborne and Williams 1973). Recent phylogenetic studies of the

subfamilies Rhododendroideae and Phyllodoceae (Kron and King 1996; Kron 1997; Kron et al. 2002) like-

wise align Kalmiopsis with Rhodothamnus, supporting the earlier conclusions by Rehder (1932) and Copeland

(1943, 1954). However, based on matK and rbcL sequence data, Kron (1997) and Kron et al. (2002) consider

Kalmiopsis most closely related to Phyllodoce, the genera also being linked by their distinctive multicellular

hairs with biseriate stalks.

The discovery of Kalmiopsis populations in the Cascade Mountains raised questions about the relation-

ship between these plants and those from the Klamath Mountains to the southwest. Well before taxonomic

questions were posed, nurserymen noticed apparent differences between the disjunct populations, recogniz-

ing the hardier nature and comparative ease in transplanting of the Cascadian plants (Love 1991). Callan

(1971) and Marquis (1977) acknowledged potential morphological differences between the populations as

well, implying that recognition as distinct taxa might be warranted. However, neither author proposed any

formal taxonomic separation.

In the last few years, opportunities have arisen for more extensive field studies of the genus in the

Klamath and Cascade Mountains. The research described here, undertaken to evaluate the suitability of

separating the groups as discrete taxa, contrasts the morphology, floral biology, and ecology of the northern

and southern populations of Kalmiopsis. Herbarium material (including the holotype of K. leachiana) and live

specimens from eleven populations (four from the Cascades and seven from the Klamath Mountains) were

utilized in the comparisons. Measurements were taken from fresh flowers, as many herbarium specimens

of Kalmiopsis tend to have shriveled corollas that underrepresent floral dimensions. Results from our studies

support the recognition of the isolated northern populations as a distinct species.

Kalmiopsis fragrans Meinke & Kaye, sp. nov. (Figs. 1-4). Tre: U.S.A. OREGON. Douctas Co.: ca. 0.7 km W of Dry Cr.

settlement, above N Umpqua R. on S exposure of cliffs along Panther Leap, Umpqua National Forest, T 26S R2E S19 NE 1⁄4, elev.

ca. 570 m, in mixed coniferous forest with Pseudotsuga menziesii, Tsuga heterophylla, Whipplea modesta, Castanopsis chrysophylla,

Polystichum munitum, and Linnaea borealis, 18 May 1993, R. Meinke and T. Kaye 6280 (noLoTYPE: OSC; isotypes: HSC, NY, RSA, UC,

US, UTC, WS, WTU).

Plantae ab Kalmiopsis leachiana habitu erecto aut serpenti, limb 11 ad 2-3 mm prope tubum, limbo et tubo corollae fere plano

et rotato ubi maturo, ovario luteolo, ciliis densis ad bases filamentorum et in tubo corollae, odore florum simili ad azaleam differt.

Evergreen shrub, + woody below (depending on age and size), usually openly branched (although internodes

will shorten and habit become condensed in full sun), mature plants tenaciously rooted in shallow soils on

the forest floor, or more often loosely attached and clambering over rocky substrates with thin organic mats,

sometimes draped over or hanging from vertical cliffs, occasionally suspended under rock overhangs, capable

of vegetative propagation by subterranean stems (often through rock fissures) or via adventitious rooting,

Meinke and Kaye, A new species of Kalmiopsis from Oregon 11

Fic. 1 Cl £ Alm: f ; . enel I: | Darb D h Mat I Area, ITI pq Maa Ir t Douglas Co., Oregon

occasionally crown-sprouting, locally forming populations consisting of one or few clones; stems few to

numerous, (22)4-12(-30) dm long, + erect to trailing and occasionally + matted, arising from a thickened

base, glabrous or rarely with sparse, fine, white pubescence, often brittle, exfoliaceous, epidermis reddish

to gray, terminally leafy and mostly naked below, the new growth + stipitate-glandular and often fragrant

with a faintly sweet scent; leaves numerous, crowded above (or less so in deep shade), thinly coriaceous

with a thick cuticle above and below, dark green, glabrous, and shining above (or reddish in anthocyanic

individuals in full sun), paler and punctately dotted with golden-crystalline, sweetly aromatic glands below,

blade (5)8-30(-45) mm long, elliptic to obovate, apiculate, petioles glabrous to finely puberulent, less than

one-fifth the length of the blades, erect to horizontally oriented; inflorescence terminal, corymbose to +

racemose, (2—)4—8(-12) per corymb, floral bud scales 2-3 mm long, membranaceous, glabrous or with scat-

tered crystalline glands, pale to reddish-pink, broadly lanceolate; flowers dimorphic, consisting of long- and

short-styled forms on separate plants, + aromatic with a spicy-sweet odor, somewhat azalea-like, the aroma

persisting and often intensifying with age; pedicels 0.5—2.5(-3.3) cm long, mostly glandular-pubescent;

calyx glabrous, urceolate to campanulate, greenish to mostly pale pink or red, sepals 3-6(-8) mm long,

overlapping at the base, thin, broadly lanceolate, margins + involute; corolla pale reddish-purple to deep

pink when fresh, deciduous, actinomorphic, lacking a defined throat, the shallow tube <2 mm deep, the limb

16-28(33) mm across, broadly cupped to campanulate in early anthesis but becoming essentially rotate

with age as the petal lobes reflex, lobes deltoid-ovate, 6-12 mm long and 4-8 mm wide, with two parallel,

ventral ridges giving petals the appearance of thickness, petal sinuses divided to within 2-3 mm of the

corolla tube, the lower edges of the petal lobes overlapping; stamens 10, nestled along the petal ridges in

fal, Dat o ID L

12 Journal of titute of Texas 1(1)

[AY

Fi ) | £1 f, sel [| X £ Val, RES ut 74 leachiana (B) | +. J cnt £1 I + L lb £ y fraarans

X r I NE ir Y J A

£4. H H Í | 2 1 £f. [| Pi EE | " E. akt. x sat * BA |

[ g glandular hairs, while K. Corollas of K, fragrans (0)

I. ES On 4 | 4L H {10 SAL £s L i} ros | I 4 - J +

and K. 24 hrs after bud break),

War katy a \

bud and as corollas open, spreading to erect and well exserted at full anthesis, those of long-styled flowers

7-13 mm long and those of short-styled flowers 11-16 mm long, filaments light pink or paler, basally dilated,

glabrous or with scattered hairs above, typically with a copious tuft of pale yellow to golden translucent hairs

present at the very base (rarely subglabrous throughout), these joining to form a dense ring of pubescence

in the floral tube that surrounds and generally conceals the base of the ovary; anthers light purple, narrow-

oblong, often slightly curved, 1.2-3.0 mm long, the terminal pore openings «0.4 mm across; pollen cream to

Meinke and Kaye, A new species of Kalmiopsis from Oregon 13

ochroleucous, tetrads, 50—60 y, no differences noted

between floral morphs; styles red to purple, usually

glabrous or rarely with a few isolated hairs, 11-15

mm long in long-styled morphs and 5-8 mm long in

short-styled morphs; stigma pale, rounded, + capitate

to shallowly bi-lobed, obviously sticky, no differences

noted between floral morphs; ovary 2-3 mm wide

and high, globose, pale yellowish gold, glandular,

o ovules numerous; capsule + depressed, 3-5 mm

4 broad, shallowly five-lobed, glandular-warty; seeds

i A minute, 0.3-0.7 mm long, oblong, shallowly pitted,

potentially over 150 per capsule (although abortion

may result in far fewer).

Distribution and Habitat.—Kalmiopsis fragrans

is endemic to a narrow area along the west slope of

the southern Cascade Mountains in Douglas Co.,

Oregon, with a known elevational range of 450 to

1325 m. It is apparently restricted to lands adminis-

tered by the Umpqua National Forest, mostly within

the drainage ofthe North Umpqua River in the vicin-

ity of Steamboat and along Ragged Ridge. Populations

normally occur in deeply shaded to partially open

sites, commonly on or closely adjacent to talus slopes,

boulder piles, or immense pillars of silicified tuff

with south-facing aspects. The new species occurs

in mixed coniferous forests variously dominated by

Pseudotsuga menziesii, Abies grandis, Tsuga heterophylla,

Fic. 3. Fl f Kalmiopsis fi (A) and K.leachiana(B)atlateanthesis Calocedrus decurrens, Arbutus menziesii, Thuja plicata,

(ca. 3-4 days after bud break), showing relative reflexing of petals and and Pinus lambertiana. Understory associates include

ge in| ( g. 2) of stamens Berberis nervosa, Holodiscus discolor, Gaultheria shal-

lon, Oxalis oregana, Whipplea modesta, Castanopsis

chrysophylla, Polystichum munitum, Linnaea borealis, Rosa gymnocarpa, Pterospora andromedea, Pleuricospora

fimbriolata, Allotropa virgata, Rhododendron macrophyllum, Acer circinatum, Rhus diversiloba, Goodyera oblongi-

folia, Thermopsis montana, Iris chrysophylla, Pyrola picta, Sanicula graveolens, Viola orbiculata, Calypso bulbosa,

Erythronium citrinum, and Luzula campestris.

Most reproductive individuals and seedling recruitment are found in areas of filtered sunlight on rocky

slopes, as well as on rock columns and other semi-moist outcrops that extend into or through the forest

canopy. A few populations occur on exposed, rocky ridges. However, K. fragrans is a shade-tolerant species,

capable of surviving for years within shallow caves and overhangs while growing from high rock ceilings or

along deeply sheltered cliffs, persisting in such sites (although with reduced reproductive output) in what

are essentially twilight conditions. Populations from deeper forests appear generally intolerant to sudden

exposure, and are apt to decline over the long term in response to habitat disturbance that results in reduced

soil moisture and increased sunlight. In particular, plants remaining after clear-cuts will persist for a short

time, but have been noted to eventually become anthocyanic, more susceptible to disease, and to suffer

higher mortality when compared to those from adjacent undisturbed sites.

Flowering —Populations typically flower from mid-April to early June, depending on elevation. Seed

production and dispersal occurs into August. The intensity of flowering in Kalmiopsis fragrans appears largely

correlated with habitat quality. Plants located in densely shaded conditions may seldom, if ever, bloom or

f*hAD o ID L

14 Journal of t titute of Texas 1(1)

ona £ Uni? *- a [A and Rì and K leachiana (€ and D) E M AO | £ tal Int

bd" Era P V 1 \ iy

F 4 fut G fa WU ES

IG. * NULAVYA y VICVYA YI JIIVIV Git I I y

produce seed, while those in areas of consistent, filtered light often flower profusely. Flowering in popula-

tions that become highly exposed due to logging or other site disturbance tends to decline over time.

COMPARISON WITH KALMIOPSIS LEACHIANA

Overall differences between Kalmiopsis fragrans and K. leachiana are summarized in Table 1. The floral and

inflorescence morphologies of the species are illustrated and contrasted in Figs. 2—4.

Habit.—Kalmiopsis leachiana is described as being a small, copiously branched shrub up to 3 dm high

(Rehder 1932; Abrams 1951; Peck 1961), but field work associated with this paper indicate that plants of this

species may occasionally trail and grow to several dm long on steep terrain. The typical habit, however, is

low and compact, with plants often flowering in dense, nearly monospecific stands on hot, barren ridges and

open slopes. Kalmiopsis fragrans, on the other hand, exhibits marked plasticity in growth form. Plants from

the few populations known to occupy exposed ridges may have a habit comparable to K. leachiana. However,

most individuals are found in forested sites, where they exist as dense, trailing mats on rock outcrops or

as loosely erect, openly branched shrubs in the more shaded areas. Plants range from a few dm to nearly

Meinke and Kaye, A new species of Kalmiopsis from Oregon 15

Taste 1. Selected morphological, geographic psis frag | K. leachiana

Trait Kalmiopsis fragrans Kalmiopsis leachiana

Habit Erect to trailing or matted Usually erect

Stem length To 12 dm, rarely up to 30 dm 2-4 dm, rarely up to 8 dm

Inflorescence size (2-)4-8(-12) flowers (5-)7-12(-15) flowers

Corolla size 16-28(-33) mm across (12-)14-20 mm across

Petal sculpting (best observed

on fresh flowers)

Petal sinuses

Corolla expansion

Corolla color

Style length (long-style)

Stamen length (long-style)

Anther size and apical pore

dimensions

Ovary color

Ridges connected within petals

Deeply cleft, to within 2-3 mm of the

floral tube

Becoming nearly flat and rotate as

petal lobes reflex

Light pink or reddish-purple

11-15 mm long

7-13 mm long

Narrowly oblong or curved linear,

1.2-3.0 mm, pore + round, «0.4 mm

Pale yellow to gold

Ridges connected between petals

Moderately cleft, to within 4-7 mm

of the floral tube

Petals reflexing but floral tube

remaining tubular-cupulate

Rose to deep pink

7-10 mm long

3-7(-9) mm long

Oblong, 0.7-1.8 mm long, pore

flared, 0.5-0.9 mm

Greenish gold

Floral pubescence Yellowish cilia densely tufted at base of

filaments surrounding ovary, evident in

floral tube (tube area rarely subglabrous)

Flowers completely glabrous, or

occasionally with fine cilia lining

basal interior of floral tube, but

not tufted or attached to filaments

Nectar pooling in floral tube and

pollen (flowers lacking pronounced

Floral pollinator attractant Spicy-sweet “azalea-like” scent and pollen

(nectar merely a trace or typically absent)

odor)

Slightly protandrous to slightly

protogynous, pollen shed about the

time of corolla expansion

Breeding system Clearly protogynous, pollen shed from a

few hours to a day after corolla expansion

Habitat Mostly on tuffaceous outcrops and within

shaded mesic coniferous forest, rooted in

rock crevices or in very shallow soil at the

base of cliffs or boulders

Plants preferring open sunny ridges,

in xeric shrub community or open

woodland, in shallow or more often

deeper soils on a range of substrates

(including ultramafics)

Endemic to the Klamath (Siskiyou)

Mountains at elevations ranging up

to 2100 m

Endemic to the southern Cascade

Mountains, at elevations ranging from

ca. 450-1325 m

Geographic distribution

3 m in length, far exceeding the largest examples of K. leachiana. Both species are capable of vegetative

propagation in nature. This trait is evidently more common in K. fragrans, where several populations ex-

ceeding 100 individuals are known that appear to consist of one or few clones, based on the presence of

only a single floral morph (i.e., long- or short-styled). However, at least two small populations of K. leachiana

(encompassing «300 m?) have been reported with only short-styled individuals (Marquis 1977). Moreover,

excavations of several apparently separate plants revealed a below ground connection of nearly 2 m in one

instance. Kalmiopsis leachiana is known to crown sprout after wildfires (Marquis 1977), and K. fragrans

reportedly may as well (Richard Helliwell, personal communication), although evidence for this was not

noted by the authors.

Floral Differences —Divergence between K. fragrans and K. leachiana is most apparent in the flowers. The

inflorescence of the new species is typically more floriferous (see Figs. 2A and 2B) and the corolla is larger

and more deeply cleft than in K. leachiana (Figs. 2C and 2D). The proportional differences between the spe-

cies are evident in corolla width and the size of floral organs, including the anthers, which are nearly twice

16 Journal of the Botanical R h Institute of Texas 1(1)

as long in K. fragrans but with much smaller terminal pores. The significance of this is unknown, but the

results may include a more prolonged release of pollen in K. fragrans (due to the larger anthers and smaller

pores)and the deposition of larger pollen loads on individual pollinators in K. leachiana. Another interesting

feature in K. fragrans is the copious pubescence (+ golden in nature but pale in dried specimens) often found

at the base of the filaments (Figs. 1, 2C), which forms a soft tuft around the base of the ovary. This is all but

lacking in K. leachiana, a notable exception being the type collection (Leach and Leach 2915, ORE), which

has some flowers with scattered cilia at the base of the filaments —see Fig. 4 in Rehder (1932). The flower

of K. fragrans is otherwise subglabrous to glabrous, while K. leachiana often exhibits a fine covering of cilia

inside the lower corolla tube.

Corollas of both species will remain open up to a week. In the greenhouse, K. leachiana flowers may be

slightly protandrous or slightly protogynous, with anthers dehiscing at or about the time of initial corolla

expansion. The flowers of K. fragrans have a more significant female phase, with anthers usually releasing

pollen from a few hours to over a day after buds begin to open. Expanded flowers of both species shift from

narrowly to broadly campanulate during the first day, with stamens initially splayed out along the corolla

away from the receptive stigma (Figs. 2C and 2D). After 1-2 days, stamens become erect and well-exserted

(Figs. 3A and 3B), with the filaments encircling the style. The corolla of K. leachiana retains a shallow, but

well-defined tube that is several mm deep throughout floral ontogeny, with the lobes eventually reflexing

perpendicular to the tube (Fig. 3B). The comparatively shallow petal sinuses of K. leachiana ensure that the

shape of the floral tube remains more or less unchanged even as the lobes reflex, and that it continues to

surround the ovary until the corolla drops. Corollas of K. fragrans have significantly deeper sinuses, and

as flowers age and petal lobes fully reflex the corolla becomes essentially flat and rotate (Fig. 3A), typically

exposing the ovary prior to corolla senescence.

Flowers of K. leachiana produce a rather viscous nectar (2-10 uL per flower per day in greenhouse

plants) and are essentially odorless. Flowers of K. fragrans are nectarless, or yield only minute traces, but

have a spicy sweet scent that increases in strength after anthers dehisce, and remains evident until corollas

are shed. The aroma is especially notable when cut branches are placed indoors, with buds allowed to bloom

and age in a confined area. The deeper floral tube of K. leachiana serves as a vessel for accumulating nectar,

whereas the dense basal ring of stamen hairs in K. fragrans may help to delay the dissipation of tiny amounts

of nectar or to trap fragrance compounds. Beeflies, syrphid flies, and bumblebees are common visitors to

flowers of both species, although foraging patterns differ depending on whether pollen or nectar is being

sought. Hummingbirds have also been noted making rapid visits to K. leachiana flowers, their foreheads

covered in pollen.

Stigma-Height Dimorphism.—Kalmiopsis has been described as heterostylous (Callan 1971; Marquis 1977;

Love 1991), although the evidence for true heterostyly is circumstantial. Kalmiopsis floral morphology (espe-

cially K. fragrans) is not typical of heterostylous species, which tend to have strongly tubular, campanulate,

or funnelform corollas (Ganders 1979). Distyly is otherwise unknown in the Ericaceae, with the possible

exception of Epigaea repens, a dioecious species that exhibits a continuum of long- and short-styled flowers

but lacks other evidence of heterostyly (Darwin 1877; Vuilleumier 1967; Clay and Ellstrand 1981).

While both K. leachiana and K. fragrans have distinct short- and long-styled floral morphs (Fig. 4), the

degree to which the two species also possess the other primary traits that define heterostyly (i.e., reciprocal

anther heights and diallelic self-incompatibility — see Ganders 1979) requires further evaluation. Barrett et

al. (2000) report that K. leachiana flowers clearly are dimorphic with respect to style length, though not for

stamens, providing evidence of a stigma-height dimorphism but not necessarily heterostyly (in the sense

of intramorph incompatibility). Observations of living and herbarium specimens during this study suggest

that reciprocal anther heights may occur in some populations of K. leachiana, but this was not confirmed.

Conversely, data for K. fragrans do show significant reciprocal differences between the floral morphs for

both stamen and style lengths (Fig. 5). Field observations of pollinators indicate that this high degree of

herkogamy apparently facilitates outcrossing in K. fragrans, but is the species functionally heterostylous?

Meinke and Kaye, A new species of Kalmiopsis from Oregon 17

Thrum anthers

16 4 Pin stigmas co e. ^ e "o

e e

à go 0 A’ e e 21 .

"P Pin anthers

Stigma and Anther Heights (mm)

10 -

8 4

6 - e e e?

«S5 oof %0

e Thrum stigmas

Fic. 5. Sy ial lati hip! | ] ig I ean forl J yl ] (pin) I sl yl I (tl fl in Kalmi f (N 41 for all

groups). D taken from 24 randomly selected ts, 12 for each floral h (estimated fl aee TUNE The mean stamen lengths

for pin (9. 46 mm an rum (14. a mm) flower (equivalent to panties au in Mae flowers) ignifi y different t (P<.0001, t=19.266,

SE=0.273 I gamy is a key

Efforts to further evaluate the floral biology of K. fragrans through a series of experimental intra- and

intermorph pollinations in wild populations were frustrated due to drought. Unseasonably warm, dry

weather coincided with sporadic fruit set and a high percentage of abortion among experimental flowers

and controls, suggesting that reproduction in our field trials may have been affected by factors other than

pollen compatibility. Flowers that were manually selfed, and then bagged, set small numbers of fruit which

contained no obvious seeds, but these results may have been similarly skewed by limited rainfall. However,

Marquis (1977) also reports evidence for self-incompatibility, albeit in a single greenhouse plant of K. fragrans.

Although fruit set occurred in 18 out of 28 undisturbed flowers, and in 34 out of 35 flowers self-pollinated

by hand, seed production was far below flowers of open-pollinated plants in nature (with an average of less

than one seed per capsule in the greenhouse versus 105 seeds from open-pollination). Finally, observations

during the current study indicate that fruit set may be depressed or absent within populations consisting

of only long- or short-styled ramets, suggesting that self-incompatibility may broadly affect seed production

within patches. This was also noted by Marquis (1977). Additional breeding system studies of Kalmiopsis

are planned.

CONSERVATION STATUS OF KALMIOPSIS

Rehder (1932) considered the collection and cultivation of K. leachiana essential to its preservation, feeling that

the beauty and apparent rarity of the species might eventually lead to its extinction. His concern was evidently

justified, as the conservation campaign he advocated quickly attracted the wrong sort of plant enthusiasts.

f*hAD o ID L

18 Journal of t titute of Texas 1(1)

Commercial collectors almost immediately began to decimate the new species (Love 1991; Kirkpatrick et al.

1994), and entire populations in the Klamath Mountains were wiped out in the 1930’s, with one collector

reportedly apprehended with 50,000 cuttings on pack horses. Another nurseryman in Portland attempted

to raise over 100,000 wild-collected plants for the retail trade (Kirkpatrick et al. 1994), but all died after a

single growing season in the wet climate of northern Oregon. Although unbridled commercial collecting

was once a legitimate threat to K. leachiana, the horticultural novelty of the species has declined and most

populations today are considered relatively secure. Moreover, the area in which K. leachiana grows is now

largely off limits to development due to federal wilderness designation. The species is the namesake for the

Kalmiopsis Wilderness Area, a region rich in endemic species and remarkable habitats (Love 1991).

There is no record of K. fragrans being mass collected in the wild, and its introduction into the alpine

and rock garden trade has evidently been less traumatic. Clones of the species identified as the *LePiniec"

form, or occasionally as the “Umpqua River” cultivar (Kruckeberg 1982; Love 1991), are still in cultivation

and available today. Recent research at Oregon State University has also demonstrated the relative ease with

which K. fragrans may be grown from seed (Kelly Amsberry, unpublished).

Of the two species, K. fragrans has the narrower geographic range, and it is significantly rarer than K.

leachiana. Most K. fragrans populations are not in designated protected areas, and the species is treated as

sensitive by the Umpqua National Forest (UNF), a status that affords it more security than most native plants

yet less than those formally listed as threatened or endangered. In a few areas the species benefits by sharing

its vertical environment with federally-managed peregrine falcons. While frequenting such habitats may

shield K. fragrans from the effects of timber harvest activity at selected sites, other areas appear less secure.

Forest Service staff report that the rocky habitat required by K. fragrans is not necessarily a deterrent to ei-

ther logging or road building, since surveys in advance of both activities have uncovered populations of the

new species (Richard Helliwell, personal communication). Fortunately, many populations reside within the

Limpy Rock Research Natural Area, which was designated with the preservation of its unique flora in mind.

In response to the on-going timber harvest program on the UNF, studies are presently underway to assess

how the removal of forest overstory along the edges of tuffaceous outcrops might impact K. fragrans.

Representative collections. Kalmiopsis fragrans. U.S.A. OREGON. Douglas Co.: Limpy Rock, Dog Cr. drainage off the N Umpqua

R., 19 Jun 1976, Chambers 4218 (ORE); same location, 17 May 1975, Chambers 4041 (OSC); Happy Camp, Umpqua R., Umpqua National

Forest, 25 May 1955, Wright s.n. (OSC); N Umpqua Hwy., steep hillsides above Horseshoe Bend, on basalt rocks and cliffs, 23 Apr 1967,

Williams s.n. (ORE). Kalmiopsis leachiana. U.S.A. OREGON. Josephine Co.: Panther Cr., T37S ROW S6, Siskiyou Mtns., 8 Jul 1950,

Whittaker s.n. (WS); Kalmiopsis Wilderness, Siskiyou National Forest, T37S R10W S1, 10 May 1997, Dennis 4838 (OSC); Illinois R., S side

of York Peak, 5 May 1957, Davis s.n. (OSC). Curry Co.: dry rocky spur of Horse Sign Butte, 8 mi S of Agnes, Applegate 7229 (OSC); Collier

Bar, T36S R11W S32, 1-2 May 1931, Leach 3180 (ORE, WTU, WILLU); the Big Craggy, 19 Jun 1938, Leach 5247 (OSC); near Game Lake

Peak, Siskiyou National Forest, 15 Aug 1938, Hanson s.n. (OSC); along Illinois R. Trail, just before the E Fork of York Cr., 24 Apr 1960,

Kezer & Faberge s.n. (OSC); Siskiyou National Forest, 1375 RIOW S34, 18 May 1939, Colville 1 (OSC); Horse Sign Butte, T36S R12W S24,

1 May 1931, Leach 3179 (OSC, WTU); higher Siskiyou Mtns. of Curry Co. (holotype), 14 Jun 1930, Leach 2915 (ORE).

ACKNOWLEDGMENTS

Matthew Carlson, Kelly Amsberry, Shannon Datwyler, Armand Rebischke, Nancy Fredricks, Sahni Burkhart,

and Steve Gisler offered valuable assistance during the field work. John Megahan provided the illustrations in

Figs. 2-4. The photograph (Fig. 1) of K. fragrans is by Melissa Carr, who also worked diligently at improving

the overall quality of the images in this paper. Kenton Chambers (OSC) graciously assisted with the Latin

diagnosis, and the Oregon State University herbarium collections and library facilities (including OSC,

ORE, and WILLU) were made available by Richard Halse. We thank USDA Forest Service staff, particularly

Richard Helliwell and Linda Mullins, for facilitating our collecting on the Umpqua and Siskiyou National

Forests. The Umpqua National Forest and the Oregon Department of Agriculture provided grant funding.

And thanks to Dan Luoma for suggesting the specific epithet.

Meinke and Kaye, A new species of Kalmiopsis from Oregon 19

REFERENCES

ABRAMS, L.R. 1951. Kalmiopsis. Illustrated flora of the Pacific States. Stanford University Press, Stanford, California.

Pp. 301-302.

BALDWIN, E.M. 1974. Eocene stratigraphy of southwestern Oregon. Oregon Department of Geology and Mineral

Industries, Bulletin 83.

Barrett, S.C.H., L.K. Jesson, and A.M. Baker. 2000. The evolution and function of stylar polymorphisms in flowering

plants. Ann. Bot. 85:253-260.

CALLAN, N.W. 1971. An investigation of the floral biology of Kalmiopsis leachiana (Hend.) Rehder. Unpublished

M.S. thesis. Southern Oregon College, Ashland.

CLAY, K. and N.C. ELLSTRAND. 1981. Stylar polymorphism in Epigaea repens, a dioecious species. Bull. Torr. Bot. Club

108:305-310.

CoPtLAND, H.F. 1943. A study, anatomical and taxonomic, of the genera of Rhododendroideae. Amer. Midl. Nat.

30:533-625.

COPELAND, H.F. 1954. Some details of the structure of Rhodothamnus chamaecistus. J. Arnold Arbor. 35:82-85.

DARWIN, C. 1877. The different forms of flowers on plants of the same species. John Murray, London.

Davis, PH. 1962. Rhodothamnus sessilifolius. Icones Plantarum 36: Tabula 3575.

GaNptRS, F.H. 1979. The biology of heterostyly. New Zealand J. Bot. 17:607-35.

HARBORNE, J.B. and P.A. Wituiams. 1973. A chemotaxonomic survey of flavanoids and single phenols in leaves of the

Ericaceae. Bot. J. Linn. Soc. 66:37-54.

HENDERSON, L.F. 1931. New plants from Oregon. Rhodora 34:203-206.

KIRKPATRICK, G., C. HOLZWARTH, and L. MuuLENs. 1994. The botanist and her muleskinner, pioneer botanists in the

Siskiyou Mountains. Leach Garden Friends, Portland, Oregon.

Kron, K.A. 1997. Phylogenetic relationships of Rhododendroideae (Ericaceae). Amer. J. Bot. 84:973-980.

KRON, K.A. and J.M. KiNc. 1996. Cladistic relationships of Kalmia, Leiophyllum, and Loiseleuria (Phyllodoceae, Erica-

ceae) based on rbcL and nrITS data. Syst. Bot. 21:17-29.

Kron, K.A., W.S. Jupp, P.F. Stevens, D.M. Crayn, A.A. ANDERBERG, PA. GADEK, C.J. QUINN, and J.L. Luteyn. 2002. Phylogenetic

classification of Ericaceae: molecular and morphological evidence. Bot. Rev. 68:335-423.

KRUCKEBERG, A.R. 1982. Gardening with native plants of the Pacific Northwest. University of Washington Press,

Seattle.

Love, R.M. 1991. The discovery and naming of Kalmiopsis leachiana and the establishment of the Kalmiopsis

Wilderness. Kalmiopsis (J. Native Pl. Soc. Oregon) 1:3-8.

Manaus, R.J. 1977. An investigation into the ecology and distribution of Kalmiopsis leachiana (Hend.) Rehder.

Unpublished M.A. thesis. Oregon State University, Corvallis.

MULLIGAN, B.O. 1973. Kalmiopsis leachiana. Quart. Bull. Alpine Gard. Soc. 41:131-134.

Peck, D.L., A.B. Grigas, H.G. ScHuckER, G. G. WELLS, and H.M. Dore. 1964. Geology of the central and northern parts of

the western Cascade Range in Oregon. U.S. Geological Survey Professional Paper 449.

Peck, M.E. 1961. A manual of the higher plants of Oregon. Binfords and Mort, Portland, Oregon.

RAMP, L. 1975. Geology and mineral resources of the Upper Chetco Drainage area, Oregon. Oregon Department

of Geology and Mineral Industries, Bulletin 88.

REHDER, A. 1932. Kalmiopsis, a new genus of Ericaceae from northwest America. J. Arnold Arbor. 13:30-34.

Stevens, PF. 1971. A classification of the Ericaceae: subfamily and tribes. Bot. J. Linn. Soc. 64:1-53.

VUILLEUMIER, B.S. 1967. The origin and evolutionary development of heterostyly in the angiosperms. Evolution

21:210-266.

WHITTAKER, R.H. 1960. Vegetation of the Siskiyou Mountains, Oregon and California. Ecol. Monogr. 30:279-338.

20 Journal of the Botanical R h Institute of Texas 1(1)

BOOK REVIEWS

RosERT H. MOHLENBROCK. Foreword by Mike Dompeck. 2006. This Land: A Guide to Western National

Forests. (ISBN 0-520-23967-9, pbk.). The University of California Press, Berkeley, CA 94704, U.S.A.

(Orders: California Princeton Fulfillment Services, 1445 Lower Ferry Road, Ewing, NJ 08618, U.S.A.

WWw.ucpress.edu, 609-883-1759, Fax 609-883-7413). $24.95, 391 pp., 63 color illustrations, 9 b/w

photographs, 71 maps, 19 figures, 6" x 9".

The Mohlenbrock family spent virtually all their | ploring U.S. national forests. Because he is a botanist, Robert H. Mohlenbrock

focused on special areas within the National Forest System, Research Natural Areas, which are "designated in perpetuity for research

and education, to maintain biological diversity on National Forest System lands..." In 1984 he began writing a monthly column in

Natural History magazine, published by the American Museum of Natural History in New York. He was asked to write in this column

about areas in the national forests which he found interesting; not always because they were pretty, rather because they had a biological

or geological story to tell. He called his column *This Land", and the theme is the same as that of his book, titled THIS LAND: A Guide

to Western Nationa

Forests.

In his Foreword, Mike Dombeck, Chief Emeritus of the U.S. Forest Service, gives the reader a sweeping history of actions taken

in the United States to honor and protect our forests:

—1864 Henry David Thoreau called for establishment of “national preserves" of virgin forests, “not for idle sport or food, but for in-

spiration and our own true re-creation."

—1864 President Abraham Lincoln signed legislation which granted Yosemite Valley and Mariposa Big Tree Grove to the care of the

state of California “for public use, resort, and recreation.”

—1891 Our first forest reserve, Yellowstone Park Timber Land Reservation, was created by President Benjamin Harrison.

—Toward the end of the nineteenth century, President Grover Cleveland established another 21 n acres of forest reserves.

fi : 1

established our first national national monuments

— President Theodore Roosevelt expanded our forest

moving the country toward an ethic of conservation. In 1905 The USDA Forest Service was established to manage the national forests

“for the greatest good for the greatest number for the long run."

The strength of this publication is its organization, which makes it easy to use by interested lay citizens as well as seasoned out-

doorsfolk and professional scientific sorts. The eight chapters focus on eight western states: Alaska, Arizona, California, Idaho, Nevada,

Oregon, Utah and Washington. Each chapter begins with a map of the state and its national forests; chapter subsections on those forests

(in alpha order) begin with a map of the national forest and it including cities, highways and ranger stations. Within

the text are descriptions of special characteristics of each of the forests - natural, political and recreational features. Mohlenbrock has

included many black and white botanical drawings throughout the book and an index of plant names. Also, there are a number of black

Pra | "t * 1 3! al

and white photos and sixty-two color plates.

Th 1 B 1 1 :

travels with him through these western forests.

practical ger nitens p terms specific to each area. Personal anecdotes add colorful interest points and contribute to the reader's sense of

sharing in h Karen Burkett, Herbarium Volunteer, Botanical Research Institute of Texas, Fort Worth, TX, 76102-4060, U.S.A.

He notes useful details, such numbers, trail names (and directions for finding them!), weather and terrain characteristics, and

ANTON Rajer. 2006. Museums, Zoos & Botanical Gardens of Wisconsin: A Comprehensive Guidebook.

(ISBN 0-9664180-0-X, pbk.). Fine Arts Publishing in cooperation with University of Wisconsin Press,

1930 Monroe Street, 3rd Floor, Madison, WI 53711-2059, U.S.A. (Orders: http://www.wisc.edu/wis-

consinpress/). $17.95, 304 pp., illustrated, maps, 6" x 9".

A lifetime of experiences have gone into the making of this guidebook. The author has grown up visiting the museums, zoos, and bo-

tanical gardens in his native state. He has described over 540 institutions is this fascinating, well-researched guidebook which covers

the full gamut of cultural and natural history institutions in Wisconsin.

The book is arranged geographically into four regions, and then by town in each region. A map accompanies each and locates

the cities in which an attraction is located. Each site is described and includes the street address, contact numbers, hours of operation,

admission fees, and collection highlights. There are indexes to help visitors find interesting sites by: institutional type, institution, and

by city. He also includes a listing of helpful contacts. The botanical garden entries are very complete and include web information and

specific opening hours, and days, where appropriate. Each entry has a detailed narrative paragraph, or two, about the site and the at-

tractions to be found there.

Well illustrated and arranged. Recommended reading for those who will be traveling in the Badger State.— Gary Jennings, Library,

Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A.

J. Bot. Res. Inst. Texas 1(1): 20. 2007

TAXONOMIC OVERVIEW OF THE

HETEROTHECA FUECRATACOMEPLEXCASTERACPAE ASTERI LAF)

Guy L. Nesom

Botanical Research Institute of Texas

509 Pecan Street

Fort Worth, Texas 76102-4060, U.S.A.

ABSTRACT

dore fulcrata (Greene) Shinners has recently been treated as ME: iu iun) sympatric varieties. These plants are viewed

1 J;

gin part to the varietal taxa: (1) H

g H. fulcrata var. senilis and H. fulcrata vars.

fulcrata and pen lifolia in pn sensu Semple), (2) Heterotheca foliosa (Nutt.) S pis M H. fulcrata vars. fulcrata and amplifolia in

part, sensu Semple), (3) Heterotheca arizonica TE Nesom, comb. et stat. nov. (= H. fulcrata var. arizonica), and (4) Heterotheca

nitidula (Woot. & Standl.) Nesom, comb. nov., end to the Mogollon Mountains of southwestern New Mexico and White Mountains

of immediately adjacent Arizona, previously treated as a synonym primarily of H. fulcrata var. amplifolia. County-level distribution maps

for these taxa are provided (based on collections examined in the current study). Heterotheca viscida (Gray) Harms is included in the

consideration because it is pa: in (GEOETdDbi ie to H. arizonica and oc Nd Dd confused with that taxon and expressions of H.

fulcrata ly as previously treated by Semple, but it is here observed

to occur more widely in New ME and to nr into trans- Pecos Texas.

RESUMEN

TT T ro NG T m | = 4 J A T hi J le

Estas

J JE P E

plantas son consideradas aquí como cuatro especies, correspondientes en parte a los taxa varietales: (1) Heterotheca fulcrata (incluyendo

H. fulcrata var. senilis y H. fulcrata vars. fulcrata y amplifolia en parte, sensu Semple), (2) Heterotheca foliosa (Nutt.) Shinners (incluyendo H.

fulcrata vars. fulcrata y amplifolia en parte, sensu Semple), (3) Heterotheca arizonica (Semple) Nesom, comb. et stat. nov. (= H. fulcrata

var. arizonica), y (4) Heterotheca nitidula (Woot. & Standl.) Nesom, comb. nov., endémica de las Mogollon Mountains del suroeste de

Nuevo México y White Mountains en la adyacente Arizona, previamente tratada como sinónimo de H. fulcrata var. amplifolia. Se aportan

A t

las colecciones examinadas en el presente i Heterotheca rd

(Gray) Harms está incluida en las S eg es similar en rango geografi H. arizonica y

Li pt.

ella y ejemplares de H. fulcrata una especie distinta tal como fue tratada previamente por

Semple, pero aquí se ha observado que aparece M en Nuevo México y se extiende por los trans-Pecos en Texas.

Heterotheca fulcrata (Greene) Shinners has been treated by Semple (1996, 2006) as comprising four strongly

sympatric varieties. My approach to the taxonomy of Heterotheca (Nesom 1997, 2006) is different from that

of Semple and the current study documents a different taxonomic interpretation of the H. fulcrata group.

Heterotheca fulcrata sensu Semple is treated here as four separate species; H. viscida (A. Gray) Harms is included

in the consideration because it is similar in geographic range and occasionally confused with expressions of

H. fulcrata. Heterotheca zionensis Semple also is included, though not closely related to H. fulcrata, to docu-

ment an increased understanding of its identity.

The current study is based primarily on study of collections from ASU, BRIT-SMU, NMC, MO, SJNM,

SRSC, and TEX-LL. Distribution maps are based on specimens examined.

KEY TO THE SPECIES OP THE HETEROTHECA FULCRATA GROUP AND E: VISCIDA

1. Leaves 5-15(-20) mm long, 2-5(-7) mm wide, with a thick-indurate, sharp-pointed, often recurving terminal

mucro Heterotheca arizonica

1. Leaves 15-40 mm long, 3-20(-25) mm wide, without a thick-indurate terminal mucro.

2. Heads not immediately subtended by foliar bracts; glands usually stipitate Heterotheca viscida

2. Heads immediately subtended by foliar bracts; glands present or absent, usually sessile if present.

3. Plants eglandular; nonglandular hairs very thin, with all cells equal in width, appressed, stem hai

appressed to closely ascending He or nitidula

J. Bot. Res. Inst. Texas 1(1): 21 — 30. 2007

£s+haD H ID L

22 Journal of t titute of Texas 1(1)

3. Plants usually glandular; sometimes sparsely so; nonglandular hairs relatively thicker, with expanded

basal cells, soreading or arching erect, stem hairs spreading or often deflexed.

4. Midstem cauline leaves (10-)15-30 mm long, 3—7(-10) mm wide, mostly oblong to oblanceolate-oblong

or lanceolate-oblong Heterotheca fulcrata

4. Midstem cauline leaves 25-60 mm long, (5-)10-18 mm wide, mostly oblanceolate-obovate

Heterotheca foliosa

Heterotheca fulcrata sensu stricto

Heterotheca fulcrata (Greene) Shinners, as treated here, is recognized by its heads immediately subtended by

foliar bracts, the bracts prominently ciliate with coarse, spreading hairs. Stems are densely hirsute-villous.

The range of the species extends from north-central Mexico into Texas, southern New Mexico, Arizona,

and Utah (Fig. 1), exclusive of the population system in Colorado and Wyoming mapped by Semple as H.

fulcrata. The latter are treated here as H. foliosa (Nutt.) Shinners (see below).

From typical Heterotheca fulcrata, Semple differentiated H. fulcrata var. senilis (Woot. & Standl.) Semple

by *distal margins of uppermost leaves with numerous long hispid-strigose hairs" [var. senilis] vs. "leaf

margins of upper leaves lacking long hairs or [with] only a few basally" [var. fulcrata] (1996, p. 28, key

couplet 17). The types of both taxa were collected in the Organ Mountains of Dona Aña Co., New Mexico,

and variability within the species accounts for the difference in vestiture. 'Senilis-like plants are the com-

mon form in Arizona, New Mexico, Texas, and Mexico; bract and leaf margins become less coarsely and

densely ciliate northward, but there does not appear to be a discontinuity. Some plants of H. fulcrata from

this area show reduced nonglandular vestiture on the leaf faces (e.g., the type of Chrysopsis cryptocephala,

below), while others produce denser nonglandular vestiture. Analogous variation in density of nonglandular

vestiture occurs in Colorado and Wyoming populations of H. foliosa (comments below). Plants identified as

H. fulcrata from northern Utah (i.e., Cache, Duchesne, Salt Lake, Summit, Utah, Wasatch counties) perhaps

represent a distinct, separately evolved population system; compared to typical, southern populations of

H. fulcrata, these plants occur at significantly higher elevations and consistently are more glandular with

reduced nonglandular vestiture.

Heterotheca fulcrata (Greene) Shinners, Field & Lab. 29:71. 1951. Chrysopsis fulcrata Greene, Bull. Torrey Bot. Club

25:119. 1898. Type: U.S.A. NEW MEXICO. LINcoLNn Co.: White Mts., near Cherokee Bills Spring, 6300 ft, 21 Aug 1897, E.O. Wooton

511 (Lectotype, Shinners 1951: ND-G; isoLecToTYPES: NY, NY [internet image!], US [internet image!])

From among ND-G syntypes Wooton 510, 511, and 512, Shinners (1951) selected 511 as the lectotype. Harms (1968a, p. 17) entered

a caveat, noting that "This choice causes some confusion since Wooton and Standley (1913, 1915) obviously considered the first cited

specimen, Wooton 510 from the Organ Mountains, as the true type of C. fulcrata, and tl

descriptions of C. cryptocephala

a, but

there was never an explicit or formal statement by Wooton and Standley regarding lectotypification of C. fulcrata. Semple (1987)

and C. senilis seem based upon this understanding.” Wooton 512 was cited as a paratype in the protologue of C. cryptocepha

stated that “I consider all three Wooton syntype collections to be members of [Heterotheca fulcrata] var. fulcrata and accept Shinners'

lectotype designation, Wooton 511 (ND-G).” Semples 1987 position is affirmed here and Shinners’ choice appears to remain valid,

notwithstanding Semple’ later citation (1996, pp. 68, 70) and 1995 and 1997 annotations (e.g., NY, US) of 510 as lectotype, in

which he apparently accepted Harms' implication that the original lectotypification was made implicitly by Wooton and Standley.

Chrysopsis senilis Woot. & Standl., Contr. U.S. Natl. Herb. 16:179. 1913. Heterotheca fulcrata (Greene) Shinners var. senilis (Woot. &

Standl.) Semple, Brittonia 39:380. 1987. Tyre: U.S.A. New Mexico. Dona Ana Co.: Organ Mts., 4800 ft, 1 Sep 1897, E.O. Wooton 509

(HoLoTYPE: US [internet image!]; isotypes: MIN, MO!, NDG, NY [internet image!]).

Chrysopsis cryptocephala Woot. & Standl., Contr. U.S. Natl. Herb. 16:179. 1913. Type: U.S.A. New Mexico. [Lincoln Co.]: White Mts., V

Pasture, sect. 23, 23 Jul 1905, E.O. Wooton s.n. (HoLoTYPE: US [internet image!]; Isotype: US).

Status of Chrysopsis nitidula

Populations recently identified mostly as Heterotheca fulcrata in the Mogollon Mountains of southwestern

New Mexico and White Mountains of immediately adjacent Arizona (Fig. 1) are recognized here as a separate

species, originally described as Chrysopsis nitidula Woot. & Standl. The protologue noted that the leaf surfaces

are "finely sericeous, the leaf as a whole appearing green and remarkably soft and smooth; ... This is very

unlike any of our other species, being strongly marked by its peculiar pubescence and long rays [10-12 mm

long].” Compared to H. fulcrata, the leaves of H. nitidula also tend to be narrow and elongate with acute apices.

Ll Llat +h

Nesom, Taxonomic overview of the

O Heterotheca fulcrata

e e

o Heterotheca foliosa

A Heterotheca nitidula

^ 8

e e

á —

e! © ONERE a0

e Se

e e ¡e - =

O E -

: ° m

$ a e E

À o [

| = _ o e i 9 @

@ ve e —

ei?

| e ° e -

2 : |. |

3 a

1 | a

= 10)

os Q |

E 0—/|A UNE NE EE

A (0)

B mm

PN RD [e] O

(0)

O O o O

[e] 7

zd 9

| al Oo

X "HO o O -l

Fic. 1. Geographi f Hotornth ful , H. foliosa, and H. nitidula. See text f itations for H. nitidula.

f4L,D o ID L

24 Journal of t titute of Texas 1(1)

Heterotheca nitidula (Woot. & Standl.) Nesom, comb. nov. Chrysopsis nitidula Woot. & Standl., Contr. U.S. Natl. Herb.

16:179. 1913. Tyre. NEW MEXICO. “Socorro Co.:” Mogollon Mts., on or near the West Fork of the Gila River, 7500 ft, 20 Aug

1903, O.B. Metcalfe 552 (HoLotyee: US [internet image!]; isotypes: GH, MO!, NMC!, RM). Redefined county boundaries now place

the type locality in Catron County.

Diagnostic description of Heterotheca nitidula: Stems 10-35 cm tall, arching upward from the base, antrorsely

strigillose. Cauline leaves mostly narrowly oblanceolate, 3-5.5 cm long, 3-8 mm wide, attentuate to the base,

apices acute. Heads solitary or usually in clusters of 2-6, usually immediately subtended by lanceolate foliar

bracts. Stems, leaves, bracts, and phyllaries eglandular, sparsely to moderate strigillose with nonglandular

vestiture of extremely thin, slightly flexuous, closely appressed hairs 0.5-1.5 mm long, without enlarged

basal cells; leaves eciliate or with a few weak, spreading cilia along the proximal margins; foliar bracts eciliate

to ciliate. Heads on some plants of the type collection are pedunculate, lacking foliar bracts, but bracteate

heads are the normal condition for the species, hence the association with H. fulcrata.

Numerous collections of Heterotheca nitidula have been made since its original description—all are

consistent in morphology and all occur within a radius of 35-40 miles. Its geographic range apparently lies

within that of H. fulcrata (Fig. 1), but no collections of typical H. fulcrata have been examined from within

the range boundary of H. nitidula. Heterotheca nitidula occurs at elevations of (7000—)7400-9200 feet; H.

fulcrata occurs at 4600—6900(-8000) feet in Texas, 2100—7200 feet in New Mexico, and 4900-7100(-8700)

feet in Arizona. Plants of H. nitidula grow on open rocky slopes and flower Aug-Sep(-Oct). A diploid chro-

mosome number (2n = 18) has been documented for H. nitidula by Semple (see citation for Semple & Heard

8021, below) and Turner (Turner 5700).

The holotype of Chrysopsis nitidula was annotated by Semple in 1985 as Heterotheca villosa var. pedun-

culata. In 1995, he re-annotated the specimen as H. fulcrata var. amplifolia, and later (1996, 2006) he cited

C. nitidula as a synonym of the latter. NMC specimens of H. nitidula were annotated by Semple as H. fulcrata

var. fulcrata and as H. fulcrata var. amplifolia. Similarly, Semple (1996) cited *Rusby 168 (MIN, NY(3), PH)"

under three different taxa: H. fulcrata aff. var. fulcrata (p. 74), H. fulcrata var. amplifolia (p. 76), and H. villosa

aff. var. pedunculata (p. 126). A duplicate of the same collection (Rusby 168, MO!) is identified here as H.

nitidula (as cited below).

Additional collecti ined: ARIZONA. Apache Co.: Apache Natl. Forest, Mount Baldy area, W Fork Little Colorado River, parking

area off State Rd 273, locally abundant in dry grassland, 2787 m, 21 Sep 1998, Brant & Stone 4128 (MO, TEX); White Mts., 6-15 Aug

1903, Griffiths 53329 (MO); along Forest Rd 275 in Stone Creek drainage, above stream in gravelly soil, 8000 ft, 11 Aug 1998, Hammond

11478 (TEX); White Mts., dry road shoulders, 8500 ft, 15 Aug 1973, Moldenke 27846 (LL); near Greer, 8300 ft, 19 Aug 1935, Peebles 12555

(LL); E edge of Alpine on US 180, road embankment, red shale substrate, limber pine forest, 8030 ft, 19 Sep 1985, Semple & Heard 8021

[voucher 2n = 18] (BRIT); 12 mi N of Alpine, common in roadside cuts, 13 Aug 1967, Turner 5700 [voucher 2n = 18] (TEX); Sitgreaves

Natl. Forest, 5.4 mi S of jct US 666 and US 180 on 666, E of road in meadow, 15 Aug 1978, Warnock 1677 (TEX). Greenlee Co.: Apache

Natl. Forest, 10 mi E of Big Lake on Forest Rd 249 and 8 mi from Alpine, near turnoff to Sierra Blanca and on hillside above lake in

rocky soil, open areas in spruce forest, 8900 ft, 16 Aug 1972, Hess 2937 (SMU). NEW MEXICO. Catron Co.: Mogollon Mts., Gila Natl.

Forest, Indian Creek drainage and Bear Wallow Mt., common perennial on E-facing slope and drier areas, 1 ft tall, 9200 ft, 4 Sep 1968,

Hess 2394 (NMC, SMU); Mogollon Mts., 1.4 road mi N of Gilita Campground, ponderosa pine, ca. 8100 ft, 7 Sep 1978, Moir & Fitzhugh

s.n. (NMC); Mogollon Mts., high rocky summits, 7 Sep 1881, Rusby 168 (MO); Gila Natl. Forest, 13 mi straight line NNE of Beaverhead

Ranger Station, near NM 163 and FS 150, montane grassland and pinon-juniper woodland, SW-facing slope, sandy, rocky ashflow, 7500

ft, 3 Oct 1995, Williams 2600 (NMC). *Socorro Co.:" Middle Fork of Gila, 7500 ft, 5 Aug 1900, Wooton s.n. (NMC). Sierra Co.: Taylor

Creek, ca. 14 air mi E of Beaverhead, rocky S-facing slope with ponderosa pine, 7400 ft, 14 Aug 1982, Spellenberg et al. 6598 (NMC).

Identity of Heterotheca foliosa

Harrington (1954) divided Colorado Heterotheca with prominent foliar bracts subtending the heads between

H. fulcrata (“involucres sparingly to definitely glandular”) and H. foliosa (Nutt.) Shinners in part (“involucres

pubescent but the glands obscure or none"). Wyoming plants were similarly identified in Dorn's key (2001).

Weber and Wittmann (1990) identified these plants primarily as H. fulcrata but later (1996) called them H.

foliosa, noting that the name H. fulcrata would be incorrectly applied.

Semple (1996, 2006) treated most of the same plants within Heterotheca fulcrata—some as var. fulcrata

(“sparsely to moderately glandular”) and some as var. amplifolia Cglandless or nearly so"). Regarding distinc-

£41 Lint 4L TB P ORPANAN A

Nesom, Taxonomic overview of the plex 25

tive features of H. foliosa (treated by him as H. villosa var. foliosa), he noted (1996, p. 117-118) that its “upper

stem leaves ... are oblong ... and its heads ... are often subtended by narrow oblanceolate bracts.” His key

couplet 15 (1996, pp. 28-29) separated H. fulcrata from H. villosa with foliar-bracteate heads by the shape

of the bracts—H. fulcrata with ovate-lanceolate bracts, H. villosa var. foliosa with oblanceolate to linear-ob-

lanceolate bracts. Even as identified and annotated by Semple, however, variation in bract shape exists in

plants that otherwise are clearly referable to the H. fulcrata var. fulcrata-amplifolia forms.

Among the plants in Colorado and southern Wyoming with bracteate heads, there is a tendency for

sessile glands to be abundantly developed on all parts and for nonglandular hairs to be reduced in density.

Such green-glandular plants are typified by Chrysopsis resinolens A. Nels. (see below). At the other extreme,

leaves and stems are eglandular or nearly so and distinctly grayish-strigose with denser nonglandular vestiture

(e.g., Chrysopsis amplifolia Rydb.). Intermediates in vestiture are common (as noted also by Semple 2006);

glands may be absent on the leaves but present on the foliar bracts. If two taxa are distinguished on the

basis of vestiture, the resulting two are similar in variability of leaf shape and size and in variability of size

and shape of foliar bracts. Further, they are ecologically similar and their geographic ranges are congruous

(see Semple 1996, Fig. 28, A and B) and disjunct from other taxa of the Heterotheca fulcrata complex (Figs. 1,

2). Their treatment here as a single taxon, Heterotheca foliosa, emphasizes the (1) geographic and ecological

coherence of the extended population system and its disjunction from related ones, and (2) intergradation in

vestiture and apparent impossibility of identifying more than a single entity without relying on a typological

concept.

The geographical coherence of Heterotheca foliosa in Colorado and Wyoming, also as recognized by

Semple (1996) in his identification of H. fulcrata in that region, substantiates its evolutionary coherence.

As identified here, the range of H. foliosa extends into southwestern Montana (Fig. 1), slightly further north

and west than mapped by Semple for H. fulcrata in the Rocky Mountains. Semple indicated the geographic

range of H. villosa var. foliosa to continue southward (into New Mexico) and much further north and west

(into Idaho, Oregon, Washington, and Canada), but as in my previous study (Nesom 2006), most plants

identified and mapped by Semple as H. villosa var. foliosa are regarded here as variants within H. villosa var.

villosa.

The morphological distinction of Heterotheca foliosa from H. fulcrata sensu stricto is not great, but they

differ conspicuously in leaf size, as noted in the key above. Further, as treated here, the two are geographi-

cally disjunct and appear to have different ecological tendencies. Closely similar but allopatric population

systems are often treated as conspecific, but recognition of both taxa at specific rank also is justified, here

emphasizing the differentiation and disjunction.

Heterotheca foliosa (Nutt.) Shinners, Field & Lab. 29:71. 1951. Heterotheca villosa var. foliosa (Nutt.) Harms, Wrightia

4:15. 1968. Chrysopsis villosa var. foliosa (Nutt.) Cronq., Bull. Torrey Bot. Club 74:150. 1947. Chrysopsis foliosa Nutt., Trans. Amer.

Philos. Soc. 2, 7:316. 1841. Tree: U.S.A. [Wyoming.] “In the Rocky Mountain plains, near the banks of the Platte. Flowering in

August” [protologue], [Jun 1834], T. Nuttall s.n. (HoLoTYPE: K; isotypes: GH 2 sheets). Nuttall, with Wyeth’s expedition, crossed the

North Platte River in southern Wyoming in June 1834, probably in Carbon County. If the collection of Chrysopsis foliosa were made in

June, the nature of his reference to “Flowering in August” is not clear, but perhaps he observed that full flowering would be reached

by that month. Nuttall’s description of C. foliosa, especially features of the leaves, clearly aligns it with the present concept of the

taxon: “About foot high, sending up many hairy stems from the same root. Nearly allied to C. villosa but far more pubescent and

hoary, with the leaves widest at the base. ... sericeously villous, and more or less canescent ... flowers fastigiate, corymbose; leaves

entire, oblong or oblong-ovate, subamplexicaule ... .”

Chrysopsis resinolens A. Nels., Bull. Torrey Bot. Club 28:232. 1901. Tyre: U.S.A. Wvowiwc. Albany Co.: Moist mountain valleys, Laramie

Peak, 13 Jul 1900, A. Nelson 7583 (HOLOTYPE: RM; isotypes: GH, MIN, MO!, NY 2 sheets [internet images!], RM, US [internet image!])

The protologue notes for the habitat and locality: “open slopes in the foothills of Laramie Peak.” A handwritten label (NY 163187)

notes ‘ ae iai slopes, Laramie Peak,” while printed labels (NY 163216, RM, US) note “Moist mountain valleys, Laramie Peak.”

225

n book has “In the open valleys.” Other label information is identical among the specimens. The plants are similar

in morphology and plausibly from the same gatherin,

ing.

Chrysopsis resinolens var. ciliata A. Nels., Bull. Torrey Bot. Club 28:233. 1901. Chrysopsis viscida var. ciliata (A. Nels.) Blake in Tidestrom,

Contr. U.S. Natl. Herb. 25:537. 1925. Type: U.S.A. WyominG. Albany Co.: sandy river bottoms, Dunns Ranch, 16 Jul 1900, A. Nelson

Journal of the Botanical R h Institute of Texas 1(1)

T rJ

M, NN

=e O (Y Heterotheca arizonica

Y A Heterotheca viscida a 1 |

qe | 582 £ Lis 4L x op I ila 4i a » a UI Eill ol e I IJ LE

Fic. 2

A "i

7566 (HOLOTYPE: RM; Isotype: NY [internet image!]). “Nelson 7560" is the coll number cited in the protologue, and the NY isotype

is labeled as “7560 .... n. var.” on a label (ex Rocky Mountain Herbarium) apparently handwritten by Nelson. The RM specimen,

however, is labeled 7566 and Nelsons original notes state that 7566 = Chrysopsis resinolens var. ciliata; Nelson 7560 is listed as a moss

(Mnium serratum). Thus the collection number in the protologue as well as the label on the NY collection are interpreted as resulting

from errors in transcription.

Chrysopsis amplifolia Rydb., Bull. Torrey Bot. Club 31:648. 1904. Chrysopsis foliosa var. amplifolia (Rydb.) A. Nels. in Coult. & Nels.,

Man. Bot. Rocky Mts. 5493. 1909. Heterotheca fulcrata var. amplifolia (Rydb.) Semple, Univ. Waterloo Biol. Ser. 37:74. 1996. Tyre:

U.S.A. Cotonapo. [Boulder Co.:] Plains and foothills near Boulder, Longmont, Jul 1902, E Tweedy 4898 (HoLotyre: NY [internet

image!]; IsoTyPE: RM).

Chr

po udata Rydb., Bull. Torrey Bot. Club 31:648. 1904. Type: U.S.A. COLORADO. [El Paso Co.: Pikes Peak,] Ruxton Dell, 2950 m, 2

Aug 1901, FE. & E.S. Clements 143 (HoLotyre: NY [internet image!]; isorvres: DH, GH, MIN, MO!, RM, US [internet image!]).

Chrysopsis imbricata A. Nels., Bot. Gaz. 37:263. 1904. Chrysopsis foliosa var. imbricata (A. Nels.) A. Nels. in Coult. & Nels., Man. Bot. Rocky

Mts. 493. 1909. Type: U.S.A. COLORADO. El Paso or Teller Co.: Pike’s Peak, open slopes, 1 Sep 1901, A. Nelson 8616 (HOLOTYPE: RM).

Chrysopsis alpicola var. glomerata A. Nels., Bot. Gaz. 40:64. 1905. Type: U.S.A. COLORADO. Larimer Co.: Estes Park, 9000 ft, Aug 1904,

WS. Cooper 174 (HOLOTYPE: RM).

Chrysopsis butleri Rydb., Bull. Torrey Bot. Club 37:129. 1910. Type: U.S.A. Montana. Gallatin Co.: Gateway, 17 Aug 1908, B.T. Butler 620

(HOLOTYPE: NY [internet image!])

Heterotheca fulcrata var. arizonica at specific rank

Semple’s description of Heterotheca ful

crata var. arizonica Semple was a useful advance in resolving problems

of identification in the H. fulcrata group. As mapped by Semple (1996), however, and as confirmed here,

var. arizonica occurs sympatrically with typical H. fulcrata (including var. senilis) in Texas, New Mexico, and

Ll E PES 4L £l + PA PR 27

Nesom, Taxonomic overview of the

xix E MS

A^ p

ECN 2 e A É

~ e |

e A

! A^ A

E E A A j

o e O O O

e OLO

j A

% e A @ i *

e o a A y

| e o ARI A a^

. o MES A ERA:

e A A a

AAA a

e AA o 2? |

A| |e a

» e 7 o. |@ [e YI^ J^ Mi^;

e A\@Al@\a| |@<@

Alo [x A ala?

O @|@

+ O Heterotheca zionensis > a »

@ @

A Heterotheca villosa var. angustifolia e J @

N O @ Heterotheca canescens 9

A se |

Fic 2 FU g I | Da g £Liln« L . > D I 4 4°, | i £ ij villoca var PIT A O I L F l g Po aL J

to South Dakota. The only collection of H. in New Mexi in this study i pped: Union Co., Clayton, 24 Sep 1907, Evans s.n. (NMC); an

to the southeast in Texas. Some of the localities (open circles) for H. inK Ided from Great Plains Flora Association (1976)

southern Arizona (Fig. 2). Intermediates, if they occur at all, are not common. Biologically, var. arizonica fits

criteria for recognition of a taxon at specific rank, and it is so treated here.

Heterotheca arizonica (Semple) Nesom, comb. et stat. nov. Heterotheca fulcrata (Greene) Shinners var. arizonica Semple,

Brittonia 39:380. 1987. Tyre: U.S.A. Arizona. Gila Co.: 0.3 km NW of East Verde River, AZ Hwy 87, NW of Payson, 13 Sep 1985,

J.C. Semple and S. Heard 7923 (HOoLOTYPE: NY; isotypes: ASU, MO, US, WAT). The herbaria cited are from the protologue; specimens

apparently have not been deposited at ASU, MO, NY, or US. The concept of the taxon is from annotations by Semple on TEX-LL

specimens.

Heterotheca arizonica is characterized as follows: (1) stems and leaves densely and prominently sessile-glan-

dular, essentially green-colored because of the sparsely to moderately hirsutulous nonglandular vestiture;

(2) cauline leaves elliptic to elliptic-oblanceolate or elliptic-lanceolate, 5-15 mm long, usually relatively

even-sized, spreading or ascending, often with only the midvein visible and distinctly thickened and raised

on abaxial surface, with a thick-indurate, sharp-pointed, often recurving terminal mucro; (3) heads often

(sometimes not) immediately subtended by small, linear to oblanceolate foliar bracts; and (4) phyllaries

glabrous to sparsely minutely glandular to sparsely strigose.

Status of Heterotheca viscida

Heterotheca viscida (A. Gray) Harms (Fig. 3) has been identified with relative consistency. It is included in

£s+haD o ID

28 Journal of t h Institute of Texas 1(1)

the consideration here because of its similarity in geographic range to H. arizonica and its occasional confu-

sion with that taxon and expressions of H. fulcrata. Heterotheca viscida is characterized by its conspicuously

glandular (usually stipitate-glandular) vestiture; because of the istic lack or paucity of nonglandular

hairs, stems and leaves are generally distinctly green. Leaves and stems of some collections from Jeff Davis

Co., Texas, are minutely and inconspicuously glandular but prominently villous; H. viscida of typical vestiture

also is common in the area and intermediates occur. Leaves are oblong-obovate to obovate-oblong, sessile

and sometimes subclasping, 15-40 mm long and (7-)10-20(-25) mm wide. Heads are on peduncles without

immediately subtending foliar bracts. Plants commonly grow on cliff faces and ledges and in crevices. Semple

(1996) mapped Heterotheca viscida in Pima Co., Arizona, but did not cite specimens from there.

Heterotheca viscida (A. Gray) Harms, Rhodora 70:302. 1968. Chrysopsis villosa (Pursh) Nuttall var. viscida A. Gray, Synopt.

Fl. N. Amer. 1(2):123. 1884. Chrysopsis viscida (A. Gray) Greene, Erythea 2:96. 1894. Type: U.S.A. ARIZONA. Santa Cruz Co.: Santa

Rita Mts., clefts of dry ledges, 7500 ft, 28 May 1881, C.G. Pringle s.n. Lectotype, Harms 1968b: GH!; isoLecTOTYPES: NY 3 sheets

internet images!], PH).

Status of Heterotheca zionensis

In an earlier study (Nesom 2006) I noted that Heterotheca zionensis Semple was difficult to distinguish from

H. villosa var. pedunculata (Greene) Semple, based on criteria provided by Semple. With further study and

perspective, H. zionensis is accepted here as a distinct species, essentially as previously described by Semple

(1996), but occurring more widely in New Mexico and extending into trans-Pecos Texas (Fig. 4). The ap-

parently disjunct plants in northwestern Colorado (e.g., Moffatt Co.: along Hwy 40, 30 Aug 1930, Baker

4833b, LL; Pat's Hole, near the confluence of Green and Yampa rivers, 8 Jul 1945, Porter 3635, SMU) also

were recorded by Semple (1996). A single collection from Idaho has been recorded in this study: Bear Lake

Co.: Bear Lake, 6 Aug 1898, Mulford 327 (MO).

Plants of Heterotheca zionensis are distinctive in their relatively large stature— stems strictly erect from

the base, commonly 4-10 dm tall (grazed or damaged plants may be smaller), up to 2-4 mm thick in the

proximal portions, and often 10—20 stems per clump—and in their silvery to silver-gray, sericeous to densely

strigose vestiture of thin-based, closely appressed nonglandular hairs. Cauline leaves are oblanceolate-ob-

ovate, spreading to ascending, and relatively even-sized up the stem; basal leaves are absent by flowering.

Capitula are pedunculate, and often numerous in a subcorymboid arrangement. Cauline vestiture is an-

trorsely appressed to ascending, rarely spreading. Semple (1987, p. 385) noted that *Non-glandular forms

of H. zionensis occur in north-central Utah; glandular and non-glandular forms occur in the Utah-Arizona

border region.” In New Mexico and Texas they are mostly non-glandular but glandular plants also have been

collected, especially near the range of H. villosa var. angustifolia.

The type of Heterotheca villosa var. pedunculata (isotypes: MO!, NMC!) was collected in Archeluta Co.,

Colorado, outside the geographic range of H. zionensis. It is similar in habit to many other plants of the

regional form of H. villosa (var. minor, as identified by Semple), but it is at the denser extreme of a variable

range in nonglandular vestiture density in the species, which gives it a silvery-gray aspect. The leaves of H.

villosa usually are basally narrowed to a petiole-like region and gradually diminished in size up the stem.

New Mexico plants treated here as Heterotheca zionensis apparently have been identified mostly as H.

villosa var. pedunculata by Semple (1996, and by annotation). Those in trans-Pecos Texas were referred by

him to a disjunct population system of H. canescens (DC.) Shinners and to H. villosa var. angustifolia. In 2003

I also identified and annotated these trans-Pecos plants as the “trans-Pecos form" of H. canescens, but later

(Nesom 2006, as mapped in Fig. 1) I included these as part of the range of H. villosa var. angustifolia. In the

current study, the ranges of Heterotheca zionensis and H. villosa var. angustifolia are recognized to overlap in

the trans-Pecos region and in central New Mexico (Fig. 4), where they sometimes are closely similar in habit

and aspect and perhaps hybridize. Warnock 6230 from Reeves Co., Texas (cited below), is densely sericeous

but like var. angustifolia, the plants produce some axillary leaves and are slightly glandular beneath the

nonglandular vestiture—they may be of hybrid origin. Some collections from San Miguel Co. and Santa Fe

Co., New Mexico, also suggest the occurrence of gene flow.

Ll Llas 4L £l + PA PERE

Nesom, Taxonomic overview of the p 29

Some New Mexico collections of Heterotheca villosa var. angustifolia were identified as H. canescens by

Semple, some as H. villosa var. pedunculata (1996, and by NMC annotation).

The following key distinguishes these taxa as they occur in Texas and New Mexico.

aay

. Heads pedunculate, without closely subtending leaves or bracts; axillary clusters of small leaves usu-

ally not produced along stems; leaves usually without sessile glands beneath the nonglandular hairs

Heterotheca zionensis

. Heads on leafy stems, often with immediately subtending, narrowly lanceolate foliar bracts; numerous axil-

lary cluster of small leaves usually produced along stems; leaves with or without sessile glands beneath the

nonglandular hairs.

ass

2. Stems thicker and obscurely lignescent, originating from ames A lex; | ly oblanceo-

late to narrowly obovate, strigose but gray-green, almost alway ge, se sile glands often sparse but

evident beneath the nonglandular hairs Heterotheca villosa var. angustifolia

2. Stems thin, distinctly lignescent, originating from relatively thin, adventitiously rooted caudex branches;

leaves narrowly oblanceolate, usually silvery-sericeous, without sessile glands beneath the nonglandular

hairs Heterotheca canescens

Because Heterotheca zionensis has not been previously recognized in Texas, specimens are cited here (below);

occurrences in New Mexico also are documented.

TEXAS. Brewster Co.: 2 mi W of Alpine, south side Hwy 90, left of first underpass to Marfa, infrequent, 4300-4500 ft, 4 Oct 1946,

Brown B95 (LL, SMU-2 sheets, SRSC); limestone soil at Althida, ca 20 mi E of Alpine, 4000 ft, 24 Sep 1935, Fletcher 487 (SRSC); Glass

Mts, arroyo at Altuda Pass, 8 Aug 1940, Warnock 279 (GH, SRSC, TEX); frequent on Alpine Golf Course, Alpine, 13 Aug 1937, Warnock

T439 (GH, TEX); Paradise Canyon, 4 mi W of Alpine, infrequent in igneous soil, 4600 ft, 8 Aug 1947, Warnoch 6656 (LL, SMU 2 sheets,

SRSC, TEX). Jeff Davis Co.: Davis Mts, Mitre Peak, 16 Aug 1927, Cory 45386 (LL). Presidio Co.: between railroad and Hwy 90, ca. 1/2

mi NW of Paisano Campground, 4900 ft, 3 Aug 1947, Hinckley 3962 (SMU, SRSC). Reeves Co.: common along hwy to Carlsbad, limestone

soil, 11 mi N of Pecos, 4000 ft, 3 Jul 1947, Warnock 6230 (SRSC, TEX); 3 mi N of Arno on Hwy 285, barren ridge with mesquite-yucca

association, 22 Aug 1942, Waterfall 4248 (MO).

MEXICO. Doña Ana Co.: Mesilla Valley, 2 mi NW of San Miguel, sandy wash, 1200 m, 18 Aug 1930, Fosberg $3850 (LL);

Mesilla Valley, sand hills, no date, Mead s.n. (NMC); Little Mt., near Las Cruces, 26 Aug 1902, Metcalfe s.n. (NMC); E Las Cruces at University

Ave. E of Telshor Drive, 2 km E of IH-25, NE side of Tortugas Mtn., roadside drain in limestone soil, 1360 m, 24 Aug 2001, Spellenberg

& Brouillet 13268 (BRIT, NMC); S edge of Las Cruces, roadside of Hwy I-10, abundant roadside weed for many miles along road edges

and borrow pits, 3900 ft, 31 Aug 1981, Ward 81-550, voucher for n = 9 (NMC); Organ Mountains, Bishop’s Cap, 4 Oct 1903, Wooton s.n.

(NMC); mesa W of the Organ Mountains, 4000 ft, 1 Oct 1907, Wooton s.n. (MO, NMC); Bishop Cap, 2 air km NNW of the top of Bishop

Cap, 1400 m, 24 Sep 1988, Worthington 17524 (TEX); Bishop Cap, 2 air km NNW of the top of Bishop Cap, limestone arroyo, 1400 m, 2

Oct 1988, Worthington 17560 (TEX); Organ Mts., Butterfield Park, soil from igneous substrate, 4600 ft, 22 Sep 1999, Worthington 28700

(TEX). Eddy Co.: Queen Quadrangle, Last Chance Canyon, canyon bottom of limestone boulders, 5000 ft, 24 Jun 1999, Baker 13358

(NMC); Guadalupe Mts., 2.6 mi by road NE of Sitting Bull Falls, 24 Jun 1981, Van Devender & Oler s.n. (BRIT); Last Chance Canyon,

1400 m, 29 Aug 1988, Worthington 27964 (TEX). Grant Co.: near Silver City, 30 Sep 1880, Greene 12857 (MO); Silver City, bank of Hwy

180 West, near corner of Hill and Mississippi streets, 6000 ft, 20 Oct 1985, Zimmerman 2984 (TEX); NW edge of Silver City, along Hwy

180, 6000 ft, 12 Sep 1987, Zimmerman 3001 (TEX). Lincoln Co. [label says “Chaves Co.”]: 35 mi W of Roswell, ca. 3800 ft, Aug 1900,

Earle 510 (MO). McKinley Co.: Zuni Mts., 8 mi E of NM Hwy 32 on road to McGaffey, sandy soil, just inside Cibola National Forest, 16

Aug 1973, Spellenberg 3527 (NMC). Otero Co.: Mayhill, low, rocky, open woods, 13 Aug 1969, 6670 ft, Demaree 60793 (SMU). Santa

Fe Co.: Santa Fe, Old Cemetery, 2120 m, 28 Sep 1935, Arsene 22126 (SMU); Santa Fe, SE part of city near Musem of Internatl. Folk Art,

dry open ground, 7200 ft, 3 Aug 1963, Bennett 8296 (TEX); Broken Hill Ranch near Rte 10, 3 mi NE from Los Cerillos, 6000 ft, 13 Aug

1963, Bennett 8298 (TEX); 4 mi N of Madrid, locally abundant on roadsides, 6800 ft, 15 Aug 2001, Neff 01-08-15-01 (TEX); Santa Fe

Creek at Santa Fe, 23 Jul 1908, Standley 4503 (NMC).

Status of Chrysopsis elata

Chrysopsis elata Osterhout was cited by Semple (1996) as a synonym of Heterotheca fulcrata var. fulcrata and

the isotype (NY) was annotated by him as such in 1997. The NY isotype had been earlier annotated by

Semple (in 1993) as H. villosa var. minor. Chrysopsis elata is identified here as H. villosa var. villosa.

Chrysopsis elata Osterhout, Bull. Torrey Bot. Club 57:560. 1931. Type: U.S.A. Coronapo. Eagle Co.: Red Cliff, 16 Aug 1906, G.E. Oster-

hout 3359 (HOLOTYPE: RM; isotype: NY [internet image!]). The protologue states that Osterhout “3335” is the type, but the RM and

NY sheets both are labeled with the collection number “3359.” The original manuscript, in Osterhouts hand, is stapled to the RM

holotype and has “3359” as the type number. On Osterhouts unique red type label, C.L. Porter in 1950 noted that publication of

the number *3335" was in error.

30 Journal of the Botanical R h Institute of Texas 1(1)

ACKNOWLEDGMENTS

I am grateful for loans of specimens from NMC, SJNM, and SRSC, for help from staff during study at MO

and TEX, to Vernon Harms and an anonymous reviewer for comments on the manuscript, to Ron Hartman

and B. Ernie Nelson for information regarding Wyoming localities and specimens at RM, and to Robert

George and Kathy Scott at BRIT for help with the maps.

REFERENCES

Dorn, R.D. 2001. Vascular plants of Wyoming (ed. 3). Mountain West Publishing, Cheyenne, Wyoming.

GREAT PLAINS FLORA ASSOCIATION (R.L. McGregor, coordinator; T.M. Barkley, editor). 1976. Atlas of the Flora of the Great

Plains. lowa State University Press, Ames.

Harms, V.L. 1968a. Nomenclatural changes and taxonomic notes on Heterotheca, including Chrysopsis, in Texas

and adjacent states. Wrightia 4:8-20.

Harms, V.L. 1968b. Application of the name Heterotheca viscida. Rhodora 70:301-303.

HARRINGTON, H.D. 1954. Manual of the plants of Colorado. Sage Books, Denver.

Nesom, G.L. 1997. Review: “A revision of Heterotheca sect. Phyllotheca (Nutt.) Harms (Compositae: Astereae)" by

J.C. Semple. Phytologia 83:7-21.

Nesom, G.L. 2006. Taxonomic overview of the Heterotheca villosa complex (Asteraceae: Astereae). Sida 22:

367-380.

Semple, J.C. 1987. New names, combinations, and lectotypifications in Heterotheca (Compositae: Astereae). Brit-

tonia 39:379-386.

SEMPLE, J.C. 1996. A revision of Heterotheca sect. Phyllotheca (Nutt.) Harms (Compositae: Astereae). Univ. Waterloo

Biol. Ser. 37:i-iv, 1-164.

SEMPLE, J.C. 2006. Heterotheca (Asteraceae: Astereae). In: Flora of North America Editorial Committee, eds. 1993+.

Flora of North America North of Mexico. 12-- vols. Oxford University Press, New York and Oxford. Vol. 20.

SHINNERS, L.C. 1951. The north Texas species of Heterotheca, including Chrysopsis (Compositae). Field & Lab.

19:66-71.

WEBER, W.A. and R.C.Whrrrman. 1990. Colorado flora: Eastern slope. University Press of Colorado, Boulder, Colorado.

WEBER, W.A. and R.C. WurrrMAN. 1996. Colorado flora: Eastern slope (rev. ed.). University Press of Colorado, Boulder,

Colorado.

Wooton, E.O and PS. StanDLeY. 1913. Descriptions of new plants preliminary to a report upon the flora of New

Mexico. Contr. U.S. Natl. Herb. 16:109-196.

Wooton, E.O and PS. STANDLEY. 1915. Flora of New Mexico. Contr. U.S. Natl. Herb. 19:9-794.

GENERIC REALIGNMENTS IN TRIBE POTEN HELEAR AND REVISION OF

DRYMOCALLIS (ROSOIDEAE: ROSACEAE) IN NORTH AMERICA

Barbara Ertter

University and Jepson Herbaria

University of California

Berkeley, California 94720-2465, U.S.A.

ABSTRACT

The convergence of morphology-based research by J. Sojak and molecular analysis by T. Erikkson et al. supports the resurrection of

Comarum, Dasiphora, Drymocallis, Sibbaldia, and Sibbaldiopsis from Potentilla s.l. These segregate genera, which were used by P.A. Rydberg

in the last continent-wide treatment of tribe Potentilleae in North America, will be used in the forthcoming treatment of Potentilleae in

Flora of North America North of Mexico. The genus Drymocallis in North America is revised to encompass 15 species and seven additional

varieties, in place of the three species of Potentilla (P. arguta, P. fissa, P. glandulosa) and 11 additional varieties recognized by D.D. Keck.

Drymocallis pseudorupestris var. saxicola Ertter (widespread), D. pseudorupestris var. crumiana D.D. Keck ex Ertter (California),

and D. deseretica Ertter (central Utah) are described as new, and several new combinations are made: D. lactea var. austiniae (Jeps.)

Ertter, D. glandulosa var. wrangelliana (Fisch. & Avé-Lall.) Ertter, D. glandulosa var. reflexa (Greene) Ertter, D. glandulosa var.

viscida (Parish) Ertter, D. campanulata (C.L. Hitchc.) Ertter, and D. cuneifolia var. ewanii (D.D. Keck) Ertter. The last combina-

tion results from the recent rediscovery of D. cuneifolia var. cuneifolia, which is currently known from a single population in the San

Bernardino Mountains of California.

Key Worbs: Rosaceae, Potentilleae, Drymocallis, Dasiphora, Comarum, generic realignment, new species, Sibbaldiopsis

RESUMEN

I gencia entre lai igación basada en la morfología realizada por J. Soják y los análisis moleculares realizados por T. Erikkson

et al. soportan] ión d ECT um, Dasiphora, Drymocallis, Sibbaldia, y Sibbaldiopsis desde Potentilla s.l. Estos géneros segregados,

que se usaron por P.A. Rydberg en el ültimo tratamiento a nivel continental de la tribu Potentilleae en Norte América, será usado en

el próximo tratamiento de Potentilleae en la Flora of North America North of Mexico. El género Drymocallis en Nore América es revisado

para incluir 15 especies y siete variedades adicionales, en lugar de las tres especies de Potentilla (P. arguta, P. fissa, P. glandulosa) y 11

variedades adicionales reconocidas por D.D. Keck. Drymocallis pseudorupestris var. saxicola Ertter (de amplia distribución), D.

pseudorupestris var. crumiana D.D. Keck ex Ertter (California), y D. deseretica Ertter (Utah central) se describen como nuevas, y

se hacen varias combinaciones nuevas: D. lactea var. austiniae (Jeps.) Ertter, D. glandulosa var. wrangelliana (Fisch. & Avé-Lall.)

Ertter, D. glandulosa var. reflexa (Greene) Ertter, D. gland var. viscida (Parish) Ertter, D. ocampanulats (C.L. Hitchc.) Ertter,

are 1 ] 15] : 1

D. cuneifolia var. ewanii (D.D. Keck) Ertter. I

de D cuneifolia var. cuneifol id,

que es conocida actualmente de una única población en las montañas de San Bernardino de California.

Tribe Potentilleae Sweet (Rosoideae: Rosaceae) is comprised of Potentilla L. and variously recognized segregate

and related genera, including Fragaria L., Ivesia Torrey & A. Gray, and Sibbaldia L. All of these genera have

at one time or another been included within Potentilla, and multiple segregate genera have been proposed.

Potentilla s.s. is widespread throughout arctic and temperate regions of the northern hemisphere, with

southward extensions along the American cordilleran system and mountains of southeast Asia and adjacent

Malesian archipelago. The last attempt at a worldwide monograph of Potentilla s.l. (excluding Ivesia, Horkelia

Cham. & Schltdl., and related segregate genera endemic to western North America) was by Wolf (1908),

who tallied over 300 species along with numerous varieties and hybrids. This number is a poor indicator

of the current status, given the significant number of taxa that have been described, reduced to synonymy,

or otherwise been subject to taxonomic realignment during the intervening century.

Two relatively recent developments have contributed to an improved understanding of worldwide

diversity and generic circumscriptions in Potentilleae, one political, one scientific. On the geopolitical front,

the collapse of the Soviet bloc and renewed relations between the United States and China made essential

comparisons of American and Asian variation significantly easier. Collaboration among researchers with

expertise in their respective geographic areas are paving the way for a modern monographic synthesis of

J. Bot. Res. Inst. Texas 1(1): 31 — 46. 2007

32 Journal of the Botanical R h Institute of Texas 1(1)

Potentilleae, including the Panarctic Flora project (Elven et al. 2007) and the Flora of China project (Potentil-

leae by Li et al. 2003). In the scientific realm, a molecular phylogenetic underpinning of Potentilleae has now

been generated by Torsten Eriksson et al. (1995, 1998, 2003) with additional efforts underway by Christoph

Dobes (pers. comm. 2006), Bente Eriksen (pers. comm. 2007), and their colleagues.

The present paper summarizes the historical and current nomenclatural framework in Potentilleae

resulting from these developments. It also provides a revision of one of the resurrected generic segregates,

—

Drymocallis Fourr. ex Rydb., including several new taxa and combinations. This synopsis serves to introduce

nomenclatural adjustments to Potentilleae that will be implemented in vol. 9 of Flora of North America North

of Mexico, scheduled for publication in late 2008.

NOMENCLATURAL AND PHYLOGENETIC OVERVIEW OF POTENTIELEAR

The last continent-wide treatments of North American Potentilleae were by Per Axel Rydberg (1898, 1908),

who argued that “Either the whole tribe, Fragaria also included, must constitute a single genus, or else both

Potentilla and Ivesia be divided into several genera” (1898, p. 16). Choosing the latter option in his 1898

monograph of North American Potentilleae, Rydberg accordingly recognized the genera Argentina Hill,

Chamaerhodos Bunge, Comarella Rydb., Comarum L., Comocarpa Rydb., Drymocallis, Duchesnea Sm., Horkelia

(encompassing Ivesia), Sibbaldia, Sibbaldiopsis Rydb., and Stellariopsis Rydb., in addition to Fragaria and a

significantly reduced Potentilla s.s. This generic framework was largely retained in his 1908 treatment of

Rosaceae for North American Flora, except for the replacement of Comocarpa with Dasiphora Raf., the resur-

rection of Ivesia as distinct from Horkelia, and the addition of Horkeliella Rydb. and the recently described

Purpusia Brandegee.

Rydberg was considered a consummate “splitter” by contemporary and later generations of botanists,

such that most of his generic segregates disappeared into synonymy in subsequent regional floras in North

America. Most European authors also adopted an inclusive circumscription of Potentilla, following the world

monograph by Theodor Wolf (1908). David D. Keck (1938) retained and revised Horkelia and Ivesia as distinct

genera, but submerged Comarella, Horkeliella, and Stellariopsis within Ivesia. Keck (in Clausen et al. 1940) also

submerged Drymocallis within Potentilla, with most of Rydberg’s species reduced to subspecies or synonyms

of a single species, P. glandulosa Lindl. In the opinion of Keck and his collaborators, “there is no justification

for excluding Drymocallis from Potentilla. Certain species undoubtedly belonging to Potentilla closely link

Drymocallis to the main body of the genus. We do believe, however, that the morphological distinctions of

the anther and the position of the style, combined with a homogeneity of form, justify the maintenance of

Drymocallis as a section or subsection of Potentilla.”

Keck’s circumscriptions of genera and species in the Potentilleae were largely adopted by subsequent

regional floras in North America, though Ivesia and Horkelia were sometimes included in Potentilla (e.g.,

Howell 1949; Kearney & Peebles 1951). An element of confusion was introduced by Hutchinson (1964), who

cited Keck's revision of Ivesia and Horhelia in his synthesis of flowering plant genera but failed to incorporate

the conclusions in his opus. Hutchinson’s indication of 35 species of Ivesia, three species of Horkeliella, two

species of Comarella, and one species of Stellariopsis conflicts with Keck's recognition of only 22 species of

Ivesia s.l., which includes the species placed in the segregate genera recognized by Hutchinson. Researchers

who rely on Hutchinson for generic delineations of these western North American genera are accordingly at

odds with current usage as reflected in recent floristic treatments (e.g., Hitchcock & Cronquist 1961; Ertter

1993; Holmgren 1997; Welsh et al. 1993). Subsequent to Keck’s revision of Ivesia and Horkelia, no flora has

used either Comarella or Stellariopsis, and when Horkeliella was resurrected it contained only two species

(Ertter 1993). Purpusia, which Keck (1938) had retained as a distinct genus, has also been submerged into

Ivesia, along with several species of Potentilla that were morphologically nearly identical to certain species

of Ivesia except for details of flower structure that apparently form an evolutionary sequence (Ertter 1989).

Further generic resurrections are now supported by the felicitous convergence of independent morpho-

logical and molecular studies. In 1989, Czech botanist Jiří Soják published his conclusion that Potentilleae

| iD in North America 33

Ertter, The tribe Potenti

was divided into two fundamental evolutionary lines. The Fragaria-line, characterized by sub-basal or lateral

styles and anthers with a single horseshoe-shaped theca that opens by a marginal slit, consisted not only

of Fragaria but also those species sometimes placed in Comarum, Dasiphora (as Pentaphylloides Duhamel),

Drymocallis, Sibbaldia, Sibbaldiopsis and two small Asian genera (Farinopsis Chrtek & Soják and Schistophyl-

lidium [Juz. ex Fed.] Ikonn.). The second evolutionary line, comprised of Potentilla s.s., Ivesia, and Horhelia,

was characterized by subterminal styles and anthers with two thecae divided by the connective apex that

opens by two lateral slits. The species potentially comprising Argentina (e.g., P. anserina L.) and the Asian

genus Tylosperma Botsch. were more problematic.

Soják was then faced with the three options of 1) lumping everything in a single swollen genus, which

in his understanding would need to be Fragaria: 2) treating the two evolutionary lines as separate genera,

with all species of Potentilleae included in either Fragaria or Potentilla; or 3) breaking the components of each

line into narrower, more homogeneous genera. Echoing Rydberg’s decision, Soják chose the last option, as

the simplest, most straight-forward solution that required the least nomenclatural disruption, and accord-

ingly recognized the genera listed above and created the new combinations needed in Eurasian Drymocallis.

Except for Drymocallis, Soják was therefore in concordance with the nomenclature used by his Russian col-

league Boris A. Yurtsev in a flora of the arctic Soviet Union (1984). Although Soják's seminal paper was not

widely available, being published in the journal of the National Museum in Prague while Czechoslovakia

was just emerging from behind the Iron Curtain, his generic conclusions are reflected in more recent papers

(e.g., Soják 2004), in which the superfluous and thus illegitimate Pentaphylloides is replaced by Dasiphora

and Argentina is sometimes treated as distinct. An unpublished outline of North American Potentilleae Soják

prepared in 1993 also reflects this generic framework, expanded by the addition of two subtribes to accom-

modate Chamaerhodos, Alchemilla L., Aphanes L., and Lachemilla Rydb.

Several years after Soják first published his generic outline, Torsten Eriksson and Michael Donoghue

(1995) presented a molecular phylogenetic analysis of the Potentilleae that provided independent confirmation

of the fundamental distinction between a potentilloid clade and a fragarioid clade, essentially correspond-

ing to Soják's evolutionary lines. Further studies (Eriksson et al. 1998, 2003) confirmed and expanded the

results, showing moreover that Chamaerhodos and the alchemilloids were also nested in the fragarioid clade

and that the position of P. anserina (Argentina) was indeed ambiguous. Comarella, Purpusia, and Stellariopsis

were specifically targeted for inclusion in further studies, indicating a reliance on Hutchinson (1964) as a

generic starting point rather than Keck (1938) and all more recent floras. As it happens, preliminary ITS

analysis of representative species of Ivesia (including Comarella, Purpusia, and Stellariopsis), Horkelia, and

Horkeliella by the Christopher Baysdorfer lab (California State University, Hayward) results in a monophyletic

polytomy (Ertter et al. 1998, unpublished data).

GENERIC DELIMITATIONS FOR FLORA OF NORTH AMERICA

There is no question that the species currently treated as Potentilla s.l. that fall into the fragarioid clade cannot

be retained in Potentilla without that genus being polyphyletic, unless Fragaria and possibly Chamaerhodos,

Alchemilla, Aphanes, and Lachemilla are also included. The two options are towards a more inclusive Poten-

tilla, or the removal of all taxa in the fragarioid clade from Potentilla. In their discussion of nomenclatural

implications, Eriksson et al. (1998, 2003) noted that if all taxa in the clades containing Potentilla s.l. were

combined in a single genus, Fragaria apparently had priority. They were understandably reluctant to do a

wholesale transfer of hundreds of species of Potentilla s.l. into Fragaria and instead, using the situation as an

example of the purported advantages of phylogenetic nomenclature, proposed several rankless names for

the best resolved clades (e.g., “Fragariinae”). Taking an alternative approach, Mabberley (2002) concluded

that Fragaria need not take precedence and accordingly provided several new combinations to allow Fragaria

to be included in Potentilla s.1.

Mabberley based his decision on a trend toward more inclusive genera in economically important taxa

(e.g., Prunus s.l., Lycopersicon combined with Solanum). In contrast, the dominant trend affecting generic

34 Journal of the Botanical R h Institute of Texas 1(1)

usage in North America has been in the opposite direction; e.g., Grimes' (1990) work on the Psoraleeae (Fa-

baceae) and Baldwin's (1999) work on the Madiinae (Asteraceae), often resultingin the resurrection of generic

segregates first proposed by Rydberg and his fellow splitters. This latter approach is easily implemented in

Potentilla, in that all of the essential generic names are already available and were used in the most recent

continent-wide treatment of the family (Rydberg 1908). Initial steps have already been taken towards floristic

implementation of the fragarioid generic segregates in both Europe (Kurtto & Eriksson 2003) and North

America (e.g., Weber 1987; Weber & Whittman 1992; Pojar 1999; Aiken et al. 2006). Full implementation

in North America will occur in volume 9 of Flora of North America North of Mexico (scheduled for publication

in 2008), in which the following fragarioid genera that have sometimes been included in Potentilla will be

recognized:

Comarum L.—Comarum palustre L. replaces Potentilla palustris (L.) Scop. One other Asian species some-

times placed in Comarum has been placed in its own genus as Farinopsis salesoviana (Stephan) Chrtek &

Soják (1984).

Dasiphora Raf. —Dasiphora fruticosa (L.) Rydb. replaces Potentilla fruticosa L. Several previous treatments

that initiated this generic change in North American (e.g., Holmgren 1997; Pojar 1999; Weber 1987) adopted

Pentaphylloides, but this name has been subsequently interpreted as an illegitimate superfluous name for

Potentilla (as noted by Kurtto & Eriksson 2003; Reveal et al. 1999).

Drymocallis Fourr. ex Rydb.—Drymocallis is the generic segregate from Potentilla with the most species,

with centers of radiation in western North America, central Asia, and southeastern Europe. My provisional

revision (below) recognizes fifteen species with an additional seven varieties in North America. Around

thirteen Old World species are currently recognized, with most of the necessary combinations established

by Soják (1989) and Kurtto & Eriksson (2003). The elevation of Drymocallis to generic status allows it to be

used more definitively in making biogeographic comparisons between the southern Rocky Mountains of

North America and the Altai of central Asia, as has recently been done by Weber (2003).

Fragaria L.—The treatment of Fragaria being prepared by Gündter Staudt will reflect his recent revision of

North American strawberries (Staudt 1999).

Sibbaldia L.—Sibbaldia procumbens L. var. procumbens replaces the nomenclaturally invalid Potentilla

procumbens (L.) Clairv. (a later homonym of the unrelated P. procumbens Sibth.), P. sibbaldi Haller f., and P.

sibbaldia Kurtz. Soják (pers. comm. 2006) recognizes only six species of Sibbaldia, with most others placed

in Potentilla.

Sibbaldiopsis Rydb.—Sibbaldiopsis tridentata (Soland.) Rydb. replaces Potentilla tridentata Soland. Soják

(ined.) is also transferring two Asian species (P. cuneifolia Bertol. and P. miyabei Makino) into this genus,

which corresponds to Wolf's (1908) “Grex Tridentatae.”

The potentilloid segregates Duchesnea, Ivesia, Horkelia, and Horkeliella will be retained as in the current

treatment for California (Ertter 1993) and the Intermountain Region (Holmgren 1997). Although it has long

been assumed that these genera most likely evolved from and are accordingly nested within Potentilla s.s., as

is now supported by molecular evidence (Erikkson et al. 1995, 1998, 2003), I strongly concur with Brum-

mitt (2002), Diggs and Lipscomb (2002), and Hórandl (2006) that paraphyly alone is insufficient grounds

for dictating generic delimitations. Comarella, Purpusia, and Stellariopsis will be retained in Ivesia, although

Soják (ined.) argues that the distinctive anthers of Stellariopsis (= Ivesia santolinoides A. Gray), which dehisce

by subterminal pores, justify recognition as a separate genus.

REVISION OF NORTH AMERICAN DRYMOCALLIS

The North American members of Drymocallis, as Potentilla glandulosa and allies, were among those studied

by Jens Clausen, David D. Keck, and William M. Hiesey in their seminal experiments on biosystematics

(Clausen et al. 1940; Clausen & Hiesey 1958). They confirmed that the complex, which they considered

Ertter, The tribe P tentill ID llis in North America 35

to be *one of the most satisfactory in the transplant investigations, and ... the most important" (Clausen et

al. 1940, p. 26), consists of a wide diversity of ecotypes, often highly localized, differing from one another

ecologically, physiologically, and morphologically. They also demonstrated that hybrids between these

ecotypes are readily generated, aided by a diploid chromosome number of n=7 shared by all species thus far

counted, and intergradation zones where ecotypes intersect are the norm. In contrast to the Potentilla gracilis

complex, which they also investigated, apomixis is not known to play a role in Drymocallis. The responsibility

of converting this biosystematic complexity into a functional taxonomic framework fell on Keck (in Clausen

et al. 1940), whose compromise approach reduced Rydberg's (1908) 28 species of Drymocallis to only three

species of Potentilla (P. arguta Pursh, P. fissa Nutt., P. glandulosa Lindl.), with most of the ecotypic variation

treated as subspecies of P. glandulosa.

My initial intent for Flora of North America was to adopt Keck's taxonomy, as modified by Ertter (1993)

and Hitchcock et al. (1969), with the requisite new combinations in Drymocallis and the description of a new

taxon from California. I reluctantly concluded, however, that Keck's outline did not adequately accommodate

current evidence of variation in North American Drymocallis, especially outside of central California where

the biosystematic experiments had taken place. For one thing, it is not clear that Keck's subspecies are more

closely related within a species than between species, as indicated, for example, by the narrow inflorescences

of P. arguta subsp. convallaria (Rydb.) D.D. Keck and P. glandulosa subsp. hansenii (Greene) D.D. Keck, or the

numerous leaflets of P. fissa and P. glandulosa subsp. arizonica (Rydb.) D.D. Keck. At the same time, there

are extremes of variation beyond those addressed by Keck's abbreviated taxonomic treatment, such as the

narrowly opened flowers of P. glandulosa subsp. ewanii D.D. Keck and P. glandulosa var. campanulata C.L.

Hitchc. Rather than perpetuate an unacceptable taxonomic framework, I accordingly have decided to take

the opportunity of a nomenclatural overhaul to introduce an alternate framework. This effort is unabashedly

provisional, in that the time frame for Flora of North America does not allow for the complete revision that

Drymocallis clearly deserves. My hope is that it is nevertheless a step forward in ting the natural

variation in Drymocallis, until such time as a more complete study can be undertaken that incorporates

molecular analysis in combination with extensive studies of variation in the field.

At heart is the question of whether the goals of taxonomy are better served by sharply defined group-

ings, however polymorphic, or by more finely parsed natural variation, even at the expense of crisp cir-

cumscriptions. I have opted for the latter, even though this approach runs counter to that of several recent

floristic works (e.g., Holmgren 1997; Welsh et al. 1993), in which many of Keck’s subspecies are relegated

to synonymy. My approach, comparable to that currently implemented in some other rosaceous genera

(e.g., Rosa and sections of Potentilla s.s.), is to give taxonomic recognition at the species or subspecies level

if a significant core of populations share a relatively cohesive suite of unique characters underpinned by a

definable ecogeographic setting. Varieties are used where the core differences are less distinct and/or the

intergradations more complex. Such a taxonomic approach can only be done with full understanding that

any attempt to recognize formal taxa in North America Drymocallis will be compromised by wide zones

of intergradation and populations that defy placement. The alternative is to accept such broadly defined

taxa that extensive variation within an ecogeographic setting is glossed over, even that which would easily

qualify as distinct species if the inconvenient zones of intergradation went extinct. Given this situation, and

the current sociopolitical setting, it would be most unfortunate if valid components of biodiversity failed

to receive the conservation attention they deserve for lack of sharp boundaries (Ertter 1997). Indeed, my

suspicion is that at least some of the purported intergradation actually represents additional unrecognized

entities, and that a cleaner taxonomy might result with the recognition of even more, not fewer, taxa. At the

very least, I believe that the resultant taxonomic framework will lead to a better appreciation of diversity

within Drymocallis and encourage more studies on a complex and fascinating genus.

Presented here is a complete synopsis of species of Drymocallis that will be recognized in Flora of North

America. Synonyms used in recent floras are listed, new combinations are provided as needed, three new taxa

are described, and some nomenclatural issues are discussed. A complete key, descriptions, and additional

36 Journal of the Botanical R h Institute of Texas 1(1)

discussion will be available in vol. 9 of Flora of North America, and the California species will be covered in

the pending new edition of The Jepson Manual. Additional synonymy can be found in Clausen et al. (1940),

Rydberg (1908), Hitchcock and Cronquist (1961), Holmgren (1997), and the upcoming treatment in Flora

of North America.

1. Drymocallis fissa (Nutt.) Rydb., Mem. Dept. Botany Columbia Coll. 2:197. 1898. = Potentilla fissa Nutt.

2. Drymocallis arizonica Rydb., N. Amer. Fl. 22:373, 1908. - Potentilla glandulosa Lindl. subsp. arizonica (Rydb.) D.D.

Keck, not Potentilla arizonica Greene

3. Drymocallis arguta (Pursh) Rydb., Mem. Dept. Bot. Columbia Coll. 2: 192. 1898. = Potentilla arguta Pursh.

In his treatment for North American Flora, Rydberg (1908) replaced D. arguta with Drymocallis agrimonioides

(Pursh) Rydb., presumably because the conspecific Geum agrimonioides Pursh (Fl. Amer. Sept. 351) was on

an earlier page than Potentilla arguta Pursh (Fl. Amer. Sept. 736). According to the International Code of

Botanical Nomenclature, page sequence has no bearing on priority in a single work that was published as a

unit, which is the situation for Flora Americae Septentrionalis since both volumes were distributed in the last

two weeks of 1813 (Reveal et al. 1999). Rydberg's (1898) initial use of D. arguta, in which Geum agrimonioides

is cited in synonymy, confirms that the more familiar epithet can be retained.

4. Drymocallis convallaria (Rydb.) Rydb., Mem. Dept. Bot. Columbia Coll. 2:193. 1898. = Potentilla arguta Pursh

subsp. llaria (Rydb.) D.D. Keck; P arguta Pursh var. convallaria (Rydberg) Th. Wolf.

2 Drymocallis micropetala Rydb., N. Amer. Fl. 22:375. 1908. = Potentilla glandulosa Lindl. subsp. micropetala (Rydb.)

D.D. Keck; P glandulosa Lindl. var. micropetala (Rydb.) S.L. Welsh & B.C. Johnst.

6. Drymocallis hansenii (Greene) Rydb., Mem. Dept. Bot. Columbia Coll. 2:200. 1898. = Potentilla hansenii Greene;

P glandulosa Lindl. subsp. hansenii (Greene) D.D. Keck.

7. Drymocallis lactea (Greene) Rydb., N. Amer. Fl. 22:369. 1908.

7a. Drymocallis lactea var. lactea - Potentilla glandulosa Lindl. var. nevadensis S. Watson; P. glandulosa subsp. nevadensis (S.

Watson) D.D. Keck; not P nevadensis Boiss. (1838).

This taxon has a well-established identity as Potentilla glandulosa var. (or subsp.) nevadensis, but the epithet

lactea has priority at species rank and its use avoids possible confusion with P. nevadensis Boiss.

7b. Drymocallis lactea var. austiniae (Jeps.) Ertter, comb. nov. Basioxvw: Potentilla glandulosa Lindl. var. austiniae Jeps.,

Fl. Calif. 2:181. 1936.

Jepson (1936) called P. glandulosa var. austiniae a nom. nov. for P. glandulosa var. fissa Jepson, the combination

he had used in his Manual (Jepson 1925). According to K. Gandhi (pers. comm. 2006), var. austiniae stands

as a legitimate taxon distinct from P. fissa Nutt., since this species was explicitly excluded by Jepson (1936),

but var. fissa Jeps. must be considered a synonym of P. fissa and is furthermore an isonym of P. glandulosa

var. fissa (Nutt.) Th. Wolf (1908).

This variety was erroneously treated as Potentilla glandulosa subsp. ashlandica (Greene) D.D. Keck by

Ertter (1993), but I now understand both to be separate entities.

8. Drymocallis ashlandica (Greene) Rydb., Mem. Dept. Bot. Columbia Coll. 2:200. 1898. = Potentilla glandulosa

Lindl. subsp. ashlandica (Greene) D.D. Keck; P ashlandica Greene, Pittonia 3:248, 1897; P ciliata Howell, Fl. N. W. Amer. 1:1715.

1898; not P ciliata Raf. (1840) or P ciliata Greene (1887).

Potentilla ashlandica Greene is sometimes said to be a nom. nov. for Potentilla ciliata Howell (e.g., Keck in Clau-

sen et al. 1940), the latter illegitimate as a result of being a later homonym, but Greene's name was published

first and makes no reference to P. ciliata. Both names are based on a collection by Howell from Ashland Butte,

the only known collection of which is in Howell’s herbarium in ORE (now housed in OSC).

9. Drymocallis pseudorupestris (Rydb.) Rydb., Mem. Dept. Bot. Columbia Coll. 2:194. 1898. = Potentilla

Ertter, The tribe P tentill ID llis in North America 37

glandulosa Lindl. subsp. pseudorupestris (Rydb.) D.D. Keck; P glandulosa var. pseudorupestris (Rydb.) Breitung; P rupestris L. var.

americana Th. Wolf.

Many populations of Drymocallis pseudorupestris from Montana and Wyoming, including the type of Potentilla

pseudorupestris Rydb. (Long Baldy, Little Belt Mountains, Montana, Flodman 598 [NY!]) are significantly larger

plants than is the norm elsewhere, with bigger petals and longer filaments. I am treating these populations as

D. pseudorupestris var. pseudorupestris, with further research needed to determine the optimum circumscrip-

tion and resultant geographic range. Two new varieties are described here: var. saxicola to accommodate the

majority of populations previously placed in Potentilla glandulosa var. pseudorupestris, and var. crumiana to

accommodate a variant from the Sierra Nevada proposed but not published by Keck.

9a. Drymocallis pseudorupestris var. pseudorupestris

9b. Drymocallis pseudorupestris var. saxicola Ertter, var. nov. (Fig. 1, M-R). Tv: UNITED STATES. Ipauo: Lemhi

Co.: Bighorn Crags ca 30 air mi W of Salmon, head of drainage W of Welcome Lake, at N end of saddle to Barking Fox Lake, ledges

in granite, T21N RI6E S28 N1/2 of SW, ca 9000 ft, 31 Jul 1990, B. Ertter 9493 with D. Atwood, R. Moseley, S. Bernatas, M. Mancuso

(HOLOTYPE: UC; isotypes: BRY, ID, NY).

A var. pseudorupestris statura minore (« 2.5 dm), floribus parvioribus, filamentis brevioribus (1-2.5 mm) differt.

Herbaceous perennial from openly branched caudex or spreading rootstocks, sometimes + rhizomatous,

forming clumps to 6 dm across. Stems few to many, erect, 0.6-2.53) dm tall, the base 1-2(-3) mm diam.

with sparse to abundant peglike glands and septate glandular hairs to 2 mm long, short eglandular hairs

sparse to abundant (lacking in some collections from Custer Co., ID). Basal leaves 3-9(-16) cm long

with (2—)3(-4) pairs of lateral leaflets; sheathing leaf-bases often sparsely strigose; terminal leaflet broadly

obovate-cuneate with a + rounded apex, 0.8-2(-4) cm long, 0.7-2(-3) cm wide, usually with abundant

peglike glands and sparse to moderately abundant eglandular hairs to 1.5 mm long, the margins single- to

+ double-toothed ca '4—'4 to base with 3-8(-12) teeth per side. Cauline leaves 0—1(-2), 2-5(-8) cm long

with 23 pairs of lateral leaflets. Inflorescence relatively open, comprising 4-7/4 of plant height, not par-

ticularly leafy; subtending bract + trilobed, 1-2.5(-3) cm long; branches diverging at a (10—)20—40(—50)?

angle; pedicels 3-15(-20) mm long, with sparse to abundant (never absent) septate glandular hairs and 0 to

abundant eglandular hairs 0.1-0.5 mm long. Flowers (2—)3—12(-20); hypanthium saucer-shaped, 2.5—4(—5)

mm diam., with highly variable proportions of peglike glands, septate glandular trichomes, and eglandular

hairs to 1 mm long; epicalyx bractlets linear-lanceolate to broadly elliptic, 2-5 mm long, + 1-1.5 mm wide,

often toothed or lobed; sepals 4—6(-7) mm long, the vestiture like that of hypanthium, the apex obtuse

(to acute), often mucronate, entire; petals cream to + yellow, not red-tinged, narrowly to broadly obovate,

4—9 mm long (longest in Washington), 3.5-6(-8) mm wide, exceeding sepals, the apex rounded; stamens

20-25, the filaments 1-2.5 mm long, not red-tinged, the anthers ovate-elliptic, 0.7-1.2 mm long; styles +

fusiform, + 1 mm long, most often golden brown. Achenes + obliquely pyriform, generally short-beaked,

ca 1 mm long, light brown.

Ecology and phenology.—Cliffs, ledges, outcrops, ridges, talus slopes, lava beds, and other generally

rocky situations, 1000-3400 m elev. Flowering May—Aug.

As circumscribed here, Drymocallis pseudorupestris var. saxicola accommodates the bulk of specimens

previously placed in Potentilla glandulosa var. pseudorupestris, minus the extremes at the northeastern and

southern ends of the range. The current circumscription encompasses significant heterogeneity, which

might be resolved into additional taxa upon further analysis. Plants from southeastern Oregon, including

the type of D. pumila Rydb. from Steens Mountain, combine the habit of D. pseudorupestris and the vestiture

of D. lactea.

REPRESENTATIVE SPECIMENS: CANADA. ALBERTA: Bellevue Hill, Waterton Lakes National Park, J. Nagy & W. Blais 682 (DS). UNITED

STATES. CALIFORNIA: Eldorado Co.: Mt. Tallac, J.T. Howell 22949 (CAS). Shasta Co.: ridge overlooking Lake Helen, Lassen Volca-

nic National Park, H. Bailey & V. Bailey 2949 (UC). Siskiyou Co.: W side of Shastina, Mt. Shasta, W.B. Cooke 15372 (UC). Fresno Co.:

Simpson Meadow, Middle Fork of Kings River, J.T. Howell 33794 (CAS). Tulare Co.: Bullfrog Lakes 6 mi SE of Mineral King, B. Rice 430

(CAS). IDAHO. Adams Co.: Black Lake, J.H. Christ 8674 (UC). Boise Co.: Middle Fork Boise River 2 1/2 mi upstream from Twin Springs,

38 Journal of the Botanical R h Institute of Texas 1(1)

MN j

W EY

Y AQ

H^ <A

Jes eo Ye

a D.

Pon if

ü VSN dl

<= Ve SANA)

Wy SON NS Y

| PA VAS

| Es nh N | a e 1

N S Hf 7 NI "rat V ¢

Vf y x AMS EN V N

p VAR

I 7

Lor

Fic. 1. Varieties of Drymocallis pseudorupestris. A-F: var. pseudorupestris. A: habit. B: basal leaf. C: hypanthi | calyx. D: vestiture of pedicel. E: flower.

F: stamen. G-L: var. crumiana. G: habit. H: basal leaf. | hy thi | ly J tit f dicel. K: flower. L: stamen. M-R: var. saxicola. M: habit.

N: basal leaf. 0: | y i lyx. P tit fp dicel. Q: flower. R: stamen

Ertter, The tribe P tentill ID llis in North America 39

B. Ertter & C. Prentice 8703 (UC). Butte Co.: 10 mi N of Craters of Moon, 13 May 1941, G. Williams s.n. (UC). Camas Co.: summit of

Featherville-Ketchum road, C.L. Hitchcock & C.V. Muhlick 10426 (UC, WTU). Custer Co.: 3.5 mi SW of Stanley Lake, C.L. Hitchcock

& C. V. Muhlick 9626 (NY, UC, WTU). Fremont Co.: Shot Gun, R.J. Davis 746 (UC). Idaho Co.: Seven Devils Ridge, J. H. Christ 12473

(ID, UO). Lemhi Co.: Sleeping Deer Mountain, B. Hayse 35 (WIS). Valley Co.: above Snowslide Lake 10 air mi NE of McCall, B. Ertter

& S. D'Alcamo 7805 (UC; to be distributed). OREGON: Deschutes Co.: Broken Top, Cascade Range, B. Ertter & S. Garrett 15052 (UC).

NEVADA: Elko Co.: 1.2 km NE of Matterhorn, Jarbidge Mts., D. Charlet 445 (UC). White Pine Co.: Schell Creek Range, A. Tiehm &

S. Crisafulli 11797 (NY, RM, WIS). UTAH: Juab Co.: divide between Toms Creek and Basin Creek drainage, Deep Creek Range, N.H.

Holmgren & P.K. Holmgren 11185 (NY, UC). Piute Co.: S of Bullion Creek Canyon, Tushar Mts., N.H. Holmgren, P.K. Holmgren, & C.S.

Keller 10897 (NY, UC). Sevier Co.: 0.75 mi NW of Mt. Terrill, S. Goodrich 24194 (BRY). WASHINGTON: Asotin Co.: Big Butte 6 mi W

of Anatone, H. St. John & R. Palmer 9594 (UC). Chelan Co.: Mission Ridge, Wenatchee Mts., J.W. Thompson 14943 (NY, UC). WYOMING:

Teton Mts., A. Nelson & E. Nelson 6578 (RM, UC).

9c. Drymocallis pseudorupestris var. crumiana D.D. Keck ex Ertter, var. nov. (Fig. 1, G-L). Tv UNITED

STATES. CALIFORNIA: Inyo Co.: Rock Creek Lake Basin, recess or cirque on east side, 11,500 ft, 21 Jul 1934, EW Peirson 11272 (Ho-

LOTYPE: UC 638452; isorvpes: DS 318001, DS 688601).

A var. saxicola foliolis plerumque 9, calycis setis conspicuis 1-1.5 mm longis, stylis plerumque vinosis differt.

Herbaceous perennial from openly branched caudex or spreading rootstock, with abundant stalked glands

that generate an abundant golden resin. Stems few, erect, (0.3—)0.8-2(-2.5) dm tall, the base 1-2 mm diam.

with sparse to abundant peglike glands and septate glandular hairs to 2 mm long, eglandular hairs generally

lacking. Basal leaves (2-)3-9-15) cm long with 3—4(—5) pairs of lateral leaflets; sheathing leaf-bases +

strigose; terminal leaflet broadly obovate to flabellate with rounded apex, 0.2-1.2(22) cm long, 0.5-1.5 cm

wide, with subsessile golden glands and sparse to moderately abundant spreading to ascending rigid setae

0.5-1 mm long, the margins predominantly single-toothed ca 3-4 to base with 2—5 teeth per side. Cauline

leaves 0-2, 23.56) cm long with 2-3 pairs of lateral leaflets. Inflorescence open, comprising 4—'2(-*/4)

of plant height, not leafy; subtending bract 0.5-2.5 cm long; branches diverging at a 10—40? angle; pedicels

5-12(-20) mm long, with + abundant subsessile glands and septate glandular trichomes, eglandular hairs

sparse to 0. Flowers 2-8; hypanthium saucer-shaped, 4-6 mm diam., with both scattered glandular tri-

chomes and + rigid spreading setae 1-1.5 mm long; epicalyx bractlets elliptic-ovate to lanceolate, 2-3.5(-4)

mm long, 1-2 mm wide; sepals 4-6(-7) mm long, often red-tinged, the vestiture like that of hypanthium,

the apex obtuse, generally mucronate, sometimes shallowly erose; petals cream to pale yellowish, often red-

tinged, narrowly to broadly obovate, 5-7(-9) mm long, 3-5 mm wide, exceeding sepals, the apex rounded;

stamens 25-30, the filaments 1.5-2(3) mm long, often red-tinged, the anthers ovate-elliptic to nearly round,

0.7-1 mm long; styles thickly to narrowly fusiform, 1.2-1.5 mm long, most often dark red (golden brown in

White Mts.). Achenes + obliquely pyriform, ca 1 mm long, pale golden-brown with reddish apex.

Ecology and phenology.—Rocky slopes, talus, and ledges, in metamorphic, granitic, and volcanic sub-

strates, 3200-3900 m elev. Flowering Jul-Aug.

The distinctiveness of this taxon was initially discussed in letters between Frank W. Peirson and Ethel

Crum in the 1930’s (archives of University Herbarium, University of California, Berkeley). Peirson, an avid

collector, had sent specimens of Potentilla from Rock Creek Lake Basin in the eastern Sierra Nevada to the

University of California at Berkeley for identification by Ethel K. Crum. Crum helped with Potentilleae for

Jepson’s Flora of California (1936), where she is described as “a scholarly assistant” who “became ardently

attached to Potentilla, wherefore certain new varieties bearing her name” (p. 174). According to Mason

(1943), Crum “assisted in the organization of the material for many of the larger genera in [vol. 2 of Flora of

California], but adopted as her own, the genus Potentilla, the manuscript of which was accepted with some

revisions.” Crum maintained this interest during her tenure as Assistant Curator of the University Herbarium

(1933-1943), during which time she was also Secretary to the Editorial Board of Madroño (Mason 1943).

In both of two drafts of her letter to Peirson of 15 November 1934, Crum identified his No. 11272 as

“Potentilla sp. nov. This very interesting specimen of the Potentilla glandulosa aggregate with reddish tinged

flowers differs in certain particulars from all segregates known to me ... . I believe that in the present state

of knowledge [of the] P. glandulosa aggregate it will be best to give this form a distinct binomial. If you

f4L,D o ID L

40 Journal of t titute of Texas 1(1)

wish, therefore, I shall be glad to prepare a description for publication, making your specimen the type of a

new species.” One draft of this letter (presumably the one sent) then adds the caveat that she first needs to

examine the recently published Potentilla brevifolia var. perserverans A. Nelson. Crum never followed through

on describing Peirson’s collection, even when supplemented with additional collections, evidently deferring

to David Keck and the contemporaneous biosystematics studies on the Potentilla glandulosa complex (letter

of 5 May 1938). Peirson gave her his support, noting that “You have probably put more work and thought

on Potentilla than anyone else in the state and it is only just and fitting that you proceed with a synopsis of

the genus in spite of publications by others” (letter of 5 Oct 1940). In response, Crum (letter of 8 Oct 1940)

indicated that she did not feel prepared to revise the genus, but agreed to return to Peirson 11272, “which I

examined several years ago and then laid aside.” She apologized for holding it up so long, and promised to

send a “definite opinion” within the next two weeks.

Instead, Crum sent the specimen to Keck, “who was very much interested considering it was quite

aberrant" (letter of 29 Oct 1940). Keck, who had just published his revision of the P. glandulosa complex (in

Clausen et al. 1940), proposed to name the new taxon after Crum, which Peirson heartily endorsed (letter

of 14 Dec 1940). Crum, however, urged Keck not to do so *not only because I don't deserve the honor but

because my name is not euphonious" (letter of 20 Dec 1940). She hoped instead that Keck would use some

variant of Peirson's name, though there already was a P. peirsonii Munz (= Drymocallis cuneifolia Rydb.).

Following Crum's untimely death in January 1943, her colleague Annetta Carter wrote to Peirson about a

letter received from Keck, in which he indicated his intent to use the epithet crumiana: *Miss Crum always

objected to any publicity, but Dave feels that considering all the work that she did on Potentilla, she certainly

deserves to have her name commemorated” (letter of 26 Oct 1944).

For whatever reason, Keck likewise never followed through on his intent to describe this distinctive

taxon. I take great pleasure in adopting the suggestion of Keck, Peirson, and Carter in giving much-deserved

recognition to one of my predecessors in Potentilla research, and finally giving a name to Peirson's discovery

over 70 years after its original collection. Subsequent collections show the taxon to be both more widespread

and perhaps less distinctive than initially believed. The red-tinged petals that first caught Crum's atten-

tion are not reliably present, but the leaflet characters provide a distinctive gestalt not found elsewhere in

Drymocallis, though approaching D. fissa in Colorado.

OTHER SPECIMENS EXAMINED: UNITED STATES. CALIFORNIA. Fresno Co.: Kings Canyon National Park, Glenn's Pass Trail, 10,580-11,900

ft, 29 Jul 1948, H. & V. Bailey 2736a (UC), Kearsarge Pass, 11,500 ft, 15 July 1900, W.L. Jepson 859 (JEPS); SE corner of Mt. Goddard, very

steep stair-step climb, 12,350-12,750 ft, 15 Aug 1957, C.H. Quibell 6933 (JEPS, RM); W toe of peak facing Goddard Canyon, 12,438 ft,

17 Aug 1957, C.F. Quibell & A. McCallum 7047 (UC). Inyo Co.: Kearsarge Pass trail, 10,500 ft, 7 Aug. 1942, A.M. Alexander & L. Kellogg

3272 (DS, NY, UQ); Seventh Lake, Big Pine Lakes, 11,200 ft, 9 Aug 1947, J.T. Howell 23963 (CAS); Inconsolable Range above Big Pine

Lakes, 12,400 ft, 14 Aug 1947, J.T. Howell 24093 (CAS, DS, UC); E side of Kearsarge Pass, 11,500 ft, 20 Jul 1948, J.T. Howell 24790 (CAS);

slope above N Fork of Oak Creek between Pk. 12606 and Pk. 10643, 11,500 ft, 6 Aug 1948, F.L. Jones s.n. (DS); head of Thibaut Creek,

vicinity of Mt. Baxter, 10,600 ft, 16 Oct 1948, F.L. Jones s.n. (DS); Kearsarge Trail, 11,000 ft, 16 Aug 1936, M. Kerr s.n. (SBBG); Rock Creek

Lake basin, cirque on east slope, 11,000 ft, 19 Jul 1931, F.W. Peirson 9515 (UC); Rock Creek Lake Basin, rocky slopes above East Recess,

11,400 ft, 11 Aug 1937, F.W. Peirson 12195 (JEPS); Taboose Pass trail, 11,200 ft, 16 Jul 1977, D.W. Taylor 6604 (JEPS). Mono Co.: White

Mountains, cirque heading N Fork of Perry Aiken Creek, 0.75 mi due E of White Mt. Peak, 11,900 ft, 25 Jul 1987, J.D. Morefield & T.S.

Ross 4701 (MO, NY, UC). Tulare Co.: near summit Arroyo-Kern Divide, region of Kaweah Peaks, 4 Aug 1897, W.R. Dudley 2417 (DS); Nine

Lakes Basin, head of Big Arroyo, cirque just north of Black Kaweah, 11,500 ft, 18 Aug 1938, C.W. Sharsmith 3792 (DS, MO, NY, UC); W

slope Kaweah Peaks Ridge above Nine Lakes Basin, head of Big Arroyo, 12,100 ft, 20 Aug 1938, C.W. Sharsmith 3805 (CAS, UC).

10. Drymocallis glabrata Rydb., Mem. Dept. Bot. Columbia Coll. 2:201. 1898. = Potentilla glandulosa Lindl. subsp.

glabrata (Rydb.) D.D. Keck; P glandulosa Lindl. var. intermedia (Rydb.) C.L. Hitchc.

Potentilla glandulosa D incisa Lindl. (Bot. Reg. t. 1973. 1837) is in all likelihood another synonym for D. glabrata,

although Keck (in Clausen et al. 1940: 48) concluded that var. incisa *is beclouded with doubt, and does not

stand for any of the preceding natural units [of P. glandulosa].” In his earlier revision of Horkelia and Ivesia,

Keck (1938, p. 83) stated that P. glandulosa B incisa was “surrounded by confusion and should doubtless be

rejected." Keck's conclusions at first seem justified: most collections labeled as corresponding to Tab. 1973

in Edward's Botanical Register, including that annotated by Keck as holotype of P. glandulosa B incisa in the

Ertter, The tribe P tentill ID llis in North America 41

Lindley Herbarium at Cambridge (CGE), consist of Horkelia californica subsp. frondosa (Greene) Ertter. This

taxon, which grows in central California, is superficially similar to the image in Tab. 1973 but differs in

significant regards, including petal color (white, not yellow). During my visit to Cambridge in 2002, how-

ever, I found a different collection of “B.R. 1973,” also from the Lindley Herbarium, filed as an unidentified

Potentilla. This sheet, which is an excellent match for the illustration and description of P. glandulosa B incisa,

has the diagnostic features of D. glabrata: i.e., leafy spreading inflorescence, large yellow petals, and septate

glandular hairs as the dominant vestiture. The purported California origin of P. glandulosa B incisa remains

a problem, but Douglas also collected within the range of D. glabrata in eastern Oregon. Given the potential

for mix-ups in collections and data following Douglas's death in 1834, as evident in the confusion with H.

californica subsp. frondosa, it is realistic to suspect that the specimens on which Tab. 1873 were based may

have originated in eastern Oregon, within the range of D. glabrata.

11. Drymocallis deseretica Ertter, sp. nov. (Fig. 2). Tv: UNITED STATES. Uran: Summit Co.: Uinta Mts., in the notch of

Notch Mt., 28 km (17.5 mi) airline distance E (75°) of Kamas, T1S R9E S30 (NE1/4), 3250 m (10,650 ft) elev.; locally common on rocky

W slope of notch, 22 Aug 1984, N.H. Holmgren & PK. Holmgren 10750 (HoLoTYPE: UC 1583251; isotypes: BRY 342384, NY, RM).

doe CAERE vs | +s ; +] TEES YE 19 4 Je TS y SED :

AD.g I g , pedicellis velutinis differt.

Herbaceous perennial from + branched caudex. Stems few, erect, (1.5-)2.5-66.5) dm, the base 2-3(-4)

mm diam., with sparse to abundant septate glandular hairs to 2(-3) mm long, inconspicuous short eglandular

hairs sparse or lacking. Basal leaves (5-)7-20 cm long with (2—)3 pairs of lateral leaflets, sometimes with

a much-reduced fourth pair; sheathing leaf-bases glabrous or + glandular, rarely sparsely strigose; vestiture

of rachis similar to that of stem; terminal leaflet + petiolulate, the blade obovate to rhombic with an acute

to obtuse apex, (1.5-)7-20 cm long, (1-)1.5-3 cm wide, both surfaces with a sparse mixture of subsessile

glands, short glandular trichomes, and simple hairs + 0.5 mm long, the margins single- to + double-toothed

with 5-9 teeth per side. Cauline leaves 1-2, sometimes equaling or exceeding basal leaves, 3-15 cm long

with 2-3 pairs of lateral leaflets. Inflorescence relatively compact, comprising (%—)'5—'2 of plant height,

relatively leafy; subtending bract trilobed, rarely pinnate, 2-7 cm long; branches diverging at a 10—20? angle;

pedicels 2—15(-20) mm long, with abundant short spreading hairs + 0.2 mm long and sparse to moderately

abundant septate glandular hairs. Flowers 3-15(-20); hypanthium shallowly saucer-shaped, 4—7 mm diam.,

with abundant short simple hairs + 0.2 mm long and + sparse glandular trichomes; epicalyx bractlets linear to

lanceolate or narrowly elliptic, (2.5-)3—8 mm long, 0.5-2(3) mm wide, rarely notched; sepals 65-26-12 (15)

mm long, the vestiture like that of hypanthium or more sparse, the apex acute (to obtuse); petals cream to

pale yellow, narrowly to broadly obovate, (4-)6-10 mm long, (2.5—)3.5-6(-7) mm wide, equaling or more

often shorter than mature sepals, the apex rounded; stamens 20-25, the filaments 1-3 mm long, the anthers

+ elliptic; styles very thickly fusiform, slightly > 1 mm long, golden-brown, attached just below middle of

achene. Achenes + obliquely pyriform with a slight beak, 1.2-1.5 mm long, light brown.

Ecology and phenology.— Openings among sagebrush, aspen, fir, and/or spruce, often where moist or

rocky, and below cliffs, 2000-3250 m elev. Flowering (May) Jun-Sep.

Drymocallis deseretica is common in the northern Wasatch and western Uintah mountains of Utah,

where it has generally been treated as Potentilla glandulosa var. intermedia (= D. glabrata) or P. glandulosa var.

pseudorupestris (2 D. pseudorupestris var. saxicola). It differs from both species in its more compact inflores-

cence, velutinous pedicels, and conspicuously enlarged acute sepals, which create a very distinctive gestalt.

Plants with a comparable gestalt, but shorter obtuse sepals, occur in the Raft River Mountains of Utah. The

species intergrades with D. glabrata to the north and D. arizonica to the south, with the exact range not yet

determined. Petal color is rarely recorded on labels and needs to be confirmed.

REPRESENTATIVE SPECIMENS: UNITED STATES. UTAH: Cache Co.: Intervale, 1 mi from mouth of Pine Canyon, Wellsville Range, 19 Jun

1932, B. Maguire 2991 (UC). Duchesne Co.: W fork of Duchesne River, 44 mi E of Heber, TIN R7E S33, 27 Jul 1978, D. Atwood 7018

(BRY); Uinta Mts., head of Duchesne River near Mirror Lake, 4 Aug 1980, S. Goodrich 14717 (BRY). Juab Co.: Mt. Nebo, jct. Willow Creek

and Cottonwood Campground trails, 23 Jul 1976, F. Peabody 645 (BRY). Salt Lake Co.: Bells Canyon, near reservoir, 20 Jul 1959, W.P.

Cottam et al. 15802 (WIS); Alta Pass above Twin Lake, 7 Aug 1939, B. Maguire 17363 (UC); above Salamander Lake, Lamb's Canyon, 10

h Institute of Texas 1(1)

J Lafthn Dat

Journal of

= zm AA

y px -

7 Zi /

ti A. habit R: infl

Ertter, The tribe P tentill ID llis in North America 4

May 1958, R.K. Vickery 2073 (UO). Sanpete Co.: top of Log Hollow Canyon, San Pitch Mts., 3 Jul 1979, R. Collins 301 (BRY); 13 km air

distance ENE of Fairview, Hwy 30, upper Gooseberry Creek drainage, 19 Jun 1985, N.H. Holmgren et al. 10880 (UO); Upper Horseshoe

Flat, 28 Jul 1962, H. Johnson s.n. (BRY). Summit Co.: Stillwater Basin, head Bear River, Uinta Mts., 18 Aug 1933, B. Maguire et al. 4172

(UC); due W of parking lot at head of Bald Mt. Trail, summit of Mirror Lake Pass, 1 Aug 1970, P.A. Replogle 200 (BRY). Tooele Co.: Mill

Fork of South Willow Canyon, Stansbury Mts., T4S R7W S14, 4 Jul 1980, A. Taye 990 (BRY). Utah Co.: Pika Cirque, Mt. Timpanogos,

12 Jul 1974, K. Allred 1027 (BRY); along Timpanogos trail between 1* and 2™ falls, 30 Jun 1964, A. C. Blauer & J. Brotherson 44 (BRY);

Little Cottonwood Canyon, Albion Basin campground, 20 Aug 1984, N.H. Holmgren & P,K. Holmgren 10721 (UC); Provo, 8000 ft., 3 Jul

1894, M.E. Jones 5575 (UC); 1 mi NE of Springville Crossing on Diamond Fork Creek, T7S R6E S31, 21 Jun 1969, J.W. Thomas 155 (BRY).

Wasatch Co.: Strawberry Valley, T3S R12W, 16 Jul 1964, V.B. Matthews 89 (BRY).

12. Drymocallis glandulosa (Lindl.) Rydb., Mem. Dept. Bot. Columbia Coll. 2:198. 1898. = Potentilla glandulosa

Lindl.

The original illustration of Potentilla glandulosa in Edwards's Botanical Register (19: pl. 1583. 1833), drawn

from living material “in the Chiswick Garden in August last,” has bright yellow petals. It is therefore prob-

able that David Douglas collected the seeds from which the specimen was grown somewhere in Oregon,

since coastal populations in California (treated here as var. wrangelliana) are predominantly white-petaled.

Variation in D. glandulosa s.s. outside of California is unclear, with no evident pattern to differences in petal

size and color; as default, these have largely been assigned to var. glandulosa. Sporadic collections have been

made in Idaho, Montana, Nevada, and Utah; disjunct populations in Arizona south of the Mogollon rim

may represent an undescribed variety.

12a. Drymocallis glandulosa var. glandulosa

12b. Drymocallis glandulosa var. wrangelliana (Fisch. & Avé-Lall.) Ertter, comb. nov. Basionvm: Potentilla

wrangelliana Fisch. & Avé-Lall., Ind. Seminum Hort. Petrop. 7:54. 1841.

12c. Drymocallis glandulosa var. reflexa (Greene) Ertter, comb. nov. Basioxvw: Potentilla glandulosa Lindl. var. reflexa

Greene, Flora Francisc. 65. 1891. - P glandulosa subsp. reflexa (Greene) D.D. Keck.

A disjunct collection from a major mining area in central Idaho (Boise Co., Clear Creek-Grimes Creek, 28

Jun 1959, L. Maydale s.n., CIC) most likely results from dispersal via the transport of mining equipment from

California, parallel to a comparable speculation for Eriogonum inerme (S. Watson) Jeps. (Ertter & Moseley

1993). The variety, with its small reflexed yellow petals, is otherwise known only from the mountains of

California and southern Oregon.

12d. Drymocallis glandulosa var. viscida (Parish) Ertter, stat. et comb. nov. Basioxvm: Drymocallis viscida Parish,

Bot. Gaz. 38:460. 1904.

This overlooked variety combines the flower and vestiture of Drymocallis glandulosa var. reflexa with the

narrow inflorescence and predominately single-toothed leaflets of D. lactea var. lactea.

13. Drymocallis campanulata (C.L. Hitchc.) Ertter, stat. et comb. nov. Basionyw: Potentilla glandulosa var. campanulata

C.L. Hitchc., Vasc. Pl. Pacific NorthW. 1:861. 1969.

The epithet *campanulata" was used provisionally as a species of Potentilla in the treatment of Rosaceae

(Hitchcock & Cronquist 1961) in volume 3 of Vascular Plants of the Pacific Northwest, “in anticipation of its

publication by Dr. Keck,” but the validating publication in “Additions and Corrections" in volume 1 (Hitch-

cock et al. 1969) treated the new taxon as a variety of P. glandulosa. This entity was referred to as the John

Day race in Clausen and Hiesey (1958).

14. Drymocallis rhomboidea (Rydb.) Rydb., Mem. Dept. Bot. Columbia Coll. 2: 203. 1898. = Potentilla rhom-

boidea Rydb.; P glandulosa Lindl. subsp. globosa D.D. Keck.

Keck (in Clausen et al. 1940) identified the type of P. rhomboidea as P. glandulosa subsp. glandulosa and

therefore described P. glandulosa subsp. globosa to accommodate this distinctive taxon.

44 Journal of the Botanical R h Institute of Texas 1(1)

15. Drymocallis cuneifolia Rydb., Mem. Dept. Bot. Columbia Coll. 2:204. 1898. = Potentilla cuneifolia (Rydb.) Th.

Wolf, not P cuneifolia Bertol.; P peirsonii Munz.

Rydberg (1898) described Drymocallis cuneifolia on the basis of a collection by Samuel B. Parish in the San

Bernardino Mountains of San Bernardino County, California, “probably near Green Lead Mines,” in June

1886 (F, GH, NY). The collection, distributed as Potentilla glandulosa var. nevadensis, was numbered and

cited as Parish 1818 (which is also the number given to a collection of Phlox dolichantha A. Gray, another rare

plant endemic to the San Bernardino Mountains). Rydberg noted that the species “differs from the others

in the small flowers with erect petals and the cuneate-flabelliform leaflets"; it also had filiform styles that

are very unusual in Drymocallis. When comparable (though much smaller) specimens were found in 1919

by F. W. Peirson in the San Gabriel Mountains in Los Angeles County, Munz and Johnston (1925) adopted

Potentilla cuneifolia (Rydb.) Th. Wolf for both forms. Since this name was a later homonym of P. cuneifolia

Bertol., however, Munz (1932) renamed the species P. peirsonii, noting that Peirson had *rediscovered a plant

that had not been collected for many years."

The continued lack of comparable new collections from the San Bernardino Mountains led Keck (in

Clausen et al. 1940) to conclude that the type of Drymocallis cuneifolia was merely an immature specimen of

Potentilla glandulosa subsp. nevadensis (= D. lactea var. lactea). As a result, he described P. glandulosa subsp.

ewanii to accommodate populations in the San Gabriel Mountains. He did, however, note that *the type of

D. cuneifolia is fragmentary, and since neither Ewanii nor cuneifolia is well represented in herbaria as yet, the

possibility exists that the differences which now appear impressive may eventually loose importance."

Serendipitously, a small population of plants comparable to the type of Drymocallis cuneifolia has

recently been discovered in the San Bernardino Mountains, not far from Parish's original collection: San

Bernardino Co., San Bernardino Mts., Grout Creek, NE of Butler Peak, 200 m W of creek crossing along FS

road 2N13, 0.3 mi W of FS road 2N70, 34?16'08"N, 116°58'40"W, 2195 m/7200 ft, 30 June 2004, Mark A.

Elvin 3555 (IRVC, UCR). These plants were identified as P. glandulosa subsp. ewanii and have the diagnostic

small narrow corollas and elongate styles of this taxon. These features, however, also characterize the type

of D. cuneifolia, which was collected in the same general area. Both Elvin 3555 and the type of D. cuneifolia

are larger plants than is the norm in the San Gabriel Mountains, with more elongate leaflets. As a result,

I am treating the San Bernardino Mountain and San Gabriel Mountain populations as varieties of a single

species, D. cuneifolia.

15a. Drymocallis cuneifolia var. cuneifolia

As indicated above, this variety is currently known from a single extant population and must therefore be

considered extremely rare and potentially threatened. Although not known from limestone substrates, its

occurrence on alluvial benches in an area known for a cohort of federally listed carbonate endemics (e.g.,

Erigeron parishii A. Gray, Astragalus albens Greene) opens the possibility that D cuneifolia var. cuneifolia might

have similar limitations.

15b. Drymocallis cuneifolia var. ewanii (D.D. Keck) Ertter, stat. et comb. nov. Basioxwx: Potentilla glandulosa Lindl.

subsp. ewanii D.D. Keck, Carnegie Inst. Wash. Publ. 520:47. 1940.

ACKNOWLEDGMENTS

Foremost I wish to thank the curatorial staff at the multiple herbaria who prepared loans used in this study,

tracked down additional information upon request, remained patient with the length of time the specimens

have been in my hands, and generally went the extra mile in their curatorial obligations: ALA, ARIZ, ASC,

ASU, BRY, CAS/DS, F, GH, ID, MICH, MO, NDG, NY, OSC, RM, RSA/POM, SBBG, UCR, US, UTC, WIS,

WTU. I also wish to acknowledge the curatorial staff at UC/JEPS, who have processed my seemingly un-

ending loans; and the courtesy offered me at other herbaria that I visited and used during the course of this

study: BM, CGE, CIC, K, LE, PR, SRP, W. I am indebted to Jiří Soják (PR) for freely sharing his extensive

knowledge and insights on Potentilleae, and to Adolf Ceska, Lenka Drabkova, and the late Otto Winkler for

Ertter, The tribe P tentill ID llis in North America 45

serving as translators and intermediaries. I wish to thank Kanchi Gandhi (GH) for deciphering a steady

stream of nomenclatural tangles; J.G. Murrell (CGE) for checking vestiture on the type of Potentilla glandulosa

B incisa; Dick Brummitt (K) and John McNeill (E) for hosting me during my visits to British herbaria; and

Mark Brunell (CPH) and Mark Elvin (IRVC) for bringing my attention to the new collection of Drymocallis

cuneifolia. The excellent illustrations by Linda Vorobik are greatly appreciated, as always, and support from

the Lawrence R. Heckard Endowment Fund of the Jepson Herbarium is gratefully acknowledged.

Note added in proof.—In a recently published paper, Soják (2006) provides new combinations in Dry-

mocallis of all subspecies of the Potentilla glandulosa complex as recognized by Keck (in Clausen et al. 1940),

with no change in rank or taxonomic arrangement.

REFERENCES

AIKEN, S.G., M.J. DaLtwrrz, L.L. CoNsAuL, C.L. MCJANNET, L J. GiLLEsPiE, R.L. Botes, G.W. Arcus, J.M. GiLLETT, PJ. Scott, R. ELVEN,

M.D. LeBLanc, A.K. BRvsriNc, and H. HoLsTAD. 2006. Flora of the Canadian arctic archipelago. [www.mun.ca/biol-

ogy/delta/arcticf/]

BALDWIN, B.G. 1999. New combinations and new genera in the North American tarweeds (Compositae-Madiinae).

Novon 9:462-471.

Brumm, R.K. 2002. How to chop up a tree. Taxon 51:31-41.

CHRTEK, J. and J. SoJAk. 1984. Farinopsis, nový rod čeledi Rosaceae. Cas Nar. Muz., Rada Pří. 153:10.

CLAUSEN, J., D.D. Keck, and W.M. Hiesey. 1940. Experimental studies on the nature of species. |. Effect of varied envi-

ronments on western North American plants. Carnegie Inst. Washington Publ. 520:1-452.

CLAUSEN, J. and W.M. Hiesey. 1958. Experimental studies on the nature of species. IV. Genetic structure of ecological

races. Carnegie Inst. Washington Publ. 615:1—312.

Dices, G.M., Jr. and B.L. Liescome. 2002. What is the writer of a flora of do? Evolutionary taxonomy or phylogenetic

systematics? Sida 20:647-674.

ELvEN, R., D.F. Murray, V. RAZZHIVIN, and B.A. Yurtsev (eds.). 2007. Checklist of the panarctic flora (PAF). Vascular plants.

Final draft version. Oslo: University of Oslo.

ERIKSSON, T. and M.J. DonoGHuE, 1995. Is Potentilla monophyletic? a phylogenetic analysis of the Rosoideae (Rosa-

ceae) using nuclear ribosomal ITS sequences. Amer. J. Bot. 82(supplement):128 (abstract 366).

Eriksson, T., M.J. DoNocHuE, and M.S. Higgs. 1998. Phylogenetic analysis of Potentilla using DNA sequences of

ribosomal internal transcribed spacers (ITS), and implications for the classification of Rosoideae (Rosaceae).

Pl Syst Evol 211: 1555179.

Eriksson, T., M.S. Hipes, A.D. Yoper, C.F. DeLwichHe, and M. DonocHue. 2003. The phylogeny of Rosoideae (Rosaceae)

based on sequences ofthe internal transcribed spacers (ITS) of nuclear ribosomal DNA and the TRNL/F region

of chloroplast DNA. Intern. J. Pl. Sci. 164:197-21 1.

ERTTER, B. 1989. Revisionary studies in /vesia (Rosaceae: Potentilleae). Syst. Bot. 14:213-244.

ErTTER, B. 1993. Horkelia, Horkeliella, lvesia, and Potentilla. In: J.C. Hickman, ed. The Jepson manual: higher plants

of California. University of California Press, Berkeley. Pp. 953-969,

ErTTER, B. 1997. Taxonomic grays vs. black and white expectations: Implications for conservation and management

of diversity. In: T.N. Kaye, A. Liston, R.M. Love, D.L. Luoma, RJ. Meinke, and M.V. Wilson, eds. Conservation and

management of native plants and fungi. Native Plant Society of Oregon, Corvallis, Oregon. Pp. 11-13.

ErTTER, B., C. BaysporFEr, and D. Alonso. 1998. Monophyly of the ivesioid Potentilleae (Rosaceae) based on ITS

sequence data [abstract]. Amer. J. Bot. 85:126-127.

ErTTER, B. and B. Mosetey. 1993 [1992]. Floristic regions of Idaho. J. Idaho Acad. Sci. 28:57-70.

Grimes, J.W. 1990. A revision of the New World species of Psoraleeae (Leguminosae: Papilionoideae). Mem. New

York Bot. Gard. 6:1-114.

HitcHcock, C.L., and A. CRoNQuisr. 1961. Rosaceae. Vasc. Pl. Pacific NorthW. 3:89- 194.

HitcHcock, C.L., A. CronquisT, and M. Ownsey. 1969. Additions and corrections. Vasc. Pl. Pacific NorthW. 1:857-861.

HOLMGREN, N. 1997. Rosaceae. Interm. Fl. 3A:64-158.

46 Journal of the Botanical R h Institute of Texas 1(1)

HORANDL, E. 2006. Paraphyletic versus monophyletic taxa-evolutionary versus cladistic classifications. Taxon

55:564—570.

HoweLt, J.T. 1949. Marin flora: manual of the flowering plants and ferns of Marin County, California. University of

California Press, Berkeley.

HUTCHINSON, J. 1964. The genera of flowering plants. Vol. 1. Clarendon Press, Oxford.

Jepson, W.L. 1925. A manual of the flowering plants of California. Associated Students Store, University of Cali-

fornia, Berkeley.

Jepson, W.L. 1936. A flora of California. Vol. Il. Capparidaceae to Cornaceae. Associated Students Store, University

of California, Berkeley.

Kearney, T.H. and RH. PeeBLEs. 1951. Arizona flora. University of California Press, Berkeley.

Keck, D.D. 1938. Revision of Horkelia and lvesia. Lloydia 1:75-142.

KurTTO, A. and T. Eriksson. 2003. Atlas Florae Furopaeae notes. 15. Generic delimitations and nomenclatural adjust-

ments in Potentilleae. Ann. Bot. Fennici 40:135-141.

Li, C.-L., H. IkeDa, and H. Onaa. 2003. Potentilla, Comarum, Sibbaldia, Chamaerhodos, Fragaria, and Duchesnea. Fl.

China 9:291-338.

Masser ey, D.J. 2002. Potentilla and Fragaria (Rosaceae) reunited. Telopea 9:793-801.

Mason, H.L. 1943. Ethel Katherine Crum. Madrono 7:33-35.

Munz, PA. 1932. Southern California plant notes—IV, Bull. S. Calif. Acad. Sci. 31:61-70.

Munz, PA. and I.M. JoHNston. 1925. The potentillas of southern California. Bull. S. Calif. Acad. Sci. 24:5-25.

Posar, J. 1999. Rosaceae. Ill. Fl. Brit. Columbia 4:258-370.

Reveal, J. L., G.E. Moutton, and A.E. ScHuvLER, 1999. The Lewis and Clark collections of vascular plants: names, types,

and comments. Proc. Acad. Natl. Sci. Philadelphia 149:1—64.

RYDBERG, PA. 1898. A monograph of North American Potentilleae. Mem. Dept. Bot. Columbia Coll. 2:1-223 + 112

plates.

RYDBERG, PA. 1908. Rosaceae. Tribe 7. Potentilleae. N. Amer. Fl. 22(3-4):268-377.

Soak, J. 1989 [1985]. Generická problematika Potentilla s.l. Cas Nar. Muz., Rada Pří. 154:117-118.

SoJAk, J. 2004. Potentilla L. (Rosaceae) and related genera in the former USSR (identification key, checklist and

figures). Notes on Potentilla XVI. Bot. Jahrb. Syst. 125:253-340.

SoJAk, J. 2006. New infraspecific nomenclatural combinations in twelve American species of Drymocallis and

Potentilla (Rosaceae). (Notes on Potentilla XVII). Thaiszia 16:47-50.

SrAUDT, G. 1999. Systematics and geographic distributions of the American strawberry species: taxonomic studies

in the genus Fragaria (Rosaceae: Potentilleae). Univ. Calif. Publ. Bot. 81:1-162.

WEBER, W.A. 1987. Colorado flora: western slope. Colorado Associated University Press, Boulder.

WEBER, W.A. 2003. The Middle Asian element in the southern Rocky Mountain flora of the western United States:

a critical biogeographical review. J. Biogeogr. 30:649-685.

Weser, W.A. and R.C. Wittmann. 1992. Catalog of the Colorado flora: a biodiversity baseline. University Press of

Colorado, Boulder.

WELSH, S.L., N.D. Arwoob, S. Goobrich, and L.C. HicaiNs. 1993. A Utah flora. 2^? Edition. Brigham Young University

Press, Provo.

Wotr, T. 1908. Monographie der Gattung Potentilla. Bibl. Bot. 16(71):1-713 + 22 plates.

YurTZEV [Jurtsev], B.A. (ed). 1984. Rosaceae. Fl. Arctica URSS 9(1):1-332.

LECTOTYPIFIGATIONS AND NEW TAXAIN POTENTILLA SECT

SUBVISCOSAE (ROSACEAE) IN ARIZONA

Barbara Ertter

University and Jepson Herbaria

University of California

Berkeley, California 94720-2465, U.S.A.

ABSTRACT

Potentilla sect. Subviscosae (Rydb.) B.C. Johnston exemplifies radiation in the montane Madrean Archipelago of Mexico and

the American Southwest, with multiple localized taxa endemic to different mountains ranges. Lectotypification of Potentilla ramulosa

Rydb. maintains usage as the variety of P. subviscosa Greene in the Santa Catalina and Rincon mountains, Arizona. Potentilla wheeleri

var. viscidula Rydb. is lectotypified on the hybrid between P. ramulosa and a trifoliate species in the Huachuca and Santa Rita mountains

of Arizona, with the latter species newly described as P. rhyolitica Ertter. Closely related plants from the Chiricahua Mountains are

recognized as P. rhyolitica var. chiricahuensis Ertter. An extremely localized Potentilla from the Hualapai Mountains, Arizona, is

described as P. demotica Ertter. The range of P. albiflora (which has yellow flowers, not white) is extended to the the nearby Mogol-

lon Rim. Potentilla cottamii is moved to P. sect. Subviscosae from P. sect. Aureae. Potentilla wheeleri is restricted to southern California,

excluding P. luteosericea (= P. pinetorum) as a separate species in Baja California, Mexico. A key to the non-Mexican species is provided

and vernacular names are suggested.

Key Wonops: Potentilla sect. Subviscosae, Madrean Archipelago, Huachuca Mountains, Hualapai Mountains pifications, new species,

Potentilla subviscosa, Potentilla ramulosa, Potentilla viscidula, Potentilla albiflora, Potentilla wheeleri, Potentilla cottamii, Potentilla luteosericea

RESUMEN

La sección Subviscosae (Rydb.) B.C. Johnston de Potentilla ejemplifica una radiación evolutiva en el Archipiélago montañoso Madrense

de México y el suroeste de Estados Unidos, representado con multiples taxa locales y endémicos en los diferentes si montañosos.

La lectotipificación de Potentilla ramulosa Rydb. mantiene su uso como variedad de P. subviscosa Greene de las montañas Santa Catalina y

Rincon en Arizona. Se lectotipifica Potentilla wheeleri var. viscidula Rydb. como un híbrido entre P. ramulosa y una especie trifoliolada en

las montañas Huachuca y Santa Rita de Arizona, A Me ha sido descrita recientemente como P. rhyolitica Ertter. Plantas cercanamente

relacionadas de las Montañas Chiricahua son recon mo P. rhyolitica var. chiricahuensis Ertter. Una Potentilla de distri

limitada en las Montañas Hualapai, Ari d ib P. demotica Ertter. I | fl ill

y no blancas) se extiende al cercano Mogollon Rim. Potentilla cott iota a ld S O ea A EE Ae E A

circunscribe su distribución al sur de California, excluyendo a P. luteosericea (= P. pinetorum) como una especie aparte en Baja California,

México. Se provee con una clave para identificar las especies que no están en México y se sugieren sus nombres comunes

INTRODUCTION

In his 1898 monograph of Potentilla L. (Rosaceae), Rydberg divided the genus into multiple “groups,” many

of which were adopted and established at formal taxonomic ranks by later workers (e.g., Johnston 1985).

Among the more intriguing is P. sect. Subviscosae (Rydb.) B.C. Johnst., initially comprised of Potentilla wheeleri

S. Watson, P. wheeleri var. viscidula Rydb., P. subviscosa Greene, and P. ramulosa Rydb. (Rydberg 1896, 1898).

These taxa and their subsequently described relatives, all endemic to the southwestern United States and

Mexico, share the following characters: 1) stems prostrate to decumbent, or even pendant on vertical rock

faces, relatively short and lax; 2) vestiture of entire plant consisting of various proportions of short glandu-

lar trichomes, short eglandular hairs ca 0.2 mm long, and long straight eglandular hairs 1-3 mm long; 3)

stipules narrowly triangular to linear, + herbaceous; 4) leaves 3—7-digitate, the leaflets neither tomentose

beneath nor notably bicolored, few-toothed; 5) pedicels relatively lax, often becoming recurved; 6) styles

(0.7-)1.5-3 mm long, usually somewhat rough-thickened basally; 7) achenes relatively few (4-25) and large

(+ 1-2 mm long), smooth to lightly ribbed.

What makes Potentilla sect. Subviscosae intriguing is the extent to which it exemplifies evolutionary

radiation in an island setting, with the “islands” in this case consisting of scattered mountain ranges isolated

J. Bot. Res. Inst. Texas 1(1): 47 — 57. 2007

48 Journal of the Botanical R h Institute of Texas 1(1)

by arid lowlands. The center of radiation for P. sect. Subviscosae, at least in the United States, is the isolated

mountains extending north from Mexico's Sierra Madre Occidentale, dubbed the Southwestern Sky Island

Ecosystem or the Madrean Archipelago (e.g., McLaughlin 1995). As a result, different unique and highly

endemic members of the section occur on the Pinaleno, Santa Catalina/Rincon, Huachuca/Santa Rita, and

Chiricahua ranges in southeastern Arizona, including taxa described in the present paper. This radiation

presumably also characterizes the more extensive Mexican portion of the “archipelago,” but collections and

taxonomic analysis of Potentilla from this region are too preliminary for a proper evaluation. Potentilla mexiae

Standl. is unquestionably a member of this section in Chihuahua and Durango, and there is evidence of

additional undescribed variation deserving taxonomic recognition.

A full-scale revision of the entire section is strongly called for, with special attention paid to species in

Mexico including those currently placed in Potentilla sections Aureae (Lehm.) Juz. and Ranunculoides (Th.

Wolf) Juz. However, the current paper focuses on those lectotypifications and new descriptions needed for

the pending treatment of Potentilla for Flora of North America North of Mexico, with some additional notes on

several species of interest in the United States.

LECTOTYPIFICATIONS

In his initial review of Potentilla “group” Subviscosae in 1896, Rydberg described Potentilla wheeleri var. viscidula

(p. 429) and P. ramulosa (p. 430 and plate 276). The former was based on four syntypes, with two from Arizona

(*C.G. Pringle, 1881; J.G. Lemmon, no. 158. 1881") and two from California (^W.G. Wright, 1879; Coville

& Funston, no. 1672. 1891"). Two syntypes were given for P. ramulosa, both from Arizona: *J.G. Lemmon,

no. 399, 1881; H.H. Rusby, 1883.” The corresponding herbarium specimens currently known to me are:

Syntypes of Potentilla wheeleri var. viscidula

* C.G. Pringle, s.n., Arizona [Santa Cruz Co. or Pima Co.], Santa Rita Mountains, alt. 8,000 ft, 3 May 1881 (distributed as Potentilla

subviscosa Greene “near P wheeleri, Watson.")

H 26877, mounted on same sheet as Lemmon 158 syntype (GH 26878), both annotated by Rydberg (“PA.R.”) as “P wheeleri var.”

* 3 sheets at NY, all annotated in Rydbergs hand as P viscidula, plus an unannotated fourth duplicate (NY 39334) mounted with an

isotype of P subviscosa Greene (NY 39333)

* 2 sheets at US, only one annotated by Rydberg

* ] sheet at MO, not annotated by Rydberg

* J.G. Lemmon 158, Arizona [Pima Co.], summit of Santa Catalina Mountains, Apr 1888

* GH 26878, mounted on same sheet with Pringle syntype (GH 26877)

* WG. Wright, s.n., California [San Bernardino Co.], Mt. San Bernardino, 12,000 ft alt., Jun 1879

* GH 26880, in packet mounted on same sheet with holotype of P wheeleri (Rothrock 324, California, southern Sierras, 8200 ft, Sept

1875) and 3 non-type collections of P wheeleri s.s. (Parry & Lemmon 100; Parish & Parish 1498)

* EV Coville & E Funston 1672, California [Tulare Co.], Whitney Meadows, Sierra Nevada, 21 Aug 1891

* GH 26879, not annotated by Rydberg

Syntypes of Potentilla ramulosa

* J.G. Lemmon 399, Arizona [Cochise Co.], Rucker Valley [in southeastern Chiricahua Mountains], 1881

* GH 244121, annotated as P ramulosa by Rydberg, originally determined as P gracilis var. rigida, on mounted half-sheet

* H.H. Rusby 591 (on 3 of 4 sheets). Arizona, Flagstaff, 1883

* 3 sheets at NY with a date of Jun 7, one annotated by Rydberg

* 1 sheet at NY with a date of September

An unnumbered Lemmon collection from the north slopes of the Santa Catalina Mountains, April and May

1881 (MO, UC), is not syntype material, even though Rydberg annotated the MO specimen as P. wheeleri

viscidula n.v.

In the upcoming treatment of Potentilla for Flora of North America, both Lemmon 158 (syntype of P.

wheeleri var. viscidula) and Lemmon 399 (syntype of P. ramulosa) fall within P. subviscosa var. ramulosa (Rydb.)

Kearney & Peebles. The Pringle syntype is a mixed collection, with P. subviscosa var. ramulosa being the

largest component. Among the multiple exsiccatae, at least one plant (i.e., on the sheet at MO) is the trifoli-

ate species described below as P. rhyolitica Ertter, and some others are probable hybrids between the two

Ertter, Lectotypificati | new taxa in Potentilla 49

species. The other syntype of P. ramulosa, Rusby 5191, consists of both early-season (June) and late-season

(September) forms of P. subviscosa var. subviscosa, which exhibits a pronounced seasonal leaf dimorphism.

Both of the remaining syntypes of P. wheeleri var. viscidula, Coville & Funston 1672 and Wright s.n,. fall within

Potentilla wheeleri s.s., which is restricted to California as currently circumscribed (and as discussed later).

Wright s.n. represents the high-elevation extreme in the San Bernardino Mountains that Jepson (1925) named

P. wheeleri var. paupercula.

From the preceding, it is clear that application of the names Potentilla wheeleri var. viscidula and P. ramu-

losa is highly dependent on which syntype is chosen as lectotype for each taxon. As it happens, first-stage

lectotypification has already been done by Kearney and Peebles (1942), who cited Pringle s.n, 1881, asthe type

of P. viscidula (Rydb.) Rydb. and Lemmon 399 as the type of P. ramulosa. In that Kearney and Peebles explicitly

state that "In order to save indexers the labor of reviewing so large a work, no new names or combinations

are published here" (p. 5), it may very well be that they likewise did not intend any new lectotypications,

but this is nevertheless what they effectively accomplished (J. McNeill, pers. comm. 2006).

Fortunately, the selection of Lemmon 399 as the type of Potentilla ramulosa preserves current usage of the

epithet for the sole, commonly encountered representative of P. sect. Subviscosae in the heavily-visited Santa

Catalina Mountains, here treated as P. subviscosa var. ramulosa. In contrast, if Rusby 591 had been selected a

new name would be needed for plants in the Santa Catalina Mountains, now that the marked seasonal di-

morphism of leaf shape in the widespread P. subviscosa var. subviscosa is better understood, as discussed later.

Potentilla ramulosa Rydb., Bull. Torrey Bot. Club 23:430. 1896. Wee: U.S.A. ARIZONA. Cochise Co.: Chiricahua Moun-

tains, Rucker Valley, 1881, Lemmon 399 (Lectotype, designated by Kearney & Peebles, Fl. pl. ferns Ariz. 403. 1942: GH 244121!; the

only specimen known) = P subviscosa var. ramulosa.

Potentilla wheeleri var. viscidula presents a more complicated situation. The name was applied broadly at its

inception (Rydberg 1896), encompassing a diversity of specimens from southern Arizona and southern Cali-

fornia. In his subsequent treatment of the genus for North American Flora, Rydberg (1908) added Chihuahua,

Mexico, to the distribution of the taxon, possibly based on specimens which would now be identified as

P. mexiae. He also raised the taxon to species rank as P. viscidula (Rydb.) Rydb., a status and circumscrip-

tion followed by Tidestrom and Kittell (1941). In contrast, Kearney and Peebles (1942) initially treated P.

viscidula as a synonym of P. wheeleri, but in their subsequent flora (1951) they likewise accepted P. viscidula

as a distinct species (with the caveat, however, that it was “Perhaps too nearly related to P. Wheeleri Wats.").

Although these Arizona and New Mexico floras continued to include California in the distribution of P.

viscidula, California floras consistently recognized only P. wheeleri, sometimes citing P. viscidula in synonymy

(e.g., Jepson 1936; Abrams 1944; Munz 1959).

In spite of incompatibility problems with published distributions and descriptions, Potentil

a viscidula

has in practice been increasingly applied to a notably sericeous, trifoliate member of P. sect. Subviscosae that

occurs on rock outcrops along popular trails near the summit of the Huachuca and Santa Rita mountains of

southeastern Arizona, corresponding to the distribution in Arizona as given in Kearney and Peebles (1942,

1951). This species is clearly distinct from the 5-foliate species P. wheeleri and P. subviscosa, with the latter

also having a very different vestiture. My initial understanding, based on herbarium studies and personal

fieldwork, was that this trifoliate species was the only member of the section in these two mountain ranges,

with a less hairy variant occurring in the Chiricahua Mountains. It was only in the process of lectotypifica-

tion that I realized that P. subviscosa var. ramulosa also occurs in the Santa Rita Mountains (as the primary

element in the existing lectotype of P. wheeleri var. viscidula), the Huachuca Mountains (Goodding 1300, May

1912, ARIZ, NY, RM), and the Chiricahua Mountains (the lectotype of P. ramulosa itself). This odd situation,

whereby most historical collections differ from more recent collections from the same mountain ranges,

may reflect the fact that early collectors worked from existing bases at lower elevations (e.g., McCleary's

Ranch, now headquarters for the Santa Rita Experimental Range), whereas more recent botanists tend to

collect along a newer network of well-constructed trails that lead to the highest peaks. Both P. viscidula and

P. subviscosa are listed in a recent flora of the Huachuca Mountains (Bowers & McLaughlin 1996).

£+sha D o ID L

50 Journal of t titute of Texas 1(1)

As it happens, the lectotype of Potentilla wheeleri var. viscidula designated by Kearney and Peebles, or for

that matter any of the syntypes, is not compatible with continued use of the name P. viscidula for the trifoliate

species. Although the International Code of Botanical Nomenclature (McNeill et al. 2006) now provides multiple

options to conserve established nomenclature in order to avoid “disadvantageous nomenclatural changes”

(Art. 14.1), including the conservation of a name using a different type (Art. 14.9), I do not believe that

usage of P. viscidula for the highly localized trifoliate species is well enough established to justify conserva-

tion. The historical conflation with P. wheeleri has never ceased, with several recent publications treating P.

viscidula as synonymous with P. wheeleri (e.g., Medina 2003; Kartesz & Meacham 1999; PLANTS database

at http://plants.usda.gov/). Furthermore, the initial protologue only allows for plants with 5-7 leaflets in

sect. Subviscosae, and the epithet *viscidula" itself is incompatible with the sericeous vestiture of the trifoli-

ate species. For these reasons a fresh slate seems preferable, with the Kearney and Peebles lectotypification

retained and the trifoliate plants described below as P. rhyolitica.

One further complication is that the dominant element in the Pringle lectotype is the same taxon as the

lectotype of P. ramulosa, with the epithet viscidula having priority at the varietal level. To preserve established

usage of P. subviscosa var. ramulosa, I am therefore selecting one of the minor elements in Pringle's collection

as a second-stage lectotypification, specifically the largest of three plants on GH 26877. From its intermedi-

ate vestiture this plant is evidently a hybrid between P. subviscosa var. ramulosa and the sympatric trifoliate

species. The selected plant, on a sheet annotated by Rydberg, is an excellent match for Rydberg's protologue,

and its designation as lectotype removes the epithet viscidula from contention with the established epithet

ramulosa at varietal rank.

Potentilla wheeleri S. Watson var. viscidula Rydb., Bull. Torrey Bot. Club 23:429. 1896. Tee: U.S.A. ARIZONA

Santa Cruz Co. or Pima Co.: Santa Rita Mountains, Alt. 8000 ft, 3 May 1881, C.G. Pringle s.n. (Lecrotype (first stage designated by

Kearney & Peebles, Fl. pl. ferns Ariz. 402. 1942; second stage herein): largest of three plants on GH 26877!) = hybrid between P

subviscosa var. ramulosa and P rhyolitica var. rhyo

itica.

DESCRIPTION OF POTENTILLA RHYOLITICA AND VARIETIES

The drawback with the preceding lectotypification of Potentilla wheeleri var. viscidula is that the trifoliate

species is left without a name, a situation that is here rectified.

Potentilla rhyolitica Ertter, sp. nov. (Fig. 1). Tr: U.S.A. ARIZONA: Cochise Co.: Carr Peak in Huachuca Mts., rhyolite

outcrops near junction of Carr Peak and Miller Canyon trails, open forest of ponderosa pine, limber pine, Douglas-fir, oak, &

Holodiscus, ca. 9000 ft elev., T23S R20E Sec. 22, 3 Jun 1993, B. Ertter 11872 (HOLOTYPE: UC; isotypes: ALA, ASC, ASU, ARIZ, GH,

MEXU, MO, NY, PR, US).

Potentilla albiflora maxime simile, sed petioli pilis longioribus (1-3 mm vs. + 1 mm) et foliolorum dentibus paucioribus.

Plants tufted to rosetted, green to grayish, abundantly glandular. Stems prostrate, 0.272 dm long, with

abundant glandular trichomes and fine spreading hairs + 1 mm long. Leaves generally ternate, 2-8(-11)

cm long; petiole 1-6(-8) cm long with abundant glandular trichomes, short hairs, and spreading longer

hairs 1-2(-3) mm long; leaflets 3(—5), obovate, often petiolulate, the central one 1-3 cm long, toothed % to

midvein with 2—3(-4) teeth per side, moderately to densely sericeous on both sides. Inflorescences 1-10-

flowered; pedicels 0.5-1(-2) cm long, becoming recurved in fruit, with abundant short glandular trichomes,

short hairs + 0.2 mm long, and scattered longer hairs ca. 1 mm long. Flowers: hypanthium 3-5 mm diam;

epicalyx bractlets lanceolate-elliptic, 2-4 mm long; sepals 3-5 mm long, acute; petals yellow, + broadly el-

liptic-obovate to obcordate, 4-7 mm long, the apex rounded to shallowly emarginate; stamens 15-20, the

filaments 1.5-3 mm long, the anthers 0.6-1 mm long; styles 5-15, slender, scarcely rough-thickened basally,

2—3 mm long. Achenes light brown to reddish brown, 1.5-2.2 mm long, smooth to lightly ribbed.

Although previously associated with Potentilla wheeleri, that strictly Californian species has 5-7 leaflets.

A closer affinity exists with P. albiflora L. O. Williams, another trifoliate member of P. sect. Subviscosae en-

demic to southeastern Arizona. Indeed, the differences between P. rhyolitica and P. albiflora are fairly subtle,

consisting primarily of differences in vestiture, number of leaflet teeth, and carpel number. Populations of P.

+ in Datantilla

LUNA IBS YALE 51

Nu Wi}:

NNI

NN S EGY

Life

;

hr.

Wii PY.

NEL) 2

B G

Fic. 1. Potentilla rhyolitica Ertter. A-G P. rhyolitica var hy litica (d f Ertter 11872). A. Habit. B. Basal leaf. C. Underside of leaf. D. Vestit fpeti

A AL lE H Act

ole. E. Flower, top view. F. Flower, bottom view. G

D vb Jigs

rhyolitica in the Chiricahua Mountains are somewhat intermediate between the two species and sufficiently

distinct so as to warrant the recognition of two varieties within P. rhyolitica, as presented here.

Potentilla rhyolitica var. rhyolitica

Stems 0.3-2 dm long. Leaves: long hairs of petiole + 1(-2) mm long; leaflets (moderately to) densely seri-

ceous, gray-green. Flowers 2-10; filaments 1.5-2.5 mm long; styles 5-15, 2-2.5 mm long. Achenes + 1.5

mm long, smooth to lightly ribbed.

Distribution and phenology.—Endemic to the summit areas of the Santa Rita and Huachuca mountains in

Santa Cruz and Cochise counties, southeastern Arizona, mostly forming dense tufts in crevices of rhyolitic

52 Journal of the Botanical R h Institute of Texas 1(1)

and quartzitic outcrops in open pine forests, 2600-2900 m elev. Flowering Apr-Jun. An anomalous collec-

tion from desert grasslands in the San Rafael Valley southwest of the Huachuca Range (Fritts & Fritts 83-143,

COLO, and reported as P. wheeleri by McLaughlin [2006]) may also belong here, though in a significantly

different ecological setting and with more oblanceolate leaflets.

As already noted, the name Potentilla viscidula has frequently been applied to existing collections of

this entity, with P. subviscosa, P. dissecta Pursh, P. concinna Richardson, and P. albiflora serving as alternate

identifications. John J. Thornber, botanist at University of Arizona and the Santa Rita Experimental Range

in the early 1900's, was the first to notice the distinctiveness of this taxon. However, although Thornber

annotated the ARIZ sheet of Goodding 110 as “Type!” of a new species, with the epithet “trifoliolata” replaced

with “pinetorum,” he did not follow through with formal publication.

The variety is sufficiently localized to warrant conservation attention, especially given its proximity to

well-used trails in popular hiking areas. *Huachuca cinquefoil" is recommended as a vernacular name.

ADDITIONAL COLLECTIONS EXAMINED: U.S.A. Arizona: Cochise Co.: Huachuca Mts., mountain tops, 8 May 1909, L.N. Goodding 102 (ARIZ);

Miller’s Canyon, moist slopes, Huachuca Mts., 8 May 1909, L.N. Goodding 102a (RM); Miller’s Cano. Huachuca Mts., dry rocky places,

usually clinging to rocks, 8 Jun 1909, L.N. ipe 110 (ARIZ, RM); Huachuca Mts., rocl its, May 1912, L.N. Goodding

1300 p.p. (BKL, RM p.p. [mixed collection with P. subviscosa var. ramulosa]); Miller Canyon, top of saddle, Huachuca Mts., 8500 ft, 23 Apr

1955, H. S. Haskell & C. F. Deaver 5177 (ASC, RSA); Huachuca Mts., Carr Peak Pass along trail to Peak, rhyolite crevices, T23S R20E S22,

ca. 8900 ft, 11 May 1984, Soreng & Muldaven 2386 (COLO). Santa Cruz Co.: Mt. Wrightson Peak along Old Baldy Trail, 100 ft below

summit, in rock crevices, 18 May 1986, D. Bertelsen s.n. (ARIZ); Mt. Baldy, Santa Rita Mts., 9400 ft, 9 May 1937, R. Darrow s.n. (ARIZ);

Mt. Wrightson (Old Mt. Baldy) in Santa Rita Mts., solid rhyolite outcrops along trail on NE side of peak, 9000 ft, T20S R15E Sec 18, 4

Jun 1993, B. Ertter 11881 (UC; to be distributed); Santa Rita Mts., 8000 ft, 3 May 1881, C.G. Pringle 13677 p.p. (MO).

Potentilla rhyolitica var. chiricahuensis Ertter, var. nov. (Fig. 1H). Te: U.S.A. ARIZONA: Cochise Co.: Flys Peak

in Chiricahua Mts. ca. 40 air mi SE of Willcox, local on summit and along trail in open forest, rocky openings in forest of limber

pine, Douglas-fir, ponderosa pine, and aspen, T18S R30E, 9660 ft, 2 Jun 1993, B. Ertter 11872 (HoLotypE: UC; isotypes: ARIZ, GH,

MEXU, MO, NY, US)

A Potentilla rhyolitica var. rhyolitica foliolis viridioribus et acheniis grandioribus (+ 2 mm vs. + 1.5 mm) differt.

Stems 0.2-1(-1.5) dm long. Leaves: long hairs of petiole 1-2(-3) mm long; leaflets moderately (to densely)

hairy, green. Flowers 1-7; filaments 2-3 mm long; styles 5-10, 22.53 mm long. Achenes + 2 mm long,

smooth.

Distribution and phenology.—Endemic to upper elevations of the Chiricahua Mountains, Cochise Co.,

Arizona, in rocky openings in mixed conifer forests, 2700-2900 m. Flowering May-Jun.

Potentilla rhyolitica var. chiricahuensis differs from the typical variety in being greener and less sericeous,

with somewhat longer and coarser hairs and larger seeds. It is also less likely to be rooted in outcrops, favor-

ing rocky flats. “Chiricahua cinquefoil” is a suitable vernacular name for this localized taxon, which merits

some level of conservation attention.

As with P. rhyolitica var. rhyolitica, Thornber was evidently the first to recognize that the trifoliate

Chiricahua plants represented a distinct taxon. He annotated Blumer 2023 (US) as the type of a new species,

using the epithet “substrigosa,” but this combination was never published.

ADDITIONAL COLLECTIONS EXAMINED: U.S.A. Arizona: Cochise Co.: Fly Park, Chiricahua Mts., 8900 ft, 7 Jul 1907, J. C. Blumer 2023 (US);

Rustler’s Park, moist slope, 9000 ft, 18-19 Jun 1930, G. J. Goodman & C. L. Hitchcock 1173 (MO, RM, UC); Chiricahua Wilderness Area

Crest Trail, !4 mi N of turnoff to Anita Park, 9480 ft, 29 May 1975, J. & A. Leithliter 7 (ASU); Chiricahua Wilderness Area, intersection

of Crest Trail and Tub Spring Trail, % mi S of Long Park, 9070 ft, 10 Jun 1975, J. & A. Leithliter 52 (ASU); Flys Peak — Cima Cabin Trail

junction, Chiricahua Mts., 9200 ft, 29 May 1959, J. McCormick & Assoc. 82 (ARIZ).

DESCRIPTION OF POTENTILLA DEMOTICA

In the opposite corner of Arizona, the rarest member of Potentilla sect. Subviscosae was discovered in 1979

as part of a floristic study in the Hualapai Mountains (Butterwick et al. 1991). It was reported as a western

range extension of P. subviscosa but differs in rock-dwelling habit, petal color, and epicalyx, among other

characters. Although the Hualapai Mountains are not considered part of the Madrean Archipelago, they

continue the theme of montane “islands” scattered across the arid Southwest.

Ertter, Lectotypificati I new taxa in Potentilla 53

Potentilla demotica Ertter, sp. nov. (Fig. 2). Tyee: U.S.A. ARIZONA: Mohave Co.: Hualapai Mts. ca. 12 air mi SE of Kingman,

pink granite knoll SE of Hualapai Peak, in open ponderosa pine forest, with Antennaria, Heuchera, Cheilanthes, Poa, T20N R15W, ca

7500 ft, 30 May 1993, B. Ertter 11864 (HoLotyeE: UC; isotypes: ARIZ, GH, MO, NY, RSA, US).

Potentilla rimicola aemulans, differt foliolis brevioribus et epicalyces bracteolis late ellipticis.

Plants perennial, rooted in rock crevices, rosetted to tufted, the caudex simple to few-branched on a thick-

ened taproot. Stems spreading, 0.2-1.5(-2) dm long, with abundant short septate glandular trichomes, short

simple hairs ca. 0.2 mm long (sparse proximally, more abundant distally), and scattered slender spread-

ing hairs 1-2 mm long. Leaves primarily basal, palmate, 2-8 cm long; stipules very narrowly triangular;

petiole 1-7 cm long, with abundant glandular trichomes and slender spreading hairs 1.5-2.5 mm long;

leaflets G—)5, oblanceolate, the central one 0.5-1.5(22) cm long, sparsely hairy on both sides, toothed ‘2-7/4

to midvein on distal 74 of blade, the rounded teeth 2-3(-4) per side; cauline leaves 1-2, highly reduced.

Inflorescences lax, 1—7-flowered; pedicels 0.5-1(-2) cm long, slender, sometimes recurved, with short

glandular trichomes, short hairs + 0.2 mm long, and longer hairs + 1 mm long. Flowers: hypanthium shal-

low, 2-3 mm diam.; epicalyx bractlets ovate-elliptic, 1-2.5 mm long, 1-1.5 mm wide; sepals 2.5-4 mm

long, obtuse; petals yellow with darker base, + obcordate with a short claw, shallowly emarginate, 3-7 mm

long, exceeding sepals; stamens 20, the filaments 1.3-2 mm long, the anthers 0.5 mm; styles 5-12, slender,

scarcely rough-thickened basally, 2-2.5 mm long. Achenes light brown with reddish apices, 1.5-1.8 mm,

smooth to lightly ribbed.

Distribution and phenology.—eastern extension of Hualapai Peak in the Hualapai Mountains of Mo-

have Co., Arizona, crevices of granite outcrop in ponderosa pine woodland, ca. 2300-2400 m. Flowering

May-Jun.

Although initially identified as Potentilla subviscosa (Butterwick et al. 1991), P. demotica is probably more

closely related to P. rimicola (Munz & I.M. Johnst.) Ertter, which grows in rock crevices in the mountains of

southern California and northern Baja California. The new species is apparently extremely rare and localized,

currently known from a single granite knoll in Hualapai Mountain Park, part of the Mohave County park

system. The label for Butterwick & Hillyard 6757 indicates that about 100 clusters of one to fifteen plants each

were present in 1980, which is compatible with my observations in 1993. Conservation attention is obvi-

ously called for, though the knoll is a long hike from the nearest trailhead and not directly on any trail.

The epithet demotica (from “demotikos,” Greek for “of the people,” suggested by J. Reveal) alludes to

the tradition among the Hualapai that they arose from the “Pai” or “the people.” *Hualapai Cinquefoil” is an

appropriate vernacular name.

ADDITIONAL COLLECTIONS EXAMINED: U.S.A. Arizona: Mohave Co.: eastern extension of Hualapai Peak, Hualapai Mts., T20N R15W Sec. 32

NW1/4, scattered in rock crevices, Ponderosa Pine Woodland, 8000 ft, 10 Aug 1979, M. Butterwick & B. Parfitt 5407 (ASU, COLO); E

portion of Hualapai Peak system, T20N R15W Sec. 32 NW1/4, granitic substrate, Ponderosa Pine Forest, 8000 ft, 6 Jun 1980, M. But-

terwick & D. Hillyard 6757 (ASU, COLO).

NOTES ON SOME OTHER SPECIES IN POTENTILLA SECT. SUBVISGOSAE

Potentilla albiflora L.O. Williams

Contrary to the specific epithet, petals of living plants are yellow, not white, though (like many Potentilla)

they tend to fade in pressed material. “Whiteflowered cinquefoil” is accordingly misleading as a common

name, with “Pinaleno cinquefoil” a recommended alternative. Although previously known only from the

Pinaleno (Graham) Mountains in Graham Co., Arizona, the following collection (distributed as P. diversifolia

Lehm.) extends the range to the Mogollon Rim and opens up the possibility of further discoveries in this

relatively sparsely botanized corner of Arizona.

Arizona: Greenlee Co.: Cienega Camp on Hwy 666, Blue Range, Apache National Forest, 20 mi SW of Alpine, mesic headwaters of the

Blue River in mixed coniferous forest, elev. ca. 2600 m, 17 Jul 1964, M. Baad 657 (MICH, WTU).

Potentilla cottamii N. Holmgren

Holmgren (1987) described Potentilla cottamii from isolated quartzite outcrops in the Pilot Range and Raft

54 Journal of the Botanical R h Institute of Texas 1(1)

acces

lashes

Fic. 2. Potentilla demotica Ertter (drawn from Ertter 11864). A. Habit. B. Basal leaf. C. Enlargement of petiole vestiture. D. Flower. E. Hypanthium,

epicalyx, calyx. F. Achene and style.

River mountains of northwestern Utah and adjacent Nevada; it has since been found in Utah's Stansbury

and Deep Creek mountains. Holmgren speculated that his new species was most comparable to P. hyparctica

Malte and P. robbinsiana Oakes, members of P. sect. Aureae occurring in arctic and alpine regions in North

America. In contrast, R. Elven (pers. comm. 2006) confirms that any resemblance with these species is

superficial, with P. cottamii having very different vestiture and stipule types. In my understanding P. cottamii

fits readily into P. sect. Subviscosae, sharing the distinctive combination of simple and glandular vestiture

types but with unusually small flowers and accordingly short styles. The trifoliate leaves, stipules, petro-

Ertter, Lectotypificati ] taxa in Potentilla 55

phytic habit, and biogeographic distribution as a montane “island endemic” are likewise compatible with

membership in P. sect. Subviscosae.

Potentilla subviscosa Greene

The two varieties of Potentilla subviscosa have generally been based on extremes of leaflet margins, with var.

subviscosa having deeply lobed leaflets and var. ramulosa having more shallowly and regularly toothed leaflets,

which are also significantly larger overall. As so defined, the varieties were largely sympatric and questionably

distinct. Multiple examples of collections from the same general area, and even variation on a single plant,

indicate that these extremes can occur as striking seasonal differences in leaf shape and petiole vestiture,

perhaps as an adaptation to the monsoonal rainfall pattern in the American Southwest. Leaves formed early

in the season and coinciding with peak flowering have deeply divided leaflets, and the petiole vestiture

often consists almost exclusively of short glandular trichomes. Later-formed leaves have progressively less

deeply divided leaflets and an increasing percentage of both short and long non-glandular hairs; the leaves

are sometimes significantly larger as well. This dimorphism is best developed in Arizona populations of P.

subviscosa, including the Rusby 591 syntype of P. ramulosa, which consists of both early-season plants with

deeply lobed leaflets and late-season plants with significantly larger, less deeply toothed leaflets. A pencil

annotation on one of the latter specimens at NY indicates “status autumnalis” in the handwriting of Jiri

Soják, a Czech expert in Potentilla based at the National Museum in Prague (PR).

Within this new context of seasonal leaf dimorphism, P. subviscosa var. ramulosa can still be distin-

guished by the absence of deeply divided leaflets even on early-formed leaves. As so defined, this variety is

the only representative of P. sect. Subviscosae that occurs in the Santa Catalina and Rincon ranges in Pima

Co., Arizona. Comparable plants have been collected on Aztec Peak in the Sierra Ancha Range of Gila Co.,

Arizona, where they overlap the range of and possibly intergrade with P. subviscosa var. subviscosa. The typi-

cal variety itself is widespread in the mountains of New Mexico and north of the Mogollon Rim in Arizona,

barely entering Colorado in the Sangre de Cristo Range of Las Animas County.

Potentilla wheeleri S. Watson.

As here circumscribed, Potentilla wheeleri is restricted to the southern Sierra Nevada, San Bernardino

Mountains, and San Jacinto Mountains in southern California. Compact plants from the alpine summit of

Mount San Gorgonio in the San Bernardino Mountains were described by Jepson (1925) as P. wheeleri var.

paupercula, but I have not found sufficient consistent differences to maintain this as a distinct variety. In

contrast, petrophytic plants from Tahquitz Peak (San Jacinto Mountains, Riverside Co., California) and the

Sierra San Pedro Mártir (Baja California, Mexico) originally described as P. wheeleri var. rimicola Munz & I.M.

Johnst. are now recognized as a distinct species, P. rimicola (Ertter 1991). Other plants of Potentilla wheeleri

s.l. from the Sierra San Pedro Mártir in Baja California Norte have more open inflorescences, leaves that are

sometimes subpalmate, and less hairy leaflets that are only shallowly and apically toothed; these populations

can probably also stand as a distinct species, P. luteosericea Rydb. (= P. pinetorum Wiggins).

KEY TO- POTENTILLA SECT. SUBVISCOSAB NORTH OF MEXICO

1. Leaflets mostly 3, rarely 5.

2. Petals < 2 mm long; styles < 2 mm long; NW Utah and adjacent Nevada P. cottamii

2. Petals 3-8 mm long; styles 2-3 mm long; SE Arizona.

3. Longest hairs on petiole + 1 mm long; central leaflet with 3-5(-6) teeth per side; Pinaleno Mountains

and nearby Mogollon Rim P. albiflora

3. Longest hairs on petiole 1-2(-3) mm long; central leaflet with 2-3(-4) teeth per side P. rhyolitica

4. Leaves gray-green, + densely sericeous; styles 2-2.5 mm long; achenes + 1.5 mm long, smooth to

lightly ribbed; Santa Rita and Huachuca mts. var. rhyolitica

4. Leaves green, + moderately hairy; styles 2.5-3 mm long; achenes + 2 mm long, smooth; Chiricahua

var. chiricahuensis

Mts.

1. Leaflets mostly 5, rarely 3 or 7.

56 Journal of the Botanical R h Institute of Texas 1(1)

5. Petals pale yellow adaxially, white abaxially, narrowly obcordate; leaflets with 2-9 teeth or lobes per side

P. subviscosa

6. Leaflets often strongly dimorphic, at least those formed early in season divided /2-%4 to midvein into

3-7 lobes per side, late-season leaflets often coarsely toothed less than 4 to midvein with 6-9 teeth

per side; long hairs on petioles 1-1.5(-3) mm long; widespread in n Arizona and New Mexico, barely

entering Colorado var. subviscosa

. Leaflets scarcely dimorphic, those formed early in season toothed 14-12 to midvein with 2-4 teeth

per side, late-season leaflets similarly toothed with up to 6 six teeth per side; long hairs on petioles

2-3(-4) mm long; Santa Catalina and Rincon mts. and Sierra Ancha in SE Arizona var. ramulosa

. Petals bright yellow adaxiall

teeth per side.

7. Plants rooted in ground; leaflets densely hairy; styles ca. 20, 1.2-2 mm long; southern Sierra Nevada

and San Bernardino Mts. of Calif. P. wheeleri

7. Plants rooted in rock crevices, often on vertical surfaces; leaflets sparsely to moderately hairy; styles

5-15(-20), 1.5-2.5 mm long.

8. Leaflets 1-3 cm long, oblanceolate to obovate-cuneate, toothed in distal Vs; epicalyx bractlets

z d Bx A

somewhat paler abaxially, narrowly to broadly obx late; leafl with 2-4(-5)

lanceolate-elliptic; San Jacinto Mts., California, & Baja California, Mexico P. rimicola

8. Leaflets 0.5-1.5(-2) cm long, oblanceolate, toothed in distal 34; epicalyx bractlets ovate-elliptic;

Hualapai Mts., Arizona P. demotica

ACKNOWLEDGMENTS

Foremost I wish to thank the curatorial staff at the multiple herbaria who provided loans used in this

study, and their patience in the length of time the specimens have been in my hands: ARIZ, ASC, ASU,

BRY, COLO, GH, MO, NY, RM, RSA, TEX, US, UTC, WTU. Special thanks are merited by Emily Wood

and Melinda Peters for their exceptional help with syntypes at GH, and the curatorial staff at UC/JEPS who

have processed my seemingly unending loans. I also wish to acknowledge John McNeill and Jim Reveal for

elucidating the Kearney and Peebles lectotypification and reviewing the manuscript; Alfonso Delgado S. for

translating the abstract into Spanish; Jifi Soják for sharing his insights on Potentilla (as well as Adolf Ceska

and Lenka Drabkova for serving as translators); Steven P. McLaughlin for sending reprints of his highly

relevant publications on southwestern biogeography; Michael Baad and Richard Rabeler for additional in-

formation on the P. albiflora range extension; Richard Beidleman and Alvin Medina for help with historical

localities; Roger Weller for geological information in the Huachuca Mountains; Les Landrum for his efforts

locating var. ramulosa and reviewing the manuscript; Reidar Elven for his evaluation of P. cottamii; and

Lindsay Woodruff for providing field assistance and companionship. As always, I gratefully acknowledge

Linda Vorobik for her excellent illustrations. Support from the Lawrence R. Heckard Endowment Fund of the

Jepson Herbarium is gratefully acknowledged.

REFERENCES

ABRAMS, L. 1944. Illustrated flora of the Pacific States. Vol. Il. Polygonaceae to Krameriaceae. Stanford University

Press, California.

Bowers, J.E. and S.P. McLAUGHLIN. 1996. Flora of the Huachuca Mountains, a botanically rich and historically signifi-

cant sky island in Cochise County, Arizona. J. Arizona-Nevada Acad. Sci. 29:66-107.

Burterwick, M., B.D. Parritt, and D. HiLLYARD. 1991. Vascular plants of the northern Hualapai Mountains, Arizona. J.

Arizona-Nevada Acad. Sci. 24-25:31-49,

ErTTER, B. 1991. New combinations in Potentilla and Horkelia (Rosaceae) in California. Phytologia 71:420-422.

HOLMGREN, N.H. 1987. Two new species of Potentilla (Rosaceae) from the Intermountain Region of Western U.S.A.

Brittonia 39:340-344.

Jepson, W.L. 1925. A manual of the flowering plants of California. Associated Students Store, University of Cali-

fornia, Berkeley.

Jepson, W.L. 1936. A flora of California. Vol. ll. Capparidaceae to Cornaceae. Associated Students Store, University

of California, Berkeley.

Ertter, Lectotypifications and new taxa in Potentilla 57

JOHNSTON, B.C. 1985. Studies in Potentilla. |. Key to North American sections. Phytologia 57:292-302.

Kartesz, J.T. and C.A. MEACHAM. 1999. Synthesis of the North American flora, Version 1.0. North Carolina Botanical

Garden, Chapel Hill.

Kearney, T.H. and R.H. PeesLes. 1942. Flowering plants and ferns of Arizona. U.S.D.A. Misc. Publ. 423:1-1069.

Kearney, T.H. and RH. Peestes. 1951. Arizona flora. Univ. California Press, Berkeley.

McLAUGHLIN, S.P. 1995. An overview of the flora of the sky islands of southeastern Arizona: diversity, affinities,

and insularity. In: L.F. DeBano, GJ. Gottfried, R.H. Hamre, PF. Folliott, and A. Ortega-Rubio (Tech. coords.), Bio-

diversity and Management of the Madrean Archipelago: the sky islands of southwestern United States and

northwestern Mexico. USDA Forest Service Gen. Tech. Rept. RM-GTR-264. Pp. 60-70.

McLAUGHLIN, S.P. 2006. Vascular floras of Sonoita Creek State Natural Area and San Rafael State Parks: Arizona's

first natural-area parks. Sida 22:661-704.

McNüiLL, J., F.R. Barrie, H.M. Bunper, V. DEMOULIN, D.L. Hawksworth, K. MARHOLD, H.H. NicoLson, J. PRADO, P.C. Sitva, J.E.

Sk0G, J.H. Wiersema, and N.J. TuRLAND (eds.). 2006. International Code of Botanical Nomenclature (Vienna Code)

adopted by the Seventeenth International Botanical Congress Vienna, Austria, July 2005. Gantner Verlag,

Ruggell, Liechtenstein.

MEDINA, A.L. 2003. Historical and recent flora of the Santa Rita Experimental Range. U.S.D.A. Forest Service Pro-

ceedings RMRS-P-30:141-148. (http://ag.arizona.edu/SRER/proceedings/Medina2.pdf)

Munz, PA. 1959. A California flora. University of California Press, Berkeley.

RYDBERG, P.A. 1896. Notes on Potentilla.—V. Bull. Torrey Bot. Club 23:429-435, plates 276-277.

RYDBERG, PA. 1898. A monograph of the North American Potentilleae. Mem. Dept. Bot. Columbia Coll. 2:1—223,

plates 1-112.

RYDBERG, PA. 1908. Potentilla. North American Flora 22:293-352.

Tipestrom, |. and T. Kite. 1941. A flora of Arizona and New Mexico. The Catholic University of America Press,

Washington, D.C.

58 Journal of the Botanical R h Institute of Texas 1(1)

BOOK REVIEWS

ERNEST SMALL and Grace DeutscH. 2002. Culinary Herbs for Short-Season Gardeners. (ISBN 0-87842-

453-9, pbk.), Mountain Press Publishing Company, Box 2399, Missoula, MT 59806, U.S.A. $20.00.

(Orders: www.mountain-press.com, info@mtnpress.com, 406-728-1900, 406-728-1635 fax). $20.00,

182 pp., 14 color photographs, 14 b/w illustrations, 128 full color illustrations, and one full color map

of plant hardiness zones of North America.

Growing up in semi-tropical Australia, writer/publisher Grace Deutsch knew she had her hands full after immigrating to Southern

Ontario, Canada with desires of having a culinary herb garden. She admired and was curious as to how her gardening friends in Mon-

tana, the Adirondacks and northern Alberta managed to have persistently beautiful and productive flower, vegetable and herb gardens

in such a relatively short warm-weather season. Fortunately, she met co-author Dr. Ernie Small by chance at a summer herb fair. Dr.

Small was raised in the Ottawa Valley and is an international authority on agriculturally important plants. He has written over 200

scientific publications and 7 books, including Culinary Herbs, the professional herb-growers bible. Together they collaborated to create

this very useful book for the novice or experienced short-season gardener who enjoys a culi garden just outside their kitchen door.

It is concise, practical, and easy-to-use.

A full color map of Canada and the USA helps you interpret your plant hardiness zone for growing perennial herbs that will sur-

vive your particular winters. Chapter: Making the Most of the Short Growing Season is full of practical advice. Chapter: Growing and Caring

for Your Herbs is a gardening primer for herb growers. Chapter: The Culinary Herb Compendium includes 50 species, with 100 cultivars.

Herbs are arranged alphabetically by common name along with scientific name and the plant family name. Icons clearly mark annuals

from perennials along with a colored “Coldest Tolerated” zone box. These boxes enable you to quickly see the coldest zone in which the

herb may overwinter successfully. These individual herb listings include the following elements:

Description of herb’s origin; life cycle; size; leaf shape, color, taste and fragrance; stem and root formation; flowering habit;

suitability for growing in pots or containers; which parts are edible; plus any additionally desirable features, for example, if the flowers

are very alluring to pecan or butterflies. Ae Notes: eu ie you nic to know about D E HEUS and moisture

requirements; pro[ pest and di ies. Harvest-

ing Notes: for maximum flavor iud how best to preserve. S cul Uses both domestic and commercial; when herb snout be added

in the cooking process; if the herb makes a tea and brewing instructions. Craft Uses: to enjoy the natural beauty of herbs. Medicinal

Uses: traditional herbal medicine uses and its value in modern medicine; nutritional information. Cautions: information on existing

known problems that susceptible individuals may have from consumption of or contact with the herbs. Cultivars and Relatives:

descriptive listing of other useful plants related to herbs and outstanding cultivars. Herbal Trivia: snippets of the myths, legends, and

folklore surrounding each herb.

I particularly enjoyed the use of the beautifully modified botanical illustration masterpieces produced in the 18' and 19" centu-

ries.—Linny Heagy, Linny/Designer, Illustrator, email: a0005835@airmail.net.

MAGGIE CAMPBELL-CULVER. 2007. A Passion for Trees: The Legacy of John Evelyn. (ISBN 1-903-91947-9,

hbk.). Eden Project Books, Eden Project penu A Bodelva Par, Cornwall PL24 2SG United Kingdom

(Orders: http://shop.edenproject hxc/gl play/splash.html). $45.00, 288 pp., color and b/w

[e]

photos and illustrations a

ks tl of the death of English horticulturi 1f fatl fmod conservation,

This copiously illustrated |

John Evelyn (1620-1706). It explores the legacy of John Evelyn through illustrated portraits of 30 of Britain best-loved trees.

Described by Samuel Pepys as ‘a man so much above others...a most excellent, humoured man and very knowing.’ Evelyn, like Pepys

is well known as a diarist, whose writing tells us much about the life and times of the 17 century, but it is in his interest in nature,

gardens, horticulture and trees that is of concern in this book. Not only a gardener, Evelyn was a royal advisor, and precursor of the

modern environmental movement, but he was also the author of exceptional works on horticulture. The greatest of these is Sylva: A

Discourse of Forest Trees, published in 1664.

Unlike other European countries, Britain became unique in the way it used trees in the landscape. British designers of the 18"

century found aesthetic considerations of prime importance over the more utilitarian aspects of silviculture. It was an idea based on

creating landscapes by consulting the ‘Genius of the Place’. Evelyn’s Sylva is among the first books in English to show an appreciation

of the aesthetic value of trees and the benefits of planting them to shape the landscape, as an artist might achieve with a painting.

From the beloved English Oaks to the Acacia, Arbutus, Bay, Box, Yew, Holly, Juniper and Laurel trees, author Maggie Campbell-

Culver walks hand in hand with Evelyn to paint loving portraits of his well-loved trees. She ends with the following:

“The management and planting of trees are exactly the same as they were in 1664. What better, therefore, could we do than follow the advice

of Sylva? The planting of trees is an essential element of life, and conservation is as necessary as it was when John Evelyn argued for it over 300

years ago.”—Linny Heagy, Linny/Designer, Illustrator, email: a0005835@airmail.net.

J. Bot. Res. Inst. Texas 1(1): 58. 2007

A NEW VARIETY OF FESTUCA ROEMERI (POACEAE) FROM THE CALIFORNIA

FLORISTIC PROVINCE OF NORTH AMERICA

Barbara L. Wilson

Carex Working Group

2/10 Emerald Street

Eugene, Oregon 97403, U.S.A.

bwilson@peak.org

ABSTRACT

The grass Festuca roemeri is a community dominant in grasslands and savannas from southwest British Columbia through central Cali-

fornia. Inland plants in southern Oregon and California differ from more northern F. roemeri in isozyme profiles, leaf pubescence, some

inflorescence measurements, and perhaps physiology. These populations are here described as F. roemeri var. klamathensis.

RESUMEN

Festuca roemeri es una gramínea dominante en la comunidad de pastizales y savanas del suroeste de Columbia Británica hasta el centro

de California. Plantas distribuidas desde el interior del suroeste de Oregón hasta el noroeste de California difieren de F. roemeri por

la pubescencia foliar, los perfiles de isoenzimas, algunas medidas de la inflorescencia y quizás en su fisiología. Estas poblaciones son

descritas en el presente trabajo como F. roemeri var. klamathensis.

A dominant bunchgrass of savannas in the Klamath Region of Oregon and California has been identified

variously as Festuca idahoensis Elmer, Festuca idahoensis x F. occidentalis Hook. hybrids, F. idahoensis var.

oregana (Hack.) C.L. Hitchc., F. ovina L., and F. rubra L. In despair, some botanists simply report Festuca sp.

The grass is F. roemeri (Pavlick) E.B. Alexeev (Alexeev 1985; Pavlick 1983; Wilson 1999), but differs slightly

from typical, more northern F. roemeri. It is described here as a variety of F. roemeri.

Festuca roemeri var. klamathensis B.L. Wilson, var. nov. (Figs. 1-2). Tv: U.S.A. Ontcox. Jackson Co.: Rogue River

National Forest, Baldy Peak Trail, T40S R3W S22, 2 Jul 1996, B.L. Wilson 8199 (HoLotyPE: OSC; isotypes: DAV, MO, NY, RSA, UC,

UTE)

A Festuca roemeri var. roemeri foliorum tricomatibus adaxialibus longioribus et numerosioribus differt.

Cespitose, the old plants sometimes dying in the center, leaving a ring of living shoots; leaf sheaths open

(sheath margins overlapping) to the base; leaves glaucous, occasionally green; leaf blades erect, somewhat

stiff, conduplicate, 5-30(-50) cm long, 0.55-1.2(-1.5) mm wide, the abaxial (outer) surface glabrous or

pubescent, the adaxial (inner) surface with numerous hairs 0.06—0.3 mm long (the longer hairs often about

as long as the leaf is thick), adaxial ribs 5-9; veins 5-7(-9). Abaxial sclerenchyma bands >2x as wide as

thick, usually forming large bands at margins and midrib, often with smaller bands opposite veins, usually

lacking adaxial sclerenchyma; flag leaf 1.5—7 cm long; culm nodes becoming exposed 1; flowering shoots

(20-)30-95 cm long, panicle 7-15(-20) cm long. Inflorescence branches at lowest node 1-2, appressed

(usually) or spreading after anthesis; spikelets 2-6 on longest branches, 7-16 mm long. Florets 3-5; glumes

unequal, the lower 3.2-5.1(-5.7) mm, the upper 4.4-6(-7.2) mm long; lemma (6-)6.2-8.5 mm long, gla-

brous, with lemma awn (1-)1.5-4.6 mm long, shorter than the lemma body; anthers 3-4.2 mm long; ovary

apex glabrous. Tetraploid.

Distribution.—inland sites from southern Douglas County in southwest Oregon to northwest California,

east of the coastal mountains (Fig. 3).

Habitat.—community dominant in mesic to dry pine or oak savanna, grasslands, and edges of grassy

balds, on a variety of substrates including serpentine.

Festuca roemeri was originally described as F. idahoensis var. roemeri Pavlick, based on populations found

in upland sites in the moist, maritime climate of southwest British Columbia and northwest Washington

J. Bot. Res. Inst. Texas 1(1): 59 — 67. 2007

5 cm

Journal of the Botanical Research Institute of Texas 1(1)

Fic. 1. Festuca roemeri var. klamathensis, habit. Left: Wilson 8199, from

Right: Whito & lillirn 210 f "m landini hinaf

tv, Oreaon

or pa

avannah on non-serpentine soils, Jackson County, Oregon.

Wilson, A new variety of Festuca roemeri 61

Fic. 2. Leaf cross cactionc of fiald llact ( ) Inl f, } H fi YA In. Lres

B and E: F. roemeri var. klamathensis. C ani F: Festuca roemeri var. roemeri.

(Pavlick 1983). It was soon recognized at the species level (Alexeev 1985). Festuca idahoensis sensu stricto

is widespread east of the Cascade Range and Sierra Nevada through the western Great Plains. The narrow

conduplicate leaves with dense hairs on the inner surface (where the stomata are located) are adaptive for the

dry to xeric continental climate in which it grows. Festuca roemeri differs from F. idahoensis sensu stricto in

its ovate to obovate-pyriform leaves that have few hairs on the inner surface. Festuca roemeri is more widely

distributed than was initially realized, growing at least as far south as Santa Cruz County, California.

Atinland sites in southwestern Oregon and northwestern California, F. roemeri grows in dry, continental

PA RT 271 O A Tp

JOUTHadl OF

ecoregion.

Wilson, A new variety of Festuca roemeri 63

environments similar to those inhabited by F. idahoensis. These inland populations differ from F. roemeri var.

roemeri in having long hairs on the inner surface, like F. idahoensis (Fig. 2). Even in a common greenhouse

environment, F. roemeri var. klamathensis had more and longer hairs than F. roemeri var. roemeri (Table 1).

The difference in leaf hairs can be detected in the field by bending the opened leaf over a finger and view-

ing the adaxial surface with a hand lens: leaves of F. roemeri var. roemeri appear glabrous whereas those

of F. roemeri var. klamathensis appear distinctly pubescent. In Festuca, the extent of abaxial pubescence is

greater at high temperatures (Aiken et al. 1994), but adaxial hair characters may be more stable that other

leaf anatomical traits, and the adaxial hair traits are species specific (Aiken et al. 1994; Dubé & Morisset

1996; Ramesar-Fortner et al. 1995; Wilson 1999).

A combination of other traits distinguishes F. roemeri var. klamathensis from F. roemeri var. roemeri and

from F. idahoensis. Festuca roemeri var. klamathensis often has leaves with a shape more typical of F. roemeri

var. roemeri, although plants growing on serpentine substrates have leaves nearly as narrow as those of

F. idahoensis (but with better defined ribs and grooves). Glumes and lemmas average slightly longer in F.

roemeri var. klamathensis than in F. roemeri var. roemeri (Table 2). In F. roemeri var. klamathensis, isozyme

band patterns as revealed on gels stained for malate dehydrogenase (following the methods of Wendel &

Weeden 1989) match those of F. idahoensis, not F. roemeri var. roemeri (Wilson 1999).

Festuca roemeri var. klamathensis is phenotypically plastic. Plants of fertile soils have lush, dense bunches

of 30 cm long, moderately glaucous leaves, and lemmas up to 8 mm long. They do not resemble the intensely

glaucous, depauperate individuals of serpentine barrens, which may have leaves up to 7 mm long, gnarled

bases, dead centers, and lemmas up to 6.5 mm long. When brought into the greenhouse the depauperate

plants produce broader leaves (Fig. 2; Wilson 1999). Plants of intermediate stature exist in the wild, but

extensive populations usually have only large or only depauperate individuals, leading biologists to treat

these variations as different taxa. A similar range of variation is observed in F. roemeri var. roemeri, but in

that variety depauperate plants are usually found on rock outcrops within populations of more lush plants.

From Lake County, California, southwards, plants of F. roemeri are intermediate and cannot always be

classified into either variety. Festuca roemeri var. klamathensis is named as a variety, rather than a subspecies,

because of these intermediates. On the north and west edges of its range, the transition zone is narrower.

The name Festuca idahoensis var. oregana (Hack.) C.L. Hitchcock has been applied dwarfed fescues of

western Washington and Oregon, including those referred to here as F. roemeri var. klamathensis (Hitchcock

et al. 1969), but its type specimen (Cusick 753, Oregon, 1884; US!), is F. idahoensis (Pavlick 1983), which

can also be dwarfed when growing extremely stressful microhabitats. The type of F. ovina var. columbiana

Beal (1886; US!) is Lake s.n. June 1882 (labeled as collected near Pullman, Washington, but apparently re-

ally collected near the head of Tukanon River, Blue Mountains, Washington, according to an annotation by

Piper). It appears to be F. idahoensis as well, and was collected well outside the known range of F. roemeri.

Conservation concerns

Application of the names Festuca idahoensis and F. ovina to populations of F. roemeri var. klamathensis has

led land managers to mistakenly plant commercial cultivars of F. idahoensis and “Sheep Fescue" as native

species in habitat restoration projects within the range of F. roemeri var. klamathensis. Festuca idahoensis

plantings within the range of F. roemeri var. roemeri usually die because F. idahoensis is adapted to be winter

dormant in a drier climate with less pressure from fungal pathogens, but F. idahoensis may survive in the

environment of F. roemeri var. klamathensis. Hybridization between the two tetraploid taxa might well occur

and could result in the loss of genetic traits typical of F. roemeri var. klamathensis, including its adaptive

phenotypic plasticity. Planting Sheep Fescue as a native grass is erroneous because F. ovina and the other

taxa sold under this name, such as F. trachyphylla (Hack.) Krajina and F. valesiaca Gaudin, are not native to

North America. A second concern is that demand for F. roemeri var. roemeri seed has grown faster than the

supply of seed. Unlike F. roemeri var. roemeri (Wilson 1997), Festuca roemeri var. klamathensis is common

and rarely grows with other fine-leaved fescue species. This simplifies collection of uncontaminated seed

and establishment of pure cultivated seed sources. Nomenclatural clarification may reduce pressure to use

64 Journal of the Botanical R h Institute of Texas 1(1)

Taste 1. Traits of £ roemeri leaf anatomy. N = number of individuals measured. Max = maximum value. Min = minimum value. s = standard deviation. Probability (p) is the

probability that the measurements for A roemeri var. roemeri and F roemeri var. klamathensis are the same.

F. roemeri var. roemeri F. roemeri var. klamathensis Probability

N Mean s Max. Min. N Mean s Max. Min. p<

Plants from the field

Number of vascular bundles 15 661 062 7.33 5:00 i 5:45 0.60 6.33 500 2977x10?

Number of fibers

(sclerenchyma strands) A EE 08 300 11 410 082 567 3.00 0.008843

Adaxial hair length (mm) 1539031 09260 117 0014 11 0.094 0.044 0.216 0.061 0.000629

Adaxial hairs/side lS eT uS pss 2007 ur 039 197 1450 8.12 153X10

Adaxial grooves 15- 5:34 -0.69 6.00 400 11 420 036 500 400 184x10°

Plants from greenhouse

Number of vascular bundles 16 659 073 75 500 11 5.98 0.98 7.50 5.00 0.9308

Number of fibers

sclerenchyma strands) 16 540 178 80 300 11 404 091 525 3.00 0.01591

Adaxial hair length (mm) 16— BOTS 0.005- 0.020 0.006 11 0.077 0.078 0.304 0.017 0.02144

Adaxial hairs/side 16 183 141 6.00 0.50 Wil 893 427 14.00 0.00 0.00018

Adaxial grooves 16- 534 073 16.50 400 11 443 079 6.00 370 0.006039

F. roemeri var. klamathensis seed at inappropriate locations. Third, F. roemeri var. roemeri is uncommon to

rare in much of its range, with most Oregon populations varying from 13 to approximately a thousand

individuals. The fate of this taxon is not tracked when it is treated as taxonomically identical to F. roemeri

var. klamathensis.

Etymology.—The name klamathensis is appropriate for this fescue variety because the ancient, varied,

and botanically complex Klamath Range (Whitaker 1960) is the center of its distribution. This region is

named after the Klamath people who live there and once used fires to manage the F. roemeri var. klamathensis

grasslands.

The following key distinguishes the native and the more common introduced fine-leaved fescues grow-

ing at low to moderate elevations in and near the range of F. roemeri.

KEY TO BINB-LBAD RESCUES OF PACIFIC COASTAL STATES AND PROVINCE

1a. Longer awns longer than the lemma bodies; ovary apices densely pubescent; plants growing in the

shade F. occidentalis

1b. Longer awns none or up to as long as the lemma bodies; ovary apices glabrous (rarely with «10 hairs); plants

growing in the sun or in partial shade.

2a. Plant rhizomatous F. rubra, sensu lato

2b. Plant cespitose.

3a. Lemmas »(5.8-)6 mm long.

4a. Leaves very narrow, smoothly rolling between the fingers, round to hexagonal in cross section;

ribs on adaxial surface usually 3(-5); native range east of the Cascade Range and Sierra Nevada

(in California wholly east of Interstate Highway 5) but occasionally planted to the west ___ F. idahoensis

. Leaves wider, not rolling between the fingers or doing so with angles that can readily be felt,

V-shaped or obovate to elliptic in cross section; ribs on adaxial surface 5 or more; native range

west of the Cascade Range and Sierra Nevada (in northern California, near or west of Interstate

Highway 5) F. roemeri

5a. Hairs on the adaxial surface of leaves short (<< thickness of leaf) and usually sparse (except

in some south coastal and Columbia Gorge populations) F. roemeri var. roemeri

5b. Hairs on the adaxial surface of leaves long (% to about = thickness of leaf) and dense

F. roemeri var. klamathensis

3b. Lemmas «5.8, usually 4-5.5 mm long.

6a. Leaf sheaths with margins fused to near the top (but readily splitting as the plant grows); leaf

Wilson, A new variety of Festuca roemeri 65

TABLE 2. Selected measurements of £ roemeri inflorescences. N= number of individuals measured. Max = maximum value. Min =

minimum value. s = standard deviation. Probability (p) is the probability that the measurements for F roemeri var. roemeri and £

roemeri var. klamathensis are the same.

F. roemeri var. roemeri F. roemeri var. klamathensis Probability

Max Min. ax Min <

Panicle length (cm) 2] “1328 229. 195 oo 9T 39:293] 180- 5,6 0.0485*

Lower glume length (mm) 21 342 066 49 20 19-959 0.69 57 32 0.0341*

Upper glume length (mm) 21 504 059 62 42 19 5.52 1.14 Y 44 9.1135

Lemma length (mm) 21 695 042 79 65 19 747. 9072 8.5 6.3 0.0097%

Longest awn length (mm) 2] 394 089 53 23 18 340 0.90 4.6 1.8 0.0692

sclerenchyma bundles « 2x as broad as thick (but sometimes fused into broader groups);

leaf

sheaths brown and shredding to reveal whitish veins. Native and introduced taxa ___ F. rubra, sensu lato

Leaf sheaths with overlapping margins; leaf sclerenchyma bundles generally > 2x as broad

as thick; leaf sheaths paler, not shredding introduced Sheep and Hard Fescues

7a. Leaves thinner (0.4-0.6 mm wide); leaf sheaths not conspicuously broader than blades;

leaves always with at least 5 ribs on adaxial surface; adaxial hairs many and long; leaf scle-

renchyma

interrupted and sometimes consisting of three discrete bands F. valesiaca

. Leaves often broader (0.5-1.2 mm wide); leaf sheaths in some cultivars conspicuously

broader than blades; leaves with 3 - many ribs on adaxial surface; adaxial hairs many and

short; leaf sclerenchyma various, sometimes forming a continuous band under the ab-

axial

epidermis F. trachyphylla

APPENDIX

SPECIMENS OF FESTUCA ROEMERI VAR. KLAMATHENSIS EXAMINED

In Oregon, all legal descriptions (TRS) are based on the Willamette Meridian.

CALIFORNIA: Alameda Co.: Upper end of Corral Hollow, 18 Apr 1941, Hoover 4847 (UC); W of Corral Hollow, on road to

Livermore, 21 May 1939, Stebbins 2710 (DAV). Del Norte Co.: Stoney Creek Bog, T17N R2E S16 (H meridian), 26 May 1978,

Alcasas 103 (HSC); T18N R1E $35 (H meridian), 20 Jun 1979, Baker 868 (HSC); Old Gasquet Road, T17N R2E $13& $24 (H meridian),

25 Jun 1975, Barker 1006 (HSC); near Gasquet, 22 May 1979, Clinton & Overton 3084 (HSC); Gasquet, Jun 1902, Davy s.n. (UC);

French Hill Road, T17N R2E S29 (H meridian), 14 Jun 1978, Nelson 4152 (HSC); Smith River, EIk Camp Ridge, May 1937, Parks

& Tracy 5833 (HSC); state line N of Monumental, at head of Shelly Creek, 17 Jun 1936, Parks & Tracy 11386 (UC); Stony Cr. Bog

near Gasquet, T17N R2E $16 (MD meridian), 13 May 1973, Smith 6749 (OSC); 1 mi SW of Patrick Creek, 19 Jun 1936, Yates 5770

(DAV); 41*59'9"N, 123*58' 16" W, 2 Jun 1980, York 928 (HSC); near High Plateau Mt., 31 May 1980, York s.n. (HSC). Humboldt Co.:

Trinity Summit, 31 Jul 1901, Goddard 134 (UC); Brannan Mountain, 10 Jul 1930, Tracy 8867 (OSC); Mail Ridge, 7 mi N of Harris, 15

Jun 1950, Tracy 18803 (UC). Lake: Snow Mountain, 15 Jun 1979, Heckard & Hickman 5040 (JEPS); Reiff, Knoxville Ridge, 11 Jun

1938, Jepson 19012 (JEPS); between Cobb Mt. & Adams Spring on the Binkley Ranch, 4 Jul 1933, Jussel 360 (UC). Mendocino

Co.: Grouse Moutain, 25 Jul 1933, Tracy 12890 (UC). Santa Cruz Co.: N end of Swanson Road, overlooking Greyhound Rock,

10 km NW of Davenport, 13 May 1983, Buck & West 265 (JEPS). Shasta Co.: T28N R10W S6 (MD meridian), 22 Jun 1980, Nelson

& Nelson 5820 (HSC). Siskiyou Co.: Paradise Lake, Marble Mountains, trail to Kings Castle, 22 Jul 1949, Alexander & Kellogg

58844 (UC); Rainbow Ridge above Sulloway Cr. about 1.5 mi W of Mt. Shasta City, 11 Jun 1936, Babcock & Stebbins 1894 (UC);

Rainbow Ridge above Sulloway Cr. about 1.5 mi W of Mt. Shasta City, 13 Jun 1936, Babcock & Stebbins 2008 (UC); Big Flat

Campound, 2 Jul 1959, Bacigalupi 7232 (JEPS); headwaters of S Fork of Salmon River near N base of Caribou Mt., just S of low

divide (Trinity Co. boundary) separating Coffie Creek drainage from that of Salmon River, 2 Jul 1959, Bacigalupi 7234 (DAV);

40?24'20"N, 123*34'00"W, 20 Jun 1980, Baker 2220 (HSC); 13 mi E of Hamburg, banks of Klamath River, 31 May 1942, Beetle

3431 (DAV); Moffitt Creek, 13 Jun 1909, Butler 833 (UC); Moffet Creek NW of Fort Jones, 6 Jun 1922, Dunning s.n. (JEPS); Big Flat,

T37N ROW S18 (MD meridian), 3 Aug 1966, Ferlatte 251 (HSC); Eastern Flank of Mt. Eddy, just below summit fell field, 13 Aug

1967, Heckard 1709 (JEPS); Eastern Flank of Mt. Eddy, just below summit fell field, 13 Aug 1967, Heckard 1711 (JEPS); toward

the head of Wagon Creek, Mt. Eddy, 17 Jul 1920, Heller 13680 (DAV); V? mi SE of Kings Castle, Marble Mts., 6000 ft, 9 Jul 1939,

Hitchcock & Martin 5321 (UC); NE face of Marble Mountain, T43N R12W S15 (MD meridian), 4 Sep 1966, Major s.n. (DAV); Summit

between Beach Creek & Toad Lake, 22 Jul 1959, Murphy 591 (DAV); 0.2 mi E of Butcher Gulch on the Cecilville-Forks of Salmon

£s+haD o ID L

66 Journal of t titute of Texas 1(1)

Road, 28 May 1972, Smith & Sawyer 5384 (HSC); junction of French Creek and Sugar Creek Road, 1/2 mi W of Parrott Mill Road,

30 Jun 1972, Smith & Sawyer 5641 (HSC); E of Cook and Green Pass, T18N R4W S8 (MD meridian), 3 Jul 1972, Smith & Sawyer

5660 (HSC); TA3N R12W 52 (MD meridian), 7 Jul 1976, Stillman 161 (HSC); TA3N R12W S15 (MD meridian), 7 Aug 1976, Stillman

s.n. (HSC); Soap Creek Ridge between Yreka and Fort Jones, 3 Aug 1949, Tofsud s.n. (DAV); Mt. Eddy, TAON RSW S7& S 18 (MD

meridian), 20 Aug 1976, Whipple 1706 (HSC); Intersection Hwys 263 & 96, TA7N R6W S18SW 1/4 of SW 1/4 (MD meridian),

30 May 1996, Wilson 8094 (OSC); Highway 263 bridge over the Klamath River, at intersection with Highway 96, SE corner of

bridge, T74N R6W S18SW 1/4 of SW 1/4 (MD meridian), 30 May 1996, Wilson 8095 (OSC); Indian Scotty Campground, T44N

R11W S26SW 1/4 of NE 1/4 (MD meridian), 1 Jun 1996, Wilson 8112 (OSC); Quartz Valley Road, S of the Charity Mission, N of

bridge over river (and N of Forest Service Road 43N21), T43N R10W S3SW 1/4 of NE 1/4 (MD meridian), 1 Jun 1996, Wilson

8115 (OSC); Idlewild Campground on the Salmon River at the intersection of Forest Service roads 41N37 and 1CO1., T40N

R10W S18SW 1/4 of SW 1/4 (MD meridian), 1 Jun 1996, Wilson 8117 (OSC); Old Edgewood-Weed road, which parallels the

l-5 on the E, 2.2 mi by road more or less W of N. Weed Blvd., and 2 mi more or less W of where a road crosses the railroad

tracks and intersects with this one from the NE, TA1N R5W S4 (MD meridian), 21 Jun 1996, Wilson 8162 (OSC); Stewart Springs

Road, T42N R5W S32SW 1/4 (MD meridian), 22 Jun 1996, Wilson 8168 (OSC); 0.9 mi from the Gazelle Road on the Stewart

Springs Road (Forest Service Road 17), at the first bend in the road, ca. 3.5 mi W and 1 mi N of Weed, T42N R5W S32SW 1/4

(MD meridian), 22 Jun 1996, Wilson 8173 (OSC). Sonoma Co.: 3 mi NW of Graton, 30 May 1937, Yates 6544 (DAV). Tehama

Co.: Tedoc Mountain, T28N R8W S29 (MD meridian), 22 Jul 1978, Smith & Nelson 10032 (HSC, JEPS); Tedoc Mountain, T28N

ROW S29 (MD meridian), 23 Jun 1979, Smith & Nelson 10162 (HSC). Trinity Co.: Morris Meadow, Stuart Fork of Trinity River,

21 Aug 1948, Alexander & Kellogg 5526 (UC); ridge road on South Fork Mtn. ca. 10 mi N of its junction with Highway 36, 16

Jun 1972, Anderson 5887 (HSC); South Fork Mt., along 1 SO 2, the road to Pickett Peak, 0.4 mi from its junction with Highway

36, 15 Jul 1971, Anderson s.n. (HSC); near Castle Rock, 25 Jun 1980, Baker 2250 (HSC); Indian Valley near Hayfork, 16 Jul 1965,

Bordon s.n. (DAV); T30N R12W S31SW 1/4 of NW 1/4 (MD meridian), 16 Jun 1980, Nelson & Nelson 5446 (HSC); T30N R12W S13

(MD meridian), 18 Jun 1980, Nelson & Nelson 5544 (HSC); Red Mountain, T26N R12W $20 (MD meridian), 9 Jun 1978, Nelson

& Sawyer 4135 (HSC); Eagle Creek Campground, Shasta-Trinity National Forest, T38N R7W S16NE 1/4 (MD meridian), 15 Jun

1979, Smith 10009 (HSC); T30N R11W 528 (MD meridian), 22 Jul 1980, Smith 10307 (HSC); Underwood Mountain Pass, 12 Jun

1965, Spellenberg 1104 (HSC, OSC); T25N R12W S11 (MD meridian), 28 Jun 1997, Sprecht 1105 (HSC); Mt. Eddy, TAON R6W S12

(MD meridian), 23 Jul 1976, Whipple 1506 (HSC).

OREGON: Douglas Co.: Beatty Creek ACEC/RNA, T30S R6W S21 and/or T30S R7W 525, 2 Jun 1994, Brainerd € Kuykendall

BLW9856 (OSC). Jackson Co.: Siskiyou Pass, junction old Hwy 99 w/ l-5 just N of Hilt, T41S R2E S8SE 1/4, 16 Jun 1998, Chambers

6130 (OSC); Mount Ashland, T40S R1E S, 6 Jul 1958, Dennis 1085 (OSC); Rogue River near Elk Creek, 9 Jun 1930, Henderson

13344 (ORE); Dutchman's Peak, summit area, 7400 ft, 6 Aug 1961, Hutchison 985 (JEPS); T40S R2W S7 E half of section, 23 Jun

1955, Jeffers 61 (OSC); Lower Applegate Creek, 18 Jul 1899, Leiberg 4096 (ORE); Grizzly Peak, 17 Jul 1913, Peck 4470 (WILLUJ; Dry

Summit of Mt. Ashland, 19 Jul 1913, Peck 9307 (WILLU); Long John Creek, T40S R1W S35, Wheeler 2987 (US); Collings Mt. near

Steamboat, T40S RAW 535, 13 Jul 1950, Whitaker 169 (WS); Observation Peak, T41S R2W S12, 14 Jul 1950, Whitaker 271 & 248

(WS); Emigrant Creek, T40S R2E S1, 20 Aug 1949, Whitaker 349 (WS); Deadman Pt. near Dutchman's Peak, T40S R2W S33, 13

Jul 1950, Whitaker 222, 224, 227 (WS); Big Red Mountain (summit), TA0S R1W S31, 15 Jul 1950, Whitaker 330, 326 (WS); Ashland

Peak, Siskiyou Mts., [395 R1E $34, 21 Aug 1949, Whitaker 55355 (WS); near Canberry Campground, 1415 RAW S10, 28 May 1994,

Wilson 6936 (OSC); Cantrall-Buckley County Park, 1.5 mi SW of Ruch, T38S R3W $33, 20 Jun 1996, Wilson 8137 (OSC); Baldy Peak

Trail, TAOS R3W S22, 2 Jul 1996, Wilson 8197 (OSC); South-facing slope E of the summit of Mount Ashland, T40S R1E $17, 6 Aug

1993, Wilson, Kuykendall, Otting & Zika 6339 (OSC); Private inholding between the Klamath and Rogue River National Forests,

0.2 mi (by road) E of Siskiyou Gap and 0.4 mi (by road) W of the intersection of forest service roads 20 and 22, where the

road crosses a small seep, T40S R1W S34, 6 Aug 1993, Wilson, Kuykendall, Otting & Zika 6381 (OSC). Josephine Co.: Woodcock

Mountain, T39S R8W S19, 20 Jun 1995, Brock 560 (OSC); Fiddler Mountain, 11 May 1974, Chambers 3947 (OSC); Illinois River

Valley, Rockydale Rd. 2 mi. N of junction with Waldo Road, T40S R8W S15NE, 15 Jun 1998, Chambers 6117 (OSC); Kalmiopsis

Wilderness Road, 1385 R8W S30, 11 Jun 1984, Fredricks 285 (OSC); Deer Creek, 4 mi from Selma, 11 Apr 1926, Henderson 5949

(ORE); Grayback Mountain, 13 Jul 1930, Henderson 13339 (ORE); Eight Dollar Mountain, near Selma, Illinois River Valley, The

Nature Conservacy Preserve, E base of the mountain, 11 May 1983, Kagan 5118301-2 (OSC); 22 mi W of Gasquet Trail on

O'Brien-Sourdough Road, Kruckeberg 1861 (WS, WTU); Siskiyou N.F., junction roads 4402 & 4402-19, 4 mi SW of O'Brien, T41S

ROW SANE 1/4, 9 May 1984, Shelly 696 (OSC); Illinois River Valley, W of Selma, 0.5 mi up Oregonite Trail from Store Gulch GS,

T37S ROW S34SW 1/4 of SW1/4, 7 Jun 1969, White € Lillico 187 (ORE); Illinois River Valley, W of Selma, hill E of Sixmile Creek,

1385 ROW S2NW 1/4 of SE 1/4, 8 Jun 1969, White & Lillico 210 (ORE); Onion Mountain Road, ca. 3 mi NW of Lookout junction,

T36S R8W 54, 20 Jun 1969, White & Lillico 289 (ORE); Hoover Gulch Trail, T38S ROW $30, 21 Jul 1969, White & Lillico 319 (OSC);

Hoover Gulch Trail, T38S ROW $30, 21 Jul 1969, White & Lillico 322 (ORE); Whetstone Butte, T38S ROW S30, 25 Jun 1969, White &

Lillico 336 (ORE); Limestone Trail, Grayback, T40S R6W S20, 10 Jul 1950, Whitaker 84 (WS); Murphy Creek, T29S R6W S2, 12 Jul

1950, Whitaker 166 (WS); Josephine Mountain, T39S ROW 520, 20 Jul 1950, Whitaker 410 (WS); No. 8 Gulch Trail, Browntown

near Holland, T40S R7W S22, 11 Jul 1950, Whitaker 119& 136 (WS); Holland-Browntown Road, 1405 R7W S4, Whitaker 145 & 147

(WS); Sucker Creek, Grayback area, T40S R7W S12, 2 Jul 1949, Whitaker 5583 (WS); Big Red Mountain, Ashland Area, Siskiyou

Mountains, T40S RTW $32, 22 Aug 1949, Whitaker s.n. (WS); Sexton Moutain, T34S R6W $23, 15 May 1994, Wilson 6834 (OSC);

Wilson, A new variety of Festuca roemeri 67

Near fen on Fiddler Mountain, 1.7 mi by road from bridge over the Illinois River, 0.7 mi from Forest Service Road 4201 on a

dirt road that intersections with 4201, 1 mi from (S of) the bridge, T38S R8W S30, 14 May 1994, Wilson, Camacho & Otting 6786

(OSC); Slope above and NE of Illinois River bridge, Eight Dollar Mountain, 24 May 1996, Zika 12866 (OSC).

ACKNOWLEDGMENTS

This research was done as part of a doctoral thesis submitted to Oregon State University, under direction

of Dr. Aaron Liston. I appreciate access to the following herbaria: DAV, HSU, ORE, OSC, UC, US, and

WILLU. I thank David Gernandt, Instituto de Biología, Universidad Nacional Autónoma de México, for

translating the abstract, and Juan N. Zavala, Beth Parsons, and Krishnawan Panashatham of the Oregon

State University Seed Lab for technical assistance with flow cytometry to determine ploidy. Funding for

part of the field research was provided by the Bureau of Land Management (BLM), Medford and Roseburg

District Offices. I thank Russ Holmes and Mark Mousseaux of the BLM and Wayne Rolle and Maria Ulloa

of the USDA Forest Service for permits and other assistance, and Carex Working Group members Richard

Brainerd, Danna Lytjen, Bruce Newhouse, Keli Kuykendal, Nick Otting, and Peter Zika for field assistance

and fruitful discussions.

REFERENCES

AIKEN, S.G., L.L. Consaut, A. SpipLE, and B. May. 1994. Allozyme and morphological observations on Festuca hy-

perborea, compared with F. baffinensis and E brachyphylla (Poaceae) from the Canadian Arctic. Nord. J. Bot.

14:137-143.

ALEXEEV, E.B. 1985. Festuca L. (Poaceae) of Alaska and Canada. Novosistematiki vysshikh rastenii [New Develop-

ments in Higher Plant Taxonomy] 22:5-34. [In Russian]

Dugé, M. and P. Morisset. 1996. La plasticité phénotypique des caractéres anatomiques foliaires chez le Festuca

rubra L. (Poaceae). Canad. J. Bot. 74:1708-1718.

HitcHcock, C.L., A. CRoNauist, M. Ownsey, and J.W. THompson. 1969. Vascular plants of the Pacific Northwest; Part 1;

Vascular Cryptogams, Gymnosperms, and Monocotyledons. University of Washington Press, Seattle.

Pavuick, L.E. 1983. The taxonomy and distribution of Festuca idahoensis in British Columbia and northwestern

Washington. Canad. J. Bot. 61:345-353.

RAMESAR-FORTNER, N.S., S.G. AikeN, and N.G. DencLer. 1995. Phenotypic plasticity in leaves of four species of arctic

Festuca (Poaceae). Canad. J. Bot. 73:1810-1823.

WENotL, J.F. and N.F. Weeden. 1989. Visualization and interpretation of plant isozymes. In D.E. Soltis and PS. Soltis,

eds. Isozymes in plant biology. Dioscorides Press, Portland, Oregon. Pp. 5-45

WHITAKER, R.H. 1960. Vegetation of the Siskiyou Mountains, Oregon and California. Ecol. Monogr. 30:279-338.

WILson, B.L. 1997. A"new" native fescue of western Oregon prairies. In: T.N. Kaye, A. Liston, R.M. Love, D.L. Luoma,

R.F. Meinke, and M.V. Wilson, eds. Conservation and management of native plants and fungi. Native Pl. Soc.

Oregon, Corvallis. Pp. 153-161.

WiLson, B.L. 1999. Fescue taxonomy in the Pacific Coast States. Ph.D. thesis, Department of Botany and Plant

Pathology, Oregon State University, Corvallis.

68 Journal of the Botanical R h Institute of Texas 1(1)

BOOK REVIEWS

Lester Rowntree. 2006. Hardy Californians: A Woman's Life with Native Plants. New Expanded

Edition. (ISBN 0-520-25051-6, pbk.). The University of California Press, Berkeley, CA 94704, U.S.A.

(Orders: California Princeton Fulfillment Services, 1445 Lower Ferry Road, Ewing, NJ 08618, U.S.A.,

www.ucpress.edu, 609-883-1759, 609-883-7413 fax). $19.95, 391 pp., 81 b/w photographs.

Like the very native California flora she sought out to find, 52 year old Lester Rowntree was tenacious. A self-proclaimed “lady-gypsy,”

who traveled California's backroads and walked its mountain trails, often alone, sometimes with a pack animal to carry her camera

and collecting equipment. Lester carved out a unique and soul nurturing authentic life—the life of a field botanist, writer, lecturer, and

gardener. In her book Hardy Californians, she describes landscaq ] places that would soon disappear, a rural and pastoral California

that would fall victim to postwar “progress

The original Hardy Californians published in 1936 became a classic over the years. Unfortunately, it also became a rare book after

the bookplates were sacrificed to the 1940s war effort. In 1980, a year after Lester’s death at age 100, Hardy Californians was reprinted

as a paperback, yet this edition also is now scarce.

This new and expanded edition is a reprint of Lester’s original version as well as new material: a biographical sketch of Lester by

her grandchildren, Lester B. Rowntree and Rowan A. Rowntree. Also included is an essay by Judith Lowrey, an award-winning writer

and native plant horticulturist who, DIC never having met Lester, captures much of Lester's essence by Dunne her prolific writ-

ings and professional contributions with added photographs of Lester at different stages of her life. The sixty-four p graphs of native

plants have been reproduced anew from Lester's original negatives, taken in the early 1930s with a large-format camera. ie included

in the back of the book is an updated species list of plants referenced by Lester in the original 1936 edition of Hardy Californians.

Often described as a female John Muir, she was both a free spirit and a recluse living for months in the mountains on beans and

bread. In her words:

“The best places of all were in the high mountains, where I knew no one was camping above me. I used to love sleeping at the edge of snow

banks during thaw time to watch the alpines open with the rising sun.

tho c after tl E ES the forest trees —l ME RU CUT Pane f A A PRU O

J J

take off your clothes and dance in the rain. Soon you know that the elements have seen you, too.”——Linny d MH Illustrator,

email: a0005835@airmail.net.

WiLLiam W. DunmirE. 2004. Gardens of New Spain: How Mediterranean Plants and Foods Changed

America. (ISBN 0-292-70564-6, pbk). University of Texas Press, PO. Box 7819, Austin, TX 78713-7819,

U.S.A. (Orders: http://www.utexas.edu/utpress, 512-471-4032). $24.95, 375 pp., illustrated, 6" x 9”.

William Dunmire discusses how plants and animals from the Mediterranean world arrived in the Americas, and how they were culti-

vated in this new setting. It is a fascinating story, for it involves the many cultures and geographic regions (the Middle East, Asia, and

Africa) that gave so much to Spain. The story also involves the many people who brought these crops and animals to the Americas. The

introduction of these new plants and animals forever changed the face, and taste, of the land.

Written in the tradition of historical and cultural geography, the reader will learn about each new crop and animal, its use and

habitat in the Old World and the challenges facing its introduction into the New World. Many crops were first introduced into Mexico,

and then later spread to New Mexico, Arizona, California, and Texas. When the Spanish left for the New World, they brought along the

plants and animals of their homeland— wheat, melons, grapes, vegetables and a cornucopia of Mediterranean fruit. Watermelon and

cantaloupe seeds were among Pueblo Indian trade items imported from Mexico, which means a European presence was felt here long

before the Spanish themselves appeared. Some of their offerings thrived and became staple crops alongside the corn, beans and squash

that had traditionally sustained the original Americans. Other imports couldn't adapt or didn't please local palates. This intermingling

of Old and New World plants gave rise to many of the culinary dishes and foods we enjoy today.

His earlier works include: Wild Plants of the Pueblo Province (1995), A Readable Guide to Southwestern Native American Ethnobotany

(1997), and Wild Plants and Native Peoples of the Four Corners (1997).

Gardens is recommended for anyone interested in plants and their uses. It is a good read and well organized with an index, glos-

sary, and extensive bibliography.— Gary Jennings, Library, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060,

U.S.A.

J. Bot. Res. Inst. Texas 1(1): 68. 2007

DESCRIPTION OF CAREX KLAMATHENSIS (CYPERACEAE) ARARE SEDGE OF

THE KEAMATH REGION OF OREGONAND CALIFORNIA, US A

Barbara L. Wilson!, Richard E. Brainerd!, Lawrence P. Janeway?,

Keli Kuykendall', Danna Lytjen!, Bruce Newhouse!,

Nick Otting!, Stephen Meyers?, and Peter F. Zika*

! Carex Working Group, 2710 Emerald Street, Eugene, Oregon 97403, U.S.A.

? Biological Sciences Herbarium, California State University, Chico, California 95929, U.S.A.

? Department of Botany and Plant Pathology, 2082 Cordley Hall, Oregon State University, Corvallis, Oregon 97331, U.S.A.

^ Herbarium, Box 355325, University of Washington, Seattle, Washington 98195, U.S.A.

Contact author: Barbara L. Wilson, bwilson@peak.org

ABSTRACT

i i d f tine f : +}

A previously undescribed sedg

t Oregon and three California sites is described here as globally rare Carex

klamathensis. This species is rhizomatous, with glaucous foliage and pale, more or less papillose, obovate perigynia with bent beaks. It

resembles and may be most closely related to the midwest North American taxa Carex meadii and Carex tetanica. It differs from both of

these in its achene surface morphology. In addition, it has smaller perigynia than C. meadii and wider staminate spikes than C. tetanica.

Itis most easily confused with a form of C. hassei that grows in serpentine fens in northwest California. That taxon usually has a mix of

flowers with two or three stigmas in the same plant. Carex klamathensis consistently has flowers with three stigmas and is a taller, more

robust plant with wider staminate spikes

Key Wonps: Carex klamathensis, Carex hassei, serpentine endemic, Paniceae

RESUMEN

Se describe en este trabajo una ciperácea previamente no descrita de ciénegas serpentinícolas del suroeste de Oregón y de tres sitios

en California como una endémica restringida, Carex klamathensis. La especie es rizomatosa con hojas glaucas y pálidas, más o menos

papilosas, periginio obovado con picos doblados. Parece estar más estrechamente relacionada a los taxa del medio oeste de América

del Norte, Carex meadii y Carex tetanica. Difiere de esas dos especies por la morfología de la superficie del aquenio. Además, posee un

periginio más pequeño que C. meadii y espigas estaminadas más anchas que C. tetanica. Se puede confundir más fácilmente con una

forma de C. hassei que ocurre en ciénegas serpentinícolas del noroeste de California. Ese taxón usualmente tiene una mezcla de flores

con dos o tres estigmas en la misma planta. Carex klamathensis consistentemente tiene flores con tres estigmas y es una planta más alta,

más robusta, con espigas estaminadas más anchas.

INTRODUCTION

A strongly glaucous sedge in Carex section Paniceae has long confused botanists studying serpentine fens

in southwest Oregon. Its long rhizomes, three stigmas, and pale, papillose, perigynia initially led to its

misidentification as Carex californica L.H. Bailey, a widespread but local plant of disturbed meadows and

roadsides west of the Cascade Range. As botanists became more familiar with the serpentine species in the

late 1980s, they realized that its indistinct perigynium beaks and pale shoot bases differentiated it from C.

californica, which has tubular perigynium beaks and red-brown shoot bases, rhizomes, and scales. Then the

species was identified as Carex livida (Wahlenb.) Willd., a species of northern bogs and known from a few

Oregon sites and one California wetland (Bolander 4745, Mendocino County, California; specimen at UC). In

the late 1990s Oregon botanists found that the serpentine plants differed from C. livida not only in habitat

but also in inconspicuous but consistent morphological traits of the leaves, inflorescence, and perigynia.

At the same time Lawrence Janeway, working independently in California, discovered Carex livida-like

sedges at three isolated springs on serpentine substrates (Fig. 1). This plant matched no other described spe-

cies, although it resembled the Midwestern species C. meadii Dewey and C. tetanica Schkuhr (A.A. Reznicek,

pers. comm.).

Further study revealed that the new species can be difficult to distinguish from a form of C. hassei L.H.

J. Bot. Res. Inst. Texas 1(1): 69 — 77. 2007

Journal of the Botanical Research Institute of Texas 1(1)

puc

ants Pass

Redding

b) X

(9 Carex Kamațhensis

"ES

Clearlake

Wilson et al., A new Carex from Oregon and California 71

Bailey which lives in serpentine fens in northwest California and has a mix of 2-stigma and 3-stigma flowers

in the same plant (Wilson et al. in preparation).

TAXONOMY

Carex klamathensis B.L. Wilson & L.P Janeway, sp. nov. (Figs. 2-3). Type: U.S.A.: Orecox: Josephine Co.: BLM Darlingtonia

fen along Fight Dollar Road, on S base of Eight Dollar Mountain, 0.9 mi from Route 199, full sun, peaty wet ground over serpentine,

elev. 420m, 42°14 N, 123?40' W, 18 May 2004, Peter E Zika 19642 (HoLotyPE: OSC; isotypes: CHSC, MICH, MO, UC, WTU).

Carex klamathensis a Carex livida differt foliis latioribus, perigyniis obovatis brevioribus. Species haec Carex meadii affinis sed differt

perigyniis brevioribus, acheniis paginis reticulatis minute. Species haec Carex tetanica affinis sed differt spicis terminalibus staminatis

latioribus, acheniis paginis reticulatis minute. A Carex hassei differt perigynibus stigmatibus 3

Description: Plant rhizomatous, the rhizomes whitish to medium brown, occasionally dull orange brown,

paler than the dark to medium brown (rarely straw-colored) scales, slender, 1-2(22.5) mm wide exclusive of

sheathing scales. Shoot bases medium to dark brown, phyllopodic. Leaves glaucous with the sheaths basally

white, with lower surface densely papillose between and sometimes over the veins, 18 to 50 cm long, the

wider leaves 2-6 mm wide (average 3.8 mm). Ventral surface of leaf sheath hyaline, the mouth shallowly

U-shaped. Culms 30-100 cm long, scabrous or not, longer than the leaves, erect at anthesis but sometimes

bending over by the time the perigynia ripen. Inflorescence 5-23 cm long (average 14.5 cm), with 1-2(-4)

lateral spikes. Bract of lowest spike 3-14 cm long (average 9.1 cm) including a sheath (0.8-)1.5-4(-8) mm

long, 0.33 to 1.5 (average 0.67) times as long as the inflorescence. Lateral spikes pistillate, usually one per

node, (0.6-)1.5-2.5 cm long, 4-7 mm wide, the uppermost usually 1.5-6 cm or more below the terminal

spike, but sometimes as close as 0.3 cm below the terminal spike. Perigynia moderately crowded in the

spike but the lowermost sometimes remote, the internodes in the middle of the spike 0.1-1.2 mm (aver-

age 0.5 mm) long. Terminal spikes all staminate in most populations, but in some populations, including

those in California and on Sexton Mountain, Oregon, these may be gynecandrous or, less often pistillate,

androgynous, or with staminate and pistillate flowers mixed. Staminate terminal spikes 1.3-2.7 cm (aver-

age 1.8 cm) long, 2-5 mm (average 3.6 mm) wide, 3.7—9.5 (average 5.3) times as long as wide, with 50-190

(average 112) flowers. Lowest staminate scale yellowish to reddish brown, paler near the midrib, acute

to obtuse, often awned, 2.2-5.1 mm (average 2.3 mm) long excluding awn, the awn if present 0.33 mm

(average 1.1 mm) long. Other staminate scales similar in color to the lowest, with the apex rounded and

sometimes mucronate. Pistillate scales 3-nerved (the lateral nerves sometimes faint), reddish brown, dark

brown, or rarely gold, the midrib and surrounding area green, white, or light brown, the edges sometimes

pale, 1.9-2.8 mm long excluding awn, the apex rounded or obtuse, less often acute, sometimes mucronate

to awned, the awn, if present, up to 1.5 mm long. Perigynia obovate to elliptic, 1.7-3.6 mm (average 2.9

mm) long, (0.8—)1.2-1.6(-1.8) mm (average 1.4 mm) wide, 1.6-2.4 (average 2.1) times as long as wide,

light green, tan, or whitish, sometimes marked with dark brown distally, papillose particularly toward the

beak or rarely smooth, the base succulent when fresh and drying withered, the beak usually curved, the

distance from beak tip to top of achene (0.1-)0.4—0.7(-1) mm. Stigmas 3, or occasionally 2 on 0—596(-1596)

of flowers that have viable achenes. Achene greenish yellow when young, ripening dark brown, trigonous,

or lenticular if stigmas 2, 1.6-2.7 mm (average 2.2 mm) long including stub at persistent base of deciduous

style, 0.7-1.7 mm (average 1.2 mm) wide. Achene width/length ratio 0.67(0.44—0.88). Anthers 2.5-3.5 mm

long when dry.

Habitat.—Fens on ultramafic (serpentine) soils, often with Darlingtonia californica, in Oregon at 400—950

m elevation in Pinus jeffreyi savannah; in California at 1000-1140 m elevation in chaparral.

Range.—Several populations in Josephine County, southwest Oregon, and also found at isolated sites

in Colusa, Lake, and Tehama counties, California (Fig. 1).

KEY TO THE NORTH AMERICAN TAXA OF CAREX SECTIONS BIGOLORES AND PANICEAE

The key for section Paniceae is modified from Rothrock and Reznicek (2002). Percents of pistillate flowers

with various stigma numbers refer to perigynia that produce hard, dark, apparently viable achenes. In all

Carex, aborted ovaries may have only two stigmas.

fal, Dat A ID hi rr

72 Journal of of Texas 1(1)

1. Stigmas prevailingly 2: 0-10%(-67% in C. hassei) of pistillate flowers with 3 stigmas Carex section Bicolores

2. Perigynia at maturity succulent throughout, orange to whitish, drying dark brown C. aurea

2. Perigynia at maturity dry throughout or succulent only at base, green, whitish, or tan.

3. Pistillate scales black with green midrib C. bicolor

3. Pistillate scales gold to dark brown.

4. Lateral spikes crowded, overlapping; terminal spike usually gynecandrous, perigynia usually crowded

with internodes between them 0.2-0.7 mm; proximal staminate scales (2-)2.5-3.7 mm, awnless;

scales rounded or obtuse; 90-10096 of pistillate flowers with two stigmas each C. garberi

4. Lateral spikes often less crowded; terminal spike usually staminate; perigynia crowded or more

distant with internodes between them 0.2-1.5 mm; proximal staminate scales 3-6(-15) mm, acute

to awned; scales obtuse to acute, often awned; 33-10096 of pistillate flowers with two stigmas each

—

C. hassei

1. Stigmas prevailingly 3; (80%-)90-100% of pistillate flowers with 3 stigmas Carex section Paniceae

5. Perigynium apex contracted to a cylindrical beak (0.4—)0.6-1.8(-2.2) mm long.

6. Bladeless basal sheaths and proximal leaf sheaths pale brown; columns, leaves, and perigynia not or very

sparsely papillose C. vaginata

6. Bladeless basal sheaths and proximal leaf sheaths strongly tinged with reddish purple; culms, leaves,

and perigynia heavily papillose.

7. Perigynia 4.2-6.8 mm long; beak 0.8-1.8(-2.2) mm long; range eastern C. polymorpha

7. Perigynia 3.4-4.2 mm long; beak 0.5-1 mm long; range western C. californica

5. Perigynium apex tapering and beakless, indistinctly beaked, or contracted to a beak less than 0.5 mm

long.

8. Lateral spikes nodding on flexible peduncles C. laxa

8. Lateral spikes erect or ascending on stiff peduncles.

9. Perigynia beak straight, cuneately tapering; leaves channeled, glaucous C. livida

9. Perigynia beak curved, concavely tapering (at least on one side); leaves flat or folded, glaucous or

10. Bladeless basal sheaths and proximal leaf hs st ly tinged with reddish | le plant

a loose clumps to extensive closed colonies of vegetative shoots from superficial rhizomes.

. Widest leaves 1.8-3(-4) mm wide; plants colonial with longest rhizomes 2.5-18 cm;

£,

habitat woodlands C. woodii

11. Widest leaves 3.5-6 mm wide; leaves loosely cespitose with longest rhizomes to 2 cm;

habitat granite balds and cliffs C. biltmoreana

Bladeless basal sheaths and proximal leaf sheaths brownish or faintly, irregularly tinged with

reddish purple; plants usually with vegetative shoots widely scattered and inconspicuous from

Ee rhizomes.

. Inflorescences usually 1.7-3.5(-4.3) times as long as bract (measured from node of proximal

al spike).

nonbas

13. Perigynia 0.6-1.4(-1.8) mm wide; achenes 0.7-1.7 mm wide; range Oregon and

California C. klamathensis

13. Perigynia 1.4-2.4 mm wide; achenes 1.8-2.9 mm wide; range Eurasia, introduced to

northeastern North America C. panicea

12. Inflorescence usually 0.9-1.6 times as long as bract (measured from node of proximal non-

basal spike).

14. Achenes (1.5-)1.7-2.2(-2.5) mm wide C. meadii

14. Achenes 07-1 .7(-1.9) mm wide.

15. Achene surface reticulate with a papilla filling each compartment outlined by

the ridges; terminal spike narrow (1.8-3 mm wide); range east of the Rocky

Mountains C. tetanica

. Achene surface reticulate but flat between ridges (or with a tiny papilla in the

center of each compartment); terminal spike wide, (2-5 mm wide); range in

Oregon and California C. klamathensis

DISCUSSION

Carex klamathensis is easily confused with other rhizomatous sedges that have glaucous foliage, pale brown

or whitish (not red-brown) plant bases, and pale, more or less papillose perigynia. Compared to C. livida,

C. klamathensis has wider leaves, more staminate flowers, and shorter, obovate (not fusiform) perigynia

(Table 1; Fig. 3 and 4).

Wilson et al., A new Carex from Oregon and California

Fic. 2. Carex klamathensis, habit.

£s+haD o ID L

74 Journal of t titute of Texas 1(1)

Fic. 3. P igy H l pistillat I £f. Ll "m

Carex klamathensis differs from C. meadii and C. tetanica of midwestern North America not only in

range, but also by its wider leaves and different achene surface. In C. klamathensis, the achene surface is

reticulate with low ridges and a flat space between the ridges, or with a minute central papilla in that flat

space. In the Midwestern taxa, the achene surface is papillate with a large papilla occupying virtually all of

the surface between the low ridges. In addition, C. klamathensis has smaller perigynia and narrower achenes

than C. meadii, and lives in a wetter habitat (Table 2; Fig. 4). Carex klamathensis has more staminate flowers

and therefore a wider staminate spike than C. tetanica (Table 2).

Carex klamathensis can be difficult to distinguish from a form of C. hassei that lives in serpentine fens

in the mountains of northwest California (Table 3). In general, C. hassei has two stigmas per pistillate flower

and C. klamathensis has three, although any three-stigma flowers that abort may have two stigmas. This

results in variation in achene shape; two-stigma flowers produce lenticular achenes and three-stigma flowers

produce trigonous achenes. In C. klamathensis, although most individuals have only 3-stigma flowers, some

plants have a few (less than 1596, usually less than 1096) flowers that have two stigmas and produce hard,

lenticular achenes. In the C. hassei from serpentine fens, few populations have only plants with 2-stigma

flowers. In most populations, the proportion of 2-stigma perigynia varies from (33-)40-90%. Two-stigma

and three-stigma flowers occur on the same plant, mixed in the same spike, and they are all capable of

Wilson et al., A new Carex from Oregon and California

Table 1. Traits distinguishing C. klamathensis from C. livida.

Trait

Carex klamathensis

Carex livida

Substrate

Leaf width (mm)

Terminal staminate spike

width (mm)

Flowers, terminal

staminate spike

Number of lateral spikes

3 or more)

Perigynium length (mm)

Perigynium shape

some perigynia)

Perigynium beak

serpentine

wider; average 3.7, (range 1.9-6 mm)

wider; average 3.5 mm (range 2-5)

average 112 (range 40-190)

average 2.2 (range 1-4, 4096 with

average 2.9 (range 1.7-3.6)

obovate to elliptic (rarely fusiform on

bent (rarely straight on some of

non-serpentine

narrower; average 2.2 (range 1.5-3.2)

narrower; average 2.2 (range 1.4-4.8)

average 50 (range 8-71)

average 1.7 (range 1-3 but only 196 with 3)

average 3.8 (range 3.1-4.8)

fusiform

straight

the perigynia)

TABLE 2. Traits distinguishing C. klamathensis from C. meadii and C. tetanica. Measurements are average and, in parentheses,

range.

Trait Carex klamathensis Carex meadii Carex tetanica

Range Pacific coast states Midwest Midwest and east

Substrate serpentine not serpentine not serpentine

Habitat fens mesic meadows wet sites

Leaf width (mm) 3.8(2-6) 2.8(2.4-3.3) 2.5(1.8-3.3)

Terminal spike width (mm) 3.6(2-5) 301.234) 2A4(1.8-3.1)

Staminate flowers 112(40-190) 112(48-174) 70(40-120)

Height (cm) 57(30-100) 32(23-47) 26(1 2-34)

Perigynium length (mm) 29(1.7-3.6) 3.6(3.3-4) 2.9(2.4-3.6)

Perigynium width (mm) 1.4(1.2-1.6) 2.0(1.6-2.5) 1.6(1.2-1.9)

Achene surface reticulate papillose papillose

TABLE 3. Selected statistically significant (p «0.05) traits distinguishing C. klamathensis from the form of C. hassei that grows in

serpentine fens in northwest California. Measurements are average and, in parentheses, range.

Trait C klamathensis C hassei

Average (range) Average (range)

Culm length (cm) 57(30-100) 33.3(15.2-45.3)

Leaf width (mm) 3.8(2-6) 2.8(1.8-3.7)

Inflorescence length (cm) 14.4(5.1-23.3) 10.0(3.9-17.8)

Inflorescence bract length (cm) 9.1(3.3-14.0) 6.9(3.2-15.2)

Terminal Spike Length (cm) 1.8(1.3-2.7) 1.4(1.1-2.3)

Terminal Spike Width (mm) 3.6(2-5) 2.8(1.4—4.5)

Staminate flowers in terminal spike 112(40-186) 81(57-152)

Lowest staminate scale, length (mm) 4,3(2.2-5.1) 3.2(1.9-5 7)

Perigynia (96 with 3 stigmas) 97%(85-100%) 37%(0-62%)

Perigynium length (mm) 29(1.9-3.5) 2.5(2.2-2.9)

Perigynium length/width ratio

Achene width/length ratio

0.48(0.37-0.58)

0.67(0.44-0.88)

0.54(0.46-0.63)

0.74(0.48-0.93)

£+sha D o ID L

76 Journal of t titute of Texas 1(1)

1 mm

Fic. 4. Perigynia of C. klamathensis and similar taxa. A. Carex hassei. B. Carex klamathensis. C. Carex livida. D. Carex meadii. E. Carex tetanica. Scale at left

is 1 mm.

producing hard, dark, apparently viable achenes. In general, C. klamathensis is a more robust plant, taller,

with longer and wider terminal spikes, longer inflorescence nodes, wider leaves, and slightly longer perigy-

nia. However, populations of delicate C. hassei may produce occasional robust plants. (The robust C. hassei

plants observed were identified as C. hassei because they had 50-91% two-stigma flowers.) Because of this

variation in C. hassei, ranges for most measured traits overlap greatly, even though average dimensions of

most traits differ significantly (Table 3). The two taxa also differ in traits such as the color and stiffness of

the foliage, which are hard to quantify.

Carex klamathensis is a globally rare species of fens and springs, endemic (Safford et al. 2005) to ser-

pentine substrates. Although 35 specimens were examined (Appendix), these represent only 3 California

and 12-15 Oregon populations. Four of the sites (10 collections) were on Eight Dollar Mountain, Josephine

County, California, and two more sites were within 3 km south of that mountain.

Populations are probably stable where habitats are stable, but habitats are threatened by road building,

recreational use of serpentine wetlands (particularly the effects of off-road vehicles), and mining. These

activities can harm C. klamathensis populations directly by killing plants or indirectly by altering water

flow. The sedge probably survives fire well, both because its rhizomes are protected underground in moist

soil and because its microhabitat does not carry intense ground fire well. However, it is vulnerable to those

fire suppression activities that involve bulldozers. The three California populations, which are somewhat

genetically distinct from the Oregon populations (Wilson et al. in preparation), are all small and isolated.

More than half of one has been destroyed in recent years by bulldozing associated with mining. We hope

that clarifying C. klamathensis taxonomy and morphology will aid in its preservation.

APPENDIX

CAREX KLAMATHENSIS SPECIMENS EXAMINED

l = from Day’s Gulch, the site of most collections, reported under various names

U.S.A. CALIFORNIA. Colusa Co.: Bear Wallow Spring; on N side of Lovelady Ridge about 0.5 mi NE of Pacific Point, T16N RO7W

$23 $1/2,5 Jul 1999, Janeway & Castro 6497 (CHSC); Lovelady Ridge, Bear Wallow Springs, 28 Jun 2002, Wilson et al. 10702 (OSC).

Lake Co.: Kanaka Glade, a spring at the head of Spanish Creek East Fork, near the top of Pacific Ridge, T16N R6W S30 SW 4

of NW %, 7 Jul 1999, Janeway & Isle 6492 (CHSC); Kanaka Glade, a spring at the head of Spanish Creek East Fork, near top of

Pacific Ridge, 3 Jul 1998, Janeway & Castro 5714 (CHSC, OSC); Kanaka Glade, 19 May 1985, Stebbins C532 (DAV); Kanaka Glade,

28 Jun 2002, Wilson et al. 10704 (OSC'). Tehama Co.: Pepperwood Springs, near top of Raglin Ridge, T25N RO7W S21 SE Ya of

NW 1⁄4, 11 Jun 2000, Janeway & Isle 6785 (CHSC, OSC); Pepperwood Springs, 20 Jun 2003, Wilson € Brainerd 10951 (CAS, DAV,

MICH, OSC); Pepperwood Springs, 28 Jun 2002, Wilson et al. 10708 (OSC, WTU). OREGON. Josephine Co.: Frank's Fen, 18

Wilson et al., A new Carex from Oregon and California 7]

Jun 2000, Brainerd & Newhouse BLW10403 (OSC, UC); Fiddler Mtn. Road, above Josephine Creek, 1.8 mi S of bridge over Illinois

River by Eight Dollar Mtn., 11 May 1974, Chambers 3958! (OSC+); BLM fen, less than 1 mile down Eight Dollar Mtn. Rod., W of

Rte. 199, 23 Mar 1996, Clery 56 (OSC); West bank of Josephine Creek, about 150 m upstream from ford, 19 Jul 1981, Greenleaf

1186 (OSC); Whiskey Creek, 17 Jun 1999, Kuykendall et al. BLW10021 (OSC); Josephine Creek, 25 Jun 1930, Leach 2836 (ORE);

TNC Bog/$8 Mtn., 13 Jun 2003, Newhouse & Kuykendall 2003-001 (WTU), 2003-002 (CHSC), 2003-003 (MO), 2003-004 (DAV,

OSC); Star Flat, 14 Jun 2003, Newhouse & Kuykendall 2003-006 (OSC) & 2003-007 (CHSC); Days Gulch Botanical Area, 14 Jun

2003, Newhouse & Kuykendall 2003-008! (OSC) and 2003-009! (NY); Mars Fen/Rough & Ready Creek, 14 Jun 2003, Newhouse

& Kuykendall 2003-010 (OSC), 2003-11 (CHSC, UC, WTU), and 2003-012 (MICH, SOC); south base of Sexton Mountain, 20 May

1948, Peck 24796 (WILLU); Eight Dollar, 18 Jun 1999, Wilson & Kuykendall 10041 (DAV, RSA, UC); Fiddler Mtn., 18 Jun 1999,

Wilson & Kuykendall 10042! (MICH, WTU) and 10044! (OSC, UC); Mike's Gulch, 2 Jul 2003, Wilson & Kuykendall 10960 (OSC);

Siskiyou NF, near fen on Fiddler Mountain, 1.7 mi by road from bridge over the Illinois River, 0.7 mi from Forest Service Road

4201 on a dirt road, 14 May 1994, Wilson et al. 6782! (OSC); Woodcock Bog, 17 Jun 1999, Wilson et al. 10013 (CHSC, OSC, WTU);

Woodcock Bog, 18 Jun 1999, Wilson et al. 10053 (OSC); Fens on East side of Eight Dollar Mtn., 18 Jun 2000, Wilson et al. 10400

(MO, NY) and 10401 (CAS, MICH); Siskiyou National Forest, 19 May 1997, Zika 13081 (WTU); BLM fen on $8 Mountain Road, 18

May 2004, Zika 19642 (CHSC, MICH, MO, OSC, UC, WTU).

ACKNOWLEDGMENTS

This project was supported financially by the Bureau of Land Management (Oregon state office, Medford

and Salem Districts, and Arcata Field Office), and by the USDA Forest Service (Shasta-Trinity and Siskiyou

National Forests). These agencies and the Mendocino, Klamath, and Six Rivers National Forests also provided

non-monetary support. We thank the following agency botanists for their assistance: Joan Seevers, Ron

Exeter, Mark Mousseaux, Linda Mazzu, and Jennifer Wheeler (all of the BLM) and Julie Kierstead Nelson,

Maria Ulloa, David Isle, Susan Stresser, and Lisa Hoover (all of the USDA Forest Service). We thank Dr.

Anton Reznicek for advice about the taxonomy of sedges. We thank Richard Halse for use of the combined

collections of the Oregon State University herbarium, and for managing loans; Barbara Ertter for loan of

specimens from JEPS and UC, Anton Reznicek for loan of specimens from MICH, and James P. Smith and

Robin Bencie for loan of specimens from HSC. Kenton Chambers edited the description. The abstract was

translated by David Gernandt, Instituto de Biología, Universidad Nacional Autónoma de México and Manuel

González Ledesma, Centro de Investigaciones Biológicas, Universidad Autónoma del Estado de Hidalgo.

Rob Fiegener of the Institute for Applied Ecology made the map. Erin Stangel photographed the perigynia.

Rena Schlac drew the illustrations.

REFERENCES

RotHrock, PE. and A.A. Reznicek. 2002. Carex Linnaeus sect. Paniceae G. Don in J.C. Loudon. In: Flora of North America

Editorial Committee, eds. Flora of North America North of Mexico. Volume 23. Magnoliophyta: Commelinidae

(in part): Cyperaceae. Oxford University Press. Pp. 426-431.

SAFFORD, H.D., J.H. Viers, and S.P. Harrison. 2005. Serpentine endemism in the California flora: a database of serpen-

tine affinity. Madrono 52:222-257.

Witson, B.L., R.E. BRAINERD, L. Janeway, K. KUYKENDALL, D. LvrJEN, B. NewHouse, N. OrriNG, S. Meyers, and PF. Zika. (in prepara-

tion). Taxonomic confusion in Carex section Bicolores on the Pacific Coast.

78 Journal of the Botanical R h Institute of Texas 1(1)

BOOK REVIEWS

Aubrey EAGLE. 2006. Eagles Complete Trees and Shrubs of New Zealand. (ISBN 0-909010-08-0, hbk.,

slipcase). Te Papa Press, Museum of New Zealand, Te Papa Tongarewa, PO Box 467, Wellington, NEW

ZEALAND. (Orders: Museum of New Zealand, Cable St., PO Box 467, Wellington, New Zealand, www.

tepapa.govt.nz). NZ $200.00, Vol. 1, 544 pages; Vol. 2, 592 pages, richly illustrated, 9" x 11".

This beautiful two-volume set brings together Eagle's botanical artworks from her award winning and best selling 1975 and 1983 pub-

lications. The new edition includes over one hundred and seventy new paintings to depict every presently known native tree and shrub

in New Zealand. The total number of plants illustrated, in color and life-size, is more than 800.

These long-awaited volumes are the result of decades of skilled draughtsmanship and loving, painstaking observation, as well as

many years of field and laboratory research by New Zealand scientists. Audrey Eagle has been painting New Zealand's flora since 1954.

She began painting the specimens she collected with the sole purpose of learning their botanical names. In 1968 she was contacted by

William Collins Publishers to write and illustrate a book about native plants. With the aim of pri Samples of every genus of New

Zealand t leting the required illustrations. After 27 years of work, her first book was published in 1975 with

228 species illustrated. Her Sod P T in 1982 with a further 405 species illustrated. The current work has 173 additional

species illustrated for a total of 806 species. With one exception every plant has been illustrated from live specimens. She is probably the

only person in the world who has had the privilege of seeing live material of all the presently know New Zealand woody flora.

Painted from live specimens, every plant is depicted at life-size in technically superb detail, and many include detailed enlargements

showing all aspects of the leaves, flowers and fruit. As many of the native plants have small flowers, these and other details helpful to the

recognition of a species are shown enlarged. In the case of leaves that are too large to fit on the page, reductions of the complete plants

are shown. Each painting is accompanied with comprehensive notes on a facing page. Written in consultation with expert botanists,

these provide accurate and up-to-date descriptions of each plant, including notes on its habitat, distribution, nomenclature and more.

Her 1983 work included short biographies of many botanists and others after whom plants have been named. In curtailed form, this

information is included with the botanical information for each species.

The botanical names of plants follow the Plant Names Database of Landcare Research, Lincoln, New Zealand. If a plant is com-

monly recognized by another name in the New Zealand Plant Conservation Network database it is noted in the text. As many formally

recognized plant species names have changed since her earlier works, the currently recognized name of each plant is indicated in the

botanical description and is made to the relevant publication. Common and Máori names have been included whenever pos-

sible. The editorial style follows the New Zealand Journal of Botany.

Passionately involved in New Zealand's botanical community, Eagle was a founding member of the Royal Forest and Bird Protec-

tion Society of New Zealand. She has also served on the Nature Conservation Council. Her illustrations have been cited in numerous

published studies of New Zealand's flora. In 2001, Audrey Eagle was appointed a Companion of the New Zealand Order of Merit for her

services to botanical art. Her latest work was short-listed for the New Zealand Booksellers Choice Award for 2007. The award is given to

the book that New Zealand booksell enjoyed reading, selling and promoting in the previous year. This set is the accumulation of

Audrey Eagle's life's work. It is an outstanding contribution to botany in New Zealand and an essential addition to any botanical library

concerned with the flora of the Pacific Rim.—Gary Jennings, Library, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX

76102-4060, U.S.A.

Dav YETMAN. 2006. The Organ Pipe Cactus. (ISBN 0-8165-2541-2, pbk.). The University of Arizona Press,

355 S. Euclid Avenue, Suite 103, Tucson, AZ 85719, U.S.A. (Orders: www.uapress.arizona.edu, orders?

uapress.arizona.edu). $ 9.95, 80 pp., 36 color photos, 1 drawing, 1 map, 8" x 10".

When you first see one, it is obvious how the organ pipe cactus got its name. Its slender vertical branches, reaching for the heavens and

perhaps 30 feet tall, bring to mind the tubes ofa pipe organ. Whether standing alone or growing in a grove, these spectacular and intrigu-

ing plants are found exclusively in the United States in a jue area of n Sonoran Desert in the southwestern corner of Arizona.

David Yetman provides an in-depth and

plants that most Americans will

E E L

never h to see in person. Seven chapters explore where fhe grow; ithe she dalicondition required for their germina-

tion, growth, E survival; their position in a genus with more than twenty species; discovery by western Europeans; early history in

the Mayan civilization; uses as a commercial crop; and the future of the organ pipe cactus. Although it is the most common columnar

cactus worldwide, it is so unusual in the United States that it is only one of two cacti (the saguaro) and the Joshua tree, to have a national

preserve established to protect it.

This is a beautifully illustrated book, well written and is a handsome addition to his other two works. He is the co-editor of Gentry’s

Río Mayo Plants: The Tropical Deciduous Forest and Environs of Northwest Mexico, also published by the University of Arizona Press, and

the author of Mayo Ethnobotany: Land, History, and Traditional Knowledge in Northwest Mexico.— Gary Jennings, Library, Botanical Research

Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A

J. Bot. Res. Inst. Texas 1(1): 78. 2007

UNA NUEVA ESPECIE DE AGAVE, SUBGENERO LITTAEA (AGAVACEAE)

DE TAMAULIPAS, MÉXICO

Abisaí García-Mendoza Cuauhtémoc Jacques-Hernández y

Jardín Botánico, Instituto de Biología, U.N.A.M. Ang el 5a d r Bravo

A.P. 70-614, Delegación Coyoacán Centro de Biotecnoloaía smica, LPN

04510 México, D.F, MEXICO Blvd. del ee S/N Esq. Elías Piña

abisaigibiologia.unam.mx Col Narciso Mendoza

88700 Reynosa, Tamaulipas MÉXICO

aguilaquecaeayahoo.com

RESUMEN

Se describe e ilustra Ag ti ti li una nueva especie de la región de la Sierra de San Carlos, Tamaulipas, México.

dcin

Esta especie pertenece al grupo Marginatae y muestra similitudes con Agave xglomeruliflora (Engelm.) A. Berger.

ABSTRACT

Agave montium-sancticaroli is described and illustrated as a new species from the Sierra de San Carlos Region, Tamaulipas, Mexico.

The new species belongs to the Marginatae group, and is similar to Agave xglomeruliflora (Engelm.) A. Berger.

La identificación de las especies de Agave utilizadas para la obtención de mezcal (bebida destilada) en Tam-

aulipas, México, llevó al descubrimiento de una nueva especie, que se describe a continuación.

Agave montium-sancticaroli García-Mend., sp. nov. (Figs. 1-3). Tro: MÉXICO. Tamautiras: Municipio de San Carlos,

6 km al NE de Los nde in carretera Ciudad Victoria a San Carlos, matorral submontano de Acacia rigidula, Cordia boissieri,

I lia pall if istra, 307 m, 27 May 2004, A. García-Mendoza, C. Jacques, A. Mora & A. Salazar

7605 o MEXU: isoriPos; ENCB, TEX, UAT).

Planta perennis, 1.5—2 m alta; folia 50—80(-100) per rosulam, 100-120 cm longa, 9-12 cm ad medium lata,

lanceolata, concava, viridi-flavida, basem versus glauscescentia, margine dentato cum fascia cornea angusta,

dentibus 4-6 mm longis, 1.5-3 mm ad basem latis, rectis vel retrorsis, 1-3(+4.5) cm inter se distantibus, inter

grandibus uno vel aliquibus dentibus minimis; spina terminali 2.5—3.5 cm longa, per 12-16 cm decurrenti.

Inflorescentia racemoso-paniculata, 5.5-7 m alta, plus minusve fusiformis, (60—)80—140 ramis lateralibus

8-13 cm. Flores 10-20 per umbellam, 4.5—5(-5.5) cm longi, viridi-flavidi; perianthii tubus 5-6(-8) mm

longus; tepala 1.5-2.2 cm longa, 3-4(-7) mm lata; filamenta 3.5—4(-5.5) cm longa, ad apicem tubi inserta.

Capsulae 3.5-4.5 cm longae, 1.7-2 cm latae, oblongae.

Plantas perennes, surculosas, rosetas compactas, 1.5-2 m de alto, 2-2.5 m de diámetro. Hojas 50-80(-

100) por planta, 100-120 cm de largo, 9-12 cm de ancho en la parte media, lanceoladas, erectas, rígidas,

fibrosas, cóncavas, verde-amarillentas, glaucescentes hacia la base, en ocasiones con bandas transversales

glaucas; margen dentado, con una delgada banda córnea; dientes 4-6 mm de largo, 1.5-3 mm de ancho en

la base, rectos a retrorsos, grisáceos, separados por 1—3(-4.5) cm, más cercanos entre sí cerca de la base, con

uno a varios dientecillos muy pequeños entre los grandes; espina terminal de 2.5—3.5 cm de largo, decurrente

por 12-16 cm, acanalada en el dorso. Inflorescencia racemoso-paniculada de 5.5—7 m de alto, más o menos

fusiforme, pedúnculo de 3-4 m, con (6080-140 ramas laterales, de 8-13 cm las inferiores, reduciéndose

gradualmente hacia el ápice hasta 2-4 cm; brácteas del pedúnculo de 11-16 cm de largo, 2.5—5.5 cm de

ancho en la base, deltoides, cartáceas, margen entero, espina terminal 7-9 mm de largo. Flores en grupos de

10-20 por umbela, de 4.5—5(-5.5) cm de largo, verde-amarillentas; pedicelos(0.5—)1-1.5 cm de largo, elon-

gándose hasta 2 cm durante la fructificación; ovario de 2-2.5(3) cm de largo, 4-6 mm de ancho, cilíndrico,

cuello del perianto de 4—5(-6) mm, tubo del perianto de 5-6(-8) mm de largo, 7-10(212) mm de ancho

en el ápice; tépalos de 1.5-2.2 cm de largo, 3-4(-7) mm de ancho en la base, ápice cuculado; filamentos

J. Bot. Res. Inst. Texas 1(1): 79 — 84. 2007

RENNES

NN zc ERG

IAS

x NW K

S A WZ

ES AU vit

: UA WAP

va i»

SINS WAG

WAVY,

aN ea Vy

——À

K W

wh Wa

» Wick sparz a

fal, Dat o ID

h Institute of Texas 1(1)

Journal of

72e.

is u naa MOR -sancticaroli. A. Roseta. B. Hoja. B Margen ue la hoja: D. Apice de la Ms E. Bráctea del pedünculo. F. Inflorescencia. G. Flor. H.

flor. | Capsula J. Semilla. Il

García-Mendoza 7605, C. Jacques, A. Mora & A. Salazar y C.

Jacques & A. Salazar 1.

García-Mendoza et al., Una nueva especie de Agave de México 81

Fic. 2. Agave montium-sancticaroli. Planta en su hábitat.

de 3.5—4(—5.5) cm de largo, insertados en el ápice del tubo, verdes con tintes púrpura, anteras 1.3-1.5(-2)

cm de largo, 1-2 mm de ancho, amarillas; estilo 5—5.5 cm, estigma trilobado. Cápsulas de 3.5-4.5 cm de

largo, 1.7-2 cm de ancho, oblongas. Semillas de 4-6 mm de largo, 3-4 mm de ancho, con una ala hasta de

0.5 mm, aplanadas, negras.

Especimenes adicionales examinados: MÉXICO. Tamaulipas: Municipio de San Carlos, 12 km al NE de Los Magueyes, carretera

Ciudad Victoria a San Carlos, 318 m, 27 May 2004, A. García-Mendoza, C. Jacques, A. Mora & A. Salazar 7608 (ENCB, MEXU, UAT);

Km 31 de la brecha “la chepina,” que une la carretera Ciudad Victoria-Matamoros a San Carlos, 289 m, 13 Abr 2003, C. Jacques & A.

Salazar 1 (MEXU).

Agave montium-sancticaroli se desarrolla en planicies y lomeríos sobre rocas calizas y suelos arenosos o de

rendzina, entre los 150 y los 800 m, en el matorral submontano y su transición hacia el bosque de Quercus;

algunas especies asociadas son: Acacia rigidula (gavia), Celtis pallida (granjeno), Cordia boissieri (anacahuita),

Ebenopsis ebano (ébano), Havardia pallens (tenaza), Helietta parvifolia (barreta), Leucophyllum frutescens (cenizo),

Opuntia engelmannii (nopal cuijo) y Prosopis glandulosa (mezquite). Su distribución se restringe a la región

centro de Tamaulipas, en el pie de monte, entre la Llanura Costera del Golfo de México y la Sierra de San

Carlos, en los municipios de Burgos, Cruillas, Jiménez, Padilla, San Carlos y San Nicolás y posiblemente en

Méndez y San Fernando. Las poblaciones se encuentran dispersas y cada planta llega a producir de dos a

ocho hijuelos estoloníferos. El epíteto específico alude al municipio y Sierra de San Carlos, donde habita de

manera natural y preponderante.

Agave montium-sancticaroli recibe el nombre común de “jarcia” y se utiliza para la elaboración de mez-

cal (nombre genérico para las bebidas destiladas de Agave). En la Sierra de San Carlos, para la elaboración

f Texas 1(1)

fth

JOUIdI OF

jemplar en floración.

Fic 2 Annvo mnntiim-eanrtirarai

García-Mendoza et al., Una nueva especie de Agave de México 83

de esta bebida (llamada localmente “vino” o “vino-mezcal”), se emplea preferentemente Agave americana L.

subsp. protamericana Gentry (*mezcal" o *maguey cenizo") y, en menor proporción, Agave univittata Haw. y

Agave funkiana K. Koch & Bouché, ambas conocidas como “lechuguilla” o “amole,” especies que por su alto

contenido de azúcares son agregadas como saborizantes.

Por sus hojas con el margen córneo continuo, espina terminal decurrente, inflorescencia racemosa, flores

con tubos muy cortos con los tépalos proporcionalmente más largos y filamentos insertados en la orilla del

tubo, Agave montium-sancticaroli se ubica en el grupo Marginatae, delimitado por Gentry (1982). Morfológi-

camente se relaciona con Agave xglomeruliflora (Engelm.) A. Berger, de la que se diferencia por tener rosetas

más grandes, con mayor nümero de hojas, proporcionalmente mucho más largas, hojas lanceoladas, dientes

del margen más pequeños, espina apical no aquillada en el envés, inflorescencia robusta con mayor número

de ramas y flores por umbela, flores con pedicelos más largos y cápsulas más grandes no rostradas. Ambas

especies comparten el tamaño de las flores, fenómeno conservador, común entre las especies del grupo y

que se utiliza poco para separar a las especies; en cambio, Gentry (1982) considera que los caracteres de

las hojas, como forma, tamafio, color y tipo de armadura, están muy diversificados y proveen los caracteres

taxonómicos suficientes para diferenciar a los taxa.

Gentry (1982) aplicó el nombre de Agave glomeruliflora a una serie de híbridos entre A. lechuguilla (subg.

Littaea) con A. gracilipes, A. havardiana y A. neomexicana (subg. Agave), los cuales tienen inflorescencias rac-

emosas y hojas intermedias en ancho entre las especies mencionadas y A. lechuguilla (fotos de estas plantas

se aprecian en Muller, 1883, como Agave chisosensis y Breitung 1968). Por su parte, Reveal y Hodgson (2002)

consideran que el nombre de Agave xglomeruliflora *should probably be more appropiately applied only to

crosses and back-crosses between Agave lechuguilla and A. havardiana.” Agave xglomeruliflora es un taxón

propio del Desierto Chihuahuense. Crece en la Sierra del Carmen, al norte de Coahuila (México) y en las

montañas Big Bend, Texas (USA); habita laderas con pastizal y bosque de Quercus-Juniperus, en altitudes

entre los 600 y los 1600 m (Gentry 1982; Reveal y Hodgson 2002).

Agave montium-sancticaroli se desarrolla a menor altitud y se localiza a más de 400 km en línea recta

de la población más cercana de Agave xglomeruliflora, en Cuatro Ciénegas, Coahuila. De manera natural, se

observó que Agave montium-sancticaroli llega a formar híbridos con Agave funkiana y A. univitatta, especies con

las que convive en su área de distribución, tales híbridos se reconocen por presentar caracteres vegetativos

intermedios; tales como el tamanio de la roseta, tamafio, forma y dentición de las hojas e inflorescencias de

menor talla, con menor nümero de ramas florales. Algunas recolectas con estas características morfológicas

se reconocen en los ejemplares de A. García-Mendoza, C. Jacques, A. Mora & A. Salazar 7607 y 7610 (MEXU

y UAT).

Consideramos que Agave montium-sancticaroli debe ser considerada una especie bajo riesgo de extinción,

ya que por su alto contenido de azücares, superior al de Agave americana subsp. protamericana (Jacques-

Hernández 2005), es altamente apreciada por los mezcaleros y se extrae en buenas cantidades de su hábitat;

así mismo, la baja densidad de sus poblaciones silvestres, el bajo numero de hijuelos vegetativos que pro-

duce y la apertura de nuevas tierras agrícolas, están disminuyendo sus poblaciones naturales, ante lo cual,

es necesario realizar los estudios biológicos pertinentes, promover su cultivo mediante semillas e hijuelos

asexuales y su propagación in vitro por cultivo de tejidos, con el objetivo de conservar las poblaciones silves-

tres y asegurar la materia prima suficiente para las próximas décadas, que permitan continuar elaborando

el apreciado vino-mezcal *San Carlos" de Tamaulipas.

AGRADECIMIENTOS

A Fernando Chiang, Arturo Mora y Raquel Galván, por leer de manera crítica el manuscrito; el primero hizo

además la descripción latina. La ilustración es obra de Elvia Esparza. La Fundación Produce Tamaulipas y la

Secretaría de Investigación y Posgrado, del Instituto Politécnico Nacional proveyeron los fondos económicos

necesarios para el presente estudio.

84 Journal of the Botanical R h Institute of Texas 1(1)

REFERENCIAS

BREITUNG, A.J. The agaves. In: The Cactus and Succulent Journal 1968 Yearbook, eds. C. Glass y R.A. Foster. Abbey

Garden Press, Reseda. (Reprint from Cact. & Succ. J., vols. 31-36).

GENTRY, H.S. 1982. Agaves of Continental North America. University of Arizona Press, Tucson.

JACQUES-HERNANDEZ, C. 2005. Identificación y evaluación del potencial productivo de las especies naturales de

agave mezcalero. Tamaulipas PRODUCE: Tecnología para el campo 2:8-10.

Mutter, C.H. 1883. A new species of Agave from Trans-pecos Texas. Amer. Midl. Naturalist 21:763-765.

Reveal, J.L. y W.C. Hopeson. 2002. Agave. In: Flora of North America Editorial Committee, eds. Flora of North America

North of Mexico, vol. 26. Oxford University Press, New York. Pp. 442—461.

RE-EXAMINATION OF MUHLENBERGIA CAPILLARIS, M. EXPANSA,

AND M. SERICEA (POACEAE: MUHLENBERGIINAE)

Danny J. Gustafson Paul M. Peterson

Department of Biology Department of Botany

e Citade National Museum of Natural History

171 Moultrie St. Smithsonian Institution

Charleston, South Carolina 29409, U.S.A. Washington, DC 20013-7012, U.S.A.

danny.gustafson@citadel.edu peterson@si.edu

ABSTRACT

Molecular genetic data [intersimple repeats (ISSR)] and morphological data support the recognition of Muhlenbergia capillaris,

M. expansa, and M. sericea as separate species. Multi-response permutation analysis show significant differences (T = -9.03, A = 0.20, P

< 0.01) among these three species indicating that individuals within a species were more genetically similar to one another than they

were to individuals of another species. Apparently, Muhlenbergia sericea and M. capillaris are derived from a recent common ancestor

and are sister to M. expansa. A key to separate Muhlenbergia capillaris, M. expansa, and M. sericea is provided.

RESUMEN

Los datos genéticos moleculares [secuencias entre repeticiones simples (ISSR)] y los datos morfológicos apoyan el reconocimiento de

Muh

= 0.20, P< 0.01) entre estas tr pecies que indican que los individuos dentro de una misma especie son genéticamente más similares

enbergia i ane M. expansa y M. sericea como especies separadas. El análisis de permutación de respuesta múltiple (T = -9.03, A

entre si que entre los individuos ze otras especies. Al parecer, Muhlenbergia sericea y M. capillaris son derivadas de un ancestro común

reciente, y M. expanda es su especie hermana. Se proporciona una clave para separar Muhlenbergia capillaris, M. expansa y M. sericea.

Muhlenbergia Schreb. is primarily a Western Hemisphere genus of 152 species, with 69 species native to

North America, north of México (Peterson 2003; Peterson et al. 2007). Members of this genus can be an-

nual or perennial, rhizomatous to cespitose, and can occur in a variety of ecological settings. The genus is

characterized by having solitary or rarely paired spikelets that are usually one-flowered; awned, mucronate

or unawned lemmas that are three-veined; and a base chromosome number of x = 10 (Peterson et al. 1997).

In recent years there has been some debate regarding the taxonomic status of Muhlenbergia capillaris (Lam.)

Trin., M. expansa (Poir.) Trin., and M. sericea (Michx.) P.M. Peterson, three perennial cespitose species na-

tive to the southeastern and gulf coast of the United States, with the outcome potentially effecting cultural

traditions and economic aspects of the Gullah peoples who have traditionally used M. sericea [synonyms:

Muhlenbergia filipes M.A. Curtis; Muhlenbergia capillaris var. filipes (M.A. Curtis) Chapm. ex Beal] as the

primary plant material to make sweetgrass baskets (Burke et al. 2003; Rosengarten 1986).

Muhlenbergia capillaris, M. expansa and M. sericea are all members of Muhlenbergia subgenus Trichochloa

section Podosemum (Soderstrom 1967; Peterson & Herrera 2001). Muhlenbergia sericea is a perennial cespi-

tose species that occurs in marginal maritime habitat along coastal barrier islands and woodlands of the

southeastern and gulf coasts (TX to NC) and is characterized by long involute leaf blades (35-100 cm long),

long-awned lemmas (8-35 mm long) and upper glumes (2725 mm long), and lemmas with long setaceous

teeth (1-5 mm long) near the apex. Muhlenbergia capillaris has a much wider ecological and geographical

range (TX to KS to MA to FL) and superficially resembles M. sericea. However, M. capillaris has shorter-

awned lemmas (2-18 mm long), unawned or shorter awned upper glumes (1-5 mm long), and usually

lacks setaceous teeth or, if present, these are less than 1 mm long. Muhlenbergia expansa grows in wet pine

savannas and pitcher plant flatwoods inland from the coastal plain (M. sericea) habitat and lacks awned

upper glumes (sometimes these can be mucronate, i.e., with a mucro less than 1 mm long) or setaceous

teeth. The lemmas of M. expansa are unawned, mucronate or have awns 1-3 mm long.

Morden and Hatch (1989) conducted a morphological study of Muhlenbergia sericea, M. capillaris, and

J. Bot. Res. Inst. Texas 1(1): 85 — 89. 2007

86 Journal of the Botanical R h Institute of Texas 1(1)

M. expansa specimens from 25 herbaria across the southeastern United States and suggested that these taxa

are three varieties of M. capillaris rather than three distinct species. In our study, we use molecular genetic

data [intersimple sequence repeats (ISSR)] to address the hypothesis that Muhlenbergia sericea, M. capillaris,

and M. expansa are three distinct species. In addition, we offer a different interpretation of Morden and

Hatch (1989) published data in support of our hypothesis that these are three distinct species.

METHODS

Plant genomic DNA was extracted from approximately 0.1 g of silica-dried leaves from field-collected plants,

herbarium sheets, and 0.5 g fresh leaf material using E.Z.N.A.9 plant DNA miniprep kit (Omega Bio-Tek,

Doraville, Georgia, U.S.A.). Muhlenbergia sericea was collected from the eastern (n = 8, collected by K. Olandt

in Charleston County, South Carolina) and western (n = 4, collected by P. Maywald in Kennedy County,

Texas) range of the species in the fall of 2004. Muhlenbergia expansa DNA was extracted from two South

Carolina herbarium sheets (Townsend 2341 & 1123) from the Clemson University Herbarium. Muhlenber-

gia capillaris extractions consisted of two individuals from Alabama (MacDonald 12080 & Allison 7225,

University of Alabama Herbarium) and three individuals from South Carolina (collected in 2005 by DJG

from Apron Island in Charleston County, South Carolina). Two individuals were extracted of Muhlenbergia

wrightii Vasey ex J. M. Coult. and M. montana (Nutt.) Hitchc. grown from seeds that were purchased from

Western Native Seed, Coaldale, Colorado.

Muhlenbergia montana and M. wrightii (outgroup) are perennial cespitose species native to the south-

western United States and occur on rocky slopes at elevations of 1100 to 3500 m. The former species has

been included in the Muhlenbergia montana complex (Herrera 1998) and the later species is probably aligned

with other genera in the Muhlenbergiinae (Peterson et al. 2004; Peterson et al. 2007).

Twenty five intersimple sequence repeat (ISSR) primers were surveyed, with six primers selected for this

study (sequence, number of bands; (GT),-RG, 8 bands; (CA)¿-RG, 8 bands; (CA),-RY, 5 bands; (GA)¿-YC, 8

bands; (CT),-G, 3 bands; (CA)¿-RG, 4 bands). ISSR polymerase chain reaction (PCR) protocol followed that

of Wolfe et al. (1998); 94? C for Imin 50sec, 40 cycles of 94? C for 40 sec, 43? C for 45 sec, and 72? C for

Imin 50 sec, followed by a final extension at 72? C for 5 min. PCR profiles were visualized in 1.596 agarose

gels and stained with ethidium bormide. Images were captured using a digital camera (Olympus C-4000

Zoom, Melville, NY), converted to a negative image, and fragment size was estimated based on a DNA marker

(Benchtop pGEM, #G7521, Promega, Madison, WI). Fragment sizes were used to assign loci for each primer

and bands were scored as diallelic for each locus (1=band present, 02 band absent). Individual ISSR profiles

were used to calculate a priori species assignment using multi-response permutation procedure (MRPP)

(PC-Ord, ver. 4.2, MjM Software Design, Gleneden Beach, Oregon, U.S.A.). Nei's genetic distance (1972)

was calculated among taxa based on band frequency data and Neighbor-Joining cluster analysis (Saitou and

Nei, 1987) using NTSYSpc 2.2d (NTSYSpc Numerical Taxonomy and Multivariate Analysis System, Applied

Biostatistics Inc., New York, NY).

RESULTS AND DISCUSSION

ISSR analysis clearly supports Muhlenbergia sericea as a separate species that shares a common ancestor with

M. capillaris and M. expansa, which is in agreement with Peterson’s (2003) recent treatment of Muhlenbergia.

MRPP analysis indicated significant differences (T = -9.03, A = 0.20, P < 0.01) among the a priori species

designation, meaning that individuals within a species were more genetically similar to one another than

they were to members of another species. If Muhlenbergia sericea and M. expansa were varieties of M. capil-

laris, then we would have expected to find members of all three a priori species forming one genetically

similar grouping.

Phylogenetic relationships among these five Muhlenbergia species revealed predictable associations, with

the more eastern Muhlenbergia species (M. expansa, M. capillaris, M. sericea) forming a monophyletic group

(Fig. 1). Muhlenbergia wrightii (outgroup) and M. montana are native to the mountain and southwestern regions

Gustafson and Peterson, Muhlenbergia capillaris, expansa, and sericea 87

M. sericea

M. capillaris

M. expansa

M. montana

: i I I l

0.00 0.02 0.04 0.06 0.08

Coefficient

Fic. 1. Neighbor-Joining cluster analysis based on Nei’ tic dist

cies (M. sericea, M. capillaris, M.

species.

g five Muhlenbergia species. The three southeastern Muhlenbergia spe-

to one another than they tain M. wrightii

of the United States and were clearly different from the three eastern species. Much of the taxonomic confu-

sion surrounding Muhlenbergia sericea, M. capillaris, and M. expansa extends from the potential geographic

and ecological overlap of these taxa and the limited number of diagnostic characters. Based on our ISSR

molecular markers, Muhlenbergia sericea and M. capillaris are more similar to one another than they are to

M. expansa (Fig. 1).

Morden and Hatch (1989) conducted a taxonomic study of Muhlenbergia sericea, M. capillaris, and M.

expansa based on morphological characters and recommended that these three taxa should be a single

species consisting of three varieties. We respectfully disagree with their conclusions and offer a different

interpretation of their results. The taxonomic, geographic, and morphological sampling was sufficient and

appropriate for the stated objectives of their study; however, they failed to use summary statistics to assess

differences among taxa. An analysis of variance or non-parametric analysis should have yielded statistically

significant differences among the three taxa for morphological characters which have been used histori-

cally to separate these species. Plotting the means + 1 standard error (Fig. 2) for blade, upper glume awn,

lemma awn, and setaceous teeth lengths are a good indication that significant differences would have been

found if the authors tested for difference among taxa. Morden and Hatch's PCA analysis does not present

key multivariate statistics, such as eigenvalues for each axis or parallel analysis indicating which axes are

appropriate for interpretation. In addition, a graph of individuals on the first two PCA axes clearly show

Muhlenbergia sericea and M. expansa as separate clusters with M. capillaris intermediate (Fig. 1; Morden and

Hatch 1989). Discriminate analysis statistics were also not presented in their manuscript; however, we would

suggest that a misclassification rate of 3.796 (13 out of 350) is not strong support for realigning these three

taxa as varieties of Muhlenbergia capillaris. In a draft (9 Aug 2006) of the Flora of the Carolinas, Georgia,

88 Journal of the Botani Institute of Texas 1(1)

E Lower Glume

Upper Glume E +

E 34 - 5 14 4

5 S

E (11-80) (10-47) (14-75) m

2 E (0-17) (0-4) (8-33)

2 <

S 277 z dom

I 5

20 y T T 0 T T T

M. capillaris M. expansa M. sericea M. capillaris M. expansa M. sericea

15 2.1

(0-10.3; 1-25) 2

z ; Eis

E z

Ss 107 Bn 1.4 4

EJ 5

oO -

H S

E 3

< E |

o 5] 2 074 (0-2) (0-0.1) (0-4.5)

£ 3

2 Q

o) (0-3.3; 0-5) (0; 0-0.6) 8

0 aA, 0.0 E ' T

M. capillaris M. expansa M. sericea M. capillaris M. expansa M. sericea

Fic. 2. hese data represent the mean 45b. itl f key morphological charact | to distinguish Muhlenbergia capillari

M. expansa, and M. sericea. C table 1 in Morden and Hatch 1989.

and Virginia, A.S. Weakley (in prep.) also comments in regards to Morden and Hatch (1989) and states that

these three taxa are undoubtedly biological species.

Based on our molecular genetic data and a more rigorous statistical interpretation of Morden and Hatch

(1989) morphological study, we conclude that Muhlenbergia sericea and M. capillaris are two closely related

species that have limited ecological and morphological overlap, but these taxa should remain as distinct

species. Morphological and ISSR analysis indicate that Muhlenbergia sericea and M. capillaris are more similar

to one another than either is to M. expansa. While proper identification of Muhlenbergia sericea, M. capillaris,

and M. expansa in the field is sometimes difficult, we recommend using a combination of ecological setting,

blade length and mature floret characters (glume awn, lemma awn, and setaceous teeth length) to distinguish

among these three closely related species. We provide a key to separate these three species below.

A KEY TO MUHLENBERGIA CAPILLARIS, M. EXPANSA, AND M. SERICEA IN NORTH AMERICA

1. Body of the glumes more than 1/2 as long as the lemmas; lemmas unawned, mucronate, or with awns only

1-3 mm long; upper glumes never awned but sometimes mucronate Muhlenbergia expansa

. Body of the glumes less than 1/2 as long as the lemmas; lemmas usually awned 2-35 mm long; upper glumes

often awned, the awns 1-25 mm long.

2. Upper glumes unawned or with awns to 5 mm long; lemmas without setaceous teeth or with teeth no

more than 1 mm long; lemma awns 2-13(-18) mm long Muhlenbergia capillaris

2. Upper glumes awned, the awns 2-25 mm long; lemmas with setaceous teeth 1-5 mm long; lemma awns

8-35 mm long Muhlenbergia sericea

Gustafson and Peterson, ia capillaris, expansa, and sericea 89

ACKNOWLEDGMENTS

We thank Patricia Gomez Bustamante for help preparing the Spanish resumen. We would also like to

thank the University of Alabama, University of Florida, and Clemson University Herbaria for providing

pressed material and Paula Maywald for material from Texas. We acknowledge the thoughtful comments

and suggestions by Drs. Mark Basinger and Barney Lipscomb which improved this manuscript. We also

acknowledge the contributions made by the Citadel Plant Ecology Laboratory undergraduate researchers

(Philip Strole, Esteban Sierra, Chris Morrow, and Garret Ryan). This research was funded by grants from

the South Carolina Sea Grant Consortium and The Citadel Foundation to DJG.

REFERENCES

Burke, M. K., A. C. HALFAcRE, and Z. Hart. 2003. Decline of sweetgrass spurs restoration of coastal prairie habitat

(South Carolina). Ecol. Restoration 21:50—51.

HERRERA ARRIETA, Y. 1998. A revision of the Muhlenbergia montana (Nutt.) Hitchc. complex (Poaceae: Chloridoideae).

Brittonia 50:23-50.

Moroen, C.W. and S.L. Hatch. 1989. An analysis of morphological variation in Muhlenbergia capillaris (Poaceae)

and its allies in the southeastern United States. Sida 13:303-314.

Ne, M. 1972. Genetic distance between populations. Amer. Naturalist 106:283-292.

Peterson, PM. 2003. Muhlenbergia Schreb. In: M.E. Barkworth, K.M. Kapels, S. Long. and M.B. Piep, eds. Magno-

liophyta: Commelinidae (in part): Poaceae, part 2. Flora of North America North of Mexico, volume 25. Oxford

University Press, New York, NY. Pp. 145-201.

PETERSON, P.M., J.T. Cotumpus and S.J. PENNINGTON. 2007. Classification and biogeography of New World grasses:

Chloridoideae. Aliso: 23: In Press.

PETERSON, PM., J.T. CotuMaus, N.F. Reruuo Ropriguez, R. Cerros TLATILPA, and M.S. Kinney. 2004. A phylogeny of the

Muhlenbergiinae (Poaceae: Chloridoideae: Cynodonteae) based on ITS and trnL-F sequences. Botany2004

abstract: http://www.2004.botanyconference.org/engine/search/index.php?func=detail&aid=38

PETERSON, P.M. and J. HERRERA Arrieta. 2001. A leaf blade anatomical survey of Muhlenbergia (Poaceae: Muhlenber-

giinae). Sida 19:469-506.

PETERSON P.M, R.D. Wesster, and J. VALDEs Reyna. 1997. Genera of New World Eragrostideae (Poaceae: Chloridoideae).

Smithsonian Contr. Bot. 87:1—50.

PorcHer, R.D. and D.A. Rayner. 2001. A guide to the wildflowers of South Carolina. University of South Carolina

Press, Columbia.

RADFORD, A.E., H.E. Aures, and C.R. Bau. 1968. Manual of the vascular flora of the Carolinas. University of North

Carolina Press, Chapel Hill.

ROSENGARTEN, D. 1986. Row upon row: sea grass baskets of the South Carolina Lowcountry. McKissick Museum,

University of South Carolina, Columbia.

Sarrou, N. and M. Nel. 1987. The neighbor-joining method: a new method for reconstructing phylogenetic trees.

Molec. Biol. Evol. 4:406-425.

SODERSTROM, T.R. 1967. Taxonomic study of subgenus Podosemum and section Epicampes of Muhlenbergia (Gra-

mineae). Contr. U.S. Natl. Herb. 34:75-189.

Worre, A.D., Q-Y, XIANG, and S.R. KEPHART. 1998. Assessing hybridization in natural populations of Penstemon (Scrophu-

lariaceae) using hypervariable intersimple sequence repeats (ISSR) bands. Molec. Ecol. 7:1107-1125.

90 Journal of the Botanical R h Institute of Texas 1(1)

BOOK REVIEWS

PATRICK Gass, edited and annotated by CAROL Lynn MACGREGOR. 1997. The Journals of Patrick Gass: Member

of the Lewis and Clark Expedition. (ISBN 0-87842351-6). Mountain Press Publishing Company,

Box 2399, Missoula, MT 59806, U.S.A. $20.00. (Orders: www.mountain-press.com, info@mtnpress.

com, 406-728-1900, 406-728-1635 fax). $20.00, 445 pp., illustrated, 6" x 9".

Captain Lewis asked the men who could write to keep journals. Patrick Gass was one of the seven known journal keepers whose journals

have survived. Patrick had only 19 days of formal education and by his own admission *never learned to read, write, and cipher till

he had come of age." His journal provides us with more details about some activities of the Expedition than do the other journals and

is more readable. Gass' journal is full of descriptions of the surrounding country and the wildlife, including a list of animals killed for

food by the expedition. He was a keen observer, and since he was a epe he promise details not included in other journals on the

construction of earth lodges and canoes of the native people. Gass g the building of Forts

Mandan and Clatsop; his records of those forts are particularly detailed and useful.

1 1

His journal was published in 1807 and proved quite popular: it went through six editions in six years. It was pub-

lished just six months after in Corps returned to St. Louis and seven years before Lewis's and Clark's were published. The inclusion

of Gass' previously unknown account book from later in his life lend new insight into Gass's work and his life. He lived until 1870 and

died when he was ninety-nine. The previous year the Pacific railroad had been completed and Patrick Gass, one of the first Americans

to cross the continent and the last survivor of the Corps of Discovery, had lived to see it.

Las TAE lie 1

The University of Nebraska Press edition of the Journals of Lewis and Clark, vol. 10, | Sergeant

Patrick Gass. The journal is to be found only in printed form; the original has been lost since its m SPICE This edition is a valu-

able supplement to it and should be purchased by libraries holding the original.—Gary Jennings, Library, Botanical Research Institute of

Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A.

RONALD D. QUINN and STERLING C. KEELEY. 2006. Introduction to California Chaparral. (ISBN 0-520-24566-

0, pbk.).The University of California Press, Berkeley, CA 94704, U.S.A. (Orders: California Princeton

Fulfillment Services, 1445 Lower Ferry Road, Ewing, NJ 08618, U.S.A., www.ucpress.edu, 609-883-

1759, 609-883-7413 fax). $19.95, 322 pp., illustrated, 4⁄2" x 714".

The California chaparral is well known to us. We have seen it hundreds, if not thousands, of times as a backdrop in movie and television

productions from early westerns to MASH to the present day. But it is not just a feature of the Hollywood Hills. It is an essential part

of the entire California landscape from the Mexican border to the Oregon border. The chaparral is a wonderfully resilient ecological

community which has adapted to recurring fires and droughts. The book’s authors, both chaparral researchers and scientists, were

interested in writing a book for a wider audience. Hee were brought together by the Press and have produced a concise, engaging, and

beautifully illustrated book. They d ib system which contains awesome and plants and animals: Fire Beetles that

mate only on burning branches, lizards that shoot blood from their eyes when threatened, Kangaroo Rats that never drink water, and

seeds that germinate only after a fire, even if that means waiting in the soil for a 100 years or more.

Part of the University of California Natural History Guide, the book follows the typical pattern for their guides. The contents

include: 1. The California chaparral. Discusses where it is found and what is the composition of vegetational communities. 2. Mediter-

ranean climate. How rainfall is affected by the unpredictable wind and the influence of temperature and microclimates. 3. Fire. The

Veces

cycle of fire and f historical fire patterns. Natural responses to the effect of fire by plants and animals. 4. Plants. Discusses

the common evergreen shrubby vegetation as well as other plant families and introduced weeds. 5. Animals. Enumerates the mammals,

rodents, birds, and insects that inhabit the chaparral. 6.Living with the chaparral. Addresses prescribed fire, threats to the chaparral,

and options for wise (human) growth. A glossary and supplemental readings and references complete the book.

Chaparral will introduce general readers to the plants and animals associated with chaparral and will be a review for biologists

and land managers its natural history, ecology, and management challenges. It is useful both as a field guide and as an introductory

overview of the ecology of chaparral. It also provides a better understanding of how we might live in harmony, safety, and appreciation

of this uni logical community.— Gary Jennings, Library, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-

4060, U.S. A.

J. Bot. Res. Inst. Texas 1(1): 90. 2007

EOEPIGYNIA BURMENSIS GEN. AND SP. NOV., AN EARLY CRETACEOUS EUDICOT

FLOWER (ANGIOSPERMAE) IN BURMESE AMBER

George Poinar Jr. Kenton L. Chambers

Department of Zoology Department of Botany and Plant Pathology

Oregon State University Oregon State University

Corvallis, Oregon 97331, U.S.A. Corvallis, Oregon 97331, U.S.A.

Ron Buckley

9635 Sumpter Road

Florence, Kentucky 41042-8355, U.S.A.

ABSTRACT

Eoépigynia burmensis gen. & sp. nov. is described from Early Cretaceous Burmese amber. The genus is characterized by small, per-

fect, actinomorphic flowers possessing a perianth with a single series of basally connate sepals, four distinct equal petals, four included

stamens alternate with the petals, an inferior ovary, a single style with a bilobed stigma, and triaperturate pollen. Flowers with similar

morphology occur in the family Cornaceae.

Key Wonps: Burma, amber, eudicot flower, Early Cretaceous, Cornaceae

RESUMEN

Eoépigynia burmensis gen. & sp. nov. se describe del ámbar birmano del Cretácico temprano. El gé 5 iza por tener flores

pequeñas, perfectas, actinomórficas que tienen un perianto con una serie sencilla de sépalos connados en la base, cuatro pétalos inde-

pendientes iguales, cuatro estambres inclusos alternando con los pétalos, un ovario infero, un estilo simple con estigma bilobulado, y

polen triaperturado. Existen flores con una morfología semejante en la familia Cornaceae.

INTRODUCTION

Burmese amber has an interesting past dating back to AD 100 when an amber trade route was established

with China. The first Europeans visited the mines in 1836 but it was not until 1896 that the amber was

noted to contain insect remains. From AD 100 until 1936, the Burmese amber mines supplied amber to

various parts of the world (Chhibber 1934). In 2001, a new amber mine was excavated in the Hukawng

Valley, southwest of Maingkhwan in the state of Kachin (26°20'N, 96°36'E) (Poinar et al. 2005). This new

amber site, known as the Noije Bum 2001 Summit Site, was dated to the Upper Albian (100 to 105 mybp)

of the Early Cretaceous (Cruickshank & Ko 2003).

The Early Cretaceous age of amber from the Noije Bum 2001 Summit Site as determined by Cruickshank

and Ko (2003) is supported by primitive insects from this deposit. For example, a bee was discovered still

possessing characters of sphecid wasps, the group considered ancestral to bees (Poinar & Danforth 2006).

An elcanid grasshopper was also found at this site, representing a group (Elcanoidea) that first appeared in

the Early Permian and continued only to the mid-Cretaceous (Poinar et al. 2007). Thus, both paleontologi-

cal data and inclusions in the amber support an Early Cretaceous age for deposits from the Noije Bum 2001

Summit Site.

Nuclear magnetic resonance (NMR) spectra of amber samples taken from that locality indicate an ar-

aucarian (possibly Agathis) source of the amber (Lambert & Wu, unpublished data 2002). While insects are

dominant, the deposits have revealed some very interesting plant fossils, including a unisexual flower with

affinities to the family Monimiaceae (Poinar & Chambers 2005) and two early bambusoid grasses (Poinar

2004). In the present paper, we describe a bisexual flower, provisionally assigned to the family Cornaceae,

from the Noije Bum 2001 Summit Site.

J. Bot. Res. Inst. Texas 1(1): 91 — 96. 2007

£+sha D o ID L

92 Journal of t titute of Texas 1(1)

MATERIALS AND METHODS

The flower is complete and well-preserved. One of the petals shows some evidence of insect damage, and

fungal hyphae are associated with one of the anthers. The flower was in anthesis at entombment, and pollen

grains occur on and adjacent to the anthers as well as on the stigma. The piece of amber containing the flower

is square, measuring 5 mm in length by 5 mm in width by 1.5 mm deep. Examination and photographs were

made with a Nikon stereoscopic microscope SMZ-10 R at 80 x and a Nikon Optiphot microscope at 800x.

DESCRIPTION

Eoépigynia Poinar, Chambers & Buckley, gen. nov. Tee serais: Eoëpigynia burmensis Poinar, Chambers € Buckley, sp. nov.

Diagnosis.—Flowers small, bisexual, regular, epigynous; perianth tetramerous; calyx comprising a short

crown (gamosepalous) at summit of ovary, sepal lobes incised, number uncertain; petals 4, separate, valvate,

regular; stamens 4, free to base, in a single whorl, alternating with the petals, filaments linear, anthers in-

trose, dorsifixed; pollen shed singly, pollen grains triaperturate (possibly tricolporate with thickened exine

adjacent to colpi); gynoecium syncarpous, ovary inferior, style 1, stigma bilobed, pericarp wall thick-textured

at anthesis, fruit type unknown; presence of floral disc not determinable.

Eoépigynia burmensis Poinar, Chambers & Buckley, sp. nov. (Figs. 1-2). Tyee: MYANMAR (BURMA): Kacuin: northern

Myanmar, amber mine in the Hukawng Valley, SW of Maingkhwan, Q620N, 96?36'E), Aug 2005, Buckley s.n. (HOLOTYPE: perfect

flower (accession 4 ab 214) deposited in the collection of Ron Buckley, Florence, Kentucky 41042-8355, U.S.A.).

Description.—Flower bisexual, glabrous, length 1.5 mm; no free hypanthium evident; calyx lobes with

incised margins, greatest length of calyx 0.34 mm; petals lanceolate-ovate, margins abaxially recurved, up

to 0.95 mm long, 0.34 mm wide; stamens with ovoid anthers up to 0.15 mm long, filaments 0.54-0.61 mm

long; stigma at level of anthers, estimated length of style 0.55 mm; length of ovary 0.54 mm; width of ovary

0.39 mm; diameter of pollen grains, 12-14 um.

Etymology.—Genus name from the Greek “eos” dawn, “epi-“ upon, and “gyne” female, from the age and

the relation of the floral perianth to the ovary. Species named for the country of origin of the fossil.

DISCUSSION:

Certain structures that would be helpful in the placement of Eoépigynia, for example the presence of a floral

disc, could not be observed due to the fossil’s orientation in the amber. However, it was possible to view the

flower from both sides to verify the characteristics described here. Based on its floral features, Eoépigynia

(Figs. 1, 2) represents a core eudicot that can provisionally be assigned to the family Cornaceae sensu lato,

in the basal asterid order Cornales. The phylogenetic position of Cornales as sister to all the remaining

asterids (perhaps excluding Ericales) is well confirmed by molecular studies (Stevens 2001 onwards; Hilu

et al. 2003; Judd & Olmstead 2004; Bremer et al. 2004). The perfect flower, inferior ovary, compound style,

4-merous perianth and androecium, and possibly tricolporate pollen of our fossil are most similar to the

modern genus Cornus (see illustrations in Wangerin 1910; Judd et al. 1999). The pollen of the fossil may be

of particular importance in this placement. In an equatorial optical section (Fig. 1D) three distinct paired

thickenings are seen in the exine marking what we assume are the three colpi (we could not focus our in-

strument clearly on a pore at this spot). This pattern is strongly reminiscent of SEM equatorial transections

of Cornus pollen illustrated by Ferguson (1977), e.g. his figure 2c of thickened endexine in this area in C.

volkensii, as well as his figures 4d of C. disciflora and 6c of Curtisia dentata. Because of the pollen orientation

in the amber, we were unable to observe a pore face-on, where the characteristic H-shaped endoaperture

thinning pattern of Cornaceae/Nyssaceae might be seen (Erdtman 1966; Ferguson 1977). The pollen grains

of Eoépigynia are smaller than in most of the types defined by Ferguson (1977) but are within the range of

his Curtisia-type, described as 12-20 x 12-17 um (p. 6). One might speculate that small pollen are related

adaptively to the small size of pollinating insects of that period, for example the 2 mm-long bee recently

described from Burmese amber by Poinar and Danforth (2006).

Poinar et al., Eoépigynia burmensis, an Early Cretaceous eudicot flower 93

Fic. 1. Foépigynia E isin B ber. A. One side of fl Scale bar = 0.34 mm. B. Opposite side of flower. Scale bar = 0.34 mm. C. Stamen:

f | hypł Scale bar 2 0.17 mm. D

J cal : I ITI L

anie vuVvVeieu vien e : y My y JI J

Scale bar = 18 pm.

94 Journal of the Botanical R h Institute of Texas 1(1)

er e . pL DEREN L

Fic. 2. Flower

Anther on right as described in Fig. 1C. Scale bar = 0.34 mm.

The generic makeup of Cornales, as well as family Cornaceae, has been ined in recent molecular

phylogenetic studies (Xiang et al. 1998; Fan & Xiang 2003; Hilu et al. 2003; Judd & Olmstead 2004). It is

proposed that Cornaceae be limited to two genera, Cornus and Alangium, and that other genera formerly

assigned here be segregated to families Nyssaceae, Mastixiaceae, and Grubbiaceae. At the ordinal level, the

once widely separated families Loasaceae and Hydrangeaceae are to be included in Cornales. If Eoépigynia is

placed in the larger context of this basal asterid clade, it shows that a simplified epigynous flower, with a single

style, 4-merous perianth parts, and four stamens, arose early in the differentiation of this evolutionary line.

As pointed out by Gustafsson and Albert (1999), epigyny is not a recent phenomenon. Examples were

found by Friis et al. (1994) in Early Cretaceous sediments in Portugal, originally dated as Valanginian or

Hauterivian but now reassigned to the early Albian (Heimhofer et al. 2005). A probable relationship to

Chloranthaceae has been established (Eklund et al. 2004). Phylogenetic studies have also shown that the

evolution of ovary position has been dynamic, with at least 64 changes from hypogyny to epigyny but only

24 changes in the opposite direction (Gustafsson & Albert 1999; Soltis et al. 2003). In neither of these papers

are Cornales specifically discussed, however. Modern Cornales are well represented in the Southeast Asian

flora, with Cornus itself having a circumpolar Northern Hemisphere distribution (Wangerin 1910; Xiang

et al. 2005). Reference fossils (fruit stones) attributed to the Cornelian-Cherry line of Cornus in the latter

paper are taken from the careful review by Eyde (1988) and are Eocene or younger in age.

In studies using molecular phylogenetic dating methods with known fossil reference points (Bremer et

al. 2004; Anderson et al. 2005), the Early Coniacian (88 mybp) cornalean genus Hironoia (Takahashi et al.

2002) has been used. Based on these papers, the age of the stem group asterids may be ca. 128 mybp, the

Cornales and Ericales diverging soon afterwards (Stevens 2001 onwards). Anderson et al. (2005) place the

separation of Cornales from remaining asterids at ca. 109 mybp. The reference fossil Hironoia consists of

Poinar et al., Eoépigynia burmensis, an Early Cretaceous eudicot flower 95

three-dimensionally preserved drupes, with characters of the endocarp wall and dehiscence valves synapo-

morphic with the genera Nyssa and Mastixia. However, it could not be placed with certainty in one or the

other genus. Another known cornalean fossil is Tylerianthus from the Upper Cretaceous Turonian Period,

ca. 90 mybp (Gandolfo et al. 1998; Crepet et al. 2004), with affinities to the Hydrangeaceae. The putative

cornacean fossil Eoépigynia would extend the age of the clade, if used in similar dating studies. It would be

well to note, however, that its generalized floral morphology would allow possible placement of this fossil in

other epigynous clades of core eudicots as well, including Saxifragales, Myrtales and Asterales. Its similarity

to Cornaceae, although highly suggestive, is not fully diagnostic of the proposed relationship. We know of

no other fossil flower from the Cretaceous with the floral syndrome of this specimen.

ACKNOWLEDGMENTS

The authors thank Elmar Robbrecht for comments comparing the fossil to members of the family Rubiaceae

and Roberta Poinar for reading earlier drafts. The helpful comments by reviewers Steven R. Manchester and

James A. Doyle are much appreciated.

REFERENCES

ANDERSON, C.L., K. Bremer, and E.M. Fris. 2005. Dating phylogenetically basal eudicots using rbcL sequences and

multiple fossil reference points. Amer. J. Bot. 92:1737-1748.

Bremer, K., E.M. Fris, and B. Bremer. 2004. Molecular phylogenetic dating of asterid flowering plants shows Early

Cretaceous diversification. Syst. Biol. 53:496-505.

CHHIBBER, H.L. 1934. The mineral resources of Burma. Macmillan € Company, London.

Crepet, W.L., K.C. Nixon, and M.A. GANDOLFO. 2004. Fossil evidence and phylogeny: the age of major angiosperm clades

based on mesofossil and macrofossil evidence from Cretaceous deposits. Amer. J. Bot. 91:1666-1682.

CRUICKSHANK, R.D. and K. Ko. 2003. Geology of an amber locality in the Hukawng Valley, northern Myanmar. J.

Asian Earth Sci. 21:441—455.

EKLUND, H., J.A. Dovie, and PS. Herendeen. 2004. Morphological phylogenetic analysis of living and fossil Chloran-

thaceae. Int. J. Plant Sci. 165:107-151.

ERDTMAN, G. 1966. Pollen morphology and plant taxonomy. Angiosperms. Hafner Publishing Company, New

York.

Eype, RH. 1988. Comprehending Cornus: Puzzles and progress in the systematics of dogwoods. Bot. Rev.

54:233-351.

FAN, C. and Q-Y. XiANG, 2003. Phylogenetic analyses of Cornales based on 265 rRNA and combined 26S rDNA-

matK-rbcL sequence data. Amer. J. Bot. 90:1357-1372.

FERGUSON, I.K. 1977. Cornaceae. World pollen and spore flora 6:1-34. Stockholm.

Fais, E.M., K.R. PEDERSEN, and PR. Crane. 1994. Angiosperm floral structures from the Early Cretaceous of Portugal.

Pl. Syst. Evol. [Suppl.] 8:31-49.

GANDOLFO, M.A., K.C. Nixon, and W.L. Cneeer. 1998. Tylerianthus crossmanensis gen. et sp. nov (aff. Hydrangeaceae)

from the Upper Cretaceous of New Jersey. Amer. J. Bot. 85:376-386.

Gustarsson, M.H.G. and V.A. ALBERT. 1999. Inferior ovaries and angiosperm diversification. In: PM. Hollingsworth,

R.M. Bateman and R.J. Gornall, eds. Molecular systematics and plant evolution. Taylor & Francis, London. Pp.

403-431.

HEIMHOFER, U., PA. HocHuu, S. BurLa, J.M.L. Dinis, and H. WelsserT. 2005. Timing of Early Cretaceous angiosperm diver-

sification and possible links to major paleoenvironmental change. Geology 33:141-144.

Hitu, K.W, T. BorscH, A. MULLER, D.E. Souris, PS. Souris, V. SAVOLAINEN, M.W. Chase, M.P. PoweLL, L.A. Auce, R. Evans, H. SAUQUET,

C. NeinHuis, T.A.B. Stotta, J.G. RoHwer, C.S. CampeeLL, and L.W. Chatrou. 2003. Angiosperm phylogeny based on

matK sequence information. Amer. J. Bot. 90:1758-1776.

Jupp, W.S., C.S. CaMPBELL, E.A. KeLLOGG, and PF. Stevens. 1999. Plant systematics: a phylogenetic approach. Sinauer

Associates, Sunderland, MA.

96 Journal of the Botanical R h Institute of Texas 1(1)

Jupp, W.S. and R.G. OLmsteAD. 2004. A survey of tricolpate (eudicot) phylogenetic relationships. Amer. J. Bot.

91:1627-1644.

POINAR, JR, G.O. 2004. Programinis burmitis gen. et sp. nov., and P. laminatus sp. nov., Early Cretaceous grass-like

monocots in Burmese amber. Austral. Syst. Bot. 17:497—504.

PoINAR, Jr., G.O., R. BuckLey, and A. Brown. 2005. The secrets of Burmese amber. Mid Amer. Paleont. Soc. 29:20-29.

Ponar JR., G.O. and K.L. CHampers. 2005. Palaeoanthella huangii gen. and sp. nov., an Early Cretaceous flower (An-

giospermae) in Burmese amber. Sida 21:2087-2092.

Ponar, JR., G.O. and B.N. DANForTH. 2006. A fossil bee from Early Cretaceous amber. Science 314:614.

POINAR, Jr., G.O., A.V. Gorochov, and R. Bucktey. 2007. Longioculus burmensis, n. gen., n. sp. (Orthoptera: Elcanidae)

in Burmese amber. Proc. Entomol. Soc. Wash. 109:649-655.

Souris, D.E., M. FisHBEN, and R.K. Kuzorr. 2003. Reevaluating the evolution of epigyny: data from phylogenetics and

floral ontogeny. Int. J. Plant Sci. 164(55):5251-5264.

Stevens, PF. (2001 onwards). Angiosperm Phylogeny Website. Version 7, May 2006 [and more or less continuously

updated since]. http://www.mobot.org/MOBOT/research/APweb

TAKAHASHI, M, PR. Crane, and S.R. MANCHESTER. 2003. Hironoia fusiformis gen. et sp. nov.: a cornalean fruit from the

Kamikitaba locality (Upper Cretaceous, Lower Coniacian) in northeastern Japan. J. Plant Res. 115:463-473.

WANGERIN, W. 1910. Cornaceae. In: A. Engler, Das Pflanzenreich. IV. 229:1-110.

Watson, L. and M.J. DatLwitz.1992 onwards. The families of flowering plants: descriptions, illustrations, identifica-

tion, and information retrieval. Version: 29th July 2006. http://delta-intkey.com

XIANG, Q-Y., D.E. Souris, and PS. Souris. 1998. Phylogenetic relationships of Cornaceae and close relatives inferred

from matK and rbcL sequences. Amer. J. Bot. 85:285-297.

XIANG, O-Y., S.R. MANCHESTER, D.T. THOMAS, W. ZHANG, and C. Fan. 2005. Phylogeny, biogeography, and molecular dating

of Cornelian Cherries (Cornus, Cornaceae): tracking Tertiary plant migration. Evolution 59:1685-1700.

RECTOIMPIFICATION OF CAULTHERIA PYROLCIFOLIA

AND G. PYROLOIDES (ERICACEAE)

Peter W. Fritsch and Debra K. Trock

Department of Botany, California Academy of Sciences

875 Howard Street

San Francisco, California 94103, U.S.A.

ptritsch@calacademy.org; dtrock@calacademy.org

ABSTRACT

Different names have been used for two eastern Asian and Alaskan species of Gaultheria in various taxonomic treatments. To resolve

this problem, a lectotype for the name G. pyroloides is selected. The name G. miqueliana, employed in ic works for the spe-

cies from Japan, Russia, and Alaska since its publication in 1918, must be relegated to synonymy under G. pyroloides. A lectotype is also

selected for the Himalayan species G. pyrolifolia. Several morphological characters are identified that differentiate the two species.

RESUMEN

Se han usado diferentes nombres para dos especies, del este de Asia Alaska, de Gaultheria en varios tratamientos taxonómicos. Para

resolver este problema, se ha seleccionado un lectotipo para el nombre G. pyroloides. El nombre G. miqueliana, empleado en la mayoría

de los trabajos taxonómicos para la especie de Japón, Rusia, y Alaska desde su publicación in 1918, debe ser relegado a la sinonimia de

G. pyroloides. Se ha seleccionado también un lectotipo para la especie del Himalaya G. pyrolifolia. Se han identificado varios caracteres

morfológicos que diferencian las dos especies.

During preparation of manuscripts on the taxonomy of Gaultheria for a treatment of the genus in Gaoligong

Shan, western Yunnan Province, China (Fritsch) and for Flora of North America (Trock), the authors de-

tected inconsistency in the application of the names G. miqueliana Takeda, G. pyrolifolia J.D. Hooker ex C.B.

Clarke, and G. pyroloides Miquel to two species from eastern Asia and the Aleutian Islands. Here we clarify

the nomenclature of these species by designating lectotypes for G. pyrolifolia and G. pyroloides.

Nomenclatural Background of Gaultheria pyroloides and G. pyrolifolia

Gaultheria pyroloides was described on the basis of several gatherings from Sikkim by John Dalton Hooker

(s.n.), two of which are now on the same sheet at K but were possibly separate when Miquel saw the material

(Fig. D, and one from Japan (collector not determined; Fig. 2). In the protologue, Miquel (1863) indicated

that he considered these specimens to represent one and the same species, as evidenced by the following

statement: “Specimina nostra capsulifera ab indices, quae in regione alpina 12-13,000 alt. Himalayae Sik-

kimensis detecta et e Museo Kewensi mihi concessa, floribusque instructa sunt, nullo modo differunt,”

which in English translates roughly to *Our [Japanese] capsuliferous specimen in no way differs from that

from the index, which, found in the alpine region of the Sikkim Himalaya between 12,000 and 13,000 feet

elevation and given to me from the Museum at Kew, is provided with flowers." The description seems to

be based solely or in large part on the Japanese specimen, because only the fruiting condition is described

and flower characters are not included. This suggests that Miquel did not have direct access to the Sikkim

specimens at the time of description and was relying at least in part on memory in his decision to include

the Sikkim and Japanese plants under the same species. Perhaps a memory lapse, then, explains why Miquel

ascribed the name G. pyroloides to *Hook. fil. et Th. herb. Ind. or." (i.e., J.D. Hooker and T. Thomson) when *G.

pyrolaefolia H f & T" in Hooker’s handwriting [as confirmed with the examples in Burdet (1975)] is written

on the Hooker and Thomson sheet, and not *G. pyroloides.”

Subsequently, C.B. Clarke [in Hooker (1882)] described Gaultheria pyrolifolia (as G. “pyrolaefolia”) based

on several gatherings of Hooker (ascribing the name to “Hook. f. ms.”), including those seen by Miquel. Clarke

appears to have been unaware that G. pyroloides was already published or at least that it was based on a

J. Bot. Res. Inst. Texas 1(1): 97 — 102. 2007

Journal of the Botanical R h Institute of Texas 1(1)

| C GARDEN:

+ ROYAL BOTANIC GARDENS KEW

E i

00442406

V ns 58 m

oen EAD, s P|

ATL

Jl tobuke ue

Lad cito. E

A 22} (2

PA. | I Baro

enne p

FLORA BRIT. IND. Vol. III. 47 Y

Named by Mr. C. D. CLARKE.

Fic. 1. Image of the lectotype (K) of Gaultheria pyrolifolia J.D. Hooker ex C.B. Clarke from Sikkim. The specimens from Lachen constitute the lectotype,

Jal | ie EA I Lage NIA | LINKEN) 1 £ 4 mM 4 1 £, 4l RAs lancha

y precisely

Fritsch and Trock, Lectotypification in Gaultheria

| HERB. LUGD. m 3:903. 13- 265 |

Wot cm

Herb.Lugd.Batay.

Gantt

Ax <s GZ.

: 72

eee eu

TY [2 E! uen Malesiana

1 [4

G -

d pa laicas dile

er r4 Len P

102330

cou CO

Fic. 2. Image of the lectot

100 Journal of the Botanical R h Institute of Texas 1(1)

Himalayan specimen in addition to one from Japan, because mention is made neither of it nor any other

potentially confounding nomenclatural issues in the protologue.

If the specimens under consideration were one and the same species, then one might argue that the only

nomenclatural issue of consequence involves the orthographic similarity between the epithets “pyrolifolia” and

“pyroloides.” Article 60.3 of the International Code of Botanical Nomenclature (ICBN; McNeill et al. 2006)

states, however, that “The liberty of correcting a name is to be used with reserve..." Because the meanings

of the epithets “pyrolifolia” and “pyroloides” are distinctly different [*Pyrola-leaved” versus “resembling Pyrola”

(the whole plant), respectively], the names are not justifiably considered as orthographic variants.

Morphological Distinctness of the Himalayan and Japanese Plants

The most recent comprehensive treatments of eastern Asian Gaultheria [i.e., those of Airy Shaw (1941) and

Middleton (1991)] have followed Takeda (1918) in recognizing two species constituting the type material of

G. pyroloides, as justified by the bluish black versus white fruiting calyx observed in Himalayan and Japa-

nese specimens, respectively (Takeda 1918). The other difference between the two species cited by Takeda

(“2-aristate anther” versus “4-aristate anther”) is incorrect and undoubtedly resulted from a misreading of

the original description of G. pyrolifolia, which states “anther-cells 2-horned at apex.” In fact both species of

Gaultheria have 4-aristate anthers when these are functional (i.e., as opposed to the highly reduced anthers

of female flowers, as observed by us on specimens from the Himalaya). The use of a single character to

distinguish species should be questioned, and to our knowledge the differences between these species have

not been documented since Takeda (1918). By examining herbarium material of both species from A, BM, E,

GH, K, KUN, and S, we have observed other features that further distinguish the bluish black-fruited species

from the white-fruited species: leaves adaxially glabrous versus densely puberulent along midvein, bracteoles

ovate versus oblanceolate, flowers 4-5 mm versus 5-6 mm long, and style 222.5 mm versus 3-3.5 mm long.

The bluish black-fruited species is distributed in the eastern Himalaya from Sikkim (India) to northwestern

Yunnan (China); the white-fruited species is distributed from central Honshu (Japan) to Sakhalin Island and

Kunashir Island, Siberia (Russia) with an isolated population on Kiska Island, Alaska (U.S.A.).

Nomenclatural Resolution

In an apparent attempt to solve the nomenclatural problem of two species comprising the original material

of the same name, Takeda (1918) gave the new name G. miqueliana Takeda to the Japanese plant (as he saw

it, the epithets *pyrolifolia" and *pyroloides" were equivalent). Inexplicably, however, G. pyroloides was cited

in synonymy, and Takeda did not designate a type for G. miqueliana. The only reference to the original mate-

rial seen by Miquel was implied by the exclusion of the Hooker material from G. pyroloides [“Syn. Gaultheria

pyroloides Miq. in Ann. Mus. Lug.-Bat. i, p. 30 (1863-64)....nec Hook. f. et Thoms.”]. Because G. pyroloides

was included in the synonymy of G. miqueliana and the type of G. pyroloides (at that time undesignated)

neither explicitly nor implicitly excluded, G. miqueliana was nomenclaturally superfluous when published

and is therefore illegitimate.

Airy Shaw (1941) later attempted to solve the problem. He asserted that the name Gaultheria pyroloides

should be credited to Miquel and can legitimately be used for the Japanese species, and he thereby used the

name *G. pyrolaefolia Hook. et Thoms. [sic]" for the blue-fruited species and G. pyroloides Miq. for the white-

fruited species. Because Airy Shaw did not clearly indicate the lectotype with the word “type” or equivalent

in accordance with Article 7.11 of the ICBN (McNeill et al. 2006), he did not lectotypify the names. Neither

apparently did Hermann Sleumer, who annotated the specimen during a revision of the genus for Flora

Malesiana in accordance with Airy Shaw's opinion (Fig. 2), but whose treatment (Sleumer 1967) makes no

mention of the issue. This is not surprising in that neither species occurs in Malesia.

Authors of major taxonomic treatments of Gaultheria that include Japan, Russia, or Alaska have not

taken up Airy Shaw's opinion; rather, they have consistently treated the white-fruited species as G. miqueliana

(e.g., Ohwi 1965; Bush 1967; Hultén 1968; Middleton 1991;Yamazaki 1993). Most authors of treatments of

Fritsch and Trock, Lectotypification in Gaultheria 101

Gaultheria that include the Himalayan region have treated the bluish black-fruited species as G. pyroloides

(e.g., Hara 1966, 1982; Fang et al. 1986; Hsu 1991; Long & Rae 1991; Middleton 1991), the only excep-

tion being Fang & Stevens (2005), who follow Airy Shaw (1941) by using G. pyrolifolia. Lectotypification is

required to clarify the application of names regarding these two species.

1. Gaultheria pyroloides Miquel, Ann. Mus. Bot. Lugduno-Batavi 1:30. 1863. (Fig. 2). Gaultheria miqueliana

TAKEDA, Bor. Mac. (Tokyo) 32:195. 1918 (wow. ILLEG. sUPERFL.). Type: JAPAN. “In insula leso” (Hokkaido) [protologue], collector

undetermined [LECTOTYPE designated here: L 102330 (Herb. Lugd. Bat. No. 903, 13-265; image)].

We have lectotypified Gaultheria pyroloides on the Japanese specimen because 1) G. pyrolifolia was described

by Clarke on the basis of the Sikkim specimens (Hooker 1882) and thus there is a legitimately published

name clearly available for it; 2) the original description of G. pyroloides is based mainly or entirely on the

Japanese specimen and published as part of a treatise on the flora of Japan; and 3) the alternative of lecto-

typifying G. pyroloides on the Sikkim specimen (with G. pyrolifolia as a taxonomic synonym) would prompt

the need for a new name for the Japanese species.

2. Gaultheria pyrolifolia J.D. Hooker ex C.B. Clarke in J.D. Hooker, Fl. Brit. India 3:457. 1882 [“pyrolae-

folia”]. (Fig. 1). Tee: INDIA. Sikxim: Lachen, 13,000 ft elev., 20 Jun 1849, J.D. Hooker s.n. (LECTOTYPE designated here: K image

catalogue number K000442406; probable duplicates: E, GH, NY-image, P).

The protologue of Gaultheria pyrolifolia cites the following gatherings, the specimens of which must be

considered syntypes: “Lachen, J.D.H.; Mon Lepcha and Jongri, J.D.H., Clarke.” Because we have not been

able to examine Clarke’s material (probably at BM), we have chosen to lectotypify on the Hooker material.

We specifically have lectotypified on the K sheet, on which Hooker’s handwriting is apparent, as follows.

Two localities are indicated on this sheet: “Lachen, 13000 ft”, on what appears to be a field label, and “Mon

Lepcha 12-14000 ft”, handwritten directly on the sheet. Even though the material is now placed on the same

sheet (we consider a third label on which is printed and handwritten “Herb. Ind. Or. Hook. fil. @ Thomson”

/ “G? [Gaultheria] / “Hab. Sikkim” “Regio. Alp” / “Alt. 12214000 ped" “Coll. JDH” to be a general label that

refers to the whole sheet), from Hooker’s journals (1854) it is clear that it represents two gatherings, made

at different times (January 1849 versus June 1849) from distinct places (about 60 km apart).

In accordance with Article 8.2 of the ICBN (McNeill et al. 2006) requiring a type to comprise a single

gathering, we have chosen the Lachen specimens on the sheet as the lectotype. The Lachen specimens were

chosen over those from Mt. Lepcha because they are in closest proximity to both Hooker’s handwritten

note *G. pyrolaefolia H f & T" and an illustration accompanied by Hooker's initials of an abortive stamen,

the flowers from which are placed among the specimens nearest the Lachen label. We do so with the caveat

that it is unclear precisely which specimens on the sheet correspond to the Lachen gathering; we assume

that at least some of the specimens directly adjacent to the label belong to this gathering and thus constitute

the lectotype. Accordingly, the E, GH, NY, and P specimens must be treated as only probable duplicates,

because there appears to be no way of knowing whether the material on those sheets is from Lachen, Mt.

Lepcha, or both.

ACKNOWLEDGMENTS

We thank the curators of A, BM, E, GH, K, KUN, and S for loaned material; L.H. Zhou for selecting specimens

for loan while at K; B. Bartholomew for helpful discussion, and G. Nesom and an anonymous reviewer for

comments on the manuscript.

REFERENCES

Airy SHAW, H.K. 1941. Studies in the Ericales: IV. Bull. Misc. Inform. Kew 1940:306-330.

Bunptr, H.M. 1975. Autographes de botanistes sous forme de fiches. VII. Candollea 30:379-410.

Bush, E.A. 1967. Gaultheria. In: Shishkin, B.K., E.G. Bobrov, eds. Flora of the U.S.S.R. S. Monson, Jerusalem. Pp.

61-62.

102 Journal of the Botanical R h Institute of Texas 1(1)

FANG, R.Z. and PF. Stevens. 2005. Gaultheria. In: Wu, Z.Y., PH. Raven, D.Y. Hong, eds. Flora of China. Volume 14. Sci-

ence Press, Beijing and Missouri Botanical Garden Press, St. Louis. Pp. 464-475.

FANG, R.Z., T.L. Mine, T.Z. Hsu, and S.H. HUANG. 1986. Ericaceae. In: Wu, CY. C. Chen, S.K. Chen, eds. Flora Yunnanica.

Volume 4. Science Press, Beijing. Pp. 336-602.

Hara, H. 1966. The flora of Eastern Himalaya. University of Tokyo Press.

Hara, H. 1982. Ericaceae. In: Hara, H., A.O. Chater, L.H.J. Williams, eds. An enumeration of the flowering plants of

Nepal. Volume 3. British Museum (Natural History), London. Pp. 4-60.

HOOKER, J.D. 1854. Himalayan journals. Two volumes. John Murray, London.

Hooker, J.D. 1882. Flora of British India. Volume 3. L. Reeve and Co., London.

Hsu, T.Z. 1991. Gaultheria. In: Fang, R.C., ed. Flora Reipublicae Popularis Sinicae. Volume 57(3). Science Press,

Beijing. Pp. 45-69.

HuLTÉN, E. 1968. Flora of Alaska and neighboring territories. Stanford University Press, Stanford, CA.

Long, D.G. and S. J. Rar. Ericaceae. 1991. In: Grierson, A.J., D.G. Long, eds. Flora of Bhutan. Volume 2, Part 1. Royal

Botanic Garden, Edinburgh. Pp. 357-404.

McNelLt, J., ER. Barrie, H.M. Bumper, V. DemouLin, D.L. HAWKsworTH, K. MARHOLD, D.H. NicoLson, J. PRADO, P.C. Silva, J.E. SkoG,

J.H. Wiersema, and N.J. TunLAND. 2006. International code of botanical nomenclature (Vienna Code). Regnum

Veg. 146.

Miooteton, D.J. 1991. Infrageneric classification of the genus Gaultheria L. (Ericaceae). Bot. J. Linn. Soc. 106:229-

258.

MiqueL, F.A.W. 1863. Annales musei botanici Lugnuno-Batavi. Volume 1. Apud C.G. van der Post 1863 1869,

Amstelodami.

Ohwi, J. 1965. Flora of Japan. Smithsonian Institution, Washington, D.C.

SLEUMER, H. 1967. Ericaceae. In: Van Steenis, C.G.G. J., ed. Flora Malesiana. Series 1, Volume 6, Part 5. N.V. Dijkstra's

Drukkerij V/H Boekdrukkerij Gebr. Hoitsema, Groningen, Netherlands.

Takena, H. 1918. Notes on far eastern plants. Bot. Mag. (Tokyo) 32:194-203.

YAMAZAKI, T. 1993. Ericaceae. In: Iwatsuki, K., T. Yamazaki, D.E. Boufford, H. Ohba, eds. Flora of Japan. Volume 3a.

Kodansha, Tokyo. Pp. 6-63.

A NEW NAME FOR THE WELL-KNOWN ASPLENIUM (ASPLENIACEAE)

FROM HALE COUNTY ALABAMA

Brian R. Keener! LJ. Davenport

University of West Alabama Samford University

Department of Biological and Environmental Sciences Department of Biological and Environmental Sciences

Livingston, Alabama 35470, U.S.A. Birmingham, Alabama 35229, U.S.A.

bkeener@uwa.edu lidavenp@samford.edu

ABSTRACT

The name of the hybrid fern Asplenium x ebenoides R.R. Scott pro sp. [= x Asplenosorus ebenoides (R.R. Scott) Wherry] has long been

misapplied to the well-known Asplenium species of reticulate origin from Hale County, Alabama. A new name, Asplenium tutwilerae,

and description are provided.

RESUMEN

Se ha aplicado mal el nombre del helecho hibrido Asplenium x ebenoides R.R. Scott pro sp. [= x Asplenosorus ebenoides (R.R. Scott)

Wherry] al bien conocido Asplenium de origen reticulado de Hale County, Alabama. Damos un nombre nuevo, Asplenium tutwilerae,

y su descripción.

INTRODUCTION

The fern name Asplenium ebenoides was originally published by R.R. Scott (Scott 1865). He based his de-

scription on a single plant, collected in 1862, from near Philadelphia, PA on the west bank of the Schuylkill

River.

As an editorial note appended to that article, Thomas Meehan suggested that the discovery might

represent a fern hybrid, since only a single specimen had been found: “Is it a hybrid or variation? or, is it a

species? Is it the last individual of a declining race, or is it the first creation of a new one?" The next year,

Berkeley (1866) declared the plant to be a hybrid and correctly identified its parents: Asplenium platyneuron

(L.) B.S.P. and A. rhizophyllum L. [2 Camptosorus rhizophyllus (L.) Link].

Seven years later, in a list of significant collections including “Asplenium eb[e]noides R.R. Scott,” D.C.

Eaton (1873) announced, “A new locality for this very rare species has been found by Miss Julia S. Tutwiler

near Havana, in Central Alabama. ... As every fact connected with this singular and disputed form [emphasis

added] will interest botanists in general and fernists in particular, we extract freely from Miss Tutwiler's

letter, which is besidels] brimful of botanical spirit:

I found it in a little magic spot, a Fairy-glen, about five miles from my home. You must know that we live in Central Alabama ... ina

hilly country of sand and red clay, with long red gullies washed everywhere into the hills, but no rocks except pudding-stones [a type of

conglomerate]. One day I happened to hear of beautiful mossy crags and cliffs some miles So and went to seek them. To my delight

and surprise, I found a little narrow glen, which seemed to have been picked up h the Blue Ridge and carried bodily through

the air to be dropped down in this odd place. ... There seemed a separate soil and climate to this little D of nature. I found there five

ferns which I had never seen in any other spot around us.

Most importantly, Tutwiler' list of ferns in the “Havana Glen" included Asplenium ebenoides and its purported

parents, A. platyneuron [“Asplenium ebeneum"] and A. rhizophyllum |*Camptosorus"].

A quarter century later, in a paper on the habitats of rare ferns in Alabama, Underwood (1896) reported

on his own visit to Havana Glen and raised doubts about the hybrid origin of Asplenium ebenoides:

The glen is a deep gorge cut in a conglomerate rock, well wooded and shaded. [Several fern species are present.] But the object of our

search is here in considerable quantity, in fact the commonest fern of the glen, Asplenium ebenoides. ... Many have regarded it a hybrid,

but the display of the species at Havana clearly demonstrates that it is not a hybrid at all. ... It appears to be multiplying, as many young

ics, Tuscaloosa, Alal 34487-0345, U.S.A.

J. Bot. Res. Inst. Texas 1(1): 103 — 108. 2007

£+sha D o ID L

104 Journal of t titute of Texas 1(1)

plants were seen in the rock crevices. This myth of hybridity may be put aside, for Asplenium ebenoides is as clearly defined a species as

we possess in the genus Asplenium. ...

Soon after, Maxon (1900) reported on his own visit to “the somewhat famous locality" of Havana Glen. His

conclusions contrasted markedly with Underwood's. He found Asplenium ebenoides to be quite abundant

and A. platyneuron to be “common in the near vicinity,” but its other supposed parent, A. rhizophyllum, was

“not in great evidence, ... though it had previously been found here in small quantity.” The sheer number

of individuals of A. ebenoides, despite the lack of one parent, raised the question in Maxon's mind “as to

whether the fact of the fern’s fertility effectually disposes of the supposition of its hybridity. May it not be

a fertile hybrid?" And, *To my mind the supposition of hybridity for A. ebenoides is not weakened by the

discovery of its evident fertility."

Slosson (1902) tested the origin of Asplenium ebenoides by forcing the hybridization of gametophytes of A.

1 +

platyneuron and A. rhizophyllum, then comparing the istics of the resulting crosses to plants collected

in various parts of the eastern United States and from Havana Glen (the latter collections by Underwood

and Maxon). Her succinct conclusion: “Surely we have here convincing proof of the origin of A. ebenoides.”

Wherry and Trudell (1930) reported on their 1929 expedition to Havana Glen, “the famous station

where this hybrid spleenwort reproduces itself.” They found Asplenium ebenoides to be “less abundant than in

former years, [with] only about 25 adult plants being seen during an hour's search. Itis, however, definitely

reproducing itself by spores ...” They also noted, “One of the parent ferns, Asplenium platyneuron, occurs

sparingly on the same rocks, but no Camptosorus could be found in the vicinity.”

Wagner (1954) determined the cytological reason behind the fertility of the Havana Glen popula-

tion—and the unimportance of the lack of one parent. His studies of sterile Asplenium ebenoides from a

Maryland population revealed “72 univalents, 36 from each of the parents.” By contrast, the fertile Alabama

population had *72 normal-appearing chromosome pairs," or 144 total chromosomes. Thus, the Havana

Glen population is a natural and self-reproducing allotetraploid

Finally, Wagner and Whitmire (1957) produced an allotetraploid by culturing the few unreduced, viable

spores from a diploid Asplenium x ebenoides originally collected in Maryland. In their words, “The culture

allopolyploid contrasts with the Alabama wild type in an ensemble of characters, both sporophytic and

gametophytic”; these differences include blade texture, blade color, frond outline, width of pinnae, number

of dwarf or abortive pinnae, form of pinnae margins, and the outline of gametophytic wings. As explanation,

“Even seemingly trivial genetic differences where the parents are as distantly related as Asplenium platyneuron

and A. rhizophyllum might be magnified in new combinations between them, and produce unexpectedly

strong differences in the allopolyploids formed from different parental varieties in different localities."

Despite the above noted differences in chromosome complement, fertility, and morphology, “the sterile

hybrid Asplenium platyneuron x rhizophyllum and its allopolyploid derivative" were treated together as Asple-

nium ebenoides in the recent Flora of North America treatment (Wagner et al. 1993).

DESCRIPTION

The description below is based on specimens from the only locality from which Asplenium tutwilerae has been

collected to date—the well-published location near Havana in Hale County, Alabama. Generally referred to

as *Havana Glen" (Walter et al. 1982), this north-south oriented ravine contains a maturing deciduous forest

of various red oaks (Quercus spp.), beeches (Fagus grandifolia Ehrh.), and hickories (Carya spp.). The sides of

the ravine are rather steep, encompassing roughly 30 m of elevation. An intermittent stream in the ravine

bottom is surrounded by dense stands of Illicium floridanum Ellis. Along the upper third of the west-facing

slope occur outcroppings of ferruginous conglomerate rocks that contain pebbles of quartz and chert (Fig.

1A). It is in the crevices of these rocks that A. tutwilerae can be found.

Asplenium tutwilerae B.R. Keener & L.J. Davenport, sp. nov. (Figs. 1B, 2). Te: U.S.A. ALasawa: Hale Co.: Havana,

growing in shaded crevices along the upper portions of sandstone conglomerate cliffs, 28 Jul 1900, CJ. Pollard & WR. Maxon 335

(HOLOTYPE: US; isotypes: MO, NY [2], PH [2], US).

ACCHEI

t Asplenium x

7 F

105

Fic. 1. A. F

106

ff^ Dat e ID L

Journal of

titute of Texas 1(1)

E TAR

A *

z(442886 )3

Nar ERST

UNITED STATES NATIONAL HERBARIUM.

DEPOSITED BY THE SMITHSONIAN INSTITUTION.

PLANTS OF ALABAMA,

COLLECTED AT HAVANA, MALE. COUNTY,

Asplenium cb ides R.R

HO YPE of: E

Asplenium tutwilerae B.R. Keener & L.J. Davenport | ; SET

det, Brian R. Keener, 2007 p PARTES LU BOLLARD a 2

a | WILMIAM R. Maxon, f to im 335.

Fi. 2. Holotype (C.L. Pollard & W.R. Maxon 335, US) of Asplenium tutwilerae.

Keener t, Asplenium x el id : M m

Planta inter A. rhizophyllo L. et A. platyneurone (L.) B.S.P., species e America boreali orientali; differt a A. rhizophyllo frondibus latioribus

ad maturitatem basi pinnatis vel pinnatifidis; differt a A. platyneurone frondibus ad maturitatem dimidio distali pinnatifido vel integro-

subintegro.

Plants epipetric. Roots numerous, filiform, to 6 cm long, monopodially branching. Rhizomes horizontal

to erect, 0.5—2 cm x 0.3-0.5 cm, imbedded by adventitious roots and bases of old and current stipes, scaly

near apices; scales narrowly triangular, 1-4 mm x 0.2-0.5 mm, pellucid, essentially one cell thick, with

thicker brownish black secondary walls forming areolate reticulum. Fronds dimorphic; smaller, mostly

sterile blades usually appressed to the substrate; larger, fertile blades, ascending to erect. Stipes castaneous

to purplish brown, lustrous, proximal 1/4 scaly, distal 1/2 pubescent; scales similar to those of rhizome,

reduced to linear-triangular to linear, reticulum also reduced to consist of midvein and/or cross hatchings;

pubescence of clavate, orange reddish hairs; stipes of smaller fronds 0.5-2.5 cm long, recurved; stipes of

larger fronds 2.0-9 cm long, ascending to erect. Blades extremely variable; herbage green, opaque, not

leathery, pubescent; pubescence of clavate, orange reddish hairs; smaller fronds lanceolate, 2-11 cm x 1-2

cm, pinnatifid in proximal 1/2-2/3, apices long attenuated in distal 1/3-1/2, with margins irregularly lobed

or entire-serrulate, occasionally pinnately compound in proximal 1/4; larger fronds lanceolate, 7-18 cm x

2-8 cm, pinnately compound in proximal 1/4-1/3, pinnatifid for middle 1/2, apices long attenuated in distal

1/4—1/3 with margins irregularly lobed or entire-serrulate, occasionally producing viable plantlets at apex.

Rachises castaneous to purplish brown, lustrous proximally, green, dull distally, pubescent; pubescence

of clavate, orange reddish hairs. Pinnae of smaller fronds 0-1 pairs, ovate, 0.5—0.9 cm x 0.4-0.5 cm, bases

truncate, apices obtuse, margins entire; pinnae of larger fronds 1-2 pairs, lanceolate, 0.7-2 cm x 0.4-1 cm,

bases truncate to auriculate, apices obtuse to acute, margins entire to crenulate-serrulate. Segments of

smaller fronds ovate, 0.3-1 cm x 0.2-0.4 cm, apices obtuse, margins entire to crenate-serrulate; segments of

larger fronds lanceolate, 0.3—5 cm x 0.3-1 cm, apices acute to obtuse, margins entire to crenulate-serrulate.

Sori of smaller fronds when fertile, 1-3 per pinna or segment in proximal and middle portion of frond, 1

on each side of the midrib corresponding with each lobe or tooth on distal attenuated apices; sori of larger

fronds, 1-18 per pinna or segment in proximal and middle portion of frond, 1 on each side of midrib cor-

responding with each lobe or tooth on distal attenuated apices. Indusia present, membraneous, attached

along one margin. Spores 64 per sporangium. 2n = 144.

Additional specimens examined: UNITED STATES. ALABAMA. Hale Co.: near Green Springs, 31 Jan 1874, Prof. Tutwiler s.n. (NY);

northern Alabama, 1877, Miss Tutwiler s.n. (MO); near Havana, 20 Mar 1878, E.A. Smith 20 (UNA [2], US [2]); Havana, 1884, JW.A.

Wright s.n. (NY); Aug 1890, J.W.A. Wright s.n. (PH); Near Havana, 16 May 1896, L.M. Underwood s.n. (MO [2], NY [3], US); rocky glen,

Havana, 21 Dec 1898, W. Trelease 326798 (MO); Havana, Jan 1905, Whatley s.n. (UNA); near Havana, Mar 1907, J.W. Moreland s.n. (PH);

one mi N of Havana, 9 May 1929, E.T. Wherry s.n. (US); Havana, 11 Jul 1953, C. O’Kelley & R. Chermock s.n. (UNA); Havana Fern Glen,

ca. 1 air mi N of Havana, 26 Jul 2006, B.R. Keener 3023 with L.J. Davenport, R. Cobb, & N. Cobb (UNA).

DISCUSSION

The name Asplenium x ebenoides has been misapplied to the Havana Glen population since its discovery in

the latter half of the 19th century. In a summary paper published 15 years after his determination of the

allotetraploid nature of that population, Wagner (1969) argued for a “single, simple approach” to the naming

of hybrids, “such as using the hybrid binomial. ... The question of whether a given hybrid is ‘fertile’ or ‘sterile,’

diploid or polyploid, is not pertinent [emphasis added].” We, however, argue just the opposite: The Havana Glen

population of Asplenium tutwilerae is sexually viable and on its own evolutionary track; therefore, it should

be recognized as a distinct species separate from A. x ebenoides.

Asplenium tutwilerae qualifies as a distinct species under the Biological Species Concept (Mayr 1963)

due to its being a sexually reproducing population that is reproductively isolated. It also qualifies under

various phylogenetic species concepts in that it represents a distinct and unique monophyletic lineage (Baum

& Donoghue 1995; Mayden 1997).

Reticulate evolution by means of a hybridization event has been well documented in the genus Asple-

nium (Smith & Levin 1963; Wagner 1954; Werth et al. 1985b). Two other Aspleniums that have arisen in a

108 Journal of the Botanical R h Institute of Texas 1(1)

fashion similar to Asplenium tutwilerae are currently designated as distinct species, A. pinnatifidum Nuttall

and A. bradleyi D.C. Eaton (Wagner et al. 1993). It is worth noting that both of these are believed to have

originated in more than one location (Werth et al. 19852) while A. tutwilerae is known currently to have a

single origin.

With a single origin and a single population, Asplenium tutwilerae immediately assumes its place as one

of the rarest fern species in the world. Efforts must begin to insure its preservation.

ACKNOWLEDGMENTS

We thank the herbarium curators of MO, NY, PH, and US for specimen loans. We greatly appreciate Richard

and Nancy Cobb for locality logistics as well as continued support and encouragement. Also, the assistance

of Guy Nesom with the Latin diagnosis was extremely helpful and generous. We are also grateful to Richard

Buckner for photographing the type specimen.

REFERENCES

Baum, D.A. and MJ. DonocHue 1995. Choosing among alternative "phylogenetic" species concepts. Syst. Bot.

20:560-573.

BenkeLEv, M.J. 1866. [Not seen. Quoted in Weatherby, C.A. 1949. Rare Scott's spleenwort: one chance in a thousand.

Horticulture 27:85, 119.]

EATON, D.C. 1873. Asplenium eblelnoides. Bull. Torrey Bot. Club 4:17-18.

Maxon, W.R. 1900. Notes on the validity of Asplenium ebenoides as a species. Bot. Gaz. 30:410-415.

MAYDEN, R.L. 1997. A hierarchy of species concepts: the denouement in the saga of the species problem. In: M.F.

Claridge, H.A. Dawah, and M.R. Wilson, eds. Species: the units of biodiversity. Chapman & Hall, London. Pp.

381-424.

Mayr, E. 1963. Animal species and evolution. Harvard University Press, Cambridge, MA.

Scott, R.R. 1865. Description of a new Amlerican] fern. Gard. Monthly 7:267-268.

SLosson, M. 1902. The origin of Asplenium ebenoides. Bull. Torrey Bot. Club 29:487-499.

Smith, D.M. and D.A. Levin. 1963. A chromatographic study of reticulate evolution in the Appalachian Asplenium

complex. Amer. J. Bot. 50:952—958.

UNDERWOOD, L.M. 1896. The habitats of the rarer ferns of Alabama. Bot. Gaz. 22:407-413.

WacNER, W.H. Jr. 1954. Reticulate evolution in the Appalachian Aspleniums. Evolution 8:103-1 18.

Waaner, W.H. Jr. 1969. The role and taxonomic treatment of hybrids. Bioscience 19:785-789.

WAGNER, W.H. JR, R.C. Moran, and C.R. WERTH. 1993. Aspleniaceae. In: Flora of North America Editorial Committee,

eds. Flora of North America Vol. 2. Oxford University Press, New York. Pp. 229-245.

WAGNER, W.H. Jr. and R.S. Whrrmire. 1957. Spontaneous production of a morphologically distinct, fertile allopolyploid

by a sterile diploid of Asplenium ebenoides. Bull. Torrey Bot. Club 84:79-89.

WALTER, K.S., W.H. WAGNER JR, and F.S. WaGner. 1982. Ecological, biosystematic, and nomenclatural notes on Scott's

spleenwort, x Asplenosorus ebenoides. Amer. Fern J. 72:65-75.

WERTH, C.R, S.l. Guttman, and W.H. EsHBAUGH. 1985a. Recurring origins of allopolyploid species in Asplenium. Sci-

Sucesos

WerTH, C.R., S.l. GUTTMAN, and W.H. EsHBAUGH. 1985b. Electrophoretic evidence of reticulate evolution in the Ap-

palachian Asplenium complex. Syst. Bot. 10:184-192.

Wuerry, E.G. and H.W. TrupeLL. 1930. The Asplenium ebenoides locality near Havana, Alabama. Amer. Fern J.

20:30-32.

RELATIONSHIPS OF HOUSTONIA PROSTRATA (RUBIACEAE) OF MEXICO AND

ARIZONA ANDA REVIEW OF HOUSTONIASUBGENERA AND SECTIONS

Edward E. Terrell!

Research Associate

Department of Botany

National Museum of Natural History

th«oninn Institution

Washington, DC 20013-7012, U.S.A.

ABSTRACT

d ecd a species discovered in 1899, has been collected in Baja California, Sinaloa, Sonora, Mexico and Cochise Co., Arizona.

e been problematical for many years. Comparisons f seed and pollen morphology in Houstonia and Hedyotis species

and in Lucya E UON support recognition of Houstonia prostrata as the type species of a new monotypic subgenus, Houstonia subgenus

Porotis, named for the unique numerous and regularly-arranged pores in the seed testa. Review of previously named subgenera and

sections in Houstonia indicates that the subgenus Chamisme should be restricted to the Houstonia purpurea L. group of four species. The

group of ten Mexican and southwestern United States species is recognized as Houstonia subgenus Ericotis.

RESUMEN

Houstonia prostrata, una especie descubierta en 1899, se ha recogido en Baja California, Sinaloa, Sonora, México y Cochise Co., Arizona.

Sus relaciones han sido problemáticas durante muchos afios. Basados en comparaciones de la morfologia de la semilla y del polen en

especies de Houstonia y de Hedyotis y en Lucya tetrandra se concluye que merece el reconocimiento como especie tipo de un nuevo sub-

género monotípico, Houstonia subgénero Porotis, nombrado así por los únicos poros testales numerosos y regularmente dispuestos de

la semilla. La revisión de subgéneros y de secciones previamente nombrados en Houstonia indica que el subgénero Chamisme se debe

restringir al grupo de Houstonia purpurea L. de cuatro especies. El grupo de diez especies mexicanas y del sudoeste de Estados Unidos

se reconoce como Houstonia subgénero Ericotis.

INTRODUCTION

Houstonia prostrata Brandegee is an annual herb native to Baja California Sur, Sinaloa, and Sonora, México,

and Cochise County, Arizona. It was first collected by T.S. Brandegee in Baja California Sur in 1899, and no

other new collections from Baja are known to the present writer. In 1904 Brandegee collected and described

another new species, Houstonia parvula, in Sinaloa. Standley (1918), in treating the North American flora,

placed Houstonia parvula in synonymy under Houstonia prostrata. Shreve and Wiggins (1964) in their Sonoran

Desert flora treated the two species as varieties: Houstonia prostrata var. prostrata with branches prostrate,

internodes shorter than leaves, leaves mostly sessile, and var. parvula with branches erect, internodes mostly

equaling or exceeding leaves, leaves mostly short-petiolate. My study of the types (Figs. 1, 2) and other col-

lections indicates that these differences are minor and overlap greatly, tly, I have not recognized

varieties and have treated the two species as one under the older species name. Houstonia prostrata has also

been found in Sonora, and was collected in 1971 in the United States by Mason, Canfield, and Gilbertson

who found it in Guadeloupe Canyon, Cochise Co., Arizona.

MORPHOLOGY AND TAXONOMY

In recent years Houstonia prostrata has been treated as Hedyotis vegrandis W.H. Lewis, a new name under

Hedyotis necessitated by a prior use of the name Hedyotis prostrata (see nomenclature below).

The question of the circumscription of Hedyotis has been a knotty problem for many years (reviewed

by Terrell 1996:16). Seed and capsule characters along with chromosome numbers and pollen morphology

are important in the classification of the tribe Hedyotideae and were used in several papers by Terrell and

"Address for correspondence: 14001 Wildwood Drive, Silver Spring, Maryland 20905, U.S.A.

J. Bot. Res. Inst. Texas 1(1): 109 — 119. 2007

110 Journal of the Botanical Research Institute of Texas 1(1)

F 1 Isnt £ LI. gene PA y J (MC 295970

IG. y \ JIOLO/ 7].

collaborators.The seed and capsule characters proved valuable in establishing tribal (Terrell and Wunderlin

2002) and other taxonomic limits (Robbrecht 1989; Terrell et al. 2005) in the Hedyotideae. Extensive varia-

tion in seed structure adds to the impression that the genus Hedyotis has been too broadly circumscribed

in the past. Hedyotis subgenus Hedyotis includes its type species, H. fruticosa L., native to Sri Lanka and

southern India. This species and a number of similar Asian and Pacific species are recognized by their cap-

sule and seed morphology (Terrell and Robinson 2003). Twenty-one Hawaiian species formerly in Hedyotis

were placed in the resurrected genus Kadua (Terrell et al. 2005). I now recognize Hedyotis species as having

seeds without ventral depressions and with a prominent hilar ridge on an otherwise level or convex ventral

face. In contrast, Houstonia prostrata has a large ventral depression containing a hilar ridge, juently, it

is here excluded from the genus Hedyotis.

In a molecular study of Houstonia Church (2003) grouped Houstonia prostrata with Stenotis, a genus

with seven species in Baja California (Terrell 2001, formerly in Hedyotis), and the monotypic genus Carterella

(Terrell 1987). The molecular data disagree with the current morphological data on Stenotis and Carterella,

which are so different in morphology from H. prostrata that I do not mention them in the present study.

Church concluded that the phylogenetic placement and taxonomic status of Houstonia prostrata “should be

reviewed more thoroughly before including it in Hedyotis or Stenotis.” I am in agreement with this statement

Terrell, H + 1 trat ; cuhnanara an rl cartinnc 111

Fic. 2. Isotype of Hi ia | la Brandegee (US-571999).

and believe that both morphology and molecular data are important and should be utilized in taxonomic

studies.

Comparison of Houstonia prostrata seeds with seeds of previously examined Hedyotideae revealed only

three genera having hilar ridges in ventral depressions. These are the following: (1) the Asian genus Neanotis,

(2) Lucya, a monotypic genus of the West Indies, (3) Houstonia, a genus of 20 species occurring only in North

America (Terrell 1996).

Neanotis was compared with Hedyotis (sens. lat. incl. Houstonia) by Lewis (1966). He found that Ne-

anotis pollen was 5-12 aperturate, whereas Hedyotis pollen was 3 or 4 aperturate. He also listed five other

differences in the pollen of the two genera and concluded that Neanotis was fully distinct from Hedyotis. His

conclusions are accepted here.

£+sha D o ID L

112 Journal of t titute of Texas 1(1)

The second genus for comparison with Houstonia prostrata is the monotypic genus Lucya (Table 1). Lucya

tetrandra occurs on rock outcrops and similar habitats in Puerto Rico, Cuba, Dominican Republic, Haiti, and

Jamaica. It is readily distinct from Houstonia prostrata in habit and aspect, and in being perennial, 4-25 cm

tall, with tubers and ovate or elliptic leaves, compared with H. prostrata, a low inconspicuous annual with

oblanceolate or linear leaves (Table 1). The seeds of Houstonia prostrata (Fig. 3) and Lucya (Fig. 4) are similar

in having thickened involute (rolled) margins, in being longitudinally bowed, and in having a large ventral

depression. They differ in the following: Lucya has (1) only a scar in the ventral depression instead of a hilar

ridge, (2) a testa lacking pores, whereas H. prostrata has numerous pores, (3)6-8 calyx lobes, a marked

departure from the usual 4 lobes in all other studied taxa of this tribe, (4)6-colporate pollen compared to

3 or 4 in Houstonia species ( based on recent data supplied by the palynologists Walter H. Lewis and Joan

Nowicke). These significant basic differences lead to the conclusion that Lucya should be maintained as a

genus distinct from Houstonia prostrata. The seed similarities suggest, however, that the two taxa are rather

closely related.

A brief description of Lucya and its nomenclatural data are added below in the taxonomic treatment.

Houstonia, the third genus with seeds similar to those of Houstonia prostrata was monographed by

Terrell (1996). A full comparison of H. prostrata with Houstonia is presented in the following review of the

infrageneric taxa in Houstonia.

INFRAGENERIC TAXA OF HOUSTONIA

This genus has two subgenera and four sections as follows: Houstonia subgenus Houstonia with sections

Houstonia and Mullera and subgenus Chamisme with sections Amphiotis (2 Chamisme) and Ericotis. A diagnostic

key to these taxa was provided by Terrell (1996: 20-21). The seeds of several of the species mentioned here

are illustrated in Terrell (1996: Figs. 1-4). The infrageneric taxa are considered below in their order.

Houstonia subg. Houstonia includes six species that are distributed throughout much of the eastern

United States, southeastern Canada, and southeastern U. S. south to Florida and eastern Texas. The type

species is H. caerulea L., the traditional Bluets. They all have a similar aspect or habit, and are small herbs,

soft-stemmed, spring-flowering, and with salverform corollas. Section Houstonia with five species has (1)

subglobose seeds each with a circular orifice opening into a subglobose hilar cavity lacking a hilar ridge, (2)

pollen 3-aperturate, (3) chromosome x = 8, except 7 in H. procumbens (J.F. Gmel.) Standl.. A sixth species,

H. rosea (Raf.) Terrell in section Mullera, differs in having (1) seeds with a hilar ridge in a shallow depres-

sion (2) pollen 4-aperturate, (3) chromosome number x = 7 (the chromosome and pollen data from Lewis

1962, 1965). Houstonia procumbens differs from other species in section Houstonia in having a chromosome

number of x = 7 and capsules widely dehiscing and sometimes separating into two halves and deflexed to

the base of the capsule. Church and Taylor (2005) in molecular studies found that H. procumbens and H. rosea

were genetically quite distinct from other species and are more closely related to each other than previously

known. Houstonia procumbens is tetraploid, H. rosea diploid. The data suggest that it would be more accurate

to include H. procumbens with H. rosea in the section Mullera, and this is done in the following taxonomic

treatment. The subgenus Houstonia with its two sections hybridizes somewhat within its own subgenus

(Church & Taylor 2005), but is quite distinct from all other Houstonia subgenera and sections.

Houstonia subg. Chamisme section Amphiotis (= section Chamisme) is typified by H. purpurea L., one of

four perennial spring- and summer-flowering species with fibrous stems, funnelform corollas, and a chromo-

some number of x = 6. All species have seeds with a low hilar ridge in a shallow depression, entire margins,

and a reticulate testa. Pollen is 3-aperturate and colpororate (Lewis in Terrell et al. 1986). The distribution

of the species includes much of the eastern U. S. and a small part of southeastern Canada. This group is

discrete and genetically distinct from other Houstonia species. Church and Taylor (2005) have provided

helpful molecular evidence about hybridization within this group.

Houstonia subg. Chamisme section Ericotis includes ten species distributed in southwestern U. S. and

Mexico. They are annual or perennial with corollas salverform, funnelform, or subrotate, and a chromosome

113

Inne

cuhnanara and cart

Terrell, H

jus2so|eoo

auou

JU9959/809

SoA

MO|Jeus

pedeus-1e»nes

NERO

sce SL

WW g0-CO

jenbeqns

uloj[auunjaJous

uu E-S |

INOJ

peAIn29J

"nd

apou Jad ouo

snojA1souJou

e1ejoe»uej|qo/A1eeui|

punsip 'snonuisAubieAs

pepunoi Jo jeuobAjod

auou

91e/n31a1

SOA

MO||eus

padeys-180Q

v 1-60

SPSC ASEC

WW € 0-70

jenbaqns

uoj[euunjaJous

WW S'7-8'0

INO}

pale

BOW JO Z| O]

SOWAD paloeMOJ[J-M9J

sno|AJsOWOU

ejejoe»uejqo ndis eu

jounsipur 'snonuis

¡guoBÁjod

auou

91e/n3H as

K|uo Je55 aeiy ¿OU

Sok

daep Ája pow

padeys-180q

Sok

WW ZO

jenbaqns

Jejngni

WW €c-oc

1uDie 0} xis

D9AJn29J

apou Jad auo

sno|Ajsowoy Ajusiedde

di||e ‘Aq 'S1eAo

jounsipur Jys

jeuo6Ajod

SoA

3jeinonal

Sok

Sak

daep Ajale1apouw

pedeus-ieoq

Sok

JaHuo| Sau €-| segni

Jejngni

WW z=

INO}

D9AJn29J

Z 01 10 ƏJISSƏS

apou Jad auo

sno|Ajsowoy Ajjuaiedde

‘dijjoueu "jeaui| ^2ue[qo

SIEM ajoue

adeys sajoae

juesaJd saJod P]59]

adÁ] 2159)

juesaJd ebpu Jelly

»piui ‘AINJOAU! SUIBIE LA

UoissaJdap

adeys

pamog K|jeuipniibuo|

uu y1Bua] spaas

Ww sajnsde)

ww y1bua] siayjuy

seqoj|/seqni

adeys

WU u1bus| sej[010)

19qUINU saqo| Xe)

Puny uSuM s|s»iped

ww u1bus| spad

SJƏMO|4 "10gu|

KIRS

adeys

S7-S LULU O£-G LULU ce-S WW p|- ww y1Bua] sanea]

0£-€ 0c-€ SZ- 6-7 WUD 3ubieu wears

quay jenuue quay jenuue SJ9QN} uin jeiuuaJed quey jenuue yqeH

DSO2SIAQns DIUOJSNOH D4OJJIA4Dd DIUOJSNOH DApupv.ja} DÁIN] DJD4JsO4d DIUOJSNOH

‘SOD BANS DIUOISNOH Jo seeds OM} pue pon] 'piDIsoJd DIUOISNOH JO s1932e4eu *| 318v].

114

Journal of the Botanical Research Institute of Texas 1(1)

Se S " es ae M

anal iG PUE ^

2 RE

; cr /

day

^" y

qa .

ARS IS

J £ LI.

by SEM. A-B, D— [rod n. id 1829 (GH), Paja! {alitoa Sur. C, Mason etal. 3061 (ARIZ), Arizona.

lled D—F, enlargements of testa

Fic. 3.5

A, seeds strongly longitudinally bowed: B, Gy ventral f.

D I

| ith | indistinct Walls

Terrell, H + H trat ; cuhnanara an rl cartinnc 115

^ ,

al H tid £

Fic. 4. Seeds of L tetrand, ined by SEM. A-D, Leonard & Leonard 12702 (US), Haiti. A,

[| | I £. T 4

1. Bar scales in microns.

d. C, same, enlarged. D

"I "E

number of x = 11 (unknown for two species). Houstonia rubra Cav. is the type species, noteworthy for its long

(8-41 mm) corollas. Nine of the species have seeds with coalescent areoles that appear as a jumbled mass

of intermixed areole walls (Terrell 1996, Figs 3d, 4d). These seeds are more complex than other Houstonia

seeds, and have the following characters: (1) boat- or cup-shaped seeds with shallow to deep depressions,

(2) margins entire or lobed and varying from thin and open to somewhat rolled and covering the edges

of the depressions, (3) hilar ridges sometimes fused with margins at one end of the seed, (4) some species

have a bilobed sinus at one end of the seed. Seven selected species are shown in Table 2. Church and Taylor

(2005) noted the genetic distinctness among these species. The first three species headed by H. rubra in

Table 2, have generally similar seed morphology. Houstonia humifusa (A. Gray) A. Gray is noteworthy for its

symmetrically lobed seed margins. Houstonia acerosa (A. Gray) Benth. & Hook. f. and H. wrightii A. Gray

have cupulate seeds, but otherwise are rather distinct from each other. The seventh species, H. parviflora

te of Texas 1(1)

fth

JOUIIdI OI

116

3je|n2naJ pa3S3/80) p93S3|20) p9sa|e0) po2s9[|Po2 po2s9|Po2 po2se|Po2 elsa}

əuou isoK ouou sok sak sak SoÁ | pue ebpu 1e snuls peqojq

OU ou Auen pau sak ou ou ou pago] suiD1euJ

Ərə pow buons Duos 9]eJopouJ 1ubijs yb IIs yb IIs peuinidn subire

MOI|8ys deep'uunipeuJ deep deep unipeul MO||eUs MO||eus MO||eus uolsseidep

ou ayelapow buons ou ou ou ou ipeuo1uo»

1e0q dn»/1eoq pedeus-dn» pedeus-1eoq Jaones Janes padeys-199nes adeys

vI-60 AO SI-S0 91-60 IG ALGO LC-90 wu u1bue| spees

S/v-S/C C/ 1-8/1 S/v-c/ CAES S/V— C9 AE SL ler Jousjur 1Ua1x3 sajnsde>

COFCO 90-10 OSO 80-€0 £0-€0 SOCO 01-€0 y1bua] seqo| seubns

COFCO € 1-90 0'C-80 S av ae) VOCO v0-SsCO Se ww u1bue| sqjeuyuy

jenbaqns jenbaqns saul £-1 seuln Z- 1 jenbəqns ¡enbagns saul p-c seqo| uey} 19buo| eqni

ullojouunj2dlous "Aesqns/'uunj "uunjqns/^|es ulloj[oeuunj uloj[ouunj-dJous uloj[euunj-dJous UllOJoAJes adeys

580 SOSE LISP OL-SE Eso MESS Lv-8 ww yua] $e]Jo4o>

ou ou Son ou ou ou ou So^e9| oxI|-i2e1g

ou ou sok ou ou ou ou pednoib pue ajissas 's|]

pale SoK pole sok sok SoA SoA peAIn291 Sja pəd ‘buy

ou ou SOK ou ou ou ou asolae

ou ou SOK ou ou ou ou 3JO[D2I256J S9AL9]

DAioyiAipd mybum DSOJ22D psnyiunu DSOISIAQNS aDI}JO1D DIQNA

‘S102117 snueb6qns DIUOISNOY JO sei»eds UaAas JO SILLY) Z Wav].

Terrell, H toni trat ; subgenera an d sections 117

Holz. ex Greenm., endemic to southeastern Texas, differs from the others in having reticulate testas and

polygonal areoles, however, its other characteristics, including chromosome number, resemble the other

nine species. Pollen in section Ericotis is 3-colporate and colpororate (Lewis in Terrell et al. 1986). The group

includes considerable variation among species but is fully distinct morphologically and genetically from the

other subgenera and sections.

The previous discussion of Lucya compared to H. prostrata (Table 1) concluded that the two taxa are

similar in two unusual seed characters, but have basic differences in pollen and calyx lobes; consequently,

Lucya remains a distinct monotypic genus.

Two of the section Ericotis species, H. subviscosa (A. Gray) A. Gray and H. parviflora, differing in their

seed morphology, are included in Table 1 for direct comparison with Houstonia prostrata and Lucya tetrandra

Several morphological characters are rather similar among these species (see also Table 2). Nine of the sec-

tion Ericotis species have seeds with coalescent areoles, a character absent in H. prostrata. The tenth species,

H. parviflora, does not have seeds like H. prostrata.

It is concluded that the general resemblances of Houstonia prostrata are to Houstonia, e.g., it has a seed

with a ventral depression containing a hilar ridge and 3-colporate pollen. It has other resemblances as

documented in the tables. The specialized characters peculiar to H. prostrata are the following: (1) seeds

longitudinally bent, (2) margins conspicuously thickened and involute, (3) testa with numerous conspicuous

regularly-arranged pores. (The first two of these characters also occur in Lucya tetrandra). The presence of

numerous pores in Houstonia prostrata is considered especially significant, as pores have never been found

in any other hedyotoid genus studied thus far. Houstonia prostrata is a clearly marked and distinctive new

subgenus of Houstonia, here termed as subgenus Porotis emphasizing its unique testal pores. The following

paragraphs document its nomenclatural and taxonomic characters.

TAXONOMIC TREATMENT

The preceding outline of the subgenera and sections follows the classification devised previously (Terrell

1996), however, it has become apparent that the sections Amphiotis (Chamisme) and Ericotis are fully distinct

and do not belong in the same subgenus. I had in 1991 treated Ericotis as a subgenus, and this combination

appears below as a restoration of an earlier combination. The four subgenera are listed below. Sections are

recognized only in Houstonia subg. Houstonia.

1. Houstonia L. subgenus Houstonia, Sp. Pl. 1:105. 1753. Lecrorve: Houstonia caerulea L. Six species. Section Houstonia

with four species, Section Mullera Terrell with two species.

2. Houstonia subgenus Chamisme Rafinesque, Ann. Gen. Sci. Phys. 5:227.1820. Lectotyrr: Houstonia purpurea

L. Four species.

3. Houstonia subgenus Ericotis Terrell, Phytologia 71:219.1991. Wee: Houstonia rubra Cav. Ten species.

4. Houstonia subgenus Porotis Terrell, subg. nov. Basionym: Houstonia prostrata Brandegee, Zoe 5:105.1901. Tee: Houstonia

prostrata.

Plantae parvae herbaceae annuae; corollae 1-2 mm longae tubulares; capsulae 3/4 inferiores longitudinaliter dehiscentes; semina

longitudinaliter curvata cymbiformia margine involuta, hilis linearibus in cavis prominentibus; testa in parietibus cellularum minute

multe porifera.

Plants small annual herbs, corollas 1-2 mm long, tubular, capsules 3/4 inferior, dehiscing loculicidally,

seeds longitudinally bent, cymbiform, margin involute, linear hilar ridge in a ventral depression, testa with

numerous minute pores.

Etymology.— Porotis is a name derived from poro, pore, and -otis, ear.

HOUSTONIA PROSTRATA

Houstonia prostrata Brandegee, Zoe 5:105.1901. Hedyotis d WH. Lewis, Rhodora 63:222.1961, nom. nov., non

Hedyotis prostrata Korthals, Nederl. Kruidk. Arch. 2, 2:1160.1851. H var. prostrata Wiggins in Shreve & Wiggins, Veg.

Fl. Sonoran Desert 2:1399.1964. Type: MEXICO. Baja California Sur: on dem sand of dry stream, resembling a prostrate Euphorbia,

La Palma, Cape Region, 25 Sep 1899, T.S.Brandegee s.n, (LECTOTYPE: UC; isorrres: GH!, NY! US-2!). (Fig. 1 isotype US-382879).

£s+haD o ID L

118 Journal of t titute of Texas 1(1)

Houstonia parvula Brandegee, Zoe 5:221.1905, non Hedyotis parvula (A. Gray) Fosb., Bishop Mus. Bull. Bot. 174:54.1943. Hedyotis

sinaloae W.H. Lewis, Rhodora 63:222. 1961, nom. nov. Houstonia prostrata var. parvula (Brandegee) Wiggins, in Shreve & Wiggins,

Veg. Fl. Sonoran Desert 2:1399.1964. Type: MEXICO. SinaLoaA: Gravel deposits of Tamazula River near Culiacan, 12 Oct 1904, T.S.

Brandegee s.n. (LECTOTYPE: UC!; isotypes: GH-2!, MO!, NY!, US-2!). (Fig. 2 isotype US-571999).

Small annual herb (Table 1). Stems 2—9 cm tall, slender, erect or prostrate and matted, minutely whitish

papillose-puberulent to glabrate. Leaves 5-14 x 0.5-2.5 mm, sessile or short-petiolate, narrowly oblanceo-

late, linear, narrowly elliptic, or narrowly oblong, minutely papillose above, glabrous or minutely papillose

beneath, margins often revolute, apices obtuse or acute. Stipules to ca. 1 mm x ca. 2 mm, scarious, deltate,

margins with 1-few sometimes gland-tipped teeth. Flowers apparently homostylous, one per node, subses-

sile or on pedicels to 2 mm long, becoming recurved at fruiting stage. Hypanthium (calyx cup) puberulent

or scaberulous; calyx lobes numbering 4, to ca.1 x ca. 0.6 mm, lanceolate or deltate. Corollas 1-2 mm long,

tubular, white or apices of lobes tinged with purple; tubes 0.5-1.5 mm long; lobes ca. 0.5 mm long, usually

shorter than tube; anthers ca. 0.2 mm long, elliptic, inserted at mouth of tube; stigmas included in tube, not

seen. Capsules 1.5—3.0 x 3-4 mm, wider than long, 3/4 inferior, thin-walled, fragile, 2-locular, glabrous or

minutely papillose, dehiscing widely loculicidally and splitting the septum. Seeds (Fig. 3) 4-10 or more per

capsule, 0.8-1.2 x 0.5-0.7 mm, black, somewhat compressed dorsiventrally, longitudinally bowed, cymbi-

form, in outline broadly elliptic, elliptic, or oblong, dorsal face rounded, ventral face with a moderately deep

elliptic depression, margin entire or slightly wavy, thickened, involute or inrolled, linear hilar ridge centered

in depression and 2/3-4/5 as long as seed, ridge ends sometimes slightly enlarged, areoles polygonal, with

low, indistinct walls, testa with numerous minute pores (Fig. 3D-F). Terrell (1986 et al., Figs. 7, 8) illustrated

the pollen of Houstonia prostrata with the numerous pores (the contribution of Joan Nowicke), and data sup-

plied by Lewis in that paper noted that H. prostrata pollen has colporate type A, the most common type in

the Rubiaceae and a generalized type from which species with more specialized pollen may have evolved.

The chromosome number for Houstonia prostrata is not known. Flowering August to October.

Distribution.—Stream beds, gravel deposits, llanos; México: Baja California Sur (Cape Region), Sinaloa,

and western Sonora; United States: Cochise Co., Arizona.

Additional collections. MEXICO. Sonora: Olneya-Prosopis-Cercidium llano, 27 mi W of Hermosillo on road to Kino Bay, 28 Aug 1941,

LL. Wiggins & R.C. Rollins 135 (ARIZ! CAS! DS! GH!, MICH!, MO!, NY!). UNITED STATES. Arizona. Cochise Co.; in gravel-filled

depression on rock outcrop above stream, Guadeloupe Canyon, in southeasternmost corner of county and state, 25 Aug 1971, C.T. Mason,

E.Canfield, R.Gilbertson 3061 (ARIZ).

LUCYA

Lucya DC., Prodr. 4:343.1830, nom. cons. (ICBN 2000). Wee species: Lucya tetrandra (L.) K.Schumann, in Engl. & Prantl,

Nat. Pflanzenf. 4(4):27.1891. Peplis tetrandra L., Amoen. Acad. 5:413.1759. Lucya tuberosa DC., Prodr. 4:434.1830, nom. illeg.

(fide ICBN 2000).

This limited description is based on 25 collections loaned from herbaria and descriptions in floras.

Small perennial herb (Table 1) with tubers to ca. 7 mm wide. Stems 4—25 cm tall, slender, erect,

spreading, or prostrate, glabrous or pubescent. Leaves with petioles to ca. 5 mm long, blades 5-32 x 4-17

mm, ovate, broadly ovate, or broadly elliptic, tapering or broadly rounded at base, glabrous or pubescent

to densely hirsute, sometimes with flattened hairs, apices usually obtuse. Stipules to ca. 1-2 mm long and

wide, apices sometimes with short teeth. Flowers apparently homostylous, usually one per node, on filiform

pedicels 1-7 mm long, erect, spreading or in fruit recurved. Hypanthium (calyx cup) glabrous to densely

hirsute; calyx lobes numbering 6-8, to ca. 1 mm long, linear or shortly lanceolate, glabrous or ciliate. Corollas

2.0-2.3 mm long, tubular, white; tubes ca.1 mm long; lobes ca.1 mm long, ovate, glabrous; anthers 0.2 mm

long, elliptic, inserted at mouth of tube; stigmas included in tube, not seen. Capsules 2.5 x 3-5 mm, wider

than long, 3/4 inferior, thin-walled, 2-locular, glabrous, sparsely pubescent, or hirsute, dehiscing widely

loculicidally and halves becoming completely deflexed. Seeds (Fig. 4) usually ca. 5-8 per capsule, 1.4-1.7 x

0.9-1.4 mm, black or dark brown, somewhat compressed dorsiventrally, longitudinally bowed, cymbiform,

in outline broadly elliptic, elliptic, oblong, or suborbicular, dorsal face rounded, ventral face with a rather

Terrell, Houstoni trat , subgenera and sections 119

deep elliptic depression, margin thickened, involute or inrolled, hilar scar centered in depression, dorsal face

and ventral rim with areole walls polygonal, sinuous or zigzag, low and distinct or indistinct, testa surface

irregularly rough, lacking pores.

ACKNOWLEDGMENTS

I express my appreciation to William Wergin, former manager of the electron laboratory, U.S. Department

of Agriculture, Beltsville, Maryland, and Walter Brown, National Museum of Natural History, Smithsonian

Institution, for providing scanning electron microscope facilities. Susann Braden provided micrographs at

the Smithsonian Institution. Marjorie Knowles formatted the illustrations for printing. I particularly thank

the curators of the herbaria cited in the text for loans of specimens. Walter Lewis and Joan Nowicke con-

tributed important pollen data. An anonymous reviewer contributed valuable suggestions. Paul Peterson

and Harold Robinson provided very helpful reviews of the manuscript. Harold Robinson also provided a

Latin translation. I thank Paul Peterson for searching (unsuccessfully) for Houstonia prostrata at the collec-

tion locality in Arizona.

REFERENCES

CHURCH, S.A. 2003. Molecular phylogenetics of Houstonia (Rubiaceae): descending aneuploidy and breeding system

evolution in the radiation of the lineage across North America. Molec. Phylogen. & Evol. 27:223-238.

CHURCH, S.A. and D.R. TavtoR. 2005. Speciation and hybridization among Houstonia (Rubiaceae) species: the influ-

ence of polyploidy on reticulate evolution. Amer. J. Bot. 92:1372- 1380.

Lewis, W.H. 1962. Phylogenetic study of Hedyotis (Rubiaceae) in North America. Amer. J. Bot. 49:855—865.

Lewis, W.H. 1965. Pollen morphology and evolution in Hedyotis subgenus Edrisia (Rubiaceae). Amer. J. Bot.

52:25/-2164.

Lewis, W.H .1966. The Asian genus Neanotis nomen novum (Anotis) and allied taxa in the Americas (Rubiaceae).

Ann. Missouri Bot. Gard. 53:32-46.

RosaRECHT, E. 1989. A remarkable new Chazaliella (African Psychotrieae), exemplifying the taxonomic value of

pyrene characters in the Rubiaceae. Adansonia 4:341—349.

SHREVE, F. and |. WicaiNs. 1964. Rubiaceae. Vegetation and flora of the Sonoran Desert. Vol. 2:1398- 1399.

STANDLEY, P.C. 1918. Rubiaceae. Oldenlandieae. In: N. Amer. Fl. 32:17-39.

TERRELL, E.E. 1987. Carterella (Rubiaceae), new genus from Baja California, Mexico. Brittonia 39:248-252.

TERRELL, E.E. 1996. Revision of Houstonia (Rubiaceae-Hedyotideae). Syst. Bot. Monogr. 48:1-118.

TERRELL, E.E. 2001. Stenotis (Rubiaceae), a new segregate genus from Baja California, Mexico. Sida 19:899-91 1.

TERRELL, E.E. and H. Robinson. 2003.Survey of Asian and Pacific species of Hedyotis and Exallage (Rubiaceae) with

nomenclatural notes on Hedyotis types. Taxon 52:775-782.

TERRELL, E.E. and R.P. WuNDERLIN. 2002. Seed and fruit characters in selected Spermacoceae and comparison with

Hedyotideae (Rubiaceae). Sida 20:549-557.

TERRELL, E.E., W.H. Lewis, H. Rosinson, and J. Nowicke. 1986.Phylogenetic implications of diverse seed types, chromo-

some numbers, and pollen morphology in Houstonia (Rubiaceae). Amer. J. Bot. 73:103-115.

TERRELL, E.E., H. Ro&iNsON, W.L. Wacner, and D.H. Lorence. 2005. Resurrection of genus Kadua for Hawaiian Hedy-

otideae (Rubiaceae), with emphasis on seed and fruit characters and notes on South Pacific species. Syst.

Bot. 30:818-833.

120 Journal of the Botanical R h Institute of Texas 1(1)

BOOK REVIEWS

Jonn O. Sawyer. 2006. Northwest California: A Natural History. (ISBN 0-520-23286-0, hbk.). The Univer-

sity of California Press, Berkeley, CA 94704, U.S.A. (Orders: California Princeton Fulfillment Services,

1445 Lower Ferry Road, Ewing, NJ 08618, U.S.A., www.ucpress.edu, 609-883-1759, 609-883-7413

fax). $75.00, 247 pp., 26 color illustrations, 17 maps, 23 tables, 6" x 9".

John Sawyer has been interested in the mountains of northwest California since before he arrived at Humboldt State College in 1966

and has studied them ever since. This book is an answer to his question “Why does this area look that way?" As a plant ecologist his

interests extended those of the conifers and the vegetation pattern.

The first two chapters cover the craggy Klamath Mountains and the rolling hills of the North Coast and highlight many specific

places, especially the national and state parks and wilderness areas, well worth visiting. A cautionary theme regarding visiting these

areas resounds through the book. Private (and public) lands may hide marijuana gardens. Stay on the roads, respect owners rights, and

heed the *No Trespassing" signs. The following chapters deal with geological history and the changing roles of fire and land use. He

illustrates how the region, in many ways the least modified portion of the state, is a place where many plants and animals have been

shielded from extinction. The last chapter concerns the biological future of northwest California. Nearly all of the plant and animal

species remain, as do the original vegetation patterns. Saving the wildlands that have been degraded and restoring them by setting them

aside can be done. The fragments of natural tapestries can be made complete again

His selected reading divided by chapter, sub-divided by EM or topic

in the text. Of interest is his inclusion of internet sources and unpublished works. The led works, theses and dissertations, are

h chapt nd ri hel; j 1

the fruit of the labors of many of his graduate students.

An index to plant names follows the bibliography. The names follow A Checklist of the Vascular Plants of Northwestern California,

John O. Sawyer and James P. Smith. The latest edition is available at the Humboldt State University Herbarium web site. His list includes

many recent nomenclatural and taxonomic changes, so the scientific names may differ from those in The Jepson Manual.

This remarkable volume is informative and engaging. It is a comprehensive natural history of the area and is recommended for all

libraries interested in the region. John O. Sawyer is Professor of Botany, Emeritus, at Humboldt State University. Among his previous

books are Trees and Shrubs of California, from University of California Press (2001), Ecology and Restoration of Northern California Coastal

Dunes (1998), Manual of California Vegetation (1995), and numerous ecological surveys of northwestern California.— Gary Jennings,

Library, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A.

Net G. SUGIHARA, JAN W. VAN WAGTENDONK, Kevin E. SHAFFER, Jo Kauman, and ANDREA E. Tuope (eds.). Fire

in California’s Ecosystems. (ISBN 978-0-520-24605-5, hbk.). The University of California Press,

Berkeley, CA 94704, U.S.A. (Orders: California Princeton Fulfillment Services, 1445 Lower Ferry Road,

Ewing, NJ 08618, U.S.A., www.ucpress.edu, 609-883-1759, 609-883-7413 fax). $75.00, 612 pp., 100

b/w photographs, 100 line illustrations, 8!2" x 11".

Pyrodiversity promotes biological diversity. Pyrodiversity is important in ecosystems where variation of fire severity provides much of

the fine-scale habitat variability. Fire is both an integral natural process in the California landscape and a growing threat to its urban

and suburban developments as they encroach on wildlands. Managing the balance between fire suppression, prevention, and use is

critical. This book provides an overview of the tools needed to manage that balance.

This text is laid out with the meat of the book in three parts. Part I introduces the basics of fire ecology. It includes an historical

overview of fire, vegetation, and climate in California; overviews of fire as a physical and ecological process; and reviews the interac-

tions between fire and the physical, plant, and animal components of the environment. Part Il explores the history and ecology of fire

in each of California's nine bioregions. Each has its own unique situation. Part IIl examines fire management in California, including

both Native American and post-European settlement; discusses current issues related to fire policy and management, including air

quality, watershed management, invasive plant species, native species, and fuel management; and considers the future of fire manage-

ment. Three appendices follow covering: Plant common and scientific names; Animal common and scientific names; and, Bioregions,

ecological zones, and plant alliances of California that occur is this text. A glossary and index complete the work.

This comprehensive volume, both a text and an authoritative reference tool, is the first to synthesize our knowledge of the science,

ecology, and management of fire in California. It will be a useful tool for biologists seeking to develop effective management measures

to maintain fire-dependent ecosystems. It will be equally useful to resource managers who are concerned with the appropriate applica-

tion of fire ecology management and with intelligent, cost-effective fire suppression.—Gary Jennings, Library, Botanical Research Institute

of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A.

J. Bot. Res. Inst. Texas 1(1): 120. 2007

A NEW VARIETY OF HUMBOLDTIA (FABACEAE: CAESALPINIOIDEAE) FROM THE

WESTERN GHATS OF INDIA

PS. Udayan, K.V. Tushar, and Satheesh George

Centre for Medicinal Plants Research

Arya Vaidya Sala, Kottakkal

Malappuram District, Kerala 676 503, INDIA

email: avscmpr@sify.com / avsc hoo.co.in

ABSTRACT

A new variety Humboldtia brunonis Wall. var. raktapushpa P.S. Udayan, KV. Tushar & Satheesh George is described and illustrated

from India.

RESUMEN

Se describe y se ilustra una nueva variedad, Humboldtia brunonis Wall. var. raktapushpa P.S. Udayan, KV. Tushar & Satheesh

George de la India.

The genus Humboldtia Vahl is known to have six species and one variety (Sanjappa 1986) mostly confined

to the Western Ghats of India with one species (Humboldtia laurifolia Vahl) extending to Sri Lanka. During

the course of floristic exploration along the Western Ghats of Kerala in South India, the authors collected

interesting specimens of H. brunonis Wall. On closer examination it turned out to be an undescribed taxon

which is described here as a new variety.

DESCRIPTION OF THE SPECIES

Humboldtia brunonis Wall. var. raktapushpa PS. Udayan, K.V. Tushar & Satheesh George, var. nov. (Figs.

1-3). Ter: INDIA. Kerara. Kozhikode District: Kakkayam, 11° 33' N 75? 55' E + 750 m elev., 08 Jan 2003 (fl), PS. Udayan, K.V.

Tushar & Satheesh George 01067 (HOLOTYPE: BRIT; isotypes: CAL, CALI, L, MH).

Differt a H. brunonis var. brunonis inflorescentiae late caramesinus rubra, sepalis erectis, petalis anguste obovatis, staminum filamentis

crasso 11 mm longis, staminodiis 3, globosis brevis alternalibus

Differs from H. brunonis var. brunonis by its much congested, bright crimson-red inflorescence, erect sepals,

narrowly obovate petals; staminal filaments 11 mm long, stout, alternating with 5, short, globose stami-

nodes.

Shrubs to small trees, 4-6 m high and to 40 cm gbh, bark grayish-black, coarsely fissured; branchlets

light brown, glabrous solid, sometimes swollen; stipules lanceolate, 2-4 x 0.5-1.5 cm, prominently parallel

veined, glabrous; appendages 2, similar, 1 x 0.5 cm, reniform, prominently veined, glabrous, persistent; leaves

alternate, pinnately 4-foliolate, subsessile, up to 25 cm long; rachis up to 5.5 cm long, obscurely winged,

shallowly canaliculated above, glabrous, young rachis brown tomentose; leaflets bijugate; lamina 8.5-16 x

2.5—5 em, chartaceous to thinly coriaceous, elliptic-lanceolate, obtusely acuminate at apex, inequilateral at

base; lateral veins 6—8 pairs, prominently reticulate below, dark green above and pale beneath, the margins

entire, grayish when dry, glabrous, young leaves drooping, coppery brown, brownish pubescent beneath,

glabrous above; inflorescence erect, 3-7 cm long, axillary racemes; peduncles 1-3 cm, brown pubescent, up

to 50-flowered, floriferous axis up to 6 cm; flowers ca 2 cm long, pedicels 5-7 mm long, pilose, bracts ovate,

acute, 3 x 2 mm, with a gland at the middle, light brown pubescent; bracteoles 2, 4 x 3 mm, ovate-obovate,

with a gland at the middle, obtuse at apex, with a pinkish midvein, brown pubescent, ciliate along margin;

calyx tube ca 1-2 mm long, brownish tomentose; lobes 4, 5 x 3 mm, ovate, concave, imbricate, obtuse at

J. Bot. Res. Inst. Texas 1(1): 121 — 127. 2007

122

v h Institute of Texas 1(1)

fal, Dat

Journal of

Shape tad He

ma dtd a

annt Oren,

le. C. Flower. D. Petal. E. Bract. E Bracteole

TE £f

boldtia L j raktapushpa. A. Fl i

F Li "I r

ith a st H. Pistil. |. | itudinal y

Udayan et al., A new variety of Humboldtia in India 123

e = Humboldtia|brunonis var. raktapushpa

O — H. brunonis|var. brunonis

20° YAA A

(

160 ANDHRA PRIADESH

KARNATAKA

INDIA

1 29 E

TAMIL NADU

T2P 76? 80? 84?

d H. brunonis var. brunonis (— O).

O PR £ LI. haldt’an h a Ls h [— a)!

Fic. 2 Fish t,

tip, brown pubescent, reddish; petals 3, bright crimson-red, obovate, 0.6—0.8 cm long, clawed, claws ca 1.5

mm long, prominently nerved, glabrous; stamens 5, fertile, alternating with 5 short, globose staminodes,

the staminal filaments ca 11 mm long, crimson red, stout, glabrous; anthers versatile, 2 x 1.25 mm, oblong;

ovary 5 mm long, stipitate, stipe 2 mm long, obliquely ellipsoid, densely pubescent, 2-ovuled; style 1 cm

long, slender, pilose towards base; stigma capitate; pods 3.5-4 x 1.5-2 cm, dolabriform, brown pubescent

when young, 1-2 seeded, 0.5 x 0.5 cm, brownish.

Distribution, habitat, and phenology.—Humboldtia brunonis var. raktapushpa is so far known only from

the type locality, Kakkayam along the foothills of the Western Ghats of Kerala (Fig. 2). This species grows

in the semi-evergreen forests at an elevation of about 750 m in moist shady locations along with tree species

such as Vateria indica L., Elaeocarpus tuberculatus Roxb., Euodia lunu-ankenda (Gaertn.) Merr., and Syzygium

laetum (Buch.-Ham.) Gandhi. Flowering from January to April, and occasionally at other seasons.

124 Journal of the Botanical Research Institute of Texas 1(1)

Fic. 3. Humboldtia t is var. raktapushpa. A. Young shoot. B. Inflorescence arising from the stem. C. Single flower. D. Infructescence. E. A portion of

the trunk showing bark characters.

Udayan et al., A new variety of Humboldtia in India 125

Fic. 4. Humboldtia brunonis var. brunonis. A. Young shoot. B. Inflorescence. C. Single flower. D. Infructescence. E. A portion of the trunk showing bark

characters.

Conservation significance.— Because of the very restricted distribution and no other known collections

of this species, a more detailed assessment of its distribution and biology would be valuable.

Etymology.—The varietal name ‘raktapushpa’ is derived from the Sanskrit, rakta meaning ‘red’ and

‘pushpa’ meaning ‘flower’ referring to the flower color.

PaRATYPE: INDIA. KERALA. KOZHIKODE DisT.: KAKKAYAM, 11? 33'N 75? 55' E ca 780 m, 7 Jan 2005 (fl & fr), P.S. Udayan et al. 03348 (CMPR,

£+sha D o ID

126 Journal of t h Institute of Texas 1(1)

TABLE 1. Comparison of two varieties of Humboldtia brunonis.

Characters var. raktapushpa var. brunonis

Bark Coarsely fissured and thinly flaky bark Not fissured, mottled gray

Inflorescence Dense, 3-7 cm long Lax, 10-15 cm long

Flowers Bright crimson-red White tinged with pink

Stamens & Staminodes Filaments stout, crimson red, to 11 mm Filaments slender, white, to 15 mm long;

long; staminodes globose staminodes filiform

Calyx Tube 1-2 mm long, brownish tomentose Tube 8-10 mm long, glabrescent

Petals Narrowly obovate, 6-8 mm long Broadly obovate, 10-15 mm long

Pod 1 or 2-seeded 3-4-seeded

DISCUSSION

Humboldtia brunonis Wall. var. raktapushpa P.S. Udayan, KV. Tushar & Satheesh George is known only by a

small population in the type locality. It resembles the typical form of H. brunonis in the vegetative form. The

most striking feature of the new variety is the short, bright crimson-red inflorescence with flowers borne in

dense spiral clusters. The vegetative feature that distinguishes var. raktapushpa is its coarsely fissured and

thinly flaky bark (Table 1).

The flower colour of Humboldtia brunonis Wall. had been interpreted variously by different authors.

Wallich (1832) while describing the flower states “flores magnitudine circiter illorum Tamarindi, coloris

forsan laeti aurantiaci Jonesiae....”, from this it appears that he was not sure whether the flowers are slightly

orange. However, the excellent plate (t. 233) drawn by Griffith in Wallich's Plant Asiatic Rarioris (1832) ac-

companying the description undoubtedly agrees with what is currently understood as H. brunonis Wall.

Brandis (1906) and Gamble (1919) probably following Wallich (1832) also described the flower as ‘orange’.

Sanjappa (1986) while revising the genus Humboldtia seems to be little confused. He described the flowers

of H. brunonis as ‘white’ and the petals of which as “white, pink or orange”. We had studied living popula-

tions of the typical variety of H. brunonis throughout its entire range of distribution and found that the

flowers are always white with slight pinkish tinge (Fig. 4). This is in corroboration with the observations

made by various other authors (Beddome 1871; Gandhi 1976; Saldanha & Singh 1984; Keshavamurthy &

Yoganarasimhan 1990). The reason for this different interpretation could be due to its deep pink sepals. It

is likely that many of the collectors mistook the sepals for the petals particularly when the tree is in a late

flowering stage with all its petals shed.

ACKNOWLEDGMENTS

Authors are thankful to M. Sanjappa and G.V.S. Murthy, Botanical Survey of India (BSD and N. Sasidharan,

Kerala Forest Research Institute (KFRI), for their comments on our specimens; curators of CAL, CALI, FRLH

and MH herbaria. The study was supported by grants from Department of Biotechnology (DBT), Government

of India. We are also thankful to the authorities of Arya Vaidya Sala, Kottakkal for providing facilities; Indira

Balachandran for helpful suggestions and encouragement; A.K. Pradeep for the review of an earlier version

of the manuscript and illustration; N.K. Janardhanan, for assisting during field visits; P. Subramanian and

V.V. Mohanan, Kerala State Forest Department, Kozhikkode for granting permission for the field visits.

REFERENCES

Beppome, R.H. 1871. Flora sylvatica. Gantz Brothers, Madras.

Branpis, D. 1906. Indian trees. Bishen Singh Mahendra Pal Singh, Dehra Dun.

GAMBLE, J.S. 1919. The flora of the Presidency of Madras. 1:410-41 1.

GANDHI, K.N. 1976. Fabaceae In: CJ. Saldanha and D.H. Nicolson, eds. The flora of Hassan District. Karnataka.

Amerind Publishing Co., New Delhi. Pp. 215—269.

Udayan et al., A new variety of Humboldtia in India 127

KESHAVAMURTHY, K.R. and S.N. YOGANARASIMHAN. 1990. Flora of Coorg (Kodagu), Karnataka, India. Vimsat Publishers,

Bangalore. Pp. 165-173.

SALDANHA, C.J. and B.G. SINGH. 1984. Fabaceae. In: CJ. Saldanha, ed. Flora of Karnataka, Vol. |. Oxford & IBH, New

Delhi. Pp. 374—506.

SANJAPPA, M. 1986. A revision of the genus Humboldtia Vahl (Leguminosae-Caesalpinioideae). Blumea 31:

329-339.

WALLICH, N. 1832. Plantae Asiaticae Rarioris. Vol. 3. London.

128 Journal of the Botanical R h Institute of Texas 1(1)

BOOK REVIEWS

Dennis W. SCHLICHT, JOHN A. Downey, and JEFFREY A. NikorA.2007. The Butterflies of Iowa. (ISBN 1-59829-533-

4, pbk.). University of Iowa Press, 100 Kuhl House, Iowa City, IA 52242, U.S.A. (Orders: http://www.

uipress.uiowa.edu/). $29.95, 233 pp., color photographs, 7" x 10".

By definition as well as size, this book is not intended as a field guide but rather as a manual to accompany comprehensive guides used in

identifying and studying the butterflies of lowa. Through the maps it is also a finding guide. Essential information is provided for each

species: status (breeding or otherwise), flight period, description, and habitat. Under the heading Natural History the authors note the

larval host plants (though often not specifically), larval and imago (adult form) | 1 other pertinent and interesting facts. A most

innovative feature is the “Questions” rubric, questions about each species, which indicate how much there is yet to learn about butterflies

in general and the potential research that even an amateur lepidopterist could undertake through careful observation and study.

Since the photographic plates are of pinned specimens, they lack brilliance. However, the views of dorsal and ventral surfaces of

male and female (sexually dimorphic species) will facilitate identification.

A well-executed study of the subject, this book should be helpful and stimulating to all levels of butterfly watchers and collectors

in Iowa and neighboring states.—Joann Karges, (TCU Library, retired), Botanical Research Institute of Texas, Fort Worth, TX, 76102-4060,

JONATHAN SILVERTON. 2005. Demons in Eden: The Paradox of Plant Diversity. (ISBN 0-226-75771-4, hbk.).

University of Chicago Press, 1427 East 60'" Street, Chicago, IL 60637-2954, U.S.A. (Orders: www.

press.uchicago.edu, 773-702-9756 fax). $25.00, 169 pp., 8 color plates, 542" x 81⁄2".

In this book the author, a professor and researcher at the Open University, Milton Keynes, asks and answers some important and

intriguing questions regarding alien, invasive plants and their past and future potential for the development of new species. He poses

a fundamental question “how to reconcile the evolution of diversity with the fact that natural selection favors individuals with demon

traits and dominating proclivities." (p.34)

He then takes the reader around the world to illustrate demon plants which he has studied intensely. With a first stop to show ue

extraordinary diversity in the plants of the Kew Gardens he proceeds to the Kew Laboratories where major molecular work i bli

ing a new evolutionary tree of knowledge. This Darwinian tree is the metaphor the reader follows in the book, as the author leads us to

the Canary Islands, to the chalk grass lands of England, to Mount Shimagare of Japan with its demon bamboo (Sasa) and the fir forests

there > in the Adirondacks, to D Mexico, to Barro Colorado, to Guanacaste, to the Florida Everglades.

plants, the colonizing tendencies of naturally or intentionally introduced aliens,

the importance of niches and gaps, soils and other Mr conditions, the costs of reproduction in some plants (the bloom-and-

die syndrome), competition among similar species, dispersal limitations, and transplanted ecosystems. An important chapter (^New

Demons?") concerns the development and use of transgenic crops, the realized and potential effects.

While the situation for biodiversity seems dire (the most dangerous demon is actually mankind), the author presents the restora-

tion of parts of Guanacaste as "good news," which should of course be emulated, and conservation efforts should expand around the

globe.

Thoroughly scientific, stimulating, and DR the book offers P NN insights into invasive plants (alien or endemic)

and biodiversity. Chapter notes include bibliograpl for further reading.

Research Institute of Texas, Fort Worth, TX, 76102-4060, USA.

Joann Karges, (TCU Library, retired), Botanical

J. Bot. Res. Inst. Texas 1(1): 128. 2007

AGASTHIYAMALAIA (CLUSIACEAE), A NEW GENUS FOR POECILONEURON

PAUCIFLORUM AN ENDEMIC AND ENDANGERED TREE

OF WESTERN GHATS INDIA

S. Rajkumar M.K. Janarthanam

Institute of Himalayan Bioresource Technology Department of Botany

PO. Box No. 6 Goa University

Palampur 176 061, INDIA Goa 403 206, INDIA

rajkumaraihbt.res.in jana@unigoa.ac.in

ABSTRACT

The taxonomic position of the Indian endemic tree genus Poeciloneuron was reassessed using morphological and anatomical data. Poe-

ciloneuron pauciflorum differs from P. indicum, the only other species of the genus, in its solitary axillary flowers, inconspicuous parallel

leaf venation and apotracheal banded wood parenchyma. Because of these differences we propose to place P. pauciflorum into a new

monotypic genus Agasthiyamalaia, gen. nov. Agasthiyamalaia pauciflora, comb. nov., is proposed with the support from morpho-

logical and anatomical characters.

ABSTRACT

La posición taxonómica del género arbóreo endémico de la India Poeciloneuron se ha reevaluado usando datos morfológicos y anatómi-

cos. Poeciloneuron pauciflorum difiere de P. indicum, la otra especie del género, por sus flores axilares solitarias, venación foliar paralela

inconspicua y parénquima del xilema apotraqueal bandeado. Por todas estas diferencias proponemos colocar a P. pauciflorum en un

nuevo género monotipico Agasthiyamalaia, gen. nov. Se propone la nueva combinación Agasthiyamalaia pauciflora, comb. nov.,

con el soporte d morfológicos y anatómicos.

INTRODUCTION

Poeciloneuron Bedd. is an endemic tree genus with two species belonging to the family Clusiaceae. It was

described by Beddome (1865) under the family Ternst | as a monotypic genus. Bentham and Hooker

(1862-67) also included Poeciloneuron in Ternstroemiaceae. Beddome (1871) later added another species viz.

Poeciloneuron pauciflorum to the genus. Beddome (1871) included this species under Poeciloneuron, with the

comment, "if this species remains in the genus, the generic character must be considerably altered." Even

though Poeciloneuron has anatomical similarities with the family Bonnetiaceae, because of its entire opposite

leaves (Seetharam 1985), and presence of secretary canals (Metcalfe & Chalk 1950; Dickson & Weitzman

1996), it was included in Clusiaceae (see also Engler 1888). Floral morphological and palynological studies

also fixed its position in the family Clusiaceae (Seetharam & Pocock 1978; Seetharam 1985). Detailed floral

morphological work suggested that Poeciloneuron belongs to the tribe Calophylleae, which also includes the

genera Calophyllum, Kayea, Mesua and Mammea (Seetharam 1985). However, a critical comparative study of

these two endemic species of Poeciloneuron is still needed.

METHODS

Plant Materials

Twigs with flowers and fruits were collected for morphological studies from southern Western Ghats in

evergreen forests. Fresh flowers, fruits and leaves were preserved in FAA solution for laboratory studies. The

wood samples were collected from mature branches for anatomical studies. The identify was confirmed in

the regional herbarium of BSI (MH) and voucher specimens are deposited in the Herbarium, Department

of Botany, Goa University, Goa, India.

Wood anatomical studies

Free hand sections (T.S., T.L.S. and R.L.S.) of wood were made. The sections were stained in safranin for

J. Bot. Res. Inst. Texas 1(1): 129 — 133. 2007

£s+haD o ID L

130 Journal of t titute of Texas 1(1)

1-2 minutes and washed and processed for permanent mount following Johansen (1940). All the stained

sections were observed under Leica MPS 32 microscope. The terminology of [AWA Committee on Nomen-

clature (1964) was followed in describing the wood anatomical characters.

RESULTS AND DISCUSSION

Morphology

The conspicuous reticulate veins of the leaves of Poeciloneuron indicum differ from distant parallel veins of

P. pauciflorum. In P. pauciflorum the flowers are axillary and solitary (or paired) whereas in P. indicum they

are in terminal or axillary panicles. The sepals are in two whorls in P. pauciflorum whereas they are in single

whorl of five in P. indicum (Table 1).

Wood anatomy

The wood of both species can be described as follows: Wood diffuse porous; vessels solitary, rounded in

outline, ca. 56 pm in diameter, mean member length 745 um (580—910 um) tailed, ca. 16 per mm’, perforation

simple; vessels to ray pits simple or bordered, alternate. Rays uniseriate, heterogenous, type III, 5-16 cells

in high, ca 280 pm high, ca 70 per mm". Parenchyma apotracheal, banded. Fibers thick walled, bordered

pits numerous, fiber tracheids present. The major difference between these two species based on wood

anatomy is wood parenchyma arrangement. In P. paciflorum the wood parenchyma is apotracheal banded

and in P. indicum it is paratracheal aliform type. Other characters such as vessel length and diameter, and

fiber length showed quantitative differences.

Pollen morphology

The characters of pollen morphology were adapted from Seetharam and Pocock (1978). Tricolporate iso-

polar, polar axis 20.8 + 2.2 um, Equatorial axis 22.6 + 2.0 um, P/E ratio 0.9, ectoaperature 7-12 x 1.5 um,

endoaperature 4-6 x 1-2 um, tectum 11-20 pm, perforate, more or less regular, bear warty projections. The

species of Poeciloneuron differ in the arrangement of their endoapertures. In P. indicum the endoaperture is

perpendicular to ectoaperture, whereas in P. pauciflorum it is parallel. Tectal perforations are irregular in

P. indicum and regular in P. pauciflorum, tectal crests are smooth in P. indicum whereas they are warty in P.

pauciflorum (Table 2.)

Poeciloneuron Bedd. is represented by two species: P. indicum and P. pauciflorum. The differences between

these two are substantial enough to necessitate placing P. pauciflorum in a separate genus.

KEY TO GENERA

Flowers in terminal or axillary panicles; sepals 5, in a single whorl; stamens 12; leaves with fine reticulation

Poeciloneuron

Flowers solitary or paired in leaf axils; sepals 4, in 2 whorls; stamens 16-22; leaves with distantly parallel

venation Agasthiyamalaia gen. nov.

Agasthiyamalaia S. Rajkumar & Janarth., gen. nov. Ter: Poeciloneuron pauciflorum Bedd., Fl. Sylv. 1:93, t. 93. 1871. Agas-

thiyamalaia pauciflora (Bedd.) S. Rajkumar & Janarth., comb. nov. herein.

Poiciloneuro proxima, floribus solitariis vel binatis, sepalis quattuor in verticillis duobus, staminibus 16-22, foliis venatione remote

parallela differt.

Agasthiyamalaia gen. nov. is very similar to Poeciloneuron Bedd. but differs in its solitary or paired axillary

flowers, four sepals in two whorls, 16-22 stamens and leaves with distantly parallel venation. Agasthiya-

malaia also differs from Poeciloneuron s. str. in certain micro-morphological characters. The apotracheal

banded wood parenchyma and perpendicular arrangement of pollen endoaperature to ecotaperature of the

former are distinct from paratracheal aliform wood parenchyma and parallel positioned pollen endo and

ectoaperatures of the latter.

Trees with clear bole. Leaves simple, opposite, petiolate, petiole rough, channeled. Flowers solitary or

paired in the axils of the fallen leaves, pedicellate; sepals 4, in two whorls, inner two larger than outer ones,

puberulous; petals 6-8, imbricate, pubescent within; stamens 16-22, attached to an elevated disc below

Rajkumar and Janarthanam, Agasthi ia, a genus in India 131

wd Y

Taste 1. Morphological differences between P indicum and Agasthiyamalaia (=P pauciflorum).

Characters P. indicum Agasthiyamalaia (—P. pauciflorum)

Leaf size Up to25x 6CM Up to 12x 4 cm

Leaf shape Ovate to oblong, acuminate at apex Oblong, bluntly acuminate at apex

Leaf surface Reticulate conspicuous venation Parallel inconspicuous venation

Inflorescence Axillary and terminal panicle Solitary or paired in leaf axils

Sepals 5, ovate, all equal in size 4, in 2 whorls, inner two larger

Petals 5, contorted 6-8, imbricate

Stamens 12 16-22

Fruit Without lobes and blunt apex 2 lobed, pointed at apex

Seed Testa smooth Testa wrinkled

4b | SES D EL \ | +

TABLE 2. Differences in pollen morphology between P indicum and A (=I adapted from Seetharam

and Pocock (1978).

Characters P. indicum Agasthiyamalaia (=P. pauciflorum)

Polar axis (56+) Sram 20.8+2.2mm

Equatorial axis 14.6+0.9mm 22.6+2.0mm

P/E ratio 1 0.9

Ecoaperature 8-12x 1mm 7-12 x 1.5mm

Endoaperature Arrangement 2x4mm, perpendicular to ectoaperature 4-6 X 1-2 mm, paraellel to ecoaperature

Tectum thickness 6-8 mm 11-20mm

Tectal crests Smooth Warty

Tectal perforation Irregular Regular

ovary, anthers lobulate, dehiscence longitudinal; ovary globose, 2-celled, with pair of ovules in each; style

2, divided halfway, undulate along the margins, greenish yellow. Fruit globose, pointed at the tip, dehiscent

in to two valves, one seeded. Seed hard, rounded, testa loose, membranaceous, striate, easily separable from

the seed; cotyledons very large, fleshy.

Distribution.—Southern parts of Western Ghats in Tamil Nadu and Kerala States of India.

Etymology.—The genus is named after Agasthiyamalai Hills in and around which it is found.

Agasthiyamalaia pauciflora (Bedd.) S. Rajkumar & Janarth., comb. nov. (Fig. 1). Basowvw: Poeciloneuron pauciflorum

Bedd., Fl. Sylv. 93, t. 93. 1871; Dyer in Hook. f., Fl. Brit. India. 1:278. 1874; Gamble, Fl. Madras 1:546. 1967 (repr. ed); Singh in

Sharma et al., Fl. India 3:146. 1993. Tyre: Bedd., Fl. Sylv. 1:93, t. 93. 1871.

Trees up to 15 m high, clear bole, bark grayish. Leaves with petiole, petiole up to 1.5 cm long, rough, chan-

neled; lamina coriaceous, oblong, up to 12 x 4 cm, rounded or acute at base, entire along the margin, bluntly

acuminate at apex. Flowers solitary or paired in the axils of the fallen leaves, pedicellate, pedicels up to 2.5

cm long, glabrous, green in colour; sepals 4, ovate, the outer two ca 2.5 x 3 mm, the inner two up to 8 x

3 mm, apically obtuse, green, puberulous; petals ovate, ca 0.3 x 0.2 cm, apically obtuse, white, pubescent

within; stamens 16-22, ca. 0.6 cm long. Ovary ca. 0.2 cm. Fruit globose, up to 2 x 1.7 cm.

Local name.—Puli-vayila, Puthangkolli.

Specimens examined: INDIA. Tamil Nadu: Mundanthurai to Kannikatti, 17 Mar 1917, s.l. 14647 (MH); way to Nagapothigai from

Inchikuzhi, 8 Feb 1989, R. Gopalan 90105 (MH); Etha river bank, 1000 m 24 Apr 1990, R. Gopalan 93232 (MH); bank of Sigapparu, way

to Nagapothigai, 750 m, 22 Jan 1991, R. Gopalan 94640 (MED; Valayar River bank, 900 m, 3 Apr 1991, R. Gopalan 96216 (MH); way to

Poonkulam, 900 m, 17 Apr 1992, R. Gopalan 99305 (MH); banks of Chittar, 8 km above Keeriparai, Kanniyakumari Dt., 23 Feb 1998,

S. Rajkumar 210 (Herbarium, Goa Univ.); 2 Nov 2000, S. Rajkumar 680 (Herbarium, Goa Univ.); Inchikuzhi to Kannikatty, 15 Aug 2002,

S. Rajkumar s.n. (Herbarium, Goa Univ.). Kerala: Travancore, s.d. & s.l. acc. No. 3224 (MH).

Distribution.—Banks of streams or rivers, in evergreen forests surrounded by grasslands. Locally dominant,

associated with Cinnamomum spp., Glochidion spp., Knema attenuate and Ochlandra spp. Young leaves are mem-

132 Journal of the Botanical R h Institute of Texas 1(1)

5mm

Fic. 1. Agasthiyamalaia pauciflora a. flowering shoot. b. flower c. calyx with stamens (corolla removed). d. petal. e. pistil. f. ovary. g. fruit. h.

seed.

branous, white, turning pinkish. Agasthiyamalaia (=P. Pauciflorum) is a narrow endemic and was relocated by

Ravikumar (pers. comm.) 70 years after its previous collections. It is listed as an endemic rare plant of Western

Ghats, India (Ahmedullah & Nayar 1990; Gopalan & Henry 2000; Mohanan & Sivadasan 2002).

IUCN Conservation Assessment.—This species has been assessed as Critically Endangered (CR Bl+2c ver.

2.3 (1994) by WCMC (1998) under Poeciloneuron pauciflorum Bedd. However, recent collections from several

populations, though from a small geographic region necessitates its reassessment. Mass multiplication using

tissue culture is being tested as part of a species recovery program by the Department of Biotechnology,

Ministry of Science and Technology, New Delhi, India.

Rajkumar and Janarthanam, Agasthi laia, a genus in India 133

wd Y

ACKNOWLEDGMENTS

We thank Peter Stevens, Missouri Botanical Garden for his critical comments on the earlier version of

manuscript; Joint Director (MH, Coimbatore) for permission to consult the herbarium; K. Ravikumar

(FRLHT, Bangalore) for help in locating the plant; D. Narasimhan (Madras Christian College, Chennai) for

pickled specimens; and to J.F. Veldkamp (Leiden, The Netherlands) for the Latin diagnosis. We also thank

reviewers John J. Pipoly III and Barney Lipscomb (BRIT) whose comments greatly helped us in revising

the manuscript.

REFERENCES

AHMEDULLAH, N. and M.P. Nayar. 1990. Poeciloneuron pauciflorum. In: M.P. Nayar and A.R.K. Sastry, eds. Red data

book of Indian plants. 3:87.

Beppome, R.H. 1865. On a new genus of Ternstroemiaceae: Poeciloneuron from Nilgiris. J. Linn. Soc. 8:267.

Beppowr, R.H. 1869-73. Flora sylvatica for South India. Madras.

BENTHAM, G. and J.D. Hooker. 1862-67. Genera plantarum. Reeve, London.

Dickson, W.C. and A.L. WErTZMAN. 1996. Comparative anatomy of the young stem, node and leaf of Bonnetiaceae,

including observations on a foliar endodermis. Amer. J. Bot. 83:405-418.

ENGLER, A. 1888. Guttiferae. In: Flora Brasiliensis 12(1):382-486.

GOPALAN, R. and A.N. Henry. 2000. Endemic plants of India: Endemics of Agasthiyamalai Hills. Dehradun.

lawa Committee ON NOMENCLATURE, 1964. Multilingual glossary of terms used in wood anatomy. Winterthur, Swit-

zerland: Verlagsanstalt Buchdruckerei Konkordia.

JoHANSEN, D.A 1940. Plant microtechnique. 1* Ed. McGrew Hill. New York.

Metcatre, C.R., and R. CHALK. 1950. Anatomy of dicotyledons. Vol. 1. Carendon Press. Oxford.

MOHANAN, N. and M. SivapasAN. 2002. Flora of Agasthiyamala. Bishen Singh mahendra Pal Singh, Dehradun.

SEETHARAM, Y.N. 1985. Clusiaceae: palynology and systematics. Trav. Sect. Sci. Tech. Inst. Fr. Pondicherry 21:1-80.

SEETHARAM, Y.N. and S.A.J. Pocock. 1978. Taxonomy and pollen morphology of Poeciloneuron (Guttiferae). Bull. Jard.

Bot. Nat. Belg. 48:359-365.

WCMC. 1998. Poeciloneuron pauciflorum. In: IUCN 2006. 2006 IUCN Red list of threatened species. www.iucnredlist.

org. Downloaded on 17 March 2007.

£+sha D o ID L

134 Journal of t titute of Texas 1(1)

BOOK REVIEWS

Bos Press (Text) and CAROL MERRYMAN (Art). 2006. Trees: Collins Wild Guide. (ISBN 0-00-719152-9, pbk.).

HarperCollinsPublishers, Ltd., 77-85 Fulham Palace Road, London, W6 8JB, UK (Orders: Collins

UK, Distributed by Trafalgar Square, No. Pomfret, VT 05053, U.S.A., www.trafalgarsquarebooks.com).

$16.00, 191 pp., color photographs, drawings, 4" x 6%".

This small book, with its illustrations, color photographs, and pertinent information, is an enormous help to anyone wishing to learn

how to identify trees most commonly found in Britain and Ireland.

The book has a brief history of trees in Britain and Ireland. It also includes, for those unfamiliar with botanical language, a short

helpful glossary.

Every page of Trees has a color photograph of a species with its scientific and common names. Accompanying each photograph is

an ID Fact File, which provides detailed descriptions of the tree, making it a valuable took for fast and accurate identification.

A compact range of general information covering specific characteristics such as the tree’s height, crown, and pattern of branch-

ing is very helpful. The leaves, their size, shape, and texture along with fruit and flowers produced by the tree are also emphasized as

important aids to correct identification

The size of the book is a definite plus. It is small enough to fit in a picket, purse, or glove compartment, age appropriate for an

adult or older child. It is a wonderful *carry along" book that imparts succinct information that both informs and educates without

overwhelming the reader.

It can be safely stated that the reader will take away from this book an increased knowledge of trees and an enhanced awareness

of how trees influence and enrich our environment.— Susan Kingeter, Volunteer, Botanical Research Institute of Texas, 509 Pecan Street, Fort

Worth, TX 76102-4060, U.S.A.

P. DaroLD BATZER and RegeccA R. SHaritz (eds.). 2007. Ecology of Freshwater and Estuarine Wetlands.

(ISBN 0-250-24777-9, hbk.). The University of California Press, Berkeley, CA 94704, U.S.A. (Orders:

California Princeton Fulfillment Services, 1445 Lower Ferry Road, Ewing, NJ 08618, U.S.A., www.

ucpress.edu, http://www.ucpress.edu/books/pages/10296.html, 609-883-1759, 609-883-7413 fax).

$59.95, 581 pp., color photos, b/w figures and photos, tables, graphs, 7" x 10".

Ecology of Freshwater and Estuarine Wetlands is designed to be a textbook and offers readers a comprehensive introduction to the ecology

of wetland systems. This well illustrated book has various chapters written by professionals whom focus on their areas of expertise.

This publication would be very suitable as a textbook for classes on wetland ecology and as a supplemental reading for exploring sys-

tems modeling or environmental engineering. Additionally, this title could also be a beneficial reference text for related classes, such as

hydrology and/or microbiology that may have a focus on the processes and uses of wetlands.

The book chapters flow in a logical order from one to another, building on previous chapters’ information. Chapter topics begin

with the nuts and bolts of the physical and biological definitions of wetlands followed by wetland geomorphology, hydrology, abiotic

constraints on flora and fauna, bacterial ecology, wetland plant communities, wetland ecosystem processes, U.S. wetland regulation

and policy, wetland restoration, flood pulsing and biodiversity in wetlands and wetlands in the global environment.

The text is very readable and is accompanied by many supportive illustrations, graphs and photos. The information presented to

readers is well balanced, providing various views and theories regarding wetlands function and definitions. Some of this text’s most

beneficial chapters include current research and views on bacterial ecology in wetlands as well as how these interactions control pro-

cesses and diversity within wetland systems. There is also a fantastic and witty chapter on wetland restoration in which the author offers

llowing reader to contemplate and learn from surprises and mistakes of others in dealing with wetland restoration.

many case studies

Chapter texts are tl ghly researched and the research studies presented and cited are all included in the extensive reference section.

Many of the chapter authors have included issues of concern or methodological steps in evaluating various wetland topics.

Ecology of Freshwater and Estuarine Wetlands, edited by Batzer and Sharitz, provides a comprehensive and timely introduction to wet-

lands ecology, including well written chapters on wetland formation processes including geomorphology, soils, hydrology and bacterial

ecology as well as legal policies and definitions. The text is full of supportive black and white illustrations, charts, tables and references

to research studies. Ecology of Freshwater and Estuarine Wetlands would serve as an excellent textbook for classes focusing on wetlands,

wetland ecology and related classes where system ecology and function may be of interest.—Lee Luckeydoo, Herbarium, Botanical Research

Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A.

J. Bot. Res. Inst. Texas 1(1): 134. 2007

NEW NAMES FOR BAMBOOS OF NEPAL (POACEAE: BAMBUSOIDEAE)

C.M.A. Stapleton

Royal Botanic Gardens, Kew

ichmond, Surrey

TW9 3AB, UK

chris_stapleton@onetel.com

ABSTRACT

The use of different generic and species concepts in Sino-Himalayan bamboos is discussed. Himalayacalamus planatus Stapleton

is separated from H. asper Stapleton, both species having now flowered. Two subspecies of Thamnocalamus spathiflorus (Trin.) Munro

are elevated to species, as T. nepalensis (Stapleton) Stapleton and T. occidentalis (Stapleton) Stapleton. Borinda chigar Stapleton is

transferred as Thamnocalamus chigar (Stapleton) Stapleton. The synonymy of two species from Nepal with those from neighboring

areas is discussed.

RESUMEN

Se discute el uso de diferent óri específicos en los bambúes chino-himalayos. Himalayacalamus planatus Stapleton

se separa de H. asper Stapleton, ER ied dm ambas especies. Dos subespecies de Thamnocalamus spathiflorus (Trin.) Munro se

elevan a especies, como T. nepalensis (Stapleton) Stapleton y T. occidentalis (Stapleton) Stapleton. Borinda chigar Stapleton se transfiere

como Thamnocalamus chigar (Stapleton) Stapleton. Se discute la sinonimia de dos especies de Nepal con las áreas próximas.

Many Sino-Himalayan bamboos are of economic or ecological importance in their natural range, but their

forest habitats are decreasing and their conservation status is of concern. Several are becoming more widely

cultivated in the US and Europe as ornamentals, which has allowed more study of their systematics and

identification. The bamboos of Nepal were first enumerated (Hara et al. 1980) according to available litera-

ture and earlier, tentative identifications in the national herbarium of Nepal, the British Museum (Natural

History), and institutions in Japan, to give a total of 10 species from 5 genera. They were studied further in

the 1980s (Stapleton 1982, 1987) and enumerated more comprehensively (Stapleton 1991, 1994a—c), to give

a total of 30 species from 11 genera, but 5696 of Himalayan bamboo taxa had no name at all at that time,

and a substantial number of new names were duly published (Stapleton 1994a-c).

At the same time, Chinese taxonomists were actively collecting and describing Sino-Himalayan bam-

boos, including those from Tibet, and a revised classification was being developed, with several new genera

(Keng 1982-84, 1987), as well as many new species, several collected in Tibet (Yi 1983, 19832). However,

a western generic classification of the grass family was also being produced (Clayton & Renvoize 1986),

and that pre-eminent global grass account applied a much broader generic concept to bamboos. Several

authorities consequently dismissed many of the new Chinese genera altogether, notably Chao and Renvoize

(1989), who relegated Pleioblastus, Oligostachyum, Bashania, Oreocalamus, Chimonocalamus, Yushania, Drepa-

nostachyum and Himalayacalamus on the grounds that they lead to confusion. Tewari (1993), Li (1997) and

Seethalakshmi and Kumar (1998) followed suit. They adopted instead a few, much larger genera, such as

Sinarundinaria Nakai. Most of the new Chinese genera were recognized by a few others (Soderstrom & El-

lis 1988; Majumdar 1989; Stapleton 1987, 1994b—c). This led to two very different approaches to bamboo

classification, especially for the subtropical to temperate clade of Asian and North American bamboos.

Along with a narrower generic concept, the new Chinese classification system (Keng 1982-84) also

applied a narrower species concept than that used in western grass taxonomy. This utilized many vegeta-

tive characteristics, without the emphasis given to floral characters usually applied in more traditionally

based treatments (Clayton & Renvoize 1986; Chao & Renvoize 1989). Keng’s classification instead followed

a different tradition (Munro 1868; Gamble 1896; Camus 1913) of using a narrower species concept in the

bamboos than in other grasses, making good use of several characters of the culm sheaths, which are well

J. Bot. Res. Inst. Texas 1(1): 135 — 142. 2007

136 Journal of the Botanical R h Institute of Texas 1(1)

differentiated in woody bamboos. Culm sheaths are usually simply referred to as culm leaves in other

grasses, where they are very similar to the foliage leaves. In this way Keng's treatment was also to conflict

with the much broader bamboo species concept applied by Chao and Renvoize (1989). In that treatment

for example, Drepanostachyum falcatum was considered to extend from Pakistan to Meghalaya, almost into

Bangladesh, and was called Sinarundinaria falcata. Himalayacalamus was treated essentially as a single spe-

cies, Thamnocalamus falconeri, extending the entire length of the Himalayas.

When Keng’s approach to bamboo classification was applied to the species found in Nepal, Sikkim and

Bhutan (Majumdar 1989; Stapleton 1994a—c), a large number of new combinations and new taxa were seen

to be required. Full use was made of vegetative characters, especially those of culm sheaths and the complex

bud and branching characters unique to woody bamboos (Stapleton 1991, 1994a—c). However, because of

the broader concepts applied in Clayton and Renvoize (1986) and Chao and Renvoize (1989), some caution

was applied in the description of new species without knowledge of floral characteristics, for example in

Himalayacalamus asper (Stapleton 1994c), in which bamboos with rather different culm sheaths from c &

w Nepal were combined. In the few cases where comprehensive floral material was available, a tentative,

somewhat broader species concept was applied, for example in Thamnocalamus spathiflorus, in which several

subspecies and varieties were recognized, rather than distinct species (Stapleton 1994b).

Recent molecular investigations (Ní Chonghaile 2002; Guo et al. 2001, 2002) have shown no support

at all for the larger genera, which appear to be polyphyletic. There has also been no support for emphasising

floral characters over vegetative ones. The major groupings within the woody bamboos, at supertribal, tribal,

or subtribal level, based mainly on differences in floral morphology, were seen to have no support whatsoever

from molecular data (Ni Chonghaile 2002). As bamboos have more vegetative characters by which they

can differ than other grasses, and there seems no reason why the species concepts applied to other grasses

should be forced onto bamboos artificially, it would appear that the recognition of genera and species on the

grounds of consistent differences in vegetative characters is now justified. Keng’s classification system has

gradually become accepted more widely around the world, and its application in the Chinese and English

language Flora of China bamboo accounts (Keng & Wang 1996; Li et al. 2006) has increased its credibility.

The recent recognition of 3 bamboo species native to the US, rather than 1, separated largely on vegetative

characters including branching (Triplett et al. 2006), is further evidence of the trend to recognize smaller

taxa with more emphasis on vegetative characters.

In consequence, it would appear that the classification system followed earlier (Majumdar 1989, Stapleton

1994a-c) is acceptable, while that applied elsewhere (Chao & Renvoize 1989; Tewari 1993; Seethalakshmi

& Kumar 1998) is unnatural and paraphyletic. Building on this support for the smaller taxa previously

established, it is realised that a few alterations are required to the names applied to the bamboos of Nepal

(Stapleton 1994a-c). In addition, further collections have since been made within Nepal and adjoining

areas, and several species have been introduced into western horticulture, allowing them to become better

known, especially as some have recently flowered. Unfortunately, for various reasons, less new botanical

fieldwork has been undertaken in Nepal than could have been hoped for, and many gaps in our knowledge

still remain. Several additional species have been recorded for Nepal (Poudyal 2006), but most are only

tentatively identified and others represent fairly recent introductions. Hopefully the return of peace to that

country, and the aspiration of compiling a Flora of Nepal account, will no doubt allow our knowledge of

Nepalese bamboos to continue to develop further.

Himalayacalamus was published as a monophyletic genus, but several new species from Nepal were later

added, and species described from Tibet and Sikkim in other genera have also been included. In broader ge-

neric treatments it was treated as a synonym of Thamnocalamus (Clayton & Renvoize 1986; Chao & Renvoize

1989) on the basis of its compressed inflorescences, but Himalayacalamus can be distinguished by its usually

solitary florets, and by reduced sheaths on inflorescence and culm branches (Stapleton 1994c). Molecular

data (Ni Chonghaile 2002) suggests that Himalayacalamus is more closely related to Drepanostachyum, from

which it differs in its fewer branches, adaxially glabrous culm sheaths, and more compressed inflorescences

with dense spikelets of fewer, usually solitary florets. The stalk of the spikelet, incorrectly termed a pedicel

Stapleton, New names for bamboos of Nepal 137

in grasses but actually a peduncle, is short in Himalayacalamus and Thamnocalamus, which has suggested

its homology with the vegetative promontory supporting culm branches (Stapleton 1997), hence use of the

term promontory as an alternative to pedicel below. Himalayacalamus and Thamnocalamus have been rede-

scribed to reflect current circumscription and terminology in Stapleton 1994b, 1994c, 2000, and in Li et

al. 2006.

Himalayacalamus planatus Stapleton, sp. nov. (Fig. 1). Tver: NEPAL. Rasuwa District: Syabru (ca. 28°12’N 85°28E), elev.

ca. 8,000 ft, 7 Oct 1984, Stapleton 328 (HOLOTYPE: K!).

Himalayacalamus asper mihi affinis, sed vaginis culmorum pilosis non asperis, auriculis et setulis oribus vaginorum foliorum absens,

nodis culmorum planatis, lemmatibus glabris differt.

Clumps dense. Culms to 2-5 m, 0.5-1.5 cm in diam., nodding to pendulous; internodes to 20 cm, surface

with little wax, soon becoming glossy, smooth with no ridges, initially with purple ring above nodes and

streaks elsewhere, becoming burgundy-red to brown if exposed, walls to 4 mm thick; nodes level, scarcely

raised, sheath scar thin, supranodal ridge absent or slightly raised; mid-culm branches 7-20, central branch

to 0.15 cm in diam., aerial roots absent. Culm sheaths quickly deciduous on early shoots, height to ligule

ca. 18 cm, similar to internodes in length, attenuating convexly in distal 1/3 to ca. 0.2 cm, basally tough

and smooth with membranous recurved margins, distally thinner and shortly hispid, distal 1/3 of both

edges densely ca. 0.1 cm white-ciliate; auricles absent; oral setae absent; ligule ca. 0.6 cm wide x 0.3 cm tall,

exterior very shortly pubescent, interior glabrous, margin serrate; blade reflexed, to 4 x 0.2 cm, proximally

scabrous, deciduous to persistent. Leaf sheath surface and edges glabrous, or overlapping edge distally

short-ciliate; auricles absent; oral setae absent; ligule short, to 0.1 cm, rounded, densely puberulous; exter-

nal ligule not pronounced, glabrous. Leaf blade to 13 x 1 cm, glabrous; petiole glabrous, often pigmented;

2ndary veins 2-3 each side; transverse veins not evident. Synflorescences fasciculated in spicate clusters of

racemes. Spikelets on ca. 2-5 mm promontories (pedicels) with 1(22) florets and a tiny «0.2 mm rudiment

ona ca. 4 mm rhachilla extension. Glumes membranous, pale, glabrous, apical ca. 0.7 mm of margins with

cilia to ca. 0.2 mm. Fertile lemma 7-9 mm, distally glabrous, not scabrous, green or purple-tinged, apical

ca. 0.5 mm with cilia to ca. 0.2 mm. Palea glabrous, keels smooth, apex blunt or very shortly bifid to ca. 0.2

mm with tuft of ca. 0.2 mm hairs. Rhachilla basally to 0.3 mm-lanate along with lemma base, proximally

scabrous, distally glabrous.

Distribution and Ecology.—This species is only known from Rasuwa District in Nepal, where it grows

in mixed temperate forest. It is also in horticultural cultivation in the UK, France and the US.

Etymology.—The epithet is derived from the level culm nodes, which are scarcely raised and have a thin

persistent culm sheath base.

Ethnobotany.—The culms are split and woven into a variety of baskets, trays and mats. Shoots are edible,

and leaves are palatable for livestock, but small. Local name is malinge nigalo (Nepali). Extensively collected

from forest areas.

Additional collections: NEPAL. Kathmandu Valley: Chalnakhel (cult.), 22 Jan 1991, Stapleton 918 (K). UNITED KINGDOM. Devon:

Dartmouth, 27 May 1997, Stapleton 1120 (K). UK (cult.). Sussex: Leigh, 9 Nov 1998, Pike s.n. (K).

Two similar bamboo species in the genus Himalayacalamus, both lacking the smooth and glabrous culm

sheaths normal for that genus were initially collected, without flowers. One was from the Seti Khola valley in

Kaski District, w Nepal in 1983, the other from near Syabru in the lower Langtang valley of Rasuwa District,

c Nepal in 1984. In the first enumeration of bamboos from Nepal (Stapleton 1991) these were presented as

2 separate species, but in the published account (Stapleton 1994c) they were conservatively combined into

one species, H. asper Stapleton, the type being the 1983 Kaski collection. Both these bamboos are now in

cultivation in the west, and both have now flowered. Better knowledge of their vegetative and floral charac-

teristics, along with a greater degree of confidence in the application of a narrower species concept, requires

a new name to be published for the species from the Langtang valley. It was introduced from the Syabru

area in 1979 by Merlyn Edwards, and grown at Kew under the misapplied name Arundinaria microphylla,

then in the US under the name Neomicrocalamus microphyllus. After this misidentification was discovered

£s+haD

138 Journal of titute of Texas 1(1)

Fic. 1. Himalayacalamus planatus with pilose culm sheath, no leaf sheath auricles or oral setae, level nodes and glabrous racemes of 1-flowered

spikelets.

Stapleton, New names for bamboos of Nepal

all sheaths present E.

2-keeled prophyll

139

I be L : L l h thy

* ££.

Fic. 2. TI g, flattened branchlet sides, sulcate culm,

leaf sheath ligules.

£s+haD o ID L

140 Journal of t titute of Texas 1(1)

(Stapleton 1999), it became known in cultivation as Himalayacalamus asper, a name that is now unfortunately

also misapplied. It has flowered in the UK, as has a plant of the real Himalayacalamus asper from Gorapani,

Kaski District, collected by Muriel Crouzet, and grown in France. The less bifid palea and shorter apical cilia

on glumes, lemma and palea distinguish H. planatus from H. asper, which, in keeping with its epithet, has

minutely scabrous lemmas and apically scabrous palea keels, as well as short, hard, bulbous-based spines

on its culm sheaths. A more recent introduction of H. asper by Jean Merret, also from Kaski District of w

Nepal, has been described as Drepanostachyum merretii by Demoly (2006). The seedlings were initially hard

to identify, although they were clearly not the same as the cultivated H. asper. Now that they have grown

larger their true identity, as the only real H. asper in cultivation, has been revealed.

Thamnocalamus chigar (Stapleton) Stapleton, comb. nov. Borinda chigar Stapleton, Edinburgh J. Bot., 51:286. 1994. Type:

NEPAL. Kaski District: Karuwa to Pipar (ca. 28°24’N 83?58'E), elev. ca. 3,000 m, 16 Nov 1983, Stapleton 315 (HoLotypE: E).

When first collected in 1983, the generic status of this species was very uncertain. As the importance of

branching and buds was not appreciated at that time, the material collected did not allow these characters

to be properly assessed. New collections of this species with better branching have since been made, and

they have revealed that it is not a species of Borinda as at first thought. It seems instead to be a rather distinct

species of Thamnocalamus. Figure 2 shows the sheath scars on the branch complement, which are consistent

with Thamnocalamus rather than Borinda (Stapleton 1994b: Fig. 1, pattern a rather than pattern b). There

is a full complement of broad enclosing sheaths, initiated by a 2-keeled prophyll. In addition, the strong

flattening on one side of the branchlets with substantial sulcation on a small culm, further characteristics

of Thamnocalamus, can clearly be seen (Fig. 2).

The length of the ligules and the long, delicate culm sheaths without a well-distinguished blade are

remarkable and obscure the affinity to better-known species of Thamnocalamus, which is however revealed

in its branch complement structure. Its flowers are still not known. Partially because of the presence of

this distinct species, and partially because of consistent vegetative differences between the taxa previously

described as subspecies, they are elevated here to species, even though it is very hard to separate them by

their flowers alone with any certainty.

Additional collection: NEPAL. Kaski District: Modi Khola, Deurali, elev. ca. 3000 m, Nov 1994, M. Edwards 206 (K).

Thamnocalamus nepalensis (Stapleton) Stapleton, stat. nov. Thamnocalamus spathiflorus Munro subsp. nepalensis Staple-

ton, Edinburgh J. Bot. 51:283. 1994.

This subspecies was distinguished from the type by its glabrous culm sheaths and leaf sheaths without oral

setae. These characters are now considered to be of importance at the species level, justifying elevation to

specific rank.

Thamnocalamus occidentalis (Stapleton) Stapleton, stat. NOV. Thamnocalamus spathiflorus Munro subsp. occidentalis

Stapleton, Edinburgh J. Bot. 51:283. 1994.

This subspecies was distinguished from the type by its glabrous but asymmetrical culm sheaths, also with

auricles and oral setae. These characters are now considered to be of importance at the species level, justifying

elevation to specific rank. This is supported by the geographic disparity between this species, from the nw

Himalayas, and T. spathiflorus from the e Himalayas. T. nepalensis, T. chigar, and T. crassinodus, all from c Nepal

are found between the two species. Although not yet collected there, it is likely to occur in w Nepal.

Bambusa jaintiana R.B. Majumdar

Bambusa alamii Stapleton

Bambusa alamii Stapleton has been considered a synonym of B. jaintiana R.B. Majumdar (Alam & Has-

san 1994). B. jaintiana was minimally diagnosed on the basis of a type collection from the Khasia Hills of

Meghalaya. The type has not been seen, but an isoparatype of B. jaintiana at K seems identical to B. tulda.

However, having now visited the Khasia Hills and the type locality for B. jaintiana, 1 am satisfied that a

Stapleton, New names for bamboos of Nepal 141

species growing there is the same as B. alamii, and it is assumed that they are synonymous. There is still a

possibility, however, that they were introduced from s China or Indochina, where several similar species of

"Weavers' Bamboo' are cultivated.

Himalayacalamus gyirongensis (T.P. Yi) Ohrnberger & Himalayacalamus porcatus Stapleton

Fargesia gyirongensis T.P. Yi was described from a collection made in Gyirong Xian, Tibet at 2400 m, which

suggests a location possibly less than 10 km north of the Nepalese border at Rasuwa Garhi, and possibly

as close as 25 km from the type locality of the later H. porcatus Stapleton, from near Syabru at 2300 m. The

description of F. gyirongensis did not mention porcate culms, and gave the culm sheath as glabrous or setose,

the persistent base initially densely setose. The culm sheath of H. porcatus is completely glabrous, although

the persistent base is lightly tomentose at first. H. porcatus also has distinct oral setae on symmetrical culm

sheaths, while F. gyirongensis was described as having no oral setae, and slightly asymmetrical culm sheaths

were illustrated. A suspicion still remains, however, that these two species could be the same, but without

inspection of the type specimen of F. gyirongensis and fieldwork in Tibet, this cannot be tested properly.

ACKNOWLEDGMENTS

Merlyn Edwards is thanked for collecting better material of T. chigar in Nepal. Muriel Crouzet and Jean Mer-

ret are thanked for collecting H. asper, and Robert Kernin is thanked for sending a sample of Jean Merret's

collection. The Keeper of the Herbarium at the Royal Botanic Gardens Kew is thanked for providing working

facilities. Two anonymous reviewers are also thanked.

REFERENCES

ALAM, M.K. and M.A. Hassan. 1994. Occurrence of Bambusa burmanica, B. cachariensis, B. jaintiana and Thyrsostachys

oliveri in Bangladesh. Bangladesh J. Pl. Tax. 1(2):21-32.

CAMUS, E.G. 1913. Les Bambusées. Paris, Lechevalier.

Chao, C.S. and S.A. Renvoize. 1989. A revision of the species described under Arundinaria (Gramineae) in Southeast

Asia and Africa. Kew Bull. 44:349-367.

CLAYTON, W.D. and S.A. Renvoize. 1986. Genera graminum: grasses of the World. Royal Botanic Gardens, Kew.

Demo y, J.-P. 2006. Une nouvelle espèce de bambou (Poacées) dédiée à son découvreur et introducteur, Jean

Merret (Drepanostachyum merretii Dem., sp. nov.). Bull. Assoc. Parcs Bot. France 41:33.

GAMBLE, J.S. 1896. The Bambuseae of British India. Ann. Roy. Bot. Gard. (Calcutta) 7(1):1-133.

Guo, Z.H., Y. Y. CHEN, D.Z. Li, and J.B. Yana. 2001. Genetic variation and evolution of the Alpine bamboos (Poaceae:

Bambusoideae) using DNA sequence data. J. Plant Res. 114:315-322.

Guo, Z.H., Y.Y. Chen, and D.Z. Li. 2002. Phylogenetic studies on the Thamnocalamus group and its allies (Gramineae:

Bambusoideae) based on ITS sequence data. Molec. Phylogen. Evol. 22: 20-30.

Hara H., W.T. STEARN, and L.H. WiLLiAMS. 1980. An enumeration of the flowering plants of Nepal. British Museum,

London.

Keng, P.C. 1982-4. A revision of genera of bamboos from the world l-V. J. Bamboo Res. 1(1):1-19; 1(2):31-46;

2(1):11-27; 2(2):137-153; 3(1):22-42.

Kena, P.C. 1987. A systematic key to the tribes and genera of subfamily Bambusoideae (Gramineae) occurring in

China and its neighbours. J. Bamboo Res. 6(3):13-28.

KENG, P.C. and Z.P. Wane (eds.). 1996. Gramineae (Poaceae), Bambusoideae, Flora Reipublicae Popularis Sinicae

9(1). (Chinese).

Li, D.Z. 1997. The flora of China Bambusoideae Project: Problems and current understanding of bamboo taxonomy

in China. In: Chapman, G.P, ed. The Bamboos. Academic Press. Pp. 61-81.

Li, D.Z., Z.P. Wane, Z.D. Zhu, N.H. Xia, L.Z. Jia, Z.H. Guo, G.Y. YANG, and C.M.A. STAPLETON. 2006. In: Wu, Z.Y., PH. Raven,

and D.Y. Hong, eds. Bambuseae. In: Flora of China, 22. Poaceae: 7-180. Science Press, Beijing & Missouri Bo-

tanical Garden Press, St. Louis.

142 Journal of the Botanical R h Institute of Texas 1(1)

Masumoar, R.B. 1989. In: Kartikeyan, S. et al. Flora Indicae, enumeratio monocotyledonae. Botanical Survey of

India, Howrah, Calcutta. Pp. 274-283.

Munro, W. 1868. A monograph of the Bambusaceae. Trans. Linn. Soc. London 26:1-157.

Ni CHONGHAILE, G. 2002. Molecular systematics of the woody bamboos (Tribe Bambuseae). Ph.D. Thesis, University

of Dublin, Trinity College, Ireland.

PoubvaL, PP. 2006. Bamboos of Sikkim (India), Bhutan, and Nepal. New Hira Books Enterprises, Kathmandu.

SEETHALAKSHMI, K.K. and M.S MuktesH Kumar. 1998. Bamboos of India: a compendium. Kerala Forest Research Insti-

tute & INBAR.

SobtERSTROM, T.R. and R.P. Eus. 1988. The position of bamboo genera and allies in a system of grass classification.

In: Soderstrom, T.R. et al., eds. Grass systematics and evolution. Smithsonian Institution Press. Pp. 225-238.

STAPLETON, C.M.A. 1982. Bamboos in Koshi Zone, East Nepal. Nepal Forestry Tech. Info. Bull. 6:12-14.

STAPLETON, C.M.A. 1987. Bamboos. In: J.K. Jackson, Manual of Afforestation in Nepal: 199-214. Forestry Research

Project, Kathmandu.

STAPLETON, C.M.A. 1991. A morphological investigation of some Himalayan bamboos with an enumeration of

taxa in Nepal & Bhutan. Ph.D. thesis, University of Aberdeen.

STAPLETON, C.M.A. 1994a. The bamboos of Nepal and Bhutan Part I: Edinburgh J. Bot. 51:1-32.

STAPLETON, C.M.A. 1994b. The bamboos of Nepal and Bhutan Part Il: Edinburgh J. Bot. 51:275-295.

STAPLETON, C.M.A. 1994c. The bamboos of Nepal and Bhutan Part Ill: Edinburgh J. Bot. 51:301-330.

STAPLETON, C.M.A. 1997. Morphology of woody bamboos. In Chapman, G.P, ed. The bamboos: 251-267. Academic

Press.

STAPLETON, C.M.A. 1999. Sino-Himalayan bamboos on the US West Coast. Amer. Bamboo Soc. Newsl. 20(6):1—6.

STAPLETON, C.M.A. 2000. Bambuseae. In: H.J. Noltie, The grasses of Bhutan Flora of Bhutan 3(2):482-515. Royal

Botanic Garden Edinburgh & Royal Government of Bhutan.

Tewari, D.N. 1993. A monograph on bamboo. International Book Distributors, Dehra Dun, India.

Triplett, J.K., A.S. WeaxLey, and L.G. Clark. 2006. Hill cane (Arundinaria appalachiana), a new species of bamboo

(Poaceae: Bambusoideae) from the Southern Appalachian mountains. Sida 22:79-95.

Yi, T.P. 1983. New taxa of Bambusoideae from Xizang (Tibet), China. J. Bamboo Res. 2(1):28-46.

Yi, T.P. 1983a. New species of Fargesia Franchet and Yushania Keng f. from Tibet. J. Bamboo Res. 2(2):18-52.

Yi, T.P. 1988. A study on the genus Fargesia Franch. from China. J. Bamboo Res. 7(2):1-119.

FOLIA TAXONOMICA E

VALIDATION OF TWO: TAXA PROM NORTHERN SOUTH AMERICA

Christian Feuillet

Department of Systematic Botany, MRC-166

Smithsonian Institution

PO. Box 3712

Washington, DC 20013-7012, U.S.A.

feuillec@si.edu

ABSTRACT

Two taxa, one species of Aristolochia (Aristolochiaceae) and one variety of Passiflora (Passifloraceae), are validated. First, Aristolochia

peltato-deltoidea, described by Hoehne in 1942, was originally published without a Latin diagnosis and therefore the name is invalid.

The species is described herein as A. kanukuensis. Second, Killip invalidly placed var. orinocensis as a variety of Passiflora foetida L. in

1938, as he did not mention that he had described it in 1930 at the rank of subspecies. Thus, Passiflora foetida var. orinocensis is

validated here by acknowledging the new rank and providing a full citation of the basionym.

RESUME

Deux taxa, une espéce d'Aristolochia (Aristolochiaceae) et une variété de Passiflora (Passifloraceae), sont validées. Premiérement, Aristo-

lochia peltato-deltoidea, décrite par Hoehne en 1942, a été publiée sans diagnose latine, le nom est donc invalide. Lespéce est décrite ici

sous le nom A. kanukuensis. Deuxiémement, Killip en 1938 a placé var. orinocensis parmi les nombreuses variétés de Passiflora foetida

L., de facon invalide car il ma pas signalé qu'il l'avait décrite en 1930 au rang de sous-espèce. Passiflora foetida var. orinocensis est

validée par la reconnaissance du nouveau rang et la référence précise au basionyme.

During the reviewing phase of the “Checklist of the Plants of the Guiana shield" (Funk et al. 2007), some invalid

names where found. This short paper addresses two of the names in use that need validation.

ARISTOLOCHIA KANUKUENSIS (ARISTOLO CHIACEAE)

Aristolochia peltato-deltoidea Hoehne (1942) was published with full Portuguese description and an illustra-

tion, but without a Latin diagnosis. Since the “Cambridge Rules” (Rendle 1935; Art. 38), a Latin diagnosis

was mandatory and it remains so including in the last “ICBN” Art. 36 (McNeill et al. 2006), therefore the

name was not validly published. When writing the treatment of the Aristolochiaceae for Flora of the Guianas

(Feuillet & Poncy 1997 & 1998), the fact that the name A. peltato-deltoidea was invalidly published was

overlooked. A name is needed for this species.

Aristolochia kanukuensis Feuillet, Sp. nov. Tyre: GUYANA. UPPER TAKUTU-UPPER EssEQUIBO: NW slopes of the Kanuku Mountains

in drainage of Moku-Moku Cr. (Takutu tributary), dense forest, on exposed rocky ledges, 150-400 m, 31 Mar-16 Apr 1938, A.C.

Smith 3385 (HoLoTYPE: US; IsoTYPES: E, G, K, MO, NY, P U)

= Aristolochia peltato-deltoidea Hoehne, nom. inval. (no Latin diagnosis), Flora Brasilica vol. XV, 2: 102-103; tab. 75. 1942.

Aristolochia kanukuensis a speciebus guianensibus folio deltato et basi peltato, perianthio glabro et flavovirenti, limbo perianthii fusco

differt.

The validating description and a key in Portuguese are given under “79. Aristolochia peltato-deltoidea” in

Hoehne (Flora Brasilica 15(2):102-103. 1942).

The epithet of the new species is derived from the name of the region where all the studied specimens

have been collected, the Kanuku Mountains of southern Guyana. Kanuku is a name of Amerindian origin

meaning “rich forest” in the Macushi language.

Other material studied: GUYANA. Upper Takutu-Upper Essequibo: S Kanuku Mountains, Maas et al. 4058 (U); Kanuku Mountains,

Nappi Cr., 130 m, 4 Nov 1987, Jansen-Jacobs et al. 705 (BRG, P, U, US, US); Kanuku Mountains, Foothills at Moco-Moco Riv., Jansen-Jacobs

et al. 4592 (BRG, P, U); SE Kanuku Mountains, Makawatta Massif, 750 m, 31 May 1996, D. Clarke & T. MacPherson 1855 (US).

J. Bot. Res. Inst. Texas 1(1): 143 — 144. 2007

£+sha D o ID

144 Journal of t h Institute of Texas 1(1)

PASSIFLORA FOETIDA VAR. ORINOCENSIS (PASSIFLORACEAE)

Passiflora foetida subsp. orinocensis Killip (1930) was included in the classic “The American Species of Pas-

sifloraceae" (Killip 1938) at the rank of variety as P. foetida var. orinocensis Killip 1930, without acknowledg-

ment of a change in rank. Therefore, the variety is not validly published. The infraspecific name is still in

use (Feuillet 1989; Tillett 2003) as a variety, but not as a subspecies.

Passiflora foetida var. orinocensis (Killip) Feuillet, stat. nov. Basiouvw: Passiflora subsp. orinocensis Killip, Gentes Herb.

2:205, fig. 107. 1930. Type: VENEZUELA. Bouvar: vicinity of Ciudad Bolivar, Isla Degrero, in the Orinoco River, about 200 ft, 6 Mar

1921, L.H. Bailey & E.Z. Bailey 1773 (HoLoTYPE: US; Isotype: NY).

In Passiflora sect. Dysosmia DC., the 20 species and ca. 30 varieties tend to produce natural hybrids when

in contact, and the taxonomy is confused. The fruits of these species range from green, yellow, to red. Red

fruits seem to represent an acquired character state and P. foetida var. orinocensis shares it with P. ciliata Ait.,

some varieties of P. foetida L., and several other species in the section from Central America, Mexico, and the

West Indies. Further molecular research on this group, which has so far been poorly sampled, may confirm

the position of var. orinocensis in P. foetida or place it in or near P. ciliata.

Other material studied: VENEZUELA. Bolivar: Bank of the lower Orinoco River, Chaffanjon 233 (P, US); Rusby & Squires 179 (K, MIN,

NY); Orinoco River, Caicara, 95 m, 12 Jun 1940, LI. Williams 13283 (US). COLOMBIA. Los Llanos: Río Meta, Curazao, 25 Oct 1938,

J. Cuatrecasas 4094 (US).

According to Tillett (2003), Passiflora foetida var. orinocensis has also been collected in Venezuela in the ter-

ritory Amazonas and the states of Apura and Barinas.

ACKNOWLEDGMENTS

I would like to thank the curators and directors of the herbaria BRG, CAY, F, G, K, MIN, MO, P, NY, B, U,

US whose collections have been made available for my work on Aristolochia and Passiflora, and used when

writing this paper. I am grateful to reviewers Shawn Krosnick, John MacDougal, and Kerry A. Barringer

who helped make this paper better.

REFERENCES

FeuiLLET, C. 1989. Diversity and distribution of Guianan Passifloraceae. In: L.B. Holm-Nielsen, I.C. Nielsen and H

Balslev, eds. Tropical forests: botanical dynamics, speciation and diversity. London. Pp. 311-318.

FeuiLLET, C. and O. Poncy. 1997. Aristolochiaceae. In: J. Boggan, V. Funk, C. Kelloff, M. Hoff, G. Cremers and C. Feuillet,

eds. Checklist of the plants of the Guianas (Guyana, Surinam, French Guiana). P. 57.

FeuiLLET, C. and O. Poncy. 1998. Aristolochiaceae. In: A.R.A. Górts-van Rijn and M.J. Jansen-Jacobs, eds. Flora of the

Guianas. Pp. 1-23.

Funk, V. T. HALLOWELL, P. Berry, C. KELLOFF, and S. ALEXANDER, 2007. Checklist of the plants of the Guiana shield (Guyana,

Surinam, French Guiana, Venezuela: Amazonas, Bolivar, Delta Amacuro). Natural History Museum, Washington.

Pp. 1—558.

HOEHNE, F.C. 1942. Flora Brasilica vol. XV, 2: 102-103; tab. 75.

Kup, E.P. 1930. Passiflora foetida subsp. orinocensis, Killip, subsp. nov. Gentes Herb. 2:205-206.

Kup, E.P. 1938. The American species of Passifloraceae. Publ. Field Mus. Nat. Hist., Bot. Ser. 19:1-613.

McNeiLt, J., ER. Barrie, H.M. Bumper, V. DemouLin, D.L. HAWKsworTH, K. MARHOLD, D.H. NicoLson, J. PRADO, P.C. SiLva, J.E. SKOG,

J.H. Wiersema, and N.J. TURLAND (eds.). 2006. International code of botanical nomenclature. Gantner V. Regnum

Veg. 146:1-568.

RENDLE, A.B. 1935. International rules of botanical nomenclature. Taylor & Francis, London. Pp. 1-29.

TiLLETT, S.S. 2003. Passifloraceae. In: PE. Berry, K. Yatskievych, and B.K. Holst, eds. Fl. Ven. Guayana, vol. 7:625-667.

NOTES ON THE DISARTICULATION OF A MEOTE AMIA

(ASTERACEAE, ASTEREAE)

Guy L. Nesom

Botanical Research Institute of Texas

509 Pecan Street

Fort Worth, Texas 76102-4060, U.S.A.

ABSTRACT

The genus Xvlothamia primarily of the Chil ]

d Sonoran deserts in northern Mexico, was originally described with nine species

Molecular evidence by Urbatsch et al. has subsequently shown that these species form two separate clades. Among species in each of

the two groups, details of phylogenetic position vary, depending on optimality criteria used in the analysis. Four species of Xylothamia,

including the type, are most closely related to the Caribbean genus Gundlachia and were transferred to Gundlachia by Urbatsch and Rob-

erts. These four, however, can be interpreted as having a sister relationship with Gundlachia and are here maintained within Xylothamia.

The remaining five species of Xylothamia are part of a clade that includes Amphiachyris, Bigelowia, Euthamia, Gutierrezia, Gymnosperma,

and Thurovia. Molecular evidence indicates that two of the five species have a sister relationship but that neither this pair nor any of the

other three are unambiguously closely related to any established genus. Segregates were proposed for these five species by Urbatsch and

Roberts as the ditypic genus Neonesomia and the monotypic genera Chihuahuana, Medranoa, and Xylovirgata. In contrast, morphological

features and M Ms proximity within the Chihuahuan Desert justify congeneric treatment id these five EX nd are here

united in 1 (with Chihuahuana, Neonesomia, and A lin synonymy). New

(G.L. Nesom) G.L. Nesom, comb. nov., Medranoa palmeri (A. Gia) G.L. Nesom, comb. nov., Medranoa purpusii (Brandes) G.L.

Nesom, comb. nov., and Medranoa pseudobaccharis (S.F. Blake) G.L. Nesom, comb. nov.

RESUMEN

listril primaria en los desiertos de Chihuahua y Sonora del norte de México, se describió originariamente

El género Xylothamia, de

con nueve especies. Los datos moleculares de Urbatsch et al. han mostrado que estas especies forman dos clados separados. Entre las

especies de cada uno de los grupos, varían los detalles de posición filogenética BAD EN de los criterios de optimización usados en

el análisis. Cuatr ] d el género caribeño Gundlachia

de Xylothamia, que incluyen el tipo, están

y fueron transferidas a Gundlachia por Urbatsch y Roberts. Estas cuatro, sin ee TBO, p MOL NN como el E hermano de

JI T de

Gundlachia y ] Xvlothamia I

ncl Amphiachyris

E L

Bigelowia, Euthamia, Gutierrezia, Gymnosperma, y Thurovia. Las pruebas moleculares indican que dos de estas cinco especies tienen una

relación de grupo hermano pero que ni este par ni ninguna de las otras tres están fuertemente relacionadas con ningún otro género

establecido. Se han propuesto segregaciones de estas cinco especies por Urbatsch y Roberts como género ditípico Neonesomia y los gé-

neros monotípicos Chihuahuana, Medranoa, and Xylovirgata. En contraste, las características morfológicas y la proximidad geográfica en

el desierto de Chihuahua justifica el tratamiento congenérico de est pecies, dque se unen aquí en Medranoa (con Chihuahuana,

Neonesomia, y Xylovirgata colocad la sinonimia). Las nuevas combinaciones son Medranoa johnstonii (G.L. Nesom) G.L. Nesom,

comb. nov., Medranoa palmeri (A. Gray) G.L. Nesom, comb. nov., Medranoa purpusii (Brandeg.) G.L. Nesom, comb. nov., y Medranoa

pseudobaccharis (S.F. Blake) G.L. Nesom, comb. nov.

The genus Xylothamia was proposed to include eight species (Nesom et al. 1990) traditionally associated with

Ericameria. A ninth was added soon after (Nesom 1992). Except for Xylothamia diffusa, which occurs in Sonora,

Baja California, and Baja California Sur, all are species of the Chihuahuan Desert. Molecular evidence by

Urbatsch et al. (2003) subsequently indicated that the nine species of Xylothamia form two separate clades.

Four species, including the type, are most closely related to the Caribbean genus Gundlachia (sensu Lane

1996). The remaining five species are part of a clade that includes Amphiachyris, Bigelowia, Euthamia, Gutier-

rezia, Gymnosperma, and Thurovia. Gundlachia and its related Xylothamia species are sister to the other group,

and this larger clade is essentially what has been termed the “Gutierrezia group" (e.g., Nesom 2000).

Parsimony analyses by Urbatsch et al. were based on combined data sets of the external transcribed

spacer (ETS) and internal transcribed spacer (ITS) DNA sequences, both with and without indels. A mor-

phological data set including ten characters was added in some of the analyses. The biphyletic nature of

J. Bot. Res. Inst. Texas 1(1): 145 — 148. 2007

146 Journal of the Botanical R h Institute of Texas 1(1)

Xylothamia is clear, but among species in each of the two lineages, details of phylogenetic positions vary,

depending on which optimality criterion was used in the analysis.

Taxonomy proposed by Urbatsch and Roberts (2004) merged the four species of Xylothamia sensu stricto

with Gundlachia. The other five Xylothamia species were apportioned into four new genera. The comments

below propose a taxonomic alternative for the species of these two groups and the taxonomic summary

shows how the original nine species of Xylothamia will be treated in the Astereae of Mexico (Nesom, expected

2007).

Expanded Gundlachia

The four species of Xylothamia sensu stricto and Gundlachia are shown by Urbatsch et al. (2003) as sister

groups in analyses including indels (Figs. 1A, a ratchet analysis, and 1B, a heuristic analysis), in a Bayesian

analysis (Fig. 2), and in a heuristic analysis including indel data and the morphological character matrix

(Fig. 3, right side). Xylothamia riskindii, the other three Xylothamia species, and Gundlachia are shown as an

unresolved trifurcation in a bootstrap analysis including indel data and a morphological character matrix

(Fig. 3, left side). The four Xylothamia species are shown as a basal grade to Gundlachia in a tree derived from

a ratchet analysis of the molecular data excluding indels (Urbatsch et al. 2003, Fig. 4). The topology shown

by Urbatsch and Roberts (2004, Fig. 1) was based on the earlier-published Figure 4.

A close relationship of Caribbean Gundlachia to Mexican Xylothamia sensu stricto is supported by the

molecular analyses, but the topology of the relationship is not resolved. Morphology of Caribbean Gundlachia

is distinctive and relatively consistent among the taxa, and the geographical and morphological contrasts

(noted below) with Xylothamia provide a rationale for recognition of these two species groups as separate

genera. Gundlachia is paraphyletic without Xylothamia in only one of the various analyses by Urbatsch et al.

Urbatsch and Roberts (2004, p. 250) noted that “Flagelliform trichomes having a subterminal ap-

pendage attachment characterize the Caribbean species [of Gundlachia] and similar trichomes are seen in

G. riskindii.” Caribbean Gundlachia and “Gundlachia” riskindii also have laminar, spatulate leaves. Neither of

these similarities, however, appears to have significantly influenced the parsimony analyses that included

morphological data. Nesom et al. (1990, p. 103) emphasized the irregularly lobed disc corollas found in

all nine of the Xylothamia species, unique among all their potentially close relatives, including Gundlachia:

“The zygomorphic disc corollas of [all of the Xylothamia] species are even more remarkable, because to our

knowledge, they do not occur in any other North American Astereae. Typically, two of the sinuses are cut

nearly to the base of the throat, one is very shallow, and the other two are intermediate in depth. The two

lobes on either side of the shallow sinus are erect, but the other three are sharply reflexed to coiling.” This

feature was not included in the morphological analysis by Urbatsch et al. (2003); it was noted by Urbatsch

and Roberts (2004) as a feature of all of the original nine Xylothamia species, but it was not mentioned as a

synapomorphy.

Xylothamia riskindii is disparate among the four species considered here as Xylothamia sensu stricto, as

evidenced by the following contrast.

1. Leaves flat, obovate-spatulate; heads solitary; ray florets 7-13; involucres 7-8 mm diam.; disc florets 30-50

X. riskindii

1. Leaves involute, appearing terete; heads in loose or compact cymes or sessile in groups of 2-3 at branch

apices; ray florets O or 1-3 hidden within the involucre; involucres 2.5-4 mm diam.; disc florets 3-7 — 1. X. diffusa,

X. triantha, and X. truncata

It also is distinct in comparison to taxa of Gundlachia.

1. Subshrubs ca 8-15 cm tall; heads solitary, sessile to subsessile; involucres broadly turbinate, 7-8 mm diam;

phyllaries without orange-resinous midveins; ray florets 7-13, corollas yellow; disc florets 30-50. X. riskindii

1. Shrubs to 200 cm tall; heads in clusters of 1-5, the clusters in racemes or corymbs, in turn borne in panicles or

flat-topped to slightly rounded corymboid clusters; involucres cylindric to narrowly obconic, 2-4 mm diam.;

phyllaries with orange-resinous midveins; ray florets 3-8, corollas white; disc florets 3-10 Gundlachia

Nesom, Disarticulation of Xylothamia 147

Urbatsch and Roberts (2004, p. 250) noted that certain evidence suggests that “G. riskindii may represent

the ancestral state for Gundlachia or may be a link connecting the Caribbean and the Mexican species." In

the evaluation here, Xylothamia riskindii remains unusual among the species placed in Xylothamia.

Chihuahuana, Medranoa, Neonesomia, and Xylovirgata

Urbatsch and Roberts (2004, p. 244) noted that “With regard to the other five species of Xylothamia, X. john-

stonii, and X. palmeri constitute a robustly supported clade (Urbatsch et al. 2003) that is herein proposed as

the new genus Neonesomia. The three remaining species of Xylothamia are each treated as monotypic genera

[Chihuahuana, Medranoa, and Xylovirgata] because they are not unambiguously supported as monophyletic

or placed within existing genera based on DNA sequence data (Urbatsch et al. 2003), and they are each

morphologically unique."

In molecular analyses including indels (Urbatsch et al. 2003: Figs. 1A, a ratchet analysis, and 1B, a

heuristic analysis), Chihuahuana, Medranoa, and Xylovirgata constitute a monophyletic group and Neonesomia

is basal to the clade that includes the three other new genera above and six more (Amphiachyris, Bigelowia,

Euthamia, Gutierrezia, Gymnosperma, and Thurovia). In the Bayesian analysis (Fig. 2), Medranoa and Chihua-

huana have a sister relationship and Neonesomia is most closely related to Thurovia. In analyses including

indel data and the morphological character matrix, the position of all four new genera is unresolved (Fig.

3, left side-bootstrap) or Medranoa and Xylovirgata show a sister relationship (Fig. 3, right side-heuristic).

In a ratchet-derived consensus tree resulting from an analysis excluding indels (Fig. 4), the positions of

Neonesomia and Medranoa are unresolved, while Chihuahuana and Xylovirgata are sister taxa.

While it is clear that each of the four taxa treated as a new genus by Urbatsch and Roberts is morphologi-

cally unique and that the molecular analyses do not provide unambiguous phyletic resolution for them (apart

from their separation from Xylothamia sensu stricto), molecular data do not provide a compelling rationale

to recognize four new genera among five species of the Chihuahuan Desert. Analogous to the position of

X. rishindii among its three congeners, X. purpusii (below as Medranoa purpusii) is relatively more distinctive

in morphology and on that basis might justifiably be treated as a monotypic genus apart from its four con-

geners. Treatment of these species within a single genus is at least as justified, based on current evidence,

as is their distribution among four. Geography and morphology provide support for their recognition as a

single lineage: geographic proximity commonly is an indicator of close evolutionary relationship, and the

zygomorphic disc corollas (discussed above) in this group of five species provide a potential apomorphy

that suggests common ancestry, as in the original delimitation of Xylothamia. While it cannot be definitively

argued that the previous treatment is flawed, a more parsimonious and conservative taxonomy is favored here.

TAXONOMIC SUMMARY

XYLOTHAMIA Nesom, Suh, Morgan & Simpson, Sida 14:106. 1990. Tee secies: Xylothamia (Aplopappus) triantha (S.E

Blake) G.L. Nesom.

1. Xylothamia diffusa (Benth.) G.L. Nesom, Sida 14:109. 1990. Ericameria diffusa Benth.; Gundlachia diffusa (Benth.)

Urbatsch & R.P Roberts.

2: Xylothamia triantha (S.F Blake) G.L. Nesom, Sida 14:113. 1990. Aplopappus [Haplopappus] trianthus S.E Blake;

Ericameria triantha (S.F Blake) Shinners; Gundlachia triantha (S.E Blake) Urbatsch & R.P Roberts.

3. Xylothamia riskindii (B.L. Turner € G. Langford) G.L. Nesom, Sida 14:113. 1990. Ericameria riskindii B.L.

Turner & G. Langford; Gundlachia riskindii (B.L. Tarner & G. Langford) Urbatsch & R.P. Roberts.

4. Xylothamia truncata G.L. Nesom, Phytologia 73:318. 1992. Gundlachia truncata (G.L. Nesom) Urbatsch & R.P

Roberts.

MEDRANOA Urbatsch & R.P Roberts, Sida 21:254. 2004. Tyee species: Medranoa (Ericameria) parrasana (S.E Blake)

Urbatsch & R.P. Roberts.

Chihuahuana Urbatsch & R.P. Roberts. Type species: Chihuahuana (Ericameria) purpusii (Brandeg.) Urbatsch & R.P. Roberts.

Neonesomia Urbatsch & R.P. Roberts. TYPE species: Neonesomia (Aster) palmeri (A. Gray) Urbatsch & R.P. Roberts.

Xylovirgata Urbatsch & R.P. Roberts. TYPE species: Xylovirgata (Haplopapus) pseudobaccharis (S.F. Blake) Urbatsch & R.P. Roberts.

f4L,D ID L

148 Journal of tani titute of Texas 1(1)

Etymology.—Medranoa is chosen here, from among the four possibilities, as the name to represent this group

of species because it honors a Mexican botanist, Dr. F.G. Medrano, which seems appropriate for this group

of primarily Mexican species.

1. Medranoa johnstonii (G.L. Nesom) G.L. Nesom, comb. nov. Neonesomia johnstonii (G.L. Nesom) Urbatsch € R.P

Roberts; Xylothamia johnstonii G.L. Nesom, Sida 14:110. 1990.

2. Medranoa parrasana (S.F Blake) Urbatsch & R.P Roberts, Sida 21:255. 2004. Ericameria parrasana S.E Blake;

Haplopappus parrasanus (S.E Blake) S.E Blake; Xylothamia parrasana (S.E Blake) G.L. Nesom.

3. Medranoa palmeri (A. Gray) G.L. Nesom, comb. nov. Aster palmeri A. Gray, Proc. Amer. Acad. Arts 17:209. 1882.;

Ericameria austrotexana M.C. Johnston (non Ericameria palmeri (A. Gray) H.M. HalD; Neonesomia palmeri (A. Gray) Urbatsch & R.P.

Roberts; Xylothamia palmeri (A. Gray) G.L. Nesom.

4. Medranoa purpusii (Brandeg.) G.L. Nesom, comb. nov. Ericameria purpusii Brandeg., Univ. Calif. Publ. Bot. 4:191.

1911; Chihuahuana purpusii (Brandeg.) Urbatsch & R.P. Roberts; Haplopappus [Aplopappus] purpusii (Brandeg.) S.E Blake; Xylothamia

purpusii (Brandeg.) G.L. Nesom.

5. Medranoa pseudobaccharis (S.F Blake) G.L. Nesom, comb. nov. Haplopappus pseudobaccharis S.F. Blake, J. Wash-

ington Acad. Sci. 40:47. 1950; Ericameria pseudobaccharis (S.E Blake) Urbatsch; Xylothamia pseudobaccharis (S.E Blake) G.L. Nesom;

Xylovirgata pseudobaccharis (S.E Blake) Urbatsch & R.P. Roberts.

ACKNOWLEDGMENTS

I am grateful to Billie Turner, John Strother, and an anonymous reviewer for their comments.

REFERENCES

LANE, M.A. 1996. Taxonomy of Gundlachia (Compositae: Astereae). Brittonia 48:532-541.

Nesom, G.L. 1992. A new gypsophilic species of Xylothamia (Asteraceae: Astereae) from the Cuatro Cienegas area

of Coahuila, Mexico. Phytologia 73:318-320.

NESOM, G.L. 2000. Generic conspectus of the tribe Astereae (Asteraceae) in North America, Central America, the

Antilles, and Hawaii. Sida, Bot. Misc. 20:i-viii, 1-100.

Nesom, G.L., Y. Sun, D.R. Morean, and B.B. Simpson. 1990. Xylothamia (Asteraceae: Astereae), a new genus related

to Euthamia. Sida 14:101-116.

UnBATSCH, L.E., R.P. Roserts, and V. Karaman. 2003. Phylogenetic evaluation of Xylothamia, Gundlachia, and related

genera (Asteraceae, Astereae) based on ETS and ITS nrDNA sequence data. Amer. J. Bot. 90:634-649.

UnBATSCH, L.E. and R.P. Roserts. 2004. New combinations in the genus Gundlachia and four new genera of Astereae

(Asteraceae) from northern Mexico and the southern United States. Sida 21:243-257.

TWO NEW SPECIES OF GRATIOLA (PLANTAGINACEAE) FROM EASTERN NORTH

AMERICACAND AN UPDATED CIRCUMSCRIPTION POR GRATIOLA NEGLECTA

Dwayne Estes Randall L. Small

University of Tennessee University of Tennessee

Dept. of Ecology and Evolutionary Biology Dept. of Ecology and Evolutionary Biology

Knoxville, Tennessee 37996 U.S.A. Knoxville, Tennessee 37996 U.S.A

Email: tnplants@yahoo.com

ABSTRACT

apes |

Nibora, a North American taxon as o circumscribed, includes six speci G. ebracteata, G. flava, G. floridana, G.

$ E 1

heterosepala, G. neglecta, and G. virginiana. 1 to western North America and the remaining

four species are mostly eastern North American. The species with the lees! range and greatest degree of morphological variability is

G. neglecta. A recent investigation of G. neglecta involving fieldwork, examination of herbarium specimens, morphological analysis, and

phytogeographic study, has resulted in the discovery of two undescribed species, G. graniticola sp. nov. and G. quartermaniae sp.

nov., both of which are endemic to rock outcrop of eastern North America. In this paper, both new species are described,

illustrated, and compared to their widespread congener, G. neglecta. An updated circumscription of G. neglecta is provided and a key

distinguishing the new species from G. neglecta is included.

RESUMEN

Gratiola sección Nibora, un taxon norteamericano como se circumscribe normalmente, incluye seis especies: G. ebracteata, G. flava, G.

floridana, G. heterosepala, G. neglecta, y G. virginiana. Gratiola ebracteata y G. heterosepala están restringidas al oeste de Norte América y

las restantes cuatro especies están principalmente en el este de Norte América. La especie con el rango más amplio y el DU grado de

variabilidad morfológica es G. neglecta. Una investigación reciente de G. neglecta con trabajo de campo, examen de esp nes de her-

bario, análisis morfológico, y estudio fitogeográfico, ha dado como resultado el descubrimiento de dos nuevas especies, G. graniticola

sp. nov. y G. dd di sp. nov., ambas endémicas de comunidades de afloramientos rocosos del este de Norte América. En este

dl d neglecta. Se aporta una circunscripción

puesta al día de G. neglecta y se A una clave para do las nuevas especies de G. neglecta.

Gratiola L. section Nibora (Raf.) Pennell (Plantagianceae) was erected by Pennell (1935) to accommodate

the annual North American species characterized by having capsules equaling or slightly exceeding the

sepals, leaves sessile to scarcely clasping and obscurely glandular punctate, and seeds yellowish and faintly

reticulate. Pennell (1935) recognized five species within the section: G. ebracteata Benth., G. flava Leavenw.,

G. floridana Nutt., G. neglecta Torr., and G. virginiana L. Mason and Bacigalupi (1954) added a sixth spe-

cies to this section when they described G. heterosepala Mason € Bacig. from northern California. Gratiola

ebracteata and G. heterosepala are restricted to western North America while G. flava, G. floridana, and G.

virginiana are mostly eastern North American (G. virginiana is also disjunct to central Mexico). Gratiola

neglecta is the most widespread and most variable species in the section, ranging across much of temperate

North America. Throughout its broad range, G. neglecta inhabits a wide diversity of wetland communities

and exhibits considerable variation in degree of branching, stem pubescence, leaf shape, flower morphol-

ogy, and capsule size.

Recent evidence from field and herbarium studies indicates that material previously referred to G.

neglecta includes two undescribed species. The first new species, G. quartermaniae D. Estes sp. nov., has a

highly fragmented distribution in eastern North America and is endemic to ephemerally wet sites associated

with calcareous outcrops (cedar glades) and prairies. The second new species, G. graniticola D. Estes sp.

nov., is endemic to north-central Georgia where it is restricted to vernal pools on granitic outcrops. In this

paper, both new species are described, illustrated, and compared to their widespread congener, G. neglecta.

Because G. quartermaniae and G. graniticola have been included within the concept of G. neglecta by previous

authors, an updated circumscription of G. neglecta is provided.

J. Bot. Res. Inst. Texas 1(1): 149 — 170. 2007

150 Journal of the Botanical R h Institute of Texas 1(1)

MATERIALS AND METHODS

In order to clarify morphological variation within and between G. neglecta, G. quartermaniae, and G. gra-

niticola, an investigation was conducted that incorporated fieldwork, examination of herbarium specimens,

morphological analyses, and phytogeography. Fieldwork was conducted in portions of 26 states in the United

States and the province of Ontario, Canada between 2001 and 2006. In addition, more than 4,000 herbarium

specimens (including some digital images), representing all taxa from sect. Nibora, were examined from the

following 49 herbaria: A, ALU, APSC, ASTC, AUA, BRIT, CAN, CITA, CLEMS, DAO, DUKE, EKY, FSU, GA,

GH, H, ILLS, ISC, JEPS, JSU, K, KANU, LL, LSU, MIN, MISS, MO, MTSU, NCSC, NCU, NLU, NO, NY, NYS,

OKL, PH, SBSC, SMU, TENN, TEX, TROY, TRT, UARK, UC, UNA, US, USCH, VDB, and VPI (herbarium

acronyms follow Index Herbariorum, http://www.nybg.org/bsci/ih/search).

From the herbarium specimens examined during this project, a subset of 87 mature and complete speci-

mens representing 55 G. neglecta, 15 G. graniticola, and 17 G. quartermaniae, was selected for use in a mor-

phometric study. Specimens were chosen to represent the full geographic distribution, range of habitat, and

morphological variation of each species. For each specimen, 10 quantitative vegetative and floral characters

were measured (Table 1); these specimens are denoted by an asterisk in the lists of representative specimens

examined. Seed measurements were taken from five of the above specimens (1 G. graniticola, 4 G. neglecta)

plus an additional 14 specimens representing a total of 10 widespread populations of G. neglecta, four of G.

graniticola, and five of G. quartermaniae. Twenty seeds from a single capsule were measured per population,

and three quantitative characters were scored per seed (Table 1). Specimens used for seed measurements are

indicated by a dagger (+) in the lists of specimens examined. For each scored character, summary statistics

including mean, standard deviation, and range were calculated; these values are presented in Table 1. In the

taxonomic key and species descriptions, measurements for characters are given as the mean + one standard

deviation with extreme values, based on additional observations, given in parentheses. In order to reveal

discontinuities in the data and to determine which characters are most useful for delimiting taxa, pairwise

comparisons of characters were conducted using scatter diagrams and box plots. Seeds and trichomes of all

three species were also examined with the aid of scanning electron microscopy (SEM) to search for useful

taxonomic characters. The geographic distribution of G. neglecta, G. graniticola, and G. quartermaniae was

determined by examining the collection data included on herbarium specimens and plotting the county-

level distribution of each species on outline maps. Each point on these maps is represented by at least one

herbarium specimen examined.

RESULTS AND DISCUSSION

Morphology

Gratiola neglecta, G. quartermaniae, and G. graniticola form a morphologically cohesive group referred to here

as the Gratiola neglecta complex. A fourth species, G. floridana, also belongs to this complex; however, it is

quite distinct morphologically in spite of sharing a suite of features uniting it with the other three species.

Gratiola floridana differs from the other members of the complex in its overall larger features including much

larger flowers 13-25 mm long (vs. 5-14 mm), longer proximal fruiting pedicels averaging 23-43 mm long

(vs. 12-25 mm), and longer seeds averaging 0./9—0.9 mm (vs. 0.4-0.6 mm). This species tends to inhabit

forested sites whereas the others mostly grow in open communities. It is also the southernmost member

of the complex ranging from northwestern Florida and southeastern Louisiana (historically) north into

southeastern Tennessee. The distribution of G. floridana only slightly overlaps with the ranges of G. neglecta

and G. quartermaniae in the northern portion of its range. Since G. floridana is one of the most distinctive

species of the genus and has rarely been confused with G. neglecta or the two new species, it will not be

discussed further.

Several characters distinguish G. graniticola from G. neglecta and G. quartermaniae (Table 2; Fig. 1, Fig.

2). Gratiola graniticola has shorter leaves (normal leaves that subtend pedicels are also referred to as bracts

or bracteal leaves in this paper) that are widest at or below the middle (Fig. 2 A), shorter pedicels that

Estes and Small, N i f Gratiola f t North America 151

Petite VI Nr atvia

Taste 1. Morphological characters measured for Gratiola graniticola, G. neglecta, and G. quartermaniae and their means +

standard deviations and ranges (parentheses). N=sample size.

G. graniticola G. neglecta G. quartermaniae

Characters (N=15) (N=55) 7

Stem height (cm) 14.8 € 5.9 19955 OSES

(7.4-29.4) (10.2-33.2) (5.8-29)

Stem diameter (mm) T2 £02 (6505 14+04

(0.7-1.4) (0.8-2.9) (0.6-2.3)

Leaf length (mm) 10 £28 30.9 £ 10.3 25.1: 50

(6.3-17 7) (11-66) (16-43)

Leaf width (mm) ZE O AS 3808

(1.1-5.2) (2.7-18) (1.8-4.5)

Leaf length/leaf width (ratio) 46+ 1.0 4.1 + 0.80 TL dd

(2.8-7 45) (2.6-6.1) (5.5-11.2)

No. teeth per leaf margin 1,020.7 3E. ]7 desi |

(0-3) (1-7) (0-3)

Proximal pedicel length (mm) 1244149 20.6 € 7.7 17+ 4.2

(5.3-22) (10.5-37) (8-22)

Bract length (mm) DUE 286295 218 £6.6

(5.3-11.2) (11.5-66) (12.5-33)

Pedicel length/bract length (ratio) 1.5 x04 (09:6 0.3 03 E03

(0.9-2.3) (0.3-1.3) (0.5-1.6)

Capsule length (mm) poss 4.3 € 0.6 4.1 € 0.6

(2.9-3.6) (2.6-5.7) (3.4-5.1)

G. graniticola G. neglecta G. quartermaniae

(N=80) N=200) N-1

Seed length (mm) 0.40 + 0.03 0.54 + 0.06 0.59 + 0.04

(0.31-0.47) (0.42-0.7) (0.43-0.71)

Seed diameter (mm) 0222002 0.24 + 0.02 0290.03

(0.17-0.27) (0.18-0.29) (0.19-0.37)

Seed length/seed width (ratio) 1.86 + 024 234977 20535027

(1.32-2.53) (1.67-3.03) (1.47-2.6)

are longer relative to their subtending bracts (Fig. 2, D-E), smaller corollas that have a purplish or pinkish

posterior lobe and beard of whitish trichomes, bracteoles that are shorter than to only slightly exceeding

the calyces, smaller more subglobose purple-tinged capsules (Fig. 2 F), smaller seeds (Fig. 2 G-H; Fig. 3 AJ,

and bulbous-based trichomes (Fig. 3). Gratiola neglecta and G. quartermaniae have longer leaves (Fig. 2 A),

longer pedicels that are mostly equal to or shorter than their subtending bracts (Fig 2, D-E), larger corollas

that usually lack purplish or pinkish coloration and that have a beard of yellow trichomes inside the corolla

orifice, bracteoles that are mostly longer than the calyces, larger more ovoid and brownish capsules (Fig. 2

F), larger seeds (Fig. 2, G-H; Fig. 3 B-C), and slender-based trichomes (Fig. 3, E-F). A scatter diagram of

leaf length vs. capsule length between G. graniticola, G. neglecta, and G. quartermaniae reveals two primary

clusters that exhibit minimal overlap (Fig. 1 A). In this scatter plot, specimens of G. graniticola mostly group

separately from the second unresolved cluster that consists of specimens of G. neglecta and G. quarterma-

niae. A scatter plot of proximal pedicel length/subtending bract length vs. leaf width also distinguishes G.

graniticola from G. neglecta (Fig. 1 B).

Gratiola quartermaniae differs from G. neglecta in having a glabrous midstem, narrower (Fig. 2 B) and

more falcate, fewer veined and fewer toothed leaves that have a larger length to width ratio (Fig. 2 C) and

seeds that average longer, thicker, and darker (Fig. 2, G-H). In comparison to G. quartermaniae, G. neglecta

has mostly pubescent (rarely glabrate in some New England estuarine populations) midstems, wider (Fig. 2

B), more veined and more toothed leaves that have a smaller leaf length to width ratio (Fig. 2 C). The seeds

152 Journal of the Botanical R h Institute of Texas 1(1)

TABLE 2. Qualitative morphological characters useful for distinguishing G. graniticola, G. neglecta, and G. quartermaniae.

Character G. graniticola G. neglecta G. quartermaniae

Stems simple-rarely branched branched-rarely simple simple-rarely branched

Leaf shape lanceolate-ovate to narrowly narrowly elliptic, rhombic, linear, linear-lanceolate,

oblong or oblanceolate or elliptic-lanceolate

Basal leaf disposition + congested not conge + congested

middle

bract 2 pedicel

gested

middle or beyond middle

bract 2 pedicel

middle or below middle

bract « pedicel

Widest point of leaf

Proximal bract to pedicel ratio

Mid-stem vestiture

Trichome shape

Ratio bracteole length/calyx

length

glandular-pubescent

bulbous-based

bracteoles < to slightly

exceeding calyx

glandular-pubescent

slender-based

bracteoles = calyx

glabrous

slender-based

bracteoles = calyx

Posterior corolla lobe color purplish or pinkish white (rarely pinkish) white (rarely pinkish)

Beard color white yellow yellow

Capsule shape subglobose ovoid ovoid

Capsule color purplish brown brown

Seed color

Habitat

grayish-brown

granite outcrops

yellowish-brown

various wetland types,

rarely on outcrops

grayish-brown

limestone/dolomite

outcrops, calcareous

prairies

of G. neglecta are lighter in color and average slightly shorter and are not as thick as those of G. quarterma-

niae (Fig. 2, G-H). In Fig. 1 C, a scatter plot of leaf length/leaf width vs. number of teeth per margin for G.

neglecta and G. quartermaniae reveals two clusters of specimens.

Distribution and Ecology

Gratiola neglecta has the largest distribution of the three species, being found throughout most of temperate

North America (Fig. 4). It ranges from Nova Scotia and British Columbia, Canada, south in the United States

to central Georgia, coastal Texas, northern Arizona, and northern California. The species is most common

in the eastern United States particularly in the lower Mississippi, Missouri, and Ohio River valleys. West of

the Mississippi River, the range of G. neglecta mostly follows the major river systems toward the Great Plains.

From the upper Missouri River watershed, G. neglecta ranges south into the southern Rocky, Cascade, and

Sierra Nevada mountains. Several populations in the western United States are associated with reservoirs;

these may represent recent introductions by migrating waterfowl. Interestingly, G. neglecta has also been

collected in France (Simon s.n. FSU; Rastetter 11653 UC) and Finland (Lampinen 5629 H; see Suominen 1984)

where probably introduced.

Gratiola neglecta grows in a broader array of wetland communities and endures a greater range of envi-

ronmental conditions than G. graniticola or G. quartermaniae. It grows from sea level to an elevation of 2400

m in the mountains of the western United States. Compared to the new species, G. neglecta occurs more

frequently in the deeper soil of agricultural fields, openings in bottomland hardwood forests, wet meadows,

mudflats, and pond margins. Rarely, G. neglecta occurs in salt marshes or on various types of shallow-soiled

rock outcrops including igneous, sandstone, limestone, and granite formations.

Gratiola quartermaniae has a fragmented range (Fig. 5) and is most common in the limestone cedar

glades of the Interior Low Plateau of middle Tennessee and northern Alabama. From this core range, it

is disjunct to the alvars of southeastern Ontario, Canada, a distance of ca. 1200 km. Most of the Ontario

populations are associated with the Napanee limestone plain but a few are found in the Dummer Moraine

and Prince Edward Peninsula physiographic regions (Chapman & Putnam 1984). Numerous other species

that are more common on calcareous outcrops in the southeastern United States also occur on Canadian

alvars including several of the species commonly associated with G. quartermaniae in Tennessee and Alabama

such as Carex granularis Muhl. ex Willd., C. crawei Dewey, C. molesta Mack. ex Bright, Isanthus brachiatus (L.)

Lr E G | £,

Estes and Small, N je of Gratiola t Narth Amarica 153

Ñ 8

E s. AAA 90

& 00 B 89595 o

$' ag eo 9? S °

PE MA SC

mal

o A

* 2-

S 4

0 1 a Li Li a a 1

0 10 20 30 40 50 60 70

Leaf Length (mm) "

20,

4 ©

716 pi

E | O O

E O

- 124

S O S S OD

z 80 O O GO

we a

9 O OQ» O A

94 Q9 &

A AR, AR

AS A DA. A

0 T T T T 1

0 0.5 1 1.5 2 2.5

Pedicel Length (mm) / Bract Length (mm) E

5 a

p 4- O ABD CO

9 7 O QGUOQOOO *

É 2 O co eee $

Ò .

$ O e. e

0 2 4 6 8 10 12

Leaf Length (mm) / Leaf Width (mm) m

Fic. 1. Scatter plots of leaf | Í le length (A) imal pedi I l length vs. leaf width (B), leaf length/leaf width vs.

number of teeth per margin (© for G. graniticola (open triangles), G. neglecta open circles), and/or 6 guareermaniae (closed circles). Note thar open

KAD pan rp en peame with pubescent mid- -stems, and solid I id-stems with ti

| dubio ypical of G. quarti j p for! ing a pub t mid-stem. This specimen (Kral 52812, VDE,

Cannon Co., TN) f | limestone at the edge of the range of G. ]

154 Journal of the Botanical Research Institute of Texas 1(1)

60-

Leaf Length (mm) Pedicel Length / Bract Length

|j, ma -=

6 -

157 Leaf Width (mm) Capsule Length (mm)

: A --

0.8:

187 Leaf Length / Leaf Width Seed Length (mm)

i EN. m

Cc G

Proximal Pedicel Length (mm) 0.41 Seed Diameter (mm)

: f on

Fic. 2. Graphical comparison of eight select ters for Gratiola graniticola (lef iude TES enter and €, quanermansde (right): leaf length

(A), eave (B), ratio of leaf length to leaf width (O, imal pedicel length (D), I I g ling bract length (E),

(F), seed length (G), (H).

B.S.P., Scutellaria parvula Michx., and Sporobolus vaginiflorus (Torr. ex Gray) Wood. Gratiola quartermaniae

is also disjunct to Will County, Illinois from its main range in central Tennessee, a distance of approx. 600

km. Two limestone glade near-endemics, Dalea foliosa (Gray) Barneby and Astragalus tennesseensis Gray ex

Chapman, share this similar distribution pattern (Baskin & Baskin 2003). Gratiola quartermaniae is also

disjunct to the Edward's Plateau of Texas, a distance of ca. 1200 km. Interestingly, Juncus filipendulus Buckley,

a species that G. quartermaniae frequently occurs with in Alabama and Tennessee, is also disjunct to the

Edward's Plateau where it occurs with G. quartermaniae. Therefore, while the disjunction patterns exhibited

by G. quartermaniae are unusual, further examination indicates that in each of these areas G. quartermaniae

occurs in similar habitat and always occurs with other calciphilous species, some of which have similar

patterns of disjunction. This species should be searched for in other regions where calcareous outcrops and

prairies occur such as the limestone glades of the southern Ridge and Valley of southeastern Tennessee and

northwestern Georgia, the Blackbelt prairies of Mississippi and Alabama, the limestone glades of central

and western Kentucky, the Ozark glades of southern Missouri and northern Arkansas, and alvar habitats

in New York, Michigan, and Ohio.

Gratiola quartermaniae is found on limestone or dolomite outcrops and calcareous prairies. In these

Estes and Small, New species of Gratiola from eastern North America

Fic. 3. Seeds of Gratiola graniticola (A), G. neglecta (B), and G. quartermaniae (C); scale bars = 100 um. Trichomes of G. graniticola (D), G. neglecta (E),

and G. quartermaniae (F); scale bars = 20 um.

ftha R * ID L

156 Journal of t titute of Texas 1(1)

r L* aS uL ius Lr. Et | I Le * Al l4 Amarirs

Fic. 4 t

habitats, the species predominantly occurs in shallow clayey soils of ephemeral pools, seasonal streambeds,

and periodically wet meadows on or immediately adjacent to outcrops. These sites are usually flat to slightly

sloping and are located in areas that receive high to moderate levels of sunlight. They are wet in late winter

157

t North America

Estes and Small, N pecies of Gratiola f

North Dakota

j )

fu |

j raras: EEE

= ===- Minnesota

| d

| ]

| South Dakota |

T d m

— A

= Nebraska ii lowa

f 1

a i

a — T E d

| — 1 | ^ P}

| & N Illinois | Indiana y im SAN HS

| ) Y ¿ d " P land. Delaware

j /

| fne | 7 AY ef, West Virginia ~ CSS Sy)

| | Missouri 4 f as’ 4 7 NE "y;

— | ) ad ^P Vi so dl

————— d < kantuck m irginia BM

7 me ver entuc y e —A

j pe st F — AN

l ——, #— e. Lon

| — | 2f p Tennessee ¿e es 5

| gue | r y Ha North Carolina S

M | Arkansas M SEE m : "TA rt

“a | J | Wtf, \ r4

^ i | “South Carolina ES

(a) |

| pet Alabama \ Georgia ^ y?

Texas E Pj y

4 É | 0

cm oh a i

os ibi, E Y \

Ty "e \

jf onda à

2. A

€

Fic. 5. Geog ic distributi ]

and early spring but become severely desiccated by late spring and early summer. Rarely, G. quartermaniae

occurs in situations otherwise more typical for G. neglecta such as low wet fields, open wet woods, and marsh

edges, but these populations are always located within close proximity to glade habitat. Limestone glade

endemics such as Leavenworthia alabamica Rollins, L. crassa Rollins, L. torulosa Gray, and Lesquerella lyrata

£+sha D o ID

158 Journal of t h Institute of Texas 1(1)

Rollins are sometimes found in disturbed non-outcrop habitats often in association with G. quartermaniae.

In central Tennessee and northern Alabama, G. quartermaniae is almost always associated with limestone

cedar glade endemics or calciphiles such as Allium cernuum Roth, Carex crawei, C. granularis, Dalea foliosa,

D. gattingeri (Heller) Barneby, Eleocharis bifida S.G. Smith, Hypericum sphaerocarpum Michx., Isoetes butleri

Engelm., Juncus filipendulus, Leavenworthia spp., Ludwigia microcarpa Michx., Mecardonia acuminata (Walt.)

Small, Sedum pulchellum Michx., Sporobolus vaginiflorus, and Talinum calcaricum Ware. In areas where G.

quartemaniae is disjunct as in Ontario, Illinois, and Texas, the species is associated with a number of ad-

ditional calciphilous taxa, including a few of those listed above.

Gratiola graniticola is restricted to granite outcrops in 13 counties on the Piedmont Plateau of Georgia

(Fig. 6). Of the approx. 17 Piedmont granite outcrop endemics (McVaugh 1943; Weakley 2007), G. graniti-

cola is one of only five species, along with Isoetes melanospora Engelmann, I. piedmontana (N.E. Pfeiffer) C.F.

Reed, I. tegetiformans Rury, and Amphianthus pusillus Torr., restricted to the ephemeral pools of the outcrops.

Interestingly, I. tegetiformans and G. graniticola are the only Piedmont granite outcrop endemics completely

restricted to Georgia.

All known populations of G. graniticola occur on granite outcrops in water-filled depressions lined

with a thin layer of soil. These depressions are filled with water during the winter and spring months but

dry out in the summer and fall. Species commonly associated with G. graniticola include Croton willdenowii

G.L. Webster, Cyperus granitophilus McVaugh, Diamorpha smallii Britt. ex Small, Eleocharis obtusa (Willd.)

Schult., Isoetes piedmontana (N.E. Pfeiffer) C.F. Reed, Juncus georgianus Coville, Lindernia monticola Muhl. ex

Nutt., Minuartia uniflora (Walt.) Mattf., Packera tomentosa (Michx.) C. Jeffrey, Pilularia americana A.Braun,

Rhynchospora sp., and Schoenolirion croceum (Michx.) A. Gray.

Gratiola quartermaniae is sympatric with G. neglecta; however, the two species generally occupy different

habitat types. They occur syntopically at a few sites in middle Tennessee and northern Alabama where the

typical glade habitat of G. quartermaniae occurs in close proximity to habitats preferred by G. neglecta. Each

of these sites is located within ca. 500 m of a cedar glade or glade-like area. Plants at these sites appeared

to belong either to G. quartermaniae or to G. neglecta with no obvious hybrids observed at most sites. One

specimen (Kral 52812 VDB, MO) collected from a seep over limestone in Cannon County, Tennessee ap-

pears to be typical G. quartermaniae in general morphology and habitat; however, the middle portion of the

stems on this specimen are slightly pubescent and more typical of G. neglecta (Fig. 1 C). It is possible that

this specimen represents a hybrid between G. neglecta and G. quartermaniae. Although G. neglecta was not

found on any cedar glades in middle Tennessee or northern Alabama, the species has been collected from

a variety of rock outcrop types elsewhere where it exhibits morphological features typical of non-outcrop

populations. Gratiola floridana is sympatric with both G. quartermaniae and G. neglecta in northern Alabama’s

Moulton Valley (Lawrence and Morgan counties). Although these three species have been found within 1

km of each other, sites supporting all three species are unknown. Gratiola floridana and G. quartermaniae

occur syntopically at one site in Lawrence County, Alabama (Whetstone et al. 16471 JSU, mixed collection

of G. floridana and G. quartermaniae). Gratiola floridana usually inhabits shaded muddy sites in forested

bottoms or ravines but in northern Alabama it rarely occurs in habitats more typical of G. quartermaniae.

No obvious hybrids between G. floridana and G. quartermaniae or between G. floridana and G. neglecta have

been discovered.

The range of G. graniticola lies near the southern edge of the range of G. neglecta and the two species

overlap only in northeastern Georgia (Elbert and Greene counties). Although the Greene County specimen

of G. neglecta (Allison 2630 GA) was collected from a granite outcrop, the two species have never been ob-

served growing syntopically and no putative hybrids have been found. A disjunct population of G. graniticola

reportedly occurs on a granite outcrop in Lancaster County, South Carolina (J. Allison, Georgia Natural

Heritage Program, pers. comm.), but specimens needed to confirm this report have not been located.

160 Etha Rataniral R :

Journal of

Institute of Texas 1(1)

KEY TO THE SPECIES OF THE GRATIOLA NEGLECTA COMPLEX

. Howers 13-25 mm ene IN surface of the corolla lobes pilose; proximal fruiting as a )23-43(-55)

mm long; seeds (0.6-)0.79-0.9 mm long, trichomes short stalked, the stalks

than the glandular head

. Howers 5-14 mm long, adaxial surface ofthe corolla lobes glal imal fruiting pedicels (5-)12-25(-37)

mm long; seeds (0.3-)0.4-0.6(-0.7) mm long, trichome stalks > 1 5 times as long as the glandular head.

2. Mid-stem leaves (11-)20-41(-66) mm long; proximal fruiting pedicels (8-)13-25(-37) mm long, (0.3-)

0.5-1(-1.6) times as long as the subtending bracteal leaves; bracteoles slightly longer to conspicuously

longer than the sepals; posterior corolla lobe white (rarely inconspicuously tinged with pink or lavender)

beard inside corolla orifice of yellow trichomes; mature capsules ovoid, brown; seeds (0.4-)0.5-0.6(-0.7)

mm long and (0.18-)0.21—0.29(-0.37) mm thick, trichomes slender-based

3. Leaves narrowly elliptic or rhombic to oblanceolate, not conspicuously falcate, 2.7-)5-11(-18) mm wide

at widest point; length to width ratio (2.5-)3.5-5(-6), each margin with (1—)3—5(-7) often conspicuous

teeth, primary veins 3-5 (7); mid-stem moderately to densely glandular pubescent (rarely glabrate)

seeds (0.18-)0.22-0.26(-0.29) mm thick

. Leaves linear, linear-lanceolate, to elliptic-lanceolate, often falcate, (1-)2.5-4(-4.5) mm wide at widest

point, length to width ratio (5.5—)6-9.5(-11), entire or each margin with 1-2(-3) inconspicuous teeth

primary vein 1(-3); mid-stem glabrous, seeds (0.19-)0.26-0.32(-0.37) mm thick G. quartermaniae

2. Mid-stem leaves (6-)7-13(-18) mm long; proximal fruiting pedicels (5-)7-17(-22) mm long, (0.9-)1-2(22.3)

times as long as the subtending bracteal leaves; bracteoles shorter than to barely exceeding sepals; poste-

rior corolla lobe conspicuously tinged with pink or purple; beard inside corolla orifice of white trichomes;

mature capsules subglobose, purplish; seeds (0.3-)0.36-0.42(-0.5) mm long and (0.17-)0.20-0.24(-0.27)

mm thick, trichomes bulbous based

I equalit M VI r shorter

G. floridana

G. neglecta

G. graniticola

TAXONOMIC TREATMENT

Gratiola neglecta Torr,, Catal. Pl. New York. 10, 89. 1819. (Fig. 7). Tyee: [no locality data on specimen, but as noted

by Stuckey (1979) this specimen was donated to the Schweinitz herbarium by John Torrey. Torrey (1819) gives the locality as “In-

undated and moist places, New York."], [no collection date provided on sheet or in Torrey (1819)], [collector not specified on sheet

but Pennell (1935) noted “it is almost certainly a plant of Torrey’ collecting. .."]. (Lectotype, here designated: PH!; ISOLECTOTYPE, here

designated: K-digital image!).

Conobea borealis Spreng., Neue Entdeck 3:26. 1822.

Gratiola missouriana Beck, Amer. Jour. Sci. 10:258. 1826.

Gratiola odorata Rat., Autik. Bot. 43. 1840.

Gratiola heterophylla Raf., Autik. Bot. 43. 1840.

Gratiola gracilis Benth., Prod. Syst. Nat. Regn. Veg. 10:402. 1846.

Gratiola officinalis Michx. fs caroliniensis Pers., Syn. Plant. 1:14. 1850.

Gratiola lutea Raf. var. glaberrima Fernald, Rhodora 34:149. 1932. Gratiola neglecta Torr. var. glaberrima (Fernald) Fernald, Rhodora

51:84. 1949

Plants annual, solitary, erect herbs, (10—)16-27(533) cm tall. Roots simple, fleshy, whitish with numerous

rootlets. Stems erect, somewhat fleshy, simple or with few to many spreading-ascending branches, terete

or slightly rounded-quadrangular in cross section, (0.8—)1.2-2.2(-2.9) mm in diameter at midstem; with

(6-)7-10(-12) leafy nodes, mid-stem internodes (17-)28-45(-48) mm long, basal internodes not conspicu-

ously shortened; stem green, usually densely short glandular-pubescent from below middle to apex, becom-

ing glabrate near the base or rarely glabrate throughout, trichomes spreading, translucent, slender-based

and gland-tipped. Leaves simple, oppositely-decussate, narrowly elliptic or rhombic to oblanceolate, or

uncommonly falcate, spreading, 3—5(-7)-veined, thin, mid-cauline blades (11-)24-44(-66) mm long and

(35-11-18) mm wide, 2.5—)3.5-5(-6) times longer than wide, median leaves usually largest decreasing

in size toward base and apex, apex acute, widest at or just distal to the midpoint, margins with (1-)2-5(-7)

remotely spaced low and inconspicuous to sharp and evident teeth per margin, base acuminate and sessile or

slightly clasping; blades green, glabrate to moderately glandular pubescent. Flowers solitary in axils of upper

median and distal bracteal leaves, erect to spreading, zygomorphic, perfect; pedicels slender, ascending to

divergent, (10—)12-30(-37) cm long, (0.27-)0.44-0.94(-1.33) times as long as the bracteal leaves, densely

to sparsely pubescent with slender-based gland-tipped trichomes. Bracteoles 2, paired, closely subtending

Estes and Small, New species of Gratiola f tern North America 161

C D

HH I 1L

Fi. 7.6 D I I. A El x

) les (scale bar = 2 mm). B. Close-up of mid-stem (scale bar = 2 mm). C. Flower,

(scale bar = 2.5 mm).

£.

lateral view (scale bar = 4 mm). D. Flower,

the calyx, lanceolate, narrowly elliptic, to oblanceolate, sometimes falcate, apex narrowly obtuse to acute,

margins with 1-2 inconspicuous teeth, bases straight or tapering, longitudinally 3-nerved, in flower 2.5-7

mm long and 0.5-1 mm wide, enlarging as fruit matures and becoming foliose and up to 15 mm long and

2 mm wide, thin, green or minutely purple-tipped, sparsely to densely covered with slender-based gland-

162 Journal of the Botanical R h Institute of Texas 1(1)

tipped trichomes on both surfaces. Calyx irregularly campanulate with 5 subequal sepals; these distinct,

lanceolate, longitudinally 3-veined, slightly fleshy, green, Q.2-)3-4.4-5) mm long and ca. 0.5 mm wide,

apex narrowly obtuse, margins entire, sparsely to densely covered with gland-tipped trichomes. Corolla

tubular, gamopetalous, zygomorphic, slightly marcescent, 6.5-12 mm long; corolla tube quadrangular,

dorsal surface with a prominent hump near midpoint, the ventral surface canaliculate, to 9 mm long, to 2.1

mm in diameter proximal to dorsal hump, pale yellow, yellowish-cream, or yellowish-green, with many

dichotomously forking brownish-violet lines extending from the tube base to the base of the corolla lobes,

sparsely to moderately glandular pubescent pubescent externally with slender gland-tipped trichomes, in-

ner surface near orifice at base of posterior corolla lobe with moderate to dense beard of clavately thickened

yellow trichomes, proximal and median inner corolla tube pilose with eglandular trichomes; corolla lobes

5, generally slightly broader than high and obtuse to emarginated, spreading, white, 1.7-2.5 mm long and

3.2-3.8 mm wide; the posterior lobe generally largest, the two lateral lobes and lower lobe equal or slightly

smaller, adaxial surfaces glabrous, abaxial surfaces glabrous or slightly glandular pubescent. Stamens 2,

inserted near the middle of dorsal surface of the corolla tube, filaments to 1.2 mm long, anthers transversely

oriented to the filaments, 0.6-0.8 mm long and 0.4—0.6 mm wide, connective greatly dilated around the two

anther sacs, whitish; staminodes inserted ca. 1.5 mm above base of corolla or absent, when present ca. 0.3

mm long and not capitate. Gynoecium 5.3-6.8 mm long, subtended at the base by an orange nectary ring,

ovary 1.4-3.3 mm long and to 2.1 mm in diameter, style 3.0-3.9 mm, stigma 2-lobed, dilated and flattened,

ca. 0.7 mm long. Capsules ovoid, apex acute to obtuse, usually widest below the middle, (2.6—)3.6—5(-6)

mm long, 3-5 mm in diameter, brown at maturity. Seeds several hundred per capsule, brownish-yellow,

10—13 ribbed, longitudinal ridges more conspicuous than the transverse ridges, asymmetrically ovoid to

cylindric, often oblique at one end, reticulate with rectangular alveolae, alveolae covered by a thin iridescent

membrane, (0.42—)0.48—0.60(-0.70) mm long and (0.18—)0.22-0.26(-0.29) mm in diameter, (1.7—)2—2.6(-3)

times longer than wide. Chromosome number: 2N=18 (Gervais et al. 1999).

Phenology.—Flowering and fruiting from March to October

Common Name.—Clammy hedge-hyssop

Specimens Measured.— CANADA. ONTARIO: Thunder Bay District, 8 km SW of Thunder Bay City, 17 Aug 1978, Garton 18549 (ISC*).

QUEBEC: Montmorency Co.: Ange-Gardien, 23 Jul 1963, Cinq-Mers et al. 69-169 (UC*). Portneuf Co.: Portneuf, 7 Jul 1941, Rouleau

1045 (PH*). SASKATCHEWAN: 8 mi E of Saskatoon, 6 Jul 1950, Ledginham 890 (SMU*).

U.S.A. ALABAMA Co.: Smith Lake ( p) ca. 14 mi due WNW of Eutaw, 1 May 1980, Haynes 7775 (UNA*). Limestone

Co.: Beaverdam Creek 0.01 mi N of US Hwy 72 / Alt. 20 n Wheeler Wildlife Refuge, 20 May 1980, Meigs 555 (UNA*). ARIZONA.

Apache Co.: River Reservoir, Greer Lakes, 1.4 mi E of AZ Hwy 373, 2 airmi NE of Greer, and 9 airmi W of Eagar, 30 Aug 1988, Ricketson

& Raechal 4415 (MO*). ARKANSAS. Union Co.: El Dorado, 3 May 1940, Demaree 22048 (PH*+). CONNECTICUT. Hartford Co.:

Suffield, 20 Jun 1923, Weatherby s.n. (NCSC*). DELAWARE. New Castle Co.: 0.5 mi W of Glasgow, 15 Jun 1929, Benner 3572 (PH*).

GEORGIA. Bartow Co.: Pig Belfry Pond, 4.8 mi E of Adairsville, 5 May 1951, Duncan 12316 (US*). Walker Co.: Chickamauga, 16 May

1900, Biltmore 3913a (US*). ILLINOIS. Johnson Co.: Ferne Clyffe State Park; floodplain of Buck Branch, 21 May 1992, Mibb 692 (NLU*).

McHenry Co.: McHenry, 15 Jun 1925, Benke 4083 (US*). INDIANA. Vanderburgh Co.: 0.5 mi S of Staser, 26 May 1926, Deam 42953

(PH*). KANSAS. Cherokee Co.: 0.5 mi W of Crestline, 6 Jun 1970, Magrath 5352 (VDB**). Greenwood Co.: 1285, R13E, sec 9, edge

of temporary pool of valley in scrub oak woodland, 13 May 1987, McGregor 38094 (GA*). KENTUCKY. Warren Co.: along Warren Co.

Rt. 1288, ca. 1 mi from intersection with Warren Co. 961, 5 Jun 1968, Nicely 1666 (NCSC*). LOUISIANA. Richland Parish: beside S

side of I-20E ca. 1.7 mi W of the Rayville Exit (La. 137), 8 May 1990, Thomas 115,966 (TENN*). MASSACHUSETTS. Berkshire Co.:

Mount Washington, 25 Aug 1923, Meredith s.n. (PH*). Worcester Co.: Boylston, 24 Jun 1962, Richardson s.n. (MO*). MINNESOTA.

Carlton Co.: between Holyoke and Foxboro, 4 Jul 1942, Lakela 4986 (SMU*). MISSISSIPPI. Carroll Co.: field beside MS 7, at Avalon,

17 May 1973, Thomas & Marx 34783 (SMU*1). Washington Co.: ca. 3.5 mi NE Leland, 12 May 1988, Bryson 7637 (VPI*). MISSOURI.

Pulaski Co.: Falls Hollow Sandstone Glade, Ft. Leonard Wood, 13 May 1994, Hays 434 (MO*). MONTANA. Lake Co.: 4 mi Sand 2 mi

W of Ronan, 8 Jul 1956, Harvey 6517 (NCU*). NEVADA. Elko Co.: 0.8 road mi E of Deeth on the road to O'Neil Basin, backwaters of

the Marys River, 4 Jul 1986, Tiehm 10727 (BRIT*). NEW JERSEY. Cumberland Co.: Maurice River W of Bricksboro, 3 Jun 1934, Long

43311 (PH*). NEW MEXICO. Rio Arriba Co.: vicinity of Chama, 9 Jul 1911, Standley 6659 (US*). NEW YORK. Clinton Co.: Rouses

Point, 7 Aug 1910, Williamson s.n. (PH*). Monroe Co.: near Rochester, 4 Jul 1913, Baxter s.n. (MO*). NORTH CAROLINA. Cabarrus

Co.: Rocky River at NC Rd. 73, 25 May 1969, Daggy 5478 (TENN?). Caswell Co.: by Hyco Creek SE of Hightowers, 22 May 1958, Bell

11947 (NCU*). Chatham Co.: 3 mi W of Mann's Chapel on Co. Rd. 1536, 22 May 1974, Massey & Levesque 3988 (NCU*). NORTH

DAKOTA. Cass Co.: Harwood, 30 Jun 1937, Stevens 246 (GA*); 7 mi W of Enderlin, 28 Aug 1968, Barker 5213 (MO*). Richland Co.:

Estes and Small,

t North America 163

Petite VI f Gratiola f

Wyndmere, 18 Jun 1965, Stevens 2775 (US*). OHIO. Champaign Co.: Thackery, 11 Jun 1914, Leonard s.n. (US*). Crawford Co.: ca. 1.5

mi NW of Lykens, 30 Sep 1979, Stuckey 9962 (PH*). Erie Co.: W of Ceylon, Berlin Township, 15 Jul 1973, Jones 73-7-15-802 (TENN?).

OKLAHOMA. Le Flore Co.: along Poteau River, near Howe, 25 May 1931, Palmer 39340 (MO*). McCurtain Co.: near Harris, ca. 2 mi

N of the Red River, 20 Apr 1946, Nelson, Nelson, & Goodman 5579 (TEX*). OREGON. Crook Co.: Farewell Bend, 17 Jul 1894, Leiberg

456 (US*). PENNSYLVANIA. Chester Co.: French Creek near Hallman, 25 Jun 1927, Stone s.n. (PH*). SOUTH DAKOTA. Brookings

Co.: T112N R52W S32 SW4 SW4, restored prairie pothole wetland, 15 Jul 1991, Galatowitsch s.n. (ISC*). Custer Co.: Custer, 25 Jul

1892, Rydberg 924 (US*). TENNESSEE. Gibson Co.: floodplain of North Fork of Forked Deer River near jct. with Hwy 104, 6 Jul 1979,

Boom, Whitten, and Wofford 529 (TENN?). Giles Co.: NW side of Ardmore, N of Hwy 7 along N side of Austin Witt Rd. E of intersection

of Austin Witt Rd. and Union Hill Church Rd., 5 May 2001, Estes 02059 (TENN*). Hardin Co.: side of Pittsburgh Landing Rd., S of

Walker Branch, 18 May 1989, Guthrie & Tennesen 2235 (NCU*). Weakley Co.: E side of TN 89 along floodplain of Cane Creek, ca. 1.5

mi N of Palmerville, 25 May 1981, Webb 3919a (VDB*). TEXAS. Franklin Co.: 3 mi E of Mount Vernon, off US 67, 3 May 1945, Lundell

13701 (LL*). Jasper Co.: 9.3 mi NE of Burkeville, 14 Apr 1960, Shinners 27909 (SMU*). VIRGINIA. Giles Co.: Flat Top Mtn. near the

upper end of Pearis Thompson Branch, NE of Holly Brook, 7 Aug 1990, Wieboldt 7368 (NCU*). Warren Co.: Waterlick, 19 Jun 1924,

Pennell 12113 (US*). WASHINGTON. Klickitat Co.: Lyle, small shoal in Columbia River on the east side of the mouth of the Klickitat

River, 26 Aug 1993, Halse 4697 (K*). Spokane Co.: margin of Newman Lake, 2 Jul 1927, St. John 8811 (MO*). Whitman Co.: wet pond

beds, Pullman, 1 Aug 1896, Elmer 163 (USt). WEST VIRGINIA. Tucker Co.: 0.25 mi Sof Burley's Camp, Cabin Mtn. Range, 8 Jul 1941,

Allard 9055 (US**). Wetzel Co.: near Littleton, 1 Jul 1961, Haught 7127 (BRIT*). WISCONSIN. Lincoln Co.: Tomahawk Twp., 18 Jul

1950, Seymour 11687 (MO*). Taylor Co.: near Rib River, 22 Jun 1957, Schlising 648 (UC*).

Gratiola quartermaniae D. Estes, sp. nov. (Fig. 8). Ter: CANADA. Ontario. Hastings Co.: Tyendinaga Township, “Tod-

dary" alvar, Daley Road, ca. 7.5 km N of Lonsdale, 44.3404 N, 77.14539 W, moist open areas on alvar, with Eleocharis compressa,

Rumex crispus, Eleocharis obtusa, 22 Jun 2006, Oldham, Norris, & Van Sleeuwen 32809 (HoLoTYPE: TENN; isotypes: BRIT, CAN, DAO,

NHIC, NY, MO).

1 : : 1 In 1 : Nel

Gratiola quar termaniae a G. ic er differt] i veli infrequentel ramosis,

lineari PEE re elliptico-lanceolatis ginibus integris vel i icue dentata,

ad medium glabris; foli il fal

laminis plerumque uni- a trinervis; seminis parum longioribus crassioribusque, magis fusce brunneis.

Plants annual, solitary, erect herbs, (6-)11-22(30) cm tall. Roots simple, fleshy, whitish with numerous

rootlets. Stems erect, fleshy, simple or with few ascending branches, terete or slightly rounded-quadrangu-

lar in cross section, (0.6—)1-1.9(-2.3) mm in diameter at midstem; with 7-10(-11) leafy nodes, mid-stem

internodes (12—)19-35(-38) mm long, basal internodes shortened, 1-7 mm long; green or suffused with

reddish or reddish-pink pigments, especially near the base or upper nodes; glabrous or nearly so from the

base to above the middle, becoming sparsely glandular pubescent among the upper flower-bearing nodes

with spreading, translucent, slender-based gland-tipped trichomes. Leaves simple, oppositely-decussate,

similar in shape but gradually reduced in size from base to apex, lowermost often congested due to the

shortened internodes and sometimes early deciduous, linear, linear-lanceolate to elliptic-lanceolate, often

falcate, spreading or ascending, mostly with one evident main vein, sometimes trinerved with two short

secondary veins, rarely the two secondary veins well-developed, slightly fleshy-thickened, mid-cauline blades

(16218—32(-43) mm long and (1-)2.5-4(-4.5) mm wide, (5.5-)6-9.5(-11) times longer than wide, apex

acute or narrowly obtuse, widest near the middle, base sessile or slightly clasping; margins entire or each

margin with 1-2(=3) remote, low, bluntly pointed teeth beyond the middle; blades green, the basal blades

sometimes suffused with red; glabrous or nearly so. Flowers solitary in the axils of middle and upper bracteal

leaves, erect to spreading, zygomorphic, perfect; pedicels slender, ascending to divergent, (8213-22 mm

long, 0.5—1.1(-1.6) times as long as the subtending bracteal leaves, sparsely pubescent with slender-based

gland-tipped trichomes. Bracteoles 2, paired, closely subtending the calyx, equaling or to 2.3 times longer

than the sepals, linear-oblanceolate to linear-lanceolate and often falcate, one-nerved or inconspicuously tri-

nerved with two small lateral nerves, in fresh material bracteoles often appearing nerveless, fleshy-thickened,

in flower 2.8-8.2 mm long, lengthening in fruit to 11.8 mm long, 0.7-1.0 mm wide, apex obtuse, margins

entire, surface green, sparsely to moderately covered with slender-based gland-tipped trichomes. Calyx

irregularly campanulate with 5 subequal, distinct, lanceolate sepals, each inconspicuously longitudinally

three-nerved, in fresh material appearing single-nerved or apparently nerveless, fleshy-thickened, green,

2./—5.] mm long and 0.7-1.0 mm wide, apex obtuse, margins entire, sparsely covered with slender-based

gland-tipped trichomes. Corolla tubular-funnelform, gamopetalous, zygomorphic, slightly marcescent,

164 Journal of the Botanical Research Institute of Texas 1(1)

"ha 7

Fic. 8. Stu queres. ii Habit of G. ramos at type locality (photo by M. Oldham, : Jun 2000): B, is Powermng spe amen showing

stes 04359TENN; scale bar = 2 cm). C. FI y ( ). D. Capsule

(from Oldham et al. 32877TENN: scale bar = 3.5 mm). E. Close-up of mid-stem (scale bar = 2 m).

6-13.7 mm long; corolla tube quadrangular, dorsal surface with a prominent hump near midpoint, the

ventral surface canaliculate, to 9.3 mm long, 1.3-2.7 mm in diameter proximal to dorsal hump, greenish-

yellow, creamy yellow, or bright yellow, with many brownish-purple lines extending the length of the tube,

Estes and Smal

I M : Lr PE peer of

, Petite VI NIauvia

t North America 165

sparsely pubescent externally with slender-based gland-tipped trichomes, inner surface near orifice at base

of posterior corolla lobe with moderate to dense beard of clavately thickened yellow trichomes, proximal

and median inner corolla surfaces pilose with eglandular trichomes up to 0.8 mm long; corolla lobes 5,

generally slightly broader than high and emarginate, white, 2.0—3.6 mm long and 2.4—5.4 mm wide; the

posterior lobe generally largest, the two lateral lobes and lower lobe equal or slightly smaller, adaxial surfaces

glabrous, abaxial surfaces glabrous or slightly glandular pubescent. Stamens 2, inserted near the middle

of the dorsal surface of the corolla tube, filaments 0.8-1.5 mm long, anthers transversely oriented to the

filaments, 0.8-1.3 mm long and 0.5-0.9 mm wide, connective whitish and greatly dilated around the two

anther sacs; staminodes inserted 1.4—1.8 mm above base of corolla or absent, when present to 0.3 mm long,

not capitate. Gynoecium 5.9-7.6 mm long, subtended at the base by an orange nectary disc, ovary 1.6-4.3

mm long and 1-2.8 mm in diameter, style 3.1—4.6 mm, stigma 2-lobed, dilated and flattened, 0.6-0.9 mm

long. Capsules ovoid, apex acute, usually widest below the middle, (3.4—)3.6—4.7(—5.1) mm long, 2.9-4.5

mm in diameter, brown at maturity. Seeds several hundred per capsule, grayish-brown to reddish-brown,

10-13 ribbed, longitudinal ridges more conspicuous than the transverse ridges, asymmetrically ovoid to

oblong-cylindric, often oblique at one end, reticulate with rectangular alveolae, alveolae covered by a thin

iridescent membrane, (0.43—)0.55-0.63(-0.71) mm long and (0.19-)0.26-0.32-0.37) mm in diameter,

(1.5-)1.8-2.3(-2.6) times longer than wide. Chromosome number unknown.

Phenology.—Flowering and fruiting from April to early June in Alabama, Tennessee, Texas and from

June to August in Illinois and Ontario, Canada

Etymology.—This species is named in honor of Dr. Elsie Quarterman, retired Vanderbilt University

plant ecologist, who has dedicated her career to the study of the ecology of the limestone cedar glades and

the species that inhabit them.

Common Name.—Quarterman's hedge-hyssop; limestone hedge-hyssop.

Conservation Status.—Gratiola quartermaniae is most common in the limestone cedar glades of middle

Tennessee where it is known from ca. 30 populations in nine counties. Although it appears to be secure

in Tennessee, the mid-state area where this species occurs is one of the most rapidly developing regions in

the southeastern U.S. and the once abundant glade habitat preferred by this species is increasingly being

destroyed. Consequently, while G. quartermaniae is not sufficiently rare in Tennessee now to warrant state

or federal conservation status, its populations should be monitored in the next few decades. In Alabama,

Illinois, Texas, and Ontario this species appears to be quite rare and is restricted to small geographic areas.

In these regions it should be afforded protection at the state or provincial level.

Representative Specimens.— CANADA. ONTARIO. Hastings Co.: Belleville, May 1861, Macoun [number illegible] (K); vicinity of

Belleville, Jun 1867, Macoun 17454 (CAN*); flats near the Iron Bridge at Belleville, Jun 1868, Macoun 41730 (CAN); Belleville, 10 Jun

1871, Macoun 123 (TRT); Belleville, 24 Jun 1871, Macoun 1261 (DAO); Pt. Anne, Belleville, Ontario, 13 Jun 1972, Morton 5091 (CAN, QK,

TRT, WAT); Hungerford Township, Larkins Alvar, ca. 9.5 km SE of Tweed, ca. 1.5 km SW of Larkins, S of Marlbank Rd., 30 Jun 2006,

Oldham, Norris, & Van Sleeuwen 32877 (DAO, MICH, MO, NHIC, TENN, US); Richmond Township, Roblin Dump alvar, ca. 1.5 km SE of

Roblin, ca. 9 km SSE of Marlbank, 30 Jun 2006, Oldham, Norris, Sutherland & Van Sleeuwen 32869 (CAN, MICH, MO, NHIC, NY, TENN,

TRTE, US, UWO). Lennox and Addington Co.: Camden East Township, ca. 10 km NW of Newburgh, ca. 15 km N of Napanee, road

to Roblin Hell Holes, off Centreville Road, 30 Jun 2006, Oldham 32868 (BRIT, CAN, DAO, HAM, MICH, MO, MT, NHIC, TENN, US,

VDB). Peterborough Co.: alvar ca. 2 mi N of Nogies Creek in Harvey Tp., 11 Jul 1974, Catling & McKay s.n. (CAN, TRT); 1.79 air mi NE

of Nogies Creek, 1.1 air mi NNW of jet. of Co. Rd. 36 and Quarry Rd., 0.37 road mi NW from jet. of Quarry Rd. and Ledge Rd., 18 Jun

2005, Estes 07955 (CAN, DAO, NY, MICH, TENN, VDB). Prince Edward Co.: Big Sand Bay, Long Point, 7 Jun 1963, Brassard & Hainault

2702 (CAN*, TRT); ca. 2 mi SE of Milford, 5 Aug 1951, Soper & Heimburger 5412 (TRT); South Marysburgh Township, Hilltop Rd., ca. 5

km SE of Milford, near South Bay, 19 Jun 2006, Oldham 32786 (DAO, MICH, MO, NHIC, NY, TENN, TRT, US).

U.S.A. ALABAMA. Franklin Co.: ca. 5-6 mi E of Russellville along N side of New Hwy 24, just W of jet. of New Hwy 24 and

County Rd. 83, 15 May 2003, Estes 04625 (TENN). Lawrence Co.: by Ala. 36 ca. 2 mi. e. jct. Ala. 157, 6 May 1978, Kral 61662 (JSU,

VDB*); approx. 4 mi NW of Mt Hope, ca. 1.5-2 mi E of Franklin County line, W of Town Creek, at Prairie Grove Glades preserve, 15

May 2003, Estes 04611 (TENN?*); ca. 0.2 to 0.4 mi ESE of Landersville, south of junction of Hwy 24 and County Rd. 55, growing in wet

ditch over limestone on west side of County Rd. 55, 34?28'09" N, 87?23'46"W, 29 Apr 2004, Estes 05928 with Webb (CAN, MO, TENN,

UNA). Morgan Co.: 5.6 mi. W of Falkville, 23 Apr 1968, Kral 30494 B (GA, VDB*); seep in sandy clay field 1 mi E jct AL 157 by AL

36, W of Danville, 14 Apr 1978, Kral 61500 (JSU, MO, VDB); N side of Morgan Co Rd 55, 0.9 mi E of Massey (McKendree Church),

f*hAD o ID L

166 Journal of t titute of Texas 1(1)

2.1 mi W of Lebanon Church and 6 mi W of int. US 31 at Falkville, 28 Apr 1989, Orzell & Bridges 9380 (TEX*). ILLINOIS. Will Co.:

Romeo, 18 Jun 1898, Umbach s.n. (US). TENNESSEE. Bedford Co.: N side Deason by US 231, 28 Apr 1974, Kral 52571 (MO, VDB);

0.2 mi N of US 41A at Rover along Bunker Hill Rd, 3 Jun 1993, Kral 82558 with Rust (VDB); approx. 5 mi NE of Unionville, ca. 0.75 mile

S of Newtown, near intersection of Longview Rd. and Putnam Well Rd., on east side of Longview Rd., 22 May 2003, Estes 04583 with

Wofford et al. (CAN, GH, TENN?). Cannon Co.: by US 71S, 0.5 mi E of Readyville, 20 May 1974, Kral 52812 (MO, VDB*). Coffee Co.:

Manchester prairie, 4 mi E of Manchester on US 41, 7 Jun 1966, Baskin & Caudle 258 (VDB). Davidson Co.: Hamilton Creek Recreation

Area, SE side of Nashville, W of Percy Priest Lake, E side of Ned Shelton Rd., 15 Jun 2003, Estes 04894 (EKY, GA, JSU, TENN*, UNA).

Giles Co.: S of Pulaski, Cedar Grove community, growing on W side of Hwy 166, south of Everly Branch and just N of Cedar Grove

Church, 18 Apr 2003, Estes 04454 (TENN*). Marshall Co.: 2.1 mi ESE Pottsville on TN 99, 2 Jun 1969, Kral 34776 (MO, VDB*); N side

TN 99, just inside W county border, 14 May 1988, Kral & Kral 74722 (VDB); approx. 4 mi NE of Chapel Hill near Beasley community,

ca. 100—200 yards east of intersection of Hwy 99 and Beasley Rd., S side of Beasley Rd., 22 May 2003, Estes 04582 with Wofford et al. (GH,

MO, NCU, NY, TENN}, TEX, UC). Maury Co.: ca. 2 mi NW of Pottsville, 1.5 mi NE of jct of Hwy 412 and Rally Hill Rd., E side of Rally

Hill Rd., 22 May 2003, Estes 04672 with Wofford et al. (TENNT). Rutherford Co.: 10 mi. E Beech Grove along US 41, 9 Jun 1970, Kral

26889 (FSU*, SMU, TENN, VDB); SE of Eagleville, 1 mile off S.R. 99, 28 May 1996, Rust 66 (VDB*); WSW of Fosterville, ca. 2 mi W of

US Hwy 231, 0.33 mi N of Squire Hall Rd., E side of Harrison Rd., 22 May 2003, Estes 04586 with Wofford et al. (NCU, TENNTt, VDB);

E of Murfreesboro, approx. 1 mi SE of Halls Hill Pike, S side of Factory Rd., Flat Rock Cedar Glade and Barrens State Natural Area, 22

May 2002, Estes 03337 (TENN*); approx. 4 mi E of Murfeesboro on Hall Hill Pike, turn S onto Smith Hall Rd. (a dead-end road), E side

of road, 22 May 2002, Estes 03336 (TENN*); N Murfeesboro, ca. 1 mile W of intersection of E Northfield Blvd. and Hwy 96, 22 May

2003, Estes 04574 with Wofford et al (TENN*); approx. 4—5 mi E of Murfreesboro, W side of Factory Rd., Flatrock Cedar Glades/Barrens

State Natural Area, 1 May 2003, Walch s.n. (TENN*); base of Garrett Knob, 29 May 2003, Bailey & Lincicome s.n. (TENN). Wilson Co.:

Lebanon, 2 Jun 1923, Pennell 11377 (PH); Cedars of Lebanon State Forest and Natural Area, N of Moccasin Rd. / Proctor Trail, 8 May

2003, Bailey s.n. (TENN). TEXAS. Bell Co.: 6 mi SE of Belton, Wolff 2317 (SMU). Llano Co.: Llano River east of Packsaddle Mountain,

4 May 1947, Whitehouse 18477 (SMU, UC, US). Williamson Co.: Round Rock, 24 March 1890, Bodin s.n. (PH, MIN-digital image); ca.

3.9 mi SSW of Liberty Hill, along CR 284, 1.3 mi W of jct CR 282, S side rd, 29 Apr 2005, Turner & Turner 122 (BRIT, MO, TENN, TEX);

southern part of co., just NW of Round Rock, FM 1431 at jct Sam Bass Rd., SE corner, 150 m S of FM 1431, 29 Apr 2005, Turner & Turner

119 (BRIT, GH, MO, TENN, TEX).

Gratiola graniticola D. Estes, sp. nov. (Fig. 9). Tye: U.S.A. Grorcia. DeKalb Co.: Rock Chapel, GA hwy 124 at Rock Chapel

County Park, gneiss flatrock, W side of highway, vernal pools, 2 May 1984, Allison 2101 (HOLOTYPE: GA).

uu 7; a be

Gratiola graniticola a G. neglecta Torr. differt herba trichomatibus brevioribus basi bulbosis , simplicibus vel

Ee Wr NE M" À

| lanceolat ti ] te obl i inil bi i l

n O

Ovat

L o L D E o o o o E

dentatis, basibus magis valde amplectentibus; pedicellis folia bractealia subtendentia aequantibus vel eos duplo longioribus; bracteolis

calycibus brevioribus vel eis vix superantibus; floribus minoribus lobis posterioribus purpurascentibus, barba in corollae orificio e

SAT -1 seseque 1 “1 " ANE 1: E +1 : 1.1.1 ` Toa Ad E A

de 2 D. eo P : O

obscure cinereis.

Plants annual, solitary, erect herbs, (7-)9-21(229) cm tall. Root simple, fleshy, whitish with numerous rootlets.

Stems erect, somewhat fleshy, simple or with few ascending branches, terete or slightly rounded-quadrangular

in cross section, (0.72)0.9-1.2(-1.5) mm in diameter at midstem; with (6—)7—10(-12) leafy nodes, mid-inter-

nodes (15—)17-30(-36) mm long, basal internodes shortened (1.5-8 mm); green or suffused with reddish or

reddish-pink pigments, especially near the base and upper nodes; glabrous or glabrate near base becoming

increasingly pubescent upward, with spreading, translucent, conical or bulbous-based, glandular trichomes.

Leaves simple, oppositely decussate, similar in shape but gradually reduced in size from base to apex, low-

ermost often congested due to the shortened internodes and sometimes early deciduous, lanceolate-ovate

to narrowly oblong usually widest at or below the middle, horizontally spreading with tips curved upward,

with one evident main vein or trinerved with two short secondary veins, slightly fleshy-thickened, blades

(6-)7-13(-18) mm long and 1—3(-5) mm wide, (2.8—)3.5—5.7(-7.4) times longer than wide, apex narrowly

obtuse, margins entire or with 1-2(3) pairs of remote, low, bluntly pointed teeth beyond the middle, base

usually amplexicaulate; blades green or leaf tips, teeth, and basal leaves often suffused with reddish pigments;

proximal leaves glabrate, median and distal leaves moderately pubescent with bulbous based trichomes.

Flowers solitary in axils of upper bracteal leaves, erect to spreading, zygomorphic, perfect; pedicels slender,

ascending, (5-)8-17(-22) mm long, (0.9—)1-2(-2.3) times as long as the subtending bracteal leaves, sparsely

to moderately pubescent with bulbous based trichomes. Bracteoles 2, paired, closely subtending the calyx,

usually shorter than or equaling the sepals, lanceolate and often falcate, longitudinally 3-nerved (sometimes

single nerved) though not often evident when fresh, fleshy-thickened, 274.5 mm long and 0.5-1.0 mm wide,

Estes and Small, New species of Gratiola f tern North America 167

Fic. 9. Gratiol. iticola. A. Habit, in Butts Co , Georgia, 10 Apr 2004 B.U [ t ith fl li t le (scalel 6 ). C. Leaf (scale

bar= > mm); D. Flower oneview scale bar = 2 mm). E. Immat psule with subtendi | | bracteoles (scale bar = 3.5 mm). F. Unopened

—4.5 mm).

Jt F

168 Journal of the Botanical R h Institute of Texas 1(1)

apex obtuse, margins entire, surface green, apex purple-tipped, abaxial surface convex, moderately covered

on both surfaces with bulbous-based trichomes. Calyx irregularly campanulate with 5 subequal, distinct,

lanceolate sepals, longitudinally 3-nerved (sometimes single nerved) though not often evident when fresh,

fleshy-thickened, green on the surface with a minute purple tip, 274.2 mm long and 0.5-1.3 mm wide, apex

obtuse, margins entire, moderately covered, especially abaxially, with conical or bulbous-based trichomes.

Corolla tubular-funnelform, gamopetalous, slightly marcescent, and zygomorphic, 6.8-9.0 mm long; co-

rolla tube quadrangular, dorsal surface with a hump near midpoint, 5.5-6.8 mm long and 1.3-1.9 mm in

diameter, outer surface pale yellowish-green or cream-colored, often purplish or pinkish dorsally, faintly

to conspicuously purple-lined exteriorly, scarcely pubescent with conical or bulbous-based trichomes, in-

ner surface near orifice at base of posterior corolla lobe with sparse beard of clavately thickened whitish to

translucent trichomes, proximal and middle inner corolla surfaces pilose with eglandular trichomes; corolla

lobes 5, each usually broader than high and often emarginated at apex, spreading, the lower three white or

cream-colored, the upper two strongly suffused with purple or pink, the lobes 1.0-1.7 mm high and 1.5-2.3

mm wide, adaxial surfaces glabrous, abaxial surfaces glabrous. Stamens 2, inserted near the middle of the

dorsal surface of the corolla tube, filaments to 1.2 mm long, anthers transversely oriented to the filament,

0.5-0.7 mm long and 0.5-0.6 mm wide, connective greatly dilated around the two anther sacs, whitish;

staminodes inserted ca. 1-1.3 mm from base of corolla tube or absent, when present minute and ca. 0.2 mm

long, not capitate. Gynoecium 4.4-4.8 mm long, subtended at the base by an orange nectary ring, ovary

1.6-2.0 mm long and 1.2-1.7 mm in diameter, style 1.9-2.2 mm long, stigma 2-lobed, dilated and flattened,

0.5-0.6 mm long. Capsules subglobose to slightly ovoid, (2.4-)2.8-3.6 mm long, 2.1-3.7 mm in diameter,

purple tinged when mature. Seeds several hundred per capsule, brown to grayish-brown, 10-13 ribbed,

longitudinal ridges more conspicuous than the transverse ridges, asymmetrically ovoid to short cylindric,

often oblique at one end, surface reticulate with rectangular alveolae, alveolae covered by a thin iridescent

membrane, (0.3-)0.36-0.42-0.47) mm long and (0.17-J0.2-0.24-0.27) mm wide, (1.321.6-2.1(-2.5)

times longer than wide. Chromosome number unknown.

Phenology.—Flowering and fruiting from April to May.

Etymology.—The ephithet graniticola was chosen to reflect the granite flatrocks that this species inhabits.

Common Name.—Granite hedge-hyssop.

History of Taxon.—Gratiola graniticola was apparently first collected in 1928 (Wherry & Benedict s.n. PH)

from “pools on granite ledges” in Gwinnett County, Georgia. A decade later, Pyron and McVaugh (2866 GA,

PH), collected a specimen of G. graniticola from granitic areas in Oglethorpe County, Georgia. McVaugh sent

a specimen of this Oglethorpe County collection to F.W. Pennell who wrote “your collection, with that of

Wherry and Benedict...differ from G. neglecta Torr. by bracts shorter relative to pedicels, capsules smaller

(3 mm long), upper corolla-lobes purple or purplish, and seeds smaller and grayer” (McVaugh 1943). He

added that these specimens seemed to match his description and photograph of G. gracilis Benth., a species

described by Bentham (1846) from Texas.

Bentham (1846) described G. gracilis Benth. from material collected by Drummond near Harrisburgh,

Texas (near present-day Houston) in ca. 1834. Unfortunately, Drummond failed to note the habitat from which

he collected the plants. A second specimen annotated by Pennell as G. gracilis was collected by Lindheimer

(43 MO) from nearby Galveston in ca. March (May?, illegible) 1842. Like Drummond, Lindheimer did not

provide specific locality or habitat information. Despite being known only from herbarium specimens, G.

gracilis was maintained as a species by Small (1903) and Pennell (1921). Later, Pennell (1935) reduced G.

gracilis to synonymy with G. neglecta noting the characters Bentham used to distinguish G. gracilis from G.

neglecta “are all variable features that occur without geographic correlation.”

During this study, a photograph of the holotype of G. gracilis (Drummond coll. 3, n. 284, K) and an isotype

(GH) were examined. As Pennell noted, these specimens do share some features with those plants from the

Georgia granite outcrops, most notably in the length of the leaves and the ratio of the length of the pedicel

and subtending bract. While of rare occurrence, G. neglecta can have relatively short leaves and bracteal

Estes and Smal

I M : Lr ie) peer of

, Petite VI NIauvia

t North America 169

leaves shorter than the pedicels (e.g., Guthrie 1002 VDB, Lake Co., TN). The three G. gracilis specimens also

differ from G. graniticola in that they lack purple coloration on the corollas and capsules, features diagnostic

for G. graniticola. In terms of habit, the stems of the G. gracilis specimens are more branched like those of G.

neglecta compared to those of G. graniticola, which are mostly simple. Lastly, G. graniticola is endemic to granite

outcrops and has not been found in non-granitic areas. Since there are no granite outcrops in southeastern

Texas, it is reasonable to assume that the plants collected by Drummond and Lindheimer likely came from

a different habitat type. Based on the evidence presented above, we follow Pennell and recognize G. gracilis

as a synonym of G. neglecta.

Conservation Status.—Gratiola graniticola should be considered a rare species in Georgia due to the small

number of populations and limited distribution.

Representative Specimens Examined.—U.S.A. GEORGIA. Barrow Co.: Winder, GA Hwy 81, roughly 0.25 mi S of junction with US

Hwy 29, E side of highway, 30 Apr 1984, Allison 2095 (GA*); same site, 19 May 2003, Estes 04590 with Allison (TENN**). Butts Co.: ca.

2.7 mi NNE of Jackson, GA Hwy 36, ca. 0.5 mi S of Cedar Rock Church, E side of highway, 13 May 1984, Allison 2175 (GA*); same site,

10 Apr 2004, Estes 05742 (TENN). Columbia Co.: ca. 4.25 mi ESE of Appling, ca. 0.45 mi NNW of confluence of Little Kiokee Creek

and Benton Branch, adjacent to Heggies Rock Preserve, 10 May 1987, Allison 2842 (GA*). DeKalb Co.: across from Rock Chapel Park,

4 mi N of railroad track in Lithonia, along State Hwy 124, 16 Apr 1978, Patrick 592 with Wofford et al. (TENN*); Lithonia, ca. 0.3 mi N

of intersection of Interstate 20 and Hwy 124, NW side of the intersection of Hwy 124 and Conyers Street on small concealed granite

outcrop, 10 Apr 2004, Estes 05733 (TENN); same site as previous, 01 May 2004, Estes 05954 (TENN*, MO, NY). Greene Co.: 8.2 mi

SSE of Greensboro, 5.8 mi W of White Plains, 2 May 1987, Allison 2834 (GA*); ca. 9 mi SSE of Greensboro, ca. 1.5 mi SW of Mosquito

Crossing, S side of Leach Flatrock Rd., 33.46738 N, 83.13214 W, 19 May 2003, Estes 04585 with Allison (NCU, TENNT). Gwinnett Co.:

6 mi SW of Grayson, 3 May 1928, Wherry & Benedict s.n. (PH); 4.25 mi E of Snellville, 2.25 mi SSE of Grayson, Langley Rd., 0.34 mi by

air NW of junction with US Hwy 78, E side of road, 13 Jun 1984, Allison 2306 (GA). Hancock Co.: 3.5 mi SE of Sparta, 11 May 1952,

Duncan 13533 (GA, digital image); ca. 1 mi or less NE of Sparta, 0.3 mi N of Hwy 16, 0.3 mi W of Twomile Creek, 33.29098 N, 82.95428

W, Estes 04659 with Allison (TENN*). Hart Co.: 5.3 mi NNE of Vanna, 1.5 mi NNE of Goldmine, ca. 0.2 mi E of county road 141 at a

point ca. 0.45 mi NW of junction with county road 140, 15 Apr 1986, Allison 2625 (GA*); same site, 19 May 2003, Estes 04588 with Allison

(NCU, TENN?). Newton Co.: ca. 3 mi NE of Covington, ca. 1.25 mi NE of the intersection of Hwy 142 and Alcovy Rd., S side of Alcovy

Rd., 19 May 2003, Estes 04584 and Allison (TENN?*); same site, 10 Apr 2004; Estes 05738 (TENN). Oglethorpe Co.: Echols’ Mill, May

1938, Pyron & McVaugh 2866 (GA, PH); ca. 0.5 mi E of Echols’ Mill, ca. 9.3 mi N 45 deg. of Lexington, 7 May 1978, Treiber & Nesom 1518

(NCU*). Pike Co.: 1.6 mi S of Hollonville on Concord Road, E side of road, 19 May 1984, Allison 2254 (GA*); same site, 01 May 2004,

Estes 05953 (MO, NY, TENN). Upson Co.: NE corner of county, ca. 0.4 mi S of Lamar County line and just E of Barnesville-Yatesville

Rd., 18 May 1984, Allison 2235 (GA*). Walton Co.: 4.9 mi WNW of Walnut Grove, Ace Moon Road (county road 197), just S of junction

with Sharon Church Road (county road 106), E side of road, 11 May 1984, Allison 2141 (GA*); by GA 138, 1 mi. NE of Walnut Grove, 17

May 1989, Kral 72517 (FSU, GH, VDB*).

ACKNOWLEDGMENTS

We appreciate the work of two anonymous reviewers whose suggestions improved this manuscript. Funding

for this research was provided by three University of Tennessee endowment funds, the Breedlove-Dennis

Fund, the Sharp Fund, and the Hesler Fund. Additional funding was received from the Southern Appala-

chian Botanical Society (Core Award) and a University of Tennessee Department of Ecology and Evolutionary

Biology summer research grant. Gene Wofford, Ron Petersen, Ken McFarland, Jeff Walck, Kurt Blum, John

Beck, and Ed Lickey provided advice or encouragement. We are also grateful to the following individuals

who assisted with field work or specimen collection: Jim Allison, Matt Turner, Billie Turner, Bill Carr, John

Beck, David Webb, Michael Oldham, Eric Ulazsek, Steve Hill, Claude Bailey, Roger McCoy, and David

Lincicome. John Dunlap assisted with the SEM work. Chris Fleming assisted in the preparation of the spe-

cies distribution maps. Mark Garland prepared the Latin diagnoses. We also wish to thank the curators of

the herbaria that loaned specimens for this study, facilitated herbarium visits, or provided us with digital

images of specimens.

REFERENCES

Baskin, J.M. and C.C. Baskin. 2003. The vascular flora of cedar glades of the southeastern United States and its

phytogeographic relationships. J. Torrey Bot. Soc. 130:101-118.

BENTHAM, G. 1846. Scrophulariaceae. In: de Candolle, A. Prod. Syst. Nat. Regn. Veg. X. Paris: Victoris Masson.

170 Journal of the Botanical R h Institute of Texas 1(1)

CHAPMAN, LJ. and D.F. Putnam. 1984. The physiography of southern Ontario. Third Edition. Ontario Geological

Survey. Special Volume 2.

Gervais, C., R. TRAHAN, and J. GAGNON. 1999. IOPB chromosome data, 14. Newslett. Int. Organ. Pl. Biosyst. (Oslo)

30:10-15.

Mason, H.L. and R. BacicaLuPi. 1954. A new Gratiola from Boggs Lake, Lake County, California. Madroño 12:

150-152.

McVaucH, R. 1943. The vegetation of the granitic flatrocks of the southeastern United States. Ecol. Monogr.

13:121-166.

PENNELL, F.W. 1921. Scrophulariaceae of the West Gulf States. Proc. Acad. Nat. Sci. Phil. 73:471-477.

PENNELL, F.W. 1935. The Scrophulariaceae of eastern temperate North America. Philadelphia: The Academy of

Natural Sciences of Philadlphia. Small, J.K. 1903.

SMALL, J.K. 1903. Flora of the southeastern United States. Published by the author, New York.

Stuckey, R.L. 1979. Type specimens of flowering plants from eastern North America in the herbarium of Lewis

David von Schweinitz. Proc. Acad. Nat. Sci. Philadelphia 131:9-51.

SUOMINEN, J. 1984. Gratiola neglecta (Scrophulariaceae), Mantsalanjoen rantakasvi. (Gratiola neglecta, a North

American wetland plant naturalized in Finland). Mem. Soc. Fauna Flora Fenn. 60:5-9.

Torrey, J. 1819. Catalogue of plants, growing spontaneously within thirty miles of the city of New-York. Lyceum

of Natural History of New York.

WEAkLEY, A.S. 2007. Flora of the Carolinas, Virginia, Georgia, and surrounding areas, working draft of 11 January

2007. University of North Carolina Herbarium, North Carolina Botanical Garden, Chapel Hill.

REVIEW OF CRATAEGUS SERIES APRICAE, SER. NOV., AND C. FLAVA (ROSACEAE)

J.B. Phipps and K.A. Dvorsky

Department of Biolog

The University of Western Ontario

London, Ontario, N6A 5B7, CANADA

jphipps@uwo.ca

ABSTRACT

This paper revises Crataegus ser. Apricae, ser. nov., remodeled from old ser. Flavae after the removal of C. flava. Twelve species are

recognized in the series, plus a number of other forms which represent possibly undescribed taxa, taxa only known from type gatherings

and one more of doubtful serial assignation. Full descriptions and synonymy as well as complete typification are provided for all taxa

fully treated. The principal species all have line illustrations and county level distribution maps. Keys distinguish all taxa and unnamed

forms recognized. Crataegus flava, although transferred to ser. Intricatae, is treated here for convenience.

Key Worbs: Crataegus series Apricae, ser. nov., taxonomic revision, C. flava, ser. Intricatae

RESUMEN

En este artícul isa Crataegus ser. Apricae, ser. nov., remodelado de la antigua ser. Fl después de eliminar C. flava. Se recono-

cen d la serie, además de cierto número de formas que probablemente representan taxa no descritos, taxa

de Tu del tipo y uno más de asignación dudosa a la serie. Se aportan descripciones completas y sinonimia así como eee

de todos los taxa tratados. Las principales especies estan ilustradas y tienen mapas de distribución a nivel de condado. En las claves se

distinguen todos los taxa y las formas no nombradas que se reconocen. Crataegus flava, aumque se haya transferido a la ser. Intricatae,

se trata aquí por conveniencia.

INTRODUCTION

This paper is the sixth in a series reviewing hawthorns of the southeastern United States, a region for a long

time lacking proper revisions after the huge burst of activity at the turn of the last century which culminated

in Beadle's (1903) seminal contribution to Small’s regional flora. Beadle's treatment of 185 Crataegus species

for the region, many described by Beadle himself, was based on the collection of hundreds of specimens for

the Biltmore Herbarium in which endeavor Beadle was ably assisted by T.G. Harbison. Beadle thus devel-

oped unparalleled personal experience although his species concepts have been considered overly narrow.

Only one later treatment covered Crataegus for the entire region, this being by Tidestrom (1933) in which he

recognized 33 species in J.K Small’s new flora of the area. As is pointed out in earlier works, e.g., Phipps and

Dvorsky (2006), Tidestrom omitted entire series and even the unique species C. triflora although it can be

found, grossly incorrectly synonymized, under C. intricata. The almost complete lack of synonymy further

reduces the value of Tidestrom's taxonomy. The Crataegus expert Palmer in Vines (1960), a work which covers

the ‘southwest’, an area deemed to reach east to the Mississippi, produced a treatment with 71 species, only

30 of which are in the southeast as routinely interpreted in my papers and which reaches west to Louisiana

and Arkansas following Cronquist (1980). Vines' work is illustrated with woodcuts that singularly fail to

distinguish any but the most dissimilar species of hawthorn and, largely lacking series Flavae in the old

sense, therefore does not significantly contribute Palmer's usually valuable insights to the matters addressed

in this paper. Later floristic workers produced treatments more in the vein of Tidestrom, as is evidenced, for

instance, by their taxon selection and routine omission of C. triflora, when it occurred in their areas. The

result has been that the southeastern United States, an area very rich in Crataegus, has become floristically

the worst-served part of the flora of North America area for this genus. Only a few floristic writers bucked

this trend, Kurz and Godfrey's (1982) Crataegus treatment in their “Trees of Northern Florida” being a good

example of this, although, even here, it appears that the authors did not consult types. Murrill’s closely

observed descriptions of northern Florida hawthorns in the early 1940s, e.g., Murrill (1942), are difficult to

match with known species and may represent extremes of variation of them and are here mainly ignored.

J. Bot. Res. Inst. Texas 1(1): 171 — 202. 2007

172 Journal of the Botanical R h Institute of Texas 1(1)

It is with this background that examination of over 10,000 specimens for the first author's studies in

the southeastern United States Crataegus flora, together with numerous field trips to the region which yielded

many personal (JBP) collections, as well as the receipt of over 500 duplicate specimens from R. Lance, in

addition to experience derived from earlier papers in this series fully confirms Beadle's position that the

southeastern United States is a region of great species richness for Crataegus. Consequently, treatments like

that of Tidestrom can at best be only accepted in part. Nevertheless, a parallel realization is that the Cratae-

gus taxonomy of the region is not particularly straightforward even though basic attention to type material

permits attaching an appropriate name to nearly all morphotypes encountered. This is so because nearly

all Beadle's names can readily be lectotypified if necessary even if those of Ashe cannot (and neotypifica-

tion of the latter's names is usually fraught with difficulty) and Beadle's names appear sufficient to account

almost completely for observed variation. This may all be observed in miniature in the current revision of

ser. Apricae.

Crataegus series Apricae represents a remodeling of series Flavae (Loud.) Rehder in the sense used in

Palmer (1925), Phipps et al. (2003) and Phipps et al. (1990). In the last two publications mentioned the

species lists for ser. Flavae were understood to be incomplete, the studies presented here being then worked

up. There proved to be more difficult taxonomic problems associated with series Apricae than for any other

series dealt with so far by the first author for the southeastern United States Crataegus flora. These may be

listed as follows.

[1 RAS. 1 a]

1. For many Flavae as a very inclusive concept, this generally speaking, being collapsed from the 81 spe-

cies of Beadle (1903) which tl thor had ized into 13 groups. Recently, however, the first author (Phipps 19882), removed most of

these groups to ser. Lacrimatae and it TUNE d became necessary to ask whether this action left the residual ser. Flavae homogeneous.

2. Crataegus flava, type species of ser. Flavae, appears to be easily the most distinct species of the series whether or not ser. Lacrimatae

is included in ser. Flavae. This paper formalizes the position, previously suggested in Phipps (19883), that C. flava, the type species,

should be removed from the remainder of old ser. Flavae.

3. Subsequently, therefore, is the new residuum, after both ser. Lacrimatae and C. flava are removed, sufficiently homogeneous to exist

as a single series?

4. If yes, the question then arises as to an appropriate name for the residual series.

5. None of the species of the series as finally demarcated below appears to be really common and several are very rare or local, limiting

the effect of insights from modern fieldwork

6. Species limits in the new series were sometimes found to be not particularly clear cut, in part b paucity of material

which always makes taxonomic decisions more difficult. This problem is heightened with several species in the series being generally

less well-marked than is often the case, particularly in the visenda cau Nevertheless, uniting such species carries its own difficulties,

especially with the phenetic breadth of th ted species and the fact that variation d flow smoothly through-

out. On the other hand, species such as C. frugiferens would be difficult to unite with anything.

Nevertheless, the study of 274 specimens, including 20 types, from 26 herbaria plus the additional un-

derstanding gained from modern fieldwork has created the opportunity to offer the following elaboration

of the first author's treatment that will appear in Flora of North America, vol. 9. It is rooted in the work of

C.D. Beadle (in Small 1903), who knew the group better than anyone else, and who remains the only guide

among the earlier generations of botanists. Also note that the first full and accurate description of Crataegus

flava is provided here for comparative purposes.

TAXONOMIC TREATMENT

The treatment presented here follows the form established for other series of the southeastern United States

hawthorns (Phipps 1988a, 1988b; Phipps & Dvorsky 2006a, etc.). This entails detailed series and species

descriptions, key to species, full typification, line illustrations of the taxa and county level distribution

maps prepared by K. Dvorsky. There is also an appendix of cited specimens. The treatment will commence

by elaborating the separation of the new series. A few extra recent collections were added to the list of cited

specimens in proof and do not appear in the maps.

The reasons for the exclusion of Crataegus flava and the separation of new ser. Apricae from ser. Lacri-

matae are most conveniently summarized as a detailed key, given below. Crataegus flava will be placed in

Phipps and Dvorsky, Crataegus ser. Apricae and C. flava 173

ser. Intricatae in the FNA treatment. A combination of plant habit, indumentum, leaf shape, and anther color

characters are emphasized.

KEY SEPARATING SER. APRICAE FROM SER. EACRIMATAE AND C. FLAVA

1. Leaves relatively large, 5-8 cm long, thin, mesomorphic; petioles highly glandular with sessile to very short

stipitate glands; twigs not flexuous at nodes; stamens 10, anthers purple; fruit narrow-pyriform (unripe) to

pyriform (ripe), yellowish to yellow- Ta with somewhat udis calyx C. flava

. Leaves smaller, 1-5 cm long, slightly to much Jr

stipitate glands (except in C. frugiferens); stamens usually 20, Aner white (cream) or pink to purple; twigs

slightly to very zigzag at nodes (except in C. frugiferens); fruit subspherical to slightly tapered at base, yellow

to red in color, lacking elevated calyx.

2. Leaves + cuneate to + parallel-sided; veins exiting near the end of the leaf, often only 1-3 per side; sides

lacking lobes for most of their length ser. Lacrimatae, p.p.

2. Leaves + rhombic, ovate or elliptic; veins exiting between the half-way point and the end of the leaf,

frequently 4-6 per side; sides clearly lobed except in a few species with very short leaves («1cm long).

3. Ultimate branches conspicuously lacrimate (except in two dwarf species «1.5 m tall); leaves and inflo-

rescence branches + tomentose young; anthers ivory to cream; leaves lobeless or with 1-2 blunt lobes

per side (except C. dispar, with sharp lobes) ser. Lacrimatae, p.p.

3. Ultimate branches not conspicuously lacrimate; leaves variably hairy to glabrous, never tomentose;

inflorescence branches glabrous to densely pubescent, never tomentose ; anthers usually pink to

purple; leaves with 3-4 distinct and usually sharp lobes per side ser. Apricae

ariably glar idular but quite witl out

—

Series Apricae J.B. Phipps, ser. nov. Wee Species: Crataegus aprica Beadle.

Frutices vel arbores parvae; truncorum interdum ater aut atrocinereus, + rimosus, sed raro commemoratus; ramuli vulgo tortuosi

sed rectiin C. frugiferenti, saepe subrufo-brunnei post unum annum; spinae vulgo 1.5-4.5 cm longae, + rectae vel leviter recurvatae. Folia

decidua, marginibus et petiolis glandulosis; laminae 1.5—5 cm longae, late-ovatae vel rhombo-ellipticae in forma generali; lobi nulli vel 1-3

per latus, vadosi vel (interdum tantum apiculi) obscure vel raro moderate profundi (et acuti in C. ignava); venatio craspedodroma, venis

3-5(6-7 in C. frugiferenti) per latus; tenues vel + chartaceae. Inflorescentiae (1-)2—6(-7) floratae; rami glabri vel pubescens (interdum

dense), f ras, caducas, lineares | glandulo-marginatas bracteolas. Flores 13-25 mm diam.; hypanthium glabrum

vel pubescente (interdum dense); lobi calycis angusto-triangulares, marginibus glandulo-serratis; petala + circularia, alba; stamina ca.

20(10), antheris roseis vel purpureis, interdum albis; styli 3-5. Fructus 8-15 mm diam., subglobosi vel globosi, aurantiaco-rubri vel

rubri, glabri vel cum pilis raris; lobi calycis reflexi; pyrenae 3—5, dorsaliter sulcatae, lateribus planis.

Shrubs or small trees; bark on trunks seldom recorded, when so, black or dark gray, + rimose; most twigs

slightly zigzag, except straight in C. frugiferens, often reddish brown after 1 yr.; thorns mainly 1.5-4.5 cm

long, + straight to slightly recurved. Leaves deciduous, margins and petioles glandular; blades 1.5—5 cm

long, broad-ovate to rhomb-elliptic in general shape; lobes none or 13 per side, if so, shallow (sometimes

mere apiculi) or obscurely to more rarely moderately deep (and quite sharply acute in C. ignava); venation

craspedodromous, 3—5(-6-7 in C. frugiferens) lateral veins per side; thin to somewhat coriaceous. Inflores-

cences (1-2)2-6(-7) flowered; branches glabrous or pubescent, sometimes densely, bearing small, caducous,

linear, membranous, gland-margined bracteoles. Flowers 13-25 mm diam.; hypanthium glabrous or pu-

bescent, sometimes densely so; calyx lobes narrow-triangular, margins glandular-serrate; petals + circular,

white; stamens usually 20(-10), anthers usually pink to purple, occasionally white; styles 3—5. Fruit 8-15

mm diam., subglobose to spherical, orange-red to red, smooth or with scattered hairs; calyx lobes usually

reflexed; nutlets 3—5, dorsally grooved, sides plane.

Series Apricae is essentially southern Appalachian/adjacent Piedmont in distribution with some exten-

sion into the coastal plain in northern Florida and South Carolina where its species constitute a character-

istic element of the Crataegus flora in sunny places. Here, 12 species are named with certainty, only a few

of which are reasonably common, for instance, C. mira. Several more are known only from their types and

receive full descriptions. An interesting Crataegus flora from the Augusta sandhills of a hundred years ago

and now unknown is treated as fully as material permits and special attention is drawn to it. Perhaps it is in

the C. flava alliance. A full and updated treatment of C. flava, except for illustrations, for which the reader

is referred to Phipps (19883), is also provided in this paper because that is where it would customarily be

sought and a parallel treatment of ser. Intricatae is not anticipated.

174 Journal of the Botanical R h Institute of Texas 1(1)

Crataegus series Apricae represents a remodeling of series Flavae (Loud.) Rehder in the sense that was

used in Phipps et al. (2003) and Phipps et al. (1990). This is due to the transfer of its type species to ser.

Intricatae. The species included here belong to Beadle's groups Euflavae (minus C. flava), Ignavae, Sororiae,

Segnes and Visendae.

Species limits in the new series were found to be not always very clear cut, in part because of a relative

paucity of material, and the present treatment takes a narrow view of specific limits generally following

Beadle (1902) and Beadle (1903) because of the potential arbitrariness of lumping in such cases. Beadle knew

the group better than anyone else, having described most of the taxa, and remains the only guide among

the earlier generations of botanists. None of the species of the series as finally demarcated here appears to

be really common and several are very rare or local, limiting insights from modern fieldwork. Some of the

taxa recognized could well be local hybrids or apomictic races and the only ploidy level recorded is 3x for

a cultivated specimen that might be C. aprica (Talent & Dickinson 2005).

KEY TO SERIES APRICAE AND CRATAEGUS FLAVA

1. Twigs + zigzag, except in C. frugiferens; leaves 2-5 cm long; anthers usually pink to purple; fruit orange-red

to red, subglobose.

2. Stamens 20

3. Inflorescence branches tomentose-canescent or scabrous-pubescent.

4. Inflorescence branches tomentose-canescent; leaf-blades + isodiametric; petioles with sessile

glands 6. C. sororia

4. Inflorescence branches scabrous-pubescent; leaf-blades ovate to rhombovate or obovate; petioles

usually with at least some glands stipitate 11. C. frugiferens

3. rana branches at most thin-pilose or pubesce

. Leaf-blades suborbiculate to ellipt-rhombic or ME lobes of leaf-blades small, mere apiculi, or

obscure or lacking.

6. Blades broad-elliptic-rhombic to suborbicular; lobes clearly present, though small.

7. Atleast some leaves usually tending to suborbiculate in shape; lobing small, neat and regular,

+ acute at anthesis, becoming more obscure later in the season; flowers 20-25 mm diam. ___ 7. C. mira

7. No leaves tending to suborbiculate in shape, larger ovate, smaller elliptic to rhombelliptic;

lobing somewhat irregular; flowers 15-20 mm diam.

8. Bis. un 8. C. leonensis

8. Anthers ivory 3. C. sp. cf. C. annosa

6. e Eu wees distally, if widest in the centre then not even approximately m with

1-several small and irregular lobes per side, or cuspidate and at the most terminally denticulate;

flowers 14-25 mm diam. (‘visenda group)

9. Blades smaller, 1.5-3 cm long, broadly or narrowly obovate to rhombovate in general shape,

seldom less than 1.5 x as long as broad; flowers 14-20 mm diam.

10. Leavesrhombic-ellipticto rhomb-obovate in general shape, the tip cuspidate; many irregularly

short-lobed 2. C. visenda

10. Leaves narrow-obovate to narrow-elliptic in general shape, nearly without discernible

lobes 3. C. galbana

9. Blades larger, 2-3.5 cm long, broadly elliptic or rnombelliptic to obovate in general shape; m

15-18 mm diam. 4. C. segnis

5. Leaf-blades + rhombic; lobes well-defined, sharp to somewhat blunt.

11. Blades 1.5-2.5 cm long; inflorescence branches pubescent 1. C. egregia

11. Blades 2.5-4.0 cm long; inflorescence branches glabrous 9. C. ignava

2. Stamens 10 (occasionally1 2-15).

12. Leaf-blades 3-5 cm long, lobes and teeth sharp; flowers 16-20 mm diam., anthers pink or purple.

13. Inflorescence branches glabrous; stamens 10 10. C. allegheniensis

13. Inflorescence branches pilose-pubescent; stamens 12-15 12. C. extraria and C. sp. cf. extraria

12. Leaf-blades 2.5-5 cm long, lobes, if any, apiculi; teeth blunt or sharp; flowers 13-16 mm diam.; anthers

pink or cream.

14. Leaves widest near the mid-point, teeth not sharp, + coriaceous at maturity; petiolar glands all

sessile; anthers cream; inflorescence branches quite densely pilose 5. C. aprica

Phipps and Dvorsky, Crataegus ser. Apricae and C. flava 175

14. Leaves ovate in general shape, sharply toothed, relatively thin at maturity, 2.5-5 cm long; peti-

oles usually with some glands stipitate; anthers pink or cream; inflorescence branches apressed

scabrous-pubescent 11. C. frugiferens

1. Twigs not zigzag; leaves 5-8 cm long; anthers pink to purple; fruit yellowish, narrow [C. flava

(ser. Intricatae), see end of treatment]

1. Crataegus egregia Beadle, Biltmore Bot. Stud. 1:82. 1902. (Fig. 1). Tye: U.S.A. Fiona. Liberty Co.: Bristol, 24 Aug

1901, T.G. Harbison 4942 (LECTOTYPE SELECTED HERE: A).

Small tree 4—6 m tall; thorns none or several, 3-4 cm long, dark, slender, straight; twigs + slender, barely

flexuous, bark brown at 1 yr., older grayish; trunk bark dark, very rough. Leaves deciduous; petioles ca. 1

cm long, 30-50% length of blade, slender, pubescent, very glandular; blades 1.5-2.5 cm long at anthesis,

23 cm long at maturity; rhombic in general shape, generally with 1 main lobe per side; margins shallowly

crenate, teeth gland-tipped; apex acute to subacute, base cuneate; venation craspedodromous, lateral veins

ca. 3/side; nearly glabrous on both sides with scattered hairs adaxially young; thin. Inflorescences ca. 3-

flowered; branches pubescent, bearing caducous, narrow-oblong, membranous, greenish, gland-bordered

bracteoles. Flowers ca.15 mm diam.; hypanthium thin-pilose; calyx lobes narrow-triangular from a wide

base, abaxially glabrous, gland-margined on barely discernible teeth; stamens 20, anthers bright purple,

styles 3-4. Fruit 9-12 mm diam., slightly pyriform, red at maturity, glabrous; sepals reflexed; nutlets 3-4,

dorsally grooved, sides plane.

Habitat and Distribution —Most specimens come from Bristol, Florida. It is also recorded from two other

locations in Florida as well as several from South Carolina and one each from Alabama and Georgia (Fig.

2). Itisa rare species that I (JBP) have not encountered in the field.

Comment.— Crataegus egregia is a distinct looking plant with a unique leaf shape which is understood

primarily from its type. Herbarium material is in some ways like a large C. egens (ser. Lacrimatae) but with

larger, more rhombic leaf-blades and, where present, longer thorns. South Carolina specimens are thorny,

typical material is thornless. Unmapped material represented by about six specimens with similar leaves

but more or less tomentose pedicels and much smaller fruit may be the same.

2. Crataegus visenda Beadle, Biltmore Bot. Stud. 1:79. 1902. (Fig. 3). Tyee: U.S.A. Fiona. Liberty Co.: Bristol, 29

Mar 1901, T.G. Harbison 4031 (Lectotype selected here: A).

Crataegus arrogans Beadle, Biltmore Bot. Stud. 1:81. 1902. Type: U.S.A. ALABAMA. Russell Co.: Phenix City, 26 Aug 1901, C.D. Beadle 4869

(LECTOTYPE selected here: US).

Crataegus sodalis Beadle, Biltmore Bot. Stud. 1:80. 1902. Tree: U.S.A. Grorcia. Burke Co.: Girard, 26 Aug 1901, C.D. Beadle 4868 (LEc-

TOTYPE selected here: NY

Crataegus tristis Beadle, Biltmore Bot. Stud. 1:84. 1902. Tyre: U.S.A. Georcia. Floyd Co.: Rome, 25 Apr 1901, C.D. Beadle 4194 (LECTOTYPE

selected here: A)

Large shrub or small tree to 10 m tall; bark on trunk rough, dark gray or brownish; 1 year old twigs dark

brown; thorns few to numerous, 1.5-3 cm long, + straight, very dark at 1 year. Leaves deciduous; petioles

slender, 25-40% length of blade, gland-margined, pilose, winged above; blades 1.5-3.0 cm long, rhomb-

elliptic to rhomb-obovate in general shape, not or shallowly 1-2-lobed per side, tip + cuspidate; margins

obscurely crenate at maturity (sharper younger); venation craspedodromous with 3—4(—5) pairs of lateral

veins; when young conspicuously pilose on the veins adaxially, thinly so abaxially, otherwise nearly glabrous.

Inflorescences 2—4-flowered; branches nearly glabrous to pilose, bearing caducous, linear, membranous,

gland-margined bracteoles. Flowers 14-18 mm diam.; hypanthium externally glabrous to pilose; calyx lobes

triangular, margins irregularly serrate and very glandular; petals + circular, white; stamens 20, anthers pale

to bright purple; styles (22)3—5. Fruit 10-12 mm diam., subglobose to pyriform, glabrous, orange or orange

flushed red to red, calyx lobes recurved; nutlets grooved dorsally, laterally smooth.

Habitat and Distribution.—Nearly all material seen is form northern Florida but there are a few records

from southeastern Alabama and southwestern Georgia. Crataegus visenda is found in dry woods, on gravelly

176

Journal of the Botanical Research Institute of Texas 1(1)

Fic. 1. Li

AODA L

TILT Y

), flowering and fruiting. Scale bars =1 cm. S. Laurie-Bourque del.

Phipps and Dvorsky, Crataegus ser. Apricae and C. flava 177

Number of Records

0 Crataegus egregia

1-3

Fic. 2. C it I l dictrihi it, £ Fd

£. SUUTTLY

ridges and in sandy soil. This species is mapped collectively with Cc. galbana and segnis as the informal

'visenda group’ (Fig. 6).

Comment.—Beadle recognized 10 species in his Visendae from which I have removed C. annosa and C.

egens. Crataegus furtiva, rather similar to C. visenda but with tomentose pedicels and hypanthia, is in ser.

Lacrimatae.

3. Crataegus galbana Beadle, Biltmore Bot. Stud. 1:74. 1902. (Fig. 4). Wee: U.S.A. Fiona. Gadsden Co.: southwest

of River Junction, 3 Apr 1900, C.D. Beadle 2083 (Lectotyre selected here: NY).

Large shrubs or small trees; twigs somewhat zigzag, at 1 year old dark brown, but partly covered with

abraded cutin, later pale to mid gray; thorns none to numerous, 1.5-2.5 cm long, straight, purple-brown at

l year, gray later. Leaves deciduous; petioles 30-45% length of blade, slender, gland-margined, pubescent;

blades 1.5-3 cm long, narrow-obovate to narrow-elliptic in general shape, apex acute to sometimes slightly

cuspidate, base + rapidly narrowed; margins + devoid of lobes except sometime a few distal half; margins

crenate-serrate, the teeth gland-tipped; venation craspedodromous with 3—4 pairs of lateral veins; when

young thin hairy adaxially especially on the mid-vein, nearly glabrous abaxially young, except along the

mid-vein and in the main axils, + glabrescent. Inflorescences 1—5-flowered; branches finely pubescent,

bearing several caducous, narrow-oblong, membranous, gland-margined bracteoles. Flowers 14-20 mm

178 Journal of the Botanical Research Institute of Texas 1(1)

y ss

3 NS

E——

Fic. 3. Line drawing of Crataegus visenda from Lance 2114 (UWO), flowering and Godfrey 79895 (UWO), fruiting. Scale bars =1 cm. S. Laurie-Bourque

del.

Phipps and Dvorsky, Crataegus ser. Apricae and C. flava

179

Te J H £f. 4 IL £. Y A me D 4 TICIACA

Fic. 4 )

Li A

Scale bars =1 cm. S. Laurie-Bourque del.

202029 /(C M £e

V 11

349 (USCH 1), flowering

180 Journal of the Botanical R h Institute of Texas 1(1)

diam.; hypanthium externally thinly pilose; calyx lobes triangular, margins glandular and strongly ser-

rate, abaxially glabrous; petals + circular, white; stamens 20, anthers pale purple; styles 3-5. Fruit 10-15

mm diam., subglobose, glabrous, orange flushed red to red; calyx lobes recurved; nutlets ca. 3-5, grooved

dorsally, laterally smooth.

Habitat and Distribution.—Locally common in the Florida Panhandle, Crataegus galbana ranges to south-

central Alabama throughout Georgia to South Carolina with two records from North Carolina. It occurs in

open woodland and scrubby places. This species is mapped collectively with Cc. visenda and segnis as the

informal ‘visenda group’ (Fig. 6).

Comment.—Crataegus galbana is the least similar to C. visenda of this group of species and might per-

haps be confused with C. aprica. Nevertheless, the smaller, differently shaped leaves and 20 pale purple

anthers will readily distinguish it from C. aprica. A good many specimens of this species were annotated ‘C.

consanguined'.

4. Crataegus segnis Beadle, Biltmore Bot. Stud. 1:32. 1901. (Fig. 5). Tyee: U.S.A. Atasama. Butler Co.: Greenville, 24

Aug 1901, C.D. Beadle 2155? (Lectotype selected here: A).

Crataegus consanguinea Beadle, Biltmore Bot. Stud. 1:34. 1901. Tyre: U.S.A. FLoripa. Leon Co.: W of Tallahassee, 28 Mar 1900, C.D.

Beadle 2044 (Lectotyre selected here: US).

Large shrubs or small trees; twigs somewhat zigzag, at 1 year old dark brown, but partly covered with

abraded cutin, later pale to mid gray; thorns none to numerous, 1.5-2.5 cm long, straight, purple-brown

at 1 year, gray later. Leaves deciduous; petioles slender, 30-45% length of blade, slender, gland-margined,

pubescent; blades 1.5-2.5 cm long, broadly elliptic or rhombelliptic to obovate in general shape, apex acute

and often somewhat cuspidate, base + rapidly narrowed; usually not but sometimes very obscurely lobed,

margins crenate-serrate, the teeth gland-tipped; venation craspedodromous with 3—5 pairs of lateral veins

except in the smaller leaves; when young thin hairy adaxially especially on the mid-vein, nearly glabrous

abaxially young, except along the mid-vein and in the main axils, + glabrescent. Inflorescences 1—5-flowered;

branches quite long pilose, bearing several caducous, narrow-oblong, membranous, gland-margined brac-

teoles. Flowers ca.15-18 mm diam.; hypanthium externally thinly pilose; calyx lobes triangular, margins

irregularly strongly glandular and serrate, abaxially glabrous; petals + circular, white; stamens 20, anthers

pale purple; styles 3—5; Fruit 10-15 mm diam., subglobose, glabrous, orange flushed red to red; calyx lobes

recurved; nutlets ca. 3—5, grooved dorsally, laterally smooth.

Habitat and Distribution.— Crataegus segnis occurs around Greenville, Alabama and in northern Florida.

This species is mapped collectively with Cc. visenda and galbana as the informal ‘visenda group’ (Fig. 6).

Comment.—This species has generally the largest and broadest leaves in the visenda group. The fruit is

subglobose and red.

5. Crataegus aprica Beadle, Bot. Gaz. 30:335. 1900. (Fig. 7). Tyee: U.S.A. Norm Cagouwa. Buncombe Co.: Biltmore, 11

May 1899, Biltmore Herb. C14 (Lectotype selected here: NY).

Shrubs, generally 2-3 m tall; branchlets somewhat flexuous; extending twigs olive-green with somewhat

sparse pubescence; l-year old twigs reddish-brown, pubescent, older dark gray-brown glabrous; 2-year old

thorns 3-4 cm long, slender, straight or recurved; dark gray-brown. Leaves deciduous; petioles 3-8 mm

long, 30-50% length of blade, pubescent, glandular; leaf blades 1.5-4 cm long, the blades rhomb-elliptic

or broad-elliptic in general outline, widest in the middle, apically blunt, sharply constricted at the base and

tapered into the winged upper petiole; extremely shallowly lobed to unlobed, lobes more prominent (mere

apiculi) young; margins crenate or obtusely serrate, the teeth gland-tipped; venation craspedodromous, with

3-4 lateral veins per side; surfaces pilose above when young but glabrescent later, glabrous below except

on the midvein; + coriaceous at maturity. Inflorescences 3-6 flowered; branches + densely pilose, bearing

caducous, linear, membranous, gland-margined bracteoles; anthesis April. Flowers 13-15 mm diam.; hypan-

thium pilose, at least near the base; calyx lobes ca. 4 mm long, narrow triangular, gland-toothed, sparsely

pubescent abaxially, with a prominent mid-vein in some; petals + circular, white; stamens 10, anthers ivory

Phipps and Dvorsky, Crataegus ser. Apricae and C. flava 181

Fic. 5. Li

del.

photo of Beadle 2044 (US), flowering and Beadle 2155? (A), fruiting. Scale bars =1 cm. S. Laurie-Bourque

182 Journal of the Botanical R h Institute of Texas 1(1)

or cream; styles 3-5. Fruit 9-15 mm diam., + orbicular, with a few hairs, red or reddish-orange; calyx lobes

patento-reflexed; nutlets 3—5, dorsally sulcate, laterally smooth.

Common Name.—Sunny Hawthorn.

Habitat and Distribution.— Crataegus aprica is found mainly around the southern end of the Appalachians

from northern Florida to Virginia with a single record each for Alabama and Tennessee (Fig. 8). It occurs at

50-3000 ft in open brushy areas where it may be quite common.

Comment.—Vegetatively, C. apricais not unlike members of the visenda group but it has 10 stamens and

ivory anthers. The record from Barbour Co., Alabama has a thinner inflorescence tomentum and generally

larger (3.0-4.5 cm long), longer petiolate (1.5-2.0 cm long), thinner leaves.

6. Crataegus sororia Beadle, Bot. Gaz. 30:336. 1900. (Fig. 9). Wee: U.S.A. Grorcia. Floyd Co.: hills above Silver Creek,

Rome, 18 Sep 1897, C.D. Beadle 1257 (LecToTYPE selected here: A).

Shrubs, generally 23 m tall; trunk bark rimose; branchlets somewhat flexuous; extending twigs olive-green

with somewhat sparse pubescence; l-year old twigs reddish-brown, pubescent, older dark gray-brown

glabrous; 2-year old thorns 3-4 cm long, slender, straight or recurved; dark gray-brown. Leaves decidu-

ous; petioles 3-8 mm long, 30-50% length of blade, pubescent, glandular; leaf blades 1.5-4 cm long, the

blades broad-elliptic to circular in general outline, widest in the middle, apically blunt, sharply constricted

at the base and tapered into the winged upper petiole; rather sharply lobed, lobes more prominent young;

margins finely serrate, the teeth gland-tipped; venation craspedodromous, with 3-4 lateral veins per side;

surfaces pilose above when young but glabrescent later, glabrous below except on the midvein; + coriaceous

at maturity. Inflorescences 3-6 flowered; branches tomentose, bearing caducous, linear, membranous,

gland-margined bracteoles; anthesis April. Flowers ca. 15 mm diam.; hypanthium densely pilose; calyx

lobes ca. 4 mm long, narrow triangular, gland-toothed, sparsely pubescent abaxially, with a prominent

mid-vein in some; petals + circular, white; stamens 20, anthers ivory, pink-purple or red; styles 4—5. Fruit

12-18 mm diam., + orbicular, with a few hairs, reddish-orange; calyx lobes patento-reflexed; nutlets 4—5,

dorsally sulcate, laterally smooth.

Habitat and Distribution.—Crategus sororia occurs around Rome, Georgia where it is still common, in

adjacent areas of Alabama and in Aiken Co., South Carolina (Fig. 10). It is found in open scrubby areas.

Comment.—This species, most similar to C. aprica, differs in leaf shape (proportionately broader), sta-

men number and anther color. Beadle (1900) says that C. sororia is found south to Florida but I have not

seen specimens of it from that state.

7. Crataegus mira Beadle, Biltmore Bot. Stud. 1:78. 1902. (Fig. 11). Tv: U.S.A. Grorcia. Cobb Co.: 9 May 1901, C.D.

Beadle 4287 (uovorvrs selected here: US).

Shrubs, generally 2-3 m tall; branchlets somewhat flexuous; extending twigs olive-green with somewhat

sparse pubescence; l-year old twigs reddish-brown, pubescent, older dark gray-brown glabrous; 2-year old

thorns 3-4 cm long, slender, straight or recurved; dark gray-brown. Leaves deciduous; petioles 3-8 mm

long, 30-50% length of blade, pubescent, glandular; leaf blades 2-3.5 cm long, the blades broad rhomb-el-

liptic to + circular in general outline, widest in the middle, apically acute, sharply constricted at the base and

tapered into the winged upper petiole; shallowly but sharply lobed, lobes more prominent young; margins

crenatoserrate to serrate, the teeth particularly prominent around anthesis, gland-tipped; venation craspe-

dodromous, with 3-4 lateral veins per side; surfaces pilose above when young but glabrescent later, glabrous

below except on the midvein; + coriaceous at maturity. Inflorescences 3-6 flowered; branches glabrous to

pilose, bearing caducous, linear, membranous, gland-margined bracteoles; anthesis April. Flowers 20-25

mm diam.; hypanthium thin-pilose to more or less glabrous; calyx lobes ca. 5 mm long, narrow triangular,

gland-toothed, sparsely pubescent abaxially, with a prominent midvein in some; petals + circular, white;

stamens 10-20, anthers cream or purple; styles 4-5. Fruit 9-15 mm diam., + orbicular, with a few hairs,

red or reddish-orange; calyx lobes patento- reflexed; nutlets 4—5, dorsally sulcate, laterally smooth.

Habitat and Distribution.—The main range of C. mira is the Florida panhandle to central Georgia but

Phipps and Dvorsky, Crataegus ser. Apricae and C. flava 183

Number of Records

0 Crataegus visenda group

1-3

[MI 4-8

= 9-14

Fic 6 € FEET A O et. bi

mu UNY

it extends to the Carolinas and Alabama (Fig. 12). It is found in open brushy areas where it may be quite

common.

Comment.—This striking species is most similar to Crataegus sororia and C. aprica but differs in its

broader and often larger leaves, thinner inflorescence indumentum and larger flower size.

8. Crataegus leonensis E.J. Palmer, J. Arnold Arbor. 13:422. 1932. Tee: U.S.A. Floripa. Leon Co.: near Tallahassee, 3

Apr 1923, T.G. Harbison 6072 (HOLOTYPE, A).

Tree 10-12 m tall, with wide-spreading intricate branches; trunk bark thick, ridged, dark gray to nearly

black: twigs somewhat zigzag, very dark at 1 yr.; thorns sparse, to 3.5 cm long, straight, very dark at 1 yr.

Leaves deciduous; petioles slender, 25-30% length of blade, extremely glandular, pubescent in the sulcus;

blade 2.5-3.5 cm long, oblong to rhombovate or narrowly obovate in general shape, tip acute; base cuneate;

sides obscurely or very shortly 1-2 lobed; margins finely to obscurely crenate-serrate, the teeth gland-tipped;

venation craspedodromous, 3—4 veins per side; nearly glabrous but with some pilosity along the main

veins adaxially and abaxially (Palmer implies glabrous abaxially); rather thin. Inflorescences 3-7 flowered;

184 Journal of the Botanical Research Institute of Texas 1(1)

Fic. 7. Line drawing of Crat ica from Phipps 5681 (UWO, cult. at K), flowering and Ulf-Hansen 119 (UWO), fruiting. Scale bars =1 cm. S. Laurie-

Bourque del.

Phipps and Dvorsky, Crataegus ser. Apricae and C. flava 185

Lo V4 H V

P la i a

Crataegus aprica

9-14

SS 15 - 24

_ -2

5-60

branches subglabrous to moderately pilose, bearing narrow-oblong, caducous, membranous, gland-bordered

bracteoles. Flowers 16-20 mm diam., hypanthium glabrous externally; calyx lobes triangular, glandular-ser-

rate, abaxially glabrous; stamens 20, anthers pink; styles 2-5. Fruit 9-12 mm diam., subglobose, glabrous,

orange-red to russet, or often green-mottled; calyx lobes somewhat elevated, spreading, prominent; nutlets

(2-)3-4(-5), dorsally grooved, laterally plane.

Habitat and Distribution —This is a local species mainly known from Leon Co., Florida. A few specimens

are also known from southern Georgia (Fig. 10). It is recorded from sandy upland woods.

Comment.—Crataegus leonensis is similar to C. mira, but it has smaller flowers than that species and

differently shaped leaves (see key). It is possibly just an extreme from of C. mira but more field study of

these two entities is needed to determine this. One of the flowering specimens on Canby & Sargent 27 (DOV)

from Chattahoochee, Georgia has a much more pubescent inflorescence but is otherwise indistinguishable.

Murrill's C. subflavida is perhaps the same but has larger yellow fruit.

186

Journal of the Botanical Research Institute of Texas 1(1)

"RS

QUE

Ba.

del iz :

F 0 LIH A P £ fvat Lyr d m* uy NAINA IRANA £1 : 4 Dh:

IG Hr Ul

Er ND H F

UN” HUTCH Y

Phipps and Dvorsky, Crataegus ser. Apricae and C. flava 187

Au ^L

eI RE

? u i

Number of Records

0 Crataegus leonensis and C. sororia

p 1-3

[II 4-8

F 10.C doc l Aictyths £f ff £L Y Iri Ow, L \ Tha + : IL L AI

Ic. IU. County g ( ) ( ) [ | y the curved tine,

9. Crataegus ignava Beadle, Biltmore Bot. Stud. 1:31. 1901. (Fig. 13). Tee: U.S.A. ALapama. De Kalb Co.: Lookout

Mountain, Valley Head, Oct 1900, C.D. Beadle 2289? (LecrorvrE selected here: US).

Shrubs, 2-45) m tall; + thorny, thorns 2-3 cm (3-5) long at 2-yr. old, straight, + fine, black; extending

shoots glabrous; 1-yr old tan to chestnut, shiny; older + dark gray, often reddish tinged. Leaves deciduous;

petioles ca. 1 cm long, glabrous, black-glandular; blades ca. 2.5-3.5 cm long in our material, narrow-ovate

to ovate-rhombic or broad-ovate in general shape; acute at the tip and wide-cuneate or somewhat rounded

at the base; with 2-3 sharp lobes on either side; margins obscurely crenate-serrate, the teeth gland-dotted;

venation craspedodromous, lateral veins 4—5 per side; thinly pilose above when young, but soon glabrescent,

+ coriaceous at maturity. Inflorescences 2—5 flowered; branches glabrous, bearing aducous, linear,

herbaceous to membranous, gland-margined bracteoles. Flowers ca. 15 mm wide; hypanthium externally

glabrous; calyx lobes 6 mm long, narrow-triangular, abaxially glabrous, the margins glandular-serrate;

petals + circular, white; stamens 20, anthers pale pink (pale purple); styles (3-5). Fruit 10-17 mm diam.,

subglobose, orange-red; calyx lobes spreading-recurved; nutlets 3-5, dorsally furrowed, laterally plane.

188 Journal of the Botanical Research Institute of Texas 1(1)

Fic. 11. Line drawing of Crat ira from L 2121 (UWO), flowering and Phipps 6684 (UWO), fruiting. Scale bars =1 cm. S. Laurie-Bourque del.

Phipps and Dvorsky, Crataegus ser. Apricae and C. flava 189

Lov

X T

d S

E da, AG

= "f i

1 -— o. Y

$ t0 AX ae

ec 1

[^F "vac M t

k 1 eS $ x bru vene

la SEV AC ES: i

$ AE Sia

BY A UA,

PORN

Number of Records

0 Crataegus mira

1-3

MID 4-8

= 9-1

Fic. 12. County | | distril ti f C y

Habitat and Distribution.—Crataegus ignava is a locally common plant of east-central Alabama and north-

central and central Georgia (Fig. 14). The senior author has collected it from rocky hills bearing stunted

trees of this species near Anniston, Alabama and red soils in central Georgia.

Comment.—If Murrill's C. subflavida is the same, this species also occurs near Gainesville, Florida.

Crataegus ignava can be a very handsome plant with its bright orange-red fruit, which is sometimes very

large, and striking foliage.

10. Crataegus allegheniensis Beadle, Bot. Gaz. 30:337. 1900. (Fig. 15). Tvrz: U.S.A. ALapama. De Kalb Co.: Lookout

Mountain, Valley Head, 7 May 1900, C.D. Beadle 2290 (Lectotype selected here: US).

Shrubs, 2-4(—5) m tall; thorns at 2 yrs 1.5-4 cm long, straight or slightly recurved, deep chestnut brown to

blackish; extending twigs reddish, glabrous, at 1 yr old reddish-gray, older gray. Leaves deciduous; petioles

0.75-1.5 cm long, very glandular, glabrous; blades (2-)3-5 cm long elliptic-ovate in general shape, acute

above, broadly cuneate below with about 3 sharp lobes per side, LII ca. 15%; margins finely toothed, the

teeth with small glands; venation craspedodromous, with ca. 5 pairs of veins; upper surface pilose when

190 Journal of the Botanical Research Institute of Texas 1(1)

Fic. 13. Line drawing of Crataegus ignava from Smith & Spaulding 4 (UWO), flowering and Phipps 7783 (UWO), fruiting. Scale bars =1 cm. S. Laurie-

Bourque del.

Phipps and Dvorsky, Crataegus ser. Apricae and C. flava 191

PA D / La

ae:

is.

apo?

0 Crataegus ignava

= 9-14

NNN 15 - 24

Fic. 14. County |

very young, soon glabrescent, below glabrous. Inflorescences 2-4 flowered; branches glabrous, bearing early

caducous, linear, membranous, gland-margined bracteoles. Flowers 20 mm diam.; hypanthium externally

glabrous; calyx lobes 5 mm long, narrowly triangular, abaxially glabrous, margins glandular-serrate; petals

+ circular, white; stamens 10, anthers pink; styles ca. 4. Fruit 8-12 mm thick, globose-pyriform, glabrous,

red; nutlets 2—5 (Beadle).

Habitat and Distribution.—Crataegus allegheniensis is known from a few locations in northeastern Ala-

bama, Tennessee and Georgia and is apparently scarce (Fig. 16). It occurs in various brushy places and in

Alabama on rocky hills.

Comment.—This species is superficially similar to C. frugiferens and may perhaps turn out to be only a

glabrous form of that species.

11. Crataegus frugiferens Beadle, Biltmore Bot. Stud. 1:30. 1901. (Fig. 17). Wee: U.S.A. Atasama. Cullman Co.:

Cullman, 24 Aug 1901, T.G. Harbison 2116 (Lectotyre selected here: US).

Journal of the Botanical Research Institute of Texas 1(1)

192

ing. Scale bars —1 cm. S. Laurie-

1546 (UWO), fruiti

ic fi Rarhor of nl

IT UI DUIUCI EL UE,

Fic. 15. Li

Bourque del.

Phipps and Dvorsky, Crataegus ser. Apricae and C. flava 193

ie CM S

" 4 č & TE 1 gU 4 A ai

2 Ios 4 fei

Number of Records

0 Crataegus allegheniensis

1-3

Fic. 16. County level distributi p of Crataegus allegt

Shrubs 3-6 m tall; thorns at 2 yr. 2-5 cm long, + fine, straight to slightly recurved shiny dark brown or black-

ish, older gray; extending shoots + appressed pubescent, reddish when young; at 1 yr. + shiny tan, older gray.

Leaves deciduous; petioles 25-40% length of blades, slender, thinly pubescent, more so in adaxial groove,

bearing a few, often stipitate, glands; blades 2.5-5 cm long, ovate to rhombovate, tip acute, base cuneate to

broadly cuneate, 0—3 lobed per side; lobes very shallow, sometimes little more than apiculi, max LII 0—596;

margins serrate with gland-tipped teeth basally, these glands sometimes stipitate; venation craspedodromous,

5—6(-7) veins per side; appressed scabrous-pubescent above young, later + glabrescent, below glabrous on

the surface, thinly scabrous on the veins; thin. Inflorescences 3-7 flowered; branches appressed scabrous-

pubescent; bracteoles very few, apparently early caducous, linear, membranous, gland-margined. Flowers

15-16 mm diam.; hypanthium externally appressed scabrous-pubescent at least below; calyx lobes 4-5 mm

long, narrow triangular, margins glandular-serrate; petals + circular, white; stamens 10 or 20, anthers cream

or pink; styles 3-4. Fruit not known to me but according to Beadle 9-12 mm thick, subglobose or slightly

pyriform, red at maturity; calyx lobes reflexed, margins glandular-serrate; pyrenes 3-5.

Habitat and Distribution.—The main range of Crataegus frugiferens is in north central Alabama and

adjacent Georgia. A single record is also known from Mississippi (Fig. 18). It is a somewhat scarce species

found both in thick woodland and rocky outcrops.

194 Journal of the Botanical Research Institute of Texas 1(1)

Fic. 17. Line drawing of Crataegus frugif from Phipps & Wells 5309 (UWO), flowering and Phipps & Spaulding 7771 (UWO), fruiting. Scale bars =1

cm. S. Laurie-Bourque del.

Phipps and Dvorsky, Crataegus ser. Apricae and C. flava 195

7 om ZV FK es ARE T]

Xx LA ^ ESN D

C : A N f mAs

ce Aa dp y: AA :

b LAS Y T TU e

XLX l w d | : | yn $ à

: —

pac

Fic. 18. County | | distril ti f fC fi if

Comment.—Crataegus frugiferens is one of the more distinct species of ser. Apricae, even to being super-

ficially similar to C. collina from which it differs in its glandular, often stipitately so, petioles and leaf bases as

well as leaf shape which is relatively broader with much more wide-cuneate base. It also has more strongly

glandular-serrate calyx lobes and usually, deeper red, softer fruit. The leaves at anthesis are no larger than

the flowers, as is common in C. collina. Its thin leaves make it less xeromorphic than other members of the

series. Specimens of this species have been annotated ‘C. rigens’.

12. Crataegus cf. extraria Beadle, Biltmore Bot. Stud. 1:73. 1902. Tee: U.S.A. Georaia. Cobb Co.: Marietta, 11 Sep

1901, C.D. Beadle 24852 (Lectotype selected here: US).

Shrub, 2.5 m tall; 1 yr. old twigs reddish-brown; older dark purple-brown. Leaves deciduous; petioles

1-1.5 cm long, glandular, thinly hairy; blades 3-4 cm long, rhomb-ovate in general shape; tip acuminate,

base cuneate; sides with 2-3 sharp to obscure lobes per side, max. LII 10-15%; margins fine-serrate, teeth

gland-tipped; venation craspedodromous with 4—5 main veins per side; at maturity leaves glabrous above,

green below, somewhat pubescent on the veins and thinly so on the surface. Inflorescences 3-5 flowered;

branches pilose-pubescent, bracteoles not recorded. Flowers 16-20 mm diam.; hypanthium externally

f4L,D o ID

196 Journal of t h Institute of Texas 1(1)

pilose-pubescent; calyx lobes narrow-triangular, margins glandular-serrate; stamens12-15(-20), anthers

dark purple, styles 2-3(-4). Fruit 9-12 mm diam., + spherical, glabrous, red but yellow-orange when not

fully ripe on pilose pedicels; calyx lobes spreading, elevated on a distinct collar; stamen remnants ca. 20 in

specimen studied; nutlets ca. 4.

Habitat and Distribution.— Crataegus extraria has always been very rare and occurred in scattered locali-

ties in North Carolina and Virginia. Somewhat similar forms (C. cf. extraria) were known from Alabama and

one such has been found recently, collected in Georgia on 5 Sep 1999 by R. Lance on the Tallulah R. Gorge,

Rabun Co. The Alabama specimen reported differs from the type description in possessing 20 stamens.

Comment.—Due to the rarity of C. extraria itself, and the very scattered distribution C. cf. extraria, plus

their collective variability, this entity, which may represent more than one taxon, is neither mapped nor

illustrated and perhaps in part constitutes sporadic hybrids between other species.

13. Crataegus cf. annosa Beadle

The following entity is known from a number of specimens. It was thought to be C. annosa during annota-

tions but it has only apricoid foliage.

Tree, 5-8 m tall; bark of trunk rough and dark; twigs somewhat flexuous at the nodes, at 1 year old

dark reddish-brown, older gray; thorns at 1 year 2-3.5 cm long, + straight, shiny dark reddish-brown, very

slightly recurved. Leaves deciduous; petioles1.5-2 cm long, slender, pubescent young, thinning older, glan-

dular; blades 3-5 cm long, broadly elliptic to rhombic in general shape; tip subacute to obtuse, base cuneate;

sides + shallowly but + sharplyl—3-lobed, margins shallowly crenate; venation craspedodromous, 4—5 lateral

veins per side; thinly hairy above at first, especially along the midvein, becoming glabrous, thinly hairy on

the abaxial surface, more densely so on the midvein and main lateral veins, glabrescent. Inflorescences 3-5-

flowered; branches pubescent, bearing caducous, linear, membranous, gland-margined bracteoles. Flowers

15-20 mm wide; hypanthium externally thinly pubescent; calyx lobes narrow triangular, margins deeply

incised with glandular teeth; petals + circular, white; stamens 20, anthers ivory; styles 3-5. Fruit 12-15

mm diam., subglobose, orange-red or red and orange when ripe, glabrous; calyx lobes erose or spreading;

nutlets 3-5, dorsally grooved, sides plane.

Habitat and Distribution —Occurs in Alabama, northern Florida, Georgia and North and South Carolina.

This is a rare plant of well-drained soils in woodland.

Comment.—This entity can be t intermediate between C. ignava and C. flava in leaf shape though

in size rather smaller than C. ignava but it isa much more hairy plant, has 20 stamens, reddish subglobose

fruit and much of the material has quite distinctly rhombic leaves. For similar reasons to C. extraria, this

taxon is neither mapped nor illustrated.

IMPERFECTLY KNOWN SPECIES

Here follows a number of forms, superficially quite distinctive but only known from a limited amount of

material. It seems important to draw attention to these forms but we consider it inappropriate to apply

definite names based on the limited knowledge. None of these entities is known today.

1. Crataegus cf. C. flava Aiton

“Small tree” in only known indication of habit (on the Harbison specimen); twigs + flexuous at the nodes,

very dark at two years; thorns 1.5-2.5 cm long, blackish, straight. Leaves deciduous; petioles 25-30% length

of blade, glandular; blades 2-4 cm long at full expansion, 1.75-3 cm at anthesis, predominantly obovate

to rhombobovate to rhombelliptic; lobes lacking on mature short-shoot leaves, mere notches or with LII

to 15%, if latter, obtuse to subacute at maturity but at anthesis lobes may be more prominent and even

acute; venation craspedodromous, 2-4 lateral veins per side; margins finely crenate-serrate almost to base;

glabrous except for hairs along veins adaxially young. Inflorescences 1—5-flowered; branches thin-pilose,

bearing plentiful, narrow, membranous, short-stipitately glandular bracteoles. Flowers 15-18 mm diam.;

hypanthium sparsely hairy; calyx lobes somewhat foliaceous, strongly glandular-serrate, nearly glabrous

(few short hairs near tip) adaxially; stamens 20, anthers “white”; styles 3-4. Fruit not known.

Phipps and Dvorsky, Crataegus ser. Apricae and C. flava 197

Habitat and Distribution.—This entity is known only from six flowering specimens from Augusta, Georgia

collected at different times from 1900-1919. The Augusta sand hills yielded many interesting hawthorns at

the beginning of the last century and similar areas in and around Augusta should be searched assiduously

for these and others by those in a position to do so.

Comment.—The leaves are extremely similar in form to those of true C. flava but the above plants cannot

automatically be placed there on account of a complete absence of fruiting material and the fact that they

have 20 stamens and white anthers. Also the twigs are generally more flexuous than in C. flava. Similar to

the above description but with more ovate leaves and purple anthers are Harbison 32 and Sargent s.n. (24

Apr 1900)—see cited specimens.

2. Crataegus calva Beadle, Biltmore Bot. Stud. 1:83. 1902. Wee: U.S.A. ALasama. Dale Co.: Ozark, 4 Sep 1901, T.G. Harbison

5004 (Lectotype selected here: US).

Shrubs 3-6 m tall; bark of trunk rough; crown irregular-spreading; twigs at 1 yr old slightly flexuous, color

not recorded; extending twigs not recorded; thorns 1.5-2 cm long, + straight, chestnut-brown in second

year. Leaves deciduous; petioles slender below, winged above, unwinged portion ca. 3096 length of blade,

minutely glandular; blades 1.5-2.5 cm long, (those of extension shoots to 4 cm) broad elliptic to obovate or

rhombobovate in general shape, sides tapered into the winged upper part of the petiole; tip + subacute (more

rarely obtuse or acute); often with 1 subacute lobe per side distally, LII 0—1596; margins crenate except near

base; venation craspedodromous, 2-3 lateral veins per side; when young with a few weak hairs on midrib

and principal veins below; somewhat coriaceous at maturity. Inflorescences 3—5 flowered; branches glabrous,

bearing deciduous, oblong-linear, membranous, gland-bordered bracteoles; flowering early to mid-April.

Flowers 16-18 mm wide; hypanthium externally glabrous; calyx lobes 3-4 mm long, narrow-triangular,

margins subentire to finely toothed, teeth glandular; petals + circular, white; stamens 20, anthers cream or

ivory; styles 3-5. Fruit typically 7-10 mm diam., globose, yellow or orange-red; calyx lobes erose or reflexed;

nutlets 3—5, dorsally grooved, sides plane.

Habitat and Distribution.—In woods and on ridges, Ozark, Alabama, not certainly known from elsewhere.

3. Crataegus arrogans Beadle, Biltmore Bot. Stud. 1:81. 1902. Tr: U.S.A. Arasama. Russell Co.: Phenix City, 26 Aug

1901, C.D. Beadle 4869 (LEctotypk selected here: US).

Tree, 4—5 m tall; bark ‘rough’, dark; thorns 1.5-2 cm long, straight to slightly recurved, + stout; twigs flexu-

ous. Leaves deciduous; petioles 2596 length of blade, glandular leaf blades 1.5—2 cm long, elliptic, pointed

at both ends; suggestions of pointed apiculi on some; veins 3-4/side; margins weakly and finely crenate

distally, teeth sometimes glandular; glabrous at maturity. Inflorescences few flowered; branches pubescent.

Flowers, diam. not recorded; hypanthium pubescent; sepals short, triangular, margins glandular-serrate;

stamens 20, anthers bright purple; styles 3-4. Fruit 9-12 mm diam., pyriform, red at maturity.

Habitat and Distribution.—Only known with certainty from the type locality.

Comment.—This is undoubtedly ser. Apricae, probably the visenda group.

4. Crataegus extraria Beadle Biltmore Bot. Stud. 1:73. 1902. Wee: U.S.A. Grorcia. Cobb Co.: Marietta, 11 Sep 1901, C.D.

Beadle 4285? (Lectotype selected here: US). Regrettably the lectotype is a very poor specimen, having lost most of its foliage.

Large shrub, 2-4 m tall; trunk bark rough or scaly, dark gray or brownish; twigs somewhat flexuous; thorns

2.5-4 cm long, slender, slightly recurved. Leaves deciduous; petioles 25-40% length of blade, winged above;

blades 2-3.5 cm long (few seen) broad rhombic to broad rhombovate in general shape; conspicuously though

shallowly acute lobed across distal end; margins with conspicuous sharp teeth; venation craspedodromous,

3—41-veined per side; glabrous; leaves on extension shoots much larger, often ovate, sharply 3-4-lobed per

side. Inflorescences 3—5-flowered; branches pilose-pubescent; bracteoles not recorded. Flower diam. not

recorded; hypanthium pilose-pubescent; sepals 4—5 mm long, margins glandular-serrate; stamens 12-15, an-

thers pale purple; styles 23. Fruit 9-12 mm diam., subglobose to ellipsoid, red at maturity, nutlets 2-3.

Habitat and Distribution.—Only known with certainty from the type locality.

Comment.—Provisionally this should be assigned to ser. Apricae though not enough is known about its

glandularity.

£+sha D o ID L

198 Journal of t titute of Texas 1(1)

DOUBTFUL SPECIES

SERIES APRICAE OR LACRIMATAE

Crataegus annosa Beadle, Biltmore Bot. Stud. 1:83. 1902. Tre: U.S.A. ALapama. Russell Co.: Phenix City, without date,

C.D. Beadle 4103 (Lectotype selected here: US; IsOLECTOTYPE: A). The lectotype is a fruiting specimen and isolectotype, C.D. Beadle

41032, with the same label data, is a flowering specimen.

Tree sometimes 8 m tall; bark of trunk rimose, dark; thorns on twigs 23 cm long, chestnut-brown or gray.

Leaves deciduous; petioles 5-20 mm long, slender, glandular, pubescent at least young; blades 2-4 cm long,

(rhomb-elliptic to obovate or obtrullate with small lobes across the apical part); venation craspedodromous,

2-3 lateral veins per side (on extension shoots 3-4 pairs); thinly hairy. Inflorescences 3-5 flowered; branches

densely pubescent. Flowers 15-20 mm diam.; hypanthium pubescent; sepals 3-4.5 mm long, margins

glandular-serrate; stamens 20, ivory anthers; styles 3-5. Fruit 10-12 mm diam., reddish-orange to yellow,

washed red; sepals reflexed; nutlets 3-5, plane-sided, grooved dorsally.

Habitat and Distribution— Known with certainty only from the type, more material is needed to come

to any definite conclusions.

Comment.—In foliage and fruit, except for the narrower leaves, C. annosa is of typical form for ser.

Apricae, but in anther number and color and the form of the narrower leaf set among the short-shoot leaves

it is a typical member of ser. Lacrimatae.

EXCLUDED SPECIES

SERIES INTRICATAE

Crataegus flava Aiton, Hortus Kewensis 1:169. 1789. Te: U.K: Kew, cultivated, 1781, Herb. Bishop Goodenough (HOLOTYPE:

K). See Phipps 1988a for a line drawing, photograph of type, and distribution map

Shrubs or small trees, 5-6 m. tall; branchlets + straight; extending twigs glabrous; l-year old twigs glabrous

Q always), purple-brown; older dark gray; 2-year old thorns 2-3 cm long, slender, purple-brown, straight

to slightly recurved. Leaves deciduous; petioles 0.75-1.5 cm long, slightly pubescent, winged above, con-

spicuously black-glandular, the glands usually sessile, sometimes short-stipitate; blades 5-8 cm long in

UK-cultivated material, rhomb-obovate in general outline, shallowly 1-3 lobed per side, lobes obtuse to

acute, at the apex subacute to obtuse, at the base cuneate, and tapered into the winged upper part of the

petiole; margins crenate to crenate-dentate, the teeth gland-tipped; venation craspedodromous, 3-4 pairs

of lateral veins; pubescent above when young, glabrescent. Inflorescences 4-6 flowered; branches slightly

villous or glabrous, bearing caducous, linear, membranous, gland-margined bracteoles; anthesis early April.

Flowers 16-18 mm diam.; hypanthium externally glabrous; calyx lobes 4—5 mm long, narrow triangular,

gland-margined; petals + circular, white; stamens 10, anthers purple; styles 3-5. Fruit 8-12 mm diam.,

+ pyriform-oblong, dull orange; calyx lobes + reflexed, slightly elevated on a collar; nutlets 3-5, dorsally

sulcate, laterally smooth.

Common Name.—Yellow hawthorn.

Habitat and Distribution.—This is a rare species of the southeast United States from southeastern Georgia

to South Carolina and northern Florida with one record from Alabama and another from North Carolina.

There are also possible records for Virginia. It is usually found in deep sandy soils.

Comment.—Long known in cultivation in England where it was described by William Aiton at Kew.

Unfortunately the name C. flava got transferred to C. lacrimata Small or a similar species due to an error by

Sargent (1902) that has been perpetuated by many workers on the flora of the southeastern United States.

Sargent's (1890) treatment of this species is, however, nearer the mark in leaf shape although typical mate-

rial is glabrous to nearly so in the inflorescence. Aiton's original interpretation was eventually resuscitated

by the first author (Phipps 19882). The true C. flava differs in significant ways from other species of series

Flavae, sens. auctt. Amerr. (= ser. Lacrimatae + ser. Apricae) as is discussed in the treatment for ser. Apricae.

In its new series (ser. Intricatae) C. flava shows most similarity to C. rubella in fruit shape and color as well

as general glandularity. It is no longer recorded in cultivation.

Phipps and Dvorsky, Crataegus ser. Apricae and C. flava 199

No recent collections match typical C. flava though several from central and southern Georgia have the

foliage characters of C. flava but possess a more globose fruit and ca. 20 stamens on which basis they would

key to an unknown form of ser. Pulcherrimae not referred to in Phipps and Dvorsky (2006). The unique leaf

shape is substantially the same as in C. sp. cf. C. flava from the Augusta sand hills, discussed above, but the

forms currently alluded to have straight twigs. Much more material is needed to sort out this assemblage of

forms.

APPENDIX

Further cited specimens (species, states, and counties arranged alphabetically)

Crataegus alleghaniensis Beadle

ALABAMA. Etowah Co.: Hind's Rock, near Noccalua Falls, 8 Apr 1998, J.B. Phipps 7655 (UWO). Jefferson Co.: along AL 150

ca. 0.5 mi S of junction with Co. Rd. 972, J.P. Barber, M.G. Bussey, R.D. Whetstone & K.E. Landers 1546 (JSU, UWO). GEORGIA.

Richmond Co.: Augusta, 15 Apr 1902, A. Cuthbert s.n. (FLAS); Augusta, 27 Apr 1903, A. Cuthbert s.n. (FLAS). Screven Co.: Blue

Springs, 25 Apr 1966, R.L. Park s.n. (NCU). TENNESSEE: Cocke Co.: within 3 mi of Wolf Creek Station, along the French Road,

24 Aug 1897, Thos. H. Kearney, Jr. 697 (UWO photo of US 313179).

Crataegus annosa Beadle

ALABAMA. Russell Co.: Phenix City, 1901, Biltmore Herb. 4103? (A); Phenix City, 26 Aug 1901, C.D. Beadle 4108? (A).

Crataegus sp. cf. C. annosa Beadle

FLORIDA. Liberty Co.: Torreya State Park, 21 Oct 1950, H. Kurz s.n. (FSU); Torreya State Park, 24 Mar 1951, H. Kurz s.n. (FSU).

Wakulla Co.: on US 98, jct. FLA 365, 11 Apr 1966, Beckner and D'Arcy 932 (LAF, FLAS). NORTH CAROLINA. Durham Co.: 9

May 1901, WW. Ashe 1949 (CM). SOUTH CAROLINA. Aiken Co.: SC 4, 7 km E of 394, 11Aug 1993, J.B. Phipps 6676 (UWO).

Camden Co.: 27 Mar 1923, T.G. Harbison 6052 (A). Chesterfield Co.: 6 Jun 1976, J. Castrale 184 (USCH); Haw Ridge, 3 Apr 1935,

V. Matthews and B.E. Smith 218 (USCH).

Crataegus aprica Beadle

ALABAMA. Barbour Co.: Hwy. 55 300 m E of Mt. Olive, 8 Jun 1992, DJ. Drennen and J. Daniel 00039 (UWO). FLORIDA. Alachua

Co.: 15 Jun 1940, WA. Murrill s.n. (GA). Gadsden Co.: Aspalaga, May, J.K. Small 297c (NY); Aspalaga, 1898, no collector s.n. (MO);

Quincy (14 mi W of), 6 Sep 1940, WB. Tisdale and W. A. Murrill s.n. (FLAS); Chattahoochee, 15 Apr 2006, J.B. Phipps $: R. Lance

9079 (UWO). GEORGIA. Burke Co.: Rte 56, N of intersection with 80, N of Lake Crystal Rd. to E, 9 Apr 1991, J.B. Phipps 6497

(UWO). Houghton Co.: Oaky Woods WMA, 14 Apr 2006, J.B. Phipps & R. Lance 9061 (UWO). Paulding Co.: 3/4 mi W of Dallas, 1

Aug 1948, W.H. Duncan 8643 (MO). Randolph Co.: SE of Coleman, 9 Apr 2001, R. Lance, 2107 (UWO). Richmond Co.: Augusta,

no date, S.F. Olney and J. Metcalf s.n. (NY); Columbia, 19 May 1934, EJ. Palmer 42408 (NY); Augusta, 11 Apr 1916, WW. Ashe 26

(NCU). NORTH CAROLINA. Buncombe Co.: (60) E of Asheville, Swannanoa River, 28 Sep 1888, W.M Canby s.n. (NY); Biltmore,

20 Sep 1902, Biltmore Herb. C. (NY); near Asheville, 18 Apr 1929, EJ. Palmer 35436 (NY, A); Biltmore, alt. 2000-2500 ft, 15-19 Sep

1908, WW. Eggleston 4125 (NY); Biltmore, 12-15 May 1897, Biltmore Herb. 297b (NY); Biltmore, 7 May 1899, WM. Canby 34 (DOV);

Biltmore, 11 May 1899, Biltmore Herb. s.n. (NY); Biltmore, 7 Sep 1899, Biltmore Herb. s.n. (NY); Biltmore, 17 Sep 1897, Biltmore

Herb. 297b (NY); Biltmore, 8 May 1896, WW. Eggleston 297 (NY); Biltmore, 12 Aug 1896, WW. Eggleston 297 (NY); Biltmore, 20 Sep

1902, C.D. Beadle C (NY); Hwy. 191 0.6 mi S of junction with Blue Ridge Pwy., Oct 1993, R. Lance 93.17 (UWO); Biltmore, 15 Sep

1908, WW. Eggleston 4125 (NY); Biltmore, 12 May 1897, C.D. Beadle 297b (NY); Biltmore Estate, by riverside on steep cliff, 7 Sep

1984, PRUIFHansen PF119 (UWO); Asheville, 15 Sep 1908, WW. Eggleston 4125 (NY); E of Asheville, 28 Sep 1888, Wm. Canby s.n.

(NY); Biltmore, 8 May 1896, C.D. Beadle 297 (NY, NCC); Biltmore, 15 May 1902, C.D. Beadle C (NY); near Asheville, 18 Apr 1929, EJ.

Palmer 35436 (A, MO); Biltmore Estate, 15 May 1902, C.D. Beadle 2335 (CM); Biltmore, 7 May 1899, C.S. Sargent s.n. (A); Biltmore,

23 Apr 1933, T.G. Harbison &Totten s.n. (NCU); Asheville, 15 Aug 1909, WW. Eggleston 4425 (MO); Biltmore, 11 May 1899, C.D.

Beadle C14 (GH, NY); near Biltmore, 11 May 1899, C.D. Beadle C13 (A); Asheville, 26 Oct 1898, C.S. Sargent s.n. (A); Biltmore, 28

Sep 1917, C.D. Beadle s.n. (A); Biltmore, 7 Sep 1899, C.D. Beadle C500 (A, NY); Biltmore, 15 Sep 1908, WW. Eggleston 4125 (UWO);

Biltmore, 11 Sep 1899, C.D. Beadle C500=C 14 (GH); Biltmore, 15 May 1897, Biltmore Herb. 297b (GH, MO); Biltmore, 24 Jul 1891,

Biltmore Herb. C42 (A); along Hwy 191, 0.6 mi S of junction with Blue Ridge Parkway, S end of “Sandy Bottoms" picnic area, Oct

1993, R. Lance RO-93.17 (UWO); 19 Sep 1908, WW. Eggleston 4125 (BH, CM, GH); Sep 1926, WW. Ashe s.n. (NCU); 10 Oct 1905,

T.G. Harbison 12 (A); no date, Biltmore Herb. 318 (A); 8 May 1896, Biltmore Herb. 297 (DOV, NCC); 15 May 1902, Biltmore Herb. 2027

(DOV). Cumberland Co.: Fayetteville, 12 Oct 1908, WW. Eggleston 4017a (NY); Fayetteville, 31 Oct 1908, WW. Eggleston 4016

(NY). Henderson Co.: 3.5 mi SE of Hendersonville, 17 May 1942, FR. Fosberg 18791 (NUC); Hendersonville, no date, H.L.B. 8819

(DUKE); Flat Rock, 9 Sep 1899, no collector, s.n. (NCU). Wake Co.: William B Umstead State Park, 22 Aug 1964, G. Sawyer & W.

Ahles 1539 (USCH). Unknown Co.: 31 May 1907, WW. Ashe 1949 (CM); Western North Carolina, 20 Aug 1901, WW. Ashe 1949

(CM); Western North Carolina, 31 May 1907, WW. Ashe 1949 (CM); 7 Jun 1900, A.R. s.n. (MO). SOUTH CAROLINA. Aiken Co.:

12-15 Sep 1909, WW. Eggleston 5031 (MO). Chesterfield Co.: On Haw Ridge, 23 Apr 1933, T.G. Harbison s.n. (NCU). Darlington

Co.: Hartsville, Swift Creek, 5 May 1941, E.B. Smith 949 (USCH); 6.5 mi S of Patrick on SC 102, 18 Jul 1958, J.A. Duke 1556 (NUC).

200 Journal of the Botanical R h Institute of Texas 1(1)

Greenville Co.: lower slopes of Caesar's Head Mt., 20 Sep 1934, E.T. Wherry s.n. (A); Caesar's Head Mt., 20 Sep 1934, ET. Wherry

s.n. (A). Kershaw Co.: Middleton, 2 mi S of Camden Junction, 20 Apr 1897, L.F. Ward s.n. (NY). Richland Co.: near Columbia;

woods, 19 May 1934, EJ. Palmer 42408 (A, NY, SC). TENNESSEE: Union Co.: Area 24 Pill, Walkers Pond Refuge, 17 Jul 1936,

L.B. Kalter 361 (TENN). VIRGINIA: Franklin Co.: Franklin, 27 Aug 1909, WW. Eggleston 4925 (NY). Southampton Co.: 29 Jun

1893, A. Heller s.n. (GH, MO); 29 Jun 1893, A. Heller 20a (A).

Crataegus egregia Beadle

ALAB Macon Co.: Tuskegee National Forest, 2 Apr 1967, T. Rankin 21 (NCU). FLORIDA. Clay Co.: Magnolia Springs, 23

Sep 1923, T.G. Harbison 5664 (A). Liberty Co.: Bristol, 24 Aug 1908, Biltmore Herb. No. 4924 (A); Bristol, 31 Mar 1907, Biltmore

Herb. No. 4924 (A). Wakulla Co.: S of Wakulla Springs, 21 Aug 1951, H. Kurz s.n. (FSU). GEORGIA. Richmond Co.: Augusta,

sandhills, 9 Sep 1902, A. Cuthbert s.n. (FLAS). SOUTH CAROLINA. Aiken Co.: no location, 20 Jun 1952, WR. Kelley & WT. Batson

s.n. (USCH); no location, 14 Jul 1952, WR. Kelley & W.T. Batson s.n. (USCH). Barnwell Co.: Barnwell State Park, 27 Aug 1956, CL.

Porter 342 (USCH); no location, no date, WR. Kelley € W.T. Batson s.n. (USCH).

Crataegus extraria Beadle and C. cf. extraria Beadle

ALABAMA. Lee Co.: Auburn, 18 Apr 1896, Farle and Underwood (NY); Auburn, 18 Apr 1894, C. Mohr (CM). GEORGIA. Lumpkin

Co.: Porter Springs, WW. Ashe (NCU). NORTH CAROLINA. Cumberland Co.: Fayetteville, 26-31 Aug 1908, WW. Eggleston 4019

(NY); Fayetteville, 16 Apr 1903, Biltmore Herb. B8066=B7272 (NY); Fayetteville, 17 Oct 1902, Biltmore Herb. B7272 (NY); Fayetteville,

17 Oct 1902, Biltmore Herb. 7272 (NY); Fayetteville, 16 Apr 1903, Biltmore Herb. 8006 (NY). Richmond Co.: Washington Rd., N of

Augusta, GA, 14 Oct 1916, WW. Ashe (NCU). VIRGINIA: Wight Co.: S of Zuni, 17 Oct 1936, M.L. Fernald & B.Long 6818 (NY).

Crataegus frugiferens Beadle

ALABAMA. Calhoun Co.: Fort McClellan Military Reservation, 8 Apr 1998, J.B. Phipps 7645 (UWO); Anniston, Natural History

Museum grounds, 30 Sep 1998, J.B. Phipps & D. Spaulding 7771 (UWO); Coldwater Mountain, woodland border next logging

road, 20 Apr 1999, D. Spaulding 10606 (UWO). Cherokee Co.: Cherokee 19, 1 mi N of 278, 8 Oct 1998, J.B. Phipps 7834 (UWO).

Chilton Co.: Bluffs above quarry at Coosa River, 4 Oct 2000, J.B. Phipps 7669 (UWO); Bluffs above quarry at Coosa River, 4 Oct

2000, J.B. Phipps hee Madison Co.: Deer Haven Rd., NE of Huntsville, 25 Apr 1983, PF Ulf-Hansen 034 (UWO). Marshall

Co.: Lake Guntersville State Park, Cutchenmire Trail near Berry Point, 21 Apr 1999, J.B. Phipps, RJ. O'Kennon & D. Spaulding 7968

(UWO). Pickens Co.: 8.3 mi E of Ethelsville, 5 Apr 1968, S. McDaniel 10447 (ALU, FSU, IBE, LAF, NO). Shelby Co.: near Pelham,

AL 33, approx 0.75 km N of Co. Rd. 52, 5 Apr 1984, J.B. Phipps & T.C. Wells 5309 (UWO); near Pelham, E side of AL 33, approx 0.75

km N of Co. Rd. 52, 5 Apr 1984, J.B. Phipps & T.C. Wells 5310 (UWO). St. Clair Co.: Pottsville, 4 Jun 1963, PE. Bostick s.n. (NCU).

GEORGIA. Floyd Co.: McGee Bend Rd., off GA 100 (WSW of Rome), 4 Apr 2000, J.B. Phipps 8207 (UWO). Houston Co.: Oaky

Woods WMA, Green Violet Prairie, 14 Apr 2006, J.B. Phipps & R. Lance 9067 (UWO). MISSISSIPPI: Chickasaw Co.: MS 41 to SE

of Natchez Trace Parkway, 13 Apr 1998, J.B. Phipps 7714 (UWO).

Crataegus galbana Beadle

ALABAMA. Conecuh Co.: Hwy. 29 at Sepulga River, 17 Aug 1985, A.R. Diamond 1529 (AUA). Geneva Co.: 7.5 mi S of Samson,

7 Jun 1969, R. Kral 35109 (AUB); near Smith Lake, 12 Sep 1967, J. Thomas 1218 (ALU). Greene Co.: near Smith Lake, 12 Sep 1967,

J.L. Thomas 1218 (ALU). Lee Co.: Auburn, 10 Oct 1900, FS. & E. Earle 23 (GH). FLORIDA. Alachua Co.: 30 Jun 1927, Manning

and Wiegand 1386 (GH); Nr. Gainesville, 14 Apr 1941, W.A. Murrill s.n. (GA); NW of Gainesville, 10 Sep 1940, W.A. Murrill s.n. (A).

Columbia Co.: Camp Oleno, 8 Sep 1940, W.A. Murrill s.n. (GA). Gadsen Co.: 8 mi SW of Chattahoochee, 20 Aug 1966, D.B.

Ward 5918 (FLAS); 1 mi S of River Junction, 6 Sep 1940, WB. Tisdale & W.A. Murrill s.n. (FLAS). Holmes Co.: 1 mi N of Westville,

17 Mar 1982, R.K. Godfrey 79427 (FSU, IBE); Nr. Westville, 31 Mar 1940, M.G. Henry 1993 (GA). Jackson Co.: 2 mi E of Grand Ridge,

27 Mar 1964, S. McDaniel 4011 (IBE, FSU); 2 mi E of Grand Ridge, 27 Mar 1964, S. McDaniel 4011 (IBE). Jefferson Co.: No local-

ity, 10 Jul 1940, WA. Murrill s.n. (FLAS). Lafayette Co.: 4 mi W of Mayo, 21 Aug 1939, WA. Murrill and W.B.Tisdale s.n. (A). Leon

Co.: Tallahassee, 3 Apr 1901, Biltmore Herb. 4052 (GH); Tallahassee, 29 Aug 1901, Biltmore Herb. 4948 (GH); Tallahassee Junior

Museum, SW of Tallahassee, 17 Aug 1982, R.K. Godfrey79926 (FSU, UWO); Tallahassee, Apr 1893, C.S. Sargent s.n. (MO). Madison

Co.: 3 mi NNE Pinetta, 2 Sep 1968, R.A. Norris 1086 (IBE); Lee (2.5 mi E of), 22 Aug 1940, WA. Murrill s.n. (FLAS). Suwanee Co.:

O'Brien (5 mi W of), 9 Aug 1946, West and Arnold s.n. (FLAS). Wakulla Co.: US 98, W of jct. FL 365, 11Apr 1966, J. Beckner s.n.

(LAF). GEORGIA. Burke Co.: On Savannah River, 20 Jun 1941, WM. Duncan 3385 (GA). Chattooga Co.: Summerville, no date,

Biltmore Herb. 6113 (NY). Floyd Co.: no location, no date, A.W. Chapman s.n. (AUB); Rome, 22 Apr 1900, C.S. Sargent s.n. (GH).

Richmond Co.: Augusta, Sand Bar Ferry, 14 Aug 1902, A. Cuthbert s.n. (FLAS). NORTH CAROLINA. Harnett Co.: 2miW of

Spout Springs, 29 Sep 1970, R.M. Downs 11614 (NCSC). Johnston Co.: Selma, 9 Apr 1912, T.G. Harbison 10834 (NCU). SOUTH

CAROLINA. Kershaw Co.: 6 mi E of Camden, 17 Jun 1933, T.G. Harbison & H.R. Totten 4052 (NCU). McCormick Co.: Clark Hill

Dam, 5 Nov 1900, C. McComb 97 (GA).

Crataegus ignava Beadle

ALABAMA. Blount Co.: Ridge of Blount Mtn., 25 Apr 1983, P.F. Ulf-Hansen 31 (UWO). E of Highland Lake on ridge of Blount

Mtn., 25 Apr 1983, PF. Ulf-Hansen 33 (UWO). Calhoun Co.: Fort McClellan Military Reservation, near Anniston, 30 Sep 1998,

J.B. Phipps & R. Smith 7773 (UWO); Fort McClellan Military Reservation, near Anniston, 8 Apr 1998, J.B. Phipps 7642 (UWO); Fort

McClellan Military Reservation, near Anniston, 2 Jul 1998, R. Smith & D. Spaulding A, B, C (UWO, 3); Fort McClellan Military Res-

Phipps and Dvorsky, Crataegus ser. Apricae and C. flava 201

ervation, near Anniston, 22 Apr 1998, R. Smith & D. Spaulding 4 (UWO); Logging Road on Coldwater Mountain, 20 Apr 1999,

D. Spaulding 10605 (UWO). Cherokee Co.: AL 9, N of Piedmont, 19 Apr 1999, J.B. Phipps & R. J. O'Kennon 7947 (UWO). Etowah

Co.: Hind's Rock, near Noccalua Falls, 8 Apr 1998, J.B. Phipps 7655 (UWO). Jefferson Co.: along AL 150 ca. % mi S of jct. with

Co. Rd. 97 and 2, 23 May 1985, J.P Barber, M.G. Bussey, R.D. Whetstone and K.E. Landers 1546 (JSU, UWO). FLORIDA. Gadsden

Co.: S of Chattahoochee, 15 Apr 2006, J.B. Phipps, R. Lance & A.. Gholson 9090 (UWO). GEORGIA. Burke Co.: NE of Waynesboro,

s of McBean 11 Apr 2006, J.B. Phipps 9040 (UWO). Screven Co.: Blue Springs, 25 Apr 1966, R.L. Park s.n. (NCU). Randolph Co.:

Few mi W of Cuthbert, 29 Mar 1948, R.F. Thorne & W.C. Muenscher 7748 (UWO, photo). Richmond Co.: Augusta, 15 Apr 1902;

A. Cuthbert s.n. (FLAS); Augusta, 23 Apr 1903; A. Cuthbert s.n. (FLAS). TENNESSEE: Cocke Co.: within 3 mi of Wolf Creek Station,

along the French Road, 24 Aug 1897, Thos. H. Kearney Jr. 697 (UWO).

Crataegus leonensis E.J. Palmer

FLORIDA. Gadsden Co.: Chattahoochee, May 1899, Wm Canby & C.S. Sargent 27 (DOV). Leon Co.: Tallahassee, Horseshoe

Plantation, 28 Mar 1914, C.S. Sargent s.n. (A); Tallahassee, Horseshoe Plantation, 16 Sep 1919, T.G. Harbison 2 (A); Tallahassee,

Horseshoe Plantation, 6 Apr 1920, 7.G. Harbison 5645, 5646 (A,2); Tallahassee, Horseshoe Plantation, Oct 6 1920, T.G. Harbison

5648 (A); Tallahassee, Horseshoe Plantation, 3 Apr 1923, T.G. Harbison 6071 (A); Tallahassee, Horseshoe Plantation, 27 Sep 1923,

T.G. Harbison 6182, 6183 (A, 2); Tallahassee, Horseshoe Plantation, 12 Apr 1931 EJ. Palmer 38557 (A); Lake Lamonia, 15 Mar 1951,

EJ. Palmer s.n. (FSU). GEORGIA. Decatur Co.: 7 mi S of Bainbridge, 1 Apr 1970, R.K. Godfrey 69336 (FSU).

Crataegus mira Beadle

ALABAMA. Cleburne Co.: Talladega Nat. Forest, 5 mi S of Fruithurst, FS. Road 2700, 18 Jun 1966, T.A. Heard & R.C. Clarke

2767 (NCU). Franklin Co.: no locality, 21 Apr 1968, J.G. South s.n. (JSU). FLORIDA. Alachua Co.: W of Gainesville, 1 Aug 1940,

W.A. Murrill s.n. (GA). Columbia Co.: 2 mi from main road towards Camp Oleno, 30 Jun 1940, WA. Murrill s.n. (FLAS); By rock

at Camp Oleno, 13 Apr 1941, WA. Murrill s.n. (FLAS). Gadsden Co.: Aspalaga, May, no day, no year, AW. Chapman 297c (NY);

Aspalaga, 1898, A.W. Chapman 2799 (MO); 6 mi S of Quincy on the Blountstown Rd., 12 Sep 1928, WW. Ashe s.n. (NCU). Leon

Co.: Tallahassee, by US 319 near Neesmith's Nursery, 11 Apr 1983, R.K. Godfrey 80460 (FSU, UWO - 5); W of Tallahassee, 1 mi E

of Ochlockonee R., 17 Mar 1982, R.K. Godfrey 78429 (FSU); Tallahassee, by US 319, near Neesmith's Nursery, 20 Jun 1983, R.K.

Godfrey 80732 (UWO). Liberty Co.: Torreya State Park, 30 Mar. 1975, R.K. Godfrey 74198 (FSU, FLAS). Suwannee Co.: Imi S of

O'Brien, 22 Aug 1940, WA. Murrill s.n. (FLAS). GEORGIA. Burke Co.: Rte. 56, 13 Apr 1993, J.B. Phipps 6709 (UWO). Floyd Co.:

Radio Springs Road, near metal shack, 24 Apr 1983, PF. Ulf-Hansen s.n. (UWO). Dodge Co.: E side of Ocmulgee River, 6 Jul

1966, J.R. Bozeman 5508 (NCC). Pulaski Co.: Jct. Wimberly Rd. and US 341, 10 Apr 2001, R. Lance 2123 (UWO). Paulding Co.:

10 mi SW of Dallas, 1 Aug 1948, WH. Duncan 8643 (GA). Randolph Co.: SE of Coleman by hwy. 160, 10 Apr 2001, R. Lance 2117

(UWO); S of Cuthbert by US 27, 10 Apr 2001, R. Lance 2121 (UWO). Richmond Co.: Augusta, 21 Apr 1903,A. Cuthbert 935 (FLAS);

Augusta, 24 Apr 1901, A. Cuthbert s.n. (FLAS); Augusta, 6 Jul 1882, M.W. Ravenel s.n. (MO); Augusta, 24 Apr 1902 C.S. Sargent 67

(UWO, photo); Augusta, 21 Apr 1902, A. Cuthbert 628 (FLAS). NORTH CAROLINA. Cumberland Co.: Fayetteville, 12 Oct 1908,

WW. Eggleston 4016a, (GH, NY); Fayetteville, Haymount, 12 Oct 1908, WW. Eggleston 401 7a (NY); Fayetteville, Haymount, 12

Oct 1908, WW. Eggleston 4016 (NY). Dublin Co.: 2.6 mi N of Magnolia, 27 Apr 1957, H.F. Ahles 24051 (NCC). Wake Co.: Camp

Crabtree waterfront, Wm. B. Umstead State Park, 22 Aug 1964, G.P Sawyer, Jr, H.E. Ahles & J.B. Whitney 1539 (USCH). SOUTH

CAROLINA. Aiken Co.: by US 302, W side, 12 Aug 1993, J.B. Phipps 6684 (UWO); Aiken, 12-15 Sep 1909, WW. Eggleston 5031

(MO). Horry Co.: Little River, Neck Road, 1 May 1966, J.F. Matthews, W.C. Williams & J.L. Kellerman s.n. (UCC). Lexington Co.:

Rte. 303, W side of road, 10 Apr 1991, J.B. Phipps 6504 (UWO).

Crataegus segnis Beadle

FLORIDA. Leon Co.: 3 mi N of Chaires, 29 Mar 1956, R. Kral 2160 (FSU). Liberty Co.: Torreya State Park, 22 Mar 1982, R.K. Godfrey

79451 (FSU). Wakulla Co.: Vicinity of Crawfordville, 29 Mar 1975, R.K. Godfrey 74196 (FSU). Walton Co.: Black Creek Rd. off US

331, 30 Mar 1975, Mr. & Mrs. H.A. Davis 16321 (FSU).

Crataegus sororia Beadle

ALABAMA. Calhoun Co.: No location, 7 Apr 1988, R. Lance s.n. (UWO). GEORGIA. De Kalb Co.: Yellow River, near Stone Mtn,,

15 Jul 1899, A.H. Curtiss 6920 (DOV). Floyd Co.: Rome, hills above Silver Creek, Sep 1899, C.D. Beadle s.n. but from ‘type tree’ (A);

Rome, Apr 1902, C.S. Sargent s.n. (A); Rome, hills above Silver Creek, cotype, 18 Apr 1899, C.D. Beadle 1257 (A); Rome, 22 Sep

1902, C.D. Beadle 7142 (A, 3); Rome, 26 Oct 1905, T.G. Harbison 2138 = tree 35 (A); Horseleg Mtn., Rome, 7 Oct 1982, J.B. Phipps

5176 (UWO); Horseleg Mtn., Rome, 4 Apr 1984, J.B. Phipps 5302 (UWO); Horseleg Mtn., Rome, 23 Apr 1983, PF. Ulf-Hansen 020,

021 (UWO). SOUTH CAROLINA. Aiken Co.: Nr. Aiken, 4 Jun 1880, G. Engelmann 14 (MO); Hwy. 47 km E of jct. 394, 16 Apr

1999, J.B. Phipps 6512 (UWO).

Crataegus visenda Beadle

ALABAMA. Dallas Co.: Selma, 11 Apr 1912, T.G. Harbison 10827 (NCU); Rte. 140, ca. 9 mi E of Selma, 10 Apr 1998, J.B. Phipps 7689

(UWO). Macon Co.: 2.3 mi E of Tuskegee, 2 Apr 1973, (ALU). FLORIDA. Clay Co.: Magnolia Springs, 7 Apr 1920, T.G. Harbison

15678 (UWO, photo). Gadsden Co.: Chattahoochee, 5 Apr 1900, C.S. Sargent s.n. (DOV); River Junction at Dolan Road, 23 Jun

1983, R.K. Godfrey 79895 (UWO). Jackson Co.: Three Rivers State Park, N of Sneads, 10 Jun 1982, R.K. Godfrey 79854 (UWO). Leon

Co.: no date, T.G. Harbison 5453 (NCU); 1.3 mi E of Micosukee, 7 Aug 1951, H. Kurz s.n. (FSU, 2); Tallahassee, 15 Apr 1920, T.G.

202 Journal of the Botanical R h Institute of Texas 1(1)

Harbison 15718 (NCU); Tallahassee, 24 Mar 1951, H.A. Kurz s.n. (FSU). Liberty Co.: Bristol, 2 Apr 1902, 7.G. Harbison 6022 (NCU);

Bristol, 29 Mar 1902, T.G. Harbison 6012 (TENN); Bristol, 30 Aug 1901, T.G. Harbison 4031 (A); Marion Co.: Rainbow Springs, 11

Nov 1945, H.R. Totten s.n. (NCU). Wakulla Co.: E of Wakulla River, 13 Aug 1951, H. Kurz s.n. (FSU); no locality, 11 Apr 1966, J.

Beckner & W. D'Arcy s.n (FLAS); near Crawfordville, 2 Apr 1955, R.K. Godfrey 53118 (ALU, FLAS, FSU, GA, NCSC, TENN, USF); 1.5 mi

S of Crawfordville, 25 Mar 1982, R.K. Godfrey 79461 (IBE). GEORGIA. Randolph Co.: SE of Coleman, W of Co. Rd., 4.2 mi N of

jet. with unknown Hwy., 9 Apr 2001, R. Lance 2114 (UWO).

ACKNOWLEDGMENTS

The authors wish to thank Susan Laurie-Bourque for her fine line illustrations, The National Sciences and

Engineering Research Council of Canada for continuing support to the first author and the curators of the

following herbaria for loan of specimens: A, ALU, AUA, AUB, BH, CM, DOV, DUKE, FLAS, FSU, GA, GM,

IBE, JSU, LAF, MARY, MO, NCC, NCSC, NCU, NO, NY, TENN, UNA, USCH, WVA.

REFERENCES

BeabLe, C.D. 1902. New species of thorns from the southeastern United States, Il. Biltmore Bot. Stud. 1:62.

Beadle, C.D. 1903. Crataegus. In: J.K. Small. Flora of the southeastern United States, ed. 2. Author, New York.

Cronauist, A. 1980. Flora of the southeastern United States. Univ. of North Carolina Press, Chapel Hill.

Kunz, H. and R.K. Goprrey. 1982. Crataegus. In: Trees of Northern Florida. Univ. of Florida Press, Gainesville.

Murri, W.A. 1942. New Florida hawthorns. Castanea 7:19-30.

PALMER, E.J. 1925. Synopsis of North American Crataegus. J Arnold Arbor. 6:5-128.

PALMER, E.J. 1960. Crataegus. In: R.A. Vines. Trees, shrubs and woody vines of the southwest. Univ. of Texas Press,

Austin. Pp. 329-387.

Puipps, J.B. 1988a. Re-asessment of Crataegus flava Aiton and its nomenclatural implications for the Crataegus

serial name Flavae (Loud.) Rehd. and its sectional equivalent. Taxon 37:108-113.

Puipps, J.B. 1988b. Crataegus (Maloideae, Rosaceae) of the southeastern United States, |. Introduction and series

Aestivales. J. Arnold Arbor 69:401-431.

Phipps, J.B. and K.A. Dvorsky. 2006. Review of series Pulcherrimae (Crataegus, Rosaceae). Sida 22:973-1007.

Phipps, J.B., R.J. O’KENNON, and R. Lance. 2003. Hawthorns and medlars. Timber Press, Portland, OR.

Puipps, J.B., K.R. RosERTsON, PG. SmitH, and J.R. Rorer. 1990. A checklist of subfam. Maloideae (Rosaceae). Canad. J.

Bot. 68:2209-2269.

SARGENT, C.S. 1890. Silva of North America. IV:113, t. 189.

SARGENT, C.S. 1902. Silva of North America. XIII, suppl.:155, t. 693.

Tarent, N. and T.A. Dickinson. 2005. Polyploidy in Crataegus and Mespilus (Rosaceae, Maloideae): evolutionary

inferences from flow cytometry of nuclear DNA amounts. Canad. J. Bot. 83: 1268-1304.

TipesTROM, |. 1933. Crataegus. In: J.K Small. Manual of the southeastern flora. Univ. of North Carolina Press, Chapel

Hill.

TAXONOMY AND NOMENCLATURE OF TAXUS (TAXACEAE)

Richard W. Spjut

World Botanical Associates

Bakersfield, California 93380-1145, U.S.A.

ABSTRACT

A taxonomic treatment of Taxus (Taxaceae) is presented, based on morphological characters. The genus is proposed to have 24 species

and 55 varieties; 24 Species and 26 varieties are b ius in a key and classified into three main groups, two subgroups, and two alli-

ances. Previously existing names were applied to 15 species and six varieties— T. baccata L. and its varieties—var. dovastoniana Leighton,

icd

var. elegantissima Hort. ex C. Lawson, var. glauca Jacques ex Carrière, var. pyramidalis Hort. ex C. Lawson, and var. variegata Watson;

T. brevifolia Nutt., T. caespitosa Nakai, T. canadensis Marshall, T. celebica (Warb.) H.L. Li, T. chinensis (Pilg.) Rehder, T. contorta Griff., T.

cuspidata Siebold & Zucc., T. fastigiata Lindl., T. globosa Schltdl., T. mairei (Lemée € H. Lév.) S. Y. Hu ex T.S. Liu, T. recurvata Hort. ex

C. Lawson, T. sumatrana (Miq.) de Laub., T. umbraculifera (Siebold ex Endl.) C. Lawson, T. wallichiana Zucc. and var. yunnanensis (W.C.

Cheng & L.K. Fu) C. T. Kuan. Six new species—T. biternata Spjut, T. florinii Spjut, T. kingstonii Spjut, T. obscura Spjut, T. phytonii

Spjut, and T. suffnessii Spjut, and four new varieties—T. brevifolia Nutt. var. polychaeta Spjut, T. brevifolia Nutt. var. reptaneta

Spjut, T. caespitosa Nakai var. angustifolia Spjut and T. contorta Griff. var. mucronata Spjut are described. Eight new combinations

are made: T. caespitosa var. latifolia (Pilg.) Spjut, T. canadensis var. adpressa (Carrière) Spjut, T. canadensis var. minor (Michx.)

Spjut, T. globosa var. floridana (Nutt. ex Chapm.) Spjut, T. mairei (Lemée & H. Lév.) S. Y. Hu ex T. S. Liu var. speciosa (Florin) Spjut, T.

TS 1c

umbraculifera var. hicksii (Hort ) Spjut, T. umbraculifera var. microcarpa (Trautv.) Spjut, and T. umbraculifera (Siebold

ex Endl.) C. Lawson var. nana (Rehder) Spjut. Taxonomy and nomenclature are discussed for each species and variety.

RESUMEN

Se presenta un tratamiento taxonómico de Taxus (Taxaceae) basado en caracteres morfológicos. Se propone que el género tenga 24 es-

pecies y 55 variedades; 24 especies y 26 variedades se presentan en una clave y se clasifican en tres grupos principales, dos subgrupos,

y dos alianzas. Los nombres existentes previamente se aplicaron a 15 especies y seis variedades—T. baccata L. y sus variedades—var.

dovastoniana Leighton, var. elegantissima Hort. ex C. Lawson, var. glauca Jacques ex Carrière, var. pyramidalis Hort. ex C. Lawson, y var.

variegata Watson; T. brevifolia Nutt., T. caespitosa Nakai, T. canadensis Marshall, T. celebica (Warb.) H.L. Li, T. chinensis (Pilg.) Rehder,

T. contorta Griff., T. cuspidata Siebold & Zucc., T. fastigiata Lindl., T. globosa Schltdl., T. mairei (Lemée & H. Lév.) S. Y. Hu ex T.S. Liu,

T. recurvata Hort. ex C. Lawson, T. sumatrana (Miq.) de Laub., T. umbraculifera (Siebold ex Endl.) C. Lawson, T. wallichiana Zucc. y var.

yunnanensis (W.C. Cheng & L.K. Fu) C. T. Kuan. Se describen seis especies nuevas—T. biternata Spjut, T. florinii Spjut, T. kingsto-

nii Spjut, T. obscura Spjut, T. phytonii Spjut, and T. suffnessii Spjut, and four new varieties— T. brevifolia Nutt. var. polychaeta

Spjut, T. brevifolia Nutt. var. reptaneta Spjut, T. caespitosa Nakai var. angustifolia Spjut y T. contorta Griff. var. mucronata

Spjut. Se hacen ocho nuevas combinaciones: T. caespitosa var. latifolia (Pilg.) Spjut, T. canadensis var. adpressa (Carrière) Spjut,

T. canadensis var. minor (Michx.) Spjut, T. globosa var. floridana (Nutt. ex Chapm.) Spjut, T. mairei (Lemée & H. Lév.) S. Y. Hu

ex T.S. Liu var. speciosa (Florin) Spjut, T. umbraculifera var. hicksii (Hort. ex Rehder) Spjut, T. umbraculifera var. microcarpa

(Trautv.) Spjut, y T. umbraculifera (Siebold ex Endl.) C. Lawson var. nana (Rehder) Spjut. Se discute la taxonomía y la nomenclatura

de todas las especies y variedades.

INTRODUCTION

The genus Taxus (Taxaceae) has long been indicated to have 7-12 species or subspecies (Cope 1998; Farjon

1998, 2001; Pilger 1903, 1916; Silba 1984, 1986; Spjut 1992); however, Spjut 2000c; www.worldbotanical.

com) has proposed recognition of 24 species and 55 varieties. The taxonomy of Taxus has been controversial

because the species do not appear reproductively isolated except by geography (Farjon 1998; Pilger 1903;

Silba 1984), although Collins et al. (2003) reported that Taxus hybrids may have impaired meiosis or less

functional pollen. The genus ranges from temperate North America into subtropical Central America, and

from temperate Eurasia to subtropical Southeast Asia (Cope 1998).

Molecular and morphological studies of Taxus have distinguished genotypes that differentiate (1) indi-

viduals within populations (Collins et al. 2003; El-Kassaby & Yanchuk 1995; Saikia et al. 2000; Spjut 2007),

(2) distinct populations within geographic regions (Doede et al. 1993; El-Kassaby & Yanchuk 1994; von

Hertel and Kohlstock 1996; Hilfiker et al. 2004; Spjut 2007), and (3) alleged geographically distinct species

J. Bot. Res. Inst. Texas 1(1): 203 — 289. 2007

204 Journal of the Botanical R h Institute of Texas 1(1)

(Collins et al. 2003; Doede et al. 1993; Florin 1948a, 1948b, 1948c; Hils 1993; J. Li et al. 2001; N. Li € Fu

1997; Krupkin unpublished, 1994; Orr 1937; Spjut 2007; Vance & Krupkin 1993). Molecular studies for

the most part have been geographically based, in which plants have been randomly selected; little attempt

has been made to correlate genetic differences or haplotypes with morphological characters (Corradini et

al. 2002), while new species have been recently described based on morphology and geographical data (N.

Li & Fu 1997).

The geographic species of Taxus that have received support from molecular and/or morphological stud-

ies include those in North America (T. brevifolia Nutt., T. canadensis Marshall, T. globosa Schltdl. var. globosa,

and var. floridana [Nutt. ex Chapm.] Spjut; Hils 1993; J. Li et al. 2001; Spjut 1992, 1993, 2007; Vance &

Krupkin 1993), the Euro-Mediterranean T. baccata L., the Sino-Japanese T. cuspidata Siebold & Zucc. (Collins

et al. 2003; J Li et al. 2001), and the tropical Southeast Asian T. sumatrana (Miq.) de Laubenfels (received

as T. chinensis (Pilg.) Rehder (Phyton Inc., Krupkin pers. comm. 1994). Jianhua Li et al. (2001) have further

shown that the North American species, T. brevifolia and T. globosa (from both Mexico and Florida), which

belong to Spjut's (1998b, 2000b) Wallichiana Subgroup, form a well-supported clade separated from a large

weakly supported clade represented by T. baccata, T. canadensis, T. chinensis, T. cuspidata and cultivars, and

that “low sequence divergence between T. floridana from Florida, and T. globosa from Mexico suggest very

recent separation between the lineages in these regions and is consistent with treating these populations as

belonging to the same species."

There is also molecular data to support distinction of less geographically separated species. Wang et

al. 2000) showed a sharp contrast between RAPD bands of T. chinensis and T. mairei (Lemée & H. Lév.) S Y.

Hu ex T.S. Liu among other species of taxads included in their study. An unpublished report on Himalayan

yews found the northwestern Himalayan yew (T. contorta Griff.) distinct from the East Himalayan T. wal-

lichiana Zucc. and the Euro-Mediterranean Baccata Alliance (Amin pers. comm.). A specimen received from

Phyton Inc., reportedly from southwestern China, was thought to be distinct from T. chinensis (Kadkade

pers. comm. 1997); indeed, it had been proposed as a new species from morphological study of herbarium

specimens (T. florinii Spjut in adnot., June 1996, A).

Despite the wealth of papers on the chemistry of Taxus in regard to developing anticancer diterpenoid

compounds (taxoids) for use in cancer chemotherapy (Appendino 1995; Kingston et al. 1990; Kingston

2005), not a single comprehensive study has emerged on the phylogeny of the genus. One might expect that

a genus with supposedly only eight species (Silba 1986) would be relatively simple to resolve taxonomically

by molecular data. Undoubtedly the traditional geographic species of Taxus have been investigated by mo-

lecular data, but lack of a definitive phylogenetic treatment would seem to reflect the need for morphological

studies to define the species as a guide for the molecular investigations.

Spjut (19982, 2006, 2007) suggested that much of the variation in leaf anatomical features of Taxus—in

the eastern Himalayas to southwestern China—could be explained by post-Pliocene hybridization between

formerly distinct Tertiary species. Taxus engelhardtii Kvacek, for example, a Tertiary species discovered from

leaves in Oligocene deposits in Europe, is much like the extant T. mairei (Lemée & H. Lév.) S. Y. Hu ex T.S.

Liu in subtropical laurophyll forests of southern China; the fossil species differs in having papillae on the

abaxial leaf midrib (Spjut 2007). Similarly, a Taxus sp. from a Middle Miocene deposit in western North

American (Kvacek & Rember 2000) has the leaf anatomical features of T. brevifolia (papillose abaxial margins

and midrib, large marginal cells, stomata in 4—7 rows), but differs slightly in the the leaves appearing flat-

tened and less mucronate; recently, it has been described as a new species (Kvacek & Rember submitted).

Two other Tertiary leaf fossils of Taxus from European deposits of late Miocene to Pliocene age are closely

similar to T. canadensis (Kvacek 1984, Spjut 2007), a species native to east-central North America but also

recognized to occur in the Euro-Mediterranean region (Spjut 2000b, 2007). Intermediates between extant

T. baccata and T. canadensis in the Euro-Mediterranean region, which have a partially papillose marginal

zone, seem best explained by hybridization (Spjut 2006).

Collins et al. (2003), in a molecular study of Taxus canadensis, T. cuspidata, and T. baccata, identified

Spjut, Taxonomy of Taxus 205

three different DNA chloroplast types, which support three stomata band types described by Spjut (2003,

www.worldbotanical.com) for the Baccata Group, and suggested that these differences indicate “a long period

of isolation.” Their study also included numerous cultivars or “hybrids” (T. x media”) between T. cuspidata

and T. baccata and between T. canadensis and T. cuspidata, none of which were found to have identical

genotypes. To explain the higher level of diversity in cultivars, they suggested that multiple hybrid events

have occurred over time, but it is also likely that other genotypes, which may belong to other species, have

contributed to the hybrid complex not present in their putatively wild samples. This reflects a conservative

view that only one species of Taxus exists within a geographic region; further, in the view here, distinct

morphological ecotypes of Taxus with parapatric distributions are considered worthy of taxonomic rank as

Species or varieties.

The IUCN (2004, 2006), in reference to a Conifer Specialist Group 1998 (Farjon & Page 1999), which

adopted the nomenclature in Farjon (1998), recently listed Taxus brevifolia as a threatened species, one that

is not rare but reportedly threatened due to the need for taxol derivatives from plant sources for the com-

mercial production of anticancer drugs. Other species of Taxus listed by the IUCN are T. globosa (including

var. floridana) and T. wallichiana. The latter name is meant to also include T. contorta, but Farjon's 2001)

treatment of T. contorta as a synonym of T. wallichiana is without taxonomic support (no types or specimens

were cited, no keys are provided, nor are there any references to such data; see also CITES (2001, 2004).

Awaiting a standard taxonomic foundation upon which species and varieties can be identified according

to the International Code of Botanical Nomenclature (ICBN, Greuter et al. 2000), all species and varieties

in the genus Taxus, whether described or undescribed, perhaps should be considered rare, threatened, or

endangered.

This paper presents a key to 24 species and 26 varieties of Taxus based on chemical (byproducts) and

morphological differences. Six new species and four new varieties are described, and eight new combina-

tions are made. Lectotypes or neotypes are designated or proposed for 23 of the species and the additional

varieties mentioned. Descriptions and specimen citations are provided primarily for new taxa in connection

with a paper on biogeographical data of Taxus (Spjut 2007). Also included in the present study are varieties

of T. brevifolia, T. caespitosa Nakai, T. canadensis, and T. umbraculifera (Siebold ex Endl.) C. Lawson, which

are not mentioned in Spjut (2007); therefore, specimen citations are also provided for these. The taxonomy

and nomenclature of each taxon are discussed.

MATERIALS AND METHODS

More than 1,000 specimens of Taxus have been studied, 845 of which are documented with details on leaf

anatomical data (Spjut 2007).

Early in the study, ca.100 representative specimens of Taxus throughout its range identified 11 species

and one variety by morphological features that best fit the traditional geographical concept (Spjut 1992,

1993; Spjut in Hils 1993). The characters were based on leaf anatomical features that included the number

of stomata rows, distribution of papillae on abaxial midrib, shape of epidermal cells, curvature of the abaxial

midrib, and the color of the stomata bands as compared to the non-stomata regions (Spjut 1992).

As the study progressed, many specimens from Eurasia could not be accommodated in the morphologi-

cal key according to the traditional geographic species concept; tly, this concept was abandoned.

Herbarium specimens were then strictly classified according to pattern recognition as evident from characters

of branching, bud-scale texture and size, change in color of branchlets from 1* to 2™ year, leaf arrangement,

leaf shape, leaf thickness, leaf color, leaf curvature lengthwise and across both surfaces, changes in leaf

curvature near margins and along midrib, shape of cones in bud and at maturity, distribution of papillae

along the abaxial leaf margin, and other characters of lesser importance (described in DELTA format, Spjut

USDA Memorandum 1995). Data on numbers of stomata rows and number of marginal cells are presented

elsewhere (Spjut 2007). Only color and phyllotaxy are further discussed (below). References for eco-geo-

graphical data on species are cited when relevant to data on herbarium specimens.

206 Journal of the Botanical R h Institute of Texas 1(1)

Results of the author's revised taxonomic concepts were presented at annual scientific meetings in

Baltimore (Spjut (1998a,b) and Portland (Spjut 2000a,b,c), which included papers on the evolution in the

Taxus leaf, phytogeography of the genus Taxus, a key to all species and varieties of Taxus (Spjut 20003),

and the occurrence of Taxus canadensis in both North America and the Euro-Mediterranean region (Spjut

2000b). Manuscripts were also submitted for peer review in 1999 and 2000; one of these was later placed

on the internet (Spjut 2003) from which data were extracted and incorporated into the present paper.

Taxonomic species concept

The species concept in Taxus in the sections that follow is based on pattern recognition employing the tra-

ditional method of defining species in keys; a taxonomic method in which my primary objective has been to

classify specimens in the fewest number of species and varieties that can be reasonably distinguished from

one another. Their character features, as presented in the following keys, can be seen in specimen photos

141 1

with data from labels at www.worldbotanical.com. My view of Taxus species is that they were most distinct

by the end of the Tertiary as a result of former geographical and ecological isolation, and that hybridization

and introgression since the Pliocene has blurred their distinction (Spjut 2007). Examples of species and

varieties that were allegedly more widespread and appear to be losing their identity through introgression

are Taxus OCR (T. ocreata Spjut ined.) (China), Taxus SCU (T. scutata Spjut ined.) (China), T. suffnessii Spjut

(Myanmar), T. wallichiana var. yunnanensis (W.C. Cheng & L.K. Fu) C. T. Kuan (NE India to SW China), and

T. contorta Griff. var. mucronata Spjut (Bhutan, Nepal).

The taxonomic value of characters was evaluated subjectively by their apparent correlation with other

characters. Species identification usually depended upon specimens having a combination of two or more

character attributes. As experience was acquired in identifying Taxus, many species could be recognized by

gross morphological features of branching, phyllotaxy, and color; however, detailed examination of a leaf

under a microscope for other character features was also necessary to confirm identification (e.g., see discus-

sion under T. kingstonii Spjut). Ideally, one might further employ chromatographic or molecular characters,

but it remains to be demonstrated whether such information can be efficiently extracted from herbarium

specimens. Needles from a number of herbarium specimens were subjected to DNA extraction, but there

was little extractable DNA (Da Cheng, pers. comm. 2007; Krupkin, pers. comm. 1994).

Variation in color as it may relate to chemotaxonomic characters

A key character for recognizing differences between species groups is the occurrence and distribution of

reddish colored cells in leaf tissues as seen in dried herbarium specimens. Differences in color are also

employed as taxonomic characters at the species level; for example, T. baccata has nearly concolorous leaf

surfaces in contrast to strongly discolorous leaf surfaces in T. recurvata Hort. ex C. Lawson, and T. hingstonii

is identified by its rusty orange color compared to a blood red color in T. mairei.

The reddish to orange cells are obviously the result of chemical byproducts. These have yet to be

identified, but they apparently are phenolics that oxidize slowly in collected and dried specimens to form

the reddish resinous substances observed in cell walls of leaf epidermal and mesophyll layers. The color

changes may occur over a period of many months or years except for species in the Sumatrana Group in

which the color change occurs usually within a week after a specimen is collected and dried. Taxus contains

cyanogenic glucosides (Khan & Parveen 1987) that break down and release benzaldehyde related componds

when plants are damaged (Seigler 1991; van Genderen et al. 1996). These include taxiphyllin, dhurrine,

triglochinine, and isotriglochinine (Khan & Parveen 1987).

Other potentially useful compounds in Taxus are biflavones, which are known to have chemotaxonomic

value in gymnosperms in addition to biological value as antifungal, anti-bacterial, and antiviral agents

(Krauze-Baranowska & Wiwart 2003). For example, in the Podocarpaceae, the presence or absence of various

flavonoid glycosides (Markham et al. 1985) has been found to correlate with recent morphological taxonomic

concepts of its genera and species (de Laubenfels 1969), and biflavones have been shown to be localized in

leaf epidermal cells of conifers with the aid of aluminum chloride-induced fluorescence (Gadek et al.1984).

Taxus biflavones include sciadopitysin, ginkgetin, kayaflavone, amentoflavone 7-O-methylamentoflavone

Spjut, Taxonomy of Taxus 207

in European and Himalayan species, and bilobetin and 4-O-methylamentoflavone in samples from Poland

(Krauze-Baranowska & Wiwart 2003). Appendino (1995) has noted uncharacterized pro-anthocyanidins,

“based on 3-flavanols of the cis and trans type,” are “probably responsible for the red color of a paste made

from bark of a Himalayan species." Also, apocarotenoids have been found in a Himalayan yew but not in

European yew (Appendino 1995).

Leaf parenchyma of Taxus also contains essential oils, but Taxus is deficient in monoterpenes that are

usually present in conifers (Appendino 1995; Jean et al. 1993). Taxus is best known for anticancer diterpe-

noid compounds (taxoids), particularly taxol (Wani et al. 1971), from which semi-synthetic derivates are

used for treating cancer (Kingston et al. 1990; Kingston 1996, 2005). Numerous novel taxoids have been

discovered in the genus (Appendino 1995); however, most reports are of little taxonomic value because

studies have focused on novel discoveries (Appendino 1995) and because the genus has been in critical

need of taxonomic study. Most taxoids of pharmacological interest are widely distributed and vary in yield

according to plant parts, location, season, drying conditions, and species (Croom 1995; Dempsey & Hook

2000; Griffith & Hook 1996; Hook et al. 1999; van Rozendaal et al. 2000). Nevertheless, Spjut et al. (1993)

reported chemotaxonomic relationships based on taxoid content between similar morphs of T. brevifolia var.

reptaneta when compared to var. brevifolia, and Chang (unpublished, pers. comm.) found chemotaxonomic

differences for taxoid hplc profiles between cultivars and between Taxus kingstonii and other Taxus species

in Taiwan. Although chemotaxonomic studies are lacking, Appendino (1995) has suggested that yews in

Europe are characterized by the presence of taxine B, those in the Pacific Northwest by the presence of

abeotaxane type alkaloids, and the Himalayan yews have 13,14-dihroxylated taxoids not found in other

regions.

Leaf phyllotaxy (frequency), arrangement (distribution), and orientation

These terms have similar but slightly different meanings in this paper.

Phyllotaxy.—refers to the frequency at which leaves develop along a twig as determined between two

leaves that occur in direct alignment. The frequency is generally expressed as a ratio of the number of leaf

cycles over the number of leaves in a cycle over the total number of leaves, the latter being a sum of the

preceding two numbers and representing what are known as Fibonacci numbers. Camefort (1956) recog-

nized four different patterns in Taxus baccata, a 2-3-5 that may spiral to the left or to the right, a 3-5-8, and

a 5-8-13. Because differences in phyllotaxy can relate to branch thickness (Camefort 1956), phyllotaxy is

generally described as dense, lax, or remote and by whether adjacent leaves along one side of the branch

overlap along their margins.

Leaf arrangement.—The phyllotaxy in Taxus may also be described as spiral, as opposed to opposite,

whorled, or alternate in other plant genera; however, a distinction is also made in regard to the directional

and distribution pattern in which leaves spread from branchlets. In the Dovaston yew (T. baccata var. dov-

astoniana Leighton), for example, leaves on the uppermost side of the branchlet point upwards and towards

the branch apex, while those along the sides and underneath spread outwards (horizontally). In the Maire's

yew (T. mairei), leaves spread horizontally along two sides of branchlet nearly in one plane and thus appear

distichous or “two-ranked” (but not truly two-ranked), in contrast to a radial arrangement of the Irish yew

(T. fastigiata Lindley, not a true whorl), in which the leaves are mostly erect except for curving downwards

along the blade. A further distinction is whether leaves along one side of a branchlet are mostly parallel to

one another (e.g., T. baccata) or crisscross (e.g., T. recurvata) as seen in pressed specimens, and whether they

appear more imbricate (e.g., T. caespitosa) or decussate (e.g., T. umbraculifera)

Leaf orientation.—This is in regard to the direction that leaves spread as a result of phototropic response.

Leaves generally twist and/or bend towards light so that the adaxial surface faces upward (Hill & Scriven

1998). Leaves in most species of Taxus tend to spread horizontally in shade and upwards in sunlight. Habit

and branching may also be correlated with leaf orientation. For example, the columnar growth form of the

Irish yew (T. fastigiata) with ascending to erect branches appears to be an adaptation to growth in open

habitats. Because the branchlets ascend upwards, the leaves would not be expected to spread much out-

208 Journal of the Botanical R h Institute of Texas 1(1)

wards; thus, their erect radial orientation is seen as a correlated feature. Taxus caespitosa, sometimes found

with prostrate branches in open habitats, exhibits a phyllotaxy that may be described as dense with a leaf

arrangement referred to as radial-imbricate, and because all the leaves on horizontal (caespitose) branchlets

point upwards in the same direction, the orientation is also referred to as erect-secund.

Species Groups, Subgroups, Alliances, and Complexes

No formal classification of sections or subsections within the genus is proposed at this time, but species of

Taxus are classified into three groups. Within the groups, two subgroups and two alliances are recognized,

based on leaf epidermal cell shape, development of epidermal papillae, and on color of stomata bands in

contrast to adjacent epidermal regions (Spjut 1998b). The Wallichiana Group includes the Wallichiana and

Chinensis Subgroups; the Baccata Group includes the Baccata and Cuspidata Alliances; the Sumatrana Group

is not subdivided. Alliances refer to taxa that share morphological features within geographical regions,

whereas taxonomic groups and subgroups are not geographically isolated. Within species alliances or groups,

species complexes are recognized to distinguish, for example, plants with radial distribution of leaves (T.

umbraculifera complex) vs. leaves appearing in a two-ranked arrangement (T. cuspidata complex).

Nomenclature, references, authorities

Species and varietal names are typified to validate the data in this study and in Spjut (2007). Descriptions of

species and full details on specimens cited are omitted for species already established; detailed descriptions

and specimen citations for all taxa can be found online at www.worldbotanical.com. Six new species are

described, Taxus biternata, T. florinii, T. kingstonii, T. obscura, T. phytonii, and T. suffnessii, because their leaf

anatomical data are relevant to taxonomic data in another study (Spjut 2007) that supports the taxonomy

in this paper. Authors for scientific names are indicated in full when the taxon name is first mentioned and

are subsequently abbreviated (fide Brummitt & Powell 1992) when there is no reference to a publication.

KEYS TO GROUPS, SUBGROUPS AND ALLIANCES OF TAXUS

1. Leaf epidermal cells tall rectangular or +angular-isodiametric in T-sect. and usually reddish in herbarium

specimens, papillose on the abaxial midrib; North America (SE Alaska and W Florida) to Central America

(Honduras, El Salvador), Himalayas, SW China Il. Wallichiana Group (IA Wallichiana Subgroup)

Leaf epidermal cells «elliptical or wide rectangular in T-sect., 1.5-3.5 times as wide as tall, or if isodiametric

not reddish in dried herbarium specimens, variable in development of midrib papillae.

2. Stomata bands in dried leaves not sharply differentiated from adjacent marginal and midrib regions, the

abaxial surface green to yellowish green, slightly darker green on midrib and along margins, or uniformly

reddish green.

. Leaves with reddish cells primarily epidermal; papillae + equally developed across stomata bands and

midrib, often more prominent along cell walls; stomata sometimes on midrib, (11—)13-19(-21) rows

per band; E pleco to China cla: va lichiana Group Chinen Subgroup)

. Leaves with reddish cells primarily i | n midrib th

bands, or more medial than marginal (T. baccata Alliance, Euro- Mediterranean, W Himalayas), or papillae

reduced along cell walls—appearing concrescent (7. cuspidata Alliance, NE temp. Asia); stomata less

than 13 rows per band in NE temp. Amer; Euro-Mediterranean, up to17 rows per band in temp. E Asia

IIl. Baccata Group

2. Stomata bands in dried leaves distinct from glossy marginal and midrib epidermal cells, the midrib entirely

smooth to papillose on outer half, or smooth from mid region to base, often discolored—blood red—in

contrast to the yellowish green to yellowish orange stomata bands.

. Leaf epidermis with cell walls meeting at sharp anges mostly smooth on midrib and marginal region

(8-36 cells wide); stomata rarely in transverse rows; leaf papillae more medial than marginal; E odds

to Indonesia t Group

. Leaf epidermis with cells meeting at rounded angles, mostly papillose to 2-6(-12) cells E margin,

sometimes less on midrib; stomata often in transverse rows; leaf papillae marginal; E Himalayas to

Indonesia IB. Chinensis Subgroup

Spjut, Taxonomy of Taxus 209

KEYS TO THE SPECIES AND VARIETIES OF TAXUS

I. WALLICHIANA GROUP: SUBGROUP IA WALLICHIANA

. Stomata 12 or more rows per band; E Himalayas, SW Chin

2. Persistent bud-scales relatively large, 2-3 mm long, Ad plane with distinct midnerve (Fig. 1); Myanmar

4. T. suffnessii

2. Persistent bud-scales nerveless, or nearly so, concave, less than 2 mm long and 1 mm wide (Fig. 2), if scales

not evident or leaves obtuse see Chinensis Subgroup; E Himalayas, Nepal to SW China (var. yunnanensis

distinguished by abaxial leaf marginal zone having long narrow cells bordered by shorter and much wider

papillose cells with medial papillae, or by a lanceolate leaf shape in SW China) 5. T. wallichiana

1. Stomata less than 12 rows per band.

SS:

3. Abaxial leaf epidermal cells gradually narrower and shorter from stomata band to margin, mostly isodia-

metric in T-sect., generally 3-10x l/w, fusiform and sinuous (wavy along cell walls) on abaxial surface, or

rarely rectangular; Taxus globosa.

4. Papillae prominent on most of the abaxial leaf surface, the marginal cells mostly sinuous 7 3, type);

El cece! to NE ses 3a. T. globosa var. globosa

4. P. idril | inal t t tal |, the marginal cells + rectangular

(Fig. 4, type); Mexico (Nuevo Leon/Tamaulipas, Veracruz), U.S. (Florida) .. Sb. T. globosa var. floridana

3. Abaxial leaf epidermal cells abruptly shorter and wider in marginal region, often taller than wide in T-sect.

5. Stomata 7-10(-12 rows); China (Yunnan, Sichuan) 2. T. florinii

5. Stomata 4-7(-9) rows; NW North America 1. T. brevifolia

6. Cones elongate, often with a narrow stipelike base 1b. T. brevifolia var. polychaeta

6. Cones appearing sessile.

7. Trees, reproducing by adventitious shoots 1a. T. brevifolia var. brevifolia

7. Layering shrubs, often forming thickets 1c. T. brevifolia var. reptaneta

I. SUBGROUP IB CHINENSIS

. Dried leaf surfaces +concolorous, or yellowish green on abaxial surface.

2. Bud-scales conspicuous at base of branchlets; Yunnan, Sichuan 11. Taxus sp. SCU

2. Bud-scales minute, vestigial or absent at base of 1*yr branchlets.

3. Leaves obtuse, usually oblong (less than 10x l/w); Vietnam, China 6. T. chinensis

3. Leaves acuminate (10-15x l/w); Vietnam, Philippines, China (Taiwan), Indonesia. 9. Taxus sp. REH

. Dried leaf surfaces discolorous, or yellowish orange on abaxial surface.

4. Leaves obtuse, dull rugose on adaxial surface; Yunnan, Sichuan 10. Taxus sp. OCR

4. Leaves acute to acuminate, smooth or glossy on adaxial surface when dried.

5. Leaves spreading parallel at nearly right angles; Nepal, NE India, Malaya, China (Taiwan, Yunnan),

Philippines 8. T. phytonii

5. Leaves spreading at oblique angles, crisscrossing in pressed specimens; Myanmar, China (Fujian, Taiwan),

Philippines, Indonesia (Sulawesi, Sumatera) 7. T. obscura

Il. SUMATRANA GROUP

. Dried leaves rusty orange, at least near apex, often rust colored on abaxial surface in contrast to a darker

green or bronze green colored adaxial surface, generally twisted obliquely to the stem axis, often recurved

along blade to sharply pointed apex, with midrib rounded on both surfaces; generally thick and rigid; mostly

1700-3000 m, NE India to China (including Taiwan) 13. T. kingstonii

. Dried leaves green to reddish, or with a blood reddish discoloration along abaxial margins and on midrib,

generally twisted nearly perpendicular to stem axis, the adjacent leaf edges often closely parallel, obtuse to

acuminate, the abaxial midrib usually elevated and truncate, or flush, usually with a channel; generally thin

and flaccid; mostly below 1200 m in China.

2. Leaves oblong to linear, obtuse to acute, or elliptical and acuminate; leaf epidermal cells larger (mam-

millose) on abaxial than adaxial midrib in T-sect., appearing short trapezoidal from surface view; S China

14. T. mairei

3. Branchlets limp, often much isodichotomous; leaves closely parallel along one side of branchlet; seed

purplish 14a. T. mairei var. mairei

3. Branchlets rigid, occasionally isodichotomous; leaves unevenly spaced along branchlets; seed tan

4b. T. mairei var. speciosa

2. Leaveslong linearto lanceolate, acuminate; leaf epidermal cells in T-sect. not larger on abaxial than adaxial

midrib, nearly rectangular in surface view; E Himalayas to Indonesia, Philippines.

210 Journal of the Botanical R h Institute of Texas 1(1)

Fic. 1. Taxus suffnessii. Close-up of persistent scales showing prominent mid ; Kingdon Ward 20902, Myanmar (isotype, BM).

4. Dried leaves plane to convex on adaxial surface 12. T. celebica

4. Dried leaves puckered, especially upper third of leaf 15. T. sumatrana

II. BACCATA GROUP

1. Abaxial surface of leaves mostly papillose between margin and stomata band—to within 8 rows of cells from

margin (e.g., Fig. 5), papillae nearly medial, the cell walls thin or smooth; leaves often curved downwards

along the blade (except T. contorta var. mucronata); Euro- Mediterranean, W & C Himalayas _ 16. T. baccata

Alliance

2. Leaf mesophyll with idioblasts (parenchyma cells with reddish walls), loosely connected; NW to central

Himalayas (var. mucronata distinguished by shorter more reflexed leaves with a wider bare marginal zone of

cells) 17. T. contorta

2. Leaf parenchyma cells often dark in color, not reddish, adhesive.

2a. Plants columnar; leaves radial, recurved 18. T. fastigiata

2a. Plants variable in habit; leaves overlapping, not distinctly radial.

2b. Leaf surfaces discolorous, blades recurved, convex, revolute 19. T. recurvata

2b. Leaves +concolorous, straight or falcate, plane or revolute 16. T. baccata

2C. Leaves spreading outwards along two sides of branchlets in one plane, appearing distichous.

2d. Leaves not overlapping, branching isodichotomous 16a. T. baccata var. baccata

2d. Leaves overlapping, branching irregular 16c. T. baccata var. elegantissima

2C. Leaves spreading outwards along two sides of branchlets in one plane, appearing distichous.

2C. Leaves erect or radial on lower parts of branchlets.

2e. Branchlet y range in part; brancl often recurved; erect near

ends of branchlets 16d. T. baccata var. glauca

2e Branchlets uniformly green, yellowish green to dark green; branchlets not recurved, spread-

ing or drooping; leaves variable.

2f Branching mostly pinnate 16f. T. baccata var. variegata

2f. Branchlets crowded terminally, appearing to arise digitately or isodichotomously.

2g. Leaves yellowish green, falling off by the third year; branching isodichotomous

16e. T. baccata var. pyramidalis

| pers

Spjut, Taxonomy of Taxus 211

icc

M een

Fic. 2. Taxus wallichiana var. wallichiana. Close-up of persistent

(P)

2g. Leaves dark green, persistent; branching subdigitate, pinnate, or subfastigiate

16b. T. baccata var. dovastoniana

1. Leaves mostly smooth between stomata band and margin, usually lacking papillae across 8-24 rows of cells,

the papillae often concrescent on Belloc a a near cell walls; leaf blades often curved upwards, or

sharply bent or curved d te E Asia and NE North America, Euro-Mediterranean

23. T. cuspidata Alliance

3. Leaves spreading more by narrow petioles, the petioles more curved than sharply bent, or petiole curving

nearer junction with branchlet than with blade; stomata (4-)5-9(-11) rows per band; leaf papillae often

obscure; plants usually low, creeping shrubs 22. T. canadensis complex

4. Leaves acute to acuminate, gradually tapering to an acute margin as seen in T-sect., usually revolute

along margins, the abaxial surface with keeled midrib and mostly rectangular epidermal cells; common

in NEN America, occasional in Euro-Mediterranean: Morocco, Portugal (Madeira), Spain, dio Estonia

22a. T. var. canadensis

4. Leaves obtuse to acute, rounded along margins in T-sect., not revolute, with flush midrib and with

trapezoidal or rectangular epidermal cells.

Sub ! l)

5. Leaves crowded, erect, recurved; NE Amer., Madeira 22c. T. canadensis var. minor

5. Leaves lax, mostly straight, spreading nearly at right angles); occasional, England, Madeley France,

Sweden, Norway, Estonia, Slovenia, U.S. (lowa). 22b. T. var. adpressa

3. Petioles more bent than curved, bending nearer junction with blade and also clasping the branchlet, the

blades often (ad)pressed to branchlet; stomata (7-)9-14(-17) rows per band; papillae always prominent

in stomata bands; plants variable in habit.

6. Leaves mostly two ranked to apex of branchlets; seed angled T. cuspidata complex

7. Branchlets short and much divided; leaves in a flat (horizontal) spray (Fig. 6), more strongly revolute

in upper third when dried; common understory tree, NE temp. Asia 20. T. biternata

7. Branchlets mostly long pinnate; leaves erect (Fig. 7), uniformly revolute along margins when ee

shrub or tree—appearing adapted to exposed habitats, Japan, Korea 23. T. cuspidata

6. Leaves in +two or more decussate ranks or radial; seed rounded 24. T. mE uh complex

8. Leaves + erect and imbricate 21. T. caespitosa

£4h,D . ID L

212 Journal of t titute of Texas 1(1)

OMA

jn |

E5045 LV OPE ISP O MA

390 502 Oh iv

A =

ca fh sel

Fic. 3. Taxus globosa var. globsoa. Illustration of abaxial surface of leaf as seen in the medial region from margin to midrib; showing stomata band

with 9 f stomata and si id Ic I f inal position of papillae on epidermal cells, drawn from

NE : " Intad +n th

, a character Teature related to

lectotype (K).

eo En

FSE

Y gal A i 7

| m i ay l

UO V fT

: A AL qu

E 3 OMS v

SUAM y j N

— . —

DARIA

Fic. 4 T, LAL floridana Wi tyatt £-^L $ | £. £1 £ ctl Pal H £, LT * dtl L : 4 Ļ J PPT)

J 1

A £, CR I tarini I)

Lot " J I ow I IT] eal. | : P" ATI H e 4 alohosa

J J 7

Fic. 5. Abaxial lef i ith 5 f th cells. followed hv 2 rows of

od L4

obscurely papillose cells and 3 rows of prominently papillose cells. Image

recurvata, Lewis 721 (BM).

9a. Branches ascending to erect with many short, crowded branchlets; leaves mostly radial (Fig. 8);

Japan 21a. T. caespitosa var. caespitosa

9a. Plants with distinct trailing or wide spreading branches; branchlets with mixed two-ranked and

erect secund leaves.

9b. Branches ascending to wide spreading; leaves oblong or linear, 2-4 mm wide; Korea, Japan,

SE Russia, NE China 21c. T. caespitosa var. latifolia

9b. Branches prostrate with erect branchlets; leaves linear, relatively narrow, ca. 2 mm wide; Korea,

21b. T. caespitosa var. angustifolia

8. Leaves + reflexed and decussate (Fig. 9) 24. T. umbraculifera

10a. Flat-topped shrub, layering 24c. T. umbraculifera var. microcarpa

10a. Hemispherical to columnar shrub or tree.

Spjut, Taxonomy of Taxus 213

. £ . *^ T. £ f + All;

6(top) Taxus biternata, cultivated, F tryR h Institute, South Korea,

Fics I branching ped tl pidat [ y

photo and specimen by Kang Hyeon Ka s.n. 7 (bottom). Taxus cuspidata, cultivated, Secrest Arboretum, Wooster, Ohio, photo by Richard Spjut.

£+sha D o ID L

214 Journal of t titute of Texas 1(1)

10b. Low rounded shrub 24d. T. umbraculifera var. nana

10b. Tall shrubs or trees.

10c. Columnar with ascending to erect branchlets 24b. T. umbraculifera var. hicksii

10c. Pyramidal, diffusely branched 24a. T. umbraculifera var. umbraculifera

TAXONOMY AND NOMENCLATURE

I. WALLICHIANA GROUP

The Wallichiana (Species) Group is recognized by the leaves having reddish epidermal cells and scarcely dif-

ferentiated stomata bands as seen in herbarium specimens, in contrast to idioblasts in the spongy mesophyll

in the Baccata Alliance, which includes the northwestern Himalayan T. contorta.

IA. Wallichiana Subgroup.—This subgroup is differentiated from the Chinensis Subgroup by leaves having

taller than wide or isodiametric angular epidermal cells in transverse section, and by the obscurely dif-

ferentiated stomata bands; the papillose epidermal cells often extend to four cells or less from the margin.

Included are three species distributed from the eastern Himalayas to Yunnan, Sichuan, and western Hubei

(2000-3700 m elevation) and two species in North America (from near sea level to 2670 m).

1. Taxus brevifolia Nutt., N. Amer. Sylva 3:86. 1849, t. 108 captioned “Taxus occidentalis," without refer-

ence to specimens. (Fig. 10). Taxus baccata L. var. brevifolia (Nutt.) Koehne, Deutsche Dendrol. 6. 1893. Taxus baccata L.

subsp. brevifolia (Nutt.) Pilger in Engler, Planzenreich 4(5):113. 1903. Tyre: U.S.A. [Orecon]: “Columbia woods,” “Nuttall Herb."

without collector, date of collection and collectors number (lectotype designated here: from uncited specimens—original material:

Nuttall Herbarium—BM! # 38752, annotated “T. occidentalis," —top left specimen of three on one sheet, with mature male cones;

ISOLECTOTYPES: K!, PH).

Taxus boursieri Carrière, Rev. Hort. sér 4, 3:228. 1854. Type: U.S.A. CALIFORNIA: Siskiyou Co.: Shasta Springs, Aug. 1894, Jepson s.n.

(NEoTYPE designated here: US!). Carriére cited no specimens and original material is unknown, but he noted that the species was

from California: “Forests near stream with Pseudotsuga, Abies grandis, Pinus lambertiana; arborescent shrub with slender branches,

leaves 1.5-1.8 cm long, glaucous below.”

Taxus lindleyana M.A. Murray, Edinburgh New Philos. J. 1:294. 1855. Also, Rev. Hort. sér 4, 4:379. 1855 (by J. Decainse). Type: U.S.A.

CALIFORNIA: N and E of San Francisco between 40° and 41? latitude [Klamath Region], “along sides of a glen.under the shade of larger

trees” ( designated here: selected from original material of two uncited specimens at E, the one with the following handwrit-

ten notations (1) “Taxus lindleyana" (in bold black ink, handwriting probably Murray) (2) “Murray,” “Ed. N. Philos. Journ.1854,”

"Legn. California" (in red faded ink, handwriting undetermined, perhaps R. Jameson) and (3) *probably only a form of T. baccata,

perhaps the T. cuspidatus [spelling?] of Sieb. & Zucc." (in thin black ink, handwriting unknown—appearing different from the other

two), and with the accession number E 00030316!). Described by Murray in his publications as a large tree, 30—40 ft high, with

a circumference 50-70 inches in diam, and with very long pendant branches. The other specimen is indicated to have come from

Vancouver, possibly collected by Murray before 1855. Judging from other plants described by Murray from his California trip in

1854, the lectotype may have been collected near Dunsmuir in Oct. 1854.

Common name.—Pacific yew.

Distribution and ecology.—SE coastal Alaska to central California—the Klamath Mts., Coast Ranges

and W Sierra Nevada to Sequoia Natl. Park—and from the S Canadian Rockies in British Columbia and

W Alberta south to W Montana. Often in shade of old-growth forests on N slopes, but also valley forests

and seral communities along streams and forest margins; from sea level to 2650 m, in mixed evergreen and

montane coniferous forests, generally Douglas fir (Eliot 1938) and lowland fir (Bolsinger & Jaramillo 1990),

western hemlock along W Cascades (Franklin & Dyrness 1969), or sitka spruce and western red cedar in

the northern range (Taylor 1932; Taylor & Taylor 1980).

This species was first discovered by David Douglas in 1825 (Sargent 1896), who after arriving in Astoria

OR that year on April 11 (Oldham 2005) had spent much of that year collecting in the Pacific Northwest.

His base of operations was the Hudson Bay Company located at Fort Vancouver just north of Portland on

the Washington state side of the Columbia River, arriving there on April 20 (Oldham 2005). However, there

apparently are no specimens of T. brevifolia by Douglas. Nuttall had also collected along the Columbia River

during 1834-1835 (Graustein 1950/51); one of the labels on the type sheet refers to T. floridana, a species

that was not discovered until 1833, a name that Nuttall had proposed sometime before 1860, the year it was

legitimately described. Hitchcock et al. (1969) indicated reference to a type “Nuttall, ‘In the dense maritime

Spjut, Taxonomy of Taxus 215

Fics. 8—9. Comparison of leaf arrangements in Taxus species of the Cuspidata Alliance. 8 (top). T pit ith imbricat ling | , culti-

vated, Secrest Arboretum, Wooster, Ohio. 9 (bottom) Taxus umbraculifera, with spreading to reflexed hat d tel

Arboretum.

, cultivated, Secrest

216 Journal of the Botanical R h Institute of Texas 1(1)

TERR nr.

Ļ : I +*-+DAA M £A Ll^sL

A B (top right), close-up of lectotype (BM), upper left

Ist yr branchlets. C (lower right). Close-up of the abaxial surface of leaves and

+L Ļ L E L 3 1 E: 10A

Fics. 10A-C. Type specimens of Taxus brevifolia. A (left). H

specimen,

Ļ £L hi H L I

persistent bud-scales at tl

forest of the Oregon" while Pilger (1903) noted that yew had been observed by Douglas in the Oregon forests

along the Columbia River. A review of the geographical distribution of T. brevifolia by Spjut (1977) reported

the occurrence of Pacific yew near Portland and other areas in the nearby Mt. Hood National Forest where

trees were known to reach a height of 20-25 m. The presence of mature male cones on the type indicates

it would likely have been collected during the spring. Thus, Oregon is likely the state locality for the type

where possibly collected by Nuttall himself during the spring of 1835; the handwriting on the type label

compares favorably with that on other Nuttall types (NY, virtual herbarium of types, particulary for speci-

mens from the “Columbia woods").

Taxus brevifolia is distinguished from the closely related T. globosa by the leaves having enlarged epidermal

cells along the abaxial surface between the stomata bands and margins, narrower stomata bands with fewer

stomata rows, and leaf blades tapering to an obtuse but mucronate apex. Leaf stomata range from 4—7(-9)

rows per band with the higher counts (7-9 rows) more frequent in specimens from the Sierra Nevada and

lower counts (4-6 rows) more often found in the Klamath Region, Coast Ranges, Cascades, and Rocky

Mountains (Spjut 2007). The relatively wide abaxial leaf margins lack papillae on 2-11 cells across.

The leaf epidermal cells in transverse leaf sections often appear tall rectangular as in the East Himala-

yan T. wallichiana, T. suffnessii, and the Southwest China T. florinii. The similar number of stomata rows per

band between T. florinii and T. brevifolia may be the result of convergent evolution (homoplasy) within the

Wallichiana Subgroup.

Spjut, Taxonomy of Taxus 217

The type has leaves spreading nearly at right angles and appearing dark metallic green on the adaxial

surface. Other specimens differ by a reddish orange color (e.g., lectotype for T. lindleyana), or by a glaucous

abaxial surface (e.g., neotype for T. boursieri). These differences may justify reevaluations of taxonomic status

in further studies.

Taxus brevifolia has been a major source of taxol (paclitaxel) employed in cancer chemotherapy, first

isolated from stem-bark (Wani et al. 1971). Historical accounts on the procurement of Taxus species for the

NCI antitumor screening are given by Croom (1995) and by Goodman and Walsh (2001); the latter draws

extensively on information from memoranda, letters, and interviews concerning availability of T. brevifolia

for supply of taxol in the short and long term.

Currently, three varieties are recognized by differences in cones, habit, and vegetative reproduction

The typical variety, a tree usually scattered in the understory of old growth forests, produces cones similar

to most other yew species. Variety polychaeta is distinguished by longer cones with persistent scales, whereas

var. reptaneta differs by the layering habit in which the plants often form thickets, in contrast to the typical

variety reproducing vegetatively by adventitious shoots from trunks or roots.

la. Taxus brevifolia var. brevifolia

Common name.—Pacific yew.

Tree, reproducing by adventitious shoots from trunks or roots, 6-13(-25) m tall, bole to 60(-130) cm

diam.; branches horizontal to upwardly ascending, sometimes drooping near ends, dividing more unequally

than equally, yellowish orange to reddish orange, or maroon; bud scales persistent on 2™-3" yr branchlets,

conspicuous, 2-3 seriate, paleaceous, chartaceous, closely to loosely adnate, plane to slightly concave, in-

durate, brownish, 1-1.522) mm long. Cone scales forming a basal hemispherical cup; male cones ellipsoidal

in bud, 4 mm long, 2 mm wide; sporangia exserted. Seed on 1“ or 2™ yr or older branchlets, longer than the

pedunculate axis, rounded, slightly 2-4 angular, ellipsoid to ovoid, tapering to apex from mid region, or

rather abruptly near apex, 5-8 mm long, to 4 mm diam.,; aril red, reddish orange, yellowish orange, or rarely

yellow, Aug—Sep.

Ib. Taxus brevifolia Nutt. var. polychaeta Spjut, var. nov. (Fig. 11). Wer: U.S.A. Wasuincron. Thurston Co.: Mud Bay,

near Tacoma, 3 Sep 1938, EG. Meyer 1589 (HoLotyee: K!).

Ab var. brevifolia, strobili elongati, lumbriciformis, squamae persistentibus.

Similar to T. brevifolia var. brevifolia in habit, differing by the longer seed cones; the seed appearing stipitate,

seed shorter than the long crooked cone axis (stalk) in the type, cone scales persistent; male cones elongate

with scales persistent to near apex.

Common name.—Worm-cone yew.

Distribution and ecology.—California (Mendocino Co., Sonoma Co.), Washington (Thurston Co. near

Mud Bay and Tacoma), Idaho (near Coeur d'Alene); observed to be rare in a redwood-grand fir-nutmeg forest

in Sonoma Co., California.

Additional specimens. CALIFORNIA. (Marin, Sonoma, or Mendocino Co.): 1854, Bigelow s.n. (US); Salmon Creek (Sonoma or Men-

docino Co.?), McMurphy 315 (US). Sonoma Co.: near Mendocino Co. line, 7 km E of Stewarts Point, Rich Spjut & Rick Spjut 16021 (BM,

BRIT, E. GH, K, US, wba). IDAHO. Without collection data, US Forest Service, received from Marion Blatch, Coeur d'Alene Nursery,

26 June 1992 (wba).

Variety polychaeta is distinct for its elongate, worm-like cones. The California specimens have shorter cones

than those specimens from further north—coastal Washington (type) and from near the Idaho/Washington

state line. In var. reptaneta, female cones may appear similarly elongate on older branchlets, while both male

and female cones on younger branchlets appear typical of the species. This is in further contrast to the male

specimen of var. polychaeta collected by John Milton Bigelow from California.

A specimen of var. polychaeta that was collected by James Ira McMurphy (1871-1943), who had lived

in Mendocino County, mentions only Salmon Creek; it may have been from the town of Salmon Creek just

£s+haD o ID L

218 Journal of t titute of Texas 1(1)

Fic. 11. Close-up of branchlet of Taxus brevifolia var. polychaeta showing three wormlike cones, one with seed; near Tacoma WA, Fred.Meyer 1589

(holotype, K).

north of Bodega Bay in Sonoma Co., or from west of the town along Salmon Creek itself, or possibly from

Big Salmon Creek just south of Albion along the Mendocino coast, or another more inland site in Mendocino

County where there is a Salmon Creek ca. 12 km E of Willits in the Mendocino National Forest. A recent

plant inventory of Big Salmon Creek does not include yew (Chanslor Wetlands Wildlife Project, website

accessed 2006), although this area had been logged.

In any case, the coastal occurrence of yew in California is rare. One record for Sonoma County was

by Milo Baker from near Annapolis where populations have since been reported from nearby Fuller Creek

and its north-facing drainages; however, in a brief survey of this area we were able to find only the one tree;

which belonged to var. polychaeta; it was growing next to Torreya californica Torr. in a redwood forest (Rich

Spjut & Rick Spjut 16021). John Bigelow was also known to have collected in Marin, Sonoma, and Mendocino

counties (CNPS, Marin Chapter, website); thus, the McMurphy specimen of var. polychaeta was probably

collected just north of Bodega Bay in view of our find (Spjut & Spjut 16021), which also seems significant

in regard tol8 endemic taxa that are recognized for Sonoma County (Best et al. 1996). Another report for

southern Mendocino County indicated that yew is rare in a ravine on Sea Ranch; however, this area was

reportedly logged in early 1900s and early 1990s; thus, yew may once have been more common there. It is

of interest to note the plant associations of two rare disjunct taxa of closely related yew in the United States

that include both US species of Torreya, one in California (Torreya californica) that is first reported here

with Taxus in a redwood forest region, and the other in Florida (Torreya taxifolia) in a white cedar forest as

described in more detail under T. globosa var. floridana.

A duplicate specimen of F.G. Meyer 1589 at the Smithsonian Institution (US) is not considered an isotype

because the seed cone in this specimen is typical for T. brevifolia. This may lead one to question whether

cone development in var. polychaeta is some sort of aberration within the ‘normal’ population; however, both

male and female cones of T. brevifolia var. polychaeta show the morphological feature of elongated cones.

Spjut, Taxonomy of Taxus 219

Also, most collections of var. polychaeta appear coastal. Molecular differences have been reported between

coastal and inland plants of T. brevifolia (El-Kassaby et al. 1994, 1995), which included a distinct genotype

on Vancouver Island in contrast to other yew populations studied in British Columbia.

lc. Taxus brevifolia Nutt. var. reptaneta Spjut, var. nov. (Figs. 12-13). Te: U.S.A. Catirornia. Siskiyou Co.: near

corner of Humboldt and Trinity Cos., Salmon Mts., 1 mi E to SE of Salmon Mt., near the NW corner of the Trinity Alps Wilderness

boundary, Klamath Natl. For., 19 mi S of the town of the Forks of the Sal n McNeal Creek Rd, West Fork of I hing Creek,

ca. 1250 m, along N-facing slopes of ravine in mixed evergreen forest of Douglas fir, white fir, ponderosa pine, tan oak, snowbush,

dogwood, hazelnut and big-leaf maple, shrub with long scandent stems ascending to 5 m or more, stems layering, forming thickets,

both male and female plants present, occasional male cones with pollen, female cones with seed but mostly without fleshy aril, one

with pale yellowish aril, 11 Sep 1990, R. Spjut & T. Spjut 11835, with seed (HOLOTYPE: US!; isotypes: BM!, BRIT!, E!, GH!, K!, waA!).

Differt a var. brevifolia frutex caulibus ascendentibus, ramis reptanibus, 0.3—5 m altis, propaginis, faciens dumetia.

Rhizomatous (layering) shrub with decumbent to ascending or contorted trunks, to 5 m high, typically

forming dense impenetrable thickets on open steep, narrow ravines, or more distantly spaced, either as low

creeping shrubs or arborescent in understory on slopes or in valleys with coniferous forests. Leaves similar

to var. brevifolia, often darker and duller on upper surface, slightly revolute along margins, more densely

papillose on abaxial midrib with papillae in 3-4 rows on each cell; midrib appearing more elevated, often

ca. 18 cells wide; stomata often in 4-6 regular rows per band. Male cones abundantly produced, ovoid in

bud, 3 mm long and 2 mm wide, scales ca. 5 in series; sporophylls ca. 8, united into a column ca. 2 mm

long, umbrelliform above, each with 5 microsporangia (0.9 mm diam.). Female bud cones ca. 1 mm long,

scales overlapping, ca. 5 seriate; seed maturing on 1* or 2-yr growth; rounded, ellipsoid to ovoid, 6-8 mm

long, 4 mm diam., often without fully developed aril. Aril red, reddish orange, yellowish orange, or rarely

yellow, maturing Aug-Sep.

Common name.—Thicket yew.

Distribution and ecology.—Rocky Mountains in British Columbia, Idaho and Montana; E Cascades in

Oregon and Washington; Klamath Mountains in Oregon and California; scattered or forming dense thickets

on steep sunny slopes of avalanche shoots, along streams, or in dense shade of valley forests, 1000—2000

m; in the Siskiyou Mountains occurring regularly between 3500 and 4000 ft on N to E slopes; probably in

the Coast Ranges, Oregon to British Columbia (Arno & Hammerly 1977), reported also from the Cascade

Ranges in California, near Mt. Shasta (Bolsinger, pers. comm. 2007).

Additional Specimens: CALIFORNIA. Siskiyou Co.: Klamath Natl. For., Marble Mountain Wilderness, Lake-of-the-Island, 1722-1820

m, forming thickets, mostly male plants on open talus and steep rocky slopes, or one female thicket observed on level soil in forest

understory along lake shore, in association with red fir, mountain hemlock, western white pine, incense cedar and other conifers, Spjut

16013-16015 (BRIT, GH, US, wba). MONTANA. Lincoln Co.: near northern panhandle of Idaho, Kootenai National Forest, Libby Mt.,

Snowshoe Mine Rd, T28N R3IW Sec. 5, 48?12' N, 115°38.34 W, 3287 ft, Spjut & Deevy 12303 (wha); Flathead Co.: Flathead Natl. For.,

4 mi N of Columbia Falls, Spur For. Rd 316G (Canyon Creek Rd), % mi W of jct. 316 and 316 G, T31N R20W Sec. 9, 3800 ft, 48?28'

N, 114? 10’ W, 3491 ft, with subalpine fir, grand fir, white pine, red cedar, hemlock, aspen, larch, plants predominantly male, Spjut

& Donner 12306 (US, wba). OREGON. Clackamas Co.: Mt. Hood Natl. For., ca.4 mi. N of Timothy Lake, FS Road 58 to Little Crater

Lake, ca. 0.5 mi from jct. with RS Road 42 (Skyline Drive), ravine or basin forest, 121%43"W, 54°08.59’N, 3400 ft, Spjut 12301 (US, wha).

WASHINGTON. Chelan Co.: N Cascades: Wenatchee Natl. For., Icicle Creek drainage, 18 mi W of Leavenworth, Black Pine Horse Camp,

scattered, somewhat scandent, at base of conifer trees in old growth forest of Douglas fir, grand fir, western white pine, and Englemann

spruce, and also forming thickets locally on flats or gentle slopes, 47°36’ N, 120?56' W, 3050 ft, Spjut 12302, 12101 (US, wba).

Taxus brevifolia var. reptaneta is recognized by its layering habit; however, as with T. canadensis, the layering

branch may die and rot away, leaving the individuals solitary (Bannan 1942). Plants vary in size, density

and branching, but trunks are similar in their ascending form (Fig. 13). In the typical habitat—along steep

open ravines—distinct trunks develop to 50 cm or more in diameter and grow within 1 m of each other,

each bearing many divaricate branches that are often longer than the main trunk. The collective growth

forms impenetrable thickets, hence the epithet reptaneta. In more shaded environments—as in the eastern

Cascades of Washington— plants are less crowded and sometimes exhibit a peculiar growth in that indi-

viduals sprawl around the base of trees such as Douglas fir, occasionally twisting and wrapping around its

trunk. In the Rocky Mountains other shrub forms grow less than 50 cm high, creeping along the ground

PiL D j ID L

220 Journal of t titute of Texas 1(1)

; gins, y aril; Sal Mt., R. Spjut & T. Spjut 11835 (isotype, wba).

B (bottom). Close-up of I hlet with mat | ; N-facing sl I Lake-of-the-Island, Marble Mt. Wilderness, Spjut 16073 (wba).

Fic. 12A—B. Taxus brevifoli reptaneta,

L £L L £.

lat, L sT

Klamath Mts., CA A (top) Herbari j i I i ictent «cal

£1 eL J

Spjut, Taxonomy of Taxus 221

Fic. 13A—C. Habit of Taxus brevifolia reptaneta. A (top). East | f the C les, E of Portland, OR ing fai ly level g lin forested

valley, trunks sharply ascending near base. B-C (bottom) Rocky Mts, b Montana; Libby Mt. B (bottom left) M I p J to by

the USDA Forester David Deevy. C (bottom right). SI yew as they grew, defoliated and debarked for extraction of the

anticancer chemical taxol (paclitaxel).

much like some T. canadensis of northeastern North America. Yew thickets may be male or female, but male

thickets were observed to be more common.

In the Klamath Mountains of California, I have observed thicket yew to occur predictably in steep ravines

and on steep banks with a north to northeast exposure at elevations from 1000-1200 m; however, I have also

found it at higher elevations in the Marble Mountain Wilderness such as near English Peak (1720-1820 m,

Lake-of-the-Island, Spjut 16013). It is interesting that the typical variety generally occurs at higher and lower

elevations in the California Klamath Region (e.g., 450 m, E fork of Willow Creek, Terrell et al. 4170; 700 m,

Tannery Gulch, Trinity Lake, Spjut 10179; 1500-2000 m, Marble Mt. Wilderness near Lovers Camp, Spjut

10721). Many of the more northern locations of var. reptaneta—in Oregon and Washington—are at slightly

f4L,D o ID L

222 Journal of t titute of Texas 1(1)

lower elevations, corresponding to the more northern latitude occurrences. These observations suggest that

var. reptaneta is a distinct ecotype within the range of var. brevifolia.

Other shrub forms in the Rocky Mountains of Canada are also evident. A specimen from the Selkirk

Range (British Columbia) differs by broader (oblong) clasping leaves with a pale glaucous green color on the

adaxial surface, while other leaves on the same plant are narrower and linear on what appears to be a layer-

ing branch. Another growing on steep exposed rocky cliffs near Golden (British Columbia) has ascending

stems from a burl-like base. In further study these plants, which are considered here as belonging to var.

reptaneta, may prove to be new varieties.

Eight samples of var. reptaneta were collected from Califonia, eastern Cascades in Oregon and Wash-

ington and from the Rocky Mountains in northwestern Montana to determine whether differences in taxoid

content were correlated with tree and shrub varieties of T. brevifolia (Spjut et al. 1993). The taxoid content

was most similar in samples that were also most similar morphologically, which were from northwestern

Montana and northern California (type locality for var. reptaneta), in comparison to samples of var. brevifolia.

These plants differed from those at other locations by the darker green thick leaves and by an abundance

of male cones on branchlets of the current season.

2. Taxus florinii Spjut, sp. nov. (Fig. 14). Tre: CHINA. YuNNaN: Litiping between Likiang and Weihai, tree 17 ft, in mixed

forests by stream,11 Oct 1939, R.C. Ching 21980 (HoLotype: A!, stomata 10 rows per band, abaxial marginal zone 5-6 cells across,

the cells thick-walled).

Similis a T. wallichiana, praecipue differt foliis decrescentes stomata et latis marginali, 7-12 seriata/zona.

Shrub or tree to 13 m high; young branchlets yellowish green, abruptly changing to reddish purple in 2

yr; bud-scales persistent on 1-2 yr branchlets in most specimens, 2-3 seriate, deltoid, concave, ca. 0.3-0.5

mm long, obscurely to prominently carinate, closely attached. Leaves arranged +distichous, homomallous,

linear-lanceolate, acuminate, straight to falcate, 1.5—3.5(-4.7) cm long, 2.0-4.0 mm wide, 200-350 um in

thick, dark glossy green above (dried leaves, similarly noted by collectors on fresh specimens), glaucous

to yellowish green below (in dried specimens, also noted as yellow green in field), slightly convex above

(adaxial) to a rounded midrib, channeled along each side below mid region to base, less concave below

(abaxial) to a flush to slightly rounded midrib, margins plane to slightly revolute; adaxial epidermal cells

nearly quadrate in transverse section, occasionally taller than wide, 20—50 pm tall, 20—40 pm wide; abaxial

epidermal cells narrower and not as tall as upper along midrib and marginal regions, rounded to partly

angular in transverse section, 5-12 um tall, 15-25 pm wide, numbering 16-20 across marginal region,

trapezoidal to short rectangular, 1-3x l/w near margins, longer nearer stomata bands, 3-5x l/w, and on

midrib, 5-10x+ l/w; papillae lacking near margins on 2-10 cells across, positioned more marginally than

medially on midrib in 1-2 alternating rows, medially to marginally on accessory cells, and along cell walls

in marginal zone; stomata bands narrower than marginal zones; stomata in 7-12 rows per band. Male cones

maturing on 1* and 2™ yr twigs, subcylindrical in bud, 3 mm wide, 4 mm long, yellowish green, the scales

4—5 seriate; microsporangia 5-6 on each sporophyll, pinkish, spongy. Female cones initiating on current

season growth, subcylindric in bud, 2-4 mm long, maturing 1* and 2™ yr, or on much older branchlets in

one specimen, the scales 5-6 seriate; seed ovoid to globose, to 5 mm long, 5 mm diam., pale (yellowish) in

color, sharply pointed at apex, covered in part by red aril, Aug.

Common name.—Florin yew.

Distribution and ecology.—Endemic to China (Xinjiang Uygur, Sichuan, Yunnan); forest or pasture or

along streams, 2500-3700 m.

Additional specimens. CHINA. Sichuan. Handwriting illegible, 3500 m, Schneider 1429 (K); Ngaitaschekou trans flumen Yalung ad

septentr oppodi. Yenyüen, 28?15'N, 2800-3500 m, Handel-Mazzetti 2602 (K); Shalie Valley on Muzhiyan Shan above Old Muli, Muli

country, ca. 2980 m, frequent in narrow, steep SW facing gulley, tree to 5 m, leaves dark green above, yellowish green below, Fliegner

et al. 1129 (K). Xinjiang Uygur (“SoC.W. Sikiang”): “NW Yunnan,” Tamichung, tree 20—30 ft, in fr., open pasture, R.C. Ching 21505

(A). Yunnan. NW, Mt. Fo Ludu, NW of Li-Kiang, W of the Yangtze, J.F. Rock 18502 (A, US); mostly illegible, 3000—3200 m, C. Schneider

2918 (A, K); no locality data, T.T. Yu 11076 (A); Dokerla, 3100 m, Yu 7848 (BM); Litiping Range, Mekong-Yangtze divide, E of Weihsi,

Spjut, Taxonomy of Taxus 223

J.F. Roch 11573 (A, US); Sikang, Me-kong, Tsa-wa-rung, 2500 m, C.W.

Wang 65475 (A); Mekong-Salween Divide, Forrest 19967 (S: C-2084);

S Chungtien, Kung-shiang-shu, Snow Mt on the way to Kai- Lou-wei,

ALA

on Yangtze bank, 2700 m, by stream in side of valley, tree 20

ft, K.M. Feng 3235 (A); same locality, 3200 m, in mixed forest, shrub

15 ft, in fruit, K.M. Feng 1809 (A); Wei-si Hsien, 2500 m, forest, C.W.

Wang 67735 (A); Zhongdian, Haba Shan, 27%22'28 N, 100?05'50 E.,

3347 m, Alpine Garden Soc. Exped. 309 (K).

Taxus florinii is recognized by the leaves having tall

angular epidermal cells and narrow stomata bands

relative to a broad region of marginal cells, the stomata

usually in 7-12 rows per band. As in T. brevifolia, the

Florin yew is variable in the number of leaf marginal

cells that lack papillae on the abaxial surface, ranging

from 2-16 cells across.

Specimens of Florin yew with fewer than 10

stomata rows per band are easily distinguished from

related species in Asia, but not necessarily from those

in North America; Schneider 2918 from Sichuan, for

example, is similar to T. brevifolia in the leaves having

7(—9) stomata rows per band bordered by short broad

irregularly shaped epidermal cells in the marginal

lull

zone, whereas Handel Manzzetti 2602—that is also from

Fic. 14. Taxus florinii, Herbari EE €

yr branchlet; Yunnan, China, Rock 18502 (US).

Sichuan—has 11 stomata rows per band—and is not

easily distinguished from T. globosa. Thus, T. florinii

and T. brevifolia could be included under T. globosa.

However, the American species generally differ by their

pale yellowish green branchlets and by the more strongly convex adaxial leaf surface—especially towards

leaf apex, in contrast to the dark purplish (rarely orange) branchlets in the 2™ yr and more plane leaves of

T. florinii. Other apparent differences are the midrib epidermal cells in T-section that in T. florinii appear

smaller, thicker-walled and less angular on the abaxial surface than the adaxial surface, whereas in the

American species there seems to be less difference in the size of the epidermal cells between the two leaf

surfaces as seen in the midrib region.

Etymology.—The epithet is in honor of Rudolf Florin who published extensively on taxads based on

leaf anatomical characteristics. Despite his controversial ideas on taxad evolution—such as proposing that

they be treated in a new class—I have found his reviews and data to be of great value.

3. Taxus globosa Schltdl., Linnaea 12:496. 1838. (Figs. 3, 15). Taxus baccata L. subsp. globosa (Schltdt.) Pilger, Planzenreich

18 (iv, 5):114. 1903. Tyre: MEXICO. HiLDAGO: Real del Monte, Cerro de las Dass [Sep] 1837, Ehrenberg s.n. (Lectotyee designated

here: ex Museum botanica Berlolinense (K!); ISOLECOTYPES: L, S, US!). No specimen ited, original material at B destroyed. Parlotore

(1868) cited an Ehrenberg collection with number from the type locality, Ehrenberg 817, which may be at FI.

Taxus globosa is characterized by leaves having quadrangular epidermal cells as seen in transverse section,

stomata in (57-11 rows per band, and alternately arranged papillae along cell walls. Higher counts of

stomata, 9-11 rows per band, were found in specimens from Central America and southern Mexico, com-

pared to lower counts, 5—7 rows per band seen more in specimens from the northern range in Mexico and

in the Florida panhandle (Spjut 2007). The abaxial leaf margins vary from 2—5 cells across (without papil-

lae) except for one specimen from Veracruz, while all five specimens from Florida have a slightly broader

marginal (epapillose) region, 5-9 cells across.

The species is traditionally known to occur from Mexico to Honduras; however, at the northern-most

range in Mexico, specimens are difficult to distinguish from those in Florida; therefore, the Florida yew is

224 Journal of the Botanical R h Institute of Texas 1(1)

Fic. 15. Herbarium specimen in part, Taxus globosa, Ehrenberg s.n. (lectotype, K).

reduced to variety of T. globosa.

3a. Taxus globosa var. globosa

Common name.—Mesoamerican yew.

Distribution and ecology.—S Mexico (Veracruz, Hidalgo, Mexico, Oaxaca), Guatemala (Baja Verapaz,

El Progreso [Sierra de las Minas, Volcán de Santa Luisa], Zacapa [Volcán Gemelos, Monte Virgen] Huehu-

etenango [Sierra de los Cuchumatanes, Cerro Cananá]) to British Honduras (Merrendón, San Idalfonso,

Omoa, Montecillos, Opalaca, Congolón) and El Salvador; montane cloud forests above 2,000 m.

3b. Taxus globosa Schltdl. var. floridana (Nutt. ex Chapm.) Spjut, comb. nov. (Figs. 4, 16). Basiony: Taxus

floridana Nutt. ex Chapman, Fl. South. U.S. 436. 1860. Taxus baccata L. subsp. floridana (Nutt. ex Chapm.) Pilger, Planzenreich

4(5):113. 1903. Taxus canadensis Marshall var. floridana (Nutt. ex Chapm.) Silba, Phytologia Mem. 7:72. 1984. Tyre: U.S.A. FLORIDA:

near Aspalaga, 1833, Croom s.n. (LEcTOTYPE designated here: PH!). No specimens cited in original description. According to Sargent

(1896), the Florida yew first discovered by Hardy Croom in 1833 near Aspalaga. He further stated that "[t]he first notice of this,

without description or specific name, was published in 1834 in the American Journal of Science (xxvi. 314)...It was next mentioned

by Nuttall in 1849 (Sylva iii: 92) who doubtfully attached to it the name of Taxus montana [synonym for Torreya taxifolia], although

Crooms specimen in the herbarium of the Philadelphia Academy was, he says, marked Taxus Floridana, the name adopted by

Chapman when the species was finally described in 1860." A specimen at PH was found with label in what appears to be Nuttall's

handwriting bearing the name Taxus floridana, indicating it was collected by Croom. However, another name, T. croomii, had been

proposed, evidently by Chapman with reference to a Chapman list (1845 fide annotation by K. Wurdach, 1987, specimen at PH)

as seen on other specimens from C.W. Short (PH), Bentham Herbarium (K), Herbarium Careyanum (K), all of which may represent

original material. Chapman indicated that Nuttall was the source for the epithet; the specimen at PH bearing the name T. floridana,

therefore, was selected as lectotype. The type material for the 1860 edition of the flora by Chapman reportedly went to Columbia

University, which is now at NY (Stafleu & Cowan 1976), but no possible type specimens for Taxus from Chapman are currently

listed among the types at NY (virtual herbarium)

Spjut, Taxonomy of Taxus 225

roe e

aye? rec

s ! "ih

Eco pupa eie ma cT o E

coenae io rit

| Bes "auo GAUL A a scc ci a LP AD,

Fic. 16. Taxus globosa var. floridana, close-up of branchlets and leaves of a herbarium specimen; handwritten notations on a label "Taxus croomii"

Chapman List, Florida (original material, K).

£+sha D o ID L

226 Journal of t titute of Texas 1(1)

Common name.— Florida yew.

Distribution and ecology.—N Mexico (Nuevo Leon, Tamaulipas, Veracruz), W Florida (rare, Apalachicola

River; Chamaecyparis swamp ca. 8 mi SE of Bristol).

The Florida yew, which differs by the abaxial leaf surface having a broader marginal area of nearly

rectangular epidermal cells with less prominent papillae, occurs in northern Mexico and in the panhandle of

Florida. In Florida it is found not much above sea level on bluffs and in ravines along 15 miles of the Apala-

chicola River in a mixed evergreen forest of Fagus grandiflora Ehrh. and Magnolia grandiflora L. with Torreya

taxifolia Arn., Kalmia latifolia L., Quercus laurifolia Michx., Pinus glabra Walter, Ilex opaca Aiton, Symplocos

tinctoria (L.) UHér., and Vaccinium elliotii Chapm., and locally in a white-cedar (Chamaecyparis thyoides [L.]

Britton, Sterns & Poggenb.) swamp eight miles southeast of Bristol (Kurz 1937). In northern Mexico, it oc-

curs between 2000 and 2500 m, which is at lower elevations than generally reported for the typical variety.

The Florida yew has allegedly retained the ancestral features of leaf epidermal papillae on the midrib,

and the angular isodiametric epidermal cells (as seen in T-section), contrary to what might be expected for

yew growing in a seasonally hot and humid climate at relatively low elevations. An example is Taxus mairei

in mixed mesophytic forests of southern China; it lacks papillae on the abaxial midrib and has elliptical

epidermal cells (T-section). Variety globosa, occurring in a subtropical montane forest with less seasonal

variation in temperature and precipitation, shows only minor differences in numbers of leaf stomata rows

and marginal cells compared to var. floridana. Perhaps there was not enough time for the Florida yew to have

evolved significant morphological differences between the oscillating periods of climate change during the

Pleistocene, or perhaps introgression has occurred between formerly distinct ecospecies, one found along

the Gulf coastal plain and another in the upland areas. The overlap of key morphological features at the

northernmost range of the species in Mexico would seem to indicate former contact in that region between

two ecotypes, whereas the plants of limited occurrence in Florida are viewed here as relicts barely surviving

instead of adapting to the present climate.

4. Taxus suffnessii Spjut, Sp. nov. (Figs. 1,1 7). Tree: MYANMAR. North Triangle (Wring Burma above Ahkail), 9000-10,000

ft, in forest and thickets, young leaves bronze, shrub or small tree, 24 May 1953, Kingdon Ward 20902 (HoLotyee: A!, with seed; leaf

with 12-13 stomata rows/band; isotype: BM!, leaf with 16 stomata rows/band, lacking papillae across 2 marginal cells).

Arbor vel frutex similis a T. wallichiana, praecipue differt perulae persistens ampliatibus, ovatis, 2-3 mm longis, folia oblongas ad leviter

ellpiticas, 1.5 cm longis, 3.0 mm latis, supra cellula epidermibus ca. 50-60 pm altis, ca. 50 um latis.

Tree or shrub; branchlets unequally divided, gray to purplish on older growth; bud scales 3—4 seriate, per-

sistent to the 3"! yr or longer, paleaceous, indurate, grayish, deltoid, closely to loosely adnate, lower scales

concave with a distinct midnerve, uppermost scales slightly cuspidate and aristate, ca. 3 mm long and 2 mm

wide. Leaves lacking on older twigs, nearly two-ranked, overlapping slightly, oblong to slightly elliptical or widest

above the mid region, straight, mostly ca. 1.5 cm long, 3.0 mm wide, 250—350 pm thick, bronze to blackish

green and concave above to a rounded or acutely keeled midrib, yellowish green and convex below to a

rounded keeled midrib, abruptly revolute near margins. Adaxial epidermal cells quadrangular in T-sect.,

or taller than wide, to 60 pm tall and 50 um wide; abaxial non-stomata epidermal cells not as large, 12-25

um high and wide, quadrangular, papillose to near leaf margins; papillae prominent, globose, in 3-4 rows,

notably medial on marginal cells and marginal on midrib cells or equally developed on accessory cells in

a stellar arrangement; stomata 12-20 rows per band. Male cones not seen. Female cones maturing on cur-

rent season growth, scales overlapping in 5-6 ranks; seed appearing succulent or deformed, conical, dark

colored with notable yellowish neck, ca. 3 mm long, 1.5 mm wide.

Common name.—Suffness yew.

Distribution and ecology.—E Himalayas: Endemic to Myanmar; 1950-3048 m.

Additional specimens. MYANMAR. Upper Burma: Hkyet, 27%45' N, 97?50' E, 9000-10,000 ft, Kingdon Ward 13003 (BM); Myintkyina,

Sumprabum, 8600 ft, Hla & Koko 4028 (K); W Central Esakan, 6400 ft, in thick forest, Kingdon Ward 21901 (A).

Taxus suffnessii stands apart from other species of Taxus in many features. These include the relatively large

Spjut, Taxonomy of Taxus 227

tall-rectangular epidermal cells, the large

persistent bud-scales with a distinct mid

nerve, prominent papillae on the abaxial

surface of leaves extending nearly to the

margin, a relatively broad leaf shape—ap-

pearing widest above the mid region,

and leaves not spreading in one plane

along two sides of a branchlet. Although

the number of stomata rows varies from

12-20, the highest number of stomata

rows seems related to the papillose cells

extending to the leaf margin, or within

2 cells from the margin; essentially, the

stomata band covers the entire abaxial leaf

surface. The poorly differentiated stomatal

bands would, therefore, seem to be an

ancestral trait.

The Suffness yew is similar to T. wal-

lichiana that differs by the smaller, more

polished, nearly nerveless bud-scales and

P "222A

sometimes by the older reddish orange

branchlets as seen in the typical variety

I! (A tarot

from northeastern India, in contrast to a Fie. 17. Taxus suffnessii,

purplish color in the Suffness yew; how- persistent scales at base; Myanmar, Kingdon Ward 20902 (holotype, A).

ever, intermediates are apparent. Examples

are Kingdon Ward 21901 from West Central

Myanmar (at “6400 ft”) and Beer 25316 from Nepal (above Sedua, 9400 ft). They have fewer and slightly

smaller bud-scales at base of branchlets but still show the characterstic midnerve, while they also differ by

having longer (linear) leaves. Kingdon Ward 21901 (BM) is referred to T. suffnessii by the entirely papillose

abaxial leaf surface (Spjut 2007). The specimen from Nepal, however, is identified T. aff. wallichiana (Spjut

2007), while similarity to T. suffnessii is further evident by the tall-rectangular epidermal cells in leaf tran-

servse sections in contrast to what is generally seen in T. wallichiana. The relatively large epidermal cells are

a striking character feature of T. suffnessii and also other species such as T. florinii and T. brevifolia. The extent

to which this character can be further employed to differentiate species of Taxus needs further study.

Taxus suffnessii is named in honor of the late Matthew Suffness in recognition of his dedication to screen-

ing natural products in the search of new drugs to treat cancer and of the strong encouragement I received

from him in this endeavor. He became Chief of the Natural Products Branch in the National Cancer Institute

after Jonathan Hartwell retired in 1976, following a brief leadership by John Douros. In 1986, Dr. Suffness

became more involved in extramural contracts, often serving as consultant to various drug discovery groups.

He focused on identifying novel leads that showed promise for development as new anticancer drugs, one

of which was taxol. It is ironic that a major compilation on taxol research to which he served as sole editor

(Suffness 1995) would not appear until just after his death from cancer in the spring of 1995.

5. Taxus wallichiana Zucc. in Siebold & Zucc., Abh. math.-phys. CL.K. Bayer. Akad. Wiss. (München) 1

(3):803, Tab. 5. 1843. (Figs. 18-21). Taxus baccata L. subsp. wallichiana (Zucc.) Pilger in Engler, Pflanzenreich IV (5):112.

1903. Taxus baccata var. wallichiana (Zucc.) C.K. Schneider ex Silva Tarouca, Freiland-Nadelgehólz. 276. 1913. No specimens cited by

Zuccarini, but he provided an illustration with reference to another earlier illustration in Wallich (1826, Tentamen florae Nepalensis 57,

Tab. 44), Wallichs name “Taxus nucifera?” (excluding synonyms) and Wallich's a that the species occurred in mountains

Í e

Wallich and/or his collectors:

around Sheopore [Shivapuri], Kathamadu, Nepal). Original material at M i

228 Journal of the Botanical R h Institute of Texas 1(1)

-- iod

pec PH

^an s

Fics. 18-19. Scanned images from sketches on 9 x 1

Ws aleaf for wo Yanenes sot fins wallichiana: ds (top). Var. walichiana, Bhutan, Ludlow Sherri uy (BM), shows abaxial leaf surface to haye An rows

ularly

alienate papillae and indicated: to have 15 rows af stomata anda RENA of 15 calls across with alternate papillae! 19 (bottom). Variety yunnanensis,

Sikkim, J. D. Hooker (K), indicating abaxial marginal zone has three rows of long cells without papillae, followed by 14 rows of stomata, without a

var.

J Es rr ret J

D £

Spjut, Taxonomy of Taxus 229

Fics. 20-21. Original material of 7 llichi 20 (left). Lecto-

type, Wallich s.n., E Indi g ts, abaxial surface

£1 J 4 I 21 (below) HI H cel "an

& Zuccarini (1843, Tab. 5).

230 Journal of the Botanical R h Institute of Texas 1(1)

(1) Wallich s.n., year 1835, eastern India; with Herb. Zuccarini label, (2) [a Min] 6054A on label from Schultes, indi d to be from

Kumaon but most likely from Nepal (see East India Co., N ical list of dried 1 1 1 f Dr. Wallich

[1828-1849]; determined by Spjut as T. wallichiana), (3) [Wallich] 6054/a om Nepal, ex Herb. Hort. Bot. ita: Geom

by Spjut as T. contorta) and (4) [Blinkworth] 6054B from Kumaon (determined by Spjut as T. contorta). LECTOTYPE e by Spjut

inadnot, 23 March 1995 and here: INDIA: eastern, communicavit Wallich, year 1835—Wallich s.n. (M!) with mature m nes, and

label *Herb. Zuccarini" Among the specimens at M, the lectotype is also the best match for the illustration in Siebold & Zuccarini

(1843)—a photocopy reproduced here (Fig. 21). Additionally, two of the four specimens at M that bear Wallich numbers 6054/a

and 6054b appear to have the wrong labels attached (or reversed) based on study of Wallich duplicates at other herbaria (GH, K, NY,

ichiana (Siebold & Zuccarini 1843), the

lectotype. Duplicates of Wallich 6054A, which might be equivalent to isosyntypes (indicated as type below), are detailed as follows

by institution: GH: label with handwriting similar to Wallich, *Taxus nucifera Wall." *Napalia." s.n. K: Four sheets. (1) with two

specimens, the larger specimen has a pasted label below it with handwriting “6054a” and no indication of locality data, the smaller

one is a single branch with mature seed, correctly annotated T. wallichiana by S. G. Harrison, but it is not 6054A, or not a type since

it is not from a male plant; (2) two specimens, one large specimen with an imprinted stamp nearby—Herb. Hookerianum, with

handwriting similar to Wallich, 6054/A, Nepal, accompanied by a smaller specimen in left corner, with a large label below, Watt 6493

—

P, PH, S); however, only one specimen is clearly associated with Zuccarinis study of T. wal

from Manipur, det. T. wallichiana by Spjut; (3) has four specimens, but only the lower left specimen is a type (T. wallichiana), below

it are several labels, one printed—ex Herb. George Gordon, presented by J. D. Hooker, 1878, the other bears handwritten annota-

tion—” Taxus wallichiana,” two largest specimens with letters a and b written nearby on left and right, respectively, and with Herb.

Benthamum imprinted in center, belong to T. contorta; uppermost annotated T. virgata, det. by Spjut to be a young shoot of T. baccata.

NY: 2 sheets, 6054A, NY accession numbers 30328 and 30329 (det. via photocopy). P: “Napalia,” 6054 with “A” inserted, annotated

Taxus nucifera Kaempf.? on label ex. Herb. Richard, and additional label ex. Herb. E. Drake. PH: *6054A Wallich."

Zuccarini was one of many recipients of specimens distributed by Wallich who generally assigned collection

numbers to species rather than to specimens in which a particular specimen number may come from different

localities and from different collectors. For example, Wallich 6054A has been reported from Central Midlands

near Kathmandu Valley in Nepal (Anonymous 1913; Hara et al. 1978), and also from “Cachemiro” (Parlatore

1868), but most likely 6054A was collected from around Kathmandu (Dan Nicolson, pers. comm.1995)

during 1822 (^Wallich's Catalogue" 1831-1832, Stafleu & Cowan 1988; Wallich 1826). Wallich was not al-

lowed to leave the Valley of Kathmandu, while his collectors were able to collect, for example, northwest of

Kathmandu Valley on the Holy mountain of Gossain (Nicolson, pers. comm., 1995). Wallich specimens of

T. wallichiana with notations of *Kumaon" appear to be an error in numbering or labeling since collections

from “Kumaon,” which generally belong to T. contorta, were collected by Robert Blinkworth as indicated in

Walllich's numerical list (^Wallich's Catalogue" 1831-1832, Stafleu & Cowan 1988). Wallich's collections

for numbers 6054a, 6054b, which were collected prior to 1826, were distributed sometime between 1831

and 1832 (Anonymous 1913).

However, the lectotype, a specimen collected by Wallich from eastern India—received by Zuccarini

in 1835 without number—would appear to be a later collection unrelated to the distribution of Wallich's

Cataogue and herbarium. Indeed, Wallich was in Assam in 1835 (Burkhill 1965). It should be further

noted that an illustration in Siebold and Zuccarini (1843, Tab. 5, reproduced here as Fig. 21) most closely

resembles the lectotype among the specimens at M as seen by the branching and distribution of male cones.

The lectotype is also characterized by reddish orange branchlets, non-inflated epidermal cells on abaxial

surface of leaves, 12-15 stomata rows per band, and marginally positioned papillae on epidermal cells.

Two varieties are recognized by differences along the abaxial leaf margin as seen by the size and shape

of epidermal cells and by the position of papillae.

5a. Taxus wallichiana var. wallichiana

Common name.—Wallich yew.

Distribution and ecology.—C Himalayas to SW China; Nepal, Bhutan, NE India (Assam, Manipur, Khasia

Hills, West Bengal), Myanmar, China (SE Tibet, Sichuan, Yunnan); montane coniferous forests with Picea,

Abies, Tsuga, or broadleaved evergreen forests of Lithocarpus, or Quercus, (1500-) 2300-3200 m. In Nepal

evidently occurring abundantly with Abies spectabilis (D. Don) Spach on limestone (Stainton 1972), and in

Bhutan apparently scattered from Ha to Mongar districts (Grierson & Long 1983).

Typical T. wallichiana is identified by the pale reddish orange branchlets, persistent cuspidate bud-

Spjut, Taxonomy of Taxus 331

scales (Pilger 1916), linear leaves arcuate near base (Orr 1937; Pilger 1903), conically shaped seeds (Orr

1937) that often mature on 2" year or older branches (in the Himalayas), and angularly shaped epidermal

leaf cells in T-section. Its leaves are further distinguished from those of T. contorta by the adhesive, bone-

like parenchyma cells, and by (11-)13-18(-21) rows of stomata per band. Plants from Nepal, West Bengal,

Khasia, and Bhutan are similar to the type.

Taxus wallichiana has been the name applied to all yews in southeastern Asia (Hu 1964; Pilger 1903 as

subsp. wallichiana); however, de Laubenfels (1988) adopted T. sumatrana for his treatment of gymnosperm

taxa in Flora Malesiana. He indicated that several species may overlap in the eastern Himalayas, suggest-

ing that T. wallichiana was outside the Flora Malesiana region. Taxus wallichiana has largely been ignored

by Rehder (e.g., Rehder 1940, 1949) and Hortus Third (Liberty Hyde Bailey Hortorium Staff 1976), while

others have mentioned it as a species confined to the Himalayas (Krüssmann 1985), or more limited to

the northwestern Himalayas (Wilson 1926), or as one of two partially sympatric species predominantly

Himalayan in distribution (Silba 1984). Since Pilger (1903, 1916) did not cite any specimens for Taxus in

the western Himalayas but indicated T. wallichiana to occur in eastern Himalayas, this omission may reflect

uncertainty on his part, as he noted there were intermediates to T. baccata.

While I do not accept all morphological variants of Taxus in southeastern Asia to belong to a single

species, Handel-Manzzetti (1929), Florin (19483), and Hu (1964) also recognized more than one sympatric

species in Asia by the lack of papillae on the abaxial leaf midrib, which I consider applicable to the Sumatrana

Group as a whole.

The taxonomic and ecological significance of midrib papillae on the abaxial epidermal surface of leaves

in Taxus has been noted by Bertrand (1874), Deryugina & Nesterovich (1981), Florin (1931, 1948b), von

Frimmel (1911), Orr (1937), and Spjut (1992, 1993, 1998a, 2000a; Spjut in Hils 1993); however, Kwei &

Hu (1974) and Cheng & Fu (1978) recognized intermediates with partially papillose midribs between T.

wallichiana (papillose midrib) and T. sumatrana (smooth midrib). The latter was treated as a variety under

two illegitimate combinations (T. chinensis var. mairei, T. wallichiana var. mairei). Spjut (1992, 1993, 1998b,

20003), however, found other features that support their distinction not only as species but as species groups,

such as epidermal cells in transverse sections appearing angular in the Wallichiana Subgroup of species (C

& E Himalayas to SW China; North America) and elliptical in the Sumatrana Group of species (E Himalayas

to Indonesia, Philippines).

Taxus wallichiana is interpreted to occur on Mt. Emei in Sichuan (China), where it intergrades with T.

chinensis. Subtle differences in size of bud-scales and color of branchlets make it difficult to consistently

separate the two species. Problematical plants may be hybrids between T. wallichiana var. yunnanensis and

T. chinensis and/or possibly another species distinguishable by slightly larger and more persistent scales at

the base of branchlets.

5b. Taxus wallichiana Zucc. var. yunnanensis (W.C. Cheng & L.K. Fu) C.T. Kuan, Fl. Sichuan. 2:215.

1983. (Fig. 19). Taxus yunnanensis W.C. Cheng & L.K. Fu, Acta Phytotax. Sin. 13(4):86, fig. 52, 4—7. 1975. Taxus chinensis (Pilg.)

Rehder var. yunnanensis (W.C. Cheng & L.K. Fu) L.K. Fu, Vasc. Pl. Hengduan Mount. 1:214. 1993. Type: CHINA. TIBET: ZavuL, 2100

m, 2 Aug 1973, Zhang 916 (noLoTYPE: CAF; isotype: PE!, leaf fragment! photocopy!). Topotyres: detached leaves, without collector or

date, “type locality" (PE); Kingdon Ward 10398 (Zayul, Rong To Valley, 8000 ft, spreading tree with brilliant green foliage, amongst

deciduous trees on slopes and in gullies, BM!).

Common name.—Yunnan yew.

Distribution and ecology.—India (Sikkim, Nagaland), Myanmar, China (Tibet, Yunnan, Sichuan); mixed

forests types, generally higher in elevation than var. wallichiana, 2100-3500 m, occurring with Larix griffi-

thiana Carriere and Picea spinulosa (Griff.) A. Henry in the Sikkim region (Rau 1974).

Taxus yunnanensis has been confused with T. wallichiana in the Flora of China (Cheng & Fu 1978). The

authors had evidently considered the type for T. wallichiana to represent the species mainly in northwestern

Himalayas; juently, they described T. yunnanensis—indicating it was found in the eastern Himalayas

(Bhutan, Tibet, Myanmar) to Yunnan and Sichuan (Cheng et al. 1975; Cheng & Fu 1978). Later, it was reduced

232 Journal of the Botanical R h Institute of Texas 1(1)

to a variety of T. wallichiana, as cited above, and more recently placed in synonymy (Li & Fu 1997), although

it had been included in synonymy by de Laubenfels (1988) under his broadly circumscribed T. sumatrana.

Li and Fu (1997), in placing T. yunnanensis in synonymy with T. wallichiana, created a new name for the yew

distributed in the northwest Himalayas, T. fuana, but this is antedated by T. contorta. The English edition

of the Flora of China (Fu et al. 1999) follows Li and Fu (1997), while Farjon (1998, 2001) maintained the

illegitimate name with a narrower geographic distribution that seemed to reflect only the type locality.

Most specimens I annotated T. yunnanensis (A, GH, July 1996; BM, Oct. 1997; Spjut 1998b) are from

Yunnan and Sichuan. They are distinguished from typical T. wallichiana by the leaves appearing slightly

wider (nearly lanceolate), more evenly distributed, less markedly curved across the adaxial surface and

paler green below than above, and having medial papillae on the abaxial epidermal cells. At the time I had

seen only leaf fragments of a type—from Tibet near the border with Myanmar and India; it differed from

the type of T. wallichiana by the abaxial surface having a broad region of large epidermal cells with medial

papillae between the margin and stomata band. These features were seen more often in yew specimens from

Yunnan and Sichuan than from northeastern India. Later, I received a B&W photocopy of a PE isotype from

Dr. Z-y. Cao, who had earlier sent me leaf fragments of topotypes, and I found that the leaf arrangement

and shape compare more closely to the type of T. wallichiana than to specimens from Yunnan and Sichuan.

Thus, plants most typical of this variety, as seen in northeastern India and nearby Tibet, are intermediate

forms distinguishable only by leaf anatomical characters. For this reason, T. yunnanensis is reduced to a

variety. Nevertheless, it is important to differentiate these and one other related species (T. florinii), in order

to distinguish the North American species (T. brevifolia, T. globosa) from their Asian relatives; otherwise,

they may have to be included under T. wallichiana.

Variety yunnanensis is distinguished by the abaxial leaf marginal zone having relatively long narrow

epapillose cells (2-4 cells wide) in contrast to the irregularly quadrate shape in var. wallichiana. This region

of narrow elongate cells, 2-4 cells wide, is usually followed by much wider zone of papillose cells (5-15

cells across), but a transitional zone of papillose cells may be absent (Fig. 19B). In transverse sections the

leaves also appear thinner and more revolute along margins, and specimens from Tibet (type locality),

Myanmar and Naja Hill in northeastern India have taller epidermal cells along the abaxial leaf marginal

zone. Another characteristic feature of var. yunnanensis is that papillae are mostly opposite and erect, best

observed on the abaxial midrib (and also marginal cells). Photomicrographs of leaf stomata bands for species

of Taxus in Jinxing and Yuxi (2000) show excellent resolution at 1000x for the medial (opposite) papillae

that characterizes T. wallichiana var. yunnanensis (referred to as T. yunnanensis), in contrast to the marginal

(alternate) arrangement seen in their T. chinensis, a species in which the papillae arrangement is similar to

that of T. wallichiana var. wallichiana. Their photos presented for T. wallichiana, however, are of T. contorta as

seen by the narrow linear cells with only a single row of papillae.

IB. Wallichiana Group, Subgroup Chinensis.—The Chinensis Subgroup is characterized by leaves hav-

ing elliptical to rarely wide rectangular epidermal cells in transverse sections, notably larger on the adaxial

surface than abaxial surface, and stomata that align both anticlinally and periclinally. Plants with angular

epidermal leaf cells, as viewed in T-section, are included if they have yellowish (ochre) tinted branches,

vestigial bud-scales at base of branchlets, and oblong leaves. The Chinensis Subgroup comprises T. chinensis

and two proposed (undescribed) species in central China, occurring between 150 m and 2500 m in elevation,

and three other species that extend into the Himalayas to Nepal and from the Pacific to the Philippines and

to Indonesia, where mostly found above 2000 m (T. obscura, T. phytonii, Taxus sp. undescribed—Taxus REH).

Introgression with species of the Sumatrana Group, Wallichiana Subgroup and Cuspidata Alliance is

evident in the northern-most and southern-most ranges of the Chinensis Subgroup. The northern range of

the Chinensis Subgroup is in central China, represented by T. chinensis. In this region, T. chinensis has sharply

divergent leaves devoid of papillae across 8-12 abaxial marginal cells (e.g., Henry 7097 from western Sich-

uan). This is also seen in T. umbraculifera (Cuspidata Alliance) of northeastern China, Korea and Japan. At the

southernmost range—in Indonesia and the Philippines—are two other species of the Chinensis Subgroup;

Spjut, Taxonomy of Taxus 233

one Taxus obscura is similar to T. chinensis in the leaf shape, texture and lack of epidermal papillae across

8—9 marginal cells, and another, Taxus phytonii, that resembles T. sumatrana in the leaves having a reddish

color along marginal and midrib epidermal cells. The latter is also similar to T. wallichiana in the linear leaf

shape, two-ranked leaf arrangement, and in the persistence of bud-scales at base of branchlets.

Although leaf stomata in Taxus develop in periclinal rows, stomata in the Chinensis Subgroup often align

transversely with the stoma anticlinally oriented. The Sumatrana Group, in contrast, has stomata arranged

+alternately to each other along adjacent rows. The anticlinal stomata in the Chinensis Subgroup, in which

the stomata bands are further differentiated from adjacent epidermal cells by color, have been observed

mostly in Taiwan and Luzon specimens. Other specimens from mainland China have less distinct stomata

bands as evident by a more uniformly yellowish orange to reddish color on the abaxial leaf surface.

6. Taxus chinensis (Pilg.) Rehder, J. Arnold Arbor. 1:51. 1919. (Fig. 22). Taxus baccata L. [subsp. cuspidata (Siebold &

Zucc.) Pilg.] var. chinensis Pilger, Pflanzenreich IV, 5:112. 1903. Taxus cuspidata Siebold & Zucc. var. chinensis (Pilg.) C.K. Schneider ex

gehólz. 276. 1913. Taxus wallichiana Zucc. var. chinensis (Pilg.) Florin, Acta Hort. Berg. 14, 8:378. 1948.

Tre: CHINA. E Sichuan: Wushanhsien, 2000-3000 m, year 1885-1888, Henry 7155, with seed (Lectotyre by Rehder & Wilson in

Sargent, Pl. Wilson. 2:8. 1914: A-18682; isoLectotypes: BM!, E, GH!, K!, S fragment, USD). The A and GH sheets were annotated by Hu

as “type” and “isotype,” respectively. SYNTYPES: Henry 6913, Farges 128. Pilger cited several collections but did not designate a type.

Silva Tarouca, Freiland-Nadel

Taxus baccata L. var. sinensis A. Henry, Elwes & Henry, Trees Gr. Brit. and Irel. 1:100. 1906. Nomen illegit. (CBN Art. 53.3, Ex. 9). Type:

CHINA. E Sicuuan: Wushanhsien, 2000-3000 m, 1885-1888, Henry 7097 (Lectotyee designated here: E; ISOLECTOTYPES: A!, BM!, P!,

US!). No specimens cited; original material at E, Henry nos. 6913 from Hubei, 7097 and 7155 from Sichuan.

Common name.— China yew.

Distribution and ecology.—Mostly China (Guangxi, Gansu, Yunnan, Sichuan, Guizhou, Hubei, Anhui,

Zhejiang), one collection from Vietnam (Hiep & Chan 405, P); forest, or forest margins, or open scrub, “under

2 «

rocky cliffs,” “often among bamboos,” generally 1000-2800 m. Reported also at elevations as low as 150 m

(Hu 1964). In Sichuan found more in the drier “mixed mesophytic forest” or “transitional zone” to an ever-

green oak forest, in contrast with T. wallichiana occurring more in hemlock-spruce-fir forests (Wang 1961).

In Vietnam “a shade tolerant” species of limestone in understory of evergreen forest of Pinus kwangtungensis

Chun & Tsiang, Podocarpus neriifolius D. Don, P. nilgeri Foxw. and other broad-leaved trees, 1000-1600 m

(Hiep 1998).

Taxus chinensis is distinguished by the pale yellowish green or “yellowish ochre” to “bronze” (“Prisma-

color” chart) color on older branchlets (“dun colored”; Orr 1937) and by the relatively short (oblong), thick

leaves, usually with conspicuous midrib papillae along epidermal cell walls. In making this distinction, I

have independently reached the same conclusion as that by Pilger (1903, 1916), Orr (1937), Florin (1948a),

and Hu (1964) for recognizing this taxon by its leaf and bud-scale characteristics, and also that by Orr (1937)

for its branch color. Although branchlets of T. wallichiana vary in color from reddish orange to purplish, they

lack this yellowish pigmentation, or are not yellowish green.

The leaf stomata bands are bordered by 4-12 marginal cells. The stomata develop in 11-19 (-21) rows

per band and are sometimes evident on the midrib.

The name T. chinensis was once used for any yew occurring naturally in China (Rehder 1940), and

also Taiwan, the Philippines, and Indonesia (Wilson 1926)—until the earlier legitimate names, which had

been classified in Cephalotaxus and Tsuga, were applied; the ICBN (Art. 11.4, 11.5) requires that the earli-

est epithet be adopted regardless of the genus it was erroneously assigned to—unless conserved. Rehder

(1936), for example, discovered one—Tsuga mairei Lemée & Lév., but continued to use his name, T. chinensis,

whereas Parlatore (1868) and Pilger (1903, 1916) had reported several earlier names (Cephalotaxus sumatrana,

Cephalotaxus celebica), whose epithets were eventually adopted, T. celebica (H.L. Li 1963), T. sumatrana (de

Laubenfels 1978); however, the correct name for a single subtropical species as applied by these authors

would have to be T. wallichiana.

Not all taxonomists accept just one species of Taxus in southern China; Florin (19483), for example, felt

there were at least two: T. chinensis, which he treated as a variety of T. wallichiana with distribution primarily

234 Journal of the Botanical R h Institute of Texas 1(1)

Jenne L IL L £ I + | £L kiai IĻ "I hi I P" I +L J nl r4 F

Fic. 22. Taxus chinensis,

Sichuan, China (isolectotype, K.

in Sichuan, and another that he considered a new species, T. speciosa Florin, which was not entirely new since

it had been earlier described, as already indicated, and Florin himself mentioned the names in synonymy

(Cheng & Fu 1978). Florin's two species were distinguished by the presence or absence of papillae on the

abaxial leaf midrib, and var. chinensis was further distinguished from var. wallichiana (in the Himalayas) by

the relatively shorter (oblong v. linear) leaves. He also indicated that T. baccata var. sinensis was synonymous

with var. chinensis according to communications he had with Orr at Edinburgh, who had sent him leaves of

Henry’s collections (Henry 7155), which Henry himself had named var. sinensis. Although I have not studied

the Edinburgh specimens of Henry's collections, the type for var.sinensis (Henry 7097) was selected based

on material at E sent to Florin, who cited Henry 7097, 7155; the latter (7155) is the type for T. chinensis.

Botanists in China have since recognized up to four species and one variety of Taxus in China (Cheng

& Fu 1978); however, they have misapplied and illegitimately combined previously known names. These

included T. chinensis var. mairei (Lemée & Lév.) W.C. Cheng & L.K. Fu (illegit.), T. wallichiana (misapplied

to T. contorta), and T. yunnanensis (superfluous for T. wallichiana). Hu (1964) followed Florin's treatment

(19482) except that she maintained T. chinensis as a species. Cheng & Fu (1978), however, reduced it again

to a variety, but not according to ICBN, as just indicated. Taxus chinensis and T. mairei were considered to

differ only as varieties because the distinguishing feature—presence or absence of papillae on the leaf midrib

(undersurface)—could not always be clearly decided due to the occurrence of intermediates (Kwei & Hu

1974; Cheng & Fu 1978), and also because the type for Cephalotaxus celebica Warb. had not been studied (Hu

1964; Cheng & Fu 1978). Hu (1964) concluded that the type for T. speciosa did not significantly differ from

that of Taxus (Tsuga) mairei (Lemée & H. Lév.) S-Y. Hu ex T.S. Liu, whereas the type for Cephalotaxus celebica

might differ because the only specimen she saw from the Celebes Islands (Neth. Ind. For. Serv. bb:19577, A)

Spjut, Taxonomy of Taxus 235

hn a

L H Wall £

Fic. 24. Taxus phytonii, g RE [ ET

: thal f+ . H g, incon- bud I +h £L hi ; Taiwan, Wilson 11154 (holotype, A).

Fic. 23 Taxus obscura, listi

rr J T T

I l PE g | hi J Hall £

in 7. phytonii; Philippines, de Laubenfels P668 (holotype, A).

had leaves with a papillose abaxial midrib, and in her opinion this was distinct from the types of T. mairei

and T. speciosa.

I have studied the same specimens that Hu studied at the Harvard University herbaria (A GH, anno-

tated by Hu, July 1955) and largely concur with her identifications of T. chinensis (Hu 1964). Additionally,

I have studied other specimens from Sulawesi, namely the type for Podocarpus (Taxus) celebicus Hemsl. (K),

Teysmann 14190 (U), and a photocopy of a holotype fragment for Cephalotaxus celebica Warb. (S); these have

similarly shaped leaves with a smooth midrib and broad marginal area of partially papillose cells on the

abaxial surface; the “Neth. Ind For.” specimen (A), in contrast, differs in having a papillose midrib as noted

by Hu (1964). Based on Florin's (19482) account and the similarity in leaf shape of the two type specimens,

in which I have recognized T. celebica by its long acuminate leaves (tapering from mid region; e.g., H. Smith

10401 [BM], Plate 6 in Florin 19483), I see no reason to disagree with Florin—that the Warburg type for

Cephalotaxus celebica lacks papillae along the abaxial midrib; thus, while Cheng and Fu (1978) could have

adopted this name in their revised treatment of Taxaceae in the Flora of China, the correct name under their

species concept is T. sumatrana; the basionym (Cephalotaxus sumatrana) was mentioned by Parlotore (1868)

and Pilger (1903,), the latter of which was cited by Cheng and Fu (1978).

7. Taxus obscura Spjut, OO (Fig. 23). Tre: THE PHILIPPINES. Luzon: Mt. Banahao, 2100 m, tree 8 m, common on ridge

in mossy forest, aril bright red, 26 Aug 1978, de Laubenfels P668 (HoLortrrE, A! [Spjut in adnot., T. phytonii var. obscura in ed.]; with

seed; leaf with 11 stomata rows/band, abaxial marginal border of 4 smooth cells across in 2 quadrate rows and 2 inflated rows, trap-

ezoidal in shape, followed by 5-6 rows of papillose cells; abaxil leaf midrib 15 cells wide, papillose on most cells, papillae covering

the entire cell surface). De Laubenfels prepared a separate specimen from a male plant from the same gathering, P-669. Type also

illustrated in de Laubenfels (1988).

Species haec ab Taxus chinensis, differt foliis linearibus, valdes discoloribus, 1-2 (C3) cm longis, 1.5-2.5 mm latis, acutibus, 0.200-0.300

mmi crassis, supra convexa, subtus concava ad plana, stomata vitta ferriginea, (1011—13(-15) seriata/zonas.

Tree 5-15 m high; ultimate branchlets crowded, long, weeping or flexuous, yellowish green, dull rusty brown

to orange with age; bud-scales scarcely persistent at base of branchlets, the scale scars similar to those of

T. chinensis, longest persistent scales ca. 1 mm long (to 2 mm long in Loher 7129). Leaves often lost by the

3rd yr, spreading obliquely and closely overlapping in + two ranks, frequently crisscrossing in herbarium

236 Journal of the Botanical R h Institute of Texas 1(1)

specimens, especially near ends of branchlets, oblong, or slightly elliptical, recurved, evenly tapered to an

acuminate sharply pointed apex, abruptly bent downwards to apex, 1-2(3) cm long, 1.5-2.5 mm wide,

200-300 um thick, dark olive green, resinous glossy and convex across adaxial surface to a recessed acute

darker midrib with channels along each basal side, rugose in dried specimens; abaxial surface slightly con-

cave across to a slightly elevated midrib, the midrib rounded to truncate, flush or slightly elevated, plane to

abruptly revolute near margins; upper adaxial epidermal cells in transverse section broad elliptical to nearly

short rectangular, thin-walled, (12—)20-—25(-30) um tall, 25-30 (-40) um wide, abaxial margin 2-4(-11)

cells across without papillae, papillose across 1-7 cells; marginal and midrib cells not as tall as those on

adaxial surface, 12-15 um tall, 15 pm wide, the epapillose marginal cells short trapezoidal to nearly quadrate,

usually inflated in 2-4 rows, + rectangular nearer stomata band, narrower and longer on midrib (3-7 l/w),

usually entirely papillose across the midrib, rarely only partly papillose on midrib, or rarely lacking midrib

papillae in lower half of leaf, papillae opposite or more often alternate in 13 irregular rows across each cell,

covering most of the cell; stomata bands abruptly differentiated by color from surrounding epidermal cells,

yellowish orange or yellowish green, broader than the maginal region; stomata diamphicyclic, anticlinal

in orientation, in (10O—)11-13(-15) continuous rows per band. Male cones subglobose in bud, 4 mm long,

scales imbricate, overlapping in 4 ranks, microsporophylls lobulate; microsporangia +8, pale pink with a

broad dark reddish center. Female cones subcylindric, 2 mm long in bud; scales overlapping in 5 ranks,

greenish, longest scale at base conduplicate; seed rounded, conical, 5-6 mm long, 4-5 mm diam., tapering

to apex from the middle.

Common name.— Obscure yew.

Distribution.—Forest margins, 2000-2450 m. Myanmar, China (Fujian, Taiwan), The Philippines

(Luzon), Indonesia (Sumatera, Sulawesi).

Additional specimens: MYANMAR (BURMA). Ruby Mines, 6500 ft, Oliver (K); Ruby Mines, no elevation data, 14 Sep 1894, Oliver (K).

CHINA. Fujian: Puchen, Chung 3866 (A). THE PHILIPPINES. Luzon: Mt. Pulog, 18°26’ N, 120°54’ E, tree 12 m, leaves dark glossy

green above, dull green beneath, Jacobs 7171 (K); Benquet Prov., Laguna, Mt. Santo Tomas, Leaño 25128 (A, US); Mt. Tonglon, Aug 1906,

Curran s.n. (US), Curran 5015 (P, PH); Benquet Prov., Merrill 839 (U, US); Mt. Pauai, 2450 m; Sulit 7582 (A); Central Luzon, Loher s.n. (US);

Mangitquiran, Loher 4850 (K, US); Laguna Prov., Mt. Banahao, 2100 m, Loher 7129 (US), 7139 (US); Luconia, summit of Mt Majayjay, 7500

ft, ex Hook. Herb. (K); Mt. Majayjay, Wilkes Exped. 1838-1842 (GH); Lepanto Dist., Mt. Data, Ramos & Edaño 40234 (K, P). Locality data

uncertain: Y. Sugilara, Ex TUS (GH). INDONESIA. Sulawesi: Celebes an Ond., Gowa Lambaja, 2000 m, Neth. Ind. For. Serv. bb:20887

(K). Sumatera: Tharolanden, 1400 ft, Boschproephakion 7709 (U).

Taxus obscura is recognized by the pendulous branchlets and by the oblong leaves that in herbarium speci-

mens show a sharp contrast in color between the adaxial and abaxial surfaces—dark glossy green above

and yellowish orange (rusty) below (dried specimens). The weeping aspect of the branchlets is evident in

herbarium specimens by their flexuous appearance.

Occasional specimens, e.g., Loher 7129 (US) from the Philippines and Lobb 461 (BM) from Malaya, have

rigid branchlets. They resemble T. kingstonii but are referred to the Chinensis Subgroup by the narrow leaves

with fusiform shaped epidermal cells on the abaxial midrib and irregularly shaped quadrate and inflated

epidermal cells along the abaxial marginal zone. Loher 7129 is assigned to T. obscura by the overlapping leaf

arrangment with leaves frequently crisscrossing, but still differs from the majority of specimens assigned to

this species by its rather long linear leaves, whereas Lobb 461 with its more parallel two-ranked leaf arrange-

ment is placed under T. phytonii, a determination that is further supported by the reddish colored epidermal

cells on the abaxial midrib. Both specimens may be hybrids between T. sumatrana and T. phytonii, or may

prove to be distinct varieties in further study.

In my 1996 annotations, a number of other specimens from Fujian, the Philippines and Indonesia

were identified T. kingstonii by the rusty orange colored leaves with relatively thick lip-like margins (e.g.,

Sulit 7582 [A], Curran 7911 [US] and Loher 4850 [US]). Taxus obscura was then distinguished as a variety of T.

phytonii by the lack of marginal papillae on the abaxial midrib. In further study, more taxonomic emphasis

was placed on branching and phyllotaxy, less on position of epidermal papillae, in distinguishing T. obscura

from T. phytonii; juently, other differences became apparent between these species. As a result the

Spjut, Taxonomy of Taxus 237

taxonomy of T. obscura broadened to include more specimens from the Philippines and Indonesia, while

that of T. kingstonii was narrowed to exclude all specimens from these areas. The specimen from Fujian also

differs by the leaves having a wider marginal zone of epiderma cells, 11 cells across, accompanied by lack

of papillae on outer midrib cells; otherwise, it agrees with T. obscura.

Taxus obscura differs from T. phytonii not only in branching and phyllotaxy but also by leaf length and

color and by the scales. The branchlets in Taxus phytonii spread more widely and scarcely overlap in pressed

specimens, and the leaves are more distinctly linear and lie closely parallel along one side of a branchlet

in pressed specimens, in contrast to a more crisscrossed crowded arrangement in T. obscura. The stomata

bands of T. phytonii show a sharper contrast in color from the marginal cells, appearing yellowish green

compared to the reddish margins and midrib, whereas stomata bands of Taxus obscura show less of a contrast

in color from the adjacent midrib and marginal zones, usually the stomata bands appear yellowish orange,

occasionally entirely green. The bud-scales of T. obscura are smaller and less persistent at base of branchlets,

and cone scales are mostly imbricate, in contrast to scales of T. phytonii appearing more conspicuous at base

of branchlets and decussate on cones.

8. Taxus phytonii Spjut, sp. nov. (Fig. 24). Tre: CHINA. Taiwan: mts. W of Karenko, 23 Nov 1918, Wilson 11154 (HOLOTYPE: Al:

ISOTYPE: US! Leaves with 12-13 stomata rows/band, abaxial marginal border of 6 smooth cells across with + 2 rows of quadrate cells

across and 4 rows of inflated trapezoidal cells, followed by a region of 10 papillose cells wide, the papillae mostly along cell walls).

Similis a T. obscura, praecipue differt folia longiora, linearis, 2-3 cm longis, complanatus, stomata zonatas flavovirescens, margines et

costa rufus, perulae persistens.

Similar to T. obscura, tree 5-20 m; ultimate branchlets crowded, weeping or horizontal, yellowish green,

dull rusty brown with age; bud-scales mostly persistent in 3-4 ranks, spreading, thick, ovate to lanceolate

or cuspidate, concave, some smooth, others carinate, often pale yellowish or brownish, longest scales ca. 1

mm long. Leaves flaccid, two-ranked, not overlapping to slightly overlapping, linear falcate, slightly recurved,

evenly tapered to an acuminate sharply pointed apex, 2-3 cm long, 1.5-2.5 mm wide, 200-300 pm thick;

dark olive green and strongly convex across the adaxial surface to a depressed acute midrib that is channeled

along base, mostly yellowish green to yellowish orange and slightly concave across the abaxial surface with

a slightly elevated and rounded midrib, curved more near margins, or margins plane; adaxial epidermal

cells in transverse section elliptical to nearly wide rectangular, thin-walled, 12-20 um tall, 25-30 (240) um

wide; abaxial marginal and midrib cells similar, or smaller, 12-15 ym tall, 15 um wide, the marginal zone

usually 9-13 cells across, usually papillose to about 4 cells from margins, papillose entirely in one speci-

men from India, or lacking in papillae across 8 cells in Alvarey 18369 from Luzon, the epapillose cells short

trapezoidal to nearly quadrate, usually inflated in 2-4 rows, rectangular nearer stomata band, narrower and

longer on midrib, 3-7 times longer than wide on midrib, midrib papillose entirely or lacking in papillae

from near the mid region to base of leaf, papillae strongly marginal in 1—3 irregular rows across each cell,

often not covering the entire cell; stomata bands abruptly differentiated from surrounding epidermal cells

by color, yellowish orange or yellowish green, broader than the marginal region; stomata diamphicyclic,

anticlinal in orientation, 10—12(-14) rows/band, the abaxial margins and midrib usually reddish in herbarium

specimens. Male cones subglobose in bud, 4 mm long, scales overlapping in 4 ranks, sporophylls lobulate,

microsporangia +8, pale pink with a broad dark reddish resinous center. Female cones subcylindric, and

2 mm long in bud; scales overlapping in 5 ranks, +decussate, greenish, longest scale at base conduplicate;

seed rounded, conical, 5-6 mm long, 4-5 mm diam., tapering to apex from the middle.

Common name.—Phyton yew.

Distribution —2000—2800 m, Nepal, NE India, Thailand, China (Yunnan, Taiwan), Philippines (Luzon).

Additional specimens: NEPAL. Williams 1014 (BM). INDIA. Assam: Pachaksihri, Laluma, 94°15’E, 27?45'N, 7000 ft, Ludlow & Sherriff

3719 (BM). THAILAND. Malaya: Lobb 461 (BM). CHINA. Yunnan: Wei-se Hsien 2800 m, tree 30 ft in forest Tsai 59874 (A). Taiwan.

Tongshi, C-j. Chang: Tongshi 1, 5 (p.p., Rt. 210/6k), 7 (p.p., Rt. 210/16k), TD-1 (wba). THE PHILIPPINES. Luzon: Mt. Banahao, Ocampo

27920 (A, P); Curran 7911 (US); Mt. Santo Tomas, Elmer 6244 (P, US); Benguet Prov., Alvarey 18369 (BM). Locality not clear—handwriting

not legible, possibly Mt. Harui, Harain Letty, 17 Oct 1802, in adnot. Taxus tosua topue, p.p. (BM, GH).

238 Journal of the Botanical R h Institute of Texas 1(1)

Taxus phytonii differs from T. obscura by the relatively longer leaves, 2-3 cm long, that spread mostly par-

allel as seen along one side of a branchlet, and by other features as discussed under T. obscura. The leaf

characteristics of T. phytonii are remarkably consistent at disjunct locations. For example, Tsai 59874 from

Yunnan and Wilson 11154 from Taiwan appear almost identical not only in their leaf shape but also in

their leaf color and in their leaf anatomy in which papillae are positioned marginally on midrib cells and

in which stomata occur in 11-12 rows per band. Ludlow & Sherriff 3719 from northeastern India also has

these features, while differing slightly in papillae extending entirely across the abaxial leaf surface, instead

of just 2-4 cells from the margin.

Taxus phytonii shows affinity to T. wallichiana by the relatively long narrow leaves that spread nearly

parallel to each other, and also to T. chinensis by the yellowish tinted branchlets and by leaves that have broad

elliptical epidermal cells as seen in transverse section. Additionally, the branchlet color and phyllotaxy of

T. phytonii are similar to that of the North American T. globosa, which differs by longer leaf epidermal cells

near the abaxial margins, also appearing quadrangular in transverse sections.

Etymology.—The epithet phytonii is in tribute to an achievement by a company named Phyton (formerly at

Ithaca, NY, now in New Jersey) for the commercial production of the anticancer drug taxol by tissue culture;

their production facility is located in Germany. Their commercial methodolgy hopefully will eliminate the

harvest of wild plants to obtain the taxoids.

9-11. Taxus spp. undescribed, aff. T. chinensis. The following proposed species need further study.

9—Taxus REH (T. rehderiana ined); Vietnam, China (Taiwan), Indonesia (Sulawesi).

Taxus “REH” (Taxus aff. chinensis, Spjut in adnot. GH, P), recognized from five specimens, differs from the

preceding two species (T. obscura, T. phytonii) by the longer leaves showing less of a contrast in color between

the two surfaces, particularly in the paler green color on the adaxial surface, and also by the abaxial leaf

surface having a wider marginal zone (smooth cells) except for two specimens from Vietnam that have a

relatively narrow leaf margin (4 cells wide). Generally, the specimens resemble T. wallichiana in the long

linear leaves but have more features in common with T. chinensis as seen in the pale yellowish green color of

the branchlets, the minute scales at the base of branchlets, the elliptical shape of the leaf epidermal cells in

transverse section, and in the tapered seeds. In my 1996-1997 annotations, the specimens from Vietnam

were identified as Taxus aff. chinensis, while those from Sulawesi and Taiwan were referred to T. phytonii.

Additionally, a specimen by Purdom s.n. (GH) from northern China: (Shaanxi: Tai-pei-shan fide Rehder &

Wilson in Sargent 1914) is similar in the gross features of branching and color, and also in the leaf anatomi-

cal feature of marginal papillae on the abaxial midrib epidermis; however, it is referred to T. biternata by

the relatively fewer stomata rows (7 rows per band), and by the persistent dark spreading scales at the base

of branchlets. A specimen from the Philippines, Loher 7129, as discussed earlier under T. obscura, may also

belong here.

10-11. Taxus spp. aff. chinensis. 10—Taxus “OCR” (T. ocreata ined. Spjut in adnot., A, BM); China

(Yunnan, Sichuan), rocks, 1500 m. 11—Taxus “SCU” (T. scutata ined. Spjut in adnot., A, BM) China (Yun-

nan, Sichuan, W Hubei), 1200-1400 m.

These proposed species are similar to T. chinensis except for conspicuous persistent bud-scales at base of

branchlets, in which they differ from one another by the bud-scale characteristics. Taxus SCU has persistent

scales, numbering 10-20 or more, loosely attached with a shape like overgrown toenails that are ready to

fall off. The scales in Taxus OCR, in contrast, tightly adhere to branchlets and look more like teeth.

Taxus OCR shows remarkable similarity at two disjunct locations, Cheng 2890 from Sichuan and Feng

11937 from Yunnan, but there are also intermediates in duplicates of Cheng 2890. It is not clear, however,

whether these came from the same plant. Additionally, Feng also collected at the same site in Yunnan on

two occasions where his first collection (Feng 11937) was from a tree and his second (Feng 12105), just nine

days later, was from a shrub that was also indicated to be common. Collectors do not always pick from the

Spjut, Taxonomy of Taxus 239

same plant when assigning collections to the same number. It is also apparent from genetic studies in other

species of Taxus that unique genotypes exist within populations (El-Kassaby & Yanchuk 1995).

Taxus SCU is also similar to T. wallichiana that differs by the fewer scales and longer leaves; however,

occasional specimens from the Himalayas and China also appear intermediate by the presence of fewer

scales and shorter leaves, or by longer leaves and more scales.

The combination of characters for Taxus OCR and SCU appear, on one hand, worthy of taxonomic

distinction, and especially necessary for taxonomic clarification of related species—T. wallichiana and T.

chinensis, while on the other hand, it would seem prudent that field studies be conducted to evaluate varia-

tion in the bud-scale character features at a particular location.

Il. SUMATRANA GROUP

The Sumatrana Group is characterized by leaves having a relatively broad region of marginal cells adjacent

to the stomata bands, usually from 8-36 cells across. This is often evident in dried specimens by a glossy

reddish discoloration. The epidermal (accessory) cells in stomata bands are further differentiated from

marginal and midrib regions by their shorter length and broader width. Leaves in the other species groups

usually have stomata bands bordered by fewer cells, or they are less differentiated by color and development

of papillae. The Sumatrana Group (Spjut 1998b, 20000) is generally found at lower elevations on mainland

Asia, below 1700 m (Hu 1964), or below 1200 m (Li & Fu 1997), than T. wallichiana, which usually occurs

above 2300 m.

The taxonomy of this Sumatrana Group is difficult due to overlapping character traits among the species

and varieties that follow. Previous taxonomists have recognized only one taxon in this group, either as a

variety (Cheng & Fu 1975, 1978; Li & Fu 1997) or as a species (Florin 1948a; Handel Manzzetti 1929; Hu

1964), but they have not applied the correct name. Taxus sumatrana has been the name applied to all yews

in southwest Asia (de Laubenfels 1978), but this is antedated by T. wallichiana. This very broad concept of

T. wallichiana leads to difficulty in distinguishing between T. cuspidata and its allies in the Baccata Group.

Farjon (1998, 2001) indicated that T. sumatrana occurs only in Indonesia and the Philippines, but his inter-

pretation, as also stated elsewhere, is without taxonomic support.

Thus, I have found it necessary to recognize more than one species in the Sumatrana Group—based

on differences in the leaves. Within the Sumatrana Group, I distinguish T. mairei by the narrow elliptic to

oblong leaves tapering to an acute to obtuse apex and by the raised midrib on the abaxial surface, appearing

truncated to channeled with relatively short trapezoidal, somewhat inflated (mammillose) epidermal cells

(Spjut 1998b; and as indicated in crude illustrations that accompanied my annotations: A, GH in June 1996;

BM in April 2005). Leaves of T. sumatrana differ by the linear to lanceolate shape tapering to an acuminate

apex and by the nearly rectangular shape of abaxial epidermal cells. Taxus mairei var. speciosa differs from

typical T. mairei in features of rigidity of branches and leaves, branching pattern, phyllotaxy, and color (in

dried specimens), but differences overlap in many of the character attributes. One new species, T. hingstonii,

is recognized by its slightly larger persistent bud-scales at base of branchlets, and by leaves that are rigid,

evenly tapered to base and apex, and by the rusty orange color in the herbarium, especially along the stomata

bands. Differences in seed shape and color are also evident among these taxa, but field studies are needed

to better evaluate the taxonomy of seed characteristics

Although the width of the abaxial leaf marginal zone varies in Taxus, this, nevertheless, appears to have

partial correlation to species within the Sumatrana Group. For example, among 15 specimens cited for T.

celebica (Spjut 2007, Fig. 6), which has the widest leaf margin (that borders the stomata band) of any species

of Taxus, 11 specimens were found to have 27 or more marginal cells. Two specimens with 22 and 24 bare

cells across the abaxial leaf margin are within the range commonly found for T. mairei, while two others

with a narrower marginal border fall within the range of T. kingstonii. Whether more taxonomic emphasis

should be placed on the number of cells across the abaxial marginal region needs further study.

A wide range in variation for shape and length of epidermal cells has been observed in juvenile foliage

240 Journal of the Botanical R h Institute of Texas 1(1)

of T. mairei, whereas the mature leaves are distinguished by the inflated, short wedge-shaped epidermal

cells (13x l/w) on the abaxial midrib. Juvenile and adult foliage may differ in conifers such as Podocarpus,

in which species cannot be differentiated by their juvenile foliage (de Laubenfels 1969). Leaf shape and

development of papillae in Taxus may also vary with exposure on the same plant as noted for plants in

Taiwan and Indonesia (de Laubenfels 1978, 1988). Differences in mature and juvenile leaves of T. mairei are

noted in this study in which mature leaves are oblong and taper to an acute apex, while immature leaves

may appear elliptical and acuminate to apex.

12. Taxus celebica (Warb.) H.L. Li, Woody Fl. Taiwan 34. 1963. (Fig. 25). Cephalotaxus celebica Warburg, Monsun.

1:194. 1900. Tyre: INDONESIA. Sulawesi (CELEBES): southern, Gipfel des Wawo-Kraeng [one of the summits of G. Bonthain], on the

forest-clad summit, 2800 m [Nov 1888], WarBuRG 16889 (HOLOTYPE: S photocopy!; IsotyPE: B-destroyed). Only one specimen was cited

by Warburg, although he did not ifically indicate that it was the type. The S specimen is a fragment of the original EN at

B. Warburg distinguished his C. celebica from C sumatrana Miq. and excluded Podocarpus celebicus Hemsl.

Podocarpus celebicus Hemsley, Kew Bull. 39. 1896. Type: INDONESIA. SuLawesi (SOUTH CELEBES): Bonthain Peak, 7000-10,000 ft, Sep

895 A.H. Everett 35 (HoLotyee: K!).

Common name.—Celebes Yew.

Distribution and ecology.—Nepal, Bhutan, NE India, South Vietnam, China (Tibet, Yunnan, Sichuan),

Indonesia (Sulawesi); forest margins below 1530-3100

Taxus celebica is recognized by the relatively large, glossy, often pale green, lanceolate plane leaves that

taper to an acuminate apex and by the narrow stomata bands relative to a broad marginal zone of long (>10x

I/w) rectangular cells, (18—)24-36 cells across. The stomata rows generally number (8-)12-14 per band,

compared to (11214-19(-21) rows in T. mairei. Variation in number of marginal cells (Spjut 2007) appears

partially correlated to the region of the leaf where sections are obtained as the leaf blade is strongly tapered

from base to apex.

Leaf stomata bands of T. celebica often appear distinct from other species in the genus by the short

wedged-shape («3x l/w) accessory cells that have relatively small (minute) medial papillae, appearing more

distantly spaced from one another, in contrast to papillae covering the entire surface of the epidermal cells

in other species. Examples are specimens from Yunnan (Forrest 7798), Sichuan (H. Smith 10401, Plate 6 in

Florin 19482), South Vietnam (Schmind s.n.), Bhutan (Cooper & Bulley 2833), Khasia (Clarke 38308), and Tibet

(Kingdon Ward 19324).

Intermediates between T. celebica and T. mairei var. speciosa, however, are apparent. My 1996-1997

annotations of T. celebica were based on leaves having most of the following taxonomic features: long linear

to lanceolate shape, acuminate apex, long epidermal cells, obscure palisade layer of parenchyma cells,

and appearing flattened in T-section. Recently, it was decided that taxonomic weight should be given to

the lanceolate shape of the leaf tapering to an acuminate apex (Spjut 2007). Examples that were formerly

annotated T. celebica—but subsequently referred to T. mairei var. speciosa—are from Fujian (Price 1258b),

Guizhou (Steward et al. 328), Sichuan (Wang 20541), Guangdong (Nanling Expedition 1838), and Ningxia

Huizu (Chao 1223). Other plants of var. speciosa with similar but more distant phyllotaxy are cultivated at

the Royal Botanic Gardens at Kew and at Edinburgh (photos in Krüssmann 1985, plate 133; van Gelderen

and van Hoey Smith 1996), and juvenile leaves received from Phyton Inc. (as T. chinensis). The cultivated

plants were reportedly grown from seed of Wilson 1265. It is interesting to note that herbarium specimens of

the wild plants collected in China and of a plant cultivated at the Kew Gardens appear very similar (www.

worldbotanical.com).

13. Taxus kingstonii Spjut, sp. nov. (Figs. 26-27). Tv: CHINA. Taiwan: Arisan Prov., Kagi, 2833 m [Mt. Alishan], tree 25

ft x 2 ft, only one seen, 2 Feb 1918, Wilson 9738 (HoLotvrE: A! [with male cones; leaves with 10-11 stomata rows per band, abaxial

margins of 8 smooth, thick-walled, trapezoidal, anticlinal to periclinal arranged cells across, followed by 5 rows of papillose cells,

midribs 12 cells wide, mostly smooth, papillose on outer 2 rows]; isotypes: BM! [leaves with 13 stomata rows and 9 marginal smooth

cells, midribs mostly smooth except upper third of leaf, papillose on outer 2 rows of cells], K!, US p.p.!).

Similis a T. sumatrana et affnibus, praecipue differt folias coloris ferruginibus vel calendulibus; folia obliquata disposita, rigida, oblonga

Spjut, Taxonomy of Taxus 241

Fic. 25. Taxus celebica, distinguished by the relatively flat leaf with a broad abaxial marginal zone and narrow stomata bands; Indonesia, Sulawesi,

Everett 35 (holotype, Podocarpus celebicus, K).

2 PA fis 1 ded 1 151 +4 Tas] : PA | : : 1; : 1 +11

au ciipuca

proparte papillosa versas de marginali, sine papillae (7-)8-12(-20) cellulae marginales latis; stomata 11-15 seriata/zona.

Shrub or tree to 12 m high, bole to 65 cm diam; branchlets unequally (type) to equally divided, yellowish

green and gradually becoming yellowish with tint of either red, orange or brown; bud scales mostly persis-

tent, 3—4 seriate, turgid, ovate, concave, carinate near apex on upper scales, tan to chestnut brown, lower

bud-scales ca. 1.5 mm long. Leaves rigid, spreading usually at less than right angles to branchlets, not parallel to one

another, more evenly tapered to base and apex than in T. mairei, lanceolate, narrowly elliptical to oblong (type),

or linear in other morphs, evenly tapered to an acute, sharply pointed apex, recurved and/or twisted downwards,

1.5-2.5 cm long, 3.0-3.5 mm wide, 0.35-0.50 mm thick, dark green and convex above to rounded midrib,

the adaxial midrib somewhat acute in lower half, often not evident near apex, paler green to yellowish

green and convex below, or plane to concave below to rounded or flush midrib, often with orange tint in the

herbarium, or dull rusty brown in the herbarium, thickened and liplike near margins, or plane and slightly

revolute near margins; adaxial epidermal cells in T-sect. wider than tall, nearly wide rectangular, or ellipti-

cal in T-sect., usually 20-25 um tall and 25-30 um wide, thin-walled, slightly inflated; abaxial epidermal

cells similar in T-sect. but not as large, 15-25 um tall, 20-30 um wide, slightly inflated near margin in 2-5

rows, more nearly rectangular in 7-11 cell rows near stomata bands, usually relatively short on midrib,

1-4(-10)x l/w, sharply 4-6 angled, often wider and more slanted at one end (trapezoidal), not inflated (in

T-sect) as in T. mairei, papillose to ca. (72)8—12(-20) rows of cells from margins, typically without papillae

on midrib (except young leaves), or partially papillose on outer midrib; papillae submarginal to medial in

2-3 irregular rows across each cell. Stomata bands broader than the non-stomatal region, olive green in

fresh material; yellowish orange in dried leaves; stomata continuous in 11-15 rows, separated by 1-2 rows

of accessory cells, stoma often with a blackish halo. Male cones globose, ca. 4 mm diam, yellowish green,

scales generally 4-seriate; pollen sacs mostly 6, pale pink with reddish mid region and patchy resinous areas.

titute of Texas 1(1)

laf tha Dat

JOUIdI OF

242

i y 1 H 113 1s I? FA r A â “~ 0

; | bli dt | | | | 5 km

| Y | | e B23

| P y] EBENEN le^ = =

a 4 4-3 ? E

2 = ©

A, >52

z r-

v A

a =

¿E ]

HESS

m too

^ S 53$

w Orn oO

<= QN =

Ra r4

zog. S

C S = 2

HERE:

" Me OS

\ ~ 2s

i sixes

Ses

* -sE¢s

(d =~ =

e Vs 55

b = ED:

4 [star]

d d £=

u i QU I

A ===

2 "2 SS

lg ZZ,

Spjut, Taxonomy of Taxus 243

Female cone in bud subcylindric, ca. 2 mm long, greenish, scales mostly 5-seriate, conduplicate at base,

maturing on 1* yr branchlets; seeds ovoid, dull, tan or purplish, stained by aril, angular where tapering to

apex, or not angled, to 7 mm long, 4 mm wide.

Common name.—Kingston yew.

Distribution and ecology.—India (Khasi Hills), Myanmar, China (Tibet, Gansu, Shaanxi, Sichuan, Yun-

nan, Taiwan); forest margins or summits, 2450-2833 m in Taiwan, to 800 m on mainland Asia.

Additional imens: INDI A. Khasia: Hooker L77 (PH), 1337 (K); Simmons 484 (P), Nungluai, 5000 ft, Mann (K, P). MYANMAR (Burma):

T Ruby Mines, 5600 ft (K). CHINA. Tibet: Oriental, Haut Mekong, Tsekou to Nekou, Soulie1411 (P: 2 sheets). Gansu: F.

N. Meyer 1790 ex USDA (P). Shaanxi (Shensi): Tsin-lin au Lao-lin, 3000 m, year 1872 Davis s.n. (P). Sichuan: Tachienlu Cheng 1001

(BM), Cheng 1475 (P). Yunnan: Shweli-Salween Divide, 10,000 ft, 25°20 N, shrub 10-20 ft, open shady thickets, Forrest 11789 (BM, K),

Salween, Forrest 12087 (K, S: C-2093), Forrest s.n. (A); Ma-Chang-Kai, valley, 25°30 N, 6000 ft., shrub, 20-30 ft., in thickets, Dec 1918,

Forrest 9462 (A, K), Forrest 15945 (BM, K); Salween E of Tengyueh, to summit of Shwell, Shwelli River, Rock 7587 (US). Taiwan: Paseian

San [Pahsienshan], Hsi 165 (PH); Liu et al. 437, (M, T, US; Fig. 3 in Li, Woody Fl. Taiwan,); Mt. Ammachan, Liu 0389 (A, K), Arizan, Nitak

(PH); Mt. Ammashan, Taichung Hsien, C.C. Tseng s.n. (BH); C-j. Chang without locality data, Tongshi #6, 4 Mar 1993 (wha).

Taxus kingstonii is recognized by its relatively turgid recurved leaves (twist and curve downwards along their

blades) that taper rather evenly to apex, and by their rusty orange color on the abaxial leaf surface (dried

specimens). In herbarium specimens, the leaves often crisscross, especially near apex of branchlets. In other

species of the Sumatrana Group, leaves are more reddish or greenish in color and evenly two-ranked and

tightly adpressed to branchlets. The Kingston yew often has tightly adhering, strongly nerved bud-scales at

the base of young branchlets, whereas scales in related species are similar to those of T. chinensis—generally

smaller, less imbricate, or often not evident. The Kingston yew generally occurs at elevations between that

of T. mairei (below 1200 m) and T. wallichiana (above 2300 m).

The Kingston yew tends to have the narrowest abaxial leaf margin among species of the Sumatrana

Group, while this does not appear to be offset by wider stomata bands as most specimens were found to

have 12-13 stomata rows per band (Spjut 2007). The width of the marginal region shows two patterns, one

occurring around (6-)8-9 cells wide as seen by plants from Myanmar and Taiwan, and another ranging

from (10—)12—16(-20) cells wide as seen in plants from Yunnan, Gansu, and Sichuan. This is in contrast to

leaves of Taxus mairei that show a wider range in variation in which most specimens havel4 stomata rows

per band and 14—24 marginal cells without papillae. This indicates a higher density of stomata in leaves of

T. mairei. Variation in leaf anatomical data for T. kingstonii may be due to hybridization with T. wallichiana

in northeastern India, with T. chinensis and T. mairei in Shaanzi, Gansu and Sichuan, and with T. celebica in

Yunnan.

Taxus kingstonii includes specimens from Yunnan, India, and Taiwan that have been considered a dis-

tinct species by their long narrow, wide spreading leaves tapering to an acuminate apex as recognized by

the epithet in regard to the acuminate leaf (Spjut unpublished, key to species of Taxus, USDA Memorandum

1995). They have the characteristic pale to rusty orange color on the abaxial leaf surface, but differ in the

broader marginal zone of trapezoidal to short rectangular epidermal cells, a feature that is seen more in T.

mairei. However, the abaxial midrib and marginal epidermal cells have thicker walls in contrast to bulging

(mammillose) cells of T. mairei; the epidermal cell walls of T. mairei are often so thin that they appear trans-

parent to the chloroplast of the spongy parenchyma cells. The character attribute of the acuminate leaf is

similar to that of T. celebica and T. sumatrana; the latter distinguished by dried leaves appearing blood red

along the midrib and marginal zones in sharp contrast to the yellowish green stomatal bands, the former by

larger and more flattened leaves with a pale green color on the adaxial surface. Some specimens from India

were annotated during 1996-1997 as T. celebica; however, in further study of a large number of specimens

obtained by Professor C-j Chang (Purdue University) from Taiwan, the concept of T. kingstonii was broadened

to include the specimens from northeastern India and Yunnan.

Despite the similarity of T. kingstonii to other species in the Sumatrana Group, it will probably be con-

fused most often with T. chinensis. A number of specimens at BM, K and P, which I annotated T. chinensis

(Oct 1997) at the time of my visit to these institutions—based on examination of leaves under a dissecting

f4L,D o ID

244 Journal of t h Institute of Texas 1(1)

scope—were later discovered to belong to T. kingstonii as a result of examining leaf sections under a compound

microscope and discovering that papillae were lacking along the abaxial midrib and marginal zones. These

specimens, including the type from Taiwan, appear closely related to T. chinensis by the leaf epiderimal cells

appearing larger on the adaxial surface than the abaxial surface as seen in transverse sections, and by the

abaxial leaf surface having a rounded keeled midrib.

Taxus kingstonii is named in honor of David G. I. Kingston, a chemist who has done extensive work on

elucidating and summarizing the taxane chemistry of the genus (e.g., Kingston 1996, 2005; Kingston et al.

1990). Taxol, from which the drug paclitaxel is marketed and used to treat ovarian and other cancers, was

originally isolated from dried bark of T. brevifolia, however, fresh leaves of many species of Taxus have proved

suitable for obtaining taxoids. Taxol was first characterized by Monroe Wall’s group (Research Triangle

Institute, Wani et al. 1971). They also discovered other significant antitumor agents such as camptothecin

and holacanthone (Wall et al. 1972).

14. Taxus mairei (Lemée € H. Lév.) S. Y. Hu ex T.S. Liu, Illustr. Native € Introd. Lign. Pl. Taiwan 1:16. 1960.

(Figs. 28-29). Tsuga mairei Lemée & H. Léveillé, Monde des-Pl. sér. 2, 16:20. 1914. Taxus chinensis Rehder var. mairei (Lemée

& H. Lév) WC. Cheng & L.K. Fu, Fl. Hupehensis 1:28. 1976 (also indicated as comb. nov. in Fl. Reipub. Pop. Sin. 7:443. 1978).

Taxus wallichiana Zucc. var. mairei (Lemée & H. Lév.) L.K. Fu & Nan Li, Novon 7:264. 1997. Tree: CHINA. YUNNAN: Dongchuan,

700—800 m, May 1912, Maire s.n. (HoLoTYPE [fide Rehder 1936]: E; isotypes: A [fragment from E]!, BM!, PD.

Rehder (1936) reportedly found a holotype for Tsuga mairei at the Royal Botanic Garden at Edinburgh (E);

however, it is not clear whether the Maire type at E was the only specimen *used" by the authors (ICBN Art.

9.1) since Lemée and Lévéille (in Léveillé 1914) did not cite specimens, and I have seen duplicates of Maire

specimens at the Museum of Natural History in London (BM) and at the Museum of Natural History in Paris

(P). The duplicate material at BM, although not from the Lévéille Herbarium, was also from E, presumably

distributed after they purchased the Léveillé Herbarium in 1919 (Stafleu & Cowan 1979). Additionally, a

fragment of the holotype of T. mairei is preserved in the Arnold Arboretum (A). The ICBN (Greuter et al.

2000) regards a fragment of a holotype as an isotype if the holotype was designated (Art. 8.3, Ex. 5), but

in my opinion a distinction such as merotype should be made because other duplicate specimens at other

institutions (isotypes) may have come from different plants.

Taxus mairei is distinguished from other species of the Sumatrana Group by the two ranked leaves

spreading at nearly right angles, by the abaxial surface of leaves having a truncated but elevated midrib

with epidermal cells larger in diameter than those on the adaxial surface (as seen in T-section), appearing

mammillose. The midrib is usually channeled, especially in the mid region of the leaf.

Taxus mairei, like T. chinensis, has a long history of nomenclatural confusion. As indicated previously,

the original authors thought they had a species of hemlock (Tsuga). Rehder (1936), upon discovering that

Tsuga mairei Lemée & H. Léveillé (Léveillé 1914) belonged to Taxus, treated it as a synonym of T. chinensis;

however, the ICBN (Art. 11.4) requires the earlier epithet, mairei, be adopted (S.Y. Hu, in Liu, Illustr. Nat.

Ind. Lign. Pl. Taiwan 16. 1960). Cheng and Fu (1978) also made the illegitimate combination—T. chinensis

var. mairei—which has since been transferred—T. wallichiana var. mairei (Lemée € H. Lév.) Fu € Li (in Li

& Fu 1997). It must be emphasized that *in no case does a name have priority outside the rank in which it

is published" (Art. 11.2); the epithet chinensis was employed at the varietal rank by Pilger (1903), and also

by Florin (19482). Rehder recognized T. chinensis as a species in 1919; whereas the epithet mairei was not

used for a variety, but as a species epithet by Lemée and Léveillé in 1914, five years earlier; therefore, mairei

has priority over chinensis in choice of epithet for a species, but has no priority whatsoever as a variety.

Regardless of the nomenclatural misapplications of the name, the taxonomic application in recent years

has been to treat yews in subtropical China under the epithet *mairei" if their leaves lack papillae on the

abaxial midrib (Fu et al. 1999). The species epithet sumatrana, an earlier available name, was not applied

because botanists in China had not seen its type (Cheng & Fu 1978; Hu 1964; Li & Fu 1997); instead, they

selected mairei for the varietal epithet—placing it under T. wallichiana.

Two varieties are recognized by differences in rigidity, branching, and phyllotaxy.

Spjut, Taxonomy of Taxus

14a. Taxus mairei var. mairei

Common name.—Maire yew.

Distribution and ecology.— China (Sichuan,

Yunnan, Anhui, Guizhou, Guangxi, Jiangxi,

Fujian, Hunan, Guangdong, Zhejiang, Taiwan);

forest margins, 300-1300 m.

Taxus mairei var. mairei is identified by zig-

zag branching, by the complanate leaves—ar-

ranged neatly in two-ranks in which the margins

of adjacent leaves along one side of the branchlet

are closely parallel—and by the mammillose

(enlarged) cells on the abaxial midrib. Leaves

also appear sessile; i.e., they are adpressed to the

branchlet at the base of the blade, and spread at

right angles as they bend and twist. Leaves of

var. speciosa are similar in shape but often larger

in size, more unequally spaced along one side of

a branchlet, differ only slightly in anatomy, but

often differ in color—by remaining green when

dried. Specimens of var. speciosa tend to have

less dichotomous branching, or a more distinct

monopodial branch.

The zigzag branching of T. mairei is perhaps

an ancestral trait. This branching is seen mostly

in southeastern China along coastal provinces

as far north and inland as Hunan. Yews further

north near Vladvostok, Russian Federation

(Palczevski 3601), South Korea (Wilson 9332),

and Japan (K. Muijabe 17 Sep 1910, A) show

other dichotomous branching treated under

T. umbraculifera var. microcarpa, which is also

recognized by crowded (overlapping) leaves

oriented in € decussate ranks. It may be noted

that leaves of T. umbraculifera, T. chinensis, and

T. kingstonii are relatively thick with a rounded

(keeled) midrib, in contrast to thinner leaves

in T. mairei that have a truncated to channeled

midrib. Taxus biternata, a common species in

temperate E Asia, easily recognized for its much-

divided branchlets, has a phyllotaxy that is more

similar to T. sumatrana than to other species of

the T. cuspidata Alliance.

Occasional specimens of T. mairei are rec-

ognized to have an abaxial leaf epidermis of the

T. chinensis type—as seen by the development of

papillae on the midrib and along the marginal

zone to ca. 8 cells from margin. Examples are

Ching 1676 (A, P) from Zhejiang at “2600 ft,” and

(os Adae. C. ESA A Y

Det. Shiu-ying Hu Vigo: )

AnNoLD Ar E H U

TYPE SPECIMENS IN HERB. LÉVENA

BOTANIC GARDEN, EDINBURGH

ALFRED REHDER. 19

ACTAS Plate No. 20 SO

HERBARIUM ™ ji

$ N 401032 /

c Y à

LD ar

] VERSA te? f 2«4 (2G D» f

VERS = “VAxnces Areca : qe Gy

- = sow MW

CSWIFITAH

t La. ]

let RR bet! 1:4 1«

Fics. 28-29. Taxus mairei var. mairei, distinguished by the isodichotomous

L Lt Jah I I Hall £ $ t

28 (top). N Yunnan, Maire

s.n., photo of specimen at E (holotype) with fragment at A (isolectotype). 29

(bottom). Maire 131 (BM).

f4L,D o ID

246 Journal of t h Institute of Texas 1(1)

Chiao & Fan 464 (A, P, US) from Sichuan at “200 m.” These specimens also lack the enlarged mammillose

midrib cells that characterize T. mairei, and the specimen from Zhejiang further differs in having rectangular

shaped epidermal cells in transverse section. These specimens may be hybrids, or perhaps they are relicts

of a T. florinii-T. chinensis complex from which T. mairei may have evolved. Leaves of T. engelhardtii from an

Oligocene deposit in Europe are remarkably similar to these specimens of T. mairei in leaf arrangement,

shape and anatomical details (Spjut 2007).

14b. Taxus mairei (Lemée & H. Lév.) S.Y. Hu ex TS. Liu var. speciosa (Florin) Spjut, comb. et stat. nov. (Fig.

30). Basionym: Taxus speciosa Florin, Act. Hort. Berg. 14(8):382. 1948. Type. CHINA. Guizaov: [Fanjing Mts.] Kiangkow, 450 m, in

light woods, tree 8 m high, fruit red, 8 Dec 1930, Y. Tsiang 7525 (HOLOTYPE: S; ISOTYPES: A!, BM!, K!, NY photocopy!, US!).

Common name.—Special yew.

Distribution and ecology.— China (S Shaanxi, Sichuan, NE Yunnan, Guizhou, N Guangxi, Hunan,

Guangdong, W Hubei, Jiangxi, Zhejiang, Fujian, Taiwan); forest margins, near streams or open areas on

hillsides, 100—750(71600) m.

Taxus mairei var. speciosa is recognized by having alternate branchlets along a common branch, and by

leaves appearing greenish and unequally spaced along branchlets in dried specimens. It appears to occur more

frequently in the interior provinces of China, in contrast to var. mairei being found more along coastal prov-

inces; however, exceptions include the type for T. mairei from N Yunnan and other specimens from Sichuan.

Taxus speciosa was described by Florin to distinguish a yew in China by the leaves having a smooth

midrib from one other in China that he recognized to have a papillose midrib, which he had determined to

represent T. wallichiana var. chinensis based on study of its type. The taxonomic feature of a smooth midrib

was also erroneously described for T. chinensis by Handel-Mazzetti (1929). Florin (19482) correctly realized

that T. chinensis was closely related to T. wallichiana even though he only mentioned the type for T. chinensis

and not T. wallichiana, although he had studied a duplicate specimen of the original material (Wallich 60544).

However, Florin incorrectly provided another name for the species that he distinguished by the absence of

leaf papillae on the abaxial midrib; he cited not only an earlier name—Cephalotaxus celebica, but also its

type (Warburg 16889). Other authorities soon realized this error (Cheng & Fu 1978), and Florin's name was

replaced by T. mairei S.Y. Hu (Liu 1960), who had also distinguished it from T. chinensis according to Florin

taxonomy as evident by her 1955 annotations of Harvard specimens (Hu 1964).

Notwithstanding, T. mairei was soon replaced by an earlier name T. celebica (Warb.) Li (1963), who

recognized only one species in China even though Hu (1964) indicated she adopted T. mairei because the

type for Cephalotaxus celebica Warb. might differ based on one specimen she saw from Sulawesi that had a

papillose midrib. She might have further assumed that the Berlin type of C. celebica.was destroyed; however,

as noted previously, Florin took a fragment of the Berlin type to Stockholm. It is surprising that an earlier

name mentioned by Pilger (1903), C. sumatrana, had been neglected, until finally applied by de Laubenfels

(1978). Although T. speciosa was founded upon nomenclatural error, it can still be recognized if its type can

be shown to belong to a distinct species; however, in this study, the epithet is retained for a distinct variety.

15. Taxus sumatrana (Miq.) de Laubenfels, Kalikasan, Philipp. J. Biol. 7:151. 1978. (Fig. 31). Cephalotaxus

sumatrana Miquel, Fl. Ind. Bat. 2:1076. 1859. Type: INDONESIA. Sumatera: western, Fort de Kock, 3000 m, without date, Teysmann

s.n. (HOLOTYPE: Ul).

Common name.—Sumatera yew.

Distribution and ecology.—E Nepal, India (Khasia) Thailand, China (Zhejiang, Taiwan), The Philippines,

and Indonesia (Sulawesi, Sumatera); mossy forests, 650-3000 m.

Taxus sumatrana is distinguished by the relatively thin (flaccid) leaves that taper to an acuminate apex

and pucker on drying, curling inward along margins, often drying dark shiny green above (adaxial surface)

and reddish green along margins and midrib below (abaxial surface). The abaxial surface has long rectangular

epidermal cells, but are shorter in about 4 rows nearest the margin, and often half of the marginal cells are

papillose towards the stomata band. The papillae develop in «several opposite rows on each cell in contrast

to alternately arranged papillae in the Chinensis Subgroup.

Spjut, Taxonomy of Taxus

I-— P (fie A

| (Ole tA cc VAR Adel

|. Det. Shiu-ying Hu à v j

| Anxorp Aasourtom, Hanvaan UsivensiTY C vt

i A MM MM 9

WUPRAAA REN

Ex Hermanns oF

Tue Mernoronrras Museum of NarumAG,

Lo s AcADEMIA Sinica. Nankina, Cros

: FLORA OF KwEICHOW

|. TAXUS CHIWSWSIS Rand.

^. | Kiemgkow, 459 meia Liga!

ie ES Bamson bt. Modo dio

247

Fic. 30. i irei speciosa, listi

hi g, Guizhou, China, Tsiang 7525, isotype (A)

£s+haD o ID L

248 Journal of t titute of Texas 1(1)

Leaves from two specimens, one from

Taiwan (de Laubenfels P670 & P671), and

another from the Philippines (de Laubenfels

P650), have relatively narrow marginal zones

without papillae. These specimens may key to

species in the Chinensis Subgroup, but they are

clearly related to T. sumatrana by the reddish

discoloration along the abaxial midrib and

marginal zones (as seen under low magni-

fication, 10x), and by the rectangular shape

of the abaxial epidermal cells and puckered

leaf blades, which are more strongly curled

à ho s. ` $ x i X , NI

, ¿SS E j NS NS inwards in upper third.

Three varieties of T. sumatrana are ap-

parent. They differ in leaf size and color. The

Fic. 31. Ti trana, distinguished by the puckered leaf tapering to an acu-

minate apex, and by the abaxial leaf surface having an elevated and truncated typical one—occurring in Indonesia, the

idrib that is darker in color (and also the marginal zones) than thestomata bands; ^ Philippines, Thailand, and Jiangsu—has rela-

Sumatra, Teysmann s.n (holotype, U). tively smaller leaves with a dark green adaxial

surface, and relatively long epidermal cells on

the abaxial midrib and marginal zones (+3-12x l/w). Another with larger paler green leaves, either with

shorter epidermal cells (+1-5x l/w), or with both longer and shorter cells, occurs in Nepal and Khasi, and

a third in Taiwan, Luzon, and Sumatera has dull olive colored leaves.

III. BACCATA GROUP

The Baccata Group is divided into two species alliances based on leaf anatomical features of the stomata

band as defined by its width and stomata density: (1) the Euro- Mediterranean Baccata Alliance and (2) the

E Asian Cuspidata Alliance.

Differences in stomata density are evident by the number of stomata rows in a band relative to the

absence of papillae on marginal cells. The Baccata Alliance has fewer stomata in slightly wider bands—at

least by 4 cells—in which the stomata bands and their adjacent papillose cells usually extend to 4 cells from

the margins, whereas the Cuspidata Alliance has a higher density of stomata in narrower bands, most often

bordered by a marginal zone of 8-18 smooth cells across.

Other differences in the leaves between these species alliances include the arrangement of accessory

cells, the prominence of papillae, and the number of the marginal papillose cells. In the Baccata Alliance,

the accessory cells are often fusiform with papillae usually covering most of the cell. Their arrangement is

diamphicyclic (Florin 1931) except for a narrow straight line of cells between stomata rows. A transitional

region of fusiform papillose cells extends to a marginal zone of shorter rectangular or trapezoidal cells that

lack papillae. Stomata bands in the Cuspidata Alliance, in contrast, usually have + pentangular to quadran-

gular or triangular accessory cells with concrescent papillae. Their stomata rows are not separated by a line

of cells; rather, stomata often share the same stomata pit.

A third type of stomata band is found in T. canadensis, which is included in the Cuspidata Alliance

even though it occurs in northeastern North America and the Euro-Mediterranean. Stomata bands of T.

canadensis are much narrower (Nicolosi 1982)—bordered by at least 11 epapillose marginal cells across. Its

stomata aperture and the irregular alignment of stomata rows are similar to T. baccata (Kvaéek 1984), while

the absence of papillae across a broad margin indicates a close relationship to T. cuspidata.

The distinction of these stomata band types may be correlated with molecular data. Collins et al. (2003)

recognized three distinct DNA chloroplast types based on specimens identified as T. baccata, T. cuspidata

and T. canadensis. They further suggested that these species probably evolved over a long period of time.

Spjut, Taxonomy of Taxus 249

These basic types of stomata bands (baccata, canadensis, cuspidata)—as measured by the width of

the band relative to the marginal region, 4, 8, and 12 cells in from the margin—are each postulated as an

ancestral type. Hybrids, however, are apparent not only between species of these band types, but also with

the Sumatrana Group as evidenced by the wider leaf margins—generally 16, 24 and 32 cells across. Various

combination of other leaf anatomical feature—position and size of papillae on cells, shape of chloroplast,

color of chloroplast and cell walls when dried, and shape and arrangement of parenchyma cells—also seem

best explained by hybridization. The anatomical features appear independently inherited of gross morpho-

logical features such as habit, branching, and phyllotaxy.

IIIA. Baccata Alliance.— Distribution and ecology—Europe, N Africa, SW Asia (Euro-Mediterranean) and

NW Himalayas; the Euro-Medierranean plants in various hardwood and mixed mesophytic forest types,

sea level to 500 m in the northern range, to 2000 m in S Europe, or to 2500 m in the Atlas Mountains of

N Africa. In hardwood forests with beech (Fagus orientalis Lipsky), basswood (Tilia spp.), elm (Ulmus spp.),

oak (Quercus spp.) and ash (Fraxinus spp.), or mixed hardwood-conifer with beech and spruce [Picea abies

(L.) Karsten] (Browicz & Zielinski 1982; Duffey 1980) and occasional black pine (Pinus heldreichii Christ;

Voliotis 1986). Around the Black Sea in beech-fir-spruce forests [Abies nordmanniana (Stev.) Spach, A. cilicica

(Ant. & Kotschy] Carr., Picea orientalis (L.) Link], or spruce absent (Browicz & Zielinski 1982), or in the

British Isles, forming single dominant woodlands with a closed canopy lacking understory shrubs and herbs

(Thomas & Polwart 2003).

Yew plants of the Baccata Alliance are characterized by falcate or recurved leaves that overlap or criss-

cross when branchlets are pressed, and by the abaxial surface of leaves having a transitional zone of papillose

epidermal cells between stomata bands and a narrow bare marginal border, usually 4-7 cells wide.

One species of Taxus has been generally recognized in the Euro-Mediterranean region, T. baccata;

however, leaf anatomical data clearly indicate two, T. canadensis and T. baccata, and additional species, T.

fastigiata and T. recurvata, and varieties are evident by other morphological differences. The species allied

to T. baccata and T. cuspidata are most distinct in Europe and eastern Asia, respectively, and least distinct

in the Caucasus Mountains. Taxus canadensis, which has been traditionally known from eastern North

America, is also recognized to occur in the Caucasus Mountains, Scandinavia, Estonia, and southwestern

Mediterranean.

Taxus baccata has included numerous varieties or forms (Beissner 1891; Carriére 1855a, 1867; Elwes

& Henry 1906; Gordon & Glendenning 1858; Gordon 1875; Knight 1850; Krüssmann 1985; Lawson et al.

1851; Loudon 1844; den Ouden & Boom 1965; Pilger 1903, 1916), recognized mostly from horticulture.

The more distinct taxa have been treated in Bailey (1923, 1933), Carrière (1855a, 1867), Elwes and Henry

(1906), Gordon and Glendenning (1858), Gordon (1875), and Pilger (1903, 1916). Prior to Beissner (1891)

and Pilger (1903), these were often considered varieties; Rehder (in Bailey 1902, 1923, 1933), for example,

initially treated them as varieties but later as *garden forms" (Rehder 1940, 1949). As evident in these pub-

lications, names for many yews in the horticultural trade were not clearly differentiated (taxonomically)

prior to 1855.

Those in Carrière (18552); therefore, are given priority in the present treatment except where earlier

legitimate names can be applied. Nothing is known about his types or the existence of his herbarium material

(Stafleu & Cowan 1976), and I would assume that his knowledge of yew taxonomy was based on descriptive

information in literature and study of living material at gardens in France. Authors who have later described

more taxa (e.g., Carrière 1867; Gordon & Glendenning 1858) have not always followed priority. And it should

be noted that although horticultural material may exist in gardens, valid names cannot be based on living

material; preserved specimens are required. Moreover, plants brought into cultivation from the wild must

retain the names that were applied to the wild specimens.

Many specimens collected in the wild compare favorably with varieties described during the 19" century,

while other varieties described later, as summarized in Chadwick and Keen (1976), Krüssmann (1985), and

den Ouden and Boom (1965), appear to be only minor variations in color and habit that probably originated

f4L,D ID L

250 Journal of tani titute of Texas 1(1)

from hybridization and selection of existing garden plants. Molecular data by Collins et al. (2003) indicate

that many cultivars are genetically distinct.

It is also conceivable that many introductions could have originated before the yew horticultural trade

developed—in the late 16'^ century (Mitchell 1974); the presence of Homo sapiens in the British Isles dates

back to 5500 yr. For instance, “the oldest wooden artifact” used by man in the British Isles is a yew spear,

which has been found in elephant remains 4600+ yr old (Godwin 1975), while in later times yews have

been planted and harvested for the construction of long bows (Hageneder 2007; Mitchell 1974; Thomas

& Polwart 2003). The existence of old yew trees in many churchyards (Loudon 1844; Lowe 1897) is also

evidence of a long history of yew associated with religious beliefs, and Tittensor (1980) has suggested that

yew woodlands in southern England originated from trees that marked “parish boundaries.”

In selecting neotypes, the available varietal names were considered based on priority of publication,

current usage, and information in the original description, but not all varietal and cultivar names are ac-

counted for in this treatment. This paper mentions those that appear more common and widely distributed.

Unfortunately, it has not been possible to evaluate their character features in the field, and to determine to

what extent yew populations may show polymorphisms and introgression, while it may be further noted

that much of the yew habitat has already been destroyed by human activities (Heinze 2004). Nevertheless,

molecular investigations are being undertaken to assess the genetic variability in various countries such as

the United Kingdom, Spain, Portugal, Austria, Germany, Slovakia, and Switzerland as evident from various

abstracts and reports on web sites.

16. Taxus baccata Linnaeus, Sp. Pl. 1040. 1753. (Figs. 32). Lecrorvr, designated by Jonsell & Jarvis in Jarvis et al. 1993:

Clifford Herb., 464, Taxus No. 1 (BM).

Of eleven varieties of Taxus baccata proposed by (Spjut unpublished, www.worldbotanical.com), five, in addi-

tion to the typical variety, are mentioned below. All are widely distributed in the Euro-Mediterranean region.

16a. Taxus baccata var. baccata

Common name.—European yew.

Distribution —Euro-Mediterranean.

Taxus baccata var. baccata is characterized by having equally divided horizontal to pendulous branchlets

with leaves + spreading in the same plane along opposite sides of a branchlet (two-ranked), the adjacent

leaves parallel to slightly overlapping or crisscrossing in dried specimens. In most other varieties leaves

curve upwards on horizontal branchlets. Taxus baccata var. washingtonii (Hort. ex Richard Smith) Beissner,

which is thought to have originated in horticulture, is possibly of hybrid origin with Taxus canadensis as

one of its parental types (Gordon 1875); however, a number of specimens from eastern Europe also appear

to be natural hybrids between T. canadensis in Europe and T. baccata.

16b. Taxus baccata L. var. dovastoniana Leighton, Fl. Shropshire 497. 1841. (Figs. 33-35). Tver: ENGLAND.

SALIK Co.: “Raised by John Dovaston in Westfelton near Shrewbury in 1777” (Loudon 1844), 1863, Westfelton ex Herb. Bidwell, anno-

tated “T. dovastonianum Leighton,” with seed (Neotyre designated here: BM! [leaves with stomata in 10 rows per band, abaxial marginal

border of 4 smooth cells]). Related material, BM: “original tree at Westfelton” communic. Jackson s.n., without seed, with galls [10

stomata rows per band, 4—5 abaxial smooth marginal cells]; other material at K. Original herbarium material unknown; however, the

neotype is from the original tree that was recognized and illustrated by Loudon in 1838 by common name, and noted to have been

“56 ft” tall in 1836. Leighton (1841) made specific reference to Loudon’ illustration (^p. 2083, fig. 1990”), which was of the whole

tree; this “cannot be critically identified for purposes of the precise application of the name of a taxon” (ICBN Art. 9.7).

The Dovaston yew is distinguished by relatively long undivided pendulous branchlets from ascending to

horizontal branches and by the oblong leaves that spread vertically and horizontally. The type has dark green

leaves with lateral and lower leaves spreading along two sides of branchlets and uppermost leaves spreading

upwards. The female cones of the Dovaston yew often develop near ends of branchlets that apparently have

terminated growth, and arillocarpia develop in pairs as noted by Leighton (1841). Taxus baccata var. glauca

can be difficult to distinguish; it is recognized by the recurved branchlets that are more evenly distributed

along the main branch.

Spjut, Taxonomy of Taxus 251

ahh md hdd

Fics. 32A—B. Taxus baccata, disti guished by the isodichot : hing and! x E oe u—

B (bottom). (loca. p fl hat ‘tL, | J 4 I n or 1

A (top) l ectotype (BM)

16c. Taxus baccata L. var. elegantissima. [Hort. ex] [Ravenscroft] [C.]Lawson et al., Abietineae—List Pl.

Fir Tribe No. 10, 82. 1851. (Fig. 36). Taxus baccata L. f. elegantissima Beissner, Syst. Eintheil. Conif. 23. 1887. Taxus

baccata L. (var.) argentea Loudon ex Gordon & Glendenning, Pinetum 312. 1858. Type: SPAIN. BaLEares: 1600 m, 15 Mar 1917,

Bianor-Maire s.n. (NEOTYPE designated here: BM! with male cones [abaxial surface of leaves with 9 stomata rows per band, lacking

papillae across 5—9 rectangular cells near margin]). Original herb rium material and origin unknown; horticultural form developed

in the Handworth Nursery (den Ouden & Boom 1965).

252 Journal of the Botanical R h Institute of Texas 1(1)

ee US

E e a Sd e d

: SC ANS

= t : TW

Fics. 33-35. Taxus baccata var. dovastoniana, distinguished by the

terminally digitate long pendant branchlets (in the type). 33 (top

left). Specimen from original tree (K). 34 (top right). Specimen at

BM (neotype), | ill ia, from the original t ith

annotation

tion reproduced from Carriére (1861)

L etl,

L J Z

J ; Leighton. 35 (lower left). Illustra-

The elegant yew is recognized by long linear leaves tapering gradually to an acute or acuminate apex, and

by the leaves often appearing in hair-like tufts near apex of branchlets. The dried leaves are dark—almost

blackish green above (adaxial surface)—and rusty orange below (abaxial surface), but on live plants—in

cultivation—leaves have been described as “striped pale yellow, later whitish” (Lawson et al. 1851; Rehder

1940), or under more shady conditions, leaves may appear green to golden in sunlight (den Ouden € Boom

1965). Occasional herbarium specimens have these features (preserved).

Spjut, Taxonomy of Taxus

Wn P A

í N

uini

wild

nnd n P Pc und li

ôi

S >

| 9

PLANTES DES

freed Pu z

Monk lagres, é

whe

BALEARES

haz. Peg Haz

1917, 16 «7 He. td,

it > &- poen plante

Fw BPiasot- Maur

253

Fic. 36. Te i t elegantissima, li

Bianor-Maire (neotype, BM).

I 41 "A I

J UNC INCO

254 Journal of the Botanical R h Institute of Texas 1(1)

The cv. 'Repandens—that is common in cultivation in the eastern United States—appears related to

this variety. Its low shrubby habit with a flat-topped crown may have been derived from a plant similar to

what has been described for T. baccata var. washingtonii.

16d. Taxus baccata L. var. glauca Jacques ex Carriére, Traité gén. Conif. 519. 1855. (Fig. 37). Taxus baccata L.

(£.) glauca (Carriére) Beissner, Handb. Nandelholzk. 175. 1891. Type: ARMENIA: Tschunakuchi, horticulture—ex Herb. Petropolitani,

[year 1899] Szovich 610 (NEoTYPE designated here: S: C-2072!), [with seed (leaf with 10 stomata rows per band, abaxial margin 4 cells

across without papillae, midrib papillose)]; isoNEorvrE: P! [with seed (leaf with 9 stomata rows per band, abaxial margin—4 smooth

cells, papillose midrib)]. Related material: Hort. Ex Hillier Arboretum, England, 22 Mar 1976, “BM (NH) 1247,” in adnot. T. baccata

‘Glauca’ (“Nigra”), “Blue John” (BM!). Original herbarium material and origin unknown.

Taxus baccata var. glauca is distinguished by the leaves that overlap parallel to each other while pointing

upwards (erect secund), especially near apex of recurved or horizontal branchlets, and by appearing yellow

in dried herbarium specimens, and by abruptly tapering to an obtuse apex. The branchlets are yellowish

orange and recurved in many specimens. The Dovaston yew, which overlaps with the glauca yew in character

attributes of color and pendulous branchlets, differs by the darker leaf color as seen in dried specimens, and

by the uppermost leaves often directed towards apex of branchlets.

Variety glauca includes at least three horticultural forms. (1) The cv. ‘Glauca’ or ‘Nigra’ or ‘Blue John’,

based on a specimen from the Hillier’s Arboretum (BM), has long recurved branchlets with oblong glaucous

(caesious) leaves, lacking in the yellowish color. (2) Asecond one, f. horizontalis (Carriére), allegedly a cultivar

that originated in France, is known for its long horizontal branches (Knight 1850; Callen 1977), while it

may be distinguished by its golden color (f. semperaurea Dallimore; Rehder 1940, 1949). (3) A third—that

is typical—has pendulous branchlets, and includes a specimen from horticulture under the name f. pendula

as recognized by C. Baenitz indicated on annotation label of a herbarium specimen (US: 1395580).

16e. Taxus baccata L. var. pyramidalis [Hort. ex] [Ravenscroft] [C.] Lawson, Abietineae—List Pl. Fir Tribe

No. 10-35 I951. (Fig. 38). Taxus baccata L. [f.] pyramidalis Beissner, Handb. Nadelholzk. 175. 1891. Taxus communis

pyramidalis Nelson, Pinaceae 172. 1866. Taxus pyramidalis Severin in Möllers Deutsch. Gárt.-Zeit. 41:227. 1926. Type: FRANCE:

Lananau Mutohir (Girandi), along marsh, 1 Sep 1930, Tidestrom 12814, with seed (Nrorvrr designated here: US!). Origin of material

in horticulture unknown; original herbarium material unknown.

This variety is recognized by the equally divided flexuous branchlets that have a yellowish orange color and

are often without leaves by the 3" year and by leaves diverging widely from younger branchlets. It appears

related to var. elegantissima by the paucity of cones. A specimen from Norway (Gamble) has darker metallic

leaves instead of the more common yellowish green color. Leaves generally lack papillae across 4-6 cells

near margins and are partly to entirely papillose across the midrib. Plants with more distinct radial orienta-

tion of leaves, which appear intermediate to var. glauca, may be distinguished as var. ericoides and var. erecta

(Spjut unpubl., worldbotanical.com).

16f. Taxus baccata L. var. variegata Weston, Bot. Univ. 1:292,347. 1770. (Fig. 39). Taxus baccata var. foli-variegata,

Loudon, Arbor. Brit. 4:2068. 1838, “T. b. 6 foliis variegates.” (nom. illeg. superfl.). Taxus baccata (var.) variegata aurea Carriere, Traité

gén. conif. 518. 1855 (nom. illeg. superfl.); Taxus baccata (var.) aurea Carriére, Traité gén. conif. 734. 1867 (nom. illeg. superfl.).

Taxus baccata subf. aureo-variegata Pilger, Pflanzenreich 18 (iv, 5):114. 1903 (nom. illeg. superfl.). Taxus baccata f. aurea (Carrière)

Pilger, Mitt. Deutsch. Dendrol. Ges. 25:11. 1916 (nom. illeg. superfl.). Taxus communis aurea [Nelson] Pinaceae 172. 1866 (nom.

illeg. superfl.). Type: FINLAND: Alandia, Lemland, Florstróm s.n. (Neotyee designated here: BM! [leaf with 11/12 stomata rows per

band and an abaxial marginal area of 4 rows of rectangular, smooth cells]). Original material and origin unknown.

Taxus baccata var. variegata is recognized by stiffly spreading branchlets with erect leaves that are mostly

dull olive green. The plants appear to be mostly shrubs. Tree forms that appear evident with more wide

spreading branches may be referred to as var. jacksonii (Spjut unpublished, worldbotanical.com). These

varieties, as well as a specimen from Iran referred to as var. subpyramidalis (Spjut unpubl., worldbotanical.

com), are similar to T. cuspidata.

17. Taxus contorta Griffith, Not. Pl. asiat. 4: 28. 1854. “Taxus contortus? Vide Itinerary Notes, p. 351, No.

116.” See also Itin. pl. Khasyah mts., 11:351. (1847—) 1848 (Book III, Chapter II, “Afghanistan Flora,

Spjut, Taxonomy of Taxus 255

Fics. 37A—B. Taxus baccata var. glauca, Armenia, ex Herb. Petropolitani,

lets, pinnately arranged. B (right). Close-up of branchlets showing

reflexed cones with seed.

Second year Kafiristhan.116. Taxus?”] (Fig. 40). Tyee: AFGHANISTAN: W of Kabul, “Bharowul, in woods, 7000-7500

ft" (locality data from Griffith nos. 112-114 in Itin. pl. Khasyah mts, collected during 1839-1841), Griffith 5002 (Lectotype designated

here: K [lower right specimen of three on a single sheet, with label indicating it was collected from Afghanistan, distributed by Royal

Gardens, Kew, 1862-3, accompanied by another label in handwriting with two words, Kafiristan, Griffith; the other two specimens

evidently belong to another single collection by another collector from another location with a handwritten note on a label indicating

that the bark was used in a tea in Ladakh, det. by Spjut as T. contorta]. ud material at K, a single specimen, Griffith 5002; however,

the same number was used by Griffith for specimens he collected at otl g., Taxus wallichiana from Assam. No specimens

cited, but locations from where specimens were reportedly collected are sequentially numbered in Griffiths Notes 1847-48.

Griffith, in his earlier publication, ips "Arbor, icu alternis m on sulcato univeniis basi Ya tortis. Brought from

] fthel from torsion of the base. The

Kafiristhan with the preceding [Pinus],

change takes place gradually judging from the slight obliquity of young lees Stomata blocked up, with a brown curious cuticular

substance." Griffin later published the name Taxus contorta in Not. Pl. Asiat. 4:28. 1854 (“Taxus contortus? Vide Itinerary Notes, p.

35:15 No :HH16*

Taxus fuana Nan Li & R. R. Mill in Li & Fu, Novon 7:263. 1997.— Tyre: CHINA. TIBET (XIZANG): Jilong, 3000 m, Qingzhang Expedition

7032 (HOLOTYPE: PE!).

Taxus orientalis Bertoloni, Mem. Acad. Sci. Bologna ser. 2, I, 229, pl. 2. 1862. No specimens cited, illustration provided, other original

material consists only of a single specimen at BOLO. Type: INDIA. Northeastern India, western Sikkim [“Stim”], 8000 ft, without

collectors name, without collection number, filed separately in the herbarium of A. Bertoloni, with two labels, handwriting with

locality data the same as that published by A. Bertoloni (HoLotyPE: BOLO [BOLO00077561], leaf fragments!). Bertoloni also decribed

this taxon in Misc. Bot. 23:17, Tab. 2. 1862.

Two varieties are distinguished by leaf spread and length.

17a. Taxus contorta var. contorta

Common name.—West Himalayan yew.

Distribution and ecology.—Mixed coniferous-hardwood forests of W Himalayas, 2300-3500 m; Afghani-

stan, Pakistan, India, W Nepal, and China (SW Tibet). Noted to be common in the Garhwal and Kumaon

regions at elevations near 8500 ft (Gamble 1922; Gordon 1875), where clouds often hang in oak-conifer

forests of Quercus semecarpifolia Sm., Abies pindrow (D. Don) Royle, and Rhododendron arboreum Sm. (Freitag

1971; Rau 1974). In the Uri Range closely associated with Abies pindrow-Picea smithiana (Wall.) Boiss. forest

(Sapru 1975), a vegetation type common to the higher ranges in the W and C Himalayas of India and Nepal

256 Journal of the Botanical R h Institute of Texas 1(1)

(Champion & Seth 1968; Rau 1974). In the Kumaon and Nepal regions mostly on N side of the Himalayas

in hemlock (Tsuga dumosa [D. Don] Eichler) forests with spruce (Picea smithiana) as a common associate (Rau

1974), especially near Rara Lake in W Nepal (Stainton 1972).

Taxus contorta is easily identified in the herbarium by the relatively long and straight narrow leaves that

are generally crowded along stems in nearly two ranks, which in the typical form generally do not spread

more than 60? from branchlets. The leaf mesophyll contains distinctive parenchyma cells (idioblasts) that

in the herbarium slowly develop a yellowish to reddish color, as determined in part from 1-year old speci-

mens (Amin 25025, 25045, 25149 from Pakistan, KIB) that show only minor discoloration of cell walls. The

idioblasts occur predominantly across the mid region of the leaf mesophyll and around the diffusion area

of the vascular bundle. The cell walls have striations that may suggest a type of sclerenchyma cell, but this

appears to be a secretory product—a terpenoid compound—in the cell that when dried becomes deposited

on the cell walls. Rao and Malaviya (1965) described what they called *osteo-sclereids" in leaves of one

of four varieties of T. baccata they reportedly studied from cultivated specimens in Sri Lanka and India;

however, their illustrations of leaf sections show thicker-walled cells that are sclereids. I have not seen these

“osteo-sclereids” in yew leaves, while I have observed similar cells in Asian species of Torreya.

The spongy parenchyma cells of many T. baccata specimens have similar idioblasts as seen by their

spherical shape and dark color, but lack striated cell walls, and do not fall out when sectioned. Species in

the Cuspidata Alliance (E Asia) generally have a leaf mesophyll largely of loose spherical to ellipsoidal cells

connected together by short cylindrical cells without idioblasts.

Taxus contorta is more related to T. baccata than to T. wallichiana by the relatively low number of leaf

stomata rows per band—usually 7-8, by features of the leaf parenchyma cells as just described, and by the

green to olivaceous color in leaves of dried specimens. The leaf mesophyll of T. wallichiana has periclinally

oriented cells connected in a skeletal-like net; in longitudinal sections these cells appear like bones. It is

interesting that T. baccata shows more variation (in Europe) than T. contorta (in the Himalayas) in branch-

ing, leaf arrangement and leaf anatomy. Of particular relevance is the occurrence of papillae on the abaxial

midrib of leaves—that in T. baccata can be densely papillose (e.g., lectotype), or entirely smooth (e.g., Curic

s.n., from Bosnia, K), whereas T. contorta always has a densely papillose midrib.

The close relationship between the European T. baccata and Himalayan T. contorta was recognized by

Handel-Mazzetti (1929) and by Florin—who, in his annotations of specimens at Stockholm (S), treated it

as a subspecies of T. baccata, adopting the epithet of T. orientalis Bertol., a later name.

Other botanists have independently recognized T. contorta as distinct from T. baccata and T. wallichiana,

but by names that are not always in accordance with the ICBN. For instance, Handel-Mazzetti (1929) cor-

rectly realized that T. wallichiana could be based on Wallich 6054A, but referred the Northwest Himalayan

yew to T. orientalis. He was also aware of T. contorta, which he considered a nomen nudum; however, Griffith

(1854) referred back to his earlier 1848 publication (see nomenclature citations above); thus, T. contorta is

not a nomen nudum (Art. 32.3, 32.4, 34.1), and predates T. orientalis Bertoloni (1862). Franco (1964), who

reviewed Taxaceae for Flora Europaea, also recognized the west Himalayan yew (T. contorta) as a distinct

species, but he annotated specimens (BM) by another name (T. angustifolia Franco, ined., dated 1956) that

if published would have been illegitimate; more recently, Nan Li and R. R. Mill (Li & Fu 1997) reached a

similar conclusion, but did publish their superfluous name, T. fuana. Occasional collections by Polunin et

al. (e.g., No. 432, BM) had been correctly determined, while most herbarium collections of this species have

been misidentified as T. wallichiana.

It is not clear to what extent Wallich and Griffith had distinguished yews in the Himalayas. Wallich's

(1826) Tentamen Florae Nepalensis recognized only one species, determined as T. wallichiana by Zuccarrini

(Siebold & Zuccarini 1843); however, Wallich specimens of Taxus numbered 6054, differentiated by letters

A-E, suggest they were distinguished—at least by collectors and location, and may include an annotation T.

virgata Wall. (nomen nudum), which I have identified as T. wallichiana (Blinkworth s.n. BM, reportedly from

Kumaon, but probably from Nepal), or T. baccata (young shoot on sheet with 3 other specimens of T. contorta,

Wallich 6054, ex Herb. Gordon, with “b” indicated lightly in pencil, K; probably added for comparison). Most

257

Spjut, Taxonomy of Taxus

Diem

ANS

A»

A Lx -

YN wc A

FAITS SSS AN — 7.

mmm r H^ o

NITED STATES NATIONAL MUSEUM

DEPOSITED BY THE t $ DEPARTMENT Ol AGRICULTURII

PLANTS OF FRANCE

A Faceata k,

Lot 1

A d y RAS [fs Ax

chee C Eoo) n :

—

IZA Dune e are ero depu) rbd o.

Fics. 38A—B. K [ pyramidalis, listinquished i I by | I | g, ft licl , branchlets, France, Tidestrom 12814 (neotype,

flaca £L Li 4 L Hi J J * | £L leralac

. VIVID F

US). A (top). Herbarium specimen. B (lower left insert)

258 Journal of the Botanical R h Institute of Texas 1(1)

Herb, HUGO GRANVIK

Purchased 1924

HERB. BRUNO FLORSTRÓM, |

E * “ 7 ae

ioe YOO ++

Finlandia: Alandia: ©. «c Been D

d r Ls t

is CAD Vv e nt e T aud

(

i rae

Fics. 394—B variegata, g y gid | g pi

i t Finland, Florström s.n. (neotvpe, BM). A (left). Herb B (top right). Close f | hl | I mal

g'eu Ye; L M JM! 7 V J I (SVP Ig

Wallich 6054A (from Nepal) belong to T. wallichiana, whereas most Wallich 6054B (from Kumaon) are T. contorta.

Griffith, who worked with Wallich on occasion (Burkhill 1965), also assigned mixed collections of Taxus

to the same number with different data; the type, for example, is from Afghanistan, but other labels with

this number (Griffith 5002) indicate the specimens were collected in the eastern Himalayas. Griffith (1854)

>» (€;

recognized possibly two species from Bhutan, distinguished by “axillary” and “terminal” “inflorescences” and

> 5

259

Spjut, Taxonomy of Taxus

lectotype, the species

Griffith 5002,

the one on right from Afghanistan

all are Taxus contorta,

Fic. 40. Herbarium sheet at K with three specimens

leaf in 7. wallichiana

£s+haD o ID

260 Journal of t h Institute of Texas 1(1)

a third “3. Taxus contortus?” by reference to his collection from Afghanistan. I have noted that three species

of Taxus are represented in Griffith collections, and also from J. D. Hooker in the Gray Herbarium (J. D.

Hooker 77, 87, GH) with three different species (T. contorta, T. kingstonii, T. wallichiana) all on one herbarium

sheet reportedly from Khasia, 5000-6000 ft, and from other locations in India.

Although T. contorta appears quite distinct from T. wallichiana, hybrids seem evident by the respective

higher and lower counts of stomata rows where geographical ranges of these species overlap in central and

east Himalayas. This includes the type for T. orientalis (BOLO!) from Sikkim. In several leaves studied of the

T. orientalis type, the parenchyma cells were found to be those of the T. contorta type, whereas the slightly

larger epidermal cells (20-25 pm tall, 25-35 um wide) along with the higher stomata counts (10-11 rows

per band) indicate affinity to T. wallichiana. Additionally, the absence of bud-scales at the base of branchlets,

and the strongly revolute linear leaves (illustrated by Bertoloni 1862) are other features I associate with T.

contorta. Specimens of T. wallichiana var. yunnanensis from Sikkim were found to have fewer stomata rows

(e.g., 13 rows, Kurz s.n. A; 14 rows, J. D. Hooker & Thomson A, GH, K) compared to specimens from other

regions.

17b. Taxus contorta Griff. var. mucronata Spjut, var. nov. (Fig. 41). Taxus mucronata Spjut ined. in adnot. (BM [Oct

1997], A [Jun 1996]). Tre: BHUTAN (Eastern). Ha: 27°22’ 89°18’, 9,000 ft, tree 15 ft, 11 Apr 1949, Ludlow et al. 16035, with male

cones (HOLOTYPE: A!; ISOTYPE: BM).

Differt a var. contorta folia breviora valdes divaricatis, 5-10 longiora quam latiora.

Tree to 3 m or more; leaves sharply bent at base of blade, 1.5-2.5 cm long, ca. 2 mm wide; abaxial margin

up to 23 cells across, the epidermal cells irregularly quadrate in up to 6 rows nearest the margin, becoming

long fusiform to rectangular towards the stomata band and on midrib, mostly 3-7x l/w, papillose on more

than half of the marginal cells—to within 8 (-5) cells across from margins and entirely on midrib; stomata

bands greenish, or yellowish green, narrower than the marginal region, with 9-11 rows of stomata; spongy

mesophyll with idioblasts. Male cone scales 4—5 seriate; sporangiophores 8, united into a ribbed column ca.

2mm long, thickened at apex of column, separating into 8 umbrellalike segments, each with 5-8-cuculately

lobed microsporangia ca. 1 mm diam. Seed in one specimen globose, reddish.

Common name.—Mucronate-leaved yew.

Distribution and ecology.—Nepal, Bhutan; upper forest region, 2300-3100 m.

Additional specimens: NEPAL. Dobremez 2106 (BM); Marayandi Valley, 3100 m, Wraber 514 (BM); ridge S of Bhahwe Sekh, 9000 ft,

Polunin et al. 1873 (BM); Dhawalagiri Zone, Mustang Dist., Ghasa, 2300 m, Mikage et al. 9550282 (BM).

I annotated one specimen from the Arnold Arboretum Herbarium (A) as “Taxus mucronata Spjut (ined.)” in

June 1996, designating it as type. Other specimens at the Museum of Natural History in London (BM) and

the Kew Herbarium (K) were later discovered and similarly annotated (Oct. 1997), including an isotype at

BM. An illustration in Cheng and Fu (1978) for T. wallichiana, redrawn for the English edition of the Flora of

China (Fu et al. 1999) as representative of T. fuana, was thought to have originally been drawn from the type

specimen for T. fuana as reported on the WBA website (2003-2006); however, upon study of the images of

the PE Taxus collections that were made available online during 2006, the holotype for T. fuana was found

to belong to the typical variety. Although Taxus fuana was not described until Nov 1997, all specimens I

saw at K (Oct. 1997) bearing the annotated name T. fuana by Nan Liand R.R. Mill belonged to var. contorta,

whereas specimens I recognized as *T. mucronata Spjut ined.” were not annotated by Nan Li and R. R. Mill.

Additionally, Farjon (1998) indicated T. fuana to occur only in Tibet, but the basis for his decision on this

and other species he recognized in Taxus is without taxonomic merit. In a review of specimen images of

Taxus on the PE virtual herbarium, other specimens of T. contorta var. contorta were found listed under T.

wallichiana, and also erroneously identified as T. wallichiana, while I might add that none of the specimens

I studied in herbaria outside China had specimens of T. contorta from China.

Variety mucronata is distinguished from the typical variety by the relatively shorter and more sharply

reflexed leaves—more at base of blade than near junction with branchlet. Leaves also differ from most

Spjut, Taxonomy of Taxus 261

Fic. 41A-B. Taxus contorta var. mucronata, distinguished in

part by t y A (left). Herbarium

specimen (holotype, A). B (right). Close-up of branchlets with

male cones.

FLORA OF EASTERN BHUTAN

Taxus wallichiana Zucc.

specimens of the typical variety by their abaxial surface having a smooth marginal border of 5-8 irregularly

quadrate epidermal cells, and 9-11 stomata rows per band. Despite these differences, occasional speci-

mens from Nepal are difficult to assign to either variety. An analysis of the leaf arrangement in T. contorta

specimens shows that leaves increasingly diverge at wider angles going from west to east in the Himalayas.

This also appears correlated also with an increase in number of stomata rows per band (Spjut 2006). The

higher stomata counts and wider angle of divergence in leaves of T. contorta, however, are characteristics of

T. wallichiana, whereas the wider leaf margin and reflexed leaves are also characteristics of T. umbraculifera

Nevetheless, the apparent hybrids between the two varieties of T. contorta would seem to justify varietal

status for var. mucronata.

18. Taxus fastigiata Lindley, Syn. Brit. Flora 241. 1829. (Fig. 42). Taxus baccata L. (var.) fastigiata (Lindl.) Loudon,

Arbor. frutic. britt. 4:2066. 1838. Taxus baccata L. f. fastigiata (Lindl.) Pilger, Planzenreich 18 (iv, 5):115. 1903. Taxus baccata L. var.

hibernica Hooker ex Henkel & Hochstetter, Syn. Nadelhólzer 356. 1865 (nom. illeg. superfl.). Type: IRELAND: from Florence Court,

ex Herb. Jackson, Sep 1890, Stewart s.n. (NEOTYPE

Original material unknown. Ireland, “mountains of Fermanagh” Parish of Killesher near Aghenteroark, George Willis, without her-

nated here: K!). No specimens cited, none reported in the Lindley Herbarium.

barium specimens; fide Veitch et al. (1881) —quoting from manuscript by Lord Kinnarid with further reference to Rossie Priory, 8

Sep 1867—transplanted to Florence Court ca. one hundred years ago; thus originally collected in 1767, or as early as 1740 fide

internet sources.

Common name.— Irish yew.

Distribution. —British Isles—United Kingdom (Ireland, Scotland, England), Sardinia?

The Irish yew (T. fastigiata) has long been considered distinct from other yews; yet, most authorities have

treated it as a synonym of T. baccata. Its radial orientation of leaves is similar to east Asian T. umbraculifera

in bending downwards (recurved) and to T. caespitosa in the imbricate arrangement, and it is also similar

262

Journal of the Botanical Research Institute of Texas 1(1)

\ é

N HERR A, B. JACRSO i

\ $ CONIFERA:

f Taxus baccata, L.

b. fastigiata, Lindi.

i arish Yow? Korby Jere in Ireland

be aalt fa LOMBA see

F 4) Tayus factiaiata ES a. L IL 4l + 1? L L IL aL H L 4 4 £ | I I Ll. Ll J L

IG. J Li À , "I L4 ws wi

] Is fi I to apex; Fl Court, ex Herb. Jackson, Stewart s.n (neotype, K)

Spjut, Taxonomy of Taxus 263

to a Euro-Mediterranean variety of T. baccata from Morocco. The Euro-Mediterranean plants with radial

spreading leaves, which occur near the western continental distribution limits of the genus, are relatively

rare. The columnar habit, associated with radial orientation of leaves, would seem to be an adaptation to

open habitats such as bogs, fens, lake shores, or areas along sea coasts.

Taxus fastigiata is known primarily from two plants that once grew in the Cuilcagh Mountains in

Fermanagh County, Ireland. A farmer named George Willis dug up two female plants; one was planted in

his garden, the other was given to his landlord, Lord Enniskillen, who planted it at Florence Court (Veitch

et al. 1881). Willis’s yew died in 1865, but the Florence yew prospered, and subsequently has become the

source for many horticultural varieties (Veitch et al. 1881). Additionally, male plants appear evident at North

Mundham in Sussex, England (Bean 1953), and specimens from Tweeddale in Scotland may also be native.

Irish yew is common in cultivation in the United States, especially in the west where it can be maintained

in hot and dry climates, temperatures often exceeding 100?F as evident at Redding, CA.

The common “English yew,” which I recognize as T. recurvata, is distinguished from typical T. baccata

by the arcuate leaves that curve downwards. Specimens from the British Isles show a more radial orienta-

tion of leaves compared to specimens from central and Eastern Europe. The leaves of T. recurvata from the

British Isles also appear to develop closer together on branchlets and are darker green in color as seen in T.

fastigiata. This includes a specimen from Neopath Castle Scotland (K) that in my opinion is a hybrid between

male T. recurvata and female T. fastigiata; the collectors Bean and Hill indicated it was a *distinct variety" by

its “plumose growth."

Several varieties of Irish yew are proposed (ww w.worldbotanical.com). One appears to be a low shrub in

the British Isles and possibly on Sardinia; the other, a tree, is known only from Scotland and England. The

Scotland plants, which are from Lock Lomond, differ from the typical Irish yew in the branches ascending

more widely. The relationships of the proposed varieties require more study.

As mentioned elsewhere in this paper, and in Spjut (2007), the Cuspidata Alliance includes plants with a

similar phyllotaxy to that of the Irish yew, particularly in specimens from the islands of Sakhalin, Hokkaido,

and Honshu. I annotated one specimen from Sakhalin Island (Dvorakovskaia & Bokina, A) as T. fastigiata in

june 1996 because the recurved leaves with a purplish green color seemed characteristic of the Irish yew;

however, I now regard it as T. caespitosa var. latifolia. Another specimen of this variety from Sakhalin Island,

Flanakan & Kirkham 203 (K), resembles T. baccata var. glauca in the upturned leaves with a yellowish green

color. The Asian T. caespitosa is usually distinguishable by the spreading leaves that bend or curve more

along their petioles instead of along their blades.

19. Taxus recurvata Hort. ex [Ravenscroft] [C.] Lawson, Abietineae—List Pl. Fir Tribe No. 10, 83 (A

handsome and distinct species with recurved branchlets and leaves mostly directed upwards"). 1851.

(Fig. 43). Taxus baccata L. [var.] recurvata (Hort. ex C. Lawson) Carrière, Traité gén. Conif. 520 (“Branches étalées, divariquées,

alongées, peu ramifies, le plus souvent réfléchies"; "feuilles longues et étroites, falquées-countour-nées, involutes." “Cette variété ne

peut étre confondue avec acune autre”). 1855. Type: ENGLAND: Sussex: Bury Hill, near summit on chalk, 24 Aug 1933, Hubbard s.n

(NEOTYPE designated here: K! [with mature seed; leaf with 8 stomata rows per band and abaxial marginal border of 4 smooth cells].

Original material unknown.

Common name.—English yew.

Distribution.—Europe, SW Asia (Caucasus Mts.).

This species is recognized by plants having most of the following characteristics in common: arcuate

leaves, i.e. the leaves are recurved (turned downwards), the leaves crisscross more than parallel each other

along one side of a branchlet, the adaxial leaf surface appearing strongly convex, leaves discolorous, and

prismatic seeds developing on branchlets of the current season. Taxus baccata differs by paler green leaves

with the blades facing in the same plane as that of branchlets, and with leaves along one side of the branchlet

overlapping more than crisscrossing each other, and also curving upwards instead of downwards.

Taxus recurvata has intermediate phyllotaxy between the radial type of T. fastigiata and the distichous-

like arrangement of T. baccata. In the British Isles, T. recurvatais common to chalk downs where *yew woods"

form a climax community as evident from specimens that correspond to literature (Watt 1926; Rodwell

264 Journal of the Botanical R h Institute of Texas 1(1)

Fics. 43A—B. Taxus recurvata, distinguished by the recurved branchlets,

crisscrossing leaves with sharp contrast in color between the abaxial and

adaxial surfaces; England, Hubbard s.n. (neotype, K). A (left). Herbarium

specimen. B (above). Close-up of leaves and seeds.

FLORA OF THE BRITISH ISLES.

Tatuo

a A

abitat Reng Ml y Mer puma

patty à T

m hath

bu 29/9995

et al. 1991—as T. baccata). Drawings and photos in Loudon (1844), Watt (1926), and Rodwell et al. (1991)

show the twisted branches that characterize this species. Specimens from Ireland identified as T. baccata var.

glauca may be an introduction as yew is generally rare in Ireland; however, one record is known to occur with

Arbutus unedo, a species more commonly found in the Mediterranean flora. The natural occurrence of yew

in Scotland has also been considered rare (Godwin 1975; Mitchell 1974) —where two different varieties of

T. fastigiata are proposed, one from Inchcailloch Island (Loch Lomond) and another from near Tweeddale.

A large tree —with a circumference of 16 m, from near Fortingall—is thought to be the oldest living yew

in Europe, estimated at 3000 yr (Voliotis 1986) to 5000 (29000) yr on internet sources. I have not studied

specimens of this plant but photos on the web show the isodichotomously divided branches from the trunk

that suggests T. baccata, and one fuzzy photo showing leaves on pendant branches that further suggest T.

baccata var. dovastoniana.

Plants with less distinct radial orientation and more distant leaves are proposed as distinct varieties

(Spjut, unpubl., www.worldbotanical.com). They differ by leaf length. One with long narrow linear leaves,

which includes specimens from Madeira, British Isles and Romania, is difficult to distinguish from T. con-

torta.

IIB. BACCATA GROUP, CUSPIDATA ALLIANCE

Yews of this alliance are recognized by leaves spreading from branchlets more by bending of their petioles

than by curving of their blades, and by having a relatively broad abaxial margin zone of epidermal cells

without papillae, usually from 8-24 cells across. Another less obvious distinction is that the epidermal cells

are slightly wider and shorter than those of the Baccata Alliance, except for T. canadensis. Five species and

15 varieties are recognized.

Spjut, Taxonomy of Taxus 265

Uy ¡27

Up f

Fics. 44A—B. Taxus biternata, disti g ishedI y t h-di

a LA Jal: L LI I | * L 1? J two-thirds

No.// 4 £ £ ARNOLD ARBORETUM. : £. 41 L 1L 4 . L PA pet £

EXPEDITION TO EASTERN ASIA, 1917-18 Mp 1 J y

Tati b E e E the current season; South Korea, Wilson 10688 (holotype,

de 2 Lum A). A (top). Herbarium specimen. B (bottom). Branchlets

Renee d me and leaves with two arillocarpia.

vb db Pr P LLLI

266 Journal of the Botanical R h Institute of Texas 1(1)

The taxa are very difficult to distinguish as they intergrade; Taxus biternata and T. caespitosa var. caes-

pitosa are most distinct.

20. Taxus biternata Spjut, sp. nov. (Figs. 6, 44). Tyee: SOUTH KOREA. Kyoc[sanc] Prov.: Kyongsan, Nemon-rei, common

or abundant, 15 Sep 1917, Wilson 10688 (HoLotyee: A! with arillocarpia (abaxial leaf with marginal zone of 8-9 smooth cells, 9 papil-

lose cells, stomata 9 rows per band, midrib lacking papillae); isotypes: K! (leaf with marginal zone of 6 cells across without papillae

followed by 8 rows of papillose cells, 7 stomata rows, and midrib with marginal papillae), US!).

Taxus microcarpa (Trautv.) Spjut ined. (in adnot.: A, BH, BM, GH, K, NA, P).

Taxus canadensis affinis, a qua imprimis differt arbor, ad 30 m alta, arillocarpia maturibus ramulus primus et stomata (7-)9-13(-16)

seriata/zona; ramis anisodichotomis horizontalibus, ramulis diffusibus biternatifidibus; folia pun Speers patentia, linearia,

1.5-2.5 em 1.5-2.5 mm b acuta ad acuminata, 0.150-0.300 mm cr Į , SUL subtus

0.8-25 um alta, 20-30 um lata, quasi elliptica i j ] 151 >

AOE cal : 1;

Is E

cellulae quasi rectangulara costa interdum maples vel laevis, proparte eine versas de Re

sine papillae (6—)8—12(-19) cellulae mar ale latis.

Trees or shrubs with erect trunks and horizontal branches, to 30 m high; branchlets often short and much-

divided, subpinnately arranged but unequally divided, appearing ternately divided or with short delicate

tertiary branchlets, horizontal or weeping, yellowish green when young, yellowish orange with age; bud-scales

closely overlapping in 3-4 ranks, mostly persistent to the 3" yr, thick, deltoid, concave, medially recurved

and incurved towards apex to form a cusp, with an obscurely thickened midnerve, ca. 1 mm long, spread-

ing from base of branchlets. Leaves persistent on older twigs, or lacking, green upon drying, in two-ranked

like arrangement to apex, linear, straight to slightly falcate, 1-2 cm long, 1-2 mm wide, 150-250 um thick,

pale green and convex above to a rounded midrib that forms a channel along the base of the midrib, pale

yellowish green and concave below to a rounded midrib, revolute near margins 30-90? in dried leaves,

more notably revolute at upper one-third of leaf; adaxial epidermal cells in T-sect. elliptical, 10-15 pm tall,

25-40 um wide; abaxial epidermal cells similar or larger, 10-15 um tall, 15-25 pm wide, numbering 11-18

between margin and stomata band, mostly rectangular, or sinuous near the stomata band, 3-7x l/w except

quadrate in 1-3 rows near margins, epapillose entirely across the marginal region, or marginal region often

partially papillose, often epapillose on (6-) 8-18 cells in from the margins, occasionally with obscure papillae

on midrib, papillae usually more prominent on marginal cells bordering stomata band, in 2 opposite rows;

stomata bands broader than the marginal region, with 7-13(-16) stomata rows per band. Male bud cones

globose, ca. 1 mm diam.; scales 4-seriate; sporophylls ca. 14, united into a terete, smooth or obscurely ribbed

column, thickened at apex, spreading shortly above, each branch bearing 8-10 lobed, cucculate sporangia.

Female cone scales 4—5 seriate; aril red or pink with tinge of white, with a deep cup, drying dark purple;

seed subglobose, obscurely angled where tapering to apex in upper half, 4 mm long, 2-4 mm diam.

Common name.—Delicate branch yew.

Distribution and ecology.—China (NE, Manchuria), Russian Federation (SE Region), North Korea, South

Korea, Japan; forests, 800-1400 m. In NE China dominant in “mixed broad-leaved deciduous and needle-

leaved evergreen forests” (Hou 1983). On N Hokkaido (Japan), “fairly common" within a mixed conifer

hardwood forest of Picea jezoensis (Siebold & Zucc.) Carriére, P. glehnii (F. Schmidt) Mast., Abies sachalinensis

(F. Schmidt) Mast., Populus maximowiczii A. Henry, Kalopanax septemlobus (Thunb.) Koidz., Ulmus japonica

Siebold, and Acer pictum Thunb. ex Murray (Wilson 1916, as T. cuspidata). Common in cultivation, includ-

ing Cv. ‘Capitata’ and shrub forms misapplied to T. media Rehder. At the Secrest Arboretum, apparently

spreading by seed among native deciduous hardwoods.

Additional specimens. RUSSIAN FEDERATION. Far East Region: Pryanyk For. Div., Zalese Village, silver fir-cedar-broad-leaved forest,

B. Cerereu (in Russian, A). Manchuria: Northern, Sochintzest, forest, small trees 20 Sep 1931, Skvortzov s.n. (A). CHINA. Jilin (Kirin), 5

Sep 1931, C. H. Chen 539 (A); Mandshuria SE, ex herb. hort. bot. Petropolitani, 1860, Maximowicz, iter secundum (A, US); Northern China

(Shaanxi: Tai-pei-shan fide Rehder & Wilson in Sargent 1914], Purdom s.n. (GH). [South] Korea. Kyog[sang] Prov.: Kyongsan, Nemon-

rei, common or abundant, Wilson 10519 (A, BM, US), Wilson 10688 (A, BM); N. Heian Prov., O.G.M. Co. Mines, Pukchin & Takkari,

833-1000 m, not uncommon in moist forests, Wilson 8685 (A, K, US); Shinkabachin Heizanchien to Ehoshin, Kankyo-N Heian divide,

rare, 5 Sep 1917, Wilson 9097 (A); Seoul, East Palace Park 24 Sep 1905, Jack s.n. (A). JAPAN. Hokkaido: Cosl Mines, Utishini, tree ,

Spjut, Taxonomy of Taxus 267

20 Sep 1892, Sargent s.n. (A); Aza-akaigawa in Morin-machi, 42?0'N 140?39'E, near stream in open woodland, 200 m, tree 20 ft, seeds

embedded in reddish aril, Meyer et al. 19261 (NA); Teshikaga-Machi, 3.2 km SE of Lake Kusharo, road 243, Kawakami-gun, Kushiro,

43.35 N, 144.23 E, Meyer et al. 19112 (NA); Hokkaido, Kitami prov., common in moist woods, tree 15 m x 1.5 m, 17 Aug 1914, Wilson

7399 (A). Honshu: Sernja prov., Yamanaka on Fuji-san, abundant, tree 6—3 m x 1.5-2.6 m, Wilson 7778 (A, K); Kai prov., around village

of Nakaihinsen, common hedge, Wilson 7544 (A); Nagano-ken, Okmachi, Uno 2611 (A, BH); Yokohama, ex Herb. Hort Petro., Maximowic

(P); Kamikawa, Nitzelius (S: C-2111); Tokyo Pref.: Oizuni, Nepymawku Makino 43775 (S: C-2111); Mt. Kiyosumi, Makino 43779 (S: C-

2122); Sapporo, Yezo, 21 Jun 1903, Arimoto s.n. (A).

Taxus biternata is easily identified by its tree habit with an erect bole and horizontal diffuse branching,

and by the much divided slender branchlets with a two ranked leaf arrangement. The tree habit not only

distinguishes it from T. cuspidata, which differs by long ascending or recurved branches, but also from a

shrub yew originally described as T. baccata var. microcarpa. A detailed study by Kolesnikov (1935) showed

that the tree variety (T. biternata)—which he referred to T. cuspidata—and shrub yew were parapatric with

distinct morphological and ecological characteristics. The shrub yew, T. umbraculifera var. microcarpa, is also

similar to T. canadensis in layering, but differs in its flat-topped radial growth—as illustrated by Kolesnikov

(1935), and by the much smaller paler seed, shaped like a *Hershey Kiss."

Occasional specimens of T. canadensis from North America (e.g., Travis 119, Maine, PH), Estonia (e.g.,

Lundstróm 742, S), and others from Europe (e.g., Handel Mazzetti, Mt. Olympus, Greece, K) and SW Asia

(e.g., Davis 13667, Turkey, K) are similar to T. biternata in the linear leaves spreading in two ranks with

more strongly revolute margins in the upper third as seen in dried specimens. While most specimens can

be distinguished by branching, the 3* yr branchlets of T. canadensis have a purplish tint, in contrast to yel-

lowish orange in T. biternata. A Maack specimen without number from Manchuria, mounted with the type

with T. baccata var. microcarpa (GH), however, has a leaf spread and color that is hard to distinguish from

T. canadensis. The female cone scales appearing on the current season growth compare more favorably with

T. biternata. Taxus canadensis seed often mature on 2™ yr or older growth as seen in herbarium specimens;

while I also recognize that field studies are needed throughout the range of the species to further substanti-

ate the taxonomic value of this character.

In southeastern Manchuria, T. biternata appears to hybridize with T. umbraculifera var. microcarpa and

var. umbraculifera. Plants with linear leaves (10x l/w or more) that are strongly recurved in upper third are re-

ferred to T. biternata. Those with relatively short leaves (oblong, 5-8x l/w) are considered var. microcarpa.

21. Taxus caespitosa Nakai, Ch[Ty]ósen Sanrin Kaihó (J. Kor. For. Soc.) 158:40. 1938. (Figs. 8, 45). Taxus

cuspidata Siebold & Zucc. var. caespitosa (Nakai) Q.L. Wang, Clavis Pl. Chinae Bor.-Or., ed. 2:73. 1995. Type: JAPAN: 15 Jul 1922,

Sawada s.n. (Lectotype designated here: TI!). The reference to the type specimen in the typographical format quoted from Nakai

(1938) is interpreted here as belonging to either of the t p ited; the other, Kimura s.n., Aug 1924 (TD, is here identified

as Taxus umbraculifera var. nana. ToroTyPES: Wilson (AD, Makino 43792 (SD.

Taxus umbraculifera subsp. latifolia (Pilger) Spjut ined. (in adnot. A).

Three varieties are recognized.

21a. Taxus caespitosa var. caespitosa

Common name.—Caespitose yew.

Distribution and ecology.—Russian Federation (Sakhalin Is.), Korea, Japan.

Additional specimens: RUSSIAN FEDERATION. Sakhalin Is.: Schmidt s.n. (GH). KOREA. In forest, Aug 1907, Faurie 1512 (A, BM,

E). JAPAN. Honshu: Niigata, Yuzawa-machi, Minamiuonuma-gun, in Pinus pumila scrub, 1650 m, evergreen shrub 0.8 m high, fr red, 3

Oct 1979, Taoda 3887 (A); Hakkada, windswept slopes, 1000-2000 m, bush 1-1.5 m, rare, 4 Jul 1914, Wilson 7133 (A); Mutsu Prov., Mt.

Hakkoda, Mizushima 1985 (A); Ohobu near Kobe Calta, 19 Mar 1955, Muroi 1058, annotated T. cuspidata var. ambraculifera (A). CULTIVA-

TION. OHIO. Secrest Arboretum: ‘Colean’, A30-131; Dwarf Japanese Yew’ A30-163; “Hill Anglo-Japanese Yew’ A30-264

Taxus caespitosa var. caespitosa is identified by the crowded branches and radial distribution of leaves that have

an erect orientation. The leaves appear more imbricate than decussate, and curve upwards along the petioles

and the blades in the same direction they spiral. Wilson (1916) described plants he saw in Japan as having

prostrate branches (from which erect branchlets apparently arise). This is evident in herbarium specimens

by the one-sided development of branchlets, which includes specimens from the type locality, Mt. Daisen,

268 Journal of the Botanical R h Institute of Texas 1(1)

Fics. 45A—B. Taxus caespitosa, distinguished by the ascending to

erect imbricate leaves; Japan, Honshu, Mt. Daisen, Makino 43792

(topotype, S). A (left). Herbarium specimen. B (right). Close-up of

leaves and seed.

“Aceh sut TE s

li r ili "n nom MAT am n iilil Fun iili

obtained by Wilson and by Makino. However, cultivars that I have seen as belonging to this variety are not

prostrate shrubs, and do not show the small seeds seen in herbarium specimens of wild plants.

In E Asia where shrub yews show evidence of considerable diversity in habit and in phyllotaxy, the one

sided branching and secund leaves of var. caespitosa are postulated as ancestral traits allegedly retained in

plants with ascending to erect basal branches; these plants are referred to var. latifolia. Kolesnikov (1935)

regarded the presence of yew in the Manchurian flora as an archaic element having little historical relationship

to the rest of the vascular flora, while shrub yews in other regions have been noted to occur in environments

that are distinct from their tree relatives such as reported by Elias and Korzhenevsky (1992) for shrub yews

in Ukraine and Georgia. Two other distinct shrub types are also recognized in North America, T. brevifolia

var. reptaneta in the Pacific NW has a characteristic habit with a distinct ecology, and T. canadensis in NE

America has long been know for its shrubby monoecious habit.

21b. Taxus caespitosa Nakai var. angustifolia Spjut, var. nov. (Figs. 46-47). Tee: JAPAN. Prov. Kozuke: Oze-ga-hara,

» &«

“shrubby habit, “leaves dark green,” “yellowish green beneath," 26 Aug 1950, Mizushima 401 (HoLotypE: A! [leaves with a double row

of palisade parenchyma cells, a marginal zone of 14 cells across lacking papillae, 9-10 stomata rows per band; seed]).

Varietas nova frutex, var. caespitosa proxima, cujus ramis primus prostratus, ramulosus numerosus, erectus; folia pectinatum disposita,

patentia, linearis, 1.5-2.0 cm longa, 1.5 mm lata, acuta, +0.200 mm crassa, «disticha; semen ovata, purpurascens, 5 mm longa.

Apparently prostrate, bearing numerous erect reddish orange branchlets, persistent bud-scales few, cuspidate,

ca. 1 mm. long; leaves mostly distichous, spreading from erect branchlets, erect on horizontal branchlets,

reportedly dark green above, yellowish below, becoming reddish green in the herbarium, revolute along

margins when dried, 1.5-2.0 mm long, ca.1.5 mm wide, lacking papillae entirely across 14 marginal cells

and on and midrib; with 9—10 stomata rows per band. Seed near base of branchlets, ovoid, purplish, taper-

ing to sharp apex.

Common name.— Ground cover yew.

Spjut, Taxonomy of Taxus 269

Fics. 46A—B. Tt

angustifolia, isti

4L sl * | IL Eor £l £.

[3 ez o id M I

4 fL LA eal AL AN DNUS IL al I 4°. | "NJ

y ,

L | AE Jel

Mizushima 401 (holotype A). A (above). Herbarium specimen,

L * 4 4 L J 4 L LI $ R

(left). Close-up of leaves and seed.

Distribution.—Korea, Japan.

This variety is recognized by the relatively thin, narrow leaves as in T. biternata, but apparently has

the habit of T. caespitosa var. caespitosa as shown in Fig. 50, reproduced from the Illustrated Encyclopedia

of Fauna & Flora of Korea (Chung 1965). In this reference, the species is reported to be a shrub that is

distinguished by layering.

Additional specimens: None. Known only from the type and illustration.

21c. Taxus caespitosa Nakai var. latifolia (Pilger) Spjut, comb. nov. (Fig. 48). Basionvm: Taxus baccata L. [subsp.

cuspidata (Siebold & Zuccarini) Pilger] var. latifolia Pilger, Pflanzenreich 4(5):112. 1903. Taxus cuspidata var. latifolia (Pilger) Nakai,

J. Kor. For. Soc. 158:39. 1938. Type: JAPAN: Hokkaipo: Hakodate, [with two labels, one dated 30 Dec 1890 and another dated 1888],

Faurie 6345 (LecToTYPE designated here: P! [with male cones (abaxial leaf with 18 smooth marginal cells followed by a stomata band

with 11 rows of stomata, and a smooth midrib] ); isoLecTotYrE: K fragment!). Syntypes: from E Russian Federation and Japan.

Additional specimens. RUSSIA FEDERATION. Sakhalin Is.: 46°37’N, 142?53'E, Prigarodne, mixed conifer (Abies, Picea)/broadleaved

(Betula, Sorbus) woodland, 150 m, to 1.8 m high, Flanakan & Kirkham 203 (k). Korea: in forest, 800 m, 19 Jul 1910, Taquet 4455 (A);

without locality data, Faurie 117 (P), 3406 (P), 5975 (P). JAPAN. Hokkaido: Shiribeshi Prov., Shiribeshi-san, branches wide spreading,

270 Journal of the Botanical R h Institute of Texas 1(1)

1-2 m, common, 1300-2000 m, 27 Jul 1914, Wilson 7265

(A). Honshu: Mutsu Prov., Mt. Hakkoda, erect tree, 8 Jul

1952, Mizushima 1989 (A). without locality data, Faurie

A fi IZ 5114 (P: 2 sheets). Locality unknown: Folley s.n., identified

| A

Y ri

T. cuspidata ‘nana’ (K).

4 À EW Variety latifolia is best distinguished by the

E YT flexuous branchlets with overlapping erect

E NS

ES if

Wa leaves; however, it varies widely in habit.

It may produce a single main branch that

creeps along the ground and layers as evident

in Faurie 5114 from Japan, Faurrie 3406 &

5975 from Korea, Flanagan & Kirkham from

Sakhalin Is., and Folley from cultivation. Erect

forms are recognized by branches and leaves

that develop primarily on one side of the

plant. Wilson (1916) described plants from

Shiribishi-san in Japan as either “prostrate

on the ground” or as a “broad shrub, 1-2 m

high.” The prostrate form could be treated as

another distinct variety. Chang (pers. comm.)

found significant differences in taxane ratios

in specimens that | identified as two forms

j of T. caespitosa that were reportedly grown

jp under similar conditions in a greenhouse.

Pilger (1903) distinguished var. latifolia

from var. cuspidata by the densely branched

Fic. 47. Taxus caespitosa angustifolia, luced fi Illustrated Encyclopedi habit, relatively wide leaves (to 2mm wide),

of Fauna & Flora of Korea (Chung 1965, as T. caespitosa) Description ofplantinthat and small, depressed globose seeds. His

| SO: description partly agrees with that given

by Trautvetter (in Maximowicz 1859) for

an earlier name T. baccata var. microcarpa; however, two lectotypes—as applied to two different varietal

names—were selected from four syntypes (Faurie 5975, 6345, Maack s.n., Schmidt s.n.) as cited by Pilger

(1903). Faurie 6345 at P was selected as lectotype for var. latifolia because it is a whole specimen mounted

alone, and because it has cones (male), compared to other Faurie specimens that were usually sterile and

mounted with other specimens on a single sheet as seen in other herbaria. Maack from Heilongjiang at GH

is the lectotype for T. baccata var. microcarpa (bottom specimen of two mounted on one sheet). A specimen

by Schmidt from Sakhalin Island at GH, referred to in the present study as var. caespitosa, is a reddish speci-

men mounted among several other greenish specimens on the same sheet—regarded as T. umbraculifera var.

nana—possibly collected by Augustiowoz. Nakai (1938) also distinguished var. latifolia by leaves 3-4.5 mm

wide and indicated that var. nana, an earlier name, was known only in horticulture; however, var. nana has

been recognized to occur naturally in Japan (Ohwi 1965).

22. Taxus canadensis Marshall, Arbust. Amer. 151. 1785. (Fig. 49). Taxus baccata L. var. canadensis (Marshall) Gray,

Man. Bot. N. United States, ed. 2, 425. 1856. Taxus baccata L. subsp. canadensis (Marshall) Pilger, Planzenreich 18 (iv, 5):113. 1903.

Type: U.S.A. VERMONT: 1877, C.G. Pringle s.n. (NEoTYPE designated here: US! [leaf with abaxial marginal region of 16-18 smooth cells,

stomata bands papillose throughout with stomata in 6 irregular rows per band]). Original material unknown.

A specimen at the Museum of Natural History in London (BM) collected by Joseph Banks in 1766 refers to “Marsh. Hb.,” which

suggests a possible type. However, the existence of material used by Marshall is unknown. His publication (Marshall 1785) was

related to his sales catalogue (Silber & deWolf 1970). A neotype was selected based on a cone-bearing specimen that showed the

most common morphology.

Spjut, Taxonomy of Taxus 271

HERB. MUS. PARIS.

ssas Kee apto dit Flas, 48A-B. Taxus caespitosa var. latifolia, distinguished by

the spreading branchlets with ascending to erect, imbricate

| DYRE leaves; Japan: Faurie 6345 (lectotype, P). A (left). Herbarium

ie specimen. B (below). Close-up of branchlets with leaves and

male cones.

Becu le 30 décembre 1890, M+ abbé Fen.

Em IR titute of Texas 1(1)

Journal of

272

NN. TULUM UU UL

Prem

M Fics. 49A—B. Taxus canadensis, neotype. U.S.A., Vermont, Pringle,

WZ yr 1877 (US). A (left).Herbarium specimen. B (below). Close-up

Ea fk hlet showing | Var lensis disti

guriieu vy

two sides of a branchlet.

LI 57

| ha AECA, PDA

H 7

«EO SI

OK an

Wim ^

PI OE

Spjut, Taxonomy of Taxus 273

Taxus canadensis is distinguished by leaves having relatively narrow stomata bands bordered by 8 or more

marginal cells. Stomata usually number 5-7(-9) rows per band in American plants, or from 4-11 rows

per band in Euro-Mediterranean plants. The marginal zone along the abaxial surface of the leaf varies

from 12-18 cells wide in specimens from North America, and from 8-24 cells wide in specimens from the

Euro-Mediterranean. American plants have papillae confined to stomata bands, sometimes only along a

stomata row. Except for one specimen from Ithaca NY, the Euro-Mediterranean plants also differ by more

conspicuous papillae in stomata bands with papillose cells sometimes extending into the adjacent marginal

zone. The European plants could be referred to fossil species T. inopinata Givulescu or T. grandis Krausel.

The consistent absence of papillae along the abaxial leaf margin in American plants clearly indicates that

development of epidermal papillae is genetically fixed.

Taxus canadensis has also been distinguished from related species by its monoecious habit in which

plants often creep along the ground and spread by layering (Allison 1991; Bannan 1942), but these characters

are rarely reported on labels of herbarium specimens. Nevertheless, T. canadensis is not always monoecious

(Allison 1991), and perhaps tree forms exist; a specimen I cited from Sainte Baume (France) is where the

oldest yew trees are known in France, “attaining a girth of 11% feet" (Elwes & Henry 1906), and where yew

was once abundant (Elwes & Henry 1906). Layering in European yew is thought to be rare, but nevertheless

known from Scandinavia (Elwes & Henry 1906) where I have also identified T. canadensis from herbarium

specimens. The occurrence of T. canadensis in Scandinavia is further evident in a photograph of a plant on

Saaremaa Island in Estonia that clearly shows a low sprawling plant in a forest understory (of spruce, Picea

abies [L.] H. Karst; Korpela's Index, website), which I have further identified as T. canadensis var. adpressa.

Two other character features that reinforce identification of T. canadensis outside North America are

bud-scales at base of branchlets and color of leaves. In the Canada yew, bud-scales remain rather loosely at-

tached at base of branchlets, and are usually keeled or folded along the mid-nerve, often appearing incurved

or cuspidate above the mid region as in T. biternata (see Cope 1998 for illustrations of bud-scale features).

This is in contrast to scales of the T. baccata and its allies that show various other combinations of character

attributes. In T. baccata var. baccata, bud-scales at base of branchlets are loosely attached, but still thick and

obtuse, not at all incurved or cuspidate, or other varieties of T. baccata have similar thickened obtuse scales

more tightly adpressed to branchlets, or in the related T. recurvata, bud-scales are thickened cuspidate but not

tightly adpressed to branchlets. The other character feature of T. canadensis is the dark green color of leaves as

determined from general observation and from under the microscope. Most specimen leaves of T. baccata differ

by a yellowish green, olive green, or glaucous green color. Exceptions occur in specimens from eastern Europe

and southwestern Asia where color and anatomical differences overlap between T. canadensis and T. baccata.

Three varieties of T. canadensis are recognized. The typical variety is frequently isodichotomously

branched with leaves in pressed specimens appearing crisscrossed in pairs along one side of a branchlet.

The leaves are broad linear and spread from branchlets at their petioles in which the blades are relatively

straight. Variety adpressa differs by irregular alternate branching and by having oblong and truncated leaves

near apex, which is acute or obtuse, while var. minor is recognized by the crowded erect (secund) leaves.

In further studies additional varieties may be distinguished; an example is Hayek & Hayek s.n. (BM) from

Styria Superior in Austria that is similar to var. minor except for the branchlets that appear more recurved

with narrower erect leaves, features that are seen more in T. cuspidata.

22a. Taxus canadensis var. canadensis

Common name.— Canada yew.

Distribution and ecology.—E North America (Manitoba near Lake Winnipeg south to Indiana, Newfound-

land south along the Appalachian Mts. to NW North Carolina and Tygarts Creek, Kentucky), NW Africa,

Europe, W Asia; shady wet places, benches above rivers, or among rocks or soil in bottomland forests of

mixed hardwoods and conifers, especially hemlock and beech, 300-1500 m.

274 Journal of the Botanical R h Institute of Texas 1(1)

22b. Taxus canadensis Marshall var. adpressa (Hort. ex

Carriére) Spjut, comb. nov. (Fig. 50). Basioxvw: Taxus [baccata var]

adpressa Hort. ex Carrière, Rev. Hort., sér 4, 4:93, fig. 8. 1855 (and Traité

gén. conif. 520. 1855). Taxus baccata L. [var.] adpressa (Carrière) Carrière,

Traité gén. conif. 731. 1867. Tyre: Described from horticulture, original

material unknown (neoTYPE designated here: Illust. Fig. 8 in Carrière, Rev.

Hort., sér 4, 4:93. 1855).

Cephalotaxus tardiva Siebold ex Endlicher, Syn. conif., 239. 1847. Taxus tar-

diva (Siebold ex Endl.) Hort. ex Knight, Syn. conif. pl. 52 (1850); T. tardiva

(Siebold ex. Endl.) Hort. ex Lindl. & Gordon, J. Hort. Soc. 5:227. 1850.

Taxus baccata f. tardiva Pilger, Pflanzenreich 18 (iv, 5):114. 1903.

Common name.—Rigid-leaf yew.

Distribution.—E North America, Europe, W Asia.

The name for this taxon has been confused in the

literature. If recognized as a species, the correct name

would be Taxus tardiva, but as a variety, the epithet adpressa

has priority. Carriere referred to it both ways in 1855, as

a species in Revue Horticole, and as a horticultural variety

in Traité Général des Coniferes under the bionomial name,

Taxus adpressa, with many synonyms listed, including

T. baccata adpressa with reference to his earlier journal

publication (Carrière 1855b). The ICBN (Art. 11.2, 11.4)

indicates that priority is determined by the “final epithet,”

which “Taxus adpressa” was indicated to be a variety, and

although in a binomial format, the epithet has priority as

variety over Cephalotaxus tardiva that was first recognized

as a species.

This variety is generally known in horticulture where

it was thought to originate as a natural seedling in a nursery

at Chester, England in 1826 (Bean 1953; Elwes & Henry

1906; Pilger 1916; Wilson 1916), but also recognized to

occur occasionally in the wild (Krüssmann 1985). Others

(Fig. 8. — Taxus adpressa.

have reported it native to China and Japan (Endlicher

1847; Koch 1873), and still other reports mentioned it as

occurring in California, in association with sugar pine,

Fic. 50. T. canadensis var. adpressa. Reproduced from Carriére

(1855b), neotype. Var. adpressa distinguished by the oblong, ponderosa pine, Douglas fir and other species (Revue

btusel | ling along two sidesofabranchlet, Horticole 1848, 1849).

overlapping slightly.

22c. Taxus canadensis Marshall var. minor (Michx.)

Spjüt, comb. nov. (Fig. 51). Basonwm: Taxus baccata B minor Michx., Fl. Bor. Amer. 2:245. 1803. Taxus minor (Michx.) Brit-

ton, Bull. Torrey Bot. Club 4:167. 1893, no specimens cited. Original Material at P, a single specimen. Type: CANADA. ex Herb. A.

NAGEL P!

iVILCTLOLU. MOLOTYPE. D.

Common name.—Minor yew.

Distribution. —NE U.S., E Canada, Portugal (Madeira), Austria.

This differs from var. canadensis by the more densely leafy branches with the leaves tending to be obtuse

and falcate-secund.

23. Taxus cuspidata Siebold & Zucc., Abh. Kónigl. Bayer. Akad. Wiss., Math.-Phys. München, III, 801.

1843 (in footnote: ref. to Siebold and Zuccarini, Fl. Jap. II. tab. 128 and to [a description] T. baccata in

Thunberg (1784, Fl. Jap. 275, “foliis solitariis, linearibus, cuspidatis, approximatis"). (Figs. 7, 52). No

specimens cited. Taxus baccata L. (var.) cuspidata Carrière, Coniferae 733. 1867; T. baccata subsp. cuspidata (Siebold & Zucc.) Pilger,

Spjut, Taxonomy of Taxus

xd,

HERB PARIS

MUS.

Herbier de VAmetique septentrionak

Xvog& Micitatx,

pentia m

P AZ

(? fca

: ewe

ax me Ld RA

Fic. 51. Te I

ing (secund), closely overlapping, leaves.

minor. Canada, ex Herb. A. Michaux (holotype, P). Var. minor disti

276 Journal of the Botanical R h Institute of Texas 1(1)

Fics. 52A—C. Taxus cuspidata, Japan, original material (M). A (left).

Herbarium specimen with lectotype, the specimen on the right

(lower) with branchlets that may have been pendant. B (top).

ain

Lei f az Farro

r r7

ES e I 11 £ H J I

AA— MERI. ZECCA RIND,

Herbar. Univ. Ludov. Maximi

Y J

Planzenreich 4(5):112. 1903. Original material at M. Type: JAPAN: (Lectotype designated here: M!, sheet with the following data on

labels *Legit in Japonia et C Siebold, anno 1842," Herbarium Regium Monacense,” “Herb. Zuccarini,” and *Herbar. Univ.

Ludov. Maximil.," the specimen on the right of two mounted on the same sheet, with mature male cones; the left specimen also

identified as T. cuspidata; ISOLECTOTYPES: M!, K!, apparently from the same plant as lectotype but with other data on labels).

Taxus cuspidata is possibly native to Hokkaido, Japan, but apparently rare except perhaps in arboreta. Two

specimens that were collected by Jack from Hokkaido (A) are clearly related to the type as seen in phyllo-

taxy, color, and leaf anatomy. The leaf anatomical data in Spjut (2007) show the specimens to have a wider

abaxial margin, 16-24 cells wide, compared to a margin of 8-15 (-18) cells across in all other specimens

from the Sino-Japanese Region. These three specimens appeared to have the most conspicuous persistent

bud-scales of all yews in that region, a character trait that reinforces the choice of the lectotype.

The authorship and publication date for T. cuspidata have been confused in the literature. The official

date of the volume for Siebold and Zuccarini's Flora Japonica, in which T. cuspidata was intended to be

described as a new species, did not appear until 1870; yet, reference to this publication, including a figure

(Tab. 129 instead of Tab. 128), was first made by Siebold and Zuccarini (1843) 27 years earlier at which time

they cited the name in footnote with further reference to a description in Thunberg (1784, Flora Japonica).

Siebold and Zuccarini's (1846) synopsis of plant taxa in Japan, which many authorities cite as the valid pub-

lication date (e.g., Farjon 1998; Fu et al. 1999), does not provide a plant description of T. cuspidata; only an

indirect reference to T. baccata in Thunberg, Flora Japonica (1784). Aside from Siebold and Zuccarini (1843),

the first direct description of T. cuspidata appeared in Endlicher (1847). Others have since also provided a

description for the species (e.g., Carrière 1855a; Lawson et al. 1851; Parlatore 1868) before Miquel published

Spjut, Taxonomy of Taxus 277

Siebold and Zuccarini (1870), in which these earlier references were noted. However, Siebold and Zuccarini’s

(1843) earlier reference to a description in Thunberg (1784) satisfies the requirement for valid publication

of “Taxus cuspidata Sieb. & Zuccar.” (Art. 32.1), indicating also the origin of the epithet (“cuspidatis”).

The leaves of T. cuspidata overlap along branchlets with an orientation that varies according to the

direction of the branchlet, appearing partially erect and slightly radial as in T. umbraculifera, or two-ranked

as in T. biternata. Indeed, the phyllotaxy of the type appears intermediate between that of T. umbraculifera

and T. biternata. Other specimens assigned to T. caespitosa var. latifolia are similar to T. cuspidata in their

long flexuous branchlets (e.g., Makino 43769 from Honshu, Maximowicz from Manchuria, Dvorakovskia &

Bokina from Sakhalin Is.).

Cultivars that I have studied at the Secrest Arboretum in Ohio (Chadwick & Keen 1976) are identified

as belonging to this species by the dome-like to pyramidal crown with a definite leader, and by the branches

also with a definite leader that ascend upwards from which hang many long simple branchlets. In other

cultivars the branchlets are stiff and recurved. The habit is reminiscent of the European Dovaston yew (T.

baccata var. dovastoniana Leighton)—recognized for its weeping branches—and to other European plants

known in horticulture as T. baccata var. glauca Carriere (US: 1396503 “f. glauca = f. subglaucescens Jacq.;” ex

Hillier’s Arboretum BM; ex Herb. Petropolitani, Szovich 610, S; C. Baenitz US, “f. pendula"). Thus, I wonder if T.

cuspidata is of horticultural origin. Wilson (1916) commented that he was not sure whether the yew plants

he saw in Japan were natural or cultivated, and since Siebold and Zuccarini (1870) noted that yew there

occurred in horticulture, in cultivation around temples, and spontaneously in the mountains, it is possible

that the original material came from horticulture. Also, it is interesting to note that Carriere (1861) thought

that the Dovaston yew was native to Japan.

This problem is complicated by naturally occurring intermediates between the Baccata and Cuspidata

Alliances as evident from anatomical data and other key characters. Examples of intermediates that are

included in the Baccata Alliance are from Sweden (Bjornstrorn, ex Mus. Stockholm), Finland (Finlandee Exsic.

419 K, p.p., bottom specimen), and the Caucasus Mountains (Busch s.n.). These are not easily distinguished

from occasional ones in the Cuspidata Alliance such as from Hokkaido (Makino 43769), Korea (Faurie s.n.

A, Wilson 9484), and Sakhalin Island (Dvorakovakaia & Bohina). Being able to discern the key differences

among these problematic Eurasian specimens requires considerable familiarity with Taxus.

Another example is Hayek & Hayek s.n. (BM), discussed earlier under T. canadensis, from Styria Superior

in Austria (Spjut 2007, appendix), that seems to differ from T. cuspidata only by its dull olive green color,

in contrast to a yellowish orange color on branchlets in the E Asian specimens. From a taxonomic point

of view, the majority of the specimens fall within the Baccata and Cuspidata Alliances. However, in further

consideration to the distribution of T. canadensis in the Euro-Mediterranean Region and eastern North

America, the Baccata and Cuspidata Alliances were possibly derived from an ancestral boreal complex that

was perhaps distributed from northeastern North America to northeastern Asia. Thus, these intermediates

could be relicts of that former complex.

24. Taxus umbraculifera (Siebold ex Endl.) [Ravenscroft] [C.] Lawson, Abietineae—List Pl. Fir Tribe 10:80.

1851. (Figs. 9, 53—54). Cephalotaxus umbraculifera Siebold ex Endlicher, Syn. Conif.239. 1847. Taxus cuspidata Siebold &

Zucc. vat. aet ifera, Makino in Makino & Nemoto, Cat. Jap. pl. 407. 1914 (Also, Illus. Fl. Nippon. 1925 typographical error,

spelling “ambraculifera,” see ICBN Art. 61 and corrected spelling by Makino, Illus. Fl. Nippon 910, fig. 2730. 1931). Type: JAPAN:

(NEOTYPE designated here: illust. in Makino, Illus. Fl. Nippon, fig. 2730. 1931.). Eprrype designated here: JAPAN. HonsHu: Hyogo

Pref., Mt. Hyonosen, 30 Aug 1948, Muroi 30, ex Herb. Hiroshi Muroi, Kobe, identified by Muroi as T. cuspidata var. ambraculifera

Makino (A!). Original material unknown.

The origin of the name T. umbraculifera may be horticultural (Gordon & Glendenning 1858, Gordon 1875);

however, Cephalotaxus umbraculifera was attributed to Siebold by Endlicher (1847) who provided a detailed

description but expressed doubt as to whether the species was distinct from Taxus cuspidata. Ravenscroft

is recognized in Stafleu and Cowan (1983) as the contributor for Lawson et al. (1851), who attributed C.

umbraculifera to Siebold and Zuccarini, and indicated it was a synonym of T. cuspidata; however, the au-

thority for the name is indicated to be Lawson because lack of internal evidence in Lawson et al. (1851) for

278

; UW. BINENSIS LOGer.) PAE / 7 AH’ Bi

ERZ,

¿5,2730 mM

3 y

Taxus cuspidata Sieb. et Zua

var. umbraculifera Moli

E Cophaltazas umbraculife

va Sieb.; T. adpressa viri)

T ice. et Z. var. nana Rod,

BK LINE - SS + B vr «x o x

i-r de re v Fe, 3. E 4

10 11 | 12 | 18 14 105

Herb. Hiroshi Muroi, Kobe, Japan.

va. JU

Taxus ‘cuspidata Sicboid «t Zuctavine ©

cbvacalifera HAKIN

EET pku

Fics. 53—54. Taxus umbraculifera var. umbraculifera. 53 (top). Neotype, Makino, Illus.

FI. Nippon (1931, Fig. 2730). 54 (above). Epitype, Japan, Honshu, Muroi 30 (A). Var.

bent downwards at petiole, the leaves also appearing in a d t

r J J

when looking down from apex of branchlet.

24a. Taxus umbraculifera var. umbraculifera

Common name.—Umbrelliform yew.

Distribution —Japan, Russian Federation—Manchuria.

fal, Dat A ID hi Pr

Journal of

of Texas 1(1)

Ravenscroft as the authority, ICBN Art. 35.5.

The epithet suggests an umbrella-like leaf

arrangement (Fig. 59), not the two-ranked

leaves as described by Endlicher (1847). Ra-

venscroft (Lawson et al. 1851), who treated

the species in section Cephalotaxus of Taxus,

indicated that branches are verticillate with

"distichous" branchlets. I have not seen any

original material, and herbaria that I have

contacted in this regard for Siebold speci-

mens reportedly have none. Nevertheless, an

illustration in Makino (1931) clearly depicts

the umbrelliform leaf arrangement that is

distinctive for this species. In later manuals

on the flora of Japan (e.g., Ohwi 1965), var.

nana Rehder (1902) became the name for

this taxon, but this is antedated by var. mi-

crocarpa Trautvetter (in Maximowicz 1859);

however, these names are applied to different

(and distinct) taxa in the present study.

The leaves of T. umbraculifera appear

in star-like (almost decussate) manner

when looking down the branch from apex,

and unlike T. caespitosa, the blades of T.

umbraculifera will face different directions.

Leaves appear perpendicular to the plane

along one side of a branchlet and in the same

plane along another side of branchlet. This is

not easily determined in pressed specimens;

however, the crisscrossed blades partly re-

flect this, which is in contrast to the radial

orientation of leaves in T. caespitosa that all

twist in the same manner, appearing imbri-

cate when observed from above the apex of

the branchlet. The leaves of T. umbraculifera

are also sharply reflexed at their petioles,

in contrast to bending upwards (erect) in T.

caespitosa; this difference accounts for the

two-ranked appearance in T. umbraculifera,

in contrast to the secund appearance in T.

caespitosa often seen on older branchlets.

Four varieties of T. umbraculifera are

recognized by differences in habit and leaf

arrangement.

Spjut, Taxonomy of Taxus 279

Additional specimens: JAPAN. Honshu: Kanagawa Pref., Mt. Ooyama, shrub 2 m high, cult., 25 Oct 1952, Suzuhi 499003 (A); Shunane

Pref., Mt. Sentsu-zan, Naito s.n. (A). RUSSIAN FEDERATION. Mandshuria SE: Ex herb. hort. bot. Petro. yr 1860, Maximowicz (S).

Variety umbraculifera appears mostly arborescent with wide spreading branches. It is recognized by the radial

distribution of leaves on erect branchlets, especially near apex, and the appearance of a decussate to nearly

two-ranked arrangement on horizontal branchlets.

24b. Taxus umbraculifera (Siebold ex Endl.) Ravenscroft var. hicksii (Hort. ex Rehder) Spjut, comb. nov.

Basionyn: Taxus media Rehder f. hicksii (Hort.) Rehder [“T. cuspidata hicksii Hort." in synon.], J. Arnold Arbor. 4:108. 1923. Taxus

cuspidata var. hicksii (Hort. ex Rehder) Bailey, Cult. Evergreens 189. 1923. Type: U.S.A. New York: horticultural specimen from Hicks

Nursery, Westbury, Long Island, 28 Sep 1922, Arnold Arboretum 8036 (HoLotyPE: A!). Bailey (1923) attributed the combination to

Rehder even though it has not been determined whether Baileys publication predates that of Rehder (8 May 1923). All conditions

for valid publication are met in Rehder (1923) but not in Bailey (1923); therefore, the name in Rehder (1923) is considered the

basionym (see ICBN Art. 45.1) even though Rehder cited a synonym used in horticulture without reference to an authority or pub-

lication and implied that it was a basionym.

Common name.—Hicks yew.

Distribution.—Endemic to Japan.

Additional specimens: JAPAN. Iwate-Pref.: Asagishi, Muroi 3593 (A). Nagano Pref.: Kamikochi, Muroi 3715 (A). Gifu- Pref.: Takayama,

Muroi 3698 (A). Hyogo Pref.: Kumatugi, Mikata-gun, Muroi 5603 (A); Mt. Hatibuse Muroi 5424 (A); Wakasugi, Muroi 5648 (A).

CULTIVATION: Maryland: Laurel, residential area, 8495 Imperial Drive, Spjut s.n. (wba); Arnold, residential area, 757 Dunberry, 1

Aug 1999, Spjut s.n. (wba).

Variety hicksii is distinguished by its columnar habit with erect branchlets and erect linear leaves that spread

in a radial arrangement. The leaves may appear two-ranked on lower branchlets, but if the branchlets are

turned over, the underside will be seen to have some of the leaves reflexed.

Although described from horticulture, this variety appears to occur naturally in Japan, based on four

specimens from there that are remarkably similar to the type. It also raises the question as to whether the

variety independently evolved there as a hybrid, or whether it may have been introduced into Japan from

North America, since the specimens cited above were collected during the mid 1950s—after the Hicks yew

was described by Rehder (1923). The Hick’s yew supposedly originated from seed of “T. cuspidata Nana”

sometime around 1900 (den Ouden & Boom 1965). Molecular data in Collins et al. (2003) placed the Hick’s

yew with T. x media Rehder; however, their study did not include morphological characters.

24c. Taxus umbraculifera var. microcarpa (Trautv.) Spjut, comb. nov. (Figs. 55-58). Basionvw: Taxus baccata L.

var. microcarpa Trautvetter in Maxim., Mem. Acad. Sci. St. Petersb. Sav. Etrang. 9:259 (Prim. Fl. Amur.). 1859. Taxus cuspidata Siebold

& Zucc. var. microcarpa (Trautv.) Kolesnikov, Bull. Far E. Branch Acad. Sci., USSR 13:43, fig. 2. 1935. Taxus cuspidata Siebold &

Zucc. var. microcarpa (Trautv.) S-Y. Hu, Taiwania 10:21. 1964 (nomen comb. supertL., illegit.). Type: CHINA. MaNcuuria: Heilongjiang,

Exped. Soc Geogr., 1 May 1855, Maack s.n. (Lectotype designated here: GH!, lower of two specimens; isolectotype: P!, upper of two

specimens on one sheet). Original type not specified; original material (syntypes) from several locations, one from Manchuria (1 May

1855, reportedly sterile, GH! P!), and another from Sakhalin Is. (Weyrich, Sep. 1853, with fruit, KFTA?).

1 PE

LAXUS

ifera subsp. laxa Spjut ined. Type: KOREA. Ooryong-too (Oagelet Island), 0-900 m, bush or small tree, common, Wilson

8538 (HOLOTYPE: A!; ISOTYPE: US!).

Common name.—Small seed yew.

Distribution.—NE temperate Asia.

Additional specimens. RUSSIAN FEDERATION. Pri ie [Pri kiy] Prov.: vicinity of Vladivostok, Palczevsky 3601 (A, K, US); Bay

of Peter, the First Sea Reserve, Island of Stenin, 26 Apr 1979, Kypehinova s.n. (in Russian, A). Manchuria Region: Korea septentrionalis

provincia Pen-nian Muorum Jahn...Frajectus Lpatan-ien, 27 Jun 1897, Komarov 88 (A); Rossica, Aultzo Prov., Ussuri [Ussuzieusis] insula

Afnold, Komarov 88 (P); Manchuria: Rossica Palczevsky [Komarov] 88 (BM, K, US [ex Herb. Baenitz]). SOUTH KOREA: Nemon-rei,

Kyongsan, tree up to 50 ft, common, 12 Oct 1917, Wilson 9332 (A); Herb. Lugd. Batv. (P); Zuccarini 593, in adnot. T. baccata (M). JAPAN.

Hokkaido: Ishikasi Prov., Apr 1884, K. Muijabe s.n. (A). Nanokwa, Tosa, 18 Apr 1888, Watanabe s.n. (A); Sapporo, Agric. College, 15 Jun

1885 (A), Jun 1878 (A); Sapporo, Siebold, ex Herb. Zuccarini (GH); Mt. Nantai, Lake Chuzenji, 20 Aug 1904, Mochizuki s.n. (A). Honshu:

Yokohama, yr 1862, ex Herb. USDA 1888 (US: top specimen); Japan, no locality or collector data (US 1311889); Hida, Takayama, 17 Sep

1910 (A); Yokohama, yr 1862, ex Herb. USDA 1888 (P, p.p.; US, lower of 2 specimens); no locality data, Kiaraboku s.n. (US 1311889).

This variety is best recognized by isodichotomous branching, the oblong leaf shape (ca. 8x l/w) and crisscross

Fics. 55-58. Comparison of illustrations with herbarium

specimens of Taxus umbraculifera var. microcarpa. 55 (upper

left). Illustration from Kolesnikov (1935, as 7. cuspidata var.

microcarpa).56 (upper right). Herbarium specimen, Manchuria,

Palczewski ex Herb, Baenitz (US). 57 (center). Illustration show-

ing habit of plant from side view (Kolesnikov 1935), appearing

radial in outline from top view (not shown). 58 (right). Type

herbarium sheetin the Gray Herl , Maack Exped. 1855 (GH)

The upper specimen annotated by S-y. Hu— T. cuspidata var. mi-

rrorcarnn.

LrULUI DU Jt Jj V 1

however, the speci that | with Trautvetter's

description of T. baccata var. microcarpa (basionym), a low shrub

1109C) icth

e lower

specimen (lectotype). The upper specimen is identified in this

publication as 7. biternata, described as a new species, a tree

J'ag

Piba Dat o ID AKI

Journal of

". eer P

ae Y ee

v VIII ZITAT T fes

PW enr AE

VoL EE m fer a ee

-—

Pl {Core

+ Di e cazpa

Herb. Acad. Petrop.

E ain A As for

amos feno DELTA TEO Fhe

: Mandshuria, ad fl. Amur. Exped. soc. geogr.

| ; leg. R. Maack 1855.

Spjut, Taxonomy of Taxus 281

leaf arrangement. Trautvetter (in Maximowicz 1859) distinguished var. microcarpa from T. baccata by the

smaller—wider than tall—seed (Pilger 1903), and Kolesnikov (1935) further indicated it was a low rounded

shrub—0.5-1.5 m high and 5-7 m in diam.—that reproduced by layering (Fig. 68). Kolesnikov (1935) had

recognized two morphological different varieties (tree and shrub yews) occurring in different habitat types

in southeastern Russian Federation.

In my 1996-1997 annotations (Taxus microcarpa [Trautv.] Spjut ined.; Taxus umbraculifera subsp. laxa

Spjut ined.), I had concluded that a specimen annotated by S-y. Hu in GH was the type for T. baccata var.

microcarpa (see Fig. 66 and Hu 1964), but upon later reviewing the illustrations in Kolesnikov (1935), repro-

duced here in Figs. 63-64, it was quite clear this was not its type but rather a specimen below it on the same

herbarium sheet. The interpretation of the taxon by Kolesnikov (1935), based on Trautvetter (in Maximowicz

1859), must be preserved; therefore, T. biternata is described as a new species, and *T. umbraculifera subsp.

laxa Spjut” will remain an unpublished name.

24d. Taxus umbraculifera var. nana (Rehder) Spjut, comb. nov. (Figs. 59). Basionvm: Taxus cuspidata var. nana

[Hort. ex] Rehder in L.H. Bailey, Cyclopedia Amer. Hort. 1773. 1902. Taxus cuspidata f. nana (Rehder) Wilson, Conif. Taxads Japan

13. 1916. Type: JAPAN. HonsHu: Pref. Hyogo, Mt. Hyonosen, 1100-1400 m, in don forest with sasa thicket, on ridge, shrub 2 m,

fr red, 11 Aug 1983, Murata 44671, det. as T. cuspidata var. nana Rehder ignated here: A!). Described from horticulture

no specimens cited and original material unclear.

Common name.—D warf yew.

Distribution.—E Russia (islands), China (Shanxi), Japan.

Additional specimens: RUSSIAN FEDERATION. Far East: Primorskiy Region, Sea Reserve, Island Bolshoy Pelis, Borzova s.n. (in Rus-

sian, A); Sakhalin, ex herbario horti Petropolitani, Augustinowiez, T. baccata var. microcarpa, Schmidt, p.p., with T. caespitosa var. caespitosa

(A). CHINA. Shanxi: (Schenhsi merid"): Taipei-schan, 1936, G. Fenzel 972 (A). JAPAN. Mt. Fujiwara Mie pref., Muroi 1969 (A); Mt.

Himekami, Iwate pref, 14 Sep 1955, Muroi 5933 (A); Japan, no additinal data, Faurie s.n. (P). CULTIVATION. U.S.A. Ohio: Secrest

Arboretum, cultivars ‘Newport,’ ‘Hatfield,

The epithet “nana” implies a dwarf plant, and Rehder (1902) described T. cuspidata var. nana as a “dwarf

compact form with shorter leaves” in regard to a horticultural plant. Rehder (1949) later considered it only a

form. Nevertheless, others applied the varietal name to native plants in Japan. The plants were characterized

as low shrubs with a radial orientation of leaves, found mostly along the seaside of Japan (Ohwi 1965).

Variety nana is a low, densely branched shrub with oblong leaves that are mostly radial and crisscross-

ing in herbarium specimens, appearing dark glossy green above (adaxial surface) and paler below (abaxial

surface).

ACKNOWLEDGMENTS

I thank Drs. Roy Vickery and Charlie Jarvis for making specimens and rare books available at BM, providing

photocopies of literature, and allowing specimens to be photographed; Ph. Morat at P for making avail-

able their specimens and allowing them to be photographed; Arne Anderberg at S for his communications,

hardcopies from the online S database of Taxus specimens, and a loan to US; Zi-yu Cao at PE for photocopies

and leaves of the isotype of T. yunnanensis and for correspondence; Umberto Mossetti at BOLO for provid-

ing leaves of their type of T. orientalis and its illustration by Bertoloni; and Annalisa Managlia for a photo of

its holotype; Amin at the Kunming Institute of Botany for providing leaf framgments of Taxus contorta and

photos of herbarium specimens of this species from Pakistan, and mention of their investigative work on

molecular data regarding T. baccata, T. contorta, T. wallichiana and other species of Taxus in China; Patricia

Holmgren for providing photocopies of types of T. wallichiana at NY; and Alfred Schuyler for allowing me to

study specimens at PH. Similarly, I am grateful for the assistance from Peter Edwards at K during my visit;

Peter Mazzeo (retired) at NA for obtaining loans from A, BH, BM, GH, K, M, and U; George Russell and

Katherine Rankin at US for obtaining and maintaining loans from S, and for study of their collections; and

curators at E who provided a loan of original material of T. lindleyana.

Thanks also to Dan Nicolson for helpful discussions on nomenclatural matters; John Thieret for his twice

282 Journal of the Botanical R h Institute of Texas 1(1)

Mh nda. VECI

{ Dare ett 3

Dee gp el

ecce P4

: ^

RU OEM A COA

PLANT OF JAPAN

Taxus cuspiónta Sieb. et Zucc

var, nana Rehder

Fics. 59A—B. 7c b life nana, Mt. Hyonosen, Japan (neotype, A). A (left)

plant. B (right). Close-up of t l ill i

L H J

T J F

generous editorial review and comments on a related manuscript, Berthold Heinze for reprints on Austrian

Taxus, including both German and English languages, Guy Nesom for his critically helpful comments, and

similarly Zlatko Kvaček; the National Agricultural Library for assistance in obtaining rare journals; the Library

of Congress in Washington, D.C.; the Missouri Botanical Garden Library staff for copies of select pages from

references on literature of China, and other journals; Aljos Farjon at the Royal Botanical Gardens at Kew for

providing copies of papers in journals not readily available in US libraries; the USDA Pacific Northwest Re-

search Station and USDA Forest Service in that region for providing fresh material of T. brevifolia and for field

assistance; Kenneth Cochran at the Secrest Arboretum for field assistance and providing specimens; and John

Wiersema at the USDA Systematic Botany Laboratory in Beltsville for discussions on nomenclature in the early

stages of this paper. Further acknowledgment is noted here to the New York Botanical Garden for providing

images of type collections online, making it possible to easily study and compare handwriting of collectors

such as David Douglas and Thomas Nuttall, in addition to being able to retrieve data on type collections.

REFERENCES

ALLISON, T.D. 1991. Variation in sex expression in Canada yew (Taxus canadensis). Amer. J. Bot. 78:569-578.

Spjut, Taxonomy of Taxus 283

Anonymous. 1913. XLIII—The Wallichian Herbarium. Kew Bull. 1913:255-263.

APPENDINO, G. 1995. The phytochemistry of the yew tree. Nat. Prod. Rep. 12:349-360.

Arno, S.F. and R.P. HauMERLY. 1977. Northwest trees. The Mountaineers, Seattle.

Baitey, L.H. 1902. Cyclopedia of American horticulture, R-Z. MacMillan Co., New York.

Bailey, L.H. 1923. The cultivated evergreens. MacMillan Co., New York.

BAILEY, L.H. 1933. The cultivated conifers of North America. MacMillan Co., New York.

Bannan, M. W. 1942. Notes on the origin of adventitious roots in the native Ontario conifers. Amer. J. Bot.

29:593-598.

BEAN, JW, 1953. Trees and shrubs hardy in the British Isles—Vol. Ill (7^ ed). John Murray, London.

BEISSNER, L. 1891. Handbuch der Nadelholzkunde. Verlag von Paul Parey, Berlin.

BERTOLONI, A. 1862. Miscellanea botanica XXIII. Mem. Acad. Sci. sér 2::228-229, tab.

BERTRAND, C.E. 1874. Anatomie comparée des tiges et des feuilles chez les Gnétacées et les Coniféres. Ann. Sci.

Nat. Bot. sér. 5, 20:6-153.

Best, C., J.T. HoweLL, W. KNIGHT, |. KNiGHT, and M. WeLLs. 1996. Flora of Sonoma County. CNPS Press.

BoLsiNGER, C.L. 2007. Personal communication. Occurrence of thicket yew (Taxus brevifolia var. reptaneta) in the

Cascade Ranges of California, near Mt. Shasta.

BorsiNGER, C.L. and A.E. JARAMILLO. 1990. Taxus brevifolia Nutt. Pacific Yew. In: R.M. Burns and B.H. Honkala, eds. Silvics

of North America, conifers, l:573-579. USDA For. Serv. Hdbk. 564.

Browicz, K. and J. ZIeELINsK! 1982. Chorology of trees and shrubs in South-West Asia and adjacent regions. Polish

Scientific Publishers, Vol. 1:17, Warszawa.

Brummit, R.K. and C.E. Powett (eds.) 1992. Authors of plant names. Royal Botanic Gardens, Kew.

BurkHILL, I.H. 1965. Chapters on the history of botany in India. Botanical Survey of India, Calcutta.

CALIFORNIA Native PLANT Society (CNPS). Website, accessed March 2006. http://www.marin.cc.ca.us/cnps/.

CALLEN, G. 1977. Les Conféres cultivés en Europe. Vol. Il, ed. J. B. Bailliére, Coll. Tech. Hort. Spéc.

Camerort, H. 1956. Etude de la structure du point végétatif et des variations phyllotaxiques chez quelques gym-

nospmeres. Ann. Des Sci. Nat. Bot. sér. 17(1):103-120.

CARRIERE, E-A. 1855a. Traité general des Coniféres. Chez l'auteur, Rue de Buffon, Paris.

CARRIERE, E-A. 1855b. Biota orientalis, Biota pyramidalis, et Taxus adpressa. Rev. Hort. (1855):93-96.

CARRIERE, E-A. 1861. If de Dowaston. Rev. Hort. (1861):175-176.

Carritre, E-A. 1867. Traité general des Coniféres. Nouvelle Ed. Chez l'auteur, Rue de Buffon, Paris.

CHADWICK, L.C. and R.A. Keen. 1976. A study of the genus Taxus. Ohio Agric. Res. Bull. 1086.

CHAMPION, H.G. and S.K. SetH. 1968. A revised survey of the forest types of India. Manager of Publications Delhi,

India Press, Nasik.

CHANSLOR WETLANDS WILDLIFE Proyect, Website. Accessed March 2006. Plants of the Salmon Creek Watershed. http://

www.sonomawetlands.org/Plants-Trees-Fix3.htm.

CHENG (ZHENG), W-c., L-K. Fu, and C-y. CHENG. 1975. Gymnospermae Sinicae. Acta Phytotax. Sin. 13(4):56-90& 66

illus., 17 plates.

CHENG (ZHENG) W-c., and L=. Fu. 1978. Taxaceae. In: Fl. Reipub. Pop. Sin. 7, Gymnospermae. Agendae Academiae

Sinicae. [English Translation].

CHUNG, T.H. 1965. Illustrated encyclopedia of fauna and flora of Korea. Vol. 5. Tracheophyta. Ministry of Education,

Plant Series, Samhwa Publ. Co. [In: Korean except for Latin names and English Forward].

CITES [CONVENTION ON INTERNATIONAL TRADE IN ENDANGERED SPECIES OF WILD FAUNA AND FLORA]. 2001. Review of the genus

Taxus. Eleventh meeting of the plants committee Langkawi (Malaysia), 3-7 Sep."Document prepared by the

United States of America" PC 11 Doc 22.5 pp. http//www.cites.org/eng/com/pc/1 1/E-PC1 1-22.pdf.

CITES [Convention on International Trade in Endangered Species of Wild Fauna and Flora]. 2004. Amendments to

Appendices | and Il of CITES. Thirteenth Meeting of the Conference of the Parties, Bangkok, Thailand, October

2-14, 2004. http://www.cites.org/eng/cop/1 3/prop/E1 3-P48.pdf; http://www.iucnredlist.org/search/details.

php/34041/all

284 Journal of the Botanical R h Institute of Texas 1(1)

Couns, D., R.R. Mitt, and M. MoLLer. 2003. Species separation of Taxus baccata, T. canadensis, and T. cuspidata (Taxa-

ceae) and origins of their reputed hybrids inferred from RAPD and cpDNA data. Amer. J. Bot. 90:175-182.

Cort, E.A. 1998. Taxaceae: The genera and cultivated species. Bot. Rev. 64:291-322.

CORRADINI, P, C. EDELIN, A. BRUNEAU, and A. BOUCHARD. 2002. Architectural and genotypic variation in the clonal shrub

Taxus canadensis as determined from random amplified polymorphic DNA and amplified fragment length

polymorphism. Canad. J. Bot. 80:205-219.

Croom, E.M., JR. 1995. Taxus for taxol and taxoids. In: M.S. Suffness, ed. Taxol. Science and applications. CRC Press.

Pp. 37-70.

Da CHENG, H. 2007. Personal communication. Unable to extract DNA from 1-15 yr old herbarium specimens of

Taxus globosa var. floridana, T. brevifolia var. brevifolia, var. polychaeta, and var. reptaneta. Chinese Academy of

Sciences, Dalian Institute of Chemical Physics, Biotechnology, Dalian, China.

Dempsey, D. and |. Hook. 2000. Yew (Taxus) species— chemical and morphological variations. Pharm. Biol.

38:274-280.

DERYUGINA, T.F. and N.D. NesterovicH. 1981. Peculiarities of the morphological and anatomical structure of conifer

needles of some Taxus L. species. Dokl. Akad. Nauk Bel. S.S.R. 25(7):652-655 [Russian with English summary].

Doene, D.L., E. CARROLL, R. WesTFALL, R. MILLER, J.H. KITZMILLER, and K.M.B. SNaDeR. 1993. Geographic variation in allozymes

and taxol, and propagation by root cuttings in Pacific yew. In: International yew resource conference. Yew

(Taxus) conservation biology and interactions, Berkeley, CA (abstract). Pp. 10.

Durrev, E. 1980. Ecological history of yew Taxus baccata in southern England. Biol. Conserv. 17:243-265.

ELIAS, T.S. and V. KonzHENEVskv. 1992. The presence of taxol and related compounds in Taxus baccata native to the

Ukraine (Crimea), Georgia, and southern Russia. Aliso 13:463-470.

Eur, W.A. 1938. Forest trees of the Pacific Coast. Putnam, NY.

EL-Kassagy, Y.A. and A.D. YancHuk. 1994. Genetic diversity, differentiation, and inbreeding in Pacific yew from British

Columbia. J. Hered. 85:112-117.

EL-KassaBy, Y.A. and A.D. YANCHUK. 1995. Genetic variation of Pacific Yew in British Columbia and its conservation.

In: Ph. Baradat, W.T. Adams and G. Müller-Starck, eds. Population genetics and genetic conservation of forest

trees. Academic Publishing, Amsterdam, The Netherlands. Pp. 227-235.

Ewes, JW. and A. Henry 1906. The trees of Great Britain and Ireland. Priv. Printer, Edinburgh.

ENDLICHER, S. 1847. Synopsis coniferarum. Scheitlin & Zollikofer, Sangalli.

FARJON, A. 1998. World checklist and bibliography of conifers. The Royal Botanic Gardens, Kew.

FARJON, A. 2001. World Checklist and Bibliography of Conifers. 2nd edition. The Bath Press, Bath, United

Kingdom.

FARJON, A. and C.N. Pace (Compilers). 1999. Conifers. Status survey and conservation action plan. Species Survival

Commission Conifer Specialist Group, International Union for Conservation of Nature and Natural Resources,

Gland, Switzerland. Frequenbtly cited as Conifer Specialist Group 1998 with further reference to data collec-

tion forms for conifer species completed by the SSC [Species Survival Commission] Conifer Specialist Group

between 1996 and 1998. IUCN, Gland, Switzerland and Cambridge, UK. 121 p.

FLORIN, R. 1931. Untersuchungen zur Stammesgeschichte der Coniferales und Cordaitales. Kongl. Svenska Veten-

skapsakad. Handl. 10:45—588.

FLORIN, R. 1948a. Enumeration of gymnosperms collected on Swedish expeditions to western and northwestern

China. Acta Horti Berg. 14:343-384.

FLORIN, R. 1948b. On the morphology and relationships of the Taxaceae. Bot. Gaz. 110:31-39.

FLORIN, R. 1948c. On Nothotaxus, a new genus of the Taxaceae, from eastern China. Acta Horti Berg. 14:385-395,

plates 1-3.

FRANCO, J.A. 1964. Taxus. Fl. Europaea 1:39.

FRANKLIN, J.F., and C.T. Dyrness. 1969. Vegetation of Oregon and Washington. USDA For. Res. Pap. PNW-80.

FREITAG, H. 1971. Studies in the natural vegetation of Afghanistan. In: PH. Davis, PC. Harper, and I.C. Hedge, eds.

Plant life of south-west Asia, Bot. Soc. Edinburgh. Pp. 89-106.

FRIMMEL, F. VON. 1911. Die untere Kutikula des Taxus-Blattesein Lichtreflektor. Oesterr. Bot. Z. 61:216-223.

Spjut, Taxonomy of Taxus 285

Fu, L-k, N. Li, and R.R. Mitt. 1999. Taxaceae. Fl. China 4:89-96, Missouri Botanical Garden Press, St. Louis.

GADEK, PA., C.J. Quinn, and A.E. AsHrorb. 1984. Localization of the biflavonoid fraction in plant leaves, with special

reference to Agathis robusta (C. Moore ex F. Muell) Em. Bail. Austral. J. Bot. 32:15-31.

Game e, J.S. 1922. A manual of Indian timbers. Sampson Low, Marston & Co, London.

GENDEREN, H. van, L. M. SCHOONHOVEN, and A. Fuchs. 1996. Chemisch-ecologische flora van Nederland en Belgiéeen

inleiding over aard en ecologische betekenis van secundaire plantestoffen, KNNV Uitgeverij, Utrecht.

Gopwin, H. 1975. History of the British flora (2? ed.). Cambridge University Press.

GOODMAN, J. and V. WatsH. 2001. The story of taxol. Nature and politics in the pursuit of an anti-cancer drug.

Cambridge University Press.

Gorpon, G. 1875. The pinetum: being a synopsis of all the coniferous plants at present known, with descriptions,

history and synonyms Qnd ed). Henry C. Bohn, London.

GORDON, G. and R. GLENDENNING. 1858. The pinetum: being a synopsis of all the coniferous plants at present known,

with descriptions, history and synonyms. Henry C. Bohn, London.

GRAUSTEIN, J.E. 1950/51. Nuttall's travels into the old Northwest. Chronica Bot. 14 (1—2). i-vii, 1-86, 9 plates and

9 pp of captions, index.

GREUTER, W. J. MCNEILL, ER. Barrie, H.M. Burbet, V. Demouuin, T.S. FiLGUEIRAS, D. NicoLson, PC. Sitva, J.E. Sk0G, P. TREHANE,

N.J. TURLAND, and D.L. HAwkswoRrH. 2000. International code of botanical nomenclature (Saint Louis Code) ad-

opted by the 16^ International Botanical Congress. St. Louis, MO, July-August 1999. Koeltz Scientific Books,

Königstein, Germany.

Grierson, AJ.C. and D.G. Lone. 1983. Flora of Bhutan. Vol 1. Pt. 1. Royal Botanic Garden, Edinburgh.

GRIFFIN, J. and |. Hook. 1996. Taxol content of Irish yews. Planta Med. 386: 370-372.

GRIFFITH, W. 1854. Notulae ad plantas asiaticas. Posthumous papers bequeathed to the Honorable The East India

Company, arranged by John M'Clelland, Bishops’ College Press, Calcutta. Vol. 4.

HAGENEDER, F. 2007. Yew. A history. Sutton Publishing Ltd., Thrupp-Stroud-Goucestershire.

HaNpEL-Mazztrr, H. 1929. Symbolae Sinicae. Julius Springer, Wien.

Hara, H., W.T. STEARN, and L.H.J. WiLLIAMs. 1978. An enumeration of the flowering plants of Nepal. Vol. 1. British

Museum, London.

Heinze, B. 2004. Zur Populationsbiologie der gemeinen Eibe (Taxus baccata). Centralblatt für das gesamte Forst-

wesen 121:47-59.

Hertel, H. von and N. Koutstock. 1996. Genetische Variation und geographicsche Struktur von Eibenvorkommen

(Taxus baccata L.) in Mecklenburg-Vorpommern. Silvae Genet. 45:290—294.

Hier, N.T. 1998. The gymnosperms of Vietnam. In: Z. Aoluo and W. Sugong, eds. Proceedings of the First Interna-

tional Symposium on Floristic characteristics and diversity of East Asian plants, July 25-27, 1996, Kunming,

Yunnan. China Higher Education Press Beijing and Springer-Verlag Berlin Heidelberg. Pp. 91-103.

HiLFIKER, K., R. HOLDEREGGER, P. Rotach, and F. Guer. 2004. Dynamics of genetic variation in Taxus baccata: local

versus regional perspectives. Canad. J. Bot. 82:219-227.

Hitt, K.D. and LJ. Scriven. 1998. The fossil record of conifers in Australia. Fl. Australia 48:527-537.

Hits, M. 1993. Taxaceae Gray. Yew family. Fl. North America 2:423-427.

HISTORYLINK.ORG. 2007. The Online Encyclopedia of Washington State History, Essay 7298. http://historylink.org/es-

says/output.cfm?file id—7298.

HitcHcock, C.L., A. CRonauist, M. Ownsey and J.W. Thompson. 1969. Vascular plants of the Pacific Northwest I. Univeristy

of Washington Press, Seattle.

Hook I., C. Poupat, A. AHOND, D. GUENARD, F. GuenrrTE, M-T. ADELINE, X-P. WANG, D. Dempsey, S. BREVILLE and P. Potier. 1999.

Seasonal variation of neutral and basic taxoid contents in shoots of European Yew (Taxus baccata). Phyto-

chemistry 53:1041-1045.

Hou, H-Y. 1983. Vegetation of China with reference to its geographical distribution. Ann. Missouri Bot. Gard.

70:509-548.

Hu, S-y. 1964. [Notes on] the Flora of China. Taiwania:1 3-62.

286 Journal of the Botanical R h Institute of Texas 1(1)

IUCN. 2004. Red List of Threatened Species. With reference to Conifer Specialist Group 1998. www.iucnredlist.

org [updated 2006].

Jarvis, C.E., ER. Barrie, D.M. ALLAN, and J.L. ReveaL. 1993. A list of Linnaean generic names and their types. Regnum

Veg. 127.

JEAN, F.-l., F.-X. GARNEAU, G.J. COLLIN, and M. BouHase. 1993. The essential oil and glycosidically bound volatile com-

pounds of Taxus canadensis Marsh. J. Essential Oil Res. 5:7-11.

JINXING, L. and H. Yuxi (eds.). 2000. Atlas of structure of gymnosperms. Science Press, Beijing.

KHAN, N. U.-D. and N. Parveen, 1987. The constituents of the genus Taxus. J. Sci. Industr. Res. 46:512-516.

KINGSTON, D.G.l. 1996. Improving on nature. Taxane J. 2:20-27.

KINGSTON, D.G.l. 2005. Taxol and its analogs. In: G.M. Cragg, D.G.l. Kingston and DJ. Newman, eds. Anticancer agents

from natural products. CRC Taylor & Francis, Boca Raton, FL. Pp. 89-122.

KINGSTON, D.G.l., G. SAMARANAYAKE, and C.A. Ivey, 1990. The chemistry of taxol, a clinically useful anticancer agent. J.

Nat. Prod. 55:1-12.

KNIGHT, J. 1850. A synopsis of the coniferous plants grown in Great Britain and sold by Knight and Perry at the

exotic nursery, Kings Road, Chelsea. Longman, Brown, Green, and Longmans, London.

Koch, K. 1873. Dendrologie. Verlag von Ferdinand Enke (Erlangen).

KoLEsNiKOv, B. P. 1935. On the shrubby kind of the spiky yew (Taxus cuspidata S. et Z.). Bull. Far E. Branch Acad. Sci.,

USSR 31-47 (In Russian with English summary).

KORPELA, |. ILKKA KORPELAS INDEX, WEBSITE. 2006 (March). Photo of Taxus canadensis var. adpressa, referred to as T. bac-

cata, indicated to have been photographed in Estonia. Cited by WBA (www.worldbotanical.com) web site

since 2003. http://honeybee.helsinki.fi/users/korpela/taxus_baccata.html.

KRAUZE-BARANOWSKA, M. and M. Wiwanr. 2003. Antifungal activity of biflavones from Taxus baccata and Ginkgo

biloba. Z. Naturf. 58c:65—69.

KRUSSMANN, G. 1985. Manual of cultivated conifers, translated by M.E. Epp. H-D. Warda and G. S. Daniels, eds.

Timber Press, Portland.

KRUPKIN, A.B. 1994 (unpublished & pers. comm). Report on PCR results for selected samples of Taxus received by

Phyton Inc. from Sweden, Korea, China and The Philippines.

Kunz, H. 1937. A new and remarkable habitat for the endemic Florida yew. Torreya 27:90-92.

Kvacek, Z. [1984 fide author notation on reprint]. Tertiary taxads of NW Bohemia. 1982 Acta Univ. Carol., Geol.,

Pokorny 4:471-491.

Kvacek, Z., and W.C. RemBER. 2000. Shared Miocene conifers of the Clarkia flora and Europe. Acta Univ. Carol. Geol.

44:75-85.

Kvacek, Z., and W.C. Rember. Under Review. Calocedrus robustior (Cupressaceae) and Taxus howardii (Taxaceae):

Two new conifers from the Middle Miocene of the western North America (Clarkia area, Latah Formation,

northern Idaho). Paleobios.

Kwe, Y-L., and S-v. Hu. 1974. [Epidermal feature of leaves of Taxus in relation to taxonomy]. Acta Phytotax. Sin.

12(3):329-334, plate 67 [Chinese with English summary].

LAUBENFELS, D.J., DE. 1969. A revision of the Malesian and Pacific rainforest conifers, |. Podocarpaceae, in part. J.

Arnold Arbor. 50:315-369.

LAUBENFELS, D.J., DE. 1978. The taxonomy of Philippine Coniferae and Taxaceae. Kalikasan 7:117-152.

LAUBENFELS, D.J., DE. 1988. Coniferales. Fl. Malesiana 10(3):337-453.

Lawson, P. and Son [C. Lawson]. 1851. The fir tribe. No. X. Abietineae. List of seeds, plants, etc. Private press, Peter

Lawson, Edinburgh [Note: Stafleu & Cowan (1979) indicate that for a later major publication, The pinetum

britannicum, Ravenscroft compiled the text and Lindley and other botanists the descriptions. Peter Lawson

was mainly an editor and Charles Lawson, the son of Peter, was the botanist; copy of relevant pages reviewed

for this paper was obtained at BM].

LEIGHTON, W.A. 1841. A flora of Shropshire. John van Voorst, London & John Davies, Shrewsbury.

LEveiLLÉ, H. 1914. Le Monde des Plantes. Sér. 2, 16:88, 20.

Spjut, Taxonomy of Taxus 287

Li, H-L. 1963. Woody flora of Taiwan. Livingston Publ. Co., Narberth, PA.

Li, J, CC. Davis, P. Det TReDIci, and M.J. DoNoGHUE. 2001. Phylogeny and biogeography of Taxus (Taxaceae) inferred from

sequences of the internal transcribed spacer region of nuclear ribosomal DNA. Harv. Pap. Bot. 6: 267-274.

Li, N. and Lk. Fu. 1997. Notes on gymnosperms |. Taxonomic treatments of some Chinese conifers. Novon

7:261-264.

Ligerty Hype Bailey HoRTORIUM, Starr. 1976. Hortus third. Macmillan Publishing Co., New York.

Liu, T-s. 1960. Illustrations of native and introduced ligneous plants of Taiwan. Vol. 1. National Taiwan Univer-

sity.

Louron, J.C. 1844. Arboretum et fruticetum Britannicum. Longman, Brown, Green and Longmans, London.

Lowe, J. 1897. The yew trees of Great Britain and Ireland. MacMillan and Co., Ltd., London.

Makino, T. 1931. Illustrated flora of Nippon, with the cultivated and naturalized plants. Honkuryukwan Co, Ltd.,

Japan. (Note: date from print on external binding, cited by others as 1940.]

MARKHAM, K.R., R.F. Wessy, L.A. WHITEHOUSE, B.J. MoLLoy, C. VILAIN, and R. Muss. 1985. Support from flavonoid glycoside

distribution for the division of Podocarpus in New Zealand. New Zealand J. Bot. 24:1-13.

MarshaLt, H. 1785. Arbustum Americanum. Joseph Crukshank, Philadelphia, PA.

Maximowicz, C.J. 1859. Primitiae florae amurensis. Buchdruchkerei der kaiserlichen Akademie der Wissenschaften,

St. Petersburg.

MircheLt, J. 1974. The yew trees of linchionaig, Dunbartonshire. Trans. Bot. Soc. Edinburgh 42:163-166.

Naka, T. 1938. Indigenous species of conifers and taxads of Korea and Manchuria, and their distribution. Chósen

Sanrin Kaihó (J. Kor. For. Soc.) 158221.

Nicolosi, R.T. 1982. Morphological features of leaves and pollen as in aid in separating selected species and

cultivars within the genus Taxus. Thesis, Ph.D., Ohio State University.

Ohw, J. 1965. Flora of Japan. English translation, eds. F. G. Meyer & E. H. Walker, Smithsonian Institution, Wash-

ington, DC.

OLDHAM, K. 2005. David Douglas arrives at Fort Vancouver to begin two years of botanical exploration on Aprl 20,

1825. HistoryLink.org. The Online Encylopedia of Washington State History, Essay 7298 (Accessed Feb. 2007,

http//www.historylink.org/essays/output.cfm?file id—7298).

Orr, M.Y. 1937. On the value for diagnostic purposes of certain of the anatomical features of conifer leaves. Notes

Roy. Bot. Gard. Edinburgh 94:256-265 & Plates.

OUDEN, P. DEN and B.K. Boom. 1965. Manual of cultivated conifers. M. Nijhoff, Hague.

PARLATORE, P. 1868. Coniferae. In: A. de Candolle, Prodromus systematis naturalis regni vegetabilis 16 (2):361—

524.

PILGER, R. 1903. Taxaceae-Taxoideae—Taxeae. Taxus. In: Engler, Das Pflanzenreich IV:110-116.

PiLGER, R. 1916. Die Taxales. Mitt. Deutsch. Dendrol. Ges. 25:1-28.

Rao, A.R. and M. Malaviya. 1965. On the distribution, structure, and ontogeny of sclereids in Taxus baccata. Linn.

Proc. Natl. Inst. Sci. India 31:114—122.

Rau, M.A. 1974. Vegetation and phytogeography of the Himalaya. In: M.S. Mani, ed. Ecology and biogeography

of India, W. Junk, Hague. Pp. 247-280.

REHDER, A. 1902. Taxus. In: L.H. Bailey and W. Miller, Cyclopedia of American horticulture. Vol. 4 (R-Z). The MacMil-

lan Co., NY

REHDER, A. 1919. New species, varieties and combinations from the herbarium and the collections of the Arnold

Arboretum. J. Arnold Arb. 1:44-60.

REHDER, A. 1923. New species, varieties and combinations from the herbarium and the collections of the Arnold

Arboretum. Taxus media hybr. nov., Taxus media f. hatfieldii forma nov., Taxus media f. hicksii, comb. nov. J.

Arnold Arbor. 4:107-109.

REHDER, A. 1936. Notes on the ligneous plants described by H. Léveillé from eastern Asia. Taxaceae. J. Arnold

Arb. 17:54.

REHDER, A. 1940. Manual of cultivated trees and shrubs (2nd ed.). MacMillan Co., NY.

288 Journal of the Botanical R h Institute of Texas 1(1)

REHDER, A. 1949. Bibliography of cultivated trees and shrubs. Arnold Arboretum, Jamaica Plain, MA.

RopweLL, J.S. (ed.), C.D. Picott, D.A. Rarcurre, A.J.C. MALLocH, H.J.B. Birks, M.C.F. Proctor, EW. SHimweLL, J.P. HUNTLEY, E.

RADFORD, M.J. WicaiNTON, and P. WiLkiNs. 1991. British plant communities. Vol. 1. Woodland and scrub. Cambridge

University Press.

RozENDAAL, E.L.M., G.P. LeLvveLD, and T.A. van Beck. 2000. Screening of the needles of different yew species and

cultivars for paclitaxel and related taxoids. Phytochemistry 53:383-389.

Sakia, D, S. PS. KnaNUJA, A.K. SHasany, M.P. Darokar, A.K. KUKREJA, and S. Kumar. 2000. Assessment of diversity among

Taxus wallichiana accessions from northeast India using RAPD analysis. IPGRI Newsletter 121:27-31.

Sapru, B.L. 1975. Vegetational studies in Jhelum Valley. Botanique 6:151-164.

SARGENT, C.E. 1896. The silva of North America. X. Houghton, Mifflin & Co., Boston and New York.

Seigler, D.S. 1991. Cyanide and cyanogenic glycosides. In: G.A. Rosenthal and M.R. Berenbaum, eds. Herbivores:

their interactions with secondary plant metabolites. Academic Press, San Diego, CA. Pp. 35-78.

SIEBOLD, P.F., von and J.G. ZUCCARINI. 1843. Plantarum, quas in Japonia collegit Dr, Ph. Fr. de Siebold. Genera Nova. Notis

characteristicis delineatiionibusque illustrata proponunt, |. Abh. Math.-Phys. CI. Kónigl. Bayer. Akad. Wiss. 3.

SIEBOLD, P.F., vou and J.G. Zuccanivi. 1846. Florae japonicae familiae naturales. Abh. Math.-Phys. Cl. Kónigl. Bayer.

Akad. Wiss. 4:123-240.

SIEBOLD, P.F., von and J.G. Zuccarini [1870]. Flora japonica, pts 6-10 [Descriptions by FA.W. Miquel and published

by Miquel]. Lugduni batavorum.

SiLBA, J. 1984. An international census of the coniferae, |. Phytologia Mem. 7:1-79.

Silea, J. 1986. Encylopaedia coniferae. Phytologia Mem. 8:1-217.

Sivger, M. and G.P DEWorr, Jr. 1970. Yews in fiction and fact. Arnoldia 30:139-147.

Spyut, RW. 1977 (July 14). Taxus brevifolia Nutt. (Taxaceae), nomenclature, description, distribution, ecology, com-

ments, uses, other, references, and attachments—table 8 from Franklin and Dyrness 1969, distribution maps.

USDA ARS Medicinal Plant Resources Laboratory, Beltsville, MD. 12 pp (circulated to USDA lab scientists, NCI;

content also incorporated into NCI solicitations for large collections of stem-bark sent to prospective bidders

in the Pacific Northwest, 1978-1990; copy available at http://www.worldbotanical.com/TAXNA.HTM).

Spyut, RW, 1992. A taxonomic key to the species of Taxus. NCI Workshop on Taxus, Taxol, and Taxotere, Rockville,

MD (Abstract only).

Spjur, RW. 1993. Reliable morphological characters for distinguishing species of Taxus (Abstract). In: International

yew resource conference. Yew (Taxus) conservation biology and interactions. Berkeley, CA. Pp. 39-40.

Spjur, RW. 1995 (May 25). Research proposal on taxonomic revision of Taxus. USDA ARS Memordandum to Thomas

S. Elias, Director, National Arboretum, with references, key to species of Taxus dated 24 Apr 1995, character

list in DELTA format. Photocopy at http://www.worldbotanical.com/TAXNA.HTM.

Spjur, RW, 1998. (a) Morphological evolution in the Taxus leaf and its significance to recognizing ecological spe-

cies within the genus. (b) Species of Taxus. Presented at the AIBS Annual Meeting, American Systematic Plant

Taxonomists, Baltimore Convention Center, MD, 5 Aug. 1998. Abstracts published on the Internet, Botanical

Society of America (URL: http://www.botany.org/).

Spjur, R.W. 2000. Three papers presented at the joint meetings of the Botanical Society of America and American

Systematic Plant Taxonomists, Portland, OR, August, abstracts published online and Amer. J. Bot.

(a) A phytogeographical analysis and classification of leaf morphological features in Taxus (Taxaceae). http://

www.ou.edu/cas/botany-micro/botany2000/section13/abstracts/27.shtml.

(b) The morphological relationships of Taxus canadensis (Taxaceae) in North America and Eurasia. http://www.

ou.edu/cas/botany-micro/botany2000/section13/abstracts/28.shtml

botany2000/section13/abstracts/166.shtml. (Note key also presented on the WBA website in 2003, extracted

from an poster document of 24 pages that included 270 figures).

Spyut, RW, 2003. Unpublished. Nomenclatural and taxonomic review of three species and two varieties of Taxus

(Taxaceae) in Asia. Prepared 1998, submitted to Sida in 2000, placed on the web, www.worldbotanical.com,

Spjut, Taxonomy of Taxus 289

2003. Additional reviews: Taxus (Overview of Taxus, 1999). Keys and descriptions for species of Taxus, Intro-

duction to taxonomy of Taxus, (2003). North America (1999) and Species groups and subgroups (2003). All

revised in 2006, 2007. www.worldbotanical.com.

Spjur, RW. 2006. Biogeographical data on putative hybrids for species of Taxus (Taxaceae) in the Himalayas and

North America. Abstract, Botanical Society of America, Chico, CA, July 29-Aug 2. http://www.botanyconfer-

ence.org/er git 1e/search/index.ph p?fu nc=detail&aid=37

Spjur, RW, 2007. A phytogeographical analysis of Taxus (Taxaceae) based on leaf anatomical characters. J. Bot.

Res. Inst. Texas: 1(1):291-332.

Spyut, R.W., J-Y. ZHou, and C-J CHANG. 1993. Comparison of taxane content between two different native yews from

the Pacific northwest. Poster/Abstract, 34th Annual Meeting, Amer. Soc. Pharmacog., San Diego, CA.

STAFLEU, F.A., and R. S. Cowan. 1976-1988. TL-2. Taxonomic literature. 7 Vols. W. Junk, Hague.

STAINTON, J.D.A. 1972. Forests of Nepal. Camelot Press, London.

SurrNEss, M (ed.). 1995. TAXOL. Science and applications. CRC Press, Inc.

TAYLOR, R.L. 1932. The successional trend and its relation to second growth forests in southeastern Alaska. Ecol-

ogy 13:381-390.

TAYLOR, R.L. and S. Taylor. 1980. Taxus brevifolia in British Columbia. Davidsonia 12:89-94.

Thomas, P.A. and A. Potwart. 2003. Biological flora of the British Isles. Taxus baccata L. J. Ecol. 91:489-524.

THUNBERG, C.P. 1784. Flora Japonica. Mülleriano, Lipsiae.

Trrrensor, R.M. 1980. Ecological history of yew Taxus baccata in southern England. Biological Conservation

17:243-265.

Vance, N.C. and A.B. KnurkiN. 1993. Using restriction fragment length polymorphisms to assess genetic variability

among yew taxa. Yew (Taxus) conservation biology and interactions: 34, Berkeley, CA (abstract).

VEITCH, J. and Sons. 1881. A manual of the Coniferae. Published by the authors, Kings Road, Chelsea.

Vouoris, D. 1986. Historic and environmental significance of the yew (Taxus baccata L.). Israel J. Bot. 35:47—52.

Watt, M.E., M.C. Wan, and H. Taylor. 1972. Isolation and chemical characterization of antitumor agents from plants.

Cancer Treat. Rep. 60:1011-1030.

WaLticH, N. 1826. Tentamen florae nepalensis illustratae. Fasc. 2. Calcutta and Serampore.

Wana, C-w. 1961. The forests of China. Maria Moors Cabot Foundation No. 5, Harvard University, Cambridge,

i-ix &1—313.

Wana, T, Y-J. Su, J-M. ZHu, G-k. FAN, and C. HUANG. 2000. RAPD analyses of Taxaceae and its related taxa. Acta Sci.

Nat. Univ. Sunyatseni 39:120-130.

Wan, M.C., H.L. Taylor, M.E. Watt, P. Coacon, and A.T. McPHAIL. 1971. Plant antitumor agents. VI. The isolation

and structure of taxol, a novel antileukemic and antitumor agent from Taxus brevifolia. J. Amer. Chem. Soc.

93:2325-2327.

Watt, A.S. 1926. Yew communities of the South Downs. J. Ecol. 14:282-310.

WiLson, E.H. 1916. The conifers and taxads of Japan. Publ. Arnold Arbor. 8:1-91, Cambridge Univ. Press, Cam-

bridge, MA.

WiLSON, E.H. 1926. Taxaceae. In: The taxads and conifers of Yunnan. J. Arnold Arbor. 7:39-42.

290 Journal of the Botanical R h Institute of Texas 1(1)

BOOK REVIEWS

Rosert H. MOHLENBROCK. 2006. The Illustrated Flora of Illinois Flowering Plants: Flowering Rush to

Rushes. Second Ed. (ISBN 0-8093-2687-6, hbk.). Southern Illinois University Press, PO. Box 3697,

1915 University Press Drive, Carbondale, IL 62902, U.S.A. (Orders: http://www.siu.edu). $65.00, 328

pp., b/w line drawings, glossary, index, 54%" x 81⁄2".

Flowering Plants: Flowering Rush to Rushes is an updated flora by Mohlenbrock that includes many of the monocot families from Illinois.

(pp 1 1

The flora includes piu on 16 families including: Butomaceae, Alismaceae Juncaginaceae, Potamogetonaceae,

Ruppiaceae, Z , Najadaceae, Araceae, Lemnaceae Sparganiaceae Typhaceae Xyridaceae, Commelinaceae, Pontederiaceae,

and Juncaceae. Flowering Plants: Flowering Rush to Rushes is a second edition, and is part of a six flora series entitled The Illustrated Flora

of Illinois. A second part of the series will be a flora containing the other half of the monocot families

The author starts out by showing the reader how to use the keys; this allows the reader to determine the family of the plant they

are interested in. The keys used for this flora are very straightforward. This flora also includes a glossary, so this book will be usable for

novices as well as those with more plant identification experience. The author has provided general descriptions of the order as well as

descriptions for all families included under that order (for Illinois). These descriptions include general morphological traits and other

notable traits for order, family and genera within the family. Dichotomous keys follow each family description and allow readers to

determine Eun followed a e within in so

intl ] i that species’ logical traits, common name, habitat, range, some

known AN data and other helpful information. There are 125 black and white lie illustrations of various species (108 from the

first edition and 17 new illustrations for this edition). These images provide readers with details of the plant habit, leaves, inflorescence

and/or fruit. Each species description also presents a distribution map by county of that species within Illinois.

This flora concludes with an explanation of why certain species were excluded from this second addition, and also contains an

appendix that discusses revisions from the previous edition (1970). Mohlenbrock has provided readers with a succinct summary of the

taxa included in this volume, a glossary, references and an index by plant names, both common and scientific

Flowering Plants: OTE Rush to Rushes by nen is a usable flora that would make a nice addition to the library of d

person interested in the id i ion of Illinoi well as anyone working with aquatic plants). This flora is a se

and has been updated over the earlier versions by the addition of new species as well as corrected plant name information. The black and

white drawings provided in the text are helpful in showing the major identification traits of the species. The keys are straightforward

1 y!

and the flora does contain a glossary for any unfamiliar terms. The description for each species is and identification hints help

the reader conclude that he/she has determined the correct species. Flowering Plants: Flowering Rush to Rushes is another well-written

and user-friendly flora from Robert Mohlenbrock.—Lee Luckeydoo, Herbarium, Botanical Research Institute of Texas, 509 Pecan Street, Fort

Worth, TX 76102-4060, U.S.A.

VB. Price and Bares H. Morrow. 2006. Canyon Gardens: The Ancient Pueblo Landscapes of the American

Southwest. (ISBN 13: 978-0-8263-3859-4, pbk.). University of New Mexico Press, MSC04 2829, 1

University of New Mexico, Albuquerque, NM 87131-0001, U.S.A. (Orders: www.unmpress.com, 800-

249-7737) $39.95, 217 pp., 36 halftones, 40 line illustrations, 2 maps, 6 */s" x 9 ?/s".

With today's concerns about global warming and climate change, this two-part anthology is very timely. Part One examines how the

Ancestral Puebloans used their vast storehouse of Ps actical a to live in, and with, the indi and to guae to changing natural

h 1 1

and historic circumstances. Part Two proposes ways in which modern learn f pragmatists. Although

aimed at contemporary landscape architects, urban "C architects and builders, this book will be thought-provoking for anyone

interested in the natural world and especially for who make laws and regulations on the local, state and national levels. As V.B. Price

states in the prologue:

We believe that in times of global climate change, water scarcities, and upheavals in energy sources and technologies, an observant

0-2

sensitivity as to how the natural environment works and a mastery of its demands could prove to be immensely useful in designing and

retrofitting urban environments in the future.

“Design with nature,” lan McHarg famously argued in the 1960's. Canyon Gardens offers some insights into how the Puebloans did

just that and some examples of how we today can also.

Contributors to this a eee include an architect/teacher/writer, two archaeologists, the curator of e at

the U. Colorado Museum l a National Park Service historian, a lyrical writer/poet, a lan

E E

architect, and a writer/architectural historian ol is a member of the Santa Clara Pueblo. The two editors—a writer/poet/columnist e

the founding director of the Master's Program in Landscape Architecture at UNM—are contributors as well. Each has his/her individual

voice and point of view. All invite the reader to consider the lessons of the past in shaping the world of tomorrow.—Penny McCook,

Volunteer, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A.

J. Bot. Res. Inst. Texas 1(1): 290. 2007

APHY TOGEOGRAPHICALANALYSIS OF TAXUS CUAXACEAE)

BASEDION. LEAP ANATOMICAL GHARAGTERS

Richard W. Spjut

World Botanical Associates

PO Box 81145

oad Quem 93380-1145, U.S.A.

j hspjut@bak ECO]

ABSTRACT

A phytogeographic analysis of 845 Taxus specimens is presented based on leaf anatomical characters for the number of stomata rows

in a stomata band and the number of epidermal cells that lack papillae between the leaf margin and stomata band. The specimens are

arranged by continent, country, state or province, and species, and represented geographically on three maps: (1) North America, (2)

the Euro-Mediterranean, and (3) Asia. Taxus is least diverse and most distinct taxonomically in North America, and most diverse and

least distinct taxonomically in southwest China. Stomata data show several clines in North America, an obvious south to north decrease

for the Mesoamerican yew (T. globosa) and Pacific NW yew (T. brevifolia) populations combined, and a less obvious reverse cline for the

Canada yew (T. canadensis). The results are discussed in review of other paleobotanical data. It is suggested that Taxus immigrated to

North America from Asia across a Pacific land connection during the Cretaceous, and from Europe to North America across North Atlantic

land bridges during the Tertiary. The low diversity of Taxus in North America is suggested to be the result of the K/T extinction event.

In the Euro-Mediterranean, evolution of Taxus is suggested to have been impacted more by extinction as a result of climatic changes

during the Tertiary, and by hybridization during the Pleistocene. The greater diversity in SW China is indicated to be the result of less

extinction there and more frequent hybridization during the Pleistocene, not only among authochtonous species, but also allochtonous

species as a result of the Himalayan uplift.

CHINESE ABSTRACT

AT ARR— AK ls A d d BA RAT FL eZ. ILR R ZA EIC H AP RP IE, ANC

Xj 845 (TEE) Taxus bw ETT f PHBL TPA. AR. MERLO, Hh

Sy ARRANGE 3 WARE: CG) dese, (E) WRN, (D EM. ALIENS ER

(RARER ARH, MAR ESL VERS BET] RE ERE APSR EDO] eH] s HAL SAN ZL UE

FBeALL, PELE T. globosa FAH AA T. brevifolia HERA mdk

AY de>, MENA T. canadensis HAG RAVER AH. RAS I WARE, EE

RSC FER. ZL AZ TE SEZ SS = 20 2) al) MA AR BR A PE Bet HAE Fe FT PU TE

MEERI. HERP — £0 ES E MA EES FEE. SSR A

ERREKERI ER AE A RST UC] HE IZ) IE ME. EH rH RR AAA

Ft, ALBARN AS BEAD AE Be TEA LE, Tm AEP A RRA Kb. MAZEE

rp Bs yu Bs S HI d FE TES I o

INTRODUCTION

The genus Taxus has included eight geographically defined species: (1) T. baccata L.—Europe, N Africa and

SW Asia (Franco 1964), (2) T. cuspidata Siebold & Zucc.—temperate E Asia (Krüssmann 1985; Ohwi 1965),

(3) T. wallichiana Zucc.—Himalayas (Krüssmann 1985), (4) T. sumatrana (Miq.) de Laub.—S China, Philip-

pines, Taiwan, Sulawesi, and Sumatera (de Laubenfels 1988), (5) T. globosa Schltdl.—N Central America

to Mexico (Ferguson 1978), (6) T. brevifolia Nutt.—NW North America (Ferguson 1978; Hils 1993), (7) T.

floridana Nutt. ex Chapm.—W Florida (Ferguson 1978; Hils 1993; Price 1990), and (8) T. canadensis Mar-

shall—NE North America (Ferguson 1978; Hils 1993; Price 1990). Except for T. sumatrana, these were also

recognized by Pilger (1903, 1916, 1926) as subspecies of T. baccata.

These eight geographically defined taxa—generally accepted as species (Bailey 1933; Farjon 1998;

Hils 1993; Krüssmann 1985; Rehder 1940; Silba 1984)—have not been clearly distinguished (Ferguson

1978; Price 1990), including several other species and varieties that have been recently recognized (Farjon

1998, 2001; Fu et al. 1999); however, I will show that this traditional geographical classification is distorted

to recognizing more species where diversity in the genus is least (North America) and less species where

J. Bot. Res. Inst. Texas 1(1): 291 — 332. 2007

292 Journal of the Botanical R h Institute of Texas 1(1)

diversity is greatest (SW China). This will be based on quantitative leaf character attributes for the number

of stomata rows (SR) in a stomata band, and the number of marginal cells (MC) across an abaxial marginal

zone without papillae. These data will be summarized on geographical maps of North America, Euro-

Mediterranean, and Asia, and analyzed in the results section of this study. This is followed by a discussion

of phytogeographical relationships. Data for all herbarium specimens studied are provided in an appendix

according to continental and political regions and taxonomy.

MATERIALS, STANDARDS AND METHODS

MATERIALS.—The materials of Taxus include 845 specimens of fresh and dried branchlets with leaves

from throughout the natural range of the genus (Appendix), and an undetermined number of specimens

from cultivated plants in Australia (1), England (~50), France (~20), and the United States (-300), and from

miscellaneous other sources, the main one was Phyton, Inc (465), now Phyton Biotech, a commercial com-

pany specializing in producing taxol from tissue culture of Taxus.

STANDARDS.—The genus Taxus Linnaeus (Taxaceae Gray) is defined by cone and leaf morphology (Florin

1931, 1948c, 1951) in relationship to other “taxad” genera, characterized by producing an arillocarpium

(Spjut 1994)—a type of cone in which the seed is subtended by a fleshy arillate bract (Airy Shaw 1973;

Cheng € Fu 1978; Florin 19482; de Laubenfels 1988).

Taxads include both extant and extinct taxa; the extant genera, in addition to Taxus, are Amentotaxus

(5-6 spp., China, Vietnam), Torreya (6 spp., E Asia, N America), Austrotaxus (1 sp., New Caledonia), Cepha-

lotaxus (8-11 spp., E Asia), and Pseudotaxus (1-2 sp., China) (Fu et al. 1999). Molecular studies employing

ribosomal RNA (Chaw et al. 1993, 1995), chloroplast DNA (Tsumura et al. 1995), or RAPD (T. Wang et al.

2000), suggested Amentotaxus and Torreya to be more closely related to each other than to Cephalotaxus or

Taxus, and that Cephalotaxus is basal to two clades, (1) Torreya/Amentotaxus and (2) Taxus/Pseudotaxus/Aus-

trotaxus (Cheng et al. 2000); however, whether these clades should be treated in separate orders, families,

subfamilies, or tribes, is controversial (Hill 1998).

Cones.—Pseudotaxus and Taxus produce a terminal seed on a lateral (secondary) short shoot (André

1956; Dupler 1920; Miller 1988) that is only partly surrounded by a loose cupular bract, whereas in other

genera the seed is more fully and tightly covered by the aril (Florin 1948b; Sahni 1920). Cephalotaxus is

distinct for its biovulate cone scales from which usually only one ovule matures (Singh 1961). Amentotaxus

differs for its terminal, “racemose” male shoots (Cope 1998; Fu et al. 1999). The Austrotaxus cone was re-

garded as isolated from other taxads based on anatomy of the seed coat (Bobrov et al. 2004).

The closely related Pseudotaxus (1—2 spp., China, Fu et al. 1999) differs from Taxus by a white aril-

locarpium (Cheng 1934), and additional sterile scales in male cones (Florin 1948c).

Leaves.—Taxus leaves are differentiated from those in other taxad genera by papillose cells that define

the “stomatic apparatus" (Dilcher 1969; Florin 1931, 1948c, 1951, 1958). This apparatus includes 4—8 small

subsidiary papillose cells that encircle each stoma (Florin ring) and adjacent (accessory) papillose cells (Figs.

1A, 1B, 1C). Stomata develop in longitudinal rows (periclinal) in a stomatal region divided into two bands

by a midrib (e.g., Fig. 1A). The midrib and marginal cells vary in size, shape, and development of papillae.

Further details—with photomicrographs—can be found in Ferguson (1978), Florin (1931, 1951), Jinxing

and Yuxi (2000), Kvatek (1984), and Kwei and Hu (1974).

Mammillae, not to be confused with papillae (Bertrand 1874), develop singly over most of the cell’s

surface as large lens like bumps. Under a dissecting scope (30x) they appear most conspicuous along leaf

margins, less so on the epidermal surfaces (adaxial or abaxial). Papillae, by contrast, are smaller and numer-

ous on a cell—like pimples. They develop in 1-3 distinct or concrescent rows, generally discernible only

under a microscope—at least 100x, and only on the abaxial surface. Papillae are always present in stomata

bands, gradually diminishing in prominence outside the bands towards the leaf margins, and may develop

entirely or partially or not at all on midrib cells.

The development of stomata in rows and their differentiation by papillose accessory cells, which together

make up the stomata band, are the most distinguishing features of Taxus relevant to data in this study, compared

Spjut, Phytogeographical analysis of Taxus 293

to other extant genera of taxads (Florin 1931, 1951). For example, Pseudotaxus has glaucous stomata bands

(Cheng 1934; Florin 1931, 1948b, 1948c, 1948d) devoid of papillae, except on subsidiary cells, and more

stomata rows per band—23-28 rows (Florin 19480), in contrast to 4-21 rows in Taxus (Appendix). Amen-

totaxus and Torreya (Amentotaxaceae) have papillose glaucous bands largely of subsidiary cells (periclinally

arranged), rather than accessory cell types (Florin 1951, 1958). Austrotaxus (Austrotaxaceae; Nakai 1938; Florin

1958), which also differs conspicuously by its long-linear leaves— comparable to some Podocarpus spp.—has

stomata evenly scattered across the entire abaxial surface without clear differentiation of rows and bands in

which the epidermal cells are similar to those on the adaxial surface—irregularly quadrate (or pentagonal)

as in Taxus. These differences, and the presence of other features such as sclereids and resin canals in leaves

of Torreya (Bertrand 1874) and Cephalotaxus, would seem to support classification of the taxads in different

families (Amentotaxaceae, Austrotaxaceae, Cephalotaxaceae, Taxaceae).

The features of the stomata band that distinguish Taxus from other extant taxads do not apply to extinct

taxads, however (Florin 1951, 1958; Harris 1976a, 1976b; Kvacek 1984; Miller 1977). Photomicrographs of

many taxad fossils from Jurassic deposits presented by Florin (1958) show remarkable detail that are strikingly

similar to extant Taxus in leaf epidermis (Kvacek 1984; Meyen 1984), except for narrower stomata bands with

fewer stomata rows (Kvacek 1984). Indeed, some leaves, which included twigs and arillocarpia, were assigned

to Taxus; these are T. bornholmiensis Florin with 4—5 stomata rows, T. harrisii Florin with 5 stomata rows,

and T. jurassica Florin with 3—5 stomata rows; however, none of these appear to belong to the genus Taxus.

Harris (19762, 1976b), for example, transferred T. jurassica to Marskea, an extinct genus characterized by op-

posite-decussate leaves (Florin 1958; Harris 1976b), which are clearly evident in Florin's (1958) photograph

of Marshea jurassica. Jurassic taxads include many other extinct genera (Florin 1958) that may have existed

since the Triassic (Florin 1951; Meyen 1984).

Epidermal cells adjacent to leaf stomata bands, the midrib and marginal areas, are usually papillose in

part. Epidermal cells on the abaxial surface nearest the leaf margin appear to have evolved in some species by

extension (folding) of the upper (adaxial) surface to the lower (abaxial) surface; leaves of many specimens are

revolute along their margins in which the abaxial epidermal cells are often more similar to those above than

to the adjacent cells below (Nicolosi 1982).

The comparative morphological relationships of Taxus to other extant taxads (Florin 1931, 1948c; Ap-

pendix) indicate that the ancestral Taxus leaf had a partially differentiated abaxial epidermis in which stomata

developed in definite rows but not in distinct bands. Evidence for this can be seen in T. wallichiana and allied

species in the E Himalayas and SW China. Their leaf stomata occur not only in stomata bands, but also on the

abaxial midrib; essentially, stomata develop across the entire abaxial leaf surface to within several cells of the

margin—in up to 21 rows. The abaxial marginal and accessory epidermal cells are all nearly rectangular and

papillose—in sharp contrast to the epidermal cells on the adaxial surface that are much shorter, +trapezoidal-

pentagonal, and without papillae.

METHODS

More than 1,000 herbarium specimens (A, BH, BM, BOLO, E, GH, K, M, NA, NY, P, PE, PH, S, U, US; Hol-

mgren et al. 1990) were studied of Taxus throughout the natural range of the genus to assess morphological

variation in characters that involve branches, bud-scales, leaves, and cones; 845 are cited in the appendix, and

additional specimens are mentioned in this paper. Each specimen was photographed with a Nikon camera

using 35 mm color film with 35 mm and 60 mm lenses. Stafleu and Cowan (1976-1988) were consulted for

location of types, other specimens of historical relevance to this study, and references.

From each herbarium specimen of Taxus, one mature leaf was selected for microscopic study of anatomi-

cal features. The Taxus leaf was soaked in water for 8-16 hrs. The leaf was then transversely sectioned in the

mid region as bryologists routinely section leaves of mosses for taxonomic identifications. With a single-edge

razor blade and dissecting needle as a guide, 5—10 transverse sections were generally made. Then an abaxial

epidermal layer was removed from both remaining leaf portions, generally 0.5-2.0 mm in length, by scraping

mesophyll parenchyma from the epidermal layer with a razor blade. Occasionally, the entire abaxial leaf

294 Journal of the Botanical Research Institute of Texas 1(1)

im -

Ad L Ini S >

P UAITI U TUR

OS PRUUN

TA Y RA

Fic. 1A. Mid leaf sections of Taxus caespitosa var. latifolia, fi Itivated plant in Maryland, U.S.A, Spjut 10485 (wba), representative of the Baccata

Group, Cuspidata Alliance. Top: transverse section (T-sect.), ~100x, sh g elliptical shaped epid | cells, phyll lay f anticlinal palisad

layer of parenchyma and spherical parenchyma cells, drawn by Karen Parker. Bottom: abaxial epidermal layer from margin (left) to midrib (right),

x, showi inal border of 8 th( illose) cells wide foll 1 by a stomata band with 13 rows of stomata, and a midrib of mostly

smooth cells, drawn by R. Spjut.

uat umm

228 cn

a. >

as ives

Fic. 1B. Mid leaf secti fT brevifoli pt f Siskiyou C , California, U.S.A., rey tative ofthe Wallichiana Group, R. Spjut & T. Spjut

11835 (wha, type). Top: T-sect. ~ 100x shows tall angular epidermal cell ( f |

to midrib (right), ~250x, shows marginal region of 10 Il

rows, and a papillose midrib, drawn by R. Spjut.

Darl Datt L al O] II! ofl nAtt\

£. A Deer O RP fall Ih,,0 £

gin Wert)

lls, 5 stomata

Spjut, Phytogeographical analysis of Taxus 295

em at ae 1 SON a E E

m TAM TD o TTA

Pr £A j zs -u = es Ti V E i E -- y me mA y; ?

GUMMI De du.

à MAA Ti : dn

Aa tang E > at ar NS AT Gi 55) | de —

POE S E x afr CS EE ee CBee.

23 eos aes x rr ra E7 ee A qm ue n Vor Pr 7 -

mmc a ERD AL pe ae E de "HITS $

Cx ^

ae PITH LULA AE SATA RANA RA TT AS

AR i 3 vV OO en ee tae, y La oe aa het oe, E,

cxx JU phe Ph Pe LA P - rr ; E a g Li CF 4 [i ge P S.

A «p.49. ME VE UA

A ar E A aL, ari X hoz

A yc TA

[o -

CIT] ody Doe ites OF) a

HN []/4778 ^v ad 1 Abe v Y yy. i A

wy, PP f. AU joy) P o0 D od

en, HOUR E^ EA PU T e

IN C6 ous e hoe e

E ge NS KO m

a Y WA DES ihe”

(TIAM 9 Uns Ys

GE Haa EN

53 4) ]

EY t

MP tamm

ELA A

B xs

Nan

cs 4

eC f e QUU YI

D OS EAS] a (TO

LINO Gas, 724 SO

54) a hes

£2 (jr 3) à

M m n ji Ais] : GN E dA ERE PEU.

EL Tt NS RDA WEE UC O dor TU Y,

Fic. 1C. Mid leaf secti tati fthe f $ t G drawn bv R. Spjut. F top to bott Ton: T. maireivar. irei fi G |

r En L4 re T E J »

(China), ee a F A INTA! L Es J J J A EE | | L 2 | £. el I J E y I Wl » A + l A * |

T r

epidermal cells on adaxial surface, the lower T-section, T. mairei var. mairei from Yunnan, Maire s.n., isotype, st g

J r * * "i "i

ahavial id tc f RETE +h sb

in width, a stomata band with 16

cells and 12-14 stomata rows.

r F J J a T 7

; upper most from Guizhou, isotype (A) of T. speciosa, showing marginal region of 23 smooth cells

| th midrib; the | tion from holotype of 7. sumatrana, showing long rectangular

surface and a medial portion of the adaxial epidermis were removed. All sections were examined under

magnifications of 100x, 250x, and 400x (Nikon binocular microscope) for cell shape, number of stomata

rows, number of cells marginal to stomata bands, and for papillae position and distribution across the abaxial

leaf surface. The results were sketched and described on small packets 3 x 5 inches. A temporary slide of

the sections and photographs of the herbarium specimen were retained for each packet. Leaves from fresh

specimens were also similarly studied throughout the range.

Figs. 1A-1C show diagrammatic leaf sections of the mid region that is representative of three species

296 Journal of the Botanical R h Institute of Texas 1(1)

Labrador

Sea

arit hen

Fic aA Al L rp" P" L IICD\ inl £ T. La f I In Is PO L L H H £, M eh A

see Appendix for specimen data. Specimens from northern Mexico indicated in yellow numbers are not easily distinguished from those in Florida;

therefore, these are considered 7. globosa var. floridana.

Nore garan

axus canadensis

Taxus boa

T. . de

ni PAA oy 5 7

Fic. 2B. Number

higher counts, 11—15 st

canadensis. Most T.

groups of Taxus. These include (1) a complete transverse section and (2) an epidermal portion of the abaxial

surface from one margin to across the midrib. As previously indicated, similar sketches were made on 3.5

x 5 inch (8 x 12.5 cm) packets for most herbarium specimens studied except only portions of the stomata

and marginal areas were drawn, while number of stomata in a band and number of marginal cells adjacent

Spjut, Phytogeographical analysis of Taxus 297

Al L £4 L I /ChV? I LT. £. 4’, » . PA. Lal D EY E r A ES

Fic. 2C

for specimen a Sumat Group, which i in SE China, is not shown to contrast the diffi bet the Wallichi Mn

Groups. The M Subgroup includes 7. suffnessii in white, T. florinii in blue, and T. wallichiana var. y is in black. The Chinensis Subgrouy

sel th TTEN i Ep P af e 1 J T ££ O L s I * a I Ari: J Bag J - s 4L KL E L ak DL I H

r r L) J

Sumatera, Sulawesi, and Taiwan? lalsot ies i inland China, in yellow. The Cuspidata Alli includes f T. cus spidata, T biternata,

and T. SUED in maroon, and E umbraculiera, in pete ote higher oo counts for T contorta d 1) and owe cani for T. wallichiana ME -14) NIE

VU OY VIGA WITS f.

florini is recognized to have 7-12 rows per band, and T suffnessii from 13-20 rows per band.

to stomata bands were recorded. The leaf margin can be difficult to pinpoint when leaves are rounded along

margins. This is determined by the smallest cell that is usually mammillose. Both types of leaf sections were

examined to determine where papillae develop on cells between the margin and stomata band.

Variation due to mechanical preparation, error in counting, and environmental factors (Deryugina &

Nesterovich 1981), were only generally assessed—for practical reasons—from duplicates that were uninten-

tionally included in this study, occasional field collections that were collected at various heights from one

or several trees of a population (top, middle, and lower branches of T. brevifolia from trees in California and

Oregon), and from test cases of selected leaves at various developmental stages from shrubs in cultivation.

Practical reasons include damage to herbarium specimens caused by removing a leaf, and the time required

to prepare leaf sections and record data, approximately one hour for each specimen.

Herbarium specimens studied are listed in an appendix with data on numbers of stomata rows per band

(SR) and numbers of marginal cells (MC) without papillae. Specimen data are arranged by continent, then by

country within continents, and finally by taxa, generally from south to north in North America, and from west

to east in Eurasia, and then east and south from the Himalayas to Indonesia. Leaf anatomical data are further

arranged by decreasing order in number of stomata rows (SR), and by increasing order in number of marginal

cells (MC) except for T. canadensis, the Sumatrana Group, and for duplicate specimens belonging to the same

species, or duplicate specimens from the same locality or collection number. Only minimal collection data

are cited, although for many specimens data were minimal. If the stomata count varied on each side of the

midrib of a single leaf, this is indicated by a slash; for example, T. canadensis frequently had 5 stomata rows

in one band and 6 in the other (5/6). A dash between numbers indicates a variable range, especially when

more than one leaf from the same specimen was studied, or a dash alone indicates absence of data. It should

f4L,D o ID L

298 Journal of t titute of Texas 1(1)

be remembered that these data are a byproduct of an overall taxonomic study of the genus Taxus (see Spjut

2007); i.e., they were not compiled with this paper in mind. Additional character features that appeared to

correlate with the findings are also noted (e.g., length of epidermal cells/width of epidermal cells or 1/w).

RESULTS

Numbers of Stomata Rows (SR).—Figures 2A, 2B and 2C show numbers of stomata rows per band on

maps of North America, Europe, and Asia, respectively, for representative specimens listed in the Appendix.

Each number represents a count from a single leaf of a herbarium or fresh specimen; the datum is plotted

at the general location where it was reported to have been collected. Where the count varied on each side

of the leaf midrib, or among duplicate specimens, the highest number was scored.

Stomata data for duplicate specimens, or among specimens from different plants at the same locality, are

summarized under five cases as follows:

(1) For 62 duplicates of herbarium specimens included in this study, the same number of stomata rows per band was found in 19 dupli-

cate specimens. Among the remaining 43 duplicates, 17 differed only by 1 row, another 17 differed by 2 rows, 6 varied by 3 rows,

and 3 varied by as much as 4 rows. Duplicates that varied by 4 stomata rows appear to have been collected from different habitats

within the same general locality. Examples are Farges 128 from Sichuan, Tsiang Ying 1425 from Guangdong, and several cases where

the same collection number was reported from multiple locations, Wilson 1265, from two sites in Sichuan and one in Hubei, and

Wilson 4053 from two locations in Sichuan (Rehder & Wilson in Sargent 1914).

(2) From a single shrub of T. caespitosa Nakai var. latifolia (Pilg.) Spjut in Maryland, ~35 yrs of age and 3 m in height, 16 leaves were

selected from branchlets with various exposures to light and age (U-2"4 yr). With one exception, 15 were found with 13 stomata

rows per band; one leaf—plucked from a well-shaded branchlet near the main trunk—had 16 rows.

(3) From trees of T. brevifolia, leaves from top, middle, and lower branches had the same count at two sites—one in California, and

another in Oregon—but varied by 1-2 rows in leaves from different trees at the same sites.

(4) In Taiwan, individuals of a population collected on five occasions (Appendix) had the same number of stomata rows at one site

(Tongshi 7), but varied from 11-14 rows per band at two other sites (Tongshi 5, Tongshi 6).

(5) The age of the leaf was apparently not a factor in 20 leaves studied of one cultivated individual of T. mairei (Lemée «Y Lév.) S.Y. Hu ex

T.S. Liu var. speciosa (Florin) Spjut; 16-18 stomata rows per band were found in all leaves from buds to 3™ yr branchlets.

Generally, the number of stomata rows per band varied by a count of 3 for about half of the individuals or

species. In NW North America (T. brevifolia), this occurred frequently among different individuals within

a population, whereas in Europe (Baccata Alliance) leaves of Taxus exhibited the same range in variation

whether obtained from the same plant or from different plants at the same site. A wider range, 13-18 stomata

rows, in the E Himalayas (T. wallichiana), and a narrower range, 5-6 or 6-7 stomata rows, in NE North

America (T. canadensis), were also evident.

Despite this range in variation, the numbers of stomata rows per band in North American Taxus (Fig.

2A) show a distinct cline from south to north for the Wallichiana Group (Central America to Florida, Cali-

fornia). Specimens from Honduras to southern Mexico had 7-11 rows per band (Bertrand 1874); those from

N Mexico and Florida had (5) 7 (-8) rows per band in further contrast to 4-7 (-9) stomata rows per band

in the Pacific NW. This northward decline in stomata rows per band is also apparent within the Pacific

NW by the highest count of 9 rows found in specimens from California, compared to the lowest count of

4 rows from specimens at more northern locations—in the Rocky Mountains. This cline was also noted to

be associated with an increase in length of epidermal cells relative to width (l/w), ranging from an average

of ca. 3x l/w to 8x l/w.

A reverse cline, however, is evident in the NE North America by 9 rows per band occasionally found in

specimens from Newfoundland, compared to the more common 6 rows per band as reported by Bertrand

(1874), which includes specimens from Quebec to Ontario, south to Kentucky (Spjut 1998a, 2000a, 2000b).

In the Euro-Mediterranean, Taxus leaves most often had 8—10 stomata rows per band (Fig. 2B); however,

notably higher counts and lower counts were found at widely scattered locations (Fig. 2B, red and white

numbers; appendix in bold type). Among the higher counts are specimens from the Caucasus Mountains,

one with 15 rows (Princeps Kascelsky, ex. Herb. Hort. Imper. Petro), another with 13 rows (Woronowa s.n.). Oc-

casional specimens with 12 rows are from Bosnia (Biol. Inst. Dubrovnik 37), Romania (Topa, Bot. Mus. Exsic),

Finland (Florstóm s.n.), Portugal mainland (Yoller 61), Portugal Azores (Goncalves 4625), and Algeria (Swingle

s.n.). Those that were noted to differ in other morphological features are indicated in bold type, including also

Spjut, Phytogeographical analysis of Taxus 299

specimens with 11 stomata rows per band, such as one specimen from England (Bowden & Hillman 433) that

had globose shaped epidermal cells, instead of the usual elliptical shape; others such as Hauti 28894 from

the British Isles and Busch s.n. from the Caucasus Mountains had a broader marginal zone of bare cells, as

indicated later under results for marginal cells. A higher number of stomata rows with a wider leaf margin

indicate a higher density of stomata as seen in the Cuspidata Alliance. The range of variation for the Baccata

Alliance was greatest in Transcaucasia.

Leaves with fewer than 8 stomata rows per band include seven specimens with 7 rows—from Germany,

Austria, Switzerland, Bosnia, and Turkey; these belong to the T. baccata Alliance. Those with fewer than 7

rows were relatively rare (1-296 of 196 specimens cited for the Euro-Mediterranean Region), two specimens

with 5 rows from Norway and Sweden, and one with 4 rows from Slovenia. These are considered T. canaden-

sis as shown later. A cline is not apparent in the Euro-Mediterranean as it is in North America; however,

the relative frequent occurrence of stomata in 8-10 rows per band appears significant when compared to a

greater range of variation in E Asia (7-21 rows per band, Fig. 2C).

In E Asia, the number of stomata rows per band ranged from 7-16 in the temperate region, and from

(5—) 7-19 (-21) in the tropical region with two patterns converging in the Himalayas, one from the west

with 5-8 (-11) rows of stomata, and another from the east with 7-21 rows per band.

Number of Marginal Cells (MC).—The absence of papillae nearest the leaf margin, as measured by

the number of marginal cells (MC) across between the margin and stomata band, is depicted geographically

in Figs. 3A and 3B, and detailed in the Appendix.

Data on marginal cells lacking papillae along the abaxial surface (MC) were recorded less often for

North American species because they were distinguishable early in the study (Hils 1993; Spjut 1992, 1993).

It was recognized that marginal cells of T. canadensis always lacked papillae (Hils 1993, Spjut 1992, 1993,

1998a, 2000b), which has since been determined to vary from 11-19 cells across in North American plants

(Appendix); the absence of papillae on the abaxial midrib is in sharp contrast to the papillose midribs of

other North American species (T. brevifolia, T. globosa).

In the Euro-Mediterranean, the abaxial surface of leaves of most Taxus specimens (27596) lacked-papil-

lae along a relatively narrow marginal zone of 4—7 cells across (Fig. 3A). This included the lectotype for T.

baccata and two specimens from the Caucasus Mountains that, unlike the lectotype, were found to have a

relatively high stomata count as noted earlier—one with 13 stomata rows per band, and one with 15 sto-

mata rows per band—and also one specimen from Bosnia with 12 stomata rows per band. A specimen from

the Caucasus Mountains—that had abaxial marginal papillae to within one cell from the margin—lacked

papillae on nearly half of the cells across the midrib in the median region (Woronowa s.n.).

Leaves of Euro-Mediterranean specimens with a relatively broad zone of bare cells between the margin

and stomata band—from 8-24 cells across—were found less frequently (~25%). These are from widely scat-

tered places. Many are indistinguishable from T. canadensis in North America—based on additional characters

of branching, phyllotaxy, and color (Figs. 4—5); therefore, are referred to T. canadensis (Spjut 2000b). In

Europe, leaves of T. canadensis may include a transitional zone of papillose cells between the stomata band

and margin (Fig. 4, specimen from Morocco). These plants may be hybrids between T. canadensis and T.

baccata.

The Cuspidata Alliance showed an intermediate range of values for abaxial marginal cells without pa-

pillae, (62)8—18(724) cells across (Figs. 3B, 6), compared to the Baccata Alliance, (134-711) cells across

(Fig. 3A) and the Sumatrana Group, 8-36 cells across (Fig. 3B, 6). The higher stomata counts in relatively

narrower stomata bands for the Cuspidata Alliance (see also Dempsey & Hook 2000) and Sumatrana Group

means in effect they have a higher leaf stomata density, recognizing also that stomata density is related to

width of the epidermal cells and the width of the stomata band (Nicolosi 1982), and that a transitional zone

of papillose cells is always present in the Baccata Alliance but not in the Cuspidata Alliance. Data on number

of papillose cells across the abaxial margin were not included in this study because this was observed to

be highly variable, although the absence of papillae in T. canadensis has taxonomic significance in North

American species as already indicated (Hils 1993; Spjut 1992, 1993, 1998a, 2000b).

300 Journal of the Botanical R h Institute of Texas 1(1)

» E

p" Norwexian

Fic. 3A. Number of epidermal marginal cells (MC) without papillae bet th in and stomata band (abaxial surf t margin) for repre-

sentative specimens of Goi from the Euro- anar Regon see SERENU for locali data. Numbers in red show tl gei

tian a laaf marni 7 ralle acrnace Marit R i N } i hit T +

Guivil Gita ma 1y" n4 / NCHS GRIVOD Y

the higl a leaf margin 6-24 cells wide that belong to 7. canadensis Data in yell bers indicate int liates that hol

cally similar to T. . biternata, T. canadensis, T. cuspidata, or T. contorta, appearing frequent in speci from the C Mountains (Apr ;

data in bold type)..

Fic. 3B. Numl f

ited in the A dix. Data for the Wallichiana Sul 17. contorta (B ta G ) i 1 apt gala Dont

FL* >. e L H ae: re £ I J IL 4 £, 4l 44l J e MET: pea Y T E All

TUTUI r contrast uie WIUC rang

cata Group) and the emanate SIUE The UNIAN Group includes 7. a in maroon, 7. mairei in black, T. kingstonii in white, and 7. sumatrana

in yellow. The C Lb T. cuspidata i , T. caespitosa and T. umbraculifera in red. Note that the widest

range in variation occurs in SW China.

Spjut, Phytogeographical analysis of Taxus 301

Approbavit

Madeira ias a

Fic. 4. Pramples of T. augue. var. canadensis as characterized uf the

iat

,arranged

+1 Ļ Ln | £. £i I ITI

in n nearly two ranks, and by

on at bd 8 bud across Eu the margin, M = Marginal callsa across sth

The specimen from Morocco pas a pantal Dance maigin, 3 9 of the

18 cells between tl | mens

above left, above right and left: Bean et al. 19634 (PH), n 570 UN

Font Quer 1928 (BM).

Nova Scotia Madeira Morocco

MC s 14 8-9(18)

SR 7 iT] 8

The absence of papillae along the abaxial mar-

| ginal surface is most variable in E Asia (0-36 cells

across, Figs. 3B, 6). This variation is related to taxo-

nomic features that define the species groups and the

species themselves as shown in Fig. 6. In Fig. 3B, the

& numerical data for the Taxus wallichiana Group, and for

the NW Himalayan species, T. contorta (Baccata Group),

were summarized because leaf stomata bands are con-

sistently bordered by a relatively narrow marginal zone of epidermal cells without papillae—most often 4

cells across (Fig. 6)—as seen also in the Baccata Alliance (Fig. 3A, Appendix). Asian plants with a marginal

border of fewer than 4 cells across generally belong to Taxus wallichiana var. yunnanensis (W.C. Cheng &

L.K. Fu) C.T. Kuan, or to T. suffnessii Spjut, whereas specimens with an abaxial leaf margin exceeding 7

cells in width usually belong to the Sumatrana Group and Cuspidata Alliance, but there are several notable

exceptions. One is Taxus chinensis with a leaf margin of 4-12 cells wide that is clearly intermediate between

the Sumatrana Group and the Cuspidata Alliance, corresponding also to its intermediate geographical posi-

302 Journal of the Botanical R h Institute of Texas 1(1)

Lor

Slovenia

| Fic. 5A. Examples of T. canademis var. Upa fom North America 2s

Europe MC ll

| and bouis pand: R refert b the Page s stomata rows in a stomata

ban

Anderson s.n. (US: 1091452),

ie (US), and Berglund s.n. (S: C- iR

lowa Norway Sweden Slovenia

MC Pas 18 2

SR 6 5 5-8 4-6

ELEELLLELELLEEEEEUEEEEEEEEEELLL LLL Í

Spjut, Phytogeographical analysis of Taxus 303

P Ni T" y "NW "M

lowa m

Fic. 5B. Shows examples of distinct alternate branching in specimens

regarded as strong evidence for the same taxon occurring at widely

disjunct localities.

tion in central China. Another is the central Himalayas T. contorta var. mucronata Spjut, a variety that is

recognized by shorter reflexed leaves as in T. umbraculifera (Cuspidata Alliance) but also with slightly more

stomata rows per band (8-11) and a wider leaf margin (8—10 cells across) than what is usually seen in the

typical variety of NW Himalayas.

Data in Figs. 3B and 6 also contrasts the wide variation in the number of marginal cells in the Sumatrana

Group and Cuspidata Alliance with other Asian taxa. The Sumatrana Group and Cuspidata Alliance share the

elliptical shape of epidermal cells as seen in T-section, in contrast to the angular leaf epidermal cells of the

Wallichiana Subgroup, and also lack of papillae on the abaxial midrib as well as along marginal regions (e.g.,

T. sumatrana, Fig. 1C). This group usually occurs at elevations below 2000 m in contrast to T. wallichiana

found mostly above 2300 m. One exception, T. kingstonii Spjut, in the Sumatrana Group, is ecologically and

morphologically intermediate between the Wallichiana and Sumatrana Groups.

Variation in the number of abaxial marginal cells recorded from the same plant, or related plants at the

same locality, was d similarly to data compiled on number of stomata rows. Among duplicate herbarium

specimens, the count was the same in nearly half of the duplicate sets. Most variation within individuals, or

within a population of individuals, occurred in the Sumatrana Group (T. celebica [Warb.] H.L. Li, T. kingstonii,

T. mairei [Lemée & H. Lév.] S. Y. Hu ex T.S. Liu, T. sumatrana; Fig. 3B, Appendix). For example, leaves from

duplicate specimens of T. mairei from Guangdong often lacked papillae along the abaxial margin on either

14 or 24 cells across. Similar dimorphic differences are evident in specimens from Sichuan and Guizhou,

and in T. kingstonii from Yunnan. These differences may be due in part to leaves from different plants, or

from different ages of shoots, or from different heights on the plant (de Laubenfels 1988), or in T. celebica,

from slightly different regions of the leaf.

In the case of Wilson 1265 (A, BM, K, S, US), which was mentioned earlier as having been collected

from three different locations (Rehder & Wilson in Sargent 1914), one site in western Sichuan at 600-650

m included duplicate specimens that was found to have either 15 (US) or 21 (A) stomata rows per band,

and either 16 (US) or 25 (A) marginal cells. Wilson also collected seed (Wilson 1265) from Sichuan near Mt.

Emei and/or Yachou Fu at 600 m from which leaves in three herbarium specimens obtained from a plant

grown from seed (of Wilson 1265) at the Royal Botanic Gardens—Kew lacked papillae entirely along an ab-

axial margin zone, 18 or 28 cells across, while all three had 8—10 stomata rows per band. This plant is not

T. chinensis as indicated in the literature (Rehder & Wilson in Sargent 1914), but Taxus mairei var. speciosa

(Florin) Spjut that appears atypical by the relatively large greenish distant leaves that are more characteristic

of T. celebica. Photographs of a very similar plant in cultivation at the Royal Botanic Garden—Edinburgh

(probably from Wilson 1265 seed) are shown in van Gelderen and van Hoey Smith (1996) and in Krüssmann

(1985). However, Wilson 1265(b) does include one specimen I identified as T. chinensis that was reportedly

obtained from western Hubei south of “Ichang,” 600-1300 m.

304 Journal of the Botanical R h Institute of Texas 1(1)

Sumatrana Group Cuspidata Alliance

sg 20 90.

É g 10 E $ 10 |

Q E 7

5 5 5 |

zo 0 20 0 To TT T ndn [Eng | T dias xl T x: T 7T T

— © O MO r- — YD BD o - + N oO N

v == N N N (09) = = N

Bare Marginal Cells Bare —— Calle

T. chinensis T. wallichiana

52 20 52 60

LT w 15 D v 40

2 E 10 | 2 E 5n |

® 5 | E

2 o 0 | t T j 1 q li LI [1 n, [Y DE M 1 2 ri 0 I o a n N m- T =T T l 1

123456 7 8 9 10111213 12845678 904121814

Bare Marginal Cells Bare Marginal Cells

Fic. 6. Number of K peci ling to the number of bare (without papill I baxial surface of |

stomata band for selected taxa from E Himalayas to China. Note that the Sumatrana Group has the widest leaf marginal zone without papillae, and

that 7. wallichiana has the narrowest zone, mostly 4 cells wide (40 specimens).

De Laubenfels (1988), commenting on his field observations of yews in Taiwan, suggested that differ-

ences in leaf shape may be seen on the same plant and further implied the same for the presence or absence

of leaf papillae.

Leaves of Taxus mairei var. speciosa that showed considerable variation were also studied from plants

cultivated in the United States. An examination of 20 leaves (Phyton s.n.)—from apical buds to 3" yr

branchlets—were found to be relatively constant in the number of marginal cells without papillae—9 cells

across—and also in having 16-18 stomata rows per band. They were notably variable in shape and length

of epidermal cells, especially juvenile foliage.

The abaxial leaf margin in T. mairei specimens obtained by C-j. Chang from near Hualien, Taiwan varied

by four cells (4—7) at four of six locations (Nos. 2, 4, 9, 10), and by only two cells (0—1 cell) at the two other

locations (Nos. 1, 5).

Finally, specimens of T. celebica from South Vietnam by Schmid (1974) were found to lack papillae on

either (23-) 24 or 32 cells across the leaf margin. Here Schmid (1974) reported that Taxus was polymorphic,

and among his specimens at the Museum of Natural History in Paris (P), is an apparent hybrid (Schmid s.n.)

between T. aff. chinensis (Poilane 4150) and T. celebica.

DISCUSSION

Phytogeography of Taxus.—Data presented for leaf character attributes of Taxus (Figs. 2 and 3) show that

stomata rows and marginal cell features are most diverse in SW China, while the same number of subspecies

(Pilger 1903), or species (Farjon 1998; Silba 1984) have been recognized to occur in both North America

and Eurasia; thus, the traditional separation of Taxus species (or subspecies) based on these geographic

discontinuities is a distorted classification. The phytogeographic data in this study support the taxonomy

of yew for only the geographical disjunct occurrences in North America.

In North America, leaf stomata of Taxus brevifolia show a cline in number of stomata rows ranging from

9 rows in California to 4 rows in the northern Rocky Mountains, and also in length of abaxial epidermal

cells relative to width (1/w) from an average of ca. 3x l/w to 8x l/w. Molecular differences have been re-

ported between coastal and inland yews (El-Kassaby et al. 1994, 1995) at more northern localities. Clinal

Spjut, Phytogeographical analysis of Taxus 305

variation in conifers has been linked to historical migrations and hybridization patterns with the advance

and retreat of glaciers since the Pliocene (Wilkinson et al. 1971); however, the cline in stomata data for the

North American Wallichiana Subgroup, including Mexico, seems best explained by loss of stomata in leaves

of Taxus as it may have migrated northwards during the Neogene, when the climate may have become in-

creasingly warmer and drier, while the differences within the Pacific Northwest may be a product of more

recent climatic changes (Graham 1999).

Also, a geographical species concept that recognizes T. sumatrana as widely distributed in SE Asia (de

Laubenfels 1988) might conclude that its distribution was achieved from long-distance dispersal by birds,

whereas geographical disjunction of Taxus in North America has been correlated with paleobotanical data

(Graham 1999). This seems paradoxical; i.e. the greater variation in leaf anatomical data of Taxus in Asia

should also be explained by evolution and paleogeography—perhaps the result of climatic and geomorphic

changes that have occurred since the Cretaceous, a period of 130 million years (my). Therefore, the discus-

sion that follows will focus on this latter hypothesis.

Although data on leaf stomata rows in Taxus are more variable in Asia than in North America and

Europe, relationships become evident when other taxonomic features are taken into consideration (Spjut

2007). For example, in the western Himalayas, the stomata counts that range from 5-8 (-10) or 8-11 stomata

rows per band (Fig. 2C) are a characteristic feature of T. contorta Griff. This species is also recognized by

the long narrow leaves that have idioblasts in the spongy mesophyll (vesicular cells appearing dark red in

herbarium specimens), a character trait not seen in the E Himalayan yews. Moreover, these features show a

closer relationship to European yews than to Asian yews. In the E Himalayas, T. wallichiana—indicated to

have 11-19 stomata rows—is recognized by leaves having large angular shaped epidermal cells as seen in

T-section, by the persistent bud-scales at the base of branchlets, by the branchlets that show a marked color

change in their 2*1 yr of growth—from yellowish green to maroon or reddish orange, and by the bone-like

parenchyma cells in the spongy mesophyll that connect in a reticulate pattern with rounded to angular

intercellular spaces. These morphological features are considered more closely related to yews of SW China

than to T. contorta of W-C Himalayas.

Data in the appendix take into account variation in T. chinensis and T. wallichiana on Mt. Emei. Speci-

mens are arranged according to increasing number of marginal cells along the abaxial surface of the leaf

without papillae. In T. wallichiana, the epapillose marginal cells, which are consistently 4 wide for numerous

specimens in the Himalayan Region (see also Fig. 10 in Spjut 2007), appear to show greater variation on

Mt. Emei where it was found that two of six specimens had a leaf margin 8 cells wide. Similarly, T. chinensis

outside of Mt. Emei was usually found to have a relatively narrow leaf margin of 4—7 cells wide, 25 of 30

specimens (8396); only 2 specimens (796) were found with a leaf margin greater than 8 cells wide, whereas

on Mt. Emei, 11 of 30 specimens (3796) had a relatively broad leaf margin (8-12 cells across). The broader

leaf margin in T. chinensis from Mt. Emei could be the result of recent hybridization with species of the

Sumatrana Group, or possibly reflects historical introgression with T. umbraculifera of NE China. Hybridiza-

tion might also account for similar variation in T. chinensis for three specimens from Guizhou, Shaanxi, and

Vietnam.

The development of leaf papillae in Taxus along the abaxial marginal zone may be partially correlated

with latitude as evidenced by the narrower range of marginal cells without papillae (7—24 cells across, Figs.

3B, 6) for the T. cuspidata Alliance in temperate NE Asia, compared to that of the more widely distributed

Sumatrana Group (8-36 cells across, Figs. 3B, 6) in SE tropical Asia. At increasingly higher latitudes, plants

with more papillae on their leaves obviously receive greater protection from ultraviolet rays of the sun—dur-

ing the longer summer days. The refractivity (protective) effect of papillae on Taxus leaves has indeed been

mathematically demonstrated (von Frimmel 1911). Nevertheless, hybridization between the tropical and

temperate species alliances in E Asia cannot be ruled out.

In the Cuspidata Alliance, I have observed that papillae are of lower stature and concrescent near cell

walls in which the cell walls appear thicker, examples of which are shown in Jinxing and Yuxi 2000). This

may be evidence of introgression with the Wallichiana Group from which T. chinensis allegedly evolved. As

306 Journal of the Botanical R h Institute of Texas 1(1)

indicated, leaves of T. chinensis often have elliptically shaped epidermal cells in T-section, a slightly wider

marginal border, ranging from 4—12 smooth cells across (Fig. 6), and midrib papillae often more conspicuous

along cell walls. Thus, the Cuspidata Alliance, which is undoubtedly related to the Baccata Alliance (Collins

et al. 2003; J. Li et al. 2001), may have acquired an expanded leaf margin as a result of hybridization with

species of the Sumatrana Group.

In cultivated individuals related to T. cuspidata and T. mairei, papillae sometimes were found on mid-

ribs of young leaves, but not the older leaves. However, the odd leaf mentioned earlier for one cultivar (T.

caespitosa) with 16 instead of 13 stomata rows/band was found with low papillae on its midrib, whereas the

other 15 leaves had smooth midribs; this odd leaf may have retained juvenile characteristics due to lack of

exposure to light. I have also completely “skinned” leaves to evaluate the distribution of papillae from base

to apex in specimens from Europe, Taiwan and the Philippines, and have found papillae to develop more

in the upper half (towards apex). The presence of midrib papillae on juvenile leaves, thus, may indicate an

ancestral trait that should not be treated as a justification for lumping all variation within a geographical

area under one species.

This alleged ancestral trait is also evident among specimens that are intermediate between T. chinensis

and T. mairei, and the extinct T. engelhardtii (Fig. 7). The characteristics of T. mairei include larger (mamillose)

epidermal cells on the abaxial midrib and marginal zones, and isodichotomous zigzag branching; those

of T. chinensis are the marginal papillae on the abaxial midrib [e.g., Ching 1676 from Sichuan; Chiao & Fan

464 (US) from Sichuan, and Tsiang Ying 1425 (P)]. A study by Kwei and Hu (1974)—that mentioned 30 of

the specimens cited in the Appendix—recognized intermediates by a partially papillose midrib; however,

Spjut (1992, 1993, 19982) has since reported other correlative taxonomic characters—such as shape of leaf

epidermal cells, development of papillae along the abaxial marginal zone and size of bud-scales—to help

further separate these species. It should also be noted that midrib papillae can be consistently present in

the W Himalayan T. contorta (Kvacek 1984), or consistently absent in the North American T. canadensis.

From Myanmar are four specimens found to have leaves almost entirely papillose within a few cells from

the abaxial margin. Three of the specimens are recognized as belonging to a distinct species (T. suffnessii) by

the relatively large and persistent bud-scales at base of branchlets, by the conspicuous papillae on epidermal

cells, and by the relatively tall-rectangular epidermal cells as seen in T-section of leaves (Spjut 2007). One

of two other specimens from NE India and Bhutan (Ludlow & Sherriff 18762, 3719)— that was recorded to be

papillose within 2 cells from the margin— differed by having elliptical instead of angular epidermal cells.

It would appear, then, that the occurrence of papillae on the abaxial surface of Taxus leaves has taxonomic

significance even when the numerical differences are relatively narrow as also seen in North American T.

globosa var. globosa and T. globosa var. floridana in which intermediates are recognized to occur in northern

Mexico.

Disjunct Relationships between Eastern Asia and Western North America.—Disjunct geographic

distributions in Taxus and other genera have long been recognized between temperate North America and

Eurasia (Axelrod 1983; Boufford & Spongberg 1983; Good 1964; Graham 1972; Hara 1972; Kornas 1972;

H. Li 1952; Tiffney 1985a; Qian 2002; Q. Wang et al. 2006); however, their rate of evolution varies. For

conifers this has been considered relatively slow (Prager et al. 1976). In the genus Abies, for example, the

subalpine fir in W North America [A. lasiocarpa (Hook.) Nutt.] appears more closely related to an endemic

species of Taiwan [A. kawakamii (Hayata) Ito] than to any of the 10 other American species (Farjon 1990;

Hunt 1993; Liu 1971). Indeed, recent molecular studies by Suyama et al. 2000) show A. mariesii Masters of

Japan to be more related to species in North America than to its relatives in Japan. Additionally, species of

Pseudotsuga (+4 spp., Farjon 1990) in Asia may have been derived from ancestors in North America (Strauss

et al. 1989).

The Wallichiana Subgroup of Taxus—characterized by angularly shaped epidermal cells in T-section—oc-

curs primarily in E Himalayas to SW China (Sichuan, Yunnan) and in North America (Spjut 1998a, 1998b,

20002). Within this subgroup, leaves of Yunnan and Sichuan plants (T. florinii, Spjut) appear indistinguish-

able from those of the American T. globosa (Spjut 1998b, 2000a, 20009). Other specimens from Myanmar

Spjut, Phytogeographical analysis of Taxus 307

Fic. 7. Comparison of | f extant 7. mairei (clear photos, isotype, P) with extinct T. engelhardtii (grainy photos, reproduced from Kvacek 1984), from

an Oligocene deposit in Bohemia.

(T. suffnessii) are similar to T. brevifolia in the relatively large bud-scales and tall rectangular epidermal cells

as seen in T-section (Spjut 20009).

An analogous disjunct relationship is seen among the white pines, Pinus monticola Douglas ex D. Don

of W North America and P. wallichiana A. B. Jackson of Myanmar (Axelrod 1986, Pinus griffithii [Hook. f. &

Thomson] Parl.). They are remarkably similar in cone morphology and needle chemistry. The antiquity of this

relationship is supported by their turpentine chemistry of saturated straight chain hydrocarbons—undecane

and heptane, the chemical structures of which are considered more archaic among the terpenoid compounds

in pines (Mirov 1953). Additionally, heptane occurs in the Mexican P. ayacahuite Ehrenb. ex Schltdl. (Mirov

1953), along with a bicyclic sesquiterpene—cardenine—that has also been found in P. parviflora Siebold &

Zucc. of Japan (Mirov 1953). The close relationship among these species, which belong to sect. Quinquefoliae

subsect. Strobus, is supported by molecular data (Liston et al. 1999) from which it has been suggested that

the ancestors probably originated in the *Old World" (Gernandt et al. 2005).

In angiosperms, it is interesting that Phipps (1983) recognized—among -145 species of hawthorns—

Crataegus mexicana Moc. & Sessé, a widely distributed species in Mexico and Guatemala, to have its closest

relative in Yunnan, C. scabrifolial (Franchet) Rehder, and that both are the *most primitive" of a taxonomi-

cally complex Laurasian genus, which has numerous species in both Mexico and in Yunnan.

For taxads and other conifers, diversity is greatest in SW China (Figs. 2, 3; Cheng & Fu 1978; Prakash

et al. 1995; Qian & Ricklefs 1999). Ancestors related to Taxus suffnessii Spjut in Myanmar (Appendix) may

have immigrated to North America across a former Aleutian (or Bering) land bridge (Hamilton 1983; Millar

1993)—as suggested for Crataegus (Phipps 1983). A logical time for this to occur would have been during

the latter half of the Cretaceous (110-100 mya), after Pangaea had fragmented (Graham 1993)—when an

epeiric sea (Wolfe 1975) possibly divided the North American continent into distinct west and east floras

(Graham 1999; Srivastava 1994; Thorne 1972, 1978). Late Cretaceous fossils related to the Alaska cedar,

Callitropsis nootkatensis (D. Don in Lambert) Florin, which includes one related sister species in North

Vietnam, and is also sister to other species in North America (Little 2006), have been found on Vancouver

Island (Mclver 1994), and an early Cretaceous fossil, Chamaecyparis eureka Kotyk, from Eureka Sound in

the Canadian Arctic, is most similar to the extant Ch. pisifera Siebold & Zucc. in Japan (Kotyk et al. 2004).

Additionally, fossil cones of Thuja smileya LePage from Late Cretaceous deposits on the North Slope of Alaska

are indistinguishable from modern species (LePage 2003).

As climate temperatures declined during the Cretaceous (Axelrod 1958; Frederiksen 1994; Graham 1999;

Novacek 1999; Srivastava 1994), Taxus might have retreated southwards, perhaps reaching southern Mexico

by the end of the Cretaceous (65 mya); similar retreats have been suggested for other genera (Phipps 1983;

Sharp 1966), but for the Tertiary Period (Phipps 1983), not the Cretaceous. A later migration and extinction

of Taxus, such as in the Tertiary near the Eocene-Oligocene boundary, may seem like a more reasonable

time frame for evolution of North American Taxus, but there also has to be ample time for diversification of

308 Journal of the Botanical R h Institute of Texas 1(1)

the Cuspidata Alliance as well as the alleged migration and extinction of the Wallichiana Group across the

Sino-Japanese Region.

The end of the Cretaceous is marked by a distinct change in the geochemical and fossil records (Mclver

1999; McIver and Basinger 1999; Novacek 1999)—indicating a rapid climatic warming— possibly due to a

meteor impact in the Caribbean Sea that might have caused massive volcanic materials to erupt and cloud

the atmosphere (O'Keefe & Ahrens 1989)—a ‘greenhouse’ calamity that could explain evidence for *eco-

logical deserts” (Tschudy et al. 1984)—-and mass extinction of major taxa (e.g., dinosaurs, Novacek 1999).

This could have extirpated yew north of Mexico; Cretaceous fossils of gymnosperms of the Taxodiaceae

(Metasequoia, Sequoia, Sequoiadendron), and Amentotaxaceae (Amentotaxus, Torreya) are known as far south

as New Mexico and North Carolina (Florin 1963) for which Taxus has had a long history in association

(Florin 1951, 1963; Kvacek 1984) but whose fossils may not always be preserved or identified.

Paleontological evidence indicates that following the Cretaceous a warmer subtropical humid climate

(Chaney 1947; Frederiksen 1994; Tiffney 19852) prevailed over much of North America until the late Eocene

(ca. 50 mya; Chaney 1947; Graham 1999; Novacek 1999; Srivastava 1994; Wolfe 1975). Assuming that an-

cestral T. globosa had survived only in Mexico, a northward migration (as the climate warmed) would account

for the cline in leaf stomata data of Taxus in W North America (Fig. 2A). Other North American conifers

with evidence of a southern ancestry include Douglas fir (Pseudotsuga menziesii (Mirb.) Franco), most likely

derived from big cone fir (P. macrocarpa (Vasey) Mayr), endemic to S California (Strauss et al. 1989), and

pines that may have drifted northwards on the San Andreas rift system (Axelrod 1986)—a system that may

have included Vancouver Island originating perhaps from as far south as *lands end" off the cape (Cabo San

Lucas) of Baja California (90 mya, Ward et al. 1997). A northward range extension of ancestral Taxus globosa

may be further correlated with the change in a Rocky Mountain flora from paleotropical (boreotropical)

to neotropical elements during the mid Eocene (Leopold & MacGinitie 1972), and later along the Gulf as

evident from biogeographical data on fishes and amphibians (Rosen 1975), maples (Acer saccharum L. Group;

Humphries 1982), and other taxa (Burnham & Graham 1999)—emphasized by Axelrod (1975, 1986).

As the climate became drier with the uplift of the W Cordillera (Chaney 1947; Wolf 1969), the range of

Taxus, like other Arcto-Tertiary genera, diminished (Axelrod 1975, 1983; Graham 1993, 1999), while other

taxa evolved (Axelrod 1958). Fossils of Taxus have been reported in Eocene (54-38 mya), Oligocene (38-27

mya), and Miocene (27-10 mya) strata of W North America (Gaussen 1979; Kvacek & Rember 2000, in

press; Manchester 1994; Meyer & Manchester 1997) in association with species of Tsuga, Abies, Lithocarpus,

Quercus, Acer, Alnus, Cornus, Carpinus, Castanea, Fagus, Liquidambar, Nyssa, Ostrya, Platanus, Tilia, Ulmus,

and Cercidiphyllum (Graham 1999; Whittaker 1961).

Extant species of Taxus are still found with these same genera today in mixed mesophytic forests of S

Appalachia (Braun 1950), China (Hou 1983), and Japan (Hayashi 1954). Taxus brevifolia allegedly evolved

from an ancestral T. globosa complex as the climate became cooler and drier during the Eocene (56-34 mya;

Graham 1999), while closer ties between the Mesoamerican yew and Florida yew were likely maintained

until the Pleistocene as evidenced by the close similarity among many shared taxa between the two regions

(Sierra Madre Oriental and S Appalachia). It is interesting that specimens of Florida yew appear indistin-

guishable from those occasionally collected in Veracruz and in Nuevo Leon/Tamaulipas, Mexico (e.g., Meyer

& Rogers 2746, BM; Mueller 1337, BM, PH), where they reportedly occur with Carpinus caroliniana Walter,

Cercis canadensis L., Frangula caroliniana (Walter) A. Gray, Hamamelis virginiana L., Liquidambar macrophylla

Oerst, Magnolia schiedeana Schltdl., Prunus serotina Ehrh, and others also found in S Appalachia, including

many lichens and mosses (Culberson et al. 1990; Graham 1973, 1999; Miranda & Sharp 1950). Moreover,

it has been shown that the Florida yew and Mesoamerican yew form a clade with the Pacific yew as a sister

species (J. Li et al. 2001), and that the Florida and Mesoamerican yew are indeed more closely related (J. Li

et al. 2001).

As previously noted, genera found with the Mesoamerican yew also occurred with Pacific yew (Graham

1999), but in the Pacific Northwest many of these genera perished—such as Carya, Disopyros, Fagus, Hama-

melis, Liquidambar, Liriodendron, Magnolia, and Morus (Axelrod 1975, 1983, 1986; Graham 1999; Manchester

Spjut, Phytogeographical analysis of Taxus 309

1999; Wood 1972). Liriodendron is one of many known from fossils in Europe and W North America with

relatives now surviving only in E North America and E Asia (Axelrod 1983; Manchester 1999; Schuster

1976). They perhaps were part of a widespread Tertiary “boreotropical” forest (Wolfe 1975; Graham 1999),

which may have included T. canadensis (Figs. 4, 5).

As conifer diversity declined and grassland vegetation expanded during the Miocene (Axelrod 1976;

Jacobs et al. 1999), Taxus possibly had attained maximum diversity in geographic and ecological species

isolation. McIver and Basinger (1989) found in Eocene deposits cones similar to western red cedar (Thuja

plicata Donn ex D. Don) that may have been derived from an earlier complex related to the extinct Thuja

polaris Mclver et Basinger, which they described from a Middle Paleocene deposit on Ellesmere Island,

whereas other cedars related to Th. occidentalis L. are not known before the Miocene (Mclver and Basinger

1989). Moreover, Th. occidentalis is recognized in the fossil record from the late Pliocene (Bennike 1990).

The redwood, Sequoia sempervirens (Lamb. ex D. Don) Endl., is hardly distinguishable from a former wide-

spread S. abietina (Brongn.) Knobloch—known from the Upper Eocene to Upper Miocene (Mai 1998). A

species of Taxus from a Middle Miocene deposit in N Idaho (Clarkia area Latah Formation) has nearly the

same abaxial epidermal features seen in the extant T. brevifolia (Kvacek & Rember 2000, in press). Klicka

and Zink (1997) concluded from DNA evidence that North American species of song birds had already

originated by early Pleistocene, and that subsequent glaciation was more of an “obstacle course" for their

survival. Yew species, by comparison, are likely to evolve more slowly as a yew trunk may live 3000 years

(Larson et al. 2000; Thomas & Polwart 2003; Voliotis 1986), and still may survive by producing trunks

from adventitious shoots (Hageneder 2007; Loudon 1844). The greater diversity of Taxus in SW China may

also be related to less species extinction there as seen in many plant genera as a result of increasingly drier

and cooler climates that had a more profound impact on the vegetation elsewhere since the Middle Miocene

(Axelrod et al. 1998; Kubitzki & Krutzsch 1998).

Relationships between Eastern North American and Eurasian Taxus.—While the Pacific floristic

element of Taxus in North America is represented by three disjunct taxa within the Wallichiana Group, the

Atlantic floristic element has only T. canadensis, a species that appears more related to the Baccata Group (J.

Li et al. 2001; Spjut 2007) than to T. globosa by its elliptical shaped epidermal cells in T-section, and by its

subcylindrical seed shape (Spjut 1998a, 2000). The lack of papillae on the abaxial leaf surface between the

margin and stomata band that characterizes the North American Canada yew is also seen more frequently

in yews of temperate NE Asia (Cuspidata Alliance) than in the Euro-Mediterranean (Baccata Alliance), and

its leaf epidermal features are most similar to T. biternata Spjut, a species closely allied to T. cuspidata. The

close relationship between T. canadensis and T. cuspidatais supported by molecular data (Collins et al. 2003).

Taxus biternata differs from T. canadensis by the 2-3 angled seeds (tapered part) developing on 1* yr branch-

lets, and by the tree habit (Spjut 2007). The complete lack of papillae along the abaxial leaf marginal zone

is a relatively rare occurrence in European yew («296), but this character trait may have once been common

in that region; for example, three species described by Kvaéek (1984) from leaves of fossil assemblages in

Europe—dating from Oligocene to Pliocene—all lacked papillae entirely between the stomata bands and

margins.

Furthermore, a “Taxus (sp. 1,” Kvatek 1984; Fig. 8) of Lower Miocene age is, in my opinion, T. canadensis.

Its leaves are more similar to American plants than to European plants, which differ by the distinctly papillose

stomata bands—except perhaps for rare North American specimens (e.g., Coy & Glen from Ithaca, New York).

These extant European variants could be referred to the extinct T. grandis Krausel or T. inopinata Givulescu

(1973)—described from Tertiary deposits in Europe (Kvacek 1984). Data for numbers of stomata rows (Figs.

2A, 2B) also support my hypothesis that the North American T. canadensis came from Europe, possibly arriving

late Paleocene or Eocene when migration across the Atlantic was possible by land (McKenna 1983; Tiffney

1985b), as suggested for the evolution of Cornus sessilis Torr. ex Durand (Xiang et al. 2005, 2006). During this

period the Gulf Coast flora shows evidence of many immigrants from Europe (Frederiksen 1994, 1995) that

included species of Fagopsiphyllum, Hydrangea, lodes, Koelreuteria, Langtonia, Nyssa, Palaeophytocrene, Pentoper-

culum, Platanites, Platycarya, Pyrenacantha, Sargentodoxa, Symplocos, Tapiscia, Tetraclinus, and Tilia (Manchester

310 Journal of the Botanical R h Institute of Texas 1(1)

Fic. 8. Comparison of T. canadensis var. minor from Madeira (Lowe ex

Barby s.n., US) and Maine (Fellows 5686, US) above with close-up on

middle left showing seed on 2" yr branchlet, and below photomicro-

Inga M5 f, il f C hR bli (“Taxus 5p. 1; Kvacek 1984)

with leaf fragment (“d” on right). This variety is recognized by the more

Wil

34 b èz

spreading obtuse leaves in var. T. canadensis var. adpressa. Note the

F r Lui a

Spjut, Phytogeographical analysis of Taxus 311

1999). It is interesting to note that 10 species (in 9 genera) of lichens recently discovered to occur in E North

America—on Mt. Katahdin in Maine—were previously known only from northern and/or central Europe,

except for one species that also occurs in Greenland and Siberia (Fryday 2006).

Relationships between the Euro-Mediterranean and Asian Taxus.—The European yews have all

been considered a single species, T. baccata; however, additional species appear evident. The lectotype (of T.

baccata) has leaves arranged mostly parallel to one other along two sides of a branchlet in a flat spray with

8-10 stomata rows/band and a papillose undersurface—from the midrib to near the margins. The typical

European yew is further characterized by isodichotomous branching, pale glaucous green leaves similar

in color on both surfaces, and cones maturing on branchlets that have terminated their growth as evident

in specimens from England, Germany, Austria, Switzerland, Czech Republic, Spain, Portugal, Algeria,

Morocco, Italy, Albania, Bosnia, and Turkey. This is in contrast to another widespread species, T. recurvata,

that I recognize by a less parallel arrangement to the leaves with a sharper contrast in color between the

leaf surfaces—notably yellowish green on the abaxial surface and dark green on the adaxial surface, and by

cones developing on branchlets that continue their growth. Both have many naturally occurring varieties

based on differences in leaf arrangement, leaf texture and branching, and leaf anatomical differences.

Intermediates include specimens similar to T. contorta in the W Himalayas. Examples are characterized

by long linear «distichously arranged leaves that in relative thickness to width (as seen in T-section) are

similar to either the W Himalayan T. contorta (1.5-2.0 mm wide, « 0.5 mm thick, e.g. Biol. Inst. Dubrovnik

from Bosnia, Barabas from Romania, Davis 13667 from Turkey), or to the E Asian T. biternata Spjut (2.0-2.5

mm wide, 0.25-0.33 mm thick, e.g., Petrak from Czech Republic, Moniz from Madeira, Handel-Mazzetti from

Greece, Anderson 42 from Bulgaria, Davis & Hedge 32208 from Turkey). The W Himalayan T. contorta is dis-

tinguished from most Euro-Mediterranean T. baccata by the presence of dark red, or sometimes yellowish,

parenchyma cells (idioblasts) in the leaf spongy mesophyll (in herbarium specimens); however, European

specimens occasionally have idioblasts in the leaf mesophyll. In fresh specimens, the intermediates would

probably be difficult to distinguish. Of further significance is that the abaxial leaf midrib of T. contorta is

always papillose (Kvacek 1984), whereas in T. baccata, the abaxial midrib varies from smooth to papillose.

I suggest that ancestral T. contorta entered the Himalayas from the north during the Miocene uplift

(Krishnan 1974), or earlier (Najman & Garzanti 2000), before it arrived in Europe (Frederiksen 1995). Its

leaves would likely have lost stomata in adapting to the rising Himalayas where environmental selection

would also likely favor the development of papillae on the abaxial leaf midrib (von Frimmel 1911).

As glaciers advanced during the early Pleistocene (2.5-1 mya), the cooling temperatures may have

led ancestral T. contorta to also retreat into Europe where it then allegedly hybridized with other species of

Taxus that may have flourished in a subtropical evergreen laurel-conifer forest (Axelrod 1975; Klaus 1989;

Kvacek 1984; Mai 1989; Palamarev 1989), but may have found refuge in ravines and coastal areas. Similar

patterns of evolution have been suggested for European species of Abies (Fady et al. 1992). The significance

of numerous European refugia for conifers was suggested by Fady-Welterlen (2005) to account for their “sig-

»&« ” LG

nificantly higher” “within species diversity” “than that of other conifer species worldwide.” In this regard, it

is interesting to note that the association of Taxus with “ancient forests" on limestone cliffs in Iowa and in

Europe is partly attributed to the topography of the habitat that offers protection from Homo sapiens (Larson

et al. 2000), whereas a severe decline in European Taxus is generally recognized (Heinze 2004).

Taxus engelhardtii Kvacek, described from a late Oligocene deposit in “NW Bohemia,” was associated

with a mixed mesophytic forest with prevailing broad-leaved componets (Kvacek 1984; Kvacek & Walther

1998) that included Laurophyllum (4 spp.), Cercidiphyllum, Liriodendron, Acer, Ostrya, Betula, Craigia and other

genera. Its leaf shape and arrangement is much like T. mairei in Sichuan, Yunnan (Fig. 10) and Guangdong

where similar forest types still occur today. Vegetation in these areas—characterized by a distinct dry

season—includes species of Cercidiphyllum and Liriodendron (Hou 1983) that have since become extinct in

Europe. In Vietnam, the closely related T. celebica occurs in a laurophyll oak forest from 1000-1600 m in

elevation in association with Cinnamomum obtusifolium Roxb. ex Nees, Michelia foveolata Merr. ex Dandy, M.

mediocris Dandy, Castanopsis fissa (Champion ex Benth.) Rehder & E. H. Wilson, Quercus bambusifolia Hance

f4L,D o ID

312 Journal of t h Institute of Texas 1(1)

and others (Dung 1996; Schmid 1974.). Extant yews in Europe, however, bear little resemblance to T. engel-

hardtii; therefore, it may not have contributed to hybrid complexes that are now apparent in Europe. Rather

it may have simply been extirpated from the Euro-Mediterranean Region due to changes in the climate.

However, another European complex appears to have been derived in part from the Irish yew (T. fasti-

giata Lindley), a possible relict of a former subtropical forest. Known initially from two trees in Ireland—that

were transplanted at the time of discovery (~1770; Veitch et al. 1881), it has always been regarded a distinct

yew even though taxonomists have included it under T. baccata (Loudon 1844; den Ouden & Boom 1965).

Its linear dark green leaves that are spirally arranged in whorls seem less evolved. Moreover, such radial

phyllotaxy is infrequent among yews—occurring at disjunct locations in the Old World, particularly coastal

regions—in Spain, Morocco, British Isles, Honshu, Hokkaido, and Sakhalin. Leaves of the Irish yew in cul-

tivated specimens from widely scattered locations were found to be remarkably similar in lacking papillae

along 6-15 cells across an abaxial marginal zone and on the midrib as well; these specimens, which are

not included in the Appendix, are from Australia (Boorman, New South Wales, A), North America (Spjut

s.n. Oregon, California, Maryland, wba), and Europe (Baker, Yorkshire, BM; Stewart Hort., Florence Court,

Ireland, K; Baenitz, Lusitania, S). Perhaps European yews during the Tertiary were more like those now

seen in E Asia but have since acquired more papillae on their leaves through introgression with ancestral T.

contorta, the alleged replacement species. Many yew specimens with dark metallic green foliage from Great

Britain appear intermediate between T. contorta and T. fastigiata; examples are the “Dovaston yew” (T. baccata

var. dovastoniana) and the English yew (in England, T. recurvata).

Evolution within the European T. canadensis complex is also evident as seen in leaves of one specimen

from the former N Yugoslavia (Slovenia, Fig. 5) by the relatively fewer (4-7) stomata rows per band and

inflated epidermal cells. Related plants in Madeira, southern France, Norway, and Sweden have more stomata

(5-9 rows/band) and less inflated epidermal cells. Characteristics of the Slovenian yew (obtuse leaf apex,

4 stomata rows/band, wedge-shaped epidermal cells) are evident in a fossil leaf from a Pliocene deposit in

Bohemia, Czech Republic (“Taxus sp. 2,” Kvacek 1984).

The increase in cell size and loss of stomata in leaves of the Slovenian yew may reflect adaptation to

changes in a climate from a warm temperate humid type with uniform distribution in rainfall towards a

climate with more pronounced warmer and drier seasons. The Yugoslavia region is also one of 33 sites in

the Euro-Mediterranean region with *Paleomediterranean" woody taxa known from Oligocene, Miocene,

and Pliocene deposits (Palamarev 1989). Thus, the Slovenian yew may be a relict of a former Mediterranean

montane flora that included the conifer genera Pinus, Juniperus, Tetraclinus, Abies, Cedrus, Cupressus, and Picea

(Palamarev 1989); some of these are reported with this Pliocene yew (Kvacek 1984).

I also distinguish T. mairei from T. sumatrana by the relatively short inflated epidermal cells on the

abaxial midrib (Appendix; Spjut in adnot. and on illustrations of packets, A, GH, Jun 1996; Spjut 1998b,

2007). The occurrence of this species in China corresponds mostly to the “broad-leaved evergreen forests of

the subtropical zone” of Hou (1983) with a climate marked by “distinct dry seasons’—“on mountains below

1100 m in the eastern humid subtropics, or on mountains between 1500 and 3000 m in the western subtrop-

ics of the Yunnan Highland” (Hou 1983). Similarly, T. brevifolia, a species confined to the North American

Mediterranean climate, has wider and taller marginal epidermal cells and fewer stomata (Fig 1B), compared

to its putative ancestor, T. globosa (Spjut 1998a, 1998b) that has evidently survived in the montane cloud

forests of Mexico and Central America. The evolution towards larger epidermal leaf cells has also been noted

between fossils and living species of Amentotaxus (Ferguson 1978).

The variation in leaf anatomical data for Taxus in SW China is also related to the convergence of differ-

ent floras in that region (Bartholomew 1999; X-w. Li & J. Li 1997; Zhengyi & Sugong 1998). These include

the “Turkmenian” in W Himalaya, Tibetan or “Indo-Chinese”, Malayan, and Sino-Japanese (Mani 1974; Rao

1974; Rau 1974). In the W Himalayas Taxus is represented by the neoendemic T. contorta, usually with 7-8

stomata rows per band, and in the eastern region by the paleoendemic Wallichiana Group with 15-18 (21)

stomata rows. The relatively lower numbers for stomata rows in T. wallichiana (11—15) and higher numbers

for T. contorta (9—11)—where these taxa overlap in their distribution—is undoubtedly due to hybridization

Spjut, Phytogeographical analysis of Taxus 313

and introgression (Fig. 2C). Further evidence for hybridization involving T. contorta is seen in the wider

marginal region of cells along the abaxial surface of leaves (MC, Appendix) and the wider angle of leaf

divergence from twigs in plants from Nepal and Bhutan (Spjut 2006).

The wide range in leaf anatomical traits for yews of SW China (Fig. 2C, 3B) is also a product of a long

evolutionary history of tropical and temperate vegetation types with possibly less extinction of taxa during

glacial climates (Hsü 1983), in comparison to greater glacial devastation to the floras of North America and

Europe. The oscillating wet and dry periods during the Pleistocene may have led to many new combinations

in Taxus between anatomical and gross morphological features that were once distinctly correlated with

eco-geographic differences prior to the Pleistocene. Introgression of character traits has been correlated with

data on the advance and retreat of glaciers for other conifers—such as between Picea rubens Sargent and P.

mariana (Mill.) B.S.P. (Bobola et al., 1996), between Picea glauca (Moench) Voss and P. engelmannii Parry ex

Engelm. (Wilkinson et al., 1971), among species of Pinus (Axelrod, 1986), and among species of Abies (Fady

et al. 1992).

The slow evolutionary rate that I have suggested for yew is perhaps not all that surprising in view of its

ability to survive almost indefinitely. Individual yew trunks can live several thousand years or more (Loudon

1844; Larson et al. 2000; Thomas & Polwart 2003), and when they fall, the plant still survives by adventi-

tious shoots, or by layering (Hageneder 2007; Loudon 1844); thus, it may continue to survive until perhaps

a change in climate forces it to either adapt or perish. Prager et al. (1976) calculated a rate of change in the

amino acid sequence for Pinaceae to occur once in every 7.5 my. In Taxus this may be longer. By extrapolation

from data in Figs. 2, and from paleoclimatic changes earlier discussed, one may hypothesize that one row of

stomata may become lost permanently in the Taxus leaf as it adapts to slight changes in climate over a period

of 10 my during which time it may also spread a distance of some 3000 km (at the rate of 300 km/my).

APPENDIX

Data for all herbarium specimens studied according to continental and political regions and taxonomy.

SR MC SR MC

NORTH AMERICA U.S.A. Florida

m Mohr (PH). Near Bristol 8 6

Wallichiana Group Croom1833. (K: type).Near Aspalaga 5-7 7

Taxus globosa var. globosa Ex Canby Herb. (PH). Rock Bluff 7 5

EL SALVADOR Blanton 7050 (PH). Rock bluff TS

Tucker 1073 (US). 2670 m 10 5 Wherry (PH). Rock bluff 7 9

HONDURAS Phyton. Florida. 7 8

Armour & Chable 6083 (US). Cerro Sta.

Barbara, 2750 m M ^ 4 Taxus brevifolia

MEXICO California

Phyton Oaxaca 11 4 Hansen 1682 (US). Sequoia gigantea Region 8/9 4

Phyton Oaxaca 10 4 Lemmon 1874 (US). Yosemite 7 2

Meisner (K). Veracruz to Orizaba 10 3 Lemmon 1874 (US). Yosemite 6/7 2

SR MC Sudworth 1899. (US). Stanislaus Forest 7 4

Sharp 52112 (GH). Tamaulipas: El Cielo to Bolander 186 (US). Forest Hills, Devil Canyon 6 -

Ojo de los Indios 10 B Leeberg 5054 (US). Lovelock, 3500 ft. 7 2

Pringle (US). Trinidad Iron Works 9 4 Stokes (US). San Mateo Co. 6 2

Harteg 438 (BM). Hildalgo: Real Monte 9 4 Heller 5941 (U5). Lake Co. DS

Ehrenberg 1837 (K: type). Hidalgo: Real Clark (US). Mendocino Co. 6 5

onte, C. Nabajas 9 3 Yager & Bozovsky (wba). Del Norte Co.:

Taxus globosa var. floridana Oregon Mt. Rd, 200 m, 3 trees:

Hernandez 01459 (BM). Veracruz 9 6 (1) HOR branch 2

Meyer & Rogers 2746 (BM). Nuevo (1) middle branch > z

Leon/Tamaulipas: 1690 m 7 2 (1) bottom branch ? i

Mueller 1337 (PH). Nuevo Leon: Sierra Madre (2) lower branch TA

O 7 2 (2) middle branch 6-7 -

Mueller 1337 (BM) ZEE (2) top branch NE

(3) lower branch 7-9 -

314

Rose 55089 (US). Trinity Co., Buckhorn

Summit, 2600 ft

Spjut 12307 (wba). Salmon Mts., 2000 m

Spjut 12307 (wba). Salmon Mts., 1500 m

Spjut 10171 (wba). Marble Mts., 2000 m

Dudley (US). Salmon Mts., Foxtail Ridge

Benson 2228 (US). Shasta Co., Hatchet Creek

Grant 1281 (US). Shasta Co., Dunsmiur

Oregon

Fisher (US). Portland

Collector, no.? (US). Jackson Co.: Wimer

Walpole 153 (US). Jackson Co.: Ashland

Coville (US). Imnaha Natl. For., Billy Meadows

USFS (wba). Josephine Co.: above Taylor

Creek, Minnow Creek Rd., 650 m

4 specimens from nearby sites:

(1)

(2)

(3)

(4)

Lankford (wba). Clackamas Co., 1060 m

Lankford (wba). Clackamas Co., 930 m

Lyall 1860 (K). Columbia River

Nuttall (K: type). Columbia River

Spjut 12301 (wba). E Cascades E of Portland

Beattie 5046 (US). Josephine Co.: 2270 ft

Cusick 3405 (US). Eastern Oregon

Washington

Spjut 12302 (wba). E Cascades E of Seattle

Horner (US). Blue Mts.

Meyer 1589 (US). Thurston Co., Mud Bay

Grant s. n. (US). Cascade Mt.

Cantwell (US). Orcas Island

Fosberg (US) King. Co., Stevens Pass

Idaho

Cronquist 6187 (US). 20 mi W of Riggins,

French Creek

Shields (wba). Idaho Co.: Allison Creek,

3400 ft.

Cochrane (wba). Idaho Co.: Nez Perce Natl.

For 5600 ft., 2 plants

Montana

Donner (wba) Flathead Natl. For., nr.

Columbia Falls, 3800 ft.

P.C. Standley 18251 (US). Glacier Natl. Park,

1400-19850 m

Thomas 11031 (US). Lake Co.: 8 mi from

Polson, 3850 ft.

Steven Wirt 100 (MRC, wba). Flathead

Co.: shrubs

Steven Wirt 100 (MRC, wba)

CANADA British Columbia

ON Ul

hh ANAANADHD BON

Orv OV

ANN

Gc» O TA

fal, Dat o ID L

Journal of

MacMillan (PH). Selkirk, 4300 ft.

Calder & Saville 9982 (US). SE of Nakusp

Macoun (US). Rocky Mts., Silver City

Macoun 2340 (US). Vancouver Is., Victoria

Baccata Group

Taxus canadensis

U.S.A.

Herb. C. W. Minott (US). Ma: Amherst

Bovin & Blain 753 (PH). Me: Cumberland

Co., 425m

True 164 (PH). Me: Ovis Island, Long Cone

Gilbert 831 (PH). Ky: Carter Co., Cascade

Caverns

Allard 12060 (US). Wv: 900-1200 m

Women's College of Baltimore (US)

Palmer & King 205 (US). Va

Taylor 424 (US). Pa: Bucks Co., Kintersville

Eames 3432 (US). Ny: Coy Glen, Ithaca

Spjut (wba) Ny: Ithaca

Spjut 11778 (wba) Nh:White Mts. Natl. For.,

Wildriver, 300 m

Stevenson (US). Vt: Willoughby Lake

Weatherby 5977 (US). Ct: Boston Hollow

Sheldon (US). Mn: Towers St. Laus

Fellows 5686 (US). Me: Rockport

Spjut 12179 (wba). Ohio: Secrest Arboretum

Shreeve 1971 (US). Md: Garrett Co., Bailing

Spring

Travis 119 (PH). Me: Cumberland Co.

CANADA: Ontario

McDonald 223 (US). Ontario: Sagastaweeki Is.

Rouleau 2700 (US). Humber Dist., Twin Lakes

Quebec

Tae hé & Lepage 332 (PH). Dartmouth River

Asselin 7212. (US). St.-Charles

Pennell 16734 (PH). La Belle Co.

Bartram & Long 649 (PH). Rimousk Co.

Fernald et al. 2404 (US). Gaspé Co,

Mt. St. Pierre

Louis-Alphonse 3547 (US). Baie Missisquo

Lucien 743 (PH). Laurentides, Bellerive

Louis-Maire 686308 (PH). Mé gantic

Bovin 1268 (US). St.- Catherine

Chas Mohr (US). Montreal

New Brunswick

Malte & Watson (S: C2153).

Allen 2528 (PH). St. John

Nova Scotia

Gorham 45139 (US). Halifax Co.: near Halifax,

St. Margaret's Bay

Bean et al. 19634 (PH). Yarmouth Co.

19015 (S: 2155).

(S: 2156). Victoria Co.

Pease & Long 19633 (PH). Cumberland Co.

Bissell et al. 19632 (PH). Digby Co.

7/7

7/8

titute of Texas 1(1)

Spjut, Phytogeographical analysis of Taxus

Newfoundland

Fernald et al. 6738 (PH). Prince Edward Is. 8/9

Palmer 1300 (US). Bay Is. 8/9

Palmer 1327 (US). Hermitage Bay, Balena 27

Buochan (S: C2130). prp

Rouleau 6545 (US): St. Barbe Distr.,

E BluePond 7/7

Banks1766 (BM). Croque WE

Fernald et al. 26201 (PH). NW Coast 6/7

Robinson & Shrenk (US). St. John's 6/7

Rouleau 5533 (US). Gander River 5-7

Fernald & Wiegand 4414 (PH). Valley

of Exploits River 6/6

Fernald & Long 27305 (PH). Pistolet Bay 6/6

Wiegland & Gilbert 27304 (PH). Highlands

of St. John 5/5

EURO-MEDITERRANEAN

Baccata Group

Specimens ranked first by number of stomata rows then

by marginal cells for

Baccata Alliance, 1. canadensis noted separately under

each country.

ALGERIA

Swingle (NA). Chria near Blida 12

Reichenbach (K). Atlas, Blida 9

Gamble (K). Atlas des Demia 8

Olaptin (S: C2070), Atlas, Blida. 7/8

Davis 52628 (BM). Cedrus forest,

1900-1950 m 9

Univ. Algeria Apr 1912 (NA).Atlas, Blida 9

MOROCCO

Trethewy 85 (K), pendula. Ifrane 1400 m 10

Lewalle 8670 (BM). Ifrane 1400 m 10

Lewalle 8670 (BM). 8

Lewalle 9670 (BM). Ifrane 1400 m 9

Davis 49209 (BM). Ifrane 1700 m 9

Lynes (BM). Mid Atlas, Azrou, 5700 ft 9

Haout 938 (BM) 9

Davis 55121 (BM). Ifrane, Cascada,

1580 m 9

Taxus canadensis

Font Quer 1928 (BM).Kaloa to Tauka, 1500m 8

PORTUGAL

Goncalves 4625 (BM). Azores 12

Yoller 61 (BM). Sierra Jerez ae 12

Goncalves 4491 (BM). Azo 10

ex Herb. Moniz (K). en 10

Cyrén (S: C-2058). Ser. Estrela 10

Fontee et al. (S: C2047). Ser. Estrela,

1400m 10

Fontee et al. (S: C2047-2). Ser. Estrela, 9

Meaden 1865 (K). Madeira 9

Moller (BM). Serra Gerez: Vidoal 8

Taxus canadensis

Lowe 5/0 (BM, bottom specimen). Madeira 11

AAU

7-14

Lowe with 570 (BM, top specimen)

Lowe ex Barby (US). Madeira

SPAIN

H. Elias 4353 (BM). Burgos: Ser. Obarenes,

1000 m

Modesto Laza Palacios (K). Ser. Tejeda €:

Almijara, Malacitana Prov.

no data 1878 (US). Laguna

Sennen 7087 (BM). Barcelona

Roivainen (S: C2075). Guipüzcoa, 900 m

Sandwith 4452 (BM), Huesca: Ser. Guara

Heywood & Davis 490 (BM). Ser. Cazoria:

Yedra

Rodriquez (K). Serrania buenia

Bianor-Maire (BM). Baleares, 1600 m

Fosberg 41055 (US). Jura Mts.

Endress Aug 1831 (K). Pyrenees

Endress Aug 1831 (S). Pyrenees

ex Herb Comby (PH)

Tidestrom 12814 (US)

Massonnet (K). Pyrenees

Herb. Hook., 1867 (K). Pyrenees

Herb. Churchillanum (K). Corsica

C. Lagerheim & G. Sjogren Jul 1844 (K).

Batsmanshus Paroeciae Elfkarl by

Rosalagiae abundans

Taxus canadensis

Herb. Gombault (S). Sainte Baume

UNITED KINGDOM

Gamble 19866 (K). Berkshire Dist

Bowden & Hillman 433 (BM, globose

epidermal cells). Nottinghamshire

Gamble 28894 (K). Weltham woods

[England SW

Albarnes 26 (K). Dorsey: churchyard

[England SW], 350 ft

Michaelstone (K). British Isles, England

Turrill 4903 (K). Leicestershire, Charwood

Forest

Ap. ys 77-390 (K). Kent

Turrill (K). Surrey, Box Hill [England SE]

Fraser (K). Surrey, Box H ill

Fraser (K). Surrey, Chalk Pits

Bean & Hill (K). Scotland: Neopath Castle,

Tweedsdale near Peebles

Ball 1838 (US). Surrey, Jumper Hill

Ex. Herb. Bidwell (BM), dovastonianum.

Westfelton

Ex. Herb. Gordon (K), dovatsonianum.

Westfelton.

Jackson (BM). Westfelton (Dovaston yew,

original)

Lewis 721 (BM). Monmouthshire

Jarrell (K). Kent: Shorehane

Bennett & Croydon 713 (US). Riddlesdown

E E IS

TARA AT

316

Hooker (PH). Kent

Barron (K). Kent, Buckland

? (K). Kent, "var. washingtonianum"

Valpy (K). Elsing, Norfolk

Boswell (BM). Shropshire, Lyth Hill

Brubaker 1960 (PH). Druids Grove

Roper 1525 (K). Bristol, Birdhamdown

Carruthers (K). Ireland: Pollawaddy

Aug. 1874 (BM). Perth Co.? [filed under

Portugal)

Turrill (K). Yorkshire: 3 mi. from Richmond

Hubbard (K). Sussex: Bury Hill

Jackson (BM). Highclere, Saddam

Turrill (K). Scotland: Loch Lomond

XO. KONO? KOPF XO XO! NO XO. Cn»

00 CO 00 00 WO

Taxus canadensis var. adpressa

Summerhayes 2581 (K). E Kent 8

SWITZERLAND

Kellermann (US: 518500) 10

Herb. A. Gray (K) 9

Fr. Castella (US). Le Pissot sur alboue, 1000 m 7

GERMANY

Reichenbach fil. (PH). Dresden 11

Martius 1831 (PH). Bavaria 9-11

Martius 1831 (K). 9/10

Milchbuder (K). Bavaria 10

Petzi 1444 (K).Bavaria 8/9

Reichenbach, ex Short Herb. (PH). Dresden 7-8

Keller (PH). Darmstadt 7

Martius 1831 (K). Bavaria Alps 7

POLAND

Baenitz (US). Silesia: Proskau, 180 m,

"f. dovastonii" 10

Baenitz (US) epacroides. Silesia: Breslau,

120 m, 'v. recurvata" 9

Baenitz (US). Silesia: Breslau, 120 m,

"f. epacroides" 10

Baenitz (US), epacroides. Silesiaca: Breslau,

Scheitniger Park 120 m 8

Baenitz (US). Silesia: Breslau, 120 m, "f. erecta" 9

CZECH REPUBLIC

Jirasek & Suza (K). Moravia Centr.: 4-450m . 10

Jirasek & Suza (US). Moravia Centr.: 4-450 m 10

Petrakm, Fl. Boeh. & Morav. exsic. 99 (BM) 8

AUSTRIA

Ex Pickler Herb. 1895 (US). 11

Ex Herb. Pichler (US: 347988, lower

specimen). Tirol

Ex Herb. Pichler (US: 347988).Tirol

Ex Shulte Herb. 1863 (K)

Hayer (S: C2034). Salzburg

Gander 1869 (K). Tirol

Gander 1869 (US: 157025). Tirol

Keck (US)

NOOO M OO)

¡(DAD c»

Taxus canadensis

A. Hayek & F. Hayek (BM). pos superior:

Kulmburg 9

hp HRA HD HAW

RA KO hh

Uo 00-4...

Ltn Dat L ID AKI +

Journal of

HUNGARY

Herb. Láng (PH)

Schónach 3084 (S: G2061A)

Schönach 3084, Austr-Hungar. (US). 445 m

Wagriesh (US: 451917). Vorarlbergia, 445 m

Wagriesh (US: 481917). Dolüa

Boros (BM). Comit. Boraod. Ohassa, 550 m

Lémke (S: C2042). Bakony: MikIlCspalhazy

Schónach, Aust.-Hungar. (US: 966290),

epacroides 445 m

Ex Herb. Mus. Nat. Hungar. (S: C2041), Bakony

Schónach 3084, (BM). 445 m

Taxus canadensis

Schónach 3084 (S: C2061 5 Specimen).

ROMANIA

Topa, Bot. Mus. Exsic. (US). Bucovina:

00 m

Topa, Bot. Mus. Exsic. (S: C2024). Bucovina,

Topa, Bot. Mus. Exsic. (US). Bucovina, 400 m 9

Anderson 102 (K). Balkan Exped., Cajan Pass

Mititleu & Barabas (BM). Bucovina:

Darmanesti, 500 m

BULGARIA

Kotschy (P)

Anderson 42 (K). Sofia: Vitorha

ITALY

Herb. Hook. 1814 (KJ. Montagnes

Levier (BM). Florentino

Lenander 1933 (S: C2008). Lago di Garda,

Riva, Sydtyrolen

McDonald: 1-37 (US). Cult.

McDonald: 1-37 (PH). Cult.

Solla (US: 280040).

Baroncini 16 Sep 1893 (US)8/9

Fireuze (BH). Cult

Martelli (PH). "Iter Sardoum", Limabara

[Sardinia]

[YUGOSLAVIA]

Biol. Inst. Dubrovnik 37 (NA). Bosnia:

Mt. Trebevic near Sarajevo, 1450 m

Kosarim (S: C2065), Macedonia, Petiska

Baldacci 169 (K). Albania

Rohleana 1908 (BM: 17197). Montenegro

Curic 1897 (K). Bosnia

Biol. Inst. Dubrovnik 136 (NA). Bosnia.

Mt. Plasma near Jablanica, 1500 m

Woloszczak (K), Tatra

Taxus canadensis

Berglund (S: C-2066; var. adpressa).

Slovenien: Bled, berget Straza

GREECE

Georginda & Izanoudakis 631 (BM). Artis:

Mt. Tzoumarka

HGT 884 (K). Hills N of Xant Is

Spjut, Phytogeographical analysis of Taxus

Heldreich (S: 2023). Oeta, 4500-6000 ft. 9

Greola (PH). "Mts. of Tyrah" 9

Guiol 2260 (BM). Mt. Olympus 7/8

Handel Mazzetti (K). Mt. Olympus,

750-850 m 8-9

TURKEY

Balonsa (BM). Taurus 9

Balonsa (P). Taurus 8

Davis & Hedge 32208 (BM). Coruh, Savval

Tepe above Murgul, 1400 m 7

Sintensis 5118 1892 (P) Paphalogonia:

Wilajet Kastanbuli 9

Sintensis 5118 (K) 7

Murray 936 (NA). Between Molla Veyis and

Meyden, S of Ardesen, 750 m 8

Davis 13667 (K). VA. Jenigli (Caira [Caria ?])

(Denizli, Boz Da, Davis 13447),

5000-5500 ft 8

IRAN

Koelz 16208 (US, distinct for obconical to

4-|obate seeds). Gozlu, Mazandaran 8-9

SYRIA

Haradjían (K). Duldul: Mt. Amanos,

5000-7000 ft 10

Haradjían 2341 (S). Duldul: Mt. Amanos,

5000-7000 ft 8

Gesbeldagh (BM) 9

Gesbeldagh (US) 10

Haradjfan 3865 (S). Duldul: Mt. Amanos,

1500-2000 m 10

Delbés (P). 1000 m 9

RUSSIAN REGION

Estonia

Taxus canadensis

Lundstróm 742 (S). Eosl near Karriland 9

Lundstróm 579 (S). Osel, Sworbe 9

Lundstróm 562 (S). Osel, Sworbe 7

ranscaucasia (Caucasus Mts.)

Elias et al. 5615 (NA) Ukraine Crimea,

Princeps Kascelsky, ex. Herb. Hort. Imper.

Petro. (S). Caucayas Hosharia

Busch (K). Caucasus, Terek

Woronowa (S: C-2027). W Transcaucasia:

Suchum, Petkir (Fl. Madshara) 13

Dmitrieva (NA). SW Georgia, Black Sea near

NE Turkey, Adzharia, 900 m 11

Inst. Bot. Acad. Sci. Armenia (US). 22 Mar 1946 10

Szovich 610 (S: C2072). Armenia:

Tschunakuchi 10

Szovich 610 (P). Armenia: Tschunakuchi

Ex. Herb. Inst.. Bot. Ac. Sc. URSS (US: 2560106).

[Transcaucasia] 9

In Russian 475 (P). Azerbaijan [Kura Mts.] 9

Herb. Komaróv (US: 1862552). Armenia 7/8

Prilipko (K). Transcaucasia 9

Goghika (NA). Caucasas: Azerbaijan, Chanlar,

1800m 10

7-10

= 00 #

Brzhezitzky & Kasumov H196 (US).

Azerbaijan 10

Taxus canadensis

Kousnetzoff 89 (US: 254512). [Russian

Federation] Kuban 10

NORWAY

Gamble 28933 (K). West Dalen 10

Taxus canadensis

Anderson (US: 1091452). Kolsas 5

DENMARK

Herb. Joh. Lange 1866 (K) 10/11

Herb. Joh. Lange 1866 (K: Right specimen) 8

SWEDEN

Thedenius (PH), Góteborg 9-10

Thedenius (US) 8

Steinvall 1872 (K). Södermanland 10

Henriksson (K). Dalsland:Gunnarans 8/9

Holmgren (US: 1276222). Blekinge 8

Lindberg 419 (K). Ekeró 11

Herb. Bot. Berjianus (S: C2177),

"washingtonianum 10

Bjornstrorn, ex Mus. Stockholm (US).

Podermanlane? [Södermanland] 10/11

Taxus canadensis

Asplund (US: long-needled specimen).

Sódermanland 9

Asplund (US: short-needled specimen). 8

FINLAND

Florstróm (BM). Alandia: Lemland 11712

Florstróm 1909 (K 10

Vidlund, Helsinki Exsic. (K). Lemland 8

EAST TEMPERATE ASIA

Cuspidata Alliance

RUSSIAN FEDERATION

Taxus biternata

B. Cerereu (A). Far East Region: Pryanyk

For. Div. 11

Taxus umbraculifera Complex

Kypehisnova (A). Primorye Prov.

Kypehinova (A) Primorye Prov., Bay of Peter,

First Sea Reserve, ls. Stenin

Dvorakovskia & Bokina (A). Sakhalin Is. 11

Palczevsky 3601 (K). Primorye Prov.: vicinity

of Vladivostok

Palczevsky 3601 (US) 9

Palczevsky 3601 (A) 8

Lyubarsky 2 (A). South, Sikhote-Alin, foothills,

Mt. Hezalaza, River Beryozovoy

Esus 203 (K). Sakhalin Is.

MANCHURIA

Taxus biternata

Ex herb. hort. bot. Petro. 1860, Maximowicz (P).

Mandshuria SE 10

318

Skvortzov 20 Sep 1931 (A). N Manchuria,

Sochintzest, forest, tree

In Russian No. 75 (P). Jilin (Kirin)

C. H. Chen 539 (A). Jilin (Kirin)

Maack 1855. (GH: top specimen)

Purdom (GH). N China [Shaanxi: Tai-pei-shan

fide Rehder & Wilson in Sargent 1914]

T. umbraculifera Complex

Palczevski [Komaróv] 88 (K). Manchuria:

Rossica

Palczevski [Komaróv] 88 (BM)

Maack 1855. (P)

Ex herb. hort. bot Petro.1860 (Bunge),

Maximowicz (P). Mandshuria SE

Ex herb. hort. bot. Petro. 1860, Maximowicz (S)

Mandshuria SE

Ex herb. hort. l.c. (P)

Ex herb. |.c.(GH)

G. Fenzel (A). Schenhsi merid., Taipei-schan

KOREA

Taxus biternata

Wilson 10519 (US). Kyongsan, Nemon-rei

Wilson 10519 (A

Wilson 9484 (A), Hallai-san down to

Mushroom House, bush

Wilson 9097 (A), Shinkabachin Heizanchien

to Ehoshin, Kankyo-N Heian divide, tree

Wilson 8685 (A). N. Heian Prov.: O.G.M. Co.

Mines, Pukchin

Wilson 8685 (US)

Wilson 8685 (K)

Komaróv 1897 (GH). Pen-nian Prov.

Wilson 10688 (US). Kyongsan, Nemon-rei

Wilson 10688 (A)

Wilson 10688 (K)

xus umbraculifera a

Tax

Wilson 8538 (US). Oagelet Island, 0-900 m

bush

Wilson 8538 (A)

Wilson 9332 (A). Kyongsan Prov.: Nemon-rei,

tree

Faurie 1512 (BM)

JAPAN

Taxus cuspidata

Ex Herb. Zuccarini (M: type, T. cuspidata)

Japan

Ex Herb. Zuccarini (K: type, T. cuspidata)

Jack (A). Hokkaido: Sapporo

Jack (GH). Hokkaido: Sapporo

Taxus biternata

Makino 43775 (S), Tokyo Pref.: Oizuni,

Nepymawku

Mujabe1884 (A). Hokkaido: Ishikasi

9/10

NO 00 00

Journal of the Botanical R h Institute of Texas 1(1)

SR MC

Makino 43779 (S). Chiba Pref: Mt. Kiyosumi 13 6-8

Uno 2611 (A). Nagano-ken, Okmachi 18 9

Mochizuki (A). Mt. Nantai, Lake Chuzenji 12 10

Wilson 7544 (A). Kai prov., Nakaihinsen,

1200 m, hedge 12 9-12

Sargent (A). Hokkaido, Cos! Mines, Utishini — 11 10

Arimoto (A). Sapporo, Yezo jj 15

Nitzelius (S: C2111), Góteborg cult., from

Hokkaido: Kamikawa, Yamabe 11 18

Wilson 7778 (A). Hondo, Sernja Prov.:

Yamanaka on Fuji-san, tree 11 -

Wilson 7778 (K) 9 14

Hatusima 13858 (A). Kagoshima Pref.,

Mt. Takahuma, tree 10 -

Ex herb. horti bot. Petropolitani,

Maximowicz. 1862 (P), Yokohama 9 6

Taxus umbraculifera Complex

Taxus caespitosa var. caespitosa

Mizushima 1985 (A). Honshu: Mt. Hakkoda — 11 13-18

Wilson 7133 (A), Honshu: Hakkoda, 1000-

2000 m, shrub 1i 8

Ex herb. horti bot. Petropolitani, Maximowic.

1862 (GH), Yokohama 10 10

Ex herb. horti bot. Petropolitani, Maximowic.

1862 (P). Yokohama 10 10

Wilson (A). Honshu: Mt. Daisen (topotype),

2000 m, shrub 10-11 9

Makino 43792 (S). Honshu: Mt. Daisen

(topotype) 10/11-10

Taxus caespitosa var. latifolia

Shimotsake 446 1888 (US, anatomy

like 7. mairei). 15 16

Faurie 6345 3 isolectotype) 14-15 12

Faurie 6345 (P, lectotype) 11 8

Bataw, Herb. Lugd (P). Honshu: Shimane 14-15 -

Faurie 5114 (P) 15 10

Faurie 5114 (P) 12 6

Makino 43769 (S), Honshu: Akita Pref. 12 7-8

Tomitar ex. Makino 43780 (S), Honshu:

Kanagawa, Mt. Imaizumi id 8

Wilson 7265 (A). Hokkaido: Shiribeshi Prov.,

1300-2000 m 11-12 10

Mizushima 401 (A). Honshu: Prov. Kozuke,

- 9-10 10

Mizushima 1989 (A). Honshu: Mt. Hakkoda | 9-10 7

K. Muijabe 17 Sep 1910 (A). Hida, Takayama 9 14

Shimotsake (P). Honshu: Nikko 8 8

Ex herb. horti bot. Petropolitani, Maximowic

1862 (P). Yokohama (with T. biternata) 11 8

Ex herb. horti bot. Petropolitani, Maximowic.

1862 (US). Yokohama (top specimen) 10 7

Ex Herb. Zuccarini, 1842, with ex Herb. Lugd.

Batav. (GH). Japan 8(-11) 12

Taxus umbraculifera

Ex Herb. Lugd. Batav. (P). Japan -14 15

Spjut, Phytogeographical analysis of Taxus

Suzuki 499003 (A), Honshu: Mt. Ooyhama,

Kanagawa-Pr, cult. 14

Faurie Dec 1904 (A), [Hokkaido], cult. and in

forest

Muroi 1969 (A). Honshu: Mt. Fujiwara

Wilson, ex. Sakurai (A), Honshu: Kyaraboken,

cult. "nana" 10

Muroi 5933 (A). Honshu: Mt. Himekami 10-12

Sapporo Agric. College (PH). Hokkaido: Kitami

Prov, Rishiri 11

Sapporo Agric. College1885 (A). Hokkaido:

Niarenai? 9-10

Sapporo Agric. College1878 (A). Hokkaido 11-13

Naito (A), ex. Herb. Kagoshima Univ.

Shimane Pref., Mt. Sentsu-zan. 11

Hatusima 13858 (A). Kagoshima Pref.,

Mt. Takahuma, tree 11

Shiota 4441 (A). Hondo, Mino Prov., hort. 11

Muroi 30 (A), Honshu: Hyogo Pref,

Mt. Hyonosen 10

Taxus umbraculifera var. hicksii

Muroi 5603 (A). Hyogo Pref.: Kumatugi,

Mikata-gun 1

Muroi 5424 (A). Mt. Hatibuse 1

Muroi 5648 (A). Wakasugi 1

Muroi 3593 (A). Iwate-Pref.: Asagishi 1

Muroi 3698 (A). Gifu-Pref.: Takayama 1

Muroi 3715 (A). Nagano Pref.: Kamikochi 1

ASIA: HIMALAYAS

Baccata Group

AFGHANISTAN-INDIA

Taxus contorta var. contorta

Sprague 730 (K). Murree 6

Aitchinson (K). Kurrum Valley, 7500-9000 ft 6

Sinnott et al. 146 (K). Between Gotchbok

and Kubkot Valley, 2750 m 7

Stewart 15343 (US). Murree, 7000 ft 7

ex Herb. Schlagintweit (PH). NW of Srinagar 5

Stewart 7374 (PH). Sonamarg, 10,000 ft 6/7

Mukinji (K). Lada Valley 6

Stewart 8414 (US). Kashmir: Pahlgam S

Stewart 8414 (A). Kashmir: Pahlgam Pj

Stewart 8414 (PH). Kashmir: Pahlgam 7

Stewart 120018 (A). Kashmir: Pahlgam 2600 m 7

Schlagintweit (P). Kashmir: Báltal to Númner 7

Kenyoer & Dugeon (PH). Bureah, 11,000 ft 7

Rau 31770 (A). Garhwal to Lake Hemkund,

3200m

ex Herb. Falconer 1000

Dwali? 8500 ft

ex Herb. Falconer 1000 (P). Kumaon 7

Koelz 10285 (A). Punjab: Kulu, above

Bandrole, 8000 ft 7-8

Schlagintweit 8941 (GH). Kashmir: Sukhi

across Bamsuru and Chaia Pass to

Khdrsali, 9000-15400 ft 7-8

S: C1994). Kumaon,

Rodin 5313 (US). Punjab Province: Rosenhiem,

Murree

Stewart 5931 (A). Kashmir: Pahlgam,

7000-10,000 ft 8

Heybrook 29 (K): Kashmir: Pahlgam 2600m 8

Lace 301 (A). Bashahr, Uri Forest 8

Stewart (PH: 829196). Dharmkat, Dharmsala,

6000 (ft?) 8

Gamble 23507 (K). Jaunsar Dist., 10,000 ft 8

Stewart 10663A (PH). Gulwarg,

7000-10,000 ft 9

Laig Raus (P). Siwalik and Jaunsar Div.,

10,000 ft 9

Pengelly (K). Chumba 10

Bertoloni (BOLO: type, T. orientalis).

Western Sikkim

NEPAL

Baccata Group

Taxus contorta var. contorta

Stainton et al. 7832 (BM). Chingnon, 10,000 ft 7/8

Polunin et al. 1353 (BM). Dhotar, 9600 ft. 9

Polunin et al. 432 (BM). Chankeli Range,

8000 ft 11

Polunin et al. 5050 (BM). W of Jumla, Belas

Gaejigeth, 10000 ft 9

Polunin et al. 1873 (BM). Chatlwe, 9000 ft. 10

Gardner 557 (BM). Shios Khola, 8500 ft 9/10

Stainton et al. 734 (BM). Lete, S of Tukucha,

8000 ft

Ottba et al. 8311066 (BM). Marayandi Khola — 11

Mikage et al. 9550282 (BM). Dhaulagiri Zone,

2405 m 10/11

Stainton et al. 5616 (BM). Chingnon, N of

Tukucha, Gadaki Valley, 10,000 ft 11

Taxus contorta var. mucronata

Dobremez 2106 (BM).

Wraber 514 (BM). Hanangi: Karayundi Valley,

3100m

Sumatrana Group

Taxus sumatrana

Herb. Banerji, 1953, in adnot. T. bounoniana

arr. (A). E Nepal: Khanigaon to Kalanti,

6,000 ft. 12

Wallichiana Group

Taxus wallichiana

Wallich 6054A (M: Original Material). [Nepal] 15

Wallich 6054A (K: Duplicate of Original

Material). [Nepal] 14

Wallich 6054A (K: Duplicate of Original

Material). [Nepal] 12

Wallich 6054A (S: Duplicate of Original

Material). [Nepal

[Wallich] (GH: Duplicate of Original

Material). Napalia. 15

319

320

Beer 25316 (BM). Above Sedua, 9400 ft

Stainton et al. 1398 (BM). Arun Valley,

N of Kutiar, 9000 ft

Stainton et al. 6601 (BM). Eastern: Duon Kosi,

Chaunrikarua, 9500 ft

Stainton et al. 4496 (BM)

Griffith 2006. 9000-10,000 ft

Stainton et al. 5102 (BM)

Tabata et al. 10585 (A). Soluhumbu Dist.:

Lamujo to Chumawa, 2450 m

Tabata et al. 10585 (BM)

Stainton et al. 8296 (BM). Pembrang?,

Williams 458 (BM). 9500 ft

Ohba et al. 8310264 (BM). Thulo Kobar

to Ran Thanti, 2600 m

Ohba et al. 8310264 (BM)

Taxus phytonii

Williams 1014 (BM). 9000 ft

BHUTAN

Baccata Group

Taxus contorta var. mucronata

Ludlow et al. 16035 (A). Eastern, Ha:

27.22' 89.18, 9,000 ft

Ludlow et al. 16035 (BM)

Sumatrana Group

Taxus celebica

Cooper & Bulley 2833 (BM). Rinchu Timakha,

Oft

Wallichiana Group

Taxus wallichiana

Ludlow et al. 18672 (BM). Tunle La. near

Kinga Rasdah, 11,000 ft

Ludlow et al. 18672 (A). Tunle La near Kinga

NI 11,000 ft

ierson & Long 4417 (A). Thimphu Dist.:

summit of Dadon La, 3110 m

Cooper & Bulley 2600 (BM). 7,500 ft

Bartholomew & Boufford 3917 (A). Above

Motithang, W of Thimphu

NE INDIA &TIBET

Sumatrana Group

us celebica

Kingdon Ward 19324 (BM)."Assam" [Tibet]:

Rima, 7000 ft

Clarke 38308 (K). Khasia: Maophlang

Taxus kingstonii

Mann 1885 (K). Khasia Hills, l.c.

Mann 1885 (BM), l.c.

Mann 1885 (P). Khasia Hills, l.c.

Kingdon Ward 18751 (A). Khasi Hills,

Mawphlang, 6000 ft

Hooker 1337 (K). Khasia: 5000 ft

10-11

fal, Dat A ID hi Pr

Journal Ul

Hooker & Thomson 1855 (P) 11

Hooker & Thomson 1855 (P) 10/11

Hooker & Thomson 1855 (P with seed) 13

Simmons 484 (P). Assam: Khasia 13

Taxus sumatrana

Mann 1885 (A). Khasia Hills: Nunghuai,

5000 ft 24

Wallichiana Group

Taxus wallichiana

Wallich (M: Lectotype). Eastern

Biswas 439 (A). E Himalaya 13

Biswas 439 (A). E Himalaya 10-11

Kurz (A). Sikkim: Tongloo 13-15

Raijada 18919 (A). Cult., Dehra Dun, Bot.

Gard. Darjeeling 14

Griffith 5002, ex Herb. Griffith. E Himal. (P) 12

Griffith 5002, ex Herb. E India Co (P) 19

Griffith 5002, ex Herb. Bunge E Himal. (P) 19

Hooker 77 (P). Khasia, 5000-6000 ft 16

Griffith 2(7)606 Assam (P) 15

Kingdon Ward 17271 (A). Sirhoi: 8000 ft 15

Kingdon Ward 17271 (BM). Sirhoi: 8000 ft 15

Vos et al. 148 (NA). West Bengal: Singalila

Range, 8400 ft

C. B. Clarke 436743 (BM). Khasia: 4500 ft,

Vale

of rocks 14

G. Watt 5955 (A). Manipur: Seriphari,

10,000 ft

157

G. Watt 5955 (P). Manipur: Seriphari,

10,000 ft 17

G. Watt 6493 (P). Manipur: Sirohifarar, 7000 ft 16

G. Watt 6208

P). Manipur: Jakpho, 11,000ft 18

Taxus wallichiana var. yunnanensis

Hooker (K). Sikkim: 7000-10,000 ft

Hooker (K). Sikkim: 7000-10,000 ft 14

Kingdon Ward 18990 (BM). Jakpho Range

Clarke 412388 (K). Jakpho, Naja Hill

Kingdon Ward 7755 (K). Barail Range, Naga,

9000-10,000 ft

Kingdon Ward 8090 (K). Assam [Tibet]:

Chiban, Delei Valley, 6000-7000 ft (K) 14

Kingdon Ward 8594 (K). Assam: [Tibet

Delei

Dann

Valley, 9000 ft (K)

Taxus phytonii

Ludlow & Sherriff 3719 (BM). Pachaksihri,

Laluma, 94?15, 27?45, 7000 ft

MYANMAR (Burma)

Sumatrana Group (Taxus kingstonii)

Oliver 4 Sep 1894 (K). Bernardmyo, Ruby

Mines

Oliver (K) 14 May 1892, 5600 ft 12

A un e

ARA RRA ROSA

Spjut, Phytogeographical analysis of Taxus

Wallichiana Group

Taxus obscura (Chinensis Subgroup)

K) Ruby Mines, 6500 ft 13-15

—

Oliver

Taxus suffnessii (Wallichiana Subgroup)

Kingdon Ward 21901 (A). West Cental Esakan,

12-

Kingdon Ward 20901 (BM). W Central 18

Kingdon Ward 20902 (A: holotype) North

Triangle, 9000-10,000 ft 12-13

Kingdon Ward 20902 (BM, isotype) 16

Kingdon Ward 13003 (BM). 27°45'N, 97*50'E,

9-10,000 ft

Hla & Koko (K) Myintkyina: Sumprabum,

8600 ft 15-16

Taxus wallichiana var. wallichiana

Kingdon Ward 9214 (BM) Northern, Adung

Valley, 97°30-98°30', 27°30-28°30', 6000 ft. 17

Kingdon Ward 9214 (A) 16

Kingdon Ward 9375 (A). N Adung Valley,

97—98?30' 27-28°30' 7000- 8000 ft

Taxus wallichiana var. yunnanensis

Kingdon Ward 22819 (BM). Mt. Viatoria,

9000-10,000 ft 16

Kernode 17205 (K). Myintkyina: Laikan-

Fenshuiling Rd, 8000 ft 12

THAILAND

Chinensis Subgroup

Taxus obscura

Lobb 461 (BM) Malaya

Sumatrana Group (T. sumatrana)

Kerr 20146 (K). Kao Kuading, 1200 m is

Kerr 20146 (BM) 12

VIETNAM

Sumatrana Group

axus celebica

Evrard 305 (P). Dalat: ravin buisé an chalet

Rimaud 9/10

Evrard 1438 (P). Lam Dóng 12

Schmind 1960 (P). Dak Tria- Manline, 1400 m 8

Schmind 1960 (P). Dalat: Dau Lamghi 9

Schmind (P). Dalat: Dak Tria, 1610 m ld

Van Cuong 12891960 (P). Dalat: Manline,

1610m 12

Wallichiana Group

Taxus chinensis

Hiép & Chan 405 (P). Hoa Binh, Mai Chôu,

Pà Co, 900-1500 m 13

Taxus aff. chinensis

Poilane 4150 (P). Phu Khanh: Nha Trang,

1500m 11

Poilane 4150 (A). Nha Trang, 1500 m

12915

12-14

Schmind (P). Dalat: Dak Tria, 1400 m 15

Soulie 1523 (P). "Haut Mekong" 15

CHINA

Tibet & Yunnan

Sumatrana Group

Taxus kingstonii

Soulie 1411 (P). “Tackou et Nekou

(Haut Mekong") 11

Wallichiana Group

Taxus florinii

R.C. Ching 21505 (A). Soc. W. Sikiang:

Tamichung 10

C. W. Wang 65475 (A). Sikang, Me-kong,

Tsa-wa rung, 2500 m 8-9

Handel-Mazzetti 2602 (K). Ngaitschekou,

2800-3500 m 11

Fleigner et al. 1129 (K). Sahlie Valley on

Muzhiyan Shan, 2980 m 10

Taxus wallichiana var. yunnanensis

Zhang 916 (PE: type). Tibet, Zayul, 2100m — 15

Kingdon Ward 6292 (BM). Zayul, 7000-8000 ft 15

Sichuan

Sumatrana Group

Tax lebica

H. Smith 10401 (BM). Huangnipu, Malingtsang,

1000m 12

Wang 20541 (A). South of Kuan-Hsien,

1160m 11-12

Farges 1895-1897 (P). Tchenkéou Tin 20

Farges 128 (P). Tchenkéou Tin 14

Taxus kingstonii

Cheng 1001 (BM). Tachienlu

Cheng 1475 (P). Tachienlu 12

Taxus mairei

Wilson 1265 (A). Western: Nin Ya-chou Fu,

2000 ft 21

Wilson 1265 (US) 15

Fang 5811 (P). Nanchuan-Hsien 17

Fang 5811 (A). Nanchuan-Hsien :

Hwa 229 (K). Metasequoia area 16

Fan & Class 91 (A). Kuan-Hsien,

Chien-Chang-Shan, 1000 m 16

Farges 1436 (P). NE 16

Farges 100 (P) 15

Law 65 (K). Pei pah 15

Hwa 27 (A). Li-chuan, Jian-Nan-Hsien,

Ta-pen-Ying, 3800 ft 15

Smith 10402 (A). W region: between

Huangnipu and Yaan (Yachou),

Malingtsang, 900 m 14

Smith 10402 (S) 15

Hwa 27 (A) 14

Hwa 27 (K) 14

fal, Dat A ID hi Pr

322

Journal of of Texas 1(1)

SR MC SR MC

Legendre 586 (P). Pao Shan NE, 600m 14 17 Chiao & Fan 464 (A) 10-11 7

Fang 3461 (A). Tienchuan Hsien, Chiao & Fan 464 (P) 12 8

Tienchuanchow, 2500-3000 ft 14 20 Chiao & Fan 464 (US) 13 9

Fang 3461 (P) 3 17 Hu 8176 (A) 14 9

Fang 12205 (A). Kuan-Hsien, Mt. Tsing-cheng, Wang 20993 (A). W of Wen-chuan Hsien,

Chengtu and Kuan-Hsien, 1390 m 14 13 2800 m 11 9

Wang 20600 (A). Wah-Hsien, Mou-tao-chi, Hu 8497 (A) = 10

Metasequoia area, 1390 m 12-14 16 Fang 18310 (A) is 10

Hu 1563 (A). Shikong: Tien-Chuan Ling-Kwan, Yu 8166 (A). 2400 m 12 10

3000 ft 14-16 - Fang 15128 (A) 16 11

Farges 128 (P). Tchenkéou Tin 14 17 Fang 15128 (A) 14 11

Farges 128 (P). Tchenkéou Tin 12 24 Wilson 624 (K). S. Wushan, ravine 12 12

Fang 3442 (A). Tienchuan-Hsien, E :

Fa aM 2500-3000 ft n mp Taxus aff. chinensis

Fang 3796 (A). UnqLing-Hsien, 5000 ft 12 E Cheng 2890 (A, Taxus OCR, in Spjut 2007)

Cao 0152 (BM). Jiabigon, Zhao Quing- iu Uo: nid RN

2500 m 10 Cheng 2890 (P) 12 4

Hu 8619 (A). Emei-Hsien, Mt. Emei 12-13 8

Hace us Taxus wallichiana

Taxus chinensis Sichuan: Mt Emei

Harry Smith 10398 (BM). Tachsiangling, Hu 8166 (A) 18 4

M l Y-s Liu 1196 (A) 5 5

Re 502 (Biol. WCUU) (A) Yachow 1600 ft 15 8 Hu 8542 (A) 15-16 4

d'Legendre (P). 2500 m 14 5 Feng 3945 (A) 14 8

Henry 7155 (US: type): E Sichuan: Lee 4465 (A) 13 8

Wushan-Hsien, 2000-3000 m IS 4 Wilson 4053 (A). W Pan-lan-shan W of Kuan

Henry 7097 (US) l4 5 Hsien, 5000-6000 ft 4 4

Henry 7097 (A) 12 Pi Gansu (Kansu)

Farges 128. NE Sichuan, Tschen-kuu-tin Dist.:

(Chenkouting), (P) 13 8 Sumatrana Group

Farges 128 (P) 16 8 Taxus kingstonii

Farges 128 (P) 11 7 Meyer 1790 (P) 14 8519

Wang 1930 (A) 12 4 Ningxia Huizu

Wang 22602 (A). Kwang-yun Hsien, 1800m 12 4 .

Sichuan (Emei-Hsien: Mt. Emei) Taxus celebica

(T. chinensis, by increasing number of bare marginal cells) Chao 1223 (A). Sikong: Lung Dung An,

Feng 1941 (A) 16-17 4 1000 m 13 — 36

Fang 16082. (A) 16-17 4 Shaanxi (Shensi)

Yu 667 (A). Mt. slope, 2600 m 15 4 See also Cuspidata Alliance, T. biternata, Purdom s.n.

Wilson 6200 1415 4 Sumatrana Group (T. kingstonii)

Lee 3237. (A) le 4 Davis 1872 (P). Tsin-lin au Lao-lin, 3000 m 12

Hu 8243. (US) 13 4

Hu 8243. (A) 14 5 Wallichiana Group

Lee 4445 (A) 13 4 Taxus chinensis

Yu 669 (A). 1000 m 12 4 Chens 1893 (P). Central 5 10

Hu 8786. (A) 14 5 Yunnan

Yu 869 (A). 2500 m 12 5

Fang 18420. (A). 2335 m PELE: Sumatrana Group

Lee 4500 (A). 21 7 Taxus celebica

Ching 1676 (A). Siachu, 2600 ft 16 Forrest (A) iil 27

Ching 1676 (P). Siachu, 2600 ft [rectangular Forrest 7798 Gaoligongshan (K) 14 18

cells, papillose midrib in upper half] 16 7 ; 3

Wang 23656 (A). 2000 m 14 7 nani

Forrest 11789 (BM). Shweli-Salween Divide,

Wilson 479 (A). 12 Z 10000 ft 13 1)

Chiao & Fan. 604 (A). 1000 m 10-11 7 Forrest 11789 (K) 13 7

Fang 10940 (A). 1200 m 13 7 e

Fang 15940 (A) 13 3 Forrest 15945 (K). Schweli-Salween Divide 13 16

Forrest 15945 (BM). Schweli-Salween Divide 12 12

Spjut, Phytogeographical analysis of Taxus

Forrest 9462 (K). Ma-Chang-Kai, valley,

25?30'N, 8000 ft

Forrest (A). Yunnan, no other data

Forrest 12087 (S). Schweli-Salween Divide

Forrest 12087 (K)

Forrest 9339 (BM)

Rankin 1913 (K). " Yung Chun"

Rock 7587 (US). Salween E of Tengyueh, to

summit of Shweli, Shweli River

—

Taxus mairei

Forrest 15053 (K)

Maire 131 (BM)

Maire 1913 (P). Tie'tchang Keol, 700 m

Maire. (A: type). Dongchuan, 700-800 m

Wallichiana Group

Taxus chinensis

Feng 630 (A). Ta-hon-shan near Ta-koo, NE

of Likiang Snow Range

Cavalerie 7823 (K)

Taxus florinii

Alpine Gard. Soc. Exped. 309 (K). Zhongdian;

haba Shan, 3347 m

Rock 18502 (A). NW: Mt Ludu, NW of Li-Kiang,

W of Yangtze

Rock 18502 (US)

Forrest 19967 (S). NW: Mekong-Salween

Divide

Schneider 2918 (A). 3000-3200 m

Schneider 1429 (K)

Yu 11076 (A). sine locality

Yu 7848 (BM); Dokerla, 3100 m

Rock 11573 (A). Litiping Range, Mekong-

Yangtze divide, E of Weihsi

Rock 11573 (A)

Ching 21980 (A). Litiping, between Likiang

and Weihai

Feng 1809 (A). S Chungtien, Kung- shiang-shu,

Snow Mt to Kai-Lou-wei, Yangtze bank

3200 m

Wang 67735 (A). Wei-si Hsien, 2500 m

Wang 67414. (A). Lung-pan la Champu fung

Taxus aff. chinensis

Feng 11937 (A: type in adnot., Taxus OCR

in Spjut 2007). Si-chour-Hsien, Faa-doou,

1500m

Feng 12105 (A) l.c.

Tsai 59874 (A, T. phytonii). Wei-se Hsien

2800m

Tsai 58464 (A: type in adnot, Taxus SCU in

Spjut 2007) Che-tse-lo, 3200 m

Isai 58464 (P)

10-11

10-12

N NON 0 Ui

Taxus wallichiana var. yunnanensis

SB 1981 Exped., Cangshan 0419 (K). W

Shangschang, above Yangbi, 2700 m

Wang 67412 (A). Champu, 2120 m

1984 SAB Exped 388 (A). Xangbi Xian, W

side of Diancang Shan Mt Range,

Malultang, Chang Shan, 2700 m,

25*46' 100°01'

Handel-Mazzetti 6408 (A). Dji-shan ad boreo-

orientem urbis Dali (Talifu), 3200 m

SB 1981 Exped., Cangshan 0419 (A). W

Shangschang, above Yangbi, 2700 m

SB 1981 Exped., Cangshan 0227 (A).

Kiemiu-ingdi above Yangbi, 3000 m

Wang 72417 (A). Chen-Kang Hsien

Yu 21036 (A). Salween, Kiukiang Divide,

Shawlongwang, 2600 m

Guizhou (Kweichow)

Sumatrana Group

Taxus mairei

Steward et al. 328 (A). Ta Ho Yen, Kianakou

Hsien, 980 m

Steward et al. 328 (US)

Steward et al. 328 (P)

Cheng 7525 (A: type, T. speciosa Florin).

Kiangkow, 450 m

Steward et al. 154 (US). Liang Feng Yah,

Tsunyi Hsien, 900 m

Steward et al. 154 (A)

Tsiang 8987 (P). Pichish

Tsiang 8987 (A). Pichish

SAG Exped. 1981 (GH). Songtao Xian,

Lengjiaba, Xiaohe and Dahe Rivers,

NE Fanjing Shan mt range, 820-1120 m

Wallichiana Group

Taxus chinensis

Cavalerie & Foriupat 2604 (P)

SAG Exped. 1981 (US).

SAGB 1986 Exped. 1854 (A). Yinjiang Xian,

Xiapingsho, W Fanjing Shan range, 1

100-1400 m

SAGB Exped.1046 (A). Jiangkou Xian,

Daiyenpeng, Kaitu River, SW Fanjing

Shan range, 750-1000 m

Hubei (Hupeh)

Wallichiana Group

Taxus chinensis

Chow 76099 (A). Shenlungkai

SA 1980 Exped. 1540 (A). S of Jiuhuping

Forest along Jizigou canyon, 1900 m

Wilson 1265b (A). Western: Nin Ya-chou Fu,

2000 ft

16517

323

ONOS

324

SR MC

SA 1980 Exped. 777 (A) Western: Shennongjia

For. Dist., NE Guanmenshan, S of Shicao

river, 1150 m

SAB 1980 Exped. 1824 (GH). Shibapan, 1850 m

SAB 1980 Exped. 1824 (A)

Wilson 716 (A)

NON W W

]

Spoovy SS

Taxus aff chinensis (Taxus sp. SCU in Spjut 2007)

SA 1980 Exped. 585 (A). Western: Shennongjia

For. Dist. 331°30’N 110*30'E, 1200-1400 m 13 8

Sumatrana Group

T i

Gressitt 2507 (A). Metasequoia Area, between

Ta-yin-pin & Chunglo, Shui-sa-pa, 900m 12 14

Anhui (Anhwei)

Sumatrana Group

Taxus mairei

Ching 3168 (A). Southern, Chanen, 300 ft 12-13 12

Wallichiana Group

Taxus chinensis

R-C Ching 2622 (A). S Anhui, Clas Hara Shan 17 5

R-C Ching 2622 (US) 13 4

Cheng 4026 (BM). Wangshan 11 6

Henan (Hunan)

Taxus mairei

Fan & Li 644 (A). Ma-Ling-Tung, Sinning Hsien,

]

600 m 5 18

Fan & Li 296 (A). Changning Hsien, Yang-Shan,

680m e -

Fan & Li 296 (BM) 14 12

Jiangsu (Kiangsu, Kiangshi)

Taxus mairei

Wang-Te-Hui 445 (A). Ningdu, Yuntungtschi 9-12 25

Y.K. Hsiung 6443 (A). NW, Si-ho, Hwang-

kong-shan Mt - 11

Wang-Te-Hui 458 (A). Lienhwa-shan, 800m - 10-12

Chow 80325 (BM). Nanking, 75 m 12 16

Guangxi (Kwangsi)

Sumatrana Group

Taxus celebica

Ching 5976 (US). Bin Long, Miu Shan, N Luchen,

border of Kweichow, 4000 ft 15-16 32

Ching 5976 (A) 14-15 31

Taxus mairei

Chiao 18795 (US). Lu Shan 16 6-10

Wallichiana Group

Taxus chinensis

Steward & Cheo 947 (P). 2110 m 15 4

Steward & Cheo 947 (BM). San Chiang Hsien,

2110m 12 4

Zhejiang (Chekiang)

Piba Dat o ID AKI

Journal of

Taxus mairei

Cheng 3617 (US). Eastern: Tien-Mu-Shang 14-16

Keng 317 (A). Taishun-Hsien 15

Hu 342 (A). Tien-Tai-Shan, 1300 m 18.

S. Chen 1063 (A) 12

(US: 145110). Tien Tai Shan 12

Ching 2489 (A). S: King Yuan, 300-800 m 11

Ching 2489 (US) 13

Taxus sumatrana

Hu 1628 (A). Lin-an Hsien, 1200 ft 14

Hu 550 (A). Y-Chien Hsien, 1000 ft 12

Guangdong

Taxus celebica

Nanling Exped. 1838 (A). Ruyuan Xian -

Taxus mairei

Tsang 20694 (US). Loh Ch'ang Dist., Chong

Uen Shan near Kau Fung 13-15

Tsang 20694 (A) -

Chiao 14510 (A). Tien-Tai-Shan, 1300 m 12

Chiao 14510 (US) 12

Tsiang Ying 1425 (A). Hung-mio to Mio- lan,

Jui-feng, Lokohong Hsien N.R. Region,

1340 m 12

Tsiang Ying 1425 (A, different label) 14

Tsiang Ying 1425 (P, specimen does not

appear to be the same plant as in A) 13

Fujian (Fukien)

Sumatrana Group

Taxus mairei

Price 1258b (K). Ing-dan E. Fookma 12

Sheng 1544 (K). Naping, 800 m 16

Chung 2865 (A). Yeuping, Shih-Sun-Keng,

650 m 18

Chung 2865 (K) 12

David (P). W: Mts 12

Chung 3581 (A). Buong Kang, mt slope,

700m 9

He-Guosheng 1544 (US). Naping, 800 m 16

Wallichiana Group

Taxus aff chinensis

H.H. Chung 3866 (A). Puchen 11

Taiwan

Sumatrana Group

Taxus kingstonii

Hsu 1651 (PH). Mt. Pasein-san, Taichang Hsien 16

Liu 389 (PH). Mt. Ammashan, Taichung Hsien 15

Liu 389 (A) 14

Hsu (PH). Mt. Pasan-shan, Taichang Hsien 14

Wilson 9738 (BM, isotype). Arisan Prov.: Kagi,

2833m 13

Wilson 9738 (A: holotype) 10-11

Liu 437 (US). Taiklang, Shih-wan-hsi,

Pa-Hsien-shan, 2250 m 12

titute of Texas 1(1)

Spjut, Phytogeographical analysis of Taxus

Nakahara (PH). Arizan Prov.

Cj Chang, Tongshi 6 (wba)

27 Sep 93

26 Nov 93

06 Dec 93

09 Dec 93

27 Jan 94.

13 Jan 94 (new growth)

13 Jan 94 (old growth)

Taxus mairei

Gj Chang 1—2,4—5, 7-10 (wba). Hua-lien

1 16 May 94

1 03 Aug 94

2 16 May 94

2 03 Aug 94

4 16 May 94

4 03 Aug 94

5 16 May 94

503 Aug 94

9 16 May 94

9 03 Aug 94

10 16 May 94

10 03 Aug 94

7 16 May 94

7 03 Aug 94

8 16 May 94

8 03 Aug 94

Taxus sumatrana

de Laubenfels P 671 (A). Tai-shu Shan For. Dist.,

2000 m (Rt. 210, 7km)

de Laubenfels P 670 (A). l.c.

Gj Chang

3 (wba) 16 May 94

303 Aug 94

Wallichiana Group

Chinensis Subgroup

Wilson 11154D (A: type in adnot). Karenko

Prov.: mts W of Karenko

Gj Chang, Tongshi 5 (wba)

27 Sep 93

26 Nov 93

06 Dec 93

09 Dec 93

13 Jan 94

Gj Chang, Tongshi 7 (wba).

27 Sep 93

26 Nov 93

06 Dec 93

09 Dec 93

27 Jan 94

Cj Chang 6 (wba). Hua-lien

16 May 94

03 Aug 94

| |

hat,

SINDEN ¿0 NO NI NS

ÑO: 00:09 LO NO: SA: NO:

Ju oO. OO Cd

Cr Ora MO: sse

Origin?

Y. Sugilara, Ex TUS (GH).[arbitrarily placed

ere]

THE PHILIPPINES

Sumatrana Group

Taxus sumatrana

Merrill 4595 (US). Lepanto Dist., Mt. Data

de Laubenfels P650 (GH). Benguet, 58 km

N of Baguio, 2100 m

Whitehead 1896 (BM). NW central Luzon:

5000-7000 ft

Williams 1002 (US). Benguet: Mt. St. Tomas

Leano 20672 (US). Benquet Prov.

Mt. St. Tomas

Vidal 623 (GH); Mt. Banahao, Pr. Tayabas

Wallichiana Group

Chinensis Subgroup

E.C. Leano 25128 (US). Luzon: Benquet Prov.,

t. St. Tomas

Elmer 6244 (P). Mt. St. Tomas

Elmer 6244 (US)

Loher 4850 (K). Luzon: central.

Loher 4850 (US)

Curran 5015 (P). Benquet Prov., Mt. Tonglan

Curran 5015 (PH)

Jacobs 7171 (K). Luzon: Mt. Pulog,

2200-2300 m

Ramos & Edaño 40234 (K). Luzon: Lepanto,

Mt. Data

Ramos & Edaño 40234 (P)

Wilkes Exped. 1838-1842 (GH). Luconia:

Mt. Mahaihai

Curran 7911 (US). Luzon: Benquet Prov.,

. Banajao

Merrill 839 (US). Luzon, Benquet Prov.

Merrill 839 (US)

Ocampo 27920 (A). Mt. Banajao

Ocampo 27920 (P)

de Laubenfels P669 (GH). Luzon: Laguna Prov.,

Mt. Banajao, 2100 m

de Laubenfels P668. (GH: type in adnot.)

Herb. Hook. (K). Luconia, 7600 ft

Loher 7139 (US)

Sulit 2350 (A). Luzon: Benguet Prov., Mt. Pauai,

2450m

Alvarey 18369 (BM). Benquet Prov.

INDONESIA

Sulawesi (Celebes)

Sumatrana Group

Taxus celebica

Everett 35 (K: type, Podocarpus celebicus

Hemsley). Bonthian Peak, 7000-1 0,000 ft

Taxus sumatrana

Teysmann 14190 (U). Bonthian

325

SR MC

12-13 14

151399

| EE ^

13, 10212

11-13 9

| E.

16 4

14 4

| NE^

14 4

12 4

14 7

l4 4

4 23

l4 2

3 è 2

12/13 5

12 894

12 4

| P

| NS

: 4

10-11 4

: 3

12-14 8

14-15 22

1-13 16

326 Journal of the Botanical R h Institute of Texas 1(1)

SR MC SR MC

Wallichiana Group, Chinensis Subgroup Sumatrana Group

Neth. Ind. For. Serv. bb:19577 (A). Ond. Malili Taxus sumatrana

1800 m 12:15: 12 Teysmann s.n. (U: type). Sumatra: western,

Neth. Ind. For. Serv. bb:20887 (K). Goua Fort de Kock, 3000 m PEAR?

Lambaja, 2000 m 11 4 de Voogd 1503 (K). Palembang, Dempo,

Sumatera 1000 m CER:

SR MC

Wallichiana Group, Chinensis Subgroup

Boschprochation 7709 (U). Tharolanden,

1400 m 14 4

ACKNOWLEDGMENTS

I thank Zlatko Kvaček of critical comments on the manuscript and Wang Qi for providing a translation of

the abstract in Chinese.

REFERENCES

ARY SHAW, H.K. 1973. A dictionary of the flowering plants and ferns. 8th ed. Revised from J.C. Willis. Cambridge

University Press, London.

ANDRÉ, D. 1956. Contribution à l'étude morphologique du cóne femelle de quelques gymnospmermes (Cepha-

lotaxacées, Juniperoidées, Taxacées). Nat. Monspliensia (Bot.) 8:3-35.

AXELROD, D.I. 1958. Evolution of the Madro-Tertiary Geoflora. Bot. Rev. 24:433-509.

AXELROD, D.I. 1975. Evolution and biogeography of Madrean-Tethyan sclerophyll vegetation. Ann. Missouri Bot.

Gard. 62:280-334.

AXELROD, D.I. 1976. History of the coniferous forests, California and Nevada. Univ. Calif. Press, Berkeley, 1-62.

AXELROD, D.I. 1983. Biogeography of oaks in the Arcto-tertiary Province. Ann. Missouri Bot. Gard. 70:629-657.

AXELROD, D.I. 1986. Cenozoic history of some western American pines. Ann. Missouri Bot. Gard. 73:565-641.

AXELROD, D.I., |. AI-SHEHBAZ, and P.H. Raven. 1998. History of the modern flora of China. In: Z. Aoluo and W. Sugong,

eds. Proceedings of the First International Symposium on Floristic characteristics and diversity of East Asian

plants, July 25-27, 1996, Kunming, Yunnan. China Higher Education Press, Beijing and Springer-Verlag Berlin

Heidelberg. Pp. 43-55.

BAILEY, L.H. 1933. The cultivated conifers of North America. MacMillan Co., New York.

BARTHOLOMEW, B. 1999. Re-evaluation of the biological effect of plate movement.—Impact of Shan—Malay plate

displacement (The movement of Burma-Malaya Geoblock) on the biota of the Gaoligong Mountains. Acta

Bot. Yunnanica 21:407-425.

BENNIKE O. 1990. The Kap København Formation: stratigraphy and paleobotany of a Plio-Pleistocene sequence in

Peary Land, North Greenland. Meddel. Grønland Geosci. 23:1-85.

BERTRAND, C.E. 1874. Anatomie comparée des tiges et des feuilles chez les Gnétacées et les Coniféres. Ann. Sci.

Nat. Bot. sér. 5, 20:6-153.

Bo8OLA, M.S., R.T. Eckert, A.S. KLEIN, K. SrAPELFE, D.E. SmitH, and D. Guenette. 1996. Using nuclear and organelle DNA

markers to discriminate among Picea rubens, Picea mariana, and their hybrids. Canad. J. For. Res. 26:433-443.

Bosrov, A.V.F. C.H., A.P. MELKIAN, S. MIKHAIL, S. Romanov, and A.N. Sorokin. 2004. Seed morphology and anatomy of

Austrotaxus spicata (Taxaceae) and its systematic position. Bot. J. Linn. Soc. 145:437-443.

Bourronp, D.E. and S.A. SPONGBERG. 1983. Eastern Asian-eastern North American phytogeographical relationships—A

history from the time of Linnaeus to the twentieth century. Ann. Missouri Bot. Gard. 70:421-439.

BRAUN, L. 1950. Deciduous forests of eastern North America. The Blankiston Co., Toronto.

BURNHAM, RJ. and A. GraHam. 1999. The history of Neotropical vegetation: New developments and status. Ann.

Missouri Bot. Gard. 86:546-589.

Chaney, RW, 1947. Tertiary centers and migration routes. Ecol. Monogr. 17:139-148.

CHAW, S-M, H. Lone, B-s Wana, A. ZHARKIKH, and W-H Li. 1993. The phylogentic position of Taxaceae based on 18s rRNA

sequences. J. Molec. Evol. 37:624-630.

Spjut, Phytogeographical analysis of Taxus 327

Chaw, S-M, H-M SUNG, H. LONG, A. ZHARKIKH, and W-H. Li. 1995. The phylogenetic position of the conifer genera

Amentotaxus, Phyllocladus, and Nageia inferred from 18S rRNA sequences. J. Molec. Evol. 41:224-230.

CHENG (ZHENG), W-c. 1934. An enumeration of vascular plants from Chekiang, Ill. Gymnospermae. Contrib. Biol.

Lab. Chinense Assoc. Adv. Sci. 9:240-241.

CHENG (ZHENG), W-c., and L-k. Fu. 1978. Taxaceae. In: Fl. Reipub. Pop. Sin. 7, Gymnospermae. Agendae Academiae

Sinicae. [English Translation].

CHENG, Y., R.G. NicoLson, K. Tripp, and S-M. CHaw. Phylogeny of Taxaceae and Cephalotaxaceae genera inferred

from chloroplast matK gene and nuclear rDNA ITS region. 2000. Molec. Phylogen. Evol. 14:353-365.

Couns, D., R.R. Mi, AND M. MOLLER. 2003. Species separation of Taxus baccata, T. canadensis, and T. cuspi-

data (Taxaceae) and origins of their reputed hybrids inferred from RAPD and cpDNA data. Amer. J. Bot.

90:175-182.

Cope, E.A. 1998. Taxaceae: The genera and cultivated species. Bot. Rev. 64:291-322.

CULBERSON, C.F., W.L. CULBERSON, and A. Johnson. 1990. The Ramalina americana complex. Bryologist 93:167-

186.

Dempsey, D. and I. Hook. 2000. Yew (Taxus) species—chemical and morphological variations. Pharm. Biol.

38:274-280.

DeRYUGINA, T.F. and N.D. NesreRovicH. 1981. Peculiarities of the morphological and anatomical structure of

conifer needles of some Taxus L. species. Dokl. Akad. Nauk Bel. S.S.R. 25(7):652-655 [Russian with English

summary].

DiLcHER, D.L. 1969. Podocarps from the Eocene of North America. Science 164:299-301.

DuNG, V., VAN (ed). 1996. Vietnam forest trees. Ten contributors, Agric. Publ. House, Hanoi.

Dupter, A. 1920. Ovuliferous structures of Taxus canadensis. Bot. Gaz. 69:492-520.

EL-KassagY, Y.A. and A.D. YaNcHuK. 1994. Genetic diversity, differentiation, and inbreeding in Pacific yew from

British Columbia. J. Hered. 85:112-117.

EL-Kassaev, Y.A. and A.D. YANCHUK. 1995. Genetic variation of Pacific Yew in British Columbia and its conserva-

tion. In: Ph. Baradat, W.T. Adams, and G. Müller-Starck, eds. Population genetics and genetic conservation

of forest trees. Academic Publishing, Amsterdam, The Netherlands. Pp. 227-235.

Fapy, B., M. ArBEz, and A. Marreau. 1992. Geographic variability of terpene composition in Abies cephalonica

Loudon and Abies species around the Aegean: hypotheses for their possible phylogeny from the Miocene.

[rees 6:162-171.

FADY-WELTERLEN, B. 2005. Is there really more biodiversity in Mediterranean forest ecosystems? Taxon 54:

905—910.

FARJON, A. 1990. Pinaceae. Koeltz Scientific Books, Kónigstein, Germany.

FARJON, A. 1998. World checklist and bibliography of conifers. The Royal Botanic Gardens, Kew.

FARJON, A. 2001. World Checklist and Bibliography of Conifers. 2nd edition. The Bath Press, Bath, United

Kingdom.

FERGUSON, D.K. 1978. Some current research on fossil and recent taxads. Rev. Palaeobot. Palynol. 26:213-

FLORIN, R. 1931. Untersuchungen zur Stammesgeschichte der Coniferales und Cordaitales. Kongl. Svenska

Vetenskapsakad. Handl. 10:45-588.

FLORIN, R. 1948a. Enumeration of gymnosperms collected on Swedish expeditions to western and north-

western China. Acta Horti Berg. 14:343-384.

FLORIN, R. 1948b. On the morphology and relationships of the Taxaceae. Bot. Gaz. 110:31-39,

FLORIN, R. 1948c. On Pseudotaxus, a new genus of the Taxaceae, from eastern China. Acta Horti Berg. 14:385—395,

plates 1-3.

FLORIN, R. 1948d. Nothotaxus or Pseudotaxus? Bot. Not. 1948:270-272.

FLORIN, R. 1951. Evolution in cordaites and conifers. Acta Horti Berg. 16:285—388, plate.

FLORIN, R. 1958. On Jurassic taxads and conifers from north-western Europe and eastern Greenland. Acta

Horti Berg. 17:257—402, 56 plates.

328 Journal of the Botanical R h Institute of Texas 1(1)

FLORIN, R. 1963. The distribution of conifer and taxad genera in time and space. Acta Horti Berg. 20:121-

FRANCO, J.A. 1964. Taxus. Fl. Europaea 1:39.

FREDERIKSEN, N.O. 1994. Paleocene flora diversities and turnover events in eastern North America and their

relation to diversity models. Rev. Palaeobot. Palynol. 82:225-238.

FREDERIKSEN, N.O. 1995. Differing Eocene floral histories in southeastern North America and western Europe: Influ-

ence of paleogeography. Hist. Biol. 10:13-23.

FRIMMEL, F. VON. 1911. Die untere Kutikula des Taxus- Blattes—ein Lichtreflektor. Oesterr. Bot. Z. 61:216-223.

Frypay, A.M. 2006. New and interesting North American lichen records from the alpine and sub-alpine zones of

Mt. Katahdin, Maine. The Bryologist 109:570-578.

Fu, L-k, N. Li and R.R. Mitt. 1999. Taxaceae. Fl. China 4:89—96, Missouri Botanical Garden Press, St. Louis.

GAUSSEN, H. 1979. Les gymnospermes actuelles et fossiles: fasc. 15. Les taxines. Toulouse, Unversité Paul-Sabatier

Faculté des Sciences.

GELDEREN, D.M. van and J.R.P. van Hoey SmitH. 1996. Conifers. An illustrated encyclopedia. 2 Vols. Timber Press,

Portland, OR.

GERNANDT, G.S., G.G. Lopez, S.O. García and A. Liston. 2005. Phylogeny and classification of Pinus. Taxon 54:29-42.

Goon, R. 1964. The geography of the flowering plants. John Wiley & Sons, New York.

GivuLescu, R. 1973. Die fossil Koniferen des Fundortes Chiuzbaia F. Inst. Géol. Mém. 19:31-34.

GRAHAM, A. 1972. Outline of the origin and historical recognition of floristic affinities between Asia and eastern

North America. In: A. Graham, ed., Floristics and paleofloristics of Asia and eastern North America, Elsevier

Publ. Co., Amsterdam. Pp. 1-18.

GRAHAM, A. 1973. History of the arborescent temperate element in the northern Latin American Biota. In: A.

Graham, ed. Vegetation and vegetational history of northern Latin America. Elsevier Scientific Publ. Co.,

Amsterdam. Pp. 301-314.

GRAHAM, A. 1993. History of the vegetation: Cretaceous (Maastrichtian)— Tertiary. Fl. North America 1:57-70.

GRAHAM, A. 1999. Late Cretaceous and Cenozoic history of North American vegetation north of Mexico. Oxford

Univ. Press, New York.

HAGENEDER, F. 2007. Yew. A history. Sutton Publishing Ltd., Thrupp-Stroud-Goucestershire.

HAMILTON, W. 1983. Cretaceous and Cenozoic history of northern continents. Ann. Missouri Bot. Gard. 70:

440—458.

Hara, H. 1972. Corresponding taxa in North America, Japan and the Himalayas. In: D.H. Valentine, ed. Taxonomy,

phytogeography and evolution. Academic Press, New York. Pp. 61-72.

Harris, T.M. 1976a. Two neglected aspects of fossil conifers. Amer. J. Bot. 63:902-910.

Harris, T.M. 1976b. The Mesozoic gymnosperms. Rev. Paleobot. and Palyn. 21:119-134.

HAYASHI, Y. 1954. The natural distribution of important trees, indigenous to Japan. Conifers Report 3. Bull. For. Exp.

Sta. Meguro, Tokyo 75:166-173 (English summary).

Heinze, B. 2004. Zur Populationsbiologie der gemeinen Eibe (Taxus baccata). Centralblatt für das gesamte Forst-

wesen 121:47-59.

HiLL, K.D. 1998. Gymnosperms—The paraphyletic stem of seed plants. Fl. Australia 48:505—526.

Hits, M. 1993. Taxaceae Gray. Yew family. Fl. North America 2:423-427.

HOLMGREN, P.K., N.H. HOLMGREN, and L.C. Barnett (eds). 1990. Index Herbariorum. Part I: The herbaria of the world.

8^ ed., W. Junk, Hague.

Hou, H-Y. 1983. Vegetation of China with reference to its geographical distribution. Ann. Missouri Bot. Gard.

70:509-548.

Hsu, J. 1983. Late Cretaceous and Cenozoic vegetation in China, emphasizing their connections with North

America. Ann. Missouri Bot. Gard. 70:490-508.

HuwPHRIES, C.J. 1982. Vacariance biogegoraphy in Mesoamerica. Ann. Missouri Bot. Gard. 69:444—463.

Hunt, R.S. 1993. Abies Miller. Fir. Fl. North America 2:354-362.

Spjut, Phytogeographical analysis of Taxus 329

Jacoss, B.F., J.D. Kinaston, and L.L. Jacogs. 1999. The origin of grass-dominated ecosystems. Ann. Missouri Bot.

Gard. 86:590-643.

JINXING, L. and H. Yuxi (eds.). 2000. Atlas of structure of gymnosperms. Science Press, Beijing.

Klaus, W. 1989. Mediterranean pines and their history. Pl. Syst. Evol. 162:133-163.

Kucka, J. and R.M. Zink. 1997. The importance of recent ice ages in speciation: A failed paradigm. Science

277:1666-1669.

Kornas, J. 1972. Corresponding taxa and their ecological background in the forest of temperate Eurasia and

North America. In: D.H. Valentine, ed. Taxonomy, phytogeography and evolution. Academic Press, New York.

Pp. 37—59.

Kotyk, M.E. A., J.F. Basincer, and E.E. Mclven. 2003. Early Tertiary Chamaecyparis Spach from Axel Heiberg Island,

Canadian High Arctic. Canad. J. Bot. 81:113-130.

KRUSSMANN, G. 1985. Manual of cultivated conifers. Translated by M. E. E, eds. H.-D. Warda €: G. S. Daniels. Timber

Press, Portland.

KRISHNAN, M.S. 1974. Ill. Geology. In: M.S. Mani, ed. Ecology and biogeography of India. W. Junk, Hague. Pp.

60-98.

Kuarrzki, K. and W. Krutzscu. Origins of East and Southeast Asian plant diversity. 1998. In: Z. Aoluo and W. Sugong,

eds. Proceedings of the First International Symposium on Floristic characteristics and diversity of East Asian

plants, July 25-27, 1996, Kunming, Yunnan. China Higher Education Press, Beijing and Springer-Verlag Berlin

Heidelberg. Pp. 56-70.

Kvacek, Z. [1984 fide author]. Tertiary taxads of NW Bohemia. 1982 Acta Univ. Carol., Geol., Pokorny 4:471-491.

Kvacek, Z. and H. WaLTHER. 1998. The Oligocene flora of Kundratice near Litoměřice, České středohoří Volcanic

Complex (Czech Republic)—a review. Acta Mus. Nat. Pragae, B, 54:1-42.

Kvacek, Z. and W.C. RemBer. 2000. Shared Miocene conifers of the Clarkia flora and Europe. Acta Universitatis

Carolinae - Geologica. 44(1):75-85.

Kvacek, Z., and W.C. Rember. Under Review. Calocedrus robustior (Cupressaceae) and Taxus howardii (Taxaceae):

Two new conifers from the Middle Miocene of the western North America (Clarkia area, Latah Formation,

northern Idaho). Paleobios.

Kwe Y-L. and S.-Y. Hu. 1974. [Epidermal feature of leaves of Taxus in relation to taxonomy]. Acta Phytotax. Sin.

12(3):329-334, plate 67 [Chinese with English summary].

Larson, D.W., U. MATTHES, J.A. GERRATH, N.W.K. Larson, J.M. GERRATH, J.C. NekoLa, G.L. WALKER, S. PoREMBSKI, and A. CHARLTON.

2000. Evidence for the widespread occurrence of ancient forests on cliffs. J. Biogeogr. 227:319-331.

LAUBENFELS, D.J., DE. 1988. Coniferales. Fl. Malesiana 10(3):337-453.

LeoroLD, E.B. and H.D. MacGinme, 1972. Development and affinities of Tertiary floras in the Rocky Mountains. In:

A. Graham, ed. Floristics and paleofloristics of Asia and eastern North America. Elsevier Publ. Co., Amsterdam.

Pp. 147-200.

LePace, B.A. 2003. A new species of Thuja (Cupressaceae) from the Late Cretaceous of Alaska: implications of

being evergreen in a polar environment. Amer. J. Bot. 90:167-174.

Li, H-L. 1952. Floristic relationships between eastern Asia and eastern North America. Trans. Amer. Philos. Soc.

42:371-429.

Li, J., C.C. Davis, PD. Trepicl, and M.J. Donochue. 2001. Phylogeny and biogeography of Taxus (Taxaceae) inferred from

sequences of the internal transcribed spacer region of nuclear ribosomal DNA. Harv. Pap. Bot. 6:267-274

Li, X-w and J. Li. 1997. The Tanaka-Kaiyong Line—An important floristic line for the study of the flora of East Asia.

Ann. Missouri Bot. Gard. 84:888-892.

Liston, A., W.A. Rosinson, D. PIÑERO, and E.R. ALvarez-BuyLLa. 1999. Phylogenetics of Pinus (Pinaceae) based on nuclear

ribosomal DNA internal transcribed spacer region sequences. Molec. Phylogen. Evol. 11:95-109.

Little, D.P. 2006. Evolution and circumscription of true cypresses (Cupressaceae: Cupressus). Syst. Bot. 31:

461-480.

Liu, T-s. 1971. A monograph of the genus Abies. Dept. For., College Agric., Natl. Taiwan Univ., Taipei.

330 Journal of the Botanical R h Institute of Texas 1(1)

LoUDON, J.C. 1844. Arboretum et fruticetum Britannicum. Longman, Brown, Green and Longmans, London.

Mal, D.H. 1989. Development and regional differentiation. Pl. Syst. Evol. 162:79-91.

Mal, D.H. 1998. Contribution to the flora of the middle Oligocene Calu Beds in Brandenburg, Germany. Rev.

Palaeobot. Palynol. 101:43-70.

MANCHESTER, S.R. 1994. Fruits and seeds of the Middle Eocene Nut Beds Flora, Clarno Formation, Oregon. Palae-

ontographica Americana 58:1-205.

MANCHESTER, S.R. 1999. Biogeographical relationships of North American Tertiary Floras. Ann. Missouri Bot. Gard.

86:472-522.

Mani, M.S. 1974. Biogeographical evolution in India. In: M.S. Mani, ed. Ecology and biogeography of India, W.

Junk, Hague. Pp. 698-722.

Mclver, E.E. 1994. An early Chamaecyparis (Cupressaceae) from the Late Cretaceous of Vancouver Island, British

Columbia, Canada. Canad. J. Bot. 72:1787-1796.

Mclver, E.E. 1999, Paleobotanical evidence for ecosystem disruption at the Cretaceous-Tertiary boundary from

Wood Mountain, Saskatchewan, Canada. Canad. J. Earth Sci. 36:775-789.

Mclver, E.E. and J.F. Basincer. 1989. The morphology and relationships of Thuja polaris sp. nov. (Cupressaceae) from

the early Tertiary Ellesmere Island, Arctic Canada. Canad. J. Bot. 67:1903-1915.

Mclver, E.E and J.F. Basincer. 1999. Early Tertiary floral evolution in the Canadian High Arctic. Ann. Missouri Bot.

Gard. 86:523-545.

McKenna, M.C. 1983. Holarctic landmass rearrangement, cosmic events, and Cenozoic terrestrial organisms. Ann.

Missouri Bot. Gard. 70:459—489.

Meven, S.V. 1984. Basic features of gymnosperm systematics and phylogeny as evidenced by the fossil record.

Bot. Rev. 5:1-111.

Meyer, H.W. and S.R. Manchester 1997. The Oligocence Bridge Creek flora of the John Day Formation, Oregon.

Univ, Calif. Publ. Geol. Sci 141:1-195, 75 pl.

MiLLAR, C.I. 1993. Impact of the Eocene on the evolution of Pinus L. Ann. Missouri Bot. Gard. 80: 471-498.

MiLLER, C.N., JR. 1977. Mesozoic conifers. Bot. Rev. 43:217-280.

MILLER, C.N., JR. 1988. The origin of modern conifer families. In: C.B. Beck, ed., Origin and evolution of gymnosperms.

Columbia Univ. Press, New York. Pp. 448-486.

MIRANDA, F. and A.J. SHarp. 1950. Characteristics of the vegetation in certain temperate regions of eastern Mexico.

Ecology S 13139323.

Mirov, N.T. 1953. Taxonomy and chemistry of the white pines. Madroño 12:81-89.

Nakal, T. 1938. Indigenous species of conifers and taxads of Korea and Manchuria, and their distribution. Chósen

Sanrin Kaihó (J. Kor. For. Soc.) 158221.

NAJMAN, Y. and E. Garzanti. 2000. Reconstructing early Himalayan tectonic evolution and paleogeography from

Tertiary foreland basin sedimentary rocks, northern India. GSA Bulletin; 112 (3):435-449.

NicoLosi, R.T. 1982. Morphological features of leaves and pollen as in aid in separating selected species and

cultivars within the genus Taxus. Thesis, Ph.D., Ohio State University.

Novacek, MJ. 1999. 100 million years of land vertebrate evolution: The Cretaceous-Early Tertiary transition. Ann.

Missouri Bot. Gard. 86:230-258.

Ohwi, J. 1965. Flora of Japan. English translation, eds. F. G. Meyer & E. H. Walker, Smithsonian Institution, Wash-

ington, DC.

O'Kerre, J.D. and T.J. AHRENS. 1989. Impact production of CO, by the Cretaceous/Tertiary extinction bolide and the

resultant heating of the Earth. Nature 338:247-249.

OUDEN, P. DEN and B.K. Boom. 1965. Manual of cultivated conifers. M. Nijhoff, Hague.

PALAMAREV, E. 1989. Paleobotanical evidence of the Tertiary history and origin of the Mediterranean sclerophyll

dendroflora. Pl. Syst. Evol. 162:93-107.

Puipps, J.B. 1983. Biogeographic, taxononmic, and cladistic relationships between east Asiatic and North American

Crataegus. Ann. Missouri Bot. Gard. 70:667-700.

Spjut, Phytogeographical analysis of Taxus 331

PILGER, R. 1903. Taxaceae-Taxoideae—Taxeae. Taxus. In: Engler, Das Pflanzenreich IV:110-116.

PiLGER, R. 1916. Die Taxales. Mitt. Deutsch. Dendrol. Ges. 25:1-28.

PILGER, R. 1926. Taxaceae. In: Engler, A. and K. Prantl, eds. Die natürlichen Pflanzenfamilien, 2nd ed., 13:199-211.

PRAGER, E.M., D.P. FowLer and A.C. WiLson. 1976. Rates of evolution in conifers (Pinaceae). Evolution 30:637-649.

PRAKASH, U., D. NA-zHENG, and PP. TRIPATHI. 1995. Fossil woods from the Miocene sediments of China with remarks

on environmental implications of Miocene floras of the region. Barbal Sahni Centenary 1995:341-360.

Price, R.A. 1990. The genera of Taxaceae in the southeastern United States. J.Arnold Arbor. 71:69-71.

Qian, H. 2002. Floristic relationships between eastern Asia and North America: Test of Gray's hypothesis. Amer.

Nat. 160 (3):31 7-332.

Qian, H. and R.E. Rickters. 1999. A comparison of taxonomic richness of vascular plants in China and the United

States. Amer. Nat. 154 (2):160-181.

Rao, A.S. 1974. The vegetation and phytogeography of Assam-Burma. In: M.S. Mani, ed. Ecology and biogeog-

raphy of India. W. Junk, Hague. Pp. 204-246,

Rau, M.A. 1974. Vegetation and phytogeography of the Himalaya. In: M.S. Mani, ed. Ecology and biogeography

of India. W. Junk, Hague. Pp. 247-280.

ReHDER, A. 1919. New species, varieties and combinations from the herbarium and the collections of the Arnold

Arboretum. J. Arnold Arbor. 1: 44-60.

REHDER, A. 1940. Manual of cultivated trees and shrubs. 2nd ed. MacMillan Co., New York.

Rosen, D.E. 1975. A vicariance model of Caribbean biogeography. Syst. Zool. 24:431-464.

SAHNI, B. 1920. On certain archaic features in the seed of Taxus baccata, with remarks on the antiquity of the

Taxineae. Ann. Bot. 34:117-133.

SARGENT, C.S. 1914. Plantae wilsonianae. Vol. 2. Cambridge University Press, Cambridge, MA.

ScHMiD, M. 1974. Végétation du Viet-nam. Mém. Orstom 74, Paris.

SCHUSTER, R. 1976. Plate tectonics and its bearing on the geographical origin and dispersal of angiosperms. In: C.B.

Beck, ed. Origins and early evolution of angiosperms. Columbia Univ. Press, New York. Pp. 48-138.

SHARP, A.J. 1966. Some aspects of Mexican phytogeography. Ciencia (México) 24:229-232.

SiLBA, J. 1984. An international census of the coniferae, |. Phytologia Mem. 7:1-79.

SINGH, H. 1961. The life history and systematic position of Cephalotaxus drupacea Sieb. & Zucc. Phytomorphol-

ogy 11:153-196.

SpJur, RW. 1992. A taxonomic key to the species of Taxus. NCI Workshop on Taxus, Taxol, and Taxotere, Rockville,

MD (Abstract only).

Spjur, RW. 1993. Reliable morphological characters for distinguishing species of Taxus (Abstract). In: International

yew resource conference. Yew (Taxus) conservation biology and interactions. Berkeley, CA. Pp. 39-40.

SpJur, RW, 1994. A systematic treatment of fruit types. Mem. New York Bot. Gard., Vol 70.

Spyut, RW. 1998. Two papers presented at the AIBS Annual Meeting, American Systematic Plant Taxonomists,

Baltimore Convention Center, MD, 5 Aug. 1998. Abstracts published on the Internet, Botanical Society of

America.

(a) Morphological evolution in the Taxus leaf and its significance to recognizing ecological species within the

genus. http://www.ou.edu/cas/botany-micro/bsa-abst/section 1 3/abstracts/107.shtml.

(b) Species of Taxus. http://www.ou.edu/cas/botany-micro/bsa-abst/section 1 3/abstracts/112.shtml.

Spyut, RW. 2000. Three papers presented at the joint meetings of the Botanical Society of America and American

Systematic Plant Taxonomists, Portland, OR, August, abstracts published online and Amer. J. Bot.

(a) A phytogeographical analysis and classification of leaf morphological features in Taxus (Taxaceae). http://

www.ou.edu/cas/botany-micro/botany2000/section13/abstracts/27.shtml.

(b) The morphological relationships of Taxus canadensis (Taxaceae) in North America and Eurasia. http://www.

ou.edu/cas/botany-micro/botany2000/section1 3/abstracts/28.shtml.

botany2000/section13/abstracts/166.shtml. (Note key also presented on the WBA website in 2003, extracted

from poster document of 24 pages that included 270 figures).

332 Journal of the Botanical R h Institute of Texas 1(1)

Spjur, RW. 2006. Biogeographical data on putative hybrids for species of Taxus (Taxaceae) in the Himalayas and

North America. Abstract, Botanical Society of America, Chico, CA, July 29-Aug 2. http://www.botanyconfer-

ence.org/er git 1e/search/index.ph p?fu nc=detail&aid=37

Spjur, R.W. 2007. Taxonomy and nomenclature of Taxus (Taxaceae). J. Bot. Res. Inst. Texas 1:203-289.

Srivastava, S.K. 1994. Evolution of Cretaceous phytogeoprovinces, continents and climates. Rev. Paleobot. Palynol.

82:197-224.

STAFLEU, F.A. and R.S. Cowan. 1976-1988. TL-2. Taxonomic literature. 7 Vols. W. Junk, Hague.

Strauss, S.H., A.H. DOERKSEN, and J.R. Byrne. 1989. Evolutionary relationships of Douglas-fir and its relatives (genus

Pseudotsuga) from DNA restriction fragment analysis. Canad. J. Bot. 68:1502-1510.

SUYAMA, Y., H. YosHimaru, and Y. Tsumura. 2000. Molecular phylogenetic position of Japanese Abies (Pinaceae) based

on chloroplast DNA sequences. Mol. Phylogen. Evol. 16:271-277.

Thomas, P.A. and A. Pouwanr. 2003. Biological flora of the British Isles. Taxus baccata L. J. Ecol. 91:489-524.

THORNE, R.F. 1972. Major disjunctions in the goegraphic ranges of seed plants. Quart. Rev. Biol. 47:365-411.

THORNE, R.F. 1978. Plate tectonics and angiosperm distribution. Notes Roy. Bot. Gard. Edinburgh 30:297-315.

TirFNEY, B.H. 1985a. Perspective on the origin of the florisitc similarity between eastern Asia and eastern North

America. J. Arnold Arbor. 66:73-94.

Tirruey, B.H. 1985b. The Eocene North Atlantic Land Bridge: Its importance in Tertiary and modern phytogeog-

raphy of the northern hemisphere. J. Arnold Arbor. 66:243-273.

TscHuby, R.H., C.L. Pitumore, C.J. ORTH, J.S. GiLmorE, and J.D. KnicHt. 1984. Disruption of the terrestrial plant ecosystem

at the Cretaceous-Tertiary Boundary, western interior. Science 255:1030-1032.

TSUMURA, Y., K. YosHIMURA, N. Tomaru, and K. Onga. 1995. Molecular phylogeny of conifers using RFLP analysis of

PCR-amplified specific chloroplast genes. Theoret. Applied Genet. 91:1222- 1236.

Verrch, J. and Sons. 1881. A manual of the Coniferae. Published by the authors, Kings Road, Chelsea.

Wang, Q., D.L. Di.cHER, X-v. Zhu, Y-L. ZHou, and T.A. Lott. 2006. Fruit and leaflets of Wisteria (Leguminosae, Papilionoi-

deae) from the Miocene of Shandong Province, eastern China. Int. J. Plant Sci. 167:1061-1074.

WANG, T., Y-J. Su, J-M. ZHU, G-K. FAN, and C. HUANG. 2000. RAPD Analyses of Taxaceae and it related taxa. Acta Sci.

Nat. Univ. Sunyatseni 39:129-130.

Warp, PD., J.M. HurTADO, J.L. Kirscuvink, and K.L. Verosus. 1997. Measurements of the Cretaceous paleolatitude of

Vancouver Island: Consistent with the Baja-British Columbia Hypothesis. Science 277:1642-1645.

WHITTAKER, R.E. 1961. Vegetation history of the pacific coast states and the “central” significance of the Klamath

Region. Madrono 16:5-21.

WILKINSON, R.C., J.W. Hanover, J.W. WricHT, and R.H. Flake. 1971. Genetic variation in the monoterpene composition

of white spruce. For. Sci. 17:83-90.

Worre, J.A. 1969. Neogene floristic and vegetational history of the Pacific Northwest. Madroño 20:83-110.

Wore, J.A. 1975. Some aspects of plant geography of the northern hemisphere during the late Cretaceous and

Tertiary. Ann. Missouri Bot. Gard. 62:264-279.

Woop, C.E., Jk. 1972. Morphology and phytogeography: The classical approach to the study of disjunctions. Ann.

Missouri Bot. Gard. 59:107- 124.

XIANG, Q-y., S.R. MANCHESTER, D.T. THomas, W. ZHANG, and C. Fan. 2005. Phylogeny, biogeography, and molecular dating

of Cornelian cherries (Cornus, Cornaceae): Tracking Tertiary plant migration. Evolution 59:1685-1700.

XIANG, Q-v., D.T. THomas, W. ZHANG, S.R. MANCHESTER, and Z. Murrett. 2006. Species level phylogeny of the genus Cor-

nus (Cornaceae) based on molecular and morphological evidence—implications for taxonomy and Tertiary

intercontinental migration. Taxon 55:9-30.

ZHENGYI, W. and W. Sucone. 1998. A proposal for a new floristic kingdom (realm). In: Z. Aoluo and W. Sugono,

eds. Proceedings of the First International Symposium on Floristic characteristics and diversity of East Asian

plants, July 25-27, 1996, Kunming, Yunnan. China Higher Education Press, Beijing and Springer-Verlag Berlin

Heidelberg. Pp. 3-42.

NOTES ON THE GAYLUSSACIA DUMOSA COMPLEX (ERICACEAE)

Bruce A. Sorrie Alan S. Weakley

University of North Carolina Herbarium University of North Carolina Herbarium

North Carolina Botanical Garden, CB 3280 North Carolina Botanical Garden, CB 3280

Chapel Hill, North Carolina 27599-3280, U.S.A. Chapel Hill, North Carolina 27599-3280, U.S.A.

ABSTRACT

Four taxa in the Gaylussacia dumosa complex are reviewed, using morphologic and habitat characters. Most morphologic characters

overlap to some degree, but all taxa are readily separable by using character suites. Gaylussacia dumosa is widespread; the other three

taxa have relatively narrow ranges and are separated geographically, but each is partly sympatric with G. dumosa. There is no overlap

in habitat; syntopy is unknown. We recognize all four of these entities as full species: G. dumosa (Andrews) Torr. & A. Gray, G. mosieri

Small, G. orocola (Small) Camp, and Gaylussacia bigeloviana (Fern.) Sorrie & Weakley comb. nov.

RESUMEN

Se revisan cuatro taxa del complejo Gaylussacia dumosa, usando caracteres morfológicos y del hábitat. La mayoría de los caracteres

morfológicos tienen algún grado solapamiento, pero todos los taxa son fácilmente separables usando series de caracteres. Gaylussacia

dumosa es una planta comün; los otros tres taxa tienen areales relativamente pequefios y están separados geográficamente, pero todos

son parcialmente simpátricos con G. dumosa. No hay solapamiento en el hábitat; la sintopía es desconocida. Reconocemos a las cuatro

entidades com pecies: G. dumosa (Andrews) Torr. & A. Gray, G. mosieri Small, G. orocola (Small) Camp, y Gaylussacia bigeloviana

(Fern.) Sorrie & Weakley comb. nov.

INTRODUCTION

The Gaylussacia dumosa complex includes four related taxa of dwarf to one-meter tall shrubs of eastern North

America. These taxa may be distinguished from other members of the genus by the persistent inflorescence

bracts longer than the pedicels (vs. early-deciduous and shorter than the pedicels) and by stipitate-glandular

sepals, pedicels, bracts, and leaves (vs. sessile-glandular). Gaylussacia dumosa (Andrews) Torr. & A. Gray was

described in 1843 (Torrey & Gray 1843); G. dumosa var. bigeloviana in 1911 (Fernald 1911); G. mosieri Small

in 1927 (Small 1927), originally named G. dumosa var. hirtella Chapman (Chapman 1860); and G. orocola

(Small) Camp in 1935, originally described as Lasiococcus orocola Small (Small 1933). While the first three

taxa have been recognized by many authors, the very locally distributed G. orocola has met with less general

acceptance and has often been synonymized within G. dumosa by authors of floristic treatments.

Camp (1935, 1941) recognized Gaylussacia dumosa (including “bigeloviana” without rank), G. mosieri,

and G. orocola at species level. In his monograph of the genus Gaylussacia, Sleumer (1967) followed Camp's

taxonomy, using morphologic characters. Authors of northeastern floras, such as Fernald (1950) and Gleason

and Cronquist (1990), recognized bigeloviana as a variety within G. dumosa. Southeastern authors, such as

Radford, Ahles, and Bell (1968), included var. bigeloviana as a synonym of G. dumosa, if they mentioned it at

all. Radford, Ahles, and Bell also synonymized “orocola” without rank within G. dumosa. In the most recent

study, Floyd (2002) analyzed DNA and morphologic characters genus-wide. She recognized three species

within the G. dumosa group: G. dumosa, G. mosieri, and G. orocola. However, she did not collect specimens

of G. orocola for DNA sampling. Also, Floyd apparently elected not to recognize var. bigeloviana; she did not

discuss it and it is unclear if she included specimens of that variety within her analyses of G. dumosa. These

omissions are unfortunate, for the inclusion of var. bigeloviana in her work may have helped to resolve its

taxonomic position, and the absence of DNA data from G. orocola weakens her conclusion that it stands

apart from G. dumosa and G. mosieri.

Overall, the G. dumosa complex ranges from Newfoundland south to Florida and west to Louisiana,

inland to the Appalachian Mountains and Cumberland Plateau. The distribution and habitats of G. mosieri

and G. dumosa have been fairly well worked out, but problems have remained regarding G. orocola, due to

J. Bot. Res. Inst. Texas 1(1): 333 — 344. 2007

334 Journal of the Botanical R h Institute of Texas 1(1)

a paucity of specimens and due to imprecise knowledge of the taxonomic limits between it and G. dumosa

var. bigeloviana. Problems also have remained between G. dumosa var. bigeloviana and var. dumosa, because of

imprecisely described taxonomic limits and alleged hybridization within a broad area of sympatry (Fernald

1950; Gleason 1952; Gleason & Cronquist 1990). In this paper we set forth criteria for distinguishing these

taxa, present a taxonomy of the group, and provide county-level distribution maps.

METHODS

We examined over four hundred herbarium specimens from CLEMS, DUKE, IBE, NCSC, NCU, USCH, VA,

VPI, and WILLI, and selected specimens from GH. We assessed morphological characters that have been

traditionally used in Gaylussacia, such as glandular hairs on the calyx, as well as new characters, such as

corolla length. Five measurements were made of each character per specimen, from at least ten specimens

across the range of each taxon. The distribution maps were prepared from herbarium specimens plus records

from the following sources: Alabama—Clark (1971), Florida—atlas of Florida vascular plants (http://www.

plantatlas.usf.edu), Georgia—Duncan and Brittain (1966) and Jones and Coile (1988), Tennessee— Chester et

al. (1997). Maps of Delaware, Maryland, New Jersey, New York, and Pennsylvania were prepared from speci-

mens plus data from state Natural Heritage Program botanists (see discussion under Distribution below).

RESULTS AND DISCUSSION

Distinguishing characters

Our analyses revealed several useful morphological characters (Table 1). These are: plant height, corolla

length, anther length, density and length of glandular hairs on the hypanthium, density and length of non-

glandular hairs on the leaf margin, and presence of sessile glands on the upper leaf surface. Habitat, when

accurately described on specimen labels, is a valuable distinguishing character. Other characters have been

used by various authors, but are not utilized here. For example, the more-or-less virgate and relatively few

branches of G. dumosa var. dumosa (vs. numerous spreading branches in the other three taxa), while useful

in the field, can be difficult to apply to herbarium specimens. Small’s description of G. orocola states that

the leaves possess stellate hairs; this is a unique character when present, but we found such hairs to be so

sparse (completely lacking on many leaves) that its use as an identification character was untenable. Size

of mature fruit may be a useful character in living plants, but the squashed fruits on dried specimens are

highly variable in dimensions.

Table 1 compares morphological characters of the four members of the Gaylussacia dumosa complex.

We briefly discuss each character.

1. Plant height. Gaylussacia dumosa var. dumosa is the only dwarf shrub of the group, seldom exceeding 0.3

m. The other three taxa usually exceed 0.5 m and may reach 1 m tall. However, var. bigeloviana may

DRM overlap var. dumosa in height, and presumably short-statured plants have caused identifica-

on problems in the purported area of overlap from Virginia to New York.

2: ON length. Gaylussacia mosieri and var. bigeloviana have distinctly longer corollas than the other two

taxa. G. orocola is unique in combining tall plant height with small corolla and anther size.

3. Anther length. This character follows that of corolla length: two taxa with long anthers and two with short anthers.

4. Hypanthium glandular hairs. The greater density and length of glandular hairs on G. mosieri (1.0-1.5 mm

twice as long as in other taxa) are obvious on dried specimens as well as on living plants. G. dumosa

var. dumosa has the least dense and shortest hairs (0.2-0.3 mm), although some individuals may be

difficult to distinguish from var. bigeloviana and G. orocola (0.3-0.5 mm).

5. Leaf margin hairs (non-glandular). Gaylussacia orocola and var. bigeloviana clearly have denser marginal hairs

than the other two taxa: 6-9 hairs per mm of leaf margin and 7—10 hairs, respectively, vs. 1-2 hairs per

mm in G. mosieri and 2—5 hairs per mm in G. dumosa. Leaf margin hairs are nearly or entirely absent

in many G. dumosa and G. mosieri plants.

6. Sessile glands on upper leaf surface. G. mosieri is unique in lacking them, while the other three taxa usually

have large numbers.

Taxonomy

Based on original morphological and ecological analyses, we recognize all four taxa at species rank. In doing

Sorrie and Weakley, Gaylussacia dumosa complex 335

TABLE 1. Comparison of characters among four North American taxa of Gaylussacia.

G. mosieri G. bigeloviana G. orocola G. dumosa

Plant height 0.5-1.0(-1.5) m (0.2-)0.4-1.0 m 0.4-1.0m 0.1-0.3(-04) m

Corolla length 7.0-8.5 mm 6.5-8.0 mm 5.5-6.5 59-95

mean 7.41 mm mean 7.06 mean 5.96 mean 5.78

SD = 0.80 SD = 0.46 SD = 0.47 Sp

Anther length 3.3-4.3 mm 3.2-42 2.5-3.2 2.8-3.1

mean 3.88 mm mean 3.73 mean 2.85 mean 3.02

SD =0.28 SD=035 SD=0.26 SEX 07

Density and length very dense; dense; 0.3-0.5 moderate to moderate; 0.2-0.3

of glandular hairs 1.0-1.5 mm dense; 0.3-0.5

on hypanthium

Density and length

of non-glandular

hairs on leaf margin

1-2 per mm of leaf margin;

up to 0.5 mm long

7-10 per mm; up

to 0.3 mm long

6-9 per mm; up

to 0.3 mm long

2-5 per mm; up

to 0.3 mm

long; often absent

Sessile glands on none numerous numerous numerous;

upper leaf surface sometimes

absent

Habitat seepage bogs, wet flat peat bogs, boggy montane sphagnous xeric to dry

woods, baygalls, ecotones cedar-maple bogs, seepage over pine-oak

of streamheads swamps, granite uplands and

peat-based sandhills, dry

pocosins within to wet pine

Carolina bays

savannas and

flatwoods, oak

barrens,

oak heaths

so, we parallel the work of Luteyn et al. (1996), who treated the three members of the Gaylussacia frondosa

(L.) Torr. & A. Gray group at species rank. In the G. frondosa group, there is a relatively widespread coastal

plain/piedmont species plus two southeastern coastal plain endemics. In the case of the G. dumosa group,

there is one widespread “core” species, G. dumosa, with three other species that, while well separated from

each other, are partially sympatric with G. dumosa (Figs. 1-4). Each of the three segregates is sharply sepa-

rated from G. dumosa by habitat, and in the case of G. orocola, mostly by elevation as well.

We have had little difficulty in identifying herbarium specimens, whether flowering, fruiting, or simply

vegetative, by utilizing a suite of characters (see Table 1 and above discussion) that renders each species

unique. Some poorly-collected specimens with vague habitat data can be difficult to identify. By employing

several characters per specimen, identification errors are greatly reduced. Moreover, we refute the assertion

that there is a broad zone of intermediacy (Long Island, New York to Virginia) involving G. dumosa and G.

bigeloviana; instead, our data suggest that identification characters used in previous works were inadequate

or were partly based on misidentified specimens.

The choice to treat the complex as four species deserves some comment. Gaylussacia mosieri has generally

been accorded specific distinction from the other three, while G. dumosa var. dumosa and var. bigeloviana

have generally been regarded as only varietally distinct from one another. Gaylussacia orocola has often been

included in G. dumosa var. dumosa. Two characters appear to separate G. mosieri from the other three taxa:

length and density of hairs on the hypanthium and absence of sessile leaf glands on the upper leaf surface.

However, other characters suggest variable and shifting groupings of the taxa (see Table 1). For example,

corolla size and anther length would ally G. mosieri and G. dumosa var. bigeloviana on the one hand and G.

orocola and G. dumosa var. dumosa on the other. Habitat and stature would suggest that G. dumosa is the outlier

from the other three, being the shortest and occupying the driest habitats. We therefore conclude that the

336 Journal of the Botanical R h Institute of Texas 1(1)

most practical treatment is to consider the four taxa to have equal taxonomic rank. Some taxonomists may

suggest that the taxa should be given varietal status, because they may view the morphological characters

as relatively subtle, because some of the taxa have traditionally been treated at that level or not recognized

at all, and because of the existence of occasional ambiguous herbarium specimens. However, the differences

exhibited between taxa is greater than that usually accorded varieties, which normally involves only one or

two minor morphological traits and often a geographical component (Grant 1981). Here, the four taxa are

differentiated by combinations of habit, corolla and anther size, vestiture, presence/absence of sessile glands

on upper leaf surface, habitat, and range. The Gaylussacia taxa treated here fit the morphological, or taxo-

nomic, species concept (Grant 1981; Stuessy 1990). While we suspect that the four taxa probably represent

distinct evolutionary lineages, thus fitting the concept of phylogenetic species, the data of Floyd (2002) are

inconclusive. Moreover, additional evidence from biochemical and crossing studies are desirable.

Names at species rank exist for three of the taxa; here we raise G. dumosa var. bigeloviana Fernald to

species status.

Gaylussacia bigeloviana (Fernald) Sorrie & Weakley, comb. nov. Basionym: Gaylussacia dumosa (Andrews) Torr. & A.

Gray var. bigeloviana Fernald, Rhodora 13:95-99. 1911. Type: U.S.A. Maine. Washington Co.: heath at base of West Quoddy Head,

Lubec, 26 Jul 1909, M.L. Fernald 2038 with K.M. Wiegand (HoLotyPE: GHU.

KEY

1. Plant < 3 dm high.

2. Corollas 6.5-8.0 mm long, averaging 7.0 mm; anthers 3.2-4.2 mm long, averaging 3.7 mm; glandular hairs

on hypanthium dense, 0.3-0.5 mm long; non-glandular hairs on leaf margin dense; plants usually 4-10

dm high, rarely less than 3 dm; plants of wet boggy habitats; northeastern range, south to DE, disjunct to

NC and SC G. bigeloviana

2. Corollas 5.3-6.5 mm long, averaging 5.8 mm; anthers 2.8-3.1 mm long, averaging 3.0 mm; glandular hairs

on hypanthium moderately dense to relatively sparse, 0.2-0.3 mm long; non-glandular hairs on leaf margin

sparse to absent; plants occasionally up to 4 dm high; plants of xeric to moist habitats; southeastern range,

north to VA (rare MD) and scattered inland to n AL, n GA, C TN, w SC, w NC, and s WV G. dumosa

. Plant » 4 dm high, ranging up to 10 dm, occasionally to 15 dm.

3. Sessile glands on upper leaf surface absent; glandular hairs on hypanthium 1.0-1.5 mm long; East Gulf

Coastal Plain endemic, sw GA-n FL-s AL-s MS-se LA G. mosieri

3. Sessile glands on upper leaf surface numerous; glandular hairs on hypanthium 0.3-0.5 mm long; ranging

from SC northward.

4. Corollas 6.5-8.0 mm long, averaging 7.0 mm; anthers 3.2-4.2 mm long, averaging 3.7 mm; plants of

peat bogs, raised bogs, peat-based pocosins, and Atlantic white cedar-red maple swamps; ranging

from Newf. to DE, and as a rare disjunct in the Coastal Plain of NC and SC G. bigeloviana

4. Corollas 5.5-6.5 mm long, averaging 6.0 mm; anthers 2.5-3.2 mm long, averaging 2.9 mm; plants of

montane bogs and seepage over rock; rare endemic of southern Appalachians of w NC G. orocola

Habitat

Gaylussacia dumosa normally inhabits much drier sites than the other three species. It is most abundant in

xeric to mesic pine-oak sandhills, pine-oak-hickory woodlands, and oak barrens, but also occurs in moist

to seasonally wet longleaf pine savannas and flatwoods.

Gaylussacia mosieri inhabits seepage bogs (often called hillside bogs), margins of streamheads and

baygalls (often with Atlantic white cedar, Chamaecyparis thyoides (L.) B.S.P.), and wet pine flatwoods. These

seepages are minerotrophic and do not accumulate peat; therefore they are best termed poor fens. Gaylus-

sacia mosieri may occur in disturbed habitats (roadside scrapes, borrow pits) that superficially appear dry,

but which are underlain by a claypan.

From Delaware northward, Gaylussacia bigeloviana inhabits peat bogs (including ombrotrophic raised

bogs), sphagnum-shrub bogs, and boggy red maple (Acer rubrum L.)-Atlantic white cedar swamps. It may also

occur in disturbed habitats (roadside scrapes, borrow pits) that superficially appear dry at some seasons, but

which are underlain by high water tables. In North Carolina, G. bigeloviana occurs in several large pocosins,

which are peat-based ombrotrophic bogs dominated by ericaceous shrubs and scattered pond pines (Pinus

serotina Michx.). These pocosins occur in the outer coastal plain within Carolina bay depressions and in

Sorrie and Weakley, Gaylussacia dumosa complex 337

extensive interstream flats (Weakley & Schafale 1992). In South Carolina, G. bigeloviana inhabits a seepage

wetland dominated by Atlantic white cedar.

The primary habitat of G. orocola is peaty montane bogs at moderate elevations in the southern por-

tion of the Appalachians, notably the bogs of the East Flat Rock area, Henderson and southern Buncombe

counties, North Carolina; these wetlands have been largely destroyed and few remnants remain (Weakley

& Schafale 1994). Specimen label data suggests that G. orocola may also occur in seepage over sloping

exposures of granitic rock. These bogs harbor endemic taxa as well as disjuncts from the Coastal Plain.

Among the endemics are Sarracenia jonesii Wherry and Sarracenia purpurea Linnaeus var. montana Schnell &

Dietermann. Coastal plain disjuncts include Chamaedaphne calyculata (L.) Moench, Myrica gale L., Helonias

bullata L., Juncus caesariensis Coville, and Eriocaulon decangulare L.

Distribution

Gaylussacia mosieri is endemic to the East Gulf Coastal Plain from Coffee County, Georgia, and Taylor

County, Florida, west to Tangipahoa Parish, Louisiana (Fig. 1). One outlying record is from Duval County,

Florida (Curtiss 1660 GH, mixed sheet with G. dumosa). All populations occur within 250 km of the Gulf

of Mexico.

Gaylussacia orocola is endemic to the Southern Appalachian Mountains of western North Carolina, in

Buncombe, Henderson, Jackson, Macon, and Transylvania Counties (Fig. 2, Appendix 2). This area supports

a concentration of “southern Appalachian bogs,” many now altered or destroyed.

Gaylussacia bigeloviana is distributed on the Atlantic seaboard from Newf 1 south to Delaware, dis-

junct to North and South Carolina (Fig. 3). Extreme inland records—maximum 120 km from saltwater—are

in York County, Pennsylvania, and Prince George's Counties, Maryland, but these are near Chesapeake Bay,

a major estuary of the ocean. Specimens at USCH collected from Atlantic white cedar swamps in Lexington

County in central South Carolina, were annotated by Wilbur and Whitehead to G. dumosa var. bigeloviana;

we concur. The specimens have unusually large leaves (ranging from 1x2.5 cm to 2x5 cm), possibly a result

of growing in shady conditions. Corolla length, anther length, and leaf margin hair density are typical for

bigeloviana, but hypanthium glandular hairs are longer than usual, ranging from 0.6-1.0 mm.

Gaylussacia dumosa occurs from Virginia and West Virginia to south Florida, west to East Feliciana Par-

ish, Louisiana (Fig. 4). We have seen one specimen from Maryland (cited below). Although predominantly a

species of the coastal plain, there are many inland records from the piedmont and even montane provinces

of northern Alabama, central Tennessee, etc. We list selected inland records in Appendix 1. Various manu-

als have ascribed a range north to Long Island, New York, but we have seen only one correctly identified

specimen of G. dumosa from north of Virginia. Fernald (1911, 1950) and Gleason (1952) suggest that there is

much intermediacy between dumosa and bigeloviana in the region from Long Island to Virginia. For example,

Gleason (1952) states that “Intermediate plants are plentiful between Va. and Long Island.” In contrast, we

have observed virtually no intermediacy in specimens from this region. Here we discuss the status of G.

dumosa in these states.

New York. Mitchell and Tucker (1997) synonymized “bigeloviana” within G. dumosa without discussion;

the inclusive G. dumosa has been documented only on Long Island and Staten Island. Stephen Young of the

New York Natural Heritage Program has observed and collected only G. bigeloviana, all in boggy habitats

(pers. comm .).

New Jersey. The inclusive G. dumosa has been documented from Monmouth County south to Cape

May and Cumberland Counties (Stone 1911). Stone stated that “I fail to distinguish the variety bigeloviana,

proposed by Prof. Fernald..." He could hardly come to another conclusion, since there is no verified speci-

men of G. dumosa sensu stricto from New Jersey; all specimens we have seen are G. bigeloviana. David Snyder

of the New Jersey Natural Heritage Program has observed and collected only G. bigeloviana, all in boggy

habitats within the Pine Barrens (pers. comm .).

Pennsylvania. Rhoads and Klein (1993) listed G. dumosa without synonymy and mapped it in Lan-

caster, Montgomery, Northampton, and York Counties. The habitat is given as *moist, acidic woods and

, arg

BEI ALO

BEES na ANA

O AO

p EP [d p nU op".

LES a, Hd PSI UT ra

EEE u l H EEH ra ag S NY

AA TH HG NR MSS S AY QA

o tee

ASE DAA RA

pup ECCLE EPA ETE LA ON)

EM AL es

| TAL POSU le ER CIERRE N

ESO SELECT CERES OS

<TC PAT eE EASTERN

VE EBENEN

CESSÉ VC ce eens

A A e ene COON

DES ROB SU AS

HU LE] AA HG PRISES

DEAR CELERE ENTRE LS

UTE E E P EE leah EE

NO N is oe eA

EDEN t

MN J j is MON

ae)

"uL MM wry

Fic. 1. County |

Sorrie and Weakley, Gaylussacia dumosa complex

oder

ere C

do AS

¡de qa aN:

A f | [PISS

| CS PAL RPT A,

mat ae AS

A OSO TON,

E OS O

ae A LG y

EE XXL SAL E

pass TAA ANA AA NN

sai NN AN rs

i DESV

SEL oC UTNE CX

COT TS POETS

a PROS

TL +

RA 0.

339

409—7|

309—7

Fic. 2. County level

SPA

EE :

HED

ESA

ien

uut

-—

He VE

e n]

TA EN MER

Journal of the Botanical Research Institute of Texas 1(1)

ee 1 j

SAO

ps en)

A 4

FON

LOE

zg E

ETE

ETT

oe MN

MES Sa p

LIN

CALM

esta

NAb,

LIL)

gro

xx D

PRH

Hee

Soa

Sp C

Buy}

om wae

wy, var PAS

NE >

FAR

NA ae

E SPOR MR |

aie

Fic. 3. County level distrik

Sorrie and Weakley, Gaylussacia dumosa complex

d OS i , ie CIA

Dm oen COH Een

rU one eL d]

we GaN Ge. n

VO GA OSC oy

EUN

MES

Fic. 4. County level d

342 Journal of the Botanical R h Institute of Texas 1(1)

swamps.” Small (1894) also reported Pennsylvania plants from wet habitats: "At the Pennsylvania localities

the plant grows in swamps, and at Smithville [Lancaster County] is actually in the water..." We believe that

all Pennsylvania records refer to G. bigeloviana.

Maryland. Brown and Brown (1992) listed G. dumosa without synonymy and state that it inhabits *Moist

to dry, sandy soils of the Coastal Zone; recorded from the Mountain Zone by Shreve.” The text description

appears to be a composite of both dumosa and bigeloviana. A specimen from Baltimore County—glade of

transmission line east of Pulaski Highway, Baltars 1824 (DUKE)—is G. dumosa. The Maryland Native Plant

Society website (www.mdflora.org/survey. data/] MPMontCoPlantsAddendum.html) lists old records from

Takoma (Montgomery County) and Powdermill Bogs (Prince George's County), adjacent to the District

of Columbia; no variety is indicated. In May of 2006, two specimens from Prince George's County were

examined by Chris Frye of the Maryland Natural Heritage Program and verified as G. bigeloviana: Suitland

Bog, 10 June 1966, Mazzeo and Dudley 1394 (NA); Airport Bog, 13 July 1945, Hermann 11547 (NA). We

believe that both G. dumosa and G. bigeloviana occur in Maryland but are very rare.

District of Columbia. We have seen one specimen apparently from this region: low woods, Steele 119

(DUKE); label pre-printed with “Plants of Washington D.C. and Vicinity.” It is G. bigeloviana due to plant

height and dense non-glandular hairs on leaf margins.

Delaware. Tatnall's (1946) species account is ambiguous, but it appears that he listed records for both

taxa from New Castle and Sussex Counties. McAvoy and Bennett (2001) listed only bigeloviana—from the

same two counties—and gave its habitat as Atlantic white cedar swamps. It is historical in the state. We

believe that all Delaware records are G. bigeloviana.

Virginia. The online Atlas of the Virginia Flora (http://www.biol vt.edu/digital_atlas) maps records of

G. dumosa sensu lato from the coastal plain plus three montane counties: Augusta, Carroll, and Page. The

Carroll County record is erroneous (T. Wieboldt pers. comm.). Carr (1938) cited specimens of G. dumosa

var. bigeloviana from two locations in Augusta County; we have examined Carr 409 (GH, VA) and it is G.

dumosa. We have not located Carr 138 (supposedly at VA), nor the Page County specimen. Nonetheless, we

believe that all current Virginia records are G. dumosa but believe that G. bigeloviana may yet be found in

the Dismal Swamp or a similar habitat in the southeastern sector of the state.

APPENDIX. I

SELECTED INLAND RECORDS OF GAYEUSSACIA DUMOSA SENSU STRICTO

There are many specimens and literature reports from the piedmont region ofVirginia, the Carolinas, Georgia, etc.; we will not

repeat them here. Instead, we focus on records from montane regions. ALABAMA. Clay Co.: Emory's Gap, 2000 ft, specimen

at NCU. DeKalb Co.: Little River Canyon parkway, specimen at IBE. The draft Atlas of Alabama Flora maps G. dumosa in several

counties in the hill country of east-central Alabama and N to Cullman, DeKalb, and Jackson Cos. GEORGIA. Bartow Co.:

specimens at FSU and NCU. Rabun Co.: sandy slopes of Thomas Bald, 2500-3000 ft, reported by Small (1894); Rock Mountain,

vicinity of Tallulah Falls, A.B. Seymour 110 (DUKE). NORTH CAROLINA. Catawba Co.: hillside near Hickory, elev. nearly 2000 ft,

reported by Small (1894). Macon Co.: Satula summit, Highlands, T.G. Harbison s.n. (NCU); top of Mt. Satulah, M.B. Wilson 1860

(DUKE) [this is 4700 ft]. Polk Co.: dry ground, Tryon, J.R. Churchill s.n. (GH). Transylvania Co.: Horsepasture Gorge, pine woods,

2000+ ft, C.L. Rogers 61341a (NCU). SOUTH CAROLINA. Oconee Co.: several collections at CLEMS, NCU, USCH, from relatively

low elevations in blackjack oak woods, xeric mixed oak woods, dry rocky slopes up to 1200 ft. TENNESSEE. Coffee Co.: Tul-

lahoma, 1070 ft, H.K. Svenson 10091 (DUKE, FSU, IBE). The Atlas of Tennessee Vascular Plants (Chester et al. 1997) maps it also

in Bledsoe, Cannon, Franklin, and Van Buren cos., all on the Cumberland Plateau. VIRGINIA. Augusta Co.: Shenandoah

Acres, vicinity of Stuarts Draft, L.G. Carr 409 (GH, VA). WEST VIRGINIA. Raleigh Co.: Flat Top Mountain. This record is discussed

in detail by Strausbaugh and Core (1977) and is from a dry habitat with other species of coastal plain affinity. Harmon et al.

(2007) map it also in Nicholas and Summers cos.

APPENDIX 2

RECORDS OF GAYLUSSACIA OROCOLA

NORTH CAROLINA. Buncombe Co.: swampy places, Biltmore, 25 May 1896, no collector (NCU), orig. det. dumosa. Henderson

Co.: King Creek Bog, end of Mine Gap Road, montane sphagnum bog with dense woody vegetation and small openings,

uncommon, 21 May 1993, B.A. Sorrie 7306 with A.S. Weakley, B. Van Eerden, M.J. Russo (NCU); near Brickton. n.d., WW. Ashe s.n.

(NCU); edge of Devil's Fork swamp, 2.5 mi E of Hendersonville, 18 Jun 1947, G.W. McDowell 408 (DUKE); in swamps near East

Sorrie and Weakley, Gaylussacia dumosa complex 343

Flat Rock, J.K. Small (NY), Tyee of Lasiococcus orocola Small, cited in Sleumer (1967); East Flat Rock, EJ. Alexander (NY), cited

in Sleumer (1967); Flat Rock, 30 May 1886, E.R. Memminger s.n. (NCU); in bog 1 1/2 mi S of East Flat Rock, on rte. 176, 10 Oct

1937, W.C. Coker and party (NCU, 2 sheets); in a bog at East Flat Rock, near Hoot's Nursery, 6 Jun 1936, D.S. Correll 5143 with

H.L. Blomquist and K.H. Garren (DUKE); Hoot's Swamp, D. Samson 719 (NY), cited in Sleumer (1967). Jackson Co.: very top of

Big Yellow Mountain, plants taller than on coast, 21 Aug 1936, W.C. Coker s.n. (NCU). Macon Co.: Satulah Mtn., Highlands, 21

Jun 1924, WW. Ashe s.n. (NCU). Transylvania Co.: oak-hickory woods on rock outcrop, 2 mi N of Cedar Mt., 2 Jun 1952, A.E.

Radford 6090 (GH, NCU) [we believe there is a mis-labeling here; there are bogs two mi N of Cedar Mountain that support

Sarracenia jonesii, S. purpurea var. montana and Arethusa bulbosa and other associates of G. orocola]; 1 mi NE of Frying Pan

Gap, 23 Sep 1957, O.M. Freeman 57831 (NCU), mixed sheet with G. baccata; by creek in bog behind Pisgah Inn, 24 Jun 1955,

L. Walton 3551 (DUKE).

ACKNOWLEDGMENTS

We wish to thank the curators of the following herbaria for specimen loans or for providing access: CLEMS,

DUKE, GH, IBE, NCSC, NCU, USCH, VA, VPI, and WILLI. Botanists from several Natural Heritage Programs,

cited above, provided critical information regarding species within their states. Two anonymous reviewers

considerably improved the manuscript.

REFERENCES

Camp, W.H. 1935. Studies in the Ericales. |. The genus Gaylussacia in North America north of Mexico. Bull. Torrey

Bot. Club. 62:129-132.

CaMP, W.H. 1941. Studies in the Ericales. A review of the North American Gaylussacieae; with remarks on the

origin and migration of the group. Bull. Torrey Bot. Club. 68:531-551.

Carr, L.G. 1938. Further notes on coastal floral elements in the bogs of Augusta County, Virginia. Rhodora

40:86-93.

CHAPMAN, A.W. 1860. Flora of the southeastern United States. American Book Company, New York, N.Y.

CHESTER, E.W., B.E. Worronp, and R. KraL. 1997. Atlas of Tennessee vascular plants. Vol. 2. Center for Field Biology,

Austin Peay State University, Clarksville, TN.

CLARK, R.C. 1971. The woody plants of Alabama. Ann. Missouri Bot. Gard. 58:99-242.

Duncan, W.H. and N.E. Brittain. 1966. The genus Gaylussacia (Ericaceae) in Georgia. Georgia Acad. Sci. 24:13-26.

FERNALD, M.L. 1911. The northern variety of Gaylussacia dumosa. Rhodora 13:95-99.

FERNALD, M.L. 1950. Gray's manual of botany. 8th edition. American Book Company, New York, N.Y.

FLovo, JW. 2002. Phylogenetic and biogeographic patterns in Gaylussacia (Ericaceae) based on morphological,

nuclear DNA, and chloroplast DNA variation. Syst. Bot. 27:99-115.

GLEASON, H.A. 1952. The new Britton & Brown illustrated flora of the northeastern United States and adjacent

Canada. New York Botanical Garden, Bronx.

GLEASON, H.A. and A. CRONQUIST. 1990. Manual of vascular plants of northeastern United States and adjacent Canada.

2"? edition. New York Botanical Garden, Bronx.

Grant, V. 1981. Plant speciation. Second edition. Columbia Univ. Press, New York, N.Y.

Harmon, PJ., D. Ford-Wentz, and W. GnarroN. 2007. Checklist and atlas of the vascular flora of West Virginia. Wildlife

Diversity Program, Elkins, W.V.

Jones, S.B., Jr. and N.C. Cole. 1988. The distribution of the vascular flora of Georgia. Department of Botany, Univ.

of Georgia, Athens.

LUTEYN, J.L., W.S. Jubb, S.P. VANDER KLOET, L.J. Dorr, G.D. WALLACE, K.A. Kron, P.F. Stevens, and S.E. CLEMANTS. 1996. Ericaceae

of the southeastern United States. Castanea 61:101-144.

McAvoy, W.A. and K.A. Bennerr. 2001. The flora of Delaware; an annotated checklist. Delaware Natural Heritage

Program, Smyrna.

MircHELL, R.S. and G.C. Tucker. 1997. Revised checklist of New York state plants. Bulletin No. 490, New York State

Museum, Albany.

RADFORD, A.E., H.A. Ahles, and C.R. Bett. 1968. Manual of the vascular plants of the Carolinas. University of North

Carolina Press, Chapel Hill.

344 Journal of the Botanical R h Institute of Texas 1(1)

RHoADs, A.F. and W.M. Klew, Jk. 1993. The vascular flora of Pennsylvania; annotated checklist and atlas. Amer.

Philosophical Soc., Philadelphia, Pa.

SLEUMER, H. 1967. Die gattung Gaylussacia H.B.K. Bot. Jahrb. Syst. 86:309-384.

SMALL, J.K. 1894. Studies in the botany of the southeastern United States— 1. Bull. Torrey Bot. Club. 21:15-20.

SMALL, J.K. 1927. A new gopherberry from the Gulf states. Torreya 27:36

SMALL, J.K. 1933. Manual of the southeastern flora. Published by the author, New York.

Stone, W. 1911. The plants of southern New Jersey. Ann. Report New Jersey State Mus. for 1910. Trenton.

STRAUSBAUGH, PD. and E.L. Core. 1977. Flora of West Virginia, 2^? edition. Seneca Books, Grantsville, W.V.

Stuessy, T.F. 1990. Plant taxonomy. The systematic evaluation of comparative data. Columbia Univ. Press, New

York, N.Y.

TATNALL, R.R. 1946. Flora of Delaware and the Eastern Shore. Soc. Nat. Hist. Delaware, Wilmington.

WEAKLEY, A.S. and M.P. ScHAFALE. 1992. Classification of pocosins and associated wetlands of the Carolina Coastal

Plain. Wetlands 11:355-375.

Weak.ey, A.S. and M.P. ScHAFALE. 1994. Non-alluvial wetlands of the southern Blue Ridge: diversity in a threatened

ecosystem. Water, Air and Soil Pollution 77:359-383. [Also published in Trettin, C.C., W.M. Aust, and J. Wis-

niewski. 1995. Wetlands of the interior southeastern United States. Kluwer Academic Publishers, Dordrecht,

The Netherlands]

REINSTATEMENT OF SAGITTARIA MACROCARPA (ALISMATACEAE)

Bruce A. Sorrie Brian R. Keener

North Carolina Natural Heritage Program University of West Alabama

3076 Niagara-Carthage Road Department of Biological and

Whispering Pines, North Carolina Lr USA, Environmental Sciences

bruce.sorrieancmail.net Livingston, Alabama 35470, U.S.A.

bkeenereuwa.edu

Adrienne L. Edwards

Illinois Natural History Survey

. Oak St.

Champaign, Illinois 61820-6970, U.S.A.

aedwards@inhs.uiuc.edu

ABSTRACT

$53 [pa P a E £44]

Sagittaria macrocarpa J.G. Sm. is here restored to full species status. A narrow distribution, the rarity of and

morphological similarity to closely related taxa has contributed to this species being overlooked or misidentified for a century. Recent

research in population genetics, molecular sequencing, and morphology support the hypothesis that Sagittaria macrocarpa is distinct

from congeners. In addition, the taxon occupies a unique range of habitats and apparently a limited distribution.

RESUMEN

Se restaura aquí a Sagittaria macrocarpa J.G. Sm. al status de especie. Una distribución restringida, la rareza de los especimenes dispo-

nibles para su estudio, y la semejanza morfológica con taxa relativamente próximos ha contribuido a que esta especie sea pasada por

alto o se identifique erróneamente durante un siglo. Investigaciones recientes en genética de poblaciones, secuenciación molecular, y

morfología apoyan la hipótesis que Sagittaria macrocarpa es diferente de sus congéneres. Además, el taxon ocupa un ünico rango de

hábitats y aparentemente una distribución limitada

INTRODUCTION

Recent field and herbarium studies have documented specimens referable to Sagittaria macrocarpa J.G. Sm.,

a name long misapplied and in synonymy under S. graminea Michx. Morphological and molecular studies

lend significant support for designating S. macrocarpa a distinct taxon. In this paper we provide evidence

for reinstating S. macrocarpa to specific status. Sagittaria macrocarpa J.G. Sm. was described in 1894 (Smith

1894). He cited only the holotype, a specimen collected by M.A. Curtis from *margin of ponds" in South

Carolina (undated, Curtis s.n. MOD. Bogin (1955) cited an isotype of S. macrocarpa at GH; while correctly

identified, this specimen is almost certainly not an isotype (see discussion ahead). Until now, these two are

the only specimens known of S. macrocarpa sensu J.G. Sm.

Small (1909) misapplied the name S. macrocarpa to plants that were later to be named S. fasciculata by

Beal (1960). This misapplication was apparently based on the large achenes of S. macrocarpa and S. fasciculata,

which are markedly larger than other taxa of the S. graminea complex. In a later publication, Small (1933)

again included S. macrocarpa in his text and provided habitat and range statements that are consistent with

S. macrocarpa of J.G. Sm. However, Small’s description does not exclude plants later to be named S. fasciculata

Beal, particularly those with 3—5 whorls of flowers.

Bogin (1955) reduced S. macrocarpa to varietal status, as S. graminea Michx. var. macrocarpa (J.G. Sm.)

Bogin. At the same time, Bogin misapplied the name macrocarpa to plants that later were named S. fasciculata

Beal.

Beal (1960) correctly recognized plants of North Carolina montane bogs and sluggish streams as S.

fasciculata Beal, based on spatulate emersed leaves, broad basal phyllodia, and narrow distribution. He

placed S. macrocarpa in synonymy with S. graminea var. graminea, stating that S. macrocarpa "differs from

J. Bot. Res. Inst. Texas 1(1): 345 — 350. 2007

346 Journal of the Botanical R h Institute of Texas 1(1)

typical S. graminea var. graminea in no way except the size of achenes which are as much as 3 mm in length

in contrast to the usual length of 1.5-2.0 mm of var. graminea. However, specimens of var. graminea collected

by the author [Beal] in the coastal plain of North Carolina exhibit a range in achene size including 3 mm.

Therefore, S. macrocarpa J.G. Sm. must be relegated to synonymy.” During our research, however, we have

not seen any S. graminea with achenes approaching 3 mm. Wooten (1973) and Haynes and Hellquist (2000)

followed Beal (1960) in placing S. macrocarpa in synonymy under S. graminea. In their treatment for Flora of

North America, Haynes and Hellquist (2000) use subspecies rather than variety to designate infraspecific

rank within Sagittaria graminea; we will do so henceforth.

REDISCOVERY AND SPECIMENS

During a botanical inventory of Fort Bragg Military Reservation, North Carolina, specimens initially deter-

mined as Sagittaria isoetiformis J.G. Sm. were collected and discussed by Sorrie et al. (1997). These plants

possess leaves with slender distal blades and/or bladeless (phyllodial) leaves; they have since been anno-

tated to S. macrocarpa based on achene and floral morphology. Other plants collected on Fort Bragg possess

somewhat wider blades and were originally determined to be S. graminea, but based on achene and floral

morphology these specimens also belong to S. macrocarpa. All of these plants occur in blackwater stream-

heads, beaver ponds, and artificial impoundments; none occur in natural depression ponds or Carolina

bays, habitats occupied by S. isoetiformis.

Once the salient characters of S. macrocarpa were determined, the senior author found additional popula-

tions and historical specimens elsewhere in the Sandhills region of North and South Carolina. The following

list represents all known vouchers. ITAM stands for the herbarium at the office of Integrated Training and

Management on Fort Bragg, North Carolina; WEWO stands for the herbarium at Weymouth Woods State

Nature Preserve, North Carolina; the other acronyms follow Index Herbariorum (2006).

NORTH CAROLINA. Hoke Co.: Fort Bragg, Field Branch, boggy portion of formerly impounded blackwater stream, S of firebreak 11

and west of Turkey Road, 18 Sep 1993, Sorrie 7558 (NCU, pers. herb); Fort Bragg, Calf Branch at Southern Pines Road, open streamside

disturbed by siltation, 11 Aug 1993, Sorrie 7545 with B.Van Eerden and R. Kral (pers. herb.); Fort Bragg, Gum Branch at Chicken Road,

margin of blackwater stream, sandy bottom with some mud and clay, 20 Jul 1993, Sorrie 7566 (NCU, pers. herb); same location, 16

Sep 1981, J. Carter III 1497 (WEWO); Fort Bragg, Puppy Creek at Plank Road Cutoff, with Scirpus etuberculatus, Sparganium americanum

Potamogeton diversifolius, 2 Jun 1995, T. Crawford and P. Crutchfield B1329 (ITAM); Fort Bragg, NE shore of McArthur Lake, with Mayaca

aubletii, Eleocharis flavescens, Nymphaea odorata, 29 Sep 1992, P. Crutchfield and M. Jones B652 (ITAM). Hoke/Moore Cos.: Fort Bragg,

Johnsons Millpond, a large beaver pond dotted with peat islets, locally common in shallow pools and where inlet streams enter, with

Sagittaria engelmanniana, Eleocharis robbinsii, 9 Oct 1991, Sorrie 5969 with A. Weakley, J. Carter III (NCU, pers. herb); same location,

sandy substrate at east shore, 18 Aug 1993, Sorrie 7557 (DUKE, GH, NCU, pers. herb); same location, peaty-sandy shore, 16 Aug 2002,

Sorrie 10981 with J. Gray (US, pers. herb.). Moore Co.: Aberdeen Recreation Lake, 4 mi NE of Pinebluff, Chloride 1.6 ppm, 16 Jun 1960,

E. Beal 5592 (DUKE, NCU); same location, southern shores, 30 May 2004, Sorrie 11262 (GH, UNA, NCU); Pinebluff Lake, impounded

blackwater stream with stressed Nyssa biflora at head end, with Panicum hemitomon, Juncus debilis, 28 May 1997, Sorrie 9252 (NCU, pers.

herb.); same location, 21 Jul 2000, Sorrie 10571 (pers. herb.); margin of Powell's Pond at intersection of Fort Bragg-Aberdeen Hwy. and

Saunders Blvd., 28 Jun 1979, T. Howard s.n. (WEWO); submerged just below beaver dam, Weymouth Woods State Nature Preserve, 23

Oct 1965, H. Ahles 63085 (WEWO); Weymouth Woods, partially submerged in old beaver pond, 9 Aug 1976, J. Carter III s.n. (NCU,

WEWO); Weymouth Woods, along Pine Island Trail near crossing of James Creek, 28 Sep 2002, Sorrie 11016 (pers. herb); Moore County

without location, 27-28 Jun 1897, W. Ashe s.n. (NCU). SOUTH CAROLINA. Aiken Co.: infrequent in wet portion of herb-dominated

sandhills seepage bogs in high voltage p g between Vaucluse and Graniteville above Flatrock Pond, Horse Creek, 26 Jun

2001, P. McMillan 5447 with R. McCartney, H. Shealy (CLEMS). Chesterfield Co.: just above hot water discharge of power plant, W margin

of Lake Robinson, 1/2 mi S of Hwy. 346 bridge, 15 Sep 1986, C. Aulbach-Smith 4200 (USCED, 4 4192 (NCU, USCH). County unknown:

“margin of ponds S. Car.", undated, M.A. Curtis s.n. (MO)-Tvrs; “S. Carolina,” undated, M.A. Curtis s.n. (GH).

STATUS OE-CITED ISOTYPE

A second Curtis specimen was cited by Bogin (1955) as an isotype of S. macrocarpa: “S. Carolina”, undated,

M.A. Curtis s.n. (GH). On the sheet J.G. Smith wrote “This plant...is intermediate between S. teres and S.

macrocarpa.” In all respects this specimen matches S. macrocarpa, except for the achenes, which are 2.0 mm

long and 1.5 mm wide. However, the achenes appear to be immature and apparently not fully developed.

The labeling of this specimen differs from the type collection in several particulars and we believe it not to

Sorrie et al., Sagittaria macrocarpa reinstated 347

be an isotype. First, the lettering is printed by machine, not in longhand as on the MO type label. Second,

the paper used for the label is different from that of the MO label. Third, there is no mention of habitat

(*margin of ponds") as on the MO label. For these reasons, we suggest that Curtis collected the GH speci-

men at a different place and date than the type at MO. Based on evidence and discussion above, we believe

this specimen is not an isotype of S. macrocarpa.

GENETIC STUDIES

Edwards and Sharitz (2000) investigated the genetic structure of S. isoetiformis and S. teres S. Wats. One popu-

lation in their study gave divergent results: *We were surprised to discover a population (NCJM) [Johnsons

Millpond, Fort Bragg, North Carolina] that was monomorphic at two loci for alleles not found in any other

population of either S. isoetiformis or S. teres. There are several possible explanations for this phenomenon:

(1) we did not sample enough populations to detect the alleles elsewhere; (2) the NCJM population has been

repeatedly misidentified as S. isoetiformis, but actually represents a known species in the Gramineae section

of the genus; (3) the NCJM population represents a cryptic species that has not been morphologically dis-

tinguished from S. isoetiformis, or (4) the NCJM population is S. isoetiformis, but contains remnant genetic

input from past introgression with another species." As a result of our research, all specimens at Johnsons

Millpond previously assigned to S. isoetiformis have now been correctly assigned to S. macrocarpa; therefore,

choices 2 and 3 of Edwards and Sharitz apply to the present situation.

Keener (2006) investigated most of the genus utilizing sequences of the nuclear ribosomal DNA non-

transcribed spacer (5S-NTS) in a systematic analysis. His findings indicated a rather strong sister relationship

of S. macrocarpa with S. cristata Engelmann, a taxon of the extreme north central United States and south

central Canada. A reasonable hypothesis is yet to be formed as to how these sister taxa came to have such

a large disjunct gap in their distributions.

MORPHOLOGICAL CHARACTERS

The following is a description of S. macrocarpa, expanded on Smith (1894). Plant: perennial, monoecious

herbs, glabrous, emersed from shallow water or stranded during flowering and fruiting, to 40 cm tall.

Rhizomes absent; corms present; stolons often extending from corms but usually broken or lost during

collecting; tubers unknown; roots septate. Leaves: narrowly bladed and petiolate, or phyllodial. Leaves

ascending to erect, (6.5—)8.0—22.2(-28.2) cm long, slender, when petiolate dilated distally into linear blades

(0.8-)1.03.1(+4.0) mm wide, lacking sagittate processes; margins entire; apex acute. Scapes: often 1 but

range up to 4, erect, (11.4-)14.3-27.2(33.8) cm long, exceeding leaves; inflorescences racemose, bearingl-3

nodes; lowest node bearing 1-2 carpellate flowers and 1-2 staminate flowers; upper nodes bearing staminate

flowers; nodal bracts scarious, connate at base for more than 1/4 to 1/3 total length, ovate, tips blunt or acute,

(1.9-)2.1-2.7(-3.0) mm long. Flowers: pedicels ascending; receptacles convex; sepals spreading to recurved,

3.0-3.5 mm long, translucent, shorter than petals; petals 3, white, (4.2—)4.6-6.4(-7.3) mm long, entire. Car-

pellate flowers: carpels numerous, apocarpous, ovules 1, styles terminal. Staminate flowers: stamens 8-10,

filaments distinct, moderately dilated basally, sparsely to fairly densely short-pubescent, 0.4-0.7 mm long,

anthers 0.9-1.0 mm long, yellow. Fruiting heads: 8-10 mm in diameter; achenes obovate, 2.0-2.7 mm x

1.3-2.0 mm, compressed, beaked, adaxially keeled, keel margin entire to scalloped; abaxially slightly keeled,

keel margin entire; faces ridged, ridges horseshoe shaped, margins scalloped to entire; resin canal 1, rarely

2; beak laterally attached, obliquely emerging, 0.5-1.0 mm long. Figure 1 depicts the salient features.

Table 1 compares several key characters among Sagittaria macrocarpa, S. isoetiformis, S. graminea ssp.

graminea, and S. fasciculata. Although no single character uniquely identifies any one of the four taxa, a

number of combinations will do so effectively. The thick rhizome of S. graminea ssp. graminea, when pres-

ent, will positively distinguish that entity from the others; however, Godfrey & Wooten (1979) state that in

young plants (through the first year or so of blooming) the rhizome is not yet well developed.

348

Journal o

fal, Dat

te of Texas 1(1)

ffcL

Fic. 1. Two plants of Sagittari pa (

facial ridge. d. beak. e. secondary facial ridge.

sel

WIU U

PHVU

.f a Pa |

UITOITI SLOTOTIJ,

ith datail nf infl

a. achene keel. b. resin canal. c

Sorrie et al., Sagittaria macrocarpa reinstated

TABLE 1. Comparison of selected morphological characters of Sagittaria

349

rocarpa, S. isoetiformis, S. graminea ssp. graminea

and S. fasciculata. Measurements in millimeters. FNA refers to Haynes & Hellquist (2000); Beal refers to Beal (1960); otherwise,

measurements are from original work. * 2 A. Edwards unpublished data.

macrocarpa isoetiformis graminea fasciculata

Achene length (2.2-)2.4-3.0 1.5-2.0(-2.5) 1.4-2.0 2.5-3.0 FNA

2.5-3.5 Beal

Achene width 1.3-2.1 0.9-1.2(-1.5) 0.9-1.2(-1.5) 1.2-1.5 FNA

1.3-2.0 Beal

Beak length 0.5-0.9 0.2-0.6 0.2-0.4 ca. 05

Achene face resin canal 1(-2), resin canal 2-3, resin canal 1, resin canal 1,

flanked by 2(-3) flanked by 2-3 surrounded by flanked by 2

low ridges low ridges 2 high ridges low ridges

No. of inflorescence 1-3 1-5 1-7 2-5

whorls

Bracts connate up to 2/5 of length at least 1/2 1/3-2/3 1/4

Leaf blade width 0-4.3 «1-22 3.0-15.0 5.0-21.0

Rhizome present no or very slender no or very slender yes, thick no

and thickness

Corms produced no yes* no yes

Stolons present yes or no yes, but no yes

usually not

collected

A KEY TO THE NARROW-LEAVED MEMBERS

OF THE SAGITTARIA GRAMINEA COMPLEX

The following key will serve to identify narrow-leaved members of the Sagittaria graminea group occurring

in the southern Atlantic region of the United States. Sagittaria graminea ssp. weatherbiana (Fernald) R.R.

Haynes & Hellquist is a much coarser plant with blunt-tipped leaves at least 1 cm wide.

1. Leaves phyllodial.

2. Achenes 1.5-2.0(-2.5) mm long; achene with 3 or more facial ridges and 2 or more resin canals; inflore-

cence bracts connate for more than half of entire length

S. isoetiformis

2. Achenes (22-)24-3.0 mm long; achene with 2-3 facial ridges and 1-2 resin canals; inflorescence bracts

connate for less than 4096 of entire length

1. Leaves petiolate.

3. Carpellate pedicels distinctly thicker in cross section than staminate pedicels, recurved in fruit _

S. macrocarpa

__S. platyphylla

3. Carpellate pedicels more-or-less equal to staminate pedicels in diameter, ascending in fruit.

4. Rhizomes coarse; achene with markedly raised facial ridges, forming an elliptical bowl-like structure.

5. Inflorescence racemose

5. Inflorescence paniculate

S. graminea ssp. graminea

S. graminea ssp. chapmanii

4. Rhizomes absent or slender; achene with slightly raised facial ridges (markedly raised in S. graminea).

6. Leaf blades at least 5 mm wide and anthers about 0.6-0.7 mm long

6. Leaf blades less than 4.5 mm wide and anthers about 0.9-1.1 mm long.

7. Achenes (2.2-)2.4-3.0 mm long, beaks 0.5-0.9 mm long

S. fasciculata

S. macrocarpa

7. Achenes 1.4—2.0(-2.5) mm long, beaks 0.2-0.6 mm long.

8. Achenes with markedly raised facial ridges, forming an elliptical bowl-like structure; leaf blades

atleast 3.0 mm wide

8. Achenes with slightly raised facial ridges; leaf blades less than 2.3 mm wide

HABITAT AND DISTRIBUTION

S. graminea ssp. graminea

S. isoetiformis

Sagittaria macrocarpa inhabits very shallow water of beaver ponds, impoundments, slow-moving streamhead

creeks, and occasionally in adjacent wet seepage slopes, all within the longleaf pine (Pinus palustris P. Miller)

350 Journal of the Botanical R h Institute of Texas 1(1)

ecosystem. These are blackwater drainages with dark tannin-stained water, low pH, and low nutrient content.

Substrates are sands or clayey sands of the Middendorf Formation (Upper Cretaceous). Peat mosses (Sphag-

num spp.) are common associates, along with members of Juncus, Eleocharis, Scirpus, Panicum, Dichanthelium,

Lachnanthes, Nymphoides, Nymphaea, Sparganium, Potamogeton, and Mayaca. Ericaceous shrubs are often

prominent around the margins of these wetlands. Within the genus Sagittaria, only S. engelmanniana J.G. Sm.

and S. graminea (rarely) have been found growing with S. macrocarpa, based on personal observations.

To date, all populations occur in the rolling Sandhills Region (inner coastal plain) of North and South

Carolina. Although documented from only four counties in two states, there is abundant potential habitat

in the region as beaver continue to reclaim former range and as humans continue to create streamhead

impoundments.

ACKNOWLEDGMENTS

We wish to thank curators of the following herbaria for providing specimens: CLEMS, DUKE, HUH, MO,

NCSC, NCU, UNA, USCH, Weymouth Woods State Nature Preserve, and Integrated Training and Manage-

ment, Fort Bragg, North Carolina. Staff at the Endangered Species Office of Fort Bragg provided access and

logistics. Staff at Weymouth Woods provided access to collecting locations. Alan Weakley and Carol Ann Mc-

Cormick of NCU provided facilities for morphological study and for securing inter-herbarium loans. Georgia

Minnich prepared the drawings. Robert Kaul and one anonymous reviewer provided helpful suggestions.

REFERENCES

BEAL, E.O. 1960. The Alismataceae of the Carolinas. J. Elisha Mitchell Sci. Soc. 76:68-79.

Boain, C. 1955. Revision of the genus Sagittaria (Alismataceae). Mem. New York Bot. Gard. 9:179-233.

EDWARDS, A.L. and R.R. SHaritz. 2000. Population genetics of two rare perennials in isolated wetlands: Sagittaria

isoetiformis and S. teres (Alismataceae). Amer. J. Bot. 87:1147-1158.

Goprrey, R.K. and J.W. Wooten. 1979. Aquatic and wetland plants of southeastern United States. Vol. 1. University

of Georgia Press, Athens.

Haynes, R.R. and C.B. Hettquist. 2000. Alismataceae. In: Flora of North America Editorial Committee, eds. Flora of

North America North of Mexico. Vol. 22, Magnoliophyta: Alismatidae, Arecidae, Commelinidae (in part), and

Zingiberidae. Oxford University Press, New York.

INDEX HERBARIORUM. 2006. Website: http://sciweb.nybg.org/science2/IndexHerbariorum.asp

KEENER, B.R. 2006. Molecular systematics and revision of the aquatic monocot genus Sagittaria (Alismataceae).

Doctoral dissertion, The University of Alabama, Tuscaloosa.

SMALL, J.K. 1909. Alismaceae. In: N. Amer. Fl. 17:43-62.

SMALL, J.K. 1933. Manual of the southeastern flora. Published by the author, New York.

Sui, J.G. 1894. A revision of the North American species of Sagittaria and Lophotocarpus. Report (Annual) Mis-

souri Bot. Gard. 6:1-38 & 29 pl. Also published in 1895:27-64 & 29 pl.

SonniE, B.A., B, VAN EERDEN, and M.J. Russo. 1997. Noteworthy plants from Fort Bragg and Camp Mackall, North

Carolina. Castanea 62:239-259.

Wooten. JW. 1973. Taxonomy of seven species of Sagittaria from eastern North America. Brittonia 25:64-74.

VARIATION IN PETRADORIA PUMILA (ASTERACEAE: ASTEREAE)

Guy L. Nesom Caleb A. Morse

Botanical Research Institute of Texas R. L. McGregor Herbarium, Division of Botany

509 Pecan Street Natural History Museum and Biodiversity Research Center

Fort Worth, Texas 76102-4060, U.S.A. University of Kansas, 2045 Constant Avenue

Lawrence, Kansas 66047, U.S.A.

ABSTRACT

Populations of Petradoria pumila with linear-filiform leaf blades, primarily in Arizona and northwestern New Mexico, are treated as P.

pumila var. graminea. They are interpreted here to be distinctive variants at the southern of anorth-south cline with the gradient

breaking sharply at the southern end. Formal taxonomic recognition can be maintained for var. graminea, although intermediates are

common and the taxon probably is artificial, especially if enclaves of linear-filiform variant populations in central Utah are identified as

var. graminea. Clusters of populations from Clark, Nye, White Pine, and Elko cos., Nevada, along e western bu of the geographic

Js 1

number. In

range, differ from others of the species (var. pumila and var. graminea) in

9 populations, i5 plants),

disc florets per head range 5-10, averaging 7.0. Over ME rest of the range of the species (n = 125 EN outside of Nevada), disc florets

range 2—4(—5), averaging 3.3. There is no evidence that the high floret number populations in Nevada represent a single lineage.

RESUMEN

Las poblaciones de Petradoria DEN con láminas cies linear-filiformes, de Arizona y noroeste de Nuevo México, se tratan como P.

PEN PI As]

pumila var. graminea. A Sur de una variación clinal Norte-Sur con el gradiente

que se rompe netamente en el extremo Sur. Se RE mantener el reconocimiento taxonómico formal de la var. pee: aunque los

TEN SS

intermedios son comunes y Į enclaves poblaciones variantes linear-filiformes

en Utah central se identifican como var. graminea. Los grupos de DE de Clark, Nye, White Pine, y Elko cos., Nevada, a lo largo

del borde Oeste de su areal geográfico, difieren de otros de la misma especie (var. pumila y var. graminea) en el número de flósculos del

disco. En estas poblaciones (9 poblaciones, 15 plantas), los flósculos del disco por capítulo varían de 5-10, con una media de 7.0. En el

resto del areal de la especie (las plantas n = 125, fuera de Nevada), los flósculos del disco varían de 2-4(-5), con una media de 3.3. No

hay pruebas de que las poblaciones con alto número de flósculos en Nevada representen una línea simple.

The genus Petradoria Nutt. was treated by Anderson (1963) to include two species: P. pumila (Nutt.) Greene

and P. discoidea L.C. Anderson (= Chrysothamnus gramineus H.M Hall). He later reinstated P. discoidea within

Chrysothamnus (Anderson 1983), but molecular evidence subsequently has shown that species to be phylo-

genetically remote from both Chrysothamnus and Petradoria (Beck et al. 2004; Roberts & Urbatsch 2004).

Chrysothamnus gramineus has now been segregated as the monotypic genus Cuniculotinus Urbatsch et al.

(Urbatsch et al. 2005), and Petradoria has reverted to its monotypic status (Urbatsch et al. 2006). Petradoria

pumila is morphologically distinct from Stenotus Nutt., its closest relative, in having many, few-flowered heads

with vertically aligned phyllaries, functionally staminate disc florets, and glabrous cypselae.

Petradoria pumila is a common plant of pine forests, pinyon-juniper woodlands, and shrub communi-

ties in the southwestern U.S.A. The distribution is centered around Utah and includes northern Arizona,

San Bernardino Co., California, western Colorado, southeasternmost Idaho, northwestern New Mexico,

Sweetwater Co., Wyoming, and eastern Nevada (Fig. 1). A single collection from the *Truckee Mts."cited and

mapped by Anderson (1963) as originating from Washoe Co., Nevada (2 May 1868, S. Watson 557, [US]) is

out of range for the species. Two other 1868 collections by Watson, however, both numbered “557,” have

locality data placing them in Elko Co., Nevada. It is probable that the specimen with the “Truckee Mt.” label

also was collected in Elko Co.

In Anderson's monograph, largely followed by Urbatsch et al. (2006), Petradoria pumila comprises two

infraspecific taxa (treated as subspecies by Anderson, as varieties by subsequent authors): var. pumila occurs

over most of the range; var. graminea (Woot. & Standl.) S.L. Welsh is mostly restricted to the southernmost

portion of the range. Urbatsch et al. (2006) distinguished the two taxa by the following contrasts:

1. Leaves usually 1(-3)-nerved, 1-2 mm wide; involucres 1.3-2 mm wide; ray florets usually 1, laminae

0.7-1.5 mm wide; disc florets 2-3 var. graminea

J. Bot. Res. Inst. Texas 1(1): 351 — 356. 2007

352 Journal of the Botanical Research Institute of Texas 1(1)

O O

O me O

XO O

"T" _ @ Olojo

2 OQ >

o PATOS

Og O

Ge O

re O Q O

» " O OQ fo

- OQ Oo

[

> % o

x O

j [NN

> A |]

E M. - O leaf width 1-2 mm

TE I @ leaf width 2.1-4 mm `

i O leaf width 4.112 mm

i trat

Fic. 1. Variation in leaf shape in Petradoria pumila. Linear-| ] plants (var. graminea) d mostly in th t t portion of the

| i thern localities. The typical fi f the species.

range but tł

Y rr vt vd 2

1. Leaves usually 3-5-nerved, 2-12 mm wide; involucres 1.9-3 mm wide; ray florets usually (1-)2-3, laminae

1-2.4 mm wide; disc florets 2-4(-5) var. pumila

The present study reexamines taxonomically significant characters. Our observations generally agree that lin-

ear-leaved variants are concentrated in Arizona and New Mexico although evidence suggests that they do not

represent a single lineage. We also document previously unrecognized variability in disc floret number.

Leaf morphology and other clinal variation

As cited and mapped by Anderson (1963), var. graminea occurs in Mohave, Coconino, and Apache cos.,

Nesom, Variation in Petradoria pumila 353

Arizona, and San Juan and McKinley cos., New Mexico. Using his criteria for identification, we add Navajo

Co., Arizona, and Sanpete and Emery cos., Utah, to the distribution of var. graminea. Welsh et al. (2003,

p. 218) also noted that *a few specimens from Emery and Garfield counties seem to be clearly allied to var.

graminea.” A county-level map of collections showing intermediate-width leaves (Fig. 1), however, sug-

gests that variation in leaf width (and number of veins, which is correlated) is continuous, if intermediates

between var. pumila and var. graminea are considered to have leaves 2.1-4 mm wide. Linear-leaved plants

tend to be concentrated in southern populations but plants of typical morphology and intermediates are

common in the same region. Disjunct enclaves of linear-leaved plants (var. graminea) in central Utah are

hardly distinguishable from southern linear-leaved plants, and it seems likely that populations with linear

leaves have arisen independently.

Anderson (1963) observed that var. pumila occurs at higher elevations where it occurs sympatrically

with var. graminea. We are not able to test this, but our data suggest that there is no overall distinction in

elevation: 20 collections from Arizona in our study referable to var. graminea occur at a range of 5000-8500

ft, averaging 7020 ft; 46 collections from Arizona referable to var. pumila or intermediates between the two

varieties occur at an elevational range of 4600—8500 ft, averaging 6350 ft. Plants of var. pumila have been

collected at a range of (4000—5000-8300(-10,000) ft in Utah, Colorado, and New Mexico. Petradoria pumila

in San Bernadino Co. (and one collection from Tulare Co.), California, occurs at 3500-7000 ft.

Anderson (1963, p. 681) noted with respect to var. pumila that *much of its variation is clinal along the

north-south axis of its range,” these trends illustrated in his figs. 44 and 45. Figure 1 of the present study

indicates that a similar clinal trend is reflected in leaf width. Figure 2 indicates that there are no disconti-

nuities in leaf width in each of four areas within the range of the species.

Anderson's Figure 45 shows geographic variation in involucral width, ray width and number, and disc

floret number—measurements for each of these features decrease in a north-to-south direction. Fig. 44

shows geographic variation in the number of involucral bracts per head; number of bracts slightly increases

southward in var. pumila (overall range 10—21) but bract number in plants identified as var. graminea ranges

11-15. We did not make a detailed analysis of involucral bract number, but whatever differences may exist

do not appear to be significant. We counted 20 bracts in several plants from northern Utah.

In summary, the linear-leaved plants may justifiably be treated without formal rank, since it seems

likely that many of the populations are independently derived, especially if those in Utah are identified as

var. graminea. But a final clinal step in reduction of leaf width gives these plants a distinctive appearance,

and they are concentrated at the southern extremity of the range of the species. The name var. graminea is

available for those who wish to use it, although the taxon probably is largely artificial.

Floret number

Over the range of the species, excluding Nevada, we confirm that numbers of florets per head are essen-

tially as reported for the species by Anderson (1963): ray florets 1-3, disc florets 2-4(—5). We also note that

heads completely lacking ray florets occur at a low frequency, sometimes on plants with rayed heads. In

counts from a total of 125 collections from all states in the range except Nevada, ray florets per head range

(0-)1-2(3, rarely), while disc florets range (1-)2—4(—5). For these same populations, ray florets average 1.4,

while disc florets average 3.3. The modal number of ray florets per head is 1 in Colorado, New Mexico, Utah,

and Arizona. Number of ray florets does not distinguish var. graminea.

In contrast to the normal situation, in a series of populations along the western extremity of the range

in Nevada (Fig. 3; 9 collections, 15 plants), disc florets per head range 5-10, averaging 7.0. Ray florets per

head range 1-3, as elsewhere in the range. In most other Nevada localities (38 collections, 66 plants), disc

florets per head range (123-5, averaging 3.8. Except for slightly broader involucres, the higher-number

plants differ in no other apparent features from the lower-number ones.

Cited here are collections mapped with high numbers of disc florets. Floret | brackets [ray,disc]; multiple plants are

separated by a backslash. NEVADA. Clark Co.: Charleston Mts., Lee Canyon, 8000 ft, 28 Jul 1913, Heller 11015 [2,8/2,6] (DS, MO, NY,

UC, US); Charleston Mts., near Griffith Peak, Windham 97-117 [2,7] (MO). Elko Co.: Angel Lake, ca. 13 km SW of Wells, 2600 m, Lowry

354 Journal of the Botanical R h Institute of Texas 1(1)

1 1.5 2 2.5 3 3.5 4 4.5 5 5.5 6 6.5 7

ARIZ (n=27) 8 5 5 4 2 2 1 0

NMEX (n=30) 4 9 6 4 2 3 0

UTAH (SnJn Co.) (n=33) 0 3 2 3 5 4 2 3 3 6 2 0

IDA-NEV (n=30) 0 2 1 4 3 7 5 3 4 1 1

[2]

£ 10

9 g-

$ sal

3 | N p

9 4- =

[e] w

z 0 T T ES T T T T | T T T

1 715 2 25 3 35 4 45 5 55 6 65 7

Avg. leaf width (mm)

=m ARIZ(n=27) «mNMEX (n-30) UTAH (SnJn Co.) (n=33) =====IDA-NEV (n-30)

Fic > M. . * LÀ | £ Lael FH. s n.a J ° oF T^ 4 [ k. 4 | 4 g * al £L Ia 4 L eal J H L

| 1. Ari (4 ties), New Mexico (5 ties), Utah (San Juan Co.), and Idal IN la (8 ties)

4630 [2,7] (MO); Ruby Mts, N slope of Verdi Peak, 9000 ft, 13 Aug 1941, Mills & Beach 1543 [3,6/2,5/3,5] (BRIT, UC). Nye Co.: between

Haystack Canyon and Mosquito Creek, 8800 ft, 22 Jul 1984, Atwood et al. 10516a [3,9/3,8] (BRY, NY); S flank of Mt. Jefferson, SE side of

Toquima Range, 10,000 ft, 2 Aug 1976, Williams 76-95-7 [1,6/3,5] (CAS, NY, UTC). White Pine Co.: Snake Range, W of Baker, ridge E

of Lehman Creek, 10 Aug 1963, Breedlove 5821 [1,7] (SMU); summit of Cave Mt., 16 mi due E of Ely, 18 Jul 1997, Hess et al. 7624 [1,9/2,7]

(BRIT, MO); Schell Creek Range, ca 1 mi S of Success Summit on NV Rte 486, 8600 ft, 16 Jul 1981, Pinzl 4406 [2,10] (MO).

In the Charleston Mountains of Clark Co. and a few other localities, considerable variation in disc floret num-

ber per head apparently occurs within some populations. Studies of population structure in the Charleston

Mountains and in the Snake Range might reveal whether such variation actually occurs continuously within

populations or whether sampling by collectors included plants from adjacent, genetically different popula-

tions. From the Prospect Ridge area of Eureka Co., other collections have been only of plants with low floret

numbers. Cited below are vouchers for collections with duplicates and multiple plants in which counts of

disc florets per head (1 count per plant) are in both low and high ranges. Clark Co.: Charleston Mts., ridge S

of Deer Creek, 2700 m, 31 Jul 1935, Clokey & Clokey 5644 |1,7/2,6/2,6/2,6/2,5/3,4/2,4/2,4/2,4/1,4/2,3] (LL,

MO, RM, RSA, TEX); Charleston Mts., gravelly wash with juniper and pine, 2270 m, 12 Jul 1936, Clokey 7345

[2,6/1,6/1,5/2,3] (MO, TEX, UC). Eureka Co.: Prospect Ridge, SW part of county, on Relay Station Rd, 8700

ft, 19 Jul 1983, Williams & Tiehm 83-108-4 [2,8/1,5] (NY). White Pine Co.: Snake Range, Mt. Washington, S

slope at head of canyon, 10400 ft, 14 Aug 1964, Holmgren & Reveal 1657 [1,6/2,4/1,4] (NY, TEX).

Vestiture variants

Plants of one population from the Clover Mountains of Lincoln Co., Nevada, produce densely hirsutu-

lous-puberulent stems and leaves (Tiehm & Crisafulli 11415 [BRY, CAS, COLO, NY, ORE, RM, RSA, TEX.

Otherwise, these specimens are similar to specimens referable to var. pumila, which mostly are completely

glabrous. Plants of another population from Lincoln Co. (Deer Lodge,” Train 2521 [MO, UC]) have very

sparsely hirsutulous leaves. Other plants from scattered localities in the range also produce a sparsely hir-

sutulous indument, e.g., Coconino Co., Arizona (Clover 4320 [TEX]; Parker 6188 [LL]); Emery Co. (Hatch s.n.

[UTC]), Garfield Co. (Richards 14962 [GH, UTC]; Foster 5399, BRY]), and Millard Co. (Brooks 20353 [KANU,

RM]; Cottam 5660 [LLD, Utah.

Nesom, Variation in Petradoria pumila 355

> e EN

O disc florets (5—)6—10 > 8

O disc florets 3-5

Fic. 3. Distribution of Petradoria pumila in Nevada. Plants with higl | f disc fl per head occur in clusters along the western boundary of

the species.

£+sha D o ID L

356 Journal of t titute of Texas 1(1)

Chromosome number

Diploids and tetraploids have been reported within Petradoria pumila (Anderson 1963). Diploids are from

Arizona, Nevada, and Utah; tetraploids are from Kane Co., Utah, and Coconino Co., Arizona. No morpho-

logical differences between diploids and tetraploids were reported by Anderson, nor do we observe any.

Taxonomy

Petradoria pumila var. pumila

Petradoria pumila (Nutt.) Greene, Erythea 3:13. 1895. Chrysoma pumila Nutt., Trans. Amer. Philos. Soc., n.s. 7:325. 1840.

Tyee: U.S.A. [Inano. Bear Lake Co.:] “In open situations, on shelving rocks toward the western declivity of the Rocky Mountains"

protologue], 1834, T. Nuttall s.n. (HoLoTYPE: BM!, photo-US!). In June of 1834, Wyeths expedition (with Nuttall as naturalist) was in

Sweetwater Co., Wyoming, but slightly north of the known localities for P pumila. After crossing the Green River, they continued to

Hams Fork in Lincoln Co., Wyoming (Graustein 1967). Leaving Ham’ Fork in early July, they traveled northwestward into Bear Lake

Co., Idaho, and continued toward Soda Springs in Caribou Co., where they stopped from 8-10 July. Because the leaf morphology

of the type specimen is closely matched in recent collections from Bear Lake Co., we believe that Nuttall collected the type there,

at the northernmost point of the range of the species, apparently the only place where he would have encountered it. Anderson

(1963) also concluded that the type collection was made in “southeastern Idaho.” In the description, Nuttall noted “Discal florets

three; rays usually two."

Petradoria pumila var. petiolaris A. Nels., Bull. Torrey Bot. Club 26:482. 1899. Type: U.S.A. WyomINc. Sweetwater Co.: North Vermilion

p , 17 Jul 1897, A. Nelson

3581 (HOLOTYPE: RM!; IsotyPEs: BM!, BRY!, COLO!, GH!, MO!, NY!, UTC!). In the protologue, Nelson noted that “The narrow leaves

and their much greater relative length give the variety a very different aspect, but I think the difference can hardly be considered

Creek, southern portion of the county, abundant on stony hillsides, growing with typical form

specific." The MO isotype has 2 ray and 5 disc florets.

Petradoria pumila var. graminea (Woot. & Standl.) S.L. Welsh, Great Basin Naturalist 43:324. 1983. Solidago

graminea (Woot. & Standl.) Blake, J. Washington Acad. Sci. 21:326. 1931. Petradoria pumila subsp. graminea (Woot. & Standl.) L.C.

Anderson, Trans. Kansas Acad. Sci. 66:682. 1964. Petradoria graminea Woot. & Standl., Contr. U.S. Natl. Herb. 16:183. 1913. Tyre:

U.S.A. New Mexico: “N. Mexico” [as on label], “northwestern New Mexico” [in protologuel, 14 Jul 1883, C.C. Marsh 209 (HOLOTYPE:

US!, internet image, photo-UC!). The type presumably was collected either in San Juan Co. or McKinley Co., New Mexico.

ACKNOWLEDGMENTS

We are grateful for loans (mostly to Morse) from ARIZ, BM, BRY, CAS, COLO, DS, GH, KANU, MO, MONTU,

NY, POM, RM, RSA, SJNM, TEX-LL, US, UTC, and WS and for help from staff at MO and TEX-LL during

study at those herbaria.

REFERENCES

ANDERSON, L.C. 1963[Feb 1964]. Studies on Petradoria (Compositae): anatomy, cytology, taxonomy. Trans. Kansas

Acad. Sci. 66:632-684.

ANDERSON, L.C. 1983. Chrysothamnus eremobius (Asteraceae): a new species from Nevada. Brittonia 35:23-27.

Beck, J.B., G.L. Nesom, PJ. Cate, G.I. Bairo, R.L. Smal, and E.E. ScHILLING. 2004. Is subtribe Solidagininae (Asteraceae)

monophyletic? Taxon 53:691-698.

GRAUSTEIN, J.E. 1967. Thomas Nuttall, naturalist: Explorations in America, 1808-1841. Harvard Univ. Press, Cam-

bridge, Massachusetts.

Roserts, R.P. and L.E. UrsatscH. 2004. Molecular phylogeny of Chrysothamnus (Asteraceae, Astereae) based on

nuclear ribosomal 3’ETS and ITS sequence data. Syst. Bot. 29:199-215.

URBATSCH, L.E., R.P. RogerTs, and K.M. Neuaic. 2005. Cuniculotinus and Lorandersonia, two new genera of Asteraceae:

Astereae and new combinations in Chrysothamnus. Sida 21:1615-1632.

UnBATSCH, L.E., R.P. Roserts, and K.M. Neusic. 2006. Petradoria (Astereae). In: Flora of North America Editorial Com-

mittee, eds. 1993+. Flora of North America North of Mexico. 12+ vols. Oxford University Press, New York and

Oxford. Vol. 21, pp. 171-172.

WELSH, S.L., N.D. Arwoop, S. Gooprich, and L.C. Hicains (eds.). 2003. A Utah flora (Third Edition, revised). Print Services,

Brigham Young University, Provo, Utah.

THE IDENTITY OF CULTIVATED PHELLODENDRON (RUTACEAE)

IN NORTH AMERICA

Jinshuang Ma Anthony R. Brach

Brooklyn Botanic Garden Missouri Botanical Garden

1000 Washington Avenue c/o Harvard University Herbaria

Brooklyn, New York 11225-1099, U.S.A. 22 Divinity Avenue

Cambridge, Massachusetts 02138-2094, U.S.A.

ABSTRACT

The identity and cultivation of the two species of Phellodendron (Rutaceae) in North America are reviewed. Phellodendron amurense is

known to have escaped and naturalized in North America, especially in the northeastern USA. Phellodendron chinense is cultivated in

gardens and arboreta, but has not escaped or naturalized yet. The differences between the two recognized species are provided, and

the variability of the bark is also discussed. In addition, representative accession records of the cultivated specimens of Phellodendron

in gardens and arboreta are provided.

RESUMEN

Se revisa la identidad y cultivo de dos especies de M MN ad (Rutaceae) en Norte América. Phellodendron amurense se sabe que ha

escapado y se ha naturalizado en Norte América, esp | noreste. Phellodendron cl lt jardines y arboreta, pero

aún no se ha escapado o naturali ado. Se aportan] dif i tre las d peci , y también se e la variabilidad de

la corteza. Además, se aportan citas de accessiones re] ivas de especimenes cultivados de Phellodendron en jardines y arboretos.

INTRODUCTION

Phellodendron, Rutaceae, is a small genus of ornamental, deciduous trees, with only two species, endemic

to East Asia (China, Japan, the Korean Peninsula and the Russian Far East, Ma et al. 2006) where the bark

and oil from its fruit are used medicinally (Huang Bo). Nearly 150 years have passed since Phellodendron

was first introduced to the West in 1850-1860 (Rehder 1940; Read 1974; Bean 1976). Nonetheless, the

nomenclatural status of the genus, especially in the horticultural field, is still confused today (Dirr 1998).

For this study, published records of cultivated species of the genus were reviewed, available specimens were

examined, major characters in cultivated specimens were analyzed, and the status of the alien species of

Phellodendron in North America was clarified.

HISTORICAL REVIEW

The genus Phellodendron was completely revised first by Sargent (1905) based on the specimens at A and

GH, and the living collections at the Arnold Arboretum of Harvard University; and a total of three species

was accepted with fine illustrations of P. amurense Rupr. (Manchuria; Mongolia; C China), P. sachalinense (F.

Schmidt) Sargent (Korea; Hokkaido, Japan) and P. japonicum Maxim. (Japan; Hubei and Sichuan, China).

Sargent's revision has been cited as the classical history of the genus. However, because not enough speci-

mens from the native area were available to his study, Sargent could do no better than earlier researchers;

and, in fact, he created further confusion by identifying specimens from central China as both P. sachalinense

and P. japonicum, a later described species (see below). Sargent also raised P. amurense var. sachalinense F.

Schmidt to specific status as P. sachalinense (F. Schmidt) Sargent because he believed that it differed from

P. amurense Rupr. in the darker color of the branchlets; the thinner, not corky, bark; the rufous, rather than

silvery pubescent, winter buds; the leaflets not lustrous adaxially and glabrous along the margins; and the

glabrous inflorescences. The species, P. sachalinense (F. Schmidt) Sargent, however, was treated as a synonym

of P. amurense Rupr. in the modern Flora of Japan (Ohwi 1984; Ohba 1999).

J. Bot. Res. Inst. Texas 1(1): 357 — 365. 2007

358 Journal of the Botanical R h Institute of Texas 1(1))

In Rehder's Manual (Rehder 1940), an authoritative guide in the field of horticulture, five species of

Phellodendron were recorded: P. amurense Rupr. from N China, Manchuria, introduced to North America

in 1856, and cultivated in Hardiness Zone III; P. sachalinense (F. Schmidt) Sargent from Sghalin, Korea, N

Japan, and W China, introduced to North America in 1877, and cultivated in Hardiness Zone III; P. lavallei

Dode from C Japan, introduced to North America in 1862, and cultivated in Hardiness Zone V; P. japonicum

Maxim. from C Japan, introduced to North America in 1863, and cultivated in Hardiness Zone IV; and P.

chinense C. K. Schneid. from C China, introduced to North America in 1907, and cultivated in Hardiness

Zone V. This work was completely adopted by another important horticultural manual (Bailey 1949), with

fine illustrations of the leaves; and also adopted by the recent monumental work, Garden Flora of Europe

(de Vries 1997). Based on these standard horticultural references for North America and Europe, different

names have been used for living specimens in various gardens and arboreta, rather than the two names

accepted here. Some were P. piriforme E.L. Wolf (e.g., Morton Arboretum record of 2004, 4539-38, Arnold

Arboretum Inventory of 2003, 41242-57 & 21607, Royal Botanic Gardens, Kew, Living Collection Database,

41938-13701, searched on March 28, 2003), which has never been effectively published (Ma et al. 2006),

or P. insulare Nakai (e.g., Holden Arboretum Living Collection Database, 4481-285, Royal Botanic Gardens,

Kew, Living Collection Database, 41988-4479, searched on March 28, 2003, and Cornell Plantations Liv-

ing Collections Database, 4482-177, searched on March 25, 2004), which has never been accepted (Ma et al.

2006), or P. lavallei Dode (e.g., New York Botanical Garden Living Collections Database, 4220/72, searched

on March 27, 2004, Royal Botanical Garden Edinburgh Catalogue of Plants 2001, 4419190039, Cornell Planta-

tions Living Collections Database, 401-130, searched on March 25, 2004, Brooklyn Botanic Garden Living

Collections Plant Inventory of 2004, 425008, 25009 & X00316, The Plants of Pennsylvania, Rhoads & Block

2000 and Garden Flora of Europe, de Vries 1997), which has never been accepted in its native flora of Japan

(Ohba 1999) and worldwide revision (Ma et al. 2006). Some were even wrongly reported as P. sachalinense

(Rhoads & Block 2000; McNamara and Pellett 2000 - the report, however, has been denied by the Junior

author (Harold Pellett, pers. comm.). Pellett stated that “We do not have any Phellodendron sachalinense that

we are confident of their identity. We think that the trees that we have in the Minnesota Arboretum are

probably hybrids with P. amurense or some other species."

DIFFERENCES BETWEEN BARK OF NATIVE AND CULTIVATED TREES

There should not be much difference between the bark of the woody plants in cultivation and in their native

habitats. However, this indeed happened in P. chinense. From measurements and observations of trees in

their native habits, the two layers of bark of the two species could be easily distinguished (Ma et al. 2006):

the outer bark of P. amurense is nearly 10 x (1.12/0.37 to 0.13/0.33) thicker than those of P. chinense. However

in this study, all of the data of P. chinense from cultivated trees in northeastern North America are basically

similar to those of P. amurense in its native habitat (see Table 1).

The ratio of outer to inter bark of P. chinense in cultivation is 1.56 to 0.92 (i.e., 1.7: 1), much larger than

natural, wild populations (0.38: 1, Ma et al. 2006), approximately one half of the bark of P. amurense in wild

(3.1: 1). Because P. chinense was from central and southwestern China where the weather is much warmer

and wetter than the sites where it is in northeastern North America, the bark likely thickened in response

to the colder temperatures, especially the outer layer, to protect the cambium. This indicates the plasticity of

bark thickness of P. chinense in response to the environment (similar to Hedge et al. 1998s finding of variable

bark thickness in response to disturbance in the western Ghats of India). However, no such change occurred

in P. amurense, with its already thick, fissured bark (insulated from extreme temperatures, see Nikolai 1986)

and grows in an area of East Asia, similar in climatic conditions to northeastern North America.

TAXONOMIC TREATMENT

Tree, deciduous, dioecious, 15-35 m high, 40-60(-100) cm in diam., usually with secretory cavities con-

taining aromatic ethereal oils scattered throughout parenchymatous tissues. Bark corky, generally in two

Ma and Brach, Cultivated Phellodendron in North America 359

layers: phloem (inner part), yellow, usually thickened with age, and cork (outer part), gray, usually thickened,

dark, deeply striped or fissured along main truck; lenticels white, slightly expanded on young branches; pith

present, white or light brown to brown, round, continuous, sometimes spongy. Buds solitary, small, always

hidden beneath leaf petiole, naked after leaves have fallen, pubescent, 2 per node, opposite. Leaf scars nearly

encircling buds, 7-8 mm in diam., vascular bundle scars 3. Leaves odd-compound, opposite; estipulate;

strongly aromatic, pellucid punctate along margin. Leaflets (7 or)9(or 11), mostly opposite, sometime alternate,

or unequal at base; leaflet blade elliptic to ovate-oblong, symmetric, 21-32 x 13-16 cm, pilose when young

or glabrous, but most becoming glabrous at maturity, base attenuate, sometimes slightly oblique, margin

subentire or with minimal and fine serrulations not easily observed, apex acute or acuminate, sometime

caudate, lateral veins pinnate, 6-11 pairs, mostly not prominent abaxially, curved forward to acute, again

divided and disappearing before reaching margin; petiole 5.5-7.5 mm long. Inflorescences, a panicle 6.5-13.5

x 4.5—9 cm, loose or compact, nearly corymbose, terminal or opposite to young stem, with many flowers

in several clusters, clusters opposite or nearly so; peduncle 4—8.5 cm long, without scales or pubescent,

branching or not. Flowers: male: 5-merous, sepals 5, petals 5, stamens 5, anthers yellow, globose, ca. 1 mm

long and in diam., 2-lobed, longitudinally dehiscent, disc small, around pistillode, pistillode clavate, white

pubescent at apex; female: 5-merous, sepals 5, petals 5, staminode clavate; carpels 5, ovary 5-locular, ovule

1 per locule, style very short or nearly absent, stigma capitate, 5-lobed, much shorter than ovary, persistent.

Fruit a drupe, black, 8-9.4 x 7.5-8.7 mm, subglobose, 5-locular, stone-like, glabrous, most with 5 grooves

and angles when dry; fruiting pedicel ca. 0.4 mm long. Seed 1 per locule, brown, sometimes with black

pits, ellipsoid, to 4.5 x 2.5-3 mm, slightly compressed, shiny; endosperm oily, cotyledons flattened, embryo

straight; germination epigeal.

KEY TO THE SPECIES OF PHEEECIOBNIDRON

1. Panicle 8.5-13.5 x 6.5-9 cm, loose, peduncle 5-8.5 cm long, branches at least 1 cm long; tree, 25-35 m high

(cultivated in North America, Europe, Australia, New Zealand, and northern Asia) 1. P, amurense

1. Panicle 6.5-9.5 x 4-6.5 cm, compact, peduncle 4-6 cm long, unbranched or nearly so; tree, 15-20(-25) m

(cultivated in North America and Europe) 2. P, chinense

1. Phellodendron amurense Rupr., Bull. Cl. Phys.-Math. Acad. Imp. Sci. Saint-Petersbourg Ser. 2, 15:353.

1857.

Phellodendron amurense Rupr var. angustifolum E.L. Wolf, P. amurense var. japonicum (Maxim. ) Ohwi, P amurense Rupr var. latifolium E.L.

Wolf, P amurense var. lavallei (Dode) Sprague, P amurense var. molle (Nakai) S.H. Li & S. Z. Liou, P amurense f. molle (Nakai) Y.C.

Zhu, P amurense var. sachalinense E Schmidt, P amurense var. wilsonii (Hayata € Kaneh.) C.E. Chang.

Phellodendron insulare Nakai

Phellodendron sachalinense (E Schmidt) Sargent, P sachalinense Rupr. var. suberosum (H. Hara) H. Hara, P sachalinense (E Schmidt) Sarg.

var. suberosum H. Hara

Phellodendron wilsonii Hayata & Kaneh.

Phellodendron amurense was introduced in the 1850s to the West (Rehder 1940). By 1874, it was cultivated

at the then two-year-old Arnold Arboretum of Harvard University (Goodale 1877; Roca-Garcia 1970; Del

Tredici 1995). Since then the tree has been reintroduced into the Arboretum many times from different

countries (China, Japan, and Russia, especially 1900-1920s) until later years of the 20" century (Rehder

1940; Anonymous 1971). By 1910, the tree was cultivated both in Europe and in North America, at such

360 Journal of the Botanical R h Institute of Texas 1(1))

TABLE 1. Bark thickness of P. chinense cultivated in northeastern North America

Arboretum Name Inventory No. CR OLB ILB Voucher*

Morton Arboretum V57-55-30 230 28 ES J.S. Ma 5101

Morton Arboretum V57-80-85 140 22 14 J.S. Ma 5100

Arnold Arboretum 6963-2A 152 0.8 0.5 J.S. Ma 5116

Arnold Arboretum 55-55C 203 1:2 0.6 DMG ou

Dawes Arboretum S.N. 138 0.8 0.6 J.S. Ma 5107

Average: 168.6 1:560 0.92 Ratio: 1.7: 1

* CR: Circumference of trunk at DBH, OLB: outer layer of bark, ILB: inner layer of bark; all voucher specimens are deposited in

BKL, all measurements are in cm.

places as Royal Botanical Garden Kew of England, Breslau, Poland, the USDA Bureau of Plant Industry,

Glenn Dale, Maryland, the Arnold Arboretum, Massachusetts, and Cornell Plantations, New York. Among

them, the Arnold Arboretum played a very important role for redistributing the species, especially in North

America. By the 1930s, it had been collected from California, Michigan, New York (Grier and Grier 1928),

Ohio, Pennsylvania, Washington D.C. and Canada, and now, it is widely planted in more than 20 states

within the USA (USDA Hardiness Zone 3-7(-8); Schopmeyer 1974; Dirr 1998; Hensley et al. 1991; Jacobson

1996; also see Fig. 1). Gardens and arboreta in Russia also played a very important role in the spread into

European gardens and arboreta, nurseries, and parks (Bean 1976).

Because P. amurense recently has been considered an invasive alien in northeastern North America

(Hao et al 2004; Glaeser & Kincaid 2005; Invasive Species Initiative 2005; Invasivespecies.gov 2005), some

cultivated plants have been removed, e.g., at Dawes Arboretum, all female trees had been cut down when

the author visited in spring 2004). This kind of action, however, may go too far because the species has only

escaped and become naturalized in only a few places in northeastern North America (http://plants.usda.

gov/), around the vicinity of gardens, arboreta, or parks in urban areas, or along residential roadsides where

they were planted, e.g., in New York City (Anonymous 1991; Anonymous 1995; Glaeser & Kincaid 2005).

Among the specimens examined for this study, all were collected in residential areas or urban habitats even

though it was reported as becoming invasive in the larger New York Metropolitan area (Greller 1977; Grel-

ler & Calhoon 1979; Mitchell 1999; 2001; Lamont & Young 2002; Glaeser 2005; Glaeser & Kincaid 2005),

e.g., New York Botanical Garden (Cruz & Nee 2003; Small & Alexander 1933) and Forest Park, Queens Co.,

New York City (Anonymous 1961). Although, seeds of P. amurense possibly require a dormant period for

germination (Starshova 1979; Zhu & Dong 1990; Mizui & Kikuzawa 1991; but see Read 1974), there are no

reports about this from the natural areas in northeastern North America. Therefore, it cannot be treated as

an invasive species if we accept recent concepts of naturalization and invasion of alien plants (Richardson

et al. 2000; Pysek et al. 2004). Furthermore, there has been no damage reported to the native flora even

though the potential exists for P. amurense to become an invasive in the future, especially in northeastern

North America (Massachusetts Invasive Plants List 2005).

Original distribution.—Mixed forests, below 2500 m: China (Beijing, Hebei, Heilongjiang, Jilin, Liaon-

ing, Nei Mongol, Shandong, Taiwan: 2,000-2,700 m), Japan, Korea, and the Russian Far East.

Cultivated distribution — CANADA: British Columbia, Ontario. USA: California, Colorado, Connecticut,

Delaware, Georgia, Illinois, Indiana, Kansas, Maryland, Massachusetts, Michigan, Minnesota, Missouri, New

a ee

Jersey, New York, North Carolina, Oregon, Pennsylvania, Tennessee, Virginia, Washingt

also in Australia, Belgium, the Czech Republic and Slovakia, Denmark, England, EUN Ende France,

Germany, Hungary, Ireland, the Netherlands, New Zealand, Norway, Poland, Russia, Scotland, and Spain.

Cultivated specimens studied: CANADA. BRITISH COLUMBIA. Vancouver: Elizabeth Park, 14 Jun 1988, G.B. Straley 4809 (MOR,

NA). ONTARIO. Ottawa: Central Experimental Farm Campus of Brockport, 8 Jun 1974, W.I. Illman cco19082 (NYS); Dominion Arbo-

Ma and Brach, Cultivated Phellodendron in North America 361

Fic. 1 M E IP d M + £ DhAllAH J H M A ecce

e d.

retum and Botanic Garden, #120646, 10 Jul 1939, J.M. Gillett X-2-257 (NA), 29 Aug 1939, fruit, G.H.M. Lawrence 778 (BH); 10 Jul 1939,

H.A. Senn 1940 (BH).

U. S. A.: CALIFORNIA. San Francisco Co.: Golden State Park, 20 May 1933, E. Walther s.n. (A); University of California Davis,

2 Aug 1972, H. Fongs.n. (CM). CONNECTICUT. Fairfield Co.: Hill Road, 9 Apr Y 20 Aug 1941, flower & fruit, E.H. Eames s.n. (CONN);

Sherman, Turner Mt., Caretakes's Lodge, 28 Jun 1989, M. Ardwin s.n. (CONN); Litchfield Co.: Salisbury Southeast side of Prospect Mt.,

23 May 1985, L.J. Mehrhoff 11350 (CONN); 10 Oct 1985, fruit, 11792 (CONN). Tolland Co.: Storrs, Mountain Road and Rte 44, 20 Oct

1999, fruit, B.A. Connolly 42 (CONN); 23 May 1969, M. Lefor & F.H. Wolfe 647 (BH, MASS). DELAWARE. New Castle Co.: Greenville,

Cuba Botanical Park, 6 Jun 1968, Cuba Botanical Park Herbarium 115 (A, K). GEORGIA. Cherokee Co.: Reinhardt College Campus, 4

Jun 1983, F.G. Meyer & P.M. Mazzeo 19708 (A, CM, MOR, NA). ILLINOIS. Champaign Co.: 31 Aug 1970, R.A. Evers 103534 (A), 18 Oct

1998, fruit; S.R. Hill 31181 (NY); 17 May 1972, B. Nelson, B. Little & C. Crist 602 (A). Cook Co.: 9 Jul 1983, fruit, L. Nee, 27451 (E). DuPage

Co.: Lisle, Morton Arboretum, 15 Jun 1995, K. Altvatter & J. Hammond 7115V95 (F); 19 Jun 1995, fruit, 7134V95 (F, seed from Bota-

nischer Garten und Botanisches Museum Berlin-Dahlem, Berlin); 11 Oct 1994, fruit, B.K. Altvatter & P. Steinhouse 6901V94, 358-25 (F,

seed from Arnold Arboretum); 3 Aug 2001, W. Hess & K. Allen 9688 (F, NY); 18 Jun 1997, G. Hickman & J. Pinkard 7926V97 (NA); 9 Jul

1998, S.K. DeMink s.n. (F); 10 Oct 1998, fruit, thick cork bark, R.D. Hyerczyk 1598 (MOR); 10 Oct 1998, fruit, 1601 (NA); 7 Sep 1998,

D.S. Kirt 475-32 (NA); S.N. Kobal 95-14 (MOR); 7 Aug 1987, F.A. Swink 7406 (MOR); 30 Sep 1992, G. Wilhelm s.n. (MOR); Will, Pilcher

Park-Joliet Park District, 10 Oct 1998, fruit, R.D. Hyerczyk 1601 (MOR). INDIANA. Monroe Co.: Bloomington, Indiana University

Campus, 29 Jul 1949, fruit, R.B. Ledin s.n. (CONN). MARYLAND. Baltimore Co.: Carroll Park, 15 May 1979, F.G. Meyer & R. Fisher

17479 (NA); 15 May 1979, 17480 (A); Charles, Bryans Farm, 27 Sep 1975, fruit, F.G. Meyer 15282 (NA); 3 Jul 2000, fruit, B.W. Steury 7036

(NA). Prince George's Co.: Glenn Dale, USDA Bureau of Plant Industry, 20 Sep 1906, I. Tidestrom 1805 (NA). Montgomery Co.: Silver

Spring, 11 Aug 1975, F.G. Meyer & P.M. Mazzeo 15262 (NA). MASSACHUSETTS. Hampshire Co.: Amherst, University of Mass., 18

Sep 1966, fruit, H.E. Ahles 64704 (MASS 2 sheets); 1 Jul 1969, fruit, A.C. Gibson 965 (A); S.R. Hill 16999 (NY); 12 Oct 1940, R.E. Torrey

s.n. (MASS); 8 Jun 1945, s.n. (MASS); 29 Aug 1946, fruit, s.n. (GH, MASS); 6 Jun 1951, R.E. Torrey & E. Putala s.n. (MASS); Holyoke

362 Journal of the Botanical R h Institute of Texas 1(1))

Range, South Hadley, 12 Sep 1999, K.B. Searcy 52 (MASS); Northampton, Smith College, 21 May 1969, A.C. Gibson 362 (A), Reading,

Jun 1879, W.H. Manning s.n. (BH). Norfolk Co.: Brookline, 1 Sep 1965, Baldini s.n. (A 2 sheets). Suffolk Co.: Jamaica Plain, Arnold

Arboretum, 16 Jun 1960, fruit, B.K. Boom 40329 (L, from Japan in 1905); 16 Jun 1960, 40331 (L, collected by Wilson from Japan in 1919);

16 Jun 1960, 40332 (L); 16 Jun 1960, fruit, 40338 (L), 27 Jun 1917, H.H. Chung 5251 (PE 2 sheets); 23 Oct 1979, fruit, K. Clausen, S.

Davis, C. Warren & M. Awolcott 79168 (BM); 20 Aug 1904, fruit, E.J. Cole s.n. (A, MICH); 8 Jun 1980, S. Davis, C. Warren & M. Wolcott

80-201-B (BH, F, PE, raised from seed of Wilson 11263, 17 Feb 1919 from Japan); 9 Jun 1982, fruit, S. Davis et al. 558 (A, BH, F, from

Arnold Arboretum £7544-C, seed from J. G. Jack, Azuma, Japan, 15, Dec 1905); 15 Sep 1982, fruit, S. Davis & M. Wolcott 566 (A, BM,

BH, F, from Arnold Arboretum #7544-C, seed from J. G. Jack, Azuma, Japan, 15, Dec 1905); 18 Aug 1960, fruit, T.R. Dudley s.n. (A); 10

Aug 1984, fruit, S. Elsik, B. Mackenzie, A. Kosmidis, & L. Stockman 1655 (A, BH, Arnold Arboretum £646-66-A); 30 May 1986, 4269 (A,

BH, Arnold Arboretum #401-56-A); 30 May 1986, 4270 (A, BH); 30 May 1986, 4280 (A, BH, Arnold Arboretum #13232-B); 25 Jun 1969,

fruit, A.C. Gibson 868 (A, Arnold Arboretum #12202-1-B); 25 Jun 1969, fruit, 869 (A, Arnold Arboretum #13232-A), 25 Jun 1969, fruit,

870 (A, Arnold Arboretum 13232-B); 25 Jun 1969, fruit, 873 (A, Arnold Arboretum #7544-c, from J. Jack 1905, Japan); 5 Sep 1985, fruit,

S. Elsik, G. Good & K, Groves 3407 (A, Arnold Arboretum #362-54-A, seed from Japan in 1952); 5 Sep 1985, 3417 (A 2 sheets, Arnold

Arboretum #1242-57-A); 21 Oct 1986, fruit, S. Elsik & R. Zinman 4866 (A, Arnold Arboretum #646-66-B, from J.G. Jack s.n. 3 Nov 1919);

18 Jul 1913 & 17 Oct 1920, C. Schneider 142-15 (BKL 2 sheets); 15 Nov 1908, fruit, 143-17 (BKL); 17 Oct 1915, fruit, 4961-1 (BKL); 17

Oct 1918, s.n. (BKL); 18 May 1926, s.n. (BKL); 15 Sep 1930, fruit, s.n. (BKL); 15 Sep 1922, H. Teuscher s.n. (A 8 sheets, MOR, 4143-6 from

Bot. Garden Petersb. in 1874); 18 Jul 1914, fruit, L.G. Hornby s.n. (MICH); 7 Jun 1927, N. Judd s.n. (A, Arnold Arboretum 41008-26); 9

Sep 1931, Kobuski & Ronsh s. n. (A 2 sheets, Arnold Arboretum 4150-28, from 143-6, 1928); 8 Sep 1931, s.n. (A, Arnold Arboretum

#1008-26, 2™ exp. Farrie, 1926); 4 Sep 1985, D. Michener & S. Elisk 3403 (A, BH, Arnold Arboretum £389-68-A); 3 Jun 1936, EJ. P. s.n.

(A, BH, Arnold Arboretum #143-9-B), Jun 1937, s.n. (A, MOR, Arnold Arboretum #7544, seed from J. G. Jack in Azuma, Japan, 1905);

3 Jun 1936, s.n. (A, BH, MOR, from #21607-A, seed of Bot. Inst., Leningrad, USSR in 1926); 17 Sep 1936, fruit, s.n. (A 4 sheets, MOR,

from #12202-1, Grafted from #12202, Arnold Arboretum in. 1919); 17 Sep 1936, fruit, s.n. (A, #19480-D, seed from Bot. Gard., Muenchen

Germany in 1925); 17 Sep 1936, fruit, s.n. (A, #19481-C, seed from L. Spath, Berlin, Germany); E.H. Wilson 876 (A); 11 May 1904, C.S.

Sargent s.n. (A); 6 Jun 1973, L. Segal & A. Thompson 71 (A, Arnold Arboretum #12202-1-A); 5 Oct 1922, H. Teuscher s.n. (MOR, Arnold

Arboretum #10724, E.H. Wilson 11263, seed collected by Wilson, from Taiwan (Formosa) in 1919); 29 May 1986, R. Zinman & J. Carey

4264 (A, BH, Arnold Arboretum #1244-57-A); Somerville, Tufts University Campus, 5 Aug 1998, fruit, NHN/EGM/APC s.n. (CONN);

Wellesley, 2 Aug 1972, fruit, S.A. Spongberg & H. Clement 72-51 (A, BM). MICHIGAN. Washtenaw Co.: Ann Arbor, University of

Michigan Arboretum, 7 Oct 1931, fruit, J.H. Ehlers 4999 (MICH). Clinton Co.: East Lansing, Michigan State University Campus, 8 Jun

1979, W.T. Gillis 15109 (CONN). MISSOURI. Boone Co.: Columbia, Missouri University Campus, Summer 1960, fruit, W. Campbell s.n.

(MOR). St. Louis Co.: St. Louis, Missouri Botanical Garden, 21 Aug 1989, fruit, M.T. Crosby 84 (TI, #U-5780); 26 Jul 1974, fruit, J.

Slama 38 (A, BH). NEW YORK. Bronx Co.: Bronx, New York Botanical Garden, 1937, L. Croizat s.n. (NY 5 sheets); 11 Jul 1923, fruit,

A.S. Foster s.n. (BH); 1909, fruit, H. Hallier s.n. (L); 26 Jul 1929, H. Moldenke 4926 (NY); 16 Aug 1995, fruit, C. Morenberg 115 (NY); 30

Aug 1980, fruit, S. Mori 13639 (NY); 29 May 1994, 23700 (NY); 5 Jul 1993, fruit, M. Nee 43585 (BH, NY); 30 Sep 2003, fruit, 52592 (NY);

3 Oct 2003, fruit, 52593 (NY); 4 Oct 2003, fruit, 52594 (NY); 5 Oct 2003, fruit, 52595 (NY); 8 Oct 2003, fruit, 52596 (NY); 10 Oct 2003,

fruit, 52597 (NY); 10 Oct 2003, fruit, 52598 (NY); 11 Oct 2003, fruit, 52599 (NY); 11 Oct 2003, fruit, 52600 (NY); 13 Oct 2003, fruit,

52601 (NY); 13 Oct 2003, fruit, 52602 (NY); 23 Sep 2003, fruit, 52950 (NY); 18 Jun 1931, fruit, P. Wilson s.n. (NY). Kings Co.: Brooklyn,

Brooklyn Botanic Garden, 6 Aug, 1979, fruit, TJ. Delendick s.n. (BKL); 25 May 1982, fruit, s.n. (BKL 5 sheets); 25 May 1982, s.n. (BKL 3

sheets); 26 May 1982, s.n. (BKL 2 sheets); 26 May 1982, s.n. (BKL 6 sheets); 26 May 1982, s.n. (BKL 5 sheets). Nassau Co.: 8 Jul 1997,

S.D. Glenn 2777 (BKL, NYS); 25 Aug 1995, D. Kunstler s.n. (BKL). Ontario Co.: Geneva, New York Agricultural Experimental Station,

19 Aug 1921, fruit, F. Blank s.n. (BH). New York Co.: Central Park, 12 Sept 1914, E.B. Jaultiruck s.n. (BKL 2 sheets). Orange Co.: Sterling

Forest, 5 Jul 2000, fruit, R.S. Mitchell 10751 (BKL); fruit, 11 Sep 2004, G. Moore, T. Delius & J.S. Ma 6864 (CONN); Scarboro, 31 Nov

1895, fruit, W.H. Manning s.n. (BH). Somerset Co.: 25 May 2000, S.D. Glenn 4161 (BKL). Suffolk Co.: Islip, along LIRR, 27 Jul 1994,

S.D. Glenn 48 (BKL). Tompkins Co.: Ithaca, Cornell University Plantation, 7 Oct 1946, fruit, M.W. Allen s.n. (BH 2 sheets); 23 Jun 1890,

fruit, L.H. Bailey s.n. (BH); May 1985, fruit, H. Banhs s.n. (BH); 9 Oct 1941, S.H. Burnham s.n. (BH 2 sheets); 24 Oct 1940, fruit, J. Comman

s.n. (BH 2 sheets); 16 Jun 1904, J. E. Coit s.n. (BH) & fruit, 25 Aug 1904, s.n. (BH); 24 Aug 1952, fruit, AJ. Eames s.n. (BH); Summer 1946,

fruit, A. Schulze s.n. (NYS). Westchester Co.: Yonkers, 20 Jun 1938, seedling, H.N. Moldenhe 10600 (BH, NY). NORTH CAROLINA.

Buncombe Co.: Biltmore, 19 Jul 1894, W.H. Manning s.n. (BH); 27 May & 2 Aug 1898, flower and fruit, S. P. 7086 (F, NY). OHIO. But-

ler Co.: Oxford, 27 May 1993, M.A. Vincent 5894 (NA). Hamilton Co.: Cincinnati, Mt. Airy Forest Park, 10 Oct 1934, fruit, E.G.

Hutchinson s.n. (BH). Cuyahoga Co.: Cleveland, Gordon Park, 5 Jun 2000, G. Wilder & M. McCombs 13672 (MICH). Franklin Co.:

Columbus, Ohio State University, 10 May 1967, J.F. Cooke & R.L. Stuckey s.n. (A, NA). OREGON. Ravalli Co.: Corvallis, Oregon State

University, 3 Jun 1968, J. Dennis 2931 (A). PENNSYLVANIA. Allegheny Co.: Pittsburgh, Highland Park, 26 Jun 1961, M. Armbruster

s.n. (CM); 17 May 2002, B.L. Isaac & J.A. Isaac 14339 (CM 2 sheets); 14 Oct 1973, V. Phelps, s.n. (CM); 27 Aug 1937, RJ. Templeton & J.R.

Steck s.n. (CM); Pittsburgh, 17 May 2002, A. Rhoads s.n. (MOAR), 24 Oct 2004, s.n. (MOAR). Berks Co.: 14 Jun 1968, fruit, W.C. Brum-

bach 6262 (A); 14 Jun 1968, fruit, 6271 (A); 22 May 1969, 6763 (A, NA, NY); 11 Aug 1971, fruit, 7644 (A, NY). Bucks Co.: 12 Aug 1998,

A.F. Rhoads & T.A. Bloch s.n. (MOAR); 6 Aug 1980, fruit, A.E. Schuyler 5481 (PH). Delaware Co.: Haverford College, 17 May 1942, P.T.

Haas s.n. (PH); 28 Aug 1995, fruit, 28 Jul 1941, s.n. (BKL, PH); 11 Oct 1933, EJ. P. s.n. (A, Arnold Arboretum 4972-34). Lancaster Co.:

Elizabeth, 16 Jul 1930, E.M. Gress s.n. (PH). Leigh Co.: Allentown, 26 May 1956, R.L. Schaeffer, Jr. 50574 (PH); 22 May 1959, s.n. (PH).

Mercer Co.: 31 Aug 1967, J.M. Fogg s.n. (MOR); 11 Jul 1968, fruit, s.n. (A); 26 May 1970, s.n. (A); 12 May 1973, S.A. & H. Spongberg 73-

Ma and Brach, Cultivated Phel

th America 363

220 (A, BM). Montgomery Co.: Melrose Park, 15 Jul 1937, C.G. Armstrong s.n. (PH); 20 Aug 1971, fruit, J.M. Fogg s.n. (NY); B. Long 39270

(PH); 29 Jun 1970, Merion, Arboretum of Barnes Foundation, Nei 1980, fruit i M. os n. (MOR); L.K. Henry s.n. (CM); 19 Feb 1933,

Fort Washington Park, A. Zakyzewski & T. Livshultz s.n. (PH). Philadel : s Arboretum, 27 Oct 1967, J.M. Fogg, Jr. s.n. (MOAR,

855-14); 12 Nov 1932, fruit, without collector 969 (MOAR 3 sheets); don 1933, 1712 (MOAR 2 sheets); 24 May 1934, 2681 (MOAR);

Philadelphia, Lakeside Avenue, 26 Jul 1950, B. Long 71831 (PH). Northumberland Co.: Snyder, 27 Sep 1927, H.N. Moldenhe 3551 (NY).

Westmoreland Co.: Greensburg, 1952, C.W. Kalbfus s.n. (CM). TENNESSEE. Knox Co.: Knoxville, Agriculture Campus, University

of Tennessee, 13 Jun 1972, fruit, P.M. Mazzeo & F.G. Meyer 12858 (MOR, NA); formerly Sanford Arboretum (now property of W. E.

Fleury, 3425 Lakeview Dr.), 15 Jun 1972, fruit, P.M. Mazzeo & P.G. Meyer 12913 (MOR, NA). Franklin Co.: Winchester, Shadow Nurs-

ery, RRI, 6 Jun 1983, F.G. Meyer & P.M. Mazzeo 19765 (CM, MOR, NA). VIRGINIA. Clarke Co.: Boyce, 25 May 1970, O. E. White Ar-

boretum, F.G. Meyer & P.M. Mazzeo 12430 (NA); Richmond, 11 Jun 1974, fruit, B.F. Kiltz 540 (NA). WASHINGTON. King Co.: Seattle,

Washington Park Arboretum, 30 May 1991, J. Canary 94 (NA); 17 Oct 1991, C. Bates 30 (NA). WASHINGTON, D.C.: Library of Congress

Grounds, Aug 1931, fruit, E.H. Walker 1737 (NA); White House, South Grounds, 22 May 1980, F.G. Meyer & P.M. Mazzeo 17804 (MOR,

NA); National Arboretum, 20 Jun 1994, fruit, F.G. Meyer & P.M. Mazzeo 17804 (NA); Tidal Basin, 10 Jun 1977, F.G. Meyer & H. Wester

15918 (MOR); Soldier's Home, University of America, 1938, fruit, F. Baeehle s.n. (NA). WISCONSIN. Dane Co.: Madison, 13 Aug 1986,

fruit, M. Nee 20818 (NY).

2. Phellodendron chinense C.K. Schneid., Ill. Handb. Laubholzk. 2:126, fig. 79 c-d, 1907.

Phellodendron amurense f. longipes Y.C. Wu

Phellodendron chinense var. falcatum Huang, P chinense var. glabriusculum C.K. Schneid., P chinense var. omeiense Huang, P chinense var.

yunnanense Huang

Phellodendron fargesii Dode

Phellodendron sinense Dode

Phellodendron sinii Y.C. Wu

Phellodendron chinense was introduced into North America much later, approximately 100 years after it was

described in 1907, and much of its introduction to North America as well as to Europe was based on E.

H. Wilson's collections from central and southwest China a century ago (Roca-Garcia 1970). This species,

however, has not been as popular as P. amurense, but cultivated in the gardens and arboreta in Hardiness

Zone V (Rehder 1940). It has never been reported as escaped or naturalized.

Original distribution Mixed forests, below 2000 m: China (Anhui, ?Fujian, ?Guangdong, ?Guangxi,

?Guizhou, Hubei, Hunan, ?Jiangsu, ?Jiangxi, Shaanxi, Sichuan, Yunnan, ?Zhejiang. Since the species has

been longtime cultivated or naturalized in some places (with “?” before the province) in China, their native

distributions or cultivation cannot be obtained with certainty.

Cultivated distribution. — CANADA: Ontario. USA: Colorado, Massachusetts, New York, Pennsylvania;

also in Belgium, England, Germany, Hungary, Ireland, Italy, and Scotland.

Cultivated specimens studied: U.S.A. MASSACHUSETTS. Suffolk Co.: Jamaica Plain, Arnold Arboretum, 9 Jun 1982, K. Clausen, S.

Davis, C. Warren & M. Awolcott 557 (A, BH, BM, Arnold Arboretum £6963-2-A); 23 Oct 1979, fruit, 79-168 (A, BH, Arnold Arboretum

#6963, seed of Wilson 161); 23 Oct 1979, S. Davis, C. Warren & M. Wolcott 79-168 (F, Arnold Arboretum #6963, from seed of Wilson 161

in Changyang Hsien, W. Hupeh, China, Feb. 1908, tree, 50); 9 Jun 1982, S. Davis et al. 557 (A, Arnold Arboretum #6963-2-A, seed

from Wilson, Changyang Hsien, W. Hupeh, China, Feb. 1908); 25 Jun 1969, fruit S. Elsik, B. Machenzie, A. Kosmidis, & L. Stockman 871

[A, Arnold Arboretum #7245); 25 Jun 1969, fruit, 874 (A, Arnold Arboretum #6963-A), 5 Sep 1985, fruit, S. Elsik, G. Good & K, Groves

3416 (A, BH, Arnold Arboretum #6963-2-A); 16 Jun 1939, E.J. P. s.n. (A, Arnold Arboretum #6963-1-A); 16 Jun 1892, R. Rehder s.n. (A 3

sheets); 5 Nov 1917, fruit, s.n. (A 2 sheets, Arnold Arboretum 46963); 15 Oct 1917 & 5 Nov 1917, fruit, s.n. (A 2 sheets, Arnold Arboretum

#7425); 6 Jun 1918, s.n. (A, Arnold Arboretum #7245); 17 Sep 1918, C.K. Schneider 161 (BKL); 18 Sep 1918, fruit, 876 (BKL) & 17 Oct

1920, fruit (BKL, from Arnold Arboretum #6963). NEW YORK. Bronx Co.: Bronx, New York Botanical Garden, 25 Sep 1922, fruit,

L.H. Bailey s. n. (BH). Tompkins Co.: Ithaca, Cornell University Plantation, 27 Sep 1995, fruit, G.M. Elston 96-3 (BH). PENNSYLVANIA.

Mercer Co.: Arboretum of Barnes Foundation, 31 Aug 1967, J.M. Fogg Jr. s.n. (BKL); 7 Sep 1970, fruit, s.n. (A, BH). Philadelphia Co.:

Morris Arboretum, 15 Sep 1959, fruit, J.M. Fogg, Jr. s.n. (MOAR); 14 Mar 1933, fruit, without collector 1351 (MOAR 2 sheets).

ACKNOWLEDGMENTS

The senior author's travel in the USA was supported by Brooklyn Botanic Garden, and he sincerely thanks

Steve Clemants, Gerry Moore, and Steve Glenn for their help and assistance. Help was provided by the fol-

lowing people: Scott Aker (US National Arboretum), Tony Aiello (Morris Arboretum), Tiffany Enzenbacher

364 Journal of the Botanical R h Institute of Texas 1(1))

and Kunso Kim (Morton Arboretum), Ethan Johnson (Holden Arboretum), Kyle D. Port (Arnold Arboretum),

Michael Ecker (Dawes Arboretum), Randall Hitchin (Washington Park Arboretum), Richard S. Mitchell (New

York State Museum), Lynsey Muir and Crinan Alexander (Royal Botanic Garden Edinburgh), Harold Pellett

(Minnesota Landscape Arboretum), Jim Pringle and Margaret Walton (Royal Botanical Garden, Canada),

Hong Qian (State Museum of Illinois), and Steve M. Young (New York Natural Heritage Program). The senior

author visited and acknowledges the following herbaria and their helpful curators: A, BH, F, GH, L, MASS,

MICH, MOAR, MOR, NY, NYS, PH, US; and sincere thanks to Carsten Glaeser (Glaeser Horticultural Con-

sulting) for his help around the New York Metro Area, and Michael Nee (New York Botanical Garden, New

York) for his primary work on the identification of the cultivated species of the genus from the New York

Botanical Garden and vicinity. We also thank Jacquelyn Kallunki (NY) and anonymous reviewers as well

as the editor for their valuable help and useful comments and suggestions.

REFERENCES

ANONYMOUS. 1961. Local flora notes—Note on Phellodendron. Bull. Torrey Bot. Club 88:123-124.

ANONYMOUS. 1971. Amur cork-tree, Phellodendron amurense. Morton Arbor. Bull. 26-27.

Anonymous. 1991. Field trip reports—Trenton-Hamilton Marshes, Mercer County, New Jersey. Bull. Torrey Bot.

Club 118:470-474.

ANONYMOUS. 1995. Field trip reports—Holmdel Park, Holmdel, Monmouth County, New Jersey. Bull. Torrey Bot.

Club 122:70-73.

BaiLev, L.H. 1949. Manual of cultivated plants (revised edition), Macmillan Co., New York, NY. pp. 372-373.

Bean, W.J. 1976. Trees and shrubs hardy in the British Isles, eighth ed., J. Murray, London. pp. 125-127.

Cruz, P. and M. Nee. 2003. The identity of the planted and of the naturalized Phellodendron in the vicinity of the

New York Botanical Garden. Typescript from the senior author, dated Nov. 12, 2003, ms. P. 6.

De Vries F.T. 1977. Phellodendron (Rutaceae). In: J. Cullen et al., eds. European Garden floras 4:110-11 1.

DeL Trevici, P. 1995. Requiem for a cork tree. Arnoldia 55(3):22-24.

Ding, M.A. 1998. Manual of woody landscape plants, their identification, ornamental characteristics, culture,

propagation and uses. 5^ ed,, Stipes Pub., Champaign, IL, pp. 703-705.

GOODALE, G.L. 1877. General notes—botany. Amer. Naturalist 11:239-241.

GLAESER, C.W. and D. Kincaip. 2005. The non-native invasive Phellodendron amurense Rupr. in a New York City

woodland. Arbor. J. 28(3):151-164.

GRELLER, A.M. 1977. A vascular flora of the forested portion of Cunningham Park, Queens County, New York, with

notes on the vegetation. Bull. Torrey Bot. Club 104:1 70-176.

GreLLER, A.M. and R.E. CALHoon. 1979. The upland, oak-dominated community of Forest Park, Queens County, New

York. Bull. Torrey Bot. Club 106:135-139.

Grier, N.M and C.R. Grier 1928. A list of plants growing under cultivation in the vicinity of Cold Spring Harbor,

New York. Amer. Midl. Naturalist 11: 307—387.

Hao, Z., Y. Wu, J.Q. Ding, D. Binion, W.D. Fu, and R. Reardon. 2004. Invasive Plants of Asian Origin established in the

United States and their natural enemies. 1:131. Forest Health Technology Enterprise Team 2004-05, Forest

Service, United States Department of Agriculture, Morgantown, WV.

HEGDE, V, M.D.S. CHANDRAN, and M. Gapa. 1998. Variation in bark thickness in a tropical forest community of Western

Ghats in India. Functional Ecology. 12:313.

HensLey, D.L., S.C. Wiest, C.E. Lone, J.C. Pair, and F.D. GiBBoNs III. 1991. Evaluation of ten landscape trees for the Mid-

west. J. Environ. Hort. 9:149-155.

INVASIVE Species INITIATIVE. 2005. Invasive plant atlas of New England, June 2, 2005 <http://invasives.uconn.edu/

ipane/index.htm>

INVASIVESPECIES.ORG. 2005. Invasive plants lists of United States, June 14, 2005 <http://www.nps.gov/plants/alien/

list/all.htm>

JACOBSON, A.L. 1996. North American landscape trees. Ten Speed Press, Berkeley, CA. Pp. 422-423.

Ma and Brach, Cultivated Phellodendron in North America 365

Lamont, E.E. and S.M. Youna. 2002. Noteworthy plants reported from the Torrey Range - 2001. J. Torrey Bot. Soc.

129367

Ma, J.S., W. Cao, Q.R. Liu, M. Yu, and LJ. HAN. 2006. A revision of Phellodendron (Rutaceae). Edinburgh J. Bot.

63:131-151.

MASSACHUSETTS INVASIVE PLANTS Lists. 2005. Results of invasive plant species Evaluations, June 14, 2005 «http://www.

newfs.org/conserve/invlist.htmst MAEVAL»

McNamara S. and H. Pettett. 2000. Cold hardiness of Phellodendron sachalinense Friedr. Schmidt seedlings increases

with age. HortScience 35:304-305.

MircheLL, R.S. 1999. A Phellodendron new to New York State found naturalized in Sterling Forest. New York Flora

Assoc. Newslett. 10(4):1-2.

MitcHett, R.S. 2001. Sterling forest flora—Summary of a four year project. New York Flora Assoc. Newslett.

12(4):4-8.

Mizul, N. and K. KikuzAwa. 1991. Proximate limitations to fruit and seed in Phellodendron amurense var. sachalinense

Pl. Spec. Biol. 6:39-46.

Nikola, V. 1986. The bark of trees: thermal properties, microclimate and fauna. Oecologia 69:148-160.

Oppa, H. 1999. Rutaceae. In: K. lwatsuki et al., eds. Flora of Japan 2c:40, Kodansha, Tokyo.

Ohw, J. 1984. Rutaceae. In: Flora of Japan. Smithsonian Institution, Washington D.C. Pp. 583-584.

Pysek, P, D.M. RICHARDSON, M. REJMANEK, G.L. WEBSTER, M. WILLIAMSON, and J. KirscHner. 2004. Alien plants in checklists and

floras: towards better communication between taxonomists and ecologists. Taxon 53:131-143.

READ, R.A. 1974. Phellodendron amurense In: C.S. Schopmeyer. Seeds of woody plants in the United States. Agri-

culture Handbook No. 450, Forest Service, USDA, Washington, DC

ReHDER, A. 1940. Manual of cultivated trees and shrubs (2nd ed.), Macmillan Co., New York, NY, pp. 528-529.

RHOADS, A.F. and T.A. Bock. 2000, The plants of Pennsylvania, An Illustrated Manual, University of Pennsylvania

Press, Philadelphia, PA, pp. 639-640.

RICHARDSON, D.M., P. Pysek, M. REIMANEK, M.G. BARBOUR, F.D. PANETTA, and C.J. West. 2000. Naturalization and invasion of

alien plants: concepts and definitions. Diversity & Distrib. 6:93-107.

Roca-Garcla, H. 1970. The cork trees. Arnoldia 30(5):161-167.

SARGENT, C.S. 1905. Trees and shrubs, illustrations of new or little known ligneous plants. 1:199, pl. XCIV, Houghton,

Mifflin and Company, Boston.

SCHOPMEYER, C.S. 1974. Seeds of woody plants in the United States. Agricultural Handbook no. 450, Forest Service,

United States Department of Agriculture, Washington D.C. Pp. 578-579.

SMALL, J.K. and E.J. ALEXANDER. 1933. Natives trees in the New York Botanical Garden, a guide for the botanist and

naturalist. Typescript, New York Botanical Garden library.

STARSHOVA, N.P. 1979. Biology of seed germination in Phellodendron amurense Rutaceae. Bot. Zhurn. 64:1159-1968

(in Russian).

Zhu, N. and D.F. Done. 1990. Seed dispersal, dormancy, seed bank and regeneration of Amur Corktree. J.N.F.

Forest. Univ. 1(1):16-22.

366 Journal of the B ical R h Institute of Texas 1(1))

BOOK REVIEWS

Jerr Lowenrets and Wayne Lewis. 2006. Teaming with Microbes: A Gardeners Guide to the Soil Food

Web. (ISBN 0-88192-775-5, hbk.). Timber Press Inc, 133 S.W. Second Avenue, Suite 450, Portland,

OR 97204-3527, U.S.A. (Orders: www.timberpress.com, mail@timberpress.com, 503-227-2878, 1-

800-327-5680, 503-227-3070 fax). $24.95, 196 pp., 74 color photos, 1 color illustration, 17 diagrams,

pns on

The new book entitled Teaming with Microbes by authors Lowenfels and Lewis is a ad resource. These authors have presented soil

science and the soil food web in a Pounds understandable way. The inter g of photographs, drawings and the easy reading style

make this a very readable text for anyone with an interest. Gardeners especially will een from reading this book; they will come away

with a comfortable (and applicable) understanding of the soil food web in relationship to their gardens and lawns.

The book is separated into two parts: basic science and applications. The first part of the book creates a terrific foundation for all

the parts of the soil food web by using explanations of soil profiles, textures, pH, and providing readers with descriptions of various

soil inhabitants from the bottom of the web up. There is a chapter dedicated to bacteria, fungi, algae, slime molds, protozoa, nematodes,

arthropods, worms, gastropods, and reptiles, mammals and birds. Within each of these chapters on soil inhabitants’ information, the

authors provide what each are, how they act within the soil web, and how these organisms contribute to nutrient ne The as

1 1

focus a good bit of time and provid information on both bacteria and fungi as is warranted based on tl

organisms in the soil food RCM

The second part of the book focuses on helping the reader apply his/her newfound soil information to creating a healthy soil

food web in their lawns and gardens. The authors demonstrate these applications via the use of nineteen separate rules, all of which are

integrated into the remaining eleven chapters. For instance, rule number two states that “Most vegetables, annuals and grasses prefer

their nitrogen in nitrate form and do best in bacterially dominated soils." Rule three relates to trees, shrubs and perennials preferring

fungal dominated soil and nitrogen in a form other than nitrate. These rules are int d into the MM of each application

and are also presented in a succinct list form in the appendix. This application section of the book inclu ..how soil food

webs apply to gardening, tools for restoring soil food webs, use of compost, mulch and compost teas, maintenance E trees and other

perennials, and growing annuals and lawns. There is also a brief chapter with seasonal activities to promote a healthy soil food web.

Teaming With Microbes is an outstanding book for those gardeners interested in a solid understanding of how to create a healthy

lawn and garden. This book would also be a beneficial read for horticulturists, soil and turf science students, agronomists, or any other

interested readers. Authors Lowenfels and Lewis have done a superb job of making soil science and soil food webs understandable; they

even provide resource sections for readers whom have further interests in the subjects covered. Definitely a recommended read!!—Lee

Luckeydoo, Herbarium, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A.

Davip Tayor (ed.). 2006. Pride of Place: A Contemporary Anthology of Texas Nature Writing (ISBN 13:

978-1-57441-208-6, pbk.). University of North Texas Press, PO. Box 311336, Denton, TX, 76203-1336,

U.S.A. (Orders: www.unt.edu/untpress, 800-826-8911, 940-565-2142, 940-565-4590 fax). $16.95,

214 pp., b/w photos, 6" x 9".

The larger-than-usual type size in Pride of Place seems apt for this collection of essays about larger-than-life Texas. David Taylor, a

professor and Honors Advisor at the University of North Texas, has assembled a group of some of the best Texas writers to put pen to

paper in this delightful contemporary anthology. Taylor hastens to say in the introduction that what binds Texans together is pride, not

geography. And pride of place is what binds together these essays. After all, “It ain't braggin' if it's true.”

Beginning with Roy Bedichek's “Still Water,” it includes Carol Cullar and Barbara “Barney” Nelson on the Rio Grande region of West

Texas, John Graves's evocative “Kindred Spirits" on Central Texas, Joe Nick Patoski's celebration of Hill Country springs, Pete Gunter

on the Piney Woods, David Taylor himself on North Texas, Gary Clark and Gerald Thurmond on the Coastal Plains, Ray Gonzles and

Marian Haddad on El Paso, Stephen Harrigan and Wyman Meinzer on West Texas, and Naomi Shihab Nye on urban San Antonio

If youre not from Texas, you'll “get it" after reading this book. If you are from Texas, reading this is like being a kid on the front

porch of a summer evening, listening to the grownups spin their tales.

Texas truly is a “State of Mind,” as the old song goes. This anthology reflects that, and Pride of Place will be a proud addition to the

Texas section of this reviewer's bookshelf.—Penny McCook, Volunteer, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX

76102-4060, U.S.A.

J. Bot. Res. Inst. Texas 1(1): 366. 2007

NOTES ON LECHEA MARITIMA VAR. VIRGINICA (CISTACEAE)

Bruce A. Sorrie Alan S. Weakley

University of North Carolina Herbarium University of North Carolina Herbarium

North Carolina Botanical Garden, CB 3280 North Carolina Botanical Garden, CB 3280

Chapel Hill, North Carolina 27599-3280, U.S.A. Chapel Hill, North Carolina 27599-3280, U.S.A.

ABSTRACT

We assess variation within Lechea maritima and provide a key to infraspecifi Specimen citati ] the range of L. maritima

var. virginica.

RESUMEN

Se valora la variación en Lechea maritima y se aporta una clave de taxa infraspecíficos. Las citas de especímenes documentan el área L.

maritima var. virginica.

Lechea maritima Legg. var. virginica Hodgdon is restricted to maritime sands, historically known from coastal

Virginia including the southern Delmarva Peninsula (Hodgdon 1938). In recent years, botanists have dis-

covered var. virginica in adjacent states (see specimens cited below). To better understand and evaluate the

status of var. virginica, we examined over 80 specimens from throughout the range of L. maritima housed at

NCSC, NCU, VPI, and WILLI. Specimens at NCU and VPI were annotated in 1984 by D. Lemke to Lechea

maritima without varietial designation. In their analysis of the genus in the southeastern United States, Wil-

bur & Daoud (1961) omitted L. maritima, because at that time it was not known from their area of coverage.

For an excellent description of the nominate variety, see Barringer (2004).

IDENTIFICATION AND KEY

We found that plants of var. virginica are generally more robust than those of var. maritima, with notably

thicker stems. Hodgdon (1938) suggested that the main above-ground stems of var. virginica may be peren-

nial, but this has not been verified. Measurements of stem thickness indicate that those of var. virginica

(2.0—4.0 mm) are significantly wider than those of the nominate variety (1.0—2.5 mm), which suggests that

southern plants may well be perennial. Whereas stems of var. maritima usually are strongly ascending and

tend to have inflorescence branches along both sides, stems of var. virginica usually are weakly ascending

to procumbent and tend to have inflorescence branches along one side.

Seed number and shape are the most consistently reliable characters that we tested; the great majority

of specimens had seeds of only one type. We rarely found capsules of var. virginica to contain more than

two seeds, whereas capsules of var. maritima contain three or four seeds, never two. One specimen at NCU

(Windler 3279, Assateague Island, Worcester County, Maryland) had three seeds in most capsules, each seed

more-or-less three-sided but mildly concave ventrally. This same specimen also had a few capsules with two

seeds, also three-sided but strongly concave ventrally. Otherwise, this specimen resembled var. virginica in

its robust size, stem 2.7 mm in diameter, and dull brown sepals. A second specimen at NCU (Ahles 57756,

north of Fenwick Beach, Sussex County, Delaware) has some capsules with three seeds and some with two;

otherwise the plants match var. virginica. Thus, near the range limits of the two varieties of L. maritima, one

may expect to encounter specimens that show evidence of hybridization. Hodgdon included measurements

of seed length in his key; we did not evaluate this character. Sepal color exhibits tendencies (tinged maroon

northward, dull brown southward), but is difficult to apply consistently due to variation.

Hodgdon's key works reasonably well in separating var. virginica from var. maritima, but there is sig-

nificant overlap in his vegetative characters, so we have de-emphasized them. We include a modified key

here in which seed number, seed shape, and stem width are stressed.

J. Bot. Res. Inst. Texas 1(1): 367 — 368. 2007

368 Journal of the Botanical R h Institute of Texas 1(1))

Seeds 3-4(-5), weakly 3-sided and more-or-less resembling sections of an orange, or 2-sided and convex

ventrally; main stems 1.0-2.5 mm diameter, strongly ascending-erect to subprocumbent; sepals strongly

tinged maroon, occasionally dull brown; southern Maine to Delaware var. maritima

Seeds 2(-3), 2-sided and flattish, concave ventrally; main stems 2.0-4.0 mm diameter, procumbent to ascen-

ing; sepals dull brown, occasionally tinged maroon; southeastern Delaware to northeastern North Carolina

var. virginica

DISTRIBUTION AND RARITY

Hodgdon (1938) stated that var. maritima ranges from southern Maine to Delaware, but did not cite speci-

mens of the nominate variety, so we do not know where in Delaware var. maritima was taken. At the time of

his monograph, var. virginica was known only from Norfolk (now City of Chesapeake), Northampton, and

Princess Ann (now City of Virginia Beach) counties in Virginia. Currently this variety is known from eight

counties in Virginia, one in Maryland, one in Delaware, and one in North Carolina. Although apparently

rare in the latter three states, var. virginica is of frequent occurrence in Virginia. lt occurs within several

national and state protected areas with much suitable habitat and is not under any acute range-wide threat,

although seashore development poses a severe threat outside of refuges. Therefore, we rank it G5T3, using

criteria developed by NatureServe. Representative specimens are cited below.

DELAWARE. Sussex Co.: sand dunes, 3.6 mi N of Delaware-Maryland line on Del. 14, N of Fenwick Beach, Ahles 57756 with Baird

(NCU). MARYLAND. Worcester Co.: Assateague Island, stabilized dunes with Hudsonia, two mi south of paved road, Hill 15741 (NCU);

sand dunes just N of Ocean City on Md. 528, Ahles 57730 with Baird (NCU). NORTH CAROLINA. Dare Co.: Nags Head, on US 158

bypass, about 2 mi N of US 64, in rear dune zone, Kindell 477 (NCSC); Jockeys Ridge State Park, sound side, maritime dry grassland,

Kirkman (report to NC Natural Heritage Program). VIRGINIA. Accomack Co.: Assateague Island, Harvill 15113 (NCU); Chincoteague

National Wildlife Refuge, Assateague Island, Fleming 12644 (WILLD. City of Chesapeake Co.: near Ocean View, Kearney, Jr. 1001 (US,

cited in Hodgdon 1938). City of Virginia Beach Co.: sandy lot, Atlantic Blvd, Ware 7371 with Kral (VPD; N of Virginia Beach, Fleming

10040 (WILLD; Back Bay National Wildlife Refuge, Chamberlain 25-11 (VPD. Lancaster Co.: cited in Terwilliger et al. 1991. Matthews

Co.: Diggs Beach, Montfrans 165 (WILLD. Middlesex Co.: N 685 (WILLD. Northampton Co.: near Kiptopeake Beach, Harvill 15363

(NCU). Northumberland Co.: cited in Terwilliger et al.1991.

REFERENCES

BARRINGER, K. 2004. New Jersey pinweeds (Lechea, Cistaceae). J. Torrey Bot. Soc. 131:261-276.

HODGDON, A.R. 1938. A taxonomic study of Lechea. Rhodora 40:29-69, 87-131.

TERWILLIGER, K. 1991. Virginia's endangered species. McDonald and Woodward, Blacksburg, VA.

WILBUR, R.L. and H.S. Daoup. 1961. The genus Lechea (Cistaceae) in the southeastern Unites States. Rhodora

63:103-118.

RECOGNITION OF LECHEA PULCI LLA VAR. RAMOSISSIMA (CISTACEAE)

Bruce A. Sorrie Alan S. Weakley

University of North Carolina Herbarium University of North Carolina Herbarium

North Carolina Botanical Garden, CB 3280 North Carolina Botanical Garden, CB 3280

Chapel Hill, North Carolina 27599-3280, U.S.A. Chapel Hill, North Carolina 27599-3280, U.S.A.

ABSTRACT

A new combination is made: Lechea pulchella Raf. var. ramosissima (Hodgdon) Sorrie & Weakley.

RESUMEN

Se hace la nueva combinación: Lechea pulchella Raf. var. ramosissima (Hodgdon) Sorrie & Weakley.

Hodgdon (1938) revised the genus Lechea and recognized three varieties within L. leggettii Britton & Hol-

lick. The nominate variety (named var. typica Hodgdon), ranges from eastern Massachusetts to northeastern

Ohio and south to northern and western Virginia. Variety moniliformis (E.P. Bicknell) Hodgdon occurs near

the coast from southeastern Massachusetts to southern New Jersey; localized disjunct populations occur

within old shorelines of the Great Lakes in Illinois, Indiana, Michigan, Ohio, and Ontario. Variety ramosis-

sima Hodgdon occurs from southeastern Virginia to central Florida and west to eastern Texas. In a revision

of Lechea in the southeastern United States, Wilbur and Daoud (1961) accepted Hodgdon’s var. ramosissima,

but without assessing its distinctiveness.

Wilbur (1966) argued that Rafinesque's older names should be used for many of the names used by

Hodgdon and others. Thus, L. leggettii became a synonym of L. pulchella Raf. However, in this paper Wilbur

declined to recognize varieties, stating that *...l am not convinced that these tendencies represent biological

varieties or subspecies.” In a study of Lechea in New Jersey, Barringer (2004) did not recognize varieties and

did not mention the southern taxon ramosissima.

MATERIALS AND METHODS

We examined over 150 specimens from NCU, VPI, and WILLI. Specimens represented a broad geographic

range from Massachusetts to Florida and Louisiana; however, we paid special attention to the potential zone

of overlap of var. pulchella and var. ramosissima in the mid-Atlantic states. Specimens represented 13 coun-

ties in Virginia, 28 in North Carolina, and 3 in Maryland. We assessed Hodgdon's (1938) key characters

of seed number, seed shape, density of capsules, panicle shape, and panicle branching. We also assessed

plant height.

RESULTS

We found that these characters, if used in combination, satisfactorily separated a high percentage of speci-

mens with mature fruit. The most useful characters were seed number, seed shape, density of capsules, and

plant height. Table 1 summarizes these characters. Density of capsules refers to the arrangement of capsules

on the ultimate branches: in var. pulchella they vary from a tight cluster (or glomerule) to a crowded row

of capsules that actually or nearly touch one another; in var. ramosissima the capsules vary from a loosely

arranged row (capsules separated by one to a few mm) to a row of capsules that may touch one another.

Panicle shape and branching, as defined by Hodgdon (1938), were difficult to apply effectively, but we in-

clude them here for completeness.

We experienced little difficulty in assigning specimens to var. pulchella or to var. ramosissima. All speci-

mens from North Carolina and southward exhibited characters of var. ramosissima, with no characters of var.

J. Bot. Res. Inst. Texas 1(1): 369 — 371. 2007

370 Journal of the Botanical R h Institute of Texas 1(1))

Taste 1. Morphological characters used to distinguish varieties in Lechea pulchella.

var. pulchella var. ramosissima

seed number 3(-4) 2(-3)

seed shape relatively narrow and 3-sided, broad and compressed, or obscurely

like sections of an orange 3-sided

density of capsules clustered at branch tip, or in a in a sparse row, sometimes in a

dense row dense row

plant height 25-55 cm 35-80 cm

panicle shape and branching ovoid to suboylindric; principle subcylindric to subglobose; principle

branches subequal, relatively short branches diminishing upward, relatively long

pulchella. Similarly, specimens from Maryland and northward exhibited characters only of var. pulchella (or of

var. moniliformis). Virginia specimens from Caroline County (Wieboldt 9538 VPI) and City of Suffolk, (Fleming

11235 WILLD had three seeds per capsule and seeds variously shaped, but had capsule density and plant

height characters typical of var. ramosissima. Virginia specimens from Fairfax County (Hunnewell 6506 VPI)

and Prince William County (Townsend 3532 VPI) had three seeds per capsule and seeds variously shaped,

but had capsule density and plant height typical of var. pulchella. These specimens apparently are examples

of hybridization. Examination of additional specimens in the zone of overlap in eastern Virginia—and in

adjacent Maryland and North Carolina—may reveal additional evidence of natural hybridization. However,

since a small percentage of collections appear to exhibit intermediate characters, we believe that recognition

of var. ramosissima is warranted, based on the concept of varieties as incompletely separated evolutionary

entities with correlated morphological and ecogeographic differences. In this regard, we stress the impor-

tance of using several key characters when identifying varieties within Lechea pulchella, since there is some

overlap in any given character.

While many species in the eastern United States exhibit little variation from north to south (Liriodendron

tulipifera L.), others exhibit a clinal pattern (Limonium carolinianum (Walter) Britton). Still other species exhibit

a more-or-less distinct shift of characters in the region of the Virginia-North Carolina boundary. Kalmia

angustifolia L. and K. carolina Small, considered species by some and varieties by others, is an example.

Lechea pulchella Raf. var. ramosissima (Hodgdon) Sorrie & Weakley, comb. nov. Basionvm: Lechea pulchella Raf.,

New Fl. N. Amer. 1:91. 1836. Lechea leggettii Britton & Hollick var. ramosissima Hodgdon, Rhodora 40:119-123, pl. 491, fig. 3. 1938.

Tyre: U.S.A. Mississippt. Jackson Co.: Ocean Springs, 29 Jul 1896, Pollard 1109 (notorvee: GH; isotypes: E, NY, US).

Lechea pulchella var. ramosissima ranges on the coastal plain from southeastern Virginia (Accomack, Caroline,

City of Suffolk, City of Virginia Beach, Greensville, and Northampton counties; specimens at NCU, VPI,

WILLD south to central Florida (reaching Hernando and Martin counties, Atlas of Florida Vascular Plants,

www.plantatlas.usf.edu) and west to western Louisiana (Beauregard Parish, specimen at NCU) and eastern

Texas (Orange County, Turner et al. 2003); disjunct in central Tennessee (Coffee County, Wilbur & Daoud

1961) and northeast Georgia (Rabun County, specimen at NCU). Habitats include dry to mesic pine-oak

woods, pine savannas, pine flatwoods, borders of shrub-tree pocosins, moist powerlines, dry fields, roadsides,

and railroad rights-of-way. In many of these habitats, fire is a recurring disturbance.

REFERENCES

BARRINGER, K. 2004. New Jersey pinweeds (Lechea, Cistaceae). J. Torrey Bot. Soc. 131:261-276.

HODGDON, A.R. 1938. A taxonomic study of Lechea. Rhodora 40:29-69, 87-131.

Sorrie and Weakley, Notes on Lechea maritima var. virginica 371

Turner, B.L., H. NicHoLs, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. Volume 1. Botanical

Research Institute of Texas Press, Fort Worth.

WILBUR, R.L. 1966. Notes on Rafinesque's species of Lechea (Cistaceae). Rhodora 68:192-208.

WILBUR, R.L. and H.S. Daoup. 1961. The genus Lechea (Cistaceae) in the southeastern Unites States. Rhodora

63:103-118.

372 Journal of the B ical R h Institute of Texas 1(1))

BOOK REVIEWS

BARRIE JUNIPER and Davip J. MarseErLEY. 2006. The Story of the Apple. (ISBN: 0-88192-784-8, hbk.). Timber

Press Inc, 133 S.W. Second Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: www.tim-

berpress.com, mail@timberpress.com, 503-227-2878, 503-227-3070 fax, 1-800-327-5680). $29.95,

240 pp., 20 color photos, 40 illustrations, 9 maps, 7" x 9".

Have you ever wondered where apples come from? Yes, those enjoyably crisp and juicy apples that we're told are as American as, well,

apple pie. The answer has been provided in the new book entitled The Story of the Apple by authors Barrie Juniper and David Mabberly.

The Story of the Apple is a richly researched book loaded with references to support the author's work. This well-documented history of

the apples' origins and geographical dissemination throughout history also includes many beautiful images of the people, places and

equipment involved; it contains a mixture of both color and black & white images.

The book begins by providing readers with the foundation of apple knowledge; i.e. the title of chapter one is *What are apples?"

Chapter two, entitled “Origin of the apple,” includes information on pollinators, scientific nomenclature, cropping phases and dispersal

methods (bears and horses especially). The authors trace the origin of the sweet apples of today to the Tian Shan fruit forest in the

mountains of Central Asia. In chapters three and four, the authors describe how early humans and animals interacted with apple fruit,

wood and seeds; and include the history of grafting of fruit and apple trees throughout history. The fifth and sixth chapters focus on

the movement of apples from Asia to Europe and into North America via land and sea routes. The fifth and sixth chapters also include

information on the cultural influence of the apple in place names and family names, as well as in various cultural folklore. In chapter

seven, an interesting history of apple cider and other apple preservation techniques used around the world is provided. The final chapter

provides readers with an overview.

The authors have been very thorough in providing referenced information in this text; the end result requires readers to pay more

attention to detail. For example readers must first keep in mind the different species the authors discuss: there are crab apples (Malus

sylvestris) and more commonly known sweet apples (Malus pumila). However, such details are Hunc to the points being made and

1 1 T3

add significantly to the valuable information provided in this text. This reader found tl interesting regarding the topics

of fruit-tree grafting, use of the word apple (and non-English lingual equivalents) and apple SOT Along with color plates of various

apple species, the book contains many other interesting images; among them are black and white woodcuts of historic cider making

equipment.

Apple historians, horticulturalists and enthusiasts will delight in The Story of the Apple. Authors Juniper and Mabberly have pro-

vided a well-researched history of the apple including the origins of the sweet apple as well the many uses of the apple fruit and wood in

various cultures worldwide. Color plates, photographs and black and white images add significantly to e the importance

of apples all over the world and throughout history. Explore The Story of the Apple and read about th i journey of the apple

from Asia to every corner of the Earth.—Lee Luckeydoo, Herbarium, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX

76102-4060, U.S.A.

Jim KimmeL (text) and Jerry ToucHsTone KimMEL (photographs). 2006. The San Marcos: A River's Story. (ISBN

13: 978-1-58544-542-4, pbk.). Texas A&M University Press, 4354 TAMU, College Station, TX 77843-

4354, U.S.A. (Orders: http://www.tamu.edu/upress/, 979-458-3982, 979-847-8752 fax). $29.95, 155

pp., 184 color photos, 26 b/w photos, 1 drawing, 1 chart, 4 maps, 85/5" x 1014".

What do mastodons, treed Gypsy bears, Texas wildrice, blind salamanders, and Ralph the Swimming Pig have in common? They are

all part of the story of the San Marcos River, an iconic river in Central Texas. Jim Kimmel affectionate and detailed text and his wife

Jerry's beautiful photos bring to life the natural and cultural history of a river whose headwaters are thought to be the site of some of the

oldest human settlements - 12,000 years old — on the North American continent.

The book examines the San Marcos River from several perspectives: geology, geography, history, biology, and botany. It covers

the river's history from the uplift that created the Balcones Fault millions of years ago, creating the conditions for the springs to issue,

to the present day efforts to protect the Edwards Aquifer, which nourishes the river, from excessive pumping. The peoples who have

lived, and live today, on its banks are c to life. Native and endangered plants and animals as well as sometimes pesky nonnative

pecies are discussed in the text and listed in the attract dices. The reader is left with an understanding and appreciation of the

San Marcos river's beauty and diversity.

Jim Kimmel, one of Texas' most distinguished geographers, is a professor at Texas State Nds in San Marcos and directs the

university's Center for Nature and Heritage Tourism. His family has lived in the San l generations. Jerry Touchstone

Kimmel is a nationally known watercolor artist and accomplished photographer.

The reviewer, a former San Marcos resident, can almost feel the tingle of the 72° water on her toes as she writes this review.

Hmmm...a leisurely float from just below the dam at Spring Lake to Rio Vista Dam seems in order for this summer.—Penny McCook,

Volunteer, Botanical Research institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A.

J. Bot. Res. Inst. Texas 1(1): 372. 2007

SEED AND CAPSULE MORPHOLOGY IN SIX GENERA

OF HEDYOTIDEAE (RUBIACEAE): THECAGONUM, NEANOTIS, DENTELLA,

KOHAUTIA, PENTODON, AND OLDENLANDIOPSIS

Edward E. Terrell ' Harold Robinson

Research Associate Curator

Department of Botany Department of Botany

National Museum of Natural History National Museum of Natural History

Smithsonian Institution mithsonian Institution

Washington, DC 20013, U.S.A. Washington, DC 20013, U.S.A.

ABSTRACT

Morphological variation in the seeds and capsules of selected species of six genera of Hedyotideae (Rubiaceae) is described and il-

lustrated by scanning electron microscopy. Oldenlandioid (trigonous) seeds occur in the genera Dentella and Pentodon. The numerous

areolar tubercles in Pentodon seeds are shown in enlarged views. The seeds in two species of Kohautia have distinctive rounded areolar

protrusions. Oldenlandiopsis has unique capsules and seeds, the latter oblate. The sulcate seeds of Thecagonum, an Asian segregate from

Oldenlandia, have several rounded or elongate depr

ssions bordered by strongly sinuous walls. Oldenlandia strigulosa, with similar seeds,

is formally transferred to Thecagonum. The Asian genus, Neanotis, a segregate from Anotis, has houstonioid seeds with a hilar ridge in a

ventral depression.

RESUMEN

] n = il E Al qd

«lx r

is géneros de Hedyotideae (Ru-

Se describe e ilustra la variación morfológica en las semillas y |

ry, £3 H D 1 3

LCILLUQGUTIL.

TE 1; ; lectore daba ridoni +1] Iden] Mod í N nlos géneros

o o

Los tubérculos areolares numerosos en las semillas de Pentodon se muestran a gran aumento. Las semillas de dos especies de Kohautia

tienen salientes areolares redondeados distintivos. Oldenlandiopsis tiene las cápsulas y semillas únicas, la última de ellas obladas. Las

semillas sulcadas de Thecagonum, un segregado asiático de Oldenlandia, tienen varias depresiones redondeadas o alargadas confinadas

por paredes fuertemente sinuosas. Oldenlandia strigulosa, con las semillas similares, se tran sfiere formalmente a Thecagonum. El género

asiático, Neanotis, segregado de Anotis, tiene las semillas houstonioid n una costilla hilar en una depresión ventral.

INTRODUCTION

This study continues investigations of seeds and capsules in the tribe Hedyotideae by means of scanning

electron microscopy (SEM), studies that began with Terrell, Lewis, Robinson, and Nowicke (1986). The

present paper describes and illustrates morphological variation in seeds and capsules of Thecagonum, Ne-

anotis, Dentella, Kohautia, Pentodon, and Oldenlandiopsis. Previous studies of Oldenlandia seeds (Terrell &

Robinson 2006) provided information about the trigonous (3-angled) seeds of that genus. Two of the genera

studied here, Dentella and Pentodon, have trigonous seeds, and the other four genera have seeds with varying

distinctive characteristics

MATERIALS AND METHODS

The herbarium of the U.S. National Museum (US), Smithsonian Institution, Washington, D.C., provided

most of the capsules and seeds for this study. One or more samples of each species were examined under a

dissecting microscope to determine sizes, shapes and other characters. Selected samples were mounted on

stubs and examined with scanning electron microscopes located at the Electron Microscope Laboratories at

the Smithsonian Institution or at the U.S. Department of Agriculture, Beltsville, Maryland. The information

provided for each genus includes names, synonyms, general distributions, and descriptions and illustrations

of seeds and capsules.

‘Address for correspondence: 14001 Wildwood Drive, Silver Spring, Maryland 20905, U.S.A.

J. Bot. Res. Inst. Texas 1(1): 373 — 384. 2007

374 Journal of the Botanical R h Institute of Texas 1(1)

RESULTS AND DISCUSSION

1. DENTELLA J.R. & G. Forst., Char. Gen. Pl. 13.1775. Wee seecies: Dentella repens (L.) J.R. & G. Forst., Char. Gen. Pl. ed.

2, 26. t. 13.1776. Oldenlandia repens L., Mant. Pl. 40.1767.

Distribution.—One or two species in S.E. Asia, Sri Lanka, Malesia, Taiwan, Micronesia, Polynesia (Ridsdale

1998). Adventive in coastal western Mexico (EJ. Lott 3116 from Jalisco and other Mexican collections in

herbarium US). Also locally adventive in Baltimore, Maryland (Reed 1970).

Chromosome number.—Dentella repens: 2n = 36 (Raghavan & Rangaswamy 1941).

Dentella repens resembles Oldenlandia species, but differs in having 5-merous flowers. The plants are

small creeping annual or perennial herbs. Leaves small, ca. 2-10 mm long, oblanceolate to obovate. Stipules

interpetiolate, to 1 mm long. Flowers solitary, axillary, isostylous. Corolla 3-4 mm long, narrowly tubular,

white.. Anthers ca. 0.5 mm long. Stigmas bifid.

Capsules 2.5-4 x 2.5-3 mm, subglobose, densely hairy, crowned with calyx lobes, fully inferior, walls

thin, fragile. They are reputed by Ridsdale (1998) to be indehiscent. Seeds (Table 1) numerous per capsule,

0.3-0.6 mm in diam.or slightly longer than wide, trigonous or obtusely angulate, basal face elliptical, lat-

eral faces larger, somewhat concave, hilum punctiform, apical, areoles polygonal, walls thick, testa surface

featureless (Figs. 1 A,B). The trigonous seeds are similar to those in Oldenlandia, thus are oldenlandioid.

Collection examined using SEM.— Clemens 3276 (US), Indo-China. Also observed were E.J. Lott 3116 (US)

and A. C. Sanders et al. 10597 (UCR), both from Jalisco, Mexico.

2. KOHAUTIA Cham.& Schltdl., Linnaea 4:156.1829. Tee species: Kohautia senegalensis Cham. & Schltdl., Linnaea

4:156.1829.

Distribution.—The genus has about 60 species in Africa, Madagascar, and tropical Asia, with 12 species

in Tropical East Africa (Verdcourt 1976). In addition, Halford (1991) described one species in Australia.

Verdcourt (1989) recorded 13 species from the Flora Zambesiaca region.

Chromosome number.—Lewis (1965) reported the chromosome number of Kohautia as x = 9. He found

pollen grains to be 3 to 8-colporate and relatively small, and stated that nearly all species of Kohautia were

easily separated by their pollen from those of Oldenlandia and other African taxa in the tribe Hedyotideae.

Our SEM study of Kohautia has been restricted to one species and, therefore, the genus was excluded

from Table 1. We found an unusual seed type which we consider worth reporting because of the areolar

protrusions.

Kohautia coccinea Royle, Ill. Bot. Himal.: 241, t. 53/1.1835. It occurs from Africa to India. It is an an-

nual herb with pink to purple flowers usually in spikes or racemes, corolla 5-11 mm. long, tubes narrowly

cylindrical. Seeds of K. coccinea examined using SEM: Burger 3545 (US), Ethiopia.

Capsules 3-5.5 mm long, oblong-ellipsoid, loculicidally dehiscent (Verdcourt 1976). Seeds 0.6-0.7

x 0.35-0.4 mm, black, obtusely angulate, hilum apical, areoles polygonal, their walls thin to rather thick,

each areole containing a conspicuous large round protrusion (Fig. 1 C,D). These protrusions may occur in

other species in Kohautia. Halford (1992) illustrated a seed of the Australian species Oldenlandia spathulata

Halford which shows round protrusions in the areoles, but he did not comment on them. He placed O.

spathulata in a group of three species including O. spermacocoides (F. Muell.) F. Muell. and O. crouchiana (F.

Muell.) F. Muell. Specimens of these three Australian species of Oldenlandia were not available to us. It is

suggested that O. spathulata may belong in Kohautia.

3. OLDENLANDIOPSIS (Griseb.) Terrell & WH. Lewis, Brittonia 42:185.1990. Tre species: Oldenlandiopsis cal-

litrichoides (Griseb. / Tus & WH. Lewis, Brittonia 42:185.1990. Oldenlandia callitrichoides Griseb., Mem. Amer. Acad. Arts n.s.

.1863. Hedy llit ides (Griseb.) WH. Lewis, Rhodora 63:222.1961.

Distribution.—One species in West Indies, Central America and Mexico (Yucatan). Adventive in Africa

(Sierra Leone), northern South America (Guyana), United States in Dade County, Florida, and Hawaiian

Islands: Oahu and Maui.

A synopsis of the United States species of Oldenlandia (Terrell 1990) treated five species. Included

Terrell and Robinson, Seed and capsule morphology in tribe Hedyotideae

Table 1. Seed characters in Thecagonum, Neanotis, Dentella, Oldenlandiopsis, Pentodon.

375

Thecagonum Neanotis Dentella Oldenlandiopsis Pentodon

biflorum (4 species)

Number per numerous 2-20 or more numerous 20-35 numerous

capsule

Length mm 0.3-0.6 0.5-1.3 0.3-0.6 0.3-0.5 0.3-0.5

Shape, type ovoid, subglobose — cymbiform or conoidal oblate trigonous

shallow cup

Hilum punctiform low, linear ridge punctiform punctiform punctiform

Hilum location centric or acentric in shallow depression apical centric apical

Ventral irregularly sulcate reticulate reticulate reticulate reticulate

surface, type

Ventral surface several small polygonal areoles polygonal polygonal polygonal

rounded or elongate areoles areoles areoles

depressions

Walls/borders sinuous, straight/curved, straight sinuous/ bearing many

conspicuous not conspicuous straight minute tubercles

among these was Oldenlandia callitrichoides Grisebach, which Terrell and Lewis (1990) soon described as

Oldenlandiopsis callitrichoides (Griseb.) Terrell & W.H. Lewis.

The plants are creeping, soft and delicate, and may be superficially confused with Oldenlandia species,

but very few Oldenlandia species are creeping (versus merely prostrate).

Capsules 1.0-2.7 x 0.5-2 mm, narrowly turbinate or obconic, thin-walled, fragile, retuse or truncate,

9/10 or fully inferior, somewhat compressed, glabrous, dehiscing loculicidally and later separating into four

narrow segments. These capsules differ entirely from the characteristically subglobose, indurate capsules of

not only Oldenlandia but also from a number of other examined hedyotoid genera.

Seeds (Table 1) 20-35 per capsule, 0.3-0.5 x 0.2-0.4 mm, oblate (depressed-ellipsoid, depressed-sub-

globose), or obtusely angulate, hilum punctiform, centric, testa reticulate, areoles polygonal, walls sinuous

or straight, surface minutely papillose (Fig.1 E,F). The seeds are distinctive in being more or less oblate.

Collection examined using SEM: F. & B. Terrell 5022 (US), Dade Co., Florida.

There are two other basic differences that also distinguish Oldenlandiopsis (Terrrell & Lewis 1990) as

a distinct genus: (1) pollen with 8-colporate apertures instead of 3(—5), 2) chromosome number 2n = 22

(x = 11) instead of x = 9; the only other Oldenlandia species with x = 11 is Oldenlandia microtheca (Terrell &

Robinson 2006).

4. PENTODON Hochst., Flora 27: 552.1844. Tver species: Pentodon pentandrus (Schumach. & Thonn.) Vatke, Oesterr. Bot.

Z. 25:231.1875. Hedyotis pentandra Schumach. & Thonn., Kongel. Danske Vidensk. Selsk. Naturvidensk. Math. Afh. 3:71.1827.

Oldenlandia y (Schumach. & Thonn.) DC., Prodr. 4:427.1830, non Retz.

Hedyotis halei Torr. & A. Gray, Fl. N. Amer. 2:42.1841. Oldenlandia halei (Torr. & A. Gray) Chapm., Fl. Southern U.S. 181.1860. Pentodon

halei (Torr. & A. Gray) A. Gray, Syn. Fl. N. Amer. 1, 2:28.1884. The name Pentodon halei was created for American plants from col-

lections prior to 1841, but plants were found to be conspecific with P pentandrus.

Distribution.—Two species in Africa (see also Verdcourt 1976, 1989, Rogers 1987). Adventive in Florida and

southeastern U.S. west to eastern Texas. Also adventive in Cuba, Nicaragua, and Brazil.

The chromosome number is x = 9 based on Pentodon halei, n 2 9, 2n = 18 (Lewis 1962); and P. pentandrus,

n = 9 (Lewis 1965). Pollen is 3-aperturate and resembles most species of Oldenlandia (Lewis 1965).

Pentodon pentandrus var pentandrus is as follows: Annual or short-lived perennial fleshy herbs. Leaves

lanceolate to elliptic, 3-8 x 0.3-2.5 mm. Stipules interpetiolate, fimbriate, 0.5—5 mm long. Flowers 5-merous,

terminal or axillary, isostylous or heterostylous. Calyx and corolla 5-lobed, corolla funnelform. Stigmas bifid.

Capsules (Fig. 2 D) 2-4 x 23.5 mm, oblong, somewhat compressed, conspicuously nerved, thin-walled,

fragile, loculicidally dehiscent. Seeds (Fig. 2 A-C) numerous, 0.3-0.5 mm in diam., trigonous or irregularly

376 Journal of the Botanical Research Institute of Texas 1(1)

Fic. 1. Seeds examined by SEM. A-B. Dentella repens, Clemens 3276 (US), Indo-China. C-D. Kohauti inea, B. 3545 (US), Ethiopia. E-F. Oldenlandi-

opsis callitrichoides, E. & B. Terrell 5022 (US), Florida. A, end view; B, F, enl ] les; C, E tral vi les showi led protrusi

bw | yey 1 Yy be |

in tribe Hedyotideae 377

Terrell and Robinson, Seed and

1 dF

and obtusely angulate, hilum punctiform, apical, testa reticulate, areoles elongate or polygonal, areole walls

with numerous minute tubercles (Fig. 2 B,C). Seed collection examined using SEM: Thieret 16435 (SMU),

Louisiana.

Pentodon capsules are distinctive in being oblong with soft, compressed walls. The seeds (Table 1) are

trigonous, thus appear oldenlandioid. The noteworthy feature of the seeds are the numerous minute tubercles

attached to the areole walls, somewhat enlarged in Fig. 2. These tubercles were noted by Lewis, but have

not been previously illustrated. Such tubercles have not been seen in any other taxon of the tribe.

5. THECAGONUM Babu, Bull. Bot. Surv. India 11:214.1969. Tyee species: Thecagonum pteritum (BL) Babu. Gonotheca Bl.

ex DC., Prodr. 4: 429.1830 (non Raf., Med. Repos. 5:352.1808, Compositae). Babu (1969) proposed the new name, Thecagonum,

to replace the illegitimate name, Gonotheca Bl. ex DC., a later homonym. Babu separated this genus from Oldenlandia by its having

a 4-angled fruit, not terete, and seeds globose or subglobose, not angular. He recognized four species in Thecagonum. Specimens of

the fourth species, T. parishii (Hook.f.) Babu, native to India and Malesia, were not available to us.

1. Thecagonum biflorum (L.) Babu, Bull. Bot. Surv. India 11:214.1969. Oldenlandia biflora L., Sp. Pl. 119.1753.

Hedyotis biflora (L.) Lam., Tabl. Encycl. 1:272.1791.

Oldenlandia paniculata L., Sp. Pl. ed. 2. 1667.1763.

The synonymy here follows Fosberg & Sachet (1991) in their treatment of Micronesian species of Hedyotis

in which they delimited Hedyotis broadly with Oldenlandia as a subgenus.

Distribution.—This well known species usually called Oldenlandia biflora has a wide distribution from

India to China, Malesia, Micronesia, and Polynesia. Flora Vitiensis (Smith & Darwin 1988) stated “crevices

of arid rocks along coasts,” and listed distribution as tropical Asia to Mauritius, throughout Malesia, eastward

to Fiji, Tonga, Niue, and Samoa.

Collections examined using SEM.—Fosberg 39194 (US), Guam; Fosberg 33792 (USF), Marshall Islands;

Anderson 2126 (US), Caroline Islands.

Chromosome number.—Oldenlandia paniculata: n = 36 (Raghavan & Ragaswamy 1941); O. biflora: n = 18

(Lewis and Oliver 1970); 2n 2 54, 72 (Selvaraj 1987).

Perennial herbs. Stems erect to prostrate. Leaves 8-40 x 3-13 mm, lanceolate, elliptic, or ovate-lan-

ceolate. Stipules 1-2 mm, interpetiolate, margins with teeth or setae. Inflorescence terminal and axillary.

Corollas broadly tubular, tubes ca. 2 mm long, lobes ca. 1 mm long.

Capsules 2-4 x 2-4 mm, subglobose, somewhat compressed, walls thin and fragile, about 4/5 to fully

inferior, dehiscing loculicidally and septicidally.

Seeds (Table 1) numerous per capsule (ca. 65 in one capsule), 0.3-0.6 x 0.3-0.6 mm, ovoid, subglobose,

or obtusely angulate, hilum centric, punctiform, surface with crowded, variously-sized, shallow, rounded or

elongate depressions, their walls thick and strongly sinuous (Fig. 3). These depressions are entirely unlike the

conventional areoles of many species, and may be described as sulcate, a term defined by Kiger and Porter

(2001), as “Having one or more elongate, relatively narrow and shallow depressions (sulci).” These distinctly

different seeds support the removal of Oldenlandia biflora from the genus Oldenlandia and the recognition of

Thecagonum as a distinct genus

2. Thecagonum strigulosum (DC.) Terrell & H. Rob., comb. nov. Basioxvw: Oldenlandia strigulosa DC., Prodr. 4:427.1830.

Hedyotis strigulosa (DC.) Fosberg, Smithsonian Contr. Bot. 45:28.1980.

Hedyotis coreana H. Lév., Repert. Spec. Nov. Regni Veg.11:64.1912.

Oldenlandia albido-punctata Merr., Philipp. J. Sci., C, 9:297.1914.

This species was not included in Thecagonum by Babu (1969), however, it was recognized as closely related

to Hedyotis biflora by Fosberg and Sachet (1991). They provided a complete description of it and in their

key to species distinguished Hedyotis strigulosa from H. biflora in characters of the leaves, inflorescence, and

capsules. They described capsules as being firm, thick-walled, and broadly ovoid, compared to H. biflora

which is thin-walled and subglobose. Hedyotis strigulosa seems not to have been transferred to Thecagonum,

so we do so here.

378 Journal of the Botanical Research Institute of Texas 1(1)

cns

(Tobi qm

ANM A pA >

Fic. 2. Seeds and capsules of Pentodon pentand ined by SEM. A-C. Thieret 16435 (SMU), Louisiana. D. Lewalle 1527 (US), Burundi. A, seed, end

I J I L . EL I n

view: B, C capsules.

1] Ney 1 y F

Our seeds are approximately 0.4-0.6 mm long and have rounded or elongate depressions like the seeds

of T. biflora, but differ in the seeds being mostly obtusely angulate.

Collections examined using SEM.—Moran 4618 (US) and Necker 362 (US), both from Guam, Marianas

Islands. These two collections were cited as Hedyotis strigulosa by Fosberg and Sachet (1991).

3. Thecagonum pteritum (Bl.) Babu, Bull. Bot. Surv. India 11:214.1969. Hedyotis pterita BL., Bijdr. 972.1826. Gonotheca

blumei DC., Prodr. 4: 429.1830, nom. illeg. Oldenlandia pterita (Bl.) Miq., Fl. Ind. Bat. 2:193.1857.

Succulent herb to 10 cm tall. Leaves 3-6 x 0.5 cm, elliptic or lanceolate. Flowers terminal and axillary.

Capsules 4-7 x 4-6 mm, oblong or obovate, somewhat compressed, walls fragile, winged, wings appar-

ently two per capsule, to ca. 1 mm wide (Fig. 4 C). The capsules resemble those of Pentodon pentandrus in

general shape and texture, thus they differ from the capsules of the two preceding species of Thecagonum.

The epithet, pterita, refers to the winged capsules.

Seeds numerous per capsule, 0.5-0.6 x 0.4-0.5 mm, broadly ellipsoid-angulate or obtusely angulate,

hilum punctiform or slightly elongate, centric, surface with variously-sized rounded or elongate depres-

sions, walls thick, sinuous, testa densely papillose (Fig. 4 A, B). The seeds are generally similar to those of

T. biflorum.

in tribe Hedyotideae 379

1 "id

L| ry)

m ue

' A

"nnm

g' t

‘

SMS

Fic. 3. Seeds of Thecagonum biflorum examined by SEM. A-D. Fosberg 39194 (US), Guam. E-F. Fosberg 33792 (USF), Marshall Islands. A, C, E, surface

J . lo: Ļ J D hil £ r4 n F I rR £4 . horders J . 4 cnn

Distribution.—India to Malesia.

Collections examined using SEM.—Ramos s.n., Apr 1909 (US), Luzon, P.I.; Merrill 6730 (US), Negros, P.I.

Oldenlandia ovatifolia (Cav.) DC., Prodr. 4:427.1830. Hedyotis ovatifolia Cav., Icon. 6:52.1801.Thecagonum ovatifolium

(Cav.) Babu, Bull. Bot. Surv. India 11:214.1969.

Oldenlandia nudicaulis Roth, Nov. Pl. Sp. 95. 1821. Hedyotis nudicaulis (Roth) Wight & Arn., Prodr. 416.1834.

This species was listed under Thecagonum by Babu (1969), however, its seeds lack the depressions typical

of T. biflora, T. strigulosa, and T. pterita, and instead are reticulate with polygonal areoles like those in other

380 Journal of the Botanical Research Institute of Texas 1(1)

Fic. 4. Seeds and capsules examined by SEM. A-C. Thecagonum pteritum, A-B. Ramos s.n., Apr 1909, pale BIG ner al (US), AOS i D- i

pie pL E, eM Sumatra; F, Belsher 558 (US), Upper Burma. A, dey

psules; D, seed, side view; E, enlargement of polygonal areoles.

genera. The capsules generally resemble those of some species of Oldenlandia (Fig. 4 F). The seeds are co-

noidal. We suggest that this species be retained in Oldenlandia.

Annual herb, stems 4-20 cm tall, leaves broadly ovate, appearing conspicuously large compared to the

entire plant, inflorescence cymose, corolla tubes 0.5-1 mm long, lobes 1.5-2 mm long. Capsules 2-2.5 x

Terrell and Robinson, Seed and capsule morphology in tribe Hedyotideae 381

2.5—3 mm, subglobose or slightly wider than long, walls fragile, thin, 7/8 to fully inferior. Seeds 30-40 per

capsule, 0.4-0.5 x 0.4-0.5 mm, conoidal or irregularly conoidal, hilum apical, punctiform, areoles polygonal,

usually longer than wide, walls thick, testa densely papillose (Fig.4 D,E).

Distribution.—India to Java and Malesia.

Collection examined using SEM.—Toroes 4448 (US), Sumatra; Belsher 558 (US), Upper Burma. Two other

collections were also studied: Toroes 1804 (US), Sumatra; Toroes 3514 (US), Sumatra.

6. NEANOTIS WH. Lewis, Ann. Missouri Bot. Gard. 53:34.1966. Te species: Neanotis indica (DC.) WH. Lewis, Ann.

Missouri Bot. Gard. 53:38.1966. Putoria? indica DC., Prodr. 4:577.1830.

Distribution.—Southeast Asia and Malesia (Ridsdale 1998). There are 28 species in genus (Lewis 1966;

Ridsdale 1998).

Lewis (1966) removed 28 species from Anotis and placed them in a new genus, Neanotis, but did not

provide a description of the new genus. The validity of the name Neanotis has been questioned, however,

we follow Saldanha and Nicolson (1976) and Ridsdale (1998) in accepting its validity. One of our reviewers

(Kirkbride) advises that the name Neanotis is valid because there is an indirect reference to a description.

Lewis (1966) found that the pollen of Neanotis differs by six important characters from that of Hedyotis

and Oldenlandia. He concluded that Neanotis pollen is distinct and differs from all other members of the

Hedyotideae.

Chromosome numbers.—x = 9. N. hirsuta var. glabricalycina (Honda) W.H. Lewis, 2n = 36, Taiwan (Lewis

et al. 1967); N. gracilis (Hook. f.) W.H. Lewis, 2n = 18, Nepal (Malla et al. 1978).

Species studied, with synonyms, and distributions are as follows:

1. Neanotis calycina (Hook.f.) WH. Lewis, Ann. Missouri Bot. Gard. 53:37. 1966. Anotis calycina Hook.f., Fl. Brit.

India 3:73.1880.

Distribution.—Himalayas, India, Burma (Myanmar), Vietnam, China.

Collection treated using SEM.—Stainton 1590 (BM), Nepal. Other examined collections not using SEM

were Stainton et al. 7107 (BM), Nepal; Henry 13512 (K, US), China.

2. Neanotis hirsuta (L.f.) WH. Lewis, Ann. Missouri Bot. Gard. 53:38.1966. Oldenlandia hirsuta L.f., Suppl. PL. Syst.

Veg. 127.1782.

Distribution.—Himalayas to China, Japan, Malaysia.

Collection examined using SEM.—Mousset 271 (US), Java.

3. Neanotis monosperma (Wight & Arn.) WH. Lewis, Ann. Missouri Bot. Gard. 53:40.1966. Hedyotis mono-

sperma Wight & Arn., Prodr. 410.1834.

Distribution.—India and Sri Lanka.

Collection examined using SEM.—Tirvangadum 514 (K, US), Sri Lanka. Collection examined but not us-

ing SEM: Fosberg 50050 (US), Sri Lanka. Approximately 30 other US collections from Sri Lanka were noted;

most of these lacked mature flowers or fruit.

4. Neanotis tubulosa (G. Don) Mabb., Taxon 29:606.1980. Oldenlandia tubulosa G. Don, Gen Hist. 3:531.1834. Neanotis

quadrilocularis (Thwaites) W.H. Lewis, Ann. Missouri Bot. Gard. 53:40.1966. Hedyotis quadrilocularis Thwaites, Enum.Pl. Zeyl.144.1859.

Anotis quadrilocularis (Thwaites) Hook.f., Fl. Brit. India 3:74.1880.

Distribution.—India, Sri Lanka.

Collection examined using SEM.—Saldanha 17811 (US), India. Neanotis tubulosa (formerly N. quadrilocularis)

differs from the other three species in having a 4-loculate instead of a 2-loculate capsule and seeds with

a slightly deeper depression surrounded by a thicker margin. This species has been generally accepted as

congeneric with other Neanotis species.

Morphological data from N. calycina are briefly as follows: Herbaceous annual; stem 5-25 cm tall; leaves

ovate-lanceolate, 0.5-3.5 x 0.2-1.5 cm, flowers in axillary or terminal cymes, corollas white, pink, or purple,

infundibular-cylindrical, 2-3 mm long.

£+sha D o ID L

382 Journal of t titute of Texas 1(1)

TABLE 2. Seed characters in four Neanotis species.

Characters calycina monosperma hirsuta tubulosa

Length mm 0.5-0.7 0.9-1.2 1.1-1.3 0.6-0.7

Thickness thick thick rather thick thick

Shape, kind ymbiform cymbiform cymbiform/saucer cymbif./shallow cup

Shape in outline suborbicular/oblong/ell. oblong, ell., suborbic. suborbic/elliptic suborbic/braly. ell.

Compression slight to moderate moderate moderate moderate

Dorsal face rounded/convex strongly rounded rounded/convex convex

Hilar ridge low, linear low, linear low, linear low, shortly linear

Depression medium to shallow shallow shallow or flattish small cup

Margin shape thickened/rather thin slightly or not thickened low/ flattish thickened

Areoles texture coarse coarse coarse coarse

Areoles walls indurate indurate indurate indurate

The apically dehiscent capsules of Neanotis species are generally of a subglobose type, but differ in being

either slightly longer than wide (as in N. richardiana (Arn.) W.H. Lewis) or wider than long (as in N. mono-

sperma). They are relatively small and have paratively few of the rather bulky seeds (ca. 2-20 per capsule).

We examined seeds of the four species by SEM (Table 2). An inclusive morphological description (Table

1) is as follows: Seeds (Fig. 5) 0.5-1.3 mm long or wide, dull black or dark brown, thick, cymbiform to shal-

lowly cup-shaped, in outline suborbicular, oblong, or broadly elliptic, compression moderate, dorsal face

strongly rounded to convex, ventral face with a low linear hilar ridge in a shallow to medium-sized depres-

sion, margin thickened or flattish, hilar ridge slightly higher to slightly lower than the bordering margin,

areoles polygonal, usually appearing coarse and indurate. The seeds usually have a “chunky” appearance

because the dorsal face (Fig. 5 F) is often rounded.

The seeds of Neanotis resemble those of Houstonia, particularly the H. purpurea group (subgenus

Chamisme) of four species in eastern North America (Terrell 1996), but are thicker and coarser. Plants of the

Neanotis species examined here also resemble the H. purpurea group in being small to medium-sized peren-

nial (or annual) herbs with lanceolate or ovate leaves. Other data, however, do not favor a close relationship:

pollen morphology is distinctly different as noted previously, and the chromosome numbers are x = 6 for

H. purpurea and x = 9 for Neanotis. If the two groups originated from the same basic stock, they underwent

long isolation on separate continents.

KEY TO SEEDS AND CAPSULES OF THE STUDIED GENERA

1. Seed surface not reticulate, lacking areoles, with several shallow rounded or elongate depressions 1 Thecagonum

1. Seed surface reticulate, areoles (cells) numerous, rounded or polygonal.

2. Seed surface with areoles each with a large rounded protrusion Kohautia

2. Seed surface with areoles lacking a rounded protrusion.

3. Seeds 3-angled (trigonous, conoidal).

4. Seeds with areole walls bearing numerous small tubercles; capsules compressed, oblong, thin ^ Pentodon

4. Seeds with areole walls lacking tubercles; capsules subglobose Dentella

3. Seeds compressed or ovoid, not 3-angled.

5. Seeds oblate, lacking a hilar ridge Oldenlandiopsis

5. Seeds somewhat compressed, with a hilar ridge in a ventral depression Neanotis

ACKNOWLEDGMENTS

We thank Scott Whittaker and Susann Braden, Electron Microscope Laboratory, Smithsonian Institution,

for preparation of SEM illustrations, and Marjorie Knowles for preparing the figures. We also thank the

curators of the herbaria cited in the text. Joseph Kirkbride and John Wiersema provided very helpful reviews

of the manuscript.

Terrell and Robinson, Seed and capsul I gy in tribe Hedyotideae 383

NT PLA

Fic. 5. Seeds of M is speci ined by SEM. A, B, N. hirsuta, Mousset 271 (US), Java. C, D, N. monosperma, Tirvangadum 514 (US), Sri Lanka. E, F,

N. tubulosa, Saldanha 17811 (US), India. A—C, E, ventral vi „enl t of areol 1 mi illae; F, dorsal vie

J ret

REFERENCES

Basu, C.R. 1969. Thecagonum Babu—a new generic name in Rubiaceae. Bull. Bot. India 11:213-214.

FosgerG, F.R. and M.-H. Sacher. 1991. Studies in Indo-Pacific Rubiaceae. Allertonia 6:191-278.

HALFORD, D.A. 1991. The genus Kohautia Cham. & Schlecht. (Rubiaceae) in Australia. Austrobaileya 3: 439-442.

384 Journal of the Botanical R h Institute of Texas 1(1)

HALFORD, D.A. 1992. Review ofthe genus Oldenlandia L. (Rubiaceae) and related genera in Australia. Austrobaileya

3:683-722.

Kicer, RW. and D.M. Porter. 2001. Categorical Glossary for the Flora of North America Project. Hunt Institute for

Botanical Documentation, Carnegie Mellon University, Pittsburgh, PA.

Lewis, W.H. 1962. Phylogenetic study of Hedyotis (Rubiaceae) in North America. Amer. J. Bot. 49:855—865.

Lewis, W.H. 1965. Cytopalynological studies of African Hedyotideae (Rubiaceae). Ann. Missouri Bot. Gard.

52:182-211.

Lewis, W.H. 1966. The Asian genus Neanotis nomen novum (Anotis) and allied taxa in the Americas (Rubiaceae).

Ann. Missouri Bot. Gard. 53:32-46.

Lewis, W.H. and R.L. Ouver. 1970. Chromosome numbers of Phanerogams. 3. Ann. Missouri Bot. Gard. 56:474.

Lewis, W.H., Y. Supa, and R.L. Ouver. 1967. Chromosome numbers of phanerogams. 2. Ann. Missouri Bot. Gard.

54:181.

Matta, S.B. et al. 1978. IOPB Chromosome number reports LIX. Presented by A. Lóve. Reports by Malla et al.

Taxon 27:55.

RAGHAVAN, T.S. and K. RANGAswAMY. 1941. Studies in the Rubiaceae. Part 1. J. Indian Bot. Soc. 20:341-356.

Reep, C.F. 1970. Dentella repens and Hedyotis corymbosa, new to the United States. Phytologia 19:311-312.

RipspALE, C.E. 1998. Rubiaceae, Tribe Hedyotideae. In: M.D. Dassanayake, ed. A revised handbook to the flora of

Ceylon. Vol. XII. A.A. Balkema, Rotterdam. Pp. 236-284.

Rocers, G.K. 1987. The genera of Cinchonoideae (Rubiaceae) in the southeastern United States. J. Arnold Arbor.

68:137-183.

SALDANHA, C.J. and D.H. Nicotson. 1976. Neanotis. In: Flora of the Hassan District, Karnataka, India. Amerind Publ.

Co. Pvt. Ltd., New Delhi. P. 583.

SELVARAJ, R. 1987. Karyomorphological studies in South India Rubiaceae. Cytologia 52:343-356.

Smith, A.C. and S.P. Darwin. 1988. Rubiaceae, vol. 4, Hedyotis. In: A.C. Smith, Flora Vitiensis Nova. Pacific Tropical

Botanical Garden, Kauai, Hawaiii. Pp. 352—359.

TERRELL, E.E. 1990. Synopsis of Oldenlandia (Rubiaceae) in the United States. Phytologia 68:125- 133.

TERRELL, E.E. 1996. Revision of Houstonia (Rubiaceae-Hedyotideae). Syst. Bot. Monogr. 48:1-118.

TERRELL, E.E. and W.H. Lewis. 1990. Oldenlandiopsis (Rubiaceae), a new genus from the Caribbean Basin, based on

Oldenlandia callitrichoides Grisebach. Brittonia 42: 185-190.

TERRELL, E.E. and H. Rosinson. 2006. Taxonomy of North American species of Oldenlandia (Rubiaceae). Sida

22:305-329.

TERRELL, E.E., W.H. Lewis, H. Rosinson, and J.W Nowicke. 1986. Phylogenetic implications of diverse seed types, chro-

mosome numbers, and pollen morphology in Houstonia (Rubiaceae). Amer. J. Bot. 73:103-115.

VERDCOURT, B. 1976. Rubiaceae (Part 1), Kohautia, Oldenlandia. In: R.M. Polhill, ed. Flora of Tropical East Africa. Crown

Agents, London. Pp. 228-242; 268-315.

VERDCOURT, B. 1989. Rubiaceae, Vol. 5, part 1. In: E. Launert, ed., Flora Zambesiaca. Flora Zambesiaca Managing

Committee.

NOTES ON PHRAGMITES AUST RALIS

(POACEAE: ARUNDINOIDEAE) IN NORTH AMERICA

Kristin Saltonstall Donald Hauber

Smithsonian Tropical Research Institute Department of Biological Sciences

Apartado 0843-03092 Loyola University

Panamd, REPUBLIC OF PANAMA New Orleans, Louisiana 70118, U.S.A.

kristin.saltonstall@aya.yale.edu hauber@loyno.edu

ABSTRACT

Three taxa of Phragmites australis have previously been reported in North America. While native P. australis subsp. americanus has been

formally described, questions remain as to the subspecific status of the introduced and Gulf Coast lineages. Here we attempt to provide

answers to commonly asked questions and describe a consistent nomenclature for one of these two lineages. While it has been treated

previously as a variety, we recognize the Gulf Coast lineage as Phragmites australis subsp. berlandieri. This subspecies is distributed

along the southernmost border of the United States and extends its distribution south through Mexico and Central America into South

America. Issues regarding the taxonomic identity of the introduced lineage are also discussed.

RESUMEN

Tres taxones de Phragmites australis han sido citados previamente en Norteamérica. Mientras la nativa P. australis subsp. americanus ha

sido descrita formalmente, aún quedan problem mo el estatus subespecífico, de las plantas introducidas y las de la Costa del Golfo.

Aquí intentamos contestar a las preguntas comunes y proponer una nomenclatura consecuente para uno de estos dos linajes. Aunque

ha sido tratada como una variedad, reconocemos el linaje de la Costa del Golfo como Phragmites australis subsp. berlandieri. Esta

subespecie está distribuida por las fronteras sureñas de los Estados Unidos, extendiéndose hacia el sur por México y Centroamérica

hasta Suramérica. Además se examina la identidad taxonómica del linaje introducido.

Phragmites australis (Cav.) Trin. ex Steud. is a widely distributed species found in marsh systems all over the

world. In 2002, Saltonstall demonstrated that it is represented in North America by three different genetic

lineages. Of these, one is native and endemic to North America, one is found in both North and South

America, and one is introduced and invasive. The native endemic has been named P. australis subsp. ameri-

canus Saltonstall, P.M. Peterson & Soreng and is widespread across North America, extending from Canada

to southern California, across the Midwest, and along the Atlantic Coast to North Carolina (Fig. 1a). The

Gulf Coast lineage is found from the Atlantic coast of Florida, around the Gulf of Mexico, across the south-

ernmost states to the Gulf of California and south through Mexico and Central America into South America

(Fig. 1b). In 2004, Saltonstall, Peterson and Soreng stated that this taxon corresponded to P. australis var.

berlandieri (Fourn.) C.F. Reed. It is not clear whether it is introduced or native to the Americas and possibly

other subtropical regions. The invasive introduced lineage of P. australis is now widespread across North

America and its distribution overlaps with both the other lineages (Fig. 1c). The use of different ranks for

the lineages and confusion over how the Gulf Coast lineage relates to native and introduced P. australis has

led to questions that we address in this manuscript.

Nomenclature of the invasive introduced lineage

Introduced Phragmites australis most likely originates from Europe (Saltonstall 2002). This creates a nomen-

clatural dilemma. The holotype of P. australis was collected near Port Jackson [Sydney Harbor], Australia in

1799. Clayton (1968) considered Australian and European specimens to be conspecific, but treated plants

from the Mediterranean region as P. australis subsp. altissimus (Benth.) Clayton. In doing so, he automatically

brought the name P. australis subsp. australis into existence. These two subspecies may be *rather imperfectly

distinguished" (p. 116) by the shape of the upper glume (Clayton 1967). Tutin (1980) did not recognize any

infraspecific taxa in his treatment of Phragmites for Flora Europaea. Phillips (1995) placed Ethiopian plants

in P. australis subsp. altissimus, but did not state how they differed from subsp. australis. She described the

J. Bot. Res. Inst. Texas 1(1): 385 — 388. 2007

386 Journal of the Botanical Research Institute of Texas 1(1)

Fic. 1. Distribution of a) Native, b) Gulf Coast, and c) Introduced lineages of Phragmites australis in North America. Reprinted from Saltonstall et al.

2004.

plants of P. australis subsp. altissimus as having culms 3-6 m tall, leaf blades 30-60 cm long by 1-3 cm

wide, and panicles 30—50 cm long. This is very similar to the description Wheeler et al. (2002) provided

for plants from New South Wales, Australia: Plants to about 6 m tall, leaf blades to 50 cm long and 3.5 cm

wide, panicles 15—30 cm long. Both descriptions provide additional details, but there is no evident distinc-

tion between the two.

The genetic relationship between European and Australian P. australis populations is poorly understood

at this time. Chloroplast DNA haplotype M, which North American introduced P. australis possesses, is

widespread across Europe and Asia. Haplotype Q, which is distinct from all haplotypes found in Europe,

was found in Australia (Saltonstall 2002). Further, in a phylogeographic study of Phragmites using AFLPs,,

1AI+) Ļ td dud *hafultf. " PET pare ieee A Y Pree *: D; Nal (Calt 4,11 20N9\ ; r 1 DH

RL H p_e

Saltonstall and Hauber, Notes on ! t t North America 387

Lambertini et al. (2006) recognized a distinct Australian/Asian clade within P. australis. The relationships of

the European taxa to the Australian taxon are unclear and need further investigation. It is also not known to

which of the named European taxa introduced P. australis belongs. Therefore, it is not possible at this time

to identify the appropriate subspecific name for the introduced lineage in North America.

Nomenclature of the Gulf Coast lineage

In treating the Gulf Coast lineage as Phragmites australis var. berlandieri, Saltonstall et al. (2004) left unan-

swered the question as to its subspecific status. Although botanical names are never shown with more than

one infraspecific rank, if both subspecific and varietal ranks are used, it is desirable to show how the taxa

relate to each other. Morphologically, the Gulf Coast strain resembles the introduced lineage more than the

native. It differs significantly from the introduced strain in only one of the four characters measured, lower

glume length, and was intermediate between the other two lineages at most characters (Saltonstall et al.

2004). Examination of microsatellite DNA variation (Saltonstall 2003) revealed that Gulf Coast populations

had unique alleles and allele phenotypes at most loci. They all share the same cpDNA haplotype I, which

was also found in some South American samples and one sample from Guam, and has several mutations

which distinguishes it from haplotype M (Saltonstall 2002). They also share similar isozyme profiles, which

differ from those of introduced P. australis (Pellegrin and Hauber 1999). Clearly, they represent a taxon that

is distinct from that of the invasive introduced lineage and from subsp. americanus. It is not as yet clear how

widely the Gulf Coast taxon is represented outside the Americas.

Saltonstall et al. (2004) recognized the Gulf Coast lineage as a variety but, in retrospect, it seems better

to name it a subspecies so as to reflect the equivalence of its genetic differentiation to that of subsp. america-

nus. The lineage has sometimes been called Phragmites karka Retz. (Jones 1997), the holotype of which was

collected in India. It is possible, though not evident, that the Gulf Coast lineage belongs to that species. It

seems best, therefore, to employ the same epithet for the subspecies as was earlier used at the varietal rank

in Saltonstall et al. (2004). If this lineage is subsequently determined to belong to the same taxon as P. karka,

then that name will have precedence at the species level, but will have no effect at the subspecies level since

there are no subspecific names in P. karka. We propose a new subspecies combination for the Gulf Coast

lineage below.

Phragmites australis subsp. berlandieri (E. Fourn.) Saltonstall & Hauber, comb. nov. Basionvm: Phragmites berlan-

dieri E. Fourn., Bull. Soc. Bot. France 24:178. 1877. Phragmites communis var. berlandieri (E. Fourn. ) Fernald, Rhodora 34:211. 1932.

Phragmites maximus var. berlandieri (E. Fourn.) Moldenke, Torreya 36:93. 1936. Phragmit bsp. berlandieri (E. Fourn.) Á.

Lóve & D. Lóve, Bull. Torrey Bot. Club 81:33. 1954. Phragmites australis var. berlandieri (E. Fourn.) C.E Reed, Phytologia 63:410. 1987.

Tre: U.S.A. Texas: entre Laredo y Bejar, Feb 1828, J.L. Berlandier 1446 (Lectotype designated by Saltonstall et al. 2004: P, [see notes by

Catling 2006]; Isolectotype: US-82049 fragm. ex P!, US-82049 fragm. ex Pitt. € Dur.! [Bruxelles]), US-82049 fragm. ex W!).

Clearly, questions remain concerning Phragmites, many of which require a global approach. The purpose of

this paper is simply to provide a consistent nomenclature, to the extent that it is possible, for the lineages

that occur in North America. The following key using morphological and genetic features is given to separate

these three lineages (from Saltonstall et al. 2004).

KEY TO THE LINEAGES OF PHRAGMITES AUSTRALIS IN NORTH AMERICA

1. Ligules 1.0-1.7 mm long; lower glumes 3.0-6.5 mm long; upper glumes 5.5-11.0 mm long; lemmas 8.0-13.5

mm long; leaf sheaths caducous with age; culms exposed in the winter, smooth and shiny; rarely occurs in

a monoculture; chloroplast DNA haplotypes A-H, S, Z, AA, AB, AC (see Saltonstall 2002, 2003) P. australis subsp.

americanus (Native lineage

. Ligules 0.4-0.9 mm long; lower glumes 2.5-5.0 mm long; upper glumes 4.5-7.5 mm long; lemmas 7.5-12.0

mm long; leaf sheaths not caducous with age; culms not exposed in the winter, smooth and shiny or ridged

and not shiny; usually occurs as a monoculture; chloroplast DNA haplotypes | or M

2. Culms smooth and shiny; southern California, Arizona, New Mexico, Texas to Florida, throughout Mexico

and Central America; chloroplast DNA haplotype | P. australis subsp

berlandieri (Gulf Coast lineage

2. Culms ridged and not shiny; southern Canada from British Columbia to Quebec south throughout the

Continental United States; chloroplast DNA haplotype M P. australis (Introduced lineage)

pan

—

WS o.

388 Journal of the Botanical Research Institute of Texas 1(1)

ACKNOWLEDGMENTS

Special thanks to Mary Barkworth for her comments and suggestions, and Paul Peterson and an anonymous

reviewer for their comments on the manuscript. We thank Dylan Craven and Adriana Sautu for providing

a Spanish translation of the abstract.

REFERENCES

CaruNG, PM. 2006. Notes on the lectotypification of Phragmites berlandieri and identification of North American

Phragmites. Bot. Electronic News 366.

CLAYTON, W.D. 1967. Studies’ in the Gramineae: XIV. Kew Bull. 21:111-117.

CLAYTON, W.D. 1968. The correct name of the common reed. Taxon 17:168-169.

Jones, S.D., J.K. WiPrr, and PM. Montcomery. 1997. Vascular plants of Texas. University of Texas Press, Austin.

LAMBERTINI, C., M.H.G. GUSTAFSSON, J. FRUDENBERG, J. LissNER, M. SPERANZA, and H. Brix. 2006. A phylogeographic study of

the cosmopolitan genus Phragmites (Poaceae) based on AFLPs. PI. Sys. Evol. 258:161-182.

PELLEGRIN, D. and D.P. Hauser. 1999. Isozyme variation among populations of the clonal species, Phragmites australis

(Cav.) Trin. ex Steudel. Aquatic Bot. 63:241-259.

PHILLIPS, S.M. 1995. Phragmites. In: |. Hedberg and S. Edwards. Flora of Ethiopia and Eritrea, vol. 7. Swedish Science

Press, Uppsala, Sweden.

SALTONSTALL, K. 2002. Cryptic invasion by a non-native genotype of the common reed, Phragmites australis, into

North America. Proc. Natl. Acad. U.S.A. 99:2445-2449.

SALTONSTALL, K. 2003. Microsatellite variation within and among North American lineages of Phragmites australis.

Molec. Ecol. 12:1689-1702.

SALTONSTALL, K., PM. PETERSON, and R.J. SoRENG. 2004. Recognition of PI p.americanus (Poaceae:

Arundinoideae) in North America: evidence from morphological and genetic analyses. Sida 21:683-692.

Turin, T.G. 1980. Phragmites. In: Tutin, T.G., V.H. Heywid, N.A.B.M. Moore, D.H. Valentine, S.M. Walters, and D.A. Webb,

eds. Flora Europaea, vol. 5. Cambridge University Press, Cambridge, England. Pp. 253.

WHEELER, D.J.B., S.W.L. Jaco&s, and R.D.B. WHALLEY. 2002. Grasses of New South Wales, 3* ed. University of New

England, Armidale, New South Wales, Australia.

T fs |

THE TAXONOMY OF CAREX TRISPERMA (CYPERACEAE)

Chad D. Kirschbaum

Wayne National Forest, Ironton Ranger District

65 te Route 93

Pedro, Ohio 45659, U.S.A.

ckirschbaumavfs.fed.us

ABSTRACT

Ora Willis Knight described Carex trisperma Dewey var. billingsii O.W. Knight in 1906, but it has not been quantitatively compared to its

den sister taxon, C. t. var. Mi d I tested the hypothesis implied by earlier taxonomic treatments that the two varieties are not

lly, ecologically. 1]

o OL

, and genetically distinct. To test these hypotheses, I analyzed DNA fingerprints, measured

PEN, characters, mapped specimen localities and recorded in situ canopy conditions. The two varieties are distinct based on

AFLP fragment data. Based on iB n analyses, the two varieties are distinguishable by leaf width, ligule length, inflorescence

length, and number of perigynia per I spike. Carex t. var. billingsii is plant of the northeastern United States and adja-

cent Canada and is confined to partially shaded/open areas in acidic bogs within deciduous forests. Carex t. var. trisperma ranges farther

west, south and east than C. t. var. billingsii and grows mostly in shaded areas of bogs and swamp forests in both deciduous and boreal

biomes. If C. t. var. billingsii were simply an open-grown morphotyp

fC. t. var. trisperma, then it would be likely that the distribution

of C. t. var. billingsii would overlap that of C. t. var. trisperma and thus, be present in open bogs west of Michigan. The absence of C. t.

var. billingsii in areas where C. t. var. trisperma is common suggests a genetic rather than an environmental basis for the differences in

morphological characters. The geographic, ecological, morphological, and genetic data strongly suggest that C. t. var. billingsii warrants

recognition at the rank of species and the new combination is effected.

Key Wonps: Carex; sedge; AFLP; morphometrics; sedge ecology; phytogeography; DNA fingerprints; sphagnum bog

RESUMEN

Ora Wills Knight describió Carex trisperma Dewey var. billingsii O.W. utente en 1906 dee no ha sido sud cuantitativamente con

su presunto taxon hermano, C. t. var. trisperma. He probado que la hipó n que las dos

variedades son morfológica, ecológica, died y genéticamente distintas. AR probar estas hipótesis, analicé re ele de DNA,

medi caracteres morfológicos, representé en mapas las localidades de los especimenes y tomé datos in situ de | ndiciones del manto.

Las dos variedades se distinguen en base a los datos de los fragmentos AFLP. Basándose en los análisis morfológicos, las dos variedades

[sca

se pueden distinguir por la anchura de las hojas, longitud de la ligula, longitud de la inflorescencia, y número de utrículos por espiga

terminal. Carex t. var. billingsii es una planta templada del Noreste de Estados Unidos y la parte adyacente de Canadá y está confinada a

áreas parcialmente sombrías/abiertas en ciénagas ácidas dentro de bosques caducifolios. Carex t. var. trisperma llega hasta más al Oeste,

más al Sur y al Este que C. t. var. billingsii y crece en áreas sombrías de ciénagas y pantanos forestales en biomas caducifolios y boreales.

Si C. t. var. billingsii fuese simplemente un morfotipo de lugares abiertos de C. t. var. trisperma, sería probable que la distribución de

C. t. var. billingsii se solapase con la de C. t. var. trisperma y estuviese presente en ciénagas abiertas del Oeste de Míchigan. La ausencia

de C. t. var. billingsii en áreas donde C. t. var. trisperma es comün sugiere una base genética en vez de ambiental para las diferencias en

los caracteres morfológicos. Los datos geográficos, ecológicos, morfológicos, y genéticos data sugieren que C. t. var. billingsii merece el

reconocimiento a nivel de especie.

During a botanical foray, in 1906, Ora Willis Knight, discovered a *peculiar little sedge" at Jewett Brook Bog

in Maine. Upon further inspection, he decided that the sedge was merely a variant of typical Carex trisperma

Dewey (Fig. 1), which was common in shaded portions of the bog. Knight named this plant Carex trisperma

var. billingsii (Fig. 1) and described the foliage of C. t. var. billingsii as *setaceous or filiform,” the inflorescence

as having 1-2 spikelets per culm, and its habitat as “sunny.” This habitat description was contrasted to that

of C. t. var. trisperma, which grew “abundantly...under the trees” (Knight 1906).

The two Carex trisperma varieties have never been thoroughly evaluated taxonomically. Gleason and

Cronquist (1991) and Toivonen (2002) briefly reported the differences between the two varieties, the most

prominent of which are narrower leaves and fewer, smaller perigynia in C. t. var. billingsii. Gleason and Cron-

quist (1991) also reported that C. t. var. billingsii ranges from New Brunswick to Vermont and Pennsylvania.

Toivonen (2002) extended that range to include Michigan and adjacent areas in Canada.

This study examines the morphology, phytogeography, ecology, and genetic relationships of the two

J. Bot. Res. Inst. Texas 1(1): 389 — 405. 2007

titute of Texas 1(1)

laf tha Dat

JOUIIdI OI

390

*SOMIAS ajewiay POMO Y Jama} ‘ I | Jamoueu sy Aq Ábojoyd b DuLadsi} "Ie *} *) WO peusinDunsip

st ISBUIIIIG "Yen 7) xo1D) “U's ‘WINDQYISATY ‘TOOT Inf YO “PUOJ UOINIDS 1eau ^0) pojjens “SUSO WEY MAN WO. suon»e|[oo uo paseq ‘(4J9]) auDruy 156u1]11q "ren pue (1461) J ^e famaq puadsy vaio) Jo JPY AU L H

zo0z Anf yo "Q'O 'unequosii

zooz Ainr vo "Q'O ‘wnequosuy,

'S991) pooneos uu ÁLOISIOAO QUUYS 9SUS "dwy

Soon PO/OHeoS uy ÁJOISIOAO qnuus asuaq E uojBuureg “peo, jo epis yuou eu; uo Bog Buneoj *pepunouns yeow *uedo Woy ssoioe

uoj&uureg “ped! jo epis yuou ay) uo oq Buneoj ‘papunouns eow *uado WO4 ssoioe [fein y pen ey nn 'peot puog OJOS uo 'puog uoynuos jo LON

ÁROBAIP si IQ peos 1103, ejqisia JOU s! Bog 'peog puog uojnias uo 'puog uojnuos jo YON

A TTSDUTTITQ "zea

aqyStuy FUISASTIZ *1€A Koneg wuredsTI3 xereD

Aomoq wurodsr13 XoIv) eeeoeradÁg

a) Aunog pioyens

Ájuno9 paoyens exysduley MeN jo Sued

ƏnysdweH MON jo SURG

! 20

| $

Kirschbaum, Taxonomy of Carex trisperma varieties 391

varieties in order to evaluate their taxonomic ranks as currently treated. I tested the hypothesis that the

two varieties are distinct species using analyses of 19 morphological characters, AFLP genetic markers,

habitat observations and the geographical distributions of the two taxa. If there are clear distinctions in

morphology, ecology, and distribution, and if sympatric plants of both varieties can be distinguished by

unique genetic markers, then raising C. t. var. billingsii to the species level will be supported. This study is

the first quantitative analysis of Carex trisperma varieties and identifies potential variations to include in

future phylogenetic studies of Carex sect. Glareosae G. Don.

MATERIAL AND METHODS

Field Sites and Collections Methods.—A total of 259 individual plant specimens were collected from 22

different sites throughout the range of C. t. var. billingsii from July to September 2002 and in New York and

Michigan in August and September 2003 (Table 1). The two varieties were syntopic at 13 of these sites. At

two sites (Dawson Ponds and Corea Heath) C. t. var. billingsii was found without C. t. var. trisperma in the

immediate vicinity of the bog (William Crins, pers. comm. and Anton A. Reznicek, pers. comm .). At all sites

where both varieties were abundant, at least one individual of each variety was collected. Voucher specimens

(MICH) were collected for morphological analysis, and several leaves of these specimens were immediately

placed in silica gel for molecular analysis.

Herbarium Specimens and Distribution Mapping.—Specimens from herbaria (MICH, DAO, MT, and

GH) were used for morphological measurements (along with field collected specimens) and distribution

mapping. Nineteen morphological characters were selected for detailed measurement (Table 2). Data were

collected from twenty specimens of each variety selected from localities throughout the Northeastern United

States. Specimen label information was used to map the distributions of the two varieties A total of 310

specimens were mapped with ArcView GIS 3.2 (Environmental System Research, Inc.). Collections of Carex

trisperma were investigated at WIS, MO, MSC, UWSP, OS and CLM to determine if C. t. var. billingsii ranged

further south than previously reported. On the edge of C. t. var. billingsii's range in Michigan, sphagnum bogs

in Allegan County and Barry County in Michigan's lower peninsula were also checked for the presence of

C. t. var. billingsii.

Specimens of both varieties were thinly sliced with a scalpel blade to prepare specimens for scan-

ning electron microscopy (SEM) (Jane Gillies, pers comm.). For SEM, the specimens were mounted onto a

polished stub with double-stick carbon-permeated tape. As necessary, further attachment to the stubs was

made with colloidal graphite. The specimens were then coated with gold and examined with an AMRay

18201 scanning electron microscope. For the anatomical cross-sectional analysis, a freezing microtone was

used to obtain sections thin enough to be photographed under a compound microscope.

Molecular analysis.—Amplified Fragment Length Polymorphisms (AFLPs) were used to investigate

species boundaries and evaluate genetic similarity within and between varieties of Carex trisperma. Ge-

nomic DNA was extracted from 8-12 mg of silica-dried leaf material from 12 syntopic populations and two

specimens (one of each variety) from different bogs in or near Algonquin Provincial Park (APP and DLB,

Table 1) for a total of 26 individuals. I isolated DNA using GenElute Plant Genomic DNA Miniprep kits

(Sigma-Aldrich) with the addition of 50 units of Ribonuclease (RNase A solution, Sigma-Aldrich) to elimi-

nate RNA contamination and 10 mg of Polyvinylpyrrolidone (PVPP, Sigma-Aldrich) to precipitate secondary

compounds. RNase and PVPP were added after cell lysis and prior to incubation.

AFLP protocols followed Vos et al. (1995) with modifications by Berres (2002) and Hipp (2004). I

used the selective amplification primers (“EcoRI + ATG”; 5' GAC TGC GTA CCA ATT CAT G 3’ and “Msel

section Ovales by Hipp (2006). The underlined bases on each primer correspond to the known sequences

of double stranded adapters ligated to the cut ends of genomic DNA subsequent to restriction digestion. The

bases in bold type are selective nucleotides employed in the AFLP process to reduce the bands amplified to

an interpretable number.

392 Journal of the Botanical R h Institute of Texas 1(1)

Taste 1. Collection localities of specimens used for AFLPs, morphometrics, and ecological information (*). AFLPs were screened

in one individual of each variety in 12 syntopic populations except for JBB. Allopatric specimens from Ontario sites; APP and

DLB were also included in the AFLP analysis. Specimen collectors include 'AAR = Anton A. Reznicek, WJC = William J. Crins,

GH = Geoffrey Hall, JH = Justin Hohn, CDK = Chad D. Kirschbaum, CJR = Carl J. Rothfels and TR = Todd Ristau (nf = not found,

**at km 8 along Hwy 60).

No. of Samples No. of Samples

Site Code Locality State/ variety variety Collector’

Province trisperma billingsii

BPB Bog Pond* ME 20 20 CDK

CH Corea Heath ME nf 1 AAR

2ABB Hwy 2A Bonus Bog ME 2 nf CDK

JBB Jewett Brook Bog ME 1 1 CDK

OPB Otter Pond Bog* ME 20 20 CDK

PMB Petit Manan ME ] 1 AAR

GLB Gorman Lake Bog MI 1 1 CDK + JH

ILB Independence Lake Bog MI 1 nf CDK + JH

MNB Minden Bog MI 2 10 CDK

ML Miner Lake Bog MI 1 nf CDK + JH

BHB Bray Hill Bog* NH 20 20 CDK

HPB Heath Bog Natural Area NH 2 2 CDK

MPB Mud Pond (Fox State Forest)* NH 20 20 CDK

SPB Scruton Pond NH 1 1 CDK

AB Allenburg Bog NY 2 nf CDK 4 TR

MSL Moss Lake Bog NY 1 1 CDK +TR

LCB Lake Carmi Bog LA 20 20 CDK

OBCI Zone 18 QBC 6 nf GH

QBC2 Zone 19 QBC 2 nf GH

APP Algonquin Prov. Park** ONT 10 nf WJC

DLB Dawson Ponds ONT nf 1 WJC

CTB Copetown Bog ONT 4 3 CJR

Totals 137 122

The final PCR product was cleaned with Centri-Sep column kits (Princeton Separations), mixed with a

fluorescent-red-labeled size standard (GeneScan™ -500 ROX™, Applied Biosystems) and Hi-Dye deionized

formamide (Applied Biosystems). Deantured and snap-chilled samples were loaded into a 96-well sample

tray for capillary electrophoresis using ABI's Prism 310 Genetic Analyzer. The operation of the 310 Genetic

Analyzer followed ABI protocols (Applied BioSystems 2000, 2001) with modifications and optimizations

described in Kirschbaum (2005).

Analytical Methods.—1 compared morphological character measurements between varieties using a

two-tailed Mann-Whitney U test. Characters that were significantly different (Table 2) were included in a

Principal Component Analysis (PCA) in order to assess relationships between the two taxa based on overall

variation in the quantitative characters. Prior to the PCA, correlation analysis between all characters was

carried out to test for character correlation, which would have heavily weighted correlated characters (Sokal

& Sneath 1963; Abbott et al. 1985). None of the variables used in this analysis was significantly correlated (p

< 0.05, r > 0.75) and, thus, all 19 were used in the analysis. To provide equal weight among characters, the

data were Z-score transformed before conducting the PCA. These procedures standardized all measurements

so that each variable had a mean of zero and a standard deviation of one (Sokal & Sneath 1963). Principal

component scores for the first three axes were graphed on a scatter plot. Discriminant Function Analysis

(DFA) was used to achieve maximum discrimination among samples on the basis of the transformed variables

and a priori designations of samples to a taxon (i.e., C. t. var. billingsii or C. t. var. trisperma). Both PCA and

Kirschbaum, Taxonomy of Carex trisperma varieties 393

Taxa um

6 bilingsii A

. $ A

A trisperma e &

= 1.00000 2"

5 eu "

c e e & n N

e . -

- A , A

E 0.00000 e m A " A

O A

E o y

=== -1.000007 e

o A

c e ® A A

à A

-2.000007 A

| I 1

-1.00000 0.00000 1.00000

Principal Component 1

Fic. 2. A scatter plot of tl f Principal Comp ts 1 and 2 for hological characters of Carex tri D arieties.

T J F y Yall

DFA were used to determine the relative importance of characters that distinguish the two taxa. SPSS 11.5

(Statistical Package for the Social Sciences, Inc.) was used for all analyses of the morphological data.

The raw AFLP fragment data (i.e., fragment size, amplitude, etc.) were copied from ABI's GeneScan™

software to a Microsoft Excel spreadsheet. A macro was used to create a matrix of ones and zeros that de-

noted the presence or absence of a peak for each specimen. This matrix was then edited with reference to a

GeneScan chromatograph. Small peaks that were present in at least 11 of the 13 samples from each variety

were added to a plant's profile if the peak in question had an amplitude within 100 RFUs of the original

amplitude threshold (800 RFU). Fragments that were present in only 1 sample out of 26 were considered to

be artifacts of PCR or electrophoresis and were eliminated from the analysis. A total of 31 fragments were

eliminated from the original GeneScan data set as a result of this editing.

The data matrix with the presence or absence of AFLP fragments for each specimen was used to cal-

culate pairwise genetic distance matrices, using Nei and Li's (1979) genetic distance index for fragments

in Phylip 3.62 (Felsenstein 2004) and Jaccard's (1908) distance measure in PC-ORD 4.32 (McCune and

Medford 1997). Nei and Li's pairwise genetic distances were compared graphically using SYSTAT 10.2.01

(SYSTAT Software Inc.).

To examine taxon boundaries with the genetic data, I performed ordination using Nonmetric Multi-

dimensional Scaling (NMS) in PC-ORD. NMS was chosen because, unlike phylogenetic or some phenetic

approaches (e.g., UPGMA or Neighbor Joining) to analyzing molecular data, NMS does not assume that any

hierarchical patterns are present in the data, an assumption that would be invalid at the start of such a study

(Lessa 1990). NMS, compared to other methods of ordination, such as Principal Component Analysis, does

not assume linearity among the variables and allows for the analysis of distance measures. Since Nei and

394 Journal of the Botanical R h Institute of Texas 1(1)

TABLE 2. Summary of quantitative characters (means, + standard error and ranges, n=20) measured and qualitative characters

observed on herbarium specimens of Carext and billingsii. Characters that were significantly differ-

ent* (two-tailed Mann-Whitney U test (MWU), asymptotic significance [as] « 0. 05) between the two varieties were analyzed

using Principal Component Analysis (PCA). Scores of the principal component (PC) axes are given for the six characters used

in the PCA. **Discriminant Function Correlation Coefficient = Pooled within-groups correlations between discriminating

variables and standardized canonical discriminant functions. Highly weighted characters are highlighted.

Quantitative Characters variety variety PC1 PC2 PC3 DECC

trisperma billingsii

Number of spikes per inflorescence * 271013 2340.10 0.38 0.69 0.06 -0.01

(MWU=117.5, as=0.01) (2-4) (2-3)

Number perigynia per lateral spike * 29402 2:2::0.15 0.57 Dus 0.65 0.03

(MWU=106, as<0.001) (1-5) (1-3)

Number of perigynia per terminal spike* 3.7+0.29 2 150 15 0.71 0.36 0.30 0.27

(MWU=59, as<0.001) (1-6) (1-3)

Achene length (mm) * 2.0+0.03 1.8+0.04 0:55 0.39 0.53 0.09

(MWU=94, as<0.001) (1.7-2.2) (1.3-2.0)

Terminal Bract (Bristle) length (mm) * 51.142.87 41.943,31 0.56 0.31 0.15 0.23

(MWU=124, as=0.04) (28-74) (15-72)

Inflorescence length (mm) * 38.82.05 22 6£1.16 0.78 0.34 012 0.37

(MWU=51, as<0.001) (14-55) (14-32)

Ligule length (mm) * 1.1+0.08 0.5+0.05 0.81 0.20 0.31 0.41

(MWU=27, as<0.001) (0.5-1.9) (0.3-1.2)

Leaf width (mm) * 122006 0.4+0.03 0.83 0.25 0.18 0.78

(MWU=0.5, as«0.001) (0.8-1.9) (0.3-0.8)

Pistillate scale width (mm) * 1.4+0.03 1.2+0.04 0.60 0.27 0.24 0.04

(MWU=99.5, as<0.001) (1.1-1.7) (0.9-1.5)

Pistillate scale length (mm) 2.70.07 274007 Not Not Not 0.03

analyzed analyzed analyzed

in PCA in PCA in PCA

(MWU=188, as=0.744) (2.2-3.2) (2.2-3.2) E li i

Staminate scale length (mm) 2220S 300,12 i , d 0.08

(MWU=167, as=0.371) (2.4—4.6) (2.2-4.3) E i :

Perigynia length (mm) 3.2+0.08 3.2+0.06 , i i -0.07

(MWU=185.5, as=0.693) (2.2-3.7) (2.7-3.9) " " i

Perigynia width (mm) 1.4+0.03 1440.04 il j i 0.24

(MWU=196.5, as=0.923) (1.2-1.7) (0.9-1.8) i i B

Achene width (mm) 1230:03 1.20.04 i 2 il 0.31

(MWU=197, as=0.933) (1.1-1.5) (0.8-1.4) ý : j

Qualitative Characters

Li's (1979) genetic distance index is not available in PC-ORD, Jaccard's Coefficient was used as a similarity

measure. Jaccard's Coefficient has been shown to perform better than other nonevolutionary-based coef-

ficients in elucidating relationships between closely related taxa (Landry & Lapointe 1996).

At sites in New England and Michigan, I randomly selected individuals at several bogs and noted the

light conditions under which the plant was growing (either completely under a tree or shrub canopy or

in a partly shaded/open condition). These two categories are based on the original habitat descriptions of

the two varieties given by Knight (1906) and habitat notes from herbarium specimens. Given these broad

habitat descriptions, 9596 of the plants visited were precisely placed in these categories. The frequencies of

169 plants growing in each condition were analyzed by variety with X? analysis.

RESULTS

Morphology.—The number of spikes per inflorescence, number of perigynia per lateral spike, achene

Kirschbaum, Taxonomy of Carex trisperma varieties 395

O variety billingsii

O variety trisperma

Count

Fic. 3. A histog fthe discriminant functi for Carex tris} D ieties, based on 14 measured morphological characters.

length, terminal bract (bristle) length, inflorescence length, ligule length, leaf width, and pistillate scale

width differed significantly between varieties (two-tailed Mann-Whitney U test, asymp. sig. < 0.05) (Table

2). No observable differences were noted for qualitative characters, such as achene shape, sheath apex, or

ligule shape and scale venation.

The overall variation of morphological characters, analyzed by PCA, is summarized by a scatter plot of

the scores of Principle Components 1 and 2 (Fig. 2). These scores resolved two groups (Fig. 2). The two taxa

separated on the first axis, which explained 44% of the total variance (Fig. 2). The second and third axes,

however, only explained 13 and 11% of the total variance, respectively. Characters with high component

(loading) scores (> 0.70) on axis one included leaf width, ligule length, inflorescence length, and number of

perigynia per terminal spikelet (Table 2). As the values on axis one increases, all four of these characteristics

increase and are associated with taxa described as C. t. var. trisperma (Fig. 2).

I assessed the relative importance of the morphological characters that distinguish the two taxa with

Discriminant Function Analysis. A distinct separation of the two taxa is evident on a histogram of the dis-

criminant function scores (Fig. 3). The three most highly weighted characters were the same as the highly

weighted characters on Principal Component 1 in the PCA (Table 2, Fig. 6). Leaf width, ligule length, and

inflorescence length will be the most useful for field recognition (Fig. 4).

The differences in leaf width originate through significant differences in leaf structure. Scanning electron

microscope and leaf cross section photos demonstrate the anatomical differences that account for difference

in leaf width (Fig. 5). The margins of C. t. var. billingsii leaves are involute above the sheath but fuse distally,

which accounts for the triangular shape of the leaf in cross section. The leaves of C. t. var. billingsii have 1-2

large areas of aerenchyma tissue on the left and right sides of the midrib. The leaves of C. t. var. trisperma

are thinly M-shaped in cross section, but appear flat apically and are deeply channeled, or keeled, on the

396 Journal of the Botanical R h Institute of Texas 1(1)

25 60

20 | 50 |

5 ka

Ez to

Q 15- E

: P

10 3 30

o D d

0.0 10

? 25

EC 20

E 5:

2 5

t ] to 15

E ^ E

B 4 | © 10

€ 2

E = ===

B 1 2

0 0.0

Var. billingsii Var. frisperma Var. billingsii Var. frisperma

Fic. 4. Box plots of the four morphological characters that best distinguish C isp varieties.

abaxial surface, thus appearing V-shaped basally (near the sheath). There are 4—5 areas of aerenchyma tissue

on either side of the midrib of C. t. var. trisperma.

Genetic Similarity.—A total of 102 loci were scored from GeneScan chromatographs. Ninety-two (90%)

of those markers were polymorphic. DNA fragments ranged from 39 to 466 base pairs in length. The aver-

age (+ standard error) fragment length was 148 +9.4 base pairs, indicating a bias towards smaller fragments.

Eighty-two markers were scored, and 8096 (66) were polymorphic in C. t. var. billingsii. For C. t. var. trisperma,

ninety-five markers were observed of which 8696 (82) were polymorphic. With the primer pairs that were

used, fragment sizes of 143, 155, and 173 base pairs were uniquely found in all specimens of C. t. var. billingsii,

and fragment sizes of 154 and 178 base pairs were uniquely found in all specimens in C. t. var. trisperma.

The scatter plot of the ordination scores calculated using Jaccard's similarity index and nonmetric

multidimensional scaling cleanly separated the two varieties (Fig. 6). The best solution (defined by the

dimensionality with the lowest final stress from a real run) is a 3-dimensional solution with a final stress

value of 13.7. All three axes explain 8596 of the variance with 2396, 4496, and 1896 of the va riance explained

on the first, second, and third axes, respectively.

The mean intraspecific pairwise genetic distance (Nei & Li 1979) across all specimens was 0.01 (+

0.0003). Intraspecific genetic pairwise distances ranged from 0.0008 to 0.024. Genetic distances within

varieties trisperma and billingsii (summarized in Fig. 7) ranged from 0.003 to 0.024 and from 0.0008 to 0.015,

respectively. The mean pairwise genetic distance within var. trisperma was 0.011 (+ 0.0004), whereas the

mean pairwise genetic distance within var. billingsii was 0.009 (+ 0.0004).

Phytogeography.— The two Carex trisperma varieties have overlapping ranges in the northeastern United

Kirschbaum, Taxonomy of Carex trisperma varieties 397

(A) (B)

F Ec H ex : (CEAM J " lE L £1 hi-h I^; 1: ££. "p £ tede WA eiu let +

t V /

(RYCERA AL 4 L ran Ļ I "n £al £f, Ff. PEL) oe hill ool nt kas (D\ CLAA AL "1 D £24 T I +. fal ££

wn) V

Carex trisperma Dewey variety trisperma. (C) C ti f ical portion of a Carex tri iety billingsii leaf. (D) C ti f a basal

PE £an r. e. Ue eis

V f f E 4 J

I £

r F Y F ` J T F , E

States, southern Ontario, and Quebec (Fig. 8). Both taxa are found as far east as Newfoundland. Carex t.

var. billingsii is found throughout New England, and its distribution follows the St. Lawrence seaway west

through the Great Lakes. The range of C. t. var. trisperma extends farther north, south, and west. Carex t.

var. billingsii ranges farther west and north than previously reported by Gleason and Cronquist (1991), but

is congruent with the distribution reported by Toivonen (2002) (Fig. 8).

Carex ttrisperma var. billingsii may be more widespread than previously described. The farthest south-

western record of C. t. var. billingsii is from Ingham County, Michigan (Parmelee 246 MSC, 134054). A speci-

men of C. t. var. billingsii collected in Wexford County, Michigan (south of Traverse City), in the northern

398 Journal of the Botanical Research Institute of Texas 1(1)

Taxa

9 —— billingsii

pi trisperma

+ inant PE IPEA . I IH "A t £, ACI

D I J

Hai ACI

Fic. 6. A 3 ter |

varieties billingsii Knight and trisperma.

Lower Peninsula, recently found at MICH, expands the taxon's range farther north in Michigan. Searches

in southwestern lower Michigan, however, did not disclose any further localitities. Specimens recently

collected from Allegany County in New York (Kirschbaum, s.n.) and Susquehanna County in southeastern

Pennsylvania (Naczi 10065.) suggest potential localities farther inland and away from coastal areas of the

eastern seaboard and Great Lakes. Ombrotrophic bogs in northwestern Pennslyvania, northern Ohio, and

Indiana are potential locations for new populations of C. t. var. billingsii. However, specimens reviewed at

WIS, MO, UWSP, OS, and CLM yielded no further range extensions for C. t. var. billingsii.

Ecology.—Carex t. var. billingsii mostly grows in full-to-partial sunlight. Often it grows in dense clumps

in ombrotrophic bogs at the base of low-growing ericaceous shrubs or along deer trails and narrow water

channels. Unlike C. t. var. billingsii, C. t. var. trisperma is not restricted to acidic sphagnum bogs and, on

the basis of herbarium label data, is commonly found in densely shaded swamp forests of various floristic,

hydrological, and edaphic compositions. At many of the sites, I also found C. t. var. trisperma in swamps that

were deeply shaded by tree canopies that were adjacent to the main bog mat. When growing in bogs C. t.

var. trispermais commonly (but not always) found in shaded portions such as dense tamarack or ericaceous

shrubs stands.

Kirschbaum, Taxonomy of Carex trisperma varieties 399

0.02r

0.01 o Taxa

e billingsii

x frisperma

Nei and Li Pairwise Genetic Distances

0 | | | | i | | | |

u^ 40 30 20 10 O 10 20 30 40 50

Count Count

Fic. 7. A summary of Nei and Li pairwise genetic distances between specimens, which were calculated from the presence or absence of AFLP

fragments.

Several individuals of C. t. var. trisperma were documented in this study and on herbarium labels as

growing in full sunlight, and several C. t. var. billingsii were noted as growing in shaded areas. These indi-

viduals retain their expected morphological characteristics despite growing in a-typical habitats. Seventy-six

percent of the 89 C. t. var. billingsii observations I collected were growing in open and slightly shaded (from

low-growing shrubs) areas and 2496 of the sp were found under tall shrub or tree canopies. Sixty-four

percent of the 80 C. t. var. trisperma observations I collected were found growing under tall shrub or tree

canopies and 3696 were growing in open and slightly shaded conditions. The proportion of plants growing

in the predicted, “typical” light conditions for C. t. var. trisperma and C. t. var. billingsii was significantly

different (? = 5.7, p < 0.02 and X?» 21.1, p< 0.001, respectively) from expected proportions of 50% of the

specimens in each condition.

DISCUSSION

Morphology.—PCA and DFA support the recognition of two distinct entities among the herbarium specimens

measured in this study (Figs. 4, 5, and 6). Both multivariate analyses place high importance on leaf width,

ligule length, inflorescence length, and the number of perigynia per terminal inflorescence in discern-

ing the two varieties (Table 2, Fig. 6). The more diminutive measurements of C. t. var. billingsii agree with

previous descriptions by Knight (1906), Gleason and Cronquist (1991), and Toivonen (2002). The lack of

morphologically intermediate specimens and the clear morphological distinctions between the two taxa,

based on analysis of several characters simultaneously (Fig. 4), do not support Knight's contention that C.

t. var. billingsii is a transitional form of C. t. var. trisperma.

400

Journal o

fth

titute of Texas 1(1)

3 ed,

ail

"lA

M MS

|; E y è .

Fa dl uM 0 600 Miles

mam:

Fic. 8. The distribution of Carex trisperma Dewey trisp (left) and billingsii Knight (right) mapped from 310 herbarium

specimens and recent collections.

Phytogeography and Ecology.— Carex trisperma var. billingsii and var. trisperma each have a unique

geographic distribution and ecology (Figs. 2-3). Carex t. var. billingsii has been found only in the northeastern

United States and adjacent Canada. Despite occurring in Sphagnum bogs, C. t. var. billingsii is a temperate

plant and is mostly confined to sites within the deciduous forest biome (Brown & Lomolino 1998). The

distribution of C. t. var. trisperma, however, spans the deciduous forest biome and reaches into the boreal

or sub-arctic biome (Brown & Lomolino 1998) and extends farther west, south, and north than that of C.

t. var. billingsii.

In many sites in the east, the two varieties are syntopic (Table 1); however, C. t. var. billingsii has never

been collected west of Michigan. If C. t. var. billingsii were simply an open-grown morphotype of C. t. var.

trisperma, then it would be likely that the distribution of C. t. var. billingsii would overlap that of C. t. var.

trisperma and thus be present in open bogs west of Michigan. However, C. t. var. billingsii has not been

collected in any of the well-studied bogs of southern Wisconsin or northern Minnesota, regions known

for peat bogs that are found in glacial lake beds (i.e., Glacial Lake Wisconsin and Agassiz Lake). Thus, the

absence of C. t. var. billingsii in areas where C. t. var. trisperma is common suggests a genetic rather than an

environmental basis for the differences in morphological characters.

Comparisons of the canopy conditions in which the two varieties were found provide evidence of some

ecological distinction between the two taxa. Carex t. var. trisperma is typically found in shaded conditions,

and C. t. var. billingsii is typically found in open conditions. This lack of morphological plasticity in oppos-

ing light conditions was also noted on herbarium specimens. For example, a specimen of C. t. var. trisperma

collected by Reznicek, 11367, Naczi and Case was “from a small colony in full sun on a hummock in the

open bog.” Each taxon retains its morphological characters across habitats with different light conditions.

These observations agree with Anderson et al. (1996), who quantified habitat characteristics of C.

trisperma in Maine. Even though they did not analyze their data by variety, they found that Carex trisperma

grew under a range of shaded-to-open conditions with a mean percent canopy cover of 53 (+ 35.8 S.D).

Genetic Analyses.——Nei and Li's (1979) genetic distances within varieties trisperma and billingsii

ranged from 0.003 to 0.024 and 0.0008 to 0.015, respectively (Fig. 8). Hipp (2004), in a study of the Carex

tenera Dewey group of section Ovales Kunth, provided the only reasonable benchmark for comparison within

Carex. Genetic distances ranged from 0.03 to 0.08 among three species in the C. tenera group (Hipp 2004).

These numbers indicate a much higher level of genetic divergence between species in the C. tenera group

Kirschbaum, Taxonomy of Carex trisperma varieties 401

than between the C. t. varieties. However, the species of the Carex tenera group can be diagnosed easily with

morphological characters (Hipp 2004), as can the varieties of Carex trisperma (Figs. 4—6). This suggests that

in Carex, morphological distinctiveness does not always correlate with genetic divergence.

Nonmetric multidimensional scaling supports genetic distinction between the two varieties. Intravari-

etal genetic distances were significantly higher in C. t. var. trisperma than in C. t. var. billingsii. This suggests

greater genetic diversity in C. t. var. trisperma, which is also consistent with its relatively more extensive

distribution compared to that of C. t. var. billingsii.

While genetic differentiation alone is not sufficient for delimiting species boundaries, the molecular

analyses combined with the consistent differences in morphological, geographical, and ecological data to

strongly suggest that C. t. var. billingsii is distinct enough from C. t. var. trisperma to warrant raising C. t. var.

billingsii to the rank of species. Indeed, phenotypic clusters based on morphological data in plants often

correspond closely to independent reproductive lineages identified by crossing studies (Rieseberg et

al. 2007). The fact that morphometric data and molecular genetic data both demonstrate strong

differences between the varieties tested in this study supports raising these varieties to species

rank

CONCLUSION

The morphological data presented in this paper suggest that Carex trisperma and Carex billingsii comb. et

stat. nov can be separated on the basis of a distinct set of morphological characters, of which leaf width,

ligule length, inflorescence length, and the number of perigynia are the most diagnostic. Based on phe-

netic analyses of the AFLP fragment data using nonmetric multidimensional scaling, the two varieties are

distinct. The combination of geographic, ecological, morphological, and genetic data strongly suggest that

Carex trisperma var. billingsii is distinct enough from C. t. var. trisperma to warrant raising C. t. var. billingsii

to the rank of species.

Future phylogenic analyses that evaluate evolutionary relationships within sect. Glareosae should include

these two species along with other putative members of this group. The author hopes that the recognition

of C. billingsii at the rank of species will bring this species to the attention of wetland scientists and conser-

vation biologists who can further study the ecological differences between it and Carex trisperma and also

further document the abundance of C. billingsii in bogs on the fringe of its distribution to determine whether

protection status is warranted.

TAXONOMIC TREATMENT AND KEY TO SPECIES

Leaves 0.8-1.9 mm wide, flat or thinly M-shaped, ligules 0.5-1.9 mm long, inflorescences (14-)23-55 mm long,

spikes per inflorescence (2-)3-4, terminal spikes with (1-)2-6 perigynia per spike Carex trisperma

Leaves 0.3-0.8 mm wide, filiform-involute, ligules 0.3-0.8(-1.2) mm long, inflorescences 14-32 mm long, spikes

per inflorescence 2-3, terminal spikes with 1-3 perigynia per spike Carex billingsii

NOMENCLATURE

Carex billingsii (O.W. Knight) C.D. Kirschbaum, comb. et stat. nov. Basionyu: Carex trisperma var. billingsii O.W. Knight,

Rhodora 8:185. 1906. Carex trisperma f. billingsii (O.W. Knight) B. Boiven, Naturaliste Canad. 94:523. 1967. Tyre: U.S.A. MAINE:

Somerset Co.: Pleasant Ridge Twp., the drier portions of upper Jewett Bog, 5 Jul 1906, O.W Knight, J. Murdoch Jr., E.B. Chamberlain,

R.A. Ware, S. Rollins 5066 (HOLOTYPE: not seen, see below; isotype: GH!).

It is not certain where Knights holotype for C. t. var. billingsii was deposited. I have contacted curators at MAINE, VT and GH but they

were not able to locate the type specimen. A duplicate of the type (cited above) is located at GH and another specimen, Knight 2021

collected at the same locality and on the same date as the specimen that Knight cited as the type specimen is also at GH.

One untypified name referable to the Carex trisperma alliance exists, Carex quaternaria Sprengel (1826),

based on material from New Jersey. The type of Carex quaternaria appears to have been destroyed at Berlin.

However, the description states “Spiculis 4 floris" (spikes four-flowered) (Sprengel 1826) (presumably the

basis for the specific epithet quarternaria). Carex billingsii is smaller and averages fewer flowered spikes than

C. trisperma, and no specimen has been observed with four perigynia in a spike. Carex quaternaria is cer-

402 Journal of the Botanical R h Institute of Texas 1(1)

tainly the same entity as Carex trisperma, which can have up to six perigynia per spike. A neotype for Carex

quaternaria from New Jersey and having some spikes with 4 perigynia is selected below.

Species Description.—Carex billingsii. Plants glabrous, loosely cespitose; rhizomes with slightly fi-

brous or non-fibrous sheaths, basal sheaths dark to light brown. Fertile culms erect at anthesis, elongating

and arching toward the ground after fruiting, 20-36 cm with 2-3 spikes per inflorescence. Leaves of the

fertile culm (1)3 or 4 located on the lower 1/3 of the culm. Leaf blades filiform-involute above the sheath,

leaf margins fusing distally, 7-13 x 0.3-0.8 mm. Leaves triangular in cross section, with 1-2 large areas of

aerenchyma tissue on the left and right sides of the midrib. Ligule 0.3-0.8(-1.2) mm long, obovate or with

rounded apex. Sheaths hyaline, apex concave. Inflorescence 14-32 mm long, proximal bract 15-63(-72)

mm, often exceeding inflorescences. Base of proximal bract sometimes elongated and expanded 1.7-3.2(-4.1)

x 0.8-1 mm. Terminal spikes 3.3-4.7 x 1.9-3.2 mm with 1-3 perigynia per spike, lateral spikes 3.5-4.5 x

1.5-2.5C3.1) mm with 1-3 perigynia per spike. Spike with 2 staminate flowers and ascending to slightly

diverging perigynia. Proximal-most and penultimate spikes 2.2-6 mm apart, penultimate and distal-most

spike (when present) 6-23 mm apart. Perigynia 2.7-3.9 x 0.9-1.6(-1.8) mm widest just below the middle.

Beak of the perigynia 0.4-0.7 mm, truncate to bi-dentulate with teeth 0.1 mm long. Abaxial nerves of the

perigynia 14-19-25), adaxial nerves 9-12. Pistillate scales 2.2-3.2 x 0.9-1.5 mm with green midrib be-

tween whitish-green margins, apex acute (mucronate), 1/2-2/3 as long as the perigynia. Stigmas 0.03-0.1

mm wide. Staminate scales 2.2—4.3 x 0.9-1.4 mm with similar coloring as pistillate scales. Anthers 0.9-1.5

mm long. Achenes 1.3-2.0 x 0.8-1.4 mm, ovate, broadly elliptical or obspatulate. Base of the achene

rounded, truncate, cuneate or attenuate, apex rounded or truncate. Fruiting mid-June-September. Found

in sunny and in shaded areas of ombrotrophic, tamarack, spruce and cedar bogs, often on tops and sides of

sphagnum hummocks, along trails and small creeks, wet, sandy swales in NJ pine barrens, Chamaecyparis

Spach swamps bordering salt marshes in MA and interdunal string bogs in Quebec. Often associated with

ericaceous shrubs such as Kalmia angustifolia L., Chamaedaphne Moench, Ledum L. and Vaccinium L.

d: CANADA. NEWFOUNDLAND: Trepassey, Avalon Peninsula, 053°24'00"W, 46°43'00"N, 16 Aug 1924, Fernald,

m Dunbar 26376 (GH). ONTARIO: 6 Jul 1915, Frere Rolland-Germain s.n. (DAO); 12 Jul 1923, Malte s.n. (MICH); Middlesex, Sifton

(Byron) Bog, UTM Grid 734575, Map 40 1/14, 081°25'00"W, 43?00'00"N, 4 Jul 1991, Reznicek, McLeod 8810 (MICH). PRINCE EDWARD

ISLAND: Kings, Murray River, N of the river on Route 24, 062°32'00"W, 46°02'00"N, 22 Jul 1953, Erskine, Smith 2103 (DAO). QUEBEC:

Berthier, Lanoraie, 073?13'00"W, 45°58'00"N, 6 Jul 1932, F. Marie-Victorin, F. Rolland- Germain 49208 (MT); Kamouraska, Peat bog 1

mi NE of Riviere Ouelle, 069°49'00"W, 47°32'00"N, 6 Aug 1947, Calder 1282 (DAO); Labelle, NE du Grand lac Nominingue, pres de

la riviere Rouge, Bellerive, 27 Jul 1939, F. Lucien, F. Eloi 577 (MT); North Bay, 3 mi N of Tomiko River bridge along Hwy 11, N of North

Bay, 23 Sep 1959, Calder, Kukkonen 24287 (DAO), Gatineau, Masham Twp., Conc. V, (Outaouais), 13 km WNW of Wakefield, 31 F/9

156575 (UTM grid), Ottawa District, 076205" W, 45?40'N, 29 Jun 1988, Reddoch 398 (DAO); Les Sillons, W side of Hwy 199, S of Pont

du Detroit, N end of Ile du Hvre aux Maisons, 14 Aug 1998, Oldham 21198 (MICH).

UNITED STATES. CONNECTICUT. Hartford Co.: Burlington, 11 Jun 1921, Weatherby D2120 (GH). Hartford Co.: Black Spruce

bog S of Rte 168 near Congamond Lake, 22 Jun 1982, Mehrhoff 6343 (MICH). Litchfield Co.: Kent, 4 Jul 1930, Torrey s.n. (MICH). Tolland

Co.: bog near railroad tracks and Rte 1-84, 4 Jul 1990, Mehrhoff 13514 (MICH). MAINE. Aroostook Co.: T8N, R5W, 25 Jul 1941, Pease,

Bean 29029 (GH); Fort Fairfield, the Aroostook River Basin, 5 Jul 1940, Chamberlain 1715 (GH); E side of Hwy 2A (Alt 2) ca. 2.7 mi S of

Forkston-TAR2WELS township line. 0577721E 5082451N UTM Zone 19T (Units: km, Datum: NAD 83), 6Jul 2002, Kirschbaum s.n. (n/a).

Hancock Co.: Corea Heath, SW side of Hwy 195 ca 1 mi NW of Corea, 068°00'W, 44°23'N, 16 Jul 1992, Reznicek 9149 (MICH); head of

Torrey Pond, Deer Isle, 7 Jul 1915, Hill 2179, (GH). Somerset Co.: Jewett Brook Bog, NE of Jewett Pond, 150 ft NW of Jewett Brook and

past the turn to Jewett Pond campsites, 4.7 mi NW of Cross rd. along Rowe Pond Rd., Pleasant Ridge Twp. 0420195E 4998596N UTM

Zone 19T (Units: km, Datum: NAD 83), 7 Jul 2002, Kirschbaum s.n. (n/a). Washington Co.: 8 Jul 1993, Reznicek 9620, (MICH). MAS-

SACHUSETTS. Barnstable Co.: Harwich, 3 Aug 1913, Weatherby s.n. (GH). Berkshire Co.: Lost Pond, Becket, 16 Jul 1909, Hoffmann

s.n. (GH). Middlesex Co.: Littleton, No date, Manning(?) s.n. (GH). Middlesex Co.: Tewksbury, no date, Gray(?) s.n. (GH). MICHIGAN.

Washtenaw Co.: Gorman Lake Bog on the S end of Gorman Lake off Lindley Rd. in Waterloo State Recreation Area, ca. 8 mi NW of

Dexter., 21 Sep 2002, Kirschbaum s.n. (n/a). NEW HAMPSHIRE. Cumberland Co.: Bog Pond, 0.75 km off Haskell Hill Rd. by way of

power line right-of-way on private property. Northern Harrison twp., 0370429E 4886298N UTM Zone 19T (Units: km, Datum: NAD

83), 7 Jul 2002, Kirschbaum s.n. (n/a). Hillsborough Co.: off hiking trail on the W side of Mud Pond in Fox State Forest. Hillsborough

twp., 0264272E 4780594N UTM Zone 19T (Units: km, Datum: NAD 83), 4 Jul 2002, Kirschbaum s.n. (n/a). Strafford Co.: between Mt.

Hussey and Mt. Chesley, Farmington, 20 Jul 1967, Hodgon 15735 (GH); N of Scruton Pond, on Scruton Pond Rd., Barrington twp., 4 Jul

2002, Kirschbaum s.n. (n/a). NEW JERSEY. Burlington Co.: Chatsworth, 1 Jul 1932, Hermann 3380, (MICH). Ocean Co.: Forked River,

No date, Churchill s.n. (GH); Pole Bridge Brook, 2 mi WSW, Whitings, 17 Jul 1914, Long 10321 (GH). NEW YORK. Allegany Co.: Moss

Kirschbaum, Taxonomy of Carex trisperma varieties 403

Lake Bog on Moss Lake off Sand Hill Rd. ca. 2.5-0.0 mi SW of Houghton, Caneadea twp. 0731623E 469767N UTM Zone 19T (Units:

km, Datum: NAD 83), 11 Jul 2002, Kirschbaum s.n. (n/a). PENNSYVANIA. Susquehaanna Co.: 2 mi N of Burnwood, W side of Ball

Lake, 24 Aug 2003, Robert F.C. Naczi 10065 (DOV). VERMONT. Franklin Co.: 055?45'00" W, 51?33'00"N, 25 Jul 1912, Woodward s.n.

(GH). Grand Isle Co.: S. Alburg, 16 Jul 1939, Knowlton s.n. (GH).

Carex trisperma Dewey, Amer. J. Sci. Arts 9:63. 1825. Basionwm: Neskiza trisperma (Dewey) Raf., Good Book. Amenit. Nat.

Philad. 27. 1840. Type: U.S.A. Massacuusetts: Berkshire Co.: Williamstown, without collection date, C. Dewey (Lectotyre designated

here: GH 63033).

U.S.A. Massachusetts. [probably Bershire Co.]: Williamstown-Deerfield, grows in the form of bogs in sphag-

nous place among hills, [no collection date], C. Dewey (GH 27464).

Carex quaternaria Spreng., Systema Vegetabilium 3:809. 1826. Tyre: U.S.A. NEw JERSEY. Sussex Co.: Culvers Gap, swamp, 30 May 1919,

Ludlow Griscom, No. 14336 (NEotveE designated here: GH-s.n.)

Species Description. Carex trisperma. Plants glabrous, loosely cespitose; rhizomes with slightly fibrous

or non-fibrous sheaths, basal sheaths dark to light brown. Fertile culms erect at anthesis, elongating and

arching toward the ground after fruiting, 15-65 cm with (2-)3-4 spikes per inflorescence. Leaves of the

fertile culm 23 or 4(5) located on the lower 1/3 of the culm. Leaf blades, appearing flat apically and deeply

channeled, or keeled on the abaxial surface, (9-)13-18(224) x 0.8-1.9 mm. Leaves thinly M-shaped in cross

section with 4-5 areas of aerenchyma tissue on each side of the midrib. Ligule 0.5-1.9 mm long, obovate or

with rounded apex. Sheaths hyaline with sheath apex concave. Inflorescence (14-)23-55 mm long, proximal

bract 28—74 mm, often exceeding inflorescences. Base of proximal bract sometimes elongated and expanded

1.5-3.7 x 0.6214 mm. Terminal spikes 4.4—6.5 x 2.5—4.4 mm with (1-)2 perigynia per spike, lateral spikes

3.6—5.0 x 2.4-4.7 mm, with (1-)2-6 perigynia per spike. Spike with 2 staminate flowers and ascending

to slightly diverging perigynia. Proximal-most and penultimate spikes 4.3-12 mm apart, penultimate

and distal-most spike (when 3 spikes are present) (14)21-33(446) mm apart. Perigynia 2.23.7 x 1.2-1.7

mm widest just below the middle. Beak of the perigynia 0.4-0.7 mm, truncate to bi-dentulate with teeth

0.1-0.2 mm long. Abaxial nerves of the perigynia 13-21, adaxial nerves 7-14. Pistillate scales 2.2-3.2 x

1.1-1.7 mm with green midrib between whitish-green margins, apex acute to mucronate, 1/3-2/3 as long

as the perigynia. Stigmas 0.05-0.1 mm wide. Staminate scales 2.4-4.6 x 0.5-1.2 mm with similar color-

ing as pistillate scales. Anthers 1-1.5 mm long. Achenes 1.7-2.2 x 1.1-1.5 mm, ovate, broadly (narrowly)

elliptical or obspatulate. Base of the achene cuneate or attenuate, apex rounded or truncate. Fruiting early

June—August. Found in deep to partial shade and occasionally in sunny areas of ombrotrophic sphagnum

bogs and cedar and spruce swamp forests and wet-mesic deciduous woods. Associated with Picea rubens

Sarg., Abies balsamea (L.) P. Mill., Taxus canadensis Marsh., Acer rubrum L. and Pinus strobus L. in bogs and

swamps of New England, Chamaecyparis Spach swamps in MA, Thuja occidentalis L., Picea mariana (P. Mill.)

B.S.P., Acer rubrum L. and Larix P. Mill. swamp forests from New York and Ontario westward and open

Sphagnum-Ericaceae bogs across its range, especially northward.

d: CANADA. ALBERTA: Swan Hills Twp., 65-R 9-W5M, 115?45'00"W, 54?45'00"N, 7 Aug 1960, Pegg 880 (DAO).

BRITISH COLUMBIA: Barkerville, 12.5 mi by road NNE of Barkerville on road to Bowron Lake, 121°23'00"W, 54°14'00"N, 9 Aug

1954, Calder, Savile, Ferguson 14328 (DAO). NEW FOUNDLAND: Virginia Water, near St. John's, No date, Robinson, Schrenk 100 (GH);

near Isthmus Cove, Pistolet Bay, 5 Aug 1925, Wiegand, Gilbert, Hotchhiss 27612 (GH); Forteau, Belle Isle, 14 Aug 1925, Long 27614 (GH);

Goose Bay, 060°21'W, 53°19'N, 10-12 Aug 1949, Schofield 749 (DAO). MANITOBA: Lac Du Bonntet, 7 Jul 1949, Breitung 7474 (DAO);

Lac du Bois, along trail to South Lake, 095°40'W50°16'N, 18 Aug 1982, Keleher 954 (DAO); near Keyhole Lake, inland from NW shor of

Tod Lake., 101°45'50"W56°34'06"N, 20294, 24 Jul 1955, Ritchie 1268 (DAO); NW side of Tulibi Lake, 19 Jun 1955, Ritchie 822 (DAO);

Taiga Biological Station, Wallace Lake., 095°20'W51°02'N, 5 Jul 1979, Keleher 737 (DAO). NOVA SCOTIA: Antigonish, vicinity of Sea-

scape Cottages, W of Bayfield, 062°00'00"W, 45°37'00"N, 20 Jun 1992, Oldham 13841 (MICH). ONTARIO. Nipissing District: 0.25

mi S of Jack Lake on old logging road, 078°33'00"W, 45°35'00"N, 12 Jun 1958, Kazdan 617 (DAO); Mowe L. Rd just S. of Plummes L. on

recently built road shoulder, 090°43'W, 48°20'N, 27 Jun 1981, Garton 20180 (MICH). Norfolk Regional Municipality: Turkey Point

Provincial Park, Wilderness Zone, Delhi Tp. Mun., UTM 545265 (401/9) (sq17NT52), 080°19'00"W, 42°42'00"N, 27 May 1987, Oldham,

Sutherland, Kirk 7206 (MICH); Bruce, Schmidt L, 29 Jul 1987, Johnson s.n. (MICH). SASKATCHEWAN: S shore of Lake Athabasca, E of

William River. Vicinity of “Little Gull” Lake., 109°00'W59°0I'N, 22826, 29 Jun 1962, Angus 295-62 (DAO); ca. 5 km SSE of Archibald

Lake, ca. 13 km NNW of Davy Lake, 108°30'W58°58'N, 29078, 8 Jul 1979, Harms, 27411 (DAO); Wollaston Lake Rd, mile 10, Hwy 105,

404 Journal of the Botanical R h Institute of Texas 1(1)

ca. 130 mi N of La Ronge., 103°37'W56°20'N, 22 Jul 1973, Ternier s.n. (DAO). QUEBEC: 075°55'51"W, 45°38'18"N, 12 Jul 1922, Malte

384/22 (MICH); Rupert House, E coast of James Bay, 18 Jul 1929, Potter 91 (GH); Red Bay, N shore of the Gulf of St. Lawrence, 26 Jul

1929, Abbe 1062 (GH); Grand Lake, Blue Grass Brook, near Camp 11, 5 Aug 1951, Rolueau 2206 (MICH); Abiti-East, Harricanaw River

Maizerets Twp. 0.75 mi E of river, 078°03'W, 49°11'N, 24 Jul 1958, Bentley 58163 (DAO).

UNITED STATES. CONNECTICUT. Windham Co.: Windham, 17 Jun 1914, Weatherby 3472 (GH). ILLINOIS. Lake Co,: 3

Aug 1906, Gleason, Shobe s.n. (MICH). MASSACHUSETTS. Plymouth Co.: Norwell, 6 Jun 1932, Knowlton s.n. (GH). Worcester Co.:

17 Jun 1938, Weatherby, Weatherby s.n. (MICH). MAINE. Grand Lake.: Blue Grass Brook, near Camp 11, 5 Aug 1951, Rolueau 2206

(MICH). Hancock Co.: Central Tract, Brooklin, 2 Aug 1914, Hill 1769 (GH). Lincoln Co.: Cathedrdal Woods, Monhegan Island, 29 Jun

1919, Jenney, Churchill, Hill 3170 (GH). Piscataquis Co.: Squaw Moosehead Station, 9 Jul 1917, Sanford 60605 (GH). Ostesgo Co.: N

side Old State R d . (F38) ca. 13.5 Km E of Otsego Lake , SE 1/4 sect. 10, T29N R2W. Lat. & Long. 44? 54' 55" N, 84° 31' 46" W, 15 Jul

2002, A.A. Reznicek 11367 (MICH). Piscataquis Co.: Greenville Junction, 8 Jul 1917, Sanford 6032 (GH). Somerset Co.: Jewett Brook

Bog, NE of Jewett Pond. 150 feet NW of Jewett Brook and past the turn to Jewett Pond campsites, 4.7 mi NW of Cross Rd. along Rowe

Pond Rd., Pleasant Ridge Twp. 0420195E 4998596N UTM Zone 19T (Units: km, Datum: NAD 83), 7 Jul 2002, Kirschbaum s.n. (n/a);

Pleasant Ridge, Upper Jewett Bog, 5 Jul 1906, Murdoch 2022 (GH). Washington Co.: salt marsh along Sandy R, behind Sandy R Beach,

3.5 mi ENE Jonesport, E side of Hwy 187, 067°32'W, 44°34'N, 7 Jul 1993, Reznicek 9615 (MICH). MICHIGAN. Baraga Co.: along US

Highway 41 near W end of Lake Michigamme, 22 Jun 1950, Richards 3184 (MICH). Berrien Co.: Buchanan Bog, 30 May 1930, Herbert

s.n. (MICH). Iron Co.: Larix Bog, near Deer Lake, 10 m NE of Crystal Falls, 088°20'W, 46°24'N, 8 Aug 1934, Grass! 8057 (MICH). Lake

Co.: Ca. 2.5 mi N of Bristol, 18 Jul 1973, Voss 14227 (MICH). Schoolcraft Co.: T42N, R16W, Sect. 11 NW1/4NE1/4, SW1/4SE1/4, 2 Sep

1971, Henson 269 (MICH). Washtenaw Co.: Gorman Lake Bog on the S end of Gorman Lake off Lindley Rd. in Waterloo State Recreation

Area, ca. 8 mi NW of Dexter, 21 Sep 2002, Kirschbaum s.n. (n/a). MINNESOTA. Aitkin Co.: 0.7 mi S of McGrath, 25 Jun 1940, Moore,

Moore 13263 (MT). Cook Co.: S of Grand Portage along Highway 61, 15 Aug 1987, Castaner 9986 (MICH). NEW HAMPSHIRE. Coos

Co.: Magalloway River, 3 Aug 1914, Pease 16207 (GH); Bray Hill Bog, NE of Whitefield of county road 116 (Jefferson Rd.) on the NW side

of Bray Hill Rd., Whitefield twp., 0297240E 4920187N UTM Zone 19T (Units: km, Datum: NAD 83), 7 Jul 2002, Kirschbaum s.n. (n/a).

Cumberland Co.: Bog Pond, 0.75 km off Haskell Hill Rd. by way of power line right-of-way on private property, northern Harrison

twp., 0370429E 4886298N UTM Zone 19T (Units: km, Datum: NAD 83), 7 Jul 2002, Kirschbaum s.n. n/a). Hillsborough Co.: off hiking

trail on the W side of Mud Pond in Fox State Forest. Hillsborough twp., 0264272E 4780594N UTM Zone 19T (Units: km, Datum: NAD

83), 4 Jul 2002, Kirschbaum s.n. (n/a). Strafford Co.: N of Scruton Pond, on Scruton Pond Rd., Barrington twp., 4 Jul 2002, Kirschbaum

s.n. (n/a). NEW YORK. Allegany Co.: Moss Lake Bog on Moss Lake off Sand Hill Rd. ca. 2.5-3.0 mi SW of Houghton, Caneadea twp.

0731623E 469767N UTM Zone 19T (Units: km, Datum: NAD 83), 11 Jul 2002, Kirschbaum s.n. (n/a). Hamilton Co.: border of The

Plains, South Branch of the Moose River, 10 Aug 1950, Smith, Weaver 7577 (DAO). Oneida Co.: Knieskern s.n. (MICH). County unknown,

Adirondack Mountains, 20 Jun 1936, Killip 31814 (MICH). OHIO. Stark Co.: thickets in center of Brewster Bog, 0.2 mi SE, jct. of M

Eaton St and Wellbrook Ave, Navarre Quad, 10 Jul 1984, Cusick, Denny, Munch 23633 (MICH). PENNSYVANIA. Elk Co.: ca. 11 km S of

town, 078°15'W, 41°L5'N, 18 Jun 1997, Grund 1918 (MICH). Tioga Co.: 4.5 mi NW of Morris, 077°20'51"W, 41°37'07"N, 29 May 1975,

Rothrock 309 (MICH). VERMONT. Franklin Co.: Berkshire, 15 Jun 1912, Underwood 2122 (GH). WISCONSIN. Jackson Co.: Bear

Bluff Twp, in the old bed of Glacial Lake, T21N, ROIE, Sect. 29, 19 Jun 1958, Hartley 3969 (DAO). Iron Co.: between Hwy 51 and Rice

L, near CNW RR, 2 mi NW of Mercer, T43N, ROSE, Sect. 26 NW1/4, 3 Jul 1976, Cochrane, Cochrane 7547 (MICH). WEST VIRGINIA.

Giles Co.: Big Good Bed at head waters of Little Stony Creek, Salt Pond Mt., 9 Jun 1946, Wood 5972 (GH). Pocahontas Co.: 0.2 mi S of

park office, Droop Mountain Battlefield State Park, NW, US Rte 219, N of Droop, 1 Jun 1990, Cusick 28918 (MICH); ultimate headwaters

of First Fork of Shavers Fork Creek, ca. 2 air mi 110 degrees from Shavers Fork crossing at Randolph County line, ca. 4.2 air mi NE of

Cass Scenic RR State Park off For Serv Rd 235, Back Allegheny Mtn, Monongahela National Forest, 27 Jul 1994, Nelson 15881 (MICH).

Tucker Co.: beyond Little Blackwater, near camp 70 near 3200f, Canaan Valley, 1 Aug 2007, Allard 10106 (GH).

ACKNOWLEDGMENTS

This project would not have been possible without the support, mentoring, and constructive criticism of

Gary L. Hannan, Marianne M. Laporte, Catherine E. Bach, and Anton A. Reznicek. Kari Jensen Kirschbaum,

Andrew L. Hipp and Anton A. Reznicek and an anonymous reviewer provided valuable comments and sug-

gestions for earlier versions of this manuscript. I also thank Paul Durica, Michael Farmer, Justin Hohn, Kari

Jensen Kirschbaum, and Ashley Wilt for their assistance with field, laboratory, and database work. I thank

William J. Crins, Geoffrey Hall, Justin Hohn, Robert F.C. Naczi, Todd Ristau, Carl J. Rothfels, Anton A.

Reznicek, and Dan Sperduto for help in collecting specimens. I thank Jane Gillies for her work and report

on the SEM. I appreciate the help of the curators at MICH, GH, DAO, and MT for sending herbarium speci-

mens. I thank Mark Berres and Andrew Hipp for technical support on AFLPs. Funding for this project was

provided by the Biology Department at Eastern Michigan University, the Office of Graduate Studies at Eastern

Michigan University, the C.F. Hanes Fund, the University of Michigan Herbarium, and Rackham School of

Kirschbaum, Taxonomy of Carex trisperma varieties 405

Graduate Studies at Michigan University. I also thank the curators at WIS, MO, MSC, TENN, UWSP, OS

and CLM who searched their collections for Carex trisperma and Carex billingsii specimens.

REFERENCES

ABBOTT, L.A., F.A. Bisev, and D.J. Rocers. 1985. Taxonomic analysis in biology: computer, models, and databases.

Columbia University Press, New York, NY.

ANDERSON, D.S., R.B. Davis, S.C. Rooney, and C.S. CampeeLL. 1996. The ecology of sedges (Cyperaceae) in Maine peat-

lands. Bull. Torrey Bot. Club. 123:100-110.

APPLIED BioSystems. 2000. AFLP Plant Mapping (#4303146). Applied BioSystems, Foster City, CA.

BERNARD, J.M., D. SOLANDER, and J. Kver. 1988. Production and nutrient dynamics in Carex wetlands. Aquatic Bot.

30:125-147.

Berres, M.E. 2002. "Studies in Avian Genetic Population Structure” University of Wisconsin Madison, http://ravel.

oology.wisc.edu/sgaap/AFLP. html/AFLP.htm. (accessed September 2003).

Bowen, B. 1967. Énumération des plantes du Canada. Naturaliste Canad. 94:523.

Braverstock, PR. and C. Moritz. 1996. Project design. In: D.M. Hillis, C. Moritz, and B.K. Mable, eds., Molecular sys-

tematics, 2"? edition. Sinauer Assoc. Inc., Massachusetts. Pp. 249-320.

Brown, J.H. and M.V. Lomotino. 1998. Biogeography, 2"? edition. Sinauer Associates, Inc., Sunderland, MA.

FELSENSTEIN, J. 2004. PHYLIP (Phylogeny Inference Package) version 3.6. Distributed by the author. Department of

Genome Sciences, University of Washington, Seattle.

GLEASON, H.A. and A. Cronauist. 1991. Manual of vascular plants of northeastern United States and Adjacent

Canada. Princeton, NJ.

Hipp A.L., A.A. Reznicek, PE. RotHrock, and J.A. Wegen. 2006. Phylogeny and classification of Carex Section Ovales

(Cyperaceae). Int. J. Plant Sci. 167:1029-1048

Hipp, A.L. 2004. Molecular Systematics of Carex section Ovales (Cyperaceae). Ph.D. dissertation. Univ. of Wisconsin,

Madison.

JACCARD, P. 1908. Nouvelles recherches sur la distribution florale. B. Soc. Vaud. Sci. Nat. 44:223-270.

KARDOLUS J.P, H.J. van Eck, and R.G. VAN DEN Bera. 1998. The potential of AFLPs in biosystematics: a first application

in Solanum taxonomy (Solanaceae). Pl. Syst. Evol. 210:87-103.

KiRSCHBAUM, C.D. 2005. Taxonomy of Carex trisperma varieties (Cyperaceae) based on morphology, phytogeog-

raphy, ecology, and Amplified Fragment Length Polymorphisms (AFLPs). M.S. thesis. Eastern Michigan Univ.,

Ypsilanti.

KNIGHT, O.W. 1906. A new variety of Carex trisperma. Rhodora 8:185

LANDRY, PA. and F.J. Lapointe. 1996. RAPD problems in phylogenetics. Zool. Scr. 25:283-290.

Lessa, E. P. 1990. Multidimensional analysis of geographic genetic structure. Syst. Zool. 39:242-252.

McCune, B. and M.J. Merrorp. 1997. PC-ORD: Multivariate Analysis of Ecological Data, Version 3.0. MjM Software

Design, Gleneden Beach, OR.

Nei, M. and W.H. Li. 1979. Mathematical model for studying genetic variation in terms of restriction endonucleases.

Proc. Natl. Acad. Sci. USA. 76:5269-5273.

RisEBERG, L.H., T.E. Woop, and E.J. Baack. 2006. The nature of plant species. Nature 440:524-527.

SokaL, R.R. and P.H. SNEATH. 1963. Principles of numerical Taxonomy. W.H. Freeman and Company, San Francisco,

CA.

SPRENGEL, C.P.). 1826. Systema Vegetabilium, edito decimal sexta 3:809. Gottingae, sumtibus Librariae Dieteri-

chianae.

Toivonen, H. 2002. Carex Linnaeus sect. Glareosae G Don. In: Flora of North America Editorial Committee, eds., Flora

of North America. Vol. 23, Magnoliophyta: Commelinidae (in part): Cyperaceae. Oxford Univ. Press, Oxford

and New York. Pp. 311-321.

Vos, P, R. Hocers, M. Bleeker, M. REUANS, T. VAN DE LEE, M. Hornes, A. FRurERS, J. Por, J. PELEMAN, M. KUIPER, and M. ZABEAU.

1995. AFLP: a new technique for DNA fingerprinting. Nucleic Acids Res. 23:4407-4414.

406 Journal of the Botanical R h Institute of Texas 1(1)

BOOK REVIEWS

Epwanp C. Situ. 2006. Incredible Vegetables from Self Watering Containers. (ISBN 978-1-58017-556-2,

pbk.). Storey Publishing, 210 Mass MoCA Way, North Adams, MA 02147, U.S.A. (Orders: www.storey.

com, sales@storey.com, 1-800-865-3429). $19.95, 254 pp., color pun maps, index, 81⁄2" x 1074".

The book, Incredible Vegetables from Self Watering Containers, is divided into tl j t that describe a new way to grow vegetables

(with containers), the materials needed to get started and how to manage a container garden, and a detailed list of the typical varieties

of edible plants that will grow in containers (this section comprises the largest portion of the boo

Many people believe that containers or pots are solely used for flowers. The first section informs the reader that this is not the

case. Growing plants in pots has been around for generations. However, growing vegetables in pots is a relatively new process. The

author mentions that there are a few key problems that one might encounter when planting a garden in pots: an increase in watering,

less productive plants compared to those found in the ground, or an inability to garden organically. The basic solution to these prob-

lems is to choose the correct container and proper soil mixture for your plant. For example, a gardener today has the ability to create a

self watering container, also known as a continuous-flow watering system. These require less time spent watering, keep the plant from

wilting or drying in between watering sessions, and allow nutrients to be kept in closer proximity to the plant’s root system. The water

in a continuous-flow Mind: yom is un up through the soil by the plant's root system and remains with the plant longer. This

| l t lly be lost through leaching. The soil mixture is also very important in growing

a productive plant. The author gives examples of the correct combination of peat moss, vermiculite or perlite, limestone, and compost

that should be used in containers. ps dn sn help n ea see pid ke itis for one to create a container garden.

In the second section, the autl d tools to create a profitable vegetable

container garden. Some important oe is “to start small" and do your research on the proper location and pot size for a particular

plant. This will make sure your gardening is successful. The author gives the reader great examples on how to mix and match several

plants in one pot, and lists the proper techniques for caring for each variety of plant. These techniques include when to water, proper

location (determined by the plants tolerance of shade or sun), management of pests, proper time to harvest or collect vegetables, and the

proper instructions on stowing containers through the winter. All of these helpful tips will help the reader become a better gardener.

The final section introduces the reader to a variety of edible vegetables, flowers, and herbs that grow well in pots. This section is well

labeled and it is easy for the reader to locate the exact plant they are looking for. The types of plants range from your basic garden varieties

such as tomatoes and cucumbers to edible flowers such as sunflowers and violas. Detailed care instructions are given for each particular

plant. These instructions include proper pot use, favorable weather conditions for each plant, particular bugs or pests that might appear

or infest the plant, and the right time to harvest your vegetables. This section will prove to be very helpful for a novice gardener.

This book would be an important tool for any gardener, ranging from an amateur to a professional. The author's use of detailed

instructions and descriptions can help motivate any reader to create their own container garden. The book was written in such a way

that those new to science and gardening would understand the importance of finding the perfect container and creating the proper soil

mixture. The reader will take away a deeper knowledge on how important soil type is to the longevity of their plants, and how one needs

to find a good location for each specific plant. I would recommend this book to any gardener that is limited on the proper space for a

garden or tempted to try their own container garden.—Keri McNew, MS Biology, Project Manager, Botanical Research Institute of Texas, 509

Pecan Street, Fort Worth, TX 76102-4060, U.S.A.

ALLAN M. ARMITAGE. 2006. Armitage’s Native Plants for North American Gardens. (ISBN 0-88192-760-0,

hbk.). Timber Press Inc, 133 S.W. Second Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders:

www.timberpress.com, mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax).

$49.95, 451 pp., 443 color photographs, 7!^" x 10%".

Native plants often require less water and maintenance than non-native plants. Although not intended for members of the “right wing

of the Native Party" according the author, this handsome book is an excellent information source for North American gardeners who

want to introduce native plants into their gardens and landscaping. Plants are included based on their availability to mainstream gar-

deners. Over 630 species and cultivars are included, arranged alphabetically by plant genera. At the back of the book are useful lists of

native plants that fill particular garden needs, including drought-tolerant plants, water-loving plants, plants that attract butterflies or

hummingbirds, deer- and rabbit-resistant plants, and plants for different growing conditions from full sun to heavy shade. The author

provides entertaining commentary about his personal experiences with these plants along with detailed information including Latin

names, plant families, common names, habitat, cultivars, hardiness zones, maintenance, and recommended propagation. Fascinating

information on the etymology of the plants' Latin and common names are also eae In addition, this book is packed with over 400

color photographs. The useful sources and resources section includes lists of native plants, local and regional native

plant societies, internet sites, and useful books for the native plant gardener. I found this book to be packed full of useful information,

while remaining enjoyable and easy-to-read.— Marissa N. Oppel, Collections and Research Assistant, Botanical Research Institute of Texas,

509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A

J. Bot. Res. Inst. Texas 1(1): 406. 2007

THOMAS WALTER 1YPIEICATION PROJECT T

EH EKNOWN WALTER TYPES

Daniel B. Ward

Department of Botany

University of Florida

Gainesville, Florida 32611, U.S.A.

ABSTRACT

Thomas Walter, a South ma plantation owner and skilled amateur botanist, in the 1780s wrote a flora that was the first treat-

ment of American plants employing the binomial nomenclature and sexual classification system of Linnaeus. Walter’s Flora Caroliniana

contained many species new to science and whose names are of modern use. So that these names shall be used in a consistent way,

provision has been made for each name to be represented by a single specimen, its type. But Walter designated no types; later authors,

following internationally agreed-upon rules, have chosen specimens to serve as lectotypes or neotypes. A compilation is provided here

of all known replacement types of Thomas Walter names.

RESUMEN

Thomas Walter, un propietario de una plantación en Carolina del Sur y botánico amateur, en los afios 1780s escribió una flora que

constituyó el primer tratamiento de las plantas americanas usando la nomenclatura binomial y el sistema de clasificación sexual de

Linnaeus. La Flora Caroliniana de Walter contenía la ciencia y cuyos nombres son de uso moderno. Para que

estos nombres estén en uso de un modo consistente, se Wan hecho las provisiones necesarias para que cada Donee esté M

por un solo peon su tipo. Pero Walter no designó tipos; autores posteriores, siguiendo das, han

que sirvan mo lectotip tipos. Se aporta una compilación de todos los tipos edo conocidos

de re NC de THORS Walter,

The Thomas Walter Typification Project is an ongoing effort to determine appropriate type specimens for

the many names of new plant species described by Thomas Walter in his Flora Caroliniana (1788). Walter

himself left no herbarium, but he had access to the collections made by John Fraser in the Carolinas and

Georgia. Fraser's specimens, now in the Natural History Museum, London, often bear Walter's identifica-

tions or comments, and are at times chosen as replacement types by later authors. Or, when no specimen

is present in the Fraser collection, specimens gathered by other persons have been chosen by later authors

to serve as replacement types for Walter's names.

The Fraser collection, because of its association with the author of the Flora, is commonly known as the

"Walter Herbarium." It is in the form of a large folio containing 690 usually very small, often fragmentary

specimens collected by John Fraser during an eighteen-month trip to the American southeast, from the fall

of 1786 through the early spring of 1788. During that time Fraser met and was befriended by Walter who

examined and partially annotated at least part of Fraser's collection. Fraser also carried Walter's manuscript

to England and saw it to publication.

The Fraser collection of the Natural History Museum—perhaps better known as the “Fraser/Walter

herbarium" to distinguish it from the presence in Paris of another part of the specimens collected by Fra-

ser—has been described in detail as an initial step in the analysis of the Walter names (Ward, 2006). Some-

what more than half of the specimens have labels that carry Walter's handwriting, either an identification

or partial identification, or a comment. Approximately half of the specimens bear similar identifications

and/or comments by Fraser, while a significant number show the handwriting of both.

Though many authors have referred to the Fraser/Walter herbarium as having been collected by Walter,

or as having been the basis for his new names, there is no credible contemporary evidence that Walter used

the collection in this way. Even where his handwriting is present, it is far more probable that he formed most

of his descriptions from plants available in vivo and that he saw the specimens only after his manuscript was

complete or essentially so. This conclusion has been documented and justified elsewhere (Ward, 2007).

Such specimens, though contemporary in time, are of course irrelevant to typification of Walter's names.

J. Bot. Res. Inst. Texas 1(1): 407 — 423. 2007

£+sha D o ID

408 Journal of t h Institute of Texas 1(1)

One small category of names given by Walter, however, cannot be separated from linkage with the Fraser

collections. Though Walter (1788) stated his observations had come from a 50-mile radius of his plantation

on the Santee River in what is now Berkeley County, South Carolina, it has long been recognized that at

least some of his species are unknown on the Carolina coastal plain (Harper 1911; Ewan 1969). It appears

certain, as Fraser has claimed (1789), that a number of species not known in Walter’s area could have come

to him only as a result of Fraser’s far wider travels (Ward 1962a, 2007). When a specimen of such a species

is found in the Fraser/Walter herbarium it is reasonable to conclude, though gathered by Fraser, either the

individual specimen or other, supplementary material of the same collection was actually used by Walter

in his writing. Supplemental materials may indeed have been available, for Walter’s descriptions commonly

contain details not shown in the Fraser specimens.

The International Code of Botanical Nomenclature (Greuter et al., 2000) dictates rules that govern

determination of types. If, as here believed, no surviving specimens can be stated to be the only materials

used by Walter, no holotype of a Walter name is possible. If a species described by Walter and known only

outside his area is represented by a specimen collected by Fraser, the Fraser/Walter specimen may reason-

ably be designated as its lectotype. If, however, either no specimen has survived, or the species is known

in Walter’s area and may well have been described by him without use of the herbarium, a neotype may be

selected.

At times authors have assigned type designation that are at variance with the origin of the materials

as described here. In such cases the Code (Art. 9.8) permits the type designation to be corrected to the

appropriate status. Such corrections have been made here where appropriate. The convention is faithfully

followed, that lectotypes are “designated” and neotypes are “selected.”

The Code (Art. 7.10) requires that a designation of lectotype or selection of neotype is without standing

unless published. However the Code, especially in former years, has not provided guidance for a standard

mode or place of publication. With this latitude, authors have used diverse outlets in publishing their type

citations. Commonly, of course, typifications appear as a component of monographic treatments or revisions.

But typifications have also been encountered in free-standing floristic notes, in proposals for conservation

of a non-legitimate name, in captions of illustrations, and even in a pre-meeting abstract of a paper to be

presented. All, if effectively published and if the type element is clearly indicated as such by the typifying

author (Art. 7.11), are valid, and are accepted here.

Names are also found to be cited as having been typified when the circumstances of their publication do

not justify type status. Notably, some members of a series of identifications by Hitchcock (1905), of grasses

in the Fraser/Walter herbarium, are simply references to certain specimens and fall short of the author hav-

ing treated them as types. Several type selections in theses and dissertations, though clearly stated, also fail

because of the non-published format of their presentation. Such nonvalid typifications are excluded here.

The present tabulation of 67 published Walter types includes 13 that are lectotypes (8 of them corrected

to that status) and 52 neotypes (34 of them corrected). Two names listed as neotypes are scheduled for

publication elsewhere (Ward, in press). The designations are of specimens in the Fraser/Walter herbarium

(43 names), as well as in other herbaria (22 names). The name used by Walter (Flora Caroliniana 1788) is

given, with appropriate page number. The modern name for each is also listed, in most cases employing

nomenclature used in the current flora of the Carolinas (Radford et al. 1968). Frequently used synonyms

may also be given. [An index of all relevant Walter names and modern equivalents will be provided later in

the present series.]

The names are presented in alphabetical order following Walter's usage. A discussion and/or justification

accompanies each name. Typifying authors, with their determination of lectotypes or selection of neotypes,

are cited. Further comments may include a simple statement of range and frequency in the Carolinas (in-

dicative of the probability that Walter knew the plant in vivo); reference to any appropriate specimens in the

Fraser/Walter herbarium (with designators as in Ward, 2006); and notation of whatever handwriting may

be on the label.

Ward, Thomas Walter Typification Project Il: known Walter types 409

TYPIFICATIONS

WALTER'S NAME: Aesculus parviflora Walter (p. 128)

MODERN NAME: Aesculus parviflora Walt.

Rare in SC (one county). Spm. 62a-A was labeled *Juglans Alba nova" by Fraser. Rembert (1984) has designated

this specimen, Fraser/Walter 62a-A, [1787] (BM), made by Fraser in South Carolina along the Savannah River

across from Augusta, Georgia, as LECTOTYPE of Aesculus parviflora Walt.

WALTER'S NAME: Andromeda ferruginea Walter (p. 138)

MODERN NAME: Lyonia ferruginea (Walt.) Nutt.

Nearly absent from SC (2 counties); frequent in adjacent northeastern GA, Judd (1981: 411) designated a

specimen, Fraser s.n., [1787] (P), from South Carolina or Georgia, as LECTOTYPE of Andromeda ferruginea Walt.,

basionym of Lyonia ferruginea (Walt.) Nutt. This specimen was among the materials Fraser sold to Charles

Louis L'Heritier in Paris (Fraser 1789). Although no indication is given that Walter saw or used that particular

Fraser specimen, the absence of the species from the area in which Walter directly obtained his materials sug-

gests that the specimen may have been part of a collection obtained by Fraser in his wider travels and utilized

by Walter in forming his description. A fragment (spm. 6-C) in the Fraser/Walter herbarium was labeled as

"Andromeda ferruginea” by Fraser, but has been identified as Lyonia fruticosa (Judd 1981: 419—420).

WALTER'S NAME: Angelica lobata Walter (p. 115)

MODERN NAME: Ligusticum canadense (L.) Britt.

Nearly absent from the SC coastal plain, but common inland; likely a Fraser discovery. Spm. 7-C, a crumpled

vegetative scrap, was labeled “Angelica” by Fraser. It was identified as Ligusticum canadense by Fernald and

Schubert (1948: 217), and called “the type.” Though Walter’s name will doubtless remain in synonymy of

Ligusticum canadense, Fraser/ Walter 7-C [1787] (BM) is corrected here to Lectotype of Angelica lobata Walt. (=

Ligusticum canadense (L.) Britt.).

WALTER'S NAME: Anonymos bracteatla] Walter (p. 181); nom. illegit.

MODERN NAME: Zornia bracteata Walt. ex Gmel.

Frequent in eastern SC. No specimen has been identified. Walter's name is illegitimate, but his description

still serves as the foundation for Gmelin's name (Ward 1962b). Mohlenbrock (1961: 30) selected Duncan

11557, [29 July 1950] (US), from McDuffie County, Georgia, as NEOTYPE for Zornia bracteata Walt. ex Gmel.,

with duplicate (GA) as ISONEOTYPE.

WALTER'S NAME: Anonymos graminifollia] Walter (p. 197); nom. illegit.

MODERN NAME: Vernonia angustifolia Michx.

Frequent throughout. Walter's name was omitted by Gmelin (1792). The name is illegitimate (Ward 1962b),

and the epithet cannot be transferred. The epithet was used by Willdenow (1803), but applied to a species

of Liatris. Willdenow's name was formed independently and is not a transfer; though he referred to Walter,

he gave a new diagnosis. A specimen (spm. 32-A) labeled “Chrysocoma affinis F 309” was identified as Liatris

graminifolia Willd. by Gaiser (1950: 414), but rejected as type of Anonymos graminifolia. [Gaiser was correct

in this rejection, though her reason was doubt as to its authenticity; Walter's Anonymos graminifolia, as de-

termined by his identification of other Fraser specimens, was Vernonia angustifolia.] Gaiser, apparently in

belief that the plant Walter had described was a Liatris, erroneously selected White s.n., 25 Oct 1948 (GH),

a plant of Liatris graminifolia Willd., from Wilmington, New Hanover County, North Carolina, as Walter's

“type.” Though Gaiser's incorrect typification of an illegitimate name is without application, in the interest

of completeness it is here listed and corrected to NEOTYPE for Anonymos graminifolia Walt.

WALTER'S NAME: Ánonymos procumbens Walter (p. 86); nom. illegit.

MODERN NAME: Houstonia procumbens (Walt. ex Gmel.) Standley [= Hedyotis procumbens (Walt. ex Gmel.)

Fosberg; Poiretia procumbens Gmel.]

Frequent to common throughout. There is no specimen. Lewis (1966) has selected Palmer s.n., 2-10 June

410 Journal of the Botanical R h Institute of Texas 1(1)

1902 (US), from Charleston County, South Carolina, as NEOTYPE for Poiretia procumbens Gmelin, basionym

of Houstonia procumbens (Walt. ex Gmel.) Standley.

WALTER'S NAME: Anonymos rotundifolia Walter (p. 181)

MODERN NAME: Crotalaria rotundifolia Walt. ex Gmel. [= Crotalaria angulata Mill.)

Common in eastern SC. Spm. 67-D has been termed “type” by authors (Fernald & Schubert 1948; Ward

1962b; Windler 1974). The label CLupinus affinis") is in Walter's hand. Since this species would surely have

been familiar to Walter near his home, and the label indicates he did not recognize it to be his "Anonymos

rotundifolia,’ there is no reason to believe this specimen was used by him. Its designation as type cannot be

dismissed, but Fraser/Walter 67-D, [1787] (BM), the foundation for Crotalaria rotundifolia Gmelin, is corrected

here to NEOTYPE. Fernald and Schubert's argument (1948: 202-203), equating Walter’s plant with Crotalaria

maritima Chapm., is incorrect; Chapman’s type came from Cape Sable (“Palm Cape”), southernmost penin-

sular Florida, and differs in leaf form and pubescence.

WALTER'S NAME: Anonymos sessifollia] Walter (p. 108); nom. illegit.

MODERN NAME: Mitreola sessilifolia (Walt. ex Gmel.) D. Don [= Cynoctonum sessilifolium Walt. ex Gmel.]

Common in eastern SC. Spm. 117-B is a slender stem of poor diagnostic character, marked with Fraser's

number 685. It was labeled *Genus nov. Pentand digyn" by Walter, not recognized by him as his Anonymos

“sessifolia.” Walter would have had living materials available; spm. 117-B could scarcely have been used by

him in forming his description. Walter's name is illegitimate, but his description still serves as the foundation

for Gmelin's name (Ward 1962b). Leeuwenberg (1974: 21) has designated no. 685 as *holotype" of Mitreola

sessilifolia. In view of Walter's failure to recognize the plant as his own, Fraser/Walter 117-B, [1787] (BM) is

here corrected to NEOTYPE for Cynoctonum sessilifolium Gmel., basionym of Mitreola sessilifolia (Walt. ex Gmel.)

D. Don. Because the specimen scarcely shows useful diagnostic features, an epitype would be welcome.

WALTER'S NAME: Anonymos setaclea] Walter (p. 170)

MODERN NAME: Agalinis setacea (Walt. ex Gmel.) Raf.

Frequent throughout. Pennell (1920: 282) stated the “type” had been identified in the British Museum by

A. B. Rendle. This presumably is spm. 51-A. Walter's name is illegitimate, but his description still serves as

the foundation for Gmelin's name (Ward 1962b). The specimen bears Walter's hand: “Gerardia...terminalis.”

Though very unlikely to have been used by Walter in preparing his description, its somewhat indirect des-

ignation as type by Pennell restricts further choice; his designation of Fraser/Walter 51-A, [1787] (BM) is here

corrected to NEOTYPE for Gerardia setacea Gmel., basionym of Agalinis setacea (Walt. ex Gmel.) Raf.

WALTER'S NAME: Anthoxanthum giganteum Walter (p. 65)

MODERN NAME: Erianthus giganteus (Walt.) Beauv.

Common throughout the SC coastal plain. Spm. 113-B bears the label “Anthoxanthum” in Fraser's distinctive

hand. Hitchcock (1905: 33) stated the specimen, since it “agrees with the description and is the only species

of the genus" that Walter described, “may be taken as the type.” Gandhi and Dutton (1993), without refer-

ence to a specific specimen, noted BM to contain the Walter “type.” The abundance of this species within

Walter's territory and the probability that he knew it in the living state (he noted the height to be 8 feet)

make it unlikely this Fraser specimen was given any weight by him. However, having been emphasized

by Hitchcock and noted by Gandhi and Dutton, Walter's specimen can continue to serve by correction of

Hitchcock's use of Fraser/Walter 113-B, [1787] (BM) to that of NeotyPe for Anthoxanthum giganteum Walt., the

basionym of Erianthus giganteus (Walt.) Beauv.

WALTER'S NAME: Arundo gigantea Walter (p. 81)

MODERN NAME: Arundinaria gigantea (Walt.) Muhl.

Common throughout. Hitchcock (1905: 53) identified a specimen (spm. 113-A) as this species, but did not

designate it as a type. McClure (1973: 26) took that step, but called the specimen the “Holotype.” He noted

the accompanying label to read “Arundo gigantea” but failed to recognize the hand as that of Fraser. Since

Ward, Thomas Walter Typification Project Il: known Walter types 411

the species is common immediately adjacent to Walter's homesite, and there is no indication Walter saw

or used Fraser's specimen, Fraser/Walter 113-A, [1787] (BM) is here corrected to NEOTYPE for Arundo gigantea

Walt., basionym of Arundinaria gigantea (Walt.) Muhl. Since 113-A consists solely of a stem apex bearing

two leaves and is marginally diagnostic, an epitype would be welcome.

WALTER'S NAME: Arundo tecta Walter (p. 81)

MODERN NAME: Árundinaria gigantea (Walt.) Muhl.

Only one Arundinaria is common in the Carolinas. Walter, under A. tecta, described the first-year stems as

"culmis tectis" stems sheathed") by young leaf-bases, and named the second-year stems A. gigantea (branches

fully developed, making the plant seem larger). There is no specimen labeled as Arundo tecta. Though (as

interpreted here) A. tecta is a synoym of A. gigantea, McClure (1973: 28) has selected McClure 22000, [1952?]

(US), from Anne Arundel County, Maryland, as NEOTYPE for Arundo tecta Walt., basionym of Arundinaria

tecta (Walt.) Muhl.

Wa ter’s NAME: Athanasia graminifolia Walter (p. 200)

MODERN NAME: Marshallia graminifolia (Walt.) Small

Infrequent in eastern SC. Spm. 16-F was identified as Marshallia graminifolia by Channell (1957: 112) and

referred to as the “type” of Athanasia graminifolia. The label (“Athanasia”) appears to be in Walter's hand.

But since materials were available near Walter's home, it is unlikely he used this specimen in preparing his

description. Fraser/Walter 16-F, [1787] (BM) is therefore here corrected to NEOTYPE for Athanasia graminifolia

Walt., basionym of Marshallia graminifolia (Walt.) Small.

WALTER'S NAME: Athanasia obovata Walter (p. 201)

MODERN NAME: Marshallia obovata (Walt.) Beadle € Boynton

Two specimens (16-C, 16-D) were identified as Marshallia obovata var. obovata by Channell (1957: 83, 88-89)

and referred to as the “type” of Athanasia obovata. He further concluded, with the aid of W. T. Stearn, that

they represented the “leafy-stemmed” taxon, which thus becomes var. obovata. That variety occurs only on

the piedmont, west of Walter’s area; the specimen therefore was probably collected by Fraser. [Var. scaposa

Channell occurs only on the coastal plain.] Walter's description may well have been based on plants of

var. scaposa accessible to him in or near Berkeley County. But Channell's assignment of the typical name to

a specimen of the western variant fixes the name in that usage. Channell did not select which of the two

specimens was to be the type and which the isotype. Both are labeled “Athanasia” in Walter's hand. Since

Walter probably based his description on var. scaposa and may not have seen these materials of var. obovata

until after completion of his text, Fraser/Walter 16-C, [1787] (BM) (marked “A. obovata," probably by Gray)

is here corrected to NEOTYPE for Athanasia obovata Walt., basionym of Marshallia obovata (Walt.) Beadle &

Boynton. Fraser/Walter 16-D (BM) (unmarked) then becomes an ISONEOTYPE.

WALTER'S NAME: Athanasia trinervia Walter (p. 201)

MoDERN NAME: Marshallia trinervia (Walt.) Trel. ex Branner & Coville

Unknown in modern SC, very rare in NC (one county) and GA (two counties). There is no specimen. Yet

once found near Walter's home; a specimen from Berkeley County, South Carolina, Cranmore Wallace s.n.,

1841 (CHARL), has been selected by Channell (1957: 68, 72-73), as NEOTYPE for Athanasia trinervia Walt.,

basionym of Marshallia trinervia (Walt.) Trel. ex Branner & Coville.

WALTER'S NAME: Carduus carolinianus Walter (p. 195)

MODERN NAME: Cirsium carolinianum (Walt.) Fern. & Schub.

Cirsium carolinianum is so rare in the Southeast (4 counties in NC, 1 in SC, 3 in GA) that Walter's plant may

not be the species that presently bears his name. But Fernald and Schubert (1948: 229, plate 1115) have

identified spm. 25-C (a near-naked scape with single head, labeled *Carduus" by Walter) as C. carolinianum,

and cited it as "Walters TYPE.” Thus, whatever the possibility Walter was writing of another species more

common near his home, his name is now locked into its present usage. Since a discovery by Fraser in his

412 Journal of the Botanical R h Institute of Texas 1(1)

wider travels would appear the only way Walter may have seen this plant, Fraser/Walter 25-C, [1787] (BM)

is here corrected to LECTOTYPE of Carduus carolinianus Walt., basionym of Cirsium carolinianum (Walt.) Fern.

& Schub. Because of the poor quality of the specimen, an epitype would be welcome.

WALTER'S NAME: Cenchrus carolinianus Walter (p. 79)

MODERN NAME: Cenchrus incertus M.A. Curtis

Common in eastern SC. There is no specimen. Hitchcock (1905: 48), perhaps not realizing there is more

than one species of Cenchrus in the Carolinas, suggested that Walter's description referred to C. tribuloides L.;

Walter's “spinosis laevibus" forecloses that possibility. Reveal (1990) selected a NeotyPe for Cenchrus carolinianus

Walt. of material from Beaufort County, South Carolina, Boufford, Bartholomew & Spongberg 23096, 12 Sept

1982 (BM), currently known as C. incertus M.A. Curtis (1837), thereby temporarily displacing that name.

But, following revision of the I.C.B.N. in 1994, Walter’s name was formally rejected (Brummitt 1995), thus

restoring C. incertus. Though Walter's diagnosis contains elements poorly compatible with C. incertus (Wilbur

1991), that common species would surely have been present in the fields of his Santee River plantation.

WALTER'S NAME: Chrysocoma gigantea Walter (p. 196)

MODERN NAME: Vernonia gigantea (Walt.) Trel. ex Branner & Coville

If this species, rare in SC (two counties). No specimen. Walter's plant is more likely to have been Vernonia

glauca (L.) Willd. or V. noveboracensis (L.) Michx. But Vernonia gigantea is often quite tall, and corresponds to

one feature of Walter's plant: *caule 8 ad 10-pedali.” Urbatsch (1972: 236), lacking any Walter type, preserved

conventional usage by selecting a collection (cited below) of V. gigantea (as customarily defined) as the neo-

type of that name. He, however, did not select a specimen (required by Art. 9.6) of the 7 cited duplicates;

that omission is remedied by selection here of Bozeman & Radford 11593, 3 Nov 1967 (FLAS), from Jasper

County, South Carolina, as NEoTyYPE for Chrysocoma gigantea Walt., basionym of Vernonia gigantea (Walt.) Trel.

& Branner. The duplicates (COLO, IND, NY, OKLA, TENN, WVA) become ISONEOTYPES.

WALTER'S NAME: Collinsonia praecox Walter (p. 65)

MODERN NAME: Collinsonia canadensis L.

Unknown on SC coastal plain, frequent westward; likely a Fraser discovery. No specimen has been identi-

fied. Spm. 96-H is this genus, but lacks flowers, and Peirson et al. (2006: 403, 406) stated it “cannot be

determined with certainty.” They then selected Newberry 1912, 6 Sept 1982 (NCU), from Chester County,

South Carolina, as NEOTYPE for Collinsonia praecox Walt.

WALTER'S NAME: Commelina caroliniana Walter (p. 68)

MODERN NAME: Commelina caroliniana Walt. [= Commelina hasskarlii C.B. Clarke]

This name has generally been disregarded or has been thought unassignable. Faden (1989) has observed that

collections from the southeastern coastal plain assumed to be of Commelina diffusa Burm. are actually of two

entities: that species; and a second one Faden equated with C. hasskarlii C. B. Clarke, an Asiatic species previ-

ously unrecognized in the United States. Faden then identified (by photo) a specimen in the Fraser/Walter

herbarium (35-C) as this second species and concluded its prior name was C. caroliniana Walt. He noted

the label to read “Commelina,” but did not recognize the hand to be that of John Fraser. Then, overlooking

the tenuous connection of Walter to these specimens, Faden designated what is here termed Fraser/Walter

35-C, [1787] (BM) as the “lectotype” of C. caroliniana. Since there is no indication that Walter saw or used

the specimen, it is here corrected to NEOTYPE for Commelina caroliniana Walt.

WaALter’s NAME: Convallaria biflora Walter (p. 122)

MODERN NAME: Polygonatum biflorum (Walt.) Ell.

Infrequent on the SC coastal plain (but incl. Berkeley Co.), common westward. Spm. 35-B was labeled

“Convallaria” by Walter, and is of fair quality. It has been annotated as “TYPE,” perhaps by J. E. Dandy at

direction of Ownbey (1944: 394) who cited this specimen as “type.” Since the plant may well have been

known by Walter and there is no evidence the specimen was seen by him prior to preparing his diagnosis,

Ward, Thomas Walter Typification Project Il: known Walter types 413

Fraser/Walter 35-B, [1787] (BM) is here corrected to NEOTYPE for Convallaria biflora Walt., basionym of Polygo-

natum biflorum (Walt.) Ell.

WALTER'S NAME: Coreopsis gladiata Walter (p. 215)

MODERN NAME: Coreopsis gladiata Walt.

Rare in SC, but known in Berkeley Co. No specimen has been identified. Smith (1976: 195-196) has selected

Godfrey 8238, 15 Sept 1939 (E), from Georgetown County, South Carolina, as NEOTYPE for Coreopsis gladiata

Walt., with duplicate (TENN) as ISONEOTYPE.

WALTER'S NAME: Coreopsis major Walter (p. 214)

MODERN NAME: Coreopsis major Walt.

Frequent in SC, though rare on the coastal plain. Spm. 37-B appears to be this; its label has no writing, which

may explain why it was not noted by Smith (1976). There is no evidence Walter saw or used the specimen.

Smith (1976: 170) has selected Tracy 4360, 10 June 1898 (NY), from Ocean Springs, Jackson County, Missis-

sippi, as NEOTYPE for Coreopsis major Walt., with duplicate (F) as ISONEOTYPE.

WALTER'S NAME: Corypha Palmetto Walter (p. 119)

MopERN NAME: Sabal palmetto (Walt.) Lodd. ex Schult. & Schult.

Infrequent along SC coast. There is no specimen. Zona (1990: 646) selected (“designated”) Curtiss 2677, July

[1894?] (NY), from Jacksonville, Duval County, Florida, as NEotype for Corypha palmetto Walt., basionym

of Sabal palmetto (Walt.) Lodd. ex Schult. & Schult., with duplicates (BH, F, GA, GH, MICH, MO, US) as

ISONEOTYPES.

WALTER'S NAME: Corypha pumila Walter (p. 119)

MODERN NAME: Sabal minor (Jacq.) Pers.

Common in coastal SC. No specimen. Though a type is scarcely needed, Zona (1990: 643) selected Hexamer

& Maier s.n., May 1855 (GH), from St. Andrews, Charleston County, South Carolina, as NEOTYPE for Corypha

pumila Walt. (= Sabal minor (Jacq.) Pers), with duplicate (CM) as ISONEOTYPE.

WALTER'S NAME: Cucubalus polypetalus Walter (p. 141)

MODERN NAME: Silene polypetala (Walt.) Fern. & Schub.

Walter's name was brought forward by Fernald and Schubert (1948: 198) as Silene polypetala (Walt.) Fern.

& Schub., on the evidence of spm. 38-E (a single crumpled flower). The specimen was surely collected by

Fraser along the Flint River, west-central Georgia, its closest location. It bears, in Walter's hand, the words

“Cucubalus polypetalus, and a 3-digit number assigned by Fraser. But Walters description of Cucubalus

polypetalus does not fit the plant. Walter’s words, “Cal. inflatus" “calyx inflated"), “petala fauce nuda” (“petals

smooth at throat”), and "floribus polypetalis” “flowers with many petals”), suggest one of the several Silene

species with expanded (“inflated”) calyces and non-auricled (=crowned), deeply bi-lobed petals. Walter may

have had an early contact with Silene cucubalus Wibel (1799), an introduced species now well established

in the mountains of NC. Lychnis alba Mill., another introduced species with an inflated calyx, is less likely

since its petals are auricled, unlike Cucubalus.

Fernald and Schubert noted the single flower (plate 1105) as “Walter’s type" of Cucubalus polypetalus.

That action, other than by conservation, is irrevocable; the error in understanding which of Walter's names

applied to the fragmentary specimen does not invalidate their action. However, since Walter's description

indicates he had access to more complete materials that were perhaps part of the same collection, Fraser/Walter

38-E, [1787] (BM) is here corrected to Lectotyre of Cucubalus polypetalus Walt., basionym of Silene polypetala

(Walt.) Fern. & Schub.

WALTER'S NAME: Echites difformis Walter (p. 98)

MODERN NAME: Trachelospermum difforme (Walt.) Gray

Common throughout. Krings (2003) identified *Walter 215" (spm. 41-C) as this species, and cited it as

“holotype” of Echites difformis. The label bears “Echites” by Walter and *Difformis" by Fraser. Since the plant

414 Journal of the Botanical R h Institute of Texas 1(1)

is common in his immediate area, Walter would have had no need for this specimen and there is no indica-

tion he used it in forming his diagnosis. Thus Fraser/Walter 41-C, [1787] (BM) is here corrected to NEOTYPE

for Echites difformis Walt., basionym of Trachelospermum difforme (Walt.) Gray.

WALTER'S NAME: Eupatorium pilosum Walter (p. 199)

MODERN NAME: Eupatorium pilosum Walt.

Common in eastern SC. Spm. 45-A was labeled “Eupatorium” by Walter; the label also bears a 3-digit num-

ber assigned by Fraser. The specimen was identified as Eupatorium pilosum by Fernald and Schubert (1948:

225-226, plate 1114), then referred to as "Walters TYPE.” The probability is high that Walter had ready

access to living materials growing near his home and did not see this specimen until shown it by Fraser.

Fernald and Schubert's typification must be given recognition, but Fraser/Walter 45-A, [1787] (BM) is here

corrected to NEOTYPE for Eupatorium pilosum Walt.

WALTER'S NAME: Gentiana Catesbaei Walter (p. 109)

MODERN NAME: Gentiana catesbaei Walt.

Infrequent in eastern SC. Spm. 50-A was labeled as “Gentiana” and spm. 50-B as “Gentiana saponaria,” both

in Walter's hand. Fernald (1939: 555-556) referred to 50-A in discussion of G. catesbaei, but his text left

open the possibility he considered it G. saponaria. Later, Fernald (1947, plate 1078) identified spms. 50-A

and 50-B as the "type" of Gentiana catesbaei. [He (1947: 176) erroneously identified the label of 50-A as hav-

ing been written by “James Britten (apparently).”] Rembert (1980) again identified spm. 50-A as G. catesbaei,

though he did not designated it as type. Of the two, spm. 50-A is of better quality; it is reasonably complete,

with leaves and flowers. Though Fernald included both specimens within his *type" of Gentiana catesbaei

Walt., his designation of Fraser/Walter 50-A, [1787] (BM) is here corrected to Lectotype. Fraser/Walter 50-B

(BM) then becomes an ISOLECTOTYPE.

WALTER'S NAME: Gratiola acuminata Walter (p. 61)

MODERN NAME: Mecardonia acuminata (Walt.) Small [2 Bacopa acuminata (Walt.) Robinson]

Frequent throughout. *[Walter's] description [is] evidently of plant here considered" (Pennell 1920: 236).

Spm. 53-B was numbered “668” by Fraser and labeled “Gratiola” by Walter. It was cited (as “Walter 668")

by Pennell (1935: 66) as “Type.” Since Walter would surely have been familiar with the plant near his home,

Fraser/Walter 53-B, [1787] (BM) is here corrected to NEOTYPE for Gratiola acuminata Walt., basionym of Me-

cardonia acuminata (Walt.) Small.

WALTER'S NAME: Gratiola ramosa Walter (p. 61)

MODERN NAME: Gratiola ramosa Walt.

Common on SC coastal plain. “Descriptive of this plant" (Pennell 1920: 240). Pennell (1935: 79) cited an

unspecified specimen as “Type,” noting “it shows well the characters of the species now considered, the

calyx lacking subtending bractlets.” He does not mention an accompanying number. Since spm. 53-D is

the only specimen on the page without such a number, and since 53-D well matches modern specimens,

it is accepted as Pennell's type. It was labeled “Gratiola” by Fraser. Since Walter would not have had use for

the specimen in forming his description, Fraser/Walter 53-D, [1787] (BM) is here corrected to NEOTYPE for

Gratiola ramosa Walt.

WALTER'S NAME: Hedysarum grandiflorum Walter (p. 185)

MODERN NAME: Desmodium cuspidatum (Muhl. ex Willd.) Loud.

Infrequent throughout SC. Not Hedysarum grandiflorum Pallas (1773). Fernald and Schubert (1948: 203) iden-

tified spm. 55-C as Hedysarum grandiflorum and referred to it as “Walter’s TYPE.” The specimen was labeled

“Hedysarum Flore magnus” by Walter, who seemed not to recall the name he had already given it. Walter's

name is a later homonym and thus illegitimate. But, having been typified by Fernald and Schubert, their

designation of Fraser/Walter 55-C [1787] (BM) is here corrected to NEOTYPE of Hedysarum grandiflorum Walt.

Ward, Thomas Walter Typification Project Il: known Walter types 415

WALTER'S NAME: Helenium aestivale Walter (p. 210)

MODERN NAME: Gaillardia aestivalis (Walt.) H. Rock

Absent from SC coastal plain, frequent on pied mont; possibly a Fraser discovery. Rock (1956) has designated

spm. 56-C as lectotype of Helenium aestivale (= Gaillardia aestivalis). The specimen is labeled *Helenium" in

Walter's hand. Rock's designation of Fraser/Walter 56-C, [1787] (BM) as LECTOTYPE is appropriate. A second

specimen (spm. 56-B), labeled *Helenium" by Fraser, was designated by Rock as “syntype.” With the tenu-

ous assumption that it is part of the same collection, Fraser/Walter 56-B [1787] (BM) is here corrected to

ISOLECTOTYPE of Helenium aestivale Walt.

WALTER'S NAME: Helenium serotinum Walter (p. 210)

MODERN NAME: Gaillardia pulchella Foug. [= Gaillardia serotina (Walt.) H. Rock]

Infrequent on SC coastal plain. Rock (1956) identified a specimen (spm. 56-D) as a Gaillardia, and designated it

as the lectotype of Helenium serotinum (and made the combination Gaillardia serotina). The typification was not

critical since Fougeroux’ name (1787) is prior to Walter’s. But Walter's hand on the label (Helenium”) makes

plausible Rock's designation of Fraser/Walter 56-D, [1787] (BM) as Lectotype of Helenium serotinum Walt.

WALTER'S NAME: Helenium vernale Walter (p. 210)

MODERN NAME: Helenium vernale Walt.

Infrequent in eastern SC (incl. Berkeley Co.). Rock (1956) has designated spm. 56-A as lectotype of Helenium

vernale. The specimen bears the hand of Fraser, who identified it only as “Helenium.” Since the species is

known from Berkeley County, it is likely that Walter prepared his description independently of this speci-

men. However, the specimen having been given type status, Rock's designation is retained, but Fraser/Walter

56-A, [1787] (BM) is here corrected to NEOTYPE for Helenium vernale Walt.

WALTER'S NAME: Hydrangea radiata Walter (p. 251)

MODERN NAME: Hydrangea arborescens L. ssp. radiata (Walt.) McClintock

Not known on the SC coastal plain, but frequent inland, thus probably a Fraser discovery. The word “Hy-

drangea” in Walter's hand on spm. 59-B indicates he saw the specimen. Fraser added the epithet “Radiata,”

written after he had access to Walter's manuscript or book. The specimen is of good quality. It was annotated

as the “Type of Hydrangea radiata” by E. McClintock in 1954; the designation was published as “Type col-

lection: Walter s.n. (BM)” for H. arborescens ssp. radiata (McClintock 1957: 172). McClintock’s designation is

retained, but Fraser/Walter 59-B [1787] (BM) is here corrected to Lectotype of Hydrangea radiata Walt.

WALTER' NAME: Kalmia hirsuta Walter (p. 138)

MopERN NAME: Kalmia hirsuta Walt.

Rare in SC (5 counties, all just south of Walter's Berkeley Co.). Probably a discovery of Fraser's; a “new vil-

lose Kalmia" was noted among other Fraser plants (letter from Walter to Forsyth-Rembert 1980: 17). Spm.

62b-C, a nearly bare twig, was labeled “Kalmia Hirsuta Nova" by Fraser. Southall and Hardin (1974) referred

to a specimen on page 62 as the “type.” Since this fragment may have been part of better materials brought

by Fraser to Walter and used by him in preparation of his diagnosis, Fraser/Walter 62b-C, [1787] (BM) is here

corrected to LECTOTYPE of Kalmia hirsuta Walt. The specimen, however, is unidentifiable without the label

and serves no useful purpose as a type. An epitype would be welcome.

WALTER'S NAME: Lobelia glandulosa Walter (p. 218)

MODERN NAME: Lobelia glandulosa Walt.

Frequent on SC coastal plain. There can be no confidence that Walter had Lobelia glandulosa, rather than L.

elongata Small which is perhaps more common. But McVaugh (1936: 288) considered a “few fragments” (GH),

taken in 1839 from the Walter herbarium, to be the “type” of L. glandulosa. [These materials have now been

returned to the Walter herbarium (correspondence attached to folio: C. A. Weatherby, 28 Jan 1936; M.L.

Fernald, 20 Jan 1936) and public apology made (Fernald 1937).] Since the specimen (65-G) bears only the

hand of Fraser, and there is no indication that Walter made use of the collection, Fraser/Walter 65-G, [1787]

(BM) is here corrected to NEOTYPE for Lobelia glandulosa Walt.

416 Journal of the Botanical R h Institute of Texas 1(1)

WALTER'S NAME: Ludwigia apetala Walter (p. 89)

MODERN NAME: Ludwigia palustris (L.) Ell.

Common throughout. Spm. 66-A was labeled “Ludwigia ?” by Walter. It was identified (from microfiche) by

Peng et al. (2005: 336) as Ludwigia palustris, and was cited as the “holotype” of L. apetala Walt. Since the

species would have been well-known to Walter and there is no indication that he made use of this speci-

men, Peng et al/s citation of Fraser/Walter 66-A, [1787] (BM) is here corrected to NEOTYPE for Ludwigia apetala

Walt. © Ludwigia palustris (L.) EIL).

WALTER'S NAME: Ludwigia decurrens Walter (p. 89)

MODERN NAME: Ludwigia decurrens Walt.

Common throughout. Spm. 66-C was labeled *Ludwigia decurrens" by Walter. An unspecified specimen

(Herb. Walter,” identified from photo) was cited as “type” of Ludwigia decurrens Walt. by Ramamoorthy

and Zardini (1987: 88). Since the species was available to Walter near his home and the specimen would

not have been needed to form his description, Fraser/Walter 66-C, [1787] (BM) is here corrected to NEOTYPE

for Ludwigia decurrens Walt.

WALTER'S NAME: Ludwigia pilosa Walter (p. 89)

MODERN NAME: Ludwigia pilosa Walt.

Common on SC coastal plain. Spm. 66-D was labeled “Ludwigia pilosa” by Walter, and appears to be that

species; the specimen also bears *658" in Fraser's hand. A specimen bearing number 658 was cited by Peng

(1989: 282) as *holotype" of Ludwigia pilosa. Since the species was available to Walter elsewhere and there

is no evidence that spm. 66-D was used by him in preparation of his text, Fraser/Walter 66-D, [1787] (BM)

is here corrected to NEOTYPE of Ludwigia pilosa Walt. Peng et al. (2005: 345) later cited the “lower left-hand

specimen" on page 66 (again, 66-D) as the *holotype" of Ludwigia arcuata Walt., clearly in gross error!

WALTER'S NAME: Ludwigia linearis Walter (p. 89)

MODERN NAME: Ludwigia linearis Walt.

Common in eastern SC. Spm. 66-E is this; it was labeled “Ludwigia” by Walter. A specimen (not specifically

designated; “Walter Herbarium, p. 66,” identified from photo) was cited by Peng (1989: 244) as “holotype”

of L. linearis. Since Walter would not have needed this specimen and there is no evidence he saw it prior to

preparing his description, Fraser/Walter 66-E, [1787] (BM) is here corrected to NEOTYPE for Ludwigia linearis

Walt.

WALTER'S NAME: Nymphaea pentapetala Walter (p. 155)

MODERN NAME: Perhaps Nelumbo lutea (Willd.) Pers.

If this species, rare in SC (4 counties). Spm. 75-F, a single petal, was labeled by Walter as “The Great Nym-

phaea.” Ward (1977) noted that, though poorly described (“corolla...pentapetala alba”), Walter's plant was

clearly a Nelumbo; he argued that it may have been either Nelumbo lutea or N. nucifera and must remain of

uncertain application. Wiersema and Reveal (1991) “with great trepidation” interpreted N. pentapetala to be

Nelumbo lutea, a later name (1788 vs. 1799); they then selected Hunt & Martin 2056, 5 June 1943 (CLEM),

from Charleston County, South Carolina, as Walter’s Nrorvrr, and simultaneously proposed rejection of his

name. With unanimous support from the Committee for Spermatophyta, Nymphaea pentapetala Walter was

then nomenclaturally rejected (Brummitt 1995), preserving Nelumbo lutea.

WALTER'S NAME: Nymphaea reniformis Walter (p. 155)

MODERN NAME: Perhaps Nymphaea odorata Ait.

There is no specimen in the herbarium. Walters name has been consistently disregarded. Ward (1977)

believed Walter's description to have been based on mixed material with elements of both Nymphaea and

Nelumbo. Wiersema and Reveal (1991) selected Godfrey & Tryon 471, 12 Jul 1939 (DUKE), a specimen of

Nymphaea odorata Ait. var. gigantea Tricker, from Berkeley County, South Carolina, as NEOTYPE, and simulta-

neously proposed rejection of Walter's name. Because of its unreconcilable ambiguity, Nymphaea reniformis

Walter was then nomenclaturally rejected (Brummitt 1995). Familiar epithets of Nymphaea and Nelumbo

Ward, Thomas Walter Typification Project Il: known Walter types 417

thus remain unchallenged. Being listed last in the genus suggests this to be a late addition from Fraser, a

frequent practice of Walter's.

WALTER'S NAME: Nymphaea sagittifolia Walter (p. 155)

MODERN NAME: Nuphar luteum (L.) Sibth. & Sm. ssp. sagittifolium (Walt.) Beal

Occasional on the lower SC coastal plain. There is no specimen in the herbarium. Walter’s description

was identified by Ward (1977). Beal (1956: 335) selected McCarthy s.n., July 1885 (NY), from eastern North

Carolina, as NEOTYPE for Nymphaea sagittifolia Walt.

WALTER'S NAME: Oenanthe filiformis Walter (p. 113)

MODERN NAME: Oxypolis filiformis (Walt.) Britt.

Frequent on the SC coastal plain. No specimen has been identified. Tucker et al. (1983: 300) have selected

Porcher s.n., 17 Sept 1981 (BM), from Berkeley County, South Carolina, as NEotyPE for Oenanthe filiformis Walt.,

basionym of Oxypolis filiformis (Walt.) Britt., with duplicates (CITA, DOV) as ISONEOTYPES.

WALTER'S NAME: Ophrys barbata Walter (p. 221)

MODERN NAME: Perhaps Calopogon barbatus (Walt.) Ames, more likely Calopogon pulchellus (Salisb.)

R. Br.

Calopogon pulchellus is frequent in eastern SC. Walter—and Fraser—would surely have known it, also perhaps

the much rarer C. pallidus Chapm. and C. barbatus. Walter’s description does not permit judgment as to which

of these species he meant by his Ophrys barbata—the lip is bearded in all. Spm. 77-E is clearly a Calopogon.

Its original label bears only the single word “Ophrys” in Fraser’s hand; it was later annotated as “Ophrys

barbata / Calopogon parviflorus” by A. A. Eaton. Goldman (1998) stated the specimen to be *Walter's type” of

C. barbatus, but he then identified it as C. multiflorus Lindl. [He was surely in error. Calopogon multiflorus is

nearly absent from the Carolinas and Georgia (a single station in NC, with unverified reports from SC and

GA), and morphology of the specimen is either of C. barbatus (viz. Eaton id.) or is inconclusive.] Goldman

then selected Orzell & Bridges 16163, 21 Mar 1991 (TEX), from Baker County, Florida, as NEOTYPE for Ophrys

barbata Walt. Goldman next proposed conservation of Ophrys barbata with this new type; his proposal was

promptly accepted by the Committee for Spermatophyta.

Goldman may not have recognized the only writing on the original label was by Fraser, not Walter,

nor that there is no indication the specimen was seen or used by Walter; these details were not brought

before the Committee. Goldman's action, however, is of value in that by selecting a neotype that continues

the historic interpretation, the name Calopogon barbatus retains its classic meaning.

WALTER'S NAME: Origanum flexuosum Walter (p. 165)

MODERN NAME: Pycnanthemum flexuosum (Walt.) BSP.

Common in eastern SC. Spm. 79-C was labeled “Origanum” by Walter. It was identified (from photo) by

Fernald and Schubert (1948: 220-222, plate 1112) as Pycnanthemum flexuosum and designated as *Walter's

TYPE.” Since the plant is common near his home it is unlikely that Walter used this specimen in preparing

his description. Even so, having been cited as “type” by Fernald and Schubert, Fraser/Walter 79-C, [1787]

(BM) has attained formal status. It is here corrected to NEOTYPE for Origanum flexuosum Walt., basionym of

Pycnanthemum flexuosum ( Walt.) BSP.

WALTER'S NAME: Panicum hirtellum Walter (p. 72)

MODERN NAME: Echinochloa walteri (Pursh) Heller

Not Panicum hirtellum L. [2 Oplismenus hirtellus (L.) Beauv.], a tropical species. Hitchcock (1905: 35) found

three specimens in the Walter herbarium that had been labeled *Panicum hirtellum." The first of these (115-A)

Hitchcock called “the long-awned form of P. crus-galli L.” [= Echinochloa crusgalli (L.) Beauv.], and the second

(115-B) *a densely flowered long-awned form [that is the] P. hispidulum of Muhlenberg, who cites P. hirtellum

Walt.” Spms. 115-A and 115-B are both labeled in Walter's hand. [The third, 115-C, is Panicum virgatum L.]

Hitchcock favored placing Walter's name in synonymy under P. crus-galli (now Echinochloa crusgalli). Pursh

(1814), however, had correctly interpreted Walter's plant as new, and named it Panicum walteri [2 Echinochloa

walteri (Pursh) Heller.] Panicum hirtellum Walter thus remains the basis for the modern E. walteri.

418 Journal of the Botanical R h Institute of Texas 1(1)

Echinochloa crusgalli is abundant throughout the Carolinas, but is *believed to be adventive from Europe

or Asia" (Gould et al. 1972), while the similar E. walteri is common in coastal areas of the Carolinas where it

is native. Hitchcock (1920: 138) referred to a specimen (115-B) as what *may be taken as the type." Though

Walter saw this specimen (as demonstrated by his hand on the label), there is no certain evidence he used

it in preparation of his text. Hitchcock's action must be acknowledged, but the status of Fraser/Walter 115-

B, [1787] (BM) is here corrected to NEOTYPE for Panicum hirtellum Walt. (2 Panicum walteri Pursh; Echinochloa

walteri (Pursh) Heller).

WALTER'S NAME: Phalaris caroliniana Walter (p. 74)

MODERN NAME: Phalaris caroliniana Walt.

Frequent on SC coastal plain. No specimen was found in the herbarium by Hitchcock (1905: 40), nor An-

derson (1961). Anderson found the description perplexing but concluded there was *no other species of the

Carolina grass flora that would fit the description" better than P. caroliniana. He then selected Duncan 9468,

[4 May 1949] (US), from McCormick County, South Carolina, as Neotype for Phalaris caroliniana Walt., with

duplicate (GA) as ISONEOTYPE.

Wa ter’s NAME: Phyllanthus caroliniensis Walter (p. 228)

MODERN NAME: Phyllanthus caroliniensis Walt.

Frequent throughout. Spm. 83-E was labeled *Phyllanthus affinis" by Walter. Webster (1970: 60) cited a

specimen on page “83” as “holotype” of P. caroliniensis ssp. caroliniensis. Since Walter neither recognized this

specimen as his new species, nor would have been in need of it for his diagnosis, Fraser/Walter 83-E, [1787]

(BM) is here corrected to NEOTYPE for Phyllanthus caroliniensis Walt.

WALTER'S NAME: Pinguicula caerulea Walter (p. 63)

MODERN NAME: Pinguicula caerulea Walt.

Frequent on SC coastal plain. Spm. 104-D was identified as Pinguicula caerulea by Fernald and Schubert

(1948: 224), then referred to as its “TYPE.” The label is misplaced; it reads “Utricularia gibba” in Walter's

hand. [Fernald and Schubert erroneously described it (plate 1113) as “mislabeled by Fraser.”] The label that

should have been with spm. 104-D is to be found with spm. 83-F (a plant of Oxalis violacea). Since the label

of “Pinguicula caerulea” that should have accompanied spm. 104-D truly was in Fraser’s hand, there is no

indication that Walter saw or used the specimen. Thus Fraser/Walter 104-D, [1787] (BM) is here corrected

to NEOTYPE for Pinguicula caerulea Walt.

WALTER'S NAME: Pinguicula lutea Walter (p. 63)

MODERN NAME: Pinguicula lutea Walt.

Infrequent on SC coastal plain, but known in Berkeley County. Spm. 83-G was identified as Pinguicula lutea

by Fernald and Schubert (1948: 224), then referred to as its “TYPE” (plate 1113). Since the label (“Pinguicula

lutea”) is in Fraser's hand and there is no indication that Walter saw or used the material, Fraser/Walter 83-

G, [1787] (BM) is here corrected to NEOTYPE for Pinguicula lutea Walt.

WALTER'S NAME: Potamogeton pinnatum Walter (p. 90)

MODERN NAME: Myriophyllum pinnatum (Walt.) BSP.

Infrequent on SC coastal plain. Aiken (1981) reported John Lewis “examined the Walter specimen [86-B]

for me.” This specimen was labeled “Potamogeton monoicum” in Walter's cramped hand. Aiken did not convey

the specimen was already annotated “Potamogeton pinnatum Walt., Fl. Carol. p. 90, M.L. Flernald].” Though

Aiken called this specimen the “holotype,” it is much more likely that Walter prepared his description from

fresh material found near his home. Appropriately, Fraser/ Walter 86-B, [1787] (BM) is here corrected to NEOTYPE

for Potamogeton pinnatum Walt., basionym of Myriophyllum pinnatum (Walt.) BSP.

WALTER'S NAME: Prasium purpureum Walter (p. 166)

MODERN NAME: Physostegia purpurea (Walt.) Blake

Common on SC coastal plain. Spm. 87-A was labeled *Prasium" by Fraser. It was identified as (a synonym of)

Ward, Thomas Walter Typification Project Il: known Walter types 419

Physostegia purpurea by Blake (1915: 134), and designated as lectotype by Cantino (1981, 1982). In absence

of evidence the specimen was seen or used by Walter, Fraser/Walter 87-A, [1787] (BM) is here corrected to

NEOTYPE for Prasium purpureum Walt., basionym of Physostegia purpurea (Walt.) Blake.

WALTER'S NAME: Quercus sinuata Walter (p. 235)

MODERN NAME: Quercus sinuata Walt.

Rare, perhaps found only along the Santee River. Believed to be a hybrid of Quercus falcata and Q. phellos. There

is no specimen in the herbarium. A NEOTYPE has been selected for Quercus sinuata Walt. (Ward, in press).

WALTER'S NAME: Rhexia Alifanus Walter (p. 130)

MopERN NAME: Rhexia alifanus Walt.

Common in eastern SC. No specimen. James (1956: 218) selected James 675, 17 June 1955 (GH), from Pineville,

Berkeley County, South Carolina, as NEOTYPE for Rhexia alifanus Walt. Noted by Kral and Bostick (1969).

WALTER'S NAME: Rhexia lutea Walter (p. 130)

MODERN NAME: Rhexia lutea Walt.

Frequent in eastern SC. No specimen. James (1956: 216) selected James 678, 17 June 1955 (GH), from St.

Stephen, Berkeley County, South Carolina, as Neotyre for Rhexis lutea Walt. Noted by Kral and Bostick

(1969).

WALTER'S NAME: Salix alpina Walter (p. 243)

MODERN NAME: Salix humilis Marsh.

Nearly absent from SC, frequent in mountains and piedmont of NC. Spm. 93-C was labeled by Fraser as

“Salix Minor of Fraser” and was identified by Blake (1915: 136) as Salix alpina Walter; Blake noted it “may

be considered the type.” Salix alpina is a later synonym (Marshall 1785 vs. Walter 1788), and designation

of its type is not needed for stability of the current name. But Blake’s reference to the specimen requires

acknowledgment of his choice. Though the specimen bears only Fraser’s hand, absence of the species from

Walter’s immediate area indicates it may have been part of materials brought to him by Fraser, which justi-

fies correction of Fraser/Walter 93-C, [1787] (BM) to LecToTYPE of Salix alpina Walt.

WALTER'S NAME: Sarracenia rubra Walter (p. 152)

MODERN NAME: Sarracenia rubra Walt.

Infrequent throughout SC. A photo of spm. 95-B is given by Rembert (1980: 24). The specimen was designated

as “type” by McDaniel (1971). However, since it was collected and labeled (as “Sarracenia Rubra”) by Fraser,

and there is no evidence it was seen or used by Walter, Fraser/Walter 95-B, [1787] (BM) is here corrected to

NEOTYPE for Sarracenia rubra Walt.

WALTER'S NAME: Sarracenia minor Walter (p. 153)

MODERN NAME: Sarracenia minor Walt.

Common in eastern SC. A photo of spm. 95-C is given by Rembert (1980: 24). The specimen was designated

as “type” by McDaniel (1971). However Walter labeled the specimen “Sarracenia lutea,” either a lapsus calami

for S. flava, or he failed to recognize it as the plant he had described as S. minor. Thus Fraser/Walter 95-C,

[1787] (BM) is here corrected to NEOTYPE for Sarracenia minor Walt.

WALTER'S NAME: Silene Catesbaei Walter (p. 141)

MODERN NAME: Silene catesbaei Walt. [= Silene polypetala (Walt.) Fern. & Schub.]

Very rare Q counties in FL, 4 counties in GA, unknown in SC and NC). No specimen in the herbarium bears

this name. Walter's description of Silene catesbaei is a near-exact match for the plant colloquially known as

Fringed Catchfly. A Neotyre has been selected for Silene catesbaei Walt. (Ward, in press).

WALTER'S NAME: Sophora villosa Walter (p. 134)

MODERN NAME: Baptisia cinerea (Raf.) Fern. & Schub.

Infrequent in eastern SC. Spm. 100-E, labeled by Walter as “Sophora” and by Fraser as “Villosa” and bearing

420 Journal of the Botanical R h Institute of Texas 1(1)

Fraser's number *360", was identified by Fernald and Schubert (1948: 200-201) as *the Walter type" of

Sophora villosa (2 Thermopsis villosa (Walt.) Fern. & Schub.). They also depicted the specimen (plate 1106)

with the label, *TYPE of Sophora villosa Walt." The specimen is indeed of a Thermopsis, but Walter's descrip-

tion is of a Baptisia, surely B. cinerea. Thermopsis villosa is known only in the Carolina mountains, where

available only to Fraser (undoubtedly the source of spm. 100-E). Walter described five legumes as Sophora,

believed by him to be congeneric. Four of his Sophora are clearly identifiable species of Baptisia that are

frequent-to-common in eastern SC. “Sophora villosa”, third in his tabulation, corresponds to Baptisia cinerea

(Raf) Fern & Schub. It is most unlikely that Walter would have inserted the morphologically different,

unfamiliar Thermopsis between the four similar Baptisia species, and then omitted treatment of a fifth spe-

cies also found in eastern SC.

But typification cannot be reversed (short of conservation), and the plant fragment, an inflorescence

branch bearing two flowers, is by the action of Fernald and Schubert (1948) the type of Sophora villosa Walt.,

the basionym of Thermopsis villosa (Walt.) Fern. & Schub. Since there is little probability that Walter saw

or used the specimen, Fernald and Schubert's unfortunate citation of Fraser/Walter 100-E, [1787] (BM) can

only here be corrected to NEOTYPE for Sophora villosa Walt.

WALTER'S NAME: Stellaria uniflora Walter (p. 141)

MODERN NAME: Arenaria uniflora (Walt.) Muhl.

Rare, on granite outcrops of the lower piedmont. Surely a discovery of Fraser. Spm. 100-K was labeled *No

Name" by Fraser. The specimen was identified (from photo) by Fernald and Schubert (1948: 195-197) as

Arenaria uniflora, then equated with Walter's Stellaria uniflora, and designated its “TYPE” (plate 1103). Since it

is most likely Walter based his description on materials brought to him by Fraser, Fraser/Walter 100-K, [1787]

(BM) is here corrected to Lectotype of Stellaria uniflora Walt., basionym of Arenaria uniflora (Walt.) Muhl.

WALTER' NAME: Utricularia inflata Walter (p. 64)

MODERN NAME: Utricularia inflata Walt.

Frequent in eastern SC. Spm. 104-E was labeled “Utricularia minor” by Walter (but is not U. minor L., of

Europe). Though (from photo) its identity is unclear, the specimen was designated by Taylor (1989: 662)

as the "lectotype" of Utricularia inflata Walt. But since it would surely have been familiar to Walter and

described by him from fresh material, Fraser/Walter 104-E, [1787] (BM) is here corrected to NEOTYPE for

Utricularia inflata Walt.

WALTER'S NAME: Viscum album ? (p. 241)

MODERN NAME: Phoradendron serotinum (Raf.) M.C. Johnst. [= Phoradendron “leucarpum” (Raf.) Reveal €

M.C. Johnst.]

Frequent throughout. Not Viscum album L., a European species. Fraser/Walter 110-F, [1787] (BM) is a scarcely

identifiable scrap, labeled simply “Viscum” in Walter's hand. The specimen was designated (“selected”) by

Reveal and Johnston (1989) as Lectotype for “V. ? album” of Walter. Then, having attached a specimen to

Walter’s description (and name?), they used the specimen as the basis for Viscum leucarpum Rafinesque (a

misspelling of “leucocarpum”!), itself the basis for Phoradendron leucarpum (Raf.) Reveal & Johnston. Their

lectotypification cannot be discarded. But since Walter had merely made a tentative error of identification and

was not publishing a new name, a simpler action would have been to recognize the irrelevance of Walter’s

name, and select a better-quality neotype for Viscum leu[co]carpum Raf.

WALTER'S NAME: Xyris caroliniana Walter (p. 69)

MODERN NAME: Xyris caroliniana Walt. [= Xyris flexuosa Muhl. ex Ell.]

No specimen. In search for the type of Walter’s Xyris caroliniana, Kral (1966: 236) located a John Fraser

specimen (at P) that corresponded to X. flexuosa. That inconspicuous dry-soil species surely is not what

Walter knew in the rice fields of his Santee River plantation, nor could it later (DBW obs., July 1990) be found

anywhere in the vicinity of Walter’s homesite. Its linear, twisted leaves conform poorly with Walter’s “fol.

gladiatis.” But, however inappropriate, the Fraser specimen having been designated by Kral as the type of X.

Ward, Thomas Walter Typification Project Il: known Walter types 421

caroliniana, Walter's name is best left assigned to the species otherwise known as X. flexuosa. Since Fraser's

collections were made quite independently of Walter, Fraser s.n., [1787] (P), from another location and by a

different collector, is here appropriately corrected to NEOTYPE for Xyris caroliniana Walt.

ACKNOWLEDGMENTS

Lectotypes and neotypes are too often published in unexpected places. There can be no assurance that

none have been overlooked in the present compilation. But for their help in pursuit of the elusive goal of

completeness, I am grateful to Bruce F. Hansen (USF), Peter C. Hoch (MO), Guy L. Nesom (BRIT), Kent

D. Perkins (FLAS), James S. Pringle (HAM), David H. Rembert (USCH), and Robert L. Wilbur (DUKE). I

have received encouragement and inspiration from Charles E. Jarvis (BM), with his long-ongoing project to

record types of all Linnaean names. I also thank Alexander Krings (NCSC) and Robert L. Wilbur (DUKE)

for their careful and most helpful review of the manuscript.

REFERENCES

AIKEN, S.G. 1981. A conspectus of Myriophyllum (Haloragaceae) in North America. Brittonia 33:57—69.

ANDERSON, D.E. 1961. Taxonomy and distribution of the genus Phalaris. lowa State J. Sci. 36:1-96.

Beal, E.C. 1956. Taxonomic revision of the genus Nuphar Sm. of North America and Europe. Elisha Mitchell Sci.

Soc. 72:317-346.

Blake, S.F. 1915. Some neglected names in Walter's Flora Caroliniana. Rhodora 17:129-137.

Brummit, R.K. 1995. Report of the Committee for Spermatophyta. Taxon 44:608.

Cantino, PD. 1981. The Physostegia purpurea-leptophylla-denticulata problem: taxonomic and nomenclatural

clarification. Rhodora 83:581—593.

Cantino, PD. 1982. A monograph of the genus Physostegia (Labiatae). Contr. Gray Herb. 211:1-105.

CHANNELL, R.B. 1957. A revisional study of the genus Marshallia (Compositae). Contr. Gray Herb. 181:41-132.

Curtis M.A. 1837. Boston Soc. Nat. Hist. J. 1:135.

Ewan, J. 1969. Historical problems for the working taxonomist. Taxon 18:194-203.

FADEN, R.B. 1989. Commelina caroliniana (Commelinaceae): a misunderstood species in the United States is an

old introduction from Asia. Taxon 38:43-53.

FERNALD, M.L. 1937. The identity of Lobelia glandulosa Walt. Rhodora 39:497.

FERNALD, M.L. 1939. New species, varieties and transfers. Rhodora 41:423-459.

FERNALD, M.L. 1947. Additions to and subtractions from the flora of Virginia. Rhodora 49:1 75-176.

FERNALD, M.L. and B.G. ScHusert. 1948. Studies of American types in British herbaria. Part IV: some species of Thomas

Walter. Rhodora 50:190-208, 217-229.

FRASER, J. 1789. A short history of the Agrostis Cornucopiae: or, the new American grass. Chelsea, England.

Gaiser, L.O. 1950. Chromosome numbers in Liatris, |l. Graminifoliae and Pauciflorae. Amer. J. Bot. 37:414-423.

GANDHI, K.N. and B.E. Dutton. 1993. Palisot de Beauvois, the correct combining author of Erianthus giganteus

(Poaceae). Taxon 42:855-856.

GMELIN, J.F. 1792. Systema Naturae. Leipzig.

GOLDMAN, D.H. 1998. Proposal to conserve the name Ophrys barbata (Orchidaceae) with a conserved type. Taxon

47:161-162.

GouLo, F.W., M.A. Au, and D.E. FairsrotHers. 1972. A revision of Echinochloa in the United States. Amer. Midl. Nat.

87:36-59.

GREUTER, W., J. MCNEILL, F.R. Barrie, H.M. Bunper, V. DEMOULIN, T.S. FILGUEIRAS, D.H. NicoLson, P.C. Siva, J.E. Scoc, P. TREHANE and

N.J. TURLAND, 2000. International Code of Botanical Nomenclature. Saint Louis Code. Koeltz, Germany.

HARPER, R.M. 1911. Early spring aspects of the coastal plain vegetation of South Carolina, Georgia, and northeastern

Florida. Bull. Torrey Bot. Club 38:223-234.

HitcHcock, A.S. 1905. The identification of Walter's grasses. Missouri Bot. Gard. Ann. Rept. 16:31—56.

HitcHcock, A.S. 1920. The North American species of Echinochloa. Contr. U.S. Nat. Herb. 22:133-153.

422 Journal of the Botanical R h Institute of Texas 1(1)

James, C.W. 1956. A revision of Rhexia (Melastomataceae). Brittonia 8:201-230.

Jupp, W.S. 1981. A monograph of Lyonia (Ericaceae). J. Arnold Arbor. 62:63-128, 129-209, 315-436.

KraL R. 1966. Xyris (Xyridaceae) of the continental United States and Canada. Sida 2:177-260.

KraL R. and PE. Bostick. 1969. The genus Rhexia (Melastomataceae). Sida 3:387-440.

Krinas, A. 2003. Typification and nomenclatural history of Trachelospermum difforme (Apocynaceae). Sida

20:1641-1644.

LEEUWENBERG, A.J.M. 1974. The Loganiaceae of Africa, XII. A revision of Mitreola L. Meded. Landbouwhogeschool

74(23):1-28.

Lewis, W.H. 1966. Typification of Hedyotis procumbens (Rubiaceae) and a new variety from southeastern United

States. Ann. Missouri Bot. Gard. 53:377-378.

McCuntock, E. 1957. A monograph of the genus Hydrangea. Proc. California Acad. Sci. IV, 29:147-256.

McCuure, FA. 1973. Genera of bamboos native to the New World (Gramineae: Bambusoideae). Smithsonian

Contr. to Botany 9:1-148.

McDan S. 1971. The genus Sarracenia (Sarraceniaceae). Bull. Tall Timbers Res. Sta., Tallahassee, Florida.

McVaucH, R. 1936. Studies in the taxonomy and distribution of the eastern North American species of Lobelia.

Rhodora 38:241-263, 276-298, 305-329, 346-362.

MOHLENBROCK, R.H. 1961. A monograph of the leguminous genus Zornia. Webbia 16:1-141.

Ownbey, R.P. 1944. The liliaceous genus Polygonatum in North America. Ann. Missouri Bot. Gard. 31:373-413.

PelRSON, J.A., PD. Cantino, and H.E. BALLARD. 2006. A taxonomic revision of Collinsonia (Lamiaceae) based on phenetic

analyses of morphological variation. Syst. Bot. 31:398-409.

Pena, C.l. 1989. The systematics and evolution of Ludwigia sect. Microcarpium (Onagraceae). Ann. Missouri Bot.

Gard. 76:221-302.

Pena, C.l, C.L. Scuuipr, PC. Hock, and P.H. Raven. 2005. Systematics and evolution of Ludwigia section Dantia (Ona-

graceae). Ann. Missouri Bot. Gard. 92:307-359.

PENNELL, FW, 1920 ["1919"]. Scrophulariaceae of the southeastern United States. Proc. Acad. Nat. Sci. Philadelphia

71:224-291.

PENNELL, F.W. 1935. The Scrophulariaceae of Eastern Temperate North America. Acad. Nat. Sci. Philadelphia

Monogr. 1.

PunsH, F. 1814. Flora Americae Septentrionalis. London.

RADFORD, A.E., H.E. AHLes, and C.R. BeLL. 1968. Manual of the vascular flora of the Carolinas. Univ. of North Carolina

Press, Chapel Hill.

RAMAMOORTHY, T.P. and E.M. Zarbini. 1987. The systematics and evolution of Ludwigia sect. Myrtocarpus sensu lato

(Onagraceae). Monogr. in Syst. Bot., Missouri Bot. Gard. 19:1-120.

RemeeRT, D.H. 1980. Thomas Walter, Carolina botanist. South Carolina Mus. Comm, Bull. No. 5.

REMBERT, D.H. 1984. Program of Abstracts of Papers, to be presented at annual meeting of the Botanical Society

of America, Fort Collins, Colorado, Aug 1984. Amer. J. Bot. 71:107.

Reveal, J.L. 1990. Proposal to reject the name Cenchrus carolinianus Walter (Poaceae). Taxon 39:353-355.

REVEAL, J.L. and M.C. Johnston. 1989. A new combination in Phoradendron (Viscaceae). Taxon 38:107-108.

Rock, H.F.L. 1956. The binomials of Helenium in Walter's Flora Caroliniana. Rhodora 58:311-317; plate 1227.

Smr}, E.B. 1976. A biosystematic survey of Coreopsis in eastern United States and Canada. Sida 6:123-215.

TayLor, P. 1989. The genus Utricularia—a taxonomic monograph. Kew Bull., Addit. Ser. 14.

Tucker, A.O., N.H. DILL, T.D. Pizzolato, and R.D. KraL. 1983. Nomenclature, distribution, chromosome numbers, and

fruit morphology of Oxypolis canbyi and O. filiformis (Apiaceae). Syst. Bot. 8:299-304,

UnBATSCH, L.E. 1972. Systematic study of the Altissimae and Giganteae species groups of the genus Vernonia

(Compositae). Brittonia 24:229-238.

WALTER, T. 1788. Flora Caroliniana. London.

Warp, D.B. 1962a. The first record of the Fraser Fir. Castanea 27:78-79.

WarD, D.B. 1962b. The genus Anonymos and its nomenclatural survivors. Rhodora 64:87-92.

Ward, Thomas Walter Typification Project Il: known Walter types 423

Warp, D.B. 1977. Nelumbo lutea, the correct name for the American lotus. Taxon 26:227-234.

War, D.B. 2006. The Thomas Walter typification project, |. Observations on the John Fraser folio. Sida 22:

dde TS.

Warp, D.B. 2007. The Thomas Walter herbarium is not the herbarium of Thomas Walter. Taxon (in press).

WEssrER, G.L. 1970. A revision of Phyllanthus (Euphorbiaceae) in the continental United States. Brittonia 22:44-

76.

WiBEL, A.W. 1799. Prim. Fl. Werth.

Wiersema, J.H. and J.L. Reveal. 1991. Proposal to reject two 1788 Thomas Walter names of American waterlilies

(Nymphaeaceae). Taxon 40:509-51 6.

WILBUR, R.L. 1991. The recent neotypification and proposal to reject the name Cenchrus carolinianus Walter. Taxon

40:326-330.

WiLDENOw, C.L. 1803. Species Plantarum, ed. 4. Berlin.

WINDLER, D.R. 1974. A systematic treatment of the native unifoliolate crotalarias of North America (Leguminosae).

Rhodora 76:151-204.

ZONA, S. 1990. A monograph of Sabal (Arecaceae: Coryphoideae). Aliso 12:583-666.

424 Journal of the Botanical R h Institute of Texas 1(1)

BOOK NOTICES

HELEN PICKERING. 2006. Wildflowers of Mombacho, Nicaragua: Flores Silvestres del Mombacho. (ISBN

1-889878-14-6, pbk.). Sida, Botanical Miscellany 28. Botanical Research Institute of Texas, 509 Pecan

Street, Fort Worth, TX 76102-4060, U.S.A. (Orders: www.britpress.org, http//www.brit.org/Sida/Sida28.

htm, 817-332-4441 ext. 32). $15.00, 217 pp., 500 + color photographs, 642" x 81^".

Volcán Mombacho and its associated National Park are in southwestern Nicaragua, about 30 miles south of Managua, near the town of

Granada. The guide covers the whole of Volcán Mombacho (now dormant but with fumarolic fields and hot springs over a large area)

including both the evergreen, humid forest between 800 meters and the crest at 1345 meters and the dry, deciduous forest below 800

meters. If the scope of the book were limited to this small area, it might be of limited interest, because relatively few will be able to

visit the park. But the majority of plants covered occur in similar habitats throughout Central America and about one-third are widely

distributed in the tropics. For the tropically challenged (including me) this is a wonderful volume—use it as a review of families and

genera (even species, since many of them are widespread) or as an introduction to Central American plant diversity.

200 species in 174 genera, 63 families, are included (below). The species are first divided by color, then alphabetically by genus

and species. For each species, there are two or three excellent photos, often including fruits and habit. *Descriptions and information

on habitat and flowering periods are taken from the Flora of Nicaragua, published by the Missouri Botanical Garden in 2001."

Acanthaceae (6 genera, 6 species), Agavaceae (1, 1), Amaranthaceae (6, 6), Apocynaceae (3, 3), Araceae (5, 6), Asclepiadaceae (2,

2), Asteraceae (24, 34), Begoniaceae (1, 1), Bignoniaceae (1, 1), Boraginaceae (3, 3), Brassicaceae (2, 2), Bromeliaceae (4, 7), Campanu-

laceae (2, 2), Cannaceae (1, 1), Caprifoliaceae (1, 1), Clethraceae (1, 1), Clusiaceae (1, 1), Commelinaceae (5, 5), Convolvulaceae (1, 2),

Costaceae (1, 1), Cucurbitaceae (2, 2), Cyperaceae (2, 3), Ericaceae (2, 2), Euphorbiaceae (5, 5), Fabaceae (s.1.) (7, 7), Flacourtiaceae (1, 1),

Gesneriaceae (3, 3), Haemodoraceae (1, 1), Heliconiaceae (1, 2), Lamiaceae (1, 2), Loganiaceae (1, 1), Malpighiaceae (1, 1), Malvaceae (3,

4), Marantaceae (2, 2), Margraviaceae (1, 1), Melastomataceae (7, 7), Meliaceae (1, 1), Menispermaceae (1,1), Moraceae (1, 1), Myrinaceae

(1,1), Myrtaceae (2, 2), Nyctaginaceae (1, 2), Orchidaceae (13, 16), Oxalidaceae (1, 1), Palmae (1, 1), Papaveraceae (2, 2), Passifl

1), Phytolaccaceae (2, 2), Piperaceae (1, 1), Poaceae (1, 1), Polygalaceae (1, 1), Rubiaceae (10, 10), Scrophulariaceae (2, 2), Solanaceae (5, 8),

Sterculiaceae (1, 1), Tiliaceae (1, 1), Turneraceae (1, 1), Ulmaceae (1, 1), Urticaceae (2, 2), Verbenaceae (6, 6), Vitaceae (1, 1), Zingiberaceae

(2, 2), Zygophyllaceae (1,1).—Guy L. Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A.

Lynn CouLtTER. 2006. Gardening with Heirloom Seeds: Tried-and-True Flowers, Fruits, and Vegetables

for a New Generation. (ISBN 0-8078-5680-0, pbk.). University of North Carolina Press, RO. Box 2288,

Chapel Hill, NC 27515-2288, U.S.A. (Orders: www.uncpress.unc.edu, 919-966-3561, 919-966-3829

fax). $22.50, 316 pp., color photographs, 8" x 9".

Gardening with Heirloom iei Tried- m -True uou Fruits, and Vegetables for a New Generation is a journey into the world of

heirloom gardening. Heirl t least 50 den old. Many are easier to grow and more disease-resis-

tant than modern cultivars. The often taste better, as well In this "sampler for heirl seeds" Lynn Coulter describes 50 heirloom seed

varieties in loving detail. Most can be grown in MM di in the United States, but some are bind suited to the warmer climate of the

South, where the author makes her home. The p ized by gardening season: spring ugh winter. This paperback edition

is generously illustrated with gorgeous color c2 E a garden plants and color images from v vintage out-of-print seed catalogs. A

favorite of mine is the photograph of “Moon and Stars” watermelons, which have blue rinds spotted with yellow stars’ aust like the night

sky. Personal stories about the joy and meaning in heirloom seed gardening, seed saving, and seed swat g. In addition,

useful gardening advice and spaces for notes are also included, along with resources for heirloom seeds e gardens. P from the joy

that gardeners derive from growing unusual heirloom varieties, heirloom seeds have another value. They broaden the genetic diversity

of the plants in our modern food supply, which is important in avoiding disasters like the Irish potato famine.—Marissa N. Oppel, MS,

Collections and Research Assistant, Botanical Research Institute of Texas, Fort Worth, TX, 76102-4060, U.S.A.

J. Bot. Res. Inst. Texas 1(1): 424. 2007

THOMAS WALTER TYPIFICATION PROJECT, III:

LECTOIYPES AND NEOT YPES FOR 20 WALTER NAMES,

MO RECOGNIZED IN THE FRASER WALTER HERBARIUM

Daniel B. Ward

Department of Botany

University of Florida

Gainesville, Florida 32611, U.S.A.

ABSTRACT

Thomas Walter’s Flora Caroliniana (1788) contained numerous species new to science and whose names are of modern use. Many of the

species he described were independently collected by John Fraser, whose specimens were seen by Walter before being taken to England.

To ensure that Walter's names shall be used in a consistent way, appropriate specimens from the Fraser collection are here chosen as

types, to represent 20 of the names published by Walter.

RESUMEN

La Flora Caroliniana (1788), de Thomas Walter, j j nuevas para la ciencia, con nombres de uso moderno.

I L

1 1 E 1;

Muchas de las especies descritas por Walter f John Fraser f vistos po

Walter antes de ser llevados a Inglaterra. Para asegurar que los e de Walter serán usados en una manera consistente, se escogen,

como tipos, los especímenes apropiados de la colección de Fraser, para representar 20 de los nombres publicados por Walter.

The Thomas Walter Typification Project is intended to bring understanding and nomenclatural precision to

the many plant names published by Thomas Walter in his Flora Caroliniana (1788). The present task is to

choose from the Fraser/Walter herbarium those specimens that are believed to be part of the material used

by Walter, or are of such quality and confident identification that though not likely used by Walter may be

selected to represent his new species.

In the 1780s Thomas Walter, an English resident of South Carolina, operated a rice plantation on the

cleared bottomlands of the Santee River, in what is now Berkeley County. A classical education and an in-

quisitive mind led Walter to observe the native plants around him, and then to attempt their identification

through use of his few books by Carl Linnaeus. Though the descriptions were brief and in Latin, Walter was

able to match many of the local plants with Linnaeus' names. Other plants, however, did not match, and

Walter wrote his own descriptions, also in Latin, of the species he thought to be new. In 1786, John Fraser,

a venturous Scot in search of plants useful for English horticulture, met Walter and shared his interest in

the native species. In 1788 Fraser returned to England, taking with him his numerous collections and the

manuscript of Walter's new flora.

Once in London, Fraser promptly published Walter's Flora Caroliniana. This small volume contained

1056 species, many identified by Walter with names from Linnaeus, the rest given names of his own (un-

imaginatively, perhaps, assigning 58 of them “caroliniana”). Of the species treated in his book, 414 (or 39%)

were given new names.

The importance and number of the species now carrying names given them by Thomas Walter is seldom

appreciated. In the decades before Walter, Linnaeus had seen collections made by his student Pehr Kalm, by

John Clayton, the city official of Gloucester, Virginia, and by Patrick Browne and others in the West Indies.

From these sources he had learned of many plants of eastern North America and had published their descrip-

tions and names. But from the Carolinas to southern Florida, very little botanical information had reached

Europe. Walter was thus in position to find and name numerous species not previously known to science.

Walter kept no herbarium. Though later writers have stated that he had done so, and that it has since

been lost or was perhaps given to Fraser, in none of Walter's surviving correspondence, nor in his book, nor

in the writings of Fraser, is there evidence that Walter prepared or used a herbarium (Ward 20072).

J. Bot. Res. Inst. Texas 1(1): 425 — 430. 2007

£s+haD o ID

426 Journal of t h Institute of Texas 1(1)

But Fraser certainly did! Though he was in the Southeast for only 18 months, with collection possible

during only a single growing season, he gathered specimens widely and with enthusiasm. His field practices

were recorded with disdain by Michaux (Sargent 1889; Ward 1977, in transl.), and the often-fragmentary

specimens now surviving in his herbarium well illustrate his haste and inexperience. Walter had opportu-

nity to review much of Fraser’s collection—over half (368, or 53.5%)) of the specimens kept by Fraser bear

labels with Walter’s handwriting (Ward 2007a)—and to name many as best he could. A portion of these

specimens, selected by Fraser after his return to England, were retained by him. Others were sold to Charles

Louis L'Heritier (Fraser 1789; Stafleu 1963) and are now in the DeCandolle Herbarium, Geneva; still others

are in the Herbarium Lamarck, Paris.

The subset of 690 specimens retained by Fraser were mounted by him and/or his sons in a folio volume

now held by the Natural History Museum, London. They form what is often called the *Walter Herbarium,"

or more appropriately, the Fraser/Walter herbarium, and are frequently consulted for suggestion as to what

Walter may have meant by certain of his new names. A previous number of this series (Ward 2006) has

addressed the characteristics and contents of this folio herbarium. Many of the species described and given

new names by Walter are not represented in the Fraser folio. And even for those that are, there is no assur-

ance that Walter used them in preparation of his Flora (Ward 20072).

A great majority of the species described by Walter occur on the Carolina coastal plain and would very

possibly have been known to him during the years he was preparing his Flora. For such species there is no

reason to believe he had need of or made use of dried specimens, either from a herbarium of his own or that

of Fraser. Walter, in his Latin introduction, reported that *for a long time he [Walter] has cultivated in his

own garden the very many plants which he describes" (Walter 1788; Rembert 1980, in transl). Thus, even

though there may also be a specimen in the Fraser collection, for coastal plain species it is best to accept that

Walter's descriptions were prepared largely or wholly from fresh materials. Such species, if not addressed

previously by other authors, are now in need of a substitute type, a neotype.

The International Code of Botanical Nomenclature (McNeill et al. 2006) provides that when a name

is unrepresented by a type specimen, or the original type specimen has been lost, a replacement specimen

may be selected from another source. Such specimens are termed neotypes, and carry the same status as

the missing type.

But it has long been recognized (Harper 1911; Ewan 1969) that a number of species treated by Walter

do not grow within the limited area (perhaps 50 mile radius, centered on his Santee plantation) specified by

Walter (1788). The means by which Walter encountered these plants is believed to have been Fraser, who

is known to have traveled into the Appalachians of the western Carolinas and as far south as the Altamaha

River, Georgia (Ward 2006). Fraser himself (1789) claimed to have provided Walter with some 200 new

species, although survey of the Flora shows only 103 species (9.796) known to grow only outside Walter's

immediate area.

Though relatively few, these species now not known within Walter's area yet included within his Flora

pose a special problem. Often Walter's description contains observations well beyond what is shown by the

specimens alone—color and other details of the flowers (when the specimen itself is fruiting or sterile), height

of the plant, etc. It is thus possible Walter based his descriptions on more generous materials— perhaps also

provided by Fraser, perhaps from another source. Yet, with the generous assumption that the extra-territorial

specimens in the Fraser collection are related to Walter's basis, they may be treated as part of the original

material, or lectotypes.

The present number of this series includes only those lectotypes and neotypes that can adequately be

based on specimens of the Fraser/Walter herbarium. Another number of this series (Ward 2007b) lists all

known lectotypes and neotypes chosen by previous authors. A future number (or series of numbers) will

encompass the many neotypes that must be selected from new materials.

It is understood that a risk accompanies selection of neotypes, in that there can never be certainty what

was intended by the original author and that distortion of the original meaning will result if the new type

differs significantly from the one once in the author's hand. Yet absence of a type carries its own potential

Ward, Thomas Walter Typification Project III: lectoty; | neoty] 427

for misrepresentation through the lack of a fixed basis against which new collections may be tested. Though

previous authors have at times selected dubious or seriously defective specimens from the Fraser/Walter

herbarium as neotypes, their actions cannot now be abrogated. But it is believed the neotypes selected here

would not have been rejected by Thomas Walter as representative of his new species.

The 20 Walter types published here include 7 specimens that are designated as lectotypes and 13

specimens that are selected as neotypes. All are based upon specimens in the Fraser/Walter herbarium,

London. (Specimens are designated as described in Ward 2006.) The lectotypes are obligatory choices, once

the thesis is accepted that materials of extra-territorial species must have been seen by Walter. The neotypes

are chosen of specimens that display adequate diagnostic characteristics and are believed to represent their

species as currently understood. These typifications are here presented in the format used previously (Ward

2007b), in alphabetical sequence, using the names given them by Thomas Walter.

DYPIBICALIONS

WALTER'S NAME: Actaea dioica Walter (p. 152)

MODERN NAME: Áruncus dioicus (Walt.) Fern.

Nearly absent from SC, common in western NC; probably a Fraser discovery. Spm. 1-G is this, but bears

only “Actea” in Fraser's hand. The specimen is of decent quality and may be part of the materials used by

Walter in forming his description. Thus Fraser/Walter 1-G [1787] (BM) is here designated Lectotype of Actaea

dioica Walt., basionym of Aruncus dioicus (Walt.) Fern.

WALTER'S NAME: Andromeda Catesbaei Walter (p. 137)

MODERN NAME: Leucothoe axillaris (Lam.) D. Don

Common in eastern SC. Spm. 6-H was labeled “Andromeda Catesbaei" by Walter. It was annotated *Holotypus

for the name A. catesbaei Walt." by N. C. Melvin in 1976, but the designation has not been published. Even

though the name will remain in synonymy under Leucothoe axillaris (2 Andromeda axillaris Lamarck, 1783),

Fraser/Walter 6-H [1787] (BM) is of respectable quality and justifies selection here as NEOTYPE for Andromeda

catesbaei Walt.

WALTER'S NAME: Anonymos paniculatla] Walter (p. 198); nom. illegit.

MODERN NAME: Carphephorus paniculatus (Walt. ex Gmel.) Hebert [= Trilisa paniculata (Walt. ex Gmel.) Cass.]

Common in eastern SC. Although Anonymos paniculata is illegitimate, Gmelin (1792: 1204) based his name on

Walter's description (Ward 1962). Spm. 32-D appears to be this. The handwriting is muddled, but includes

“paniculata” in Walter's hand. Walter would not have needed this specimen. But its quality is fair (though

its basal leaves are lacking), and Fraser/Walter 32-D [1787] (BM) is here selected as NEOTYPE for Chrysocoma

paniculata Gmel., basionym for Carphephorus paniculatus (Walt. ex Gmel.) Hebert.

WALTER'S NAME: Anonymos tinctori[a] Walter (p. 68), nom. illegit.

MODERN NAME: Lachnanthes caroliniana (Lam.) Dandy [= Lachnanthes tinctoria (Walt. ex Gmel.) Ell.]

Common on the SC coastal plain. This plant was long known as Lachnanthes tinctoria (Walt.) Ell. After the

decision by the 1950 International Botanical Congress that combinations made under “Anonymos” were il-

legitimate, Ward (1962) proposed Gmelin (1791: 113) as the first validating author. Wilbur (1962) found that

Lamarck had done so five months earlier, but made the unneeded combination, L. caroliniana (Lam.) Wilbur.

Fraser/Walter 117-C [1787] (BM), labeled “nova genera” in Fraser’s hand and named to genus by A. Gray, is here

selected as NEOTYPE for Heritiera tinctorium Gmel., basionym of Lachnanthes tinctoria (Walt. ex Gmel.) Ell.

WALTER'S NAME: Arethusa racemosa Walter (p. 222)

MODERN NAME: Ponthieva racemosa (Walt.) Mohr

Occasional in coastal SC. The label of spm. 8-D ("Arethusa racemosa”) is in Walter's hand. But since the

species would have been available to Walter near his home and the accompanying 3-digit number (“??9”)

indicates the specimen to be a Fraser collection, it may not have been seen by Walter until after completion

of his manuscript. Blake (1915) correctly called it “an excellent specimen.” Fraser/Walter 8-D [1787] (BM) is

thus here selected as NEOTYPE for Arethusa racemosa Walt., basionym of Ponthieva racemosa (Walt.) Mohr.

428 Journal of the Botanical R h Institute of Texas 1(1)

WALTER'S NAME: Asclepias polystachia Walter (p. 107)

MODERN NAME: Asclepias exaltata L.

Westernmost NC and SC, unknown on the SC coastal plain. Spm. 10-H was labeled “Asclepias Novum" by

Fraser. The specimen was identified (from photo) by Fernald & Schubert (1948: 218-220) as this species, but

was not called its type. The specimen is mediocre, with three leaves and a largely destroyed inflorescence.

But since Walter must have relied on Fraser for material of this common but exclusively montane species,

Fraser/ Walter 10-H [1787] (BM) is here designated Lectortyee of Asclepias polystachia Walt.

WALTER'S NAME: Carpinus caroliniana Walter (p. 236)

MODERN NAME: Carpinus caroliniana Walt.

Common throughout. Spm. 26-A bears “Carpinus” in Walter's hand. No type is known to have been des-

ignated elsewhere. Since this tree is found on what was Walter's Santee River property, there would have

been no need for him to have relied on this specimen. Even so, the specimen is of adequate quality, and

Fraser/Walter 26-A [1787] (BM) is here selected as NEoTYPE for Carpinus caroliniana Walt.

WALTER'S NAME: Cinna glomerata Walter (p. 59)

MODERN NAME: Andropogon glomeratus (Walt.) BSP.

Common throughout the SC coastal plain. Hitchcock (1905: 32) identified a specimen (spm. 113-E) as

Andropogon macrourus Michx., presently treated as a synonym of Andropogon glomeratus. He associated the

specimen with Walter's name on the basis of its label, “Cinna glomerata,” but did not refer to it as a type.

Though he did not comment on the handwriting, it is that of Walter. Since Andropogon glomeratus is com-

mon, this specimen may have been obtained by Fraser anywhere in his travels and need not have been used

by Walter in preparing his description. It is however of reasonably good quality. Having been addressed by

Hitchcock, Fraser/Walter 113-E [1787] (BM) is here selected as NEOTYPE for Cinna glomerata Walt., basionym

of Andropogon glomeratus (Walt.) BSP.

WALTER'S NAME: Cinna lateralis Walter (p. 59)

MODERN NAME: Andropogon virginicus L.

Common throughout. Hitchcock (1905: 33) interpreted a specimen (spm. 113-C), bearing the label “Cinna”

in Fraser's hand, as “probably the basis” of Cinna lateralis Walt., and identified it as “one of the forms of

Andropogon virginicus L.” Though the name will surely remain in synonymy, Fraser/Walter 113-C [1787] (BM)

is here selected as NEOTYPE for Cinna lateralis Walt.

WALTER'S NAME: Cypripedium reginae Walter (p. 222)

MODERN NAME: Cypripedium reginae Walt.

Very rare: in the Carolinas, known only in NC (2 counties). Spm. 39-B is labeled *Cypripedium Reginae" in

Fraser’s hand. It has been marked as “type” (by O. Ames?), but the designation has not been published.

Since Walter could have seen this species only through the agency of Fraser, this specimen (or another of

the same gathering) was probably used by him. Thus Fraser/Walter 39-B [1787] (BM) is here designated

LECTOTYPE of Cypripedium reginae Walt.

WALTER'S NAME: Eupatorium compositifolium Walter (p. 199)

MODERN NAME: Eupatorium compositifolium Walt.

Common in eastern SC. Spm. 46-B, an excellent specimen, was identified (from photo) as Eupatorium

compositifolium compositum," writ in haste] by Fernald & Schubert (1948: 227). Its label, “Eupatorium,” is

in Walter's hand, but there is no evidence it was used by him in preparation of his text. Its quality justifies

selection here of Fraser/Walter 46-B [1787] (BM) as NEOTYPE for Eupatorium compositifolium Walt.

WALTER'S NAME: Iva imbricata Walter (p. 232)

MODERN NAME: Iva imbricata Walt.

Frequent along SC coast. Spm. 61b-B appears to be this. It was labeled “Iva ? nova” by Walter. It has been

annotated as “Lectotype” (presumably of Iva imbricata) by Lillian T. Gillis (date unknown); this designation

is believed not to have been published. The specimen is of good quality, and merits recognition as a type.

Ward, Thomas Walter Typification Project III: lectoty, | neotyj 429

But, in light of Walter's failure to recognize his own species, it can scarcely be ranked as lectotype. Thus

Fraser/Walter 61b-B [1787] (BM) is here selected as NEOTYPE for Iva imbricata Walt.

WALTER'S NAME: Limodorum praecox Walter (p. 221)

MODERN NAME: Spiranthes praecox (Walt.) Watson

Infrequent on SC coastal plain. Spm. 65-D appears to be this. It was labeled “Limodorum” by Fraser, and

has been annotated as “praecox” (by O. Ames?). There is no indication it was seen or used by Walter. But

Fraser/Walter 65-D [1787] (BM) is of good quality and is here selected as NEOTYPE for Limodorum praecox Walt.,

basionym of Spiranthes praecox (Walt.) Watson.

WALTER'S NAME: Lysimachia lanceolata Walter (p. 92)

MODERN NAME: Lysimachia lanceolata Walt. [= Steironema lanceolatum (Walt.) Raf.]

Not known in eastern SC, frequent westward, notably along Savannah River where Fraser traveled; surely

his discovery. Spm. 68-E is this; it was labeled “Lysimachia nov" by Walter. It is a decent specimen of an

entire plant. The probability is high that Walter relied on Fraser's material. Thus Walter 68-E [1787] (BM) is

here designated Lectotype of Lysimachia lanceolata Walt.

WALTER'S NAME: Magnolia Fraseri Walter (p. 159)

MODERN NAME: Magnolia fraseri Walt.

Found only in mountains of western NC and SC; a John Fraser discovery. Spm. 70-A was labeled by Fraser

as “Magnolia Fraseri." The specimen has been annotated as “type specimen" (presumably by J.E. Dandy 1929),

though this choice appears not to have been published. The fold-out plate in Flora Caroliniana cannot be the

type, in that it was prepared in England and not seen by Walter until after publication. Since Walter only

had access to material of this species through the efforts of Fraser, it is appropriate that Fraser/Walter 70-A

[1787] (BM) is here designated Lectotype of Magnolia fraseri Walt.

WALTER'S NAME: Prenanthes autumnalis Walter (p. 193)

MODERN NAME: Prenanthes autumnalis Walt.

Frequent in eastern SC. Spm. 87-C (no label) was annotated as Prenanthes autumnalis by S.F. Blake. The

epithet when published in the Flora was not italicized, as is usual for Walter's names. Although there is no

indication this specimen was seen or used by Walter, itis of good quality, and Fraser/Walter 87-C [1787] (BM)

is here selected as NEOTYPE for Prenanthes autumnalis Walt.

WALTER'S NAME: Rhamnus carolinianus Walter (p. 101)

MODERN NAME: Rhamnus caroliniana Walt.

Absent from SC coastal plain, frequent inland; probably a Fraser discovery. Spm. 91-F was labeled *Rham-

nus novus" by Walter, with *Carolinianus" added by Fraser. The specimen is of poor quality, but is readily

identifiable. Since Walter could scarcely have seen this species without the agency of Fraser, Fraser/Walter

91-F [1787] (BM) is here designated Lectotype of Rhamnus caroliniana Walt.

WALTER'S NAME: Sium suave Walter (p. 115)

MODERN NAME: Sium suave Walt.

Occasional along SC coast. Spm. 42-F has been identified as Sium suave by Blake (1915: 131) from direct

exam, and by Fernald & Schubert (1948: 217) from photo. Neither, however, called it the type. The label

(“Eryngium”) is by Fraser and there is no evidence the specimen was seen or used by Walter. Still, it is of fair

quality and, having been confirmed by the above authors, Fraser/Walter 42-F [1787] (BM) is here selected

as NEOTYPE for Sium suave Walt.

WALTER'S NAME: Statice caroliniana Walter (p. 118)

MODERN NAME: Limonium carolinianum (Walt.) Britt.

Occasional along the SC coast. Spm. 101-D, a quite good specimen, is this species. Although the generic

name (“Statice”) is in Walter's hand, the specific name (“Caroliniana”) was assigned by Fraser. It is unlikely

that this specimen was used by Walter in writing his Flora. Even so, Fraser/Walter 101-D [1787] (BM) is

430 Journal of the Botanical R h Institute of Texas 1(1)

judged adequate to be selected here as NEOTYPE for Statice caroliniana Walt., basionym of Limonium carolin-

ianum (Walt.) Britt.

WALTER'S NAME: Vincetoxicum acanthocarpos Walter (p. 104)

MODERN NAME: Matelea carolinensis ( Jacq.) Woodson

Rare on SC coastal plain, common inland; perhaps a Fraser discovery. The epithet was not italicized in the

Flora, though the name is by Walter. Walter noted two varieties: one, “corollis purpureis", is surely M. caroli-

nensis; the other, “corollis nigricantibus" (“blackish”), is unassignable. Spm. 109-B was labeled “Vincetoxicum”

by Walter, and has been annotated (by S.F. Blake?) as *V. acanthocarpos Walt." It was suggested by Drapalik

(1970: 59) from photo, to be a *probable lectotype" of Walter's species. Jacquin's epithet is earlier (1787),

and typification of Walter's name is not imperative. However, in support of Drapalik's tentative suggestion,

Fraser/Walter 109-B [1787] (BM) is here designated Lectotype of Vincetoxicum acanthocarpos Walt. (= Matelea

carolinensis (Jacq.) Woodson).

ACKNOWLEDGMENTS

I am grateful to Bernice G. Schubert (GH), whom I never knew, for her 1946 photographs of the Fraser/

Walter herbarium, to Carroll E. Wood (GH) for his generosity in lending me her negatives, to Christine M.

Housel (ABT) for composition of the Spanish abstract, to Alexander Krings (NCSC) for his careful reading

of the manuscript, and to Charles E. Jarvis (BM) for his encouragement and advice in development of the

Thomas Walter Typification Project.

REFERENCES

BLAKE, S.F. 1915. Some neglected names in Walter's Flora Caroliniana. Rhodora 17:129-137.

DnaPAuk, D.J. 1970. A biosystematic study of the genus Matelea in the southeastern United States. Ph.D. diss.,

Univ. of North Carolina, Chapel Hill.

Ewan, J. 1969. Historical problems for the working taxonomist. Taxon 18:194-203.

FERNALD, M.L. and B.G. SCHUBERT. 1948. Studies of American types in British herbaria. Part IV: some species of Thomas

Walter. Rhodora 50:190-208, 217-229.

FRASER, J. 1789. A short history of the Agrostis Cornucopiae: or, the new American grass. Chelsea, England.

GMELIN, J.F. 1791, 1792. Systema Naturae. Leipzig.

HARPER, R.M. 1911. Early spring aspects of the coastal plain vegetation of South Carolina, Georgia, and northeastern

Florida. Bull. Torrey Bot. Club 38:223-234.

HitcHcock, A.S. 1905. The identification of Walter's grasses. Missouri Bot. Gard. Ann. Rep't. 16:31-56.

McNüiL, J., F.R. Barrie, H.M. Bunper, V. DemouLin, D.L. HAWKsworTH, K. MARHOLD, D.H. NicoLson, J. PRADO, PC. SILVA, J.E. SkOG,

N.J. TURLAND, and J. Wiersema, eds. 2006. The international code of botanical nomenclature (Vienna Code), July

2005. Regnum Veg. 146:1-568.

REmMBERT, D.H. 1980. Thomas Walter, Carolina botanist. South Carolina Mus. Comm, Bull. No. 5.

SARGENT, C.S. 1889. Portions of the journal of Andre Michaux, botanist, written during his travels in the United

States and Canada, 1785-1796. Proc. Amer. Philos. Soc. 26:1-145.

STAFLEU, F.A. 1963. ['Heritier de Brutelle: the man and his work. Preface to facsimile, Sertum Anglicum, 1788. Hunt

Botanical Library, Pittsburg, PA.

WacrtR, T. 1788. Flora Caroliniana. London.

Waro, D.B. 1962. The genus Anonymos and its nomenclatural survivors. Rhodora 64:87-92.

Warp, D.B. 1977. Nelumbo lutea, the correct name for the American lotus. Taxon 26:227-234.

War, D.B. 2006. The Thomas Walter typification project, |. Observations on the John Fraser folio. Sida 22:

1111-1118.

Warp, D.B. 2007a. The Thomas Walter herbarium is not the herbarium of Thomas Walter. Taxon (in press).

Ward, D.B. 2007b. The Thomas Walter typification project, Il. The known Walter types. J. Bot. Res. Inst. Texas

1:407-423.

WILBUR, R.L. 1962. The identity of Walter's species of Anonymos. J. Elisha Mitchell Sci. Soc. 78:125-132.

CHROMOSOME NUMBER OF LAUBERTIA CONTORTA (APOCYNACEAE:

APOCYNOIDEAE) AND ITS PHYLOGENETIC IMPORTANCE

Justin K. Williams and Dawn P. Derr'

Department of Biological Sciences

Sam Houston State University

Huntsville, Texas 77341-2116, U.S.A.

ABSTRACT

The mitotic chromosome count for Laubertia contorta (2n = 18) is the first reported chromosome count for the genus. A discussion of

the relationship of Laubertia to other genera and the use of chromosome numbers in constructing non-molecular phylogenies in the

Apocynaceae is presented. A distribution map and color photo of the species is also provided.

Key Wonps: Laubertia, Apocynaceae, chromosome, Forsteronia, Prestonia, Mexico

RESUMEN

Se realiza el recuento de cromosomas en mitosis de Laubertia contorta (2n = 18) que es el primero el género. Se presenta una discusión

de la relación de Laubertia con otros géneros y el uso de los números cromosomáticos en la construcción de filogenias no-moleculares

en Apocynaceae. Se aporta también un mapa de distribución y fotografia en color de la especie.

A cursory review of chromosome numbers in the Apocynaceae (Van der Laan & Arends 1985; Goldblatt &

Johnson 2003) indicates that 73 of the 179 genera in the Apocynaceae have been counted. Few chromosome

counts in the Apocynaceae are meiotic because plants characteristically produce only between 3-8 flowers

per inflorescence. In addition, because the anthers produce little pollen, it is difficult to fix the anthers at

the proper meiotic phase for counting. Therefore most counts in the Apocynaceae are mitotic counts made

from the region of cell division in root tips using the “squash” technique (Witkus 1951; Raffauf 1964). The

majority of taxa counted are those commonly cultivated and therefore with readily accessible root tips.

Van der Laan and Arends (1985) discussed in detail the systematic utility of chromosome numbers in the

Apocynaceae. Based on their observations, Van der Laan and Arends (ibid) suggested that chromosome

numbers have the potential for resolving some important relationships in the Apocynaceae. In addition,

they pointed out that only eight of the 55 genera in the Americas have had chromosome counts reported,

and they strongly suggested that more counts be pursued.

Laubertia A. DC. comprises three species of Neotropical lianas native from Central Mexico to northern

South America (Morales 2002). One species, Laubertia contorta (Mart. & Gal.) Woodson, is endemic to Mexico,

where it ranges from Sinaloa to Chiapas (Fig. 1) and is readily distinguished from the other species of the

genus by its twisted corolla tube and deep maroon corollas lobes (Fig. 2a). The relationship of Laubertia is

disputed (Williams 1999), with some authors relating it to Echites P. Browne and others to Prestonia R. Br.

(Morales 2002). A morphological cladistic analysis (Fig. 3, Williams 2004) places Laubertia basal to a clade

of four species of Prestonia (here referred to as the “Prestonia” clade). Although Laubertia was nested with

Prestonia, the bootstrap support for this relationship was below 5096. Chromosome numbers were not used

for one of the character states in this analysis. Since the time of this analysis chromosome counts in the

Apocynaceae have accumulated, including the one presented here. This paper provides the first chromosome

count for Laubertia and examines the utility of chromosome numbers in morphological cladistic analyses.

MATERIALS AND METHODS

Voucher and fruiting specimens (Fig. 2b) were collected during January 2003 in Mexico (Fig. 1) by the first

author. Voucher specimens were mounted and deposited at SHST and GH. MEXICO. CHIAPAS: 25 ft inside

'Current address: 14 Twin Feather Place, The Woodlands, Texas 77381, U.S.A.

J. Bot. Res. Inst. Texas 1(1): 431 — 435. 2007

432

f4L,D

Journal of tanical R h Institute of Texas 1(1)

ms! "o * LN 100 * 325 LJ a

I q I ' 1 t

sb \ »

À ;

4 à

a A

` '

Bs ` E 7

| =

/ i ^ \ 5 E

. Y Ñ y ec ER Fa M

n T e i i Sj qm

\ $ Doy N

N TTL EK y []

* 4 1 i | P, = | i

` a / X

A ‘a P ad i à

"5 ln k g "

2. $5,

Ep v

, * H

x DES ge i/o

20h. Dn ES - / PEN M Pd —|M

s i " Net"

«f YA. Y? i

_ ——À H—Ó f

o 200 AOI m es & e py

J 4

d MP Ns P

rove ar ed 3

ee V Mi

e ¡e

to is

] | l i i

i" "o 105 100 Lo N +

Fic. 1 Nieto ds 1£, 1999) TL p Lalit] I T d.

the park from the entrance to *El Choreoderro" waterfall park ca. 10 mi N of Tuxtla Gutiérrez, 2 Jan 2003,

Williams 2003-1 (SHST, GH).

The collected seeds (Fig. 2c) were planted in small starter pots with commercial potting soil in July

2003 and were maintained in the greenhouse at the Department of Biological Sciences at Sam Houston

State University. Twenty-four seeds were planted with no additional measures performed, such as soaking

or scarification. Fifty percent of the seeds germinated (Fig. 2d) over a three month period. Once the plants

developed several leaves (Fig. 2e) and an extensive root system, actively growing root tips (Fig. 2f) were

collected, fixed, and analyzed using standard procedures (Van der Laan & Arends 1985).

Finally, using the same taxa, character matrix, and methodology in Williams (2004), we re-analyzed

the data adding an additional character: chromosome number. Chromosome numbers for 29 of the 45 taxa

included in Williams (ibid) were identified from the present study and a literature search (Van der Laan &

Arends 1985; Goldblatt & Johnson 2003). The genera identified with chromosome counts were then scored

based on their respective base number: 0: x = 11 (Adenium Roem. & Schult., Apocynum L., Nerium L.); 1:

x = 10 (Cerbera L., Strophanthus DC., Thevetia L.); 2: x = 9 (Forsteronia G. Mey, Laubertia, Parsonsia R. Br.,

Prestonia); 3: x = 6 (Echites, Odontadenia Benth., Pentalinon Voigt); 4: x = 8 (Mandevilla Lindl.).

RESULTS

A mitotic chromosome number of 2n = 18 or a base number of x = 9 was observed for Laubertia contorta. Van

der Laan and Arends (1985) reported chromosome length in the Apocynaceae between 0.5—4 um, with the

average chromosome length between 1-2 um. The length of the chromosomes in L. contorta varies between

0.5-1.5 um, consistent with other chromosomes in the Apocynaceae.

MEIL in Ch 1

of Laubertia contorta 433

Fic. 2. Phot hs of vari t ] ts of Lauberti tort Corolla: b. follicles: c. Seed witt d i lli iuvenile plant

1 1 JI J F

with leaves; f. root tips.

434 Journal of the Bot R h Institute of Texas 1(1)

Laubertia contorta

Prestonia acutifolia

46/52 Prestonia mexicana

76/73 66/68 Prestonia tomentosa

Prestonia portobellensis

ap values. The first number indicates values

Fic. 3. Dendogram of “Prestonia” clade (from Williams 2004). N

letermined in this study and tl 1 number is from tl 2004 “Only the value for the “lgubertig Preston” branch

LN,

The heuristic search in Williams (2004) yielded a total of 48 equally parsimonious trees of 159 steps.

The new search with chromosome counts added yielded a total of 98 equally parsimonious trees of 164

steps. The bootstrap value for the Prestonia clade (Fig. 3) was below 50% (46%) when chromosomes were

not included (Williams 2004) and over 50% (52%) when chromosomes were included.

DISCUSSION

Van der Laan and Arends (1985) reported x = 11 as the base chromosome number for the Apocynaceae based

on its prevalence in the family and on the observation that many of the plesiomorphic taxa possess a base

number of x = 11. A review of the chromosomes numbers in the Apocynaceae presented by Van der Laan

and Arends (ibid) indicates that a base chromosome number of x = 9 is found in eight genera represented

in four of the 21 tribes recognized in the Apocynaceae (Endress @ Bruyns 2000): Mesechites (Forsteronia),

Echiteae (Laubertia, Parsonsia, Prestonia), Malourtieae (Pachypodium Lindl.), and Plumerieae (Allamanda L,

Plumeria L.). The Plumerieae is in the subfamily Rauvolfioideae (anthers free from pistil head and aestivation

of corolla bud sinistrorse) and differs from the other three tribes which are in the Apocynoideae (anthers fused

to the pistil head and aestivation of corolla bud dextrorse). Based on subfamilial and tribal circumscription

(Endress & Bruyns 2000) along with phylogenetic evidence (Williams 2004) it is suggested here that the

evolution of x = 9 evolved independently in potentially four different clades within the Apocynaceae.

A cursory review of chromosome counts of the Apocynaceae (Van der Laan & Arends 1985; Goldblatt

& Johnson 2003) shows that at present the only chromosome counts for Apocynoideae genera suggested

and potentially related to Laubertia are for Echites (x = 6), Forsteronia (x = 9), Parsonsia (x = 9), Pentalinon (x

= 6), and Prestonia (x = 9). The increased bootstrap value for the Laubertia/Prestonia (Fig. 3) clade calculated

with the additional character state (chromosome numbers) provides further support for the close relationship

between the two genera. And although the re-sampling of the data matrix, only increased the bootstrap value

of the Laubertia/Prestonia by 696, this increased the value of the clade to over 5096, which is often utilized

as the lower threshold value for the beginning of support in phylogenies constructed from large scale data-

sets (Sanderson & Wojciechowski 2000) like this one. In addition, the results indicate that chromosome

numbers are useful in constructing phylogenies in the Apocynaceae. Based on the phylogenetic significance

of chromosome counts as potential tribal synapomorphies, it is suggested that further attempts be made to

secure chromosome counts for other neotropical genera of the Apocynoidae, specifically Allotoonia Morales

& J.K. Williams, Angadenia Miers, Fernaldia Woodson, Rhabdadenia Muell-Arg., Mesechites Muell-Arg., The-

nardia H.B.K., Thoreauea J.K. Williams and additional species of Echites and Prestonia.

ACKNOWLEDGMENTS

We thank Tami Cook for providing us with access to her digital light microscope, Andrew Dewees for veri-

fying the different mitotic stages and the chromosome count, Guy Nesom and an anonymous reviewer for

Williams and Derr, Chromosome number of Laubertia contorta 435

making valuable edits to the manuscript and Bob Rhodes for mixing the Carnoy's solution and aceto-orcein.

We also thank Jon Plum for providing photographs of Laubertia contorta flowers (Fig. 2a).

REFERENCES

ENDRESS, M.F. and P. Bruyns. 2000. A revised classification of the Apocynaceae s.l. Bot. Rev. 66:1-56.

GoLbsLarr, P. and D.E. Johnson. 2003. Index to plant chromosome numbers 1998-2000. Monogr. Syst. Bot. Mis-

souri Bot. Gard. 94.

Morales, J.F. 2002. Studies in Neotropical Apocynaceae |: A revision of the genus Laubertia. Rhodora 104:

170-185.

RAFFAUF, R.F. 1964. Some chemotaxonomic considerations in the Apocynaceae. Lloydia 27:288-298.

SANDERSON, M.J. and M.F. WojciecHowski. 2000. Improved bootstrap confidence limits in large-scale phylogenies,

with an example from neo-Astragalus (Leguminosae). Syst. Biol. 49:671-685.

VAN DER LAAN, EN. and J.C. AnENbs. 1985. Cytotaxonomy of the Apocynaceae. Genetica 68:3-35.

WiLLIAMS, J.K. 1999. A phylogenetic and taxonomic study of the Apocynaceae subfamily Apocynoideae of Mexico

with synopsis of subfamily Plumerioideae. Ph.D. dissertation. Austin: The University of Texas, Austin.

WiLLIAMS, J.K. 2004. Polyphyly of the genus Echites (Apocynaceae: Apocynoideae: Echiteae): evidence based on

a morphological cladistic analysis. Sida 21:117-131.

Witkus, E.R. 1951. The chromosome number of several species of Strophanthus. Bull. Torrey Bot. Club 78:

80-82.

436 Journal of the Botanical R h Institute of Texas 1(1)

BOOK REVIEWS

ETHAN B. Russo and FRANJO GROTENHERMEN (eds.). 2006. Handbook of Cannabis Therapeutics from Bench

to Bedside. (ISBN 0-7890-3097-7, pbk.). Haworth Press, Inc., 10 Alice Street, Binghamton, New York,

NY 13904-1580, U.S.A. (Orders: www.haworthpress.com, orders@haworthpress.com, 1-800-429-6784,

607-722-5857). $39.95, 471 pp., 6" x 8Y2".

+] 1 il

Despite regulatory problems, lot of research on C bis sati 1 the pl hemical ds d d from it. Humans

produce endocannabinoids that act on cannabinoid receptors 1 and 2 (CB, and CB). CB ‚receptors are found mainly in the brain, spinal

chord, and peripheral nervous system and are associated with the psychoactive properties of Cannabis sativa. They are also associated

with motor control, memory processing, and pain control, making them targets for drug discovery efforts. CB, receptors are found in

the immune system and are associated with Lau release. They are not associated with the psychoactive effects of cannabinoids. This

makes the CB, recep ] fi 1 efforts, particularly for analgesic, anti-inflammatory, and antineoplastic

drugs. The md duce of phytochemical ae un Mn psychoactive effects like A?- a as well as non-

psychoactive compounds like cannabinol. T ] 1 many others derived from C

by researchers, ene the pone giant Phizer, as new drugs to treat conditions ME chronic pain, nausea, and mm

side effects from cancer chemotherapy and radiotherapy, as well as glaucoma, obesity, Parki 's

ase, and schizophrenia.

The Handbook of Cannabis side From Bench to Bedside is an informative guide to the world of cannabis research. It begins

with historical ethnobotanical studies of ancient Arab and Greek texts. Three chapters are devoted to the pharmacokinetics and phar-

macodynamics of the cannabinoids, including synergistic effects. This is followed by two chapters on the endocannabinoids and their

receptors. A total of seven chapters are devoted to the medicinal uses of Cannabis sativa, including the treatment of nausea from cancer

chemotherapy, loss of appetite in AIDS patients, multiple sclerosis, and crack cocaine addiction. The last section of the book is titled

"Side Effects." It includes an interesting chapter by a registered nurse on Cannabis sativa and harm reduction, including the effect that

legalization would have on the duties of health care professionals. The last chapter focuses on the positive and negative consequences

of legal Cannabis sativa in a clinical setting.

Many of the chapters in this book wer

p culled from the Journal of Cannabis Therapeutics, which is no longer in existence

and hard to obtain. In addition, the book's editors and contributors are experts in the field of cannabis research. Therefore, this book

is a useful source of hard-to-find scientifically validated information on Cannabis sativa as a medicine, making it a wonderful antidote

to the massive amounts of misinformation and rumors about this culturally significant plant.—Marissa N. Oppel, MS, Collections and

Research Assistant, Botanical Research Institute of Texas, Fort Worth, TX, 76102-4060, U.S.A.

Thomas J. ELreL. 2006. Botany in a Day: The Patterns Method of Plant Identification, 5* edition. (ISBN

1-892784-15-7, pbk.). Hops Press, LLC, 12 Quartz Street, Pony, MT 59747-0697, U.S.A. (Orders:

www.hopspress.com, www.wildflowers-and-weeds.com, www.mountain-press.com). $25.00, 221 pp.,

b/w illustrations, 815" x 11".

Botany in a Day: The Patterns Method of Plant Identification is an excellent learning tool. It is focused on plants that grow in the northern

parts of North America, however it is still useful for those of who live in the southern United States. The first part is a textbook on

plant identification. It is intended to be completed in one day, although it took me longer than that. Unlike a field guide, which is a list

of individual species, this guidebook is focused on recognizing visual patterns of plant groups. The book is illustrated with copious

black-and-white line drawings that are incredibly helpful visual aids. The second part is focused on the edible and medicinal uses of

wild plants, including an introduction to phytochemistry. This book is an excellent introduction to plant identification in the field. It

has helped me to learn how to identify plants that grow here in North Central Texas.— Marissa N. Oppel, MS, Collections and Research

Assistant, Botanical Research Institute of Texas, Fort Worth, TX, 76102-4060, U.S.A.

J. Bot. Res. Inst. Texas 1(1): 436. 2007

VASCULAR PLANT TYPE SPECIMENS AN THE

UNIVERSITY OF BRITISH COLUMBIA HERBARIUM (UBC)

Jeffery M. Saarela' Linda Lipsen

Department of Botany, and Department of Botany

UBC Botanical Garden and Centre for Plant Research University of British Columbia

University of British Columbia Vancouver, British Columbia, CANADA

Vancouver, British Columbia., CANADA lindaj@interchange.ubc.ca

Cindy M. Sayre Jeannette Whitton

VanDusen Botanical Garden Department of Botany

tree University of British Columbia

Vancouver, British Columbia, CANADA Vancouver, British Columbia, CANADA

jwhitton@interchange.ubc.ca

ABSTRACT

ls TEN b Js

We provide a brief a history of the University of British Columbia

plant type specimens in UBC.

RESUMEN

Se aporta una breve historia del herbario de la Universidad de la Columbia Británica y su colección de plantas vasculares, y un catalogo

de tipos de plantas vasculares en UBC.

INTRODUCTION

General Collection History

The University of British Columbia Herbarium (UBC; Holmgren & Holmgren 1990) is the largest herbarium

in western Canada, with approximately 600,000 accessioned specimens, including 242,000 bryophytes,

222,000 vascular plants, 67,000 algae, 35,000 lichens, and 15,000 fungi. John Davidson initiated the col-

lection in 1912, when he became the Provincial Botanist of British Columbia. The original herbarium (not

affiliated with UBC at the time) was located in the botanical offices in downtown Vancouver, and subsequently

moved to a number of downtown locations before being moved to a temporary UBC location at Fairview in

1916, when John Davidson became “the demonstrator in charge of UBC herbarium and botanical garden.”

The collections were moved to the university campus around 1925, and were housed in various buildings

closest to their curators, with the vascular plant collection located in what is now the Geology building (D.

Brownstein, pers. comm.). In 1973, all five collections were moved to their present location in the Biological

Sciences building. The collection will soon move again to the Beatty Biodiversity Museum, along with all

the other non-anthropological UBC natural history collections.

History of the Vascular Plant Collection

Although details of the early growth of the vascular plant collection are unknown, it is well documented

that many early British Columbia collectors donated their entire collections to John Davidson. Substantial

early additions to the herbarium collection were donated by: Albert J. Hill (UBC database collecting years:

1875-1912; 2,500 specimens from various families, mostly from British Columbia); Eli Wilson (1903-1913;

21,000 specimens from various families, all from BC); W. Taylor (1912-1937; 900 specimens from various

families, all BC); and A.E. Baggs (1910-1929; 1,000 specimens from BC and England) (The University of

1C pondi tl 1C t Add R h Division, Canadian M f Nature, P.O. Box 3443, Station D, Ottawa, Ontario, CANADA, jsaarela@mus-nature.ca

J. Bot. Res. Inst. Texas 1(1): 437 — 448. 2007

438 Journal of the Botanical R h Institute of Texas 1(1)

British Columbia Calendar 1922-1923; UBC database). Much of the early collection was destroyed during

the 1930s by insect damage (W. Schofield, D. Brownstein, pers. comm., September 2006).

The size of the vascular plant collection increased substantially over the following decades, as a result

of the efforts of many collectors and exchanges with other institutions. The collection included ~8,000

accessioned sheets in 1915, ~37,000 by 1952, 177,000 in the 1980s, and 212,000 in 1990. At present, the

collection includes more than 222,000 accessioned specimens. Many of the largest donated collections of

vascular plants have been from former directors of the UBC herbarium [J.W. Eastham (UBC database col-

lecting years: 1948-1965; ~10,000 specimens from various families, all from B.C., with an emphasis on

Poaceae and Cyperaceae); T.M.C. Taylor (1966-1971; ~7,000 specimens from various families, collected in

Canada, Europe, and Japan, with an emphasis on pteridophytes); K.I. Beamish (1972-1974; ~6,500 speci-

mens from various families, from Canada (B.C. and Yukon), Australia, and Tasmania, with an emphasis on

Saxifragaceae and Dodecatheon); G.B. Straley (1990-1996; «6,000 specimens, mainly from B.C. and U.S.A,

with an emphasis on cultivated species)]. Additional large collections have been provided by U.B.C. faculty

members [V.J. Krajina (1949-1975; ~10,000 specimens, mainly from B.C. and Hawaii, with an emphasis on

pteridophytes), W.B. Schofield (1964—present; 1,500 specimens, mainly from western Canada (B. C. and

Yukon) and Alaska], herbarium staff [J. Pinder-Moss (1968-1976; ~1,000 specimens from various families,

all B. C.)], local botanists [e.g., J.A. Calder (1954-1964; 2,200 specimens, mainly from B.C.)], past graduate

students [J. Pojar (1969-1975; «800 specimens, mainly from B.C.)], and herbarium demonstrators, such

as John Davidson (1916-1948; ~2,500 specimens from B.C. and Scotland). Current active collectors for

the UBC herbarium are: H. Kennedy (1969—present; 1,000 specimens, mainly from South America, with

an emphasis on Marantaceae); T. Goward (1979—present; 1,500 specimens, mainly from B.C; A. Ceska

(1978—present; 1,200 specimens, mainly from B.C., with an emphasis on Cyperaceae, Juncaceae, and aquatic

species “whose flowers measure « 1 cm in diameter" (Adolf Ceska, pers. comm., December 2006); and F.

Lomer (1988—present; 2,000 specimens, mainly from B.C., from various families).

Of the 222,000 vascular plant specimens in UBC, ~45% are from British Columbia, and 22% are from

the rest of Canada, with the Northwest Territories and Yukon Territory especially well represented. Speci-

mens from the United States make up about 1696 of the collection, with 996 from the five Pacific Coast states

(Alaska, California, Hawaii, Oregon, and Washington) and 796 from the remainder of the United States.

Hawaiian plants are especially well represented. About 1796 of the collection originates from the rest of

the world, with the largest numbers from Great Britain, Finland, China, Australia, Denmark, Japan, South

Africa, Taiwan, Russia, Greenland, and Sweden.

UBC Type Collection

Until recently, most vascular plant type specimens in UBC were integrated with the general collection. We

have searched the collection in an attempt to find all type specimens, though it is possible that additional

type specimens (not clearly labeled as such) remain in the general collection. The composition of the vascular

plant type collection largely reflects the geographic location of the herbarium and taxonomic groups of inter-

est to researchers that are or have been associated with the UBC herbarium le.g., studies of the neotropical

genus Calathea (Marantaceae) by Dr. Helen Kennedy]. We have located 89 type specimens (excluding para-

types and topotypes), including seven holotypes, one isoneotype, and 81 isotypes. The geographic regions

represented by the type specimens include 11 countries, eight states in the U.S.A (mostly western), three

Canadian provinces, and one Canadian territory.

In this paper, we present comprehensive data for all known type specimens in UBC, including family,

taxonomic name, authority, place and date of original publication, type status, country, collection locality

information, habitat, collector(s), collection number, and UBC herbarium accession number. Specimens are

listed alphabetically under their respective families. We have carefully examined all of the original proto-

logues and specimens to confirm the status of type specimens. In many instances, protologue data differ

from specimen label data. To provide maximum information, we therefore present all relevant protologue

and specimen label data; in most cases, morphological data included on labels or in protologues are not

Saarela et al., University of British Columbia Herbarium: vascular plant t i 439

vt L)

included here. To maintain consistency throughout the manuscript, we have altered the formatting of some

of the original data. Additional isotypes for several collections presented here are deposited in the United

States National Herbarium (US); we include this information with the protologue data, identified with an

asterisk (*) if US was not listed as a specimen repository in the original protologue. In an appendix, we

provide an index to collectors and collector numbers, based on data from the UBC specimens. Authority

abbreviations and publication citations follow Lawrence et al. (1968), Brummitt and Powell (1991), and

Bridson and Smith (1991). This is the first published account of the UBC vascular plant type specimens, and

it makes information on these historical collections available to the global botanical community to facilitate

taxonomic work. All type specimens have been scanned, and high resolution images are available on the

UBC herbarium webpage (http://www.botany.ubc.ca/herbarium

CATALOGUE OF THE VASCULAR PLANT TYPE SPECIMENS IN THE UBC HERBARIUM

APIACEAE

Lomatium stebbinsii Schlessman & Constance, Madrono 26:41. 1979. ebd d U.S.A. CALIFORNIA. Tuolumne Co.: 4 mi (6.4 km)

E of Long Barn on NE side of Bald Mountain, S2 T3N RITE, gravelly open vol t with Allium, Calyptridium, Arabis

Crepis, Penstemon, and Ceanothus; Pinus ponderosa association; peduncles spreading, ee mM fruits shining, heavily grazed by

rabbits, 4100 ft (1242 m), 26 May 1978, Constance 3895 (HOLOTYPE: WTU; IsotypEs: NY, UC, WS, US*). T-Specimen: CALIFORNIA. Tu-

olumne Co.: Sierra Nevada, 4 mi E of Long Barn on NE side of Bald Mountain (Sect. 2, Twns. 3N, R 17E), ca. 5600 ft elev., gravelly open

volcanic slope and ridge crest with Allium, Calyptridium, Arabis, Crepis, Penstemon and Ceanothus, Pinus ponderosa-Calocedrus association,

26 May 1978, L. Constance 3895 (isotype: UBC-V170207).

ASTERACEAE

Blennosperma bakeri Heiser, Madrono 9:103. 1947. T-Protologue: CALIFORNIA. Sonoma Co.: W outskirts of Sonoma in “hog wallow” ca.

0.25 mi S of Napa Street in field on E side of street, 2 Apr 1946, M.S. Baher 11307 (Herbarium of the University of California no. 72576;

isotypes are to be distributed widely; US*). T-Specimen: CALIFORNIA. Sonoma Co.: W outskirts of Sonoma in “hog wallow” ca. 0.25 mi

S of Napa Street in field on E side of street, 2 Apr 1946, M.S. Baker 11307 (isotype: UBC-V95320).

Helianthus nuttallii Torr. & A. Gray subsp. canadensis R.W. Long. Brittonia 18:77. 1966. T-Protologue: ManrroBA: District de Souris,

Mont Tortue, 1 mi de Mountain Side Fosse, Boivin and Alex 10245 (notorvre: DAO). T-Specimen: Manitoba: District de Souris, Mont

Tortue, 1 mi au norde de Mountain Side Fossé, 25 Aug 1952, Bernard Boivin and J.F. Alex 10245 (isotype: UBC-V155305).

Microseris laciniata (Hook.) Sch. Bip. subsp. detlingii K.L. Chambers, Sida 21:200. 2004. EProtolpgue U.S.A. OREGON. Mm Co.:

Siskiyou Pass, S side on the old highway where it joins Hwy. 5, 2.1 mi N of Hilt exit, grassy of ir pallida

brushland, in heavy clay soil on slope above road, 22 Jun 1967, K. L. Chambers 2868 (HOLOTYPE: OSC: ISOTYPES: BRIT-SMU, CAS, MO, NY,

RSA, UC, US, WS, WTU). T-Specimen: U.S.A. OREGON. Jackson Co.: Siskiyou Pass, slope to E of the old highway, where it joins the new

freeway (Hwy. 5), 2.1 mi N of the Hilts exit, R2E, T41S, Sec. 8, 3750 ft, 22 Jun 1967, K. L. Chambers 2868 (isotype: UBC-V220903).

Solidago xl dii B. Boivin, Phytologia 23:21. 1972. T-Protologue: Kleefeld, 1.5 mi au sud-est, prairie, 16 Aug 1958, Boivin, Bernard

& Perron 12942 (HoLoTYrE: DAO). T-Specimen: Manitosa: District de Provencher, Kleefeld, 1.5 mi au sud-est, prairie, ligules jaunes,

devenant blanches, 16 Aug 1958, B. Boivin, J.-P. Bernard, and J.M. Perron 12942 (isotype UBC-V155279).

BRASSICACEAE

Cardamine oligosperma Nutt. var. lucens G.S. Torr., Rhodora 17:157. 1915. T-Protologue: Klickitat Co.: damp places, common at

Bingen, Suksdorf 7452 (HoLotyre: GH). T-Specimen: WasuiNcrON. Klickitat Co.: Bingen, near my house, 10, 12 Apr 1912, Wilhelm N.

Suksdorf 7452 (sotyPe: UBC-V150824).

CARYOPHYLLACEAE

Silene andersonii Clokey, Bull. S. Calif. Acad. Sci. 38:2. 1939. T-Protologue: NevaDa. Clark Co.: Charleston Mountains, on a steep, N

slope in Lovell Canyon, associated with Cercocarpus lepidofolius Nutt. and Artemesia tridentata Nutt., 2,600 m, 9 Aug 1937, Clokey 7514

(HoLotyPE: Clokey Herbarium; isotypes: CAS, US*). T-Specimen: Nevara. Clark Co.: Levell [sic] Canyon, steep, N, gravelly slope with

Artemesia tridentata Nutt and Cercocarpus ledifolius Nutt., 2600 m, 9 Aug 1937, LW. Clokey 7514 (isotype: UBC-V111762).

CHENOPODIACEAE

Corispermum hookeri Mosyakin, Novon 5:349. 1995. T-Protologue: CANADA. SaskaTCHEWAN: District de Moose Jaw, palier sablonneux

au pied du Coteau du Missouri, 5-6 mi à l'est de Mortlach, dune éventrée, 11 Sep 1960, B. Boivin & G.F. Ledingham 14079 (HOLOTYPE:

DAO; IsotyPes: NY, TRT, UBC). T-Specimen: CANADA. SaskaTCHEWAN: District de Moose Jaw, palier sablonneux au pied du Coteau du

Missouri, 5-6 mi à l'est de Mortlach, dune éventrée, 11 Sep 1960, B. Boivin and G.F. Ledingham 14079 (isotype: UBC-V155271).

Corispermum hookeri Mosyakin var. pseudodeclinatum Mosyakin, Novon 5:350. 1995. T-Protologue: CANADA. BRITISH COLUMBIA:

beside Burnaby Lake, sand dune, 3 Oct 1965, I. Bayly 83 (HoLotYre: UBC). T-Specimen: CANADA. BRITISH COLUMBIA: beside Burnaby

Lake, sand dune, 3 Oct 1965, I. Bayly 83 (HoLoTYPE: UBC-V190492).

440 Journal of the Botanical R h Institute of Texas 1(1)

CYPERACEAE

Carex raymondii Calder, Rhodora 54:246. 1952. T-Protologue: Manitosa: Gillam, moist soil along margins of wood-road, Schofield

1270 (HoLotTyPE: DAO). T-Specimen: CANADA. Manitosa: Churchill District, Gillam, 27 Jul 1950, W.B. Schofield 1270 (isotype: UBC-

V40731).

Fimbristylis spathacea Roth var. depauperata T. Koyama, J. Jap. Bot. 30:1. 1955. T-Protologue: Honshu, cóté d'Ichinomiya dans la

prov. de Kadzusa, T. Koyama 11120 (HOLOTYPE: TSM). T-Specimen: JAPAN: Honshu, Kadzusa, Ichinomiya coast, scattered in stable dune

sands sheltered by Pinus thunbergii, 11 Sep 1954, T. Koyama 11120 (isotype: UBC-V102843).

DRYOPTERIDACEAE

Woodsia phillipsii Windham, Contr. Univ. Michigan Herb. 19:50. 1993. T-Protologue: U.S.A. Arizona. Cochise Co.: Rucker Canyon,

Chiricahua Mts, canyon sides in pine woods, 6500 ft, 7 Oct 1945, Phillips 2854 (HoLotype: GH; isotypes: ARIZ, ASC, UBC, US). T-Speci-

men: Arizona. Cochise Co.: Rucker Canyon, Chiricahua Mts, 6500 ft, 7 Oct 1945, Walter S. Phillips 2854 (isotype: UBC-V5012).

EQUISETACEAE

Equisetum variegatum Schleich. var. nelsonii A.A. Eaton, Fern Bull. 12:41. 1904. T-Protologue: ILLINOIS: E Chicago, along a canal,

N.L.T. Nelson s.n. (HOLOTYPE: not specified; Isotype: US*). T-Specimen: ILLINOIS: east Chicago, Jun 1901, N.L.T. Nelson 85 (isotype: UBC-

V3681).

FABACEAE

Oxytropis campestris DC. var. wanapum Joyal, Great Basin Naturalist 50:373. 1990. T-Protologue: U.S.A. WASHINGTON. Grant Co.:

Saddle Mountain, above Lower Crab Creek and E of Beverly, T15N, R24E, S2, N1/2, elev. ca. 550 m, NNE aspect at crest of ridge, in

sandy (volcanic ash) soils above steep basalt talus, 25 May 1987 (flower and early fruit), Joyal 1264 (HoLotYrE: US; isotypes: BRY, CAN,

CAS, ISC, K, MO, MONTU, NY, OSC, S, UBC, WS, WTU). T-Specimen: WasuiNGTON. Grant Co.: Saddle Mountain, above Lower Crab

Creek, sandy ridge above steep N-facing basalt talus, T15N, R24E, Sec 2 NY, ca. 600 m, common on sandy (volcanic ash) soils, 25 May

1987, Elaine Joyal 1264 (isorvee: UBC-V201206).

Oxytropis sordida (Willd.) Persoon subsp. murrayi Jurtsev, Arctic Fl. U.S.S.R 9:179. 1986. T-Protologue: Yukow: St. Elias Mts.,

Observation Mt. and vicinity, at terminus of Kaskawulsh Glacier, D. F. and B. M. Murray 522, 1966 (HoLoTYPE: LE; isotypes: ALA, BRY).

T-Specimen: CANADA. Yukon TERR.: St. Elias Mts., Observation Mt. and vic., at terminus, Kaskawulsh Glacier, ridge crest, 5500-7000

ft. 60? 40' N, 138? 44' W., 10 Jul 1966, David F. and Barbara M. Murray 522 (isotype: UBC-V173554).

GESNERIACEAE

Cyrtandra campaniformis St. John, Bernice P. Bishop Mus. Bull. 229:54. 1966. T-Protologue: Oahu, Laie Trail, Kahawainui Gulch,

in moist Metrosideros forest, 1,250 ft, 25 Mar 1956, H. St. John 25960 (HoLotyre: BISH). T-Specimen: U.S.A. Hawaiian Islands, Laie Trail,

Kahawainui Gulch, 381 m, in moist Metrosideros forest, 25 Mar 1956, St. John 25960 (isoTYPE: UBC-V38956).

Cyrtandra kailuaensis St. John, Bernice P. Bishop Mus. Bull. 229:172. 1966. T-Protologue: Oahu, Kailua, Fourth South Fork of the South

Fork of Kahanaiki Stream, open scrub of guava, Perrotettia, kukui, Rubus, 800 ft, 18 Sep 1955, G. Pearsall 2 (HoLotype: BISH). T-Specimen:

U.S.A. HAWAIIAN IsLaNDs: Kailua, Fourth South Fork of Kahanaiki Stream, 244 m, 18 Sep 1950, Pearsall 2 (isotype: UBC-V45207).

Note.—The collection date on the UBC specimen is incorrectly listed as 18 Sep 1950.

Cyrtandra pearsallii St. John, Bernice P. Bishop Mus. Bull. 229:269. 1966. T-Protologue: Oahu, Kailua, fourth south fork of the South

Fork of Kahaniki Stream, 800 ft, open scrub of guava, Clermontia, Rubus, at base of large rock, 18 Sep 1955, G. Pearsall 4 (HoLoTYPE: BISH;

ISOTYPE: US*). T-Specimen: U.S.A. Hawaiian IstANDs: Fourth South Fork of the South Fork of Kahanaiki Stream, Kailua, open scrub of

guava, Clermontia, Rubus, at base of large rock, 244 m, 18 Sep 1950, Pearsall 4 (isorvee: UBC-V45209).

Note.—The collection date on the UBC specimen is incorrectly listed as 18 Sept 1950.

JUNCACEAE

Juncus laccatus Zika, Preslia 74:261. 2002. T-Protologue: U.S.A. WasHiNcTon. Clallam Co.: low W ground, S side of Route 101 near

Dry Creek and Dry Creek Road, 6 air km SSE of Angeles Point, Olympic Peninsula, 140 m, 48?06'N, 123°31'W, 27 Sep 2001, P.F. Zika

16611 (HOLOTYPE: WTU; isotypes: CAN, GH, MICH, MO, NY, OSC, PRA, UBC, UC, US). T-Specimen: Wasuincton. Clallam Co.: S side

of Route 101 near Dry Creek and Dry Creek road, 6 air km SSE of Angeles Point, Olympic Peninsula, low wet ground, full sun, with

Juncus hesperius, J. ensifolius, Lotus corniculatus, Alnus rubra, 140 m, T30N R8W S22, 48%6' N, 123?31' W, 27 Sep 2001, P.F. Zika 16611

(IsoTYPE: UBC-V219821).

LAURACEAE

Licaria applanata van der Werff, Novon 4:65. 1994. T-Protologue: ECUADOR: Pichincha, Reserva Río Guajalito, near Chiriboga,

along old road Quito-Sto. Domingo, 1,850 m, van der Werff et al. 12194 (HOLOTYPE: MO; Isotypes: AAU, GB, HBG, K, NY, P, QCNE, QRS,

US). T-Specimen: ECUADOR: Pichincha, Reserva Río Guajalito, near Chiriboga, along old road Quito-St. Domingo, montane forest,

Lauraceae collected in pature, 1850 m, 5 Jul 1991, H. van der Werff, B. Gray, G. Tipas, and J. Campaña 12194 (sotyPe: UBC-V211491).

Ocotea rugosa van der Werff, Novon 4:70. 1994. T-Protologue: ECUADOR: Bolivar, in small patches of disturbed cloud forest along

first 15 km of road Chillanes-El Tambo, elev. 2,100 m, van der Werff et al. 12429 (male fl) (HoLotypE: MO; tsotyres: AAU, GB, NY, QCNE,

QRS, US). T-Specimen: ECUADOR: Bolivar, small patches in disturbed cloud forest, rather dry, along first 15 km of road Chillanes-El

Tambo, 2400 m, 18 Jul 1991, H. van der Werff, B. Gray, and G. Tipas 12429 (Isotype: UBC-V211489).

Saarela et al., University of British Columbia Herbarium: vascular plant t i 441

vt L)

LILIACEAE

Erythronium quinaultense G.A. Allen, Syst. Bot. 26:269. 2001. T-Protologue: U.S.A. WasHINGTON. Grays Harbor Co.: Higley Peak, W

side of summit, openings in coniferous forest, T23°N R10°W sect. 1, elev. 800 m, 24 May 1996, Allen 9603 (HoLotyPE: UVIC; ISOTYPES:

WTU, OSC). T-Specimen: WasHINGTON: Grays Harbor ee See National Forest, W side of Higley Peak along FS road #2190, S of

short trail to it, rocky ledges and openings in 1 adjacent road cutbacks on W facing slope, 2620 ft, T23N RLOW

sect. 1, 24 May 1996, G.A. Allen 9603 "rin ‘UBC -V218119).

MALVACEAE

Hibiscus newhousei Roe, Pacific Sci. 15:22. 1961. T-Protologue: Kauai, Moloaa Forest Reserve, ca. 500 ft altitude, 10 Nov 1958, LE.

Lane 44 (HOLOTYPE: not specified). T-Specimen: HAWAII: Kauai Island, Moloaa Forest Reserve, medium forest, “headwall,” 150 m, 22.2N

159.6W, 10 Nov 1958, LE. Lane 44 (isotype: UBC-V183389a).

MARANTACEAE

Calathea amazonica H. Kenn., Selbyana 15:63. 1994. T-Protologue: PERU. Dept. AMAZONAS: Rio Cenepa vicinity of Huampami, ca.

5 km E of Chávez Valdivia, en bosque secundario, 200-250 m, ca. 04?30'S, 78°30'W, 1978, E. Ancuash 1142 (HOLOTYPE: MO; ISOTYPE:

UBC). T-Specimen: PERU. Dept. AMAZONAS: Río Cenepa, vicinity of Huampami, ca. 5 km E of Chávez Valdivia, ca. 78°30' W, 4? 30'W,

200-250 m; gruesa hierba 30 cm, hojas pubescentes, flores blancas, en bosque secundario, no tiene usa; 1978, Ernesto Ancuash 1142

(isoTYPE: UBC-V208550).

Calathea anderssonii H. Kenn., Canad. J. Bot 63:1145. 1985. T-Protologue: ECUADOR: MONA 15 km W of Tenaalong the Tena-Pamo

road (road to Salcedo), along stream bank, 550 m, 19 Jul 1982, Kennedy, B L E: NY; ISOTYPES: AAU, BH, GB, QCA,

SEL, UBC). T-Specimen: ECUADOR. Prov. Naro: 15 km W of Tena along the Tena-Pano n" TM to Salcedo), along stream bank, 550

m, 19 Jul 1982, Helen Kennedy, Libby Besse, and Ray Baker 4383 (isotype: UBC-V218518, 1 of 4; UBC-V183369, 2 of 4, UBC-V218519, 3

of 4; UBC-V218520, 4 of 4)

Calathea annae H. Kenn. & Marcelo, Phytologia 82:96. 1997. T-Protologue: Cultivated at Sítio Roberto Burle Marx, Barra de Guara-

tiba, Edo. Rio de Janeiro, Brazil, accession number MAR-1, from Brazil, Edo, unknown, 11 Jan 1991, H. Kennedy and M. de F. Gomes

de Souza 4696 (HOLOTYPE: RB; isoTYPES: K, RB, Sitio RBM, UBC). T-Specimen: BRASIL: cultivado no Sitio Roberto Burle Marx, Barra de

Guaratiba, Edo. Rio de Janeiro, 11 Jan 1991, Helen Kennedy and M. de Fátima Gomes de Souza 4696 (Isotype: UBC-V207963, 1 of 2; UBC-

V207964, 2 of 2).

Note.—The UBC specimen is labeled Calathea annae H. Kenn. and Braga, but the taxon was described as Calathea annae H. Kenn

& Marcelo.

Calathea anulque H. Kenn., Fl. Ecuador 32:38. 1988. T-Protologue: ECUADOR: Carchi, environs of Maldonado, 1450-1650 m, 2 Jun

1978, Madison, Plowman, Kennedy and Besse 4949 (HOLOTYPE: SEL; isotypes: AAU, F, QCA, UBC). T-Specimen: ECUADOR. Prov. CARCHI:

environs of Maldonado, wet montane forest, elev. 1450-1650 m, 2 Jun 1978, M.T. Madison, T.C. Plowman, H. Kennedy, and L. Besse 4949

(isotype: UBC-V190301, 1 of 3; UBC-V190299, 2 of 3; UBC-V190300, 3 of 3).

Calathea attenuata H. Kenn., Nord. J. Bot. 6:146. 1986. T-Protologue: COLOMBIA: Amazonas, Leticia, 8 km out of Leticia along the

road past airport, near Finca Vega, rain forest, 7 Feb 1972, Kennedy and Andrews 1343 (HOLOTYPE: MO; isotypes: AAU (sterile), COL, GB,

NY, UBC). T-Specimen: COLOMBIA: Amazonas, Leticia, ca. 5 km out of Leticia on the road past the airport, disturbed rain forest, 7

Feb 1972, Helen Kennedy and Robin Andrews 1343 (isotype: UBC-V184199).

Calathea caquetensis S. Suárez & Galeano, Caldasia 9:12. 2000. T-Protologue: COLOMBIA: Amazonas, río Yari, margen oriental,

cerca asu desembocadura en el río Caquetá, 0? 34'S, 72? 20' W, ca. 200 m, 13 Jun 1991, G. Galeano et al. 2829 (HoLotTypE: COAH; ISOTYPES:

COL, UBC). T-Specimen: COLOMBIA. DEPARTMENTO DE AMAZONAS: Río Yari, margen izquierdo bajando, cerca a su desembocadura en el

Río Caquetá, 13 Jun 1991, G. Galeano, X. Martínez, and S. Suárez 2829 (isotype: UBC-V215100).

Calathea clivorum H. Kenn., Canad. J. Bot. 63:1147. 1985. T-Protologue: ECUADOR: Napo, 20 km W of Tena along the Tena-Pano

road from Tena (road to Salcedo), 550 m, 19 Jul 1982, Kennedy, Besse and Baker 4368 (HOLOTYPE: K; isotypes: AAU, GB, M, MO, QCA, SEL,

U, UBC, US). T-Specimen: ECUADOR. Prov. Naro: 20 km W of Tena along the Tena-Pano road from Tena (road to Salcedo), growing on

a nearly vertical cliff face, 550 m, 19 Jul 1982, Helen Kennedy, Libby Besse and Ray Baker 4368 (isorvree: UBC-V183370).

Calathea compacta S. Suárez & Galeano, Caldasia 22:12. 2000. T-Protologue: COLOMBIA: Amazonas, río Caquetá, margen sur,

Puerto Asai, frente a Dos Islas, 0°55' S, 71?38' W, ca. 200 m, 4 Jun 1991, G. Galeano et al. 2711 (notorvee: COAH; isotype: COL, UBC).

T-Specimen: COLOMBIA. DEPARTMENTO DE AMAZONAs: Río Caquetá, margen sur, Puerto Asai frent a Dos Isla, en cercania de la casa de

Alfonso Rodríguez, G. Galeano, X. Martínez, and S. Suárez 2711 (isorvre: UBC-V215101).

Calathea contrafenestra H. Kenn., Canad. J. Bot. 62:18. 1984. T-Protologue: Cultivated at Marie Selby Botanical Garden, Sarasota,

Florida, from rhizomes collected in Limnococha, Ecuador (M.T. Madison, T.C. Plowman, and L. Besse 5400), 18 Jun 1982, H. Luger [sic]

800 (HoLoTYPE: SEL; isotypes: BH, US). T-Specimen: Cultivated, Marie Selby Botanical Garden, Sarasota, Florida, from rhizomes col-

lected in Ecuador, Limoncocha, Napo, Ecuador (Madison, Plowman, and Besse 5400), 18 Jun 1982, Luther 800 (isotype: UBC-V184263, 1

of 2; UBC-V184262, 2 of 2).

Calathea curaraya H. Kenn., Fl. Ecuador 32:128. 1988. T-Protologue: ECUADOR: Napo, road (under construction) Coca-Cononaco,

c. 30 km S of Río Tiputini, 200 m, 23 Jan 1982, Harling, Bohlin, Lindström & Roth 19833 (HOLOTYPE: GB; isorvee: UBC). T-Specimen: EC-

UADOR: Coca, Coca-Cononaco Road (in construction), ca. 30 km S of Rio Tiputini, 200 m, 23 Jan 1982, G. Harling, J.-E. Bohlin, Marie

Lindstrom, and Suzanne Roth 19833 (isotype: UBC-V202966).

Qnm

442 Journal of the Botanical R h Institute of Texas 1(1)

Calathea ecuadoriana H. Kenn., Canad. J. Bot. 62:15. 1984. T-Protologue: U.S.A. Hawai: Honolulu, cultivated at Lyon Arboretum,

1 Jun 1982, H. Kennedy 4224 (HoLotype: US; isotypes: BH, CAN, HLA, K, M). T-Specimen: U.S.A. Hawai: Honolulu, Lyon Arboretum, 1

Jun 1982, Helen Kennedy 4224 (isotype: UBC-V220043).

Calathea fatimae H. Kenn., Phytologia 82:94. 1997. T-Protologue: BRAZIL: Cultivated at Sitio Roberto Burle Marx, Barra de Guaratib

Munic. Rio de Janeiro, Edo. Rio de Janeiro, Brazil, accession number MAR-101, from rhizomes collected by Fátima Gomes de Souza from

Morro do Coco, ca. 40 km N of the city of Campos, Munic do Campos, Edo. De Janeiro, Brazil, flowered in cultivation 11 Jan 1991, H.

Kennedy and M. de F. Gomes de Souza 4700 (HOLOTYPE: RB; Isotypes: K, Sitio RBM, UBC). T-Specimen: Cultivado no Sitio Roberto Burle

Marx, Barra de Guaratiba, Edo. Rio do Janeiro, Brasil, rhizomes collected by M. F. Gomez de Souza from Morro do Coco, Munic do

Campos, Edo. Rio de Janeiro, Brasil, 19 Jan 1991, Helen Kennedy and M. de Fatima Gomes de Souza 4700 (isotype: UBC-V207965, 1 of 2;

UBC-V218521, 2 of 2).

Calathea fucata H. Kenn., Fl. Ecuador 32:133. 1988. T-Protologue: ECUADOR: Napo, Río San Miguel opposite San Miguel, 4 Feb

1971, Kennedy 808 (HOLOTYPE: SEL; isotypes: DUKE, QCA, UBC). T-Specimen: ECUADOR. Naro Prov.: Río San Miguel, across river from

San Miguel, Putumayo, Colombia, 4 Feb 1971, Helen Kennedy 808 (isotYrE: UBC-V194073).

Calathea gandersii H. Kenn., Fl. Ecuador 32:110. 1988. T-Protologue: ECUADOR: Napo, road from Tena to Pano, trail through primary

forest from road's end to the sawmill, 760 m, Davis 422 (HoLoTyPE: UBC). T-Specimen: ECUADOR. Provincio Naro: road from Tena to

Pano, trail through primary forest from roads end to the sawmill, 759 m, 14 Dec 1976, E. W. Davis 422 (HoLoTyPE: UBC-V193731).

Calathea gloriana H. Kenn., Selbyana 18:35. 1997. T-Protologue: COSTA RICA. Prov. HEREDIA: Estación Biológica La Selva, Lindero

Sur por las cabeceras de la Quebrada Sura, bosque secundario viejo, ca. 100 m, 19 Jul 1990, H. Kennedy and B. Hammel 4545 (HOLOTYPE:

CR; isotypes: BM, MO, U, UBC, US). T-Specimen: COSTA RICA. Prov. HEREDIA: Cantón d piquí, estación Biológica La Selva, Lindero

Sur por las cabeceras de La Quebrada Sura, bosque secondario, viejo, 100 m, 10?25' 30" N, 84°01'30" W, 19 Jul 1990, Helen Kennedy

and Barry Hammel 4545 (isotype: UBC-V211858)

Calathea grazielae H. Kenn. & Marcelo, Phytologia 82:101. 1997. T-Protologue: BRAZIL: cultivated at Sitio Roberto Burle Marx, Barra

de Guaratiba, Munic. Rio de Janeiro, Edo. Rio de Janeir, Brazil, from rhizomes collected in Brazil, Edo. unknown, flowered in cultiva-

tion 19 Jan 1991, H. Kennedy and M. de. F. Gomes de Souza 4699 (HOLOTYPE: RB; IsoTYPEs: K, MO, Sitio RBM, UBC). T- Specimen: BRAZIL:

cultivado no Sítio Roberto Burle Marx, Barra de Guaratiba, Edo. Rio de Janeiro, 19 Jan 1991, Helen Kennedy and M. de Fátima Gomes de

Souza 4699 (isOTYPE: UBC-V208250, 1 of 2; UBC-V218522, 2 of 2).

Note.—The UBC specimen is labeled Calathea grazielae H. Kenn. and J. M. Braga (the latter authority written by hand after the

label was printed). The taxon was described as Calathea grazielae H. Kenn. & Marcelo.

Calathea hylaeanthoides H. Kenn., Canad. J. Bot. 75:1356. 1997. T-Protologue: COSTA RICA. PUNTARENAS Prov.: Cantón de Osa, Osa

Peninsula, Reserva Forestal Golfo Dulce, in Rancho Quemado Valley along Quebrada Quebradón and the Río Riyito, 200 m, 8?40' N,

83°34' W, 11 Sep 1990, H. Kennedy, B. Hammel, and J. Solomon 4664 (HOLOTYPE: CR; Isotypes: BM, COL, DUKE, F, G, GB, INB, K, MA,

MICH, MO, P, U, UBC, US). T-Specimen: COSTA RICA. PUNTARENAS Prov.: Cantón de Osa, Osa Peninsula, Reserva Forestal Gulfo Dulce,

in Rancho Quemado Valley along Quebrada Quebradón and the Rio Riyito, 200 m, 8°40'N, 83°34'W, 11 Sep 1990, Helen Kennedy, Barry

Hammel, and James Solomon 4664 (isoTYPE: UBC-V211818, 1 of 2; UBC-V211817, 2 of 2).

Calathea incompta H. Kenn., Canad. J. Bot. 75:1361. 1997. T-Protologue: COSTA RICA. PUNTARENAS Prov.: Cantón de Osa, Osa

Peninsula, 2.5 mi SW of Rincón, by Río Agua Buena, above the airfield, tropical wet forest, 15-30 m, 8?32' N, 83?29' W, 4 Sep 1971, H.

Kennedy 1139 (HoLoTvYPE: UBC; isotypes: CR, K, MO, US). T-Specimen: COSTA RICA. PUNTARENAS Prov.: Cantón de Osa, Osa Peninsula,

2.5 mi SW of Rincón, by Río Agua Buena, above the airfield, 15-30 m, 8?32' N, 83?29' W, tropical wet forest, 4 Sep 1971, Helen Kennedy

1139 (HoLotyPE: UBC-V204768, 1 of 3; V218523, 2 of 3; V218524, 3 of 3).

Calathea lagoagriana H. Kenn., Nord. J. Bot. 6:148. 1986. T-Protologue: ECUADOR. Napo Prov.: 33 km S of Rio Aguarico on the road

from Lago Agrio to Coca, wet tropical forest, 2 Jul 1982, Kennedy, Besse, and Baker 4288 (HOLOTYPE: NY; IsoTyPes: GB, K, MO, SEL, UBC,

WIS). T-Specimen: ECUADOR. Naro Prov.: 33 km S of Río Aguarico on the road from Lago Agrio to Coca, wet tropical forest, ca. 350

m, 2 Jul 1982, Helen Kennedy, Libby Besse, and Ray Baker 4288 (isorvee: UBC-V189968).

Calathea lanicaulis H. Kenn., Canad. J. Bot. 63:1143. 1985. T-Protologue: ECUADOR: Napo, 5.6 km along the Tena-Pano road from

Tena, disturbed tropical rain forest, ca. 500 m, 17 Jul 1982, Kennedy, Besse and Baker 4373 (HoLotyPE: NY; isotypes: AAU, BH, GB, K,

MO, QCA, SEL, U, UBC). T-Specimen: ECUADOR. Prov. Naro: 5.6 km along the Tena-Pano road from Tena, disturbed tropical rain

forest, ca. 500 m, 17 Jul 1982, Helen Kennedy, Libby Besse, and Ray Baker 4373 (isotype: UBC-V218525, 1 of 4; UBC-V183368, 2 of 4;

UBC-V218526, 3 of 4; UBC-V218527, 4 of 4).

Calathea latrinotecta H. Kenn., Fl. Ecuador 32:42. 1988. T-Protologue: ECUADOR: Carchi, Penas Blancas, 20 km below Maldonado on

Río San Juan, 900-1000 m, Madison, Plowman, Kennedy, and Besse et al. 4646 (HOLOTYPE: SEL; isotypes: AAU, F, QCA, UBC). T-Specimen:

ECUADOR. Prov. Carcht: Pénas Blancas, 20 km below Maldonado on the Río San Juan, wet montane forest, elev. 900-1000 m, 27 May

1978, M.T. Madison, T.C. Plowman, H. Kennedy, and L. Besse 4646 (isotype: UBC-V 218579, 1 of 4; UBC-V190304, 2 of 4; UBC-V218578,

3 of 4; UBC-V218580, 4 of 4).

Calathea liesneri H. Kenn., Novon 3:49. 1995. T-Protologue: VENEZUALA: Territorio Federal Amazonas, Depto. Atabapo, in saddle

between Cerro Duida and Cerro Marahuaca near base of Cerro Duida, medium height forest, 1,000 m, 03?34'N 65?32'W, 25 Oct 1988,

Liesner 25333 (HoLoTYPE: MO; Isotypes: K, UBC, VEN). T-Specimen: VENEZUALA: Amazonas, Atabapo, in saddle between Duida and

Marahuaca near base of Duida, 1000 m, 03°34'N 65°32'W, 25 Oct 1988, Ronald Liesner 25333 (isotype: UBC-V218528).

Calathea maasiorum H. Kenn., Brittonia 47:156. 1995. T-Protologue: FRENCH GUIANA: Saúl, vicinity of Eaux Claires, Sentier

Saarela et al., University of British Columbia Herbarium: vascular plant type speci 443

Botanique, from entrance to 450 m from entrance, non-flooded moist forest, ca. 250-350 m, 03?37'N 53°12'W, 10 Feb 1993, Mori, Maas,

& Mass et al. 22909 (HoLotyPE: NY; isotypes: CAY, NY, U, UBC). T-Specimen: FRENCH GUIANA: Saul, vicinity of Eaux Claires, Sentier

Botanique, from entrance to 450 m from entrance, ca. 250-350 m, 3°37'N, 53°12'W non-flooded moist forest, 10 Feb 1993, S. Mori, C.

Gracie, T. Croat, H. Maas, P. Maas, T. Pennington, and D. Reed 22909 (isotype: UBC-V208553).

Calathea neblinensis H. Kenn., Phytologia 69:373. 1990. T-Protologue: VENEZUELA: Amazonas, Dept. Río Negro, near Cerro de

La Neblina Base Camp which is on Río Mawarinuma, in forest near stream, 140 m, 0? 50' N, 66? 10' W, 5 Feb 1984, R. L. Liesner 15662

(HOLOTYPE: MO; isotypes: F, NY). T-Specimen: VENEZUELA: Territorio Federal Amazonas, Dept. Río Negro, near Cerro de La Neblina

Base Camp which is on Río Mawarinuma, elev. 140 m, 0?50' N, 66°10' W, 5 Feb 1984, Ronald Liesner 15662 (isotype: UBC-V217000).

Calathea pallidicosta H. Kenn., Nord. J. Bot. 6:143. 1986. T-Protologue: ECUADOR. Pastaza Prov.: 3 km S from junction with Puyo-

Macas road towards Canelos (31 km total from Puyo Cemetery), disturbed tropical rain forest, 870 m, 21 Jul 1982, Kennedy, Besse and

Baker 4404 (HOLOTYPE: NY; isotypes: GB, SEL, UBC). T-Specimen: ECUADOR. Prov. Puyo: 3 km S from junction with Puyo-Macas road

towards Canelos (31 km total from Puyo Cemetery), disturbed tropical rain forest, 21 Jul 1982, Helen Kennedy, Libby Besse and Ray Baker

4404 (isotype: UBC-V189965, lof 3; V218529, 2 of 3; V218530, 3 of 3).

Calathea paucifolia H. Kenn., Fl. Ecuador 32:86. 1988. T-Protologue: ECUADOR: Pastaza, Río Pacayacu, tributary to Río Bobonaza,

SE of Canelos, 10 March 1971, Lugo 1605 (HoLoTYPE: GB). T-Specimen: ECUADOR: Pastaza, Río Pacayacu, in the vicinity of Canelos, 10

Mar 1971, Holguer Lugo S. 1605 (isotype: UBC-V 202967 [2 separate sheets]).

Calathea plurispicata H. Kenn., Fl. Ecuador 32:33. 1988. T-Protologue: ECUADOR: Napo, road in construction Coca-Curaray, 20-30

km S of Coca, c. 250 m, Harling and Andersson 11940 (HoLoTYPE: GB; isotype: UBC). T-Specimen: ECUADOR: Napo, road Coca (Puerto

Francisco de Orellana) to Curaray, 20-30 km S of Coca, primary rain forest, alt. ca. 350 m, 13 Nov 1974, G. Harling and L. Andersson

11940 (isotype: UBC-V190302, 1 of 2; UBC-V218531, 2 of 2).

Calathea poeppigiana Loesner ex Kennedy, Fl. Ecuad. 32:77. 1988. T-Protologue: ECUADOR: Pastaza, tributary to Río Curarary,

c. 21 km E of Curaray (Jesus Pitishka), temporary shallow pools in virgin rain forest, c. 200 m, 22 March 1980, Harling and Andersson

17592 (HOLOTYPE: GB; Isotype: UBC). T-Specimen: ECUADOR: Pastaza, Río Namoyacu ca. 21 km E of Curaray (Jesus Pitishka), virgin

rain forest, ca. 200 m, G. Harling and L. Anderson 17592 (isorvee: UBC-V203042).

Calathea retroflexa H. Kenn., Canad. J. Bot. 75:1357. 1997. T-Protologue: COSTA RICA. San José Province: Cantón de Pérez Zaledón,

San Isidro del General, above La Ese, 3.3 km N of kilometer marker no. 126 on CR route no. 2, ca. 1380 m, 9°26'40" N, 83?43'00" W,

15 Aug 1990, H. Kennedy 4555 (HOLOTYPE: CR; ISOTYPES: BM, F, GB, INB, K, MO, U, UBC). T-Specimen: COSTA RICA. SAN José PROVINCE:

Cantón de Pérez Zaledón, San Isidro de General, above La Ese, 3.3 km N of km marker #126 on CR route #2, ca. 1380 m, 9?26'40" N,

83?43'00 W, 15 Aug 1990, Helen Kennedy 4555 (isotype: UBC-V211816).

Calathea robinae H. Kenn., Ann. Missouri Bot. Gard. 60:419. 1973. T-Protologue: COSTA RICA. Limón PROVINCE: old forest ca. 2 mi

from La Lola on the road to Siquirres near stream bed, 26 Feb 1971, H. Kennedy, H. Andrews & H. L. Dressler 1379 (HOLOTYPE: F; ISOTYPES:

BM, COL, CR, DAV, DUKE, GH, K, MO, NY, U, US). T-Specimen: COSTA RICA: Limón, old forest ca. 2 mi from La Lola on the road to

Siquirres, near stream bed, Helen Kennedy, Robin Andrews, and Robert Dressler 1379 (isotype: UBC-V 204804).

Calathea steyermarkii H. Kenn. & Nagata, Brittonia 41:164. 1989. T-Protologue: U.S.A. HAWAII: Honolulu, cultivated at Lyon Ar-

boretum, accession # L 76.1230 (from rhizomes collected by H. Kennedy in Parque Nacional Henri Pittier, Edo. Maracay, Venezuala),

5 Apr 1984, K. Nagata 2886 (HoLotyPE: MO; Isotypes: HLA, MAY, UBC). T-Specimen: Hawaii, Honolulu, cultivated at Lyon Arboretum

accession # L-76.1230 (from rhizomes collected by H. Kennedy in Parque Nacional Henri Pittier, Edo. Maracay, Venezuala), 5 Apr 1984,

Kenneth M. Nagata 2886 (isorvee: UBC-V203398).

Calathea striata H. Kenn., Fl. Ecuador 32:60. 1988. T-Protologue: COLOMBIA: Amazonas, Leticia, 8 km out of Leticia on the road

past the airport, forest on N side of road, Finca Misiones, rain forest, 11 Feb 1971, Kennedy 813 (HoLotYPE: GB; IsoTypes: AAU, COL,

DUKE, F, U, UBC). T-Specimen: COLOMBIA: Amazonas, Leticia, 8 km out of Leticia on the road past the airport, forest on N side of

the road, Finca Misiones, rainforest, 11 Feb 1971, Helen Kennedy 813 (isotype: UBC-V204750).

Calathea tinalandia H. Kenn., Canad. J. Bot. 63:1141. 1985. T-Protologue: ECUADOR: Pinchincha, Tinalandia Hotel grounds, km

112 on the road Quito to Santo Domingo de los Colorados, tropical moist forest, ca. 700 m, 28 Jul 1982, Kennedy 4400 (HoLotyeE: NY;

ISOTYPES: AAU, GB, K, MO, QCA, SEL, U, UBC). T-Specimen: ECUADOR. Prov. PicHINcHa: Tinalandia Hotel grounds, km 112 on the

road Quito to Santo Domingo de los Colorados, tropical moist forest, 28 Jul 1982, Kennedy 4400 (isorvee: UBC-V183373).

Calathea utilis H. Kenn., Nord. J. Bot. 6:457. 1986. T-PRororocur: ECUADOR. Prov. Napo: Sardinas, 6 km from Baeza on road to Lago

Agrio, montane wet forest, 1900 m, 16 Jul 1982, Kennedy, Besse, and Baker 4363 (HoLotyPE: MO; Isotypes: AAU, GB, K, NY, QCA, S, SEL,

UBC). T-Specimen: ECUADOR. Prov. Naro: 6 km from Baeza on the road to Lago Agria, Sardinas, montane W forest, 1900 m, 16 Jul

1982, Helen Kennedy, Libby Besse, and Ray Baker 4363 (isotype: UBC-V203560, 1 of 3; UBC-V218581, 2 of 3; UBC-V218582, 3 of 3).

Monophyllanthe araracuarenses S. Suárez, Galeano & H. Kenn., Novon 11:356. 2001. T-Protologue: COLOMBIA: Amazonas, río

Caquetá, margen sur, Pena Roja, 0°39'S, 72°05'W, ca. 200 m, 9 Jun 1991, G. Galeano, X. Martínez & S. Suárez 2767 (HOLOTYPE: COAH;

ISOTYPES: COL, MO, UBC). T-Specimen: COLOMBIA. DEPARTAMENTO DE Amazona: Pena Roja trocha al monte, en la margen sur del rio

Caquetá, 9 Jun 1991, G. Galeano, X. Martínez, and S. Suárez 2767 (isotype: UBC-V218577).

MELANTHIACEAE

Trillium ovatum Pursh forma hibbersonii T.M.C. Taylor & Szczaw., Syesis 7:250. 1974. T-Protologue: British CoLumBla: W coast

of Vancouver Island, near Boat Basin, Hesquiat Harbour, 49°20'N, 136°30'W, alt. 2000 ft, 1938, J. Hibbersonii s.n. (HoLOTYPE: UBC).

T-Specimen: CANADA. BritisH COLUMBIA: near Boat Basin, Hesquiat Harbour, W coast of Vancouver Island, 1938, J. Hibberson s.n.

(HOLOTYPE: UBC-V73131).

444 Journal of the Botanical R h Institute of Texas 1(1)

POACEAE

Achnatherum wallowaensis J.R. Maze & K.A. Robson, Madrono 43:401. 1996. T-Protologue: U.S.A. OREGON. Wallowa Co.: Wallowa-

Whitman National Forest, ca. 34 km N of Enterprise, near Boner Gulch along Forest Service Road 46, 45? 43' 41.16" N, 117? 08' 10.32"

W (SW !4 of SE !4, section 24, T3 N, R 45 E), 1481 m, 26 Jun 1993, J. and E. Maze, K.A. Robson & T. Henn 1007 (HOLOTYPE: US; ISOTYPES:

COLO, DAV, ID, NMC, OCS, UBC, UC, UTC, WTU]. T-Specimen: U.S.A. OrecoN. Wallowa Co.: Wallowa-Whitman National Forest,

near Boner Gulch along Forest Service Road 46, ca. 34 km N of mepeg SW 14 of SE 14, section 24, T 3 N, R 45 E, pos les in

Ri o n

E TELE

, Epilobium sp., Poa sandbergi,

shallow rocky soil at 1279 m with Bromus tectorum, B. japonicus

Agropyron spicatum, L ti Sed t talam, Allium tolmei, Eriophyllum lanatum, Grindellia sp., Achillea millefolium, 26

Jun 1993, J. and E. Maze, K.A. Robson, m T. Henn 1007 e UBC-V209875).

Bromus ayacuchensis Saarela & P.M. Peterson, Sida 22:919. 2006. T-Protologue: PERU. DEPARTAMENTO ÁYACUCHO: Provincia Lucanas,

12 km E of Puquio on road towards Cuzco, (14%41'19.2 S Lat, 74°04'28.3 W Long), 3730 m, among large boulders with Berberis (Ber-

beridaceae), above meadow with small creek, 11 Mar 2002, P.M. Peterson 16452 & N.F. Refulio-Rodriquez (HOLOTYPE: US; IsOTYPES: K, MO,

UBC, USM). T-Specimen: PERU. DEPARTAMENTO AYACUCHO: Provincia Lucanas, 12 km E of Puquio on road towards Cuzco, (14?41'19.2 S

Lat, 74?04'28.3 W Long), 3730 m, among large boulders with Berberis (Berberidaceae), above meadow with small creek, 11 Mar 2002,

P.M. Peterson 16452 & N.F. Refulio-Rodriquez (isotype: UBC-V222906

Elymus ambiguus Vasey € Scribn. var. salmonis C.L. Hitchc., Vasc. Pl. Pacific NorthW. 1:558. 1969. T-Protologue: [DAHo: Custer Co.:

shale cliff 9 mi S of Challis, along E side of Salmon R., 31 May 1962, Hitchcock & Muhlich 22305 (HoLotYrE: UW). T-Specimen: IDAHO.

Custer Co.: shale cliffs 9 mi S of Challis, along E side of Salmon River, 31 May 1962, C.L. Hitchcock and C.V. Muhlick 22305 (isoTYPE:

UBC-V95381).

Oryzopsis swallenii C.L. Hitchc. & Spellenb., Brittonia 20:164. 1968. T-Protologue: [pAHo. Clark Co.: open adv T just N of

Birch Creek, 2 mi NW of Blue Dome, along Highway 28, near the Lemhi Co. line, ca. 5600 ft, 7 Jul 1965, C.L. Hitchcock 2

WTU; Isotype: US). T-Specimen: IpaHo. Clark Co.: open sagebrush slope N of Birch Creek, 3 mi NW of Blue Dome, elev. ca. 5600 n

on decomposed limestone, semi-barren gentle slope with dwarfed sagebrush and very dwarfed Oryzopsis hymenoides; plants tufted, the

herbarium specimens representing ca. 1/10-1/2 of the whole plant, 7 Jul 1965, C.L. Hitchcock 23868 (isorYrE: UBC-V116845)

Paspalum eitenii Swallen, Phytologia 14:385. 1967. T-Protologue: BRAZIL: Maranháo, Municipio de Loréto, “Ilha de Balsas" region,

between Rios Balsas and Parnaíba, collected in ravine of dry brook in extensive “caatinga” forest, 6 Apr 1962, George Eiten and Liene T.

Eiten 4091 (uoLorvre: US). T-Specimen: BRAZIL: Maranhao, Municipio de Loréto, “Ilha de Balsas” region, between the Rios Balsas and

Parnaiba, ca. 30 km S of Loreto, ca. 0.5 km N of main house of Faz. São Raimundo on trail to Fazenda Santa Rita, ca. 7°19'S, 45°7-8'W,

in ravine of now dry brook in extensive “caatinga” forest, soil light brown fine sand with clay, 200-300 m, 6 Apr 1962, George Eiten and

Liene T. Eiten 4091 (isotype: UBC-V117490).

POLYGONACEAE

Dedeckera eurekensis Reveal & J.T. Howell, Brittonia 28:246. 1976. T-Protologue: UNITED STATES. CALIFORNIA. Inyo Co.: Last

Chance Range, in a rocky canyon ca. 3 airline mi (4.8 km) SE of Eureka Valley sand dunes and 3.5 airline mi (5.6 km) NW of Marble

VABM 7559, in T10S, R40E, on steep limestone, N-facing, rocky slopes in the canyon, associated with Atriplex, Eriogonum and Prunus at

ca. 4000 ft (1200 m) elev., 29 Jul 1975, J. L. Reveal, M. C. DeDecker & P. W. DeDecker 3909 (HOLoTYPE: US; isotypes: 35 to be distributed).

T-Specimen: CALIFORNIA, Inyo Co.: Last Chance Range, along the jeep corridor between Saline Valley and Eureka Valley, ca. 3 airline

mi SE of Eureka Valley sand dunes and 3.5 airline mi NW of Marble VABM 7559, in T.10S., R.40E., on steep limestone, N-facing rocky

slopes associated with Atriplex, Buddleja, Prunus and Eriogonum, at ca. 4000 ft elev., 29 Jul 1975, James L. Reveal, Mary C. DeDecher, and

Paul. W. DeDecker 3909 (isotype: UBC-V173364).

Eriogonum natum Reveal, Great Basin Naturalist 35:363. 1975. T-Protologue: Uran. Millard Co.: along U.S. Highway 50-6, 46.2 mi

E of the Nevada state line and ca. 43 mi W of Delta, on low white alkaline clay outcrops 50-300 meters N of the highway, ca. 0.2 mi E of

the dirt road junction to the Antelope Spring-Black Hill Well roads, NNW of Sevier Lake, 13 Aug 1975, Reveal & Reveal 3924 (HOLOTYPE:

US: sotypes: ARIZ, ASU, BRY, CAS, COLO, GH, ISC, K, MARY, MO, NY, OKL, OSC, PH, RM, RSA, SMU, TEX, UC, UTC, WTU). T-

Specimen: Utan. Millard Co.: along U.S. Highway 50-6, 46.2 mi E of the Nevada State line and ca. 43 mi W of Delta, ca. 0.2 mi E of

the dirt road junction of Antelope Spring and Black Hill Well with the highway NNW of Sevier Lake, 13 Aug 1975, James L. Reveal and

Mark L. Reveal 3924 (isotype: UBC-V173366).

Eriogonum umbellatum Torr. var. furcosum Reveal, Great Basin Naturalist 45:278. 1985. T-Protologue: CALIFORNIA. El Dorado

Co.: along California Highway 89, 2.2 mi S of U.S. Highway 50, on sandy granitic soil, associated with Arctostaphylos, Artemisia, and

juniper-pinyon, 23 Aug 1975, Reveal 3971 (HoLoTYPE: US; isotypes: ARIZ, BRY, CAS, DUKE, F, GH, K, MARY, MEXU, MICH, MO, NY,

OKL, RENO, RM, RSA, TEX, UC, UTC and elsewhere). T-Specimen: CALIFORNIA. El Dorado Co.: along California Highway 89, 2.2 mi S

of U.S. Highway 50, on sandy granitic soils just E of the highway, associated with Artemisia, pinyon-juniper, and Arctostaphylos, 23 Aug

1975, James L. Reveal 3971 (isotype: UBC-V173374).

POLYPODIACEAE

Pellaea gastonyi Windham, Contr. Univ. Michigan Herb. 19:36. 1993. T-Protologue: CANADA: BritisH COLUMBIA: Kinbasket Moun-

tain by Kinbasket River, NW of Golden, 17 Aug 1953, Calder and Savile 11976 (HoLoTyPE: US; isotypes: DAO, UBC, WTU). T-Specimen:

CANADA. BritisH CoLumBra: Kinbasket Mountain by Kinbasket River, NW of Golden, very common in rock crevices, rocky gullies and

slopes at base of mountain, 17 Aug 1953, J.A. Calder and D.B.O. Savile 11976 (isotype: UBC-V75464).

Saarela et al., University of British Columbia Herbarium: vascular plant t i 445

vt L)

Dal

F.A. Lang, Madrono 20:57. 1969. T-Protologue: BritisH CoLumsia: Cheakamus River, Lang 211 (HOLOTYPE: UBC).

T-Specimen: BRITISH Co_umBia: Basalt Columns at McGuire, Cheakamus River, in crevices in basalt, 50°N, 123°W, 2n = 74, 6 Nov 1964,

F.A. Lang 211-B (HoLotYPE: UBC-V137746; Isotypes: UBC-V137749, UBC-V137745).

Polystichum kwakiutlii D.H. Wagner, Amer. Fern J. 80:50. 1990. T-Protologue: CANADA. British COLUMBIA: coast, Alice Arm, 10-9-

(19)34, A.D. York s.n. (HOLOTYPE: UBC). T-Specimen: CANADA. BRITISH COLUMBIA: coast, Alice Arm, 10-9-1934, A.D. York s.n. (HOLOTYPE:

UBC-V4859).

PORTULACACEAE

Claytonia perfoliata Donn ex Willd. var. angustifolia Greene, Fl. Francisc. 2:179. 1891. T-Protologue: U.S.A. CALIFORNIA: Santa

Clara Co.: foothills near Stanford University, 1 Apr 1902, C.F. Baker 487 (Neotyre: NDG [designated by Miller and Chambers, Syst. Bot.

Mongr. 78:120. 2006]; Isonsotyres: CAN; CAS; DS; F; GH; LL; MICH; MO; NY; POM; UBC; UC; WIS). T-Specimen: CALIFORNIA. Santa

Clara Co.: foothills near Stanford University, 1 Apr 1902, C.F. Baker 487 (IsoneoTYrE: UBC-V34942).

PTERIDACEAE

Adiantum peas var. rangiferinum E .S. Burgess, Proc. & Trans. Roy. Soc. Canada 4, Sect. 4:11. 1886. T-Protologue: ...on

thickly shaded ging the waters of Gold Stream, at the base of Mount Finlayson, twelve mi from Victoria, B.C., J. R. Anderson

s.n. (HOLOTYPE: not specified). T-Specimen: Goldstream [sic], rocky streams, J.R. Anderson s.n. (isotype: UBC-V4181).

RANUNCULACEAE

Delphinium caprorum Ewan, Bull. Torrey Bot. Cl. 69:145. 1942. T-Protologue: WASHINGTON: Cascade Mts., from Goat Rocks, 7000

ft, JW. Thompson 15206 (HOLOTYPE: COLO; IsOTYPES: to be distributed; US*). T-Specimen: WasuiNGTON. Lewis Co.: Cascade Mountains,

alpine rock slides on Goat Rocks, 7000 ft, J. William Thompson 15206 (isotype: UBC-V46005)

Aconitum variegatum L. subsp. variegatum var. carniolicum Starm., Fritschiana 10:1. 1997. T-Protologue: Slowenien, Krain, ENE

Laibach (Ljubljana), etwa 3 km NW Sagor (Zagorje), im Tal vom Kotredesch-Bach (Kotredestica) E der Ruine Gallenberg (Gamberk),

460 m, GF 9855/2 Gebüsch, 31 Aug 1996, A. Podobnik s.n. (HOLOTYPE: LJU; isotypes: CL, GJO, GZU, IBF, JACA, KL, KRA, LE, LI, LG, LJM,

M, MEL, NY, OSC, PE, Herb. Podobnik, Herb. Starmühler, TBI, TNS, TK, W, WU, Z). T-Specimen: Slowenien, Krain, ENE Laibach

(Ljubljana), etwa 3 km NW Sagor (Zagorje), im Tal vom Kotredesch-Bach (KotredeScica) E der Ruine Gallenberg (Gamberk), 460 m, 31

Aug 1996, A. Podobnik s.n. (isotype: UBC-V222228).

ROSACEAE

Crataegus atrovirens J.B. Phipps & O'Kennon, Sida 20:141. 2002. T-Protologue: CANADA. BRITISH CotuMBIA: Northern Okanagan,

Spallumcheen Municipality, Otter Lake Cross Road, 22 Aug 2000, J.B. Phipps 8171 & S.R. Phipps (HOLOTYPE: UWO; IsoTYPES: BRIT, CAN,

MO, TRT, UBC, US). T-Specimen: CANADA. BRITISH COLUMBIA: N Okanagan, Spallumcheen Municipality, Otter Lake Cross Rd., E side,

ca. quarter mi S of Hales Rd., roadside hedge under natural trees, 1300 ft, 50?22.5'N 119?13'W, 22 August 2000, J.B. Phipps and S.R.

Phipps 8171 (isotype: UBC-V217344).

Crataegus castlegarensis J.B. Phipps & O'Kennon, Sida 20:121. 2002. T-Protologue: UNITED STATES. IpaHo. Lemhi Co.: US 93, E

side, 14.5 rd. mi N of Salmon, roadside ditch next to irrigated field, alt. ca. 3950 ft, bush, 5 m tall, foliage dull, dark, coriaceous, fruit

strongly clustered, red-burgundy, with short pedicels, 20 Aug 1996, J.B. Phipps & O'Kennon 7396 (HOLOTYPE: UWO; isorvpes: BRIT, CAN,

MO, MONTU, TRT, UBC, V, WS). T-Specimen: IpAuo. Lemhi Co.: US 93, 14.5 rd mi N of Salmon, E side, roadside ditch next to irrigated

field, ca. 3950 ft, 45?22'N 113%5712'W, 20 Aug 1996, J.B. Phipps and R.J. O'Kennon 7396 (isotype: UBC-V217347).

Crataegus enderbyensis J.B. Phipps & O'Kennon, Sida 20:136. 2002. T-Protologue: CANADA. BRITISH COLUMBIA: just E of Enderby

after bridge over Shuswap R., S side of road, rough grass near water; alt. ca 1250 ft, large bush 5 m tall, burgundy-red fruit, 25 Sept

1993, J.B. Phipps & R.J. O'Kennon 6808 (HOLOTYPE: UWO; isotypes: BRIT, CAN, DAO, UBC). T-Specimen: CANADA. BRITISH COLUMBIA:

just E of Enderby after bridge over Shuswap River, rough grass near water, S side of road, ca. 1250 ft, 50°33'N 119°08'W, 25 Sep 1993,

J.B. Phipps and RJ. O’Kennon 6808 (isotype: UBC-V217345).

Crataegus okanaganensis J.B. Phipps & O’Kennon, Sida 18:178. 1998. T-Protologue: CANADA. BRITISH COLUMBIA: rough grass

with hawthorns on Hwy. 97 near entrance to Kelowna airport, alt. 1800 ft, equals JBP 6907, bush, 4 m tall, 17 Aug 1994, Phipps, J.B.,

& O'Kennon, RJ. 6974 (HOLOTYPE: UWO; isotypes: CAN, TRT, UBC, US). T-Specimen: CANADA. BRITISH COLUMBIA: rough grass with

hawthorns on Hwy. 97 near entrance to Kelowna airport, 1800 ft, 119?24"W 49°53'N, equals JBP 6907, bush 4 m tall, 17 Aug 1994, J.B.

Phipps 6974 & RJ. O’Kennon (isorvee: UBC-V213139).

Crataegus okanaganensis J.B. Phipps & O'Kennon var. wellsii J.B. Phipps & O'Kennon, Sida 20:132. 2002. T-Protologue: U.S.A.

WASHINGTON. Okanogan Co.: Palmer Lake, N side; large bush, 5 m tall, (equals J.B.P. & R.J.O'K. 6875), 4 May 1994, J.B. Phipps 6905 (uo-

LOTYPE: UWO; isotypes: BRIT, CAN, DAO, UBC, US, WS). T-Specimen: WasHiNcTon. Okanogan Co.: Palmer Lake, N side, open hawthorn

thicket at upper level of pebble beach at picnic/campsite, just E of Chopata Lodge, 1150 ft, 119°37'W 48°54'N, 4 May 1994, J.B. Phipps

6905 (isoTYPE: UBC-V217342).

Crataegus orbicularis J.B. Phipps & O’Kennon, Sida 20:138. 2002. T-Protologue: CANADA. British CoLumbla: Northern Okanagan,

Spallumcheen Municipality, Back Enderby Rd., ca. 4 mi SE of Enderby, ca. 200 m N of creek crossing, alt. ca. 1450 ft, dense hedges E

side of road, bush 4 m tall, 21 Aug 1994, J.B. Phipps & RJ. O’Kennon 7039 (HoLotyPe: UWO; isotypes: BRIT, CAN, MO, TRT, UBC). T-

Specimen: CANADA. BritisH COLUMBIA: ‘Back Enderby’ Rd, ca. 4 mi SW of Enderby, ca. 200 m N of creek crossing, ca. 1450 ft, 119°07'W

50°31'N, 21 Aug 1994, J.B. Phipps and R.J. O’Kennon 7039 (isotype: UBC-V217343).

446 Journal of the Botanical R h Institute of Texas 1(1)

Crataegus shuswapensis J.B. Phipps & O'Kennon, Sida 20:128. 2002. T-Protologue: CANADA. BritisH COLUMBIA: Northern Okanagan,

Enderby, ca. 200 m E of bridge over Shuswap River on Mabel Lake Rd., bush, 3.5 m tall, fruit deep purple (2JBP 6910), 20 Aug 1994, J.B.

Phipps & RJ. O'Kennon 7009 (HoLotyPe: UWO; isotypes: BRIT, CAN, DAO, TRT, UBC). T-Specimen: CANADA. BRITISH COLUMBIA: ca. 200

woods, 1250 ft, 119%08' W, 50°33'N,

m E of bridge over Shuswap River on road Enderby to Mabel lake, N side of road, edge

20 Aug 1994, J.B. Phipps and R.J. O’Kennon 7009 (isotype: UBC-V217346).

eed

SAXIFRAGACEAE

x Heuchera easthamii Calder & Savile, Brittonia 11:54. 1959. T-Protologue: British CoLumsia: 14 mi N of Hazelton, edge of thicket

by Kispiox River, 19 Aug 1954, Calder, Savile & Ferguson 14748 (HoLoTYPE: DAO). T-Specimen: CANADA. BRITISH COLUMBIA: at bridge

crossing Kispiox River 8 mi by road N of Kispiox village, N of Hazelton, 19 Aug 1954, J.A. Calder, D.B.O. Savile, and J. M. Ferguson 14748

(Isorvre: UBC-V72935).

Saxifraga codyana Zhmylev, Byull. Moskovsk. Obshch. Isp. Prir. Otd. Biol. 97:95. 1991. T-PROTOLOGUE: YUKON TERRITORY: British Mts.,

tributary of Fith [sic] R., 69°07'N, 140°15'W, N271667, Jul 1980, W. J. Cody 27166 : DAO; IsotyPe: MW). T-Specimen: CANADA.

Yukon Territory, British Mountains, tributary of Firth River, 69°07'N, 140°15'W, 7 Jul 1980, W. J. Cody 27166 (isotype: UBC-V212762).

Saxifraga taylori Calder & Savile, Brittonia 11:248. 1959. T-Protologue: BRITISH CoLuMBIA: Queen Charlotte Islands, Mt. de la Touche

near head of Fairfax Inlet, Tasu Sound, Calder & Taylor 23511 (HoLotyre: DAO). T-Specimen: CANADA. BRITISH COLUMBIA: Queen Charlotte

Islands, below Mt. de la Touche near head of Fairfax Inlet, Tasu Sound, W coast of Moresby Island, very common on boulders on coarse

talus slope and rock slides from 1400'-2000', rare on cliffs, 16-17 Aug 1957, J.A. Calder and R. L. Taylor 23511 (isotype: UBC-V 124231).

SCROPHULARIACEAE

Castilleja praeterita Heckard & Bacig., Madrono 4:209. 1970. T-Protologue: CALIFORNIA. Tulare Co.: N edge of Horse Meadow on

Salmon Creek, ca. 7 air mi E SE of Fairview, southern Sierra Nevada, elev. 7400 ft, 10 Aug 1966, Bacigalupi and Heckard 9190 (HOLOTYPE:

JEPS; isotypes: to be distributed). T-Specimen: CALIFORNIA. Tulare Co.: edge of Horse Meadow, ca. 10 air mi NE of Kernville, 7400 ft, 10

Aug 1966, Rimo Bacigalupi and L.R. Heckard 9190 (isotype: UBC-V 206957).

THYMELACEAE

Daphne x mantensiana J. Manten ex T.M.C.Taylor & Vrugtman, Baileya 12:39. 1964. T-Protologue: BRITISH CoLumBIA: Vancouver,

garden of Mr. H. Eddie, 7 Aug 1963, F. Vrugtman 1644 (HoLoTYPE: UBC). T-Specimen: CANADA. BRITISH COLUMBIA: ex hort garden of Mr

H. Eddie, Vancouver, B. C., 49? 16 N, 126? 15 W, 7 Aug 1963, F. Vrugtman 1644 (HoLoTYPE: UBC-V115036).

VISCACEAE

Phoradendron insignis Steyerm., Bol. Soc. Venez. Ci. Nat. 26:415. 1966. T-Protologue: VENEZUELA: Estado Bolivar, vicinity of road

campamento 150 at km 150 in valley of savanna of Río Uarama below Uarama-tepui, NE of Luepa, 1200 m, 9 Mar 1962, Julian A. Steyermark

and Leandro Aristeguieta 59 (HOLOTYPE: VEN). T-Specimen: vicinity of road campamento 150 at km 150 in valley of savanna of Río Uarama

below Uarama-tepui, NE of Luepa, 1200 m, 9 Mar 1962, Julian A. Steyermark and Leandro Aristeguieta 59 (isotype: UBC-V132586).

ADDITIONAL NOTES

Several UBC specimens are incorrectly labeled as isotypes:

A specimen [Heller 2818a (UBC-V181080)] of Cibotium st.-johnii Krajina var. typicum Krajina (Stud. Bot. Cechosl. 1:94. 1938) is labeled

(as C. st.-johnii) as an isotype. This collection is listed in the protologue of C. st.-johnii as a paratype; the type specimen of C. st.-johnii is

Krajina 26. No collections of Krajina 26 are present in UBC.

A specimen [St. John 20114 (UBC-V47651)] of Cyrtandra calpidicarpa (Rock) St. John & Storey (Occ. Pap. Bernice Pauahi Bishop Mus.

20:81. 1950) is labeled as an isotype. This name was a combination based on (the basionym) Cyrtandra longifolia Hbd. var. calpidicarpa

Rock (Amer. J. Bot. 4:618. 1917), thus the UBC specimen (collected in 1940) is not an isotype because the basionym is based on a 1909

collection according to its protologue.

A collection (Runyon 2655) of Cyperus aristatus Rottb. var. runyoni O'Neill (Rhodora 44:56. ae in UBC is labeled isotype. Ac-

cording to the protologue, the type series for this taxon is based on Runyon 1933. Since the collection number iffer, the UBC specimen

is clearly not an isotype.

A specimen [J.W. Eastham s.n. (UBC-V26885)] of Spiranthes romanzoffiana Cham., collected on 18 Aug 1938 at Lucile Lake, Garib-

aldi, British Columbia, includes has a handwritten note: “Type collected at Unalaska presumably on Kotzebue 1st Voyage. Also found at

Bantry Bay, Co. Cork, Ireland.” Based on this information, this specimen has been confused in the UBC herbarium as a type specimen,

but it clearly is not, as the species was described over a century earlier [Linnaea 3:32(-33). 1828].

A specimen (B. Boivin 6789, Loon Lake, district of North Battleford, Saskatchewan, 4 Aug 1949) with the name Geum aleppicum

Jacq. var. cuneatum is labeled isotype; however, this variety was never published and is therefore a nomen nudum. According to online

databases, duplicates of this collection (each also labeled as isotypes) are d ited in UC/JEPS and NY. There is also a duplicate labeled

isotype in MO, as well as additional collections with this name (Joseph Rohrer, pers. comm.). Boivin (Phytologia 15:354. 1967), in a

brief discussion of the taxonomic history and morphological variability of Geum aleppicum in the Canadian prairie provinces, noted “A

substitute name was prepared in 1949 and used extensively on herbarium sheets but was never actually published because the reputed

distinguishing characters proved to be elusive.” Boivin unfortunately did not include the invalid name in his discourse, but it seems

likely that he was referring to this collection of Geum aleppicum var. cuneatum, collected in the same year (by him) in which the substitute

name was prepared.

£D Li. LI L

Saarela et al., University of British Col

vascular plant ty] I i 447

APPENDIX 1

INDEX BY COLLECTORS AND NUMBER

Allen 9603

Ancuash 1142

Anderson s.n.

Bacigalupi & Heckard 9190

Baker 487

Baker 11307

Bayly 83

Boivin & Alex 10245

Boivin & Ledingham 14079

Boivin, Bernard, & Perron 12942

Calder & Savile 11976

Calder & Taylor 23511

Calder, Savile, & Ferguson 14748

Chambers 2868

Clokey 7514

Cody 27166

Constance 3895

avis 4

Eiten & Eiten 4091

Galeano, Martínez, & Suárez 2711

Galeano, Martínez, & Suárez 2767

Galeano, Martínez, & Suárez 2829

Harling & Andersson 11940

Harling & Anderson 17592

Harling, Bohlin, Lindstróm, & Roth 19833

Hibberson s.n.

Hitchcock 23868

Hitchcock & Muhlich 22305

Kennedy & Andrews 1343

Kennedy & de Fátima Gomes de Souza 4696

Kennedy & de Fátima Gomes de Souza 4699

Kennedy & de Fátima Gomes de Souza 4700

Kennedy & Hammel 4545

Kennedy, Andrews, & Dressler 1379

Kennedy, Besse, & Baker 4373

Kennedy, Besse, & Baker 4383

Kennedy, Besse, & Baker 4404

Kennedy, Hammel, & Solomon 4664

Koyama 11120

Lane 44

Lang 211

Liesner 15662

Liesner 25333

Lugo 1605

Luther 800

Madison, Plowman, Kennedy, & Besse 4646

Madison, Plowman, Kennedy, & Besse 4949

Maze, Maze, Robson, & Henn 1007

Mori, Gracie, Croat, Maas, Maas, Pennington, & Reed 22909

Murray & Murray 522

Nagata 2886

Nelson 85

Pearsall 2

Pearsall 4

Peterson & Refulio-Rodriquez 16452

Phillips 2854

Phipps 6905

Phipps & O'Kennon 6808

Phipps & O'Kennon 6974

Phipps & O'Kennon 7009

Phipps & O'Kennon 7039

Phipps & O'Kennon 7396

Phipps & Phipps 8171

Podobnik s.n.

Reveal 3971

Reveal & Reveal 3924

Reveal, DeDecker, & DeDecker 3909

Schofield 1270

St. John 25960

Steyermark & Aristeguieta 59

Suksdorf 7452

Thompson 15206

van der Werff, Gray, & Tipas 12429

van der Werff, Gray, Tipas, & Campana 12194

Vrugtman 1644

Kennedy, Besse, & Baker 4288 York s.n.

Kennedy, Besse, & Baker 4363 Zika 16611

Kennedy, Besse, & Baker 4368

ACKNOWLEDGMENTS

We are grateful to Iain Taylor for providing information on the history of the UBC herbarium; Helen Kennedy

and Fred Ganders for providing information on the status of some specimens in UBC; David Brownstein

for information on John Davidson; John Pinder-Moss and Adolf Ceska for information on specializations

of past collectors; Joseph Rohrer for information on Geum aleppicum var. cuneatum; Jason Grey for scan-

ning the type specimens (supported by the Government of Canada Summer Career Placement Program);

and Derek Tan for making the type images available on the UBC herbarium web site. We also thank Rusty

Russell (US) for comments that improved the manuscript, and for information on isotypes in US for taxa

448 Journal of the Botanical R h Institute of Texas 1(1)

included here. JMS was supported by the Natural Sciences and Engineering Research Council of Canada, a

Department of Botany Herbarium Research A

of British Columbia).

ip, and a University Graduate Fellowship (University

REFERENCES

BnipsoN, G.D.R. and E.R. SurrH (eds.). 1991. Botanico-Periodicum-Huntianum/Supplementum. Hunt Institute for

Botanical Documentation, Carnegie Mellon University, Pittsburgh, PA.

Brummit, R.K. and C.E. Powel. 1992. Authors of plant names. Royal Botanic Gardens, Kew.

HotMwcREN PK, and N.H. Hot wcntN. 1990. Index herbariorum. New York Botanical Garden Press, New York.

LAWRENCE, G.H.M., A.F.G. BUCHHEIM, G.S. DANIELS, and H. DoLezAt (eds.). 1968. Botanico-Periodicum-Huntianum. Hunt

Botanical Library, Pittsburgh, PA.

THE UNIVERSITY OF BRITISH COLUMBIA CALENDAR, EIGHTH SESSION. 1922-1923. The Sun Publishing Company, LTD., Vancouver,

British Columbia.

ARAU ARIAN SOURGE OF FOSSIEIFEROUS BURMESE AMBER.

SPECTROSCOPIC AND ANATOMICAL EVIDENCE

George Poinar Jr. Joseph B. Lambert

Department of Zoology Department of Chemistry

Oregon State University 2145 Sheridan Road

Corvallis, Oregon 97331, U.S.A. Northwestern University

Evanston, Illinois 60208-3113, U.S.A.

Yuyang Wu

Department of Chemistry

2145 Sheridan Road

Northwestern University

Evanston, Illinois 60208-3113, U.S.A.

ABSTRACT

Recent fossil discoveries show ia Burmese a is one E the most p HM sites from the Early Cretaceous. We have used

both nuclear mag ] f amber taken from the Noije Bum 2001

Summit Site in the Hukawng Valley, Myanmar. All spectra were identified as een to Group A, which on the basis of a previous

analysis of New Zealand amber and copal, is related to members of the Araucariaceae, especially Agathis. Bi- to multiseriate, angular,

alternate, contiguous 5-6-sided intertracheal pitting on the fossil wood is LH of araucarioid pitting and only occurs in wood of

extinct or extant members of the Araucariaceae. The amber from th is considered to be derived from araucarioid (especialy

Agathis) trees in the Araucariaceae

Key Wonps: Burma, fossilized resin, tracheid pits, Nuclear magnetic spectroscopy

RÉSUMÉ

De récentes découvertes chez les fossiles montrent que l'ambre birman est l'une des plus importantes sources de Crétacé ancien. Nous

avons utilisé à la fois la résonnance magnétique nucléaire (NMR) et les w a pour pA E source si de

l'ambre trouvé à Noije Bum 2001 Summit Site dans la vallée de Hukawng, Myanm

au groupe A, qui sur la base d'analyses antérieures sur l'ambre et le copal de ene Zélande est lié à des membres des Apaucanlacees

spécialement Agathis. Des trous trachéaux à 5-6 cótés, bi- à multisériés, angulaires, alternés, contigus, sur le bois fossile sont typiques

des alvéoles des Araucarias ou d'araucarioides et se rencontrent dans le bois des membres des Araucariaceae. Lambre de ce site minier

est donc considéré comme venant d'Agathis, un genre actuel d'Araucariaceae.”

INTRODUCTION

Burmese amber was traded with China as early as AD 100, but it was not until 1896 that fossils were reported

in these deposits. In 1999, a new amber site was discovered in the Hukawng Valley in Myanmar (Poinar

et al. 2005), and palynomorphs from this site were assigned to the Upper Albian of the Early Cretaceous

(97-110 million years ago) (Cruickshank & Ko 2003). The same deposits have yielded the oldest bee (Poinar

& Dancroft 2006), the oldest angiosperm flowers in amber (Poinar & Chambers 2005; Poinar 2004) and the

first fossil evidence of vector borne diseases (Poinar & Poinar 2004), making it one of the most significant

Cretaceous amber deposits in the world. Due to the scientific importance of this amber, we have used both

spectroscopic and anatomical analyses to determine the plant source.

Clues to the determination of source trees of amber deposits can be provided by plant macrofossils or

microfossils found in the amber or in sedimentary beds containing the amber and by spectroscopic analysis

of the amber. A combination of these methods can then be used to describe source trees, as was done with

Agathis levantensis, the araucarian tree responsible for the production of Middle East amber (Poinar & Milki

2001). Up to the present, the only spectroscopic evidence for the tree source of Burmese amber has been the

nuclear magnetic resonance (NMR) analysis of three samples of Burmese amber (Lambert & Poinar 2002).

The results of these analyses were conflicting with one sample unassignable (Lambert & Frye 1982), one

J. Bot. Res. Inst. Texas 1(1): 449 — 455. 2007

450 Journal of the Botanical R h Institute of Texas 1(1)

related to the family Dipterocarpaceae (Lambert et al. 19992) and the third related to the Araucariaceae

(Lambert & Wu, unpublished research, 2002). Therefore, further analysis was undertaken with additional

Burmese amber samples collected from this new site that was first mined in 2001. Tracheid fibers in one

of the amber samples from this new site are characterized and used to provide anatomical evidence of the

plant source.

MATERIALS AND METHODS

The amber samples analyzed in this study were collected from lignitic seams in sandstone-limestone deposits

in the Hukawng Valley in Myanmar. The mine site was located on the slope of the Noije Bum hill about

a mile (1.5 km) SSW of the old Khanjamaw mine site and southwest of Maingkwan (26?20'N, 96?36 E).

Apparently this site had never been mined previously (Chhibber 1934) (Doug Cruickshank, pers. comm.,

December 20, 2006), and we refer to it as the “Noije Bum 2001 Summit Site.”

Wood Fiber Analysis

A square piece of amber measuring approximately 23 mm on all sides and 6 mm thick contained numerous

strips of wood fibers. This piece of amber was cut with a diamond saw along the flat side, leaving two nar-

row pieces of amber, each approximately 3 mm in thickness. The surfaces of these were polished and the

wood fibers examined with a Nikon Optiphot optical microscope (with magnifications up to 800x). The

amber pieces containing the wood fibers are deposited in the Poinar amber collection (accession B-P-16)

maintained at Oregon State University.

Spectroscopic Studies

Solid-state nuclear magnetic resonance spectroscopy with cross polarization and magic angle spinning

(CP/MAS) was used to characterize the amber samples (Lambert et al. 1996). Both normal decoupling and

interrupted decoupling modes were used to examine the solid-state "C NMR spectra.

Solid state ^C NMR data were recorded on a 400 MHz Varian NMR System. The Direct Drive console

had a clean rf architecture, a powerful digital receiver, and utilized advanced phase, amplitude modula-

tion. The system had a 5 mm T3 PENCIL probe. The magic angle spinning rate was set to 5000 Hz. The

cross polarization pulse sequence called tancpx was used for normal proton decoupling. For interrupted

decoupling, the pulse sequence tancpxidref was used, in which a 50 ps delay was applied in the 'H channel

directly before the 180? pulse in the PC channel. A typical parameter set was as follows: spectrum frequency

100.544 MHz, spectral width 50 kHz, pulse width 3.4 us for the 90° pulse for both 'H and PC, delay time

5 ys, contact time 2 ms, acquisition time 20.5 ms, and scan number 256. Spectra were referenced to an

external adamantane peak at 6 38.3 and were converted to tetramethylsilane at 6 0.0. Data were collected

and processed with software Vnmr] 2.1B.

This procedure was performed on 5 separate samples (Nos.276, 375, 376, 422, 441) from the Noije Bum

2001 Summit Site. Four of the samples were clear (Nos. 276, 375, 422, 441) and one was opaque (376).

RESULTS

Wood Fiber Analysis

The tracheid surfaces contained 2-3 rows (2-3 seriate) of alternately arranged contiguous, angular, 5-6

sided tracheid pits (Fig. 1). Most of the bordered tracheid pits were hexagonal with diameters ranging from

11 tol4 um. None of these biseriate and triseriate pits possessed thickenings, such as crassulae or bars of

Sanio between them. The polygonal pit cavities, which did not extend beyond the pit borders, varied from

round to 5-6 sided (Fig. 1C). Resin globules occurred within some of the tracheids (Fig. 1C).

Spectroscopic Analysis

In all samples, the spectra with normal decoupling have the largest peak at 6 38 and a second, smaller peak

at 6 18-20, with a broad grouping in the unsaturated region. With interrupted decoupling, the largest peak

again was at 6 38, with nothing in the unsaturated region. All spectra were identified as belonging to Group

Poinar et al., Araucarian source of fossiliferous B b 451

Fic.1. A. Bi- and tri-seriate tracheid pit wood fragment in Burmese amber collected from the Noije Bum 2001 Summit Site. Scale bar = 14 um.

B. Contiguous, alternate, 5—6 sided pit ly referred t ioid pitti wood fragment in B ber. Scale bar = 8pm. C. Pit

Mat +

I^ f l

1). Scale bar = 15um.

£+sha D o ID L

452 Journal of t titute of Texas 1(1)

ES IS A ECES SN ECI A EE EEC] EEEE ETE EE E E TE ET CET EEE] CEET EET EEEE TET | T FTVTTTTT]

200 180 160 140 120 100 80 60 40 20 ppm

Fic. 2. PC NMR spectra of B l le 376 collected from the Noije Bum 2001 Summit Sit

srr MN FI

A (Fig. 2), which on the basis of a New Zealand series reported earlier (Lambert et al. 1993), is related to

members of the Araucariaceae, especially Agathis, a genus commonly known as kauri pines now restricted

to the Southern Hemisphere.

DISGUSSION

The arrangement of intertracheary pits and their cavities can be used to characterize familes and genera of

conifers (Core et al. 1979; Patel 1968). Bi- to multiseriate, angular, alternate, contiguous, 5-6 sided longi-

tudinal tracheid pits as seen on the radial walls of the tracheids, as found here, are commonly referred to

as araucarioid pitting and only occur in wood of members of the Araucariaceae (Patel 1968; Tidwell 1998).

Diameters of tracheid pits in members of the Araucariaceae vary from 12 to 16 um (Core et al. 1979; Tidwall

1998) which is within range of the pits in the Burmese amber wood samples (11-14 um). The presence of resin

globules or plugs, as found in the fossilized wood, is characteristic of araucaroid tracheids (Patel 1968).

There are currently three extant genera in the family Araucariaceae: Agathis, Aruacaria and Wollemia.

While the wood structure of Agathis and Araucaria is similar, (Langenheim 1995), recent resin of these two

genera often can be distinguished by NMR analyses (Lambert & Poinar 2002), primarily because most Ar-

aucaria species produce gum resins (Lambert et al. 2005). In addition, resin deposits from Araucaria trees

are small and do not polymerize and fossilize, as does Agathis (Langenheim 1995). While the NMR spectra

Poinar et al., Araucarian source of fossiliferous B b 453

of Wollemia and Agathis are similar (Lambert et al. 1999b), there is no evidence that Wollemia produces large

deposits of resin that polymerize and form amber deposits.

Under normal decoupling conditions in NMR analyses, signals are obtained from all carbons present in

the sample. With interrupted decoupling, signals are selected for quaternary carbons and carbons that are

moving rapidly in the solid while other signals are edited out. These two spectral modes served as fingerprints

to analyze and identify the Noije Bum 2001 Summit site amber samples as belonging to the worldwide Group

A, which is most similar to members of the genus Agathis in the family Araucariaceae. Determination of this

taxonomic grouping was based on comparisons with previously obtained NMR spectra of Agathis resin, copal

and amber from New Zealand (Lambert et al. 1993). It has been found, however, that the PC fingerprint of

modern resins (as opposed to gum resins) from the family Araucariaceae are indistinguishable from that of

modern resins from the family Cupressaceae (Lambert et al. 2005). The presence of araucarioid wood fibers

in the amber from the Noije Bum 2001 Summit site is collaborative evidence that the NMR spectra reported

been made since it is difficult to obtain samples of fossilized resin associated with present day members of

the Cupressaceae. There apparently are no localities that contain semi-fossilized or fossilized resin deposits

of species of the Cupressaceae in soil surrounding the trees, swamps containing the stumps of former forests

or in coal deposits, as occurs in New Zealand with Agathis (Poinar 1991; Lambert et al. 1993).

While some fragments of amber have been attributed to members of the Curpessaceae, (which now

includes genera of the former family Taxodiaceae), there is no evidence that extant Cupressaceae produce

copious amounts of resin under normal growth conditions. Members of this family only have a limited capacity

to produce trunk resins (Langenheim 1995), the source of the majority of fossiliferous amber and then only

when they experience severe traumatic stress. This is in contrast to species of Agathis, that produce not only

copious amounts of trunk resins, but also were the source of fossilized resins (copal and amber) extending

over thousands and millions of years and under a wide range of stratigraphic conditions (Halkett & Sale

1986; Poinar 1991; Lambert & Poinar 2002). While wood of the extinct conifer family Cheirolepidiaceae

may also contain tracheids with araucaroid radial pitting, the absence of resin ducts and canals is a feature

typical of these wood types (Axsmith & Jacobs 2005; Taylor & Taylor 1993), thus they could not have been

the source of Burmese amber.

Determining araucarioid (very likely Agathis) trees as the source of Burmese amber is congruent with

large amounts of pollen of the araucarian, Araucaricites australis, recovered in a palynological analysis of the

sedimentary formation that contained the amber at the Noije Bum 2001 Summit Site (Davies 2001).

Commonly known as kauri or kauri pines, species of Agathis can become quite large and long-lived

(from 500 to 1000 years) and have been compared with the Giant Sequoias of North America (Halkett & Sale

1986). By bulk alone, they would have formed a significant part of the Burmese amber forest. The climate

of Burma was tropical- subtropical during the Early Cretaceous (Boucot et al. 2007), which is the preferred

climate for extant Agathis species, all of which are now confined to the Southern Hemisphere (Poinar &

Milki 2001).

Previous spectroscopic analyses of Burmese amber have been conducted with material from other

sites. The first sample of Burmese amber tested (#7) was provided by Curt Beck and came from the Musée

National d'Histoire Naturelle, Paris, (No. 101.604). The label stated it was collected from Mangotaimaw Hill,

Myanmar. Its spectrum was broad and could not be assigned to any present group (Lambert & Frye 1982).

In 1992, an analysis was performed on a piece of amber presented to the senior author by a commercial

trading company located in Taiwan. This sample, (4154), which was dark brown, nearly opaque and lacked

fossils, was represented as coming from the Hukong Valley in Burma. It produced a spectrum typical of

the worldwide Group B, which is related to the extant angiosperm family Dipterocarpaceae (Lambert et al.

1999a; Lambert & Poinar 2002). In 2002, a sample of Burmese amber (#276) collected from the Noije Bum

2001 Summit Site, where samples were also taken by Cruickshank & Ko (2003) for their dating studies,

454 Journal of the Botanical R h Institute of Texas 1(1)

showed a definite placement in the worldwide Group A, similar to the samples presented here. This result

was simply cited in publications as Lambert & Wu, unpublished research (2002).

The present study shows that the amber from the Noije Bum 2001 Summit Site is produced from an

araucarioid, quite probably a member of the genus Agathis. If sample #154 did originate from Burma, it

indicates that at least two separate plant families (Araucariaceae and Dipterocarpaceae) were sources of

amber in Myanmar, quite possibly at two distinct geological periods.

ACKNOWLEDGMENTS

The authors acknowledge financial support provided by the Graduate Research Institute of Gallaudet Uni-

versity and by the National Science Foundation (Grant No. CHE-03-49412) and thank Roberta Poinar for

comments on earlier drafts of this manuscript and Pierre Jolivet for providing the French abstract. Com-

ments of Sherwin Carlquist, Eugene Mazzola, and an anonymous reviewer are greatly appreciated. We also

acknowledge an instrument grant from the National Science Foundation (Grant DMR-05-21267).

REFERENCES

AxsmirH, B.J. and B.F. Jacoss. 2005. The conifer Frenelopsis ramosissima (Cheirolepidiaceae) in the Lower Cretaceous

of Texas: systematic, biogeographical, and paleoecological implications. Int. J. Plant Sci. 166:327-337.

Boucor, A., Xu CHEN, and C.R. Scotese. 2007. Preliminary compilation of Cambrian through Miocene climatically

sensitive deposits (submitted).

CHHIBBER, H.L. 1934. The mineral resources of Burma. Macmillan and Company, London.

Core, H.A., W.A. Cote, and A.C. Dav. 1979. Wood structure and identification. 2" ed. Syracuse University Press,

Syracuse.

CRUICKSHANK, R.D. and K. Ko. 2003. Geology of an amber locality in the Hukawng Valley, northern Myanmar. J.

Asian Earth Sci. 21:441—455.

Davies, E.H. 2001. Palynological analysis and age assignments of two Burmese amber sample sets. Unpublished

report by Branta Biostratigraphy Ltd. for Leeward Capital Corp.

HALKETT, J. and EV. Sate. 1986. The world of the Kauri. Reed Methuen Ltd, Auckland.

LAMBERT, J.B. and J.S. Frye. 1982. Carbon functionalities in amber. Science 217:55-57.

LAMBERT, J.B., S.C. JOHNSON, G.O. POINAR, JR, and J.S. Fave. 1993. Recent and fossil resins from New Zealand and Aus-

tralia. Geoarchaeology 8:141-155.

LAMBERT, J.B., S.C. JoHNson, and G.O. Pomar, Jr. 1996. Nuclear magnetic resonance characterization of Cretaceous

amber. Archaeometry 38:325-335.

LAMBERT, J.B., C.E. SHAWL, S.C. JOHNSON, and G.O. Poinar, Jr. 1999a. Fossil resin from Asia. Ancient Biomolecules

3:29-35.

LAMBERT, J.B., C.E. SHaw, G.O. Ponar, JR., and J.A., SANTIAGO-BLAY. 1999b. Classification of modern resins by solid state

nuclear magnetic resonance spectroscopy. Bioorg. Chem. 27:409-423.

LAMBERT, J.B. and G.O. Pomar, Jr. 2002. Amber: the organic gemstone. Acc. Chem. Res. 35:628-636.

LAMBERT, J.B., Y. Wu, and J.A. SANTIAGO-BLay. 2005. Taxonomic and chemical relationships revealed by nuclear mag-

netic resonance spectra of plant exudates. J. Nat. Prod. 68:635-648.

LANGENHEIM, J.H. 1995. Biology of amber-producing trees: focus on case studies of Hymenaea and Agathis. In: K.B.

Anderson and J.C. Crelling, eds. Amber, Resinite, and fossil Resins, American Chemical Society, Washington,

DC. Pp. 1-31.

PateL R.N. 1968. Wood anatomy of Cupressaceae and Araucariaceae indigenous to New Zealand. New Zealand

J. Bot. 6:9-18.

Polar, Jr, G.O. 1991. Resinites, with examples from New Zealand and Australia. Fuel Process. Technol. 28:

135-148.

POINAR, JR., G.O. and R. Milki. 2001. Lebanese amber. Oregon State University Press, Corvallis.

Poinar, JR, G.O., R. Buckley, and A. Brown. 2005. The secrets of Burmese amber. Midl. Amer. Paleont. Soc. 29:

20-29.

Poinar et al., Araucarian source of fossiliferous B b 455

POINAR, JR, G.O. and R. Pomar. 2004. Paleoleishmania proterus n. gen., n. sp., (Trypanosomatidae: Kinetoplastida)

from Cretaceous Burmese amber. Protista 155:305-310.

Pomar, JR, G.O. 2004. Programinis burmitis gen. et sp. nov., and P. laminatus sp. nov., Early Cretaceous grass-like

monocots in Burmese amber. Austral. Syst. Bot. 17:497—504.

Ponar JR., G.O. and K.L. CHampers. 2005. Palaeoanthella huangii gen. and sp. nov, an early Cretaceous flower (An-

giospermae) in Burmese amber. Sida 21:2087-2092.

Ponar, JR., G.O. and B.N. DANForTH. 2006. A fossil bee from Early Cretaceous amber. Science 314:614.

Tavlor, T.N. and E.L. TayLoR. 1993. The biology and evolution of fossil plants. Prentice Hall, Englewood Cliffs, NJ.

TioweLL, W.D. 1998. Common and fossil plants of Western North America, 2" Ed. Smithsonian institution Press.

456 Journal of the Botanical R h Institute of Texas 1(1)

BOOK REVIEWS

Peter HeDDEN and STEPHEN G. Thomas (eds.). 2006. Annual Plant Reviews, Volume 24: Plant Hormone

Signaling. (ISBN 1-4051-3887-4, hbk.). Blackwell Publishing. 2121 State Ave., Ames, IA 50014-8300,

U.S.A. and 9600 Garsington Road, Oxford, OX4 2DQ, UK. (Orders: www.blackwellprofessional.com,

lers@ blackwellpublishing.com, 515-292-0140, 515-292-3348 fax, 1-800-862-6657). $199.99,

348 pp., illustrations, T x oy",

1 ji

The 24" volume of Blackwell Publishing’s Annual Plant Review series focuses on plant hormones, signaling that do not always

conform to the classic definition of a hormone. For example, ethylene, a hormone exploited by our ancestors to hasten fruit ripening

in the form of wound ethylene, is actually a gas. It was once used to light street lamps and was one of the first anesthetics. The gas's

anesthetic effect was first discovered when a group of Chicago scientists investigating why flowers in a greenhouse would not bloom

discovered that the culprit was the ethylene used to light the greenhouse. They decided to investigate the gas's effect on animals, and the

world's first anesthetic was discovered. Salicylic acid is another important plant hormone with medical applications. It is used today as a

topical acne and wart treatment and is the basis of the salicylate cl f drugs, which include aspirin. In plants, salicylic acid is involved

in protection against pathogens, thermogenesis, flowering, leaf abscission, and stomatal closure. Technological advancement has led to

many recent advances in the understanding and identification of plant hormones and their receptors, actions, and interactions with the

environment and each other. This volume is dedicated to recent advances in the relatively new field of plant hormone research, includ-

ing eight chapters that focus on the biosynthesis and metabolism of the “classic” plant hormones: ethylene, salicylic acid, cytokinin,

the gliberellins, abscisic acid, the brassinosteroids, the oxylipins, and auxin. Auxin is the first plant hormone to be discovered. It was

identified as a plant growth promoter in 1926, although its effects were first observed in the 1800's. Chapter 2 covers auxin metabo-

lism and “how plants perceive and respond to auxin.” It includes a discussion of auxin receptors. The first chapter focuses on abscisic

acid, which is a key hormone in regulating the drought response in plants, including mediation of *seed maturation processes such as

desiccation tolerance and dormancy." Chapter 4 is focused on cytokinin, including its biosynthesis and metabolism. Its roles include

the regulation of shoot and root growth, leaf senescence, stress response, and pathogen resistance. Cytokinin is believed to effect two

types of signaling local (paracrine or Md ae long-distance, or endocrine- de IE incus 9 focuses on plant hormone

n £f

distribution and transport. Thel lant hormones rol flower, and seed

development. The role of plant! till being elucidated, along with how Mer ganaling occurs aun plants. It is believed that

future research will reveal the importance of protein Sen in this process.— Marissa N. Oppel, Collections and Research Assistant,

Botanical Research Institute of Texas, Fort Worth, TX, 76102-4060, U.S.A.

WILLIAM C. Plaxton and MICHAEL T. McManus (eds.). 2006. Annual Plant Reviews, Volume 22: Control of

Primary Metabolism in Plants. (ISBN 1-4051-3096-2, hbk.). Blackwell Publishing. 2121 State Ave.,

Ames, IA 50014-8300, U.S.A. and 9600 Can on Road, Oxford, OX4 2DQ, UK. (Orders: www.

blackwellprofessional.com, orders? blackwellpublishing.com, 515-292-0140, 515-292-3348 fax,

1-800-862-6657). $219.99, 386 pp., 614" x 915".

Primary metabolism in plants is defined by the editors of this volume *...as the primary auxotrophic pathways in plants...and those in

common with the primary pathways in mammalian cells." This edition of the Annual Plant Reviews series covers the control of primary

metabolism in plants. Metabolic control, as opposed to metabolic regulation, is the adjustment of a metabolic pathway output “...in

response to an external signal." The editors write in the preface that the ability to control the rates of metabolic processes is so essential

to the survival of living cells that it must be as old as life itself. This book includes reviews covering the genomics, proteomics, and

metabolomics of the control of primary metabolism, as well as the control of specific metabolic pathways and enzymes. There have been

many advances in these fields over the last few years, and many discoveries related to plant metabolic control have been made. Aimed

at scientists in the fields of plant biochemistry, physiology, molecular biology, and cell biology, the reviews gathered in this volume are

excellent sources of information about the basics, as well as recent advances in our understanding of the control of primary metabolism

in plants.— Marissa Oppel, Collections and Research Assistant, Botanical Research Institute of Texas, Fort Worth, TX, 76102-4060, U.S.A.

J. Bot. Res. Inst. Texas 1(1): 456. 2007

STATUS OF SCHOPNOPLEGTUS HAL (HALES BULRUSED

(CYPERACEAE IN TAE UNITED TAIE.

Paul M. McKenzie S. Galen Smith

U.S. Fish & Wildlife Service Department of Biology

101 Park DeVille Dr, Suite A University of Wisconsin-Whitewater

Columbia, Missouri 65203, U.S.A. Whitewater, Wisconsin 53190, U.S.A.

Marian Smith

Department of Biology

Southern Illinois University

Edwardsville Illinois 62026, U.S.A.

ABSTRACT

New information is provided on the distribution, ecology and life history, threats, research needs and conservation status for Hall’s

bulrush (Schoenoplectus hallii). ne information is presented on taxonomy, ue status of Hall’s bulrush in each state where it has

been recorded, management ,and the adequacy of any l

eto protect or manage the species’

habitat. A history is provided for previous evaluations of Hall’s bulrush by the DU: s Fish and UE Service, and the species' potential

as a candidate for official listing under the Endangered Species Act of 1973.

RESUMEN

Se aporta nueva información sobre la distribución, ecología y historia vital, amenazas, necesidades de investigación y estado de conser-

vación de la espadaña (Schoenoplectus hal i Se presenta información adicional sobre su taxonomía, el estatus de la espadaña en todos

l tados donde se ha citado ] b manejo, y lo apropiado de cualquier mecanismo de regulación para proteger o

manejar el hábitat de la especie. Se aporta una historia de las evaluaciones previas de la espadaña por el U.S. Fish and Wildlife Service,

y el potencial de la especies como candidata para estar colocada en la lista oficial de la Ley de Especies Amenazadas de 1973.

INTRODUCTION

McKenzie (1998) provided an initial range-wide conservation status assessment of Hall's bulrush (Schoenoplec-

tus hallii) (A. Gray) S.G. Smith, which included information on distribution, ecology and life history, threats,

research needs, and conservation status. Beatty et al. (2004) included a summary of some new information

on S. hallii in a conservation assessment for the species in Region 2 of the U.S. Forest Service (USFS), but the

main focus of their report was Kansas and Nebraska. Since McKenzie's (1998) analysis, new information on

the species distribution, germination requirements, and threats to its continued existence has been reported.

The purpose of this paper is to summarize information received from published and unpublished reports

that have been completed since the 1998 assessment (McKenzie 1998). Due to the potential confusion of

S. hallii with the closely related S. erectus (Poir.) Palla ex J. Raynal and S. saximontanus (Fernald) J. Raynal,

reports of S. hallii throughout the United States were re-evaluated by Ernie Schuyler of the Philadelphia

Academy of Natural Sciences or S.G. Smith (Schuyler 1969; S.G. Smith, pers. comm).

Taxonomy

Taxonomic nomenclature for plants used in this report follows Yatskievych (1999) or Yatskievych & Turner

(1990) except for Lesquerella fendleri which follows Great Plains Flora Association (1986), S. erectus which

follows Smith (2002b), Nymphoides peltata which follows Gleason and Cronquist (1991), and Clarkia spring-

villensis, Lolium multiflorum, and Gilia achilleifolia which follow Hickman (1993).

Schoenoplectus hallii belongs to Schoenoplectus section Supini (Cherm.) J. Raynal, in the Cyperaceae.

Until recently the genus Schoenoplectus (Rchb.) Palla was generally included in the genus Scirpus L. sensu

lato (e.g., Gleason & Cronquist 1991; Tucker 1987; Schuyler 1969). Schoenoplectus sect. Supini has been

confused with S. sect. Actaeogeton, but it can be distinguished from that group by the frequent occurrence

J. Bot. Res. Inst. Texas 1(1): 457 — 481. 2007

458 Journal of the Botanical R h Institute of Texas 1(1)

of amphicarpy (the production of solitary pistillate flowers enclosed in sheaths of basal leaves in addition to

bisexual flowers in spikelets at the culm tips), which occurs in all North American species in the section, the

lack of perianth (except rarely in basal flowers), and, in North American species, the presence of a cauline

leaf (Smith 2002b; Smith & Hayasaka 2001, 2002). Recent investigations have led to division of Scirpus

sensu lato into various segregate genera (e.g., Wilson 1981; Weber & Wittmann 1992; Strong 1994; Smith

1995, 2002b; Smith & Yatskievych 1996; Yatskievych 1999; Smith 2002b; Flora of North America Editorial

Committee 2002). Schoenoplectus hallii is currently recognized by most taxonomists as the correct name for

the species (Smith 1995). Raynal (1976) placed Schoenoplectus hallii in synonymy with S. erectus, but Smith

(1995) provided convincing evidence why Raynal's treatment was invalid. Today most taxonomists agree

that S. hallii is a valid, distinct species.

Schoenoplectus hallii is most similar to S. erectus, which is known from FL, GA, SC, TX, Mexico, South

America, Asia, Africa and Australia, and S. saximontanus, which is known from British Columbia, CA, CO,

KS, MO, NE, OH OK, SD, TX, UT, and Mexico (Smith: 2002b)

Description

A small to medium-sized, tufted annual, with short rhizomes hidden among the aerial stem bases; culms

4-45 cm long, to 1 mm wide, stiff to flaccid, cylindric, leaves 3-4 basal and one cauline, blades 1-2, 0.1-20

cm long, to 1 mm wide; lowest involucral bract sometimes resembling a continuation of the culm, 3-15

cm long, about one-half as long as the culm; inflorescence consisting of 1-7 sessile spikelets in a head-like

cluster, or rarely with 1 or 2 short branches; spikelets ovoid to lanceoloid, 5-20 mm long, 2.0-3.5 mm wide;

spikelet scales 2.5 to ca. 4.0 mm long, tan or pale orange-brown to straw-colored as they age, ovate, midrib

region often green, midrib projecting as a short cusp (mucro) past the body of the scale; solitary pistillate

flowers sometimes present at the base of the culm, enclosed by an encircling leaf sheath (most frequent late

in the flowering season); perianth bristles absent; stigmas 2 or 3 in basal flowers; achenes ovoid to obovoid

or nearly circular, base abruptly contracted to a short neck, apex with an abrupt beak 0.1 mm long; concave

(rarely nearly flat) on one side and convex on the other, transversely rugose (corrugated) with about 15-18

conspicuous cross-wrinkles on each side, 1.3-2.0 mm long, brown when young, dark brown to black at

maturity; basal achenes significantly larger (Smith et al. 2006), unequally 3-angled; 2n = 22.

Schoenoplectus hallii is very similar to the other two species of Schoenoplectus sect. Supini that occur

in North America: S. erectus and S. saximontanus (Smith 2002b). In contrast to the achenes of S. hallii as

described above, the achenes of S. erectus are strongly convex on one side and slightly convex on the other

side and have 10—15 ridges on the most convex side. Both S. hallii and S. erectus can be distinguished from

S. saximontanus by their two-branched styles (three-branched in S. saximontanus) and by their two-sided

achenes (3-sided in S. saximontanus). Schoenoplectus erectus differs from S. hallii and S. saximontanus in the

color of its spikelet scales, which are bright orange on their bases and toward their tips.

Some specimens from a mixed population of S. hallii and S. saximontanus in Oklahoma have a high

percentage of aborted achenes or achenes that are morphologically intermediate between the two species

and may be hybrids (Magrath 2002; Smith et al. 2004). Some specimens from southwestern Georgia may

be S. erectus x S. hallii hybrids (Smith 2002b).

Reported range

There is controversy in the literature concerning the historical range (pre-1981) of S. hallii. Previous reports

from Alabama (U.S. Fish & Wildlife Service 1993), Colorado (Harrington 1954; Small 1972; U.S. Fish &

Wildlife Service 1993), Florida (Beetle 1947; Koyama 1962; Radford et al. 1964; Mohlenbrock 1976; Great

Plains Flora Association 1986; Robertson et al. 1994), South Carolina (Radford et al. 1964; U.S. Fish &

Wildlife Service 1993), South Dakota (Great Plains Flora Association 1977), and Texas (Correll & Johnston

1970; Small 1972; Mohlenbrock 1976; Great Plains Flora Association 1986; U.S. Fish & Wildlife Service

1993; Beatty et al. 2004; O'Kennon & McLemore 2004) are either based on misdeterminations of S. saxi-

montanus along the western edge of the species range, or misidentifications of S. erectus from the southern

United States (Rolfsmeier 1995; McKenzie 1998; Smith 2002b). Further, many county records from states

McKenzie et al., C ti t of Scl lectus hallii in the United States 459

where S. hallii has been documented, as reported by the Great Plains Flora Association (1977), were based

on misdeterminations of S. saximontanus (Rolfsmeier 1995; McKenzie 1998; Smith 2002b). A report for Or-

egon (Koyama 1962) was due to the misreading of the label on a specimen from near the village of Oregon,

Dane County, WI. A record for Decatur County, GA, based on a voucher specimen at the State of Georgia

Herbarium (T. Patrick, Georgia DNR, pers. comm.), had been previously re-determined by Schuyler (1969)

to be S. erismanae [= S. erectus (Schyuler, Philadelphia Academy of Natural Sciences, pers. comm.)], and four

collections from Baker and Decatur counties, GA and originally identified as S. hallii were redetermined by

S.G. Smith to be S. erectus (S.G. Smith, pers. comm.).

Based on verification of specimens by Schuyler or S.G. Smith, the documented pre-1981 range for S.

hallii is GA, IL, IA, KS, MA, MI, MO, NE, and WI (Table 1) (McKenzie 1998; Smith 2002b; Beatty et al.

2004). Schoenoplectus hallii has been extirpated from Massachusetts where it was last collected in 1908, and

the lack of documentation since 1946 suggests that the species may have been extirpated from Georgia

(McKenzie 1998; O'Kennon & McLemore 2004; NatureServe 2006). Schuyler or S.G. Smith has confirmed

the identification of specimens of S. hallii collected from 10 states within the last 25 years (Table 1) (Smith

2002b; O'Kennon & McLemore 2004). Texas has recently been included in the range of S. hallii based on

the discovery of the species in the state in 2003 (O'Kennon & McLemore 2004).

Apparent changes in distribution and status of S. hallii in the United States

For the following reasons, it is extremely difficult to assess any apparent changes in the distribution and

status of S. hallii: 1) population numbers of this species vary widely from year to year depending on the

availability of suitable wetland habitat, 2) populations may fail to emerge at some sites for many years, but

re-emerge when conditions are favorable for germination, and 3) the species is frequently confused with S.

saximontanus and S. erectus.

Due to the persistence of the achenes in the seed bank, the Nature Conservancy considers any site where

S. hallii has been recorded within the last 25 years to be extant (Ostlie 1990; Ostlie & Gottlieb 1992). To

determine if there has been any apparent change in the distribution and status of this species, we followed

the Nature Conservancy’s criterion and first analyzed all documented county and single locality records

of S. hallii older than 25 years (Table 1, Table 2; Fig. 1). We compared that information with documented

records of the species within the last 25 years (Table 1, Table 3; Fig. 2) and within the last five years (Table

1, Table 4; Fig. 3). We found that determining the exact number of records was problematic due to: 1) the

frequent misidentification of specimens, 2) the lack of specific locality data given for some collections, 3)

over-counting of populations that resulted when sites that were within 1 km of one another were recorded

as separate sites, and 4) the failure to monitor some known sites, especially during years when drought con-

ditions persist or when there was a lack of flooded conditions needed for germination and growth. Despite

these limitations, we obtained sufficient information for comparing records within the last five and 25 years

and what was known and confirmed historically.

Prior to 1981, S. hallii was documented at approximately 30 sites distributed across 15 counties in

nine states (Table 1, Table 2). Within the last 25 years, S. hallii has been confirmed from approximately

84 sites scattered across 26 counties in 10 states (Table 1, Table 2). The increased number of sites docu-

mented between 1981 and 2006 largely reflects recent surveys that were conducted in IL, MO, NE and

OK during years when conditions were optimal for the species (Table 1, Table 2). Over 7996 of records

confirmed within the last 25 years have been from IL, MO, and NE (Table 1). Within the last five years,

S. hallii has been collected at 25 sites from 11 counties in only six states: eight sites in Missouri, six sites

in Nebraska, but only one site each in IL, IN, KY, MI, OK, and TX, and none in GA, IA, KS, or MA (Table

1, Table 4). The paucity of records within the last five years may be due to drought conditions that re-

duced available habitat or reflect a failure to monitor sites. In Wisconsin, abnormally high rainfall flooded

the only known site and prevented a survey in 2006 (S.G. Smith pers.comm.). Despite the fewer docu-

mented sites within the last five years, new populations have been recently confirmed for Indiana (M.

Homoya, braska (Steinauer 2001a,b), Oklahoma (Magrath 2002; Smith 2002b) and Texas (O'Kennon &

460

fal, Dat o ID L

Journal of Institute of Texas 1(1)

TABLE 1. Approximate number of sites where Schoenoplectus hallii was documented historically (i.e., prior to 1981), or where

the species has been confirmed within the last 25 years, and within the last 5 years.

Number of sites

State Historical Last 25 years Last 5 years

GA ] 0 0

IA 1 0 0

IN 0 6 3

IL ~13 ~29 1

KS 7 5 0

KY 0 1 1

MA 2 0 0

MI 1 5 1

MO 2 10 10

NE 2 27 6

OK 0 13 q^

TX 0 1 1

WI 1 1 1

Total ~30

*Includes 14 ponds

TagLE 2. Counties with historical records of Schoenoplectus Taste 3. Counties where Schoenoplectus hallii has been

hallii prior to 1981. documented within the last 25 years.

State County State County

GA Dougherty IL Alexander, Cass, Kankakee, Mason, Morgan

IL Cass, Mason, Menard IN Daviess, Lake, Porter

IA Muscatine KS Harper, Reno

KS Harvey, Reno KY Christian

MA Essex, Middlesex MI Allegan, Muskegon

MI Muskegon MO Howell, Scott

MO Howell, St. Louis NE Brown, Garfield, Holt, Loup, Rock, Wheeler

NE Holt, Rock OK Atoka, Comanche, Johnston

WI Dane TX Wise

WI Dane

TABLE 4. Counti

ith A

ITO

hallii within the past five years.

umented records of Schoenoplectus

State County

IL Mason

IN Daviess, Lake

MO Howell, Scott

NE Brown, Holt, Rock, Wheeler

OK Comanche

TX Wise

Indiana DNR, pers. comm.), Ne McLemore 2004). It is likely that S. hallii has been extirpated from Massachu-

setts and possibly Georgia (McKenzie 1998; Smith 2002b; O’Kennon & McLemore 2004; NatureServe 2006).

Documented records of S. hallii suggest that it has always been a rare species in most states as reported

by Schuyler (1969; pers. comm.) and S.G. Smith (pers. comm.). While this still may be the case, the lack

McKenzie et al., C ti t of Scl lectus hallii in the United States 461

| TET RE

e»

f£

Legend

0 130 260 520 780 1,040 MJ] Historical Records

A [ |] State boundary

[ |] Counties

Fic. 1 Documented, LEE . | ty J zr. p J $ L Hi: aL Hi LI ICh 4 I . to 1981.

of extensive surveys for the species in states where there is abundant available habitat, and during years

when habitat conditions are suitable, prevents an accurate picture of the distribution of S. hallii in North

America.

Habitat and Life History/ Ecology

Schoenoplectus hallii is an obligate wetland species (Reed 1988; Swink & Wilhelm 1994; McClain et al. 1997;

McKenzie 1998; Penskar & Higman 2003; Beatty et al. 2004). Reported habitats are often characterized by

fluctuating water levels (Ostlie 1990; Ostlie & Gottlieb 1992; McKenzie 1998; Beatty et al. 2004; O'Kennon

& McLemore 2004). Schoenoplectus hallii has been described as a “specialized, primary successional plant

with a narrow niche" (G. Yatskievych, MOBOT, pers. comm.) that colonizes “sandy pioneer habitat” (Schuyler

1969), and it is usually found on bare soil where fluctuating water levels may prevent the establishment

of competing perennials (Schuyler 1969). It has been reported from the edges of ephemeral pools, sink-

hole and sand ponds (Schuyler 1969; Robertson et al. 1994; McClain et al. 1997; McKenzie 1998; Smith

2002b; Beatty et al. 2004), sandy clay ponds (O'Kennon & McClemore 2004) and sand prairies (Schuyler

1969; McKenzie1998), sand pits (Bowles et al. 1990), ditches (Steinauer 2001a; Smith 2002b), wet places

in cultivated fields (Smith 2002b), “silty, muddy flats" (Chester 1988), and “cattle trails that lead through

shallow-water wetlands and other depressions" (Beatty et al. 2004). Occasionally, the species can be found

in rocky or cobble habitat. Rocks are along the edges of the sinkhole pond habitat in Missouri (Schuyler

1969; McKenzie 1998) and the species was associated with *many cobbles" at the recent rediscovery site in

Wisconsin (S.G. Smith, pers. obs.).

During years when spring rainfall or high river levels create suitable wetland habitat, S. hallii can often

462 Journal of the Botanical R h Institute of Texas 1(1)

[ITTI O OO T ð IJ O IO IS

[].]

| EEE

|_| | | |

| | iT]

| | :

[ |

| | | tT

Legend

0 95 190 380 570 760 —— [X] within last 25 years

A [| State boundary

[| Counties

"n J £ C.L lart hall: Ilnitnel Cat

Fic. 2. Documented, last 25 years

be found in abundance in the low depressions of cultivated fields in IL, KY, and MO (Chester 1988; Robert-

son et al. 1994; McClain et al. 1997; Missouri Natural Heritage Program 2006a). In drier years, these areas

lack wetland habitat and are cultivated for crops (Chester 1988; Robertson et al. 1994; McClain et al. 1997).

Although the species is usually associated with sandy soil, it can be found on exposed mud with a high silt

content (Chester 1988). At one site in southern Missouri, Smith (2001) determined that S. hallii does not

survive below 14% soil moisture and inhabits soils with a range of organic matter from 0.3-2.6%.

Schoenoplectus hallii germinates sporadically from year to year depending on the availability of wet, ex-

posed habitat (Schuyler 1969; Penskar @ Higmann 2003; Robertson et al. 1994). In some areas, the species

can disappear for long periods only to reappear when conditions are favorable for germination and growth

(Robertson et al. 1994; Chester 1988; McKenzie 1998; McClain et al. 1997; Penskar @ Higman 2003). In

Illinois, populations can vary from hundreds of thousands of plants covering extensive areas in one year to

being entirely absent in other years (Robertson et al. 1994; McClain et al. 1997; McKenzie 1998; Beatty et

al. 2004).

The exact mechanisms necessary to initiate seed germination and development of mature plants are

only beginning to be understood. Smith (2001, 2002a, 2003) and Baskin et al. (2003) concluded that the

species requires a combination of flooding, ethylene and light for germination, and that dormancy, which

occurs in mature seeds, is more readily broken if flooded conditions occur in late spring and summer

rather than late winter or early spring. Smith & Houpis (2004) investigated gas exchange in response to

vapor pressure deficit in S. hallii and determined that the stomates of the species do not close in response

McKenzie et al., C ti t of Scl lectus hallii in the United States 463

{| [| [Tit |

Legend

EN Within last 5 years

[| State boundary

==] Counties

0 87.5 175 350 525 700

Kilometers

" J ECL | se hAll::tta +b Ilnitnsl Cert OP re | I er

Fic. 3. Documented,

to increasing drought. This condition imposes a constraint on the species, requiring that it take advantage

of optimal transient conditions to complete its lifecycle before conditions become unsuitable.

Studies by Smith (2001, 2002a, 2003) and Penskar & Higman (2003) have determined S. hallii can

produce a seed bank containing thousands of achenes. These achenes apparently remain dormant for de-

cades until conditions are optimal for germination and growth (Robertson et al.1994; McClain et al. 1997;

McKenzie 1998; Beatty et al. 2004; NatureServe 2006). This is a strategy noted for other species of sedges

(Baskin et al. 2000) and plants associated with desert environments (Venable & Lawlor 1980).

The biological and ecological significance of amphicarpy in S. hallii and other members of Schoenoplectus

Section Supini have received little attention in the literature. Browning (1992) suggested that amphicarpy

in the genus Schoenoplectus was environmentally induced due to a decrease in water levels. Others have

postulated that amphicarpy has evolved to protect fruits from herbivory or changes in microclimate (Bruhl

1994; Magrath 2002).

The dispersal mechanism for the transport of achenes of S. hallii is not known, but some have suggested

that the species is spread by migratory waterfowl (McClain et al. 1997; Beatty et al. 2004) that have been

found to transport the seeds of other plant species long distances (deVlaming & Proctor 1968; Dunn and

Knauer 1975; Powers et al. 1978; Kantud 1996). Magrath (2002) postulated that large herbivores such as

cattle and bison were the likely dispersal agents for achenes of S. hallii on the Wichita Mountains Wildlife

Refuge in Oklahoma, and Mike Homoya of the Indiana DNR hypothesized that the discovery of S. deltarum

(Schuyler) Soják in Indiana was due to migrating waterfowl.

464 Journal of the Botanical R h Institute of Texas 1(1)

Associated Species

Schoenoplectus hallii is usually associated with other wetland-adapted plants. Although associates have not

been provided for many sites, detailed data collected at others (e.g., Voss 1967; Robertson et al. 1994; Swink

& Wilhelm 1994; McClain et al. 1997; Steinauer 2001a; Penskar & Higman 2003; O'Kennon & McLemore

2004) reveal the following commonality of wetland-adapted taxa: Agrostis spp., Alisma spp., Ammania spp.,

Bacopa spp., Cyperus spp., Echinochloa spp., Eleocharis spp., Fimbristylis autumnalis (L.) Roemer & Schultes,

Heterantha spp., Hypericum spp., Isoetes spp., Juncus spp., Leersia spp., Lindernia spp., Lipocarpha micrantha

(M.Vahl) G. Tucker, Ludwigia spp., Lycopus spp., Polygonum spp., Rhexia spp., Rhynchospora spp., Rorippa

spp., Rotala ramosior (L.) Koehne, Sagittaria spp., Schoenoplectus spp., Scirpus spp., Typha spp., and Xyris spp.

Echinodorus tenellus (Mart.) Buchenau var. parvulus (Engelm.) Fassett, a species for which the U.S. Fish &

Wildlife Service (USFWS) has concern and which is listed as endangered in most mid-western states, has

been recorded at S. hallii sites in GA, KY, MI, and at five sites in Missouri (Voss 1967; Chester 1988; Robertson

et al. 1994; McKenzie 1998; Penskar & Higman 2003; Missouri Natural Heritage Program 20063).

State Accounts

Alabama.— Previous reports of S. hallii in Alabama are referable to S. erectus (S.G. Smith, pers. comm.)

and recent searches for the species have failed to document its occurrence in the state (A. Schotz, Alabama

Heritage Program, pers. comm .).

Colorado.— Previously published accounts of S. hallii’s occurrence in the state (Harrington 1954; Small

1972; U.S. Fish & Wildlife Service 1993) are based on misdeterminations of S. saximontanus (Schuyler, pers.

comm.; Smith 2002b).

Florida.—There are no documented records of this species in Florida and investigations by Smith

(2002b) have determined that previous records of S. hallii from Florida are referable to S. erectus.

Gual

Georgia. oplectus hallii has been documented solely from Dougherty County (Schuyler 1969;

S.G. Smith, pers. comm.), and the species has not been collected in the state since 1966 (Patrick, pers.

comm.). Previous reports from Decatur and Baker counties have proven to be S. erectus (Schuyler 1969). A

specimen taken in Decatur County by Thorne [collection number 6553 and identified as S. hallii, has not

been located (Patrick, pers. comm.)] nor examined by S.G. Smith (pers. comm.). Given that the specimen

was collected close to a site (and apparently in the same year) where Thorne collected (collection number

6536), a specimen of S. erectus that was misidentified as S. hallii (S.G. Smith, pers. comm.) it is likely that

the material is also S. erectus. Without knowing the correct identification of Thorne's collection number

6553 and with no recent surveys being conducted for S. hallii, the exact status of the species in this state is

unclear (McKenzie 1998; O'Kennon & McLemore 2004).

Illinois.—The type specimens for Schoenoplectus hallii were taken in Illinois (Gray 1863; Winterringer

1959; Schuyler 1969; Mohlenbrock 1976), and more records of the species have been documented here than

in any other state. It was historically recorded from Cass (Winterringer 1959; Schuyler 1969; Mohlenbrock

& Ladd 1978), Mason (Winterringer 1959; Schuyler 1969; Mohlenbrock 1976; Mohlenbrock & Ladd 1978),

and Menard (Winterringer 1959; Mohlenbrock 1976; Mohlenbrock & Ladd 1978) counties (Table 2). The

species was discovered in Alexander County in 1993 (T. Kleninger, Illinois Natural Heritage Database, pers.

comm .), and it was documented at 27 sites in Cass, Kankakee, Mason and Morgan counties following surveys

in 1993 (Robertson et 21.1994). Approximately 29 sites have been documented in the state within the last 25

years (Kleninger, pers. comm.) (Table 1). Populations ranged in size from *one plant to thousands of plants

forming nearly pure stands and covering several acres" (Robertson et al. 1994). In 1995, when habitat was

optimal for the species, some populations covered several acres and included hundreds of thousands of

plants that were observed in flooded agricultural fields that were too wet for farm equipment (McClain et

al. 1997; McKenzie 1998). Searches for S. hallii in Illinois in 1996 at the same sites yielded only one plant

and none in 1997 (B. McClain, Illinois DOC, pers. comm .). It has apparently been observed at only one site

in Mason County within the last five years (Bill McClain, pers. comm. 2006).

Indiana.—Schoenoplectus hallii was first discovered in Porter County in 1981 by Dritz et al. (Bowles et al.

McKenzie et al., C ti t of Scl lectus hallii in the United States 465

1990; Swink & Wilhelm 1994). It was later discovered at one site in Lake County and at two additional sites

in Porter County (Swink & Wilhelm 1994; Homoya, pers. comm.; R. Hellmich, Indiana DNR, pers. comm .).

A sixth population was found in Daviess County in 2002 (Hellmich, pers. comm.). Population estimates at

these sites range from “a few plants” to *10,000- plants at the new site in Daviess County (Homoya, pers.

comm.; Hellmich, pers. comm.). In 2006, the species was observed at the Daviess County site on 14 June,

which is the earliest date recorded for this species in the Midwest. Because there are no active searches for

S. hallii, itis not known if the species is more widely distributed in Indiana.

Iowa.—lowa was mistakenly omitted from the range map for S. hallii in Smith (2002b). An 1890 record

for Muscatine County (Davidson 1959; Guldner 1960; Roosa et al. 1989) was recently confirmed (S.G. Smith,

pers. comm.). A very immature but identifiable Reppert s.n. collection taken from Muscatine County in

1890 was discovered and examined by S.G. Smith in 1997 at the Putnam Museum of History and Natural

Science in Davenport, lowa (S.G. Smith, pers. comm.). The specimen that is the basis for a 1960 report from

Louisa County (Roosa et al. 1989), however, has not been located nor verified (S.G. Smith, pers. comm.; John

Pearson lowa DNR, pers. comm.). Although Roosa et al. (1989), Robertson et al. (1994), McKenzie (1998),

and Beatty et al. (2004) reported that S. hallii was probably extirpated from the state, it is listed as a species

of *Special Concern" (Iowa Department of Natural Resources 2007) and Pearson believes that there is still

sand prairie habitat in Muscatine and Louisa counties that should be searched (pers. comm.).

Kansas.— Historically, S. hallii was reported from five counties (Great Plains Flora Association 1977); but

it has only been documented from three counties based on correctly-identified voucher specimens: Harper,

Harvey and Reno (R. McGregor, C. Freeman and C. Morse, University of Kansas, pers. comm.; S.G. Smith,

pers. comm.). Population estimates range from “a few plants" to “abundant” (McGregor and Freeman, pers.

comm .). The species has not been observed in the state since 1997 despite intensive surveys of sand prairie

communities by several investigators (Freeman, pers. comm .).

Kentucky.—Schoenoplectus hallii was first discovered in Kentucky by Chester in 1983 in Christian

County (Chester 1988). Since its original discovery, Chester has annually monitored the population at the

only known site in the state, where the numbers of plants have been estimated in the thousands (Chester,

Austin Peay State Univ., pers. comm.).

Massachusetts.—The occurrence of S. hallii in Massachusetts is based solely on historical collections

made in Middlesex and Essex counties. The species was recorded from along the shoreline of Winter Pond

in Middlesex County between 1876 (Schuyler 1969) and 1931 (Ostlie 1990). A second, undated specimen

taken from Essex County by Horner at S. Georgetown and reported by Sorrie (1987) and Ostlie (1990) is

housed at the New England Botanical Club herbarium and has been confirmed by Schuyler (pers. comm.).

Sorrie (1987) reported that S. hallii has been extirpated from Massachusetts and stated that recent efforts to

find suitable habitat for the species had failed. Consequently, the species is believed to have been extirpated

in the state (McKenzie 1998; Smith 2002b).

Michigan.—First recorded at Carr Lake in Muskegon County in 1900 (Voss 1967), S. hallii has since

been documented at four additional sites: two in Muskegon County and two in Allegan County (Brodowicz

1990; Penskar, Michigan Natural Features Inventory, pers. comm.; Penskar & Higman 2003). The species

was last collected in Muskegon County in 1988 and in Allegan County in 2002 (Penskar, pers. comm.). In

favorable years, S. hallii can be abundant at Michigan sites. Brodowicz (1990) estimated “hundreds of plants"

at a Muskegon County site he visited in 1988 and Penskar (pers. comm.) reported that numbers at Michigan

sites range from a few stems to tens of thousands of plants. Several years of intensive searching by experts

have failed to yield any additional populations, but Penskar & Higman (2003) reported that suitable habitat

exists that has not yet been surveyed. The failure to observe the species in Michigan since 2002 may be due

to the drought that persists at some localities in the Midwest (Penskar & Higman 2003).

Missouri.—Historically, S. hallii was known from Howell and St. Louis counties (Steyermark 1963).

The inscription “hills west of St. Louis" on a collection by Englemann in 1845 (five specimens in different

herbaria), which was cited in the original description, provides documented evidence of the species' oc-

466 Journal of the Botanical R h Institute of Texas 1(1)

currence in St. Louis County (Yatskievych, pers. comm.). There are currently 10 extant sites in the state:

three are located along the edges of sinkhole ponds in Howell County and seven are along the edges of sand

depressions, swales, or ponds in Scott County (Missouri Natural Heritage Program 20062). Population size

fluctuates widely from year to year at most of the extant sites. As few as four plants have been documented

at one of the sinkhole pond sites (T. Smith, Missouri DOC, pers. comm.), but hundreds of thousands of

plants were estimated in 1998 and 2002 at sites in Scott County during years when conditions were optimal

for the species (Missouri Natural Heritage Program 20063). While sink-hole pond habitats in the state have

been intensely surveyed, additional searches in appropriate sandy swale habitat in Butler, Clark, Lewis, Mis-

sissippi, New Madrid, Pemiscott, Ripley, Scott, and Stoddard counties are warranted. Only one population

was located during a search of extant sites in Scott County in August 2006 (McKenzie, pers. obs.). The lack

of occurrence at additional sites in 2006 is undoubtedly due to the dry conditions prevailing at the sites.

Nebraska.—Although The Great Plains Flora Association (1977) listed S. hallii from nine counties in

Nebraska, Schuyler (1969) and Rolfsmeier (1995) determined that the species was historically known only

from Holt and Rock counties. Other reported county records for Nebraska were based on misdetermina-

tions of S. saximontanus (Rolfsmeier 1995; Schuyler, pers. comm.; S.G. Smith, pers. comm.). In 1999, two

populations were discovered in Holt and Brown counties (Steinauer 20012). Steinauer (20012) subsequently

conducted a survey for S.hallii in 2000 and found an additional 18 populations, where plant numbers ranged

trom 2 10-9,000+,

In 2001, Steinauer (2001b) discovered another four populations of Schoenoplectus hallii. Steinauer

(2001a, b) established new county records for the species in Garfield, Loup, and Wheeler counties (Table 3).

Twenty-seven extant sites of S. hallii have been documented in the state within the last 25 years, scattered

throughout six counties in the sandhills region of northcental Nebraska (Table 1) (R. Schneider, Nebraska

Natural Heritage Program, pers. comm.). Because it has been estimated that there are 19,300 square miles

(Knue 1997) of sandy habitat within the sandhills region of the state, ongoing surveys are likely to yield

additional new populations of this species (Steinauer 20013). Further surveys are necessary before the status

of this species in the state can be more accurately assessed.

Ohio.—A report that the species occurs in Ohio (NatureServe 2006) is in error.

Oklahoma.—Prior to 2000, there were five reports of S. hallii recorded for Oklahoma from Atoka,

Comanche, Johnston, and Woods counties (Watson 1993; L. Watson, Oklahoma Biological Survey, pers.

comm .; P. Hernandez, pers. comm.; L. Magrath, University of Science and Arts of Oklahoma, pers. comm.).

Of these, three have been confirmed by S.G. Smith as S. hallii; one each from Atoka, Comanche, and Johnston

counties (S.G. Smith, pers. comm.). The others, one from Comanche County and one from Woods County,

were determined to be S. saximontanus (S.G. Smith, pers. comm.). In 2000, Magrath and refuge staff of the

Wichita Mountains Wildlife Refuge conducted surveys for S. hallii and S. saximontanus at 134 ponds on the

refuge (Magrath 2002). Schoenoplectus hallii was documented at 14 ponds, S. saximontanus at 10, and both

species at four ponds. Plants were typically found on ponds that were subject to seasonal drawdowns, and

population estimates ranged from one plant to several thousand (Magrath 2002). Despite being documented

at 14 ponds, personnel ofthe Oklahoma Natural Heritage Inventory in Norman consider the Wichita Moun-

tains Wildlife Refuge to have only one site until further examination of the populations on the Refuge can

be undertaken (B. Hoagland, pers. comm .).

Magrath (2002) noted the destruction of some populations of S. hallii on the refuge by livestock, but

acknowledged that such animals may benefit the species by transporting achenes to other sites. In 2001, the

sites on the refuge were revisited to assess the status of S. hallii and S. saximontanus, and the predominance

of abnormal and aborted achenes was noted in several plants of both species (Smith et al. 2004). In 2002,

specimens of S. hallii, S. saximontanus and potential hybrids were collected (Smith et al. 2004) and those with

unusual achenes were independently determined by S.G. Smith and Schuyler as the first putative hybrids

recorded between S. hallii and S. saximontanus (Smith et al. 2004).

Further studies on the potential of hybridization of the two species on the Wichita Mountains Wildlife

McKenzie et al., C ti t of Scl lectus hallii in the United States 467

Refuge are warranted, as the possibility of hybridization (Seehausen 2004) may threaten the continued

existence of S. hallii in Oklahoma. Additional surveys should be conducted in Oklahoma populations and

confirmed sites should be monitored. Until such investigations are conducted and the extent of hybridiza-

tion is determined, the status of S. hallii in Oklahoma will remain unknown.

Oregon.— Koyama (1962) erroneously reported S. hallii from Oregon by misreading the label on a

collection made by J. Zimmerman (3444) near the small town of Oregon in Dane County, WI (Schuyler

1969).

South Carolina.—Previous reports for this species in South Carolina were based on misdeterminations

of S. erectus (S.G. Smith, pers. comm.).

South Dakota.—Previously published reports of S. hallii in South Dakota are based on misdetermina-

tions of specimens of S. saximontanus (S.G. Smith, pers. comm.; Schuyler, pers. comm.).

Texas.—Prior to 2003, it was concurred that all historical records of S. hallii for Texas were referable

to either S. erectus or S. saximontanus (S.G. Smith, pers. comm.; Schuyler, pers. comm.). The species was cor-

rectly reported in the state in 2003, when it was discovered in Wise County (O'Kennon & McLemore 2004).

The species is known to persist at three ponds on the Lyndon B. Johnson National Grasslands between April

and December (O'Kennon & McLemore 2004), but it may be present throughout the year (O'Kennon, pers.

comm .). O'Kennon & McLemore (2004) noted that S. hallii occurred along the margins of sandy clay ponds

that have widely fluctuating water levels. They estimated that there were ca. 200 individuals at each of the

small ponds, which merge into one site during periods of high water. The site is managed by the Forest

Service (O'Kennon & McLemore 2004).

Wisconsin.—Schoenoplectus hallii was first collected in Wisconsin in 1950 by J. Zimmerman at Lake

Barney in Dane County and it was rediscovered at Lake Barney in 1996 by S.G. Smith and J. Dobberpuhl

(S.G. Smith, pers. comm.). There were few plants at this small site in 1996 and they were apparently dwarfed

due to grazing by cattle or horses. Without some means of properly managing this site, S. hallii is threatened

with extirpation (S.G. Smith, pers. comm .). S.G. Smith, with J. Laatsch of the Wisconsin DNR, searched

the shore of Lake Barney in 2006 and did not find S. hallii, perhaps because of turbid water that obscured

much of the shore vegetation (S.G. Smith, pers. comm ).

Previous evaluations by the USFWS

Schoenoplectus hallii was listed as a category 2 candidate species in the USFWS's 1993 Plant Candidate Review

for Listing as Endangered or Threatened Species (58 FR 51143-U.S. Fish & Wildlife Service 1993). Category

2 candidate species comprised taxa for which information indicated that a proposal to list as endangered or

threatened was possibly appropriate, but for which conclusive data on biological vulnerability and threats

were not currently available to support proposed rules. In 1995, the USFWS's Columbia, Missouri Field Of-

fice began a status review of S. hallii and solicited information from species experts and botanists throughout

the range of the species. Data were summarized in an initial range-wide status assessment completed in

1998 (McKenzie 1998).

On 5 Dec 1996, the Director of the USFWS established new policy on the definition of candidate spe-

cies (formerly category 1) and outlined how the USFWS would consider species for which they remained

concerned (formerly category 2 or C2) (61 FR 64481-U.S. Fish and Wildlife Service 1996). Under the new

policy, candidates are defined as those species for which the USFWS has on file sufficient information on

biological vulnerability and threats to support issuance of a proposed rule to list as endangered or threatened,

but issuance of the proposed rule is precluded by other listing actions.

Former C2 species for which the USFWS lacks sufficient information to classify as candidate species will

no longer be enumerated on an official list. Nonetheless, other agencies that have developed extensive data

bases on former C2 species (e.g., The Nature Conservancy; state natural heritage programs) will continue

to monitor these species and maintain communication with the USFWS to help determine when sufficient

information is available to warrant their addition to the USFWS' list of candidate species.

468 Journal of the Botanical R h Institute of Texas 1(1)

Summary of Threats

A. The present or threatened destruction, modification, or curtailment of the species’ habitat or

range.—Although a clear picture of the overall distribution of S. hallii is currently not possible without

additional survey work being conducted during years when habitat is suitable, several known populations

are subject to a multitude of threats. The greatest threats to Hall's bulrush involve the destruction and/or

alteration of its wetland habitat (Ostlie & Gottlieb 1992; Robertson et al. 1994; McKenzie 1998; Penskar &

Higman 2003; Beatty et al. 2004; NatureServe 2006). Range-wide, the lowering of water tables, depletion

of ground water for irrigation, and changes to the hydrology supporting S. hallii habitat may be the most

important factors that threaten the continued existence of the species (McKenzie 1998; Steinauer 2001a;

Beatty et al. 2004). Any alteration to the hydrology that continually supports this habitat would disrupt

the cycles necessary for this species’ germination and establishment (Ostlie 1990; Robertson et al. 1994;

Steinauer 20013). Freeman (pers. comm. in Ostlie 1990; Morse, pers. comm.) postulated that groundwater

depletion in Kansas may have lowered the water table such that wetland habitat favorable for the species is

now being created less often than occurred historically, and Knue (1997) implied that ditching and pumping

water from groundwater sources within the Nebraska sandhills for agriculture and domestic livestock may

negatively impact wetland habitat. Many records of S. hallii for Nebraska are from this region of the state

(Rolfsmeier 1995). The depletion of groundwater aquifers has also been identified as a threat to S. hallii in

Illinois (S. Horn, Illinois TNC, pers. comm.). In Missouri, center-pivot irrigation may be lowering under-

ground aquifers that contribute to flooded conditions in sand prairie swales (Bob Gillespie, Missouri DOC,

pers. comm .). The alluvial aquifer in Missouri provides more than 7 billion gal of water per day for row crop

agriculture in the area (Kleiss et al., 2000), resulting in a recession of groundwater from the shallow ponds

in the region. Smith & Houpis (2004) determined that S. hallii did not respond physiologically to drought

conditions, but continued to transpire until plants were desiccated. Because of this failure to respond to

dry conditions, Smith (2003) concluded that for the species to complete its life cycle, groundwater levels

must remain within 1 m of the surface throughout the growing season in Missouri, and Steinauer (20012)

suggested that elevated ground water levels in excess of 1 m above normal water levels were necessary to

provide habitat in the Nebraska sandhills.

Schoenoplectus hallii thrives when fluctuating water levels create the temporary, wet habitat that prevents

the establishment of competing perennials (Ostlie 1990; Robertson et al. 1994; McClain et al. 1997; Schuyler

1969). Robertson et al. (1994), however, noted that inundation will kill flowering plants of S. hallii if rising

waters overtop the plants following germination and establishment. Schuyler (pers. comm.) postulated that

the permanent inundation of the only known historical site in Massachusetts is responsible for the disap-

pearance of the species there. Schuyler (1969) stated, *It appears that S. hallii grows in unstable habitats of

sandy substrates, pioneer habitats from the standpoint of plant succession, which are well-suited for the

growth of S. hallii but few other species of flowering plants. The restriction of S. hallii to this kind of unstable

sandy pioneer habitat probably accounts for its unusual localized distribution."

The habitat of S. hallii is threatened by residential, commercial, agricultural, and recreational develop-

ment (McClain et al. 1997; McKenzie 1998; Beatty et al. 2004). In Kentucky, the largest portion of the only

known site in the state was recently destroyed during activities associated with a new truck stop adjacent

to a major highway (D. White, Kentucky State Nature Preserves Commission, pers. comm.). The remaining

portion of the site is also threatened from the tillage associated with planting crops (Chester 1988). Although

Illinois supports the largest number of populations of S. hallii within its range, the species’ existence in that

state is threatened with the continued and widespread alteration of its habitat for agricultural and residen-

tial development (McKenzie 1998; Beatty et al. 2004; NatureServe 2006). The largest populations known

for the species in 1995 were destroyed in Illinois due to tillage in 1996 and 1997, when drier conditions

enabled farmers to plow wetland habitat (McClain et al. 1997; McKenzie 1998). Some sites in the state are

threatened due to the filling of sand ponds for development and agriculture and the drainage of wet sand

habitat to increase agricultural production (McClain et al. 1997).

McKenzie et al., C ti t of Scl lectus hallii in the United States 469

Schoenoplectus hallii in Michigan is subject to threats in the state including dredging and filling operations

associated with residential development (Crispin & Penskar 1990), off-road vehicle use and trail bikes, (Ostlie

1990; Penskar & Higman 2003), and the potential alteration of the natural hydrology of wetland habitat

by local industries (Crispin & Penskar 1990; Penskar & Higman 2003). The only site in Porter County, IN

is also threatened due to heavy recreational use (Homoya, pers. comm.), and Steinauer (20012) identified

ditch maintenance as a potential threat to populations in Nebraska that occur in road-side ditches. Penskar

(pers. comm.) noted that one site on private property in Michigan is negatively impacted through shoreline

modifications by heavy equipment.

Heavy grazing has been noted at sites in KS, MO, NE and WI, but it is not known whether this type of

disturbance negatively impacts the species (Freeman, pers. comm.; Steinauer 2001a; Magrath 2002). Some

populations of S. hallii are threatened from various chemical contaminants or herbicides. Portions of one of

the larger Illinois’ populations were destroyed in 1995 due to application of post-emergent herbicides (B.

Meyers-Croteau, Illinois State University, pers. comm.). A historical site in Massachusetts was negatively

impacted by storm-water runoff, septic effluent, and herbicides that were used to control unwanted species

(Sorrie 1987; B. Sorrie, pers. comm., in Ostlie 1990). The application of numerous chemicals associated with

agricultural practices throughout the species’ range may inhibit achene germination (Rojas-Garciduenas et

al. 1962; Kozlowski and Sasaki 1968), seedling development (Eliasson & Palen 1972) and growth (Musarrat

& Haseeb 2000), or prevent sexual reproduction (Nartvaranant et al. 2004).

Woody encroachment and the spread of exotic plants have been identified as threats to S. hallii in various

portions of the species’ range (Sorrie 1987; McKenzie 1998; Steinauer 2001a; Beatty et al. 2004; Rolfsmeier

& Weedon 2005). It is threatened by late-successional perennials in areas where early to mid-successional

habitat is not maintained or regulated (Bowles et al. 1990; Robertson et al.1994). Purple loosestrife (Lythrum

salicaria L.) was identified as a threat to S. hallii in Indiana, Massachusetts and Nebraska (Sorrie 1987; McK-

enzie 1998; Steinauer 2001a; Beatty et al. 2004; Rolfsmeier & Weedon 2005). Sorrie (pers. comm. in Ostlie

1990) suggested that the establishment of Lythrum salicaria at a historical Massachusetts site prevented any

reestablishment of S. hallii there, and Bowles et al. (1990) asserted that a Lake County, IN site was also

threatened by this exotic species. Rolfsmeier & Weedon (2005) noted that leafy spurge (Euphorbia esula L.)

may be a potential threat to S. hallii in the future in Nebraska as this aggressive exotic is near extant sites.

The species is also threatened by competition from reed-canary grass (Phalaris arundinacea L.) and various

buckthorn (Rhamnus) species (S.G. Smith, pers. comm .).

An analysis was recently conducted in Missouri that outlined all known threats to sand prairies in the

state, which included seven of the ten known S. hallii sites. Threats identified that were not discussed above

included: 1) changes in ownership that could result in a lack of management, 2) land leveling associated

with agricultural and industrial operations that modify or destroy sandy swale habitats, 3) disruption of

overland flood events by flood control infrastructures (levees, ditches, berms, etc.), 4) reductions in the

frequency of overland flood events necessary to create suitable habitat for germination and plant growth,

5) destruction of sandy habitats due to quarrying for sand or disposal of garbage or refuse, 6) pollution of

groundwater or surface waters that support the species, 7) the loss of landowner agreements or management

and support capabilities, 8) disinterest of landowners and conservation land managers in the development

and implementation of management techniques necessary to maintain the habitat, and 9) the lack of outreach

support to the public on the importance of maintaining sandy swale habitat (Gillespie, pers. comm .).

While the distribution of S. hallii is not well understood, the threats to many of the populations are of

such magnitude that proper maintenance and management of habitat is needed to ensure the continued

existence of this species.

B. Over-utilization for commercial, recreational, scientific, or educational purposes.—There is little

evidence that this species is being negatively impacted due to over- utilization for commercial, recreation,

scientific or educational purposes.

470 Journal of the Botanical R h Institute of Texas 1(1)

C. Disease or predation.—1t has been postulated by S.G. Smith (pers. comm.) that S. hallii may be threatened

by predation from increasing populations of mute swans (Cygnus olor) and Canada geese (Branta canadensis)

These waterfowl species readily feed on the vegetation and achenes of bulrushes (Martin et al. 1951). In ad-

dition, climate models for the Midwest predict that the increasing incidence of extreme weather events will

cause an increase in the number of insect pests that damage native vegetation (Rosenzweig et al. 2000).

eC]

D. The inadequacy of existing regulatory mechanisms. tus hallii currently has a NatureServe

ranking of G2 (globally imperiled because of rarity or some factor(s) making it very vulnerable to extinction

or elimination) and is listed as S1 (critically imperiled) in all states where it is considered extant, except for

Michigan, Missouri and Nebraska where it is listed as S2 (imperiled) (Missouri Natural Heritage Program

2006b; NatureServe 2006). These rankings, however, do not provide any regulatory protection for S. hallii

or its habitat.

In some states, S. hallii is given special designations separate from the Nature Conservancy ranking.

Schoenoplectus hallii is listed as endangered in Kentucky (Kentucky State Nature Preserves Commission

2007), a species of special concern in Iowa (Iowa Department of Natural Resources 2007), threatened in

Illinois (Herkert & Ebinger 2002; Illinois Endangered Species Protection Board 2007), endangered in In-

diana (Indiana Department of Natural Resources 2007), threatened in Michigan (Michigan Department of

Natural Resources 2007; Michigan Natural Features Inventory 2007), status unknown in Oklahoma and

Texas (Oklahoma Biological Survey 2007; The Nature Conservancy of Texas 2007), and endangered in

Wisconsin (Wisconsin Department of Natural Resources 2007).

In Illinois, some protection is provided to any plant species that is state listed by the Illinois Endangered

Species Protection Board (2007) following regulations outlined under the Illinois Endangered Species Pro-

tection Act (Illinois Department of Natural Resources 20072; Michigan State University 2007). Under this

statute, individuals are prohibited from: 1) taking state listed plants without the expressed written permis-

sion of the landowner or 2) selling or offering for sale plants or plant products of endangered species on the

Illinois list (Illinois Department of Natural Resources 20072; Michigan State University 2007). Additionally,

consultation is required for any state funded project that could adversely affect state listed species (Illinois

Department of Natural Resources 2007b).

Little protection is afforded S. hallii under Indiana law, but personnel with the Indiana Department

of Natural Resources have an opportunity to provide input on state-funded projects that could negatively

impact the species (Homoya, pers. comm.). Endangered or threatened plants can not be taken in Michigan

without a permit (Penskar, pers. comm.), as they are protected under the Endangered Species Act 451 of

1994 (Part 365; Michigan Legislature 2007) of Michigan (Michigan Department of Natural Resources 2007;

Michigan Natural Features Inventory 2007; Michigan State University 2007).

In Nebraska, S. hallii is listed as a Tier 1 At-Risk Species as part of the Nebraska Game and Parks

Commission's Natural Legacy Project (Nebraska Game & Parks Commission 20072). This program is part

of the state's development of a Comprehensive Wildlife Strategy that has been initiated in all 50 states (Ne-

braska Game & Parks Commission 2007b). Nebraska’s listing of S. hallii as Tier 1 At-Risk Species does not

provide any regulatory protection status to the species, but the designation has heightened awareness of the

species management needs and the Nebraska Department of Roads proactively consults with the Nebraska

Game and Parks Commission for highway projects that may impact the species (Schneider, pers. comm.).

Schoenoplectus hallii can not be collected in the state without a permit (Schneider, pers. comm.).

A state listing as endangered provides little protection for S. hallii in Wisconsin under state statutes

29.604 and NR(Natural Resources) 27.03-NR27.07 (Wisconsin Legislature 20072, b; Michigan State Univer-

sity 2007). Under Wisconsin law, the taking of S. hallii is prohibited without a permit under section 27.05.

Permits are not required, however, for persons who want to take this or other state listed species: 1) on

property which they own or lease or for which they have been granted landowner permission, except if the

plants or their progeny are sold or processed, 2) on property that is being used for agriculture, construction,

McKenzie et al., C ti t of Scl lectus hallii in the United States 471

or forestry practices, or 3) on property that is being operated or maintained as a utility facility [Michigan

State University 2007; Wisconsin Legislature 2007b- 27.05 (3)].

State endangered species statutes exist for IN, KS, MO, NE, OK, and TX but plants are not covered

under state laws for these states. A summary of all state statutes is available at the Animal Legal & Historical

Web Center of the Michigan State University College of Law (Michigan State University 2007).

E. Other natural or man-made factors affecting its continued existence.—Population isolation.—Because

of human modification to the landscape (Robertson et al. 1994; McKenzie 1998; NatureServe 2006), suitable

habitat for the species has been reduced to small, isolated transient wetlands. As a result, populations of S.

hallii are small and often widely separated, some by several hundred miles. Reduction in population size is

accompanied by loss of genetic variation, which reduces the ability of the population to adapt to changing

environments and increases the risk of extinction (Barrett & Kohn 1991; Newman & Pilson 1997). Ellstrand

& Elam (1993) concluded that in small populations, fitness is likely to decrease because of the fixation of

deleterious recessive alleles. This was confirmed in a field study of Lolium multiflorum Lam., which had

reduced vigor and reproductive capacity with decreasing population size (Polans & Allard 1989). The loss

of fitness may not become evident until later stages in the species' life cycle. In small populations of Gilia

achilleifolia Benth., inbreeding depression was expressed as decreased survivorship of seedlings, rather than

in seed production or germination (Schoen 1983). Others (Frankel & Soule 1981; Holtsford & Ellstrand

1991) determined that inbreeding depression, and a greater threat of extinction, was higher in populations

with an increased rate of selfing. Although the mating system of S. hallii has not been studied, the terminal

flowers are perfect and have the potential for self fertilization. This subject warrants further study.

Loss of seed bank integrity.—While populations of S. hallii are frequently isolated, in years optimal

for germination, they may be extremely large, presumably due to the regeneration of the population from

a persistent seed bank (McClain et al. 1997). A large, persistent seed bank, however, does not necessarily

result in a restoration of the genetic variation or the vigor of the original population. Wienhold and van der

Valk (1989) determined that the number and density of seeds in a seed bank decreases after 10 or more years

when wetlands are drained for extended periods. It is likely that the same losses apply to habitats occupied

by S. hallii during extended dry years and that such declines are accelerated by such agricultural practices

as plowing, disking, or harrowing that damage seeds in the soil (Smith 2001).

It is generally accepted that older seeds have reduced viability (Roberts 1973) and increased chromo-

some breakage and gene mutation (Murata et al. 1982; Murata et al. 1984). Levin (1990) maintained that

genetic and chromosomal changes associated with a long-lived seed bank provide novel genetic variation for

the evolution of a species and are inherently beneficial. Others disagree with this assessment (e.g., Hamilton

1994) and empirical evidence is inclusive. Mean heterozygosity among plants derived from the seed bank

was lower than that existing in surface plants of Lesquerella fendleri (A. Gray) S. Wats. (Cabin et al. 1998),

while in Clarkia springvillensis Vasek, it was concluded that the seed bank could act as a buffer against the

small population effect by supplying plants with greater heterozygosity than that existing in the above-

ground population (McCue & Holtsford 1998). Populations regenerated from a seed bank sometimes show

evidence of inbreeding depression and a decline in performance of seedlings. After restoration of a popula-

tion of Nymphoides peltata (Gmelin) O. Kuntze from a seed bank, Takagawa et al. (2006) reported that there

was a negative effect of inbreeding depression and noted that genetic variation and growth performance

of seedlings derived from the seed bank were significantly reduced. It is, therefore, important to examine

genetic makeup and chromosomal damage and their effects on survival and genetic composition of popula-

tions of S. hallii, as well as the longevity of seeds in the soil, before it can be assumed that the seed bank is

representative of a viable population.

Hybridization.—A threat of hybridization between S. hallii and S. saximontanus exists in Oklahoma (Smith

et al. 2004) and one site in Kansas where the two species are sympatric (Freeman, pers. comm. 2006). Smith

(2002b) also reported that a specimen taken from the coastal plain of Georgia was intermediate between S.

472 Journal of the Botanical R h Institute of Texas 1(1)

hallii and S. erectus. It is not known if anthropogenic practices have potentially altered the distribution of S.

hallii in relation to its closely related congeners. Magrath (2002) noted that S. hallii and S. saximontanus co-

occurred at four ponds on the Wichita Mountains Refuge and Smith et al. (2004) reported the first putative

hybrids between the two species from material collected on the refuge. Although S. hallii and S. saximontanus

were documented in 1997 from the same locality in Harper County, KS, a cursory examination of specimens

of both species from the locality indicated no evidence of hybridization (Freeman, pers. comm. 2006). A

more organized and thorough examination of individuals in the population at this locality may also reveal

the presence of hybrids as occurred in Oklahoma (Smith et al. 2004). Because waterfowl are known dis-

persal agents for various sedges, including S. hallii (Dunn & Knauer 1975; McClain et al. 1997; Beatty et al.

2004), it is possible that the management of various wetlands/impoundments for waterfowl may increase

the potential of S. hallii mixing with its more common congeners, S. saximontanus and S. erectus.

According to conservation geneticists, hybridization poses a serious threat to the survival of a rare

species that hybridizes with a closely related congener (Levin et al. 1996). Although the extinction of rare

species typically is attributed to systematic environmental change that renders the habitat unsuitable (Har-

rison 1991; National Research Council 1995), hybridization may have a profound effect on the persistence

of a species (Rieseberg 1991; Ellstrand 1992; Rieseberg & Linder 1999).

Hybridization promotes the extinction of rare species by reducing the potential for plants to replace

themselves, thereby inhibiting the growth of their populations—the lower the rate of growth, the greater the

potential for extinction in a variable environment (Menges 1992). Hybridization may reduce a population's

growth rate by adversely affecting its reproductive effectiveness, its competitive status and its interactions

with herbivores (Levin et al. 1996).

The growth rate of a population may be retarded by the production of hybrid seed, which is produced

in place of the rare species, i.e., resources are limited and an investment in hybrid seed reduces the amount

of energy that can be allocated to conspecific seed. The outcome is the same whether the hybrid seeds abort

or are viable. When hybrid seeds are produced by both a rare species and its abundant congener, a numeri-

cally small population will produce a higher percentage of hybrid seed than the more numerous related

species when the two are intermixed (Felber 1991; Levin et al. 1996). Species have a number of defenses

against interbreeding; however, closely related congeners often have weaker barriers to hybridization and a

greater minority disadvantage (Levin et al. 1996).

The numerical disadvantage of a rare species is compounded by the proliferation of fertile hybrids. The

addition of these plants to a population containing two related plants decreases the proportional representa-

tion of the rare species. In time, this backcrossing can result in the assimilation of the rare species whose

genetic identity will become extinct (Rhymer & Simberloff 1996). Over evolutionary time, the DNA of the

former rare species may be lost from the gene pool altogether (Rieseberg et al. 1996).

lectus hallii can disappear from sites during periods of drought, only to

Climate change.—Schoen

reappear when flooded conditions exist (Robertson et al. 1994; McClain et al. 1997; Penskar & Higman

2003; McKenzie 1998; Beatty et al. 2004). Global warming and climate change could contribute to loss of

wetland habitat required by this species by causing droughts to be more extensive and persistent, especially

in the Midwest where drought conditions have persisted for several years, and are predicted to continue with

increasing frequency in the future (Hansen 1989; Rosenzweig et al. 2000). Climate models indicate that

high temperatures and an increase in pests (Rosenzweig et al. 2000) and other invasive species (Vitousek

1994) will accompany the increased frequency of droughts and other extreme events (Dai et al 1996). Global

warming and climate change have increasingly been identified as factors which may contribute to the loss

of biodiversity and extinction of imperiled species (Wilcove et al. 1998; Thomas et al. 2004; Maschinski et

al. 2006), but only Beatty et al. (2004) have addressed the issue for S. hallii.

Changes in crop production and ethanol demand.—Habitat for S. hallii is threatened due to projected

changes in agricultural development, especially corn production that is needed for increasing demands for

ethanol production (Keeney and Mueller 2006; U.S. Department of Agriculture 2007). Keeney and Mueller

McKenzie et al., C ti t of Scl lectus hallii in the United States 473

(2006) estimated a 25496 increase in volume of water used in ethanol production from 1998 to 2008 for

only one state in the Midwest. Corn production for ethanol plants is projected to increase in 2007 by 14.296

in Illinois (U.S. Department of Agriculture 2007), the state with largest number of extant sites for S. hallii

(Table 1). Nebraska has the second largest number of extant sites for the species (Table 1) and large areas

of unsurveyed habitat, but Keeney and Mueller (2006) postulated that increasing demand for ethanol pro-

duction may result in competing water uses in that state. Increased demands for water from above ground

sources and underground aquifers could prevent the formation of important wetland habitat needed by S.

hallii throughout the Midwest.

Ownership, current protective status, habitat protections or management

Few of the extant sites of S. hallii in the United States receive protective status and only a small percentage

of sites documented within the last 25 years are actively managed to maintain habitat for the species. Of

the 84 sites recorded for the species within the last 25 years, approximately 16 (~ 19%) are under public

ownership. The remaining 8196 are privately owned, which could make them potentially vulnerable to

habitat change that may threaten the species at these sites. The only known sites in Kansas and Kentucky

are under private ownership, as are the majority of sites in Illinois, Missouri and Nebraska. The existence of

S. hallii populations on public land, however, does not guarantee that the habitat will be properly managed

and maintained to ensure the persistence of the species. The S. hallii site at Horseshoe Lake Conservation

Area in Illinois is also under public ownership, but there is no management plan for the species (McClain,

pers. comm. 1996).

Schoenoplectus hallii is protected and actively managed at four of the five sites in Michigan. Two sites

on the Allegan State Game Area are under the jurisdiction of the Wildlife Division of the Michigan DNR.

They are protected by barriers that control access, and are regularly patrolled and managed (Penskar, pers.

comm. 2006). One site in Muskegon County is located within a dedicated Research Natural Area on the

USFS's Huron-Manistee National Forest, and a second site is protected as part of a nature preserve owned

by the Michigan Nature Association (Penskar, pers. comm. 2006). Three sites at the Indiana Dunes National

Lakeshore are owned by the National Park Service; however, Bowles et al. (1990) reported that at least one

site is threatened by succession and the spread of purple loosestrife.

The one site for S. halliiin Oklahoma is on a USFWS Wildlife Refuge, but the species is not covered under

a management plan (C. Kimball, Wichita Mountains Wildlife Refuge, pers. comm .). Specific management

actions may be included for the species when the refuge develops a 15-year Comprehensive Conservation

Plan (CCP) (Kimball, pers. comm .), but the initiation of such a plan has not yet been scheduled (U.S. Fish

& Wildlife Service 2007).

Two of the sinkhole pond sites in Howell County, MO are registered under the Nature Conservancy's

Registry Program (T. Smith, pers. comm.), but the program is not regularly maintained (D. Ladd, Missouri

TNC, pers. comm.). Landowner agreements that exist for two sites in Scott County, MO will likely maintain

habitat for the species into the foreseeable future, but continued support for the agreement is unpredictable

if there is a change in ownership (Gillespie, pers. comm.).

In Nebraska, two sites (South Pine WMA and Twin Lakes WMA) are on public land, but one of the sites

is within a few miles of a population of purple loosestrife (Steinauer 20012). The listing of S. hallii as a Tier 1

At-Risk Species will ensure that it is a priority for conservation efforts and a focus for various management

plans (Schneider, pers. comm.; Nebraska Game & Parks Commission 2007c).

The only known S. hallii site in Wisconsin is on prison grounds, which is under public ownership, but

the site has limited access (Anderson, pers. comm.). There is no management plan in place to benefit the

species at this site (Anderson, pers. comm .).

The only known site in Texas is on the Lyndon B. Johnson National Grasslands, but no information on

management actions for the species was identified by O'Kennon and McLemore (2004).

Management actions needed

While the management of S. hallii sites is probably necessary for its continued existence, the success of such

474 Journal of the Botanical R h Institute of Texas 1(1)

efforts may be difficult due to the specific habitat requirements for this wetland-adapted plant (Bowles et al.

1990). Ostlie (1990) recommended that extant sites of the species should be protected from human distur-

bance and that the hydrological cycles necessary for the creation and maintenance of its habitat should be

preserved. Robertson et al. (1994) suggested that the acquisition of extant sites should be a priority. In areas

where land acquisition is not feasible, easements that incorporate various management agreements may be

the best approach for maintaining and protecting S. hallii habitat. The Nature Conservancy’s Registry Program

could be an efficient method of managing populations on private land, but the lack of active participation

due to limitations in time and personnel will limit the usefulness of the program. Various state-initiated

landowner contact programs may also be helpful in managing the species over a long-term basis such as

the Landowner Incentive Program (LIP) implemented in many states. The USFWS's Partners for Fish and

Wildlife Program could be helpful in assisting landowners in the management of S. hallii on private land.

Perhaps the greatest potential for managing habitat to benefit the species will be through the development

and implementation of each state's comprehensive wildlife strategy similar to the one in Nebraska (Nebraska

Game & Parks Commission 2007b).

Management plans should be developed for the few sites that are publicly owned. Such plans should

include the following recommendations: 1) protect and maintain the hydrology essential to the species, 2)

retard plant succession, 3) control competing perennials, especially aggressive exotics, 4) control off-road

vehicle use, 5) develop active public outreach and education programs, and 6) support the development of

long- term monitoring programs and active research on the species.

Research needs

Additional research on the life history and ecological requirements of S. hallii, as undertaken by Baskin et

al. (2003), Penskar & Higman (2003), Smith (2001, 2002a, 2003), Smith & Houpis (2004), Smith et al.

(2006), and as identified by Beatty et al. (2004), are needed to better understand the population dynamics

of this species. Investigations that compare reproduction from achenes versus perennial growth from the

species’ short rhizomes need to be undertaken. Although initial germination and life history dynamics for

terminal achenes have been established (Baskin et al. 2003; Penskar & Higman 2003; Smith 2001, 2002a,

2003; Smith et al. 2006), further research is warranted to more clearly pinpoint the environmental factors

necessary for germination and establishment of S. hallii achenes. The role of amphicarpy in the life history

of the species needs evaluation, including studies of the mating systems of both terminal and basal florets.

Additional research should be initiated to determine the relationship of wetland hydrology to the population

dynamics of S. hallii, especially the role of surface flooding and underground aquifers and groundwater in

the creation of habitat for the species. Studies to identify the dispersal mechanisms that enable the species

to colonize new areas would be valuable. The germination of achenes of Schoenoplectus spp. can apparently

be enhanced when they pass through the digestive tracts of birds (deVlaming & Proctor 1968; Powers et

al. 1978; Kantud 1996). Similar studies involving the achenes of S. hallii should be initiated to determine if

birds help facilitate dispersal of this species.

Examinations of additional specimens are necessary to clarify the historical and current distribution of

the species (e.g., specimens from GA, IA, and OK) (S.G. Smith, pers. comm .). Additional surveys are necessary

to better delineate the range of S. hallii, provided they are conducted during years when habitat conditions

are favorable to the species and specimens can be examined by experts who can accurately distinguish the

species from its close relatives. An examination of precipitation patterns may be useful in predicting what

years the species is more likely to emerge. Voucher specimens should be confirmed by either S.G. Smith or

Ernie Schuyler. The communities and ecological associates that occur with S. hallii should be further char-

acterized, which will enable researchers in the future to develop adequate search-images for the species.

Studies should be initiated that examine the effects of grazing, controlled burning and other types of

disturbance (e.g., off-road vehicle use, trail bikes, burning, mowing, disking) on the species. Population

genetics studies need to be conducted to determine the amount of variation and gene flow within and among

populations. Studies of the population and genetic consequences of recruitment from a long-lived seed

McKenzie et al., C ti t of Scl lectus hallii in the United States 475

bank need to be conducted to clarify the current definition of “extant sites" as represented by the presence

of achenes in the soil. Monitoring efforts need to be initiated for all extant populations, especially those

populations that appear annually and are not as cyclic in their occurrence as at other sites within the species’

range. The report of a putative hybrid between S. hallii and S. saximontanus at the Wichita Mountains Wildlife

Refuge in Oklahoma (Smith et al. 2004) suggests that hybridization is a threat at sites where conspecifics are

sympatric. Thus, further investigations that examine the extent of hybridization at Oklahoma and Kansas

sites should be initiated to determine if S. hallii is threatened from genetic swamping or genetic dilution.

Schoenoplectus hallii would benefit from additional seed viability studies and from research that examines

genetic differences between and among populations. Finally, the impacts of global warming and climate

change on the loss of wetland habitat for the species should be evaluated by conducting studies similar to

analyses for other species (e.g., Hannah et al. 2002; Pearson & Dawson 2003) and incorporating the results

into long-term management plans.

ACKNOWLEDGMENTS

We are grateful to the numerous individuals throughout the range of S. hallii who provided information

contained in this report. We appreciate the excellent input and support of this project provided by Ernie

Schuyler. Mike Penskar of the Michigan Natural Features Inventory and Charles Bryson of the USDA-ARS

Southern Weed Science Research Unit provided constructive reviews. Jill Utrup of the USFWS graciously

produced Figures 2, 3, and 4 for this report. Elderine Milligan of the USFWS edited a final draft of the

document.

REFERENCES

Barrett, S.C.H. and J.R.K. KoHN.1991. Genetic and evolutionary consequences of small population size in plants:

implications for conservation. In: D.A. Falk and K. Holsinger eds. Genetics and conservation of rare plants.

Oxford Univ. Press, New York. Pp. 3-31.

Baskin, C.C., J.M. Baskin, and E.W. Chester. 2000. Effect of flooding on the annual dormancy cycle and on germina-

tion of seeds of the summer annual Schoenoplectus purshianus (Cyperaceae). Aquatic Bot. 67:109-116.

Baskin, C.C., J.M. Baskin, E.W. CHester, and M. SwrrH. 2003. Ethylene as a possible cue for seed germination of Schoeno-

plectus hallii (Cyperaceae), a rare summer annual of occasionally flooded sites. Amer. J. of Bot. 90:620-627.

Beatty, B.L., W.F. JENNINGS, and R.C. RAWLINSON. 2004. Schoenoplectus hallii (Gray) S.G. Sm. (Hall's bulrush): a technical

conservation assessment. Prepared for the USDA Forest Service, Rock Mountain Region, Species Conservation

Report. Peer review administered by Center for Plant Conservation, St. Louis, Missouri.

BEETLE, A.A. 1947. Scirpus. N. Amer. Flora 18(8):481-504.

Bow es, M.L., M.M. DeMauro, N. PAvLovic, and R.D. Hiesert. 1990. Effects of anthropogenic disturbances on endan-

gered and threatened plants at the Indiana Dunes National Seashore. Nat. Areas J. 10:187-200.

Bropowicz, W. 1990. Noteworthy collections. Scirpus hallii A. Gray. Michigan Botanist 29:31.

BRowNING, J. 1992. Hypogynous bristles and scales in basal florets in amphicarpous Schoenoplectus species (Cy-

peraceae). Nordic J. Bot. 12:171-175.

BruHL, J.J. 1994. Amphicarpy in the Cyperacae, with novel variation in the wetland sedge Eleocharis caespitosis-

sma. Austral. J. Bot. 42:441—448.

CABIN, R.J., R.J. MircH&LL, and D.L. MitcHett. 1998. Do surface plant and soil seed bank populations differ genetically? A

multipopulation study of the desert mustard Lesquerella fendleri (Brasicaceae). Amer. J. Bot. 85:1098-1109.

CHESTER, EW, 1988. An unusual habitat type for three rare Kentucky plants. Abstract. Trans. Kentucky Acad. Sci.

49(102):56.

Correll, D.S. and M.C. Johnston. 1970. Manual of the vascular plants of Texas. Texas Research Foundation,

Renner.

Crispin, S. and M. Penskar. 1990. Scirpus hallii Gray (Hall's bulrush). Unpubl. abstract, Michigan Natural Features

Inventory, endangered species manual.

476 Journal of the Botanical R h Institute of Texas 1(1)

Dal, A, LY. FUNG, and A.D. Det Genio. 1996. Surface observed global land precipitation variations during 1900-1988.

J. Climate 10:2943-2962.

DAVIDSON, R.S. 1959. The vascular flora of southeastern lowa: Univ. lowa Stud. Nat. Hist. 20(2):1-102.

DEVLAMING, V. and V.W. Proctor. 1968. Dispersal of aquatic organisms: viability of seeds recovered from the drop-

pings of captive killdeer and mallard ducks. Amer. J. Bot. 55:20-26.

Dunn, D.B. and D.F. Knauer. 1975. Plant introductions by waterfowl to Mingo National Wildlife Refuge, Missouri.

Trans. Missouri. Acad. Sci. 9:27-28.

ELIASSON, L. and K. Paten. 1972. Effect of light on the response of pea seedling roots to 2,4—D. Physiol. Pl. 26:

206-209.

ELSTRAND, N.C. 1992. Gene flow by pollen: implications for plant conservation genetics. Oikos 63:77-93.

ELLSTRAND, N. and D. Eram. 1993. Population genetic consequences of small population size: implications for plant

conservation. Ann. Rev. Ecol. Syst. 24:217-242.

FELBER, F. 1991. Establishment of a tetraploid cytotype in a diploid population: effect of the relative fitness of the

cytotypes. J. Evol. Biol. 4:195-204.

FLORA OF NORTH AMERICA EDITORIAL Committee, eds. 2002. Vol. 2. Magnoliophyta: Commelinidae (in part): Cyperaceae.

Oxford University Press, New York.

FRANKEL, O.H. and M.E. Soule. 1981. Conservation and evolution. Cambridge University Press. Cambridge.

GLEASON, H.A. AND A. CRONQUIST.1991. Manual of vascular plants of northeastern United States and adjacent Canada,

ed. 2. New York Botanical Garden, Bronx.

Gray, A. 1863. Manual of botany of the northern United States. Third edition. New York.

GREAT PLAINS FLORA AssociarioN. 1977. Atlas of the flora of the Great Plains; R.L. McGregor, Coordinator; T.M. Barkley,

ed. lowa State University Press, Ames.

GREAT PLAINS FLORA ASSOCIATION. 1986. Flora of the Great Plains; R.L. McGregor, Coordinator; T.M. Barkley, R.E. Brooks,

and E.K. Schofield, eds. University Press of Kansas, Lawrence.

GULDNER, L.F. 1960. The vascular plants of Scott and Muscatine counties. Davenport Mus. Publ. Bot. No. 1.

HAMILTON. M.B. 1994. Ex situ conservation of wild plant species: time to reassess the genetic assumptions and

implications of seed banks. Conserv. Biol. 8:39-49.

HANNAH, L., G.F. Mipa.ev, and D. Mitter. 2002. Climate change-integrated conservation strategies. Global Ecol.

Biogeogr. 11:485-495.

HANSEN, J.E. 1989. The greenhouse effect: Impacts on current global temperature and regional precipitation. In:

The challenge of global warming. Island Press, Washington, DC. Pp. 35-43.

HARRINGTON, H.D. 1954. Manual of the plants of Colorado. Sage Books, Denver, CO.

HARRISON, S. 1991. Local extinction in a metapopulation context: an empirical evaluation. Biol. J. Linnean Soc.

42:7 3-88.

HERKERT, J.R. and J.E. EBINGER. 2002. Endangered and threatened species of Illinois: status and distribution. Vol. 1.

Plants. Illinois Endangered Species Protection Board, Springfield.

HICKMAN, J.C. 1993. Ed. The Jepson manual: higher plants of California. University of California Press, Berkeley.

HoLTSFORD, T.P. and N.C. Ettstrand. 1991. Inbreeding effects in Clarkia tembloriensis populations with different

natural outcrossing rates. Evolution 44:2031-2046.

ILLINOIS ENDANGERED SPECIES PROTECTION BOARD. 2007. 2006 endangered and threatened species list. Springfield. http://

www.dnrstate.il.us/espb/datelist.htm. Accessed Jan 2007.

ILLINOIS DEPARTMENT OF NATURAL Resources 2007a. Illinois Endangered Species Board. Wildlife- 520 Illinois Compiled

Statute (ILCS) 10/IIlinois Endangered Species Protection Act. Springfield. http://www.ilga.gov/legislation/ilcs/

ilcs3.asp?ActID=1 730&ChapAct=520%26nbsp%3BILCS%26nbsp%3B 1 0%2F&ChapterID=43&ChapterName=

WILDLIFE&ActName-zIllinois--Endangered--Species--Protection--Act962E. Accessed Apr 2007.

ILLINOIS DEPARTMENT OF NATURAL Resources 2007b. Consultation requirements associated with the Illinois Endangered

Species Protection Act. Springfield. http://dnrstate.il.us/orep/nrrc/briefhtm. Accessed Apr 2007.

McKenzie et al., C ti t of Scl lectus hallii in the United States 477

INDIANA DEPARTMENT OF NATURAL RESOURCES 2007. Endangered, threatened, rare and extirpated plants of Indiana.

Indianapolis. http://www.in.gov/dnr/naturepr/endanger/etrplants.pdf. Accessed Jan 2007.

lowA DEPARTMENT OF NATURAL Resources. 2007. Threatened and endangered species. DesMonies. http://www.iowadnr.

com/other/files/chapter77.pdf. Accessed Jan 2007.

KaNTUD, H.A. 1996. The alkali (Scirpus maritimus L.) and saltmarsh (S. robustus Pursh) bulrushes: a literature review.

National Biological Service, Information and Technology Report 6. Jamestown, North Dakota. Northern Prairie

Wildlife Research Center web database. Available at http://npwrclib.cr.usgs.gov/starweb/pubsearch/servlet.

starweb. Accessed Sep 2006.

Keeney, D. and M. Mutter. 2006. Water use by ethanol plants: potential challenges. Institute for Agriculture and

Trade Policy. Minneapolis, Minnesota.

Kentucky STATE NATURE PRESERVES COMMISSION. 2007. Kentucky rare plant database. http://eppcapps.ky.gov/nprare-

plants/details.aspx?species=Schoenoplectus%20hallii. Accessed Jan 2007.

Kleiss, B.A., R.H. Coupe, G.J. GONTHIER, and B.G. Justus. 2000. Water quality in the Mississippi embayment, Missis-

sippi, Louisiana, Arkansas, Missouri, Tennessee and Kentucky from 1995-1998. Circular 1208, U.S. Geological

Survey, Denver.

KNuE, J. 1997. Nebraskaland magazine wildlife viewing guide. Nebraskaland Mag. 75(1):1-96. Nebraska Game

and Parks Commission, Lincoln.

KovAMA, T. 1962. The genus Scirpus Linn. Some North American aphylloid species. Canad. J. Bot. 40:913-937.

KozLowski T.T. and S. Sasaki. 1968. Germination and morphology of red pine seeds and seedlings in contact with

EPTC, CDEC, CDAA, 2,4-D and Picloram. Amer. Soc. Hort. Sci. 193:655-662.

Levin, D.A. 1990. The seed bank as a source of genetic novelty in plants. Amer. Nat. 135:563-572.

Levin, D.A., J. Francisco-ORTEGA, and R.K. Jansen. 1996. Hybridization and the extinction of rare plant species. Con-

serv. Biol. 10:10-16.

MacRath, L.K. 2002. Schoenoplectus hallii and S. saximontanus. 2000 Wichita Mountain Wildlife Refuge survey.

Oklahoma Native Pl. Rec. 2(1):54-62.

Martin, A.C., H.S. Zim, and A.L. Netson. 1951. American wildlife & plants: A guide to wildlife food habits. Dover

Publ., New York.

MascHINSKI, J., J.E. Bacas, P.F. QUINTANA-AscENCIO, and E.S. Menaes. 2006. Using population viability analysis to predict

the effects of climate change on the extinction risk of an endangered limestone endemic shrub, Arizona

cliffrose. Conserv. Biol. 20:218-228.

McCain, W.E., R.D. McCain, and J.E. EsinGER. 1997. Flora of temporary sand ponds in Cass and Mason counties,

Illinois. Castanea 62:65-73.

McCue, K.A. and T.P. HoLTsFORD. 1998. Seed bank influences on genetic diversity in the rare annual Clarkia springvil-

lensis (Onagraceae). Amer. J. Bot. 85:30-36.

Mckenzie, PM. 1998. Hall's bulrush (Schoenoplectus hallii) status assessment. Unpublished report to the U.S. Fish

& Wildlife Service, Columbia, MO.

MENGES, E.S. 1992. Stochastic modeling of extinction in plant populations. In: Fiedler and Jain, eds. Conservation

Biology Chapman and Hall, New York. Pp. 253-275.

MICHIGAN DEPARTMENT OF NATURAL Resources. 2007. Wildlife Division- endangered and threatened species. Rule 99.1028.

Plants. Lansing. http://www.state.mi.us/orr/emi/admincode.asp?AdminCode=Single&Admin_Num=299010

21&Dpt=NR&RngHigh. Accessed Apr 2007.

MICHIGAN LEGISLATURE. 2007. Natural Resources and Environmental Protection Act 451 of 1994. Part 365.

Endangered species protection. Sections 324.36501-36507. Lansing. http://www.legislature.mi.gov/

(S(jppokk3m3hgmug345wl1pedq5))/mileg.aspx?page-getobject&objectname-mcl-451-1994-iii-1-endan-

gered-species-365. Accessed Apr 2007.

MICHIGAN NATURAL FEATURES INVENTORY. 2007. Michigan's special plants. Lansing. http://web4.msue.msu.edu/mnfi/

data/specialplants.cfm. Accessed Apr 2007.

478 Journal of the Botanical R h Institute of Texas 1(1)

MICHIGAN STATE University. 2007. Animal Legal €. Historical Web Center. Statutes and Laws. East Lansing, MI. http://

www.animallaw.info/statutes/stusmi324_365.htm. Accessed Apr 2007.

Missouri NATURAL HERITAGE PRocRAM 2006a. Missouri species and communities of conservation concern database.

Missouri Department of Conservation. Jefferson City.

Missouri NATURAL HERITAGE PRocRAM 2006b. Missouri species and communities of conservation concern checklist.

Missouri Department of Conservation. Jefferson City.

MoHLENBROCK, R.H. 1976. The illustrated flora of Illinois: Sedges-Cyperus to Scleria. Southern Illinois University Pres.,

Carbondale and Edwardsville.

MonH.LENBROCK, R.H. and D.M. Lapp. 1978. Distribution of Illinois vascular plants. Southern Illinois University Press,

Carbondale.

Murata, M, T. Tsuchiya, and E.E. Roos. 1982. Chromosomal damage induced by artificial seed aging in barley. Il.

Types of chromosomal aberrations at first mitosis. Bot. Gaz. 1413:111-116.

Murata, M., T. Tsuchiya, and E.E. Roos. 1984. Chromosomal damage induced by artificial seed aging in barley

behavior of chromosomal aberrations during plant growth. Theor. Appl. Genet. 67:161-170.

Musarrat, J. and A. Hasees. 2000. Agrichemicals as antagonist of lectin-mediated Rhizobium-legume symbiosis:

paradigms and prospects. Current Sci. 78:793-797.

NARTVARANANT, P.S. HAMILL, J. LEONARDI, A.W. WHILEY AND S. SUBHADRABANDHU. 2004. Seasonal effects of foliar application of

phosphonate on phosphonate translocation, in vitro pollen viability and pollen germination in Hass avocado

(Persea americana Mill). J. Hort. Sci. Biotech. 79:91-96.

NATIONAL RESEARCH COUNCIL. 1995. Science and the endangered species act. National Academy Press, Washington,

DC.

NatureServe. 2006. NatureServe Explorer: An online encyclopedia of life [web application]. Version 5.0 NatureServe,

Arlington, Virginia. Available at: http://www.natureserve.org/explorer. Accessed Sep 2006.

NEBRASKA GAME AND PARKS COMMISSION. 2007a. The Natural Legacy Project: a blueprint for conserving wildlife and

their habitats. Appendix 8: Tier 1 At-Risk Species. Lincoln. http://www.ngpc.state.ne.us/wildlife/programs/

legacy/pdfs/appendix8.pdf. Accessed Jan 2007.

NEBRASKA GAME AND Parks COMMISSION. 2007b. The Natural Legacy Project: a blueprint for conserving wildlife and their

habitats. Lincoln. http://www.ngpc.state.ne.us/wildlife/programs/legacy/about.asp. Accessed Jan 2007.

NEBRASKA GAME AND Parks Commission. 2007c. The Natural Legacy Project: a blueprint for conserving wildlife and

their habitats. Chapter 3: Methodology: identifying ecological communities, at-risk species and biologically

unique landscapes. Lincoln. http://www.ngpc.state.ne.us/wildlife/programs/legacy/pdfs/chpater3.pdf. Ac-

cessed Jan 2007.

NEWMAN, D. and D. PiLsoN. 1997. Increased probability of extinction due to decreased genetic effective population

size: experimental populations of Clarkia pulchella. Evolution 51:354-362.

O'KENNoN, R.J. and C. McLemore. 2004. Schoenoplectus hallii (Cyperaceae), a globally threatened species new for

Texas. Sida 21:1201-1204.

OKLAHOMA BIOLOGICAL SURVEY. 2007. Oklahoma Natural Heritage Inventory: Rare and vulnerable plant species of Okla-

homa. Norman. http://www.oknaturalheritage.ou.edu/scirpus_hal.htmitdistribution. Accessed Jan 2007.

Osrue, W.R. 1990. The Nature Conservancy's element stewardship abstract for Scirpus hallii-Hall's bulrush. Ar-

lington, Virginia.

OsTLIE, W.R. and S. Gorruts. 1992. The Nature Conservancy's element global ranking form for Scirpus hallii.

Pearson, R.G. and T.P. Dawson. 2003. Predicting the impacts of climate change on the distribution of species: are

bioclimate envelope modes useful? Global Ecol. Biogeogr. 12:361-371.

PENskAR, M.R. and PJ. Hicman. 2003. Hall's bulrush habitat characterization and monitoring project: 1999-2002

Report. Region 3 of the U.S. Fish & Wildlife Service Section 6 Endangered Species Grant Report No. 2003-26.

Michigan Natural Features Inventory. Lansing, MI.

PoLANs, N.O. and R.W. ALLARD. 1989. An experimental evaluation of the recovery potential of ryegrass populations

from genetic stress resulting from restriction of population size. Evolution 43:1320-1323.

McKenzie et al., C ti t of Scl lectus hallii in the United States 479

Powers, K.D., R.E. Nose, and R.H. CHasreck. 1978. Seed distribution by waterfowl in southwestern Louisiana. J.

Wildlife Managem. 42:598-605.

RADFORO, A.E., H.E. Ahles, and C.R. Bett. 1964. Manual of the vascular flora of the Carolinas. University of North

Carolina Press, Chapel Hill.

Rayna, J. 1976. Notes Cyperologiques: 26. Le Genre Schoenoplectus Il. 'amphicarpie et la sect. l'amphicarie et

la sect. Supini. Adansonia, ser. 2, 16:119-155.

REED, PE. Jk. 1988. National list of plants that occur in wetlands: national summary. U.S. Fish € Wildlife Service

Biol. Rep. 88(24):1-244. Washington, D.C.

RHYMER, J.M. and D. SimserLOFF. 1996. Extinction by hybridization and introgression. Ann. Rev. Ecol. Syst. 27:

83-109.

RIESEBERG, L.H. 1991. Hybridization in rare plants: insights from case studies in Cerocarpus and Helianthus. In: Genetics

and conservation of rare plants. D.A. Falk and K.E. Holsinger, eds. Oxford University Press, NY. Pp. 171-181.

RIESEBERG, L.H. and C.R. Linder. 1999. Hybrid classification: insights from genetic map-based studies of experimental

hybrids. Ecology 80:361-370.

RIESEBERG, L.H., B. Sinervo, C.R. LINDER, M.C. UnGerER, and D.M. Arias. 1996. Role of gene interations in hybrid speciation:

evidence from ancient and experimental hybrids. Science 272:741-745.

Roserts, E.H. 1973. Loss of seed viability: chromosomal and genetical aspects. Seed Sci. Tech. 1:515-527.

ROBERTSON, K.R., L.R. PHILLIPPE, and S.M. GEHLHAUSEN. 1994. The current status of Scirpus hallii A. Gray, Hall's bulrush, in

Illinois. Illinois Dept. of Conservation Report, Springfield.

RoJAS-GARCIDUENAS, M., M.A. Ruiz, and J. CaniLLo. 1962. Effect of 2,4-D and MCPA on germination and early growth.

Weed Sci. 10:69-71.

ROLFSMEIER, S.B. 1995. Keys and distributional maps for Nebraska Cyperaceae, Part 1: Bulbostylis, Cyperus, Dulichium,

Eleocharis, Eriophorum, Fimbristylis, Fuirena, Lipocarpha, and Scirpus. Trans. Nebraska Acad. Sci. 22:27-42.

ROLFSMEIER, S.B. and R.R. Weebon. 2005. A survey of potential habitat for western prairie fringed orchid (Platanthera

praeclara) along U.S. Highway 20 from Long Pine to Ewing, NE. High Plains Herbarium unpub. Rep., Chadron,

NE.

Roosa, D.M., M.J. Leoscuke, and L.J. Eiters. 1989. Distribution of lowa's endangered and threatened vascular plants.

lowa Dept. of Nat. Res., Des Moines.

ROSENZWEIG, C., A. IGLESIAS, X.B. YANG, PR. EPSTEIN, and E. Chivan. 2000. Climate change and U.S. Agriculture: the impacts

of warming and extreme weather events on productivity, plant diseases and pests. Center for Health and the

Global Environment, Harvard Medical School, Boston, MA.

ScHoEN, D.J. 1983. Relative fitness of selfed and outcrossed progeny in Gilia achilleifolia (Polemoniaceae). Evolu-

tion 37:292-301.

SCHUYLER, A.E. 1969. Three new species of Scirpus (Cyperaceae) in the southern United States. Notul. Nat. Acad.

Nat. Sci. Philadelphia 423:1-12.

SEEHAUSEN, O. 2004. Hybridization and adaptive radiation. Trends Ecol. Evol. 19:198-207.

SMALL, J.K. 1972. Manual of the southeastern flora. Hafner Publ. Co., New York.

SmitH, M. 2001. An assessment of the biological and ecological requirements of Schoenoplectus hallii (Hall's bulrush)

population in Scott County, Missouri. Region 3 ofthe U.S. Fish &Wildlife Service Section 6 Endangered Species

Grant. First Year Progress Report. Submitted to the Missouri Department of Conservation, Jefferson City.

SurrH, M. 2002a. An assessment of the biological and ecological requirements of Schoenoplectus hallii (Hall's

bulrush) population in Scott County, Missouri. Region 3 of the U.S. Fish & Wildlife Service Section 6 Endan-

gered Species Grant Second Year Progress Report. Submitted to the Missouri Department of Conservation,

Jefferson City.

SurrH, M. 2003. An assessment of the biological and ecological requirements of Schoenoplectus hallii (Hall's bul-

rush) population in Scott County, Missouri. Region 3 of the U.S. Fish & Wildlife Service Section 6 Endangered

Species Grant. Final Report. Submitted to the Missouri Department of Conservation, Jefferson City.

SMITH, M. and J.L.). Houeis. 2004. Gas exchange responses of the wetland plant Schoenoplectus hallii to irradiance

and vapor pressure deficit. Aquat. Bot. 79:267-275.

480 Journal of the Botanical R h Institute of Texas 1(1)

SMITH, M., S. AMMANN, N. Parker, and P. MErTTLER-CHERRY. 2006. A quantitative study of styles and achenes of terminal and

basal flowers of Schoenoplectus hallii, a rare plant species of transient wetland habitats. Sida 22:1159-1173.

SMITH, M, P. McKenzie, P. Merr.ER- CHERRY, and G. SurrH. 2004. A putative hybrid of Schoenoplectus saximontanus and

S. hallii (Cyperaceae) from Oklahoma. Sida 21:475-479.

Smith, S.G. 1995. New combinations in North American Schoenoplectus,

(Cyperaceae). Novon 5:97-102.

Smith, S.G. 2002b. Schoenoplectus. Vol. 2. Magnoliophyta: Commelinidae (in part): Cyperaceae. Flora of North

America Editorial Committee, eds. Oxford University Press, New York. Pp. 44-60.

SMITH, S.G. and G. YarskievvcH. 1996. Notes on the genus Scirpus sensu lato in Missouri. Rhodora 98:168-179.

SuirH, S.G. and E. Hayasaka. 2001. Delineation of Schoenoplectus sect. Malacogeton (Cyperaceae) new combination

and distinction of species. J. Jap. Bot. 76:339-343.

Smity, S.G. and E. Hayasaka. 2002. New combinations within Schoenoplectus smithii and S. purshianus (sect. Ac-

taeogoton, Cyperaceae) and comparison with eastern Asian relatives. Novon 12:106-111.

Sonne, B.A. 1987. Notes on the rare flora of Massachusetts. Rhodora 89(858):1 39.

SrEINAUER, R.F. 2001a. 2000 Survey for Schoenoplectus hallii (Gray) S.G. Sm. in the eastern sandhills of Nebraska.

Final report to the Nebraska Game and Parks Commission, Lincoln.

STEINAUER, R.F. 2001 b. 2001 Survey for Platanthera praeclara Sheviah & Bowles in Nebraska & Gaura neomexicana

ssp. coloradensis (Rydb.) Raven & Gregory, Lodgepole Creek, Kimball & Cheyenne County, Nebraska. Final

report to the Nebraska Game and Parks Commission, Lincoln.

STEYERMARK, J.A. 1963. Flora of Missouri. lowa State University, Ames.

STRONG, M. 1994. Taxonomy of Scirpus, Trichophorum, and Schoenoplectus (Cyperaceae) in Virginia. Bartonia

58:29-68.

Swink, F. and G. WitHELM. 1994. Plants of the Chicago region. 4th Ed. Indiana Academy of Science, Indianapolis.

TAKAGAWA, S., l. WASHITANI, R. UEsuai, and Y. TsuMURA. 2006. Influence of inbreeding depression on a lake population

of Nymphoides peltata after restoration from the soil seed bank. Conserv. Gen. 7:705-716.

THe NATURE CONSERVANCY OF TEXAS. 2007. An annotated list of rare plants of Texas. Austin.

THomas, C.D., A. CAMERON, R.E. GREEN, M. BAKKENES, L.J. BEAUMONT, Y.C. COLLINGHAM, B.F.N. ERASMUS, M.F. DE SIQUEIRA, A. GRAINGER,

L. HANNAH, L. HUGHES, B. HUNTLEY, A.S. VAN JAARSVOID, G.F. MiDGLEY, L. Mites, M.A. ORTEGA-HuertA, A.T. PETERSON, O.L. PHILLIPS,

and S.E. WiLLiAMS. 2004. Extinction risk from climate change. Lett. Nature 427:145-148.

Tucker, G.C. 1987. The genera of Cyperaceae in the southeastern United States. J. Arnold Arbor. 68:361—445.

U.S. DEPARTMENT OF AGRICULTURE. 2007. National Agricultural Statistics Service. Prospective plantings. Washington,

D.C. http://www.nass.usda.gov/Newsroom/2007/03_30_2007.asp. Accessed Apr 2007.

U.S. Fisu & WILDLIFE Service. 1993. Endangered and threatened wildlife and plants; review of plant taxa for listing

as endangered or threatened species. Notice of review; 50 CFR Part 17. Federal Register Vol. 58. No. 188, Sep

30, 1993. Pp. 51143-51190.

U.S. Fisu & WiLpure Service. 1996. Endangered and threatened wildlife and plants; notice of final decision on

identification of candidates for listing as endangered or threatened. Notice of review; 50 CFR Part 17. Federal

U.S. Fisu & WILDLIFE Service. 2007. America's National Wildlife Refuge System- refuge planning- by Region. Wash-

ington, D.C. http://www.fws.gov/refuges/refugePlanning/. Accessed Jan 2007.

VENABLE, D.L. and L. LawLor. 1980. Delayed germination and dispersal in desert annuals: escape in space and time.

Oecologia 46:272-282.

Vitousek, PM. 1994. Beyond global warming: ecology and global change. Ecology 75:1861-1876.

Voss, E.G. 1967. The status of some reports of vascular plants from Michigan. Michigan Bot. 6:13-24.

WATSON, L.E. 1993. Monitoring of plant candidate species in Oklahoma-year two. Unpubl. Report to U.S. Fish &

Wildlife Service. Oklahoma Biological Survey, Univ. of Oklahoma, Norman.

Weeer, W.A. and R.C. Wittmann. 1992. Catalog of the Colorado flora: A biodiversity baseline. Univ. of Colorado

Press, Niwot.

DATL L

, Isolepis and Trichophorum

McKenzie et al., C ti t of Scl lectus hallii in the United States 481

WiENHOLD, C.E. and A.G. VAN DER VALK. 1989. The impact of duration of drainage on the seed banks of northern

prairie wetlands. Canad. J. Bot. 67:1878- 1884.

WiLcove, D.S., D. RorusrEN, J. Du&ow, A. PHILLIPS, and E. Losos. 1998. Quantifying threats to imperiled species in the

United States. BioSci. 48:607-615.

WiLson, K.L. 1981. A synopsis of the genus Scirpus (Cyperaceae) in Australia. Telopea 2:153-172.

WINTERRINGER, G.S. 1959. Notes on Cyperaceae from Illinois. Rhodora 61:290-292.

Wisconsin DEPARTMENT OF NATURAL Resources. 2007. Working list: vascular plants. Madison. http://dnr.wi.gov/org/land/

er/wlist/index.asp?mode-detail&Grp-20. Accessed Jan 2007.

WISCONSIN LEGISLATURE. 2007a. Department of Natural Resources. Statute 29.604. dames (ed and threatened spe-

cies protected. Madison. http://nxt.legis.state.wi.us/nxt/gat II?f=templates&fn=default.htm&vid=WI:

Default&d=stats&d=29.604. Accessed Apr 2007.

WISCONSIN LEGISLATURE. 2007b. Department of Natural Resources. Statute NR 27 (03-07). Endangered and threatened

species. Madison. http://www.legis.state.wi.us/rsb/code/nr/nr027.pdf. Accessed Apr 2007.

YATSKIEVYCH, G. 1999. Steyermark' flora of Missouri.Vol 1. Revised Ed. Missouri Department of Conservation, Jef-

ferson City.

YATSKIEVYCH, G. and J. Turner. 1990. Catalogue of the flora of Missouri. Monogr. Syst. Bot. Missouri Bot. Gard.

37:1-345.

482 Journal of the Botanical R h Institute of Texas 1(1)

bOOK REVIEWS

Markus RIEDERER and CAROLINE MULLER (eds.). 2006. Annual Plant Reviews, Volume 23: Biology of the Plant

Cuticle. (ISBN 1-4051-3628-X, hbk.). Blackwell Publishing. 2121 State Ave., Ames, IA 50014-8300,

U.S.A. and 9600 P des Road, Oxford, OX4 2DQ, UK. (Orders: NM NT E com,

lers@ blackwellpublishing.com, 515-292-0140, 515-292-3348 fax, 1-800-862-6657). $249.99

hbk., 438 pp., 614" x Qv".

This is another installment from Blackwell Publishing's noteworthy Annual Plant Review Series. As always, it includes chapters on the

latest research on its subject written by experts in the field. This volume examines the biology of the plant cuticle in depth. The first

chapter is an introduction written by one the book's editors: Professor Markus Riederer of the Julius-von-Sachs Institute in Wurzburg,

Germany. Dr. Riederer opens with an interesting question put to him by a University search committee: “Does it make sense, and is it

fun at all, to spend so much time with the outermost micrometer of a plant?" Dr. Riederer replies, *...it is fun indeed to study the plant

cuticle and the plethora of processes related to it." He goes on to state that it is his hope and that of the volume's contributors that readers

..will find that it is worthwhile to invest time, brains and iran) into this endeavor."

The plant cuticl gapl face tl mposed of waxes and other extracellular substances.

The plant cuticle has b tial component of pl for millennia. The oldest cuticle found a back to the late Silurian

and early Devonian periods, about 400 million years ago. It eae plants from ultraviolet (UV) radi pathogens like fungi, and

it can even control the behavior of hungry herbivores. The cuti ls transpiration in cooperation with the stomata by preventing

water loss. The importance of this function, especially i ic environments, is obvious. In addition to preventing water loss, the cuticle

protects the plant by preventing rainwater and its microbes and unwanted molecules from passing into the plant. Some cuticles even

cause microbes, dust, and other small molecules to be removed along with the rainwater in a process known poetically as the Lotus

effect. The resistant cuticle also prevents the loss and uptake of polar molecules like salts through the plant’s surface, and controls the

uptake of organic compounds, including environmental pollutants. During the time when the stomata are closed, the cuticle assumes

complete control over the exchange of gases and vapors through the plant-atmosphere interface. In addition, the cuticle often provides

important structural support to the plant, an essential factor in the dreaded tomato fruit cracking familiar to farmers and gardeners.

Some 2,300 publications about the Dn cone bu been published in the last ten years according to the database BIOSIS. This

is the first experiment-based devoted the plant nda since the 1970's that is not a compilation of confer-

d, including new research on cut iti d biosynthesis, quantitative

ence proceedings. Many d

assessment of the transport of postal organic competes across the cuticle, and the cuticle’s a in UV protection. Scientists have

discovered new roles for the cuticle, including a role in pollen stigma-interactions. The cuticle’s role in regulating interactions between

plants and their environment is of interest to ecologists, environmental scientists, entomologists, and phytopathologists. In addition,

practical information for horticultural and agricultural scientists is in included. This volume is excellent and long overdue source of

information about the current research on the plant cuticle.—Marissa N. Oppel, MS, Collections and Research Assistant, Botanical Research

Institute of Texas, Fort Worth, TX, 76102-4060, U.S.A.

Kent J. BRADFORD and Hiroyuki NONOGAKI (eds.). 2007. Annual Plant Reviews, Volume 27: Seed Develop-

ment, Dormancy, and Germination. (ISBN 1-4051-3983-8, hbk.). Blackwell Publishing. 2121 State

Ave., Ames, IA 50014-8300, U.S.A. and 9600 ON Road, Oxford, OX4 2DQ, UK. (Orders: www.

blackwellprofessional.com, orders? blackwellpublishing.com, 515-292-0140, 515-292-3348 fax,

1-800-862-6657). $199.99, 367 pp., 614" x 915".

Seeds are not only vital to the life cycles of gymnosperms and angiosperms, they are also important staples in the human diet. The 27"

volume of Blackwell Publishing's Annual Plant Revi covers “seed biology from the point of view of the developmental and regulatory

processes that are involved in the transition from a developing seed through dormancy and into germination and seedling growth." Ten

years have passed since a book on this subject has been published, and there have been many advances in this field of research. The

model system Arabidopsis thaliana has been the subject of much recent research and is featured prominently throughout this book and

on its cover. The first two chapters cover the genetic control of seed development, including seed mass and seed coat development and

dormancy. These chapters are followed by six chapters devoted to seed dormancy, including seed dormancy models, genetic aspects,

lipid metabolism, and the roles of nitric oxide and abscisic acid. The final four chapters are focused on seed development and germina-

tion, including the roles of the abscisic acid and the gibberellins and the mechanisms, regulation, and genes involved in germination

and the transition to seedling growth. The “current challenges and remaining questions for future research" are discussed, as well. This

book is a resource for seed biologists, plant breeders, geneticists, plant developmental biologists, and graduate students. —Marissa N.

Oppel, MS, Irving, Texas, U.S.A.

J. Bot. Res. Inst. Texas 1(1): 482. 2007

A NOTE ON THE TYPE LOCALITY OF OENOTHERA ARIZONICA (ONAGRACEAE)

Kathryn Mauz

University of Arizona Herbarium

O Box 210036, University of Arizona

Tucson, Arizona 85721, U.S.A.

km arizeearthlink.net

ABSTRACT

The type locality for Oenothera arizonica (Munz) W.L. Wagner (Onagraceae), in the valley of the Santa Cruz River, north of Tucson, Ari-

zona, USA, is elucidated based upon historic property records. Today, the area of the historic farm is the site of residential and industrial

development and a highway interchange.

RESUMEN

La localidad del tipo Oenothera arizonica (Munz) W.L. Wagner (Onagraceae), en el valle del Río Santa Cruz, al norte de Tucson, Arizona

EEUU tad tad 1 t Taree as EP — 1 1] id 1

} ndictrial

Hoy, la área del

y también el lugar de una intersección de autopista.

The desert evening primrose, Oenothera arizonica (Munz) W.L. Wagner, is a sprawling winter-spring annual

that grows in sandy and floodplain soils at lower elevations of the Sonoran Desert in northwestern Sonora,

Mexico, and southwestern Arizona, USA (Fig. 1A & 1B). This note presents geographical and historical

context for its type locality:

Oenothera arizonica (Munz) W.L. Wagner, Novon 8:308. 1998. Oenothera deltoides Torr. & Frém. var. arizonica Munz,

Amer. J. Bot. 18:315. 1931; Oenothera californica (S. Watson) S. Watson ssp. arizonica (Munz) WM. Klein, Aliso 5:179. 1962; Oenothera

avita (WM. Klein) WM. Klein ssp. arizonica (Munz) WM. Klein, N. Amer. Fl. II, 5:116. 1965. Type: U.S.A. Arizona. Pima Co.: Grosetta

[sic] Ranch near Tucson, Arizona, 2400 ft elev., 28 Apr 1903, J.J. Thornber 509 (HoLoTYPE: UC; isotypes: ARIZ!, MO, US).

In addition to the type, cited above, four collections of Oenothera arizonica are known from the type locality:

“along ditches,” 5 Mar 1901, Griffiths 2393a (ARIZ!); 29 Apr 1902, Mr. & Mrs. Thornber s.n. (MO!); 20 Apr

1903, Thornber 4931 (ARIZ!); 27 Apr 1903, Mrs. Thornber 2983 (ARIZ!). Specimens at ARIZ document other

species at Grossetta's ranch in 1902-1903: Bromus diandrus Roth, Hordeum arizonicum Covas, Helianthus

petiolaris Nutt. var. canescens A. Gray, Machaeranthera asteroides Greene var. asteroides, Malacothrix glabrata

(A. Gray ex D.C. Eaton) A. Gray, Stephanomeria exigua Nutt. ssp. exigua, Eremalche exilis (A. Gray) Greene,

and Gaura parviflora Lehm.

Type localities for new species or infraspecific taxa described from historical collections are often poorly

defined, primarily the result of limited data on specimen labels. In some cases, a place name is idiosyncratic

to the era of the collection, whether or not it ever appeared on a map. The type locality of Oenothera arizonica

is an example of the latter: a farm whose owners were known to the collector and in the community, but

which did not become an enduring geographic entity. In the spring of 1891, the Tucson newspaper drew

attention to local farming efforts, including “S.V. Grossetta, one among the few enterprising men of this val-

ley, is preparing to try the efficacy of irrigating his lands down the Santa Cruz by means of pumps. If others

have been successful Mr. Grossetta certainly will be" (Arizona Daily Star, 14 Apr 1891, pp.3). The paper

undoubtedly meant A. (Antonio, or Anthony) V. Grossetta, who had moved to Tucson in 1880 and became

a prominent proprietor and promoter of culture in the town (G. Grossetta, pers. comm., Dec 2003). The

boundaries of Grossetta's ranch were legally recorded, even if the particular collection sites of O. arizonica

within that parcel remain unknown.

The 120-acre Grossetta parcel included portions of sections 34-35, T.13S, R.13E (Pima County Recorder

1906), extending east across the floodplain of the Santa Cruz River for three-quarters of a mile, at about

2310-2340 ft (704-713 m) elevation (Fig. 1C). The northern boundary of the one-time parcel became the

J. Bot. Res. Inst. Texas 1(1): 483 — 485. 2007

484 PE TM

Journal of

of Texas 1(1)

110° QW

30* 0'N

w TYPE LOCALITY

e OTHER COLLECTION LOCALITY

O 125 250Km

HEN — |

0 100 — 200 mi

32° 1534 N

Fic. 1. Oenothera arizonica. A—Location of the type locality in Pima Co., southern Arizona, within th i ] pecimens

at University of Arizona Herbarium (ARIZ) (modified from records available through the id database, seine asu. edu). B— Trailing stems ii me

living plant (photo by He Sur 3 às ee 2 Eon Pinal Co., Arizona). C— Type loca

J J da

[ (y ) g Santa Cruz Ri SCR) an intestate 100 10)i I Tucson. Section li I

in red, and nearby major roads are labeled. Imag porti f USGS digit q q les (section 34, Jaynes SE; section 35, Tucson

North SW) from Ari Regional Image Archive (aria.arizona.edu). May displayed in Uni Transverse Mercator projection (NAD83/zone 12);

hi I: P "T I £, thn WiCCod dat

y y'ap F a VIC WYUJOT UGLUITT.

alignment for what is now called “Miracle Mile,” originally part of the two-mile dogleg, built in 1937, joining

U.S. Highway 80/89 and State Highway 84 (Anonymous 1937), and today the southern terminus of State

Highway 77 where it meets U.S. Interstate 10. The modern interchange at Interstate 10 fills the western

portion of the former ranch, near the University of Arizona Experimental Farm that also takes up part of

the Grossetta’s original parcel. East of the interstate, an industrial site, warehouses, and housing occupy the

land.

Conservation concerns.—As a farm and orchard in the Grossetta's time, the type locality of Oenothera

Mauz, Type locality of Oenothera arizonica 485

arizonica was near the eastern and upper elevation limits of the species' range as it has since been docu-

mented by other collections (Fig. 1A). This range corresponds closely to the Lower Colorado River Valley

biogeographic subdivision of the Sonoran Desert (Shreve 1951). The river valleys associated with this biotic

community, including those near Tucson, have in the past century been intensively utilized for agriculture

1 ahr

where irrigation is available; in the past thirty years, the lowlands izing this gion have experi-

enced among the highest rates of population growth in the binational Sonoran Desert (Nabhan & Holdsworth

1998). Although the desert evening primrose may grow in fallow fields and along farm roads in agricultural

areas (author’s observations), widespread disturbance and replacement of low desert landcover—and of

agricultural fields—with urban land uses may present a conservation concern for O. arizonica populations

as the region continues to endure unprecedented development.

ACKNOWLEDGMENTS

I thank the curators at ARIZ, MO, and UC, whose collections were consulted in the preparation of this note,

and colleague Hugo Rodriguez, who made suggestions for the Spanish abstract. The maps for this note were

prepared, with permission, at the Arizona Remote Sensing Center, University of Arizona. The comments

of reviewers are also appreciated.

REFERENCES

ANONYMOUS. 1937.'Miracle Mile, safety-plus thoroughfare. Arizona Highways 8(6):14-15.

NABHAN, G.P. and A.R. HoLosworTH. 1998. State of the desert biome: uniqueness, biodiversity, threats and the

adequacy of protection in the Sonoran Bioregion. The Wildlands Project, Tucson.

Pima COUNTY RECORDER. 1906. Deed, SY. of NEW sec. 34 & SWW of NWM% sec. 35, T.13S, R.13E, Gila & Salt River Meridian

& Base Line [Bessie H. Grossetta, 8 Oct 1891; recorded 12 Oct 1906]. Book of deeds no. 39:693-694, viewed

on microfilm at Pima County Recorder, Tucson.

SHREVE, F. 1951. Vegetation of the Sonoran Desert. In: F. Shreve and |.L. Wiggins. Vegetation and flora of the Sonoran

Desert, vol. 1. Publication 591, Carnegie Institution of Washington, Washington, DC. Pp. 1-191.

486 Journal of the Bot titute of Texas 1(1)

bOOK REVIEWS

Ropert H. Wess, STANLEY A. LEAKE, and RayMonpD M. TURNER. 2007. The Ribbon of Green: Change in Ripar-

ian Vegetation in the Southwestern United States. (ISBN 0-8165-2588-9, hbk.). The University of

Arizona Press, 355 S. Euclid, Suite 103, Tucson, AZ 85719-6654, U.S.A. (Orders: www.uapress.arizona.

edu, 520-621-3920, 520-621-8890 fax). $75.00, 462 pp., b/w photos, 9!4" x 1214".

From the outside, this large book, with its gorgeous color photograph of the San Juan River on the cover, looks like a coffee-table book. It

is actually a fascinating study of the riparian vegetation of the southwestern United States. Riparian vegetation accounts for one third of

the vascular plants in this region and is home to many species of wildlife. Unlike many ecological books, this one actually contains some

very welcome good news- riparian vegetation is increasing in many areas. This increase is due to many factors, including winter floods

that allow the establishment of new species on disturbed areas, flood control, and favorable climatic conditions. The authors focus on

factors that affect the woody riparian vegetation of major river valleys in Arizona, Utah, southern Nevada, and southeastern California.

They use repeat photography and hydrological research to analyze changes in riparian vegetation in these areas. This long-term study

encompasses 140 years of Poe E history, and black-and-white photographs illustrate the book. The authors present an analysis

of thousands of photographs, incl ly photographs by surveyors, scientists, explorers, and tourists, as well as modern-day repeat

photographs of the same locations. Woody plants and identifiable perennials are the focus of the authors' photographic analysis. This

book paints a surprising portrait of the changes in southwestern riparian ecosystems over the last century and a half.—Marissa N. Op-

pel, MS, Irving, Texas, U.S.A.

CERIDWEN TERRILL Foreword by Gary PauL NasHan. 2007. Unnatural Landscapes: tracking invasive spe-

cies. (ISBN 0-8165-2523-4, pbk.). The University of Arizona Press, 355 S. Euclid Avenue, Sutie 103,

Tucson, AZ 85719, U.S.A. (Orders: www.uaapress.arizona.edu). $17.95, 220 pp., 5 maps, 26 b/w

photos, 61⁄2" x 8".

Terrill's book on unnatural landscapes is a coe din puc ad various islands that have been impacted by invasive species in

many id Ceiridwen picked four North A dy pecies of birds, fishes, plants and mammals since *islands

y e T2

l for exploring the problem of invasive species because their native plants and animals are highly specialized, isolated

organisms, sien few in number, and ent susceptible to the negative effects of introduced species.”

In reading this book, one can better understand why an exotic or an introduced species may become invasive and how it becomes

detrimental to oe d ud and even more so to the entire ecosystem. For instance, s author points out that a horticultural flat of the

African I dulis) is sold for $12.99 without any ] fthe plant. Another

concern is the dra of exotic pets that can provoke a chain reaction A the rM of unknown foreign parasites and

diseases.

"Invasive species haven't just peted natives for nutrients, space, and water. They also can change fire regimes, alter streams

courses, prevent native plants from regenerating, negatively affect human health and local economies." One good example given by

Mrs. Terrill is a crevice-dwelling freshwater crayfish in warm pools of Ash Meadows, Nevada. After introduction, the crevice-dweller

(Procambarus clarkii) became a top carnivore and keystone species in the community. A keystone species has a key role in an ecosystem,

affecting many other species, and whose removal leads to a series of extinctions within the ecosystem.

Even if a good question is raised, “extinction is a natural process, so what's the big deal?,” Terrill states that one of the methods to

deal with invasive species is to eradicate them. It is simpler on an island because there is less risk of reinvasion. Even after eradication,

and exotic animals such as sheep are removed, it can sometimes take more than a decade for the landscape to fully recover. Eradication

of certain plants may be achieved by reducing or eliminating essential minerals elements. For instance, Cheatgrass (Bromus tectorum)

was successfully removed on Anaho Island National Wildlife Refuge (Nevada) by the removal of nitrogen. Invasive pigs on Santa Cruz

Island (California) have been controlled through large-scale hunting, relocation on and off the island, and contraceptive measures.

Ceiridwen Terrill hopes people will begin to take personal action to stop or at least slow down the spread of exotic species, and in

a larger political arena, she encourages people to influence public policy with their vote. I think this book is more about letting people

know of the potential problems and dangers of invasive species. Education can go along way in prevention. An “Ounce of prevention

is worth a pound of cure." Guidelines need to be implemented and prominently displayed and preventive equipment installed in more

parks and natural areas for people to follow. Examples include,“Boot brushes on the mainland pier provide an opportunity for visitors

to remove invasive seeds from their footwear before their trip to Santa Cruz Island." Also, visitors to this Island are encouraged not to

hike through weed-infested areas, but to stay on trails to avoid disturbing soils and making it easier for invasive plants to establish.

We definitely need more people exposing the problem of invasive species to the general public. Terrill has done a marvelous job

describing in simple terms, the problems and potential solutions of invasive species.— Virginie H. Raquet, Botanical Research Institute of

Texas, 509 Pecan Street, Fort Worth, TX 76102-4068, U.S.A., vraquet@brit.org.

J. Bot. Res. Inst. Texas 1(1): 486. 2007

NOTES ON CONOCARPUS ERECTUS (COMBRETACEAE)

IN THEBAA GALIPOR NTS PENINSULA, MEXICO

José Luis León-de la Luz and Raymundo Domínguez-Cadena

Centr dol x jones Ri lógi acA | Noroeste (CIBNOR)

Mar Bermejo 195, Col. Playa Palo de Santa Rita

La Paz, B.C.S. 23090, MÉXICO

jlleon04@cibnor.mx

ABSTRACT

Mangrove stands of Conocarpus erectus in the semi-arid southern part of the Baja California Peninsula are rare, since the species reaches

here its northern limit of distribution in the northern and western hemispheres. Four populations are described. Occurrence of these

stands is relevant because this species seems to have responded positively to heavy rains over the last two decades. The coastal areas

that have been favored maintain suitable levels of soil moisture so that populations of this species have been able of colonize new areas

or showing a vigorous growth.

Key Worbs: Conocarpus erectus, Baja California, mangrove, global change

RESUMEN

Las agrupaciones del mangle botoncillo Conocarpus erectus en el semi-árido sector sur de la Peninsula de Baja California, México, son

vue

que en esta región la especie encuentra el límite norte de su distribución geográfica, tanto en los hemisferios norte y

occidental. La presencia estas agrupaciones es relevante dado que los autores consideran que la especie a respondido positivamente a

eventos que han dejado elevada humedad en el suelo de la franja costera de la zona en las últimas dos décadas, hecho que ha permitido

a pequeñas poblaciones de esta especie colonizar nuevos puntos o reiniciar un activo crecimiento.

PALABRAS CLAVE: Conocarpus erectus, Baja California, manglar, cambio global

Primary environmental conditions required for establishing mangroves include water currents of low

kinetic energy and a relatively high water temperature, i.e., higher than 20?C as the mean annual tempera-

ture (Lugo 1998). Along the west coast of the Baja California Peninsula, low winter temperatures and cold

ocean currents seems act as the main controls for the spread of mangroves. Hence, mangroves on the Gulf

of California side of the peninsula occur to 29?N at Bahía de Los Angeles (BA in Fig. 1) and near 27?N on

the Pacific coast side at Laguna San Ignacio of the Estero El Coyote (EC in Fig. 1).

Mangrove species in the Baja California Peninsula are Rhizophora mangle L. (red mangrove, Rhizopho-

raceae), Laguncularia racemosa (L.) Gaertn. (white mangrove, Combretaceae), and Avicenia germinans (L.) L.

(salty mangrove, Aviceniaceae). These three species typically grows as an association. Typically R. mangle

and L. racemosa are permanently in touch with seawater and A. germinans grows landward in wet soil (Turner

et al. 1995).

The button mangrove (Conocarpus erectus L., Combretaceae) is a common member of the mangrove

association in tropical and subtropical zones in the Western Hemisphere. The species extends from the

Caribbean Islands, including Bermuda and The Bahamas, through central Florida and northeastern Mexico

and southward along the Atlantic coast to Brazil. On the Pacific coast, it extends from northern Mexico to

northwestern Peru, including the Galapagos Islands. Also, it is found in western tropical Africa from Senegal

to Zaire. The button mangrove is one of the more terrestrial mangrove species because it grows landward

from the typical mangrove stands where soils are occasionally flooded. As with other mangroves, it is also

present where the annual mean isotherm is above 20?C, avoiding near freezing temperatures (Robertson &

Alongi 1992; Dawes 1998).

Typically, this species grows in brackish or saline silt along depositional coasts, behind mangrove stands

immediately above the intertidal belt, as well as in coastal marshes, estuaries, inlets, and mudflats. It usually

grows to a height of 8-10 m as an arborescent form (Tomlinson 1986). On the Pacific coast, button man-

J. Bot. Res. Inst. Texas 1(1): 487 — 490. 2007

488

grove extends marginally towards northwestern

Mexico. Wiggins (1980) and Turner et al. (1995)

recorded solitary individuals or small clumps on

the Baja California Peninsula, south of 24?N at

sites designated with cross marks (+) in Figure 1.

Compiling information about locations

from vouchers of C. erectus in our herbarium and

visiting these locations in recent years, as well

as populations that were seen by informants, we

present the following account of four sites where

well structured populations of this species occur.

In general terms, analysis is interesting because

the southern part of the Peninsula (especially

the Cape Region) is the northern limit of the

geographical distribution in the northern and

western hemisphere.

In the last decade, this area has received

larger-than-typical volumes of rain (four hurri-

canes in six years: two 2001, one 2003, one 2006),

which brought a few months of flooding and high

water to the mouths of arroyos, estuaries, and

salt marshes, typical habitats where the species

has been collected. Our findings and analyses

suggest that this species is currently undergoing

population and regional expansion.

Site 1: Isla Espíritu Santo (24?30'57.7" N,

110°23'11.4" W; IES in Fig. 1).

In the summer of 2002 (El Meztefio Cove), Felix

Pico et al. (2005) found a stand of tree-like button

mangrove on a narrow strip of land (130 m long)

fal, Dat o ID L

Journal of

titute of Texas 1(1)

À ne "o 106°

H a

| a8 1

C ER. \

ee i b

L 99° 3 AS a

X 1 wA d

- 28° A £ E “an, =

bod Es GULF OF a MEXICO

wd “y CALIFORNIA Bs j^

EC "|^ % ae

| po is x^ | »

y e E " pe

v |

& Pies YS

Oe E E TS

ET px S N

24 Se c EE

PEC) AN

pm MIG `

PACIFIC OCEAN

Fic. 1. Limits of four species of mang long the coast of North

America. Gulf of California st line: BA = Bahia del les (Rhizopt

mangle, Avicenia germi „and Lag TA ET

EC = Estero El C yot (R mangle and | racemosa). St fr

(ele A a

| to, PEC = Plutarco Elías Calles,

MIG = Migrifio, and LSA = La Salina. Previously, isolated trees of C. erectus

were reported on sandy beaches along the southwestern tip of the Baja

California Peninsula (+).

between a mudflat of sand and alkaline, clayey silt on the sea side and a colluvial, rocky soil with desert

scrub on the land side. On a recent survey, the site contained a population of 87 plants. Measurements of

height, two canopy diameters, and the basal perimeter of the trunk are shown in Table 1.

This is an old population because the community contains a majority of mature specimens with basal

trunk circumference of up to 0.45 m, which suggests an age of several decades. No other mangrove species,

of any size or age, is present near this cove.

Hurricanes strike this area irregularly. During summer 2001, Hurricane Juliette brought high seas that

flooded the coves and bays on the west side of the island. At El Mezteño Cove, a break in the barrier beach

allowed continuous ebb and flow of seawater through the new channel, which initiated expansion of the

mangrove population, a condition that is clearly noticeable to visitors. Before 2001, the population seemed

inconspicuous, probably from lack of an adequate water supply. During the summers of 2003 and 2006,

Hurricanes Ignacio, Marti, and John renewed the tidal channel flow system, but the button mangrove did

not show the vigorous growth seen in 2001-02.

Site 2: Plutarco Elias Calles (23°12'36.9"N, 110?08'24.8"W; PEC in Fig. 1)

This site is on the relatively cold Pacific coast side of the Baja California Peninsula in a salt marsh bordered

with a strand of coastal dunes. This population is composed of tall individuals of shrubby habit with basal

branching occupying an area bordering a mudflat. According to informants from the early 1990s, this popu-

lation appears to be relatively young. This aspect is supported by the homogeneous basal perimeter of the

León de lal 1 Domi AE tus in the Baja California Peninsula 489

Taste 1. Structural characteristics of the four stands of button mangrove Conocarpus erectus known in the southern Baja California

Peninsula, Mexico. Name of each stand is followed by the estimated surface area and number of individuals.

Name of button mangrove stand max min Population Canopy

Cover (m?)

1. Isla Espíritu Santo (1,420 m?; 87 plants)

Height (m) 4.30 0.90

Canopy cover (m?) 12 0.12 795.83

Basal trunk circumference (m) 0.45 0.04

2. Plutarco Elias Calles (16,362 m7; 280 plants*)

Height (m) 9.10 1230

Canopy cover (m?) 51.62 0.56 9,100.60

Basal trunk circumference (m) 0.62 0.08

3. Migriño (7,500 m?; 225 plants*)

Height (m) 5.10 1.40

Canopy cover (m?) 12:37 0.30 8,257.50

Basal trunk circumference (m) 0.94 0.09

4. La Salina (27,000 m?; 2,160 plants*)

Height (m) 4.30 0.90

Canopy cover (m?) 15.58 0.66 38,017.20

Basal trunk circumference (m) 0.59 0.11

* |nference based on 20 x 10 m? sampling and Google Earth (2007) view analysis

main stem in most of the population (the larger mangroves) and the meager accumulation of deadwood and

few dead plants. Freshwater draining from a neighboring area used for cattle grazing seems to contribute

to suitable soil humidity. Measurements of height, two canopy diameters, and the basal perimeter of the

trunk are shown in Table 1.

Site 3: Migriño (23?01'48.7"N, 110?04'54.9"W; MIG in Fig. 1)

This is a peculiar site, located 45 m above sea level along the banks of a sandy arroyo in a strip of pure man-

grove vegetation that is 250 m long. A freshwater spring is located immediately above the upper tip of the

strip, and this is probably the basis for this patch of vegetation. The shore is almost 1 km west of this site.

According to informants, this is an ancient population. An accumulation of litter and deadwood prevails

throughout the patch and some dead trunks are >0.60 m in circumference. Young mangrove grows along

the borders of the stand and in the downstream edge of the stand. The stand is actively growing and the

thick canopy, which blocks sunlight reaching the interior, prevents seedlings from growing. Main trunks

are continuously being buried by bed load sandy sediments deposited after heavy rains.

Site 4. La Salina (23?35'08.1"N, 109°32'12.3"W; LSA in Fig. 1)

This is the biggest and undoubtedly the oldest button mangrove stand on the Peninsula. It is located in a

salt marsh on the Gulf of California coast that is irregularly flooded. The salt marsh has a perimeter of about

2.2 km with a surface area of about 6 hectares. No other species of mangrove is present. This population

receives a small inflow of freshwater to this micro-basin. Occasionally, sea swells break through the dune

strand and introduces seawater to the marsh. Deposits of detritus border the deeper part of the lagoon.

During a recent flood, water reaching 2 m in depth remained for 2-3 months and killed the foliage of the

covered button mangrove. This strip of mangrove is 900 m long.

In summary, this species has responded positively to the recent events of heavy rains and the decade

of the 1990s with two “El Nifio” events that impacted the Peninsula. Even more recently, four hurricanes

have brought flooding. The coastal areas have been favored with suitable levels of soil moisture that enabled

this species to colonize new areas or trigger vigorous growth.

490 Journal of the Botanical R h Institute of Texas 1(1)

ACKNOWLEDGMENTS

We thank CIBNOR authorities for funding our trips to the Cape Region coasts where we found these man-

grove populations, mainly Proyecto Manglares. We appreciate the detailed and constructive reviews of three

anonymous reviewers.

REFERENCES

Dawes, C.J. 1997. Marine botany, 2nd. Ed. John Wiley & Sons. New York, NY.

FELIX-Pico, E.F., O.F. HotaufN-QuiRÓNEZ, L. GobiNez-Orta, and N.A. SANTA-MARÍA-GALLEGOS. 2005. 3. Estructuras de las

comunidades de Mangles de Bahía de Loreto, Isla de La Paz, Isla Espíritu Santo y Costa Oriental de la Bahía

de La Paz. Libro de Resúmenes del I Taller sobre Manglares de la Península de B.C., La Paz, B.CS., Pp. 9-10.

GOOGLE EARTH. 2007. Google Earth™ Mapping Service.

Luco, A., S. Brown, and M.M. Brinson. 1988. Forested wetlands in freshwater and saltwater environments. Limnol.

Oceanogr. 33:894-909.

ROBERTSON, A.l., D. M. ALonal. 1992. Coastal and estuarine studies: Fourth tropical mangrove ecosystems meeting.

American Geophysical Union, Washington, DC.

TOMLINSON, P.B. 1986. The botany of mangroves. Cambridge University Press, Cambridge, U.K.

Turner, R.M., J.E. Bowers, and T. Buraess. 1995. Sonoran desert plants: an ecological atlas. University of Arizona

Press, Tucson.

Wiaains, I.L. 1980. Flora of Baja California. Stanford University Press, Stanford, CA.

REDESCUBRIMIENTO DE AXINIPHYLLUM SAGITTALOBUM (ASTERACEAE)

BBLEXSIBSRSSNESIDRBTDBESURS Y NOMS DE PAS BSPBCIBSDETESEB

GÉNERO QUE HABITAN EN EL ESTADO DE GUERRERO, MÉXICO

Alberto González-Zamora Isolda Luna-Vega José Luis Villasenor

Departamento de Biología Evolutiva Departamento de Biología Evolutiva Departamento de Botánica

Facultad de Ciencias, UNAM Facultad de Ciencias, UNAM Departamento de Botánica

Apartado Postal 70-399 Apartado Postal 70-399 Apartado Postal 70-233

Coyoacán, 04510 MÉXICO, D.F. Coyoacán, 04510 MÉXICO, D.F. Coyoacán, 04510 MÉXICO, D.F.

z mU :

IVenp.rcrencias. UNGM.MX

RESUMEN

Axiniphyllum sagittalobum, especie endémica a la Sierra Madre del Sur de Guerrero, México y que fue descrita

por B.L. Turner con base en una sola colecta de Hinton (1939), fue recolectada con flores, por primera vez

desde su descripción, en el bosque de pino del municipio de San Miguel Totolapan. Otras dos especies de

Axiniphyllum se registran habitando en el estado de Guerrero, A. corymbosum y A. pinnatisectum. Se ofrecen

claves de identificación de estas tres especies, descripciones mas amplias, datos de distribución geográ-

fica, información acerca del hábitat y referencias de herbario. Durante nuestro reciente trabajo de campo

no encontramos a A. pinnatisectum, especie también muy rara con solo dos especímenes representados en

herbarios del extranjero; esta especie también es endémica a la Sierra Madre del Sur. Se sugiere emprender

recolectas de campo para buscar esta especie con detenimiento, ya que en el caso de que no sea colectada

durante los próximos afios, podrá considerarse como extinta. Ninguna de las especies antes mencionadas

está considerada dentro de la Norma Oficial Mexicana vigente (NOM-059-ECOL-2001).

ABSTRACT

Axiniphyllum sagitalobum, a strict endemic of the Sierra Madre del Sur, Guerrero, Mexico, was described by B.L. Turner on basis of a single

collection of Hinton in 1939. It has been recently collected with flowers, in the pine forests of San Miguel Totolapan municipality, for the

first time since is description. Two other species of Axiniphyllum are dE NU from the Mexican state i Guerrero: A. corymbosum and A.

pinnatisectum. An identification key for habitat information

and herbarium, which is known only from the typ ion; T P is also Bend to ne e Madre del Sur. None of the three

species of Axiniphyllum is considered in the recent Norma Oficial Mexicana (NOM-059-ECOL-2001).

Los estudios florísticos de la familia Asteraceae en México se han abordado a nivel regional o estatal (e.g.,

Villasetior 1982, 1987, 1989; McVaugh 1984; Villarreal et al. 1996; Ortiz & Villasenor 1998; Balleza & Villa-

señor 2002; Villarreal & Villaseñor 2004; Villaseñor et al. 2004). Uno de los estados con mayor diversidad

florística de México es Guerrero; dentro de los límites de su territorio se estima que existen aproximada-

mente 666 especies, incluyendo taxones subespecificos (Villasenor et al. 2004) incluidos en 143 géneros,

14 de ellos endémicos a México (Villaseñor et al. 1998). Existen pocos estudios florísticos de la familia en

el estado, aunque se ha intentado realizar un inventario de las especies que lo habitan. Villasenor (1987)

publicó uno de los inventarios más completos de Asteraceae a nivel genérico para la Cuenca del Río Balsas,

donde incluye un total de 131 géneros para el área.

El género Axiniphyllum está integrado por cinco especies, con una distribución restringida a los bosques

templados del occidente de México, en los estados de Durango, Guerrero, Nayarit y Oaxaca. La descripción de

Axiniphyllum fue hecha por Bentham (1872), en la cual incluyó dos especies que carecían de flores radiadas,

A. corymbosum Benth. distribuida en Guerrero y Oaxaca y A. tomentosum Benth., con un área de distribución

restringida a las zonas montañosas de Oaxaca; la última especie había sido descrita anteriormente bajo el

nombre de Polymnia scabra por Zuccarini (1832), sin embargo Blake (1930) realizó la transferencia de esta

ültima al género Axiniphyllum.

J. Bot. Res. Inst. Texas 1(1): 491 — 498. 2007

£s+haD o ID L

492 Journal of t titute of Texas 1(1)

Desde que el género fue descrito por Bentham y por más de un siglo, no se descubrieron nuevas espe-

cies y Axiniphyllum se mantuvo como un género con solo dos especies. Más tarde, Turner (1978) adicionó

dos especies a partir de colectas hechas en el estado de Guerrero: A. sagittalobum B.L. Turner y A. pinnati-

sectum (P.G. Wilson) B.L. Turner, ésta ultima descrita al principio como Rumfordia pinnatisecta P.G. Wilson.

Por ultimo A. durangense B.L. Turner, del sur del estado de Durango y Nayarit, es la especie descrita más

recientemente (Turner 1987).

El interés en Axiniphyllum se basa principalmente en el pobre conocimiento que se tiene de él, ya que

las colectas realizadas de las especies de este género son escasas y por lo tanto su representación en los

herbarios es pobre, lo que hace pensar que sus poblaciones son pequeñas. Dado lo anterior, es necesario el

estudio de las especies de este género, sobre todo de aquellas con una distribución restringida. El objetivo

de este trabajo es ofrecer una descripción detallada de las especies de Axiniphyllum en el estado de Guerrero,

así como datos de su distribución geográfica conocida, datos ecológicos y poblacionales y una clave tax-

onómica, con la finalidad de contribuir al conocimiento florístico del estado y ahondar en el conocimiento

de las Asteraceae mexicanas. Todas las especies del género tienen una distribución restringida al país.

MATERIAL Y MÉTODOS

Se recopilaron los datos de distribución de ejemplares de herbario del género Axiniphyllum depositados en

los siguientes herbarios: Herbario Nacional de México, Instituto de Biología, UNAM (MEXU), Herbario de la

Escuela Nacional de Ciencias Biológicas, IPN (ENCB), Herbario de la Facultad de Ciencias, UNAM (FCME) y

Herbario de la Universidad Autónoma de Guerrero (UAGC). Las especies de este género no están representados

en herbarios extranjeros a excepción de algunos ejemplares de A. corymbosum y los ejemplares tipo.

Debido a la escasez de ejemplares de herbario de A. pinnatisectum y A. sagittalobum, se realizaron dos

salidas a campo a las localidades tipo durante los meses de septiembre y octubre de 2005, con el objetivo de

obtener mas datos sobre su morfología externa y llevar a cabo observaciones de las poblaciones y su hábitat.

RESULTADOS

El trabajo realizado en campo se enfocó a la recolección de ejemplares de Axiniphyllum pinnatisectum, el cual

solo cuenta con dos ejemplares colectados por Hinton en 1936 y 1937, depositados en los herbarios K, NY

y US y A. sagittalobum, el cual solo era conocido a partir del ejemplar tipo colectado por Hinton en 1939,

depositado en el herbario TEX-LL y cuyos isotipos se encuentran en MICH, NY y US. Sólo se encontró una

pequeña población de A. sagittalobum (menos de 100 individuos), por lo que se recolectó un único ejemplar

de respaldo que está en proceso de ser depositado en los herbarios FCME, UAGC, MEXU, TEX-LL y NY.

Axiniphyllum pinnatisectum no fue encontrada en campo.

Las localidades tipo de estas dos especies se encuentran en poblados relativamente cercanos entre sí.

Estas zonas actualmente están sujetas a fuerte acción antropogénica, por lo que el bosque ha sido conver-

tido, en la mayoría de los casos, en campos de cultivo. La colecta de A. sagittalobum permitió elaborar una

descripción más completa de la especie, en comparación con la realizada por Turner (1978).

En el estado de Guerrero se encuentran representadas tres especies: A. corymbosum, con una distribución

amplia dentro del estado y A. pinnatisectum y A. sagittalobum, ambas endémicas al estado, con una distribución

conocida restringida a la Sierra Madre del Sur, en lo que se conoce como la región administrativa conocida

como “tierra caliente" (Fig. 1).

La revisión de las colecciones de los herbarios permitió corroborar que existe un escaso numero de

ejemplares de las especies de Axiniphyllum; MEXU es el herbario que contiene una mayor cantidad de espe-

címenes con 31 ejemplares, seguida de ENCB con cinco.

A continuación se presenta la descripción de las tres especies registradas para el estado de Guerrero.

TRATAMIENTO TAXONÓMICO

Axiniphyllum Benth., Hookers Icon. Pl. 12-16; t. 1118. 1872. Especie tipo: Axiniphyllum corymbosum Benth.

González-Zamora et al., Redescubrimiento de Axiniphyll gittalobum en Guerrero, México 493

-102 -101 -100 -99

e + € *Chilpancingo €

e e.

cr + +

«€ Axiniphyllum corymbosum

+ Axiniphyllum pinnatisectum

O Axiniphyllum sagittalobum

50 0 50 100 Kilómetros

oJ S| de +s

ET 01 ET) E 3

Fic. 1 Ni pa Pa | A hall fr

Hierbas perennes erectas, de 0.5 a 1.5 m alto; rizoma fibroso tipo cormo; tallo rojizo, piloso en la base volvién-

dose glabro, 1.5—3.6 mm diam.; hojas opuestas, en ocasiones concentrándose en la parte basal formando

un tipo de roseta, triangulares, sagitadas o sagitadas tripartitas, 2.7-16 cm longitud, 3-9.1 cm ancho; haz

con pubescencia pilosa de tricomas multicelulares septados uniseriados, en ocasiones mezclados con glan-

dulares en toda la superficie; envés con pubescencia pilosa de tricomas multicelulares septados uniseriados

£+sha D o ID L

494 Journal of t titute of Texas 1(1)

o glandulares principalmente a lo largo de las venas; lámina entera, trilobulada o pentalobulada, márgenes

enteros, aserrados o lobados a veces ligeramente revoluto, lóbulo terminal entero, ondulado, lobulado, aserrado

o crenado; base sagitada a atenuada decurrente, ápice agudo, nervación acródroma imperfecta basal, con 3

venas de primer orden más una en cada lóbulo; pecíolo alado connado-perfoliado, con pubescencia y tipo

de tricomas iguales que en la lámina, 1-9.1 cm longitud. Sinflorescencia politélica, un dicasio, el internodo

basal 15-58 cm longitud, pedúnculos 2.1-13.6 cm longitud, glabros o con pubescencia pilosa de tricomas

glandulares mezclados con no glandulares multicelulares septados uniseriados, brácteas con las mismas

características de las hojas delimitando la sinflorescencia y los paraclados, 3-33 mm longitud, 0.7215 mm

ancho; involucro 4.8-17 mm longitud, 9.4-28 mm ancho; brácteas involucrales imbricadas en 2-3 series,

las más externas con margen entero, coriáceas, ápice agudo, pubescencia pilosa de tricomas multicelulares

septados uniseriados o glandulares mezclados con no glandulares 4-7.8 mm longitud, 0.5-2.7 mm an-

cho, las internas 4.2-8.8 mm longitud, 0.5-2.1 mm ancho, papiráceas, ápice agudo, amarillentas. Flores

radiadas presentes o ausentes, cuando están presentes 5-8, pistiladas y fértiles, amarillo pálido, ligulas

3-lobadas en el ápice, 7-12 mm longitud, 3-6 mm ancho, tubo densamente pubescente piloso, 0.8-3 mm

longitud, 0.4—2 mm ancho, ramas del estilo amarillas, glabras, apéndices agudos, ca. 1.1 mm longitud.

Aquenios negros, glabros, 4-costillas, cuadrangulares, 1.5-2 mm longitud, 0.9 mm ancho. Flores del disco

hermafroditas, fértiles, amarillas, 5-lobuladas, 1.5-7 mm longitud, 0.4-2.3 mm ancho, pubescencia pilosa

en toda la superficie concentrándose en la base del tubo, tubo 0.2-2.1 mm longitud, limbo 2.3-5.1 mm

longitud, lóbulos 0.2-1.3 mm longitud, ápice agudo. Anteras amarillas, 1.6—4.9 mm longitud, glabras,

base truncada, apices del estilo apendiculado; ramas del estilo amarillo ocre, 0.5-5.3 mm longitud, pilosas

en la superficie exterior comenzando desde un poco debajo de la bifurcación, con tricomas unicelulares,

apéndices del estilo agudos. Aquenios glabros, negros, epaposos, 4-costillas, 1.1-3.3 mm longitud, 0.3-1.4

mm ancho, cuadrangulares.

CLAVE PARA LAS ESPECIES DE AXINIPHYLLUM EN GUERRERO, MÉXICO

1. Flores radiadas presentes.

2. Lámina pentalobulada, lóbulo terminal cuando no entero aserrado A. pinnatisectum

2. Lámina entera o trilobulada, lóbulo terminal cuando no entero crenulado A. sagittalobum

1. Flores radiadas ausentes A. corymbosum

1. Axiniphyllum corymbosum Benth., Hooker’ Icon. Pl. 12:17. 1872. Two: MÉXICO. Oaxaca: “woods in the province

of Oaxaca, at an elevation of 7500 ft," Sep 1840, H. Galeotti 2089 (HoLotipo: K; foto en MEXU!).

Hierba de 0.55-0.85 m alto; tallo 1.6-3.4 mm diám.; hojas triangulares en ocasiones formando una roseta

basal, 2.7-8.5 cm longitud, 3.5-9.1 cm ancho; lámina entera a ligeramente trilobulada, margen entero a

aserrado, lóbulo terminal aserrado o entero; pecíolos 1-8.1 cm longitud. Internodo basal de la sinflorescencia

15-20.5 cm longitud, pedúnculos 2.1-12.1 cm longitud; brácteas 9-33 mm longitud, 1.4-15 mm ancho;

involucro 4.8—7.8 mm longitud, 9.4-13.7 mm ancho; brácteas involucrales externas 4.1-7.6 mm longitud,

0.5-2.5 mm ancho, las más internas 4.2-8.6 mm longitud, 0.5-1.7 mm ancho. Flores radiadas ausentes.

Flores del disco 4.1-7 mm longitud, 0.4-2.3 mm ancho. Anteras 1.6-3.4 mm longitud; ramas del estilo

0.5-2.3 mm longitud. Aquenios 1.1-2.6 mm longitud, 0.4-1.4 mm ancho.

Distribución y altitud.—bosque de pino-encino de Guerrero y Oaxaca, desde los 1800 hasta 2500 m

snm.

Floración.—septiembre a noviembre.

Especímenes examinados: MEXICO. Municipio de Alcozauca: “Cerro Azul,” 3 km al SO de Alcozauca, Oct 1989, FJ. Díaz P. SN

(MEXU). Municipio de Atlixtac: km 13 del camino que va de Alzacoalaya a Heuycatenango, 4 Oct 1981, F.G. Lorea 1680 (MEXU;

FCME). Municipio de Chilapa de Álvarez: km 70-71 de la carretera Chilpancingo-Chilapa-Tlapa, 13 Oct 1991, J.L. Panero 2458

(MEXU). Municipio de Chilpancingo: Cima del Cerro Alquitrán, 21 Nov 1968, H. Kruse 2032(b) (MEXU); aproximadamente 3 km

al E de Amojileca, camino Chilpancingo-Omiltemi, 30 Oct 1982, F.G. Lorea 2628 (MEXU); Mazatlán, falda E del Cerro Alquitrán, 7

Nov 1969, H. Kruse 2666 (MEXU; ENCB); Mazatlán, falda E del Cerro Alquitrán, 8 Oct 1968, H. Kruse 2032 (MEXU); near Huapango

on road to San Vicente S of Omiltemi, 18 Oct 1984, D.E. Breedlove 61873 (MEXU); km 70-71 of the road Chilpancingo-Chilapa-Tlapa,

González-Zamora et al., Redescubrimiento de Axiniphyll gittalobum en Guerrero, México 495

across from microwave station (S side) along an intermittent creek approximately 200 m from the road, 7 Nov 1990, J.L. Panero 2040

(MEXU); Microondas road up Cerro Alquitrán, marked “El Tejocote" on hwy 95 W of Mazatlán, ca. 7.2 km W of hwy 95, 30 Sep 1983,

W.R. Anderson et al. 12905 (MEXU; CHAPA); 3 km antes de Chilpancingo a Omiltepec, J. Gutiérrez y A. Terán 32 (MEXU; FCME). Mu-

nicipio de Quechultenango: 3 km al S de Quechultenango al O del camino a Tlayolapa, 28 Sep 1982, A. Nuñez 1101 (MEXU; FCME).

Municipio de San Luis Acatlán: aproximadamente 1 km al SE de Mixtecolapa, sobre el camino a Tres Cruces-Pascala de Oro, 26 Oct

1989, F.G. Lorea 4869 (MEXU; FCME).

2. Axiniphyllum pinnatisectum (P.G. Wilson) B.L. Turner, Madrofio 25:50. 1978. Rumfordia pinnatisecta PG. Wilson,

Kew Bull. 1958. 164. Tipo: MÉXICO. Guerrero: Mina Dist Agua Zarca-File, pine forest, 30 Nov 1937, Hinton et al. 11289 (HoLotipo:

lu/l I j I cfm US!

K; isoriPos: foto en http://www.nybg.org/bsci/hcol/vasc/ NY!; foto en http://ravenel si

foto en http://www.biosci.utexas.edu/prc/databases.html TEX-LL!).

Hierba perenne de 1-1.5 m alto; hojas triangulares 3-16 cm longitud, 5 cm ancho; envés con tricomas glan-

dulares; lámina pentalobulada, margen irregularmente dentado a casi lobado, por lo que Wilson (1958) y

Turner (1978) la citan como pinnatilobada, la mitad inferior más angosta, lóbulo terminal aserrado o entero.

Internodo basal de la única sinflorescencia visible 20 cm longitud; brácteas 3-10 mm longitud; involucro

ca. 10 mm longitud, 20-28 mm ancho; brácteas involucrales externas ca. 4 mm longitud, las más internas

no vistas. Flores radiadas 5-8, lígulas ca. 11 mm longitud, 5 mm ancho, tubo piloso ca. 2 mm longitud.

Aquenios ca. 2 mm longitud; flores del disco ca. 1.5 mm longitud, el limbo ca. 4 mm longitud. Anteras ca.

4 mm longitud; ramas del estilo ca. 2 mm longitud. Aquenios ca. 2.5 mm longitud, 1.1 mm ancho.

Distribución y altitud.—conocida sólo de los ejemplares tipo provenientes del estado de Guerrero, a los

2300 m snm.

Floración.—octubre a noviembre.

3. Axiniphyllum sagittalobum B.L. Turner, Madroño 25:50. 1978. (Fig. 2). Tiro: MÉXICO. Guerrero: Distrito Mina,

Municipio de San Miguel Totolapan, Toro Muerto, 2800 m, 30 Oct 1939, G.B. Hinton et al. 14761 (uoLotiro: foto en http://www.

biosci.utexas/edu/prc/databases.html TEX-LL!; isoriros: MICH, foto en http:/Awww.nybg.org/bsci/hcol/vasc/ NY!; foto en http://ravenel.

si.edu/botany/types/jstaxa/typeFrames.cfm US!).

Hierba hasta de 1.5 m alto; diámetro del tallo 1.5-3.6 mm; hojas triangulares, sagitadas, las basales arrose-

tadas y grises cuando muertas, 4.4—8.3 cm longitud, 3-6.7 cm ancho; lámina entera a trilobulada; margen

ligeramente revoluto aserrado cada serración con un mucrón y una glándula apical color pardo claro, lóbulo

terminal crenulado; pecíolos 4.7-9.1 cm longitud. Pedúnculos 3-13.6 cm longitud; brácteas 4.1-9.4 mm

longitud, 0.7-4 mm ancho; involucro 9-17 mm longitud, 11-17 mm ancho; brácteas involucrales externas

6.3-7.8 mm longitud, 0.9-2.7 mm ancho, las internas 6.1-8.8 mm longitud, 1.4-2.1 mm ancho. Flores ra-

diadas presentes, lígulas 8, tubo ca. 0.4 mm ancho. Aquenios ca. 1.5 mm longitud. Flores del disco 4.9-6.8

mm longitud, 0.9-1.3 mm ancho, el tubo 1.1-2.1 mm longitud, el limbo 3.8-4.7 mm longitud, los lóbulos

0.8-1.1 mm longitud. Anteras 4.5-4.9 mm longitud; ramas del estilo 4.5-5.3 mm longitud. Aquenios 2.7-3.3

mm longitud, 0.3-0.7 mm ancho.

Distribución y altitud. —endémica del noreste de Guerrero, en bosque de pino, 2800 m snm.

Floración.—octubre a noviembre.

Especímenes examina dos adicionales: MEXICO. Municipio de San Miguel Totolapan: Toro Muerto, 200 m después del aserradero,

camino a Puerto del Gallo, 24 Oct 2005, A. González-Zamora et al. 117 (MEXU; FCME; UAGC; TEX-LL; NY).

El conocimiento florístico de las plantas mexicanas, en especial de las especies endémicas y amenazadas,

siempre ha sido considerado como importante para los planes de conservación de su diversidad biológica.

Turner & Nesom (1998) calculan que 31% de las especies mexicanas de Asteraceae presentan áreas de

distribución restringidas, por lo que muchas de ellas pueden considerarse amenazadas. Tal es el caso de

Axiniphyllum, el cual no ha sido objeto de una evaluación del riesgo por las instancias adecuadas, lo cual puede

constatarse ya que no está listada dentro de la Norma Oficial Mexicana vigente (NOM-054-ECOL-2001) y

menos aún dentro de las listas rojas de especies amenazadas de la IUCN (2006). Las especies de este género

están pobremente representadas en los herbarios porque sus poblaciones son muy pequeñas. El trabajo de

campo permitió observar el deterioro y la destrucción del hábitat donde viven estas especies (bosques de

496

Journal of the Botanical Research Institute of Texas 1(1)

AG p

D. Flor del disco: E. Brá

| pl ta; B. Flor radiada; C. Bra

Zamora et al. 117, MEXU, FCME, UAGC, TEX-LL, NY).

HEN A Y | HP p E

González-Zamora et al., Redescubrimiento de Axi g en Guerrero, México 497

pino), lo cual permite asegurar que al menos dos de las especies reportadas para el estado de Guerrero se

encuentran en peligro de desaparecer a corto y mediano plazo si siguen operando los factores que inciden

desfavorablemente en su supervivencia. Es urgente que un estado tan rico en diversidad biológica establezca

zonas de protección que tomen en cuenta la rareza de la flora en estas regiones de difícil acceso. El caso

de A. pinnatisecta merece atención inmediata, ya que después de casi 70 años de haberse colectado los dos

únicos ejemplares existentes, el tipo de vegetación donde habita parece haber desaparecido por completo

(bosques de Pinus herrerai Martínez, P. ayacahuite Schltdl. var. ayacahuite y P. douglasiana Martínez) y si en

un futuro cercano no se vuelve a colectar deberá considerarse como una especie extinta más, por lo que se

sugiere emprender recolectas de campo para buscar esta especie con detenimiento. Ambas especies deben

incluirse dentro de la Norma Oficial Mexicana vigente (NOM-059-ECOL-2001) y en las listas rojas de la

IUCN (2006) y deben emprenderse estudios demograficos de A. sagittalobum.

AGRADECIMIENTOS

A los curadores de los herbarios CHAPA, ENCB, FCME, MEXU y UAGC por las facilidades para la revisión

de las colecciones. Guy Nesom y un árbitro anónimo hicieron valiosas sugerencias al manuscrito final. Jaime

Jiménez hizo valiosas observaciones. Se agradece al personal del Herbario de la Universidad Autónoma de

Guerrero, en especial a Mario Alberto Morlet y Hamlet Santa Anna, por su ayuda en el trabajo de campo.

Francisco Maradiaga nos ayudó a comprender la regionalización político-administrativa del estado de

Guerrero. Ramiro Cruz hizo la ilustración de la planta. El primer autor contó con una beca del CONACYT

(898976) para la realización de sus estudios de Maestría.

REFERENCIAS

BALLEZA, JJ. y J.L. ViLLAsENoR. 2002. La familia Asteraceae en el estado de Zacatecas (México). Acta Bot. Mex.

59:5-69.

BENTHAM, G. 1872. Compositae. In: J.D. Hooker, ed. Hooker' Icon. PI. 12:16-17.

BLAKE, S.G. 1930. Notes on certain types specimens of American Asteraceae in European herbaria. Contr. U.S.

Natl. Herb. 26:227-263.

INTERNATIONAL UNION FOR CONSERVATION OF NATURE AND NATURAL Resources. 2006. UICN Red List of threatened species.

www.iucnredlist.org. Gland. Switzerland.

McVaucH, R. 1984. Compositae. In: W.R. Anderson, ed. Flora Novo-Galiciana. 12:1-1157. The University of Michigan

Press. Ann Arbor.

Ortiz, E. y J.L. VILLASEÑOR. 1998. La familia Asteraceae en el estado de Nayarit (México). Acta Bot. Mex. 44:25-57.

SECRETARÍA DE MEDIO AMBIENTE Y RECURSOS NATURALES. 2002. Norma Oficial Mexicana NOM-054-ECOL-2001. Protección

Ambiental-Especies nativas de México de flora y fauna silvestres-Categorías de riesgo y específicas para su

inclusión, exclusión o cambio-Lista de especies en riesgo. Diario Oficial de la Federación, 6 de Marzo de 2002.

(2? sección):1-85.

Turner, B.L. 1978. Taxonomy of Axiniphyllum (Asteraceae-Heliantheae). Madroño 25:46-52.

Turner, B.L. 1987. A new species of Axiniphyllum (Asteraceae: Heliantheae) from Durango, Mexico. Madroño

34:165-167.

Turner, B.L. y G. Nesom. 1998. Biogeografía, diversidad y situación de peligro o amenaza de Asteraceae de México.

In: T. Ramamoorthy, R. Bye, A. Lott y J. Fa, eds. Diversidad biológica de México: orígenes y distribución. Instituto

de Biología, UNAM. Pp. 167-192.

VILLARREAL, J.Á. y J.L. VitLAsEROR. 2004. Compositae. Tribu Tageteae. Fl. Veracruz 135:1-67.

VILLARREAL, J.Á., J. VALDÉS y J.L. VILLASEÑOR. 1996. Corología de las Asteraceae de Coahuila, México. Acta Bot. Mex.

36:29-42.

VILLASENOR, J.L.1982. Las Compositae del Valle de Tehuacán-Cuicatlán. Flora genérica. Tesis de Licenciatura. Facultad

de Ciencias, UNAM.

VILLASEÑOR, J.L. 1987. Clave genérica para las compuestas de la Cuenca del Río Balsas. Bol. Soc. Bot. Mexico

47:65-86.

498 Journal of the Botanical R h Institute of Texas 1(1)

VILLASENOR, J.L. 1989. Manual para la identificación de las Compositae de la Península de Yucatán y Tabasco. Rancho

Santa Ana Bot. Gard. Tech. Rep. 4:1-122.

VILLASEÑOR, J.L., E. Ortiz y V. JUAREZ. 2004. Asteraceae. In: A. García, M.J. Ordonez y M. Briones, eds. Biodiversidad de

Oaxaca. Instituto de Biología, UNAM-Fondo Oaxaqueno para la Conservación de la Naturaleza-World Widlife

Found. México. Pp. 177-192.

VILLASEÑOR, J.L., G. IBARRA y D. Ocaña. 1998. Strategies for the conservation of Asteraceae in Mexico. Conservation

Biol. 12:1066-1075.

ZUCCARINI, J.G. 1832. Plantarum novarum vel minus cognitarum. Abh. Math.-Phys. Cl. Kónigl. Bayer. Akad. Wiss.

1:289-396.

INVENTORY AND DISTRIBUTION OF AGAVE (AGAVACEAE)

SPECIES TN JALISCO. MEXICO

Gerardo Hernandez-Vera Miguel Chazaro Basanez Ericka Flores-Berrios

Centro de Investigacion y Centro Universitario de Ciencias Sociales y Centro de Investigacion y

Asistencia en Tecnologia y Humanidades, depart to de Geogratia Asistencia en Tecnologia y

Diseno del Estado de Jalisco Universidad de Guadalajara Diseno del Estado de Jalisco

A.C. Av. Normalistas Guanajuato 1045 A.C. Av. Normalistas

800 S.H. Colinas de la Normal Guadalajara Jalisco, MEXICO 800 S.H. Colinas de la Normal

44270 Guadalajara Jalisco, MEXICO 44270 Guadalajara Jalisco,

MEXICO

ABSTRACT

In order to review and update the list of Agave species previously reported for Jalisco, Mexico, an extensive botanical exploration was

done throughout ret ive areas of the state. Twenty-three species were registered; from which 7 represent new records for Jalisco,

and 3 proposed as new species. Fifty per cent of the listed species were found in pine and pine-oak forests, whereas 2996 were found

in the tropical deciduous forest. Among the many factors that might affect the distribution of agaves, edaphic factors and moreover

physiography, can play an important role.

RESUMEN

Con la finalidad de hacer una revisión y actualizar el listado de especies de Agave previamente reportadas para Jalisco, se realizó una

extensa exploración botánica por el estado. Se registraron 23 especies, de las cuales 7 representan nuevos registros para Jalisco y 3 se

proponen como nuevas especies. El cincuenta por ciento de las especies enlistadas se encontró en los bosques de pino y pino-encino,

mientras que el 2996 prospera en bosque tropical caducifolio. Entre los diversos factores que pueden afectar la distribución de los agaves,

se plantea que los edáficos y en especial la fisiografía pueden jugar un papel importante

INTRODUCTION

In Mexico, the genus Agave L. is of great importance since many of its species are used for food, in the

manufacture of fibers for threads and textile industry, as ornamental plants, and mainly in the production

of different types of alcoholic beverages such as tequila and mezcal (Gentry 1982; Valenzuela 1997; Nobel

1998.). This genus comprehends more than 200 species, from which 7596 are in Mexico, which is considered

as the center of origin (Granick 1944; García-Mendoza 2002).

In Jalisco, the study of the genus Agave has focused basically on Agave tequilana Weber var. azu

. Because

of its importance as the only species allowed by the Official Mexican Standard for tequila production, the

alcoholic beverage traditionally associated with Mexico, and in particular Jalisco (Cedeño 1995; DOF 1997).

In contrast, the study of other non-cultivated agaves many of which are useful species that occur throughout

the State, has been neglected.

For Jalisco, Gentry (1982) reports 14 species, whereas McVaugh (1989) reports 18, considering some

of them as “doubtful” and “excluded” due to the lack of herbarium specimens and information about wild

populations. Thus, based on our collections, the aim of this study was to update the list of species, corroborate

in the field McVaugh and Gentry's reports, and contribute to the knowledge of Agave species distribution in

jalisco.

Site Description

Located in the western region of Mexico, Jalisco is one of the largest states consisting of 80,000 Km? and

4 physiographic provinces: a) Western Sierra Madre, in the northern part of the state, b) Central Plateau,

northeast region, c) Trans-Volcanic Belt (Eje Neo-volcanico), central region of the state, and d) Southern Sierra

Madre, southwest region (INEGI 2003). Igneous rock soils constitute 7996 of its surface, which is favored by

several vegetation types, such as conifer and oak forests, tropical subdeciduous forest, tropical deciduous

J. Bot. Res. Inst. Texas 1(1): 499 — 509. 2007

500 Journal of the Botanical R h Institute of Texas 1(1)

forest, tropical savanna, gallery forest, alpine tundra, thorn forest, and cloud forest. The topography ranges

from 0—4360 m above sea level (INEGI 2003). AII of these geographical and geological characteristics con-

tribute to its floristic diversity, which is estimated to be 7000 plant species (Cházaro & Lomelí 1995).

MATERIALS AND METHODS

Twenty field trips were carried out throughout representative areas of the state of Jalisco in 2002 and 2003.

Identification of the species was done according to Gentry's (1982) and McVaugh's (1989) descriptions and/or

taxonomic keys. Vegetation types were registered as well as altitude and geographical coordinates by means

of a global positioning system (GPS Magellan 320). Plants were collected and documented for herbarium

specimens deposited at IBUG and IEB herbaria.

RESULTS AND DISCUSSION

Out of 102 collected specimens 23 species were identified, from which seven of them represented new re-

cords for the state and three were treated as new species (Table 1). Description of the species Agave sp. nov.

ined. (“Colimilla” ravine) and Agave sp. nov. ined. (Tequila municipality) are in preparation, whereas Agave

vazquezgarciae is in press (Cházaro et al. in press).

From Gentry’s (1982) and McVaugh's (1989) reports, Agave stringens Trel., A. hookeri Jacobi, A. cantala

Roxb., A. impressa Gentry and A. longisepala Todaro, were not located. A. stringens was reported by Gentry

based on a Trelease collection in Rio Blanco (Zapopan municipality) (Gentry 1982). At the cited location we

only found A. guadalajarana Trel., (Fig. 1), however according to the vegetation type and Trelease original

plant description (Trelease 1920), we assume A. stringens could be a form of A. angustifolia Haw.

McVaugh (1989) cites a Diguet's collection for A. hookeri from Cerro Viejo (Tlajomulco and Jocotepec

municipalities). It is likely that Diguet identified incorrectly the specimen, since the only species found there,

was the closely related A. inaequidens Koch which spreads throughout the oak and pine-oak forests in the

"Sierra del Tigre” (Fig. 2). Gentry (1982) reported a collection for A. hookeri south of Jalisco near Jiquilpan

(Michoacan state); it appears this species has been introduced from Michoacan state as a cultivar.

With regard to Agave cantala, we could not find it near Villa Guerrero as reported by Gentry (1982) who

was unsure about the identification of the plant he collected. Perhaps this is a species related to A. tequilana

Weber, as suggested by Trelease (1920) and Valenzuela and Nabhan (2003), who treat it as a synonym of A.

vivipara L. Agave impressa was not found in the “Sierra de los Huicholes” (Bolaños) as Gentry was informed

(Gentry 1982). Probably there was confusion with the common name since he was given the name *mas-

parillo” for this agave in the type locality (Escuinapa, Sinaloa); the same name is used for A. maximiliana

Baker, abundant in the “Sierra de los Huicholes”. For A. longisepala, McVaugh (1989) concluded there was

uncertainty about its identity and taxonomic position, and Gentry (1982) knew the species only from an

illustration. This could be a synonymous for A. tequilana since Trelease (1920) reported it as a related species

cultivated near Tequila, Jalisco.

Whereas Agave americana L. var. expansa (Jacobi) Gentry and A. tequilana were found as cultivated

species, A. salmiana Otto ex Salm, ssp. crassispina (Trel.) Gentry, was found as wild populations in north-

east Jalisco (Sierra Cuatralba) (Fig. 3). McVaugh (1989) had reported it as a cultivated species. Considered

by Gentry (1982) as a relatively rare species, A. attenuata Salm-Dyck was found in “Rincón de Manantlan,”

south of Jalisco (Fig. 2). We could corroborate the presence of large stems in this species, clearing the un-

certainty about this agave's feature reported by McVaugh (1989) (Fig. 4).

The higher number of species was found in the pine and pine-oak forests, being the tropical deciduous

forest the second one in harboring the highest diversity (Fig. 5). Hence, the distribution of Agave species

based on physiographic provinces shows that they are mainly concentrated in southern Sierra Madre and

Trans-Volcanic Belt (Eje Neovolcanico), which are two areas whose main vegetation types are tropical de-

ciduous forest and pine and pine-oak forests, respectively (Figs. 1-3, 6).

These results are similar to those presented by García-Mendoza (2002) who reports that in Mexico,

Hernández-Vera et al., Agave species in Jalisco, Mexico 501

Taste 1. Agave species in Jalisco, México. New record for Jalisco 8. New species u.

Group Species

Rigidae Agave angustifolia Haw.

$ Agave rhodacantha Trel.

Agave tequilana Weber

Crenatae bon juidens Koch.

Agav imiliana Baket

Marmoratae o e Gentry

Agave valenciana Cházaro & A. Vazquez,

Parryanae Agave guadalajarana Trel.

§ Agave parryi ae

Ditepalae $ Agave wocomahi Gentry

Salmianae $ Agave salmiana Otto ex Salm-Dyck

Total for subgenus Agave 11

Amolae Agave ulifera Trel.

Agave attenuata Salm-Dyck

oo vilmoriniana Berger

aoe Cházaro, Valencia €: Lomelí, sp. nov.

Filiferae pee eee Gent

Agave schidigera je (sensu Gentry)

§ Agave filifera Salm-Dyck (sensu Gentry)

5 Agave geminiflora (Tagl.) Ker-Gawler

Marginatae § Agave angustiarum Trel.

nAgave sp. nov. ined. (“Colimilla” ravine, municipalities of

Tonala and Zapotlanejo)

a Agave sp. nov. ined. (municipality of Tequila)

Striatae Agave rzedowskiana P. Carrillo, R. Vega & R. Delgad.

Total for subgenus / ¡ttaea 12

Total species 23

tropical deciduous forests and coniferous and oak forests are among the vegetation types with the highest

number of Agave species, only surpassed by deserts and chaparrales. Nevertheless, it was found that several

species are not exclusive or restricted to a certain vegetation type since they can either thrive in two or more

types or can be found in ecotones (Table 2). Soil characteristics can play an important role in the distribution

of agaves as has been suggested by Nobel & Berry (1985), who conclude that although soil types have not

been systematically related to the distribution of agaves, edaphic factors may affect their seedling establish-

ment. In this study, the two physiographic provinces with the highest number of Agave species (Southern

Sierra Madre and Neo-Volcanic Belt) are conformed mainly by igneous rocks, a soil type previously reported

as favored by these plants (Alvarez de Zayas 1989; García-Mendoza 2002).

Agave angustifolia Haw., A. schidigera Lem. (sensu Gentry) and A. maximiliana Baker, were the most

widely distributed species, thriving in many different vegetation types in a broad range of altitudes and

latitudes throughout the state (Table 2; Figs. 1, 3 & 6). This could reflect their ample tolerance limits to each

of the several environmental factors, based on high levels of genetic diversity as reported for A. angustifolia

(Colunga-GarciaMarin et al. 1999). In contrast, A. gypsophyla Gentry and A. geminiflora (Tagl.) Ker-Gawler,

were found in small areas associated to specific geographical and ecological conditions such as soil type

and vegetation. Only a small population of A. geminiflora was observed, thriving in an oak woodland along

a rocky brook 1814 m above sea level, in the central region of Jalisco (municipality of Tequila). In the other

hand, A. gypsophyla could be found only in rocky gypsum soils (whence the name) south of the state, mu-

nicipality of Pihuamo (Fig. 2).

Agaves in the subgenus Agave were found likely to thrive in flatlands, whereas agaves in the subgenus

502 Journal of the Botanical R h Institute of Texas 1(1)

-105° -104° -103° -102°

| F) | |

SAN LUIS

POTOSI

E] N

N- AGUASCALIENTES N

NAYARIT q

ZACATECAS 2

o hr

e A

N- ' yy

AS o

%, GUANAJUATO

eL. N

x Q

^to

a MICHOACÁN

o Y, COLIMA

Oo. Kilometers -m

s 40 0 40

eS es

| | | |

-105? -1049 -103° -102°

Æ A gave maximiliana ~ Trans-Volcanic Belt

€ Agave guadalajarana Central Plateau

O Agave colimana Southern Sierra Madre

Western Sierra Madre

Fic. 1. Distributi f Ag imiliana, A guadalajarana, and A. colimana

Littaea, appear to grow mainly in sharp rocky cliffs (66% of this subgenus specimens were found exclusively

in this habitat). Gentry (1982), suggests that the subgenus Littaea could represent the phylogenetic and geo-

logically older form of agaves, predominantly diploid with more “primitive” leaf and habit characteristics

(Granick 1944). Hence, thriving in sharp rocky slopes could represent a “primitive” growth habit, relying

more on their toxic substances in the leaves and inaccessible cliff sites for survival than on defensive armor.

Most in the subgenus Littaea, lacked of marginal spines or “teeth,” or if present, they were fragile and brittle

Hernández-Vera et al., Ag in Jalisco, Mexico

503

-1059 -104° -103° -102°

5 | e | |

/ \ \ — SANLUIS

E ^ o " ET POTOSI

Ea m X b e 3

V Ben z^

| d

N | / AL

NI // AGUASCALIENTES E N

el os

| GUANAJUATO

i 6^

2, 7 ds

NT des

EUR

MICHOACÁN

E Y E. COLIMA

Oo _ Kilometers n=

= wu 40 ^0. 40 ©

| LON d | |

-105° -104° -103° -102°

O Agave parryi

A Agave ineaquidens

B Agave angustiarum

A Agave valenciana

E Agave vazquezgarciae

Y Agave gypsophila

Y Agave sp. nov. ined.

("Colimilla" ravine, municipalities

of Tonala and Zapotlanejo)

^ Agave attenuata

E Agave gemimiflora

— - Trans-Volcanic Belt

Central Plateau

Southern Sierra Madre

Western Sierra Madre

Fic. 2. Distribution of Agave parryi, A. inaequidens, A. angustiarum, A. valenciana, A. vazquezgarciae, A. gypsophila, A. attenuata, A. geminiflora, and Agave

sp. nov. ined. ("Colimilla" ravine).

504 Journal of the Botanical R h Institute of Texas 1(1)

-105? -104° -103° -102°

| T | |

SAN LUIS

POTOSI

CN AGUASCALIENTES N

NAYARIT

&- Fo dg

E GUANAJUATO

R} 8

^t.

MICHOACÁN

o Y, COLIMA um

O) —

e. 40 Le: ir 4©

| | r | |

-105? -104° -103° -102°

€ Agave angustifolia A Agave sp. nov. ined.

0 Agave rhodacantha (municipality of Tequila)

$ Agave wocomahi Central Plateau

Æ Agave pedunculifera Southern Sierra Madre

O Agave salmiana Western Sierra Madre

Fic. 3. Distribution of Agave angustifolia, A. rhodacantha, A. vilmoriniana, A. wocomahi, A. pedunculifera, A. salmiana, and Agave sp. nov. ined. (munici-

pality of Tequila).

Hernández-Vera et al., Ag ies in Jalisco, Mexico 505

L I linc: In RA 414 +) flisi TL L "I I 4 J I J

Fic. 4. Agave attenuata thriving in rocky cliffs at 1 700

by this species. Photo: Piet Van der Meer.

based on collected specimens. As suggested by Gentry (1982), the ecological importance of these cliff-dwell-

ing species appears to be their contribution to soil building and soil-holding capabilities on the steep rocky

slopes.

Richness and Conservation Status

With 23 species of Agave known so far as wild plants, Jalisco state stands second place in agaves diversity

in Mexico, only surpassed by Oaxaca state where García-Mendoza (2004) reported 30 species. We believe

2 more species could be present in Jalisco, namely Agave durangensis Gentry and Agave striata Zucc. The first

one in Huejuquilla region, since we have collected it in the adjacent Valparaiso municipality in Zacatecas

state, whereas Agave striata has been seen and photographed at Sierra del Laurel (Esperanza Quezada, pers.

comm.), a mountainous range at Calvillo municipality in Aguascalientes state. The eastern slopes of these

mountains belong to Jalisco; hence, we predict it occurs at Sierra del Laurel in the Jaliscan side, a region

which has remained unexplored. On the other hand, the botanist Pablo Carrillo recently found Agave orni-

thobroma (pers. comm.) in Mezquitic municipality, into de Huichol indians region north of Jalisco. Further

explorations in the zone will be necessary.

The geographic distribution of Agave species in Jalisco is uneven; some cover an ample range, such

as the Agave angustifolia complex, whereas others like A. valenciana, Agave sp. nov. ined. (Municipality of

Tequila), A. vazquezgarciae, and A. geminiflora are endemic to small areas. Among the 18 Agave species in

L e

Mexico that are considered within some status of protection by the Federal Government (DOF 1994), only

A. gypsophyla and A. ornithobroma occur in our study area; however with the data gathered so far, we consider

£s+haD o ID L

506 Journal of t titute of Texas 1(1)

ld Oak and Pine-Oak forests

@ Tropical deciduous forest

td Thorn forest

lá Cloud forest

Fic. 5. P. t fA ies I tation type in Jalisco, Mexico

J J T 4 J at

the following taxa should be added to the list (the first three of them endemic to Jalisco): a) Agave valenciana,

restricted to small areas of the Mascota river canyon and the Talpa river, b) Agave sp. nov. ined., restricted to

the Santiago river basin ravines next to Tequila, c) A. vazquezgarciae, from Sierras Manantlán and Cacoma,

and d) A. geminiflora, previously known only from the type locality (Ocotillo, Nayarit); we found it near “El

Salvador,” municipality of Tequila.

CONGLUSIONS

A thorough botanical exploration of Jalisco resulted in the corroboration and update of previous reports

of Agave species in the state. The great diversity of Agave species in Jalisco is related to the geographical

and geological characteristics of the state. To the authors’ knowledge, this is the first extensive botanical

exploration of Agave species in Jalisco. Documented specimens with exact geographical coordinates will

provide a reliable source of information on these plants for further research. We consider that basic botani-

cal research is essential as a preliminary step for a better understanding and rational management of plants

and thus, habitat.

ACKNOWLEDGMENTS

This research was funded by Consejo Estatal de Ciencia y Tecnologia, Jalisco (COECYTJAL), Project 48-

2001. Gerardo Hernandez-Vera was supported by a CONACYT graduate fellowship. We also express our

gratitude to MSc. Manuel Rodriguez for his great support and assistance during the entire period of field

work. Thanks to Centro de Investigación y Asistencia en Tecnologia y Diseño del Estado de Jalisco A. C.

(CIATEJ) for publication support.

REFERENCES

ÁLVAREZ DE Zayas, A. 1989. Distribución geográfica y posible origen de las Agavaceae. Rev. Jard. Bot. Nac. Univ. Habana.

1(10):25-36.

Hernández-Vera et al., Agave species in Jalisco, Mexico 507

-105° -104° -103° -1029

| 7] | |

SAN LUIS

POTOSI

OC AGUASCALIENTES

NAYARIT

3 j N

NT p Tes

p GUANAJUATO

6 | N

da =

MICHOACÁN

o y COLIMA K à;

O) i x

=|- 2 40 a Sg 0

a eee

| | | |

105° -104° -1039 -102°

3K Agave schidigera — — Trans-Volcanic Belt

A Agave filifera Central Plateau

€ Agave rzedowskiana Southern Sierra Madre

Western Sierra Madre

Fic. 6. Distributi f Ag h idig , A filifera, and A. rzedowskiana

CARRILLO-REvES, P, R. VEGA Avia, and R. RAMIREZ-DELGADILLO. 2003. Agave rzedowskiana, a new species in subgenus Littaea (Agavaceae)

from western Mexico. Brittonia 55:240-244,

CEDEÑO, C.M. 1995. Tequila production. Crit. Rev. Biotechnol. 15(1):1-11.

CHÁZARO-BasaREZ, B.M. y E. LomeLí M. 1995. Las Agavaceas del Estado de Jalisco. In: M. Cházaro-Basañez, E. Lomeli

M., R. Acevedo R. y S. Ellerbracke, eds. Antología Botánica del Estado de Jalisco. Universidad de Guadalajara.

Guadalajara, Jalisco. México. Pp. 87-90.

508

fal, Dat

Journal of

Taste 2. Ecological distribution of Agave in Jalisco, Mexico.

h Institute of Texas 1(1)

Species Vegetation Type Altitude (m above

sea level)

Agave angustifolia Tropical deciduous forest/Pine-oak forest 707-1,845

and the ecotone from both/Thorn forest/

Agave rhodacantha Oak forest/Tropical deciduous and 250-1,133

subdeciduous forests/Ecotone oak

forest- Tropical subdeciduos forest

Agave tequilana var. azul Only cultivated

Agave inaequidens Oak and pine-oak forests 1,910-2,319

Agave maximiliana Oak and pine-oak forests/Ecotone 1,086-1,987

oak forest-Tropical deciduous forest

Agave gypsophila Tropical deciduous forest 590-600

Agave valenciana Ecotone oak forest-Tropical 1182

deciduous forest

Agave guadalajarana Oak forest and ecotone with Tropical 1,689-1,850

deciduous forest

Agave parryi Thorn forest, Yucca sp. forest 2,364

Agave wocomahi Thorn forest 1,975

Agave salmiana Oak forest 2:532

gave pedunculifera Oak forest 1,910

Agave attenuata Ecotone oak forest-Tropical deciduous 1,699

forest

Agave vilmoriniana Tropical deciduous forest 1,275

Agave vazquezgarciae, Sp. NOV. Ecotone oak forest-cloud forest 1613

Agave colimana Tropical deciduous forest/Oak forest/ 801-1,167

Ecotone pine-oak forest-cloud forest

Agave schidigera Oak and pine-oak forests/Tropical 875-2,178

deciduous forest/Ecotone oak

forest- Tropical deciduous forest

Agave filifera Thorn forest/Yucca forest/ 2,341-2,450

Oak-Juniperus forest

Agave geminiflora Oak forest 1,814

Agave angustiarum Oak forest/Tropical deciduous forest 914-979

Agave sp. nov. ined. ("Colimilla" Tropical deciduous forest 1,270-1,300

ravine, municipalities of

Tonalá and Zapotlanejo)

Agave sp. nov. ined. (municipality Tropical deciduous forest 1,055-1,531

of Tequila)

Agave rzedowskiana P. Carrillo, Oak forest 1,689

R. Vega & R. Delgad.

CHAZARO-BASAREZ, M., A. VazQUEZ-GARCÍA, and Y. VARGAS-RODRÍGUEZ. 2005. Agave valenciana (Agavaceae) a gigantic new

species from Jalisco, Mexico. Novon 15:525-530.

CHÁZARO-BasAREZ, M., O. VALENCIA-PELAYO, J.A. LOMELI-SENCION, and Y. VARGAS-RODRÍGUEZ. (In Press). Agave vazquezgarciae

(Agavaceae), a new species from Jalisco, Mexico. Novon.

COLUNGA-GARCÍAMARIN, P, J. CoeLLo-CoeLLo, L.E. EGUIARTE, and D. Piñero, 1999. Isozymatic variation and phylogenetic

relationships between henequén (Agave fourcroydes) and its wild ancestor A. angustifolia (Agavaceae). Amer.

J. Bot. 86115125

[DOF] Diario OriciAL DE LA FEDERACIÓN. 1994. Norma Oficial Mexicana, Protección Ambiental-Epecies nativas de

México de Flora y Fauna Silvestres-Categorías de riesgo y Especificaciones para su Inclusión, Exclusión o

Cambio-lista de Especies en Riesgo. NOM-059-ECOL-1994. May 16, modified on March 22, October 16, 2000

and September 7, 2001. México, D.F.

Hernández-Vera et al., Agave species in Jalisco, Mexico 509

[DOF] Diario OFICIAL DE LA FEDERACION. 1997. Norma Oficial Mexicana Bebidas Alcohólicas-Tequila-Especificaciones.

NOM -006-SCFI-1994. September 3, modified on December 24, February 1 and March 1, 2000. México, D.F.

GaRCÍA-MENDOZA, A. 2002. Distribution of Agave (Agavaceae) in Mexico. Cact. Succ. J. 4 (74):177-188.

Garcia-MENDOZA, A. 2004. Agaváceas. In: A. García-Mendoza, M.J. Ordóñez y M. Briones S., eds. Biodiversidad de

Oaxaca. Instituto de Biología UNAM. México, D. F. Pp. 141-148.

GenTRY, H.S. 1982. Agaves of Continental North America. The University of Arizona Press, Tucson.

GRANICK, E.B. 1944. A karyosystematic study of the genus Agave. Amer. J. Bot. 31:283-298.

[INEGI]. Instituto NACIONAL DE GEOGRAFÍA ESTADÍSTICA E INFORMÁTICA. 2003. Aspectos geográficos de Jalisco. México.

McVaucH, R. 1989. Liliaceae. In: W.R. Anderson, ed. Flora Novo-Galiciana. Vol. 15:120-293. The University of Michi-

gan Herbarium, Ann Arbor.

Nose, PS. 1998. Los incomparables agaves y cactos. Primera edición en español. Ed. Trillas. México.

NostL, PS. and W.L. Berry. 1985. Element responses of agaves. Amer. J. Bot. 72:686-694,

TRELEASE, W. 1920. Agave. In: PC. Standley. Trees and shrubs of Mexico. Contr. U.S. Natl. Herb. 23:107-142.

VALENZUELA, Z.A. 1997. El Agave tequilero, su cultivo e industria. Segunda edición. Monsanto-Litteris editores.

México.

VALENZUELA, Z.A. and G.P. Na&HAN. 2003. Tequila! A natural and cultural history. The University of Arizona Press,

Tucson.

510 Journal of the Botanical R h Institute of Texas 1(1)

BOOK REVIEWS

RicHarD WiLFORD. 2006. Tulips: Species and Hybrids for the gardener. (ISBN 13: 978-0-88192-763-4,

hbk.). Timber Press Inc, 133 S.W. Second Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders:

www.timberpress.com, mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax).

$34.95, 211 pp., 103 color photos, 2 b/w illustrations, 2 maps, 614" x 944",

Windmills, Dutch girl with a white hat over her blond pigtails, a blue skirt, wooden shoes and TULIPS! Childrens flower drawings often

depict a cup shaped single flower on a straight stem with two long thin leaves—a tulip, the well known flower from the Netherlands?

Hybrid tulips are the ones most of us know and love for springtime gardens and Easter gift giving.

The author, Richard Wilford, currently the Collections Manager at the Royal Botanic Gardens, Kew, responsible for alpines, bulbs

and herbaceous perennials, introduces the reader to many species tulips.

Chapters include geography, the tulip plant, cultivation, history and classification and species descriptions. The subject of each

chapter is thoroughly discussed leaving very little to the imagination.

Tulips are found growing in the wild on three continents—Asia, Europe and Africa. By trying to replicate the conditions found

where the species are growing in the ya gardeners in many parts of the world can have success with tulips. The ideal climactic condi-

tions can be as diverse as the sq nenvironment that has cold winters, wet springs and dry summers is ideal for good bulb growth

and fl LIDWEI pr oduction.

Tulip species are found from western Himalaya through the Caucasus Mountains, Iran, and Turkey and as far as the Iberian Pen-

insula. They are found in southern Siberia, west to Crete and northern Africa. They were introduced to Europe in the sixteenth century

and their cultivation began among gardeners and botanists. Some tulips were scattered accidentally and readily became naturalized in

areas where they are now erroneously considered natives

The tulip plant characteristics are fully described by the author. The diff intl fl , leaves, and bulbs provide ongo-

ing discussion among botanists as well as the naming and renaming of various plants. There is still confusion in the genus/species names.

r. Wilford botanists who lin classifying and describing the tulip—starting with Linnaeus. Swiss

French, German and Russian botanists—de Candolle, Reboule, Koch and Regel were involved in naming the species.

Cultivation—by bulbs and seeds is described by the author in great detail—from the types of pot m soil, to the watering and

pots as] }

temperature best for growing tulips. Mr. Wilford considers planting the | 1. As the plants

bloom and then go dormant, the pots are easily transported in the garden. Depending on you particular climate, watering is a constant

concern. Too much water can cause soggy soil and rotting bulbs, too little water does not provide the moisture necessary to make new

bulbs. Good drainage is essential for tulips. After you have read the chapter on cultivation, you will be an expert. Your neighbors will

be astounded and envious!

In the chapter on species descriptions, Mr. Wilford provides a plethora of species descriptions with beautiful color photographs.

You will be amazed at the variations in flower shapes and colors.

The garden hybrid tulips were bred in Persia and Turkey in the fifteenth and sixteenth centuries. In the past 400 years, Holland

has made a made a name for itself in the hybridizing and purveying of tulips (bulbs and plants).

Tulips: species and hybrids for the gardener is a text full of information, for botanical explanation, exploration and practical garden-

ing—and is a visual delight.—Ann Schrader, Herbarium Volunteer, Botanical Research Institute of Texas, Fort Worth, TX, 76102-4060, U.S.A.

CELESTINE L. Duncan and Janer K. Crank (eds.). 2005. Invasive Plants of Range and Wildlands and Their

Environmental, Economic, and Societal Impacts. ISBN 1-891276-42-5, pbk.). Weed Science Society

of America, http://www.wssa.net/. (Orders: Weed Science Society of America, 810 10' Street, Lawrence,

KS 66044-8897, U.S.A.). $20.00, 222 pp., illustrated, 6" x 9".

This study assesses the economic and environmental losses caused by invasive plants on range and wildlife areas. The authors have

focused on 16 key plants in the contiguous 48 states and estimate the current area infested by them. Fourteen of the species occur on

range, pasture, or open woodland habitats while two occur in riparian or wetland habitats

The sixteen species include: Russian knapweed, musk thistle, diffuse knapweed, yellow starthistle, spotted knapweed, Canada

thistle, hawkweed, perennial pepperweed, leafy spurge, Sericea lespedeza, purple loosestrife, downy brome, medusahead, Dalmatian

toadflax, tropical soda apple, and Tamarisk or saltcedar. Taxonomy, botanical characteristics, susceptible habitats, plant distribution,

and spread and reproduction ized for each species.

Each species has its own chapter which summarizes eight categories ofimpacts, including; livestock and wildlife; plant commu-

nities; rare, sensitive, and threatened species; community function; soil and water resources; human health; economics; and value and

use. An extensive Literature cited’ section concludes each chapter.

Scientists, weed control specialists, resource planners and policy makers will welcome the comprehensive information provided

in this valuable reference work.—Gary Jennings, Library, Botanical Research Institute of Texas, Fort Worth, TX, 76102-4060, U.S.A.

J. Bot. Res. Inst. Texas 1(1): 510. 2007

DIVERSIDAD Y DISTRIBUCIÓN DE LA FLORA VASCULAR ACUÁTICA

DE TAMAULIPAS, MÉXICO

Arturo Mora-Olivo José Luis Villasenor

Instituto de Ecologia y Alimentos Departamento de Botdnica

Universidad Autónoma de Tamaulipas Instituto de Biologia

13 Blvd. Adolfo López M 928 Universidad Nacional Autónoma de México

Cd. Victoria, Tamaulipas, MÉXICO MÉ | DF.

amorao@uat.edu.mx vrios@ibiologia.unam.mx

RESUMEN

Los ambientes acuáticos (h dales) uno de los ecosistemas más los y con su biodiversidad pobremente conocida. Como

una primera fase hacia el conocimiento de la flora vascular acuática del noreste de México, en este trabajo se presenta información

sobre la diversidad y distribución de las hidrófitas (plantas acuáticas) de Tamaulipas, uno de los tres estados que conforman la región

noreste de México y que contiene la mayor cantidad de cuerpos de agua. Las especies se clasificaron por tipo de planta acuática, forma

biológica y forma de vida. En los humedales de Tamaulipas se registraron 426 especies pertenecientes a 213 géneros y 85 familias de

plantas vasculares; tal riqueza representa 5796 del total conocido para México. Un 22.8% de las especies son acuáticas estrictas, 43.7%

son subacuáticas y 33.596 tolerantes. Por su forma biológica, 88.596 de las especies son herbáceas, 6.196 arbustos y 5.496 árboles. Por su

forma de vida dominan las hidrófitas enraizadas emergentes (376 especies, 88.396). Un alto d de dd son de amplia distri-

bución, conocidas en toda América o alcanzando regiones del Viejo Mundo; sol id endémicas

de México (una de ellas d de RR El estado destaca como un reservorio importante de plantas ere a humedales y

constituye biogeog , al itorio el limite de distribución geográfica de un buen número

de elementos boreales y meridionales.

ABSTRACT

The aqua ic environments (wetlands) are among the more threatened ecosystems, with their biodiversity poorly known. As a first step

D a better knowledge of the aquatic vascular plant diversity of northeastern Mexico, in this paper the diversity and distribution of

the aquatic plants (hydrophytes) recorded and documented in the state of Se are xd It is one of the three states com-

prising the northeastern of Mexico and includes the largest area OMM a The sp ] d by type, biological form and life

form. A number of 426 species, belonging to 213 genera and 85 ili ded; such a fig 57% of the total vascular

plants species recorded in all Mexico. True aquatics represented 22.8% of the species, as marginal aquatics 43.7%, and wetland tolerants

33.5%. By habit, 88.5% of the species are herbs, 6.1% shrubs and 5.4% trees; on the other hand, the dominant life form was emergent

rooted hydrophytes (376 species, 88.3%). A high percentage of species are widespread, distributed either all along America or reaching

part of the Old World; only nine species were recorded as endemic to Mexico (one endemic to Tamaulipas). The state of Tamaulipas is an

important reservoir of species associated with wetlands and constitutes a relevant biogeographic area, due to the fact that in its territory

the geographic limit of a number of boreal and meridional elements is found

INTRODUCCIÓN

Los humedales (ambientes acuáticos), no obstante su importancia ecológica y económica, constituyen uno

de los ecosistemas más amenazados, principalmente por su destrucción, conversión, fragmentación y con-

taminación (Jain 1990; Amezaga et al. 2002; Santamaría y Klaassen 2002; Saunders et al. 2002; Abellán et

al. 2005; Fitzsimons & Robertson 2005). Desafortunadamente, su biodiversidad es poco conocida, lo que

dificulta el establecimiento de estrategias para su conservación, tanto a escala regional como global. Hasta

ahora, no existe un inventario de las plantas acuáticas del mundo, a pesar de que Cook et al. (1974) y Cook

(1990) han realizado esfuerzos encaminados a revertir esta deficiencia. Existen pocos países con inventarios

de plantas que crecen en humedales; Estados Unidos es uno de ellos, donde se reportan un poco más de

6,000 especies (Reed 1988, 1997).

En México, los magros esfuerzos por inventariar la flora vascular acuática se han enfocado a las familias

de angiospermas estrictamente acuáticas, aquellas en las que todas sus especies son hidrófitas (Lot et al.

1986, 1999); tal grupo consta de 24 familias estrictamente acuáticas, que incluyen 118 especies. Esfuerzos

J. Bot. Res. Inst. Texas 1(1): 511 — 527. 2007

f4L,D o ID L

512 Journal of t titute of Texas 1(1)

adicionales han permitido documentar 747 especies de plantas vasculares acuáticas en México (Lot et al.

1999). Un trabajo regional, realizado por Bonilla-Barbosa (2004) registró 134 especies de plantas vasculares

acuáticas en la Sierra Madre Oriental. A nivel estatal solamente dos entidades federativas han publicado

información exclusivamente sobre su flora vascular acuática, Aguascalientes (Siqueiros 1989) y Morelos (Bo-

nilla-Barbosa et al. 2000). Hasta ahora solamente 15 estados cuentan con inventarios publicados de su flora

(Villaseñor 2004); sin embargo, poco o ningún énfasis han puesto en las especies de plantas acuáticas.

El Noreste de México, región que comprende los estados de Coahuila, Nuevo León y Tamaulipas (Fig.

1), representa una porción del país florísticamente importante, que de manera natural extiende sus límites

hasta el suroeste de Texas en Estados Unidos (Villaseñor 1990; Rzedowski 199la, 1991b, 1993). Al cruzar

en su territorio el Trópico de Cáncer, en esta porción se combinan tanto especies del trópico como de las

regiones boreales, especialmente de Norteamérica. La flora vascular registrada hasta la fecha en esta región

alcanza 6,062 especies (Villaseñor datos no publicados), lo que representa casi la cuarta parte de la Flora

de México (Villaseñor 2003); en el caso de la flora que se ha colectado en humedales, esta región contiene

unas 844 especies, de las cuales 50.5% se encuentra en el estado de Tamaulipas.

De los tres estados que conforman el Noreste de México, Tamaulipas contiene la mayor proporción

de cuerpos de agua (Palacio-Prieto et al. 2000). Esto se debe a que en su territorio, que colinda en la parte

oriental con el Golfo de México, llegan escurrimientos de las zonas montañosas que constituyen la Sierra

Madre Oriental, aunado al deficiente drenaje que presentan los suelos de la planicie costera que ha per-

mitido la formación de numerosos humedales, tanto continentales como costeros. De hecho, Tamaulipas es

únicamente superado por Tabasco (1.9% de su superficie total), en proporción de superficie de cuerpos de

agua (1.6%). Varios de estos humedales (algunos fuertemente afectados por problemas de contaminación

y eutroficación, Cruickshank y Tamayo 1976), son considerados sitios prioritarios a nivel nacional por la

CONABIO (Arriaga et al. 1998, 2002) e internacional por la Convención Ramsar (Ramsar Bureau 2001),

como es el caso de la Laguna Madre y el sistema lagunario del río Tamesí. La posición estratégica de Ta-

maulipas ha permitido la conformación de una flora compleja, repartida en diferentes regiones y provincias

florísticas que forman parte tanto del Reino Holártico como del Reino Neotropical (Rzedowski 1978). En su

superficie, por ejemplo, se asienta prácticamente todo el territorio de la Provincia Florística de la Planicie

Costera del Noreste (Rzedowski 1978) y diversos autores han ubicado a parte del estado como miembro de

una región biogeográfica particular, la Provincia Biótica o Biogeográfica Tamaulipeca (Dice 1943; CONABIO

1998; Morrone et al. 2002).

En Tamaulipas se han realizado trabajos vegetacionales y florísticos generales (Puig 1968, 1970, 1976;

González-Medrano 1972; Martínez & González-Medrano 1977; Johnston et al. 1989; Briones 1991; Valiente-

Banuet et al. 1995; Hernández et al. 2005) aunque pocos son los estudios que hacen referencia a su flora

acuática vascular. Lot et al. (1993) reconocen para el estado 34 especies de familias estrictamente acuáticas;

Martínez & Novelo (1993), al estudiar la vegetación de los cuerpos de agua de Tamaulipas, registran 167

especies y Mora-Olivo & Novelo (2005) mencionan la existencia de 175 especies de ambientes acuáticos en

la Reserva de la Biosfera El Cielo. En conjunto, dichos trabajos registran 278 especies de plantas asociadas a

humedales. Trabajo de campo llevado a cabo en los últimos años ha revelado la existencia de un número muy

superior al reconocido hasta ahora, por lo que se considera necesario presentar un inventario actualizado y

lo más completo posible de esta riqueza vegetal.

Dado que muy probablemente la mayor riqueza de flora acuática del noreste de México se concentre en

Tamaulipas, como una primera fase del estudio de las plantas propias de ambientes acuáticos de esta región,

se desarrolló el presente trabajo, cuyo objetivo es proporcionar una lista actualizada de las especies de plantas

vasculares acuáticas de este estado. Un objetivo adicional es la evaluación de su distribución geográfica,

con el fin de determinar la importancia que los humedales de Tamaulipas tienen en la conservación de la

riqueza vegetal especializada a vivir en ambientes acuáticos.

Mora-Olivo y Villasenor, Flora acuática de Tamaulipas, México 513

22°0'N 28°0'N 34°0'N

1 1 1 1

16?0'N

LI y y LI LI LI y L LI

119°0"W 113°0'W 107*0"W 101°O"W 95°0'W 89°0'W

Fic. 1 | ls ee dal + P M Avi I D ^l: M H

C= Coahuila, NL = Nuevo León, T = Tamaulipas.

MATERIALES Y MÉTODOS

Área de estudio

Tamaulipas tiene una superficie de 78,380 km? y se localiza entre los paralelos 22? 12' 31" y 27? 40' 42" de

latitud Norte y los meridianos 97? 08' 38" y 100? 08' 52" de longitud Este. Colinda al norte con el estado

de Texas de los Estados Unidos, al sur con los estados de Veracruz y San Luis Potosí, al este con el Golfo de

México y al oeste con el estado de Nuevo León (Fig. 1). La mayor parte del territorio tamaulipeco se eleva

poco sobre el nivel del mar, con excepción de la Sierra Madre Oriental, que en su parte más alta alcanza

los 3,000 m. Por su localización geográfica y orografía, Tamaulipas presenta una diversidad de climas, que

va desde los sub-húmedos y húmedos, con lluvias en verano en la zona sur-sureste, hasta los templados en

el Altiplano Tamaulipeco y serranías, que varían de húmedos a secos según la altitud (INEGI 2001). Se ha

reportado para el estado la presencia de manglares, tulares y carrizales (Miranda & Hernández 1963), además

de la vegetación característica de ríos, presas, canales, lagunas y charcos (Martínez & Novelo 1993).

Concepto de planta acuática

Aunque se han propuesto muchas definiciones y clasificaciones de plantas acuáticas (Raunkiaer 1934; Weaver

& Clements 1938; Muenscher 1944; Cook et al. 1974; Cook 1990; Daubenmire 1979; Novelo & Gallegos

1988; Lot et al. 1993), para efectos de este estudio, las especies consideradas como hidrófitas o plantas

acuáticas son aquellas que se desarrollan en agua o sobre un substrato que está al menos periódicamente

anaerobio debido al exceso de agua (Tiner 1991). Dado que las condiciones de humedad pueden variar y

las plantas que viven en estos ambientes húmedos tienen diferentes adaptaciones, se dividió a las hidrófitas

f4L,D o ID L

514 Journal of t titute of Texas 1(1)

en tres categorías: acuáticas estrictas, subacuáticas y tolerantes (Lot et al. 1993). Sólo que en este caso, cada

categoría representa la zona que ocupan las plantas en un cuerpo de agua, en una adaptación a la clasificación

de humedales de Tiner (1991) (Fig. 2). Es posible que, dentro del concepto de planta acuática utilizado en

este trabajo, se incluyan algunas especies que para otros autores no sean acuáticas, especialmente en el caso

de las tolerantes, que pueden incluir plantas que soportan el disturbio, frecuentemente como malezas. Las

especies también se clasificaron por su forma biológica (hierba, arbusto o árbol) y siguiendo a Dalton &

Novelo (1983) y a Sculthorpe (1985), por su forma de vida (enraizada emergente, enraizada de hojas flotantes,

enraizada de tallos postrados, enraizada sumergida, libre flotante y libre sumergida).

Métods

El catálogo de especies es el resultado de un intenso trabajo de campo, realizado en la mayor cantidad de

ambientes acuáticos presentes en el estado, como ríos, lagunas, presas, canales, cuerpos de agua temporales

y áreas con suelos saturados. El primer autor ha lectado alrededor de 4,000 números de plantas acuáticas

en los últimos veinte años, material que está depositado en los herbarios UAT, MEXU, ENCB, XAL y TEX

(abreviaciones de acuerdo a Holmgren et al. 1990). Adicionalmente se revisaron ejemplares depositados en

los herbarios mencionados. La información obtenida con estas actividades se complementó con la consulta

a bases de datos públicas (como la REMIB y W3TROPICOS) y personales (Novelo, inédito) y la revisión de

literatura especializada, como son revisiones y monografías taxonómicas, además de estudios florísticos.

Entre los trabajos nacionales que se consultaron destacan los de Potamogeton (González 1989), Cyperus

(Tucker 1994), Utricularia (Olvera 1996), Podostemaceae (Novelo & Philbrick 1997) y Marsileaceae (Pérez-

García et al. 1999). El arreglo del catálogo se hizo siguiendo las clasificaciones utilizadas por Mickel &

Smith (2004) para helechos y plantas afines, por Brummitt (1992) para las gimnospermas, y las propuestas

por Dahlgren et al. (1985) para monocotiledóneas y por Cronquist (1981) para dicotiledóneas. De los taxa

se obtuvo información sobre su distribución a nivel mundial, para lo cual se utilizó la misma bibliografía

y bases de datos mencionadas.

RESULTADOS

Diversidad

Un total de 426 especies (con 48 taxa infraespecíficos), pertenecientes a 213 géneros y 85 familias de plantas

vasculares se registraron en los humedales de Tamaulipas (Tabla 1, Anexo). La riqueza de monocotiledóneas

y dicotiledóneas más o menos está balanceada, siendo las segundas un poco más diversas (49.396), con

54 familias, 128 géneros y 210 especies. Las monocotiledóneas comprenden 46.596, con 23 familias, 76

géneros y 198 especies. Los helechos y gimnospermas acuáticas son raras, representando menos del 596 de

la riqueza.

Las diez familias con mayor riqueza en el estado (Tabla 2) incluyen en conjunto 43.296 de los géneros y

54.596 de las especies. De ellas, solamente Alismataceae es una familia con miembros estri te acuáticos.

Otras familias estrictamente acuáticas que se encuentran en México están bien representadas en el estado de

Tamaulipas. Por ejemplo, en su territorio se registran todas las especies mexicanas conocidas de Cymodocea-

ceae (2), Najadaceae (3), Zannichelliaceae (1), Nelumbonaceae (1), Salviniaceae (2) y Taxodiaceae (1). Familias

que tienen la mitad o más de sus especies en el estado, son Equisetaceae (6696, 2 especies), Potamogetonaceae

(6096, 6), Hydrocharitaceae (57%, 4), Cabombaceae (50%, 1), Ceratophyllaceae (5096, 1) Menyanthaceae (50%,

1), Nymphaeaceae (5096, 5), Parkeriaceae (5096, 1), Ruppiaceae (5096, 1) y Typhaceae (5096, 1).

La mayoría de las plantas acuáticas registradas son herbáceas (377, 88.596), un porcentaje menor son

arbustos (26, 6.196) o árboles (23, 5.496). Como acuáticas estrictas se reconocen 97 especies (22.896), 186

(43.796) son subacuáticas y 143 (33.596) son tolerantes (Tabla 3). La mayoría de las plantas herbáceas son

subacuáticas (4496), siguiendo en orden decreciente las tolerantes (31.696) y, en menor cantidad, las acuáticas

estrictas (24.496). Entre los arbustos no se registran acuáticas estrictas; se observa en cambio una repartición

homogénea entre las subacuáticas y las tolerantes. Finalmente, sólo cinco especies de árboles se consideran

hidrófitas estrictas, una gimnosperma (Taxodium mucronatum), una monocotiledónea (Acoelorraphe wrightii)

Mora-Olivo y Villasenor, Flora acuática de Tamaulipas, México 515

las plantas

Tabla de agua estacional

Tabla de agua permanente

Alto nivel de agua

Bajo nivel de agua

o 77 |

Zona Inundada Zona Permanentemente Zona Periódicamente Zona Sin Saturación

Agua Profunda Agua Somera Periódicamente Saturada Saturada

Acuáticas m -—

: Subacuáticas Tolerantes No Acuáticas

Estrictas

( PLANTAS

PLANTAS ACUATICAS TERRESTRES

Fic. 2. Esquema que ilustra los tres tipos de hidrófitas consideradas en este trabajo, de acuerdo con su nivel de inundación y saturación (Basado en

Tiner 1991).

y tres dicotiledóneas (Rhizophora mangle, Laguncularia racemosa y Annona glabra), las otras especies son sub-

acuáticas (9) o tolerantes (9).

La forma de vida dominante en la flora acuática de Tamaulipas es la de enraizadas emergentes (376

especies, 88.3%), siendo las dicotiledóneas las más frecuentes (Tabla 4). Un pequefio numero de especies se

registró como enraizadas de hojas flotantes (10, 2.396), enraizadas de tallos postrados (3, 0.796), enraizadas

sumergidas (23, 5.4%), libres flotantes (11, 2.696) y libres sumergidas (3, 0.796). En el Anexo a cada taxón

se le indica su forma de vida, su forma biológica y si es acuática estricta, subacuática o tolerante.

Distribución

La mayoría de las especies de hidrófitas que se registran en el estado de Tamaulipas presenta una amplia

distribución geográfica (Tabla 5). Más del 6096 de las especies se distribuyen tanto a lo largo de todo o casi

todo el continente Americano, así como en el Viejo Mundo. Las especies con distribución neotropical (México

a Centro y Sudamérica) ocupan el tercer lugar; de ellas, el elemento sudamer icano (México hasta Sudamérica)

supera ligeramente (8.996) al elemento mesoamericano (México a Centroamérica, 5.296). El elemento neártico

(especies distribuidas de México a Norteamérica) es ligeramente menor (12.9 96) que el neotropical, aunque

su influencia en la flora vascular acuática de la región es significativa. Un 9.396 de las especies registra una

distribución desde Norteamérica (principalmente el sur de los estados Unidos) a Centroamérica; el 396 de

tales especies alcanza además islas del archipiélago de la región del Mar Caribe.

El elemento Mexicano (especies endémicas de México) es notablemente escaso (2.196), representado

únicamente por 9 especies; de ellas, una restringe su distribución al estado de Tamaulipas (Carex fructus). Si se

ampliara el concepto de endemicidad y se incluyera hacia el norte de México las regiones vecinas del suroeste

de los Estados Unidos y hacia el sureste porciones de Belice y Guatemala, el número de especies de distribución

restringida aumentaría en 18 especies más (6.396). Entre las especies que exceden ligeramente los límites políti-

cos al norte de México se pueden citar a Eleocharis brachycarpa, conocida solo de Tamaulipas y Texas o Baccharis

neglecta, Helenium elegans var. amphibolum, Justicia runyonii o Marsilea macropoda, que se localizan también en

Texas y en porciones de los estados mexicanos de Chihuahua, Durango o Hidalgo Otras especies se distribuyen

hacia el sur, hasta Guatemala y Belice, como Cyperus megalanthus, Helenium quadridentatum o Pluchea salicifolia

En los humedales de Tamaulipas predominan las especies subacuáticas y las tolerantes (Tabla 6); en

516 Journal of the Botanical R titute of Texas 1(1)

TABLA 1. Riqueza de la flora vascular acuática de Tamaulipas por grupos taxonómicos.

Grupo Familias Géneros Especies

Helechos y plantas afines 7 8 (3.7%) 17 (4.0 96)

Gimnospermas 1 1 (0.5%) 1 (0.2%)

Dicotiledóneas 54 128 (60.1%) 210 (49.3%)

Monocotiledóneas 25 76:1553-700) 198 (46.5%)

Total 85 213 (100%) 426 (10096)

TABLA 2. Las 10 familias con mayor nümero de especies en la flora vascular acuática de Tamaulipas.

Familias Géneros Especies

Cyperaceae 13 (6.1%) 80 (18.896)

Poaceae 25 (11.796) 56 (13.196)

Asteraceae 23 (10.8%) 33 (7.796)

Polygonaceae 2 (0.996) 14 (3.396)

Scrophulariaceae 8 (3.796) 11 (2.6%)

Lythraceae 6 (2.896) 10 (2.396)

Fabaceae 5 (2.3%) 8 (1.9%)

Acanthaceae 4 (1.9%) 7 (1.6%)

Alismataceae 2 (0.9%) 7 (1.6%)

Mimosaceae 4 (1.9 96) 6 (1.496)

| 92 (43.2%) 232 (54.5%)

TABLA 3. Distribución de la flora vascular acuática de Tamaulipas por grupo taxonómico, tipo de planta acuática y forma bi-

ológica. H = Hierbas, R = Arbustos, B = Árboles.

Acuáticas Estrictas Subacuáticas Tolerantes

Grupo H B H R B H R

Helechos y plantas afines 9 0 0 2 0 B e 0 0

Gimnospermas 0 0 | 0 0 O0 0 c. 0

Monocotiledóneas 54 0 0 94 2 2 45 1 0

Dicotiledóneas 29 0 4 70 9 7 68 14 9

Tota 92 0 5 166 11 9 119 15 9

TABLA 4. Distribución de la flora vascular acuática de Tamaulipas por grupo taxonómico y forma de vida. EE = Enraizada emer-

gente, EH = Enraizada de hojas flotantes, ET = Enraizada de tallos postrados, ES = Enraizada sumergida, LF = Libre flotante,

LS z Libre sumergida.

Grupo EE EH ET ES LF LS

Helechos y plantas afines 10 3 0 0 4 0

Gimnospermas ] 0 0 0 0 0

Monocotiledóneas 175 0 1 16 6 0

Dicotiledóneas 190 7 2 7 1 3

Total 376 10 3 23 11 3

Mora-Olivo y Villasenor, Flora acuática de Tamaulipas, México 517

TABLA 5. Distribución geográfica de las especies de hidrófitas vasculares presentes en Tamaulipas.

Región Especies 96

México (endémicas) 9 2]

México y Norteamérica 55 129

México y Centroamérica 22 5.2

México a Sudamérica 38 8.9

Norteamérica, México y Caribe 18 30

Norteamérica a Centroamérica 27 63

Norteamérica a Sudamérica 181 30.8

México y Viejo Mundo 181 30.8

Total 426 100.0

TABLA 6. Distribución geográfica de las especies de hidrófitas vasculares presentes en Tamaulipas de acuerdo con su a su

grado de tolerancia al agua.

Región Acuáticas Estrictas Subacuáticas Tolerantes

México (endémicas) 2 4 3

México y Norteamérica y 18 30

México y Centroamérica 2 12 8

México a Sudamérica 5 16 i

Norteamérica, México y Caribe 2 9 2

Norteamérica a Centroamérica 6 14 7

Norteamérica a Sudamérica 33 58 40

México y Viejo Mundo 40 55 36

Total 97 186 143

conjunto constituyen 77.296 de su flora. El otro 22.896 lo conforman las acuáticas estrictas. En las tres

categorías se observa una predominancia de especies de amplia distribución (América y Viejo Mundo),

siguiendo en importancia el elemento neotropical. Destaca también entre las acuáticas estrictas el impor-

tante nümero de especies neárticas (7), mesoamericanas (6) y neotropicales (5), lo que pone de relieve el

papel biogeografico de los humedales del estado como zona de transición entre las regiones biogeográficas

templadas y tropicales.

A] evaluar la distribución geográfica de las especies por formas de vida, se observa un amplio predo-

minio de las enraizadas emergentes en todas las divisiones geográficas utilizadas (Tabla 7); nuevamente,

las especies de amplia distribución muestran el mayor espectro de formas de vida. Con una distribución

más restringida solamente se determinaron 9 especies (2.196), ocho enraizadas emergentes y una enraizada

sumergida (Tabla 7).

En Tamaulipas se encuentra el límite meridional de 13 especies (o taxa infraespecíficos) neárticas y el

límite septentrional de 39 neotropicales o paleotropicales. Los taxa norteamericanos que encuentran su límite

sur de distribución en Tamaulipas son Ambrosia trifida, Cyperus acuminatus, Cyperus erythrorhizos, Cyperus

floribundus, Eleocharis coloradoensis, Fimbristylis puberula, Funastrum cynanchoides, Justicia runyonii, Lythrum

californicum, Nuphar advena subsp. advena, Phyla lanceolata, Polygonum hydropiperoides var. opelousanum y

Ulmus crassifolia. Los taxa neotropicales que encuentran su límite norte de distribución en Tamaulipas son

Acmella oppositifolia var. oppositifolia, Aeschynomenne scabra, Crinum erubescens, Cuphea hyssopifolia, Cyperus

humilis, C. laxus, C. lundellii, C. manimae var. manimae, C. megalanthus, C. tenuis, Equisetum myriochaetum,

Fleischmannia arguta, Fuirena camptotricha, Habenaria pringlei, Habenaria quinqueseta, Helenium mexicanum,

Hymenachne amplexicaulis, Hypoxis decumbens, Juncus ebracteatus, Lythrum gracile, Nymphaea amazonum, Pas-

palum arundinaceum, Phyla dulcis, Pithecellobium lanceolatum, Polygonum acuminatum, Polygonum ferrugineum,

518 Journal of the Botanical R h Institute of Texas 1(1)

TABLA 7. Distribución geográfica de la flora vascular acuática de Tamaulipas por forma de vida. EE = Enraizada emergente, EH

= Enraizada de hojas flotantes, ET = Enraizada de tallos postrados, ES = Enraizada sumergida, LF = Libre flotante, LS = Libre

sumergida.

Grupo EE EH ET ES LF LS

México (endémicas) 8 0 0 1 0 0

México y Norteamérica 51 4 0 0 0 0

México y Centroamérica 2] 0 0 1 0 0

México a Sudamérica 36 j 0 0 1 0

Norteamérica, México y Caribe 11 0 0 2 0 0

Norteamérica a Centroamérica 24 0 0 3 0 0

Norteamérica a Sudamérica 114 4 1 8 4 0

México y Viejo Mundo Al 1 2 8 6 3

Total 376 10 3 23 11 3

Polygonum persicarioides, Ruellia paniculata, Rhynchospora contracta, Salvinia auriculata, Solanum diphyllum,

Spermacoce confusa y Steinchisma laxa, mientras que los taxa paleotropicales son Cyperus tenuis, Eleocharis

mutata, Fimbristylis complanata, Hyptis capitata, Neptunia natans y Nymphoides indica.

Entre las hidrófitas de los humedales de Tamaulipas se han identificado 34 especies no nativas (intro-

ducidas). Algunas de ellas se comportan como malezas acuaticas agresivas, causando serios problemas en

presas, canales de riego y otros cuerpos de agua (Novelo & Martinez 1989; Mora 1997). Entre ellas se pueden

citar a Eichhornia crassipes (lirio acuático), nativa de Sudamérica y actualmente distribuida en las regiones

tropicales y subtropicales del mundo e Hydrilla verticillata, hidrófita enraizada sumergida originaria de Asia

y ahora ocupando amplias regiones templadas y tropicales del planeta (Langeland 1996).

DISCUSIÓN

De los tres estados que conforman la región noreste de México, Tamaulipas es el que cuenta con la mayor

riqueza de hidrófitas con 426 especies registradas hasta la fecha. Tal cifra constituye en este momento 5796 de

toda la riqueza vascular acuática reportada por Lot et al. (1999) para México, lo que indudablemente refleja la

necesidad de incrementar el estudio de la flora vascular acuática de todo el país. Su flora es representativa de

los humedales de esta región biogeográfica de México y es un reservorio potencial de biodiversidad para ser

considerado en futuras acciones de conservación de este importante habitat mundialmente amenazado.

Tamaulipas ocupa el segundo lugar a nivel nacional, después del estado de Tabasco, por la superficie

de humedales en su territorio, tanto naturales como artificiales (Palacio-Prieto et al. 2000). Por su extensión

y posición geográfica, algunos humedales del estado han sido considerados a nivel nacional e internacional

como áreas prioritarias para la conservación (Arriaga et al. 1998, 2002; Ramsar Bureau 2001); hoy se puede

justificar su importancia de acuerdo con su biodiversidad de hidrófitas presentes. Estimaciones recientes

(Mora € Villaseñor, resultados no publicados) indican que por su riqueza de hidrófitas, Tamaulipas ocupa

el quinto lugar nacional (solamente superado por Chiapas, Oaxaca, Veracruz y Tabasco); sin embargo,

también a nivel nacional, el estado ocupa el lugar catorce (de un total de 32) por su riqueza florística total

(Villasefior 2003, datos no publicados). Con la excepción de Tabasco (lugar 21), los otros tres estados son

los que ocupan en México la mayor riqueza total a nivel estatal.

Rzedowski (1991b) estimó que 396 de la flora vascular de México está asociada a humedales. Datos

preliminares (Mora & Villasefior, resultados no publicados) muestran que la proporción es mayor, estimando

un valor a nivel nacional de 5.496 y una media por estados de 14.196 (desviación estándar 5.296). Para Tam-

aulipas se ha determinado hasta la fecha que alrededor del 1596 de su flora está asociada con humedales,

cifra cercana a la media estatal. Para Coahuila y Nuevo León, los otros dos estados que constituyen la región

noreste de México, se ha estimado que su flora asociada a humedales es menor a 996. Los altos valores de

riqueza que presenta Tamaulipas con respecto a Nuevo León y Coahuila se pueden explicar, porque además

Mora-Olivo y Villasenor, Flora acuática de Tamaulipas, México 519

de su mayor cantidad de cuerpos de agua continentales, el estado cuenta con amplios ambientes costeros, los

cuales incluyen poco más del 1496 (62 especies) de plantas halófitas características de humedales salobres,

como los mangles, pastos marinos y otras hidrófitas.

Un alto porcentaje de especies asociadas a humedales son cosmopolitas o pantropicales, por lo que no

es sorprendente advertir la amplia distribución geográfica de la mayoría de las hidrófitas encontradas en

Tamaulipas. Aunque poco se ha discutido acerca de los factores históricos que determinaron la distribu-

ción actual de muchas hidrófitas, es evidente que un gran porcentaje de ellas deben su actual distribución

a factores antropogénicos (Stuckey 1993; Sawada et al. 2003) o a la dispersión a larga distancia por aves

(Figuerola & Green 2002; Green et al. 2002). Será interesante abordar en un futuro preguntas encaminadas

a explicar los patrones de distribución de las plantas asociadas a humedales de Tamaulipas, especialmente

con un enfoque histórico.

A nivel de país, en otros estudios se ha señalado el bajo nivel de endemismo de las plantas propias de

humedales, como ha sido el caso de Nueva Zelanda (McGlone et al. 2001) y México (Rzedowski 1991b).

Para el caso particular de Tamaulipas, aquí se dan a conocer valores igualmente bajos en sus ambientes

acuáticos (0.296 de endemismo, una especie), el cual se incrementa a 2.196 si se considera al elemento en-

démico de México. En contraste, altos niveles de endemismo vegetal se han registrado en algunas regiones

de Tamaulipas, como son sus zonas montañosas y sus zonas áridas (Rzedowski 1991b; Hernández et al.

2005; Martínez-Ávalos & Jurado 2005). Sin embargo, hasta la fecha ningün otro estado de México reporta

un mayor nümero de endemismos acuáticos; de acuerdo con Lot et al. (1999), solamente Jalisco y Nayarit

tienen una especie acuática estricta endémica dentro de sus límites políticos, Oserya longifolia y Echinodorus

virgatus respectivamente.

La riqueza y endemismo de la flora acuática tamaulipeca se hace más relevante al considerar que en su

territorio se registra un buen nümero de especies que encuentran su distribución marginal en el estado. En

Tamaulipas encuentran su límite de distribución boreal o meridional poco más del 1096 de las hidrófitas

registradas; esto indica que sus humedales constituyen una zona de transición importante para las hidró-

fitas de las regiones templadas y tropicales. El estudio de las hidrófitas de la región apoya las propuestas

de considerar a la región noreste de México como una zona biogeográfica de particular interés, claramente

diferenciada de otras regiones del país, como lo han hecho Dice (1943), Rzedowski (1978), CONABIO (1998)

o Morrone et al. (2002). Continuar con el estudio de las plantas asociadas a los humedales de Coahuila y

Nuevo León permitirá conocer mejor la flora vascular acuática que se desarrolla en esta interesante región del

país conocida como el noreste de México y ayudará a proponer mejores estrategias para su conservación.

ANEXO

CATÁLOGO DE LAS PLANTAS VASCULARES ACUÁTICAS DEL ESTADO DE TAMAULIPAS

A= Acuática estricta, S= Subacuática, T= Tolerante. H = Hierba, R = Arbusto, B = Árbol. EE = Enraizada

emergente, EH = Enraizada de hojas flotantes, ET = Enraizada de tallos postrados, ES = Enraizada sumergida,

LF = Libre flotante, LS = Libre sumergida. ET = Endémica de Tamaulipas, EM = Endémica de México, NA =

México y Norteamérica, MC = México y Centroamérica, MS = México a Sudamérica, NM = Norteamérica,

México y Caribe, NS = Norteamérica a Sudamérica, VM = Viejo Mundo.

* = [ntroducidas

Helechos y plantas afines

Adiantaceae Equisetaceae

Adiantum capillus-veneris L. T/H/EE/VM Equisetum hyemale L. var. affine (Engelm.) A.A. Eaton S/H/

Acrostichum aureum L. A/H/EE/VM EE/NC

Acrostichum danaeifolium Langsd. & Fisch. A/H/EE/VM Equisetum myriochaetum Schltdl. & Cham. S/H/EE/MS

Azollaceae Marsileaceae

Azolla microphylla Kaulf. A/H/LF/NS ee ancylopoda A. Braun A/H/EH/NS

rsilea macropoda A. B A/H/EH/NA

hows vestita Hook. & ae subsp. vestita A/H/EH/NA

520

Parkeriaceae

Ceratopteris thalictroides (L.) Brongn. A/H/LF/VM

Salviniaceae

Salvinia auriculata Aubl. A/H/LF/MS

Salvinia minima Baker A/H/LF/NS

Thelypteridaceae

Thelypteris kunthii (Desv.) Morton T/H/EE/NS

Thelypteris ovata R.P. St. John var. lindheimeri (C. Chr.) A.R. Sm.

T/H/EE/NC

Thelypteris pilosa (M. Martens & Galeotti) Crawford T/H/EE/

Thelypteris puberula (Baker) C.V. Morton var. puberula T/H/

EE/MC

Thelypteris tetragona (Sw.) Small T/H/EE/NS

GIMNOSPERMAS

Taxodiaceae

Taxodium mucronatum Ten. A/R/EE/NA

ANGIOSPERMAS

MONOCOTILEDÓNEAS

Alismat

Echi Mo . (Spreng.) Fassett. A/H/EE/NS

Echinodorus cordifolius (L.) Griseb. subsp. cordifolius A/H/

EE/NS

Echinodorus grandiflorus (Cham. €: Schltdl.) Micheli subsp.

aureus (Fassett) R.R. Haynes & Holm-Niels. A/H/EE/MS

Sagittaria lancifolia L. subsp. media (Micheli) Bogin A/H/

EE/NS

Sagittaria latifolia Willd. A/H/EE/NS

Sagittaria longiloba Engelm. ex J.G. Sm. A/H/EE/NC

Sagittaria platyphylla (Engelm.) J.G. 5m. A/H/EE/NC

Amaryllidaceae

Crinum erubescens Sol. A/H/EE/MS

Hymenocallis littoralis (Jacq) Salisb. A/H/EE/VM

Zephyranthes pulchella J.G. Sm. A/H/EE/NA

Araceae

Pistia stratiotes L A/H/LS/VM

Xanthosoma robustum Schott A/H/EE/NS

Arecaceae

Acoelorraphe wrightii (Griseb. & H. Wendl.) H. Wendl. ex Becc.

A/B/EE/NS

Cannaceae

Canna glauca L. A/H/EE/NS

Commelinaceae

Callisia micrantha (Torr.) D.R. Hunt S/H/EE/NA

Commelina communis L. T/H/EE/NA

Commelina diffusa Burm. f. S/H/EE/VM

*Murdannia nudiflora (L.) Brenan T/H/EE/VM

Cymodoceaceae

Halodule wrightii Asch. A/H/ES/NS

Syringodium filiforme Kutz. A/H/ES/NS

Cyperaceae

Bolboschoenus robustus (Pursh) Soják A/H/EE/NS

fal, Dat M ID hi Pr

Journal of

of Texas 1(1)

Bulbostylis capillaris (L.) C.B. Clarke T/H/EE/NS

Bulbostylis juncoides (Vahl) Kük. ex Osten T/H/EE/NS

Carex fructus Reznicek T/H/EE/ET

Cladium jamaicense Crantz A/H/EE/NS

Cyperus acuminatus Torrey & Hooker S/H/EE/NA

Cyperus aggregatus (Willd.) Endl. T/H/EE/NS

Cyperus articulatus L. A/H/EE/VM

Cyperus haspan L. S/H/EE/VM

Cyperus canus C. Presl. S/H/EE/NS

Cyperus compressus L. S/H/EE/VM

Cyperus digitatus Roxb. subsp. digitatus S/H/EE/VM

Cyperus elegans L. S/H/EE/NS

Cyperus flavicomus Michx. S/H/EE/VM

Cyperus entrerianus Boeckeler S/H/EE/NS

Cyperus erythrorhizos Muhl. S/H/EE/NA

*Cyperus esculentus L. T/H/EE/VM

Cyperus floribundus (Kük.) R.N. Carter & S.D. Jones T/H/EE/NA

Cyperus hermaphroditus (Jacq.) Standl. T/H/EE/NS

Cyperus humilis Kunth S/H/EE/MS

*Cyperus involucratus Rottb. S/H/EE/VM

*Cyperus iria L. S/H/EE/VM

Cyperus laevigatus L. S/H/EE/VM

Cyperus lanceolatus Poir. S/H/EE/VM

Cyperus laxus Lam. S/H/EE/VM

Cyperus ligularis L. T/H/EE/VM

Cyperus lundellii O'Neill S/H/EE/MS

Cyperus manimae Kunth var. asperrimus (Liebm.) Kük. T/H/

EE/NS

Cyperus manimae Kunth var. manimae (Liebm.) Kük. T/H/

EE/NS

Cyperus megalanthus (Kük.) G.C. Tucker S/H/EE/MC

Cyperus niger Ruíz López & Pavón S/H/EE/VM

Cyperus ochraceus Vahl S/H/EE/NS

Cyperus odoratus L. S/H/EE/VM

*Cyperus oxylepis Steud. S/H/EE/NS

Cyperus polystachyos Rottb. S/H/EE/VM

Cyperus prolixus Kunth S/H/EE/VM

*Cyperus rotundus L. T/H/EE/VM

Cyperus squarrosus L. S/H/EE/VM

Cyperus surinamensis Rottb. S/H/EE/NS

Cyperus tenuis Swallen S/H/EE/VM

Cyperus virens Michx. S/H/EE/NS

Eleocharis acicularis (L.) Roem. €: Schult. A/H/EE/VM

Eleocharis albida Torr. A/H/EE/NA

Eleocharis atropurpurea (Retz.) J. Presl & C. Presl S/H/EE/VM

Eleocharis brachycarpa Svens. A/H/EE/NA

Eleocharis cellulosa Torr. A/H/EE/NS

Eleocharis coloradoensis (Britton) Gilly A/H/EE/NA

Eleocharis flavescens (Poir.) Urban A/H/EE/NS

Eleocharis geniculata (L.) Roem. & Schult. S/H/EE/VM

Eleocharis interstincta (Vahl) Roem. & Schult. A/H/EE/NS

Eleocharis macrostachya Britt. A/H/EE/NS

Fleocharis minima Kunth A/H/EE/NS

Eleocharis montevidensis Kunth S/H/EE/NS

Eleocharis mutata (L.) Roem. & Schult. A/H/EE/VM

Eleocharis parvula (Roem. & Schult.) Link ex Bluff, Nees &

Schauer A/H/EE/VM

Eleocharis radicans (A. Dietr.) Kunth A/H/EE/NS

Mora-Olivo y Villasenor, Flora acuática de Tamaulipas, México

Eleocharis rostellata (Torr.) Torr. A/H/EE/NS

Fimbristylis annua (All) Roem. & Schult. S/H/EE/VM

Fimbristylis caroliniana (Lam.) Fernald. S/H/EE/NM

Fimbristylis castanea (Michx.) Vahl S/H/EE/NC

Fimbristylis complanata (Retz) Link A/H/EE/VM

Fimbristylis cymosa (Lam .) R. Br. S/H/EE/VM

Fimbristylis dichotoma (L.) Vahl S/H/EE/VM

Fimbristylis puberula (Michx.) Vahl S/H/EE/NA

Fimbristylis spadicea (L.) Vahl S/H/EE/MS

Fimbristylis db (Lam ) Link S/H/EE/NS

Fuirena cat icha C. Wright S/H/EE/MC

Fuirena Sd Vahl S/H/EE/NS

Killinga brevifolia Rottb. S/H/EE/VM

Killinga odorata Vahl S/H/EE/VM

Kilinga pumila Michaux S/H/EE/VM

Lipocarpha micrantha (Vahl) G. Tucker S/H/EE/VM

Oxycarium cubense (Poepp. & Kunth) Lye A/H/EE/VM

Rhynchospora colorata (L.) H. Pfeiffer T/H/EE/NS

Rhynchospora contracta (Nees) Raynal S/H/EE/NS

Schoenoplectus americanus (Pers.) Volkart ex Schinz & R. Keller

H/EE/NS

Schoenoplectus californicus (C.A. Mey.) Soják A/H/EE/NS

Schoenoplectus erectus (Poir.) Palla ex J. Raynal subsp. raynalii

(Schuyler) Lye A/H/EE/VM

Schoenoplectus pungens (Vahl) Palla A/H/EE/VM

Schoenoplectus saximontanus (Fernald) J. Raynal A/H/EE/NA

Schoenoplectus tabernaemontani (C.C. Gmel.) Palla A/H/

Hydrocharitaceae

MODI engelmannii Asch. A/H/ES/NC

Hydrilla verticillata (L.£) Royle A/H/ES/VM

Thalassia testudinum Banks ex Kónig A/H/ES/NS

Vallisneria americana Michx. A/H/ES/VM

Hypoxidaceae

Hypoxis decumbens L. var. decumbens T/H/EE/MS

Iridaceae

Cipura campanulata Ravenna S/H/EE/MS

Cipura paludosa Aublet S/H/EE/MS

Sisyrinchium angustifolium Mill. T/H/EE/NA

Sisyrinchium biforme E.P. Bicknell T/H/EE/NA

Juncaceae

nos arcticus Willd. var. mexicanus (Willd.) Baslev S/H/EE/

Juncus dichotomus Elliott S/H/EE/NS

Juncus ebracteatus E. Mey S/H/EE/MS

Juncus nodosus L. S/H/EE/NA

Lemnaceae

Lemna

Lemna gibba L. A/H/LF/VM

Spirodela polyrhiza (L ) Schleid. A/H/LF

tialis Welw. A/H/LF/VM

——

Wolffiella lingulata (Hegelm.) Hegelm. A/H/LF/NS

Marantacea

eae

Thalia geniculata L. A/H/EE/VM

521

Najadaceae

Najas guadalupensis (Spreng.) Magnus var. guadalupensis

A/H/ES/NS

Najas marina L. A/H/ES/VM

Najas wrightiana A. Braun A/H/ES/NC

Orchidaceae

Bletia purpurea (Lam.) DC. T/H/EE/NS

Habenaria pringlei B.L. Rob. S/H/EE/MC

Habenaria quinqueseta (Michx.) Sw. T/H/EE/MC

Habenaria repens Nutt. A/H/EE/NS

Poaceae

Andropogon glomeratus (Walter) Britton, Sterns & Poggenb.

S/H/EE/NS

Arundinella berteroniana (Schultes) A. Hitch. & Chase T/H/

EE/MS

*Arundo donax L. T/R/EE/VM

Distichlis spicata (L) Greene var. spicata S/H/EE/NS

*Echinochloa colona (L) Link S/H/EE/VM

*Echinochloa crus-galli (L.) P. Beauv. S/H/EE/VM

Echinochloa crus-pavonis (Kunth) Schult. S/H/EE/VM

Echinochloa polystac hya (Kunth) Hitchc. S/H/EE/VM

*Echinochloa pyramidalis (Lam.) Hitchc. & Chase S/H/EE/VM

Eragrostis cilianensis (All.) Vignolo ex Janch. T/H/EE/VM

Eragrostis hypnoides (Lam.) Britton, Sterns & Poggenb.

SEE

Eragrostis reptans (Michx.) Nees T/H/EE/NA

Eragrostis secundiflora C. Presl. T/H/EE/NS

Eriochloa acuminata (J. Presl) Kunth S/H/EE/NA

Eriochloa punctata (L.) Desv. S/H/EE/NC

Guadua angustifolia Kunth subsp. angustifolia T/B/EE/MS

*Hemarthria altissima (Poir.) Stapf € C.E. Hubb. T/H/EE/VM

Hymenachne amplexicaulis (Rudge) Nees A/H/EE/VM

Leersia hexandra Sw. A/H/EE/VM

Leptochloa fusca (L.) Kunth subsp. fascicularis (Lam.) N.W.

Snow S/H/EE/NS

Leptochloa fusca (L.) Kunth subsp. uninervia (Presl.) Hitchc. &

Chase S/H/EE/NS

Leptochloa panicea (Retz.) Ohwi subps. brachiata (Steud.) N

Snow T/H/EE/NS

Leptochloa nealleyi Vasey S/H/EE/NM

Lithachne pauciflora (Sw.) P. Beauv T/H/EE/MS

Monanthochloé littoralis Engelm. S/H/EE/NM

Panicum hirsutum Sw. T/H/EE/NS

Panicum trichoides Sw. T/H/EE/NS

Panicum virgatum L. S/H/EE/NS

Paspalidium geminatum (Forssk.) Stapf A/H/EE/VM

Paspalum linaceum Poir. S/H/EE/MS

Paspalum conjugatum PJ. Bergius T/H/EE/VM

Paspalum conspersum Schrad. S/H/EE/NS

Paspalum denticulatum Trin. S/H/EE/NS

Paspalum distichum L. A/H/EE/NS

Paspalum harwegianum Fourn. T/H/EE/NC

Paspalum langei (Fourn.) Nash T/H/EE/NS

Paspalum monostachyum Chase T/H/EE/NA

Paspalum pubiflorum Rupr. T/H/EE/NC

Paspalum setaceum Michx. var. setaceum S/H/EE/NS

Paspalum squamulatum Fourn. S/H/EE/MC

522

*Paspalum urvillei Steud. S/H/EE/VM

Paspalum virgatum L. S/H/EE/NS

Paspalum vaginatum Sw. S/H/EE/VM

*Pennisetum purpureum Schum. T/H/EE/VM

Phragmites australis (Cav.) Trin. subsp. australis S/R/EE/VM

*Polypogon monspeliensis (L.) Desf. T/H/EE/VM

*Polypogon viridis (Gouan) Breistr. S/H/EE/VM

Setaria magna Griseb. S/H/EE/NS

Setaria parviflora (Poir.) Kerguélen T/H/EE/VM

Spartina patens (Aiton) Muhl. S/H/EE/NC

Spartina spartinae (Trin.) A. Hitchc. S/H/EE/NS

Sporobolus airoides (Torr.) Torr. T/H/EE/NA

Sporobolus pyramidatus (Lam.) Hitchc. T/H/EE/NS

Sporobolus virginicus (L.) Kunth S/H/EE/VM

Sporobolus wrightii Munro ex Scribn. T/H/EE/NA

Steinchisma laxa (Sw.) Zuloaga S/H/EE/MS

*Urochloa mutica (Forssk.) T.O. Nguyen S/H/EE/NS

Pontederiaceae

*Fichhornia crassipes (C. Martius) Solms-Laub. A/H/LF/VM

Heteranthera dubia Jacq.) MacMill. A/H/ES/NS

Heteranthera limosa (Sw.) Willd. A/H/EE/NS

Heteranthera mexicana S. Watson A/H/EE/NA

Heteranthera rotundifolia (Kunth) Griseb. A/H/EE/NS

Potamogetonaceae

Potamogeton foliosus Raf. subsp. foliosus A/H/ES/NS

Pot illinoensis Morong A/H/ESNS

Potamogeton nodosus Poiret A/H/EH/VM

Potamogeton pusillus L. var. pusillus A/H/ES/VM

Stuckenia pectinata (L) Bórner A/H/ES/VM

Stuckenia striata (Ruiz & Pav.) Holub A/H/ES/NS

Ruppiac

Tee maritima L. A/H/ES/VM

Typhaceae

Typha domingensis Pers. A/H/EE/VM

Xyridaceae

Xyris ambigua Beyr. ex Kunth S/H/EE/NC

Xyris jupicai Rich. S/H/EE/NS

Zannichelliacea

Zannichellia oct L. A/H/ES/VM

DICOTILEDONEAS

Acanthac

Blechum DU (Lam.) Urb. T/H/EE/NS

Dicliptera sexangularis (L) Juss. T/H/EE/MS

Justicia runyonii Small T/H/EE/NA

Ruellia coerulea Morong T/H/EE/NS

Ruellia inundata Kunth T/H/EE/MS

Ruellia malacosperma Greenm. T/H/EE/MS

Ruellia paniculata L. T/H/EE/MS

Aizoaceae

Sesuvium maritimum (Walter) Britton, Sterns & Poggenb.

/MC

T/H/EE/M

Sesuvium portulacastrum L. T/H/EE/VM

Sesuvium verrucosum Raf. T/H/EE/NA

Trianthema portulacastrum L. T/H/EE/VM

fal, Dat A ID hi Pr

Journal of

of Texas 1(1)

Amaranthaceae

Alternanthera obovata (M. Martens & Galeotti) Standl. S/H/

EE/MC

Amaranthus australis (Gray) Sauer SH/EE/NC

Blutaparon vermiculare (L.) Mears T/H/EE/NS

Tidestromia lanuginosa (Nutt.) Standl. subsp. lanuginosa

T/H/EE/NM

Annonaceae

Annona glabra L. A/B/EE/VM

Apiaceae

*Centella erecta (L. f.) Fernald S/H/EE/NS

Eryngium nasturtiifolium Juss ex Delar f. S/H/EE/NC

Hydrocotyle bonariensis Lamarck S/H/EE/NS

Hydrocotyle mexicana Schltdl. & Cham. T/H/EE/MC

Hydrocotyle umbellata L. S/H/EE/NS

Hydrocotyle verticillata Thunb. var. triradiata (A. Rich.) Fernald

S/H/EE/VM

Hydrocotyle verticillata Thunb. var. verticillata S/H/EE/VM

Apocynaceae

biflora Jacq.) Müll. Arg. T/H/EE/MS

Asclepiadaceae

*Cryptostegia grandiflora Roxb. ex R. Br. T/H/EE/VM

*Cryptostegia madagascariensis Bojer ex Decne. T/H/EE/VM

Funastrum clausum (Jacq.) Schltr. S/H/EE/NS

Funastrum cynanchoides (Dcne.) Schltr. var. cynanchoides

S/H/EE/NA

Asteraceae

Acmella oppositifolia (Lam.) R.K. Jansen var. oppositifolia

Ambrosia trifida L. T/H/EE/NA

Aster subulatus Michx. var. subulatus SH/EE/NS

Baccharis neglecta Britton & A. Brown T/R/EE/NA

Baccharis salicifolia (Ruiz & Pav.) Pers. S/R/EE/NS

Barkleyanthus salicifolius (Kunth) H. Rob. & Brettell T/R/EE/

Borrichia frutescens (L.) DC. S/H/EE/NA

Chloracantha spinosa (Benth.) G.L. Nesom S/H/EE/NC

Conoclinium betonicifolium (Mill) R.M. King €: H. Rob. S/H/

Coreopsis tinctoria Nutt. T/H/EE/NS

Eclipta prostrata (L.) L. S/H/EE/VM

Egletes liebmanni Sch. Bip. S/H/EE/MC

Egletes viscosa Less. S/H/EE/NC

Flaveria trinervia (Spreng.) C. Mohr T/H/EE/VM

Fleischmannia arguta (Kunth) B.L. Rob. T/H/EE/MC

Heischmannia porphyranthema (A. Gray) R.M. King & H. Rob.

T/H/EE/EM

Gymnocoronis latifolia Hook & Arn. A/H/EE/MC

Helenium elegans DC. var. amphibolum (A. Gray) Bierner

S/H/EE/EM

Helenium mexicanum Kunth S/H/EE/MC

Helenium microcephalum DC. var. microcephalum S/H/EE/

Helenium microcephalum DC. var. ooclinium (A. Gray) Bierner

Helenium quadridentatum Labill. S/H/EE/MC

Mora-Olivo y Villasenor, Flora acuática de Tamaulipas, México

Jaegeria hirta (Lag.) Less. T/H/EE/MS

Laennecia coulteri (A. Gray) G.L. Nesom T/H/EE/NA

Melanthera nivea (L.) Small T/H/EE/NS

Mikania cordifolia (L. f) Willd. S/H/EE/NS

Mikania micrantha Kunth S/H/EE/NS

Mikania scandens (L.) Willd. S/H/EE/NS

Packera tampicana (DC.) C. Jeffrey T/H/EE/NA

Pluchea carolinenesis (Jacq.) G. Don T/R/EE/VM

Pluchea odorata (L.) Cass. S/H/EE/NS

Pluchea salicifolia (Mill.) S.F. Blake S/H/EE/MC

Solidago velutina DC. T/H/EE/NA

Trichocoronis wrightii (Torr. & A. Gray) Gray var. wrightii

A/H/EE/NA

Bataceae

Batis maritima L. S/H/EE/VM

Bignoniaceae

Chilopsis linearis (Cav.) Sweet T/H/EE/NA

Boraginaceae

Heliotropium curassavicum L. var. curassavicum T/H/EE/NS

Brassicaceae

Cakile geniculata (Robins.) Millsp. T/H/EE/NA

Cakile lanceolata (Willd.) O.E. Schulz subsp. pseudoconstricta

Rodman T/H/EE/NC

*Cardamine hirsuta L. T/H/EE/VM

*Nasturtium officinale R. Br. A/H/ET/VM

Rorippa teres (Michx.) Stuckey S/H/EE/NC

Cabombaceae

Cabomba paleaformis Fassett A/H/ES/MC

Capparidacea

Crateva tapia L. z B/EE/MS

Ceratophyllaceae

Ceratophyllum demersum L. A/H/LS/VM

Chenopodiaceae

vii Torr. S/H/EE/NC

Salicornia virginica E S/H/EE/VM

Suaeda conferta (Small) |. M. Johnston S/H/EE/NM

Suaeda linearis (Elliott) Moq. S/H/EE/NM

Suaeda nigra (Raf) J.F. Macbr. S/H/EE/NM

Suaeda tampicencis (Standl.) l. M. Johnst. S/H/EE/NM

Coli E

3d cornia OIgel

Chrysobalanaceae

Chrysobalanus icaco L. S/R/EE/VM

Combretaceae

Conocarpus erectus L. T/R/EE/VM

Laguncularia racemosa (L.) C.F. Gaertn. A/B/EE/VM

Convolvulaceae

Ipomoea carnea Jacq. subsp. fistulosa (Mart. ex Choisy) D.

Austin T/H/EE/MS

Ipomoea rubens Choisy T/H/EE/VM

Elatinaceae

Bergia texana (Hook.) Seub. ex Walp. S/H/EE/NA

Euphorbiaceae

Caperonia castaneifolia (L.) A. St.-Hill. S/H/EE/MS

Caperonia palustris (L.) A. St.-Hil. S/H/EE/VM

523

Fabaceae

Aeschynomene indica L. S/H/EE/VM

Aeschynomene rudis Benth. S/H/EE/MS

Aeschynomene scabra G. Don S/H/EE/MS

Dalbergia brownei Jacq.) Urban S/R/EE/NS

Desmodium triflorum (L) DC. T/H/EE/VM

Sesbania drummondii (Rydb.) Cory T/R/EE/NA

Sesbania herbacea (Mill. McVaugh S/H/EE/NS

Vigna luteola (Jacq.) Benth. S/H/EE/VM

Gentianaceae

Centaurium calycosum (Buckley) Fernald T/H/EE/NA

Eustoma exaltatum (L.) Salisb. subsp. exaltatum S/H/EE/NC

Eustoma exaltatum (L.) Salisb. subsp. russellianum (Hook.)

rtez S/H/EE/NA

DN aL (Kunth) G. Don T/H/EE/EM

Sabat j enm. T/H/EE/NA

nean stellaris Pursh S/H/EE/NM

Haloragaceae

*Myriophyllum aquaticum (Vell.) Verdc. A/H/ES/VM

Myriophyllum hippuroides Nutt. ex Torr & A. Gray A/H/ES/NC

Hydrophyllaceae

Hydrolea spinosa L. A/H/EE/NS

Juglandaceae

Carya illinoinensis (Wangenh.) K. Koch T/B/EE/NC

Carya myristiciformis (F. Michx.) Nutt. T/B/EE/NA

Carya palmeri W.E. Manning T/B/EE/EM

Lamiaceae

Clinopodium brownei (Sw.) Kuntze S/H/EE/NS

Hyptis capitata Jacq. T/H/EE/VM

Hyptis verticillata Jacq. T/H/EE/NS

pee pel atlace ae

Utricularia foliosa L. A/H/LS/VM

Utricularia gibba L. A/H/LS/VM

Lobeliaceae

Lobelia cardinalis L. subsp. cardinalis S/H/EE/NS

Lobelia purpusii Brand. A/H/EE/EM

Loganiaceae

Mitreola petiolata (J.F. Gmel) Torr. & A. Gray S/H/EE/NS

Lythraceae

Ammannia auriculata Willd. S/H/EE//M

Ammannia coccinea Rottb. S/H/EE/VM

Ammannia robusta Heer & Regel S/H/EE/NS

Cuphea hyssopifolia Kunth T/H/EE/NS

Heimia salicifolia Link S/R/EE/NS

Lythrum alatum Pursh. var. lanceolatum (Elliott) Rothr. S/H/

EE/NM

Lythrum alatum Pursh. var. linearifolium A. Gray S/H/EE/EM

Lytrum californicum Torr. & A. Gray S/H/EE/NA

Lythrum gracile Benth. S/H/EE/MC

Nesaea palmeri S.A. Graham S/H/EE/EM

Rotala ramosior (L.) Koehne A/H/EE/VM

Malvaceae

Kosteletzkya depressa (L) OJ. Blanch., Fryxell & D.M. Bates

T/H/EE/MS

524

Malachra alceifolia Jacq. T/H/EE/MS

Malachra capitata (L.) L. T/H/EE/MS

Menyanthaceae

Nymphoides indica (L) Kuntze A/H/EH/VM

Mimosa

Inga vera "Wild S/B/EE/NS

Mimosa pigra L. S/R/EE/VM

Neptunia natans (L. f.) Druce A/H/LF/VM

Neptunia plena (L.) Benth. S/H/EE/VM

Neptunia pubescens Benth. var. pubescens S/H/EE/NS

Pithecellobium lanceolatum (Humb. & Bonpl.) Benth. T/B/

EE/MS

Molluginaceae

Glinus radiatus (Ruiz & Pav.) Rohrb. T/H/EE/NS

*Mollugo verticillata L. T/H/EE/VM

Moraceae

Ficus insipida Willd. S/B/EE/MS

Nelumbonaceae

Nelumbo lutea (Willd.) Pers. A/H/EH/NS

Nymphae

Nuphar b E W.T. Aiton subsp. advena A/H/EH/NA

Nymphaea amazonum Mart. & Zucc. subsp. amazonum

A/H/EH/MS

ibas ampla (Salisb.) DC. A/H/EH/NS

phaea elegans Hook. A/H/EH/NA

nO Planchon A/H/EH/NS

Oleaceae

Fraxinus berlandieriana DC. S/B/EE/NA

Onagra

ms eee (Nutt.) H. Hara S/H/EE/VM

Ludwigia octovalvis (Jacq.) PH. Raven S/H/EE/VM

udwigia palustris (L.) Elliott A/H/ES/NM

Ludwigia peploides (Kunth) PH. Raven A/H/ET/NS

Ludwigia repens J.R. Forst. A/H/ES/NM

Plantaginaceae

*Plantago major L. T/H/EE/VM

Platanaceae

Platanus rzedowskii K, Nixon & Poole S/B/EE/EM

Podostemaceae

Oserya coulteriana Tul. A/H/ES/EM

Tristichia trifaria (Bory ex Willd.) Spreng. A/H/ES/VM

Polygonaceae

Polygonum acuminatum Kunth A/H/EEMS

Polygonum ferrugineum Wedd. S/H/EE/MS

Polygonum glabrum Willd. S/H/EE/VM

Polygonum hydropiperoides Michx. var. hydropiperoides

S/H/EE/NS

Polygonum hydropiperoides Michx. var. opelosanum (Riddell

ex Small) J.S. Wilson S/H/EE/NS

*Polygonum lapathifolium Willd. S/H/EE/VM

Polygonum pensylvanicum L. S/H/EE/VM

*Polygonum persicaria L. S/H/EE/VM

Polygonum persicarioides Kunth S/H/EE/MS

fal, Dat A ID hi Pr

Journal of

of Texas 1(1)

Polygonum punctatum Ell. S/H/EE/NS

Polygonum segetum Kunth S/H/EE/MS

Rumex chrysocarpus Moric. S/H/EE/NA

*Rumex crispus L. T/H/EE/VM

*Rumex obtusifolius L. S/H/EE

*Rumex pulcher L. subsp. pulcher S/H/EE/VM

Primulaceae

Anagallis minima (L.) E.H.L. Krause S/H/EE/VM

Samolus ebracteatus Kunth var. ebracteatus T/H/EE/MC

Samolus parviflorus Raf. S/H/EE/NS

Rhizophoracea

Rhizophora Ut L. A/B/EE/NS

Rubiaceae

Cephalanthus occidentalis L. S/R/EE/NC

Cephalanthus salicifolius Bonpl. S/R/EE/MC

*Pentodon pentandrus (Schumach. & Thonn.) Vatke T/H/EE/

Spermacoce confusa Rendle T/H/EE/MS

Spermacoce glabra Michx. T/H/EE/NS

Spermacoce tenuior L. T/H/EE/NS

Salicaceae

Populus j Wesmael var. mexicana S/B/EE/EM

Populus tremuloides Michx. T/H/EE/NA

Salix thurberi Nutt. S/R/EE/NA

Salix humboldtiana Willd. S/B/EE/NC

Salix nigra Marshall S/B/EE/NA

Salix taxifolia Kunth S/R/EE/NC

Scrophulariaceae

Bacopa monnieri (L.) Wettst. A/H/EE/VM

Calceolaria mexicana Benth. T/H/EE/MS

Capraria biflora L. T/H/EE/VM

Capraria mexicana Moric. ex Benth. T/H/EE/MC

Lindernia dubia (L.) Pennell S/H/EE/NS

Mecardonia procumbens (Mill.) Small T/H/EE/VM

Mecardonia vandellioides (Kunth) Pennell S/H/EE/NS

Mimulus glabratus Kunth var. glabratus S/H/EE/NS

modia durantifolia (L.) Sw. S/H/EE/NS

media schottii Holz. S/H/EE/NA

Veronica peregrina L. S/H/EE/NS

Solanaceae

Calibrachoa parviflora (Juss.) D'Arcy T/H/EE/NS

Lycium carolinianum Walter var. quadrifidum (Dunal) Hitchc.

lanum hiense L. T/H/EE/NS

Solanum diphyllum L. T/H/EE/MC

Solanum tampicense Dunal A/H/EE/NC

Tamaricaceae

*Tamarix gallica L. T/R/EE/VM

*Tamarix chinensis Lour. T/R/EE/VM

Ulmaceae

Celtis laevigata Willd. T/B/EE/NA

Ulmus crassifloia Nutt. T/B/EE/NA

Urticaceae

Boehmeria cylindrica (L.) Sw. T/H/EE/NS

Mora-Olivo y Villasenor, Flora acuática de Tamaulipas, México 525

Verbenaceae Phyla lanceolata (Michx.) Greene S/H/EE/NS

Avicennia germinans (L) L. S/B/EE/VM Phyla nodiflora (L. Greene T/H/EE/VM

Clerodendrum ligustrinum (Jacq.) R. Br. S/R/EE/MC Phyla stoechadifolia (L.) Small S/H/EE/NS/NS

Lippia alba (Mill) N.E. Br. T/H/EE/VM Phyla strigulosa (M. Martens & Galeotti) Moldenke T/H/EE/

Phyla dulcis (Trevir) Moldenke T/H/EE/VM NS

AGRADECIMIENTOS

El primer autor agradece al Programa de Mejoramiento del Profesorado (PROMEP) de la Universidad Autónoma

de Tamaulipas por el apoyo para realizar estudios de posgrado en la Universidad Nacional Autónoma de

México. Enrique Ortiz y Abril Angeles gentilmente elaboraron las figuras. Se agradece a Hilda Flores, Mahinda

Martínez y Fernando Chiang la revisión crítica del manuscrito y sus atinados tarios. El doctor Alejandro

Novelo Retana (1951-2006) colaboró estrechamente en el desarrollo de este trabajo; desafortunadamente

su muerte prematura nos impidió seguir contando con su amplia experiencia en la flora acuática de México.

Esta contribución se dedica cordialmente a su memoria.

REFERENCIAS

ABELLÁN, P, D. SANCHEZ-FERNANDEZ, J. VELASCO y A. MILLAN. 2005. Conservation of freshwater biodiversity: a comparison

of different area selection methods. Biodivers. & Conservation 14:3457-3474.

AMEZAGA, J.M., L. SANTAMARÍA y A.J. GREEN. 2002. Biotic wetland connectivity-supporting a new approach for wetland

policy. Acta Oecol. 23:213-222.

ARRIAGA, L., E. VAZQUEZ, J. GONZALEZ, R. Jiménez, E. Muñoz y V. AGUILAR. (coords.). 1998. Regiones prioritarias marinas de

México. Comisión Nacional para el Conocimiento y Uso de la Biodiversidad, México, D.F.

ARRIAGA, L., V. AGUILAR y J. ALCOCER. 2002. Aguas continentales y diversidad biológica de México. Comisión Nacional

para el Conocimiento y Uso de la Biodiversidad, México, D.F.

BoNiLLA-BARBOSA, J. 2004. Flora acuática vascular. En: Luna l., J.J. Morrone y D. Espinosa (eds... Biodiversidad de la

Sierra Madre Oriental. Las Prensas de Ciencias, México, D.F. Pp. 149-159.

BoniLLA-BARBOSA, J.R., J.A. VIANA-LASES y F. SALAZAR-VILLEGAS. 2000. Listados Florísticos de México. XX. Flora acuática de

Morelos. Instituto de Biología, Universidad Nacional Autónoma de México, México, D.F.

Briones V., O.L. 1991. Sobre la flora, vegetación y fitogeografia de la Sierra de San Carlos, Tamaulipas. Acta Bot.

Mex. 16:15-43.

BRuMMrrT, R.K. 1992. Vascular plant families and genera. Royal Botanic Gardens, Kew, Great Britain.

Conasio. 1998. La diversidad biológica de México: estudio de País, 1998. Comisión Nacional para el Conocimiento

y Uso de la Biodiversidad. México, D.F.

Cook, C.D.K. 1990. Aquatic plant book. SPB Academic Publishing, The Hague, Netherlands.

Cook, C.D.K., BJ. Gur, E.M. Rix , J. SCHNELLER y M. Seitz. 1974. Water plants of the world. A manual for the identification

of the genera of freshwater macrophytes. Dr. W. Junk b.v., Publishers, The Hague, Netherlands.

Cronauist, A. 1981. An integrated system of classification of flowering plants. Columbia University Press, New

York.

CRUICKSHANK, G.G. y L. Tamayo. 1976. Atlas del agua de la Republica Mexicana. Secretaría de Recursos Hidráulicos,

México, D.F. P. 247.

DAHLGREN, R.T.M., H.T. Curronp y PF. Yeo. 1985. The families of monocotyledons. Structure, evolution, and taxonomy.

Springer-Verlag, New York.

DALTON, P. y A. Novelo. 1983. Aquatic and wetland plants of the Arnold Arboretum. Arnoldia 43(2):7-44.

DAUBENMIRE, R.F. 1979. Ecología vegetal. Tratado de autoecologia de las plantas. Tercera edición. Limusa, México,

D.F.

Dice, L.R. 1943. The biotic provinces of North America. University of Michigan Press, Ann Arbor.

FIGUEROLA, J. y A. GREEN. 2002. Dispersal of aquatic organisms by water-birds: a review of past research and priorities

for future studies. Freshw. Biol. 47:483-494.

526 Journal of the Botanical R h Institute of Texas 1(1)

FITZSIMONS, J.A. y H.A. ROBERTSON. 2005. Freshwater reserves in Australia: directions and challenges for the develop-

ment of a comprehensive system of protected areas. Hydrobiologia 552:87-97.

GoNzÁALEZ, M. 1989. El género Potamogeton (Potamogetonaceae) en México. Acta Bot. Mex. 6:1-43.

GONZALEZ-Meprano, F. 1972. La vegetación del nordeste de México. Anales Inst. Biol. UNAM, Ser. Bot. 43:11-50.

GREEN, A.J., J. FIGUEROLA y M.I. SANCHEZ. 2002. Implications of waterbird ecology for the dispersal of aquatic organ-

isms. Acta Oecol. 23:177-189.

HERNÁNDEZ, L.G., J. Treviño C., A. Mona-Ouvo y M. Martinez. D. 2005. Diversidad florística y endemismos. En: Sánchez-

Ramos, G., P. Reyes-Castillo & R. Dirzo (eds.). Historia natural de la Reserva de la Biósfera El Cielo, Tamaulipas,

México.Universidad Autónoma de Tamaulipas, Cd. Victoria. Pp. 244-253.

HOLMGREN, P, N.H. HOLMGREN y L.C. Barnett. 1990. Index herbariorum. Part |: The herbaria of the world. 8th edition.

New York Botanical Garden, Bronx.

INEGI. 2001. Síntesis geográfica del estado de Tamaulipas. Secretaría de Programación y Presupuesto, México,

Bas

JA, S.K. 1990. Conservation of aquatic plants. In: Gopal, B., ed. Ecology and management of aquatic vegetation

in the Indian Subcontinent; 237-241. Kluwer Academic Publishers, Dordrecht.

JoHNsTON, M.C., K. Nixon, G. Nesom y M. Martinez. 1989. Listado de plantas vasculares conocidas de la Sierra de

Guatemala, Gómez Farías, Tamaulipas, México. Biotam 1(2):21-33.

LANGELAND, K.A. 1996. Hydrilla verticillata (L.F) Royle (Hydrocharitaceae), The perfect aquatic weed. Castanea

61:293-304.

Lor, A., A. NoveLo y P. RAMfREz-GaRCÍA. 1986. Listados florísticos de México V. Angiospermas acuáticas mexicanas 1.

Instituto de Biología. Universidad Nacional Autónoma de México, México, D.F.

Lor, A., A. NoveLo y P. Ramirez-Garcia. 1993. Diversity of Mexican aquatic vascular plant flora. In: Ramamoorthy, T.P,

R. Bye, A. Lot y J. Fa (eds) Biological diversity of Mexico: 577—591. Oxford University Press, New York.

Lor, A., A. Novelo, M. Otvera y P. RAMfREZ-GARCÍA. 1999, Catálogo de angiospermas acuáticas de México. Hidrófitas

estrictas emergentes, sumergidas y flotantes. Cuadernos 33. Instituto de Biología. Universidad Nacional

Autónoma de México, México, D.F.

Martinez-Avatos, J.G. y E. JuraDO. 2005. Geographic distribution and conservation of Cactaceae from Tamaulipas,

México. Biodivers. & Conservation 14:2483-2506.

Martinez y O., E. y F. GONZALEZ-Meprano. 1977. La vegetación del sudeste de Tamaulipas, México. Biótica 2(2):

1-45.

Martínez, M. y A. Novelo. 1993. La vegetación acuática del Estado de Tamaulipas, México. Anales Inst. Biol. UNAM,

Ser. Bot. 64:59-86.

McGtone, M.S., R.P. DUNCAN y PB. Heenan. 2001. Endemism, species selection and the origin and distribution of the

vascular plant flora of New Zealand. J. Biog. 28:199-216.

Mickel, J.T. y A.R. SMTH. 2004. The pteridophytes of Mexico. Mem. New York Bot. Gard. 88:1-1054.

MIRANDA, F. y E. HERNANDEZ X. 1963. Los tipos de vegetación de México y su clasificación. Bol. Soc. Bot. Méx.

28:29-179.

Mora O., A. 1997. Bases para el control y aprovechamiento de las malezas acuáticas en canales de riego de

Tamaulipas, México. Tesis de Maestría, Unidad Académica Multidisciplinaria Agronomía y Ciencias, Universidad

Autónoma de Tamaulipas, Cd. Victoria.

Mora-Outvo, A. y A. Novelo. 2005. La vegetación acuática y semiacuática. En: Sánchez-Ramos G, P Reyes-Castillo &

R Dirzo (eds.) Historia natural de la Reserva de la Biósfera El Cielo, Tamaulipas, México. Universidad Autónoma

de Tamaulipas, Cd. Victoria, Tamaulipas, México. Pp. 106-115.

Morrone, J.J., D. ESPINOSA O. y J. LLorENTE B. 2002. Mexican biogeographic provinces: preliminary scheme, general

characterizations, and synonymies. Acta Zool. Mex. (n.s.) 85:83-108.

MutNscHER, W.C. 1944. Aquatic plants of the United States. Comstock Publishing Co., Inc. Ithaca, N.Y.

NoveLo, A. y M. Gallegos M. 1988. Estudio de la flora y la vegetación acuática relacionada con el sistema de chi-

nampas en el sureste del Valle de México. Biótica 3 (1-2):121-139.

Mora-Olivo y Villasenor, Flora acuática de Tamaulipas, México 527

NoveLo R., A. v C.T. PuiLgnick. 1997. Taxonomy of Mexican Podostemaceae. Aquatic Bot. 57:275-303.

Novelo, A. y M. Martinez. 1989. Hydrilla verticillata (Hydrocharitaceae): problemática maleza acuática de reciente

introducción en México. Anales Inst. Biol.. UNAM, Ser. Bot. (Número ünico):97-102.

Orvera, M. 1996. El género Utricularia (Lentibulariaceae) en México. Anales Inst. Biol.. UNAM, Ser. Bot. 67:

347-384.

PALACIO-PRIETO, J.L., G. Bocco, A. VELASQUEZ, J.F. Mas, F. TAKAKiI-TAKAKI, L. LUNA-GONZALEZ, G. GOMEZ-RODRIGUEZ, J. LOPEZ-GARCIA,

M. PALMA M, |. Trejo V., A. PERALTA H., J. PRADO-MOLINA, A. RODRIGUEZ-AGUILAR, R. MAYORGA-SAUCEDO y F. GONZALEZ M. 2000.

La condición actual de los recursos forestales en México: resultados del Inventario Forestal Nacional 2000.

Investigaciones Geográficas, Bol. Instit. Geograf., UNAM 43:183-203.

Perez-Garcla, B., R. RIBA y D.M. Johnston. 1999. Marsileaceae. Flora Méx. 6(5):1-17.

Pus, H. 1968. Notas acerca de la flora y la vegetación de la Sierra de Tamaulipas, México. Anales Esc. Nac. Cien.

Biol.17:49-123.

Puic, H. 1970. Etude phytogéographyque de la Sierra de Tamaulipas, Mexique. Bull. Soc. d'Hist. Nat. Toul.

106(12):60-78.

Pula, H. 1976. Végétation de la Huasteca, Mexique. Mission Archéologique et Ethnologique Francaise au Mexique.

Collection d' Etudes Mésoamericaines. Vol. 5. México, D.F.

RAMSAR BUREAU. 2001. The Ramsar list of wetlands of international importance. Ramsar Bureau. Gland, Switzerland.

RAUNKIAER, C. 1934. Life forms of plants and statistical plant geography. Clarendon Press, Oxford.

REED, PB., JR. 1988. National list of plant species that occur in wetlands: 1988 national summary. Biol. Rep. 8824),

U.S. Fish and Wildlife Service, Washington, D.C.

Reep, P.B., Jr. (comp.) 1997. Revision of the national list of plant species that occur in wetlands. In cooperation

with the National and Regional Interagency Review Panels. Department of Interior, U.S. Fish and Wildlife

Service, Washington, D.C.

RzeDowskl, J. 1978. La vegetación de México. Limusa, México, D.F.

Rzepowski, J. 1991a. Diversidad y orígenes de la flora fanerogámica de México. Acta Bot. Mex. 15:3-21.

Rzepowskl, J. 1991b. El endemismo en la flora fanerogámica de México: una apreciación analítica preliminar. Acta

Bot. Mex. 15:47—64.

RzEpowskI, J. 1993. Diversity and origins of the phanerogamic flora of Mexico. En: Ramamoorthy, T.P., R. Bye, A. Lot

y J. Fa (eds) Biological diversity of Mexico. Oxford University Press, New York. Pp. 129-144.

SANTAMARÍA, L. y KLAASSEN. 2002. Waterbird-mediated dispersal of aquatic organisms: an introduction. Acta Oecol.

23:115-119.

SAUNDERS, D.L., J.J. Meeuwie y A.C.J. VincenT. 2002. Freshwater protected areas: strategies for conservation. Conserva-

tion Biol. 16:30-41.

SAWADA, M, A.E. Viau y K. GAJEwskI. 2003. The biogeography of aquatic macrophytes in North America since the

last glacial maximum. J. Biog. 30:999-1017.

ScuLTHORPE, C.D. 1985. The biology of aquatic vascular plants. Edward Arnold. Ltd., London.

Siqueiros, M.E. 1989. Contribución al conocimiento de la flora acuática y subacuática de Aguascalientes. Univer-

sidad Autónoma de Aguascalientes, Aguascalientes.

Stuckey, R.L. 1993. Phytogeographical outline of aquatic and wetland angiosperms in continental eastern North

America. Aquatic Bot. 44:259-301.

Tiner, R. 1991. The concept of a hydrophyte for wetland identification. BioScience 41:236-247.

Tucker, G.C. 1994. Revision of the Mexican species of Cyperus (Cyperaceae). Syst. Bot. Monog. 43:1-43.

VALIENTE-BANUET, A., F. GONZALEZ y D. Pinero. 1995. La vegetación selvática de la región de Gómez Farías, Tamaulipas,

México. Acta Bot. Mex. 33:1-36.

VILLASEÑOR, J.L. 1990. The genera of Asteraceae endemic to Mexico and adjacent regions. Aliso 12:685-692.

VILLASEÑOR, J.L. 2003. Diversidad y distribución de las Magnoliophyta de México. Interciencia 28:160-167.

ViLLASENOR, J.L. 2004. Los géneros de plantas vasculares de la flora de México. Bol. Soc. Bot. Méx. 75:105-135.

Weaver, J.E. y F.E. Clements. 1938. Plant ecology. Second edition. McGraw-Hill, New York.

528 Journal of the Botanical R h Institute of Texas 1(1)

BOOK NOTICES

SANDRA PERRIN. 2002. Organic Gardening in Cold Climates (rev. ed.). (ISBN 0-87842-451-2, pbk.). Mountain

Press Publishing Company, PO. Box 2399, Missoula, MT 59806, U.S.A. (Orders: www.mountain-press.

com, info@mtnpress.com, 406-728-1900, 406-728-1635 fax, 800-234-5308). $12.00, 142 pp., b/w

illustrations, 514" x 815".

The author, an organic gardener in Missoula, Montana, says that she wants this book to be a “simple explanation of good organic garden-

ing." The map indicates that the northern half of the U.S.A., Canada, and Alaska are included in the purview of the discussion.— Guy

Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A.

Susan H. MUNGER (Text), CHARLOTTE STUAB Thomas (Illustrations), and VerLYN KLINKENBORG (Foreword). 2003.

Common to this Country: Botanical Discoveries of Lewis and Clark. (ISBN 1-57965-224-7, hbk.).

Artisan, 708 Broadway, New York, NY 10003, U.S.A. (Orders: www.artisanbooks.com, 212-254-5900,

212-677-6692 fax). $22.95, 128 pp., 25 watercolor illustrations, color photos, 7" x 9".

The author gives interesting and quickly readable discussion us history and significance of 25 of the speci llected by Meriwether

Lewis of the Lewis & Clark Expedition: Amelanchier alnifolia, Angeli t l ursi, Camassia quamash, Clarkia pulchella,

KU US tinctoria, Echinacea angustifolia, Erythronium grandiflorum, e triflorum, Velia rediviva, Lilium philadelphicum, Linus lewisii,

pomifera, Mahonia aquifolium, Mi guttat bd cespitosa, Opuntia polyacantha and

Dn a f£. 4 Tb E ae i T * T nn re

> E

other species, ii, Pinus ponderosa,

TOT :T1 E - 1

“This illustrated botanical guide will me to e m buffs and dudes alike."

With the nicely written Du eed and excellent map of the expedition's route, it is a fine, popular introduction to this fascinating

piece of history.— Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A.

W. Date NELSON. 2003. Interpreters with Lewis and Clark: The Story of Sacagawea and Toussaint

Carbonneau. (ISBN 1-57441-165-9, hbk.). University of North Texas Press, PO Box 311336, Denton,

TX 76203-1336, U.S.A. (Orders: www.unt.edu/untpress, 940-565-2142, 940-565-4590 fax, 1-800-

826-8911). $24.95, 184 pp., 22 illustrations, 2 maps, 644" x 944",

The author has written “a well-documented account of the lives of two near-mythical figures in Western history, Sacagawea [Shoshone

Indian] and her husband Toussaint Charbonneau [French Canadian fur trader]. From the Lewis and Clark accounts and from all other

available resources, he has extracted key references in recreating the fascinating life stories of these two intriguing people. ... [He] fol-

lows the couple from their diverse origins through their history-making travels with Lewis and Clark and on through the rest of their

lives. ... In mi the story, the author carefully examines the numerous N E to a readers a balanced account of these

interpreters w ith Lewis and Clark” (comments by Philip J. Roberts

of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A.

Guy Nesom, Botanical Research Institute

Barry G. Hatt. 2008. Phylogenetic Trees Made Easy: A How-to Manual, Third Edition. (ISBN 978-0-

87893-310-5, pbk.). Sinauer Associates, Inc., 23 Plumtree Road, Sunderland, MA 01375-0407, U.S.A.

(Orders: orders@sinauer.com, www.sinauer.com, 413-549-4300, 413-549-1118 fax). $39.95, 233

p., illustrated, 9" x 914”,

Contents.—1) Read Me First! 2) Tutorial: Create a Tree! 3) Acquiring and Aligning the Sequences. INTERLUDE I. Major Methods for

Estimating Phylogenetic Trees. 4) Neighbor-Joining Trees. 5) Drawing Phylogenetic Trees. 6) Parsimony. 7) Maximum Likelihood. 8)

Bayesian Inference of Trees with MrBayes. INTERLUDE II. Which Method Should You Use? 9) Reconstructing Ancestral Sequences.

10) Detecting Adaptive Evolution. 11) Working with Various Computer Platforms. Appendices, Literature Cited, Index.

J. Bot. Res. Inst. Texas 1(1): 528. 2007

THE CACIA EAE OF THE NATURAL MUNICIPAL EARIGOR PRAIN Bla.

RIO DE JANEIRO, BRAZIL: TAXONOMY AND CONSERVATION!

Alice de Moraes Calvente Regina Helena Potsch Andreata

Universidade Santa Ursula Universidade Santa Ursula

Laboratorio de Angiosp ICBA Laboratório de Angiosr |

Rua Fernando Ferrari 75 Rua Fernando Ferrari 75

Botafogo, Rio de Janeiro Botafogo, Rio de Janeiro

RJ, BRAZIL, CEP 2223 1-040 RJ, BRAZIL, CEP 22231-040

alicecalventeayahoo.com

*Corresponding author

ABSTRACT

This paper contains the floristic inventory for the Cactaceae occurring in the Natural Municipal Park of Prainha, located at 23? 02' 41"S

43? 30' 40"W, in the Rio de Janeiro Municipality, Rio de Janeiro State, Brazil. The area is a threatened native Atlantic Forest remnant

with no floristic inventory. Ten Cactaceae genera occur in the Park, which are Pereskia Mill. (1 sp.), Brasiliopuntia (K. Schum.) A. Berger

(1 sp.), Opuntia Mill. (1 sp.), Epiphyllum Haw. (1 sp), Hylocereus (A. Berger) Britton & Rose (1 spJ, Lepismium Pfeiff. (1 sp), Rhipsalis

Gaertn. (5 spp.), Cereus Mill. (1 spJ, Coleocephalocereus Backeb. (1 sp.) and Pilosocereus Byles & Rowley (1 sp.). For the 14 taxa UD in

the survey we p t morphology, taxonomy, ecology and ion that were obtained through field and li

Among the studied species, five are classified under threat categories and 54% are endemic to Brazil. A large floristic similarity was

found between this Park and other coastal areas that have been surveyed in the Rio de Janeiro State.

Key wonps: Brazil, Cactaceae, Atlantic Forest, Conservation Units, Taxonomy

RESUMEN

Este papel contiene el inventario florístico para las Cactaceae del Parque Natural Municipal de Prainha, situado en 23° 02' 41"S 43° 30!

40"W, enel municipio de Río de Janeiro, estado de Rio de Janeiro, Brasil. El área es un amenazado remanente del Bosque Atlántico nativo,

con la ausencia de i ios florísticos. Diez géneros de Cacta urren en el Parque, los cuales, Pereskia Mill. (1 sp.), Brasiliopuntia

(K. Schum.) A. Berger (1 spJ, Opuntia Mill. (1 spJ, Epiphyllum Haw.(1 sp.), Hylocereus (A. Berger) Britton & Rose (1 spJ, Lepismium Pfeiff.

(1 sp.), Rhipsalis Gaertn. (5 spp.), Cereus Mill. (1 spJ, Coleocephalocereus Backeb. (1 sp.) e Pilosocereus Byles & Rowley (1 spJ. Para los 14

taxones encontrados en el examen proporcionamos datos en morfología, taxonomía, ecología y conservación, que fueron obtenidos a

través de las actividades científicas a campo y del estudio de la literatura. Entre las especies estudiadas, cinco presentan algún grado de

amenaza y 5496 son endémicos al Brasil. Una semejanza florística grande fue encontrada entre este parque y otras áreas costeras que se

han examinado ya en el estado de Río de Janeiro.

INTRODUCTION

The Cactaceae family comprises about 100 genera and 1500 species distributed in four subfamilies: Opun-

tioideae, Pereskioideae, Maihuenioideae and Cactoideae (Hunt 1999). The species are restricted to the new

world, with the exception of Rhipsalis baccifera (Mill.) Stearn, and can be recognized for the presence of the

areole with spines and trichomes, succulent photosynthetic stems, absence of conspicuous leafs and flowers

with receptacular epiginy (Barthlott & Hunt 1993). Cacti have a great economic importance and are widely

spread as ornamental plants, but they also have many other uses as for medicine, as food, as cattle fodder,

as fences, as fiber and filling, as furniture and as building material (Anderson 2001).

In Brazil occur 3096 of the Cactaceae genera distributed in three of the four Cactaceae subfamilies:

Opuntioideae, Pereskioideae and Cactoideae (Taylor & Zappi 2004). The species inhabit various vegetation

formations as the Cerrado (Savanna), the Caatinga (Savannic Steppe) and the Atlantic Rain Forest, the latter

being the main vegetation at the Natural Municipal Park of Prainha (Joly et al. 1999).

The Atlantic Forest has an estimated floristic diversity of 20000 species, and 6000 of them are endemic,

1Th; rp + nf +h AAC? Ai tati J I 1I 4t tha D Aa DA A = fiAnri DialAni IDatanirca\ nf tho M Mari lilas APTA Eau ld

Rio de Janeiro.

J. Bot. Res. Inst. Texas 1(1): 529 — 548. 2007

530 Journal of the Botanical R h Institute of Texas 1(1)

however, this vegetation today is restricted to only 696 of the originally occupied area (Mittermeier et al.

2000). At the Rio de Janeiro State the original Atlantic Forest formations have been devastated by an intense

human interference, especially by the coast, where are located the most populated areas in Brazil.

This study provides the floristic inventory for the Cactaceae at the Natural Municipal Park of Prainha

(NMPP), an Atlantic Forest remnant located at the Rio de Janeiro city coast. Morphologic, taxonomic and

ecological data on species and information that can be used to help the conservation of the NMPP and of

the species are provided.

STUDY AREA

The Area named Prainha integrated the rural properties known as Camorim, Vargem Pequena e Vargem

Grande Farms. In 1990 the Area of Environmental Protection (APA) of Prainha was created to save the

natural scenario and to implement an ecological park at the region with the objectives of promoting the

sustainable use through ecological tourism and recreation associated to environmental education activities.

The Natural Municipal Park of Prainha (NMPP), originally named Ecological Municipal Park of Prainha

was created in 1999 and today is under guard of the Municipal Secretariat of Environment—SMAC (Rio de

Janeiro 1999). The park is located at S 23? 02' 41"W 43? 30' 40", at the Rio de Janeiro City, Brazil, between

the Recreio dos Bandeirantes district and the APA of Grumari, and has 126.30 hectares (Fig. 1). In the park,

a short extension beach is delimited by the coastal sides of the Caeté and Boa Vista hills and the Pedra dos

Cabritos rock, building a natural amphitheater with the hillside varying from 0 to 460 m of altitude (Fig.

2).The part of the park located above 100 ms.m. is also included inside the area of the Pedra Branca State

Park (Prefeitura da Cidade do Rio de Janeiro 1998).

The region is occupied by the Brazilian Atlantic Rain Forest (i.e. Dense Ombrophylous Atlantic Forest)

in the hillsides and for the restinga vegetation (sandy coastal plane with pioneer communities) in areas near

the beach (Ministério das Minas e Energia 1983; Joly et al. 1999). The existing forest at the area of study is

mainly secondary in advanced process of natural regeneration with some scattered small more preserved

areas of putative primary origin.

The steep hillsides and the hilltops are frequently dominated by gneiss-granite rock outcrops and those

are colonized by a diverse rupicolous flora composed mostly from Cactaceae, Bromeliaceae and Velloziaceae

taxa. The rock outcrop referred on the bibliography (Prefeitura da Cidade do Rio de Janeiro 1998) and also

in this study as Pedra dos Cabritos (Cabritos Rock) is popularly named Morro da Boa Vista (Boa Vista Hill)

and is the most visited site inside the park (Fig. 2).

METHODS

Botanical material was collected during 17 field excursions to 18 localities at the NMPP from April of 2003

to October of 2004 (Fig. 2). The specimens collected were prepared according with the usual herbarium

techniques (Fidalgo & Bononi 1984) and incorporated in the Santa Ursula University Herbarium (RUSU)

and in the National Museum Herbarium (R). The following herbaria collections of the Rio de Janeiro State

were consulted to obtain data: FCAB, GUA, HD, R, RB, RBR, RFA e RUSU (Holmgren et al. 1990). The ba-

sinonyms were cited when applied and the synonyms were only cited when published recently (after 2000)

or when they are names still being used erroneously in herbaria collections.

The conservation statuses for the Rio de Janeiro State species were obtained in Calvente et al. (2005).

The categories were accessed according with the IUCN 3.1 version (2001). The floristic similarity between

the NMPP and the other areas in the Rio de Janeiro State was analyzed with the software BioDiversity Pro

Version 2/1997 (The Natural History Museum & Scottish Association for Marine Science) that was used for

the production of a presence and absence matrix, the calculation of the Jaccard distance equation and for

the creation of a dendogram using the Group Average.

Calvente and Andreata, Cactaceae of the Natural Municipal Park of Prainha (Brazil)

0 23 5000

531

Lal

Fic. 1. Location of the Natural Municipal Park of Prainha, RJ (IPP, 1999a).

KEY TO CACTACEAE TAXA IN THE NATURAL MUNICIPAL PARK OF PRAINHA

1. Branches with well-developed leafs; pedunculated flowers. (Pereskioideae)

1. Branches leafless or with minute scale-like, persistent or deciduous leafs; sessile flowers.

2. Glochids present. (Opuntioideae - Opuntieae)

3. Tree with di

1. Pereskia aculeata

Ius stems, primary segments cylindrical and S5GCOI \dary g PORE n ES |

segments all yellow; fruit globose

2. Brasiliopuntia brasiliensis

3. Shrub with monomorphic stems, primary and secondary segments flattened; perigonium segments

externally deep red and internally yellow; fruit turbinate 3. Opuntia monacantha

2. Glochids absent. (Cactoideae)

4. Columnar tree or shrub with 4-15 ribs. (Cereeae)

5. Stem segments with 9-15 ribs; fruit dehiscent by a basal pore

. 13. Coleocephalocereus fluminensis

subsp. fluminensis

5. Stem segments with 4-6 ribs; fruit dehiscent by a longitudinal or lateral slit.

6. Flowers 17-23 cm long; fruit dehiscent by a longitudinal slit —

.. ..... 12. Cereus fernambucensis subsp.

ernambucensis

6. Flowers 6-9 cm long; fruit dehiscent by a lateral slit 14. Pilosocereus arrabidae

532 Journal of the Botanical Research Institute of Texas 1(1)

KM y

Pre

F im 2

UE.

HEL 333.9 MAN

fy // hf ye TH D

I— |

MO 1000 |

yl | ] \

pu,

OCEANO

ATLÁNTICO

um M

Fic. 2. Areas visited at the Natural Municipal Park of Prainha, RJ. 1. Pedra dos Cabritos; 2. B las Mirtáceas; 3. Cruzeiro do Sul; 4. Bosque das Brasi-

liopuntias; 5. Afl t tral; 6. Restinga; 7. Afl to Leste. (IPP, 1999b).

4. Non-columnar epiphyte or scandent plants with stems cylindrical, flattened or with 3-5 wings.

7. Flower-tube 10-20 cm long, pericarpel and flower-tube with bract-scales. (Hylocereeae)

8. Stem segments 3-winged, areoles with 3-4 conical spines; flowers infundibuliform, 25-35 cm long

5. Hylocereus setaceus

8. Stem segments flattened, areoles glabrous; flowers hypocrateriform, 19-24 cm long

4. Epiphyllum phyllanthus subsp. phyllanthus

7. Flower-tube less than 0.5 cm or absent, pericarpel and flower-tube glabrous. (Rhipsalideae)

9. Stem segments flattened or 3-5 winged.

10. Branching mesotonic; segment areoles with copious pilose hairs 6. Lepismium cruciforme

10. Branching acrotonic or subacrotonic; segment areoles glabrous or with scarce pilose hairs,

tomentose hairs or bract scales.

. Wings discontinuous in the same stem segment 9. Rhipsalis paradoxa subsp. paradoxa

11. Wings continuous in the same stem segment.

12. Stem segments flattened, rarely 3-wi 1; perigoni tsgold

m=i

—

vvellow,fruitoblong

2 - > un ,

purple 7. Rhipsalis elliptica

12. Stem segments 3-5-winged; perigoni g ts white; fruit depressed-globose white

or pinkish 11. Rhipsalis triangularis

9. Stem segments cylindrical.

13. Stem segments all of determinate growth; flowers 2.5-3 cm diam. 8. Rhipsalis grandiflora

13. Primary stem segments of indeterminate growth and secondary segments of determinate

growth; flowers 0.9-1 cm diam. 10. Rhipsalis teres f. heteroclada

1. Pereskia aculeata Mill., Gard. dict. ed. 8. 1768. (Fig. 3 A-C).

Shrub, scandent, terricolous or rupicolous, glochids absent. Stem brown or purplish, woody, cylindri-

cal, ca. 0.5 cm diam. Leafs alternate, elliptic, rarely obelliptic, 6-9 x 2.5-3 cm, slightly bicolor, purplish

Calvente and Andreata, Cactaceae of the Natural Municipal Park of Prainha (Brazil) 533

Fic. 3. A-C. Pereskia aculeata Mill. A. Branch with sar B. pe ie ÓN of DON C. Fruit. D. Brasiliopuntia brasiliensis (Willd.) A. Berger,

flattened branch with Towers and fruits, E-F. 0 , F. Fruit. G-H. Epiphyllum phyllanthus (L.)Haw. subsp.

phyllanthus; G. Branct fruits, H. Longitudinal section of fl

£s+haD o ID L

534 Journal of t titute of Texas 1(1)

green, venation inconspicuous. Areoles in the axils of leafs; spines 2, geminate, deltoid, retrorse, curve, ca.

2 cm long, sometimes absent. Flower diurnal, solitary or in inflorescences formed by the growing of one

flower at the previous flower pedicel, rotate, 4-6 x 4 cm; pedicel 2-5.5 cm long; pericarpel ca. 0.5 x 0.5

cm. Pericarpel and pedicel areoles with 1 fleshy lanceolate basal bract-scale; hairs tomentose; spines 2-3

fine, acicular, 0.5-1.0 cm long, stiff. Perigonium external segments 5, deltoid, ca 0.2 x 0.2 cm, fleshy, green,

apex acuminate; internal segments 13, obdeltoid to oblong, ca. 1.5 x 0.8-1.0 cm, membranaceous, white or

greenish, apex mucronate or emarginate; filament white, anther golden yellow; ovary semi-inferior, stigma

with 3—5 lobes; nectary a ring around the style base. Fruit globose, ca. 1.0 x 0.8 cm, orange, indehiscent,

with persistent perigonium; fruit areoles with 1 fleshy lanceolate basal bract-scale (deciduous when ripe),

tomentose hairs, (1-23 acicular spines.

Heliophyte species commonly inhabiting the rock outcrops at the NMPP. The stigma frequently can

be anomalous, exhibiting the fusion of two lobes or the short development of one lobe that becomes black

and wilted. Flowers annually from February to April and fruits from April to August. Conservation status:

Least Concern (LO).

Specimens examined: BRAZIL. Rio de Janeiro: Mun. Rio de Janeiro, Parque Natural Municipal da Prainha, 23° 05'S 43° 29'W, elev. 5

ms.m., 20 Jul 1995, Scheinvar 6272 (RB); Pedra dos Cabritos (topo do morro da Boa Vista), ca. 359 ms.m., 10 Apr 1996, Braga & Bovini

3320 (RUSU, RB); ibidem, 12 Apr 2003, Calvente et al. 34 (RUSU); ibidem, 29 Jan 2004, (25 Mar 2004, fl. cult), Calvente & Versieux 85

(RUSU).

2. Brasiliopuntia brasiliensis (Willd.) A. Berger, Entwicklungslin. Kakt. 94. 1926. (Fig. 3 D). Cactus brasiliensis

Willd., Enum. pl. suppl. 33. 1814. Opuntia brasiliensis (Willd.) Haw., Suppl. pl. succ. 79. 1819.

Tree, 3-6 m, terricolous or rupicolous. Trunk woody, fleshy at apex. Areoles borne around the trunk;

spines 6-25, acicular, 1.5-4.0 cm long. Stem segments dimorphic; primary segments cylindrical, 22-40 x

0.8-2.5 cm, fleshy, green; secondary segments flattened, elliptic to oblong, 6-15(-18) x 3-5(—5.5) cm, fleshy,

bicolor, green, sometimes yellowish. Stem segments areoles with short tomentose hairs; glochids numer-

ous, deciduous; spines 0-1(-2), acicular, 1-3 cm long. Young stem segments areoles with 1 minute deltoid

deciduous fleshy bract-scale; spines 1-2, acicular, 0.1-0.5 cm long. Flower diurnal, 0-1 per areole, sessile,

rotate, 2.5-3.5 x 2.5-4.5 cm; pericarpel depressed-obovate, 1.2-1.6 x 0.7-1.4 cm, with areoles similar to

the stem segments areoles but with 1 fleshy minute deltoid basal bract-scale; perigonium segments 15-17,

obelliptic, 1.0-1.6 x 0.5-0.7 cm, spreading, yellow; ovary inferior, stigma with 4-5 lobes. Fruit globose,

ca. 3.5 cm diam., greenish yellow, indehiscent; fruit areoles with copious short tomentose hairs, numerous

glochids; seed 2, 1 cm long, with copious, long, lanate hairs.

At the NMPP this species is sciophyte inside the forested areas where it can reach greater heights but

itis shorter while growing as heliophyte on forest gaps. Many individuals are found growing close together,

if one falls horizontally usually it will produce new shoots that grow vertically. Anomalous styles were fre-

quently observed presenting a protuberance close to their apex, probably caused by the fusion of stamens

parts or the fusion of the anther with style regions during the floral development. The flowers are visited

by bees which are supposed to be the pollinators and by hemipterans which possibly feed perforating the

stems. Brasiliopuntia was previously considered a synonym of Opuntia but today it is considered a separate

genus based on molecular evidences (Wallace & Dickie 2002) and morphological apomorphies such as

the heteromorphic stems (Taylor et al. 2002) and pollen grains with distinct ornamentation (Leuenberger

1976). Flowers annually during October, and usually the flowering is abundant and synchronic among the

specimens. Immature fruits were observed from April to October. The fruits last many months to mature

and were found semimature fallen to the ground during the following year flowering season. Conservation

status: Least Concern (LC).

Specimens examined: BRAZIL. Rio de Janeiro: Mun. Rio de Janeiro, Parque Natural Municipal da Prainha, Morro da Boa Vista, 30

Sep 1996, Braga & Bovini 3519 (RUSU); Bosque das Mirtáceas, 12 Apr 2003, Calvente et al. 33 (RUSU); Bosque das Brasiliopuntias, 17

Oct 2003, Calvente et al. 62 (RUSU); ibidem, 28 Oct 2003, Calvente & Bocayuva 65 (RUSU); ibidem, 10 Oct 2004, Calvente & Versieux

109, 110, 111 (RUSU).

Calvente and Andreata, Cactaceae of the Natural Municipal Park of Prainha (Brazil) 535

3. Opuntia monacantha Haw., Suppl. pl. succ. 81. 1819. (Fig. 3 E-F).

Shrub non columnar, erect, 1-3 m, terricolous. Trunk woody formed by the secondary growth of the basal

stem segments. Areoles borne around the trunk; spines 3-12 acicular, ca 4 cm long, stiff. Stem segments

monomorphic, flattened, obovate to elliptic, (7—)15—46 x (3.5-)8-15 cm, fleshy, green. Stem segments ar-

eoles with 1 minute deciduous fleshy basal bract-scale; hairs tomentose, short; glochids numerous; spines

1-2 acicular, 2-4(—5.5) cm long, stiff. Flower diurnal, 0—2(—3) per areole, sessile, rotate, 6-8 x 5-6 cm,

many grouped in poorly defined floriferous regions; pericarpel turbinate, 4.0—6.0 x 2.0-2.5 cm, with areoles

similar to the stem segments areoles but with only 1 acicular, ca. 0.7 cm long spine; perigonium segments

ca. 29, deltoid to obovate, 1.5—3.0 x 1.0-1.5 cm, spreading, apex mucronate; external segments fleshy,

purplish; internal segments membranaceous, yellow; stamens sensitive; ovary inferior, stigma with 6-7

lobes. Fruit turbinate, ca. 7 x 4 cm, reddish green, indehiscent; fruit areoles with short tomentose hairs,

numerous deciduous glochids.

This is a frequent heliophyte species on the open restinga vegetation near the beach. The population

occurring at the NMPP is not of great size and the number of young specimens around well established

mature individuals suggests that this population is on a recovering stage and that this species may have a

great clonal reproduction capacity. The flowers are visited by bees which are supposed to be the pollina-

tors, by hemipterans and are predated by coleopterans that feed on the stamens after the anthesis. Opuntia

monacantha was found sometimes erroneously misidentified as O. vulgaris Mill. on herbaria collections. This

name is considered today a synonym of O. ficus-indica (L.) Mill. (Taylor et al. 2002). Flowering is annual,

abundant and synchronic among individuals at the NMPP during February and from May to October. Im-

mature fruits occur during almost the whole year and were found ripe in January. Conservation status:

Least Concern (LC).

Specimens examined: BRAZIL. Rio de Janeiro: Mun. Rio de Janeiro, Parque Natural Municipal da Prainha, 27 Feb 1987, Giordano &

Ferreira 230 (RB); vegetacáo de restinga próxima a praia, 06 Jun 2003, Calvente et al. 43 (RUSU); ibidem, 28 Oct 2003, Calvente & Bo-

cayuva 69 (RUSU); ibidem 15 May 2004, Calvente & Versieux 92 (RUSU); Sede do Parque, próximo aos sanitários, 10 Oct 2004, Calvente

& Versieux 101 (RUSU).

4. Epiphyllum phyllanthus (L.) Haw. subsp. phyllanthus, Syn. pl. succ. 197. 1812. (Fig. 3 G-H). Cactus

phyllanthus L., Sp. pl. 1:469. 1753.

Epiphyte, glochids absent, branches ca. 1.5 m long, erect or with pendent apex. Stem segments flattened,

somewhat oblong with lateral strangulations, 20-60 x 0.5-5 cm, leafless, fleshy, green, margin serrate or

crenate. Areoles borne at the stem segments margins indents, glabrous. Flower nocturnal, 0-1 per areole,

sessile, hypocrateriform, 19-24 x 3-4 cm; pericarpel oblong, ca 2.0 x 0.7 cm, with few sparse deltoid bract-

scales; flower-tube long, 16-20 x 0.4 cm, with few sparse bract-scales; perigonium segments oblong, 1.0-1.2

x 0.3-0.4 cm, spreading, apex acute; external segments ca. 6, green; internal segments ca. 16, pinkish white;

filament white, anther brown; ovary inferior, stigma with 9 lobes; nectar-chamber along the interior of the

flower-tube. Fruit oblong, ca. 8.5 x 3 cm, green when immature, with few sparse oblong bract-scales.

This species can be semiheliophyte to sciophyte. It is probably pollinated by moths because of the long

and slender flower-tube. According with Bauer (2003) the fruit when ripe is magenta and dehiscent by a

lateral slit. It is a plant very frequent in its area of occurrence, inhabiting tropical forests and dryer forests

such the ones in the northeast and middle west of Brazil. It flowers during October also when immature

fruits were found. Kimnach (1964) disting varieties for this species (var. phyllanthus, rubrocoronatum,

hookeri, guatemalense, pittieri, and columbiense), however, Bauer (2003) recognizes for Epiphyllum phyllanthus

only the typical subspecies and the E. phyllanthus subspecies rubrocoronatum (Kimnach)Ralf Bauer, which

are differentiated principally by the color of stamens, white on the former and red or orange to purple on the

latter. The remaining varieties of Kimnach (1964) are considered subspecies of E. hookeri Haw. on Bauer's

synopsis (2003), due to morphological and geographical aspects. Conservation status: Least Concern

(LC):

£s+haD o ID L

536 Journal of t titute of Texas 1(1)

Specimens examined: BRAZIL. Rio de Janeiro: Mun. Rio de Janeiro, Parque Natural Municipal da Prainha, subida para o Morro da

Boa Vista, ca. 200 ms.m., 09 Oct 1996, Braga & Lira Neto 3559 (RB); Bosque das Brasiliopuntias, 28 Oct 2003, Calvente & Bocayuva 66

(RUSU)

5. Hylocereus setaceus (Salm-Dyck ex DC.) Ralf Bauer, Cactaceae Syst. Init. 17:29. 2003. (Fig. 4 A-B).

Cereus setaceus Salm-Dyck ex DC., Prodr. 3:469. 1828. Selenicereus setaceus (Salm-Dyck ex DC.) Werderm., Bras. Saulenkakt. 87.

1933. Selenicereus rizzinii Scheinvar, Rev. Brasil. Biol. 34:249. 1974.

Shrub, scandent or prostrated, rupicolous, glochids absent. Stem segments 3-winged, (20—)35-120 x Q-)3-5

cm, leafless, fleshy, sometimes with woody parts, green, margin entire, lobed or faintly serrate. Areoles borne

at the wings margins indents with short tomentose hairs; spines 3-4, lateral, conic, short, 0.4-0.5 cm long,

stiff, pungent. Flower nocturnal, 0-1 per areole, sessile, infundibuliform, 25-35 x 6-8 cm, lateral; pericarpel

ca. 4 x 2 cm. Pericarpel areoles borne on top of tubercles, with 1 oblong basal bract-scale with acute apex;

hairs short, tomentose; spines 4—5, acicular, 6-8 cm long, slightly stiff, purplish. Flower-tube long, ca. 10

x 3 cm with areoles similar to the pericarpel areoles but gradually towards the flower apex with hairs and

spines scarcer and longer bract-scales; perigonium segments ca. 40, erect to suberect, oblong; external seg-

ments 2-9 x 1 cm, fleshy, green, apex acute; internal segments ca. 9 x 2 cm, membranaceous, white, apex

rounded; ovary inferior, stigma with 18 lobes; nectar-chamber along the interior of the flower-tube. Fruit

ovoid, ca. 7.0 x 3.5 cm, purplish, indehiscent, perigonium black persistent. Fruit areoles with pilose hairs;

spines 6-14, acicular, 0.5-0.8 cm long.

It is a heliophyte species very frequent on rock outcrops. Sometimes the stem segments have expanded

conspicuous wings. It is probably pollinated by moths because of its long flower-tube. Recently, Bauer (2003)

transferred this species from Selenicereus (L.) Britton & Rose to Hylocereus so it is common to find specimens

at the herbaria still identified as S. setaceus. It flowers annually in October and fruits were observed from

January to March. Conservation status: Least Concern (LC).

Specimens examined: BRAZIL. Rio de Janeiro: Mun. Rio de Janeiro. Parque Natural Municipal da Prainha, Pedra dos Cabritos, 28

Oct 2003, Calvente & Bocayuva 68 (RUSU); Cruzeiro do Sul, 29 Jan 2004, Calvente & Versieux 80 (RUSU); ibidem, 10 Oct 2004, Calvente

& Versieux 105, 106 (RUSU).

6. Lepismium cruciforme (Vell.) Miq., Bull. Sci. Phys. Nat. Néerl. 49. 1838. (Fig. 5 A). Cactus cruciformis Vell., Fl.

flum. 207. 1829. icon. 5:tab.29. 1831. nom. cons.

Epiphyte, prostrated, mesotonically branched, glochids absent. Stem segments all of indeterminate growth

3-winged, 9-24 x 2-3 cm, leafless, fleshy, green, wings continuous in the same stem segment, margin crenate

or serrate, extremes attenuate. Areoles borne at the wings margins indents, with copious long white pilose

hairs. Flower diurnal, 0—2(—3) per areole, sessile, ca. 1 x 0.7 cm, lateral or subapical; pericarpel immerse

in the areole, glabrous; flower-tube short, less than 0.5 cm, glabrous; perigonium segments 5-7, yellowish

green to whitish, purple spotted; stamen pale yellow; ovary inferior, style purplish, stigma white with 3-4

lobes. Fruit globose, 0.5—0.6 cm diam, magenta, glabrous, indehiscent, perigonium deciduous.

Only one specimen was collected within the NMPP, growing as sciophyte without flowers, so its de-

scription here is based on additional specimens. Fruits were observed in October. Conservation status:

Least Concern (LC).

Specimens examined: BRAZIL. Rio de Janeiro: Mun. Rio de Janeiro, Parque Natural Municipal da Prainha, 10 Oct 2004, Calvente &

Versieux 108 (RUSU); Mun. Maricá, Restinga de Maricá, 24 Oct 1986, Giordano et al. 179 (RB). Sáo Paulo: Mun. Londrina, entre a represa

Chavantes e Jurumim, 20 Oct 1966, Linderman & Haas 3163 (RB).

7. Rhipsalis elliptica G. Lindb. ex K. Schum. in Martius, Fl. Bras. 4(2):293. 1890. (Fig. 5 B-C).

Epiphyte, branches ca. 60 cm long, acrotonically or rarely subacrotonically branched, glochids absent. Stem

segments all of determinate growth, leafless, fleshy, green, margin crenate or incised; primary segments

1-3 subsequent, flattened, 9.5—23 x 0.4-2.0 cm, base semicylindrical; secondary segments 1-2(3) in each

branching axis, flattened, elliptic, oblong or obelliptic, rarely 3-winged, 4—16 x 1—5 cm, wings continuous

at the same segment, margin sometimes undulate. Areoles borne at the wings margins indents, glabrous;

Calvente and Andreata, Cactaceae of the Natural Municipal Park of Prainha (Brazil)

Fic. 4. A-B. IE ical (Salm-Dyck ex oe ) Ralf Bauer; A. B

Lem. subsp. fe

N NI

N LI

SN SN |

EX i

a Wo

un Wo My

RANN | WU.

with fruit, F. Longitudinal section of flower G- H. Pil

FOl bal

VRylac 2 Rawlar

-i L4

rf R

£I

(RAS

££

C-D. Cereus fernambucensis

J.) Backeb subsp fluminensis; E Branch

t, H.I git

{fl

537

f4L,D o ID L

538 Journal of t titute of Texas 1(1)

when on reproductive phase with scarce pilose hairs; 0—3(—5) bract-scales. Flower diurnal, 0-1(-2) per

areole, sessile, rotate, 1.0 x 1.4 cm, lateral or subapical, flower-tube absent; pericarpel 0.5—0.6 x 0.3-0.4 cm,

glabrous; perigonium segments golden yellow, membranaceous, conspicuously reflexed at anthesis involving

the pericarpel; external segments 3, deltoid to oblong, 0.05—0.4 x 0.05-0.3 cm; internal segments 5, oblong

to obelliptic, 0.6-0.7 x 0.2-0.5 cm; ovary inferior, stigma with 5 lobes; nectary a ring around the style base.

Fruit oblong, ca. 0.6 x 0.4 cm, purple, glabrous, indehiscent, perigonium deciduous.

This species is sciophyte and the margin of the stem segments can be lacerate by predation or wearing.

Flowers annually and synchronically among the individuals in August. Fruits in November. The flowering

period is short (ca. 1 week) and few fruits develop completely. It is endemic of the Brazilian Atlantic Forest.

Conservation status: Least Concern (LC).

Specimens examined: BRAZIL. Rio de Janeiro: Mun. Rio de Janeiro, Parque Natural Municipal da Prainha, trilha para a pedra dos

cabritos (morro da Boa Vista), 09 Oct 1996, Braga 3554 (RUSU); ibidem, 22 Aug 2003, Calvente et al. 50 (RUSU); ibidem, 11 Jun 2004,

Calvente & Versieux 96 (RUSU); trilha para o Cruzeiro do Sul, 22 Aug 2003, Bocayuva et al. 46 (RUSU).

8. Rhipsalis grandiflora Haw., Suppl. pl. succ. 83. 1819. (Fig. 5 D-E).

Epiphyte, branches ca. 2.5 m long, acrotonically branched, glochids absent. Stem segments all of deter-

minate growth, leafless, fleshy or woody, grayish green, cylindrical, 6.5-13 x 0.5-0.8 cm, 13 (-4) in each

branching axis. Areoles borne around the stem segments, glabrous. Young stem segments areoles purplish

with 0-1 deltoid minute bract-scale. When on reproductive phase stem segments areoles with 1-2 falcate

bract-scales. Flower diurnal, 0—2 per areole, sessile, rotate, 1.5 x 2.5-3.0 cm, lateral, perpendicular to the

stem segments, flower-tube absent; pericarpel ca. 0.2 x 0.4 cm, glabrous, slightly immersed in the areole;

perigonium segments spreading or reflexed; external segments 5-6, deltoid, 0.1-0.6 x 0.1-0.2 cm, semifleshy,

greenish, apex acute; internal segments 8—9, oblong, 0.7-1.2 x 0.4-0.6 cm, membranaceous, pinkish or

yellowish white, apex rounded slightly involute; ovary inferior, stigma with 5 lobes; nectary a ring around

the style base. Fruit globose, white, glabrous, indehiscent, with persistent black perigonium.

This species is heliophyte or semiheliophyte. The stem segments frequently have lacerated regions.

The flowers have a sweet smell and are visited by bees which are supposed to be the pollinators. Flowers

in October and fruits in November. Few fruits were observed on specimens from NMPP. Rhipsalis grandi-

flora when sterile is similar to R. teres f. heteroclada (Britton & Rose) Barthlott & N.P. Taylor but can be

distinguished because it has all the stem segments of determinate growth and lacks composite terminal

areoles. The purplish areoles in its young stem segments is a feature also present in other species such as

R. neves-armondii K. Schum. which can be distinguished by its pinkish flowers with golden yellow stamens

and conspicuously erumpent flower buds borne in wooly areoles. In general R. grandiflora is stouter than all

the similar species and its flowers are conspicuously large and showy. It is endemic of the Brazilian Atlantic

Forest and its habitat is reducing due to the urbanization. Conservation status: Near threatened (NT).

Specimens examined: BRAZIL. Rio de Janeiro: Mun. Rio de Janeiro, Parque Natural Municipal da Prainha, Bosque das Mirtáceas, 12

Apr 2003, Calvente et al. 32 (RUSU); Trilha para o Cruzeiro do Sul, 11 Jun 2004, Calvente & Versieux 95 (RUSU); ibidem, 10 Oct 2004,

(17 Oct 2004, fl. cult), Calvente & Versieux 117 (RUSU).

9. Rhipsalis paradoxa (Salm-Dyck ex Pfeiff.) Salm-Dyck subsp. paradoxa in Cact. Hort. Dyck. 1849:228.

1850. (Fig. 5 F).

Lepismium paradoxum Salm-Dyck ex Pfeiff. in Enum. Diagn. Cact. 140. 1837.

Epiphyte, branches ca. 2 m long, acrotonically branched, glochids absent. Stem segments all of indeterminate

growth, leafless, fleshy, green, 9-21(-26) x 1-2 cm, 1-3 in each branching axis, 3-winged in transversal

section; wings 3.0—7.5 x 0.4—0.6(-1.0) cm, discontinuous in the same segment, subsequent to the areoles.

Areoles borne around the stem segments, glabrous. Flower diurnal, sessile, rotate, 1.5 x 2.0 cm, white,

lateral, flower-tube absent; pericarpel ca. 0.4 x 0.4 cm, glabrous; perigonium segments ca. 8, oblong, 0.7-1.2

x 0.15-0.3 em, apex rounded; ovary inferior, stigma with 5 lobes. Fruit depressed-turbinate, 0.4-0.8 cm

diam., white, glabrous, indehiscent, perigonium deciduous.

Calvente and Andreata, Cactaceae of the Natural Municipal Park of Prainha (Brazil) 539

Fic 5 A I | s 2 of. (V H ) Miq ; L L "4E a T BC DL ip | lli a Y aij Lindh ey K Schum ;B D L 1 d 4 ,C I |

of flower. D—E. Rhipsalis grandiflora Haw.; D. Branch with fl E. Longitudinal section of fl F. Rhipsalis | (Salm-Dyck ex Pfeiff.) Salm-Dyck

subsp. paradoxa, branch. G-H. Rhipsalis teres f. heteroclada (Britton & Rose) Barthlott & N.P. Taylor; G. Branch with fl | fruits, H. Longitudinal

J | D L ‘esh £l If. | I *4.

section of flower. l-J. Rhipsalis triangularis W ; f flower.

540 Journal of the Botanical R h Institute of Texas 1(1)

This is a sciophyte species that in the NMPP was not found fertile so the flowers and fruits are described

here according with additional specimens. It is endemic of the Brazilian Atlantic Forest and its habitat is

reduced due to the deforestation and urbanization. Rhipsalis paradoxa subsp. septentrionalis N.P. Taylor &

Barthlott occurs in Pernambuco, Bahia, Minas Gerais and Espírito Santo states and is distinguished from

the typical subspecies due to its darker yellow flowers and slender stem segments with 0.7-1.1 cm (Barthlott

& peas 1995). Conservation status: Near Threatened (NT).

d: BRAZIL. Rio de Janeiro: Mun. Rio de Janeiro, Parque Natural Municipal da Prainha, Cardoso (s.n., RUSU 14317);

Due das Brasiliopuntias, 10 Oct 2004, Calvente & Versieux 116 (RUSU); Mun. Mangaratiba: Reserva Rio das Pedras, Rio Grande, 21-22

Dec 1996, Lira Neto et al. 522 (RUSU); trilha para o cambucá, 200—300ms.m, 20 Oct 1996, Bovini et al. 1090 (RUSU).

10. Rhipsalis teres f. heteroclada (Britton & Rose) Barthlott & N.P. Taylor, Bradleya 13:65. 1995. (Fig. 5

G-H). Rhipsalis heteroclada Britton & Rose, Cact. 4:224. 1923.

Epiphyte, branches 1-1.5 m long, acrotonically or subacrotonically branched, glochids absent. Stem segments

cylindrical, leafless; primary segments long, of indeterminate growth, 8.0-22 x 0.3 cm, 1-2 subsequent,

usually woody, grayish; secondary segments short, of determinate growth, 2.0-7.0 x 0.1-0.3 cm, 2-7 in each

branching axis, fleshy, green. Areoles borne around the stem segments, usually glabrous, areoles terminal

in the stem segments composite and with pilose hairs. Young stem segments areoles with scarce pilose

hairs and 1 deltoid minute bract-scale. When on reproductive phase stem segments areoles with 1-2 falcate

bract-scales ca. 0.1 cm long. Flower diurnal, 0—2 per areole, sessile, rotate, 0.6—0.8 x 0.9-1.0 cm, lateral,

sub-apical or apical, slightly oblique to perpendicular to the stem segments, flower-tube absent; pericarpel

ca. 0.3 cm diam, glabrous; perigonium segments suberect to spreading, membranaceous, greenish to red-

dish; external segments 4, deltoid, ca. 0.1 cm long or shorter, the apex slightly reflexed; internal segments

5, triangular to oblong, 0.4-0.6 x 0.1-0.2 cm, apex rounded, sometimes with the margins involute; ovary

inferior, stigma with 4 lobes; nectary a ring around the style base. Fruit depressed-globose, ca. 0.4 cm diam,

greenish white, glabrous, indehiscent, with persistent black perigonium.

This species is frequent at the NMPP as sciophyte to semisciophyte. It is distinct from the other forms

of Rhipsalis teres (Vell.)Steud. due to the stouter stem segments and larger flowers. The young apical shoots

are slender then the mature basal segments. The Rhipsalis teres f. heteroclada is very hard to be differentiated

from the R. teres f. teres in herbaria, especially in those cases when only the slender apical shoots are collected.

For that reason, more detailed studies must be done to establish a more precise morphological differentiation

among them. Rhipsalis teres f. heteroclada is endemic and frequent in the Brazilian Atlantic Forest. It flowers

from June to October and fruits from July to November. The fruit mature rapidly. Conservation status:

Least Concern (LC).

Specimens examined: BRAZIL. Rio de Janeiro: Mun. Rio de Janeiro, Parque Natural Municipal da Prainha, trilha para a pedra dos

Cabritos (morro da Boa Vista), ca. 150 ms.m. 09 Oct 1996, Braga & Lira Neto 3557 (RUSU, RB); ibidem, 29 Oct 2003, Braga et al. 7232

(RB); ibidem, 15 May 2004, Calvente & Versieux 86, 91 (RUSU); Bosque das Mirtáceas, 23? 02' 41,2"S 43? 30' 40,5"W, 09 Jul 2003, Calvente

et al. 49 (RUSU); trilha para o Cruzeiro do Sul, 03 Oct 2003, Bocayuva 70 et al. (RUSU); ibidem, 17 Oct 2003, Calvente et al. 61 (RUSU),

Bosque das Brasiliopuntias, 11 Jun 2004 (10 Oct 2004, fl. cult), Calvente & Versieux 94 (RUSU)

11. Rhipsalis triangularis Werderm., Feddes Repert. Spec. Nov. Regni Veg. 1071-1080: 3. 1937. (Fig. 5 I-J).

Shrub non columnar, rupicolous or epiphyte, acrotonically branched, glochids absent. Stem segments all

of determinate growth, leafless, fleshy, 6-24 x 1-4(-6) cm, 1-5 in each branching axis, pale green or yel-

lowish green, margin serrate or crenate usually reddish, 3—-4(—5) winged, wings continuous at the same

segment. Areoles 1,5-2,5 cm apart at the wings margins indents, with short tomentose hairs; bract-scales

0—5, stiff, vestigial from the reproductive phase, similar to spines. When on reproductive phase or young

stem segments areoles with 0—8 linear bract-scales. Flower diurnal, 0—3 per areole, sessile, rotate, ca.

1.5 x 2.5 cm, lateral, sub-apical or apical, flower-tube absent; pericarpel 0.4—0.6 x 0.4—0.6 cm, glabrous;

perigonium segments reflexed or spreading, white; external segments 3-4, deltoid, 0.2-0.4 x 0.4-0.6 cm,

slightly fleshy, apex rounded; internal segments 9-10, oblong, 0.5-0.9 x 0.2-0.4 cm, membranaceous,

Calvente and Andreata, Cactaceae of the Natural Municipal Park of Prainha (Brazil) 541

apex rounded slightly involute; ovary inferior, stigma with 5-7 lobes; nectary a ring around the style base.

Fruit depressed-globose, 0.6-0.7 x 0.7-0.8 cm, pinkish when immature and white when ripe, glabrous,

indehiscent, with persistent grayish black perigonium.

This is a heliophyte or semiheliophyte species which forms large populations at the rock outcrops

and probably reproduces vegetatively. The basal stem segments sometimes have the margins lacerated.

The flowers are visited by bees which are supposed to be the pollinators, ants and moths. The flowering is

abundant and synchronic among the individuals during April to June and scarce in October. Fruits were

observed in November and April. Werdermann (1937) described Rhipsalis triangularis from a cultivated

material which was sent from the Rio de Janeiro Botanical Garden to the Dahlem Botanical Garden and the

original habitat of this species was unknown for the author. Rhipsalis triangularis during many years was

considered a problematic taxa because of the absence of the type material and new collections with the

features described by Werdermann (Barthlott & Taylor 1995). However the specimens found at the NMPP

fit perfectly in the original description of R. triangularis with the exception of few differences which can be

attributed to different environmental conditions or to the fact that Werdermann (1937) based his descrip-

tion in one specimen with only 3 stem segments. These few differences are that the specimen described by

Werdermann had faintly indented serrate margins, absence of aerial roots and pale green pericarpel while

the NMPP populations have in general crenate or serrate margins, presence of aerial roots and brownish

green or pinkish magenta pericarpel. Rhipsalis agudoensis N.P. Taylor is similar to R. triangularis but is distinct

for its magenta pink fruits and is referred to a distant site, the city of Agudo at the Rio Grande do Sul State

(Taylor 2003). Further field studies are necessary to better evaluate the relation between Rhipsalis agudoensis

and R. triangularis. Conservation status: Critically Endangered (CR).

Specimens examined: BRAZIL. Rio de Janeiro: Mun. Rio de Janeiro, Parque Natural Municipal da Prainha, Pedra dos Cabritos, 340

ms.m., (mirante Boa Vista), 06 Jun 1996, Braga & Bovini 3357 (RUSU); ibidem, 17 Apr 2003, Calvente & Bocayuva 37, 38 (RUSU); ibidem,

12 Dec 2003, Cardoso & Zaldini 51 (RB); ibidem, 15 May 2004, Calvente E Versieux 87, 89, 90 (RUSU); Cruzeiro do Sul, 06 Jun 2003,

Calvente et al. 42 (RUSU); Bosque das Brasiliopuntias, 10 Oct 2004, Calvente & Versieux 107a, 112 (RUSU).

12. Cereus fernambucensis Lem. subsp. fernambucensis, Cact. gen. sp. nov. 58. 1839. (Fig. 4 C-D).

Cereus obtusus Haw., Rev. pl. succ. 70. 1821.

Shrub, columnar, prostrated or erect, terricolous or rupicolous, glochids absent. Stem segments leafless,

fleshy, 12-26 x 3-5 cm, 2-3(-4) in each branching axis, grayish green; ribs 4-5, 1-2 cm wide, ca. 1 cm thick,

margin entire, faintly crenate or incised. Areoles borne at the ribs margins indents, with short tomentose

hairs; central spines 1-3, acicular, 1.0—3.2 cm long, stiff; lateral spines 2-5, acicular, 0.4-1.5 cm long, stiff.

Flower nocturnal, 0—1 per areole, sessile, infundibuliform, 17-23 x 13-17 cm, lateral; pericarpel ca. 1.5

x 1.2 cm; flower-tube 9-12 x 1 cm; pericarpel with bract-scales deltoid, ca. 0.2 cm long, apiculate, green,

towards the flower-tube apex gradually becoming purple, oblong, ca. 1.5 cm long with apiculus absent;

perigonium external segments ca. 14oblong, 2.0—5.5 x 0.7-0.9 cm, slightly fleshy, purplish green, spread-

ing, apex acute; internal segments ca. 30, obelliptic or oblong, 3.8-5.0 x 1.0-2.0 cm, membranaceous,

pinkish white, suberect, base attenuate, apex rounded; ovary inferior, stigma with 13 lobes; nectar-chamber

ca. 7 cm long. Fruit depressed-oblong, ca. 6 x 5 cm, magenta, glabrous, dehiscent by a longitudinal slit,

perigonium deciduous.

In the NMPP this species is frequent as heliophyte in the open restinga vegetation and on the rock out-

crops close to the beach and its altitudinal distribution reaches 100 ms.m at the most. Small coleopterans

feed on the androecious after the anthesis. It flowers annually from September to October and fruits from

October to March. Few fruits are produced after flowering and those were found ripe on the following year

flowering season. It is endemic of the Brazilian Atlantic Forest. In the Rio de Janeiro State occur Cereus fer-

nambucensis subsp. sericifer (F.Ritter) N.P. Taylor & Zappi and the typical subspecies, which is distinguished

by its coastal distribution and magenta pink fruits, while C. fernambucensis subsp. sericifer has an inland

distribution and yellows fruits (Taylor & Zappi 2004). Conservation status: Least Concern (LC).

542 Journal of the Botanical R h Institute of Texas 1(1)

Specimen mined: BRAZIL. Rio de Janeiro: Mun. Rio de Janeiro, Parque Natural Municipal da Prainha, sob pedra na praia, 10 Sep

2003, Calvente & de Paula 51 (RUSU); ibidem, 1 Oct 2004, Calvente & Versieux 102, 103, 104 (RUSU).

13. Coleocephalocereus fluminensis (Miq.) Backeb. subsp. fluminensis, Jahrb. Deutsch. Kakteen-Ges.

1941(2):53. 1942. (Fig. 4 E-F). Cereus fluminensis Miq., Bull. Sci. Phys. Nat. Neérl. 48. 1838.

Shrub, columnar, 50-130 cm, erect or semidecumbent, rupicolous, glochids absent. Stem segments leaf-

less, fleshy, sometimes woody at base, 60-180(2240) x 10 cm, green; ribs 9-15, ca. 1.5 cm wide, 1-2 cm

thick, margin entire. Areoles borne at the ribs margins, with short tomentose hairs; spines 3-7, acicular,

1.5-3.0 cm long, flexible, gray, yellowish at the apex, usually 1 central e the others lateral. Cephalium lateral,

10-15-30) x 5-7 cm, with copious long, lanate hairs; spines acicular, 3-5 cm long, sometimes undulate or

curve, flexible, golden yellow with brownish apex. Flower nocturnal, borne on the cephalium, 1-2 in each

flowering, sessile, infundibuliform, ca. 7 x 5 cm, pink; pericarpel ca. 0.8 x 0.8 cm, glabrous; flower-tube

with minute bract-scales sometimes involute; perigonium segments ca 65, triangular to oblong, 1.0-2.0

x 0.3-0.6 cm, membranaceous, spreading, apex acute sometimes involute; ovary inferior, stigma with 13

lobes; nectar-chamber 1.2 cm long. Fruit obconic, ca. 2.6 x 2.1 cm, magenta, glabrous, dehiscent by a basal

pore, with persistent black perigonium.

In the NMPP this taxon is frequent on the rock outcrops as heliophyte commonly growing on mosses.

The flowers are supposed to be bat pollinated because of its morphology and the fruits are frequently visited

by lizards and ants which are supposed to be the dispersers. It flowers synchronically in January, April,

june, August and October. The fruits were observed from March to November and are ripe almost during

the whole year, being released gradually by the cephalium. The specimens are threatened by the fire on

the rock outcrops and inhabit areas with hiking and climbing tracks where they are regularly destroyed.

This is a Brazilian Atlantic Forest endemic taxa vulnerable at the Rio de Janeiro State. It is distinguished of

Coleocephalocereus fluminensis subsp. decumbens (F.Ritter)N.P. Taylor & Zappi, which is a Minas Gerais State

endemic species, mainly due to its green grayish stem segments and dark brown spines (Taylor & Zappi

2004). Conservation status: Vulnerable (VU).

d: BRAZIL. Rio de Janeiro: Mun. Rio de Janeiro, Parque Natural Municipal da Prainha, Pedra dos Cabritos (morro

da Boa Vista), 359 ms.m., 06 Jun 1996, Braga & bovini 3342 (RUSU, RB); ibidem, 17 Apr 2003, Calvente & Bocayuva 36 (RUSU); ibidem,

29 Jan 2004, Calvente & Versieux 81 (RUSU); Cruzeiro do Sul, 28 Oct 2003, Calvente & Bocayuva 67 (RUSU).

14. Pilosocereus arrabidae (Lem.) Byles & Rowley, Cact. Succ. J. Gr. Brit. 19:66. 1957. (Figs. 4 G-H).

Pilocereus arrabidae Lem., Rev. Hort. 34:429. 1862.

Shrub, columnar, 1-2 m, erect, rupicolous, glochids absent. Stem segments 30-150 x 5-7 cm, leafless,

fleshy, rarely woody at base, green; ribs 5-6, 1.4-2.0 cm wide, ca. 2.5 cm thick, margin entire. Areoles

borne at the ribs margins, with short tomentose hairs; lateral spines 5-10, acicular, 0.2-1.0 cm long; central

spines 13, acicular, 1.0-2.0(3.0) cm long; areoles at the stem segment apex with scarce, long, pilose hairs.

Flowers nocturnal, 0-1 per areole, sessile, infundibuliform, 6—7(-9) x 3-4 cm, lateral; pericarpel ca. 0.9

x 1.9 cm, with 2-3 sparse bract-scales; flower-tube with 0—2 sparse fleshy deltoid bract-scales; perigonium

segments deltoid, apex acuminate; external segments (0.5—)1.0—1.4 x 0.7-1.0 cm, fleshy , greenish, suberect;

internal segments 1.4-2.2 x 0.5-1.0 cm, membranaceous, white, spreading; ovary inferior, stigma with 9

lobes; nectar-chamber ca. 0.9 cm long. Fruit depressed-globose, 2-4 x 3—4.5 cm, magenta red, glabrous,

dehiscent by a lateral slit, with persistent black perigonium.

It is a heliophyte species rare at NMPP but common at the adjacent restinga of Grumari. It is probably

pollinated and dispersed by bats due to its flowers and fruits morphology. It flowers and fruits in October.

Pilosocereus arrabidae is a Brazilian Atlantic Forest endemic usually inhabiting the coastal restingas. The Rio

de Janeiro State is the southern limit of its geographic distribution where its natural habitat is threatened

by human interference. Conservation status: Near Threatened (NT).

Specimens examined: BRAZIL. Rio de Janeiro: Mun. Rio de Janeiro, Parque Natural Municipal da Prainha, Cruzeiro do Sul, 28 Oct

2003, Calvente & Bocayuva 70 (RUSU).

Calvente and Andreata, Cactaceae of the Natural Municipal Park of Prainha (Brazil) 543

PDLORISTIC ASPECTS

The Cactoideae subfamily was the most diverse at the NMPP, with 7996 of the inventoried species and is

followed by Opuntioideae (1496) and Pereskioideae (796). Four tribes of Opuntioideae and Cactoideae occur

at the area of study: Opuntieae, Hylocereeae and Rhipsalideae with 2 genera each and Cereeae, the largest

one with 3 genera. The Pereskioideae is monogeneric and only one species belonging to this subfamily oc-

cur at the NMPP, Pereskia aculeata. The richest genus is Rhipsalis, with five species, while the other genera

present only one species each (Fig. 6, Table 1).

In the NMPP, 5096 of the taxa are strictly terrestrial and among them 4396 are only rupicolous, 4396

are rupicolous or terricolous and 1496 are only terricolous. The exclusive epiphytes correspond to 3696 of

the inventoried species and they belong mostly to Rhipsalideae. The 1496 remaining species can be either

epiphytic or terrestrial (Figs. 7, 8).

Three main vegetation physiognomies can be recognized in the NMPP, the coastal ombrophilous forest

(Atlantic Rain Forest), the restinga vegetation (beach or scrub and herbaceous vegetation at plain sands dunes)

and the rock outcrops. Most of the taxa (3696) occur only in forested areas as epiphytes, however, 29% of

the species can be found either in forested areas or on rock outcrops and 2196 inhabit exclusively the rock

outcrops. Opuntia monacantha is restricted to the restinga and Cereus fernambucensis subsp. fernambucensis can

occur either in the restinga or on rock outcrops, both cases correspond to 796 of the studied species (Fig. 9).

It can be observed that the species present habit plasticity, what is especially true concerning the taxa

present at the rock outcrops. Taxa as Pilosocereus arrabidae that are usually found as terricolous in restinga

vegetation or others as Rhipsalis elliptica that are preferably epiphytes at forested areas are found as rupicolous

at the rock outcrops, this also indicates that some particular conditions, such as high solar irradiation, water

stress and temperature could be common both to those terrestrial, rupicolous and epiphytic habitats, however,

more accurate studies are necessary.

The comparative analyses of the Cactaceae floristic results of this work with other areas at the Rio de

Janeiro State demonstrate greater similarity of the NMPP with other coastal areas represented by APA of

Cairucu, APA of Maricá, National Park of the Restinga of Jurubatiba and APA of Massambaba (Table 2, Fig.

10). These areas have taxa that are characteristic of restinga pioneer formations as Pereskia aculeata, Brasilio-

puntia brasiliensis, Opuntia monacantha, Hylocereus setaceus e Cereus fernambucensis subsp. fernambucensis. In

spite its location inside the Rio de Janeiro City, the area of the Vista Chinesa Forest Reserve is less similar

with the costal areas (Table 2, Fig. 10). This area comprises only Atlantic Forest formations lacking the restinga

vegetation characteristic taxa. The same applies to the Macaé de Cima Reserve, located in the mountains of

the Órgãos range in the Rio de Janeiro State, which can be even more differentiated in floristic aspects due to

altitudinal and climate peculiarities (e.g. Lima & Guedes-Bruni 1997; Rocha et al. 2003; Calvente et al. 2005).

CONSERVATION

The taxa occurring in the NMPP are grouped under four threat categories: Critically Endangered (CR),

Vulnerable (VU), Near Threatened (NT) and Least Concern (LC).

Rhipsalis triangularis is considered Critically Endangered because of its restricted populations only

currently known to occur in small areas at the NMPP and because of the fragility of its habitat. Coleocepha-

locereus fluminensis subsp. fluminensis inhabits exclusively rock outcrops at the Rio de Janeiro State and is

the only taxa evaluated as Vulnerable (VU) in the NMPP, corresponding to 7% of the total number of taxa

(Fig. 11). This species suffer with the urbanization, pollution and human interference manifested through

vandalism, fire and lack of planning for the use of rock outcrops areas. This results in the decline of their

population and in the decrease of habitat quality and for that reason the conservation of populations of this

species in the NMPP must be strongly assured.

Rhipsalis grandiflora, R. paradoxa subsp. paradoxa, and Pilosocereus arrabidae are considered Near Threat-

ened (NT) corresponding to 21% of studied taxa (Fig. 11). Their habitat is decreasing because of the great

fal, Dat o ID L

544 Journal of

titute of Texas 1(1)

TABLE 1. Cactaceae taxa occurring at the Natural Municipal Park of Prainha, RJ.

Subfamily Tribe Genus Specific & Conservation

infraspecific status

taxa

Pereskiodeae — Pereskia P aculeata LC

Opuntioideae Opuntiaea Brasiliopuntia B. brasiliensis LC

Opuntia O. monacantha LC

Cactoideae Hylocereeae Epiphyllum E. phyllanthus LC

subsp. phyllanthus

Hylocereus H. setaceus LG

Rhipsalideae Lepismium L. cruciforme Le

Rhipsalis R. elliptica ES

R. grandiflora NT

R. paradoxa NT

subsp. paradoxa

R. teres f. heteroclada ES

R. triangularis CR

Cereeae Cereus C. fernambucensis LC

subsp. fernambucensis

Coleocephalocereus C. fluminensis VU

subsp. fluminensis

Pilosocereus P arrabidae NT

Total 3 4 10 14

TABLE 2. Similarity values between the inventoried areas for the Cactaceae family at the Rio the Janeiro State (Freitas 1990/92,

1996,1997; Rizzini et al. 1990; Scheinvar et al. 1996; Moura & Costa 2001), greater values are shown in bold.

PNMP Macaé de Cairucu Maricá Vista Jurubatiba Massambaba

Cima Chinesa

PNMP * 9.5238 59.26 56 42.8571 60.8696 59.2593

Macaé de Cima * i 0 28.5714 0 0

Cairucu Š = 66.67 59.2593 63.6364 69.2308

Marica $ È n i

Vista Chinesa * i s iS Es 43.4783 44.4444

Jurubatiba id D à i is b 72.7273

Massambaba * 2 a = i ^ z

pressure caused by the deforestation and urbanization expansion. If their conservation is not guaranteed

they may be more threatened in the future, this is particularly important to P. arrabidae because it inhabits

the restinga vegetation, which is more threatened every day because of the aggressive urbanization and

tourism development in the coast of the Rio de Janeiro State (Joly et al. 1999). Conservation units located

in those coastal regions like the NMPP and the APA of Grumari must be in greater number, more protected

and controlled by the government to assure the preservation of their biological diversity.

Taxa under Low Risk (LC) are Pereskia aculeata, Brasiliopuntia brasiliensis, Opuntia monacantha, Epiphyl-

lum phyllanthus subsp. phyllanthus, Hylocereus setaceus, Lepismium cruciforme, Rhipsalis elliptica and R. teres f.

heteroclada, corresponding to 65% of sampled taxa. They are usually more frequent or widely distributed in

the Rio de Janeiro State and their populations are in a less priority situation in spite the reduction of their

habitat (Fig. 11).

Many Coleocephalocereus fluminensis subsp. fluminensis specimens were seen burned and dead inside

Calvente and Andreata, Cactaceae of the Natural Municipal Park of Prainha (Brazil) 545

PERESKIOIDEAE

OPUNTIOIDEAE

14%

CACTOIDEAE ^v

79%

Fic. 6. Percentage of each subfamily of Cactaceae at the Natural Municipal

Park of Prainha, RJ.

EPIPHYTE

36%

j| TERRESTRIAL

7 50%

TERRESTRIAL/

EPIPHYTE

Fi. 7. Percentage of each Cactaceae habit at the Natural Municipal Park

of Prainha, RJ.

RUPICOLOUS

4396

Y RUPICOLOUS

43%

Fic Q n de £ L Y 4 4 x * IL | 441 Al 4 TAA I

i F

Park of Prainha, RJ.

OMBROPHILOUS RESTINGA

FOREST/ ROCK 7%

PE RESTINGA/

° ROCK

~~ OUTCROPS

omBROPHILOUS (IHE T e

FOREST

36%

ROCK

OUTCROPS

21%

il hki£e4 A M +L T d. + +1 N

Fic. 9. P t the Natural

Municipal Park of Prainha, RJ.

areas damaged by fire on the rock outcrops in the

NMPP. The Cactaceae taxa are highly resistant to

water stress and to high solar irradiation, however

they do not have any structures that can protect

them against fire and once they are burned they

hardly regenerate (Gibson & Nobel 1986). The

invasion of alien gramineous species of Panicum

L. and Melinis Beauv. can aggravate the situation

because they are easily flamed, dispersing the fire

to the individuals around them.

The occurrence of climbing paths and tracks on

all the extension of the rock outcrops can also rep-

resent a threat to the inhabitant taxa of these areas.

The Cactaceae individuals are particularly disturbed

by this because they are frequently damaged or

removed for their prickly stems that are consider a

threat by the eyes of visitors. The ideal action was

to restrict and signalize specific areas at the rock

outcrops to be used for ecological tourism and at

the same time develop an education work with the

visitors teaching the importance of these species and

how to act on the behalf of their conservation.

Porembski et al. (1998) observed the advan-

tages on the colonization of the rock outcrops by

the Cactaceae, Bromeliaceae and Velloziaceae. The

harsh environmental conditions in these habitats

make difficult the establishment of seedlings origi-

nated of sexual reproduction, however, the high

capacity of clonal growth of these plants raises the

chance of a perpetual colonization because the

establishment of only one individual result on the

formation of great populations in isolated areas.

According with Meirelles et al. (1999) effec-

tive actions have to take place in order to protect

the rock outcrops. These ecosystems are naturally

characterized by high rarity, endemism and fragil-

ity values that alone can justify their conservation.

Besides that, factors caused by human interference

such as pollution, vandalism and fire increase their

vulnerability which can result in local extinction.

Banana plant populations were observed in

forest areas inside the NMPP and on its surround-

ings, especially at the APA of Grumari. These

populations should be controlled and even removed

if necessary because they can disturb the integrity

of the original vegetation.

Rocha et al. (2003) listed the main pressures

against the conservation of the Pedra Branca State

£4h,D . ID L

546 Journal of t titute of Texas 1(1)

Jaccard Cluster Analysis (Group Average Link)

Macae de Cima

Vista Chinesa

Massambaba

Jurubatiba

Marica

Cairucu

PNMP

0. 96 Similarity 50. 100

Fic. 10. (Above) Similarity between the inventoried areas for the

Cactaceae family at the Rio de Janeiro State (Freitas 1990/92, 1996,

1997; Rizzini et al. 1990; Scheinvar et al. 1996; Moura & Costa 2001)

SNS

NT So

21% AD

o Ss

Fic. 11. Percentage of each conservation statuses evaluated for the

f Prainha, RJ..

Park and among them is the removal of the original forests for the establishment of monocultures as the

banana.

Visitors and people that work at the NMPP reported de occurrence of the extraction of ornamental

plants including cacti probably with the objectives of commercialization or cultivation in private properties.

Traps and camping signs suggest the action of hunters at the NMPP what compromises not only the animal

species but also the plant species because the opening of many new tracks disturbs the regenerating native

vegetation. Authority measures must be taken to eliminate these noxious activities at the NMPP.

It is possible to observe that the NMPP is still under many threats regardless of being a Conservation

Unit. One important step for the resolution of these problems would be the creation of a management plan

capable of assuring that the resolutions listed on the NMPP creation decree are effectively implemented.

Among them, the planning of a well directed ecological tourism at the area and the precise delimitation of

Calvente and Andreata, Cactaceae of the Natural Municipal Park of Prainha (Brazil) 547

the zone for environmental regeneration and of the intangible zone, which is to be designated exclusively

for the preservation of the natural resources would be critical.

ACKNOWLEDGMENTS

The authors thank the administration of the NMPP and the Secretaria Municipal de Meio Ambiente do Rio

de Janeiro for the plant collection permit; the Coordenacáo de Aperfeicoamento de Pessoal de Nivel Supe-

rior (Capes) for the graduate student fellowship and to the Conselho Nacional de Pesquisas (CNPq) for the

research productivity grant to the first and second authors respectively; to Beat Leuenberger and Ralf Bauer

for kindly providing their bibliography; to the curators of the consulted herbaria; to Maria Helena Pinheiro

for the ink drawings; to Eduardo Luís Martins Catharino and one anonymous reviewer for valuable com-

ments; and for all people that contributed for this work especially to Melissa Bocayuva, Leonardo Versieux,

Leandro Cardoso, Carlos Alberto Zaldini and to Joáo Marcelo Braga.

REFERENCES

ANDERSON, E.F. 2001. The cactus family. Timber Press, Portland, Cambridge.

BanrHLorT, W. and N.P. Tavon. 1995. Notes towards a monograph of Rhipsalideae (Cactaceae). Bradleya 13:

43-79.

BarTHLOTT, W. and D.R. Hunt. 1993. Cactaceae In: K. Kubitzki, J.G. Rohwer and V. Bittrich, eds. The families and genera

of vascular plants. Springer-Verlag Berlin, Heidelberg, Germany. 2:161-196.

Bauer, R. 2003. A synopsis of the tribe Hylocereeae F. Buxb. Cactaceae Syst. Init. 17:3-63.

Britton, N.L. and J.N. Rose. 1919. The Cactaceae: descriptions and illustrations of plants of the cactus family, volume

1. The Carnegie Institution of Washington, Washington D.C., U.S.A.

CALVENTE, A.M., M.F. Freitas, and R.H.P. ANDREATA. 2005. Listagem, distribuição geográfica e conservação das Cactaceae

no Estado do Rio de Janeiro. Rodriguésia 56(87):141-162.

FipaLGO, O. and V.L.R. Bononi (Coord.). 1984. Técnicas de coleta, preservação e herborizacáo de material botánico.

Instituto de Botanica, Sao Paulo.

FnerrAs, M.F. 1990/92 [1996]. Cactaceae da Área de Proteção Ambiental da Massambaba, Rio de Janeiro Brazil.

Rodriguésia 42/44:67-91.

Frerras, M.F. 1996. Cactaceae. In: M.P.M. Lima and R.R. Guedes-Bruni, eds. Reserva Ecológica de Macaé de Cima,

Nova Friburgo, RJ: Aspectos florísticos das espécies vasculares, volume 2. Jardim Botánico do Rio de Janeiro,

Rio de Janeiro. Pp. 153-161.

Frerras, M.F. 1997. Cactaceae Juss. In: M.C.M. Marques, A.S.F. Vaz and R. Marquete, org. Flora da APA Caiurucu,

Parati, RJ: espécies vasculares (Série estudos e contribuições n? 14). Jardim Botánico do Rio de Janeiro, Rio

de Janeiro. Pp. 105-117.

GiBSON, A.C. and PS. Noset. 1986. The cactus primer. Harvard University Press, Cambridge, Massachusetts.

HoLmGren, PK., N.H. HoLmGrEN, and L.C. Barnett. 1990. Index herbariorum Part I: The herbaria of the world. 8 ed.

New York Botanical Garden, New York.

Hunt, D.R. (Compiler). 1999. CITES Cactaceae Checklist. 2.ed. Royal Botanical Gardens Kew & International Orga-

nization for Succulent Plant Study (IOS), UK.

IUCN. 2001. IUCN Red List categories and criteria: Version 3.1. IUCN Species Survival Commission: Gland, Swit-

zerland and Cambridge, UK.

Jory, C.A., M.PM. Alpar, C.A. Kunk, D.G. McGrath, A.G. Moreira, P. Moutinho, D.C. NePsrAD, A.A. Olivera, A. Pott, M.J.N.

RopaL, and E.V.S.B. Sampaio. 1999. Evolution of the Brazilian phytogeography classification systems: Implications

for biodiversity conservation. Ci. & Cult. J. Brazil. Assoc. Advancem. Sci. 51(5/6):331-348.

Kimnach, M. 1964. Epiphyllum phyllanthus. Cact. Succ. J. (Los Angeles) 36:105-115.

LEUENBERGER, B.E. 1976. Pollen morphology of the Cactaceae. Cact. Succ. J. Gr. Brit. 38:79-94,

Lima, H.C. and R.R. Guebes-Bruni (Organizers). 1997. Serra de Macaé de Cima: diversidade florística e conservação

em Mata Atlántica. Jardim Botánico do Rio de Janeiro, Rio de Janeiro, RJ.

548 Journal of the Botanical R h Institute of Texas 1(1)

Meirettes, S.T., V.R. PiveLLo, and C.A. Joly. 1999. The vegetation of granite rock outcrops in Rio de Janeiro, Brazil, and

the need for its protection. Environm. Conservation 26:10- 20.

MINISTÉRIO DAS Minas E ENERGIA. 1983. Projeto RadamBrasil: folhas SF-23/24, levantamento de Recursos Naturais,

volume 32. Secretaria Geral, Rio de Janeiro.

MirrERMEIER, R.A., H.C.G. Mittermeler, and N. Myers. 2000. Hotspots: Earth's biologically richest and most endangered

terrestrial ecoregions. Conservation International/ CEMEX, México City.

Moura, R.L. and A.F. Costa. 2001. Cactaceae. In: A.F. Costa and I.C. Dias, eds. Flora do Parque Nacional da Restinga

de Jurubatiba e arredores, Rio de Janeiro, Brazil: listagem florística e fitogeografia. Angiospermas, pteridófitas

e algas. Museu Nacional, Rio de Janeiro. Pp. 50-51.

POREMBSKI, S.G. MARTINELLI, R. OHLEMULLER, and W. BarTHLOTT. 1998. Diversity and ecology of saxicolous vegetation mats

on inselbergs in the Brazilian Atlantic Rain forest. Diversity & Distrib. 4:107-119.

PREFEITURA DA CIDADE DO Rio DE JANEIRO. 1998. Guia das Unidades de Conservacáo Ambiental do Rio de Janeiro.

IBAMA/ DUMA/ SMAC, Rio de Janeiro.

Rio DE JANEIRO (Municipio). 1999. Decreto n? 17.445, de 25 de marco de 1999.

RocHA, C.F.D., H.G. BerGALLo, M.A.S. Alves, and M.V.A. Sluys. 2003. Biodiversidade dos Grandes Remanescentes

Florestais do Estado do Rio de Janeiro e nas Restingas da Mata Atlantica. Ed Rima, Sao Carlos, SP.

Rizzini, C.M., C. PEREIRA, E.M.L. OccuioNi, and F.V. Acarez. 1990.Consideracóes sobre a ocorrência de Cactaceae na

APA de Maricá, Rio de Janeiro, Brazil. Acta Bot. Brasil. 4:171-182.

SCHEINVAR, L., A.A. Canposo, D.C.P. Sıva, and J.C. ELeuTéRIO. 1996. Cactaceas da Reserva Florestal da Vista Chinesa.

Albertoa 4(1 1):117-136.

TayLor, N.P. 2003. A new Rhipsalis from Rio Grande do Sul, Southern Brazil. Cactaceae Syst. Init. 16:12.

Tavlor, N.P, W. Stuppy, and W. BarTHLOTT. 2002. Realignment and revision of the Opuntioideae of Eastern Brazil. In:

D. Hunt and N. Taylor, eds. Studies in the Opuntioideae (Cactaceae). David Hunt, England, UK. Pp. 99-132.

TayLor, N.P. and D.C. Zapri. 2004. Cacti of Eastern Brazil. The Royal Botanic Garden, Kew, Richmond, UK.

Wattace, R.S. and S.L. Dickie. 2002. Systematic implications of chloroplast DNA sequence variation in Opuntioi-

deae. In: D.R. Hunt. and N.P. Taylor, eds. Studies in the Opuntioideae (Cactaceae). David Hunt, England, UK.

Pp. 9-24.

WERDERMANN, I.E. 1937. Neue Sukkulenten aus dem Botanischen Garten Berlin-Dahlem. Il. Feddes Repert. Spec.

Nov. Regni Veg. 42:1-7.

ASSESSMENT OF PLANT BIODIVERSITY IN

WECHIAU COMMUNITY HIPPOPOTAMUS SANCTUARY IN GHANA

Alex Asase* Alfred A. Oteng-Yeboah

Department of Botany Council for Scientific and Industrial Research (CSIR) Ghana

University of Ghana P O. Box M32

20) n5 Legon- Accra, GHANA

Legon-Accra, GHANA

dim edu.gh

rresponding author

ABSTRACT

A study was conducted in the Wechiau Community Hippopotamus Sanctuary in Ghana in order to assess the plant biodiversity found in

the sanctuary. The study was conducted over a period of three years (2000-2003) through botanical inventories and using sample plots.

A total number of 227 species of plants belonging to 72 families were identified and three vegetations types: Riverine forest, Floodplain

and Upland flora were found in the sanctuary. Many of the species of plants were Mesophanerophytes and Thereophytes whereas a

paucity of the species was Heleophytes, Hydrophytes, Hemicryptophytes and Chamaephytes. The results of the study are discussed and

recommendations made for further research to support the conservation and sustainable use of plants in the sanctuary.

Key Worbs: Plant biodiversity, Wechiau, Hippopotamus, Ghana, conservation

RESUMEN

Se realizó un edd" en el M oa Community Hippopotamus Sanctuary en Ghana para valorar la biodiversidad vegetal presente en

el santuario in. El est 5 durante un periodo de tres años 2000-2003) mediante inventarios botánicos y usando parcelas de

muestreo. Se identificaron un total de 227 especies de plantas pertenecientes a 72 familias y tres tipos de vegetación: bosque de rivera,

llanuras de inundación y tierras altas en el santuario. Muchas de las especies de plantas fueron mesofanerófitos y terófitos mientras

que las especies minoritarios fueron helófitos, hidrófitos, hemicriptófitos y caméfitos. Se discuten los resultados del estudio y se hacen

recomendaciones para investigaciones posteriores con vistas a la conservación y el uso sostenible de las plantas en el santuario.

INTRODUCTION

The Wechiau Community Hippopotamus Sanctuary is a community-based initiative in the Upper West Region

of Ghana to protect the remaining unprotected hippopotamus (Hippopotamus ampbibius) population in the

Black Volta River, which would help to develop the ecotourism potential for the area. The other population

of hippopotamus in Ghana is found in the Bui National Park, which is about 56 km from the present sanc-

tuary. This population of hippopotamus in the park is under threat with the government's plan to develop

a dam for hydro-electrical power generation. It is anticipated that the present population of hippopotamus

in the Bui National Park will migrate to the sanctuary when the dam is finally developed.

The chiefs and people of Wechiau traditional area with technical assistance from a local non-gov-

ernmental organization, Nature Conservation and Research Centre (NCRC) have decided to conserve

the populations of hippopotamus and other biological resources in their area. They also hope to generate

income through the promotion of ecotourism focused to improve the livelihood of the rural communities

The sanctuary is one of the two examples in Ghana where the local people are taking full control of the

management of their biological resources through a Sanctuary Management Board (SMB). There is however

no baseline information on the plant biodiversity in the sanctuary that will assist in making management

decisions for the conservation and sustainable use of the plant resources in the sanctuary.

The present study was therefore conducted in order to assess the plant biodiversity in the Wechiau

Community Hippopotamus Sanctuary in Ghana.

MATERIALS AND METHODS

Study area

The study was conducted at Wechiau about 40 km southwest of Wa in the Upper West Region of Ghana.

The study area is positioned between latitudes 02%41'N and 02°49'N and between longitudes 09°43'W and

J. Bot. Res. Inst. Texas 1(1): 549 — 556. 2007

f4L,D o ID

550 Journal of t h Institute of Texas 1(1)

09°53'W. The area has been demarcated and conserved as Wechiau Community Hippopotamus Sanctuary

and occupies an area of about 40 Km? along the banks of the Black Volta River. The vegetation of the sanctu-

ary is primarily Guinea savanna (Oteng-Yeboah & Asase 2002) and the terrain is mostly flat.

METHODS

The plant biodiversity in the sanctuary was assessed through botanical inventories and using sample plots

over a period of three years (2000-2003). Botanical inventories entailed walking through different areas of

the vegetation of the sanctuary whiles picking and identifying plants with identification aids. Each species

of plant identified was monitored for its life form (sensu Raunkiaer 1934) throughout the period of the study.

The botanical inventories were conducted through twelve field visits to the sanctuary and voucher specimens

of the species of plants encountered were collected and have been deposited at the Ghana Herbarium (GC)

at the Department of Botany, University of Ghana. The preliminary field identifications of the plants were

checked using the Flora of West Tropical Africa (Hutchison & Dalziel 1954-1972) and by comparison with

identified specimens at the GC herbarium.

In order to determine the species richness in the sanctuary, sample plots measuring 25 mx 25m, 5m

x5mand Im x Im were randomly demarcated in the sanctuary. Forty-one of each plot size was studied.

The 25 m x 25 m plot was used to study trees and 5 m x 5 m used to assess small trees and shrubs whereas

the 1m x 1m plots was used to study the ground flora (herbaceous and grass species). The species richness

was evaluated from the average number of species count for all the plots examined for each plot size.

RESULTS

Species richness and composition

A total of 227 species of plants belonging to 71 families were identified in the sanctuary. The mean numbers

of species of trees was 7.62 in 5.3 families from the 25 m x 25 m plots and that of small trees and shrubs was

3.33 species in 2.62 families from the 5 m x 5 m plots whiles that of ground flora species was 4.10 species

in 3.92 families from the 1 m x 1 m plots.

Many of the plant species in the sanctuary (85.596) belong to the dicotyledonous group (Appendix 1).

The monocot group contributed 14.196 of the species and only one species of Pteridophyte (Ophioglossum

costatum) was identified from the sanctuary (Appendix 2 and 3). Within the dicotyledonous group, the family

Papilionaceae contributed the largest number of genera (9.296) and species (8.296). The other dicotyledonous

families that contributed many species and genera to the flora of the sanctuary were Rubiaceae, Asteraceae,

Euphorbiaceae, Ceasalpinaceae, Mimosaceae, Verbanaceae, and Combretaceae.

The monocotyledonous group contained many genera and species of the families Poaceae, Liliaceae,

Cyperaceae, Araceae, and Commelinaceae. However, the families Hypoxidaceae, Orchidaceae, and Zingib-

eraceae contributed only one genus and species each to the flora of the sanctuary.

Vegetation types and species composition

The vegetation of the sanctuary changes gradually from Riverine forest vegetation through Floodplain vegeta-

tion to Upland flora vegetation as one moves inland from the Black Volta River. The three vegetation types

found in the sanctuary were largely based on their physiognomy and species composition.

Upland flora vegetation.—The appearance and species composition of this vegetation type is similar

to the general characteristics of the Guinea Savanna vegetation. The species diversity is very high in this

vegetation type compared to that of the other two vegetation types with species of trees from the families

Sapotaceae (Vitellaria paradoxa), Mimosaceae (Entada africana, Parkia biglobossa), Caesalpinaceae (Afzelia

africana, Daniellia oliveri and Tamarindus indica), Anacardiaceae (Haematostaphis barteri, Lannea acida and

Lannea kerstingii), Bombacaceae (Adansonia digitata, Bombax costatum), Meliaceae (Khaya senegalensis and

Pseudocedrela kostchyi), Simaroubaceae (Hannoa undulata), Papilionaceae (Pterocarpus erinaceus) and Com-

bretaceae (Terminalia avicennoides and Terminalia species).

Smaller trees and shrubs were from the families Euphorbiaceae (Bridelia ferruginea), Cochlospermaceae

Asase and Oteng-Yeboah, Flora of the Wechiau Community Hi | Sanct 551

(Coclospermum planchonii and C. tinctorium), Combretaceae (Combretum ghasalense, Combretum molle and

Combretum spp.), Celastraceae (Maytenus senegalensis), Chrysobalanaceae (Parinari curattefollia and Parinari

polyandra), Olacaceae (Ximenia americana), Caesalpinaceae (Piliostigma thonningi and Swatzia madacasgarensis)

and Polygalaceae (Securidaca longepedunculata).

The herbaceous flora also included species from the families Liliaceae s.l, but currently in Asparaga-

ceae (Asparagus flagellaris), Hypoxidaceae (Curculigo pilosa), Euphorbiaceae (Sapium grahamii, Euphorbia baga,

Euphorbia ladermannniana) and Zingiberaceae (Kaempferia aethiopica). Grass species were mainly Andropogon

gayanus, Hyparhennia subplumosa, Hyperthelia spp., and Ctenium newtonii.

In areas where the vegetation has previously been intensively farmed, Vitellaria paradoxa and Parhia

biglobossa were the main trees found and the herbaceous species consisted of ruderal plants from the fami-

lies Asteraceae (Ageratum conyzoides, Emilia sp., Tridax procumbens), Rubiaceae (Mitracarpus scaber, Borreria

radiata) and Euphorbiaceae (Euphorbia hirta, Euphorbia heterophylla).

Floodplain vegetation.—This vegetation was found next to the Upland flora vegetation towards the

Black Volta River. The vegetation area is often flooded during the rainy season and the ground becomes

patchy in the dry season. Trees and climbers were few but with a very dense grass cover in the rainy season.

The main tree species were from the families Rubiaceae (Mitragyna inermis) and Mimosaceae (Acacia nilotica

and Acacia gourmaensis). Other species of trees were from the families Balanitaceae (Balanities aegyptiaca),

Rhamnaceae (Ziziphus mauritianum), Rubiaceae (Crossopteryx febrifuga), Combretaceae (Terminalia macrop-

tera), Anacardiaceae (Lannea acida and L. kerstingii), Papilionaceae (Pterocarpus erinaceus) and Meliaceae

(Pseudocedrela kotschyi).

The ground flora was predominated by members of the family Poaceae (including Vetiveria fulvibarvis,

Andropogon gayanus, Brachiaria jubata, and Hyparhhenia subplumosa). Herbaceous species of the families Mi-

mosaceae (Cassia mimosoides), Papilionaceae (Crotalaria goorensis and Tephrosia platycarpa), Amaryllidaceae

(Crinum humile) Polygonaceae (Polygonum senegalensis), Commelinaceae (Aneilema sp., Floscopa africana, and

Murdannia simplex), and Spenocleaceae (Sphenoclea zeylanica) were also common.

Riverine forest vegetation.—This vegetation type was found on either side of the Black Volta River.

The vegetation was very dense as a result of large number of climbers and relatively tall trees that formed

a canopy. Species of tree from the families such as Sterculiaceae (Cola laurifolia), Papilionaceae (Pterocarpus

santaliniodes), and Bignoniaceae (Kigelia africana) that are found along permanent rivers were common. Other

common trees were from Combretaceae (Anogeissus leiocarpus), Ulmaceae (Celtis integrifolia), Bombacaceae

(Ceiba pentandra), and Ebenaceae (Diospyros mespiliformis).

Among the smaller trees and shrubs were members of the families: Caesalpinaceae (Cassia sieberiana), Papili-

onaceae (Milletia zechiana), Mimosaceae (Endata abyssinica), Flacourtiaceae (Cassipourea congoensis, Dissomeria

crenata), Rutaceae (Afraegle paniculata), and Verbanaceae (Vitex chrysocarpa).

Climbers and herbaceous species were mainly from Sapindaceae (Paullinia pinnata), Passifloraceae

(Passiflora sp.), Colchicaceae (Gloriosa superba), Annonaceae (Monanthotaxis sp.), Capparidaceae (Capparis

erythrocarpus), Araceae (Achomanes welwitchii), Liliaceae s.l, but strictly in the family Anthericaceae (Chloro-

phytum pusilium), and Papilionaceae (Aeschynomene afraspera).

Life forms and species composition

The 227 plants identified in the sanctuary belong to 9 life-forms categories (Table 1). Many of the species

of plants in the sanctuary were Mesophanerophytes and Thereophytes with few species of Heleophytes,

Hydrophytes and Hemicryptophytes in the sanctuary. Many of the species of Thereophytes in the sanctuary

were members of the families Rubiaceae, Asteraceae and Euphorbiaceae. Also the Mesophanerophyte and

Microphanerophyte were made up of species from Caesalpinaceae, Meliaceae, Anacardiaceae and Mimosa-

ceae. The Chamaephytes made up of species from the families Annonaceae, Aristolochiaceae and Cochlo-

spermaceae whereas the species of Nanophanerophtes were from members of Sapindance and Verbenaceae.

Geophytes were commonly members of the Amaryllidaceae and Commelinaceae.

552 Journal of the Botanical R h Institute of Texas 1(1)

TABLE 1. Life form composition of species of plants in Wechiau Community Hippopotamus Sanctuary in Ghana.

Life-form Total number of Contribution of life form to the

species contributed flora of the sanctuary (96)

Thereophyte herb (ThH) 60 2115

Thereophyte climbing herb (Thl) 3

Chamaephyte herb (ChH) 5 7.05

Chamaephyte climbing herb (ChHI) 6

Chamaephyte shrub (ChS) 5

Hydrophyte herb (HyH) 3 1:32

Helophyte herb (HeH) ] 0.045

Geophyte herb (GrH) 26 11.45

Hemicryptohyte (Hcr) 8 3:52

Mesophanerophyte tree (MPT) qu 28.21

Mesophanerophyte shrub (MPS) 36

Microphanerophyte shrub (mpS) 15 8.37

Microphanerophyte epiphye (mpE) 2

Microphanerophyte liana (mpWl) 2

Nanophanerophyte shrub (NpS) 21 11.89

Nanophanerophyte liana (NpWI) 6

DISCUSSION

The present study is the first attempt to compile an authoritative checklist of the plant biodiversity in the

Wechiau Community Hippopotamus Sanctuary and the study has shown that the composition of plant

species in the sanctuary is similar to that of Guinea savanna vegetation (Hopkins 1974; Lawson 1985). The

species richness and diversity in their life-forms of the plants in the sanctuary was also similar to Guinea

savanna vegetation studied by Oteng-Yeboah (1996)

Bushfires impact on plant biodiversity and therefore it is important for the Sanctuary Management

Board to make strategic management plans that will protect core areas of the sanctuary from bushfires and

other human activities such farming. With the exception of few species of plants such as Afraegle paniculata

(Schum. & Thonn.) Engl., Commiphora dalzielli Hutch. and Ceiba pentandra (L.) Gaertn, many of the species

of plants identified in the sanctuary were generally very common. The Sanctuary Management Board could

initiate species specific management programs such as the collection of seeds and other propagules of these

uncommon species of plants for their subsequent cultivation in the sanctuary.

It is hoped that this information on the plant biodiversity in the sanctuary will serve as a baseline

data for making management decisions for the conservation of the sanctuary resources and promotion of

ecotourism activities in the area. It is however recommended that further research should investigate the

quantitative abundance and distribution of the species of plants as well as the indigenous uses of the plants

in sanctuary.

APPENDIX

Checklist of dicotyledonous plants in Wechiau Community Hippopotamus Sanctuary in Ghana. See Table

1 for definition of life-form.

Amaranthaceae Lannea kerstiigii Engl. & K. Krause; MPT

Celosia trigyna L; ThH Spondias mombin L.;

Pandiaka involucrata (Moq.) B.D. Jackson; ThH Annonaceae

Anacardiaceae Annona gl Schumach. & Thonn,; ChS

Haematostaphis barteri Hook.f; MPT Annona senegalensis Pers.; ChS

Ozoroa insignis Delile; MpS Hexalobus petalus Engl. & Diels., NpwWI

Lannea acida A. Rich. MPT

Ll VAL kL: ‘al . Lis P c

Asase and Oteng-Yeboah, Flora of the Wechiau Community

Apocynaceae Celastraceae

Saba senegalensis (A. DC.) Pichon; mPWI Hippocratea africana Loes. ex Engl.; ThH

Strophanthus hispidus DC., NpS Maytenus lensis (Lam.) Exell; npS

Aristolochiacea Chrysobalanaceae

Aristolochia un Duch.; ChHI Parinari curatellifolia Planch. ex Benth.; mpS

Asclepiadaceae Parinari polyandra Benth.; mpS

Calotropis procera (Aiton) W.T. Aiton; npS Cochlospermaceae

Pachycarpus lineolatus (Decne) Bullock; npS Cochlospermum planchoni Hook. f; ChS

Asteraceae Cochlospermum tinctorum A. Rich.; ChH

Acanthospermum hispidum DC.; ThH Combretaceae

Ageratum conyzoides L.; ThH Anogessius leiocarpus Guill. & Perr; MPT

Aspilia africana (Pers.) C.D. Adams; ThH Combretum aculeatum Vent; mpS

Blumea aurita DC.; ThH Combretum ghaselense Engl. & Diels; mpS

Echinops longifolia A. Rich.; GrH Combretum hypopilinum Diels; mpS

Gomphrena celosoides Mart.; ThH Combretum molle L. Br. ex G. Don; mpS

Synedrella nodiflora Gaertn.; ThH Combretum nigricans Leprieur ex Guill. & Perr; mpS

Tridax procumbens L.; Combretum paniculatum Vent.; Now!

Vernonia amygdalina Delile; npS Combretum sericeum G. Don.; ChH

Vernonia perrottetii Sch.Bip. ex Walp.; ThH Pteleopsis suberosa Engl. & Diel; mpS

Vernonia purpurea Sch.Bip.ex Walp.; ThH Quisqualis indica L; MPS

Vicoa leptoclada (Webb) Dandy; ThH Terminalia avicenoides Guill. & Perr.; mpS

Terminalia laxiflora Engl.; MPS

Terminalia macroptera Guill. & Perr; mPT

Balanitaceae

Balanites aegyptiacus Delile; MPS

Convolvulaceae

Bignoniaceae ON ance ere

Kigelia africana (Lam.). Benth.; MPT o eR

Ipomoea sp.; NoWI

tereospermum kunthianum Cham.; MPS

Cucurbitaceae

td DEPTS f ThEI

insonia digitata L; MPT Zehneria halli Hook. f; ThH

Bombax costatum Pellegr. & Vuillet; MPT Ebenaceae

Ceiba pentandra (L.) Gaertn; MPT Diospyros mespiliformis Hochst. ex A. DC; MPT

Boraginaceae Euphorbiaceae

Heliotropium indicum L.; ThH Bridellia ferruginea Benth.; MPS

Bülterscade Euphorbia baga ^. Chev.; ThH

Euphorbia convolvuloides Hochst ex Benth.; ThH

Commiphora dalzielli Hutch.; mpS

Euphorbia ladermaniana Pax & K. Hoffin,; TAH

Caesalpinaceae Euphorbia macrophylla Pax.; ThH

Burkea africana Hook.; MPS Euphorbia poissonii Pax.; npS

Cassia absus L; ThH Euphorbia sp.; npS

Cassia mimmosoides L.; ThH Hymenocardia acida Tul.; MPS

Cassia sieberiana DC.; MPS Jatropha curcas L; npS

Cassia tora E ThH . Jatropha gossypiifolia L; npS

Dalium guineensis Willd.; MPS Phyllantus amarus Schum & Thonn.; ThH

Daniellia oliveria (Rolfe) Hutch. & Dalz; MPT Sapium grahamiPrain.; ThH

iid Uli Guill. & Perr. MPS Securinega virosa (Willd) Baill.; Chs

rlina doka Craib & Stapf; MPT Tragia vogelii Keay.; ThH

Piliostigma thoni ngii i (Schumach) Delile-Redh.; mps .

Flacourtiaceae

Caesalpinaceae Oncoba spinosa Forssk.; MPS

Tamarindus indica L MPT

enna occidentalis (L) Link; ThH Lamiaceae

Ocimum canum Sims; ThH

Capparaceae i ;

Cleome viscosa L.; ThH AR ;

Cadaba tnoceEorsskm Strychnos innocua Delile; MPS

Capparis erythrocarpos Isert; NDWI Strychnos spinosa Lam. MP5

Ritchiea reflexa Gilg & Benedict; NpWI

554

Loranthaceae

Tapinanthus dodeneifolius (DC.) Danser mpE

Tapinathus bangwensis (Eng. & Krause) Danser; mpE

Malvaceae

Hibiscus asper Hook.f; ThH

Sida alba L; TAH

Sida sp.; ThH

Meliaceae

Khaya senegalensis A. Juss; MPT

Pseudocedrela kotschyi Harms.; mpT

Trichilia emetica Vahl; mpS

Menispermaceae

Cissampelos mucronata A. Rich.; ChHI

Mimosaceae

Acacia gourmaensis A. Chev.; MPS

Acacia hockii De Wild.; mpS

Acacia nilotica L. (Delile). MPS

Acacia sp.; np

Afzelia africana Sm.; MPT

Dicrostachys glomerata Chiov.; mpS

Entada abyssinica Steud.; mp

Entada africana Guill. €: Perr; mpS

Mimosa pigra L.; ThH

Parkia biglobosa (Jacq) R. Br. ex G. Don; MPT

Proposis africana Taub.; MPT

Moraceae

Ficus abutilifolia (Miq) Miq.; MPS

Ficus capensis Thunb.; MPS

Ficus platyphylla Del.; MPS

Moringaceae

Moringa oleifera Lam.; MPS

Myrtaceae

Eugenia subherbacea A. Chev; ThH

Nyctaginaceae

Boerhavia diffusa L.; ThH

Olacaceae

Boerhavia diffusa L.; ThH

Opiliaceae

Opilia celtidifolia (Guill. & Perr.) Endl. ex Walp.; mPWI

Papilionaceae

Abrus precatorius L.; ThHI

Aeschynomene afraspera J. Leonard.; ThH

Alysicarpus ovalifolius J. Leonard.; ThH

Canavalia sp.; TRH

Crotalaria goreensis Guill. & Perr; ThH

Erythrina senegalensis DC.; mpS

Lonchocarpus laxifl Guill. & Perr; MPS

Lonchocarpus sericeus (Poir) H.B.K.; npS

Millettia zechiana Harms; npS

Ostryoderris stuhlmannii (Taub.) Dunn ex Harms; MPS

Pericopsis laxiflora (Benth. ex Baker) Meeuwen; MPS

Pterocarpus erinaceus Poir; MPT

Pterocarpus santalinoides UH'er ex DC; MPT

Swarztia madagascarensis Desv.; MPS

f*hAD o ID L

Journal of t titute of Texas 1(1)

Tephrosia platycarpa Guill. & Perr.; ThH

Uraria picta (Jacq) DC; ThH

Passifloraceae

Passiflora foetida L.; ThH

Pedaliaceae

Sesamum alatum Thonn. ThH

Sesamum indicum L.; Th

Polygalaceae

Securidaca longepedun: ulata Fresen.; MPS

Polyonoaceae

Polygonum senegalensis Meisn.; Hy

Portulacaceae

Talinum trigulare Jacq.) Willd.; ThH

Rhamnaceae

Ziziphus mauritiana Lam.; ChS

Rhizophoraceae

Cassipourea congoensis R. Br ex DC; MPS

Rubiaceae

Borreria radiata DC.; ThH

Borreria scaber (Schum & Thonn.) K. Schum; ThH

Chassalia sp.; mpS

Chrysanthellum americanum Rich.; ThH

Crossopteryx febrifuga (Afzel. ex G. Don) Benth.; MPT

Fadogia agrestis Schweinf. ex Heirn.;; npS

Feretia apodanthera Del.; ThH

Gardenia ternifolia Schum. & Thonn.; npS

Mitracarpus scaber Zucc.; ThH

Mitragyna inermis (Willd). O. Ktze; mPT

Nauclea latifolia Sm.; mpS

Oldelandia corymbosa L.; ThH

Polysphaeria arbuscula K. Schum.; npS

Rutaceae

Afraegle paniculata Engl.; mPT

Zanthoxylum xanthoxyloides (Lam.) Waterman.; nPS

Samydaceae

Dissomeria crenata Hook. f. ex Benth.; MPS

apindaceae

Allophyllus africanus P. Beauv.; NDWI

Blighia sapida Kónig; MPT

Cardiospermum grandiflorum SW.; ThHI

Paullinia pinnata L; NDW

Sapotaceae

Malacantha alnifolia Pierre; MPT

Vitellaria paradoxa C.F. Gaertn. f; MPT

Scrophulariaceae

Striga hermonthica Benth ; ThH

Striga linearifolia (Schumach. & Thonn.) Hepper.; ThH

Simaroubaceae

Hannoa undulata Planch.; MPT

Solanaceae

Datura metel L; TAH

Asase and Oteng-Yeboah, Flora of the Wechiau Community Hi | Sanct 555

Sphenocleaceae Ulmaceae

Sphenoclea zeylsnica Gaertn.; Hy Celtis integrifolia Lam. MPT

Starculiaceae Trema orientalis Blume.; NpS

Cola laurifolia Mast; MPT Verbena

Sterculia setigeria Delile; MPS Mud capitatum Schum. & Thonn,; NpS

Waltheria indica A. L. ; ChH Lantana trifolia L; ThH

Taccaceae MA Med Has ChH

Tacca leontopetaloides (L.) Kuntze.; GrH nie.

del md Planch.; NpS

Tiliaceae Vitex doniana Sweet; MPS

Corchorus C. ThH T

Grewia carpinifolia Juss.; MpS bod TET

S Hutch. & Dalziel; ChH Cissus E CNH

lis Juss; MpS Cissus populnea Guill. & Perr; ChHI

APPENDIX

Checklist of monotyledonous in Wechiau Community Hippopotamus Sanctuary in Ghana. See Table 1 for

definition of life-form.

Amaryllidaceae Anthericace

Crinum humile A. Chev; GrH E M Schweinf. ex Baker; GrH

Araceae Hyacinthaceae

Haemanthus rupestris Baker; GrH Scilla picta A. Chev. ex Hutch & Dalziel.; GrH

Amorphophallus dracontioides N.E. Br.; GrH

Anchomanes welwitchii Rendle; GrH

Stylochiton lancifolium Kotschy & Peyr.; GrH

Orchidaceae

Eulophia cristata Lindl.; GrH

Poaceae

Andropogon gayanus Kunth; Hcr

Brachiaria lata (Schumach.)C.H. Hubb.; Hcr

Brachiaria sp. Hcr

Ctenium villosum Berhaut.; GrH

Commelinaceae

Aneilema setiferum A. Chev.; GrH

Floscopa africana C.B. Clarke; Cl.; GrH

Murdannia simplex (Vahl) Brenan; GrH

Cyperaceae Cymbopogon giganteus Chiov.; Hcr

Cyperus ambilis Vahl; GrH Cynodon dactylon (L.) Pers.; Hcr

Fimbristylis so., Hcr all aegyptium Willd.; GrH

Kyllinga sp.; GrH ET a Sp; GrH

Mariscus foliosus C.B. Clarke; GrH Hyperrhenia sp.; Hcr

und Speer

Panicum maximum Jacq.; GrH

Saccharum spontaneum L.; HyH

Liliaceae s.l. Aloaceae Setaria sp.; GrH

Aloe buetteneri A. Berger; GrH

Hypoxidaceae

Curculigo pilosa (Schum. €: Thonn.) Engl; GrH

Zingiberaceae

Asparagac Kaempferia aethiopica Solms ex Engl.; GrH

Asparagus decl Baker; ChHI

Pteridophyte species in Wechiau Community Hippopotamus Sanctuary in Ghana. See Table 1 for definition

of life-form.

Ophioglossaceae

Ophioglossum costatum R. Br; GrH

ACKNOWLEDGMENTS

We are thankful to the communities living in the sanctuary and the SMB for their hospitality and logistic

support throughout the study. Earthwatch Institute provided the financial assistance for the study. We are

also grateful to NCRC for logistics support. Sebsebe Deimissew (ETH) and Hilary O. Edeoga (Michael Okpara

Univ. of Agriculture, Umuahia, Nigeria) provided critical reviews.

556 Journal of the Botanical R h Institute of Texas 1(1)

REFERENCES

Hopkins, B. 1974. Savanna and forest. Heinemann, Ibadan and London.

DaLzieL HUTCHINSON, J. and J.M. DaLziEL. 1954-1972. Flora of West Tropical Africa. Vols.1, 2 & 3. Crown Agents, Lon-

don.

Lawson, G.W. 1986. Plant life in West Africa. Ghana Universities Press.

OTENG-YEBOAH, A.A. 1996. Biodiversity in three traditional groves in Guinea Savanna, Ghana.

OrtNG-YEBOAH, A.A. and A. Asase. 2002. Wechiau Community Hippopotamus Sanctuary-Preliminary data on floristic.

Ghana Science Asssociation Book of Abstracts. P. 57.

RauNktER, C. 1934. The life forms of plants and statistical plant geography. Clarendon Press, Oxford.

BOTANICAL COMPOSITION AND MULTIVARIATE ANALYSIS

OP VEGETATION ON THE POTHOWAR PLATEAU. PAKISTAN

Altaf A. Dasti Shehzadi Saima Mohammad Athar!

Institute of Pure & Applied Biology Institute of Pure & Applied Biology California Dept. of Food and Agric.

Bahauddin Zakariya University Bahauddin Zakariya University 1220 N Street

Multan, PAKISTAN Multan, PAKISTAN Sacramento, California 95814, U.S.A.

Attiq-ur-Rahman Saeed A. Malik

Institute of Pure & Applied Biology Institute of Pure & Applied Biology

Bahauddin Zakariva University ahauddin Zakariva University

Multan, PAKISTAN Multan, PAKISTAN

ABSTRACT

Vegetation of Pothowar Plateau (32? 32' and 34? north latitude and 70? 17' and 73? 5' east longitude, annual rainfall 250 to 750 mm) was

analyzed using ordination (DECORANA) and classificatory cluster ca NUM Five us associations were cce on the

basis of the cluster analysis. The most noticeable feature indicated by this analyses

from the flat lowland community. Clearly disjunct p merged from these analyses The major axes brought out Bs the ordination

were related to broad soil types. The application of the classification to the ordination allowed an interpretation of the vegetation varia-

tion in terms of topography, redistribution of rainwater, the nature of the bedrock and soil depth. The vegetation patterns revealed are

discussed in relation to geo-morphological factors and problems of plant assemblage in vegetation of widely scattered plants.

Key Worbs: Vegetation analysis, plant communities, Pothowar Plateau, Pakistan

RESUMEN

La vegetación de la meseta de Pothowar (32? 32' y 34? latitud Norte 70? 17' y 73? 5' longitud Este, precipitación anual de 250 a 750

mm) se analizó mediante ordenación (DECORANA) y técni lasificatorias de análisis cluster. Se reconocieron cinco asociaciones de

plantas en base al análisis cluster. La i A ble señalad álisis fue la separación del complejo d

de montaña de la comunidad de las zonas llanas bajas. De los ollo surgieron patrones dane a Los ejes principales

que salen de la ordenación están relacionados con los grandes tipos de suelo. La aplicación de la clasificación a la ordenación permitió

una interpretación de la variación de la vegetación en términos de topografía, redistribución de la precipitación, la naturaleza de la roca

madre y la profundidad del suelo. Se discuten los patrones de vegetación en relación con los factores geo-morfológicos y problemas de

inserción de plantas muy diseminadas en la vegetación.

INTRODUCTION

Studies on the arid and semi-arid areas of Pakistan and India have mainly been floristical and/or phyto-

geographical (Athar 2005; Champion & Sethi 1968; Chaudhuri 1960; Hussain 1969; Gupta 1975; Shaukat

et al. 1976; Malik et al. 1988). The use of numerical approaches has been rare (Shaukat et al. 1990; Shaukat

& Ahmad 1989), although these may be useful for summarizing the major gradients in data sets, assisting

the formulation of hypotheses and testing their validity (Birks 1992; Odgaard & Rasmussen 2000). Modern

synecological methods have developed techniques for use at local and regional level, seeking to reduce the

complexity of field data sets by classification and ordination of floristic data and then relating the results

to environmental information (ter Braak 1987). Such objective approaches have rarely been applied to the

vegetation data of Pakistan.

The primary objective of this study was to explore the factors that determine the boundaries and

composition of plant communities on the Pothowar Plateau. This was achieved by sampling all common

species present within a complex vegetation mosaic coinciding with local gradients in topography and soil

distribution. Numerical techniques were used to summarize the floristic data.

! Corresponding author: atariq@cdfa.ca.gov

J. Bot. Res. Inst. Texas 1(1): 557 — 568. 2007

558 Journal of the Botanical R h Institute of Texas 1(1)

MATERIALS AND METHODS

Study Area

The Pothowar Plateau is situated between 32? 32' and 34 north latitude and 70? 17' and 73? 5' east longitude.

It covers an extensive area of 1.5 million hectares. In altitude it rises from about 250 m near the Indus River

to 1,076 m in the west. The hills are detached and isolated from each other. They run mostly in an east west

direction and consist of a series of serrated ridges. The slopes are gentle at first and then climb more steeply.

The steepness of the slopes varies from 15° to 50? and is precipitous in some places. Elsewhere the gradient

is flat to gently undulating. It runs across the northern part of the area as a wedge with its base resting the

Indus, where it is 22 km wide. It tapers gradually over a length of 72 km to the east, and ends about 24 km

north-west of Margalla Range.

Climate

The climate of the area is of an extreme nature. The western portion of tract is hotter and drier than the

eastern and northern parts. The winter is bitterly cold while the summer is unbearably hot. The temperature

rises first in April, than remains almost steady, due to windstorms from Balochistan, up to the middle of

May when it shoots up again. June and July are the hottest months (average maximum temperature 42? C),

while December and January are the coldest months (average minimum temperature 1.7? C). The monsoon

starts by the third week of July and continues till the beginning of September when the nights get cooler.

The cold weather sets in by the middle of October. During winter the days are bright, and the nights are

clear. Early spring frosts are common and sufficiently severe to cause wide spread injury to plants, even to

the indigenous tree growth in a fairly advanced stage of development. Preliminary analyses of weather data

indicate that mean temperature decreases in a linear fashion with altitude (Champion et al. 1965). Frost

may occur especially at the upper altitudinal limits and in valley bottoms.

Rainfall is scanty and uncertain, and its annual distribution is very uneven. The annual rainfall varies

from 250 to 750 mm. Monsoons start late in July, and most of the annual rain is before September. Spring

and fall rains are rare and uncertain. Winter rains start by the end of December. They stop by the end of

February when the windstorms set in. Winter rains generally extend over a shorter period than the mon-

soons, followed by a prolonged period of dry weather. Humidity is also generally low, falling to about 1596

in the summer afternoons, the annual mean being about 5096 (Hussain & Ilahi 1991).

The ground water resources are limited. The sources of water for humans and livestock are wells and

dugout ponds. The drainage is very satisfactory. The entire area is drained off to the west into the Indus

River, largely through Soan River. Springs are mostly seasonal and flow only during the rains. Perennial

springs are very few, and are found at only 14 places.

Geology, soils and topography

Geologically the area is divisible into Attock slate and limestone. Attock slates are Precambrian and contain

gritty layers of an arenaceous type. On weathering they give rise to fertile loamy clay, which collects only in

sheltered places. The major part of the area (Kala-Chitta forest area) is composed of limestone in age from

Triassic to Liassic. It contains marls, ferr and bleached shales and sandstones belonging to the Eocene

suit, ferruginous pisolite, variegated sandstone, soft sandy carbonaceous clays, and shales varying in age

from Albian to Tithorian. Tertiary freshwater formations of sandstone, alternating with red and purple clays

and shales, are also very common in the Pothowar Plateau. Soils are universally shallow and dry and may

overlie a great variety of geological formations from limestone, shales and quartzites to crystalline rocks.

The soil derived from sandstone is poor and less fertile than soil from limestone. It supports nothing except

the most xerophyllous vegetation (Hussain & Ilahi 1991)

Vegetation

Champion et al. (1965) described the study area as dry sub-tropical broad-leaved forest. Rafi (1973), Beg

(1975), and Hussain and Ilahi (1991) called it dry sub-tropical semi-evergreen forest. The trees and shrubs

Dasti et al., Vegetation of Pothowar Plateau, Pakistan 559

are mostly thorny and have leaves of moderate size resembling Mediterranean vegetation elsewhere in the

world (Naqvi 1974). There is usually little ground-layer vegetation most of the year, but during the monsoon

a fairly complete cover of grass and herbs may develop. Floristic composition varies continuously in response

to small-scale altitudinal differences on the plateau, although most species show fairly broad distribution

patterns at that scale. The limestone region has only scrub forest composed of odd stunted and mutilated

Acacia with its associates. A few diminutive Olea and Rhamnus can also be observed on the northern slopes.

The southern slopes are almost entirely devoid of tree growth. The shrubs include a great many genera and

families, many of central Asian origin. Unpalatable species are favored by heavy grazing, and Dodonaea,

Rhazya, and Withania become conspicuous with thorny Carissa and Gymnosporia.

Pothowar Plateau is especially interesting because its vegetation merges at lower elevations with tropical

thorn forests and at higher elevations with subtropical pine and temperate forests. Because of its elevation,

precipitation on the plateau can reach, or even some times exceed, 600 mm, thus making Pothowar a sort

of enclave of the fringes of the arid areas in which the diversity of natural sites (wooded wadies, humid

canyons, springs and rock pools) has allowed a number of relict species to persist. The very situation of the

plateau thus makes it a sort of bio-geographical crossroad.

Vegetation Analysis

Attock district is representative of Pothowar Plateau. Twenty study sites were selected in Attock district to

cover the range of vegetation variation on the plateau (Fig. 1). Selection of these sites was based on repeated

surveys. Plant species were inventoried in 5 x 2 m quadrates. Field sampling procedures involved the random

placement of quadrates within each of the eight 25 x 50 m cells into which each study site was divided. On

each site, presence or absence of all species was recorded from a sample of 24 quadrates (three per cell).

Presence or absence of plant species was converted to frequencies and used to classify and ordinate both

sites and species by the reciprocal averaging (RA) procedure of Hill and Gouch (1980). Rare species were

eliminated from the analyses since these species can severely distort ordinations produced by RA (Hacker

1983; Dasti & Agnew 1994). Rare species occurred only in a single stand with frequencies of 196 or less.

Detrended correspondence analysis (DCA) was selected as an appropriate ordination method based on

gradient length and preliminary correspondence analyses (Jongman et al. 1995). The default options of the

program DECORANA were used for the analyses (Causton 1988; Hill 1979). DCA axes 1 and 2 were used

to interpret the data.

Species frequencies were clustered using the monothetic information statistic procedure incorporat-

ing the Spearman rank order dissimilarity coefficient (Causton 1988; Hill 1979). Scatter of classification

groups were plotted on overlays of the ordination axes to assess the compatibility of the two methods of data

simplification (Dargie & El Demerdash 1991; Dasti & Agnew 1994; Dasti & Malik 1998). The relationships

between soil characters and DCA axes 1 and 2 were determined using Spearman rank correlation (Causton

1988)

Soil Analysis

Soil samples (0—10 cm depth) were taken from each site at three different points and mixed into a composite

sample. Soil depth to 2 cm was sampled in the hard clays because the main root zone occurred within the

top 2 em layer. Sandy soils were sampled up to 10 cm depth because the top layer of soil was very mobile

and the rooting zone was deeper. The samples were air-dried and passed through a 2 mm sieve. Three sub-

samples were drawn from this composite sample. Soil texture, water holding capacity, and soil moisture

content was determined using standard methods (Hussain 1989; Richards 1954). Soil pH was recorded from

a pH meter (HM-1OK Digital, England), and conductivity was determined by CM-30 ET digital conductivity

meter. Organic carbon was estimated following Jackson (1958).

The Duncan multiple range test was used to detect and compare any significant difference between

the means of different communities at the 596 level of significance. The percent data obtained from particle

size analyses were normalized by an arcsine transformation and subjected to analyses of variance between

the communities for each variable.

560 Journal of the Botanical Research Institute of Texas 1(1)

C NG

e O MM 5

4 us. ond = M

HA H UU cake € a.

Mu E ie e ge y a

(7 (f. if pa tange SR o =

^w K «aet a BEES a) A

L vd X Ve

: O daos cm "s Fatah Jung’

Jand

" $

z $

2 Shahpur

Fic. 1. Attock district map st g tl pling the Pott Plateau. The study region i E eT RON lary

RESULTS

Classification

Five plant associations were recognized in the cluster analyses. The botanical composition of each associa-

tion is presented in Table 1. These associations were delineated based on specifying three hierarchical levels

(Fig. 2). The most noticeable feature indicated by this analysis was the separation of mountain-complex

communities (Associations A-D) from the flat lowland community (Association E). The 55 samples of group

E were separated from the other samples at the first level by Chrysopogon aucheri, and are characterized also

by Cyperus niveus and Dicanthium annulatum. In the mountain-complex region, the limestone plateau com-

munities (Associations A and B) were separated from those of the sandstone massif (Associations C and D)

at level two by Dodonaea viscosa. As a result of three hierarchical levels four communities in the mountain

complex were recognized:

1) Calcareous hilltops (Association A).

2) Interior rocky basins in the calcareous massif (Association B).

3) Sandstone massif (Association C).

4) Interior rocky basins on the sandstone massif (Association D).

The interior rocky basins on calcareous plateau (limestone strata) are characterized by Olea cuspidata that

is altogether absent from such basins in sandstone strata. The enclosed basins with moderately to slightly

Dasti et al., Vegetation of Pothowar Plateau, Pakistan 561

Taste 1. Mean relative frequency values (96) of species in the 5 associations detected by normal cluster analyses.

Species A B C D E

Acacia modesta Wall. 12:8] 25.48 23.64 28.90 16.83

Acacia nilotica (L.) Debile 1:63 0.70 - - 0.70

Anagalis arvensis L. 1.39 3.69 1.96 - 1.09

Asparagus gracilis Royle 0.40 - - - -

Boerhaavia coccinea Mill. - 237 - - 341

Capparis decidua (Forssk.) Edgew. 3.66 - 1555 [epee -

Carissa opaca Stapf ex Harines 0.64 - - - 3.09

Chrysopogon aucheri (Boiss.) Stapf. - - - - 49.60

Cymbopogon jawarancusa (Jones) Schult. 15:05 4.59 - - -

Cynodon dactylon (L.) Peris. 6.96 14:56 2,39 2.80 233

Cyperus niveus Retz. - - - - Ll

Desmostachya bipinnata (L.) Stapf. 14.01 - 6.65 9.34 -

Dichanthium annulatum (Forssk.) Stapf. - - - - 555

Dodonaea viscosa (L.) Jacq. 23.34 12.01 - - 19.26

Ehretia obtusifolia Hochst ex DC. 0.88 - - - 1:85

Eremopogon foveolatus (Del.) Stapf. 2.34 - - - -

Erianthus griffithii (Munro) Hk. f. 29.87 5.84 0.80 - 1.94

Erodium cicutarium (L.) UHerit ex Ait. - 1.19 16.21 - -

Evovulus alsinoides L. - 0.97 - - 3.62

Gnaphalium pulvinatum Del. 0.14 6.50 z = 2.51

Grewia damine Gaertin - - - - 2.66

Grewia hirsuta Vahl 1.04 - - 39.25 -

Grewia oppositifolia Roxb. 4.74 = = 19.62 3

Grewia tenax (Forsk.) Aschers & Schweinf. 2.16 0.96 7,22 20152 0.84

Grewia villosa Willd. 4.92 - 14.39 -— -

Gymnosporia royleana (Wall.) Lawson 9.75 21.24 21.08 8.09 18.67

Justicia peploides (Nees in Wall.) T. Anders 1.04 244 36.42 2 =

Linum strictum L. 7.30 - - - 447

Malcomia africana (L.) R. 0.25 0.92 4.28 19.62 112

Medicago laciniata (L.) Mill. 223 6.29 1282 - 1.85

Melhania futteypoyensis Munro ex Mast 2.95 - 5.8] - -

Myrsine africana L. 3.66 i 2.92 0.86 0.35

Olea cuspidata Wall. ex DC. 20.86 15:92 4.04 - 20522

Oxalis corniculata L. -— 512 0.80 - 1.93

Periploca aphylla Dcne. 1.94 - 5.63 2.89 -

Plantago ciliata Desf. - 2:37 - - 1.93

Prosopis juliflora Swartz 0.09 Lf 12.64 - -

Reptonia buxifolia Dcne. 123 7.57 - - -

Rhamnus pentapomica Parker 1:58 - 4.63 20.53 -

Rhazya stricta Dcne. 0.09 3:18 - 6.06 -

Sageretia theezans (L.) Brongn - - - - 6.24

Taraxacum officinale Wigg. 0.43 2.80 - - 0.99

Tetrapogon villosus Desf. - B - = 26]

Trachynia distachya (L.) Link 4.97 232.52 - 1.06 -

Ziziphus jujuba Mill. 3:69 9,93 i - ce

iziphus nummularia (Brum. f.) Wight €. Arn. 1E 2 0.23 16.47 299

impeded run-off are dominated by various species of Grewia. Further subdivisions at lower information

gains were regarded as minor variants, and were not considered. The vegetation communities are briefly

described below in the context of the major discriminating species. For this purpose the area has been

subdivided into five broad regions based on geomorphology.

£s+haD o ID L

562 Journal of t titute of Texas 1(1)

100% A | T

3 12

75%

EN EN ES EN

Dissimilarity

Coefficient

E D C B A

Fic. 2. Dendrogram for the cluster analyses of sampling sites, with 20 sites divided into five groups. The number of sites in each plant association is

given in boxes.

1. Mountain-Complex Plant Communities

The mountain-complex communities represent the vast extensive rocky hillocks and scarps with a thin

veneer of sediments. Skeletal soils of these habitats have no clear-cut profile. However, some minor rocky

hollows (enclosed basins) strewn with stony material exist in hills and scarps where run-off accumulates

during summer.

1.1. Limestone massif: Erianthus griffithii community—Association A.—This association dominates ex-

tensive areas in the limestone plateau and represents the characteristic vegetation of skeletal soils with no

clear-cut profile. Such soils develop generally on calcareous parent material and support xerophytic species.

Acacia modesta, Dodonaea viscosa, Erianthus griffithii, Gymnosporia royleana, and Olea cuspidata are the major

species of this type of vegetation. The dominance of E. griffithii, along with other perennial grass such as

Cymbopogon jwarancosa and Desmostachya bipinnata, gives this community a superficial resemblance to gra-

mineous steppe.

1.2. Enclosed basins in calcareous massif: Trachynia distachya community—Association B.—This association

represents the vegetation of depressions or hollows strewn with stony material with a lime incrustation.

This association is dominated by Acacia modesta, Dodonaea viscosa, Gymnosporia royleana and Olea cuspidate

Compared with the former association, the contribution of Cymbopogon jwarancosa, Eremopogon foveolatus and

Erianthus griffithii decreased remarkably. This association is marked by high average frequency of Trachynia

distachya and a ground cover of Cynodon dactylon, Malcomia africana, and Plantago ciliata.

1.3. Enclosed basins in sandstone massif: Justicia peploides—Association C.—Association C includes the

stands belonging to the northern inter-mountain depressions in sandstone strata. This association is distinc-

tive because of the high dominance of Acacia modesta, Capparis decidua, Erodium cicutarium, Grewia villosa,

Gymnosporia royleana, Justicia peploides and Prosopis juliflora and a few plants of Olea cuspidate

1.4. Sandstone plateau: Acacia modesta—Association D.—This association represents the southern slope

Dasti et al., Vegetation of Pothowar Plateau, Pakistan 563

of sandstone strata (Kala-Chitta range). The general vegetation is of the xeromorphic woodland type; the

chief tree and shrub species are Acacia modesta, Capparis decidua, Grewia hirsuta, G. oppositifolia, G. tenax,

Rhamnus pentapomica, and Ziziphus nummularia. The herbaceous vegetation is extremely short and includes

annuals like Malcomea africana that appears during the short period of the rainy seasons.

2. Lowlands or playes (Wadi beds) communities

Lowlands receive run-off during rains and become dry thereafter. Wadi soils are formed from alluvial

sediments carried down the slopes by run-off water. They are characterized by their heavier texture, high

water-holding capacity, deeper profile and slightly to poor drainage. The alluvial playes are dominated by

shrubs of Olea cuspidata associated with Chrysopogon aucheri (Association E) in more favorable habitats (oa-

ses), like runoff-fed depressions and runnels. Beside the dominant species (Cyperus niveus and Dichanthium

annulatum), Acacia modesta, Dodonaea viscosa, and Gymnosporia royleana are common associates. The woody

components of this association have large ranges and are generally found in oases as well as the plateau and

scarps under the most arid conditions.

Gradient Analysis

Site and species ordination in the plane of first two axes are presented in Figures 3 and 4. The first DCA

axis of the normal data set had an eigenvalue of 0.392 (1396 of the variance explained). The eigenvalue for

the second axis was 0.207 (896 of variance explained). Further axes each explained less than 796 of total

variances.

Site ordination (Fig. 3) reveals a marked relationship between the first axis and the soil factors. There

was a highly significant correlation (P « 0.01) between the sample scores along DCA axis 1 and the compo-

nents of soil texture (sand, silt and clay) and moisture, 1? values exceeded 0.4. Regarding axis 2, r values of

these edaphic factors were considerably low (« 0.03). Besides the soil physical characters, soil pH showed

a significant positive correlation (P«0.05) with DCA axis 1, but not with axis 2. The availability of organic

matter in the sample plots showed a significant correlation with both DCA axes (Table 2). In addition to

these factors, the ordination axes (1 and 2) appeared markedly influenced by topography, redistribution of

rainwater and soil depth. It is evident from Figure 3 that sites of the Wadi-beds (16-20, stands belonging

to Association E) are clearly grouped at one end (high score) and those of mountain complex (1-15 sites

belonging to Associations A-D) on the other end (low score) of DCA axis 1. However, among the mountain

complex, several sites are separated from the others suggesting a degree of floral diversity within this group.

Site 15 is strongly separated from rest of the sites suggesting again the operation of site-specific factors. Sites

of the enclosed basins (2, 3 and 12) that collect the run-off water occupy an intermediate position along

axis 1 reflecting the fair conditions of water and soil availability for plant growth. The distribution of the

species along DCA axes 1 and 2 is presented in Figure 4. Several species had configurations similar to that

evident for sites. However, some species do not follow the site distribution, suggesting that they are largely

unaffected by the underlying topographic or edaphic factors.

Comparison of the site and species ordinations reveals that the separation of Wadi sites (16-20) as a

distinct group at the resource-rich end of the gradient is due to the relative abundance of Chrysopogon aucheri

and the presence of Cyperus niveus, Dicanthium annulatum, Grewia damine, and Sageretia theezans. These spe-

cies are virtually absent from the plateau sites. The ordination positions of Grewia tenax, Justicia peploides,

Medicago laciniata, Prosopis juliflora, and Trachynia distachya suggest that these species are characteristic of

moderately xeric habitats. Both Cymbopogon jawarrancusa and Desmostachya bipinnata have maximum fre-

quencies on the limestone massif. Capparis decidua defines the most xeric end of the gradient, while Grewia

hirsuta and Rhamnus pentapomica display marked peaks in frequency on the eroded sandstone sites.

The remaining species occupy ordination positions that suggest a lack of any association to the site

configuration along the gradient. These are identified as a group within the marked boundary of Figure 4.

The distributions of Acacia modesta, Dodonaea viscosa, Erianthus griffithii, Gymnosporia royleana, Olea caspidata,

and Reptonia buxifolia are particularly noteworthy. Among these species G. royleana and O. cuspidata show

marked increases in frequency from plateau to lowland communities (Table 1). While A. modesta is able to

564 Journal of the Botanical R h Institute of Texas 1(1)

205.007] 8.

182.227]

159.447

— 1.

136.67 6.

113.8977 l 18.

91.11 7

19.

68.33 7

45.56 —

0.00 — 12.

| | | | | | | | | | | |

0.0 24.7 494 74.1 98.91 123.64 148.36 173.09 197.82 222.55 24727 272.00

Fic. 3. Sites biplot of detrended pond lysis (DCA) 1 and 2 for the qualitative vegetation data obtained from Pothowar Plateau. The

dictrihiutions nf 2 cit | tha NCA

maintain relatively constant frequencies in all the sites, D. viscosa, E. griffithii, and R. buxifolia showed a pat-

tern of increasing frequency of occurrence when extended to limestone plateau sites. In fact these species

are absent from sandstone massif.

DISCUSSION

Plant assemblage and geomorphology

The results indicate that the landscape, nature of the rock and redistribution of rainfall water by run-off

are the main sources of spatial variation in the study area. These geomorphological factors determine the

boundaries and the composition of the plant communities. The species Chrysopogon aucheri, Cyperus niveus,

Dicanthium annulatum, Grewia damine, and Sageretia theezans in Association E (Table 1) are most common in

the flat or rolling lowlands that receive sufficient runoff water, and virtually absent from plateau sites. The

species Boerhaavia coccinea, Carissa opaca, Ehretia obtusifolia, Evovulus alsinoides, and Olea caspidata reached

maximum abundance in valley bottom sites and showed declining frequency of occurrence when extended

into the plant communities on shallower soils of plateau sites (A-D). This pattern of distribution provides

strong circumstantial evidence that dip and scarp slopes (topography) determine the differences in vegeta-

tion types through the distribution of run-off generated by rainwater. Depth and moisture of the soil again

are important factors that exert influence on plant assemblages. On scarp slopes, the soil is shallow and

quickly dries out representing the more xeric habitat as indicated by DCA.

In upper plateau sites, the boundaries and composition of plant communities are determined by a

Dasti et al., Vegetation of Pothowar Plateau, Pakistan 565

454 —

Dichanthium

h F

Linum

320.32 — Acacia

Rhamnus

Dodonaea

Periploca

253.33 — Asparagus

Myrsine Desmostachya Cynodon

186.44 7 Cymb Acaci icio d

come ym epee cacia -Plantago

l Reptonia Gymnosporia

Zizyphus !

119.56 7 , , Olea Sageretia

Erianthus Zizyphus

= Chrysopogon

92.67 . Grewia

Anagalis

-14.22 ;

Boerhaavia Prosopis Medicago . l

C Oxalis Grewia

-81.11 — epee Gnaphalium

Rhazya

Erodium Taraxacum

-108 —

| | | | | | | | | |

-230 -84 -39 6.36 1.8 97.2 142.7 188.1 233.6 279 324

Fic AR l+ f T lar J liete (INCA\ L L TO O £4 A AL

le u

complex of factors including moisture, geology, soil and adjacent topography (Champion et al. 1960, 1965).

The most influential factor in distribution of the vegetation appeared to be the geological substrate. For

example, the species Acacia nilotica, Asparagus gracilis, Cymbopogon jawarencusa, Dodonaea viscosa, Erodium

cicutarium, Eremopogon foveolatus, Erianthus griffithii, Gnaphalium pulvinatum, Olea caspidata, Prosopis juliflora,

Reptonea buxifolia, and Ziziphus jujuba are associated with limestone strata (Association A). These species

are altogether absent from the eroded sandstone plant communities (Association D). A fairly high available

calcium status is possibly associated with the frequent occurrence of these species. Such species may be

considered as lime tolerant (Hussain 1969). Besides these differences in species composition, species abun-

dance also varies between the lime and sandstone sediments (Table 1). The majority of the plant species

that are distributed over calcareous or non-calcareous rocks of the mountain complex showed a pattern of

increasing frequency of occurrence when extended to sandstone plateau. This trend is common in Acacia

modesta, Capparis decidua, Grewia hirsuta, G. oppositifolia, G. tenax, Periploca aphylla, Rhamnus pentapomica,

Rhazia stricta, and Ziziphus nummularia.

Enclosed depressions in the limestone plateau form a marked transitional zone where the soil is mixed

with rock fragments. Clay minerals of limestone-derived soils are largely inherited from the parent material.

566 Journal of the Botanical R h Institute of Texas 1(1)

TABLE 2. Spearman rank correlation coefficients between the detrended correspondence analysis (DCA) first and second

axes.

Axes

Soil Parameters 1 2

Sand -0.609** 0.089

Silt -0.618** 0.161

Clay 0.744*** 20,176

ES 0313 0.393

pH -0.543* 0.041

Organic matter -0.407* -0.456*

Water holding capacity 0.075 -0.619**

Moisture content 0.425* eile

Level of significance: * P<0.05, ** P< 0.01, *** P«0.001.

The soils in the enclosed depressions remain exposed to continuous leaching, physical erosion and physico-

chemical changes without any rule by which a precise pattern of plant assemblage can be determined.

However, Acacia modesta, Gymnosporia royleana, Prosopis juliflora, Trachynia distachya and other species are

more frequent in depressions over limestone massifs while Acacia modesta, Capparis decidua, Erodium cicu-

tarium, Justicia peploides, and Prosopis juliflora are frequent on sandstone massif (Association C). This may be

interpreted as the variation in floristic composition on these strata that commonly reflects variations in the

substrate which in-turn depends on the nature of the parent materials from which the soil is derived.

The vegetation environmental correlation

Apart from the fact that the landscape variables are indeed relevant for explaining the main vegetation types,

the correspondence between the results of cluster analyses and DCA planes permit a direct interpretation

of scores of stand data in DCA plane in relation to soil variables. The five associations produced by cluster

analysis are plotted on first two axes as a scattered diagram (Fig. 5). The ordination axes may represent in

some way the major substrate influences, which affects the stands in these data, and have been used as the

plant and soil characteristics of the associations to discuss the most significant features of the environment

(Table 2).

The importance of water holding capacity and organic matter as the environmental factors affecting

plant species associations is not surprising, but has close relationship with water absorption and its reten-

tion. The present analyses and assessment of pattern and zonation along the first ordination axis suggest

that the most important environmental gradient and boundaries across the landscape are associated with

organic matter. The distribution of species along ordination axis 2 is significantly related not only to organic

matter, but also to water holding capacity. However, it is difficult to assess the relative importance of these

factors, but consistent negative interaction terms between these factors suggest that some combined effect

is important. The reasons for these correlation must be hydrological, particularly the pattern of run-off

generation and redistribution of organic matter of the study area, and probably are common to many arid

mountain terrains (Dasti & Malik 1998).

Vegetation is a complex collection of substrate specialists and generalists. It may be concluded that

both classification and ordination are able to delimit the plant associations according to their environments.

Topographic heterogeneity at the local scale also is an important factor that governs the community struc-

ture in plateau habitat. The complex gradients in edaphic conditions associated with topography provide

an opportunity to conduct further research, both in the laboratory and the field.

Dasti et al., Vegetation of Pothowar Plateau, Pakistan 567

203.00 7

182.227]

159.447

136.677

113.897

91.11 7

68.33 7

45.56 —

22.18: 1

0.00 —-j

| | | | | | | | | | | |

0.0 24.7 49.4 74.1 98.9 123 148 173 197 222.5 247 272

Fic 5 D I+. Yan Par | J eli n Lal +. J} +l S xt J 1L I 4 I

F u allaly

ACKNOWLEDGMENTS

The manuscript was prepared when Mohammad Athar (Tariq) was a visiting professor at the Institute of

Pure and Applied Biology, Bahauddin Zakariya University, Multan under the foreign faculty hiring program

of the Higher Education Commission, Islamabad. The authors are grateful to Joseph H. Kirkbride, Jr. and

two anonymous reviewers for providing valuable suggestions for improving the manuscript.

REFERENCES

ATHAR M. 2005. Nodulation of native legumes in Pakistani rangelands. Agric. Conspectus Scientificus, 70:49-

54.

Bec, A.R. 1975. Wildlife habitats of Pakistan. Bull. No. 5. Pakistan Forest Institute, Peshawar.

Birks, H.J.B. 1992. Some reflections on the application of numerical methods in quaternary palaeoecology. Publ.

Karelian Inst. 102:7-20.

CAUSTON, D.R. 1988. Introduction to vegetation analysis. Allen and Unwin, London.

568 Journal of the Botanical R h Institute of Texas 1(1)

CHAMPION, H.G. and S.K. SEHI. 1968. A revised survey of the forest types of India. Government Manager of Publica-

tions, Dehli.

CHAMPION, H.G., S.K. SETHI, and G.M. Khattak. 1960. Forest types of Pakistan. Pakistan Forest Institute, Peshawar.

CHAMPION, H.G., S.K. SETHI, and G.M. Khattak. 1965. Forest types of Pakistan. Pakistan Forest Institute, Peshawar.

CHAUDHURI, 1.1. 1960. The vegetation of Kaghan Valley. Pakistan J. Forest. 10:285-294.

Darcie, T.C.D. and M.A. EL DEMERDASH. 1991. A quantitative study of vegetation-environmental relationship in two

Egyptian deserts. J. Veg. Sci. 2:3-10.

Dast, A.A. and A.D.Q. Acnew. 1994. The vegetation of Cholistan and Thal desert. J. Arid Environ. 27:193-208.

Dast, A.A. and S.A. Maux. 1998. A transect of vegetations and soil on the Indus valley scarp slope. Pakistan J.

Plant Sci. 4:73-84.

Gupta, R.K. 1975. Plant life in Thar. In: R.K. Gupta and |. Prakash, eds. Environmental Analysis of the Thar Desert.

English Book Depot, Dehra Dun. Pp. 202-236.

Hacker, R.B. 1983. Use of reciprocal averaging ordination for the study of range condition gradients. J. Range

Manage. 36:25-30.

Hitt, M.O. 1979. DECORANA - FORTRAN program for detrended correspondence analysis and reciprocal averag-

ing. Ecology and systematic, Cornell Univ., Ithaca, New York.

Hitt, M.O. and H.G. GoucH 1980. Detrended correspondence analysis: an improved ordination technique. Veg-

etation 42:47-58.

Hussa, F. 1989. Field and laboratory manual of plant ecology. Univ. Grants Commission, Islamabad.

Hussan, F. and H. ILaHI. 1991. Ecology and vegetation of lesser Himalayas, Pakistan. Department of Botany, Univ.

of Pehawar, Pakistan.

Hussain, S.S. 1969 Vegetation survey of Ayub National Park, Rawalpindi. Pakistan J. Forest. 19:339-348.

JACKSON, A.B. 1958. Soil chemical analysis. Pretnice-Hall, Englewood Cliffs, New Jersey.

JONGMAN, R.H.G., C.J.F. TER BRAAK, and D.F.R. van TONGEREN. 1995. Data analysis in community and landscape ecology.

Cambridge Univ. Press, Cambridge.

Mauk, S.A, M.H Bokhari, A.A. Dast, and A. HANAN. 1988. Phyto-ecological studies of sand dune vegetation. In:

Modern Trends of Plant Science Research in Pakistan. Univ. of Peshawar, Peshawar. Pp. 29-34.

Naavi, H.H. 1974. Some observations on the vegetation of Peshawar and Khyber Pass, Pakistan. Biologia

20:151-162.

ODGAARD, R.V. and P. Rasmussen. 2000. Origin and temporal development of macro-scale vegetation pattern in the

cultural landscape of Denmark. J. Ecol. 88:733-748.

Rari, M. 1973. Vegetation types of Quetta-Kalat regions. Punjab Forest Department, Lahore.

RICHARDS, L.A. 1954. Diagnosis and improvement of saline and alkali soils. Agriculture Handbook No. 60, Depart-

ment of Agriculture & JBH Publishing, Bombay.

SHAUKAT, S.S. and M. AHMAD. 1989. A comparison of principal component and factor analysis as ordination model

with reference to desert ecosystem. Coenoses 4:15-28.

SHAUKAT, S.S., M.A. KHAIRI, and R. AHMAD. 1976. A phyto-sociological study of Garden area, southern Sind, Pakistan.

Pakistan J. Bot. 8:133-146.

SHAUKAT, S.S., M.A. KHAIRI, D. KHAN, and J.A. QuresHi. 1990. Multivariate approaches to the analysis of the vegetation

of Gadap area, southern Sind. Trop. Ecol. 21:61-102.

TER BRAAK, C.J.F. 1987. CANOCO—a FORTRAN program for (canonical) correspondence analysis, principal com-

ponents analysis and redundancy analysis (version 201). TNO, Eageningen.

SEED DISPERSALAND SOIL SEED BANK OF SERIPHIDIUM- OUBTTENSEB

(ASTERACEAE) IN HIGHLAND BALOCHISTAN, PAKISTAN

Sarfraz Ahmad Shamim Gul

Arid Zone Research Center Department of Botany

Pakistan Agricultural Research Council University of Balochistan

Quetta, PAKISTAN Quetta, PAKISTAN

Muhammad Islam Mohammad Athar!

Arid Zone Research Center California Dept. Food &Agric.

Pakistan Agricultural Research Council 1220 N Street, Rm 325

Quetta, PAKISTAN Sacramento, California 95814, U.S.A.

ABSTRACT

1 ters

Seed dispersal and soil seed | f Seriphidium quettense were studied over two years in highland Balochistan. Seed dispersal

started in early winter and continued till early spring. Primary dispersal agents were wind and water. Dispersal unit was capitulum,

most often without outermost whirl of phyllaries. There was generally one achene per capitulum but occasionally two to three achenes

may also be present. Mean dispersal distance was 23 to 40 cm from parent plant to ground surface. Majority of seeds (59-65%) fell

under canopy of parent plants. Sampling of soil for seed bank was done before and after seed dispersal of S. quettense. Soil samples

were collected from two depths (0—2 cm, 2-4 cm) and from two microhabitats (under adult individuals and open interspaces) by using

soil cores. Seed bank status was determined by observing S. quettense seedling emergence in the pots. Seriphidium quettense exhibited

persistent soil seed bank attribute. High seed densities were found under canopy of adult individuals and in upper soil layer. Persistent

soil seed bank may help seeds to occupy safe sites as they develop over time.

Key Wonps: Seriphidium quettense, seed bank, seed dispersal

RESUMEN

Se estudió la dispersión de las emillas y el banco de semillas del suelo de Seriphidium quettense durante dos afios en las tierras altas

de Balochistan. La dispersión de semillas comenzó a principios del invierno y continuó hasta principios de primavera. Los agentes

dispersantes primarios son el viento y el agua. La unidad de dispersión es el capítulo, la mayoría de las veces sin los filarios externos.

Normalmente hay un aquenio por capítulo pero ocasionalmente pueden estar presentes ods o tres. La distancia media de dispersión fue

de 23 a 40 cm desde la planta madre hasta la superficie del suelo. La mayoría de las semillas (59-65%) cayó bajo el dosel de las plantas

madres. El muestreo del suelo para el banco de semillas se realizó antes y después de la dispersión de las semillas de S. quettense. Se

colectaron muestras de suelo a dos profundidades (0—2 cm, 2-4 cm) y de dos microhábitats (bajo individuos adultos y en los espacios

abiertos entre ellos) mediante núcleos de suelo. El estatus del banco de semillas se determinó observando la la aparición de plántulas

de S. quettense en los puntos de muestreo. Seriphidium quettense muestra una persistencia de su banco de semillas. Se encontraron altas

densidades de semillas bajo el dosel de los individuos adultos y el la capa alta del suelo. La persistencia del banco de semillas del suelo

puede ayudar a las semillas a ocupar lugares seguros y desarrollarse tiempo después.

INTRODUCTION

About 93% of the total area of Balochistan is classified as rangelands and 90—9596 feed requirements of sheep

are met by these rangelands (Quraishi et al. 1993). About 87% of people of Balochistan derive their liveli-

hood directly or indirectly from livestock rearing (Heymell 1989). However, rangelands of Balochistan are

degrading very rapidly by overgrazing and removal of vegetation for fuel wood. Re-establishment of native

plant species is one of the options to restore the productivity of degraded rangelands.

In arid and semiarid rangelands, re-establishment of native plant species is vital to maintain function,

structure, diversity, and stability of the landscape. Native species have evolved under the prevailing stresses

of the region and have the ability to exploit the limited available resources. Seriphidium and Artemisia species

are the dominant dwarf shrubs of Balochistan. The genus Seriphidium comprises of 125 species distributed

! Corresponding author: atariq@cdfa.ca.gov

J. Bot. Res. Inst. Texas 1(1): 569 — 575. 2007

570 Journal of the Botanical R h Institute of Texas 1(1)

in North America, temperate Asia and Europe (Ghafoor 2002). Seriphidium quettense (Podlech) Ling. (Syn.

Artemisia quettensis Podlech) provides forage for small ruminants when other range species are in short sup-

ply particularly under drought conditions. Shrubs provide many benefits to humans and animals including

fodder for livestock and wildlife, functioning of rangeland ecosystems, erosion control, industrial products

and medicines (McKell 1989).

Regeneration of most range species depends on seed production, seed dispersal, seed predation, soil

seed bank, suitable environmental conditions, and presence of suitable microsites (Chambers & MacMahon

1994; Watkinson 1978). Seed dispersal can determine size, structure and composition of plant population

and diversity of plant community (Askew et al. 1997; Peart 1979). Soil seed bank enables plant populations

to maintain their genetic variability, withstand adverse periods, and persist through time (Baskin & Baskin

1978; Thompson & Grime 1979). Soil seed bank have both horizontal and vertical dimensions as a result of

phase I and phase II dispersal (Chambers & MacMahon 1994; Guo et al. 1999; Simpson et al. 1989). Such

distribution of seeds has a great impact on population growth and its structure (Chambers & MacMahon

1994). Seeds are present in high densities under plant canopies in arid habitats representing horizontal

pattern of seed banks in soil (Guo et al. 1999). The present study was conducted to determine the pattern

of seed dispersal of Seriphidium quettense and evaluate temporal and spatial status of its soil seed bank.

MATERIALS AND METHODS

Study site

The experiment was conducted in Chiltan National Park Hazarganji, Balochistan, Pakistan in an extended

area of the park that was protected from livestock grazing since 1998. Climate of the area is Mediterra-

nean type. Área receives rainfall mostly during the winter months. Occasionally rainfall occurs during the

summer as well. Mean annual rainfall of over 19 years is 200 mm. Soil of the area is sandy-to-sandy loam.

Dominant species of the site are Seriphidium quettense, Cymbopogon jawarancusa, Chrysopogon aucheri and

annual grasses (Marwat et al. 1992).

Seed dispersal

Experiments were conducted for two-year to determine the pattern of seed dispersal in S. quettense. In As-

teraceae (Compositae) the dispersing unit is a fruit, an achene or cypsela. However, the term seed is used

in this study for the fruit dispersal. Fifty meters long transects were established during December 2002

and December 2003. Ten points were marked along the transect during each year and the nearest plant

was selected on each point. Inflorescences of other plants within 3 meters radius were clipped to prevent

confounding of the seeds. Fluorescent powder was applied to the selected plants prior to seed dispersal. Seed

dispersal was observed at five days interval until the end of dispersal period. Distance of seed dispersal was

measured from the edge of the base of the marked plants. Seeds were collected after measuring the distance.

Percentage of dispersed seeds was determined in two microhabitats under parent plant canopy and open

interspaces. The data were subjected to linear regression analysis between number of seeds dispersed and

dispersal distance.

Soil seed bank

Temporal and spatial status of S. quettense seed bank in the soil was examined for two years from two depths

(0-2 cm, 2-4 cm) and from two microhabitats (under canopy of adult individuals, and open interspaces).

Samples were collected before and after fresh seed dispersal to assess temporal aspect of soil seed bank. Fifty

meter long transects were established at each sampling date. Ten points were marked on transect lines. Five

samples were collected under adult individuals and from open interspaces at 0—2 cm and 2-4 cm depths at

each marked point by using 10 cm diameter soil core. First sampling before the seed dispersal period was

done in mid November during 2002 and 2003. Second sampling was done in late February during 2003 and

2004. As seeds of S. quettense are 0.8 mm in size (Ghafoor 2002), 0.5 mm mesh sieve was used to remove

gravel and unnecessary soil, to ensure maximum retention of seeds in the sample for their germination. The

sub samples were pooled and filled in pots for observing seedling emergence. Pots were placed in open air

Ahmad et al., Seed bank of Seriphidium quettense 571

70 -

- 604

m oO

2 404

2 30-

Z 20-

10 -

2003 2004

Years

Fic. 1. Percentage of dis ] seeds (capitula) under | t plant pies (2) and in open inters (E) during 2003 and 2004

and watered regularly. The emerging seedlings were identified, counted and removed. Seedling emergence

was observed for three months. First year samplings were germinated in mid March and the second year

samplings in the last week of February.

RESULTS

Seed dispersal

Seed dispersal of S. quettense started from mid to late December and continued till late February. Primary

dispersal agents were wind and water. Dispersal unit was a capitulum, most often without outermost whorl

of phyllaries. There was generally one achene per capitulum but occasionally two to three achenes may also

be present. Maximum dispersal distance was 130 cm and 210 cm for first and second year respectively. The

mean dispersal distance from parent plants to ground surface was 23 to 40 cm for first and second year

respectively. Seeds distributed in all directions around the plants. However, majority of seeds were found

in northeast side according to wind direction. Majority of the seeds (59-65%) fell under canopy of parent

plants (Fig. 1). There was a negative correlation between dispersal distance and number of the seeds (Fig.

2a, b). Number of the seeds decreased as the distance increased and created a negative exponential curve

(Fig. 2a, b).

Soil seed bank

Seed densities of S. quettense varied with microhabitats and soil depths for each sampling date (Fig. 3a,b,c,d).

High seed densities were found in the upper 0-2 cm soil layer than the lower 2-4 cm and under the adult

individuals than in the open interspaces. In the first year sampling, microhabitats and depths were signifi-

cant (p<0.05) for the second sampling while non-significant for the first sampling (Fig. 3a,b). In second

year samplings, differences in numbers of seedlings between microhabitats were significant (p«0.05) for

both samplings as a higher number of seeds were observed under canopy of adult individuals. Differences

between soil depths were non-significant for both samplings (Fig. 3c,d). A high number of seeds were found

in upper soil layer.

572 Journal of the Botanical R h Institute of Texas 1(1)

s y = -2.9228x 292.44

400 +- R? = 0.6521

Number of dispersed seeds

Distance (cm)

900 4

n %.

9 7504 » :

D R = 0.3975

2 6004

Sum

= 450 1e

E 3004

5 1504

0 4 e"

0 50 100 150 200 250

Distance (cm)

Fic. 2. Patt £ 1 4: ie Cc iphidi ] Lf £. I t pl fed +L g KI £. J H gy 2003 (a) 1 2004 (b). Li

regression equations, where Y is number of d

DISCUSSION

Seriphidium quettense had a prolonged period of seed dispersal which started from early winter and lasted

till early spring. Prolonged dispersal has also been reported for fringed sagebrush (Bai & Romo 1997).

Although predation on S. quettense seeds is unknown, but prolonged dispersal is believed to be helpful in

avoiding predation (Willson 1993). Longer periods of seed dispersal in S. quettense seems to be a tactic for

contemporary seed germination as well as for the chances of seed burial in the soil (Davlaeminck et al. 2005;

Mahmood et al. 2005; Olano et al. 2002). Though seeds of S. quettense are very light, dispersal was limited

with a majority of seeds falling under the canopy of adult plants. Restricted dispersal of seeds around the

Ahmad et al., Seed bank of S

+ First Sampling a

3 e a a

E E 4.0 4

S 8 307 a a

z 300:

s 20 — —

> Canopy Interspace 0-2 cm 2-4 cm

Microhabitat Depth

Year 2003

Second Sampling b

5. 20 a

g E 154

A EE : :

8*5 3. mamn -

> Canopy Interspace 0-2 cm 2-4 cm

Microhabitat Depth

Year 2003

First sampling c

ES = a

E ES 40 “|

2 Y

g 207

S A

> i

Canopy Interspace

Microhabitat

Year 2004

Second sampling d

[Sun

(D)

> Canopy Interspace 0-2 cm 2-4 cm

Microhabitat Depth

Year 2004

Fic 3 T | I J [ e: f 44 £ HI Y) L RA LA] y £: ££. DANAC

573

574 Journal of the Botanical R h Institute of Texas 1(1)

parent plants signifies safe sites for seed germination (Davlaeminck et al. 2005; Mahmood et al. 2005).

Characteristics like seed weight, size, shape, height of parent plants, and the nature of surrounding vegeta-

tion are important determinants when wind and gravity are the primary agents for seed dispersal (Willson

1993). The limited seed dispersal has also been reported for other species of Artemisia (Bai & Romo 1997;

Bauer et al. 2002; Friedman & Orshan 1975) and has been attributed to low release height of the parent

plants (Bauer et al. 2002). Phase II dispersal of seeds in horizontal and vertical direction may have greater

influence on patterning of plants in ecosystems than their initial dispersal (Chambers & MacMahon 1994;

Russel & Schupp 1998). There are greater chances of rain in the area during January and February that may

help farther movement of the seeds and can greatly affect seed germination and seedling establishment.

Densities of emerged seedlings of S. quettense in pots ranged from 3 to 68 per m". Seed densities of

fringed sagebrush in mixed prairie in Alberta have been reported as 75 to 183 per m? (Johnston et al. 1969).

In second year, seedling densities were higher due to variation in sampling site and early placement of samples

in the pots than the first year. Increasing temperature may hinder germination of these species (Bai & Romo

1994; Booth & Bai 1999). Roberts (1981) reported that use of seedling emergence techniques to quantify

soil seed bank status may under estimate the soil seed bank density if germination requirements are not

met. Soil seed bank of S. quettense showed variable spatial and temporal patterns across different samplings.

Both increasing and decreasing trends have been reported for total number of seeds of S. tridentatum with

distance from canopy center (Guo 1998). Canopies of S. quettense retained more seeds than open interspaces

in all samplings. It may be attributed to the limited dispersal of the seeds and comparatively high quantity

of litter under canopies as compared to bare interspaces (Russell & Schupp 1998).

Higher seed densities were found in upper soil layer (Guo et al. 1999). Small seeds are more prone

to move downward in loose textured soils in arid environment as compared to larger ones (Chambers &

MacMahon 1994). The period of seed dispersal of S. quettense immediately follows spring season. That can

increase germination due to greater chances of rainfall during late winter or early spring.

Seriphidium quettense has a persistent soil seed bank and seedlings emerged from first samplings before

the seed dispersal period. Our findings are in conformity with previous studies that small seeds and seeds

of disturbed habitats generally form persistent soil seed bank (Bai et al. 1995; Bai & Romo 1997; Mahmood

et al. 2005). Prolonged seed dispersal and persistent soil seed bank attributes of S. quettense may help their

seeds to occupy safe sites as they develop over time.

ACKNOWLEDGMENTS

The study was funded by a research grant from Pakistan Science Foundation, Islamabad which is gratefully

acknowledged. Sincere thanks are due to Safdar Ali Kayani, Professor of Botany Department, University

of Balochistan, Quetta for valuable suggestions and technical assistance during this research. The authors

thank Sarwat N. Mirza (Univ. of Arid Agric., Rawalpinidi, Pakistan) and A. Mahmood (KUH) for helpful

suggestions to improve the manuscript.

REFERENCES

Askew, A.P, D. Corker, D.J. Hopkinson, and K. THomson. 1997. A new apparatus to measure the rate of fall of seeds.

Funct. Ecol. 11:121-125.

Bal, Y. and J.T. Romo. 1994. Germination of previously buried seeds of fringed sage (Artemisia frigida). Weed Sci.

42:390-397.

Bal, Y. and J.T. Romo. 1997. Seed production, seed rain and the seed bank of fringed sagebrush. J. Range Manage.

50:151-155.

BA, Y., J.T. Romo, and J.A. Youne. 1995. Influences of temperature, light, and water stress on germination of fringed

sage (Artemisia frigida). J. Range Manage. 43:219-225.

Baskin, J.M. and C.C. Baskin. 1978. The seed bank in a population of an endemic plant species and its ecological

significance. Biol. Conserv. 14:125-130.

Ahmad et al., Seed bank of S iphidi juett 575

Bauer, K.M, E.L. BerLow, and C.M. D'Antonio. 2002. The relationship between climate and Rothrock sagebrush

colonization patterns. J. Range Manage. 55:620-625.

Boot, D.T. and Y. Bal. 1999. Imbibition temperature affects on seedling vigor in crops and shrubs. J. Range Man-

age. 52:534-538.

CHAMBERS, J.E. and J.A. MacMahon. 1994. A day in the life of a seed: movements and fates of seeds and their im-

plications for natural and managed systems. Annu. Rev. Ecol. Syst. 25:263-292.

DEVLAEMINCK, R., B. Bossuvr, and M. Hermy. 2005. Seed dispersal from a forest into adjacent cropland. Agric. Ecosyst.

Environ. 107:57—64.

FRIEDMAN, J. and G. OrsHAN. 1975. The distribution, emergence and survival of seedlings of Artemisia herba in the

Negev desert of Israel in relation to distance from the adult plant. Ecology 63:627-632.

GHAFOOR, A. 2002. Asteraceae (1)-Anthemideae. In: S.I. Ali and M. Qaiser, eds. Flora of Pakistan. Department of

Botany, University of Karachi, Karachi.

Grime, J.P. 1979. Plants strategies and vegetation. John Wiley & Son, Chichester, UK.

Guo, Q. 1998. Microhabitat differentiations in Chihuanuan desert plant communities. Ecology 139:71-80.

Guo, Q., PW. RuNbEL, and D.W. GoobALL. 1999. Structure of desert seed banks: comparisons across four North

American desert sites. J. Arid Environ. 42:1-14.

HevmeLL, V. 1989. Evaluation of pasture conditions in Nichara Union Council in relation to construction of water

storage tanks. Studies 09/89. Pak-German self- help project for rural development, Balochistan, Quetta,

Pakistan.

JOHNSTON, A., S. Suouak, and PW. STRINGER. 1969. Viable seed population in Alberta Prairie top soils. Canad. J. Plant

Sci. 49:75-82.

MAHMOOD, S., M. ATHAR, and A.D.Q. Acnew. 2005. Soil seed banks and dispersal patterns of sea rush, Juncus maritimus

in spatially and temporally disturbed coastal habitats. Phytologia 87:225-226.

Marwar, Q.U.D., M. Nisar, and F. Hussain. 1992. Vegetation studies of Chiltan National Park Hazarganji, Quetta.

Pakistan J. Agric. Res. 13:71-79.

McKett, C.M. 1989. The biology and utilization of shrubs. Academic Press, San Diego, California.

OLANO, J.M., |. CALLALLERO, N.A. LASKURIAN, J. Loin, and A. Escupero. 2002. Seed bank spatial pattern in temperate

secondary forest. J. Veg. Sci. 17:775—784.

Peart, M.H. 1979. Experiments on the biological significance of the morphology and seed-dispersal units in

grasses. Ecology 67:843-863.

QuralsHi, M.A.A., G.S. KHAN, and M.S. Yaqoos. 1993. Range management in Pakistan. Qazi Publications, Lahore.

RoserTs, H.A. 1981. Seed banks in the soil. Adv. Appl. Biol. 6:1-55.

RUSSELL, S.K. and E.W. ScHupp. 1998. Effects of microhabitat patchiness on patterns of seed dispersal and seed

predation of Carcocoarpus ledifolius (Rosaceae). Oikos 81:434-443.

SIMPSON, R.L., M.A. Leck, and V.T. Parker. 1989. Seed banks: general aspects and methodological issues. In: M.A. Leck,

V.T. Parker, and R.L. Simpson, eds. Ecology of soil seed banks. Academic Press, San Diego. Pp. 3-8.

Thompson, K. and J.P. Grime. 1979. Seasonal variation in the seed banks of herbaceous species in ten contrasting

habitats. Ecology 67:893-921.

WATKINSON, A.R. 1978. The demography of a sand dune annual Vulpia fasciculati. |l. The dispersal of seeds. J. Ecol.

66:483-498.

WiLLSON, M.F. 1993. Dispersal mode, seed shadows and colonization patterns. Vegetatio 107/108:261-280.

576 Journal of the Botanical R h Institute of Texas 1(1)

BOOK NOTICES

Bui Witt. 2006. Orchids In Your Pocket: A Guide to the Native Orchids of Iowa. (ISBN 1-58729-499-0,

laminated fold-out guide). University of Iowa Press, 100 Kuhl House, Iowa City, IA 52242-1000, U.S.A.

(Orders: www.uipress.uiowa.edu, 319-335-2000, 319-335-2055 fax, 1-800-621-2736). $9.95, 437

pp., 32 color photos, 444" x 9".

A plastic-laminated, fold-up pocket guide (9 x 4 inches) with color photos of thirty-one species of orchids plus two hybrids. Common

and scientific names, habitat and distribution, height, phenology, and rarity status are given for each species. Eighteen of the species are

listed as threatened, endangered, or of special concern.—Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth,

TX 76102-4060, U.S.A.

Jonn TuLLock. 2005. Growing Hardy Orchids. (ISBN 0-88192-715-5, hbk.). Timber Press Inc., The Hasel-

tine Building, 133 SW Second Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: www.

timberpress.com, 503-227-2878, 503-227-3070 fax, 1-800-327-5680). $29.95, 244 pp., 100 color

illustrations, 6" x 9".

“Part of the reason I first ł i lin growing i hids st

d directly from their reputation for cultivation.

In no reference book could I find any encouragement for including native orchids in my wildflower garden; in fact, most of the books

I consulted contained numerous exhortations to avoid even es to a them. ... I developed my orchid growing techniques

f } 1 E

thr ugh di tol t

thirty years of excursions

up and down the [Tennessee] valley and into ue mountains on either side a it. MASS Sen by the insights of other orchidists whose works

appear in the bibliography, I now grow [in Knoxville] twelve North American orchid species and three exotic ones." Although there

apparently is no explicit list of these species, the following are specifically mentioned in the text and photos as growing in his garden:

B ae striata, ie ae ca por nen D Goodyera pubescens, Platanthera ciliaris, Platanthera integrilabia, Platanthera flava,

Plat p] Sniranthe

nua vat. odorata, and Tipularia discolor

The eee is an ardent conservationist and notes the value of “bringing wild species into cultivation and developing effective

techniques for their large-scale propagation" and that transplants of species in immediate threat of the “bulldozer” are a valuable way of

conserving germ plasm. Among the appended resources is a list with addresses of orchid suppliers and organizations.

Chapters include: 1) Native orchid conservation: One view; 2) General principles of hardy orchid cultivation; 3) Propagation of

hardy orchids; 4) Mycorrhizal associations and hardy orchids; 5) Hardy orchids through the seasons; 6) Bletilla: The ideal beginner's

plant; 7) A catalog of hardy and half-hardy orchids.— Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX

76102-4060, U.S.A.

PauL Martin Brown (Text) AND STAN Fotsom (Original Artwork). 2006. Wild Orchids of the Prairies and

Great Plains Region of North America. (ISBN 0-8130-2975-9, flexibind). University Press of Florida,

15 Northwest 15th Street, Gainesville, FL 32611-2079, U.S.A. (Orders: www.upf.com, 352-392-1351,

352-392-7302 fax, 1-800-226-3822). $29.95, 376 pp., 316 color plates, 94 b/w illustrations, 81 maps,

The beauty and compact taxonomy of North American orchids has stimulated production of a plethora of regional identification guides

and accounts. This is a particularly fine one, plus it’s relatively inexpensive and easily of size to be carried around in the field. Included

are 21 genera, with 64 species, 9 varieties, and 8 hybrids, over a range from southern Saskatchewan and Manitoba south to the north-

eastern corner of New Mexico, the northern half of Texas, and the northwestern corner of Louisiana. The author notes that “recent work

in volume 26 of Flora of North America (2002) has greatly fine-tuned the identification and distribution of these orchids.”

The book is divided into four parts: (1) a brief discussion of prairies and orchids, an introduction to the format for Part 2, and keys

to the genera, (2) treatments of the genera and species, (3) references and resources, and (4) orchid hunting. Two aid give distribu-

tion by state for each species and a chart of flowering times. Part 2 has s to PUE within each g descriptions ea

detailed range maps, several color photos (and often line drawings) for ing habit “at floral details, and

the author’s personal experience on ecology, phenology, abundance, critical US of nce and hybrids. Formally described

color and growth forms are listed for each species. Part 3 has a checklist for the region, provincial and state species lists, *some regional

orchid statistics,” rare, threatened, and endangered species, synonyms and misapplied names, recent literature references for new taxa,

combinations, and additions, a commentary on use of Luer's 1975 compendium on orchids of the USA and Canada, and commentaries

Gwith comparative photos) on distinctions in 8 cases of cryptic species, species pairs, and varietal pairs. Part 4 give details suggestions

on where to look for orchids in each of seven subregions.— Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth,

IX 76102-4060, U.S.A.

J. Bot. Res. Inst. Texas 1(1): 576. 2007

VASCULAR PLANT PECI SAREA RELATIONSHIPS (SPECTES RICHNESS)

INTHE WEST GULF COASTAL PEAIN. AFIR T APPROXIMATION

Michael H. MacRoberts, Barbara R. MacRoberts Robert G. Kalinsky

Bog Research and Biology Department

740 Columbia Herbarium Louisiana State University-Shreveport

Shreveport, Louisiana 71104, U.S.A Museum of Life Sciences Shreveport, Louisiana 71115, U.S.A.

Louisiana State University-Shreveport

Shre\ eport, Louisiana 71115, U.S.A.

ABSTRACT

We surveyed the West Gulf Coastal Plain literature for information on vascular plant species/area relationships (species richness) and

produced curves for small (micro-) areas (< 5 ha) and large (macro-) areas (> 50 ha). A brief preliminary comparison of species richness

between the West Gulf Coastal Plain and the Atlantic and East Gulf Coastal Plain indicates that while some micro-scale areas may be

richer in species in the East Gulf Coastal Plain, macro-scale areas have about the same richness in both areas.

RESUMEN

Se revisó la bibliografía sobre la llanura costera del West Gulf en busca de inf ión sob ies de pl l /relación con

el área (riqueza de especies) y se hicieron curvas para micro-áreas (< 5 ha) y macro-áreas (> 50 ha). Una breve comparación preliminar

de la riqueza de especies entre la llanura costera del West Gulf y la llanura costera del Golfo Atlántico y del Este indica que mientras

algunas micro-áreas pueden ser mas ricas en especies en la llanura costera del East Gulf, las macro-áreas tienen aproximadamente la

misma riqueza en ambas áreas.

INTRODUCTION

We have noted that many papers seeking to compare an area's species richness find few relevant studies

with comparable information. For example, addressing the long-held belief that the Big Thicket is species

rich, Diggs et al. 2006:169) ask: *How does [the Big Thicket] compare with other areas of similar size?"

But, lacking “the needed comparative data,” they were unable to answer the question. Being interested in

species richness for some time and having collected considerable information on it for various communities

and locations in the West Gulf Coastal Plain over the past two decades (MacRoberts & MacRoberts 1992b),

we decided to survey the literature to see if we could locate enough data to draw a species/area curve for

the West Gulf Coastal Plain.

Whittaker (1972) defined alpha, beta, and gamma diversity (richness). In this paper, we will be concerned

with alpha diversity (within habitat or community, such as for a pitcher plant bog, prairie, or baygall) and

gamma diversity (geographic scale diversity, such as for a county or state, which include many communi-

ties and habitats). We will not use Whittaker's terms in this paper but will speak only of species richness to

cover both alpha and gamma diversity; all references to areas smaller than 5 ha in this paper indicate alpha

diversity and all references to areas larger than 50 ha indicate gamma diversity.

METHODS/RESULTS

We surveyed the West Gulf Coastal Plain—eastern Texas, western Louisiana, southeastern Oklahoma,

and southern Arkansas (Fig. 1)—literature for species richness information. Our criterion for accepting or

rejecting a study was its thoroughness. This unfortunately left out most county/parish checklists done as

Masters theses since few are thorough and virtually none are published. Nonetheless, we found data for both

large and small areas including many community types (Peterson & Brown 1983; Matos & Rudolph 1985;

MacRoberts € MacRoberts 1988,1990, 1991, 1992a, 1992b, 1993a, 1993b, 1993c, 1994, 1995a, 1995b,

1996a, 1996b, 1997, 1998a, 1998b, 2001, 2003, 2004a, 2004b, 2004c, 2005, 2006, in prep.; MacRoberts

J. Bot. Res. Inst. Texas 1(1): 577 — 583. 2007

578 Journal of the Botanical Research Institute of Texas 1(1)

F 1 M L e. + + LAAJ af, If. tal DIA:

IG. WEI UUI VUda»tadi r laid

et al. 2002a, 2002b, 2002c, 2003, 2004; Thomas & Allen 1993-1998; Nesom & Brown 1998; Diggs et al.

1999, 2006; Neill 2000; Fleming et al. 2002; Neyland 2002; Singhurst et al. 2003; Brown et al. 2005, 2006;

Philipps et al. in prep.; Thomas et al. in prep.) We plotted the data on a log/log scale. We did this separately

for areas smaller than 5 ha and for areas larger than 50 ha. The results are shown in Figures 2 and 3.

DISCUSSION

There is wide variation in number of taxa reported for the same and for different sized areas in the West

Gulf Coastal Plain. The reasons are many. Community differences account for some of the difference be-

AA D L a Fu | c a LM t bad 4L VAL ff" Lr 4 IDI x

dl., UIC WeSt QUIT COdStdl r idin 579

West Gulf Coastal Plain Species Area Curve

[Micro-areas — area « 5 ha]

2.5

W154

0 *

E :

5 i $

Z, | "

e *

Ar e

= .

0.5 -

R^ = 0.8031

0 T T T T T

-5 -4 -3 -2 -1 0 1

Log, )Area (ha)

Fic. 2. West Gulf Coastal Plain species/ (Micro-areas = area <5 ha). Open boxes are Big Thicket sites.

cause different communities have different species numbers. For example, baygalls have fewer species per

unit area than pitcher plant bogs (MacRoberts & MacRoberts 2001; MacRoberts et al. 2004). Thoroughness

of collecting differs considerably. Some areas have been collected extensively for years, while others have

not. For example, while Caddo Parish has been the subject of three checklists, few counties/parishes in the

West Gulf Coastal Plain, notably in Texas, have been the subject of one (MacRoberts & MacRoberts 2006).

Taxonomic preferences also affect numbers. Some investigators report species, subspecies, and varieties

while others report only species (MacRoberts & MacRoberts 2006). Finally, inclusion criteria differ. Some

investigators collect extensively in urban parks and lawns, transportation routes, and other highly disturbed

ruderal areas and report many more non-natives than do others.

Nonetheless, allowing for these sources of variation and potential error, Figures 2 and 3 allow for

general comparisons. For example, the question posed by Diggs et al. (2006) in our introductory remarks

is now answerable. According to Diggs et al. (2006) the Big Thicket consists of 17,974 sq km and has 1826

580 Journal of the Botanical R h Institute of Texas 1(1)

West Gulf Coastal Plain Species Area Curve

[Macro-areas = area > 50 ha]

" d

Q J

epum

Q d

un J

& 34

= ]

Z. E

= J

7 *

- R^ = 0.9105

2 La a a a a a TT] TT TTTTTT TT] TT TT TT TT TT] TT TT TT T TT T[ TT TT TT TT TT] T TT TT TT TT ] T TT TT TT TT

-] 0 l 2 3 4 5 6

Log;oyArea (km)

Fic. 3. West Gulf Coastal Plain sj / (Macro-areas = area > 50 ha). Open boxes are Big Thicket sites.

taxa. Located on the curve in Figure 3, the Big Thicket is therefore average. At micro-scale levels, although

the data are scanty, the Big Thicket is typical: two 3.0 ha wetland pine savannas had 106 and 117 taxa, and

four 0.0001 ha wetland pine savanna plots had 19, 20, 21, and 24 taxa (Fig. 2). At intermediate macro-scale

levels: 2.84 sq km, 8.81 sq km, 31.5 sq km, and 58.1 sq km, the Big Thicket also appears to be unexceptional

with 401, 544, 738, and 693 taxa respectively (Matos & Rudolph 1985; MacRoberts & MacRoberts 1998a;

MacRoberts et al. 2002c; Brown et al. 2005, 2006)(Fig. 3).

While the aim of this paper is to develop species richness curves for the West Gulf Coastal Plain, it is

almost impossible not to compare our figures with those farther east, since it has been suggested that the

West Gulf Coastal Plain may not be as rich as the Atlantic and East Gulf Coastal Plain (Currie & Paquin

1987; Currie 1991). For example, Fort Bragg Military Reservation, North Carolina, consists of 738 sq. km

and has an impressive 1206 taxa (Sorrie et al. 2006). Platt (1999) summarized some of the small area plots

in frequently burned pine savannas in the Atlantic Coastal Plain: up to 30-40 species in 0.0001 ha plots,

MacRoberts et al., Species richness in the West Gulf Coastal Plain 581

70—90 species in 0.01 ha plots, and 100-150 species in 0.1 ha plots, making these the most species rich

sites reported in North America (Walker & Peet 1983; Peet & Allard 1993; Brewer 1998).

However, state and county sized areas appear to be less spectacular: Florida, consisting of 170,305 sq

km, has 3834 species of which 3196 are exotics (Wunderlin 1998); the Florida panhandle, consisting of

38,628 sq km, has 2359 species of which 1696 are exotics (Clewell 1985); Oktibbeh County, Mississippi,

consisting of 1185 sq km, has 1148 species of which 1596 are exotics (Leidolf et al. 2002); and Pike County,

Alabama, consisting of 1759 sq km, has 1190 species of which 2096 are exotics (Diamond 2003). Comparable

areas in the West Gulf Coastal Plain appear to be as species rich. For example, east Texas, consisting of

162,134 sq km, has 3660 taxa (3402 species) of which 1896 are exotic (Diggs et al. 2006); the Gus Engeling

Wildlife Management Area in Anderson County, Texas, consisting of 45 sq km, has an impressive 930 taxa

(Singhurst et al. 2003); the Walter B. Jacobs Memorial Nature Park, Caddo Parish, Louisiana, consisting of

65 ha, has a staggering 587 taxa (Thomas et al. in prep); and Caddo Parish, Louisiana, consisting of 2,284

sq km, has an impressive 1405 species of which 1796 are exotic (MacRoberts & MacRoberts 2006). At the

micro-scale level, bogs, xeric sandylands, and upland longleaf pine savannas in the West Gulf Coastal Plain,

while not reaching the higher numbers reported east of the Mississippi River, have been reported with up

to 28 species in 0.0001 ha plots, 46 in 0.001 ha plots, and 113 in 0.1 ha plots (MacRoberts & MacRoberts

1993b, 2004c; MacRoberts et al. 2002b; Philipps et al. in prep).

These are preliminary comparisons. This paper is a first imation at a West Gulf Coastal Plain

LUE

species-area curve. It is based on a wide variety of sources, most of which were not prepared for the pur-

pose of measuring species richness. Additionally, the data are relatively few. More information is needed on

both micro- and macro-scale areas. Undoubtedly, further research, notably in frequently burned West Gulf

Coastal Plain open habitats such as bogs, pine savannas, upland longleaf forest/savanna, xeric sandylands,

and prairies, will alter these curves and make species-richness comparisons truly possible within the Gulf

Coastal Plain and between the Gulf Coastal Plain and areas to the east, north, and west.

ACKNOWLEDGMENTS

We thank James H. Peck and Stephen P. McLaughlin for their helpful reviews.

REFERENCES

Brewer, J.S. 1998. Patterns of plant species richness in a wet slash-pine (Pinus elliottii) savanna. J. Torrey Bot.

Society 125:216-224.

BROWN, L.E., B.R. MacRoserts, M.H. MacRoserts, PA. Harcomee, W.W. Pruess, I.S. Ersik, and D. JoHNson. 2005. Annotated

checklist of the vascular flora of the Turkey Creek Unit of the Big Thicket National Preserve, Tyler and Hardin

counties, Texas. Sida 21:1807-1827.

BROWN, L.E., B.R. MAcRoserts, M.H. MacRoserts, PA. HARcoMBE, W.W.Pruess, l.S. Esik, and S.D. Jones. 2006. Annotated

checklist of the vascular flora of the Big Sandy Creek Unit, Big Thicket National Preserve, Texas. Sida 22:

705-723.

CueweLL, A.F. 1985. Guide to the vascular plants of the Florida panhandle. Univ. Presses of Florida, Florida State

Univ. Press, Tallahassee.

Currie, D.J. 1991. Energy and large-scale patterns of animal- and plant-species riches. Amer. Naturalist 137:

27-49.

Currie, D.J. and V. Paquin. 1987. Large-scale biogeographical patterns of species richness of trees. Nature

329:326-327.

DiAMOND, A.R, 2003. A checklist of the vascular flora of Pike County, Alabama. Castanea 68:143-159.

Dicas, G.M., B.L. Liescome, and R.J. O'Kennon. 1999. Shinners & Mahler' illustrated flora of north central Texas. Sida,

Bot. Misc. 16:1-1626.

Dices, G.M., B.L. Lipscome, M.D. Reep, and RJ. O'Kennon. 2006. Illustrated flora of east Texas. Sida, Bot. Misc. 26:

1-1594.

582 Journal of the Botanical R h Institute of Texas 1(1)

FLEMING, K.M., J.R. SiINcHURST, and W.C. HoLmes. 2002. Vascular flora of Big Lake Bottom Wildlife Management Area,

Anderson County, Texas. Sida 20:355-371.

Leipote, A., S. McDanteL, and T. Nume. 2002. The flora of Oktibbeha County, Mississippi. Sida 20:691-765.

MacRoserts, B.R. and M.H. MacRostnrs. 1988. Floristic composition of two west Louisiana pitcher plant bogs.

Phytologia 65:184-190.

MacRostRrs, B.R. and M.H. MacRoserts. 1990. Vascular flora of two west Louisiana pitcher plant bogs. Phytologia

68:271-275.

MacRostRrs, B.R. and M.H. MacRoserts. 1991. Floristics of three bogs in western Louisiana. Phytologia 70:

135-141.

MacRoserts, M.H. and B.R. MacRoserts. 1992a. Floristics of a sandstone glade in western Louisiana. Phytologia

72:130-138.

MacRoserTs, B.R. and M.H. MacRoserts. 1992b. Floristics of four small bogs in western Louisiana with observations

on species/area relationships. Phytologia 73:49-56.

MacRostRrs, B.R. and M.H. MacRoserts. 1993a. Floristics of two Louisiana sandstone glades. Phytologia 74:

431-437.

MacRostnrs, B.R. and M.H. MacRoserts. 1993b. Floristics of a bog in Vernon Parish, Louisiana, with comments on

noteworthy plants in western Louisiana. Phytologia 75:247-258.

MacRogerTs, M.H. and B.R. MacRoserts. 1993c. Vascular flora of sandstone outcrop communities in western Loui-

siana, with notes of rare and noteworthy species. Phytologia 75:463-480.

MacRoserts, M.H. and B.R. MacRoserts. 1994. Floristics of a xeric sandyland in western Louisiana. Phytologia

77:414-424.

MacRostnrs, B.R. and M.H. MacRoserts. 1995a. Vascular flora of two calcareous prairie remnants on the Kisatchie

National Forest, Louisiana. Phytologia 78:18-27.

MacRoserts, B.R. and M.H. MacRostnrs. 1995b. Floristics of xeric sandhills in norhwestern Louisiana. Phytologia

79:123-131.

MacRoserrs, B.R. and M.H. MacRoserts. 1996a Floristics of xeric sandhills in eastern Texas. Phytologia 80:1-7.

MacRoserts, B.R. and M.H. MacRoserts. 1996b. The floristics of calcareous prairies on the Kisatchie National Forest,

Louisiana. Phytologia 81:35-43.

MacRoserts, B.R. and M.H. MacRostnrs. 1997. Floristics of beech-hardwood forest in east Texas. Phytologia

82:20-29.

MacRostnrs, B.R. and M.H. MacRoberTs. 1998a. Floristics of wetland pine savannas in the Big Thicket National

Preserve, southeast Texas. Phytologia 85:40-50.

MacRoberts, B.R. and M.H. MacRostnrs. 1998b. Floristics of muck bogs in east central Texas. Phytologia 85:

61-73.

MacRoserrs, B.R. and M.H. MacRoserts. 2001. Bog communities of the West Gulf Coastal Plain: a profile. Bog Res.

Pap. Bot. Ecology 1:15 15 T.

MacRoserrs, M.H. and B.R. MacRoserts. 2003. The east-west transition of flora in Texas: a biogeographical analysis.

Sida 20:1693-1700.

MacRoserts, M.H. and B.R. MacRoserts. 2004a. The Big Thicket: typical or atypical? East Texas Hist. J. 42:42-51.

MacRoserts, M.H. and B.R.MacRoserts. 2004b. Understory composition in shortleaf/loblolly oak/hickory landscapes.

Unpublished report. Kisatchie National Forest, Louisiana.

MacRostnrs, B.R. and M.H. MacRoserts. 2004c. Floristics of upland longleaf pine savanna in central Louisiana. Proc.

Louisiana Acad. Sci. 65:1-13.

MacRoserTs, M.H. and B.R. MacRoserts. 2005. The ecology of Trillium texanum (Trilliaceae) on the Angelina National

Forest, Texas. Sida 21:1893-1903.

MacRogerTs, M.H. and B.R. MacRoserts. In Prep. Vascular flora and community structure of the C. Bickham-Dickson

Park/Red River Education and Research Park, Caddo Parish, Louisiana, Red River floodplain. 25 page ms.

AA D L a Fu | c a LM t bad 4L VAL ff" Lr 4 IDI x

dli., UIC WeSt uui COdStdl r idin 583

MacRoseets, B.R. and M.H. MacRoserts. 2006. An updated, annotated vascular flora of Caddo Parish, Louisiana,

with notes on regional phytogeography and ecology. Sida 22:1191-1219.

MacRostRrs, B.R., M.H. MacRoserts, and L.S. Jackson. 2002a. Floristics and management of pitcher plant bogs in

northern Natchitoches and Winn Parishes, Louisiana. Proc. Louisiana Acad. Sci. 64:14-21.

MacRostRrs, B.R., M.H. MacRoserts, and J.C. CATHEY. 2002b. Floristics of xeric sandylands in the post oak savanna

region of east Texas. Sida 20:373-386.

MacRostRrs, B.R., M.H. MacRoserts, and L.E. Brown. 2002c. Annotated checklist of the vascular flora of the Hickory

Creek Unit of the Big Thicket National Preserve, Tyler County, Texas. Sida 20:781-795.

MacRoserts, M.H., B.R. MacRoserrs, and L.S. Jackson. 2003. Louisiana prairies. Pp. 80-93. In E, Peacock and T. Schau-

wecker (eds.) Blackland prairies of the Gulf Coastal Plain. Univ. Alabama Press, Tuscaloosa.

MacRoserts, B.R, M.H. MacRoserts, and L.S. Jackson. 2004. Floristics of baygalls in central Louisiana. Phytologia

86:1-22.

Matos, J.A. and D.C. RupoLpH. 1985. The vegetation of the Roy E. Larsen Sandylands Sanctuary in the Big Thicket

of Texas. Castanea 50:228-249.

NELL, A.K. 2000. The vascular flora of Madison County, Texas. MS Thesis, Texas A & M University. College Station.

Nesom, G.L. and L.E. Brown. 1998. Annotated checklist of the vascular plants of Walker, Montgomery, and San

Jacinto counties, east Texas. Phytologia 84:107-153.

NEYLAND, R. 2002. An update to the vascular flora of Calcasieu Parish, Louisiana. Sida 20:431-433.

Peet, R.K. and D.J. ALLARD. 1993. Longleaf pine vegetation of the southern Atlantic and eastern gulf coastal regions:

a preliminary classification. Proc. Tall Timbers Fire Ecol. Conf. 18:45-80.

PETERSON, C.D. and L.E. Brown. 1983. Vascular flora of the Little Thicket Nature Sanctuary, San Jacinto County, Texas.

Brunswick Press, Houston.

PuiuiPPs, T., S.B. Wacker, B.R. MacRoserts, and M.H. MacRoserts. In Prep. Fox Hunter's Hill floristic survey.

Piatt, W.J. 1999. Southeastern pine savannas. In: R.C. Anderson, J.S. Fralish, and J.M. Baskin, eds. Savannas, barrens

and rock outcrop plant communities of North America. Cambridge Univ. Press., Cambridge. Pp. 23-51.

SiNGHURST, J.R.. J. Carey, D. ProcHaska, H. Haucke, G.C. KroH, and W.C. HoLmes. 2003. Vascular flora of Gus Engeling

Wildlife Management Area, Anderson County, Texas. Southeastern Naturalist 2:347—368.

SonniE, B.A., J.B. Gray, and PJ. CRUTCHFIELD. 2006. The vascular flora of the longleaf pine ecosystem of Fort Bragg and

Weymouth Woods, North Carolina. Castanea 71:129-161.

Thomas, R.D. and C.M. ALLEN. 1993-1998. Atlas of the vascular flora of Louisiana. 3 vols. Louisiana Department of

Wildlife and Fisheries and The Nature Conservancy. Baton Rouge.

THomas, R.D., L. Raymonp, B.R. MacRoserts, and M.H. MacRoserts. In Prep. Documented and annotated checklist

of the vascular flora of the Walter B. Jacobs Memorial Nature Park, Caddo Parish, Louisiana. Proc. Louisiana

Acad. Sci. 27 page ms.

WALKER, J. and R.K. Peet. 1983. Composition and species diversity of pine-wiregrass savannas of the Green Swamp,

North Carolina. Vegetatio 55:163- 179.

WHITTAKER, R.H. 1972. Evolution and measurement of species diversity. Taxon 21:213-251.

WUNDERLIN, R.P. 1998. Guide to the vascular plants of Florida. Univ. Presses of Florida, Tallahassee.

584 Journal of the Botanical R h Institute of Texas 1(1)

BOOK NOTICES

ELIZABETH L. Horn. 1994. Coastal Wildflowers of the Pacific Northwest. (ISBN 0-87842-291-9, pbk.).

Mountain Press Publishing Company, PO. Box 2399, Missoula, MT 59806, U.S.A. (Orders: www.

mountain-press.com, info@mtnpress.com, 406-728-1900, 406-728-1635 fax, 1-800-234-5308). $14.00,

179 pp., color photos, 1 map, 515" x 6",

This handy volume includes 164 species of the most common herbs and shrubs ( sedges, etc.) along the coast from Mendocino,

California, to the northern tip of Vancouver Island, British Columbia, illustrated iy: 200 0 photos The photos are grouped by family within

T "i J

hl tal forests. The textincludes notes

each of five designated habitats—beaches and dunes; d grasslands;

on ethnobotanical uses and personal observations on natural history. It’s a nicely done, low-priced guide, the fourth printing (2002) of

this 1993 volume, obviously continuing to be useful for students and visitors to the Pacific coast —Guy Nesom, Botanical Research Institute

of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A.

RONALD J. TAYLOR AND GEORGE W. DouaLas. 1995. Mountain Plants of the Pacific Northwest. (ISBN 0-87842-

314-1, pbk). Mountain Press Publishing Company, PO Box 2399, Missoula, MT 59806, U.S.A. (Orders:

www.mountain-press.com, info@mtnpress.com, 406-728-1900, 406-728-1635 fax, 1-800-234-5308).

$25.00, color photos, b/w line drawings, 6" x 9".

A field guide to Washington, western British Columbia, and southeastern Alaska, covering more than 450 species of “wildflowers, ferns,

trees, and grasses." Subsequently published field guides for roughly the same region have covered many more species (Plants of Western

Oregon, Washington & British Columbia -Kozloff 2005, 2500 species; Wildflowers of the Pacific Northwest -Turner & Gustafson 2006,

1200 species; see reviews in Sida 22(1); Plants of the Pacific Northwest Coast, Pojar & MacKinnon 2005, 794 species), but the coverage

of the Taylor & Douglas book is excellent and representative, and the large, beautiful, and informative photos of Ron Taylor are worth

having on any account.—Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A.

CALIFORNIA COASTAL COMMISSION. 2005. Experience the California Coast: A Guide to Beaches and Parks in

Northern California. (ISBN 0-520-24540-7, pbk). University of California Press, 2120 Berkeley Way,

Berkeley, CA 94704-1012, U.S.A. (Orders: www.ucpress.edu, 1-800-777-4726, 1-800-999-1958 fax).

$24.95, 320 pp., 315 color illustrations, 9 b/w photos, 1 line illustration, 47 maps, 6" x 9".

Excellent maps and photos for natural areas in Del Norte, Humboldt, Mendocino, Sonoma, and Marin counties. With this detailed guide

and one of the excellent, recently published field guides to flowering plants, botanists studying northwestern California plants are set

to go.— Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A.

KRISTIN HuisiNGA, LORI MAKARICK, KATE WATTERS. 2006. River and Desert Plants of the Grand Canyon. (ISBN

0-87842-523-3, pbk.). Mountain Press Publishing Company, PO. Box 2399, Missoula, MT 59806, U.S.A.

(Orders: www.mountain-press.com, info@mtnpress.com, 1-800-234-5308, 406-728-1900, 406-728-

1635 fax). $22.00, 276 pp., 263 color photos, 80 line drawings, 5 color figures and maps, 6" x 9".

More than 300 low elevation species are illustrated by 310 excellent color photos and 92 line drawings. The species are arranged by fam-

ily and genus within each of four categories: Ferns & fern allies, Grasses & grass-like plants, Trees, and Shrubs & forbs. Comments on

natural history accompany each illustration. A brief introduction to Grand Canyon Ecology is up front, and a novel thumbnail ID guide

is helpful: each of 5 pages shows 25 species, arranged by flower color. This is a great book, probably to become a huge seller, because it

will be useful and accessible to students, naturalists at all levels, and even tangential tourists.

Asteraceae taxonomy is difficult: “Cirsium mexicanum” on p. 198 is Carduus nutans; “Pseudognaphalium stramineum" on p. 217 is

Pseudognaphalium luteoalbum.— Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A.

J. Bot. Res. Inst. Texas 1(1): 584. 2007

DISTRIBUTION OF HANGING GARDEN VEGETATION ASSOCIATIONS

ON TEE CODE EST 5s

James F. Fowler! N.L. Stanton Ronald L. Hartman

USDA Forest Service Department of oology and Physiology Rocky Mountain Herbarium

Rocky Mountain Research Station University of Wyoming Department of Botany

2500 South Pine Knoll Drive Laramie, Wyoming 82071-3265, U.S.A. University of Wyoming

Flagstaff, Arizona 86001, U.S.A. Laramie, Wyoming 82071-3265, U.S.A.

ABSTRACT

Hangi ] island-like habitats dominated by meso phytic-hydrophytic plant communities, g i p the xeric canyon

1 E f

SIS añ Le Colorado Plateau in the American West. We measured the db nie of species and physical aan abitat

73 individual hanging gardens during the growing seasons of 1991-1993. Cluster analysis of a simplified Morisita community similar-

ity index matrix produced four distinct clusters of herbaceous vegetation: Adiantum capillus-veneris, Aquilegia micrantha, Calamagrostis

scopulorum, and Cirsium rydbergii Herbaceous Associations. These associations are based on species abundance and are floristically

described via summary tables showing dominance, constancy, and presence of hanging-garden endemic species. Univariate compari-

sons of species richness, hanging garden size, and dominance show that the A. capillus-veneris and A. micrantha associations tend to be

smaller, less species rich, and have higher dominance values than the C. scopulorum and C. rydbergii asssociations. Two associations were

dominated by species endemic to hanging gardens: Aquilegia micrantha and Cirsium rydbergii. These four hanging garden associations

contrast with previous local vegetation surveys which place all hanging garden communities into one vegetation type despite the lack

of acommon dominant or constant species.

RESUMEN

1 AERA ] + ld e Se ] ; 1 EE] 1 1 af

hidrófitas, que crecen en las filtraciones

de ls eee secas de los cafiones de la Meseta de eee en el oeste de ie Estados Unidos. Medimos la abundancia de especies y

características físicas del microhábitat de 73 jardines colgantes individuales durante las épocas de crecimiento de 1991-1993. El cluster

análisis de una matriz simplificada del indice de similitud de la comunidad de Morisita poro cuatro dead distintos de ML AG

herbácea asociada: Adiantum Capillus-veneris, Aquilegia micrantha, Calamagrostis scopulorum, y Ci

en la abundancia de las especies y son descritos florísticamente en una tabla que muestra la dominancia P constancia, y la presencia de

las PR endémicas en los iae MM I i de la riqueza de especies, el tamaño del jardín colgante

L E

y su A. capillus-veneris y de A. micrantha tienden ser más pequeñas, menos ricas en especies,

y tienen valores más oe en la dominancia que las asociaciones de Calamagrostis scopulorum y de Cirsium rydbergii. Dos asociaciones

están dominadas por especies endémicas de los jardines colgantes: Aquilegia micrantha y Cirsium rydbergii. Estas cuatro asociaciones de

jardines colgantes contrastan con los estudios locales previos de vegetación que colocan todas comunidades del jardines colgantes en

un solo tipo de vegetación a pesar de la falta de una especie común, dominante o constante.

The lush plant growth of hanging gardens on canyon walls in the entrenched drainage system of the Colo-

rado Plateau attracted the attention of early botanists and pioneers (Welsh & Toft 1981). Powell (1895), the

first scientific explorer of the Colorado River canyon, noted the “oak glens,” ferns and mosses on moistened

sandstone, and rich vegetation associated with seeps “in marked contrast to the general appearance of naked

rock” within what is now Glen Canyon NRA (Fig. 1). Many subsequent visitors have noted the island-like

boundary between hanging gardens and their xeric canyon-wall surroundings (Woodbury 1933; Malanson

& Kay 1980; Welsh 1989). The microclimate of the larger hanging gardens often contrasts sharply with the

surrounding high desert: water is abundant, soils are moist, and canyon walls offer periodic shade (Malanson

1980; Welsh & Toft 1981). Hanging gardens are also relatively small, mostly less than one hectare, further

contributing to the island-like effect, visually similar to the inselberg (granitic and gneissic rock outcrop)

communities found in North and South America, Africa, and Australia (Barthlott & Porembski 2000).

However, in contrast to the much better studied inselberg systems (Groger & Barthlett 1996; Porembski et

Dazlr M tata D h Ctt

'Correspondence: James F. Fowler, USDA Forest Service

t 1 1

86001, U.S.A.; jffowler@fs.fed.us

, Southwest Forest Science Complex, 2500 S. Pine Knoll Drive, Flagstaff, Arizona

J. Bot. Res. Inst. Texas 1(1): 585 — 607. 2007

586 Journal of the Botanical R h Institute of Texas 1(1)

Dinosaur NP 23

EIS LA

Arches NP

Capitol Reef NP

1 ire Canyonlands NP

n P.

e Natural Bridges NM

E ui diis

Glen Canyon NRA

Fic. 1. Study locations of hangi len plant iti pled on the Colorado Plateau during tl 1199121995.

al. 1996; Sarthou & Villers 1998), hanging gardens are moist vegetation islands rather than xeric vegetation

islands (Szarzynski 2000) embedded within bedrock expanses.

Hanging gardens are dominated by mesophytic and hydrophytic herbaceous vegetation (Malanson 1980;

Welsh & Toft 1981; Welsh 1989) which develops at and downslope from ground-water seepage zones within

the exposed aquifer along crossbedded sandstone canyon walls. Ground-water “sapping” erosion processes

(Higgins 1984; Laity & Malin 1985; Baker 1990; Dunne 1990; May et al. 1995) produce the physical hanging

garden habitat: perennially wet rock walls and/or wet, subirrigated colluvial soils (May et al. 1995). They are

often located in wet theater-headed valleys formed *by weakening or removal of basal support as a result of

enhanced weathering and erosion" by water seepage (Laity & Malin 1985, p. 203). These diffuse-discharge,

Fowler et al., Distribution of hanging gardens in the Colorado Plateau 587

seepage erosion areas are the result of ground water moving through highly transmissive aquifers, down the

dip planes to where impermeable layers within or below an aquifer intersect a canyon wall, as well as along

deep fractures within the aquifer (Laity & Malin 1985; Dunne 1990; May et al. 1995), and are distinctly

different from point-source erosion processes caused by discharge at springs (Higgins 1984). Ground-water

sapping processes provide both the raw material for colluvial soil development and water for subirrigation

of vegetation growing on that habitat.

Floristic work has elucidated the flora associated with hanging garden habitats (Eastwood, 1896; Clover

& Jotter 1944; Harrison et al. 1964; Welsh & Moore 1968; Welsh 1986a, b; Shultz et al. 1987; Welsh 1989),

including several species endemic to hanging gardens (Harrison et al. 1964; Welsh 1989; Fowler et al. 1995;

Welsh et al. 2003). Microhabitat affinities for several hanging garden plant species have also been described

(Welsh & Toft 1981; Welsh 1989).

Woodbury (1933), Welsh and Toft (1981), and May et al. (1995) have proposed hanging garden classifica-

tion systems based on microhabitats and geomorphology. Woodbury (1933) briefly described hanging garden

seral associes in Zion NP as part of a park-wide vegetation classification monograph. Malanson (1980, 1982)

and Malanson and Kay (1980) measured frequency of species occurrence, species richness, and calculated

similarity indices to investigate the effects of flooding frequency, dispersal distance, plant dispersal types,

and physical habitat characteristics on species assemblages in 29 small hanging gardens in Zion NP. These

studies did not measure species abundance or take a phytosociological approach. In this study, we measured

canopy coverage for each vascular plant species on 73 hanging gardens in a regional study, classified them

into herbaceous associations, and examined the regional distribution of those associations.

STUDY AREA

The study area was located on the Colorado Plateau in Utah and Colorado (Fig. 1), a major portion of the

known geographic range of hanging gardens. Field work was conducted on hanging gardens at seven National

Park Service units during the summers of 1991-1993: Arches NP, Canyonlands NP, Capitol Reef NP, Dinosaur

NP, Glen Canyon NRA, Natural Bridges NM, and Zion NP (Fig. 1). We selected most of the larger, known

hanging gardens in each park unit and many smaller ones that we were able to locate. Hanging gardens that

were too small to use the vegetation sampling methods described below were excluded from this study.

METHODS

We defined hanging garden habitats as predominately herbaceous vegetation growing on permanently wet

soil and wet rock surfaces, originating from seeps on canyon walls in sandstone aquifers (Malanson & Kay

1980). Hanging gardens delineated from the surrounding bedrock and xeric soils by the presence of wet

rock surfaces and/or wet soils (May et al. 1995). The hanging garden boundary with a riparian community,

when present, was defined by where colluvial soils stop and alluvial soils begin (May et al. 1995).

Each hanging garden was visually separated into the following microhabitats: seepline, wet wall, ledge,

and wet colluvial soil to stratify vegetation sampling. Seeplines are drier, linear microhabitats that develop

at fractures in the sandstone or, more often, at horizontal impervious bedding planes on canyon walls and

at the back of small alcoves. Wet walls included inverted, vertical, or sloping rock surfaces below seeps that

moisten the sandstone surface or have sufficient discharge to produce thin sheet flows of water with occa-

sional drip points. Ledge microhabitats are of sufficient horizontal width to support strips of hydrophytic

vegetation in depressions with wet, saturated soil. Wet colluvial soil microhabitats are complex in shape

and composed of wet, subirrigated colluvial soils located downslope of seepage zones, sometimes within

and/or below a larger alcove created by groundwater sapping. Since these soils develop directly from the

weathering of sandstone, they are obviously very sandy and may be virtually saturated near seeps.

We collected data on the following physical parameters: length of major axis, aspect, elevation, map

location, and geologic formation. Hanging garden size was calculated from major axis length and the mean

of ten equally spaced widths taken during vegetation sampling.

£+sha D o ID

588 Journal of t h Institute of Texas 1(1)

Vegetation

On each hanging garden, the herbaceous, vascular plant vegetation stratum was systematically sampled

(Cochran 1977; Krebs 1989) using 20 x 50 cm quadrats and six cover classes to estimate canopy cover for

each species (Daubenmire 1959) in each microhabitat. Woody plants less than two m tall were included in

the herbaceous stratum canopy estimates; the few occurrences of isolated trees were not included. The major

axis of each microhabitat was divided into ten equal segments. Widths were measured beginning with a

random starting point along the initial segment and at subsequent equal-segment lengths. A random point

was chosen along each width to determine placement of the lower left corner of the sampling frame Q0 cm

side). Edge quadrats were proportionally weighted to include only the portion within the hanging garden.

Total floristic composition (Daubenmire 1959, 1961), vascular plants only, was recorded during canopy cover

sampling and during an additional visual search (approximately one hr) on each hanging garden. Voucher

specimens for each species were collected. These were identified by B. E. Nelson, Herbarium Manager and

R. L. Hartman, Curator of the Rocky Mountain Herbarium at the University of Wyoming and are deposited

there and at the respective parks. Vascular plant nomenclature follows Flora of North America Editorial

Committee (1993+) and Welsh et al. (2003) in that order of priority.

Floristics of each herbaceous association are described in summary stand tables by floristic criteria

(Meuller-Dombois & Ellenberg 2002) and follow the definitions and analysis techniques of the U. S. Na-

tional Vegetation Classification (ESA 2006). Diagnostic classes for this study were dominance determined

by absolute canopy cover (Daubenmire 1959), constancy (frequency of occurrence) (Daubenmire 1952;

Meuller-Dombois & Ellenberg 2002), and endemism.

Indices

Plant community diversity (heterogeneity) was measured as two separate components (Krebs 1989): species

richness (McIntosh 1967) and dominance. Dominance in a community was determined by the proportion d

=N,,,./N, (Berger & Parker 1970) where N,,.. = number of individuals in the most abundant population and

N, = the total number of individuals in the community. Canopy coverage estimates were used as N,,, /N,

since both are proportional measures of species community importance.

Classification of hanging garden vegetation was based on cluster analysis of simplified Morisita (Horn

1966) community similarity indices for 73 hanging gardens calculated from species-level canopy coverage

estimates of existing vegetation. An exception is the genus Carex. Our canopy coverage estimates for six spe-

cies in this genus were combined since consistent, vegetative separation of species was not possible. These

indices were placed in a community similarity matrix for cluster analysis in SYSTAT 9 (SYSTAT 19993) using

the complete linkage method (farthest neighbor) on maximum Euclidean distances between indices.

Statistics

Statistical analyses were performed with SYSTAT 9 (19992). Plant species richness, species dominance,

and hanging garden size data were analyzed by General Linear Model ANOVA. Dominance and hanging

garden size were log transformed (SYSTAT 1999b) to achieve normal distributions prior to ANOVA tests.

Homogeneity of variances was checked post hoc and Tukey HSD tests were used for pairwise comparisons

of ANOVA results to detect significant differences between vegetation associations.

The statistical association between geologic formation and the putative herbaceous associations was

described using the Pearson Chi-square goodness-of-fit test to detect the presence of an association (Loether

& McTavish 1976; Wilkinson et al. 1996), and a proportional reduction in error measure (PRE), Goodman-

Kruskal's lambda, to measure the strength of statistical association (Wilkinson et al. 1996). PRE statistics

describe how much error is reduced in predicting the column variable when the row variable is known for

association tables (Wilkinson et al. 1996).

RESULTS

Vegetation Classification

Species lists with abundance estimates for species are in Appendix I. Cluster analysis of the simplified

Fowler et al., Distribution of hanging gardens in the Colorado Plateau 589

Morisita community similarity index (Krebs 1989) matrix classified the 73 hanging garden vascular plant

communities into four associations plus one dissimilar cluster (Fig. 2): Adiantum capillus-veneris, Aquilegia

micrantha, Calamagrostis scopulorum, and Cirsium rybergii Herbaceous Associations. Euclidian distance clas-

sification thresholds (Fig. 2) ranged from 1.03 for separating the Cirsium rydbergii Herbaceous Association

from the dissimilar group to 1.31 for separating the C. rydbergii and the Adiantum capillus-veneris Herbaceous

Associations. The dissimilar cluster (n=11) had hanging garden communities with little similarity within

the cluster (mean similarity index of 0.069) and had no plant species consistently in common.

The distribution of the four herbaceous associations across the seven geologic formations was different

than expected by chance alone (p=0.0098), the lambda statistic indicating a 25% improvement in being

able to predict the herbaceous association when the geologic formation is known (Table 1). The A. micrantha

and C. scopulorum Herbaceous Associations were widespread on the Colorado Plateau (Table 1). Both were

found in five parks and five sandstone geologic formations, differing in only one park and one geologic for-

mation; the C. scopulorum Herbaceous Association was found on the Humbug formation omitted from Table

1. The C. rydbergii Herbaceous Association was found only in the central part of the Colorado Plateau: two

parks and three formations (Table 1). The A. capillus-veneris Herbaceous Association was found on the same

geologic formation as the C. rydbergii Herbaceous Association but was more widespread (Table 1). The A.

capillus-veneris and C. rydbergii Herbaceous Associations were notably absent from the Cedar Mesa and Weber

formations, which were found only in our Natural Bridges NM and Dinosaur NP study sites respectively.

However, there were significant, positive statistical associations between geologic formations and herbaceous

associations (Table 1). The strongest ones being between the A. micrantha Herbaceous Association and the

Weber formation and between the C. rydbergii Herbaceous Association and the Navajo formation.

Adiantum capillus-veneris Herbaceous Association

Floristics of the Adiantum capillus-veneris Herbaceous Association are defined in Table 2. The dominant

species in this association was Adiantum capillus-veneris which covered from 32-9396 of the herbaceous

canopy in the 11 hanging gardens classified here (Fig. 2). Pairwise simplified Morisita community similar-

ity indices ranged from 0.48 to 0. 99. Both A. capillus-veneris and Epipactis gigantea had high constancy.

Adiantum capillus-veneris was found in seepline and wet wall microhabitats, whereas E. gigantea was found

in seepline microhabitats and on the upper portion of wet soil microhabitats next to vertical canyon walls

with wetter seeplines. Epipactis gigantea and the Genus Carex also had high canopy cover on some hanging

gardens. The vegetation was dominated by ferns, forbs, and graminoids (Table 2), but woody plants were

occasionally present as isolated individuals or at the drier edges of the wet soil/rock habitats characteristic

of hanging gardens.

Species diversity was relatively low in this association as shown by high dominance values, a measure

of community evenness, as well as low species richness (Table 3). Dominance values were significantly

higher than in Calamagrostis scopulorum and Cirsium rydbergii Herbaceous Associations (ANOVA F = 5.02,

p = 0.0036; Tukey HSD p = 0.0167 and 0.044 respectively). Five species endemic to hanging gardens were

found in this association (Table 2). The overall size of hanging gardens with this association was compara-

tively small (Table 3), significantly smaller than hanging gardens in the Cirsium rybergii and Calamagrostis

scopulorum Herbaceous Associations (see below) (ANOVA F = 7.21, p = 0.0003; Tukey HSD p = 0.0003 and

0.0266 respectively).

Aquilegia micrantha Herbaceous Association

The Aquilegia micrantha Herbaceous Association was dominated by the hanging garden endemic A. micrantha,

a constant species with an average canopy cover of 29% (Table 4). Aquilegia micrantha predominately grew

on wet colluvial soil microhabits. Calamgrostis scopulorum was also a constant species in this herbaceous

association, but with less than 5% average cover. Pairwise simplified Morisita community similarity indices

ranged from 0.28 to 1.00 for this association. With the exception of small Pinus edulis plants, which occurred

in trace amounts of cover on 47% of these hanging gardens, woody plants rarely occurred (Table 4).

In addition to A. micrantha, four other hanging garden endemic species were found on these sites, but

590 Journal of the Botanical R h Institute of Texas 1(1)

TABLE 1. Percentage of hanging garden vegetation associations found on each of five geologic formations (row percentages).

Pearson Chi Square (X? = 34.8798, p = 0.0098) and Goodman-Kruskal's lambda (A 20.2564 PRE) statistics indicate the existence

and strength of statistical association respectively. Positive “+” and negative "-" indicate the statistical association between a

geologic formation and a vegetation association. Cell Chi Square contributions are shown below cell percentages for each

row followed by two letter park codes (shown in Fig. 2) to indicate geographic distribution. Observed geologic formations

are Jurassic Entrada sandstone, Jurassic Navajo sandstone, Jurassic Navajo sandstone/Jurassic Kayenta formation contact

zone, Permian Cedar Mesa sandstone, Pennsylvanian-Permian Weber sandstone. Vegetation associations are Adca, Adiantum

capillus-veneris; Aqmi, Aquilegia micrantha; Casc, Calamagrostis scopulorum; Ciry, Cirsium rydbergii. N = 60, note that 2 geologic

formations with only 1 hanging garden each were omitted from this table.

Herbaceous Vegetation Association

Geologic Formation Adca Aqmi Casc Ciry

n=11 md Mio n=11

Entrada 29% 43% 0% - 29%

n=7 0.4002 9521] 2.4500 0.4002

AR AR AR

Navajo 3596 + 696 - 1896 41% +

n-17 2.6675 3.0243 1.4626 4.8386

CR GC, ZI GC GC Z| GC

Navajo/Kayenta 18% 18% 53% 12%

= 0.0044 0.6852 1.5634 0.4001

Ge Z| GL GE Cle Z| GC

Cedar Mesa 0% 50% 50% 0%

n=8 1.4667 13255 0.5143 1.4667

NB NB

Weber 0% 67% + 33% 0%

n=9 1.6500 4.6676 0.0071 1.6500

DI DI

at lower constancy (Table 4). This association was significantly less species rich than the C. scopulorum

and C. rydbergii Herbaceous Associations (see below) (ANOVA F = 3.79, p = 0.015; Tukey HSD p = 0.0248

and 0.0372 respectively). Dominance values averaged 0.59, not significantly different than the other three

associations (Table 3). Hanging gardens in this association were also significantly smaller in average size

than hanging gardens classified in the C. rydbergii association (see below)(ANOVA F = 7.21, p = 0.0003;

Tukey HSD p = 0.0009, Table 3).

Calamagrostis scopulorum Herbaceous Association

Floristics of the Calamagrostis scopulorum Herbaceous Association are defined in Table 5. As the dominant

species, C. scopulorum had an average cover of 31% and a range of 13-66% (Table 5). Calamagrostis scopu-

lorum grew in larger clumps on wet colluvial soil microhabitats and as smaller patches on seepline and wet

wall microhabitats. Several Carex species also occurred in these hanging gardens (Table 5). The hanging

garden endemic A. micrantha was moderately constant and ranged up to 1196 total cover. Simplified Moristia

community similarity indices ranged from 0.27 to 0.99 among the 23 hanging gardens classified into this

association.

Species diversity in this association was relatively high as shown by high species richness and low

dominance (Table 3). This association was significantly richer than the A. micrantha Herbaceous Association

(ANOVA F = 3.79, p = 0.0003; Tukey HSD p = 0.0248), and its dominance values were significantly lower

than the A. capillus-veneris Herbaceous Association (ANOVA F = 5.02, p = 0.0036; Tukey HSD p = 0.0167).

Three hanging garden endemic taxa were found on the 23 hanging gardens classified in this association

(Table 5). The C. scopulorum Herbaceous Association tended to contain large hanging gardens (Table 3),

significantly larger than hanging gardens in the A. capillus-veneris Herbaceous Association (ANOVA F = 7.21,

p = 0.0003; Tukey HSD p = 0.027).

Fowler et al., Distribution of hanging gardens in the Colorado Plateau

Backarch —AR

Swallow — GC

Overlook | — Zl

Horseshoe— CR

Crossbed— GC

Zephyr— GC

Lower Three— GC

Zigy—GC

Upper Three—GC

Hook— GC

Dune— GC

Weeping Rock—Zl

Fall —21

Court —Zl

Trail's End—Zl

Adiantum capillus-veneris

Association

——àÀ

— S

——————] RE

A) Cirsium

-j ry

—— d ee

m Association

Dissimilar

uster

Aquilegia micrantha

J Association

Calamagrostis scopulorum

J Association

Maximum Euclidean Distance Between Clusters

591

Fic. 2. Cluster tree showing four hanging garden herbaceou

names r

IND fi

Bridges WM DI= Dinosaur (NP).

associations and a dissimilar cluster based on cluster analysis of simplified

Moristia ae mia anaes Names refer to 73 individual hanging garcen plant communitie: Two letter Ende idus nanging areen

Arches (NP); CL = Canyonlands (NP); GC (NRA); ZI = Zion (NP); CR f (NP)

Vapiivi nevcri

TJ, NB = Natural

592 Journal of the Botanical R h Institute of Texas 1(1)

TABLE 2. Stand table for the Adiantum capillus-veneris Herbaceous Association showing canopy cover, diagnostic classes, and

constancy for all species with a constancy 20.40 and/or species endemic to hanging gardens. Constant species are defined

as constancy 20.60, and 't"is defined as less than 0.5 96 canopy cover. N = 11.

Species Diagnostic Constancy Average Minimum Maximum

dass 96 cover 96cover — ?ocover

Adiantum capillus-veneris dominant, constant 1.00 57 32 93

Epipactis gigantea constant 0.82 2 0 16

Aquilegia micrantha endemic 0.27 1 0 8

Cirsium rydbergii endemic 0.36 2 0 8

Mimulus eastwoodiae endemic 055 t 0 2

Primula specuicola endemic 0.27 t 0 2

Zigadenus vaginatus endemic 0.09 t 0 t

0.45 3 0 18

Petrophytum caespitosum 0.45 t 0 2

TABLE 3. Mean values (+ 1 SE) for hanging garden size, species richness, and dominance of four hanging garden herbaceous

associations. Also shown are number of sites (N) and number of hanging garden endemic plants species found (HGE) in each

association. Values followed by the same letter within each column do not differ at P«0.05 in Tukey's HSD post hoc tests.

Hanging garden vegetation Size m? Species Dominance N HGE

association Richness

Adiantum capillus-veneris 6820 ? 1442 *b O 710,05" 1] 5

Aquilegia micrantha 16745 3b as 05920069" 17 5

alamagrostis scopulorum 314x80* 1825 0.48+0.04 ^ 23 3

Cirsium rydbergii 5674163" (920 0.41+0.03 * 11 5

TABLE 4. Stand table for the Aquilegia micrantha Herbaceous Association showing canopy cover, diagnostic classes, and con-

stancy for all species with a constancy 20.40 and/or species endemic to hanging gardens. Constant species are defined as

constancy 20.60, and "t" is defined as less than 0.5 96 canopy cover. N = 17.

Species Diagnostic Constancy Average Minimum Maximum

dass 96 cover % cover 96 cover

Aquilegia micrantha dominant, constant, 1.00 29 9 54

endemic

Calamagrostis scopulorum Constant 0.82 4 0 32

Cirsium rydbergii Endemic 0.24 1 0 8

Mimulus eastwoodiae Endemic 025 1 0 11

Primula specuicola Endemic 0.24 t 0 2

Zigadenus vaginatus Endemic 0.35 2 0 19

Carex sp. 0.41 2 0 10

Pinus edulis 0.47 t 0 t

Cirsium rydbergii Herbaceous Association

The hanging garden endemic Cirsium rydbergii dominated this association with an average of 23% canopy

coverage (Table 6). These hanging gardens also had nine additional constant species (Table 6). Two of these

were co-dominants, A. capillus-veneris and Schyzacharium scoparium, and three were endemic to hanging

garden habitats: A. micrantha , Mimulus eastwoodiae, and Primula specuicola. One additional hanging garden

endemic species was also found at low constancy in these hanging gardens: Zigadenus vaginatus. Cirsium ry-

dbergii and A. micrantha were found throughout the wet colluvial soil microhabitat and Z. vaginatus was often

Fowler et al., Distribution of hanging gardens in the Colorado Plateau 593

TABLE 5. Stand table for the Calamagrostis scopulorum Herbaceous Association showing canopy cover, diagnostic classes, and

constancy for all species with a constancy 20.40 and/or species endemic to hanging gardens. Constant species are defined

as constancy 20.60, and 't"is defined as less than 0.5 96 canopy cover. N = 23.

Diagnostic Constancy Average Minimum Maximum

dass 96 cover 96 cover % cover

Calamagrostis scopulorum dominant 1.00 3l 15 66

Carex sp. constant 0.65 E 0 20

Aquilegia micrantha endemic 0.43 2 0 11

Zigadenus vaginatus endemic 0.22 1 0 14

Mimulus eastwoodiae endemic 0.22 t 0 t

Adiantum capillus-veneris 0.57 4 0 30

Epipactis gigantea 0.48 1 0 16

Maianthemum stellatum 057 3 0 20

Toxicodendron rydbergii 0.48 t 0 1]

TABLE 6. Stand table for the Cirsium rydbergii Herbaceous Association showing canopy cover, diagnostic classes, and constancy

for all species with a constancy 20.40 and/or species endemic to hanging gardens. Constant species are defined as constancy

70.60, and 't"is defined as less than 0.5 96 canopy cover. N = 11.

Species Diagnostic Constancy Average Minimum Maximum

dass % cover % cover % cover

Cirsium rydbergii dominant, constant, endemic 1.00 23 1 48

Adiantum capillus-veneris co-dominant, constant 0.91 10 0 29

Schizachyrium scoparium co-dominant, constant 0.64 [5 0 50

Carex sp. constant 0.91 7 0 19

Aquilegia micrantha constant, endemic 0.82 3 0 14

Primula specuicola constant, endemic 0.91 1 0 3

Mimulus eastwoodiae constant, endemic 0.64 3 0 8

Calamagrostis scopulorum constant 0.64 2 0 15

Petrophytum caespitosum constant 0.64 2 0 12

Epipactis gigantea constant 0.73 ] 0 5

Zigadenus vaginatus endemic 927 1 0 13

Castilleja linearifolia 0.45 t 0 2

Dichanthelium acuminatum 955 4 0 26

found in the upper portions of the wet colluvial soil microhabitat next to canyon walls. Mimulus eastwoodiae

and P. specuicola were often attached directly to vertical or inverted, wet rock surfaces. Simplified Morisita

community similarity indices ranged from 0.06 to 0.90 among the 11 hanging gardens in this association.

Species richness was relatively high (Table 3), significantly higher than in the A. micrantha Herbaceous

Association (ANOVA F = 3.79 p = 0.015; Tukey HSD p = 0.0372. Dominance values were low (Table 3),

significantly lower than in the A. capillus-veneris Herbaceous Association (ANOVA F = 5.02, p = 0.0036,

Tukey HSD p = 0.0044). Similar to the C. scopulorum Herbaceous Association, this association occurred on

large hanging gardens; the average size was significantly larger than in the A. capillis-veneris and A. micrantha

Herbaceous Associations (ANOVA F = 7.21, p = 0.0003; Tukey HSD p = 0.0003 and 0.0009 respectively).

DISCUSSION

The A. capillus-veneris and A. micrantha Herbaceous Associations tend to grow on smaller hanging gardens

that are less diverse; i.e., have low species richness and high dominance values (Table 3). The A. capillus-

veneris Herbaceous Association is dominated by relatively dry seepline microhabitats and by rhizomatous

594 Journal of the Botanical R h Institute of Texas 1(1)

stands of A. capillus-veneris which grow on and near narrow seeplines formed in the Navajo sandstone. The

A. micrantha association is dominated by A. micrantha which grows on the less wet colluvial soil microhabi-

tats, especially on Cedar Mesa and Weber sandstone formations. Note that these two geologic formations

and the Entrada formation were found in only one park each and thus also act as surrogates for geography.

In contrast, the hanging gardens with C. scopulorum and C. rydbergii Herbaceous Associations tended to

be larger and, by implication, wetter: i.e., large wet colluvial soil microhabitats. They also tend to be more

diverse: i.e., more species rich and low species dominance (Table 3).

The dissimilar cluster (Fig. 2) may represent a collection of unique plant communities which are not

part of any herbaceous association. Alternatively, with further sampling on the Colorado Plateau, we may

find that some of the hanging gardens within this cluster do represent undiscovered herbaceous associa-

tions. Further research is needed.

Neither Tuhy and MacMahon (1988) nor Romme et al. (1993) attempted to classify the vegetation of

individual hanging gardens but rather placed all hanging gardens into one vegetation type. We found no

species that all hanging gardens had in common (much less a common dominant) which precludes a single

hanging garden herbaceous association. The alliance name on NatureServe (2006), Aquilegia micrantha

saturated hanging garden herbaceous vegetation, would obviously work for the A. micrantha Herbaceous

Association but not for the remaining three associations since neither A. micrantha nor any other species was

constant or dominant across all four associations described in this paper (Tables 2, 4, 5, 6). Four separate

alliances would seem to be necessary in spite of a common distinct habitat.

NatureServe (2006) also lists two hanging garden herbaceous associations inside our study area: A.

micrantha - M. eastwoodiae and A. micrantha - C. scopulorum Herbaceous Vegetation, each association being

named for its dominant species. Aquilegia micrantha and C. scopulorum are shown as characteristic species

for both associations, but no data are shown for canopy coverage or constancy. In addition, no publications

are cited that have this type of data and the link to VegBank (2006) indicates that no plot-observations are

available, precluding further comparison. Woodbury's (1933) fern-columbine associes in Zion NP hanging

gardens is habitat-based and seems to overlap the A. capillus-veneris and C. scopulorum Herbaceous Associa-

tions described in this paper.

Outside of Woodbury's (1933) study in Zion NP, there are no previously published studies that describe

hanging garden associations. Most inselberg studies also classify vegetation by habitat (e.g., Burbanck & Platt

1964), but we found one with a phytosociological focus. Sarthou and Villiers (1998) conducted a regional

vegetation study on three inselbergs in French Giana using relevé quadrats and tabular analysis. In spite

of different techniques, their results were comparable to our study. They found six vegetation associations

versus our four, and they found two French Gianan inselberg endemics versus our five hanging garden

endemics. More interestingly, their most widespread vegetation association (with three subassociations)

was dominated by Pepinietum geyskesii, a French Gianan inselberg endemic. In our study, two vegetation

associations were dominated by hanging garden endemics: the A. micrantha and the C. rydbergii Herbaceous

Associations (Tables 4 & 6). Both studies illustrate the concept of insular vegetation on continental islands

at a regional scale. Our study also looked at the regional distribution of these four herbaceous associations

and found that each has a unique distribution on the Colorado Plateau (Table 1).

APPENDIX 1

Vascular plant species lists and species richness (R) for each hanging garden sampled. Canopy coverage estimates (0-100906)

are shown to the right of the scientific name for ee which fell within community sampling quadrats with T indicating less

than 0.5%; no estimates were made for taxa not falling wit! mpling quadrats: ARCH = Arches NP, CANY = Canyonlands NP,

CARE = Capitol Reef NP, DINO = Dinosaur NM, GLCA = Glen m NRA, NABR - Natural Bridges NM, ZION - Zion NP.

ARCH Backarch HG R = 24 Cirsium rydbergii 7 Heterotheca villosa var. Cyperaceae

Anacardiaceae Pseudognaphalium micro- minor 1 Carex aurea 2

ee rydbergii 7 cephalum T Solidago velutina subsp. Euphorbiaceae

Astera sparsiflora Euphorbia brachycera T

Fowler et al., Distribution of hanging gardens in the Colorado Plateau

Fabiaceae

Fagaceae

Quercus turbinella T

Liliaceae

Zigadenus vaginatus T

Onagraceae

Oenothera longissima

Orchidaceae

Epipactis gigantea 2

Platantherera zothecina

Poaceae

Elymus canadensis T

Muhlenbergia andina 3

Dichanthelium

acuminatum 1

Phragmites australis 2

Schizachyrium scoparium 1

Pteridaceae

Adiantum capillus-veneris 48

Primulaceae

Primula specuicola T

Ranunculaceae

Aquilegia micrantha 9

Rosaceae

Rosa s

Scrophulariaceae

Castilleja linariifolia

Mimulus eastwoodiae T

ARCH Bake-oven Wall HG

Asteraceae

Cirsium rydgbergii 1

Heterotheca villosa var.

minor 3

Cyperaceae

Carex aurea 4

Euphorbiaceae

Euphorbia brachycera T

ciao

ale hll 1

phyli

Kiliaceae

igadenus vaginatus T

Orchidaceae

Epipactis gigantea 3

Poaceae

Dichanthelium

acuminatum 7

Schizachyrium scoparium 32

Pteridaceae

Adiantum capillus-veneris 23

Primulaceae

Primula specuicola 2

Ranunculaceae

A HI H » +)

Aquilegia micrantha 14

Clematis ligt isticifolia T

Santalaceae

Comandra umbellata var.

pallida 3

Scrophulariaceae

Mimulus eastwoodiae T

ARCH Scaly HG R = 8

Anacardiaceae

Rhus aromatica var.

simplicifolia

Toxicodendron rydbergii 1

Asteraceae

Artemisia ludoviciana var.

ludoviciana

Cyperaceae

Carex aurea 1

Orchidaceae

Platantherera zothecina T

Poaceae

Dichantheli

Pteridaceae

Adiantum capillus-veneris 72

Scrophulariaceae

Mimulus eastwoodiae 2

acuminatum

ARCH Banded HG R = 6

Pinaceae

Pinus edulis

Poaceae

Dichantheli

Pteridaceae

Adiantum capillus-veneris 6

Primulaceae

Primula specuicola

Ranunculaceae

Aquilegia micrantha 33

Scrophulariaceae

Mimulus eastwoodiae 11

acuminatum

ARCH Winter Camp HG

R=19

Anacardiaceae

Rhus aromatica var.

simplicifolia T

Asteraceae

Cirsium rydbergii

Heterotheca villosa var.

minor 16

Solidago velutina subsp.

sparsiflora

Cyperaceae

Carex aurea 1

Euphorbiace

Euphorbia e 2

Liliaceae

Zigadenus vaginatus 7

Oleaceae

Fraxinus anomala

Onagraceae

Oenothera longissima T

Orchidaceae

Epipactis gigantea 15

Poaceae

Dichanthelium

acuminatum 1

Phragmites australis 9

Pteridaceae

Adiantum capillus-veneris 19

Pellaea breweri

Primulaceae

Primula specuicola 2

Ranunculaceae

Aquilegia micrantha 29

Santalaceae

Comandra umbellata var.

pallida 2

Scrophulariaceae

Castilleja exilis

Mimulus eastwoodiae

ARCH Solitaire HG R = 16

Anacardiaceae

Rhus aromatica var.

simplicifolia T

Toxicodendron rydbergii

Asteraceae

Cirsium rydbergii 1

Heterotheca villosa var.

minor 2

Solidago velutina subsp.

sparsiflora 1

Euphorbiaceae

Euphorbia brachycera T

Liliaceae

igadenus vaginatus 19

Orchidaceae

Epipactis gigantea 5

Poaceae

pcne thurberi 3

Dichantheli iu

Adiantum capillus-veneris 4

Pellaea glabella subsp.

simplex

Primulaceae

Primula specuicola 2

Ranunculaceae

A " ; si th

PAGUTICGIC FHHCIFEOTIUIG 38

Scrophulariaceae

Mimulus eastwoodiae 1

595

ARCH Dead Tree HG R= 18

Apocynaceae

Apocynum cannabinum

Asteraceae

Cirsium rydbergii 5

Heterotheca villosa var.

minor 2

Cyperaceae

Carex aurea 4

Euphorbiaceae

Euphorbia brachycera T

Fagaceae

Quercus havardii

Juncaceae

Juncus ensifolius var.

montanus

Liliaceae

Zigadenus vaginatus 1

Orchidaceae

Epipactis gigantea 5

Poaceae

Muhlenbergia andina 1

Dichanthelium

acuminatum 8

Schizachyrium scoparium 19

Pteridaceae

Adiantum capillus-veneris 11

Primulaceae

Primula specuicola 3

Ranunculaceae

Aquilegia micrantha 8

Santalaceae

Comandra umbellata var.

Scrophulariaceae

Castilleja exilis T

Mimulus eastwoodiae 8

CANY Box HG R = 17

Apocynaceae

Apocynum cannabinum 1

Asteraceae

Artemisia ludoviciana subsp.

ludoviciana

Cirsium rydbergii 16

Packera multilobata

Cyperaceae

Carex aurea 2

Liliaceae

Maianthemum stellatum

igadenus vaginatus

Orchidaceae

To d gigantea

rera zothec ina

PANNE

Pinus edulis

596

Poaceae

Calamagrostis scopulorum

Phragmites australis

Ranunculaceae

Aquilegia micrantha 10

Rhamnaceae

Rhamnus betulifolia

Rosaceae

Rosa sp.

Scrophulariaceae

Castilleja linariifolia

Mimulus eastwoodiae

CANY Northface HG R = 13

Asteraceae

Cirsium rydbergii 8

Betulaceae

Betula occidentalis

Cornaceae

Cornus sericea

Fagaceae

Quercus gambelii

Liliaceae

Maianthemum stellatum

Zigadenus vaginatus

Orchidaceae

Platantherera zothecina

Poaceae

Calamagrostis scopulorum

Ranunculaceae

Aquilegia micrantha 54

Rosaceae

Amelanchier sp.

Rosa sp.

Scrophulariaceae

Mimulus eastwoodiae

CANY Raven HG R = 16

Anacardiaceae

Rhus aromatica var.

simplicifolia

Toxicodendron rydbergii

Apocynaceae

Apocynum cannabinum T

Asteraceae

Cirsium arizonicum var.

bipinnatum

Berberidaceae

Berberis fendleri 3

Cyperaceae

Carex aurea 10

Orchidaceae

Platantherera zothecina

Pinaceae

Pinus edulis T

Poaceae

Calamagrostis scopulorum

Ranunculaceae

Aquilegia micrantha 10

Rhamnaceae

Rhamnus betulifolia 3

Rosaceae

Holodiscus dumosus

Rosa sp.

Rubiaceae

Galium sp.

Scrophulariaceae

Castilleja linariifolia

Mimulus eastwoodiae

CANY Gate HG R = 12

Anacardiaceae

Toxicodendron rydbergii

Asteraceae

Cirsium arizonicum var.

Platantherera zothecina

Pinaceae

Pinus edulis

Poaceae

Calamagrostis scopulorum

Muhlenbergia thurberi

Piptatherum micranthum

Ranunculaceae

Aquilegia micrantha 37

Rosaceae

Amelanchier sp.

Cercocarpus sp.

Rosa sp.

CANY Rocky HG R=7

Asteraceae

Cirsium arizonicum var.

bipinnatum 8

Erigeron sparsifolius 1

Pinaceae

Pinus edulis

Poaceae

Calamagrostis scopulorum

Ranunculaceae

Aquilegia micrantha 6

Santalaceae

Comandra umbellata var.

pallida 4

Scrophulariaceae

Mimulus eastwoodiae

fal, Dat

Journal of

CANY Tier HG R= 18

Agavaceae

Yucca sp. T

Anacardiaceae

Toxicodendron rydbergii 1

Apocynaceae

Apocynum cannabinum T

Asteraceae

Cirsium calareum 1

Cyperaceae

Carex aurea 9

Fagaceae

Quercus gambelii T

Liliaceae

Maianthemum stellatum 4

Orchidaceae

Epipactis gigantea 3

Platantherera zothecina T

Pinaceae

Pinus edulis

Poaceae

Calamagrostis scopulorum

Polemoniaceae

Gilia aggregata var.

maculata 2

Ranunculaceae

Aquilegia micrantha 11

Rhamnaceae

Rhamnus betulifolia

Rosaceae

Rosa sp. 2

Santalaceae

omandra umbellata var.

pallida T

Scrophulariaceae

Castilleja linariifolia 1

Mimulus eastwoodiae T

CARE Sidewall HG R = 12

Anacardiaceae

Toxicodendron rydbergii 19

Asteraceae

Cirsium arizonicum var.

bipinnatum 4

Heterotheca villosa var.

minor

Sonchus sp.

Equisetaceae

Equisetum laevigatum 1

Oleaceae

Fraxinus anomala

Orchidaceae

Epipactis gigantea

Poaceae

Elymus canadensis T

Dichanthelium

acuminatum 7

R h Institute of Texas 1(1)

Pteridaceae

Adiantum capillus-veneris 56

Scrophulariaceae

Castilleja scabrida

Ulmaceae

Celtis reticulata 5

CARE Horseshoe HG R = 8

Apocynaceae

Apocynum cannabinum

Asteraceae

Brickellia longifolia var.

Epipactis gigantea T

Oleaceae

Fraxinus anomala

Poaceae

Elymus canadensis

Elymus trachycaulus

Pteridaceae

Adiantum capillus-veneris 95

Scrophulariaceae

Mimulus eastwoodiae T

DINO Ponderosa HG R = 17

Asteraceae

Cirsium ownbeyi 2

Erigeron nematopyllus

Heterotheca villosa var.

minor T

Cupressaceae

Juniperus osteosperma

Hydrangiaceae

Fendlerella utahensis

Liliaceae

igadenus vaginatus 14

Pinaceae

Pinus edulis

Calamagrostis scopulorum

Muhlenbergia thurberi

Piptatherum micranthum

Poa fendleriana

Pteridaceae

Pellaea glabella subsp.

simplex

Ranunculaceae

Aquilegia micrantha 1

Rosaceae

Amelanchier sp.

Petrophytum caespitosum

Rubiaceae

Galium sp.

Scrophulariaceae

Castilleja linariifolia

Fowler et al., Distribution of hanging gardens in the Colorado Plateau

DINO Yampa HG R= 18

Asteraceae

Cirsium ownbeyi T

Erigeron nematophyllus 6

Í leterotheca villosa Val.

minor 2

Packera multilobata

Taraxacum officinale

Caprifoliaceae

Symphoricarpos sp.

Hydrangiaceae

Fendlerella utahensis

Liliaceae

Zigadenus vaginatus 1

Linaceae

Linum lewisii

Poaceae

Calamagrostis scopulorum

Elymus repens 1

Hesperostipa comata var.

Pellaea glabella subsp.

simplex

Ranunculaceae

Aquilegia micrantha 2

Rosaceae

Cercocarpus intricatus 7

Holodiscus dumosus T

Petrophytum caespitosum 7

crophulariaceae

Castilleja linariifolia T

DINO Bull HG R = 16

Aceraceae

Acer negundo 2

Anacardiaceae

Rhus aromatica var.

trilobata 1

Toxicodendron rydbergii 2

Asteraceae

Achillea millefolium T

laraxacum officinale

Betulaceae

Betula occidentalis

Caprifoliaceae

Symphoricarpos sp. 2

Cornaceae

Cornus sericea

Liliaceae

Maianthemum stellatum 7

Orchidaceae

Platantherera zothecina 7

Pinaceae

Pseudotsuga menziesii

Poaceae

Calamagrostis scopulorum

Primulaceae

Dodecatheon pulchellum

var. pulchellum

Ranunculaceae

Clematis ligusticifolia

Rubiaceae

Galium sp.

Salicaceae

Salix sp.

DINO Snow HG R = 8

Anacardiaceae

Rhus aromatica var.

trilobata 2

Cupressaceae

Juniperus osteosperma

Loasaceae

Mentzelia multicaulis 2

Orchidaceae

Epipactis gigantea 19

Pinaceae

Pinus ponderosa

Poaceae

Calamagrostis scopulorum

Primulaceae

Dodecatheon pulchellum

var. pulchellum 12

Ranunculaceae

Aquilegia micrantha 15

DINO Bench HG R = 17

Asteraceae

Achillea millefolium

Ericameria nauseosa

Cirsium ownbeyi 2

Erigeron nematophyllus

Heterotheca villosa var.

minor

Cupressaceae

Juniperus osteosperma

Cyperaceae

Carex aurea 2

Ephedraceae

Ephedra viridis

Hydrangiaceae

Fendlerella utahensis 1

Liliaceae

Zigadenus vaginatus 7

Pinaceae

Pinus edulis

Poaceae

Calamagrostis scopulorum

Elymus elymoides var.

brevifolius

Polypodiaceae

Pellaea glabella subsp.

simplex

Ranunculaceae

Aquilegia micrantha 35

Rosaceae

Cercocaarpus intricatus T

Salicaceae

Populus deltoides subsp.

wislizenii

DINO Rimrock HG R = 9

Asteraceae

Erigeron nematophyllus

Heterotheca villosa var.

minor

Liliaceae

Zigadenus vaginatus 2

Pinaceae

Pinus edulis

Poaceae

Calamagrostis scopulorum

Ranunculaceae

Aquilegia micrantha 19

Rosaceae

Cercocarpus intricatus

Petrophytum caespitosum

Scrophulariaceae

Castilleja lineariifolia

DINO Signature HG R = 19

Asteraceae

Ericameria nauseosa

Cirsium ownbeyi 4

Heterotheca villosa Val.

minor

Brassicaceae

Lepidium montanum var.

jonesii

Cyperaceae

Carex aurea 2

Euphorbiaceae

Euphorbia brachycera

Liliaceae

igadenus vaginatus 2

Orchidaceae

Epipactis gigantea 16

Poaceae

Calamagrostis scopulorum

Muhlenbergia thurberi

Dichanthelium

acuminatum T

Pteridaceae

Adiantum capillus-veneris T

Pellaea glabella subsp.

simplex

Primulac

Dodecatheon pulchellum

var. pulchellum

597

Ranunculaceae

Aquilegia micrantha 3

Clematis ligusticifolia 34

Rosaceae

Cercocarpus intricatus

Petrophytum caespitosum

Scrophulariaceae

Castilleja linariifolia

DINO Marbles HG R = 16

Asteraceae

Erigeron nematophyllus 18

Heterotheca villosa var.

minor 2

Xanthisma grindelioides T

Taraxacum officinale T

Berberidaceae

Berberis repens

Boraginaceae

Cryptantha sp. 1

Caprifoliaceae

Symphoricarpos sp. 1

Hydrangiaceae

Fendlerella utahensis 2

Pinaceae

Pinus edulis T

Poaceae

Achnatherum hymenoides T

Pteridaceae

Pellaea glabella subsp.

simplex

Ranunculaceae

Aquilegia micrantha 1

Rosaceae

Cercocarpus intricatus 8

Holodiscus dumosus 3

Petrophytum caespitosum 5

Scrophulariaceae

Castilleja sp. T

DINO Buzz HG R= 7

Asteraceae

Artemisia ludoviciana subsp.

ludoviciana 6

Melilotus officinalis

Poaceae

Agrostis stolonifera 6

Calamagrostis scopulorum 2

Primul e

Dodecatheon pulchellum

var. pulchellum 2

Ranunculaceae

Aquilegia micrantha 37

598

DINO Redrock HG R = 18

Asteraceae

Toxicodendron rydbergii

Artemisia ludovi

lude and 2

} leterotheca villosa Val.

minor 4

Taraxacum officinale 4

Brassicaceae

Lepidium latifolium 15

Cyperaceae

Carex aurea 5

Fabaceae

Melilotus albus

Melilotus officinalis

Liliaceae

Maianthemum stellatum

Orchidaceae

Platantherera zothecina 11

Poaceae

Agrostis stolonifera

Calamagrostis scopulorum 2

Muhlenbergia andina

Aquilegia micrantha 18

Clematis ligusticifolia

Rosaceae

Petrophytum caespitosum 7

Ulmaceae

Celtis reticulata 4

DINO Limestone HG R = 29

Apocynaceae

Apocynum cannabinum

Asclepiadaceae

Asclepias speciosa

Asteraceae

misia lt "2 PT

pror s 2

Ericameria nauseosa

Cirsium ownbeyi 15

Brassicaceae

Lepidium montanum var.

jonesii

Cyperaceae

Carex parryanna 2

Carex aquatilis var. aquatilis

Carex aurea

Equisetaceae

Equisetum laevigatum 1

Fabaceae

Melilotus officinalis T

Juncaceae

Juncus ensifolius var.

montanus

iciana subsp.

viciana subsp.

Lamiaceae

Mentha arvensis 2

Liliaceae

Maianthemum stellatum T

Onagraceae

a Sp.

Oenot th

ra caespitosa T

Poaceae

Calamagrostis scopulorum

Elymus trachycaulus 1

Glyceria striata T

Koeleria macrantha

Phragmites australis 8

Poa pratensis

Pseudoroegneria spicata

Polygonaceae

Erigonum corymbosum var.

corymbosum

Ranunculaceae

Aquilegia micrantha 1

Clematis ligusticifolia 12

Rosaceae

Rosa woodsii 6

Violaceae

Viola sp.

DINO Ely HG R =

Asteraceae

Crepis runcinata var. glauca

Erigeron nematophyllus

Betulaceae

Betula occidentalis

Euphorbiaceae

Euphorbia brachycera T

Hydrangiaceae

Fendlerella utahensis

Loasaceae

Mentzelia sp. 3

Pinaceae

Pinus edulis

Poaceae

Calamagrostis scopulorum 4

Muhlenbergia thurberi 1

Achnatherum hymenoides 1

Pteridaceae

Pellaea glabella subsp.

Dodecatheon pulchellum

var. pulchellum 1

Ranunculaceae

Aquilegia micrantha 13

Rosaceae

Cercocarpus intricatus

Petrophytum caespitosum 7

fal, Dat

D hil rers

Journal of

Santalaceae

Comandra umbellata var.

pallida 2

Scrophulariaceae

Castilleja linariifolia

GLCA Camp HG R= 19

Apocynaceae

Apocynum cannabinum 3

Asteraceae

Artemisia ludoviciana subsp.

mexicana

Brickellia longifolia var.

longifolia T

Taraxacum officinale

Berberidaceae

Berberis repens

Betulaceae

Ostrya knowltonii

Cyperaceae

Carex aurea 10

Fabaceae

Cercis occidentalis var.

orbiculata 3

Fagaceae

Quercus gambelii

Orchidaceae

Platantherera zothecina 1

Poaceae

Calamagrostis scopulorum 4

Polypogon interruptus

Polypogon viridis

Pteridaceae

Adiantum capillus-veneris 15

Ranunculaceae

Aquilegia micrantha 9

Clematis ligusticifolia

Rhamnaceae

Rhamnus betulifolia

Rosaceae

Rubus neomexicanus 3

Scrophulariaceae

Mimulus eastwoodiae 3

GLCA Channel HG R= 11

Anacardiaceae

Toxicodendron rydbergii T

Asteraceae

Cirsium rydbergii 1

Cyperaceae

Carex sp. T

Fagaceae

Quercus gambelii 2

Poacea

eT scopulorum 6

Pterida

foras capillus-veneris 7

of Texas 1(1)

Primulaceae

Primula specuicola 1

Ranunculaceae

Aquilegia micrantha 18

Rosaceae

Petrophytum caespitosum 5

Scrophulariaceae

Mimulus eastwoodiae 1

Vitaceae

Parthenocissus vitacea 2

GLCA Corner HG R= 13

Agavaceae

Yucca sp. T

Anacardiaceae

Artemisia ludoviciana subsp.

ludoviciana

Cirsium rydbergii 2

Cyperaceae

Carex aurea

Fagaceae

Quercus gambelii

Orchidaceae

Epipactis gigantea

Poaceae

Calamagrostis scopulorum

Pteridaceae

Adiantum capillus-veneris 54

Rosaceae

Petrophytum caespitosum

Scrophulariaceae

Mimulus eastwoodiae

Ulmaceae

Celtis reticulata

Vitaceae

Parthenocissus vitacea 4

GLCA Crossbed HG R = 23

Agavaceae

Yucca sp. |

Anacardiaceae

Rhus aromatica var.

simplicifolia

Asteraceae

Brickellia longifolia var.

longifolia 1

Cirsium rydbergii 11

Erigeron sparsifolius

Tetraneuris ivesiana

Gutierrezia sarothrae T

Solidago velutina subsp.

sparsiflora 2

Cyperaceae

Carex curatorum 3

Fowler et al., Distribution of hanging gardens in the Colorado Plateau

Loasaceae

Mentzelia cronquistii

Oleaceae

Fraxinus anomala

Orchidaceae

Epipactis gigantea

Poaceae

Andropogon glomeratus var.

scabriglumis

Calamagrostis scopulorum 2

Dichanthelium acuminatum

Schizachyrium scoparium 26

Polygonaceae

Eriogonum corymbosum

var. corymbosum T

Pteridaceae

Adiantum capillus-veneris 2

Primulaceae

Primula specuicola 1

Rhamnaceae

Rhamnus betulifolia

Rosaceae

Petrophytum caespitosum

Scrophulariaceae

Castilleja linariifolia

Ulmaceae

Celtis reticulata

GLCA Dune HG R = 20

Agavaceae

Yucca sp. 7

Anacardiaceae

Rhus aromatica var.

simplicifolia 3

Asteraceae

Symphotric hum chilense T

Cirsium rydbergii 9

Solidago velutina subsp.

sparsiflora 2

Cyperaceae

Carex sp. 19

Oleaceae

Fraxinus anomala

Orchidaceae

Epipactis gigantea

Poaceae

Andropogon glomeratus var.

scabriglumis 1

Bromus tectorurm ab

Calamagrostis scopulorum 7

Muhlenbergia andina T

Dichanthelium acuminatum

Pteridaceae

Adiantum capillus-veneris 6

Primulaceae

Primula specuicola 1

Rhamnaceae

Rhamnus betulifolia 9

Rosaceae

Petrophytum caespitosum 4

Salicaceae

Salix ligulifolia 8

Scrophulariaceae

Castilleja linariifolia 1

Ulmaceae

Celtis reticulata

GLCA Fence HGR 19

Anacardiaceae

Rhus aromatica var.

trilobata

Toxicodendron rydbergii 1

Asteraceae

Cirsium sp. T

Heterotheca villosa var.

minor 2

Solidago canadensis

Solidago velutina subsp.

sparsiflora 1

Campanulaceae

Lobelia cardinalis subsp.

graminea

Cyperaceae

Carex sp. 9

Fagaceae

Quercus gambelii

Liliaceae

Zigadenus vaginatus T

Oleaceae

Fraxinus sp. 2

Orchidaceae

Epipactis gigantea

Poaceae

Andropogon glomeratus var.

scabriglumis 69

Dichanthelium acuminatum

Phragmites australis

Polemoniaceae

Gilia aggregata var.

maculata 2

Pteridaceae

Adiantum capillus-veneris 4

Primulaceae

Primula specuicola

Rosaceae

Petrophytum caespitosum

GLCA Graffiti HG R = 8

Asteraceae

Brickellia longifolia var.

longifolia

Orchidaceae

Epipactis gigantea 5

Poaceae

Muhlenbergia sp. T

Pteridaceae

Adiantum capillus-veneris 93

Ranunculaceae

Aquilegia micrantha

Rhamnaceae

Rhamnus betulifolia

Rosaceae

Petrophytum caespitosum

Ulmaceae

Celtis reticulata

GLCA Hardwood HG R = 29

Anacardiaceae

Baccharis emoryi 7

Cirsium rydbergii

Erigeron sparsifolius

Brassicaceae

Thelypodium integrifolium

Cladium californicum 6

Fabaceae

Cercis occidentalis var.

orbiculata

Fagaceae

Quercus gambelii 3

Juncaceae

Juncus arcticus 13

Oleaceae

Forestiera pubescens 6

Orchidaceae

Epipactis gigantea

Poaceae

Calamagrostis scopulorum 3

Elymus canadensis T

Muhlenbergia andina

Dichanthelium

acuminatum-T

Panicum virgatum 7

Phragmites australis

Schizachyrium scoparium T

Sphenopholis obtusata

Pteridaceae

Adiantum capillus-veneris 9

Primulaceae

Primula specuicola 1

Ranunculaceae

Clematis ligusticifolia 1

Rhamnaceae

Rhamnus betulifolia 1

Rosaceae

Petrophytum caespitosum 1

599

Scrophulariaceae

Castilleja linariifolia

Mimulus eastwoodiae T

Ulmaceae

Celtis reticulata

Vitaceae

Parthenocissus vitacea 3

GLCA Hawk HG R= 12

Asteraceae

Brickellia longifolia var.

longifolia

Cirsium rydbergii 36

Fagaceae

Quercus gambelii T

Poaceae

Calamagrostis scopulorum 1

Muhlenbergia andina

Polypogon viridis T

Pteridaceae

Adiantum capillus-veneris 20

Primulaceae

Primula specuicola T

Ranunculaceae

Aquilegia micrantha T

Rosaceae

Petrophytum caespitosum

Scrophulariaceae

Mimulus eastwoodiae 1

GLCA Hook HG R - 28

Asteraceae

Artemisia ludoviciana subsp.

ludoviciana 4

Herrickia glauca var.

glauca 4

Brickellia longifolia var.

longifolia

Cirsium rydbergii 49

Solidago sp.

Sonchus asper

Cyperaceae 6

Carex aurea

Carex curatorum

Equisetaceae

Equisetum hyemale 2

Equisetum laevigatum

Fagaceae

Quercus gambelii 2

Juncaceae

Juncus arcticus 9

Liliaceae

Maianthemum stellatum 5

Orchidaceae

Epipactis gigantea T

600

Poaceae

Agrostis exarata

Andropogon glomeratus var.

scabriglumis T

Bromus tectorum T

Calamagrostis scopulorum T

Eh Imie canadencca

Muhlenbergia andina 2

Muhlenbergia thurberi

Dichanthelium

acuminatum T

Phragmites australis 2

Sphenopholis obtusata

Pteridaceae

Adiantum capillus-veneris 2

Primulaceae

Primula specuicola

Ranunculaceae

Aquilegia micrantha

Scrophulariaceae

Mimulus eastwoodiae

GLCA Ice Ha R = 17

Anacardiaceae

Toxicodendron rydbergii

Asteraceae

Aster sp.

Artemisia ludoviciana subsp.

ludoviciana

Brickellia longifolia var.

longifolia

Cirsium rydbergii 29

Cyperaceae 19

Carex lai Yugit JOAO!

Equisetaceae

Equisetum sp.

Oleaceae

Fraxinus anomala

Orchidaceae

Platantherera zothecina

Poaceae

Calamagrostis scopulorum T

Pteridaceae

Adiantum capillus-veneris 22

Primulaceae

Primula specuicola

Ranunculaceae

Aquilegia micrantha 3

Rosaceae

Petrophytum caespitosum

Scrophulariaceae

Castilleja linariifolia

Scrophulariaceae

Mimulus eastwoodiae

Vitaceae

Parthenocissus vitacea 7

GLCA Lower Cow HG

Anacardiaceae

Toxicodendron rydbergii 1

Asteraceae

Cirsium rydbergii

Solidago veluti! HA subsp.

sparsiflora 1

Campanulaceae

Lobelia cardinalis subsp.

graminea 1

Cyperaceae

Carex curatorum 14

Fagaceae

Quercus gambelii T

Orchidaceae

Epipactis gigantea T

Poaceae

Calamagrostis scopulorum

Phragmites australis

Pteridaceae

Adiantum capillus-veneris 4

Primulaceae

Primula specuicola 10

Ranunculaceae

Clematis ligusticifolia 2

GLCA Lower Three Garden

Asteraceae

Cirsium rydbergii 21

Cyperaceae 8

Carex aurea

Carex curatorum

Poaceae

Dichanthelium

acuminatum 26

Muhlenbergia sp. 2

Primulaceae

Primula specuicola 3

Rosaceae

Petrophytum caespitosum

Scrophulariaceae

Castilleja linariifolia 2

GLCA Pedestal HG R = 14

Agavaceae

Yucca sp. 4

Asteraceae

Cirsium rydbergii 7

Heterotheca villosa var.

minor T

Cyperaceae 14

Carex aurea

Carex curatorum

fal, Dat

Journal of

Juncaceae

Juncus arcticus T

Onagraceae

Oenothera longissima 1

Orchidaceae

Epipactis gigantea T

Poaceae

Calamagrostis scopulorum T

Schizachyrium scoparium 32

Pteridaceae

Adiantum capillus-veneris 17

Ranunculaceae

Aquilegia micrantha 1

Clematis ligusticifolia 1

Rhamnaceae

Rhamnus betulifolia 1

GLCA Pyro HG R = 22

Asteraceae

Artemisia ludoviciana subsp.

Ibula 2

Brickellia longifolia var.

longifolia T

Cirsium rydbergii 8

nod rannrion

za canadensis

Pseudognaphalium

luteoalbum

Sonchus arvensis

Carex aurea 9

Scirpus sp.

Fabaceae

Cercis occidentalis var.

orbiculata

Fagaceae

Quercus gambelii

Orchidaceae

Epipactis gigantea T

Poaceae

Bromus rubens

Eh rniicprannden si

Muhlenbergia andina

Dichanthelium

acuminatum 1

Polypogon monspeliensis

Pteridaceae

Adiantum capillus-veneris 68

Primulaceae

Primula specuicola

Scrophulariaceae

Mimulus eastwoodiae

Tamaricaceae

Tamarix ramosissima

Typhaceae

Typha sp. 6

titute of Texas 1(1)

GLCA Rattlesnake HG

R=20

Asteraceae

Brickellia longifolia var.

longifolia T

Cirsium rydbergii T

Solidago sp. 6

Cyperaceae

Carex aurea 1

Fagaceae

Quercus gambelii 1

Juncaceae

Juncus ensifolius var.

montanus

Liliaceae

Maianthemum stellatum 4

Oleaceae

Forestiera pubescens T

Fraxinus anomala

Orchidaceae

Epipactis gigantea 3

Platantherera zothecina T

Poaceae

Agrostis exarata

Calamagrostis scopulorum

Elymus canadensis T

Pteridaceae

Adiantum capillus-veneris 7

Ranunculaceae

Aquilegia micrantha 3

Rhamnaceae

Rhamnus betulifolia

Rosaceae

Petrophytum caespitosum

Scrophulariaceae

Mimulus eastwoodiae T

Vitaceae

Parthenocissus vitacea 4

GLCA Stone Basin HG

Anacardiaceae

Toxicodendron rydbergii 1

Apocynaceae

Apocynum cannabinum 11

Asteraceae

Conyza canadensis T

Pseudognaphalium sp. 2

Solidago sp. 4

Campanulaceae

Lobelia cardinalis subsp.

graminea

Cyperaceae

Carex sp. 4

Poaceae

Andropogon glomeratus var.

scabriglumis

Fowler et al., Distribution of hanging gardens in the Colorado Plateau

Bouteloua cul tipendula Var.

caespitosa

Calamagrostis scopulorum T

Elymt ic ranadencic

Dichanthelium

Adiantum capillus-veneris 4

Scrophulariaceae

Castilleja linariifolia

Vitaceae

Parthenocissus vitacea 10

GLCA Swallow HG R= 11

Asteraceae

Cirsium rydbergii

Sonchus sp. T

Platantherera zothecina 3

Poaceae

Calamagrostis scopulorum T

Pteridaceae

Adiantum capillus-veneris 42

Primulaceae

Primula specuicola 2

Ranunculaceae

A Ai " i PI

A\GUHCGIC FEHOIOTICHIO 6

Clematis ligusticifolia

Rosaceae

Rubus neomexicanus

Scrophulariaceae

Mimulus eastwoodiae

GLCA Upper Three HG

Asclepiadaceae

Asclepias latifolia

Asteraceae

Solidago velutina subsp.

sparsiflora

Cirsium rydbergii 40

Cyperaceae

Carex aurea 15

Fabaceae

Cercis occidentalis var.

orbiculata

agaceae

Quercus gambelii 1

Orchidaceae

Epipactis gigantea 1

Poaceae

Calamagrostis scopulorum

]

Muhlenbergia sp. 1

Dichanthelium acuminatum

Schizachyrium scoparium! 5

Pteridaceae

Adiantum capillus-veneris 1

Primulaceae

Primula specuicola T

Rosaceae

Petrophytum caespitosum

Scrophulariaceae

Castilleja linariifolia

Mimulus eastwoodiae T

GLCA Wrong HG R = 10

Anacardiaceae

Toxicodendron rydbergii 11

Apocynaceae

Apocynum cannabinum

Asteraceae

Artemisia ludoviciana subsp.

mexicana T

Solidago velutina subsp.

sparsiflora

Fagaceae

Quercus gambelii 1

Poaceae

Andropogon glomeratus var.

scabriglumis 25

Calamagrostis scopulorum

Dichanthelium

acuminatum 2

Polemoniaceae

Gilia aggregata var.

maculata

Pteridaceae

Adiantum capillus-veneris 30

GLCA Zephyr HG R = 17

Agavaceae

Yucca sp. 12

Anacardiaceae

Rhus aromatica var. simpli-

cifolia 2

Asteraceae

Cirsium rydbergii 31

Cyperaceae

Carex curatorum 4

Fagaceae

Quercus gambelii

Orchidaceae

Epipactis gigantea T

oaceae

Andropogon glomeratus var.

Schizachyrium scoparium 50

Primulaceae

Primula specuicola T

Pteridaceae

Adiantum capillus-veneris 2

Ranunculaceae

Aquilegia micrantha 2

Rhamnaceae

Rhamnus betulifolia 2

Rosaceae

Petrophytum caespitosum 2

Scrophulariaceae

Castilleja linariifolia 2

Ulmaceae

Celtis reticulata

Urticaceae

Parietaria pennsylvanica T

GLCA Zigy HG R = 30

Agavaceae

Yucca toftiae

Apocynaceae

Apocynum cannabinum T

Asteraceae

Artemisia ludoviciana subsp.

ludoviciana

Sonchus asper

Cyperaceae 3

Carex aurea

Carex curatorum

Fleocharis rostellata 12

Scirpus Sp.

Fabaceae

Oxytropis sp.

Liliaceae

Zigadenus vaginatus 13

Onagraceae

Oenothera longissima

Orchidaceae

Epipactis gigantea T

Poaceae

Agrostis exarata

Bromus tectorum

Calamagrostis scopulorum 6

Muhlenbergia asperifolia

Muhlenbergia thurberi

Dichanthelium acuminatum

Phragmites australis T

Schizachyrium scoparium 10

Pteridaceae

Adiantum capillus-veneris 1

Primulaceae

Primula specuicola T

Ranunculaceae

Aquilegia micrantha 1

Clematis ligusticifolia

Rosaceae

Petrophytum caespitosum

Rosa woodsii

Scrophulariaceae

Mimulus eastwoodiae

601

Typhaceae

Typha sp.

NABR AM HG R= 5

Pinaceae

Pinus edulis T

Poaceae

Calamagrostis scopulorum

Ranunculaceae

Aquilegia micrantha 27

Rhamnaceae

Rhamnus betulifolia

Rosaceae

Cercocarpus montanus

NABR Fir HG R=11

Asteraceae

Cirsium undulatum

Heterotheca villosa var.

minor

Gentianaceae

Swertia radiata 2

Hydrophyllaceae

Phacelia sp.

Liliaceae

Zigadenus vaginatus 6

Pinaceae

Abies bifolia

Poaceae

Calamagrostis scopulorum

Poa fendleriana

Rosaceae

Amelanchier sp. 2

Holodiscus dumosus 2

Rubiaceae

Galium multiflorum var.

coloradoense

NABR Kachina HG R = 19

piaceae

Aletes macdougalii subsp.

breviradiatus 2

Asteraceae

Cirsium undulatum 5

Cirsium sp.

Erigeron kachinensis 5

Heterotheca villosa var.

minor T

Celastraceae

Pachystima myrsinites T

Cyperaceae

Carex aurea T

Ephedraceae

Ephedra viridis 12

Gentianaceae

Swertia radiata T

Juncaceae

Juncus arcticus T

602

Liliaceae

igadenus vaginatus 19

Orchidaceae

Epipactis gigantea T

Pinaceae

Pinus edulis

Poaceae

Calamagrostis scopulorum 4

Muhlenbergia thurberi 5

Ranunculaceae

Aquilegia micrantha T

Clematis ligusticifolia

Rhamnaceae

Rhamnus betulifolia 4

Rubiaceae

Galium multiflorum var.

coloradoense

Santalaceae

Comandra umbellata var.

pallida 1

NABR Liverwort HG R = 9

Brassicaceae

Lepidium montanum var.

jonesii T

Lesquerella rectipes

Cupressaceae

Juniperus osteosperma

Elaeagnaceae

Shepherdia rotundifolia

Pinaceae

Pinus edulis T

Poaceae

Calamagrostis scopulorum

Pteridaceae

Cheilanthes feei

Ranunculaceae

Aquilegia mi I AI tha 49

Rosaceae

Petrophytum caespitosum

NABR Long HG R = 10

Berberidaceae

Berberis repens

Gentianaceae

Erigeron kachinensis

Liliaceae

igadenus vaginatus

Pinaceae

Pinus edulis

Poaceae

Calamagrostis scopulorum 4

Ranunculaceae

Aquilegia micrantha 14

Rhamnaceae

Rhamnus betulifolia

Rosaceae

Cercocarpus montanus

Rubiaceae

Galium multiflorum var.

coloradoense

NABR Micro HGR = 4

Asteraceae

Cirsium sp. 3

Erigeron kachinensis 6

Poaceae

Calamagrostis scopulorum 3

Ranunculaceae

Aquilegia micrantha 48

NABR Petro HG R = 9

Asteraceae

Cirsium undulatum 11

Erigeron kachinensis 7

Heterotheca villosa var.

minor T

Cyperaceae

Carex aurea 11

Gentianaceae

Swertia radiata

rchidaceae

Platantherera sp. 1

Poaceae

Calamagrostis scopulorum

Pteridaceae

Adiantum capillus-veneris T

Ranunculaceae

Aquilegia micrantha T

NABR Slickrock HG R = 14

Asteraceae

Cirsium sp. 13

Cirsium arizonicum var.

bipinnatum

Cirsium undulatum

Heterotheca villosa var.

minor

Cyperaceae

Carex aurea 2

Fagaceae

Quercus gambe

Gentianaceae

Swertia radiata

Juncaceae

Juncus arcticus 12

Liliaceae

Zigadenus vaginatus T

Orchidaceae

Platantherera sp.

—

Pinaceae

Pinus edulis 2

Poaceae

fal, Dat

Journal of

Bromus tectorum

Calamagrostis scopulorum

Muhlenbergia thurberi

Rubiaceae

Galium multiflorum var.

coloradoense 5

NABR Tuwa HG R = 10

Apiaceae

Aletes macdougalii subsp.

breviradiatus

Asteraceae

Cirsium undulatum 5

Cyperaceae

Carex aurea 8

Gentianaceae

Swertia radiata 4

Juncaceae

Juncus arcticus

Pinaceae

Pinus edulis T

Poaceae

Calamagrostis scopulorum

Muhlenbergia thurberi 3

Ranunculaceae

Aquilegia micrantha 2

Rhamnaceae

Rhamnus betulifolia

ZION Canyon Overlook I

Aceraceae

Acer negundo var. violaceum

Asteraceae

Brickellia californica 2

Ageratina herbacea

Perityle tenella T

Stephanomeria pauciflora

Fagaceae

Quercus turbinella T

Orchidaceae

Epipactis gigantea 16

Poaceae

Muhlenbergia thurberi 5

Poa fendleriana

Pteridaceae

Adiantum capillus-veneris 34

Rosaceae

Holodiscus dumosus

Petrophytum caespitosum T

Saxifragaceae

Heuchera rubescens

Scrophulariaceae

Castilleja sp.

titute of Texas 1(1)

ZION Canyon Overlook Il

Asteraceae

Artemisia ludoviciana subsp.

albula 3

Brickellia californica

Brickellia grandiflora 6

Erigeron sionis T

Heterotheca villosa var.

Perityle tenella

Solidago velutina subsp.

sparsiflora

Sonchus asper

Stephanomeria pauciflora 1

Taraxacum officinale T

Celastraceae

Pachystima myrsinites 1

Fagaceae

Quercus turbinella

Zauschneria latifolia 2

Orchidaceae

Epipactis gigantea 1

Poaceae

Bromus ciliatus

Bromus tectorum 1

Muhlenbergia thurberi T

Poa fendleriana 1

Poa pratensis

Pteridaceae

Adiantum capillus-veneris 33

Rosaceae

Cercocarpus intricatus

Petrophytum caespitosum 2

Salicaceae

Populus fremontii 3

Salix gooddingii

Scrophulariaceae

Castilleja scabrida var.

scabrida

ZION Court of the

Patriarchs HG R = 20

Aceraceae

Acer negundo var. negundo T

Agavaceae

Yucca utahensis

Asteraceae

Artemisia ludoviciana subsp.

ludoviciana

Perityle tenella

Solidago missouriensis

Sonchus asper

Cyperaceae

Carex aurea 5

Fowler et al., Distribution of hanging gardens in the Colorado Plateau

Equisetaceae

Equisetum laevigatum 5

Liliaceae

Maianthemum stellatum 2

Oleaceae

Fraxinus velutina

Orchidaceae

Epipactis gigantea

Poaceae

Agrostis exarata

Calamagrostis scopulorum

Muhlenbergia sp. 14

Poa pratensis 5

Adiantum capillus-veneris 13

Rosaceae

Petrophytum caespitosum

Rubiaceae

Saxifragaceae

Parnassia palustris var.

montanensis

Scrophulariaceae

Mimulus cardinalis

ZION Fall HGR = 14

Araliaceae

Aralia racemosa subsp.

bicrenata

Asteraceae

Erigeron sionis

Taraxacum officinale

Brassicaceae

Nasturtium officinale

Juncaceae

Jur JE GI»SCT isifolius val.

montanus T

Liliaceae

Maianthemum stellatum 6

Orchidaceae

Epipactis gigantea

Poaceae

Agrostis exarata

Pteridaceae

Adiantum capillus-veneris 40

Primulaceae

Dodecatheon pulchellum

var. zionense 32

Ranunculaceae

Aquilegia chrysantha 2

Aquilegia formosa var.

ormosa

Rosaceae

Petrophytum caespitosum

Scrophulariaceae

Mimulus cardinalis

ZION Falling Water HG

Aceraceae

Acer negundo var. violaceum

Anacardiaceae

Toxicodendron rydbergii

Araliaceae

Aralia racemosa ssp.

bicrenata

Asteraceae

Symphotric hum lanceola-

tum var. hesperium 30

Brickellia longifolia var.

longifolia

Cirsium sp.

Chrysothamnus scopulorum

Sphaeromeria ruthiae

Cyperaceae

Carex aurea 1

Hydrangiaceae

lamesia icana var.

Liliaceae

Maianthemum stellatum

Oleaceae

Fraxinus velutina

Poaceae

Bromus ciliatus

Calamagrostis scopulorum

Pteridaceae

Adiantum capillus-veneris 2

Ranunculaceae

Aquilegia chrysantha 22

Clematis ligusticifolia

Rosaceae

Petrophytum caespitosum

Rubiaceae

Galium triflorum

crophulariaceae

Mimulus cardinalis

Violaceae

Viola sp. T

ZION Grotto HG R = 28

Aceraceae

Acer negundo var. violaceum

Anacardiaceae

Toxicodendron rydbergii

Araliaceae

Aralia racemosa subsp.

bicrenata 2

Asteraceae

Artemisia ludoviciana

Symphotric hum lanceola-

tum var. hesperium

Brickellia longifolia var.

longifolia

Cirsium neomexicanum

Berberidaceae

Berberis repens T

Brassicaceae

Erysimum capitatum

Caprifoliaceae

Pachystima myrsinites

Hydrangiaceae

Jamesia americana var.

Zionis

Liliaceae

Maianthemum stellatum 2

Poaceae

Bromus ciliatus

Calamagrostis scopulorum

Pteridaceae

Adiantum capillus-veneris 1

Adiantum aleuticum

Primulaceae

Dodecatheon pulchellum

Aquilegia chrysantha 4

Aquilegia formosa var.

formosa

Rosaceae

Amelanchier alnifolia

Cercocarpus intricatus

Holodiscus dumosus

Petrophytum caespitosum

Rubiaceae

Galium triflorum T

Saxifragaceae

Heuchera rubescens

Scrophulariaceae

Mimulus cardinalis 6

Violaceae

Viola nephrophylla 2

ZION Kaye's HG R = 30

Anacardiaceae

Toxicodendron rydbergii 1

Asteraceae

Artemisia ludoviciana subsp.

ludoviciana T

Cirsium arizonicum

laraxacum officinale T

Cyperaceae 19

Carex aurea

Carex curatorum

Equisetaceae

Equisetum hyemale

Fagaceae

Quercus turbinella

603

Hydrophyllaceae

Phacelia heterophylla

Juncaceae

Juncus ensifolius var.

montanus T

Liliaceae

Maianthemum stellatum T

Oleaceae

Fraxinus velutina

Orchidaceae

Epipactis gigantea

Poaceae

Agrostis exarata

Calamagrostis scopulorum

Muhlenbergia andina 1

Dichanthelium

acuminatum 2

Phragmites australis 1

Poa pratensis 2

Danthonia californica T

Pteridaceae

Adiantum capillus-veneris 3

Ranunculaceae

Aquilegia chrysantha 2

Aquilegia formosa var.

formosa

Clematis ligusticifolia

Rosaceae

Petrophytum caespitosum

Rubiaceae

Galium trifidum

Saxifragaceae

Heuchera rubescens

Parnassia palustris var.

montanensis

Scrophulariaceae

Mimulus cardinalis T

Violaceae

Viola sp. 1

ZION Lower Emerald HG

Anacardiaceae

Toxicodendron rydbergii

Asteraceae

Herrickia glauca var.

glauca 1

Cyperaceae

Carex aurea

Liliaceae

Maianthemum stellatum 3

Orchidaceae

Epipactis gigantea 1

Poaceae

Agrostis exarata 5

Calamagrostis scopulorum

604

Pteridaceae

Adiantum capillus-veneris 1

Primulaceae

Dodecatheon pulchellum

var. zionense 25

Ranunculaceae

Aquilegia chrysantha T

Scrophulariaceae

Mimulus cardinalis

Vitaceae

Vitis arizonica

ZION Menu Falls HG R = 23

Aceraceae

Acer negundo var. negundo

Anacardiaceae

Toxicodendron rydbergii

Apocynaceae

Apocynum cannabinum

Asteraceae

Baccharis salicina

Cirsium arizonicum

Sonchus asper

Cyperaceae T

Carex aurea

Carex curatorum

Equisetaceae

Equisetum laevigatum

Fagaceae

Quercus turbinella

Liliaceae

Maianthemum stellatum

Oleaceae

Fraxinus velutina

Poaceae

Agrostis exarata

Bromus diandrus

Calamagrostis scopulorum

Polypogon viridis

Muhlenbergia sp. 6

Pteridaceae

Adiantum capillus-veneris 16

Ranunculaceae

Aquilegia chrysantha 9

Aquilegia formosa var.

ormosa

Clematis ligusticifolia

Rosaceae

Petrophytum caespitosum

Scrophulariaceae

Mimulus cardinalis 15

ZION Narrows Trail HG

R26

Aceraceae

Acer negundo var. negundo 4

Anacardiaceae

Toxicodendron rydbergii T

Asteraceae

Artemisia p ud subsp.

ludov.

Brickellia ee var.

ongifolia

Cirsium neomexicanum

Cirsium wheeleri

Sonchus asper

Berberidaceae

Berberis repens

Cannabaceae

Humulus lupulus var.

ceae

Maianthemum stellatum 19

Orchidaceae

Epipactis gigantea

Poaceae

Bromus ciliatus

Bromus diandrus

Calamagrostis scopulorum

Elymus glaucus

Muhlenbergia andina

Achnatherum hymenoides

Pteridaceae

Adiantum d veneris 8

Primulace

D pulchellum

var. Zionense 13

Ranunculaceae

Aquilegia chrysantha 1

Aquilegia formosa var.

formosa

Clematis ligusticifolia T

Rosaceae

Petrophytum caespitosum

Scrophulariaceae

Mimulus cardinalis T

Vitaceae

Vitis arizonica 3

ZION Pine Creek HG R = 6

Asteraceae

Herrickia glauca var.

glauca 8

Solidago velutina subsp.

sparsiflora

Poaceae

Calamagrostis scopulorum

Poa fendleriana

Pteridaceae

Adiantum capillus-veneris

fal, Dat

Journal of

Primulaceae

Dodecatheon pulchellum

var. zionense 6

ZION Snail HG R = 13

Anacardiaceae

Toxicodendron rydbergii

Asteraceae

Herrickia glauca var. glauca

Liliaceae

Maianthemum stellatum T

Oleaceae

Fraxinus velutina

Orchidaceae

Epipactis gigantea

Poaceae

Bromus ciliatus

Calamagrostis scopulorum

Polypogon viridis

Pteridaceae

Adiantum capillus-veneris 32

Ranunculaceae

Aquilegia chrysantha 1

Rosaceae

Petrophytum caespitosum

Rubiaceae

Galium aparine var.

echinospermum

Scrophulariaceae

Mimulus cardinalis

ZION Trail’s End HG R = 13

Asteraceae

Cirsium arizonicum var.

arizonicum

Sonchus arvensis

Sonchus asper

Berberidaceae

Berberis repens 1

Liliaceae

Maianthemum stellatum 3

Orchidaceae

Epipactis gigantea

Poaceae

Bromus diandrus

Calamagrostis scopulorum

Pteridaceae

Adiantum capillus-veneris 19

Primulaceae

Dodecatheon pulchellum

var. zionense 12

Ranunculaceae

Aquilegia chrysantha

Scrophulariaceae

AAiremidliic emrdinnlic

D hil rers

of Texas 1(1)

Vitaceae

Vitis arizonica

ZION Upper Emerald

Anac cardiaceae iac

CN m 2

Apiace

re: a T

Apocynaceae

Apocynum cannabinum 3

Asclepiadaceae

Asclepias hallii

Asclepias speciosa

Asteraceae

Artemisia ludoviciana subsp.

ludoviciana

Cirsium sp. T

Brickellia grandiflora

Sonchus arvensis

Sonchus asper

Berberidaceae

Berberis repens

Cyperaceae 3

Carex aurea

Carex curatorum

Equisetaceae

Equisetum arvense

Liliaceae

Maianthemum stellatum 20

Oleaceae

Fraxinus velutina 1

Onagraceae

Oenothera longissima

Orchidaceae

Epipactis gigantea T

Poaceae

Bromus tectorum

Calamagrostis scopulorum

Elymus canadensis 3

Muhlenbergia andina 1

Adiantum capillus-veneris

Primulaceae

Dodecatheon pulchellum

var. zionense 20

Ranunculaceae

Aquilegia chrysantha T

Aquilegia formosa var.

formosa

Rosaceae

Petrophytum caespitosum

Saxifragaceae

Heuchera rubescens

Scrophulariaceae

Mimulus cardinalis

Fowler et al., Distribution of hanging gardens in the Colorado Plateau

Mimulus guttatus

Violaceae

Viola sp. 3

Vitaceae

Vitis arizonica

ZION Weeping Rock HG

Berberidaceae

Berberis repens

Cyperaceae 7

Carex aurea

Carex curatorum

Fagaceae

Quercus gambelii

Quercus turbinella

Poaceae

Bromus diandrus T

Calamagrostis scopulorum

Muhlenbergia thurberi

Panicum sp.

Phragmites australis 2

Polypogon interruptus 8

605

Aquilegia formosa var.

formosa

Clematis ligusticifolia

Rosaceae

Petrophytum caespitosum 1

Rubiaceae

Galium multiflorum var.

multiflorum

Apocynaceae Liliaceae Pteridaceae Scrophulariaceae

Apocynum cannabinum 1 Maianthemum stellatum 7 Adiantum capillus-veneris 7 Mimulus cardinalis 30

Asteraceae Linaceae Primulaceae Tamaricaceae

Artemisia ludoviciana subsp. — Linum lewisii Dodecatheon pulchellum Tamarix ramosissima

ludoviciana 1 Oleaceae var. zionense 2 Ulmaceae

Cirsium arizonicum var. Fraxinus velutina Ranunculaceae Celtis reticulata

arizonicum Aquilegia chrysantha 7 Violaceae

Viola sp. T

ROSSI AO , x

ACKNOWLEDGMENTS

We thank C.L. May and J. Hak for their assistance and insight during the 1991-1993 field seasons. We

also thank the National Park Service staff at each of the administrative units for their logistical support,

especially V. Vieira, R. Harris, J. Belnap, and S. Petersburg. This research was supported by NPS-32752 to

Stanton. John Spence and an anonymous reviewer provided helpful review comments.

REFERENCES

Baker, V.R. 1990. Spring sapping and valley network development; with case studies by R.C. Kochel, V.R. Baker,

J.E. Laity, and A.D. Howard. In: C.G. Higgins and D.R. Coates, eds. Groundwater geomorphology; the role of

subsurface water in earth-surface processes and landforms. Special Pap. Geol. Soc. Amer. 252, Boulder, CO.

Pp. 235-244.

BanrHLorr, W. and S. Poremeski. 2000. Vascular plants on inselbergs: systematic overview. In: W. Barthlott and S.

Porembski, eds. Inselbergs, Ecological Studies Vol. 146. Springer-Verlag, Berlin. Pp. 103-116.

Bercer, W.H. and F. L. Parker. 1970. Diversity of planktonic Foraminifera in deep-sea sediments. Science 168:

1345-1347.

BungANck, M.P. and R. B. Platt. 1964. Granite outcrop communities of the Piedmont Plateau in Georgia. Ecology

45:292-306.

Cuover, E.U. and L. Jotter. 1944. Floristic studies in the canyon of the Colorado and tributaries. Amer. Midl. Natu-

ralist 32:591-642.

CocHRAN, W.G. 1977. Sampling techniques, 3rd ed. John Wiley, New York.

DAUBENMIRE, R. 1952. Forest vegetation of northern Idaho and adjacent Washington, and its bearing on concepts

of vegetation classification. Ecol. Monogr. 22:301-330.

DAUBENMIRE, R. 1959. A canopy-coverage method of vegetational analysis. Northw. Sci. 33:43-64.

Dunne, T. 1990. Hydrology, mechanics, and geomorphic implications of erosion by subsurface flow. In: C.G. Higgins

and D.R. Coates, eds. Groundwater geomorphology; the role of subsurface water in earth-surface processes

and landforms. Special Pap. Geol. Soc. Amer. 252, Boulder, CO. Pp. 1-28.

EAsTWOOD, A. 1896. Report of a collection of plants from San Juan County, in southeastern Utah. Proc. Calif. Acad.

Sci. I| 6:271—329.

ECOLOGICAL SOCIETY OF AMERICA VEGETATION CLASSIFICATION PANEL. 2006. Description, documentation, and evaluation of

associations and alliances within the U.S. National Vegetation Classification, ver. 4.5. Ecological Society of

America, Washington, DC.

FLORA NORTH AMERICA EDITORIAL Committee (eds.). 1993+. Flora of North America North of Mexico 13+ vols. Oxford

Univ. Press, New York and Oxford.

606 Journal of the Botanical h Institute of Texas 1(1)

Fow er, J.F., N.L. STANTON, R.L. HARTMAN, and C.L. May. 1995. Level of endemism in hanging gardens of the Colorado

Plateau. In: P.G. Rowlands, C. van Riper Ill and M.K. Sogge, eds. Proceedings of the second biennial conference

on research in Colorado Plateau National Parks. National Park Service Transactions and Proceedings Series

NPS-NRNAU-NRTP-95/1 1. Pp. 215-223.

Grocer, A. and W. BartHLOTT. 1996. Biogeography and diversity of the inselberg (Laja) vegetation of southern

Venezuela. Biodiversity Letters 3:165-179.

Harrison, B.F., S. L. Welsh, and G. Moore. 1964. Plants of Arches National Monument. Brigham Young University

Sci. Bull. Bio. Ser. 5:1-23.

HicalNs, C.G. 1984. Piping and sapping; development of landforms by groundwater outflow. In: R.G. Lafleur, ed.

Groundwater as a geomorphic agent, Allen and Unwin, Boston, MA. Pp. 18-58.

Horn, H.S. 1966. Measurement of “overlap” in comparative ecological studies. Amer. Naturalist 100:419-424,

Kreps, C.J. 1989. Ecological methodology. Harper and Row, New York.

Larry, J.E. and M.C. Matin 1985. Sapping processes and the development of theater-headed valley networks on

the Colorado Plateau. Bull. Geol. Soc. Amer. 96:203-217.

LoetHer, HJ. and D.G. McTavish. 1976. Descriptive and inferential statistics, an introduction. Allyn and Bacon,

Boston, MA.

MALANSON, G.P. 1980. Habitat and plant distribution in hanging gardens of the narrows, Zion National Park, Utah.

Great Basin Naturalist 40:178-182.

MaLanson, G.P. 1982. The assembly of hanging gardens: effects of age, area, and location. Amer. Naturalist

119:145-150.

Malanson, G.P. and J. Kay. 1980. Flood frequency and the assemblage of dispersal types in hanging gardens of

the Narrows, Zion National Park, Utah. Great Basin Naturalist 40:365-371.

McIntosh, R.P. 1967. An index of diversity and the relation of certain concepts to diversity. Ecology 48:392-404,

May, C.L., J.F. Fowver, and N.L. Stanton. 1995. Geomorphology of the hanging gardens on the Colorado Plateau.

In: PG. Rowlands, C. van Riper Ill and M.K. Sogge, eds. Proceedings of the second biennial conference on

research in Colorado Plateau National Parks. National Park Service Transactions and Proceedings Series NPS-

NRNAU-NRTP-95/11. Pp. 3-24.

Muetter-Domeois, D. and H. Ettenserc. 2002. Aims and methods of vegetation ecology. The Blackburn Press,

Caldwell, NJ.

NatureServe. 2006. NatureServe Explorer: An online encyclopedia of life. NatureServe, Arlington, VA. «http://www.

natureserve.org/explorer> Accessed March 2006.

POREMBSKI, S., J.SZARZYNSKI, J-P. Munp, and W. BarTHLOTT. 1996. Biodiversity and vegetation of small-sized inselbergs

in a west African rain forest (Tai, lvory Coast). J. Biogeogr. 23:47-55.

PowtLL, J.W. 1895. Canyons of the Colorado. Argosy-Antiquarian Ltd., New York.

Romme, W.H., K.D. Heit, J.M. Porter, and R. FLEMING. 1993. Plant communities of Capitol Reef National Park, Utah.

Technical Report NPS/NAUCARE/NRTR-93/02, National Park Service, Cooperative Park Studies Unit, Northern

Arizona University, Flagstaff, AZ.

SanrHOU, C. and J-F. Vittiers. 1998. Epilithic plant communities on inselbergs in French Giana. J. Veg. Sci. 9:

847-860.

SHuLTz, L.M., E.E. Ne&Lv, and J.S. Tuuv. 1987. Flora of the Orange Cliffs of Utah. Great Basin Naturalist 47:287-298.

SYSTAT. 1999a. SYSTAT" 9 Software. SPSS, Inc., Chicago, IL.

SYSTAT. 1999b. SYSTAT" 9 Statistics |. SPSS, Inc., Chicago, IL.

SZARZYNSKI, J. 2000. Xeric islands: environmental conditions on inselbergs. In: W. Barthlott and S. Porembski, eds.

Inselbergs, Ecological Studies Vol. 146, Springer-Verlag, Berlin. Pp. 37-48.

Tuy, J.L. and J.A. MacMahon. 1988. Vegetation and relict communities of Glen Canyon National Recreation Area.

Final report NPS contract CX1200-6B076. Glen Canyon National Recreation Area, National ParkService, United

States Department of the Interior, Page, Arizona, USA.

VEGBANK, 2005. The Ecological Society of America. <http://vegbank.org/vegbank> Accessed March 2006.

Fowler et al., Distribution of hanging gardens in the Colorado Plateau 607

WELSH, S.L. 1986a. New taxa and combinations in the Utah flora. Great Basin Naturalist 43:254-260.

WELSH, S.L. 1986b. New taxa in miscellaneous families from Utah. Great Basin Naturalist 43:261-264.

WELSH, S.L. 1989. On the distribution of Utah's hanging gardens. Great Basin Naturalist 49:1—30.

WetsH, S.L., N.D. Arwoob, S. GoobRicH, and L.C. Hiaains. 2003. A Utah flora, 3rd edition. Brigham Young Univ. Press,

Provo, Utah.

WeLsH, S.L. and S.L. Moore. 1968. Plants of Natural Bridges National Monument. Proc. Utah Acad. Sci. 45:

220-248.

WetsH, S.L. and C.A. Torr. 1981. Biotic communities of hanging gardens in southeast Utah. Res. Rep. Nat. Geogr.

Soc. 13:663-681.

WILKINSON, L., G. BLANK, and C. Gruser. 1996. Desktop analysis with SYSTAT. Prentice Hall, Upper Saddle River, NJ.

Woobsury, A.M. 1933. Biotic relationships of Zion Canyon, Utah with special reference to succession. Ecol. Monogr.

3:1147-245.

608 Journal of the Botanical R h Institute of Texas 1(1)

BOOK NOTICES

Rav E Evert WITH CONTRIBUTIONS BY SUSAN E. EICHHORN. 2006. Esau's Plant Anatomy: Meristems, Cells, and

Tissues of the Plant Body: Their Structure, Function, and Development, 3rd Edition. (ISBN

978-0-471-73843-5, hbk). John Wiley & Sons Inc., One Wiley Drive, Somerset, NJ 08875, U.S.A.

(Orders: www.wiley.com, 877-762-2974, 1-800-597-3299 fax). $159.95, 624 pp., b/w photos, line

drawings, 8/4" x 111A".

This is a complete redo of Esau's second edition, which is hardly recognizable except for the general organization of the chapters. The

last few chapters of edition 2 (Stem, Leaf, Root, Flower, Fruit, Seed) are gone, having been integrated into earlier discussion or in part

T PAY |

probably relegated to treatments of plant “morphology.” The format is modern and easily readable—with excellent illustrations. “A major

goal of this book is to provide a firm foundation in the meristems, cells, and tissues of the plant body, while at the same time noting

some of the many advances being made in our understanding of their function and development through molecular research. ... [It has

been] planned primarily for advanced students in various branches of plant science, for researchers (from molecular to whole plant),

and for teachers of plant anatomy."

General References include: 1) Structure and Development of the Plant Body—An Overview. 2) The Protoplast: Plasma Mem-

brane, Nucleus, and Cytoplasmic Organelles. 3) The Protoplast: Endomembrane System, Secretory Pathways, Cytoskeleton, and Stored

Compounds. 4) Cell Wall. 5) Meristems and Differentiation. 6) Apical Meristems. 7) Parenchyma and Collenchyma. 8) Sclerenchyma.

9) Epidermis. 10) Xylem: Cell Types and Developmental Aspects. 11) Xylem: Secondary Xylem and Variations in Wood Structure. 12)

Vascular Cambium. 13) Phloem: Cell Types and Developmental Aspects. 14) Phloem: Secondary Phloem and Variations in Its Structure.

15) Periderm. 16) External Secretory Structures. 17) Internal Secretory Structures. Addendum: other pertinent references not cited in

the text. Glossary; Author Index; Subject Index.

It's remarkable to see a new text for plant anatomy, as several recent publications have risen and relatively quickly gone out of

press. But in addition to Esau's classics and Evert's revision, those below are apparently now available.

Dickison, W.C. 2000. Integrative plant anatomy. Harcourt/Academic Press. (hardback)

Fahn, A. 1990. Plant Anatomy (ed. 4). (587 pages) Pergamon Press, Oxford. (used paperback on Amazon)

Mauseth, J.D. 1988. Plant Anatomy. (560 pages). Benjamin/Cummings. Menlo Park, California. (paperback on Amazon.com)

— Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A.

Davip AND SHREY Bassett. 2007. Delphiniums. (ISBN 978-0-88192-800-6, hbk.). Timber Press Inc., The

Haseltine Building, 133 SW Second Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders:

www.timberpress.com, 503-227-2878, 503-227-3070 fax, 1-800-327-5680). $29.95, 160 pp., 80

color photos, 712" x 934".

This is a DUAE done book T ao DS (Delphinium) and larkspurs o ida), mostly from the perspective of their use as

TX 1 J t 2n +] ni

garden plants I ] perennial) primarily i Hemisphere; Consolida includes about

40 species (mostly annual). The inherent native pus along with several hundred years of intensive cultivation have provided a huge

diversity in flower form and color (purple, blue, red, pink, white, yellow) and plant habit.

The book is divided into two parts: 1) Knowing Delphiniums (Introducing D; A survey of wild D; D in cultivation; Breeding D).

2) Growing Delphiniums (D in the Garden; Cultivation of D; D propagation; Growing D in containers; Pests and diseases; Hybridizing

D; Growing species).—Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A.

PETER GOLDBLATT AND DALE E. JOHNSON (eds.). 2006. Index to Plant Chromosome Numbers: 2001-2003.

(ISBN 978-1-930723-54-2, pbk.). Missouri Botanical Garden Press, PO. Box 299, Saint Louis, MO

63166-0299, U.S.A. (Orders: www.mbgpress.org, orders@mbgpress.org, 314-577-9547, 314-577-

9594 fax). $40.00, 242 pp., 7" x 10".

The latest summary in this invaluable series of chromosome counts of naturally occurring and cultivated plants, taken from publica-

tions throughout the world. Counts for this volume were extracted from 238 serials. Reviewers are from the Missouri Botanical Garden,

University of North Carolina, Hiroshima University, Komarov Botanical Institute, Royal Botanic Gardens (Kew), and the University

of Istanbul. “Chromosome counts for bryophytes and vascular plants published in the IPCN (from 1975 onward) may be accessed

through the Missouri Botanical Garden's wTROPICOS at the Garden's website, www.mobot.org.” The counts from 2001-2003 had not

been posted to the online index, as of 15 June, 2007.—Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX

76102-4060, U.S.A.

J. Bot. Res. Inst. Texas 1(1): 608. 2007

BEESASCUEARTEORAOBIBHELBANCOSGKDBIOEOGICAL STATION.

MURRAY STATE UNIVERSITY C AL TOWA C OUNT KENTUCKY

Ralph L. Thompson

Hancock Biological Station

Murray State University, Murray, KENTUCKY 40271, U.S.A.

Berea College Herbarium

Department of Biology, Berea College, Berea, Kentucky 40404, U.S.A.

ralph_thompson@berea.edu

ABSTRACT

Th 1 £1 IT SR a EST

l Station, Murray State University, was surveyed throughout the growing seasons of 1998-1999

and during June 2000, 2001, 2002, He 2006. The 37.5-ha tract lies 23 km from Murray, Kentucky, in northeastern Calloway County

contiguous to Kenlake State Resort Park to the north and adjoins the Kentucky Lake shoreline to the east. The study site is situated

within the Jackson Purchase of western Kentucky. Vegetation is predominately upland dry and dry-mesic oak-hickory forest. Burned

warm-season grassland, early to mid-successional areas, culturally-disturbed areas, and wetland areas are other diverse habitats. Vascular

plants consist of 573 specific and infraspecific taxa in 334 genera from 121 families. A total of 469 are native and 104 are exotic species.

Of the exotics, 47 are Kentucky invasive pest plant species. Taxonomic representations are one Lycopodiophyta, one Equisetophyta,

eight Polypodiophyta, four Pinophyta, and 559 Magnoliophyta.

Key Wonps: Hancock Biological Station, vascular flora, habitats, oak-hickory forest, field station; invasive exotics, Kentucky Lake

RESUMEN

La flora vascular del Centro Biológico Hancock de la Universidad Estatal de Murray tudió durante las temporadas de crecimiento de

1998-1999 y durante junio de 2000, 2001, 2002, y 2006. El terreno de 37,5 hectáreas está ubicado a 23 kilómetros de Murray, Kentucky

en la parte noreste del condado de Calloway contiguo al centro recreativo Kenlake State Resort Park hacia el norte y junto a la ribera

del lago Kentucky hacia el este. El terreno que se ha investigado está situado dentro del Jackson Purchase del oeste de Kentucky. La

vegetación que predomina es el bosque seco-húmedo de roble-nogal americano de las tierras altas. Paraderas quemadas en la estación

templada, áreas de sucesión temprana o mediana, áreas afectadas por cultivos y áreas húmedas son otros de los hábitats. Las plantas

vasculares son 573 grupos taxonómicos específicos e infraespecíficos de 334 géneros de 121 familias. Un total de 469 son nativas y

104 son especies exóticas. De las exóticas, 47 son especies de plantas invasoras en Kentucky. Representaciones taxonómicas son una

Lycopodiophyta, una Equisetophyta, ocho Polypodiophyta, cuatro Pinophyta y 559 Magnoliophyta.

INTRODUCTION

Hancock Biological Station (HBS), a 37.5-hectare tract of upland Oak-Hickory Forest adjacent to Kentucky

Lake, is the biological field station of Murray State University, Murray, Kentucky (Fig. 1). HBS is a member of

the Organization of Biological Field Stations, a consortium of 220 biological field stations in North America

(OBFS 2006) and a member of the Association of Ecosystems Research Centers. HBS was founded in 1966

through the efforts of Hunter M. Hancock, former Murray State University professor and chair of the Depart-

ment of Biological Sciences (White 2002). Since 1972, HBS has served as a year-round facility for aquatic and

terrestrial biology research and service programs, and it has presented students with opportunities for field

classes, independent research, and faculty-directed undergraduate and graduate research (White 2002).

In the early 1980s, HBS and the Land Between the Lakes (LBL) were designated as an Experimental

Ecological Reserve by the National Science Foundation and the Institute of Ecology. As an Experimental

Ecological Reserve, HBS and LBL serve as an important natural system for long-term ecological research

(White 2002). HBS currently serves as the primary field research facility for the Center for Reservoir Research

(CRR) established in 1987 by the Commonwealth of Kentucky. Among the CRR’s goals is the improvement

in knowledge needed to manage, protect, and preserve the environmental quality of reservoir resources. The

CRR has become nationally recognized in basic and applied aquatic research and education by providing

facilities and a permanent research technical staff (White 2002).

J. Bot. Res. Inst. Texas 1(1): 609 — 630. 2007

610 Journal of the Botanical Research Institute of Texas 1(1)

o 250

meters

V + I Lal AA tran £ +L D Le lal LN J I 75 :

Fic. 1. Hancock Biological Station, Calloway County, Kentucky, y

series, 1950, United States Geological S Washington, D.C., and Mid-America Remote Sensing Center (20032).

J y!

To further serve education, in the mid-1980s the Ecological Consortium of Mid-America (ECOMA) was

formed among several colleges and universities. The purposes of ECOMA were to utilize the HBS facilities,

the LBL resources, and the Kentucky Lake-Barkley Lake complex in undergraduate and graduate teaching,

to facilitate service programs, and to serve as a base of operation for field trips and research throughout the

year (White 2002).

The Hancock Biological Station was one of five major collection sites for a master's thesis of the vascu-

lar flora of Calloway County by Woods (1983). After additional collections were added to the Murray State

University Herbarium, the vascular flora of Calloway County was published by Woods and Fuller (1988).

To do a thorough floristic survey of just HBS, the objectives of the current descriptive study were to 1)

document the HBS vascular flora with voucher specimens, 2) depict the physical site, 3) describe the plant

Thompson, Flora of H k Biological Station in Kentucky 611

habitats, and 4) present a complete annotated list of the vascular plants with origins, habitats, and relative

abundance values.

THE STUDY- SITE

History and Facilities

Hancock Biological Station lies between latitudes 36*4424" and 36*44'00" N and between longitudes

8807730" and 88*0652"W within the 7.5-minute series Rushing Creek Quadrangle (Fig. 1). Prior to the

1920s, the oak-hickory forests west of the Tennessee River were completely harvested with much of the

wood used as fuel for the iron furnaces in the LBL region. Through the 1940s, a large floodplain existed on

the west bank of the Tennessee River in front of the present station, and most of the terrain in the uplands

was pastures, cultivated fields, or scattered woodlands. The completion of Kentucky Dam on the Tennessee

River in 1944 formed Kentucky Lake and impounded much of the upland terrain. In the past 80 years, the

present vegetation has evolved from a combination of secondary forest succession and human activities that

have limited natural habitats at HBS through the creation of Kentucky Lake .

In 1966, the original station grounds consisted of 16.2 ha of abandoned fields, pastures, and oak-hickory

forest stands to the south of Kenlake State Resort Park. An agreement in the late 1960s between Murray

State University and the Tennessee Valley Authority (TVA) provided an additional 13.3 ha of land from the

114 m TVA boundary upward. In 1988, Kenlake State Resort Park transferred an additional 8.0 ha to HBS

(White 2002). The total HBS tract is currently estimated at 37.5 ha.

HBS facilities currently consist of 26 buildings. Facilities include the main laboratory and classroom

building, glasshouse/mesocosm building, a resource building, boat house, bath house, 15 student cabins,

four faculty cabins, maintenance shop, and well house, as well as a picnic area and wastewater wetland

complex (Fig. 2). The station is reached at the end of the asphalt-paved Emma Drive that leads to Lancaster

Road, KY 497, and then to KY 94. Watersport Road leads from the Pacer Point Recreation Area, passes

through the western portion of the HBS property, and connects with Emma Drive. Elevation at HBS ranges

from 107.9 m at the Kentucky Lake shoreline to the 114.3 m Tennessee Valley Authority boundary to a 143

m ridge crest just west of the junction of Emma Drive and Watersport Road (Fig. 1).

Physiography

HBS is located in the Jackson Purchase or the Mississippi Embayment Section of the East Gulf Coastal Plain

based on Fenneman (1938). Keys et al. (1995) classified the area west of the Tennessee River (the Kentucky

Lake impoundment) as belonging to the Deep Loess Hills and Bluffs Subsection of the Upper Gulf Coastal

Plain Section of the Eastern Broadleaf Forest Province. Woods et al. (2002) designated the hilly terrain west

of Kentucky Lake as a part of the Western Highland Rim extending eastward through the Tennessee River

and Cumberland River Valleys.

Geology

The geology at the study site includes alluvium, loess, and cherty limestone bedrock of the Quaternary, Cre-

taceous, and Mississippian Carboniferous Series (Seeland and Wilshire 1965; Fig. 3). The exposed flattened

ridges and rolling hills are covered with unstratified, clayey, silty loess from the Quaternary Pleistocene

that covers continental sand and gravel deposits. Sand, gravel, and clay of the Upper Cretaceous McNairy

Formation are found in the southwestern corner of HBS. In a small area by Kentucky Lake, gravel, clay, and

clayey silts of the Upper Cretaceous Tuscaloosa Formation overlie Mississippian bedrock. Thick-bedded

cherty limestone of the Mississippian Warsaw Limestone Formation is in the extreme northwestern part of

the boundary. The largest amount of bedrock at HBS is composed of cherty, fine-grained limestone of the

Mississippian Fort Payne Formation. The Fort Payne System bedrock is exposed along the steep cliff line at

Kentucky Lake. Quaternary Pleistocene and Recent alluvium comprised of silt, sand, and stratified cherty

gravel is found in three lowland valley coves adjacent to Kentucky Lake (Fig. 3).

612 Journal of the Botanical Research Institute of Texas 1(1)

PIE nia BIOLOGICAL

as Mi anto STATION

— Welt 7". J € Psocosm area

qe. To KY oe ie y ssn) a Shed

rol z ° A. n al Kentucky

Student Cabins

E # Main Building

* $

a €

M Ert

x ey a

. * : ML $ id EN

Pay a Se: h " +

OS " be PicniciArea ^.

+ A JB j; Wetland

t, bi $ ¿o Complex

br.»

Boathouse,” ~N AN

e P

Fic. 2. The physical site facilities of the Hancock Biological Station, directly f

not to scale). See Figure 1 for facilities location on the property

http:// ky.edu/hbs (Murray State University 2006). (Map

Soils

The principal soil association of HBS and vicinity is the Bodine-Brandon Association (Humphrey et al. 1973;

Fig. 4). This soil association predominates on steep to sloping, well-drained to excessively drained, silty

cherty uplands. Bodine series are acid to strongly acid (4.5—5.0 pH), well-drained or excessively drained

residual cherty limestone soils from the Warsaw Limestone and Fort Payne Formations. These soils are

located on 12-60 percent upper to middle slopes and side slopes leading to the Kentucky Lake shoreline.

Bodine topsoils are brown cherty silt loams to 13 cm and subsoils are yellowish-brown, cherty silty loams

from 15-57 cm, and yellowish-red, very cherty, silty clay loams from 58-157 cm deep. The Brandon series

occupy 6—30 percent rolling upper elevation side slopes and flattened ridges at HBS (Fig. 4). These soils are

acid to strongly acid (4.5—5.5 pH), well-drained, and are developed in 0.6—1.2 m of loess. Brandon topsoils

consist of brown silty loams to 24 cm deep, subsoils of yellowish-red silty clay loams from 25-69 cm, and

Coastal Plain gravelly brown loams from 71-127 cm in depth (Humphrey et al. 1973).

A thin band of Saffell series lies between Bodine and Brandon soils on a ridge with 6-12 percent slopes

in the west central portion of HBS. Saffell topsoils are acid to strongly acid (4.5—5.5 pH), well-drained, yel-

lowish-brown, very gravelly silt loams 25 cm deep. The subsoils are yellowish-red gravelly loams from 26-88

cm and very gravelly brown sandy loams from 90-150 cm. The Ochlockonee series lies on the 0—4 percent

sloping alluvial valley floodplain in the southernmost part of the study site near Pacer Point Recreation Area.

Ochlockonee topsoils are strongly acid (5.1-5.5 pH), well-drained, brown silt loams to 18 cm. Subsoils are

Thompson, Flora of Hancock Biological Station in Kentucky 613

A x, ot

" Qal

o 250

1 i

meters

Fic 2 ge | Lal Li Il. D? | . ICa PR RA Pe 1 £. c I J DA and H G LT YH PN e LARAL EN JAA? A . n 4 c . r "1 AAAS LY

3. 3

234 J 7 V 7 J V 1

lal J A fr | nh l'a t Ia Fal 4 RA_AI * r Va Ia T r

gy Qal =Q y ;Q y ; Upper y Formation; Kt = Upper

+ AA is r pn r

Mv ppian Warsaw Limestone; Mfp

brown sandy loams from 18-89 cm and gravelly sandy loams from 90-127 cm in depth (Humphrey et al.

Vegetation

The forest vegetation in the Jackson Purchase is predominantly Oak-Hickory Forest (Küchler 1964; Bryant

and Held 2001; Woods et al. 2002). Braun (1950) included the Jackson Purchase (Mississippi Embayment

Section) in her Western Mesophytic Forest Region based on the mixed mesophytic vegetation composition of

the western loess bluffs. Braun (1950) noted that she would have placed the vegetation in her Oak-Hickory

Forest Region except for these western loess bluffs. The forest vegetation of Hancock Biological Station is

currently a mixture of dry oak-hickory forest and dry-mesic oak-hickory forest (Fig. 5).

614 Journal of the Botanical Research Institute of Texas 1(1)

meters

Fic. 4. Soils of the Hancock Biological Station. Modified from Humphrey et al. (1973) ang iii Ameria Remote Sensing Center (2003c). Soil code: Bo

= Bodine cherty silt loams, 12-60% slopes; Br = Brandon silt loam, 6—3096 slopes; Oc gravelly loams, 0—496 slopes; Sgc — Saffell very

gravelly silt loams, 6-12% slopes.

Climate

Climate of the Jackson Purchase is a humid temperate continental type characterized by warm to hot sum-

mers and cool to moderately cold winters. Climatic data (1971-2000) are from the United States Department

of Agriculture, Forest Service Weather Station at Golden Pond, 13 km east-northeast from HBS. The mean

annual temperature is 14.9° C. January is the coldest month at 1.2? C, and July is the warmest month at

26.0” C. The length of the growing season averages 209 days from the median first frost on October 27 to the

median last freeze on April 6. Mean annual precipitation is 127 cm and is fairly well distributed throughout

the year. August is the driest month at 8.1 cm of precipitation and December is the wettest at 12.9 cm. The

mean annual snowfall in January, February, and March is 10.8 cm (Kentucky Climate Center 2006).

Thompson, Flora of Hancock Biological Station in Kentucky 615

9 meters

Fi. 5. Habitats of the H k Biological Station. Habitat code: 1 = dry oak-hickory forest, 2 = dry-mesic oak-hickory forest, 3 = upland early and mid-

successional areas, 4 — burned old-field grassland, 5 — culturally disturbed , 6 = pond and roadside ditches, 7 = wetland complex,

8 = riparian forest, 9 gent h and wetland low, 10 = seasonal dewatered shoreline.

METHODS

A floristic survey was conducted during the growing seasons from March—November 1998 and 1999 with

additional collections in June 2000, 2001, 2002, and 2006. Vascular plants were identified using Mohlen-

brock (1986), Gleason and Cronquist (1991), and Jones (2005). Arrangement of families and nomenclature

follows Jones (2005). Vernacular names are derived from a combination of Jones (2005) and USDA, NRCS

(2006). Plants were collected in duplicate with the master set deposited into the Berea College Herbarium

(BEREA) and the second set placed in the herbarium of Hancock Biological Station, a part of the Murray

State University Herbarium (MUR). Plant habitats were delineated through field reconnaissance and field

collections in conjunction with topographic-moisture features, soil type, underlying geology, vegetation

616 Journal of the Botanical R h Institute of Texas 1(1)

(dominant and associated species of the canopy, subcanopy, shrub, and herbaceous layers), and anthropo-

genic disturbances.

A relative abundance value is assigned each taxon inclusive throughout all HBS habitats. Relative abun-

dance categories modified from Thompson and Poindexter (2006) are Rare—1 to 4 individuals or colonies,

Scarce—5 to 10 individuals or colonies, Infrequent—11 to 30 individuals or colonies, Occasional—31 to

100 individuals, Frequent—101 to 1000 individuals or colonies, and Abundant—1000s of individuals or

colonies.

RESULTS AND DISCUSSION

Taxonomic Summary

The annotated list from the boundaries of HBS includes 573 specific and infraspecific taxa in 334 genera

from 121 families (Table 1). Taxonomic representation is one Lycopodiophyta, one Equisetophyta, eight

Polypodiophyta, four Pinophyta, and 559 Magnoliophyta (411 Magnoliopsida and 148 Liliopsida). A total

of 104 (18.2 %) were exotic taxa (Table 1). Thirty species, native or exotic, were deliberately planted on the

study site. The largest families in species richness are Asteraceae (72), Poaceae (68), Fabaceae (41), Cyperaceae

(36), Lamiaceae (21), Rosaceae (17), and Scrophulariaceae (16). The largest genera are Carex (20), Quercus

(11), Dichanthelium (10), Juncus (8), Lespedeza (8), Polygonum (8), and Hypericum (7). One hundred-five are

woody plants (64 trees, 22 shrubs, and 19 vines), and 468 are herbaceous plants (148 annuals, 17 bienni-

als, and 303 perennials).

Woods (1983) listed 912 species, 428 genera, and 119 families from his M.S. thesis on the vascular

flora of Calloway County. After more collections, Woods and Fuller (1988) increased the number to 1018

species, 462 genera, and 129 families. In this study at HBS, 573 specific and infraspecific taxa account for

56.396 of the total Calloway County flora based on Woods and Fuller (1988). This number also comprises

22.096 of the 2600 known vascular plant species of Kentucky based on Jones (2005).

Thirty-nine new Calloway County records were documented in the present study from a search of MUR,

the vascular plant atlas of Campbell et al. (2006), and the distribution maps from the USDA, NRCS (2006).

Trepocarpus aethusae, occasional in relative abundance along the Kentucky Lake shoreline, was state-listed

“Threatened” in 2000 (KSNPC 2000) and state-listed “Special concern" in 2005 (KSNPC 2006). Aphanes

microcarpa, an exotic European annual, was first documented in Kentucky from Madison County (Abbott

et al. 2001). The collection from HBS is the second county recorded for Kentucky.

Invasive Exotic Species

In Kentucky, 94 invasive exotic plants belong to the “severe threat” (29 species), “significant threat” (33

species), and “lesser threat" (32 species) classifications (Kentucky Exotic Pest Plant Council 2000). The 104

naturalized species at Hancock Biological Station have a definite impact upon the native flora, vegetation,

and habitats. Forty-seven (45.2%) of the 104 exotics are invasive pest plants. Twenty-nine species (27.9%)

belong to the “severe threat" (14 species) and “significant threat" (15 species) categories. Several of these

invasive exotics have become naturalized on the Hancock premises thorough deliberate plantings.

The most notable "severe threat" exotics affecting HBS native vegetation are the abundant and widespread

Festuca arundinacea, Lespedeza cuneata, Lonicera japonica, and Microstegium vimineum. Among other “severe

threat" taxa are Coronilla varia, Elaeagnus umbellata, Ligustrum sinense, Rosa multiflora, Sorghum halepense,

and Stellaria media. *Significant threat" exotics include Daucus carota, Eleusine indica, Glechoma hederacea,

Hedera helix, Lespedeza stipulacea, L. striata, Poa pratensis, Polygonum caespitosum, P. persicaria, Seteria faberi,

and Vinca minor. The 104 exotic species will continue to have deleterious effects of displacing and replacing

native species, disrupting nutrient cycles, and changing the pattern of plant succession.

Plant Habitats

Hancock Biological Station is delineated into 10 habitats: five terrestrial and five wetland. Terrestrial habitats

are dry oak-hickory forest, dry-mesic oak-hickory forest, upland early and mid-successional areas, burned

old field warm-season grassland, and culturally disturbed areas (Fig. 5). The dry oak-hickory and dry-mesic

Thompson, Flora of H k Biological Station in Kentucky 617

TABLE 1. Taxonomic distribution of vascular plants at the Hancock Biological Station, Kentucky.

Division Families Genera Species Native Exotic Percent Species

Composition

Equisetophyta ] 1 1 1 0 0.17

Lycopodiophyta 1 1 1 1 0 eI

Polypodiophyta 4 7 8 8 0 1.40

Pinophyta 2 3 4 4 0 0.70

Magnoliophyta 101 322 559 455 104 97.56

Magnoliopsida 87 248 411 237 74 MS

Liliopsida 15 74 148 Ne 30 25:83

Totals: 121 334 nz 469 104 100.00

oak-hickory forests are representative of the two types of oak-hickory forest in the Jackson Purchase Area

described by Bryant and Held (2001).

The five wetland areas are pond and roadside ditches, wetland complex, riparian forest, emergent marsh

and wetland meadow, and seasonally dewatered shoreline (Fig. 5). Transitional ecotones tend to exist between

adjacent terrestrial and terrestrial, terrestrial and wetland, and wetland and wetland habitats. These 10 plant

habitats are described and characteristic species or indicator species within each habitat are listed.

Dry oak-hickory forest —The woody vegetation of open dry flat to rolling topography on southern and

western trending upper slopes, ridge tops, and higher elevations is characterized by dry oak-hickory forest

(Fig. 3). Forest soils include Bodine cherty silt loams covered by Quaternary loess and some areas of Bran-

don silt loams. Quercus stellata, Q. marilandica, Q. falcata, and Carya glabra are canopy indicator trees. Other

important overstory trees include Acer rubrum, Carya tomentosa, Nyssa sylvatica, Q. velutina, and Ulmus alata.

The understory layer is somewhat scrubby and sparse. Among the shrubs and small trees are Amelanchier

arborea, Vaccinium arboreum, V. stamineum, and Viburnum rufidulum. A single clump of the hemiparasitic shrub,

Phoradendron leucarpum, was observed in one Carya glabra. Woody vines include Parthenocissus quinquefolia,

Smilax bona-nox, S. glauca, Toxicodendron radicans, and Vitis aestivalis.

Several native herbaceous perennials are found beneath the dry oak-hickory forest. Characteristic pe-

rennial herbs include Antennaria plantaginifolia, Asplenium platyneuron, Aureolaria flava, Carex muehlenbergii,

Comandra umbellata, Coreopsis major, Cunila origanoides, Danthonia spicata, Dichanthelium acuminatum, D. boscii,

D. laxiflorum, Euphorbia corollata, Galium circaezans, Houstonia canadensis, Hypoxis hirsuta, Lechea tenuifolia,

Orbexilum pedunculatum, Porteranthus stipulatus, Pteridium aquilinum var. latiusculum, Sericocarpus linifolius,

Scutellaria parvula, Symphyotrichum patens, Tephrosia virginiana, and Viola pedata. In certain exposed areas,

fruticose lichens (Cladina spp. and Cladonia spp.) and cushiony mosses are abundant.

Dry-mesic oak-hickory forest.—These forest stands occur on more mesic, steeper northern, western and

eastern trending middle to upper side slopes of shallow valleys and valley coves, and adjoin the Kentucky

Lake shoreline (Fig. 5). Forest soils are Brandon silt loams from the Warsaw Limestone and the Fort Payne

Formations. Topographic-moisture conditions, slope aspect, and soil types are extremely important in the

transitional mosaic of dry-mesic oak-hickory forest to dry oak-hickory forest relative to species composition.

A considerable intergradation of woody and herbaceous species exists between these two oak-hickory forest

types. Quercus alba is the dominant canopy tree. Other indicator canopy trees of dry-mesic oak-hickory

forest include Quercus rubra, Q. velutina, Carya ovata, C. tomentosa, Acer rubrum, Fraxinus americana, Nyssa

sylvatica, Prunus serotina, and Ulmus rubra. Diospyros virginiana, Juniperus virginiana, Morus rubra, Sassafras

albidum, and Ulmus alata are scattered in the stand. Subcanopy trees are Cercis canadensis and Cornus florida.

Characteristic woody vines and shrubs are Aralia spinosa, Asimina triloba, Parthenocissus quinquefolia, Rosa

carolina, Smilax bona-nox, S. glauca, Staphylea trifolia, Toxicodendron radicans, Vaccinium stamineum, Vitis aes-

tivalis, and V. rotundifolia.

£s+haD o ID L

618 Journal of t titute of Texas 1(1)

Many perennial herbs are especially evident during spring and summer. Characteristic herbs include

Anemonella thalictroides, Brachyelytrum erectum, Bromus pubescens, Desmodium nudiflorum, Dioscorea villosa,

Galium triflorum, Luzula bulbosa, Scutellaria elliptica, Spigelia marilandica, and Viola palmata. Other perenni-

als of more mesic, shaded side slopes and valley bottoms are Carex albicians, C. blanda, C. digitalis, Dentaria

laciniata, Iris cristata, Maianthemum racemosum, Podophyllum peltatum, Polygonatum biflorum, Polystichum

acrostichoides, Solidago caesia, and Uvularia sessilifolia. Representative herbs of the three alluvial valley coves

adjoining the riparian forest include Boehmeria cylindrica, Carex grayi, C. typhina, Chasmanthium latifolium,

Elymus virginicus, Iris virginica, Phegopteris hexagonoptera, Phryma leptostachya, Pilea pumila, Polygonum virgin-

ianum, and the abundant “severe threat” Microstegium vimineum.

Upland early and mid-successional areas.—Most of the area in early and mid-successional stages have

developed from a large abandoned pasture of native grasses, forbs, and dry woodland species. Shade intoler-

ant, successional woody species are prevalent in the old pasture and the other disturbed habitats that include

dry and dry-mesic oak-hickory forest edges, a narrow power line corridor-cut bordering Emma Drive, and

a small 25 year old planted loblolly pine stand (Fig. 5). If succession continues without significant distur-

bance, these areas will progress toward an oak-dominated forest with a hickory component. Invading trees

present are Acer rubrum, Albizia julibrissin, Diospyros virginiana, Juniperus virginiana, Nyssa sylvatica, Sassafras

albidum, and Ulmus alata. The undergrowth is variable and ranges from sparse to densely vegetated areas.

Characteristic successional shrubs include Rhus copallina, R. glabra, Rosa setigera, Rubus argutus, R. flagellaris,

and Symphoricarpos orbiculatus. The ubiquitous woody vines are abundant Lonicera japonica, Toxicodendron

radicans, and Vitis rotundifolia.

A combination of annuals, biennials, and perennials in the summer and fall aspects are present in the

successional areas including several from the Asteraceae, Fabaceae, and Poaceae. Several tall grass prairie

species are persisting in the old pasture. Characteristic species in these successional areas are Ambrosia

artemisiifolia, Andropogon virginicus, Daucus carota, Dichanthelium polyanthes, Diodia teres, Festuca arundina-

cea, Erigeron annuus, Eupatorium serotinum, Galium pilosum, Lespedeza cuneata, L. intermedia, L. procumbens,

Monarda fistulosa, Pyncnanthemum tenuifolium, Potentilla simplex, Schizachyrium scoparium, Setaria parviflora,

Solidago canadensis, Symphyotrichum dumosum, S. pilosum, Verbesina helianthoides, and Vernonia missurica.

Burned old field warm season grassland.—This open dry habitat has several tall grass prairie species and

forbs; but, it never was a part of the Midwestern Tall Grass Prairie Region. The habitat (1450 m?) was initially

derived from an abandoned field with upland dry forest soils, and it is now completely enclosed by dry and

dry-mesic oak-hickory forest (Fig. 5). The habitat is representative of the warm season grassland barrens

described for Land Between the Lakes by Martin and Taylor (2002). HBS personnel prescribed-burn the site

every 1-2 years to preserve a non-forested or grassland habitat and to enhance the persisting warm season

prairie elements. Secondary successional woody invaders suppressed by fire in the old field are Acer rubrum,

Diospyros virginiana, Juniperus virginiana, Liquidambar styraciflua, Liriodendron tulipifera, Nyssa sylvatica, Rhus

copallina, R. glabra, Rubus argutus, Smilax bona-nox, and S. glauca.

Species composition in this fire-maintained habitat is similar to the warm season grassland of the Elk

and Bison Prairie of Land Between the Lakes National Recreation Area (Thompson and Poindexter 2006).

Indicator species present are Andropogon ternarius, Asclepias tuberosa, Carex hirsutella, Ceanothus americanus,

Coreopsis major, Crotalaria sagittalis, Hypericum denticulatum, Euphorbia corollata, Linum medium var. texanum,

Lobelia puberula, Parthenium integrifolium, Polygala sanguinea, P. ambigua, Pycnanthemum tenuifolium, Oenothera

fruticosa, Rudbechia hirta, Scleria pauciflora, S. triglomerata, Schizachyrium scoparium, Sorghastrum nutans,

Stylosanthes biflora, and Tripsacum dactyloides.

Culturally disturbed areas —Anthropogenic-influenced habitats include the mowed irregular-shaped

station yard, the mowed Emma Drive road shoulder, the Boy Scout trail, faculty cabins trail, the ruderal

graveled area around the glasshouse/mesocosm and the gravel roads to student cabins, Wolfson House, and

boat house (Figs. 2, 5). Many exotic and native annuals and perennials have become established in these

disturbed grassy and gravelly areas. The preeminent taxon is the “severe threat” Festuca arundinacea. Other

characteristic species include Bromus commutatus, Cardamine hirsuta, Cerastium glomeratum, Cynodon dactylon,

Thompson, Flora of H k Biological Station in Kentucky 619

Dactylis glomerata, Dichanthelium laxiflorum, Digitaria sanguinalis, Eleusine indica, Gamochaeta purpurea, Juncus

tenuis, Lespedeza cuneata, L. stipulacea, L. striata, Medicago lupulina, Oxalis stricta, Plantago lanceolata, P. rugelii,

Poa pratensis, Stellaria media, Taraxacum officinale, Trifolium dubium, T. repens, and Veronica arvensis.

Pond and roadside ditches —In 2000, Hancock Pond (1300 m?) was created within an old successional

pasture 20 m from the HBS entrance gateway and 25 m north of Emma Drive (Fig. 5). The borrowed soil was

used as fill for the glasshouse/mesocosm (Fig. 2). The pond readily filled with water and hydrosere succes-

sion has progressed rapidly for the last six years. Invading emergent species include Cyperus pseudovegetus,

Eleocharis ovata, Hypericum mutilum, Juncus acuminatus, J. brachycarpus, J. diffusissiimus, J. effusus var. solutus,

Ludwigia alternifolia, Scirpus cyperinus, and Typha latifolia. Salix nigra is currently the only woody volunteer.

The roadside ditches along Emma Drive and Watersport Road have a few wetland plants established in-

cluding Carex lurida, Eleocharis ovata, Juncus biflorus, J. effusus var. solutus, Ludwigia alternifolia, Salix nigra,

Scirpus cyperinus, and Typha latifolia. These wetland plants from the ditches undoubtedly provide a viable

seed source for hydrosere pond succession.

Wetland complex.—An artificial-designed gravel-covered wetland (5 m by 40 m or 200 m?) was built

in 1990 for the station wastewater (Fig. 5). This wetland complex was initially planted with several native

wetland species, and other native and exotic wetland species have volunteered. Established wetland species

include Boehmeria cylindrica, Carex crinita, C. franhii, C. lupulina, C. vulpinoidea, Diodia virginiana, Equisetum

hyemale, Impatiens capensis, Hemerocallis fulva, Iris virginica, I. pseudoacorus, Justicia americana, Leersia oryzoides,

Onoclea sensibilis, Pontederia cordata, Polygonum sagittatum, Schoenoplectus tabernaemontani, Scirpus atrovirens,

S. cyperinus, Thalia dealbata, and Typha latifolia.

Riparian forest.—Riparian forest is scattered along the Kentucky Lake shoreline. It abuts dry-mesic oak

hickory forest borders including three low relief, mesic valley bottoms or coves at Pacer Point cove, boat

dock cove, and the north peninsula cove (Fig. 5). The Kentucky Lake shoreline is composed of Fort Payne

cherty limestone gravel, sand, and silt. Shoreline habitats are annually flooded during the winter and spring

months. Riparian indicator trees are Acer negundo, A. saccharinum, Betula nigra, Liquidambar styraciflua,

Platanus occidentalis, and Salix nigra with a few Populus deltoides, Taxodium distichum, two Quercus lyrata, and

one Nyssa aquatica. At the edge of alluvial valley coves and the gravelly shoreline, Alnus serrulata, Amorpha

fruticosa, Cephalanthus occidentalis, Cornus amomum, Ilex decidua, and Styrax americana are characteristic

shrub-swamp species. Entangled woody vines on trees and shrubs consist of Bignonia capreolata, Brunnichia

ovata, Campsis radicans, Smilax rotundifolia, Toxicodendron radicans, Vitis palmata, V. rotundifolia, and Wisteria

frutescens. Characteristic herbs of the riparian forest include many from the dry-mesic oak-hickory forest

valley coves and the seasonally dewatered shoreline gravel, sand, and mudflats.

Emergent marsh and wetland meadow.—With nearly level relief and saturated soils, a seasonally flooded

emergent marsh intermixed with a sedge-grass wetland meadow has developed between HBS Pacer Point

cove and Pacer Point Recreation Area (Fig. 5). Riparian shrubs and trees are typically missing. A combina-

tion of emergent marsh and meadow species include Alternanthera philoxeroides, Ammannia coccinea, Carex

frankii, C. lupulina, C. tribuloides, Cyperus strigosus, Eleocharis acicularis, Fimbristylis autumnalis, Hibiscus laevis,

Hypericum mutilum, Juncus effusus var. solutus, Justicia americana, Leersia oryzoides, Lindernia dubia, Lycopus

virginicus, Mimulus alatus, Panicum rigidulum, Phyla lanceolata, Polygonum sagittatum, Rotala ramosior, and

Rhynchospora corniculata.

Seasonal dewatered shoreline —The Kentucky Lake shoreline habitat ranges from steep cherty limestone

erosion areas several meters high connecting to oak-hickory forest, to a nearly level or slightly level band of

seasonally dewatered gravel, sand, silt, and clay mudflats. The sparsely vegetated shoreline is most prominent

in the fall when the water level of Kentucky Lake is lowered to 107.9 m (winter pool) and then disappears

when the lake is raised to 109.4 m in spring (summer pool). Among the many native and exotic characteristic

annuals are Acalypha virginica, Amaranthus rudis, A. tuberculatus, Bidens frondosa, B. vulgata, Diodia virginiana,

Echinochloa crusgalli, Eclipta alba, Euphorbia maculata, E. nutans, Ipomoea lacunosa, Myosurus minimus, Panicum

dichotomiflorum, Polygonum caespitosum var. longisetum, P. persicaria, P. pennsylvanicum, and Sida spinosa.

620 Journal of the Botanical R h Institut

of Texas 1(1)

ANNOTATED LIST OF PLANTS

The annotated list of the vascular flora is arranged alphabetically by family, genus, and species in the Pteri-

dophyta, Pinophyta, and Magnoliophyta (Magnoliopsida and Liliopsida). Nomenclature follows Jones (2005).

An asterisk (*) preceding a scientific name indicates an exotic or non-indigenous taxon. A double asterisk

(**) indicates an invasive exotic plant for Kentucky from the Kentucky Exotic Pest Plant Council (2000). A

dagger (+) represents a planted native or exotic taxon at HBS. A diesis or double dagger (t) indicates a new

Calloway County distribution record. After the scientific name, plant habitat(s) are given in a numbered

code: l=dry oak-hickory forest, 2=dry-mesic oak-hickory forest, 3=upland early and mid-successional areas,

4=burned old field warm season grassland, 5=culturally disturbed areas, 6=pond and roadside ditches,

7=wetland complex, 8=riparian forest, 9=emergent marsh and wetland meadow, and 10=seasonal dewatered

shoreline. Relative abundance values, Rare (R), Scarce (S), Infrequent (D, Occasional (O), Frequent (E), and

Abundant (A), follow habitat(s). An italicized representative voucher number by the author or other collector

ends the entry for each species.

EQUISETOPHYTA

Equisetaceae

tEquisetum hyemale L. var. affine (Engelm.) Calder & R.L. Taylor,

Common scouring-rush, 7; |; 98-87

LYCOPODIOPHYTA

Lycopodiaceae

Lycopodium digitatum Dill., Southern ground cedar, 3; R;

99-318

POLYPODIOPHYTA

Aspleniaceae

Asplenium platyneuron (L.) B.S.P., Ebony spleenwort, 1; |;

Dennstaedtiaceae

Pteridium aquilinum (L.) Kuhn var. latiusculum (Desv.) Underw.,

Western bracken fern, 1, 2; O; 99-237

Dryopteridaceae

tOnoclea sensibilis L., Sensitive fern, 7; O; 02-213

Polystichum acrostichoides (Michx.) Schott, Christmas fern,

2; O; 99-227

Woodsia obtusa (Spreng. Torr., Bluntlobe cliff fern, 2; |;

98-190

Ophioglossaceae

Botrychium dissectum Spreng., Cutleaf grape fern, 2; |; 98-718

Botrychium virginianum (L.) Sw., Rattlesnake fern, 2; |; 06-230

Thelypteridaceae

Phegopteris hexagonoptera (Michx.) Fee, Broad-beech fern,

2:5: 06-25]

PINOPHYTA

Cupressaceae

Juniperus virginiana L., Eastern redcedar, 1, 2, 3; O; 98-101

Taxodium distichum (L.) Rich., Bald cypress, 8; S; 98-406

Pinaceae

tPinus taeda L., Loblolly pine, 3; O; 00-157

tPinus virginiana Mill., Virginia pine, 5; R; 99-423

MAGNOLIOPHYTA—MAGNOLIOPSIDA

Acanthaceae

Justicia americana (L.) Vahl, American water-willow, 7, 9; A;

99-309

Ruellia caroliniensis (J.F. Gmel.) Steud., Carolina wild petunia,

2, 35 0; 98-71

Aceraceae

tAcer barbatum Michx., Southern sugar maple, 2; |; 02-226

Acer negundo L., Box-elder, 7, 8; O; 98-11

Acer rubrum L. var. rubrum, Red maple, 1, 2, 3, 4; A; 99-29

Acer saccharinum L., Silver maple, 8; O; 99-02

Amaranthaceae

*Alternanthera philoxeroides (Mart.) Griseb., Alligator-weed,

9, 10; A; 01-241

*Amaranthus rudis J.D. Sauer, Water-hemp, 10; O; 99-451

*Amaranthus retroflexus L., Redroot amaranth, 10; S; 99-449

Amaranthus tuberculatus (Moq.) J.D. Sauer, Roughfruit ama-

ranth, 10; |; Fuller 3000

Anacardiaceae

Rhus copallina L., Winged sumac, 1, 3, 4; F; 98-324

Rhus glabra L., Smooth sumac, 3, 4; O; 98-111

Toxicodendron radicans (L.) Kuntze, Eastern Poison lvy, 1, 2,

3598; A; 02-209

Annonaceae

Asimina triloba (L.) Dunal, Pawpaw, 2; O; 98-415

Apiaceae

Angelica venenosa (Greenway) Fernald, Hairy angelica, 4;

R; 98-426

Chaerophyllum tainturieri Hook., Hairyfruit chervil, 3, 5; F;

Cicuta maculata L., Spotted water hemlock, 9, 10; |.; 98-440

**Daucus carota L., Queen Anne’s lace, 3, 5; F; 06-247

Eryngium prostratum Nutt., Creeping eryngo, 5; |; 98-366

Sanicula canadensis L., Canadian snakeroot, 2; O; 99-274

t*Torilis arvensis (Huds.) Link, Spreading hedge-parsley, 5;

Trepocarpus aethusae Nutt. ex DC., White nymph, 2, 8; O;

£L LD:AI s IC

Thompson, Flora of

n Mantir]

Apocynaceae

Amsonia tabernaemontana Walter, Eastern bluestar, 3; R;

Apocynum cannabinum L., Indian-hemp, 3, 5; |; 98-01

t**Vinca minor L., Common periwinkle, 5; O; 01-214

Aquifoliaceae

llex decidua Walter, Deciduous holly, 2; R; 02-236

Araliaceae

Aralia spinosa L., Hercules-club, 2; O; 06-225

t**Hedera helix L., English ivy, 5; |; 98-56

Aristolochiaceae

Aristolochia serpentaria L., Virginia snakeroot, 2; R; 99-338

Asclepiadiaceae

e due Duo Britton, Honeyvine, 2, 3; S; 98-567

Asclepia Sm., Clasping milkweed, 4; R; 99-232

Que perennis Walter, Aquatic milkweed, 8, 9; S; 99-386

Asclepias syriaca L, Common milkweed, 1, 3; S; 98-168

Asclepias tuberosa L., Butterfly milkweed, 3, 4; |; 01-178

Asclepias variegata L., Redring milkweed, 2; |, S; 98-44

tMatelea gonocarpos (Walter) Shinners, Angularfruit milkvine,

Astera

*Achi le UE L., Common yarrow, 3, 4; |; 00-166

Ageratina altissima (L.) R.M. King & H.E. Rob., White snakeroot,

Ambrosia artemisiifolia L., Annual ragweed, 3, 5; F; 01-150

Ambrosia trifida L., Giant ragweed, 3; O; 98-431

Antennaria plantaginifolia (L.) Richardson, Plantain pussytoes,

1; O; 99-

Bidens aristosa (Michx.) Britton, Bearded beggar-tick, 6; S;

-455

Bidens bipinnata L., Spanish needles, 5; S; Woods 818

Bidens frondosa L., Devil's beggar-tick, 9, 10; F; 01-601

Bidens vulgata Greene, Big devil's beggar-tick, 9; |; 01-608

Boltonia asteroides (L.) L'Her. var. recognita (Fernald & Griscom)

Cronquist, White doll's daisy, 3; |; 98-690

**Chrysanthemum leucanthemum L., Ox-eye daisy, 3, 5; |;

**Cichorium intybus L., Chicory, 5; S; 98-384

Cirsium discolor (Muhl. ex Willd.) Spreng. Field thistle, 3; |;

t*Cirsium vulgare (Savi) Ten., Bull thistle, 3; R; 98-583

Conoclinium coelestinum L., Blue mistflower, 9; |; 01-634

Conyza canadensis (L.) Cronquist, Horseweed, 5; O; 99-442

tCoreopsis auriculata L., Lobed tickseed, 2; R; 98-395

tCoreopsis lanceolata L., Lanceleaf tickseed, 3; |; 99-179

Coreopsis major Walter, Greater tickseed, 1, 4; O; 01-120

Coreopsis tinctoria Nutt. var. tinctoria, Golden tickseed, 3, 5;

S; 01-222

Coreopsis tripteris L., Tall tickseed, 1, 2; |; 98-552

t*Cosmos bipinnatus Cav., Garden cosmos, 3; p

*Eclipta prostrata (L JL, False daisy, 9, 10; S; 98-6

Elephantopus Raeusch., Carolina aes -foot,

2; |; 98-707

Erechtites hieraciifolia (L) Raf. ex DC., American burnweed,

5; |; 01-614

621

Erigeron annuus (L.) Pers., Annual fleabane, 3, 4, 5; F; 01-105

Erigeron philadelphicus L., Philadelphia fleabane, 5; |; 06-227

Erigeron strigosus Muhl ex Willd., Prairie fleabane, 2, 4; F;

-160

Fupatorium fistulosum Barratt, Trumpetweed, 3, 4; O; 98-550

Eupatorium perfoliatum L., Common boneset, 3, 9; O; 99-419

Eupatorium serotinum Michx., Lateflowering thoroughwort,

6, 9; O; 01-603

Eupatorium sessilifolium L., Upland boneset, 2; S; 98-587

Euthamia graminifolia (L.) Nutt. ex Cass., Flat-top goldenrod,

3, 6; |; 99-411

Gamochaeta purpurea (L.) Cabrera, Spoonleaf purp

ing, 2, 5; F; 02-204

Helenium flexuosum Raf., Purplehead sneezeweed, 6; S;

everlast-

-594

Helianthus angustifolius L., Swamp sunflower, 3; |; 01-625

Helianthus divaricatus L., Woodland sunflower, 2, 3; |; 98-405

Helianthus hirsutus Raf., Hairy sunflower, 1; O; 99-412

Helianthus microcephalus Torr. & A. Gray, Small woodland

sunflower, 2, 3; O; 98-548

Hieracium gronovii L., Beaked hawkweed, 1; O; 98-623

Krigia biflora (Walter) S.F. Blake, Twoflower dwarf-dandelion,

2, 3; O; 99-51

Krigia caespitosa (Raf.) K.L. Chambers, Weedy dwarf-dande-

lion, 5; O; 06-229

Krigia dandelion (L.) Nutt., Potato dwarf-dandelion, 1; |;

Lactuca canadensis L., Canada lettuce, 3; O; 98-347

Lactuca floridana (L.) Gaertn., Woodland lettuce, 2, 3; |;

-545

*Lactuca serriola L., Prickly lettuce, 3; S; 99-312

Liatris squarrosa (L.) Michx., Plains blazing-star, 1, 3; |; 99-520

Liatris squarrulosa Michx., Southern blazing-star, 1, 3; O;

+Matricaria discoidea DC., Disc mayweed, 3; R; 99-244

Mikania scandens (L.) Willd., Climbing hempvine, 8; R;

Packera glabella (Poir.) C. Jeffrey, Yellowtop, 10; F; 99-03

Parthenium integrifolium L., Wild quinine, 1, 4; O; 01-121

Pluchea camphorata (L.) DC., Marsh fleabane, 8; R; 99-460

Pseudognaphalium obtusifolium (L.) Hilliard & B.L. Burtt.,

Fragrant cudweed, 3; |; 98-639

Pyrrhopappus carolinianus (Walter) DC., Carolina desert-

Chicory, 3, 5; l; 99-283

Rudbeckia hirta L., Black-eyed susan, 3, 4; O; 98-67

Sericocarpus linifolius (L.) B.S.P., Narrowleaf white-topped

aster, 1,3, 4; O; 01-245

Solidago caesia L., Axillary goldenrod, 2; O; 98-706

Solidago canadensis L., Canada goldenrod, 3, 4; F; 01-610

Solidago juncea Aiton, Early goldenrod, 3; F; 99-328

Solidago nemoralis Aiton, Gray goldenrod, 3; O; 98-643

Solidago odora Aiton, Anise-scented goldenrod, 4; |; 01-612

Solidago speciosa Nutt. var. erecta (Pursh) McMillan, Showy

goldenrod, 1; O; 01-622

*Sonchus asper (L.) Hill, Spiny sow-thistle, 5; R; 98-456

Symphyotrichum dumosum (L.) G.L. Nesom, Longstalk aster,

3,4; F; 01-619

—

622

Symphyotrichum lateriflorum (L.) A. Love & D. Love, Calico

aster, 22 1-01-6017

Symphyotrichum ontarione (Wiegand) G.L. Nesom,

Bottomland aster, 10; O; 01-611

Symphyotrichum patens (Aiton) G.L. Nesom var. patens,

Clasping aster, 1; O; 01-633

Symphyotrichum pilosum (Willd.) G.L. Nesom, Hairy white

old-field aster, 3; F; 01-615

*Taraxacum officinale G.H. Weber ex Wiggers, Common

dandelion, 5; F; 99-12

Verbesina helianthoides Michx., Ozark wingstem sunflower,

3: O; 00-163

Vernonia missurica Raf., Missouri ironweed, 3; O; 01-604

Xanthium strumarium L., Rough cocklebur, 10; F; 98-657

Balsaminaceae

Impatiens capensis Meerb., Orange jewelweed, 7, 8, 9, 10;

; 32

Berberidaceae

Podophyllum peltatum L., May-apple, 2; O; 99-20

Betulaceae

Alnus serrulata (Aiton) Willd., Hazel alder, 8; O; 99-445

Betula nigra L., River birch, 8; O; 99-242

Corylus americana Walter, American hazelnut, 2; S; 98-224

tOstrya virginiana (Mill.) K. Koch, Hop-hornbeam, 3; |;

01-224

Bignoniaceae

Bignonia capreolata L., Cross-vine, 8; F; 98-163

Campsis radicans (L) Seem. ex Bureau, Trumpet-creeper, 8,

10; O; 01-204

Boraginaceae

Cynoglossum virginianum L.Wild comfrey, 2; R; 02-234

Myosotis macrosperma Engelm., Largeseed forget-me-not,

3; |; 99-183

Brassicaceae

*Arabidopsis thaliana (L.) Heynh, Mouse-ear cress, 5; O;

*Capsella bursa-pastoris (L.) Medik., Shepherd's purse, 5; R;

*Cardamine hirsuta L., Hairy bittercress, 5, 10; A; 99-16

Cardamine parviflora L., Dryland bittercress, 2; |; 99-98

Cardamine pensylvanica Muhl. ex Willd., Pennsylvania bit-

tercress, 10; l; 99-05

Dentaria laciniata Muhl. ex Willd., Cutleaf toothwort, 2; I;

99-18

Draba brachycarpa Nutt. ex Torr. & A. Gray, Shortfruit whit-

low-grass, 5; |; 99-33

*Draba verna L., Whitlow-grass, 5; F; 99-13

Lepidium virginicum L., Wild peppergrass, 5; |; 00-167

Rorippa sessiliflora (Nutt) Hitchc., Marsh yellowcress, 9; O;

*Sisymbrium officinale (L.) Scop., Hedge-mustard, 5; S;

99-175

Callitrichaceae

tCallitriche terrestris Raf., Terrestrial water starwort, 5; S;

Poindexter 06-133

fal, Dat o ID L

Journal of

titute of Texas 1(1)

Campanulaceae

Campanulastrum americanum (L.) Small, American bellflower,

2; R; 98-325

Lobelia inflata L., Indian tobacco, 5, 10; |; 99-336

Lobelia puberula Michx., Downy lobelia, 3, 4; O; 01-621

Lobelia spicata Lam., Spiked lobelia, 3; R; Hunter & Austin

1807

Triodanis perfoliata (L.) Nieuwl. var. biflora (Ruiz & Pav.) Bradley,

Venus’ looking glass, 1, 5; |; 98-69

Triodanis perfoliata (L.) Nieuwl. var. perfoliata, Venus’ looking

glass, 5; F; 00-165

Caprifoliaceae

**Lonicera japonica Thunb., Japanese honeysuckle, 2, 4, 5,

8; A; 98-03

Sambucus canadensis L, Common elderberry, 2; 8; |; 01-164

Symphoricarpos orbiculatus Moench, Coralberry, 2, 3; O;

98-375

Viburnum rufidulum Raf., Rusty blackhaw, 1, 2; |; 99-111

Caryophyllaceae

**Arenaria serpyllifolia L, Thymeleaf sandwort, 5; F; 01-101

*Cerastium brachypetalum Desportes ex Pers., Gray mouse-

ear chickweed, 5; R; 99-104

*Cerastium glomeratum Thuill, Clammy mouse-ear chick-

weed, 5; F; 99-56

Cerastium nutans Raf, Nodding mouse-ear chickweed, 5;

*Cerastium vulgatum L., Common mouse-ear chickweed,

5; O; 98-66

**Dianthus armeria L., Deptford pink, 5; |; 00-151

Silene antirrhina L., Sleepy catchfly, 5; S; 99-138

Silene stellata (L.) W.T. Aiton, Starry campion, 2; S; 98-414

Silene virginica L., Fire pink, 2; |; 99-22

**Stellaria media (L.) Vill., Common chickweed, 5; F; 99-53

Chenopodiaceae

**Chenopodium album L., Lamb's-quarters, 10; S; 01-168

Cistaceae

Lechea mucronata Raf., Hairy pinweed, 1; S; 98-511

+Lechea tenuifolia Michx., Narrowleaf pinweed, 1; F; 01-218

Clusiaceae

Hypericum denticulatum Walter, Coppery St. John's-wort, 1,

3, 4; 0; 99-332

Hypericum drummondii (Grev. & Hook.) Torr. & A. Gray, Nits-

and-lice, 1; R; 98 493

Hypericum hypericoides (L.) Crantz subsp. hypericoides, St.

Andrew's-cross, 2; O; 01-184

Hypericum mutilum L., Marsh St. John's-wort, 6, 9; O; 99-390

tHypericum prolificum L., Shrubby St. John's-wort, 2; S;

99-314

Hypericum punctatum Lam., Dotted St. John's-wort, 2, 3, 4;

Hypericum stragulum W.P. Adams & N. Robson, St. Andrew's-

cross, 1, 3, 4; O; 98-409

Convolvulaceae

*Calystegia sepium (L.) R. Br. var. sepium, Hedge bindweed,

8; l; 99-379

£L LD:AI s IC

Thompson, Flora of

in Mantel

**Inomoea hederacea Jacq., lvyleaf morning-glory, 10; R;

99-439

Ipomoea lacunosa L., White morning-glory, 10; F; 98-665

Cornaceae

Cornus amomum Mill., Silky dogwood, 8; |; 01-117

Cornus florida L., Flowering dogwood, 2, 3, 5; O; 99-26

Cucurbitaceae

Melothria pendula L., Creeping cucumber, 5; R; 98-604

Sicyos angulatus L., Bur cucumber, 2, 8; S; 01-643

Cuscutaceae

Cuscuta pentagona Engelm., Field dodder, 5; O; 99-302

Ebenaceae

Diospyros virginiana L., Persimmon, 2, 3, 8; F; 98-130

Elaeagnaceae

t**Flaeagnus umbellata Thunb., Autumn-olive, 5; S; 99-32

Ericaceae

Vaccinium arboreum Marshall, Sparkleberry, 1; F; 00-156

Vaccinium stamineum L., Deerberry, 1, 2; O; 01-216

Euphorbiaceae

Acalypha rhomboidea Raf., Rhomboic copperleaf, 5, 10; |;

Acalypha virginica L. Virginia copperleaf, 5, 10; F; 98-561

Croton capitatus Michx., Woolly croton, 5; R; 98-621

Croton glandulosus L. var. septentrionalis (L) Muell.-Arg.,

Toothleaf croton, 3; |; 98-711

Croton monanthogynus Michx., Prairie-tea, 5; O; 98-370

Euphorbia corollata L., Flowering spurge, 1, 3, 4; O; 98-356

Euphorbia maculata L., Spotted sandmat, 5; 10; F; 99-431

Euphorbia nutans Lag., Eyebane spurge, 5, 10; O; 98-684

Phyllanthus caroliniensis Walter, Carolina leaf-flower, 10; R;

Fabaceae

**Albizia julibrissin Durazz., Mimosa, 3, 5; O; 01-165

Amorpha fruticosa L., False indigo, 8; S; 99-224

Cercis canadensis L., Eastern redbud, 2, 3, 5; O; 99-31

Chamaecrista fasciculata (Michx.) Greene, Partridge-pea, 3,

4; O; 98-596

Chamaecrista nicitans (L.

Moench., Sensitive-pea, 3, 4; |;

Clitoria mariana L., Butterfly pea, 2, 3; |; 00-321

**Coronilla varia L., Crown-vetch, 5; O; 01-128

Crotalaria sagittalis L., Weedy rattlebox, 4; R; 99-325

tDesmodium glabellum (Michx.) DC., Smooth tick-trefoil, 3,

pO, 98097

+Desmodium glutinosum (Muhl. ex Willd.) A. Wood, Clustered

tick-trefoil, 2; S; 98-398

Desmodium marilandicum (L) DC., Maryland tick-trefoil, 2,

3; |; 98-588

Desmodium nudiflorum (L.) DC., Naked tick-trefoil, 2; O;

-320

Desmodium paniculatum (L.) DC., Panicled tick-trefoil, 3; O;

Desmodium rotundifolium DC., Roundleaf tick-trefoil, 1; S;

8-5,

623

tDioclea multiflora (Torr. & A. Gray) C. Mohr, Cluster-pea, 1;

Galactia volubilis (L.) Britton, Hairy milk-pea, 3; O; 99-326

Gleditsia triacanthos L., Honey locust, 2; R; 98-326

**Lespedeza cuneata (Dum.-Cours.) G. Don, Sericea lespedeza,

3:5 01-633

Lespedeza hirta (L) Hornem., Hairy lespedeza, 1; O; 01-618

Lespedeza intermedia (S. Wats.) Britton, Wand lespedeza, 1,

3; O; 98-703

Lespedeza procumbens Michx., Downy trailing lespedeza,

3; E: 96-699

Lespedeza repens (L) Barton, Smooth trailing lespedeza, 3;

**| espedeza stipulacea Maxim., Korean clover, 3, 5; A; 98-379

+**/ espedeza striata (Thunb.) Hook. & Arn., Japanese clover,

3,5; A; 01-636

Lespede a virgit ica (L) Britton, Virginia lespedeza, 3: |; 98-626

t*Lotus corniculatus L., Birdsfoot-trefoil, 3; R; 01-203

**Medicago lupulina L., Black medic, 5, 10; F; 06-246

** Melilotus alba Medik., White sweet-clover, 3; 5; |; 01-221

**Melilotus officinalis (L) Lam., Yellow sweet-clover, 5; R;

Orbexilum pedunculatum (Mill.) Rydb., Sampson's snakeroot,

1; O; 98-18

Robinia pseudoacacia L., Black locust, 2; O; 99-91

Strophostyles umbellata (Muhl. ex Willd.) Britton, Perennial

woolly-bean, 3; S; 98-595

Stylosanthes biflora (L.) B.S.P., Pencil-flower, 3, 4; O; 99-234

Tephrosia virginiana (L.) Pers., Virginia goat's-rue, 1; O; 01-124

*Trifolium campestre Schreb., Pinnate hop-clover, 5; S; 98-46

+*Trifolium dubium Sibth., Little hop-clover, 5; A; 01-102

t*rrifolium pratense L., Red clover, 3, 5; O; 02-237

*Trifolium repens L., White clover, 5; A; 01-139

*Vicia sativa L., Common vetch, 5; S; 99-215

*Vicia villosa Roth var. varia (Host) Corb., Winter vetch, 3; R;

Wisteria frutescens (L.) Poir., American wisteria, 8; O; 98-412

Fagaceae

Fagus grandifolia Ehrh., American beech, 2; S; 99-290

Quercus alba L., White oak, 1, 2; A; 98-540

Quercus coccinea Muenchh., Scarlet oak, 1, 2; R; 98-369

Quercus falcata Michx., Southern red oak, 1, 2; F; 98-564

Quercus imbricaria Michx., Shingle oak, 2; |; 98-449

Quercus lyrata Walter, Overcup oak, 8; R; 99-424

Quercus marilandica Muenchh., Blackjack oak, 1; F; 99-200

tQuercus muhlenbergii Engelm., Chinkapin oak, 2; R; 98-41

Quercus rubra L., Northern red oak, 2; F; 98-345

tQuercus shumardii Buckley, Shumard oak, 5; R; 98-603

Quercus stellata Wangenh., Post oak, 1, 2; A; 98-123

Quercus velutina Lam., Black oak, 1, 2, 3; A; 02231

Gentianaceae

Sabatia angularis (L.) Pursh, Rose marsh-pink, 3, 6; |; 99-324

Geraniaceae

Geranium carolinianum L., Carolina crane's-bill, 3, 5; O;

624

Haloragaceae

t*Myriophyllum spicatum L., European water-milfoil, 9; I;

Fuller 3004

Hamamelidac

Liquidambar TUE L., Sweetgum, 2, 4, 8; O; 98-192

Hippocastanaceae

tAesculus pavia Aiton, Red buckeye, 2; R; 99-334

Hydrangeaceae

Hydrangea cinerea Small, Wild hydrangea, 2; |; 01-207

Juglandaceae

Bue E (Wangenh.) K. Koch, Bitternut hickory,

2: d

Carya ie de Mill.) Sweet, Pignut hickory, 1, 2; A; 02-223

Carya ovata (P. Mill.) K. Koch, Shagbark hickory, 2, 3; A; 98-

2:51

tCarya pallida (Ashe) Engl. & Graebn., Pale hickory, 2; S;

98-197

Carya tomentosa (Poir.) Nutt., Mockernut hickory, 2, 3; A;

Lamiaceae

**Glechoma hederacea L., Ground-ivy, 5; O; 01-255

Cunila origanoides (L.) Britton, Maryland dittany, 1; O; 98-717

Hedeoma pulegioides (L.) Pers., American false pennyroyal,

2; R; Woods 819

**Lamium amplexicaule L., Henbit, 5; |; 99-11

*Lamium purpureum L., Purple dead-nettle, 5; O; 99-14

Lycopus virginicus L., Virginia water-horehound, 9; S; 98-682

t**Mentha x piperita L., Peppermint, 7; R; 00-322

Monarda fistulosa L. subsp. mollis (L.) Benth., Wild bergamot,

3; 4; |: 01-170

*Perilla frutescens (L.) Britton, Beefsteak plant, 10; R; 98-710

Prunella vulgaris L. var. lanceolata (W. Barton) Fernald, Self-

heal, 3, 5; O; 98-64

Pycnanthemum pycnanthemoides (Leavenw.) Fernald,

Southern mountain mint, 3, 4; |; 01-623

Pycnanthemum tenuifolium Schrad., Slender mountain mint,

3, 4; O; 99-285

tPycnanthemum virginianum (L.) Durand & A.B. Jackson,

Virginia mountain mint, 3; S; 98-519

Salvia lyrata L., Wild sage, 2, 5; O; 99-151

Scutellaria elliptica Muhl., Hairy skullcap, 2; O; 00-158

Scutellaria incana Biehler, Downy skullcap, 2; |; 98-294

Scutellaria integrifolia L., Largeflower skullcap, 2; |; 99-167

Scutellaria ovata Hill, Heartleaf skullcap, 2; R; 98-93

Scutellaria parvula Michx., Little skullcap, 1; |; 99-87

Stachys tenuifolia Willd., Smooth hedge-nettle, 6; R; 00-320

Teucrium canadense L., Canada germander, 10; R; 98-208

Lauraceae

Sassafras albidum (Nutt.) Nees, Sassafras, 1, 2, 3, 4; O; 98-223

Linaceae

Linum medium (Planch.) Britton var. texanum (Planch.)

Fernald, Common yellow flax, 3, 4; O; 06-239

Loganiaceae

Spigelia marilandica L., Indian-pink, 2; |; 01-112

fal, Dat A ID hi Pr

Journal of

of Texas 1(1)

Lythraceae

Ammannia coccinea Rottb., Valley redstem, 9; S; 98-659

Rotala ramosior (L.) Koehne, Tooth-cup, 9, 10; A; 99-383

Magnoliaceae

Liriodendron tulipifera L., Tuliptree, 2, 3; S; 98-293

tMagnolia grandiflora L., Southern magnolia, 5; R; 06-224

Malvaceae

Hibiscus laevis All., Smooth rose-mallow, 9, 10; O; 99-387

*Sida spinosa L., Prickly sida, 10; S; 98-539

Melastomataceae

Rhexia virginica L., Wingstem meadow-beauty, 8; R; Woods

Menispermaceae

Cocculus carolinus (L.) DC., Carolina coralbeads, 2; |; 98-496

Molluginaceae

Mi verticillata L., Carpetweed, 5, 10; O; 01-162

onotropacea

chs om. L., Pine-sap, 2; R; 98-217

Moraceae

Maclura pomifera (Raf.) C.K. Schneid., Osage-orange, 3; R;

Morus rubra L., Red mulberry, 2, 3; O; 98-195

Nyssaceae

Nyssa aquatica L., Swamp tupelo, 8; R; 02-151

Nyssa sylvatica Marshall, Blackgum, 2, 3, 4; A; 01-119

Oleaceae

Fraxinus americana L., White ash, 2, 3; |; 98-382

Fraxinus pennsylvanica Marshall, Green ash, 8; R; 98-357

t**Ligustrum sinense Lour., Chinese privet, 5; R; 99-393

Onagraceae

Ludwigia alternifolia L., Square-pod water-primrose, 6, 9;

O; 98-516

Ludwigia decurrens Walter, Wingstem water-primrose, 9; R;

Oenothera biennis L., Common evening-primrose, 3, 5; O;

Oenothera fruticosa L. subsp. fruticosa, Common sundrops,

4; |; 99-188

Oenothera speciosa Nutt., White evening-primrose, 3; R;

98-02

Oxalidaceae

Oxalis stricta L, Common yellow wood-sorrel, 5; S; 01-108

Oxalis violacea L., Violet wood-sorrel, 1, 2; |; 99-80

Passifloraceae

Passiflora incarnata L., Maypop passion-flower, 2, 3; R; 00-325

Passiflora lutea L. var. glabriflora Fernald, Yellow passion-

flower, 2; S; 98-373

Phrymac

Phyrma URN L., Lopseed, 2; |; 98-279

Phytolaccaceae

Phytolacca americana L., American pokeweed, 5; |; 01-113

£L LD:AI s I C4

Thompson, Flora of

n Kantıırl

Plantaginaceae

Plantago aristata Michx., Bracted plantain, 5; l; 01-223

*Plantago lanceolata L., English plantain, 5; F; 98-74

Plantago rugelii Decne., Rugel's plantain, 5; F; 98-180

Plantago virginica L., Hoary plantain, 3, 5; O; 99-126

Platanaceae

Platanus occidentalis L., American sycamore, 7, 8; O; 98-201

Polemoniaceae

Phlox divaricata L., Forest phlox, 2; O; 99-23

tPhlox paniculata L., Summer phlox, 5; R; 99-321

Polygalaceae

Polygala ambigua Nutt., Loose milkwort, 4; |; 98-404

Polygala sanguinea L., Blood milkwort, 4; S; 98-124

Polygonaceae

Brunnichia ovata (Walter) Shinners, Buckwheat vine, 8; O;

Polygonum amphibium L. var. emersum Michx., Water smart-

weed, 9; |; 98-652

*Polygonum aviculare L., Knotweed, 5, 10; O; 99-462

**Polygonum caespitosum Blume var. longisetum (Bruijn)

Steward, Asiatic smartweed, 5, 10; F; 01-638

Polygonum hydropiperoides Michx., False water-pepper, 9;

O; 99-438

Polygonum pensylvanicum L., Pennsylvania smartweed, 1

|; 98-664

e persicaria L., Spotted lady's thumb, 5, 10; O;

Bp sagittatum L., Arrowleaf tearthumb, 7, 9; O;

-447

Polygonum virginianum L., Jumpseed, 2; |; 98-568

**Rumex acetosella L., Sheep sorrel, 5; F; 98-77

*Rumex crispus L., Curly dock, 5; |; 98-83

*Rumex obtusifolius L., Bitter dock, 5; |; 98-446

Portulacaceae

Claytonia virginica L., Spring-beauty, 5; O; 99-38

Primulaceae

Lysimachia ciliata L., Fringed loosestrife, 2; S; 99-254

Lysimachia lanceolata Walter, Lanceleaf loosestrife, 2; R; Hunter

and Austin 1806

Ranunculaceae

Anemone virginiana L., Tall anemone, 2; |; 98-102

Anemonella thalictroides (L.) Spach., Rue-anemone, 2; O;

Clematis virginiana L., Virgin's-bower, 2; |; 98-447

Myosurus minimus L., Mouse-tail, 10; R; 99-49

Ranunculus abortivus L., Smooth smallflower crowfoot, 5;

O; 99-08

+Ranunculus micranthus (A.Gray) Nutt. ex Torr. & A. Gray, Hairy

smallflower crowfoot, 5; |; 99-59

+**Ranunculus parviflorus L., Stickseed buttercup, 5; R;

Ranunculus recurvatus Poir., Hooked buttercup, 2; |; 99-150

*Ranunculus sardous Crantz., Hairy buttercup, 5; O; 99-61

Rhamnaceae

Ceanothus americanus L., New Jersey tea, 1, 4; |; 98-149

625

Rosaceae

Agrimonia rostellata Wallr., Beaked agrimonia, 2; |; 98-549

Amelanchier arborea (F. Michx.) Fernald, Downy serviceberry,

1, 2; F; 99-25

+*Aphanes microcarpa (Boiss. & Reut.) Rothm., Slender parsley

piert, 5; S; Poindexter 06-152

Crataegus mollis (Torr. & A. Gray) Schelle, Downy hawthorn,

(PROS

Fragaria virginiana Duchesne, Wild strawberry, 3; l; 99-140

Geum canadense Jacq., White avens, 2; O; 98-209

Porteranthus stipulatus (Muhl. ex Willd.) Britton, Indian-physic,

1; F; 01-100

Potentilla simplex Michx., Old-field cinquefoil, 3, 4; F; 99-79

Prunus americana Marshall, American plum, 1; R; 02-218

Prunus angustifolia Marshall, Chickasaw plum, 1; |; 00-164

Prunus serotina Ehrh., Wild black cherry, 2, 3; F; 99-99

Rosa carolina L., Pasture rose, 1, 3; O; 98-560

**Rosa multiflora Thunb., Multiflora rose, 3, 4; S; 01-176

Rosa setigera Michx., Prairie rose, 3; |; 01-1

Rubus allegheniensis Porter, Common blackberry, 3; |; 98-244

Rubus argutus Link, Southern blackberry, 3, 4; F; 98-169

Rubus flagellaris Willd., Northern dewberry, 3, 4; F; 99-229

Rubiaceae

Cephalanthus occidentalis L., Buttonbush, 8; O; 01-206

Diodia teres Walter, Rough buttonweed, 3; F; 01-246

Diodia virginiana L., Virginia buttonweed, 7, 9, 10; F; 99-346

Galium aparine L., Cleavers, 3, 5; O; 99-119

Galium circaezans Michx., Forest bedstraw, 1; O; 02-217

**Galium pedemontanum (Bellardi) All., Pedmont bedstraw,

5; S; 98-78

Galium pilosum Aiton, Hairy bedstraw, 1, 3; O; 99-316

Galium tinctorium L., Swamp bedstraw, 9; |; 98-133

Galium triflorum Michx., Fragrant bedstraw, 2; O; 98-425

Houstonia caerulea L., Spring bluets, 5; F; 99-50

Houstonia canadensis Willd. ex Roem. & Schult., Canada

bluets, 1; O; 99-135

Houstonia pusilla Schoepf, Small bluets, 5; O; 99-57

+*Sherardia arvensis L., Field-madder, 5; R; Poindexter 06-142

Salicaceae

Populus deltoides W. Bartram ex Marshall, Eastern cotton-

wood, 8; S; 98-327

Salix humilis Marshall, Upland willow, 3; |; 99-415

Salix nigra Marshall, Black willow, 6, 8; O; 99-239

Santalaceae

Comandra umbellata (L.) Nutt. subsp. umbellata, Bastard

toadflax, 1; l; 99-139

Sapindaceae

*Cardiospermum halicacabum L., Balloon-vine, 10; R; Fuller

3006

Saururaceae

Saururus cernuus L., Lizard's-tail, 9; |.; 99-344

Saxifragaceae

Heuchera americana L., American alumroot, 2; S; 98-13

626

Scrophulariaceae

Agalinis tenuifolia (Vahl) Raf, Common false foxglove, 3; |;

01-616

Aureolaria flava (L.) Farw., Smooth foxglove, 1; S; 98-615

Aureolaria pedicularia (L.) Raf. var. pectinata (Nutt) Gleason,

Annual foxglove, 1;0; 01-624

tLeucospora multifida (Michx.) Nutt., Cleftleaf Conobea, 5;

99-361

Lindernia dubia (L.) Pennell var. anagallidea (Michx.) Cooperr.,

False pimpernel, 9, 10; O; 99-457

Lindernia dubia (L.) Pennell var. dubia, False pimpernel, 9,

10; |; 99-389

Mecardonia acuminata (Walter) Small, Axilflower, 10; R;

Mimulus alatus Aiton, Sharpwing monkey-flower, 9; |; 98-515

Mimulus ringens L., Alleghany monkey-flower, 7; R; 98-494

Penstemon digitalis Nutt. ex Sims, Foxglove beardtongue,

99-134

Penstemon pallidus Small, Eastern white beardtongue, 3;

*Verbascum blattaria L., Moth mullein, 3; R; 00-168

*Verbascum thapsus L., Common mullein, 3; R; 98-146

*Veronica arvensis L., Corn speedwell, 5; F; 98-55

Veronica peregrina L. subsp. peregrina, Purslane speedwell,

5; O; 99-48

Veronicastrum virginicum (L.) Farw., Culver's root, 1; R; 99-418

Simaroubaceae

**Ailanthus altissima (Miller) Swingle, Tree-of-heaven, 2; R;

Solanaceae

Physalis pubescens L., Downy ground-cherry, 10; |; 99-426

Solanum carolinense L., Bull-nettle, 3, 5; S; 98-339

Solanum ptycanthum Dunal ex DC., Black nightshade, 5; R;

Staphyleaceae

Staphylea trifolia L., Bladdernut, 2; R; 98-266

Styracaceae

Styrax americanus Lam., American snowbell, 8; |; 99-343

Ulmaceae

Celtis occidentalis L., Common hackberry, 2; |; 022211

Ulmus alata Michx., Winged elm, 1, 2, 3, 4; F; 98-151

icana L., American elm, 2; O; 98-220

Ulmus rubra Muhl., Red elm, 1, 3; F; 98-164

limi amori

Urticaceae

Boehmeria cylindrica (L.) Sw., False nettle, 2, 7, 8; F; 98-292

Pilea pumila (L) A. Gray, Clearweed, 2; O; 98-441

Valerianaceae

Valerianella radiata (L.) Dufr., Beaded corn-salad, 3; O; 99-74

Verbenaceae

Phyla lanceolata (Michx.) Greene, Frogfruit, 7, 9; O; 01-240

Verbena simplex Lehm., Narrowleaf vervain, 3; S; 98-50

Verbena urticifolia L., White vervain, 6, 9; |; 98-436

Violaceae

Viola palmata L., Threelobe wood violet, 2; O; 99-65

fal, Dat o ID L

Journal of

titute of Texas 1(1)

Viola pedata L., Bird'sfoot violet, 1; |; 99-78

Viola rafinesquii Greene, Field pansy, 3, 5; O; 99-36

Viola sororia Willd., Common blue violet, 5; F; 99-09

Viscaceae

Phoradendron leucarpum (Raf.) Reveal & M.C. Johnst.,

American mistletoe, 1; R; 06-757

Vitaceae

Ampelopsis cordata Michx., Raccoon-grape, 2; S; 99-364

Parthenocissus quinquefolia (L.) Planch., Virginia-creeper, 1,

2; A; 01-208

Vitis aestivalis Michx., Summer grape, 1, 2; F; 98-118

tVitis palmata Vahl, Red grape, 8; R; 01-217

Vitis rotundifolia Michx., Muscadine grape, 1, 2, 8; A; 01-114

MAGNOLIOPHYTA—LILIOPSIDA

Agavaceae

Mantfreda virginica (L.) Salisb. ex Rose, False aloe, 1; R; 98-344

tYucca filamentosa L., Spanish-bayonet, 5; R; 02-224

Alliaceae

Allium canadense L., Wild onion, 3; O; 99-240

*Allium vineale L., Field garlic, 3; |; 02-207

Amaryllidaceae

t*Narcissus pseudonarcissus L., Daffodil, 2; |; 99-17

Araceae

Arisaema dracontium (L.) Schott, Green-dragon, 2; R; 01-226

Commelinaceae

**Commelina communis L., Asiatic dayflower, 5; O; 06-231

*Commelina diffusa Burm. f, Creeping dayflower, 10; |; 98-

671

Commelina virginica L., Virginia dayflower, 8; O; 98-669

Convallariaceae

Maianthemum racemosum (L.) Link var. racemosum, False

Solomon 's-seal, 2; |; 98-09

Polygonatum biflorum (Walter) Elliott, Smooth Solomon's-

seal, 2; S; 99-272

Cyperaceae

+Carex albicans Willd. var. albicans, Whitetinge sedge, 2; O;

99-159

Carex blanda Dewey, Eastern wood sedge, 2; O; 99-162

Carex caroliniana Schwein, Carolina sedge, 2; R; 99-123

Carex cephalophora Muhl. ex Willd., Oval-leaf sedge, 5; |;

tCarex crinita Lam. var. crinita, Fringed sedge, 7; O; 99-149

Carex debilis Michx. var. debilis, White-edge sedge, 2; O;

Carex digitalis Willd. var. macropoda Fernald, Slender wood

sedge, 2; O; 99-163

+Carex frankii Kunth, Frank's sedge, 9; S; 98-271

Carex glaucodea Tuck., Blue sedge, 9; S; 98-19

Carex grayi J. Carey, Gray's sedge, 2; |; 06-222

Carex hirsutella Mack., Hairy green sedge, 3, 4; F; 01-134

Carex laxiflora Lam. var. laxiflora, Broad looseflower sedge,

2009-139

tCarex lupulina Muhl. ex Willd., Hop sedge, 7, 9; O; 01-233

Carex lurida Wahlenb., Yellow green sedge, 6; S; 99-213

£L LD:AI s I C4

Thompson, Flora of

in Kantırl

Carex muehlenbergii Schkuhr ex Willd., Muhlenberg's sedge,

1; F;01-130

Carex nigromarginata Schwein. Blackedge sedge, 2; R;

Carex retroflexa Muhl. ex Willd., Reflexed sedge, 2; O; 99-161

Carex tribuloides Wahlenb., Blunt broom sedge, 9; S; 01-215

Carex typhina Michx., Cattail sedge, 2, 9; S; 01-232

Carex vulpinoidea Michx., Fox sedge, 7; S; 98-75

Cyperus echinatus (L.) A. Wood, Globe flatsedge, 3, 6; S;

Cyperus esculentus L., Yellow nutsedge, 9; S; 99-444

Cyperus pseudovegetus Steud., Marsh nutsedge, 6, 9; S; 01-

238

Cyperus squarrosus L., Bearded nutsedge, 8, 10; A; 99-436

Cyperus strigosus L., Strawcolor nutsedge, 6, 9; |; 01-234

+Fleocharis acicularis (L.) Roem. & Schult., Needle spikerush,

10; A; 99-455

Eleocharis ovata (Roth) Roem. & Schult., Blunt ovate spikerush,

6; |; 98-444

Fimbristylis autumnalis (L.) Roem. & Schult., Slender fimbry,

9; A; 99-380

Isolepis carinata Hook. & Arn. ex Torr., Keeled bulrush, 6; S;

Poindexter 06-161

tKyllinga gracillima Miq., Pasture spikesedge, 5; |; Poindexter

06-162

Rhynchospora corniculata (Lam.) A. Gray, Shortbristle horned

beakrush, 9; |; 99-376

tSchoenoplectus tabernaemontani (K.C. Gmel.) Palla, Softstem

bulrush, 7; S; 98-522

tScirpus atrovirens Willd., Green bulrush, 7; |; 99-214

Scirpus cyperinus (L.) Kunth, Woolgrass, 6, 9; O; 98-554

Scleria pauciflora Mulh. ex Willd., Fewflower nutrush, 4; O;

99-235

Scleria triglomerata Michx., Whip nutrush, 4; O; 98-137

Dioscoreaceae

t**Dioscorea oppositifolia L, Chinese yam, 7; R; 99-250

Dioscorea villosa L., Wild yam, 2; O; 01-220

Hemerocallidaceae

t**Hemerocallis fulva (L.) L., Orange day-lily, 5, 7; O; 99-258

Hydrocharitaceae

Najas guadalupensis (Sprengel) Magnus, Southern water-

nymph, 9; A; 99-457

Hypoxidaceae

Hypoxis hirsuta (L.) Coville, Yellow star-grass, 1; |; 99-77

Iridaceae

Iris cristata Soland. ex Aiton, Dwarf crested iris, 2; O; 99-69

t*iris pseudoacorus L., Yellow flag, 7; S; 99-147

tiris virginica L., Southern blue flag, 7, 8; O; 99-148

Sisyrinchium angustifolium Mill., Narrowleaf blue-eyed-grass,

Juncaceae

Juncus acuminatus Michx., Taperpoint rush, 6; |; 98-

Juncus biflorus Elliott, Bog rush, 3, 6; O; 01-225

Juncus brachycarpus Engelm., Whiteroot rush, 6; |; 98-256

Juncus diffusissiimus Buckley, Slimpod rush, 6; S; 99-287

—

627

Juncus effusus L. var. solutus. Fern. & Wieg., Soft rush, 6, 9;

O; 02-202

Juncus marginatus Rostk., Grassleaf rush, 6; |; 98-153

+Juncus nodatus Coville, Stout rush, 6; S; Poindexter 06-140

Juncus tenuis Willd., Slender path rush, 3, 5; A; 99-245

Luzula bulbosa (A. Wood) Rydb., Bulbous woodrush, 1, 2;

F; 99-86

Marantaceae

tl halia dealbata Fraser ex Roscoe, Powdery alligator-flag,

7; S; 98-716

Melanthiaceae

Chamaelirium luteum (L.) A. Gray, Devil's-bit, 2; R; 99-208

Orchidaceae

Spiranthes cernua (L.) Rich., Nodding labies’-tresses, 3; R;

98-641

Spiranthes lacera (Raf.) Raf. var. gracilis (Bigelow) Luer, Slender

adies'-tresses, 5; R; 98-611

Spiranthes vernalis Engelm. & A. Gray, Spring ladies'-tresses,

6; R; 98-212

Tipularia discolor (Pursh) Nutt., Crane-fly orchid, 2; S; 98-526

Poaceae

*Agrostis gigantea Roth., Redtop, 6, 7; O; 00-326

*Agrostis stolonifera L., Creeping bent grass, 3; |; 99-133

Alopecurus carolinianus Walter, Carolina foxtail, 2; R; 99-47

Andropogon ternarius Michx., Splitbeard bluestem, 3, 4; O;

01-626

Andropogon virginicus L, Broom-sedge, 3, 4; F; 01-637

Aristida dichotoma Michx., Churchmouse threee-awn, 3; F;

Aristida longespica Poir., Slimspike three-awn, 3; O; 01-640

Brachyelytrum erectum (Schreb) P. Beauv., Bearded shorthusk,

2; O; 98-181

*Bromus commutatus Schrad., Hairy chess, 3, 5; O; 00-159

Bromus pubescens Muhl., Woodland brome, 2; O; 99-219

Chasmanthium latifolium (Michx.) H.O. Yates, Wood oats,

2; F; 99-378

*Cynodon dactylon (L.) Pers., Bermuda grass, 5; A; 98-352

*Dactylis glomerata L., Orchard grass, 3, 5; O; 02-200

Danthonia spicata (L.) P. Beauv., Poverty oat grass, 1; A;

Dichanthelium acuminatum (Sw.) Gould & C.A. Clark subsp.

fasciculatum (Torr.) Freckmann & Delong, Hairy panic

grass, 1,3; A; 01-138

Dichanthelium acuminatum (Sw.) Gould & C.A. Clark subsp

lindheimerii (Nash) Freckmann & Delong, Hairy panic

grass, |, 3107:2017

Dichanthelium boscii (Poir.) Gould & C.A. Clark, Bosc's pani-

cum, 1, 2; F; 02-215

Dichanthelium clandestinum (L.) Gould, Deer-tongue pani-

cum, 2, 6; O; 98-430

Dichanthelium commutatum (Schult) Gould, Variable panic

grass, 1, 3; F; 99-145

Dichanthelium depauperatum (Muhl.) Gould, Starved panic

grass, 1; O; 01-133

Dichanthelium dichotomum (L.) Gould, Forking panic grass,

2; F; 98-17

628

Dichanthelium laxiflorum (Lam.) Gould, Looseflower panic

grass, T, 5) A; 995-23

Dichanthelium polyanthes (Schult) Mohlenb., Manyflower

panic grass, 2; O; 01-115

Dichanthelium villossimum (Nash) Freckmann subsp. villosis-

simum, Longhair panic grass, 3; S; 99-203

*Digitaria ischaemum (Schreb. Schreb. ex Muhl., Smooth crab

grass, 5, 10; |; 98-668

*Digitaria sanguinalis (L) Scop., Hairy crab grass, 5, 10; F;

**Echinochloa crus-galli (L.) P. Beauv., Common barnyard

grass, 6, 10; F; 99-446

**Fleusine indica (L.) Gaertn., Yardgrass, 5; O; 98-300

Elymus glabriflorus (Vasey) Bush, Smooth wildrye, 3; O;

L/D

Elymus virginicus L. var. virginicus, Virginia wildrye, 2; O;

01-205

+**Fragrostis cilianensis (All) Vignolo ex Janch. Tufted love

grass, 5; R; 98-605

Eragrostis hypnoides (Lam.) B.S.P., Teal love grass, 10; O;

-666

+Fragrostis pectinacea (Michx.) Nees, Tufted love grass, 5;

Eragrostis spectabilis (Pursh) Steud., Purple love grass,3, 4;

**Festuca arundinacea Schreb., Tall fescue, 3, 5, 6; A; 99-172

**Holcus lanatus L., Velvet grass, 3, 6; S; 02-229

Hordeum pusillum Nutt., Little barley, 5; O; 06-234

Leersia oryzoides (L.) Sw., Rice cut grass, 7, 9; F; 98-655

Leersia virginica Willd., Virginia cut grass, 2; O; 98-598

t**Lolium perenne L. var. aristatum Willd., Perennial rye grass,

3; |; 06-245

Melica mutica Walter, Twoflower melic grass, 2; S; 06-238

**Microstegium vimineum (Trin.) A. Camas, Nepalese eulalia,

2; A; 01-631

Muhlenbergia schreberi J.F. Gmel., Nimblewill, 5; F; 98-693

Muhlenbergia sobolifera (Muhl) Trin., Rock muhly, 2; |; 98-

Panicum anceps Michx., Beaked panic grass, 6, 9; F; 99-382

Panicum dic if] ichx. subsp. dichotomifl Fa

panicum, 10; O; 98-679

Panicum rigidulum Bosc ex Nees, Redtop panicum, 9; F;

*Paspalum dilatatum Poir., Dallis grass, 5; S; 00-327

fal, Dat o ID L

Journal of

titute of Texas 1(1)

Paspalum floridanum Michx., Florida paspalum, 9; |; 98-715

Paspalum laeve Michx., Smooth beadgrass, 5; F; 01-186

*Phleum pratense L., Timothy, 3, 5; |; 02-220

**Poa annua L., Annual bluegrass, 5; F; 06-226

*Poa compressa L., Canada bluegrass, 5; O; 01-172

**Poa pratensis L., Kentucky bluegrass, 5; A; 02-216

Poa sylvestris A. Gray, Woodland bluegrass, 2; |; 00-328

Saccharum alopecuroides (L.) Nutt., Silver plume grass, 2, 3;

|; 01-632

Schizachyrium scoparium (Michx.) Nash, Little bluestem, 3,

4; A; 01-641

**Setaria faberi R.A. Herrm., Nodding foxtail, 5; |; 07-788

Setaria parviflora (Poir.) Kerguelen, Knotroot foxtail, 3, 4; F;

*Setaria pumila (Poir.) Roem. & Schult., Yellow foxtail, 5; |;

Sorghastrum nutans (L.) Nash, Indian grass, 3, 4; F; 98-642

**Sorghum halepense (L.) Pers., Johnson grass, 3; O; 01-194

Sphenopholis nitida (Biehler) Scribn., Shiny wedge grass, 2;

O; 99-102

Tridens flavus (L.) Hitchc., Purpletop, 3, 4, 5; F; 01-639

Tripsacum dactyloides (L.) L, Eastern gama-grass, 4; O; 98-97

t*Triticum aestivum L., Wheat, 3; R; 01-135

Urochloa platyphylla (Nash) R.D. Webster, Signal grass, 3;

R; 01-605

Vulpia octoflora (Walter) Rydb., Common six-weeks fescue,

2; A; 01-110

Ponteridiaceae

tPontederia cordata L., Pickerel-weed, 7; R; 98-66

Potamogetonaceae

tPotamogeton pusillus L., Slender pondweed, 6; F; 99-422

Smilacaceae

Smilax bona-nox L., Catbrier, 2, 3, 4; O; 98-117

Smilax glauca Walter, Glaucous greenbrier, 2, 3, 4; F; 98-239

Smilax hispida Raf., Bristly greenbrier, 2; R; 99-223

Smilax herbacea L. var. pulverulenta (Michx.) A. Gray, Carrion-

flower, 2; R; 99-370

Smilax rotundifolia L., Common greenbrier, 2, 3, 4; F; 99-273

Typhaceae

Typha latifolia L, Common cat-tail, 6, 7; F; 01-252

Uvulariaceae

Uvularia sessilifolia L., Sessileleaf bellwort, 2; O; 01-229

ACKNOWLEDGMENTS

My appreciation is extended to David S. White, Hancock Biological Station, Murray State University, for HBS

descriptive data, encouragement, critical review, and support, Robert F.C. Naczi, Delaware State University,

Dover, for identification and verification of the Cyperaceae, Ross C. Clark, Eastern Kentucky University,

for verification of woody plants, David D. Taylor, USDA Forest Service, for a critical review, and Melanie G.

Bentley, Eastern Kentucky University, for the five figures.

REFERENCES

BRAUN, E.L. 1950. Deciduous forests of eastern North America. Macmillan Publishing Co., New York.

Bryant, W.S. and M.E. H& b. An ordination of the plant communities of the Jackson Purchase Region of Kentucky.

Thompson, Flora of H k Biological Station in Kentucky 629

In: EW. Chester and A.F. Scott , eds. Proceedings of the ninth symposium on the natural history of Lower

Tennessee and Cumberland River Valleys. The Center for Field Biology, Austin Peay State University, Clarksville,

TN. Pp. 11-18.

CAMPBELL, J. and M. MebLev. 2006. Illustrated atlas of vascular plants in Kentucky: a first approximation. July 2006

draft. Privately printed, Lexington, Kentucky.

FENNEMAN, N.M. 1938. Physiography of eastern United States. McGraw-Hill Co., New York, NY.

GLEASON, H.A. and A. Cronauist. 1991. Manual of vascular plants of northeastern United States and adjacent Canada,

2nd Edition. The New York Botanical Garden, Bronx.

HUMPHREY, M.E., F.L. ANDERSON, R.A. Hayes, and J.D. Sims. 1973. Soil survey of Calloway and Marshall Counties, Kentucky.

United States Department of Agriculture, Soil Conservation Service, Washington, D.C.

Jones, R.L. 2005. Plant life of Kentucky: an illustrated guide to the vascular flora. The University Press of Kentucky,

Lexington.

Kentucky CLIMATE CENTER. 2006. The Kentucky Climate Center at Western Kentucky University. Climate Summa-

ries—Golden Pond 8 N, Kentucky (1971-2000). (http:// kyclim.wku.edu/ climate.stations/153295_viewer.

html). Accessed 17 Oct 2006.

Kentucky Exotic Pest PLANT Counci. 2000. Southeast Exotic Pest Plant Council: Kentucky invasive exotic plant list.

(http://www. invasive.org/seweeds.cfm). Accessed 15 Nov 2006.

KENTUCKY STATE NATURE PRESERVES Commission (KSNPC). 2000. Rare and extirpated biota of Kentucky. J. Kentucky Acad.

Sel 611157132.

Kentucky STATE NarURE PRESERVES COMMISSION (KSNPC). 2005. Rare and extirpated biota of Kentucky. (http://www.

naturepreserves.ky.gov/inforescources/reports pubs.htm). Accessed 25 Nov 2006.

Keys, JR., J., C. CARPENTER, S. Hooks, F. Lienic, W.H. McNas, W.E. Russett, and M-L Smith. 1995. Ecological units of the

eastern United States—first approximation (map and booklet of map unit tables). United States Department

of Agriculture, Forest Service, Southern Region Geometronics, Atlanta, GA.

KücHLER, A.W. 1964. Potential natural vegetation of the conterminous United States (map and accompanying

manual). American Geographical Society, Special Publication No. 36. New York, NY.

ManriN, W.H. and T.H. Tav Lon. 2002. Grasslands and grassland management in Land Between the Lakes. In: E.W.

Chester and J.S. Fralish, eds. Land Between the Lakes, Kentucky and Tennessee: four decades of Tennessee

Valley Authority Stewardship. The Center for Field Biology, Austin Peay State University, Clarksville, TN. Pp.

267-283.

MID-AMERICA REMOTE SENSING CENTER. 2003a. MSU Hancock Biological Station: Wetlands NWI (map). College of Sci-

ence, Engineering & Technology, Murray State University, Murray, KY.

MID-AMERICA REMOTE SENSING CENTER. 2003b. MSU Hancock Biological Station: Geologic formations (map). College

of Science, Engineering & Technology, Murray State University, Murray, KY.

MID-AMERICA REMOTE SENSING CENTER. 2003c. MSU Hancock Biological Station: Soils (map). College of Science, Engi-

neering & Technology, Murray State University, Murray, KY.

MoHLENBROCK, R.H. 1986. Guide to the vascular flora of Illinois. Revised and enlarged edition. Southern Illinois

University Press, Carbondale.

OBFS. 2006. Organization of Biological Field Stations. (http://www.obfs.org/). Accessed 2 Dec 2006.

SEELAND, D.A. and H.G. WiLsHIRE. 1965. Geologic map of part of the Rushing Creek Quadrangle in southwestern

Kentucky, Map GO-445. United States Geological Survey, Washington, D.C.

THoMPSON, R.L. and D.B. PoiNpExrER. 2006. Vascular flora of the Elk and Bison Prairie, Land Between the Lakes National

Recreation Area, Trigg County, Kentucky. Castanea 71:105-123.

USDA, NRCS. 2006. The PLANTS database. National Plant Data Center, Baton Rouge, LA. (http.:// plants.usda.

gov/). Accessed 7-11 Dec 2006.

Wurre, D.S. 2002. The Hancock Biological Station and the Center for Reservoir Research. In: EW. Chester and J.A.

Fralish, eds. Land Between the Lakes, Kentucky and Tennessee: four decades of Tennessee Valley Authority

stewardship. The Center for Field Biology, Austin Peay State University, Clarksville, TN. Pp. 205-215.

630 Journal of the Botanical R h Institute of Texas 1(1)

Woops, A.J., J.M. Omernik, W.H. Martin, G.J. Ponp, W.M. ANpnEWs, S.M. Cau, J.A. Comstock, and D.D. TavLor. 2002. Ecore-

gions of Kentucky (color poster with map, descriptive text, summary tables, and photographs): U.S. Geological

Survey, Reston, VA (map scale 1:000,000).

Woops, M. 1983. The vascular flora of Calloway County, Kentucky. M.S. thesis, Murray State University, Murray,

KY.

Woops, M. and MJ. Futter. 1988. The vascular flora of Calloway County, Kentucky. Castanea 53:88-109.

CHECISHTISIDSOPSTBBSVASCUIEABRSDINNSESSOE

GRAWLDORDCOUBCTYSPENINSYEIEVAINDS

Cynthia M. Morton Loree Speedy

Section of Botany Section of Botany

Carnegie Museum of Natural History Carnegie Museum of Natural History

4400 Forbes Avenue 4400 Forbes Avenue

Pittsburgh, Pennsylvania 15213, U.S.A. Pittsburgh, Pennsylvania 15213, U.S.A.

James K. Bissell

Section of Botany

Cleveland Museum of Natural History

Wade Oval Drive, University Circle

Cleveland, Ohio 44106, U.S.A.

ABSTRACT

Crawford County has been botanized since the early 1800s; however, a checklist of the county's vascular flora has not previously been

compiled. A search of herbaria augmented by targeted fieldwork has resulted in a list of 1168 taxa of native and naturalized plants

comprising 485 genera and 134 families. The five families, with the largest number of species, are Asteraceae, Poaceae, Cyperaceae,

Rosaceae, and Liliaceae. Carex, Potamogeton, Polygonum, Salix, and Rubus, are the five largest genera. There are approximately 208 non-

native species that have been introduced mainly from Europe and Eurasia. This checklist recognizes 10 species of Lycophytes, 46 species

of Pteridophytes, 7 species of Gymnosperms and 1105 species of Angiosperms. Ninety-seven species have global or state ranking.

RESUMEN

El Condado de Crawford, se localiza en el estado de Pennsylvania. Este condado es considerado de las áreas más diversas del estado,

sin embargo hasta ahora no ha sido De un Pa Este catálogo contiene familias, géneros, y taxa específicos que están orde-

nados alfabéticamente en los grup p l Pteridofitas, Gimnospermas y Angiospermas. Esta lista incluye los

1; 1

n el Condado de Crawford. Se encontraron un total de al menos 134

familias, 485g géneros, y 1168 especies. Las cinco DE mayores, por el número de especies, fueron Asteraceae, Poaceae, Cyperaceae,

Rosaceae y Liliaceae. Los seis géneros mayores, por el numero de especies, fueron Carex, Potamogeton, Polygonum, Salix y Rubus.

INTRODUCTION

Even though Pennsylvania lacks checklists for most of its counties, the vascular flora known thus far is quite

diverse. Rhoads and Klein (1993) reported 3318 taxa of vascular plants for the state, including 2076 native

and 1242 introduced. Approximately 25 of Pennsylvania's 67 counties have floras completed, but some

are unpublished theses and surveys and therefore unavailable to the public. The exact number of genera

and species in the flora of Crawford County was not previously known because a checklist had never been

compiled. Such checklists provide baseline information that can be used to monitor environmental changes

and guide conservation decisions.

Site Description

Crawford County is located in the northwestern corner of Pennsylvania, adjacent to Ohio (Fig. 1). It is

bordered by the counties of Erie to the north, Warren to the east, Venango to the southeast, and Mercer to

south. Named for Colonel William Crawford on March 12, 1800, the county contains approximately 2,690

sq. km. Crawford County receives total annual precipitation of approximately 97 cm in its southwestern

region and 112 cm in its northwestern region; 5596 of this precipitation usually falls between April and

September. Average season snowfall is 178 cm in the southwestern area, 230 to 240 cm in the central area,

and more than 300 cm in the northeastern area. The last frost of the season is usually around May 15th

and the first frost of the season around October 6th. The length of the growing season, from the last killing

J. Bot. Res. Inst. Texas 1(1): 631 — 653. 2007

Journal of

Ea DE p . ID

titute of Texas 1(1)

632

raus E Spartansburg

Beaver pn Venango "P Rockdale M —

Cussewago|Tw a wp

TWD [Springboro Twp g P Gambridge TWP Sparta Twp

Venango Ath

Conneautville_ J ens

Woodcock Twp Centerville jos

BODEN Saegertown Richmond

Conneaut wp Hayfield W k Twp

Twp Twp o Steuben

S ; Blooming unid WP Oil Creek

ummit Valley Twp

i Tun dvi Hydetown

Twp inesville Meadville East Randolph Troy Twp A

Vernon Mea Twp Titusville

North Pu Twp eer Twp

Shenango WP Teonneaut Twp

d Labs Union

East

South West |. TWP \eairfield Wayne

poenae Fallowfield TwpL Twp

n~ Tw Twp

West Ea t ai h irfi Cochranton

Shenango |Fallowfield wp Fairfield

Twp Twp Twp

0 5 10 20 Kilometers

0 5

10 Miles

Crawford County, Pennsylvania

T *" £, va $ D. I s .

Fic. 1 Top y, y p

frost in spring to the first killing frost in autumn, ranges from 145 to 150 days in the southwestern region

to 130 to 135 days in the northeastern region (USDA 1979).

The soils of the county consist mainly of the following 7 series, covering at least 1,963 sq. km., or 75%,

of Crawford County: Venango, Cambridge, Frenchtown, Holly, Red Hook, Sheffield, and Alden.

The Venango Series consists of nearly level upland flats, to knobs and side slopes of valleys that make

up about 29% of the county. These soils are weathered glacial till that contain sandstone, siltstone, and shale.

They have very slow permeability and a moderate available water capacity. Most areas are cleared and used

for crops or pasture or are idle.

The Cambridge Series consists of nearly level areas to upland knobs, side slopes of valleys, and crests

of slopes that make up about 13% of the county. These soils are weathered from glacial till that contains

sandstone, siltstone, and shale. They have slow permeability and a moderate available water capacity. Most

of the area is used for crops or pasture, but some land is wooded or idle.

The Frenchtown Series consists of nearly level and gently sloping areas that make up about 12% of

the county. These soils are weathered from glacial till that contains sandstone, shale, and some crystalline

rocks. They have slow permeability and a moderate available water capacity. Most of the area is wooded but

some areas are idle or used for crops or pasture land.

The Holly Series consists of nearly level soils that are on floodplains of major stream valleys that make

up about 9% of the county. These soils are weathered from recent stream deposits. They have moderate slow

Morton et al., Flora of Crawford County, Pennsylvania 633

permeability and a moderate to high available water capacity. Most of the area is used for pasture land but

some areas are woodland or idle or used for crops.

The Red Hook Series consists of nearly level to gently sloping soils on terraces in stream valleys that

make up about 496 of the county. These soils are weathered from glacial outwash. They have moderate per-

meability and a moderate to high available water capacity. Most of the areas are used for woodland, pasture,

or crops or are idle.

The Sheffield Series consists of poorly drained, nearly level soils that make up about 496 of the county.

These soils are weathered from glacial till that contains sandstone, shale, and some limestone. They have

very slow permeability and a moderate available water capacity. Most areas are woodlands but some contain

crops, pasture, or are idle.

The Alden Series consists of nearly level soils with slight depressions that make up about 496 of the

county. These soils are weathered from glacial till and local alluvium. They have low permeability and high

available water capacity. Most of the areas are woodlands, but pasture and idle lands are also present.

Crawford County is part of the Northwestern Glaciated Plateau section of the Appalachian Plateau

Province. The Northwestern Glaciated Plateau section consists of broad, rounded uplands cut by long, linear

valleys. The uplands are transected by flat-floored, narrow to wide valleys and are separated from adjacent

uplands by steep slopes on one or both sides of the valley. The valleys are oriented northwest-southeast and

are linear with the valley floors often containing wetlands. Elevation ranges from 900 to 2,200 feet (USDA

1979).

Crawford County contain stands of old growth, timber that has been logged very little or never. Old

growth makes up less than 596 of the forests remaining in the United States and contains unique trees that

make a significant contribution to forest biodiversity. One site is the Tryon-Weber Woods, which has 40

acres of rare hardwood old growth, containing American beech and sugar maples. This type of forest is the

only remaining stand in Pennsylvania. During early settlement, most of Pennsylvania was covered with

Eastern hemlock, white pine, and mixed hardwoods. From then until the early 1900s forests were logged

and as the forests were clear-cut, fast growing species quickly regenerated. The forest structure since then

has shifted to mixed hardwoods.

The effects of herbivory on individual species and isolated populations have been documented for

Crawford County; region wide effects on the flora are evident but not well documented. Over time, selective

browsing by densely populated deer can result in reduced species richness and altered species composition,

with dominance by a few non-preferred and browsing-resilient species. Unpalatable and browsing-resilient

species become abundant and can interfere with the reestablishment of preferred and less browsing-resilient

species. Overall, heavy browsing by deer in woody plant communities can change the trajectory of forest

vegetation development. Whether these changes for Crawford County are permanent is a matter of scientific

debate and will most likely be determined in due time and with additional studies.

Crawford County contains several major watersheds, which contain large wetland areas. Several major

watersheds include the Shenango River Watershed, French Creek Watershed, Oil Creek Watershed and the

Conneaut Creek Watershed.

The Crawford County area of the Shenango Creek Watershed comprises 120,700 acres. The tributaries

of Shenango Creek in Crawford County include Paden Creek, Linesville Creek and Crooked Creek. The

watershed includes a major impoundment, Pymatuning Lake, which is the largest man-made reservoir in

Pennsylvania. Pymatuning Lake drains into the Shenango system from two outlets. The first is Crooked

Creek in Crawford County, which flows southward through the Pymatuning Marsh and the second is the

Shenango River, which flows southward out of the western lobe of the lake between South and West Shenango

Townships. Two other features within the Crawford County portion of the Shenango River Watershed are

Crystal Lake and Dollar Lake. Both Crystal and Dollar Lakes are glacial kettle lakes formed during the last

lce Age,

The French Creek Watershed encompasses 1270 sq. mi. in area; with approximately 4196 of Crawford

634 Journal of the Botanical R h Institute of Texas 1(1)

County lying therein. Wetlands, such as marshes, swamps, and fens are all found within the French Creek

watershed. Conneaut Marsh is an example of a wetland that works to control flooding, filter excess nutrients

in the water and provide habitats to several aquatic species. French Creek contains major waterways and

many smaller streams. The area within the watershed has much overland flow, which picks up pollutants.

Consequently one of the major threats to the French Creek watershed is non-point source pollution. However,

to date it has been characterized as having very good to excellent water quality. The watershed is hailed as

one of the most ecologically diverse systems in Pennsylvania.

The Oil Creek Watershed is located in the eastern part of Crawford County. Oil Creek flows mainly

within Venango, Crawford and Warren Counties and is approximately 340 sq. mi. Crawford County comprises

167 sq. mi. of the watershed with dramatic topography of forested hillsides and clean clear streams.

The Conneaut Creek Watershed flows northward into Lake Erie near Conneaut, Ohio. The watershed

takes in approximately 96 sq. mi. Some streams include Main Branch, Middle Branch, and East Branch of

Conneaut Creek and Stone Run. Conneaut Creek Watershed is located in Beaver, Spring, Conneaut, Sum-

merhill and Summit Townships in Crawford County.

History

Initially the Seneca Indians dominated the region. The French arrived in 1748 and formed a short portage

between Presque Isle and Le Boeuf until 1758. Europeans did not settle the land again until 1788, when a

small party came out from Northumberland County and began to build an establishment where Meadville

now stands.

On March 12th of 1800 the Allegheny Legislature passed an act separating Crawford, Mercer, Venango,

Warren, and Erie counties from the territory of Allegheny County. Crawford County was named in honor

of Col. William Crawford, a staunch defender of the white settlers against Indian attacks.

French Creek, which flows from north to south, was the chief route for shipping local lumber to

Pittsburgh. The only other means of transportation was horse and wagon. In 1826 Major Douglass made

surveys for the development of a canal crossing to link the north to the south; this was the first constructed

transportation route in the county. The canal, called the Feeder Canal, was approximately 22 miles long. It

was completed in 1834, and canal boats began operating between Bemustown Dam and Conneaut Lake.

Before the 1800s there were no roads at all. Until the Erie and Waterford Turnpike Company was

chartered in 1805, the settlers would just make their way through the woods. The Meadville-Waterford-

Erie Turnpike was constructed between 1806 and 1809 and used for thirty-five years until it became the

Susquehanna and Waterford Turnpike. By 1820 this road extended from Waterford to Bellefonte in Centre

County and by 1824 to Philadelphia. The Turnpike was used from 1824 until it was abandoned in 1845. The

Mercer and Meadville Turnpike was completed in 1821 and is still known to county residents as the Mercer

Pike. In 1859 Col. E.L. Drake successfully drilled a well that produced oil in eastern Crawford County. This

development brought prosperity and railroads to Titusville.

The lack of adequate shipping facilities had hindered the development of local manufacturing as well as

the marketing of local produce outside the county. The completion of Atlantic and Great Western Railways of

Pennsylvania (now the Erie Railroad) to Meadville in 1862 stimulated economic growth. In 1913 Col. Lewis

Walker purchased a small enterprise, first called the Automatic Hook and Eye Company and later Talon,

Inc., which eventually made Meadville the zipper capital of the world. Railroad construction boomed from

1860 to 1866. By 1871 tracks ran from Bloomfield, Steuben, Troy, and Oil Creek to Titusville and became

part of the Buffalo, New York, and Philadelphia Line. Today Crawford County has more miles of railroad

than any county in Pennsylvania.

Industry developed gradually, and today most boroughs have manufacturing enterprises that provide

substantial employment. The five principal industries are agriculture, gas and oil production, textiles and

textile products, and metals and metal products, and allied products. Today about 4096 of Crawford County

is farmed, 4696 is used for commercial woodland, and the rest is being developed for urban use or recre-

ation. Dairying is the primary farming; Crawford is one of Pennsylvania's top 10 milk-producing counties.

Morton et al., Flora of Crawford County, Pennsylvania 635

Principal crops are corn, wheat, oats, and hay. Lumber, pulpwood, maple syrup, and Christmas trees from

the woodland areas also generate income. Besides agricultural and industry, a thriving resort business at-

tracts thousands of summer visitors to the recreational facilities of Conneaut Lake, Pymatuning Reservoir,

Cambridge Springs, and Canadohta Lake (Bates 1885).

In 1800 the county population was approximately 2,345; according to the census of 2000, Crawford

County has 90,366 residents (USDA (1979, 2000).

Major Collectors

Collecting in Crawford County for the Carnegie Museum Herbarium began in the early 1800s and has

continued to the present. Most of the collectors have been members of the Botanical Society of Western

Pennsylvania and the staff of the Section of Botany at Carnegie Museum.

Botanists who have made significant contributions to the knowledge of the Crawford County are: J.K.

Bissell, J.A. Isaac, L. Speedy, R.C. Leberman, S.P. Grund and O.E. Jennings.

METHODS

This checklist was compiled by searching the herbaria of the Carnegie Museum of Natural History (CM) and

the Cleveland Museum of Natural History (CLM) for Crawford County specimens. In addition, fieldwork

was conducted targeting underrepresented areas of the county. Most of the collections were made during the

last two years, but several specimens date back as far as 1881 and 1882. The majority of the specimens are

deposited at the Carnegie Museum of Natural History (CM). Herbarium collection information was obtained

from the Morris Arboretum (MOAR) and the Cleveland Museum of Natural History (CLM) for 189 samples.

Other herbaria that were examined for specimens include the Academy of Natural Sciences of Philadelphia.

Rhoads and Block (2000) was the primary source for plant identification. For generic and species names, we

have generally followed the Synthesis of North American Flora (Kartesz 1999). Authorities are abbreviated

for the majority of taxa according to Brummitt and Powell (1992).

RESULTS AND DISCUSSION

This list includes the names of all native and naturalized species known to occur in Crawford County. It

includes a total of 134 families, 485 genera, and 1168 species. The five families with the largest number

of species, are Asteraceae, Poaceae, Cyperaceae, Rosaceae, and Liliaceae. Carex, Potamogeton, Polygonum,

Salix and Rubus, are the largest genera. This checklist recognizes 10 species of Lycophytes, 46 species of

Pteridophytes, 7 species of Gymnosperms and 1105 species of Angiosperms. There are approximately 208

non-native species that have been introduced mainly from Europe and Eurasia. Ninety-seven species have

global or state ranking.

Of the 97 plants that have global or state ranking only three taxa have a global ranking of G3 or vul-

nerable status (Potamogeton hillii, Poa paludigena, and Platanthera leucophaea). The remaining 94 plants are

either a G4 or G5 status, indicating an apparently secure or secure condition globally.

The genus Potamogeton is in the family Potamogetonaceae. Potamogeton hillii is one of the smaller linear-

leaved pondweeds and can be difficult to recognize. This herbaceous perennial is totally submerged except

for the flowering spike. The leaves range from 1.5-2.0 mm wide and are bristle-tipped, and 3-veined. The

flowers are arranged in very short, few flowered spikes which are nearly globose. It occurs in cold, clear,

slow-moving water in streams, ponds, and beaver ponds with a muddy substrate.

Poa is part of the grass family (Poaceae). Most members of this genus are very similar and require

microscopic identification of the flower parts to identify. Poa paludigena is a perennial with culms forming

loose clumps. Culm internodes are terete with leaf-sheaths scaberulous. Leaf blades are erect and 0.3-2 mm

wide. The inflorescence is a panicle, with branches in pairs. Spikelets are comprised of 2-5 fertile florets

and are oblong and laterally compressed. Florets are slightly woolly. It typically grows in moist, open areas

such as meadows or moist woods and can be found among sphagnum or other mosses. This species can be

easily overlooked or misidentified due to its similarity to closely related species.

fal, Dat o ID L

636 Journal of

titute of Texas 1(1)

Platanthera is a member of the Orchid family. The specific epithet leucophaea refers to the off-white color

of the flowers. This species forms a cluster of fleshy thickened roots at the base. The 2-5 leaves are lanceolate

with the bases sheathing the stem. The inflorescence is a raceme of 5-20 flowers, creamy-white and strongly

fragrant at dusk. The basal petal, the labellum, is tripartite and fringed. It is found almost exclusively in

moist prairies or open sphagnum bogs. Currently it is found in less than sixty sites in the U.S., many with

only a few individuals.

There are four taxa in the Crawford County flora that are listed by the Pennsylvania Department of

Agriculture (2004) as noxious weeds: Cirsium arvense (canadian thistle), Cirsium vulgare (bull or spear thistle),

Lythrum salicaria (purple loosestrife), and Rosa multiflora (multiflora rosa). Other species considered invasive

in our native ecosystems are Acer platanoides (Norway maple), Lonicera japonica (Japanese honeysuckle),

Phragmites australis (Common reed), Polygonum cuspidatum (Japanese knotweed), Elaeagnus umbellata (Au-

tumn olive), Lonicera morrowii (Morrow's honeysuckle), and Lonicera tatarica (Tatarian honeysuckle) (DCNR

2000).

While this checklist is probably not all-inclusive of every species in Crawford County, it is the most

comprehensive list presently available.

ANNOTATED CHECKLIST OF THE SPECIES OF

CRAWFORD COUNTY, PENSYLVANIA

Taxa are listed according to the following format: taxon name, author(s), ((year) collector and number} global:

state ranking [Synonyms] and non-native source. We follow the state (S) and global (G) ranking systems

developed by The Nature Conservancy (1996 version). The global numbers are designated from 1 (critically

imperiled) to 5 (secure). Other notations include SH, which denotes historical occurrence, and SR, which

indicates reported without persuasive documentation. Synonyms are included for names not in common

usage in the state or regional manuals. In cases where there was more than one specimen present in the

collection, recent collections of current known collectors for the western Pennsylvania region were cited.

Families, genera, and specific and infraspecific taxa are arranged alphabetically within vascular plant

groups Angiosperms, Gymnosperms, Lycophytes, and Pteridophytes.

ANGIOSPERMS Amaranthaceae

Amaranthus albus L., 1918 E.M. Gress s.n.

Amaranthus blitum L., 2005 R. Thompson 05-504 Tropical

erica

Acanthaceae

Justicia americana (L.) Vahl, 1988 J.K. Bissell et al. 88:101

Aceraceae Amaranthus hybridus L., 1947 H.A. Wahl 4076

Acer negundo L., 1997 R.C. Leberman. s.n. Amaranthus retroflexus L., 1948 S. Tisherman s.n. Tropical

Acer nigrum Michx. f, 1922 O.E. Jennings. s.n. America

Acer pensylvanicum L., 2003 R Coxe and M. Bradburn s.n.

Acer platanoides L., 1997 R.C. Leberman s.n. Europe

Acer rubrum L. var. rubrum, 2005 L. Speedy LSB329

Acer rubrum L. var. trilobum Torr. & A. Gray ex K. Koch, 1952

L.K. Henry and W.E. Buker s.n.

Acer saccharinum L., 2003 S. Ernst et al. 71

Acer saccharum Marsh. var. saccharum, 1955 L.K. Henry s.n.

Acer spicatum Lam., 1932 W.R. Van Dersal 1866

Acoraceae

Acorus americanus (Raf.) Raf., 2002 J.K. Bissell 2002:134 G5:51

Acorus calamus L., 1999 J.K. Bissell and B. Danielson 1999:194

Furope

Alismataceae

Alisma subcordatum Raf., 2003 J.K. Bissell 2003:148

Sagittaria latifolia Willd., 2004 S.P.Grund and L. Miller 3487

Sagittaria rigida Pursh, 2001 J.K. Bissell 2001:149

Anacardiaceae

Rhus glabra L., 2004 J.A. Isaac 17601

Rhus typhina L., 1997 R.C. Leberman s.n.

Toxicodendron radicans (L.) Kuntze, 1997 J.A. Isaac 9878

Toxicodendron vernix (L) Kuntze, 2005 L. Speedy LSB1565

Annonaceae

Asimina triloba (L.) Dunal, 1955 D.L. Pearth s.n.

Apiaceae

Angelica atropurpurea L., 1928 J.A. Murray s.n.

Angelica triquinata Michx., 1980 C.E. Jenkins et al. 2777

Angelica venenosa (Greenway) Fern, 1901 J.A. Shafer s.n.

Chaerophyllum procumbens (L.) Crantz, 2004 S.P. Grund and

L.Miller 3534

Cicuta bulbifera L., 2005 L. Speedy, M. Bowers and M. Fodse

LSB1608

Cicuta maculata L., 2000 J.A. Isaac and C.F. Chuey 13168

Morton et al., Flora of Crawford County, Pennsylvania

Conium maculatum L., 1980 F.Q. Jenkins and C.E. Jenkins

2836 Europe

Cryptotaenia canadensis (L.) DC., 2004 J.A. Isaac 17468

Daucus carota L., 2000 J.A. Isaac and C.F. Chuey 13170

Eurasia

Erigenia bulbosa (Michx.) Nutt, 2004 J.K. Bissell 2004:006

(95:52

Heracleum maximum Bartr., 2000 J.A. Isaac and C.F. Chuey

13167

Hydrocotyle americana L., 2004 J.K. Bissell 2004:114

Osmorhiza claytonii (Michx.) C.B. Clarke, 1972 W.E. Buker s.n.

Osmorhiza longistylis (Torr.) DC., 1972 W.E. Buker s.n.

Pastinaca sativa L., 1968 R.C. Leberman s.n. Eurasia

Sanicula canadensis L. var. canadensis, 2004 J.A. Isaac 17678

Sanicula marilandica L., 1989 J.K. Bissell and B. Danielson

Sanicula odorata (Raf.) K.M. Pryer & L.R. Phillippe, 1984 J.K.

Bissell 84:120

Sium suave Walt., 2005 L. Speedy LSB1557

Taenidia integerrima (L.) Drude, 1972 W.E. Buker s.n.

Thaspium barbinode (Michx.) Nutt, 1997 R.C. Leberman s.n.

Zizia aurea (L.) W.D.J. Koch, 1962 W.E. Buker s.n.

Apocynaceae

Apocynum androsaemifolium L., 1997 R.C. Leberman s.n.

Apocynum cannabinum L., 2005 L. Speedy 761

Vinca minor L., 1919 H.W. Mossman s.n. Europe

Aquifoliaceae

llex montana Torr. & A. Gray ex Gray, 1975 R.C. Leberman s.n.

llex verticillata (L.) A. Gray, 2005 L. Speedy LSB1091

Nemopanthus mucronatus (L.) Loes., 2004 J.A. Isaac 17593

Araceae

Arisaema dracontium (L.) Schott, 1995 R.C Leberman. s.n.

Arisaema triphyllum (L.) Schott ssp. stewardsonii (Britt.)

Huttleston, 1994 J.D Wagner et al. 114

Arisaema triphyllum (L.) Schott spp. triphyllum, 2004 S.P. Grund

and. L. Miller 3642

Arisaema triphyllum (L.) Schott ssp. pusillum (Peck) Huttleston,

.A. Darling s.n.

Calla palustris L., 2004 J.A. Isaac18474

Peltandra virginica (L.) Schott, 2005 L. Speedy, M. Bowers, and

M.F B1607

Symplocarpus foetidus (L) Salisb. ex Nutt., 2005 L. Speedy

LSB337

—

Araliaceae

Aralia hispida Vent., 1962 L.K. Henry s.n.

Aralia nudicaulis L., 1996 R.C. Leberman s.n.

Aralia racemosa L., 1996 R.C. Leberman s.n.

Aralia spinosa L., 2005 R. Thomspson 05-858

Panax quinquefolium L., 1962 R.C. Lebermans.n.

Panax trifolius L., 2005 L. Speedy LSB349

Aristolochiaceae

Asarum canadense L., 1972 W.E. Buker s.n.

Asclepiadaceae

Asclepias exaltata L., 1981 R.C. Leberman s.n.

Asclepias incarnata L. ssp. incarnata, 2005 L. Speedy, J. Bissell,

M. Bowers and T.Pearson LSB1583

637

Asclepias syriaca L., 2004 J.A. Isaac 17609

Asteraceae

Achillea millefolium L., 2004 J.A Isaac and. R. Coxe 17590

Furasia

Ageratina altissima (L.) King & H.E. Robins. var. altissima, 1998

R. Leberman s.n.

Ambrosia artemisiifolia L. var. elatior (L.) Descourtils, 1928 J.A.

Ambrosia trifida L. var. trifida, 1928 J.A. Murray Sh:

Anaphalis margaritacea (L.) Benth., 1968 W.E. Buker s.n.

Antennaria howellii Greene ssp. neodioica (Greene) Bayer, 1950

L.K. Henry and W.E. Buker s.n.

Antennaria neglecta Greene, 1984 AW. Cusick 23365

Antennaria pani Fern. ssp. parlinii Fern., 1971 W.E. Buker

S.n.

Anthemis arvensis L., 1980 R.C. Leberman s.n. Europe

Anthemis cotula L., 2004 J.A. Isaac 17602 Europe

Arctium lappa L., 2000 J.A. Isaac and C.F. Chuey 13172

Eurasia

Arctium minus Bernh., 1970 W.E. Buker s.n. Eurasia

Artemisia vulgaris L., 1969 W.E. Buker s.n. Eurasia

Bellis perennis L., 2004 S.P. Grund and L. Miller 3588 Europe

idens cernua L., 1991 R.C. Leberman s.n.

Bidens connata Muhl. ex Willd., 2005 L. Speedy, J. Bissell, M.

Bowers and T. Pearson LSB1558

Bidens coronata (L.) Britt., 2004 J.A. Isaac 18456

Bidens discoidea (Torr. & A. Gray) Britt., 2003 J.K. Bissell

2003) 725555

Bidens frondosa L., 1969 W.E. Buker s.n.

Bidens laevis (L.) B.S.P., 1868 Thomas C. Porter s.n. G5:53

Bidens tripartita L., 2004 J.A .Isaac et al. 17722

Bidens vulgata Greene, 1944 O.E. Jennings s.n.

Centaurea biebersteinii DC., 2003 S. Ernst 32 Exotic

Cichorium intybus L., 1928 J.A. Murray s.n. Europ

Cirsium arvense (L.) Scop., 1980 C.E. Jenkins M 7 Jenkins

2883 Eurasia

Cirsium muticum Michx.,

1987 J.K. Bissell and B. Danielson

Cirsium pumilum (Nutt.) Spreng., 1928 J.A. Murray s.n.

Cirsium vulgare (Savi) Ten., 1969 W.E. Buker s.n. Eurasia

Conyza canadensis (L.) Cronq. var. canadensis, 1997 W.E.

Tomon W.E. s.n.

Coreopsis tripteris L., 1994 K. Marsh et al. s.n.

Crepis capillaris (L.) Wallr., 2004 J.A. Isaac and R. Coxe 17589

Europe

Doellingeria infirma (Michx.) Greene, 2005 L. Speedy et al.

1569 s.n. [Aster infirmus Michx.]

Doellingeria umbellata (P. Mill.) Nees var. umbellata, 2004 S.P.

Grund and L. Miller 3668

Erechtites hieraciifolia (L.) Raf. ex DC. var. hieraciifolia, 1992 K.

Marsh et al. s.n.

Frigeron annuus (L.) Pers., 2004 J.A. Isaac 17699

Erigeron philadelphicus L. var. philadelphicus, 1972 W.E. Buker

S.n.

Erigeron pulchellus Michx. var. pulchellus, 1982 W.E. Buker and

Grisez s.n

Erigeron strigosus Muhl. ex Willd. var. strigosus, 1928 J.A.

Murray s.n.

638

Fupatorium altissimum L., 1998 K. Marsh s.n.

Eupatorium fistulosum Barratt, 1984 J.K. Bissell 1984:119

Eupatorium maculatum L. var. maculatum, 2005 L. Speedy, J.

Bissell, M. Bowers and T.Pearson LSB1566

Eupatorium perfoliatum L. var. perfoliatum, 2005 L. Speedy

and L. Armstrong LSB1598

Eurybia chlorolepis (Burgess) Nesom, 1997 J.D. Wagner and G.

Podniesinski 1780 [Aster chlorolepis Burgess]

Eurybia macrophylla (L.) Cass, 1991 R.C. Leberman s.n.

Eurybia schreberi (Nees) Nees, 1997 J.K. Bissell et al. 1997182

[Aster schreberi Nees]

Euthamia graminifolia (L) Greene var. graminifolia, 2004 S.P.

Grund and L. Miller 3601

Galinsoga quadriradiata Cav., 1994 K. Marsh and G. Marsh s.n.

Central & South America

Gamochaeta purpurea (L.) Cabrera, 1901 J.A. Shafer s.n.

Gnaphalium uliginosum L., 1962 L.K. Henry. s.n. Europe

Hasteola suaveolens (L.) Pojark., 1998 J.K. Bissell 1998:074

Helenium autumnale L., 1951 L.K. Henry and W.E. Buker s.n.

Helenium flexuosum Raf., 2006 S.P. Grund 4137

Helianthus microcephalus Torr. & Gray, G.W. Clinton s.n. G5:

Helianthus tuberosus L., 1970 W.E. Buker s.n.

Heliopsis helianthoides (L.) Sweet, var. helianthoides, 1983 R.C.

e n

Hieracium aurantiacum L., 2004 J.A. Isaac 17489 Europe

Hieracium caespitosum Dumort., 2004 J.A. Isaac 17460

Furope

Hieracium gronovii L., 1965 D.L. Pearth s.n.

Hieracium paniculatum L., 1887 B.H. Patterson s.n.

Hieracium pilosella L. var. pilosella, 2004 J.A. Isaac 17487

Europe

Hieracium scabrum Michx. var. scabrum, 1928 J.A. Murray

Hieracium venosum L., 1996 R.C. Leberman s.n.

Hypochaeris radicata L., 1980 R.C. Leberman s.n. Eurasia

Inula helenium L., 2004 J.A. Isaac 17686 Europe

Krigia biflora (Walt.) Blake var. biflora, 2004 J.A. Isaac 17481

Lactuca biennis (Moench) Fern., 1951 L.K. Henry and W.E.

Lactuca canadensis L., 1959 L.K. Henry and F.H. Beer s.n.

Lactuca floridana (L.) Gaertn. var. floridana, 1970 W.E. Buker

Lactuca saligna L., 1970 W.E. Buker s.n. Europe

Lactuca sativa L., 2003 S. Ernst 117 Exotic

Leucanthemum vulgare Lam., 2004 J.A. Isaac and R. Coxe

17585 Exotic

Matricaria discoidea DC., 2004 J.A. Isaac 17453 Exotic

Megalodonta beckii (Torr. ex Spreng.), Greene 2003 S. Ernst

et al. 102 G4G5:S1

Packera aurea (L.) A. & D. Léve, 2005 L. Speedy LSB346 [Senecio

aureus L.

Packera obovata (Muhl. ex Willd.) W.A. Weber & A. Lóve, 1971

W.E. Buker s.n. [Senecio obovatus Muhl.ex Willd.]

Petasites hybridus (L.) P.G. Gaertn., B. Mey. & Scherb., 1968 M.

Bayliss s.n. Europe

Prenanthes alba L., 1994 K. Marsh et al. s.n.

fal, Dat o ID L

Journal of

titute of Texas 1(1)

Prenanthes altissima L., 2005 R. Thompson 05-1041

Pseudognaphalium macounii (Greene) Kartesz, 2002 J.A.

Isaac 15721

Pseudognaphalium obtusifolium (L.) Hilliard & Burtt ssp. obtu-

sifolium, 1997 T.J. Tomon s.n.

Rudbeckia hirta L. var. hirta, 1980 C.E. Jenkins 2851

Rudbeckia hirta L. var. pulcherrima Farw., 1952 L.K. Henry and

F.H. Beer s.n.

Rudbeckia laciniata L. var. laciniata, 2004 J.A. Isaac 18457

Sericocarpus asteroides (L.) B.S.P., 1928 J.A. Murray s.n.

Silphium perfoliatum L. var. perfoliatum, 1994 K. Marsh et

al. s.n.

Solidago altissima L., 1997 J. Long s.n.

Solidago bicolor L., 1970 W.E. Buker s.n.

Solidago caesia L., 1970 W.E. Buker s.n.

Solidago canadensis L. var. canadensis, 2005 L. Speedy

LSB1571

Solidago flexicaulis L., 1969 W.E. Buker s.n.

Solidago gigantea Ait., 2005 L. Speedy LSB1119

Solidago juncea Ait., 1959 L.K. Henry and F.H. Beer s.n.

Solidago nemoralis Ait., 1991 R.C. Leberman s.n.

Solidago patula Muhl. ex Willd. var. patula, 2004 J.A. Isaac

18440

Solidago rugosa P. Mill var. aspera (Ait) Cronq., 1949 O.E.

Jennings and C.M. Boardman s.n.

Solidago rugosa P. Mill var. rugosa, 2005 L. Speedy LSB1559

Solidago uliginosa Nutt. var. uliginosa, 1914 J. Bright s.n.

G4G5:53

Solidago uliginosa Nutt. var. linoides (Torr. & A. Gray) Fern,

1940 J. Wurdack Jr. s.n. G4G5:SH

Sonchus arvensis L. ssp. arvensis, 2004 J.A. Isaac 17706

Europe

Sonchus arvensis L. ssp. uliginosus (Bieb.) Nyman, 1968 W.E.

Buker s.n. Europe

Sonchus asper (L.) Hill, 2005 R. Thompson 05-1094 Europe

Symphyotrichum boreale (Torr. & A.Gray) A. & D. Lóve, 1993

J.K. Bissell and B. Danielson 1993:182 G5:51

Symphyotrichum lanceolatum (Willd.) Nesom ssp. /anceola-

tum, 2002 J.A. Isaac 15713 [Aster lanceolatus Willd. ssp.

lanceolatus var. lanceolatus]

Symphyotrichum lateriflorum (L.) A. & D. Love var. lateriflorum

2004 J.A. Isaac 18478 [Aster lateriflorus (L.) Britton]

Symphyotrichum lowrieanum (Porter) Nesom, 1994 K. Marsh

et al. s.n.

Symphyotrichum novae-angliae (L) Nesom, 2004 J.A. Isaac

18477 [Aster novae-angliae L.

Symphyotrichum pilosum (Willd.) Nesom var. pilosum, 2004

J.A. Isaac 18479 [Aster pilosus Willd]

Symphyotrichum pilosum (Willd) Nesom var. pringlei (Gray)

Nesom, 1971 W.E. Buker s.n.

Symphyotrichum praealtus (Poir.) Nesom var. praealtus, 1997

J.K. Bissell et al. 1997:077 G5:S3

Symphyotrichum prenanthoides (Muhl. ex Willd.) Nesom, 2004

J.A. Isaac 18461 [Aster prenanthoides Muhl. ex Willd]

Symphyotrichum puniceum (L.) A. & D. Lóve var. puniceum,

2005 L. Speedy LSB1617 [Aster puniceus L.]

urophyllum (Lindl.) Nesom, 1971 W.E. Buker

Symphyotricl

Morton et al., Flora of Crawford County, Pennsylvania

Symphyotrichum x longulum (Sheldon) Nesom [boreale x

puniceum], 1993 J.K. Bissell and B. Danielson 1993:188

Tanacetum vulgare L., 1935 A.G. Dickey s.n. Europe

Taraxacum officinale G. Weber ex Wiggers ssp. officinale, 1997

J. Long s.n. Eurasia

Tragopogon dubius Scop., 2004 J.A. Isaac and R. Coxe 17576

Furope

Tragopogon porrifolius L., 1947 W.E. Bukers.n. Europe

Tragopogon pratensis L., 2004 J.A. Isaac 17716 Europe

Triadenum fraseri (Spach) Gleason, 1994 J.D. Wagner 664

Triadenum virginicum (L.) Raf., 2005 L. Speedy LSB1623

Tussilago farfara L., 2004 J.A. Isaac 17317 Eurasia

Verbesina alternifolia (L.) Britt. ex Kearney, 1946 N. Russell

Vernonia gigantea (Walt.) Trel., 1997 J.A. Isaac 9872

Balsaminaceae

Impatiens capensis Meerb., 2005 L. Speedy LSB1114

Impatiens pallida Nutt., 1940 R.W. Little s.n.

Berberidaceae

Berberis thunbergii DC., 2005 R. Thompson 05-1051 Japan

Berberis vulgaris L., 2004 J.A. Isaac 18463 Europe

Caulophyllum thalictroides (L.) Michx., 2004 J.A. Isaac 17357

Podophyllum peltatum L., 1934 O.E. Jennings s.n.

Betulaceae

Alnus incana (L.) Moench ssp. rugosa (Du Roi) Clausen, 2005

L. Speedy, M. Bowers and M. Fodse LSB1618

Alnus serrulata (Ait.) Willd., 2005 L. Speedy, M. Bowers and

M. Fodse LSB1609

Betula alleghaniensis Britt. var. alleghaniensis, 2005 L. Speedy

LSB341

Betula alleghaniensis Britt. var. macrolepis (Fern) Brayshaw,

.E. Jennings s.n.

Betula lenta L., 1950 L.K. Henry and W.E. Buker s.n.

Betula pendula Roth, 1952 L.K. Henry and F.H. Beer s.n.

Eurasia

Betula populifolia Marsh., 1997 R.C. Leberman s.n.

Corylus americana Walt., 1998 R.C. Leberman s.n.

Ostrya virginiana (P. Mill.) K. Koch var. virginiana, 1997 R.C.

Leberman s.n.

Boraginaceae

Cynoglossum officinale L., 1915 O.E. Jennings and G.K.

Jennings s.n. Eurasia

Cynoglossum virginianum L. var. virginianum, 1925 O.E.

Jennings s.n.

Echium vulgare L., 2004 J.A. Isaac and R. Coxe 17575 Europe

Hackelia virginiana (L.) 1.M. Johnst., 2005 R. Thompson 05-

Lappula squarrosa (Retz.) Dumort., 1914 J. Bright s.n. Eurasia

Myosotis laxa Lehm., 2004 S.P. Grund and L. Miller 3488

Myosotis scorpioides L., 2005 L. Speedy LSB741 Europe

Symphytum officinale L., 1928 J.A. Murray s.n. Eurasia

Brassicaceae

Alliaria petiolata (Bieb.) Cavara & Grande, 2005 R. Thompson

05-653 Europe

Arabidopsis thaliana (L.) Heynh., 2004 J.A. Isaac 17454

Furope

639

Arabis glabra (L.) Bernh., 1905 O.E. Jennings s.n.

Arabis lyrata L., 1930 W.R. VanDersal s.n.

Armoracia rusticana P.G. Gaertn., B. Mey. & Scherb., 1909 O.E.

Jennings s.n. Eurasia

Barbarea vulgaris Ait. f., 2005 L. Speedy LSB325 Eurasia

Brassica juncea (L) Czern., 1971 R.C. Leberman s.n. Eurasia

Brassica nigra (L.) W.D.J. Koch, 2004 J.A. Isaac 17713 Eurasia

Brassica rapa L. var. rapa, 1949 L.K. Henry s.n. Europe

Capsella bursa-pastoris (L.) Medik., 2004 J.A. Isaac 17352

Eurasia

Cardamine angustata O.E. Shulz, 1989 J.K. Bissell et al.

89:0

:021

Cardamine bulbosa (Schreb. ex Muhl.) B.S.P,, 2005 L. Speedy

LSB3360

Cardamine concatenata (Michx.) Sw., 2004 J.A. Isaac 17337

Cardamine diphylla (Michx.) Wood, 2005 L. Speedy LSB351

Cardamine douglassii Britt., 2005 L. Speedy LSB330

Cardamine pensylvanica Muhl. ex Willd., 2005 L. Speedy

LSB305

Cardamine pratensis L. var. pratensis, 1994 J.K. Bissell et al.

1994:044 G5T5:51

Cardamine rotundifolia Michx., 1993 J.K. Bissell et al. 93:148

Draba verna L., 1958 W.E. Buker s.n. Europe

Erysimum cheiranthoides L., 1909 O.E. Jennings s.n. Eurasia

Erysimum inconspicuum (S. Watson) MacMill.ill. var. incon-

spicuum, 1951 L.K. Henry s.n.

Hesperis matronalis L., 2004 S.P. Grund and L. Miller 3632

Europe

Lepidium campestre (L.) Ait. F., 1982 R.C. Leberman s.n.

Eurasia

Lepidium virginicum L. var. virginicum, 1981 W.E. Buker s.n.

Raphanus raphanistrum L., 1949 L.K. Henry s.n.

Mediterranean

Raphanus sativus L., 1914 J. Bright s.n. Mediterranean

Rorippa nasturtium-aquaticum (L) Hayek, 2005 L. Speedy

743

—

LSB

Rorippa palustris (L.) Bess. ssp. fernaldiana (Butters & Abbe

Jonsell, 1995 R.C. Leberman s.n.

Rorippa palustris (L.) Bess. ssp. hispida (Desv.) Jonsell, 1901

Rorippa sylvestris (L.) Bess., 1995 R.C. Leberman s.n. Europe

Sisymbrium altissimum L., 1960 W.E. Buker s.n. Eurasia

Sisymbrium officinale (L.) Scop., 1901 J.A. Shafer s.n. Europe

Thlaspi arvense L., 1930 O.E. Jennings s.n. Europe

Cabombaceae

Brasenia schreberi J.F. Gmel., 2004 L. Miller and S.P. Grund 22

Cabomba caroliniana Gray var. caroliniana, 2003 S. Ernst et

Caesalpiniaceae

Senna hebecarpa (Fern.) Irwin & Barneby, 1947 H.A. Wahl

4184

Callitrichaceae

Callitriche heterophylla Pursh ssp. heterophylla, 1991 J.K. Bissell

et al. 1991:058

Callitriche palustris L., 1988 J.K. Bissell et al. 88:130

640

Campanulaceae

Campanula aparinoides Pursh, 1988 J.K. Bissell et al.

1988:135

Campanula rapunculoides L., 1970 W.E. Buker s.n. Eurasia

Campanulastrum americana (L.) Small., 2002 J.K. Bissell and B.

Danielson 2002:156 [Cyanococcus fascatus Small)

Lobelia cardinalis L., 1988 J.K. Bissell et al. 1988:135

Lobelia inflata L., 1959 L.K. Henry and F.H. Beer s.n.

Lobelia siphilitica L. var. siphilitica, 2004 J.A. Isaac 18460

Lobelia spicata Lam. var. spicata, 1922 O.E. Jennings s.n.

Capparaceae

Polanisia dodecandra (L.) DC. ssp. dodecandra, 1950 W.E.

Buker s.n.

Caprifoliaceae

Diervilla lonicera P. Mill, 1997 R.C. Leberman s.n.

Lonicera canadensis Bartr. ex Marsh., 1997 J.D. Wagner and

G. Podniesinski 1776

Lonicera dioica L., 2004 S.P. Grund and L. Miller 3573

Lonicera morrowii Gray, 2004 S.P. Grund and L. Miller 3616

Japan

Lonicera oblongifolia (Goldie) Hook., 1993 J.K. Bissell and B.

Danielson 1993:119 G4:S1

Lonicera tatarica L., 1980 C.E. Jenkins and F.Q. Jenkins 2873

Eurasia

Lonicera villosa (Michx.) J.A. Schultes, 1994 J.D. Wagner 905

Sambucus nigra L. ssp. canadensis (L.) R. Bolli, 2005 L. Speedy

LSB736

Sambucus racemosa L. var. racemosa, 1995 R.C. Leberman

STE

Triosteum perfoliatum L., 1940 O.E. Jennings s.n.

Viburnum acerifolium L., 2004 J.A. Isaac 17461

Viburnum lentago L., 2005 L. Speedy LSB1129

Viburnum nudum L. var. cassinoides (L.) Torr. & A. Gray, 2004

J.A. Isaac 18437 G5:S1

Viburnum opulus L. var. americanum Ait., 2004 J.A. Isaac

17503 [Viburnum opulus L. ssp. trilobum (Marsh.) Clausen]

G5T5:5354

Viburnum opulus L. var. opulus, 2004 J.A. Isaac 17599 Eurasia

Viburnum rafinesquianum J.A. Schultes, 1980 C.E. Jenkins and

F.Q. Jenkins 2862

Viburnum recognitum Fern., 1994 J.D. Wagner et al. 93

Caryophyllaceae

Agrostemma githago L., 1922 O.E. Jennings s.n. Europe

Arenaria serpyllifolia L., 2004 J.A. Isaac and R. Coxe 17586

Furope

Cerastium fontanum Baumg., 1904 O.E. Jennings s.n. Eurasia

Cerastium glomeratum Thuill., 2000 A.W. Cusick 35372

Eurasia

Cerastium nutans Raf. var. nutans, 2004 J.A. Isaac 17709

Dianthus armeria L., 2004 J.A. Isaac and R. Coxe 17583

Europe

Dianthus barbatus L., 1954 W.E. Buker s.n. Eurasia

Moehringia lateriflora (L.) Fenzl, 1993 J.K. Bissell et al.

19935151

Myosoton aquaticum (L.) Moench, 2004 J.A. Isaac and R. Coxe

17571 Europe

fal, Dat o ID L

Journal of

titute of Texas 1(1)

Paronychia canadensis (L) Wood, 1914 C.A. Darling s.n.

Saponaria officinalis L., 1994 K. Marsh et al. s.n. Europe

Silene antirrhina L., 1951 L.K. Henry and W.E. Buker s.n.

Silene dioica (L.) Clairv., 1970 R.C. Leberman s.n. Europe

Silene latifolia Poir. ssp. alba (P. Mill.) Greuter & Burdet, 1972

W.E. Buker s.n. Europe

Silene noctiflora L., J.A. Murray s.n. Europe

Silene vulgaris (Moench) Garcke, 1980 R.C. Leberman s.n.

Europe

Stellaria corei Shinners, 1932 W.R. VanDersal 1923

Stellaria graminea L., 2004 J.A. Isaac 17697 Europe

Stellaria longifolia Muhl. ex Willd. var. longifolia, 1994 J.D.

Stellaria media (L.) Vill. var. media, 1919 H.W. Mossman s.n.

Europe

Vaccaria hispanica (P. Mill.) Rauschert, 1868 T.C. Porter s.n.

Europe

Celastraceae

Celastrus scandens L., 1973 R.C. Leberman s.n.

Euonymus alata (Thunb.) Sieb., 2005 R. Thompson 05-519

China & Japan

Euonymus obovata Nutt., 2001 J.A. Isaac and C.F. Chuey

13648

Ceratophyllaceae

Ceratophyllum demersum L., 2004 S.P. Grund and L. Miller

3579

Ceratophyllum echinatum Gray, 2002 J.K. Bissell 2002:128

Ceratophyllum muricatum Cham., 1984 J.K. Bissell 1984:119

Chenopodiaceae

Atriplex prostrata Bouchér ex DC., 1887 B.H. Patterson s.n.

Chenopodium album L. var. album, 1966 L.K. Henry s.n.

Chenopodium berlandieri Moq. var. bushianum (Aellen

1947 H.A. Wahl 4073

Chenopodium botrys L., 1941 W.C. Grimm s.n. Europe

Chenopodium capitatum (L.) Ambrosi, 1909 O.E. Jennings

Sce s

Chenopodium simplex (Torr.) Raf., 1923 E.M. Gress s.n.

Clusiaceae

Hypericum ascyron L., 1998 R. Leberman s.n.

Hypericum ellipticum Hook., 1868 Thomas C. Porter s.n.

Hypericum mutilum L., 1997 J.A. Isaac 9870

Hypericum perforatum L., 2004 S.P. Grund and L. Miller 3516

Furope

Hypericum prolificum L., 2005 L. Speedy LSB1120.

Hypericum punctatum Lam., 2004 S.P. Grund and L. Miller

3626

Commelinaceae

Tradescantia virginiana L., 1914 C.A. Darling s.n.

Convolvulaceae

Calystegia sepium (L.) R. Br. ssp. angulata Brummitt, 2005 L.

Speedy, J. Bissell, M. Bowers and T. Pearson

Calystegia sepium (L.) R. Br ssp.erratica Brummitt, 1950 L.K.

Henry and W.E. Buker s.n.

Cornaceae

Cornus alternifolia L. f., 1998 R. Leberman s.n.

Morton et al., Flora of Crawford County, Pennsylvania

Cornus amomum P. Mill, 2005 L. Speedy LSB720

Cornus canadensis L., 1988 J.K. Bissell and B. Danielson

1988:035

Cornus florida L., 1988 J.K. Bissell 1988:034

Cornus obliqua Raf., 2005 L. Speedy LSB1128

Cornus racemosa Lam., 2005 L. Speedy LSB1127

Cornus sericea L. ssp. sericea, 2005 L. Speedy, M. Bowers and

M. Fodse LSB1613

Crassulaceae

Sedum acre L., 1995 R.C. Leberman s.n. Eurasia

Sedum ternatum Michx., 1932 W.R. Van Dersal 1940

Cucurbitaceae

Echinocystis lobata (Michx.) Torr. & A. Gray, 1991 R.C. Leberman

s.n.

Sicyos angulatus L., 1997 J. Long s.n.

uscutaceae

Cuscuta cephalanthi Engelm., 1988 J.K. Bissell et al. 88:232

G5:SU

Cuscuta gronovii Willd. ex J.A. Schultes, 2005 J.K. Bissell

Cuscuta obtusiflora Kunth, 1988 J.K. Bissell and B. Danielson

1988:271

Cuscuta polygonorum Engelm., 1988 J.K. Bissell et al. 88:271

G5:SU

Cyperaceae

Bulbostylis capillaris (L.) Kunth ex C.B. Clarke ssp. capillaris,

1988 J.K. Bissell et al. 88:270

Carex aestivalis M.A. Curtis ex Gray, 1907 O.E. Jennings s.n.

Carex alata Torr., 2003 J.K. Bissell 2003:082 G5:52

Carex albicans Willd. ex Spreng. var. albicans, 1930 O.E.

Jennings s.n.

Carex albursina Sheldon, 2004 J.A. Isaac 17464

Carex amphibola Steud., 2004 J.A. Isaac 17450

Carex annectens (Bickn.) Bickn., 2004 S.P. Grund and L. Miller

3498

Carex appalachica J. Webber & P.W. Ball, 2005 L. Speedy

Carex arctata Boott ex Hook., 1984 J.K. Bissell 84:103

Carex atlantica Bailey var. atlantica, 1991 J.K. Bissell and B.

Danielson 1991:071

Carex atlantica Bailey ssp. capillacea (Bailey) Reznicek, 1997

J.K. Bissell and G. Buckley 1997:102

Carex baileyi Britt., 1980 R.C. Leberman s.n.

Carex bebbii Olney ex Fern., 1988 J.K. Bissell et al. 88:076

G5:S]

Carex blanda Dewey, 2004 J.A. Isaac and R. Coxe 17572

Carex bromoides Schkuhr ex Willd., 2004 J.A. Isaac 17540

Carex brunnescens (Pers.) Poir, 1991 J.K. Bissell and B. Danielson

Carex bushii Mack., 1997 R.C. Leberman s.n.

Carex buxbaumii Wahlenb., 2004 J.A. Isaac and M. Fiely

17668 G5:53

Carex canescens L. var. canescens, 2004 J.A. Isaac 17475

Carex canescens L. var. disjuncta (Fern.) Toivonen, 1952 W.E.

Buker s.n.

Carex caroliniana Schwein., 2000 J.K. Bissell and B. Danielson

2000:038

641

Carex cephalophora Muhl. ex Willd., 1980 R.C. Leberman s.n.

Carex communis Bailey, 1991 J.K. Bissell and B. Danielson

1991:070

Carex comosa Boott, 2005 L. Speedy and L. Armstron

LSB1594

Carex crinita Lam. var. crinita, 2005 L. Speedy 742

Carex cristatella Britt., 2004 J.A. Isaac 17694

Carex cryptolepis Mack., 1993 J.K. Bissell and B. Danielson

1993:092 G4:51

Carex debilis Michx. var. pubera Gray, 1886 B.H. Patterson

S.N.

Carex debilis Michx. var. rudgei Bailey, 2004 J.A. Isaac 17441

Carex diandra Schrank, 2004 J.A. Isaac 17534 G5:S2

Carex digitalis Willd. var. digitalis, 2004 J.A. Isaac 17465

Carex disperma Dewey, 2004 J.K. Bissell 2004:193 G5:53

Carex echinata Murr. ssp. echinata, 1993 J.A. Isaac 4761

Carex flava L., 2004 J.A. Isaac 17674 G5:S2

Carex folliculata L., 2004 J.K. Bissell 2004:121

Carex gracilescens Steud., 1985 J.K. Bissell 85:55

Carex gracillima Schwein., 2005 L. Speedy LSB1121

Carex granularis Muhl. ex Willd., 1994 J.D. Wagner and J.K.

Bissell 352

Carex grayi Carey, 2004 J.A. Isaac 17532

Carex gynandra Schwein., 2004 S.P. Grund and L. Miller

Carex hirsutella Mack., 1952 L.K. Henry and W.E. Buker s.n.

Carex hystericina Muhl. ex Willd., 2004 S.P. Grund and L.

Miller 3607

Carex interior Bailey, 2004 J.A. Isaac 17672

€ j ens Rudge, 2005 L. Speedy 1132

Carex lacustris Willd., 2004 S.P. Grund and L. Miller 3556

Carex laevivaginata (Kükenth.) Mack., 1988 J.K. Bissell and B.

Danielson 1988:035

Carex lasiocarpa Ehrh. var. americana Fern., 1994 J.D. Wagner

and J.K. Bissell 392 G5:S3

Carex laxiculmis Schwein. var. laxiculmis, 2004 J.A. Isaac

Carex laxiflora Lam., 1908 O.E. Jennings s.n.

Carex leptalea Wahlenb. ssp. leptalea, 2004 J.A. Isaac 17538

Carex leptonervia (Fern.) Fern, 2004 J.A. Isaac 17434

Carex lupulina Muhl. ex Willd., 2005 L. Speedy 739

Carex lurida Wahlenb., 2005 L. Speedy 1103

Carex mitchelliana M.A. Curtis, 2004 J.A. Isaac 17600 G3G4:

Carex normalis Mack., 1922 J. Bright s.n.

Carex novae-angliae Schwein., 1992 J.K. Bissell and B.

Danielson 92:063

Carex oligocarpa Schkuhr ex Willd., 1929 J. Bright 1766

Carex ormostachya Wieg., 1985 J.K. Bissell 85:052 G4:S2

Carex pallescens L., 1997 S.P. Grund and J.A. Isaac 1873

Carex pedunculata Muhl. ex Willd., 1989 J.K. Bissell et al.

1989:021

Carex pensylvanica Lam., 2004 J.A. Isaac 17455

Carex plantaginea Lam., 2004 J.A. Isaac 17495

Carex platyphylla Carey, 1980 R.C. Leberman s.n.

Carex prairea Dewey ex Wood, 2004 J.K. Bissell 2004:085

G5?:S2

Carex prasina Wahlenb., 2001 J.A. Isaac 13638

642

Carex projecta Mack., 2004 S.P. Grund and L. Miller 3554

Carex pseudocyperus L., 2004 J.A. Isaac 17669 G5:S1

Carex radiata (Wahlenb.) Small, 2004 J.A. Isaac 17448

Carex rosea Schkuhr ex Willd., 2004 S.P. Grund and L. Miller

3553

Carex sartwellii Dewey, 1947 O.E. Jennings s.n. GAG5:SX

Carex scabrata Schwein., 2001 J.A. Isaac 13639

Carex scoparia Schkuhr ex Willd. var. scoparia, 2004 S.P. Grund

and L. Miller 3608

Carex seorsa Howe, 2004 J.A. Isaac 17471

Carex sparganioides Muhl. ex Willd., 1929 O.E. Jennings s.n.

Carex squarrosa L., 1942 C.E. Wood 2209

Carex stipata Muhl. ex Willd. var. maxima Chapman, 1906

O.E. Jennings s.n.

Carex stipata Muhl. ex Willd. var. stipata, 2004 J.A. Isaac

Carex straminea Willd. ex Schkuhr, 1991 J.K. Bissell et al.

1991:061

Carex stricta Lam., 2004 S.P. Grund and L. Miller 3555

Carex swanii (Fern.) Mack., 2005 L. Speedy 1087

Carex tenera Dewey, 1991 J.K. Bissell et al. 91:061

Carex torta Boott ex Tuck., 2004 J.A. Isaac 17426

Carex tribuloides Wahlenb., 1993 J.K. Bissell and B. Danielson

1993:090

Carex trichocarpa Muhl ex Willd., 1994 J.K. Bissell et al.

94:068

Carex trisperma Dewey, 2004 J.A. Isaac 17539

Carex tuckermanii Dewey, 2002 J.K. Bissell 2002:161

Carex typhina Michx., 1998 J.K. Bissell 1998:101 G5:S2

Carex utriculata Boott, 1993 J.K. Bissell et al. 93:093

Carex vesicaria L. var. monile (Tuck.) Fern., 1952 L.K. Henry

and F.H. Beer s.n.

Carex virescens Muhl. ex Willd., 1930 H.E. Stone s.n.

Carex vulpinoidea Michx., 2004 L. Miller and S.P. Grund 44

Cladium mariscoides (Muhl.) Torr., 1994 J.D. Wagner and L.L.

Smith 653 G5:52

Cyperus bipartitus Torr., 2004 S.P. Grund and L. Miller 3614

Cyperus erythrorhizos Muhl., 2004 J.A. Isaac and J. Wagner

18542

Cyperus esculentus L., 1901 J.A. Shafer s.n. Exotic

Cyperus houghtonii Torr., 1941 W.C. Grimm s.n. G4?:S1

Cyperus odoratus L., 2001 J.K. Bissell 2001:150

Cyperus strigosus L., 2004 S.P. Grund and L. Miller 3598

Dulichium arundinaceum (L.) Britt., 2005 L. Speedy LSB1105

Eleocharis acicularis (L.) Roemer & J.A. Schultes, 2004 S.P.

Grund and L. Miller 3499

Eleocharis compressa Sull., 1993 J.K. Bissell and B. Danielson

93:92 G4:51

Eleocharis elliptica Kunth, 1994 J.D. Wagner and J.K. Bissell

Eleocharis erythropoda Steud., 1988 J.K. Bissell et al. 88:035

Eleocharis intermedia J.A. Schultes, 1992 J.K. Bissell and B.

Danielson 92:130 G5:S2

Eleocharis obtusa (Willd.) J.A. Schultes var. obtusa, 2005 J.K.

Bissell 2005:074

Eleocharis ovata (Roth) Roemer & J.A. Schultes, 2004 J.A.

Isaac 17696

fal, Dat o ID L

Journal of

titute of Texas 1(1)

Eleocharis palustris (L) Roemer & J.A. Schultes, 1995 R.C.

Leberman s.n.

Eriophorum virginicum L., 2004 J.A. Isaac 18471

Eriophorum viridicarinatum (Engelm.) Fern., 1994 J.D. Wagner

and J.K. Bissell 358 G5:S2

Rhynchospora alba (L.) Vahl, 1994 J.D. Wagner 662

Schoenoplectus acutus (Muhl. ex Bigelow) A. & D. Lóve var.

acutus, 1988 J.K. Bissell et al. 1988:129 G5:G2

Schoenoplectus pungens (Vahl) Palla, Verh.K.K., 1974 R. Fulton

s.n.1974 R. Fulton s.n.

Schoenoplectus fluviatilis (Torr.) M.T. Strong, 1997 J.A. Isaac

Schoenoplectus tabernaemontani (K.C. Gmel.) Palla, 2004 L.

Miller and S.P. Grund 27

Scirpus atrocinctus Fern., 1984 J.K. Bissell 84:119

Scirpus atrovirens Willd., 2004 S.P. Grund and L. Miller 3670

Scirpus cyperinus (L.) Kunth, 2005 L. Speedy LSB1107

Scirpus georgianus R.M. Harper, 2004 S.P. Grund and L. Miller

Scirpus hattorianus Makino, 2004 J.A. Isaac 17695

Scirpus lineatus Michx., 1972 R.C. Leberman s.n.

Scirpus microcarpus J. & K. Presl, 1993 J.K. Bissell and S. Lor

1993:206

Scirpus pendulus Muhl., 2004 J.A. Isaac 17704

Scirpus polyphyllus Vahl, 2005 L. Speedy LSB1112

Dioscoreaceae

Dioscorea villosa L., 1904 O.E. Jennings s.n.

Droseraceae

Drosera rotundifolia L. var. rotundifolia, 1994 J.D. Wagner and

L.L. Smith 652

Elaeagnaceae

Elaeagnus umbellata Thunb., 1991 R.S. Fox s.n. Asia

Shepherdia canadensis (L.) Nutt., 1992 J.K. Bissell and B.

Danielson 92:064 G5:51

Ericaceae

Arctostaphylos uva-ursi (L.) Spreng,. 1930 W.R. Van Dersal

Chamaedaphne calyculata (L.) Moench, 1991 R.C. Leberman

Epigaea repens L., 2004 J.A. Isaac 17325

Gaultheria procumbens L., 1990 J.K. Bissell and B. Danielson

1990:197

Gaylussacia baccata (Wangenh.) K. Koch, 2004 J.A. Isaac

Lyonia ligustrina (L.) DC. var. ligustrina, 1884 FT. Aschman

s.n.

Rhododendron maximum L., 1962 L.K. Henry s.n.

Rhododendron periclymenoides (Michx.) Shinners, 1942 O.E.

Jennings s.n.

Eli 1G IA NAGI

is, 1927 O.E. Jennings

et al. s.n.

Vaccinium corymbosum L., 2005 L. Speedy LSB302

Vaccinium fuscatum Ait., 1983 R.C. Leberman s.n. [Cyanococcus

atrococcus (Gray) Small]

Vaccinium macrocarpon Ait., 1999 J.K. Bissell 1999:073

Vaccinium myrtilloides Michx., 2004 J.A. Isaac 18436

Morton et al., Flora of Crawford County, Pennsylvania

Vaccinium pallidum Ait., 2004 J.A. Isaac 17463

Vaccinium stamineum L., 2004 J.A. Isaac 17457

Eriocaulaceae

Eriocaulon aquaticum (Hill) Druce, 1933 W.R. Van Dersal s.n.

Euphorbiaceae

Acalypha rhomboidea Raf., 1970 W.E. Buker s.n.

Chamaesyce maculata (L.) Small, 1964 W.E. Buker s.n.

Chamaesyce nutans (Lag.) Small, 1964 W.E. Buker s.n.

Euphorbia corollata L., 1901 J.A. Shafer s.n.

Euphorbia cyparissias L., 1933 O.E. Jennings s.n. Eurasia

Fabaceae

Amphicarpaea bracteata (L.) Fern. var. bracteata, 1984 J.K.

Bissell 1984:120

Amphicarpaea bracteata (L.) Fern. var. comosa (L) Fern, 1939

O.E. Jennings s.n.

Apios americana Medik., 2004 S.P. Grund and L. Miller 3604

Baptisia tinctoria (L.) R. Br. ex Ait. f., 1968 R.C. Leberman s.n.

Chamaecrista fasciculata (Michx.) Greene var.fasciculata, 1994

K. Marsh et al. s.n.

Desmodium canadense (L.) DC., 1968 W.E. Buker s.n.

Desmodium glutinosum (Muhl. ex Willd.) Wood, 2004 J.A.

Isaac 17682

Desmodium marilandicum (L.) DC., 1994 K. Marsh et al. s.n.

Desmodium nudiflorum (L.) DC., 1907 J.A. Shafer s.n.

Desmodium paniculatum (L.) DC. var. paniculatum, 1973 R.C.

Leberman s.n

Desmodium perplexum Schub., 1994 K. Marsh et al. s.n.

Desmodium rotundifolium DC., 1907 J.A. Shafer s.n.

Gleditsia triacanthos L., 1909 O.E. Jennings s.n.

Lathyrus japonicus Willd. var. maritimus (L.) Kartesz & Gandhi

1914 J. Bright s.n. G5:52

Lathyrus ochroleucus Hook., 1901 J.A. Shafer s.n. G4G5:S1

Lespedeza frutescens (L.) Hornem., 1994 K. Marsh et al. s.n.

Lespedeza hirta (L.) Hornem. var. hirta, 1994 K. Marsh et al. s.n.

Lotus corniculatus L., 1980 F.Q. Jenkins and C.E. Jenkins 2820

Furope

Lupinus perennis L. ssp. perennis, 1930 W.R. VanDersal 1271

(35153

Medicago pads L., 2004 J.A. Isaac 17707 Eurasia

Medicago a L. ssp. sativa, 2004 J.A. Isaac 17691 Eurasia

Melilotus Re (L) Lam., 1922 O.E. Jennings s.n. Eurasia

Strophostyles helvola (L.) Elliott, 1933 O.E. Jennings s.n.

Trifolium aureum Pollich, 1952 L.K. Henry and W.E. Buker

Trifolium campestre Schreb., 1914 C.A. Darling s.n. Europe

Trifolium dubium Sibth., 2005 R. Thompson 05-650 Europe

Trifolium hybridum L., 1914 C.A. Darling s.n. Eurasia

Trifolium pratense L., 1938 O.E. Jennings s.n. Europe

Trifolium repens L., 1904 O.E. Jennings s.n. Europe

Vicia sativa L. ssp. nigra (L.) Ehrh., 1928 J.A. Murray s.n.

Furope

Vicia tetrasperm (L.) Schreb., 2005 R. Thompson 05-600

Eurasia

Vicia villosa Roth. ssp. villosa, 1972 W.E. Buker s.n. Europe

Fagaceae

Castanea dentata (Marsh.) Borkh., 2004 J.A. Isaac 18419

643

Fagus grandifolia Ehrh., 1997 R.C. Leberman s.n.

Quercus alba L., 1996 R.C. Leberman s.n.

Quercus bicolor Willd., 1909 O.E. Jennings s.n.

Quercus coccinea Muenchh. var. coccinea, 1963 P.B. Monk

S.N.

Quercus imbricaria Michx., 1997 R.C. Leberman s.n.

Quercus macrocarpa Michx. var. macrocarpa, 1919 O.E.

Jennings and J.C. Fettermann s.n.

Quercus palustris Muenchh., 1997 R.C. Leberman s.n.

Quercus prinus L., 1996 R.C. Leberman s.n.

Quercus rubra L. var. rubra, 2005 L. Speedy LSB1080-2

Quercus shumardii Buckl., 2006 J.K. Bissell 2006:007 G5:S1

Quercus velutina Lam., 2003 S. Ernst 28

Fumariaceae

Corydalis flavula (Raf) DC., 1932 W.R. Van Dersal 1921

Gentianaceae

Bartonia virginica (L.) B.S.P., 1996 S. Grund 1741

Centaurium pulchellum (Sw.) Druce, 2004 J.A. Isaac 18406

Europe

Gentiana clausa Raf., 1988 J.K. Bissell and B. Danielson

1990:197

Geraniaceae

Geranium maculatum L., 2005 L. Speedy LSB755

Haloragaceae

Myriophyllum sibiricum Kom., 2003 J.K. Bissell 2003:091 G5:

Myriophyllum spicatum L., 2004 L. Miller and S.P. Grund 18

Eurasia

Myriophyllum verticillatum L., 2001 J.K. Bissell 2001:148 G5:S1

Proserpinaca palustris L. var. crebra Fern. & Grisc., 1993 J.A.

Isaac 4762

Hamamelidaceae

Hamamelis virginiana L., 1997 R.C. Leberman s.n.

Hippocastanaceae

Aesculus glabra Willd. var. glabra, 1997 R.C. Leberman s.n.

Hydrocharitaceae

Elodea canadensis Michx., 2004 S.P. Grund and L. Miller

3586

Elodea nuttallii (Planch.) St. John, 1923 O.E. Jennings s.n.

Vallisneria americana Michx., 2004 S.P. Grund and L. Miller

3587

Hydrophyllaceae

Hydrophyllum virginianum L., 1960 L.K. Henry s.n.

Iridaceae

Iris pseudacorus L., 2004 J.A. Isaac and J. Wagner 18543

Europe

Iris versicolor L., 2004 L. Miller and S.P. Grund 39

Sisyrinchium angustifolium P. Mill., 2004 J.A. Isaac 17483

Juglandaceae

g cordiformis (Wangenh.) K. Koch, 1997 R.C. Leberman

Ts Mom (P. Mill.) Sweet, 1996 R.C. Leberman s.n.

Carya laciniosa (Michx. f£) G. Don, 1997 J.K. Bissell et al.

1997-183 G5:53

644

Carya ovalis (Wangenh.) Sarg., 1915 O.E. Jennings and G.K.

Jennings s.n.

Carya ovata (P. Mill.) K. Koch, 1907 O.E. Jennings s.n.

Carya x laneyi Sarg. [cordiformis x ovata], 1938 O.E. Jennings

s.n.

Juglans cinerea L., 1924 O.E. Jennings s.n.

Juglans nigra L., 1963 P. Monk s.n.

Juncaceae

Juncus acuminatus Michx., 1997 J.K. Bissell and B. Gordon

1997:155

Juncus articulatus L., 2004 L. Miller and S.P. Grund 47

Juncus bufonius L. var. bufonius, 2004 J.A. Isaac 17710

Juncus canadensis J. Gay ex Laharpe, 2004 J.A. Isaac 18470

Juncus dudleyi Wieg., 1997 J.A. Isaac 9859

Juncus effusus L. var. pylaei (Laharpe) Fern. & Wieg, 2004 S.P.

Grund and L. Miller 3495

Juncus effusus L. var. solutus Fern. & Wieg., 2005 L. Speedy

LSB31106

Juncus marginatus Rostk., 1973 R.C. Leberman s.n.

Juncus nodosus L., 1996 S. Grund et al. 1750

Juncus subcaudatus (Engelm.) Coville & Blake var. subcaudatus

1932 W.R. Van Dersal 51

Juncus tenuis Willd., 2004 L. Miller and S.P. Grund 48

Luzula acuminata Raf. var. acuminata, 2005 L. Speedy

LSB344

Luzula echinata (Small) F.J. Herm., 1981 R.C. Leberman s.n.

Luzula multiflora (Ehrh.) Lej. ssp. multiflora, 2005 L. Speedy

LSB301

Lamiaceae

Agastache scrophulariifolia (Willd. Kuntze, 1947 H.A. Wahl

4082A

Blephilia ciliata (L.) Benth., 1909 B.H. Patterson s.n.

Clinopodium vulgare L., 2005 L. Speedy LSB757

Collinsonia canadensis L., 1904 O.E. Jennings s.n.

Galeopsis tetrahit L. var. tetrahit, 2004 J.A. Isaac 18420

Eurasia

Glechoma hederacea L., 1932 R. Van Dersal 1922 Eurasia

Hedeoma pulegioides (L.) Pers., 1955 D.L. Pearth s.n.

Lamium amplexicaule L., 1887 B.H. Patterson s.n. Eurasia

Leonurus cardiaca L. ssp. cardiaca, 1973 R.C. Leberman s.n.

Asia

Lycopus americanus Muhl. ex W. Bart., 2004 S.P. Grund and

L. Miller 3589

Lycopus uniflorus Michx. var. uniflorus, 2005 L. Speedy and L.

Armstrong LSB1099

Lycopus virginicus L., 1904 O.E. Jennings s.n.

Mentha arvensis L., 1984 J.K. Bissell 1984:105

Mentha spicata L., 1901 J.A. Shafer s.n. Europe

Mentha x piperita L. (pro sp.) [aquatica x spicata], 1984 J.K.

Bissell 84:119 Eurasia

Mentha x villosa Huds. (pro sp.) [spicata x suaveolens], 2005

R. Thompson 05-869 Eurasia

Monarda clinopodia L., 1998 R. Leberman s.n.

Monarda didyma L., 1966 L.K. Henry and W.E. Buker s.n.

Monarda fistulosa L. var. fistulosa, 1994 K. Marsh et al. s.n.

Monarda fistulosa L. var. mollis (L.) Benth., 1975 R.C. Leberman

s.n.

fal, Dat o ID L

Journal of

titute of Texas 1(1)

Monarda media Willd., 1996 J.K. Bissell et al. 1996:091

Nepeta cataria L., 1901 J.A. Shafer s.n. Europe

Prunella vulgaris L. ssp. lanceolata (W. Bart.) Hultén, 2004 S.P.

Grund and L. Miller 3515

Pycnanthemum incanum (L.) Michx. var. incanum, 1901 J.A.

Shafer s.n.

Scutellaria galericulata L., 2005 L. Speedy LSB1102

Scutellaria lateriflora L. var. lateriflora, 2004 J.A. Isaac 18415

Stachys tenuifolia Willd., 1973 R.C. Leberman s.n.

Teucrium canadense L. var. canadense, 1998 R. Leberman s.n.

Teucrium canadense L. var. occidentale (Gray) McClintock &

Epling, 1932 W.R. Van Dersal 1806

Trichostema dichotomum L., 1901 J.A. Shafer s.n.

Lauraceae

Lindera benzoin (L.) Blume var. benzoin, 2005 L. Speedy

LSB334

Sassafras albidum (Nutt.) Nees, 1997 R.C. Leberman s.n.

Lemnaceae

Lemna minor L., 2005 L. Speedy LSB731

Lemna trisulca L., 1997 J.K. Bissell and B.W. Danielson

1997:067

Lemna turionifera Landolt, 1951 O.E. Jennings s.n. G5:SU

Spirodela polyrhiza (L.) Schleid., 2004 L. Miller and S.P. Grund

Wolffia borealis (Engelm. ex Hegelm.) Landolt ex Landolt &

Wildi, 1951 O.E. Jennings s.n.

Wolffia brasiliensis Wedd., 2004 J.A. Isaac and J. Wagner

Wolffia columbiana Karst., 1986 J.K. Bissell et al. 1986:285

Wolffiella gladiata (Hegelm.) Hegelm., 1997 J.K. Bissell and B.

Danielson 1997:067 G5:52

Lentibulariaceae

Utricularia gibba L., 1988 J.K. Bissell and B. Danielson

Utricularia intermedia Hayne, 2004 J.A. Isaac 17675 G5:52

Utricularia macrorhiza Le Conte, 2004 S.P. Grund and L.

Miller 3609

Utricularia minor L., 2004 J.K. Bissell and B. Danielson

Liliaceae

Allium canadense L. var. canadensis, 1950 L.K. Henry and W.E.

Buker s.n.

Allium tricoccum Ait., 2004 J.A. Isaac 17683

Asparagus officinalis L., 1957 W.E. Buker s.n. Europe

Chamaelirium luteum (L.) Gray, 1900 D. Bard s.n.

Clintonia borealis (Ait.) Raf., 2004 J.A. Isaac 17421

Clintonia umbellulata (Michx.) Morong, 1981 R.C. Leberman

Erythronium albidum Nutt., 2001 J.K. Bissell 2001:006 G5:S3

Erythronium americanum Ker-Gawl ssp. americanum, 2004

J.A. Isaac 17315

Hemerocallis fulva (L.) L., 1914 C.A. Darling s.n. Asia

Hyacinthoides nonscripta (L.) Chouard ex Rothm., 1998 G.

Shaffer s.n. Exotic

Hypoxis hirsuta (L.) Coville, 1889 J. Whitesides s.n.

Leucojum aestivum L. ssp. aestivum, 1998 C. Stelter s.n.

Exotic

Morton et al., Flora of Crawford County, Pennsylvania

Lilium canadense L. ssp. editorum (Fern) Wherry, 1997 R.C.

Leberman s.n.

Lilium philadelphicum L. var. philadelphicum, 1882 J.

Whitesides s.n.

Lilium superbum L., 1998 R. Leberman s.n.

Maianthemum canadense Desf., 2005 L. Speedy LSB1086

Maianthemum racemosum (L.) Link spp. racemosum, 1997

R.C. Leberman s.n.

Maianthemum stellatum (L.) Link, 1919 O.E. Jennings s.n.

Maianthemum trifolium (L.) Sloboda, 1937 O.E. Jennings s.n.

Medeola virgit vana L., 1981 R.C. Leberman s.n.

Polygonatum biflorum (Walt, Ell. var. commutatum (J.A. & J.H ),

1997 R.C. Leberman s.n.

Pol pubescens (Willd.) Pursh, 2004 J.A. Isaac 17435

Prosartes lanuginosa (Michx.) D. Don, 1981 R.C. Leberman s.n

[Disporum lanuginosa (Michx.) Nichols.]

Stenanthium gramineum (Ker-Gawl.) Morong var. gramineum

ca, 1890 J.E. Whiteside s.n. G4G5:5152

Streptopus lanceolatus (Ait.) Reveal var. roseus (Michx.) Reveal,

1985 J.K. Bissell and M. Hoberecht 1985:048

Trillium erectum L., 2004 J.A. Isaac 17335

Trillium flexipes Raf., 1941 O.E. Jennings s.n. G5:52

Trillium grandiflorum (Michx.) Salisb., 2004 J.A. Isaac 17339

Trillium undulatum Willd., 1971 R.C. Leberman s.n.

Uvularia grandiflora Sm., 1974 G. Williamson 94

Uvularia perfoliata L., 1919 O.E. Jennings s.n.

Uvularia sessilifolia L., 1981 R.C. Leberman s.n.

Veratrum viride Ait., 2005 L. Speedy LSB752

Limnanthaceae

Floerkea proserpinacoides Willd., 2005 L. Speedy LSB347

Linaceae

Linum striatum Walt., 1992 J.K. Bissell et al. 92:170

Linum usitatissimum L., 1901 J.A. Shafer s.n. Europe

Lythraceae

Decodon verticillatus (L.) Elliott, 2005 L. Speedy LSB1109

Lythrum salicaria L., 2005 L. Speedy LSB727 Europe

Magnoliaceae

Liriodendron tulipifera L., 1917 C.A. Darling s.n.

Magnolia acuminata (L.) L., 1997 R.C. Leberman s.n.

Malvaceae

Abutilon theophrasti Medik., 1994 K. Marsh and G. Marsh

s.n. Asia

Malva moschata L., 2004 J.A. Isaac et al. 17723 Europe

Malva neglecta Wallr., 1948 D.H. Krouse s.n. Eurasia & N.

Africa

Menispermaceae

Menispermum canadense L., 1914 C.A. Darling s.n.

Menyanthaceae

Menyanthes trifoliata L., 1994 J.D. Wagner and J.K. Bissell 393

Moraceae

Morus alba L., 1997 R.C. Leberman s.n. Asia

Myricaceae

Morella pensylvanica (Mirb.) Kartesz, 1994 J.D. Wagner and

J.K. Bissell 404

645

Najadaceae

Najas flexilis (Willd.) Rostk. & Schmidt, 2004 S.P. Grund and

L. Miller 3576

Najas gracillima (A. Braun ex Engelm.) Magnus, 2004 S.P.

Grund and L. Miller 3581

Najas guadalupensis (Spreng.) Magnus ssp. guadalupensis,

2004 S.P. Grund and L. Miller 3584

Najas minor All., 2004 S.P. Grund and L. Miller 3637 Exotic

Nelumbonaceae

Nelumbo lutea Willd., 1992 J.K. Bissell 1992:109

Nymphaeaceae

Nuphar lutea (L.) Sm ssp. advena (Ait.) Kartesz & Gandhi, 2003

>, Ernst 116

Nuphar lutea (L.) Sm ssp. variegata (Dur.) E.O. Beal, 2004 J.K.

Bissell 2004111

Nymphaea odorata Ait. ssp. odorata, 2004 L. Miller and S.P.

Grund 26

Nymphaea odorata Ait. ssp. tuberosus (Paine) Wiersma &

Hellquist, 1923 O.E. Jennings s.n.

Nyssaceae

Nyssa sylvatica Marsh., 2004 J.A. Isaac 17545

Oleaceae

Fraxinus americana L., 2004 S.P. Grund and L. Miller 3540

Fraxinus nigra Marsh., 2004 S.P. Grund and L. Miller 3535

Fraxinus pennsylvanica Marsh., 2001 J.A. Isaac 13668

Fraxinus profunda (Bush) Bush, 2003 R. Coxe and M. Bradburn

Ligustrum obtusifolium Sieb. & Zucc., 1998 K. Marsh s.n.

Japan

Onagraceae

Chamerion angustifolium (L.) Holub ssp. cirrumvagum

(Mosquin) Kartesz, 1967 W.E. Buker s.n.

Circaea alpina L. ssp. alpina, 1972 R.C. Leberman s.n.

Circaea lutetiana L. ssp. canadensis (L.) Asch. & Magnus, 2004

SP Grund.3552

Epilobium ciliatum Raf. ssp. ciliatum, 1928 J.A. Murray s.n.

Fpilobium coloratum Biehler, 2005 L. Speedy and L. Armstrong

LSB1593

Epilobium hirsutum L., 1994 K. Marsh and G. Marsh s.n.

Europe

Epilobium leptophyllum Raf., 1994 J.D. Wagner 678

Epilobium strictum Muhl. ex Spreng., 2004 J.A. Isaac 18426

525593

Ludwigia palustris (L.) Elliott., 2005 J.K. Bissell 2005:074

Oet othera biet H WS L., 1964 W.E. Bu ker Sans

Oenothera fruticosa L. ssp. glauca (Michx) Straley, J.A. Murray

Oenothera nutans Atk. & Bartlett, Curtis 39

Oenothera perennis L., 1981 R.C. Leberman s.n.

Orchidaceae

Arethusa bulbosa L., 1905 O.E. Jennings s.n. G4:S1

Calopogon tuberosus (L.) B.S.P. var. tuberosus, 1994 J.D. Wagner

and J.K. Bissell 390

Corallorhiza maculata (Raf.) Raf. var. maculata, 1998 R.

Leberman s.n.

646

Corallorhiza maculata (Raf) Raf. var. occidentalis (Lindl.) Ames,

Esse

Corallorhiza trifida Chatel., 2004 J.A. Isaac 17417

Cyp ile Ait., 1962 L.K. Henry s.n.

EE Kn dd Salisb. var. pubescens (Willd.) Knight,

1969 W.E. Buker and W.C. Buker s.n. G5:S1

Cypripedium reginae Walt., 1962 L.K. Henry s.n. G4:S2

Epipactis helleborine (L.) Crantz, 1998 R. Leberman s.n.

Furope

Galearis spectabilis (L.) Raf., 1949 L.K. Henry s.n.

Goodyera pubescens (Willd.) R. Br. ex Ait. f, 2004 J.A. Isaac

17443

Isotria verticillata Raf., 1997 R.C. Leberman s.n.

Liparis loeselii (L.) L.C. Rich., 1969 W.E. Buker s.n.

Malaxis brachypoda (Gray) Fern., 1968 W.E. Buker s.n. G4O:S1

Platanthera clavellata (Michx.) Luer., 2005 L. Speedy

LSB1085

Platanthera dilatata (Pursh) Lindl. ex Beck var. dilatata, 1994

J.S. Shriver and C. Smith 216 G5:S1

Platanthera flava (L.) Lindl. var. herbiola (R. Br. ex Ait. f.) Luer,,

1975 R.C. Leberman s.n.

Platanthera grandiflora (Bigelow) Lindl.,

M huronensis (Nutt) Lindl.,

Stull- sri.

TORUM hyperborea (L.) Lindl. var. hyperborean, 1997 J.A.

Isaac 9881 G5:51

Platanthera lacera (Michx.) G. Don, 1980 R.C. Leberman s.n.

Platanthera leucophaea (Nutt.) Lindl., 1881 C.W. C. s.n. G3:SX

Platanthera macrophylla (Goldie) P.M. Brown, 1962 R.C.

vnrinerluiim

1922 J. Bright s.n.

1965 J. Stull and D.

Leberman s.n.

Platanthera orbiculata (Pursh) Lindl., 1982 R.C. Leberman

STI:

Platanthera psycodes (L) Lindl., 1962 L.K. Henry s.n.

Pogonia ophioglossoides (L.) Ker-Gawl., 1994 J.D. Wagner and

J.K. Bissell 391

Spiranthes cernua (L.) L.C. Rich., 2002 J.K. Bissell 2002:176

Spiranthes lucida (H.H. Eat) Ames, 1982 W.E. Buker s.n. G5:S3

Spiranthes ochroleuca (Rydb.) Rydb., 1970 R.C. Leberman

S.N.

Spiranthes romanzoffiana Cham., 1995 J.S. Shriver and C.

Smith 423 G5:S1

Orobanchaceae

Conopholis americana (L.) Wallr. F., 2004 J.A. Isaac 17456

Epifagus virginiana (L.) W. Bart., 2005 L. Speedy, M. Bowers

and M. Fodse LSB1605

Oxalidaceae

Oxalis dillenii Jacq., 1904 O.E. Jennings s.n.

Oxalis montana Raf., 1985 J.K. Bissell and M. Hoberecht

Oxalis stricta L., 1905 O.E. Jennings s.n.

Papaveraceae

Chelidonium majus L. var. majus, 2004 J.A. Isaac 17355

Furope

Phrymaceae

Phryma leptostachya L., 1901 J.A. Shafer s.n.

Phytolaccaceae

fal, Dat A ID hi Pr

Journal of

of Texas 1(1)

Phytolacca americana L. var. americana, 1969 W.E. Buker s.n.

Plantaginaceae

Plantago aristata Michx., 1904 O.E. Jennings s.n.

Plantago lanceolata L., 2004 J.A. Isaac 17711 Europe

Plantago major L., 1901 J.A. Shafer s.n.

Plantago rugelii Dcne., 1901 J.A. Shafer s.n.

Platanaceae

Platanus occidentalis L., 1928 J.A. Murray s.n.

Poaceae

Agrostis gigantea Roth, 2004 J.A. Isaac 17703 Europe

Agrostis perennans (Walt) Tuck., 1907 O.E. Jennings s.n.

Agrostis scabra Willd., 2003 J.K. Bissell 1999:073

Agrostis stolonifera L., 2004 J.A. Isaac et al. 17

Alopecurus aequalis Sobol. var. aequalis, 2001 J.A. Isaac and

C.F. Chuey 13659 G5:53

Andropogon gerardii Vitman, 2002 J.A. Isaac 15714

Andropogon virginicus L. var. virginicus, 2002 J.A. Isaac 15717

Anthoxanthum odoratum L. spp. odoratum, 2004 J.A. Isaac

17350 Eurasia

Arrhenatherum elatius (L.) P. Beauv. ex J. & C. Presl var. elatius,

1952 W.E. Buker s.n. Europe

Bouteloua curtipendula (Michx.) Torr. var. curtipendula, 2002

J.A. Isaac 15715 G5:52

Bouteloua gracilis (Willd. ex Kunth) Lag. ex Griffiths, 2002

J.A. Isaac 15723

Brachyelytrum erectum (Schreb. ex Spreng.) Beauv., 1984 J.K.

Bissell 1984:103

Bromus ciliatus L. var. ciliatus, 1992 J.K. Bissell et al. 1992:102

Bromus inermis Leyss. ssp. inermis, 2004 J.A. Isaac 17715

Furope

Bromus japonicus Thunb. ex Murr., 2004 J.A. Isaac and R. Coxe

asia

Bromus latiglumis (Shear) A.S. Hitchc., 1994 K. Marsh et al.

s.n.

Bromus racemosus L., 2004 J.A. Isaac 17692 Europe

Bromus secalinus L., 1968 W.E. Buker s.n. Europe

Bromus tectorum L., 1922 O.E. Jennings s.n. Europe

Cinna arundinacea L., 2005 L. Speedy LSB1110

Cinna latifolia (Trev. ex Goepp.) Griseb., 1998 J.K. Bissell

1998:104

Dactylis glomerata L. ssp. glomerata, 1980 R.C. Leberman

s.n. Europe

Danthonia compressa Austin ex Peck, 1984 J.K. Bissell

1984:103

Danthonia spicata (L.) Beauv. ex Roem. €: J.A. Schultes, 1882

Jj uu Siti:

Dicl acuminatum (Sw.) Gould & C.A. Clark var. fas-

ciculatum (Torr.) Freckm ann, 2005 J.K. Bissell 2005:074

Dichanthelium acuminatum (Sw.) Gould & C.A. Clark var.

lindheimeri (Nash) Gould & C.A. Clark, 1933 O.E. Jennings

Dichanthelium clandestinum (L.) Gould, 1981 R.C. Leberman

Dichanthelium dichotomum (L.) Gould var. dichotomun, 1901

J.A. Shafer s.n.

Dichanthelium latifolium (L) Gould & C.A. Clark, 1993 J.K.

Bissell et al. 1993:065

Morton et al., Flora of Crawford County, Pennsylvania

Dichanthelium linearifolium (Scribn.ex Nash) Gould, 1922 J.

Bright s.n.

Dichanthelium sabulorum (Lam.) Gould & C.A. Clark var.

thinium (A.S. Hitchc. & Chase) Gould & C.A. Clark, 2002

J.A. Isaac 15712

Dichanthelium spl pon (Elliott) Gould var. sphaerocar-

pon, 1922 J. Bright s.n.

Digitaria ischaemum (Schreb.) Schreb. ex Muhl., 2002 J.A.

Isaac 15718 Eurasia

Digitaria sanguinalis (L.) Scop., 1881 J. Whitesides s.n.

Echinochloa crus-galli (L.) Beauv., 1984 DW. Vogler and CW.

Bier 6-4 Eurasia

Echinochloa muricata (Beauv.) Fern. var. muricata, 2005 J.K.

Bissell 2005:074 Exotic

Eleusine indica (L.) Gaertn.,

Tropics

Elymus hystrix L. var. hystrix, 2005 L. Speedy LSB754

Elymus repens (L.) Gould, 2004 J.A. Isaac 17693 Exotic

Elymus riparius Wieg., 1994 K. Marsh et al. s.n.

Elymus villosus Muhl. ex Willd., 1882 J. Whitesides s.n.

Elymus virginicus L. var. virginicus, 1997 J.A. Isaac 9876

Eragrostis cilianensis (All.) Vign. ex Janchen, 1964 W.E. Buker

s.n. Europe

Eragrostis hypnoides (Lam.) B.S.P., 2005 J.K. Bissell and B.

Danielson 2005:074

Eragrostis pectinacea (Michx.) Nees ex Steud. var. pectinacea,

1952 H.A. Wahl 13813

Eragrostis spectabilis (Pursh) Steud., 1994 K. Marsh et al. s.n.

Festuca subverticillata (Pers.) Alexeev, 1922 J. Bright s.n.

Glyceria acutiflora Torr., 1992 J.K. Bissell et al. 92:108

Glyceria canadensis (Michx.) Trin., 2005 L. Speedy and L.

Armstrong LSB1599

Glyceria grandis S. Watson var. grandis, 1932 W.R. Van Dersal s.n.

Glyceria laxa (Scribn.) Scribn., 1909 B.H. Patterson s.n.

Glyceria melicaria (Michx.) F.T. Hubb., 2005 L. Speedy

LSB1116

Glyceria septentrionalis A.S. Hitchc., 2004 S.P. Grund and L.

Miller 3520

Glyceria striata (Lam.) A.S. Hitchc., 2005 L. Speedy LSB1098

Holcus lanatus L., 2004 J.A. Isaac 17714 Europe

Hordeum vulgare L., 1916 C.A. Darling s.n. Eurasia

Leersia oryzoides (L.) Sw., 2004 S.P. Grund and L. Miller 3669

Leersia virginica Willd., 2004 J.A. Isaac 18408

Lolium perenne L. spp. multiflorum (Lam.) Husnot, 2004 J.A.

Isaac 17605 Europe

Lolium perenne L. ssp. perenne, 1949 Edgar T. Wherry s.n.

Furope

Lolium pratense (Huds.) S.J. Darbysh., 1994 J.D. Wagner and

J.K. Bissell 363 Europe

Milium effusum L. var. cisatlanticum Fern., 2004 J.A. Isaac

1882 J. Whitesides s.n. Old World

17544

Muhlenbergia frondosa (Poir) Fern, 2002 J.A. Isaac 15726

Muhlenbergia glomerata (Willd.) Trin., 1994 J.D. Wagner 661

Muhlenbergia mexicana (L.) Trin., 1994 J.D. Wagner 695

Muhlenbergia schreberi J.F. Gmel., 1946 W.E. Buker s.n.

Muhlenbergia sobolifera (Muhl. ex Willd.) Trin., 1887 J.

Whitesides s.n.

Muhlenbergia sylvatica Torr. ex Gray, 1997 J.A. Isaac 9856

647

Oryzopsis asperifolia Michx., 1980 AW. Cusick and B. Andreas

Panicum capillare L., 1964 W.E. Buker s.n.

Panicum dichotomiflorum Michx. var. dichotomiflorum, 2002

J.A. Isaac 15725

Panicum gattingeri Nash, 1998 J.K. Bissell 1998:159

Panicum rigidulum Bosc ex Nees var. elongatum (Pursh)

Lelong, 1988 J.K. Bissell and B. Danielson 1988:273

Panicum virgatum L. var. virgatum, 2002 J.A. Isaac 15722

Pennisetum glaucum (L.) R. Br., 2002 J.A. Isaac 15711 Exotic

Phalaris arundinacea L., 2005 L. Speedy LSB737 v

Phleum pratense L., 2004 J.A. Isaac 17604 Europe

Phragmites australis (Cav.) Trin. ex Steud. ssp. americana, 2005

L. Speedy et al. 1579 s.n.

Poa alsodes Gray, 2004 J.A. Isaac 17449

Poa annua L., 2004 J.A. Isaac 17452 Eurasia

Poa compressa L., 2004 J.A. Isaac 17708 Europe

Poa paludigena Fern. & Wieg., 2002 J.K. Bissell 2002:061 G3:

Poa palustris L., 1952 L.K. Henry and W.E. Buker s.n.

Poa pratensis L., 1952 L.K. Henry and W.E. Buker s.n. Europe

Poa saltuensis Fern. & Wieg., 1994 J.K. Bissell et al. 94:106

Poa trivialis L., 1991 J.K. Bissell et al. 1991:059 Europe

Schizachne purpurascens (Torr.) Swallen, 1925 S.S. Dickey s.n.

Setaria faberi Herrm., 1997 J.A. Isaac 9851 Asia

Setaria parviflora (Poir.) Kerguélen, 1994 K. Marsh et al. s.n.

Setaria pumila (Poir.) Roemer & J.A. Schultes, 1949 Edgar T.

Wherry s.n. Europe

Sorghastrum nutans (L.) Nash, 2002 J.A. Isaac 15716

Spartina pectinata Bosc ex Link, 1933 O.E. Jennings s.n.

Sphenopholis intermedia (Rydb.) Rydb., 1993 J.K. Bissell et

al. 93:102

Sphenopholis ay (L) A.S. Hitchc., 1994 J.D. Wagner

d J.K. Bissell 3

Sporobolus Ha (Poir) Merr. var. compositus, 1996 S.

Grund 1747

Sporobolus cryptanarus (Torr.) Gray, 2002 J.A. Isaac 15724

Sporobolus vagit iflorus (Torr. ex Gray) Wood var. vaginiflorus,

1949 O.E. Jennings and C.M. Boardman s.n.

Torreyochloa pallida (Torr. Church var. pallida, 2001 J.A.

Isaac 13661

Tridens flavus (L.) A.S. Hitchc. var. flavus, 1994 K. Marsh et

Triticum aestivum L., 1916 C.A. Darling s.n. Eurasia

Podostemaceae

Podostemum ceratophyllum Michx., 1997 J.K. Bissell

Polemoniaceae

Collomia linearis Nutt., 1914 J. Bright s.n.

Phlox divaricata L. ssp. divaricata, 1971 W.E. Buker s.n.

Phlox maculata L. ssp. maculata, 1935 A.G. Dickey s.n.

Phlox paniculata L., 2004 S.P. Grund and L. Miller 3618

Phlox stolonifera Sims, 2004 J.A. Isaac 17356

Polemonium reptans L. var. reptans, 2005 L. Speedy LSB759

Polygalaceae

Polygala polygama Walt., 2006 J.K. Ms 2006:190 G5:S1

Polygala sanguinea L., 1970 R.C. Leberm

Polygala verticillata L. var. verticillata, oe a Shafer s.n.

648

Polygonaceae

Fagopyrum esculentum Moench, 1993 J.A. Isaac 4896 Asia

Polygonum amphibium L. var. emersum Michx., 2004 J.A.

Isaac 18427

Polygonum amphibium L. var. stipulaceum Coleman, 1999

J.K. Bissell 1999:073

Polygonum arifolium L., 2005 L. Speedy LSB1113

Polygonum aviculare L., 2004 J.A. Isaac 17717 Europe

Polygonum caespitosum Blume, 1997 J. Long s.n. Asia

Polygonum cilinode Michx., 1993 J.K. Bissell and B. Danielson

Polygonum convolvulus L. var. convolvulus, 1914 C.A. Darling

s.n. Europe

Polygonum cuspidatum Sieb. & Zucc.,

Japan

Polygonum hydropiper L., 2004 J.A. Isaac 18422 Europe

Polygonum hydropiperoides Michx., 2004 J.A. Isaac and J.

Wagner 18540

Polygonum lapathifolium L., 1988 J.K. Bissell and B. Danielson

¡Oo uo IER

Polygonum pensylvanicum L., 2005 J.K. Bissell 2005:074

Polygonum persicaria L., 2004 J.A. Isaac 17712 Europe

Polygonum punctatum Elliott var. confertiflorum (Meisn.)

Fassett, 1904 O.E. Jennings s.n.

Polygonum punctatum Elliott var. punctatum, 1996 S. Grund

1748

Polygonum robustius (Small) Fern., 2003 J.K. Bissell and J.

Bish 2003:125

Polygonum sagittatum L., 2005 L. Speedy LSB1101

Polygonum scandens L., 1994 K. Marsh and G. Marsh s.n.

Exotic

Polygonum setaceum Baldwin, 2005 L. Speedy et al. 1580

G5T4:52

Polygonum virginianum L., 2005 L. Speedy LSB1136

Rumex acetosella L., 2004 J.A. Isaac 17444 Europe

Rumex altissimus Wood, 1930 H.E. Stone s.n.

Rumex crispus L. ssp. crispus, 1952 L.K. Henry and F.H. Beer

s.n. Europe

Rumex obtusifolius L., 2004 S.P. Grund and L. Miller 3518

Furope

Rumex orbiculatus Gray var. orbiculatus, 1988 J.K. Bissell et

al. 1988:135

Rumex verticillatus L., 1995 R.C. Leberman s.n.

Pontederiaceae

Heteranthera dubia Jacq.) MacM., 2003 J.K. Bissell 2003:074

Pontederia cordata L., 2004 S.P. Grund and L. Miller 3592

Claytonia c a Michx. var. caroliniana, 2005 L. Speedy

LSB332. 2005

Claytonia virginica L. var. virginica, 2004 J.A. Isaac 17312

Portulaca oleracea L., 1949 Edgar T. Wherry s.n.

Potamogetonace

Potamogeton a Tuck., 2004 S.P. Grund and L. Miller

3509

Potamogeton crispus L., 2004 J.A. Isaac and J. Wagner 18538

Europe

Potamogeton diversifolius Raf., 1994 J.K. Bissell et al. 1994:173

£sthka Dat M ID hi rrt

Journal of

of Texas 1(1)

Potamogeton epihydrus Raf., 2004 J.A. Isaac and J. Wagner

18545

Potamogeton foliosus Raf., 2004 S.P. Grund and L. Miller

3583

Potamogeton friesii Rupr., 2003 S. Ernst et al. 122 G4:S1

Potamogeton gramineus L., 1933 O.E. Jennings s.n. G5:SH

Potamogeton hillii Morong, 1865 Thomas C. Porter s.n. G3:

Potamogeton illinoensis Morong, 2003 S. Ernst and L. Speedy

Potamogeton natans L., 2004 J.A. Isaac and J. Wagner 18539

P g nodosus Poir., 2002 J.K. Bissell and B. Danielson

2002:154

perfoliatus L., 2003 S. Ernst et al. 79

Potamogeton praelongus Wulfen, 2003 S. Ernst et al. 78 G5:

P pulcher Tuck., 1909 O.E. Jennings s.n. G5:S1

pusillus L. ssp. pusillus, 1907 J.A. Shafer s.n.

Potamogeton pusillus L. ssp issi (Mert. &Koch) Haynes

C.B. Hellq., 2004 S.P. Grund and L. Miller 3577

Potamogeton richardsonii (Benn.) Rydb., 2003 S. Ernst et al.

Potamogeton robbinsii Oakes, 2004 S.P. Grund and L. Miller

3591

Potamogeton spirillus Tuck., 2004 S.P. Grund and L. Miller

3580

Potamogeton vaseyi J.W. Robbins, 2004 L. Miller and S.P.

Grund 19 G4:51

Potamogeton zosteriformis Fern., 2003 S. Ernst et al. 142 G5:

Stuckenia pectinata (L.) Boerner, 2002 J.K. Bissell and B.

Danielson 2002:154

Primulaceae

Lysimachia ciliata L., 2005 L. Speedy LSB733

Lysimachia nummularia L., 2005 L. Speedy LSB724

Lysimachia quadrifolia L., 1972 R.C. Leberman s.n.

eode terrestris (L.) B.S.P., 2005 L. Speedy, M. Bowers and

e LSB1625

Lysimachia e L., 1992 J.K. Bissell et al. 1992:064

Lysimachia vulgaris L., 2004 J.A. Isaac et al. 17667 Eurasia

Samolus valerandi L. ssp. parviflorus (Raf) Hultén, 2003 R. Coxe

and M. Bradburn s.n.

Trientalis borealis Raf. sop. borealis, 1985 J.K. Bissell 1985:048

Pyrolaceae

Chimaphila maculata (L.) Pursh, 1998 R. Leberman s.n.

Chimaphila umbellata (L.) W. Bart. ssp. cisatlantica (Blake)

Hultén, 1901 J.A. Shafer s.n.

Monotropa hypopithys L., 2004 J.A. Isaac 18416

Monotropa uniflora L., 2005 L. Speedy LSB1094

Pyrola americana Sweet, 1995 R.C. Leberman s.n.

Pyrola chlorantha Sw., 1890 J.S. Ogden s.n. G5:51

Pyrola elliptica Nutt., 2004 J.A. Isaac 17614

Ranunculaceae

Actaea pachypoda Elliott, 1982 R.C. Leberman s.n.

Actaea racemosa L. var. racemosa, 1909 O.E. Jennings s.n.

Actaea rubra (Ait.) Willd., 1970 R.C. Leberman s.n.

Morton et al., Flora of Crawford County, Pennsylvania

Anemone canadensis L., 1990 J.K. Bissell and B. Danielson

1990:192

Anemone quinquefolia L. var. quinquefolia, 2005 L. Speedy

LSB342

Anemone virginiana L. var. virginiana, 1997 R.C. Leberman

STE

quilegia canadensis L., 1972 W.E. Buker s.n.

Caltha palustris L. var. palustris, 2005 L. Speedy LSB338

Clematis virginiana L., 2005 L. Speedy LSB1125

Coptis trifolia (L.) Salisb., 2004 J.A. Isaac 17331

Hepatica nobilis Schreb. var. acuta (Pursh) Steyermark, 2004

J.A. Isaac 17318

Hydrastis canadensis L., 1883 J. Whitesides s.n.

Ranunculus abortivus L., 2005 L. Speedy LSB333

Ranunculus acris L., 2005 L. Speedy LSB762 Europe

Ranunculus ambigens S. Watson, 1918 E.M. Gress Field No.

Ranunculus aquatilis L., 2001 J.K. Bissell 20011146 G5T5:S3

Ranunculus flabellaris Raf., 1909 O.E. Jennings s.n. G5:S2

Ranunculus hispidus Michx. var. hispidus, 1971 W.E. Buker s.n.

Ranunculus hispidus Michx. var. nitidus (Chapman) T. Duncan,

1950 L.K. Henry and W.E. Buker s.n.

Ranunculus longirostris Godr., 1988 J.K.Bissell et al. 1988:134

Ranunculus pensylvanicus L. f., 1984 J.K. Bissell 1984:105

Ranunculus recurvatus Poir. var. recurvatus, 1960 L.K. Henry

S1.

Ranunculus repens L., 2004 J.A. Isaac 17348 Europe

Thalictrum dioicum L., 1972 W.E. Buker s.n.

Thalictrum pubescens Pursh, 2004 S.P. Grund and L. Miller

3483

Thalictrum revolutum DC., 1885 none s.n.

Thalictrum thalictroides (L.) Eames & Boivin, 1919 O.E. Jennings

S.N.

Rhamnaceae

Ceanothus americanus L., 1986 R.C. Leberman s.n.

Frangula alnus P. Mill., 2003 S. Ernst et al. 145 Exotic

Rhamnus alnifolia UHér., 2004 J.A. Isaac 18439

Rhamnus cathartica L., 1998 K. Marsh s.n. Europe

Rosaceae

O d Wallr., 2005 L. Speedy LSB1122

a parviflora Ait., 2005 R. Thompson 05-714

2 pubescens Wallr., 1962 R.C. Leberman s.n.

Agrimonia striata Michx., 1904 O.E. Jennings s.n.

Amelanchier arborea (Michx. f) Fern. var. arborea, 2005 L.

Speedy LSB310

Amelanchier laevis Wieg., 2004 J.A. Isaac 17594

Amelanchier stolonifera Wieg., 2004 J.A. Isaac 17485

Amelanchier x intermedia Spach (pro sp) [arborea x canaden-

sis], 1972 R.C. Leberman s.n.

Comarum palustre L., 1997 J.K. Bissell and G. Buckley

1997104

Crataegus chrysocarpa Ashe var. chrysocarpa, 1963 P.B.

Monk s.n.

Crataegus crus-galli L., 1929 J. Bright 1779

Crataegus dissona Sarg., 1905 O.E. Jennings s.n.

Crataegus macrosperma Ashe, 2001 J.A. Isaac 13645

Crataegus mollis Scheele, 1922 J. Bright s.n. G5:SU

649

Crataegus monogyna Jacq., 1933 O.E. Jennings s.n. Eurasia

Crataegus prona Ashe, 1907 O.E. Jennings tree 82

Crataegus pruinosa (Wendl. f) K. Koch, 1907 O.E. Jennings

tree 84

Crataegus punctata Jacq., 1997 R.C. Leberman s.n.

Dalibarda repens L., 1994 J.D. Wagner and L.L. Smith 642

Filipendula rubra (Hill) B.L. Rob., 1970 R.C. Leberman s.n.

Filipendula ulmaria (L) Maxim., 1968 R.C. Leberman s.n.

Eurasia

Fragaria vesca L. ssp. americana (Porter) Staudt, 1904 O.E.

Jennings s.n.

Fragaria vesca L. ssp. vesca, 1950 L.K. Henry and W.E. Buker

s.n.

Fragaria virginiana Duchesne ssp. grayana (S.Watson) Staudt,

1904 O.E. Jennings s.n.

Fragaria virginiana Duchesne ssp. virginiana, 1930 W.R. Van

Dersal 1273

Geum aleppicum Jacq., 1952 L.K. Henry and W.E. Buker s.n.

Geum canadense Jacq. var. canadense, 2005 L. Speedy

Geum laciniatum Murr. var. laciniatum, 1951 L.K. Henry and

W.E. Buker s.n.

Geum rivale L., 2004 J.A. Isaac 17689

Malus coronaria (L.) P. Mill, 2005 L. Speedy LSB1117

Malus floribunda Sieb. ex Van Houtte, 1995 R.C. Leberman

s.n. Asia

Malus pumila P. Mill, 2005 R. Thompson 05-573 Eurasia [Pyrus

malus

Photinia MODEM (Lindl.) Robertson & Phipps, 1994 J.D.

Wagner et al. 119

Photinia melanocarpa (Michx.) Robertson & Phipps, 1991

R.C. Leberman s.n.

Photinia pyrifolia (Lam.) Robertson & Phipps, 1993 J.A. Isaac

4635

Physocarpus opulifolius (L) Maxim. var. opulifolius, 1997 R.C.

Leberman s.n

Potentilla norvegica L. ssp. monspeliensis (L.) Aschers. &

Graebn, 2001 J.A. Isaac and C.F. Chuey 13649

Potentilla recta L., 2004 J.A. Isaac 17688 Europe

Potentilla simplex Michx. var. simplex, 1907 O.E. Jennings s.n.

Prunus americana Marsh., 1924 O.E. Jennings s.n.

Prunus pensylvanica L. f. var. pensylvanica, 2005 L. Speedy

[SB327

Prunus serotina Ehrh. var. serotina, 2005 L. Speedy LSB1081

Prunus virginiana L. var. virginiana, 2004 J.A. Isaac 17591

Rosa carolina L. var. carolina, 1973 R.C. Leberman s.n.

Rosa eglanteria L., 1907 J.A. Shafer s.n. Europe

Rosa micrantha Borrer ex Sm., 1954 W.E. Buker s.n. Europe

Rosa multiflora Thunb. ex Murr., 2005 L. Speedy LSB1124

Asia

Rosa palustris Marsh., 1997 J.A. Isaac 9866

Rubus allegheniensis Porter var. allegheniensis, 2004 S.P. Grund

and L. Miller 3617

Rubus canadensis L., 1927 J. Bright s.n.

Rubus flagellaris Willd., 1955 H.A. Davis and T. Davis 10981

Rubus hispidus L., 2005 L. Speedy LSB763

650

Rubus idaeus L. ssp. strigosus (Michx.) Focke, 1992 J.K. Bissell

et al. 1992:064

Rubus ithacanus Bailey, 1955 H.A. Davis and T. Davis 10982

Rubus laudatus Berger, 1955 H.A. Davis and T. Davis 10971

Rubus occidentalis L., 1980 C.E. Jenkins and FQ. Jenkins

2861

Rubus odoratus L. var. odoratus, 1980 R.C. Leberman s.n.

Rubus pensilvanicus Poir., 2004 S.P. Grund and L. Miller 3527

Rubus pergratus Blanch., 1955 H.A. Davis and T. Davis 10962

Rubus plicatifolius Blanch., 1955 H.A. Davis and T. Davis

Rubus pubescens Raf. var. pubescens, 2005 L. Speedy

ESBIISS

Rubus roribaccus (Bailey) Rydb., 1904 O.E. Jennings s.n.

Rubus uvidus Bailey, 1955 H.A. Davis and T. Davis 10972

Rubus wheeleri (Bailey) Bailey, 1955 H.A. Davis and T. Davis

973

Sanguisorba canadensis L., 1987 J.K. Bissell and B. Danielson

Sorbus americana Marsh., 1915 J. Bright s.n.

Sorbus decora (Sarg.) Schneid., 1992 J.K. Bissell and B.

Danielson 92:173 G4G5:S1

Spiraea alba Du Roi var. latifolia (Ait.) Dippel, 1991 R.C.

Leberman s.n.

Spiraea tomentosa L., 2005 L. Speedy LSB1602

Waldsteinia fragarioides (Michx.) Tratt. ssp. fragarioides, 2004

J.A. Isaac 17334

Rubiaceae

Cephalanthus occidentalis L., 2005 L. Speedy LSB1093

Galium aparine L., 1980 C.E. Jenkins et al. 2812

Galium asprellum Michx., 2004 S.P. Grund and L. Miller 3623

Galium boreale L., 1996 R.C. Lebermans.n.

Galium circaezans Michx. var. circaezans, 1981 R.C. Leberman

s.n.

Galium circaezans Michx. var. hypomalacum Fern., 1932 W.R.

VanDersal s.n.

Galium concinnum Torr. & A. Gray, 1952 L.K. Henry and W.E.

Galium labradoricum (Wieg.) Wieg., 1994 J.K. Bissell et al.

94:024 G5:S1

Galium lanceolatum Torr., 1942 Richard W. Pohl 3976

Galium obtusum Bigelow ssp. obtusum, 2005 L. Speedy

LSB1088

Galium palustre L., 2004 L. Miller and S.P. Grund 25

Galium pilosum Aiton, 1969 W.E. Buker s.n.

Galium tinctorium (L.) Scop., 2005 L. Speedy LSB734

Galium trifidum L. ssp. trifidum, 1993 J.K. Bissell and B.

Danielson 1993:188 G5:S2

Galium triflorum Michx., 2005 L. Speedy LSB319

Houstonia caerulea L., 1905 O.E. Jennings s.n.

Mitchella repens L., 2005 L. Speedy LSB748

Salicaceae

Populus alba L., 1985 J.K. Bissell 85:055 Eurasia

Populus deltoides Bartr. ex Marsh. ssp. deltoids, 1930 O.E.

Jennings and G.K. Jennings s.n.

Populus grandidentata Michx., 1933 O.E. Jennings s.n.

Populus tremuloides Michx., 1997 R.C. Leberman s.n.

fal, Dat o ID L

Journal of

titute of Texas 1(1)

Populus x jackii Sarg., [balsamifera x deltoides] 1990 A.W.

Cusick 29024

Salix alba L., 2004 L. Miller and S.P. Grund 54 Eurasia

Salix amygdaloides Andersson, 2005 L. Speedy LSB1574

Salix bebbiana Sarg., 2004 J.A. Isaac 17541

Salix discolor Muhl., 2005 L. Soeedy LSB307-1

Salix eriocephala Michx., 2003 R. Coxe and M. Bradburn s.n.

Salix exigua Nutt., 2003 S. Ernst et al. 74

Salix fragilis L., 2003 S. Ernst et al. 72 Europe

Salix humilis Marsh. var. humilis, 1907 O.E. Jennings s.n.

Salix interior Rowlee, 2004 L. Miller and S.P. Grund 42

Salix lucida Muhl. ssp. lucida, 1988 J.K. Bissell and B. Danielson

1988:86

Salix myricoides Muhl. var. albovestita (Ball) Dorn, 1933 O.E.

Jennings s.n. G4:52

Salix nigra Marsh., 2003 S. Ernst et al. 73

Salix petiolaris Sm., 1994 J.K. Bissell et al. 94:048 G5:SNA

Salix purpurea L., 1933 O.E. Jennings s.n. Europe

Salix sericea Marsh., 2003 S. Ernst et al. 96

Salix serissima (Bailey) Fern., 2001 J.A. Isaac and C.F. Chuey

13650 G4:52

Sarraceniaceae

Sarracenia purpurea L. var. gibbosa (Raf) Wherry, 1929 O.E.

Jennings s.n.

Saururaceae

Saururus cernuus L., 2003 S. Ernst et al. 144

Saxifragaceae

Chrysosplenium americanum Schewein. ex Hook., 2004 J.A.

saac 17328

Heuchera americana L. var. americana, 1979 R.C. Leberman

S Fh

Hydrangea arborescens L., 1966 R.C. Leberman s.n.

Mitella diphylla L., 2005 L. Speedy LSB760

Penthorum sedoides L., 2004 S.P. Grund and L. Miller 3624

Ribes americanum P. Mill., 2005 L. Speedy LSB721

Ribes cynosbati L., 1997 R.C. Leberman s.n.

Ribes glandulosum Grauer, 1915 J. Bright s.n.

Ribes hirtellum Michx., 2004 J.K. Bissell 2004:089

Ribes lacustre (Pers.) Poir., 1966 R.C. Leberman s.n. G5:S1

Ribes rotundifolium Michx., 1924 O.E. Jennings s.n.

Ribes rubrum L., 2004 J.A. Isaac 17345 Eurasia

Ribes triste Pall., 2004 J.A. Isaac 17365 G5:S2

Saxifraga pensylvanica L., 2004 J.A. 1saac17349

Saxifraga virginiensis Michx. var. virginiensis, 1971 W.E. Buker

Tiarella cordifolia L. var. cordifolia, 2004 J.A. Isaac 17362

Scrophulariaceae

Aureolaria virginica (L.) Pennell, 1969 R.C. Leberman s.n.

Chaenorhinum minus (L) Lange, 2004 J.A. Isaac and R. Coxe

17587 Europe

Chelone glabra L., 1994 J.D. Wagner 674

Gratiola neglecta Torr., 1995 R.C. Leberman s.n.

Linaria vulgaris P. Mill., 1928 J.A. Murray s.n. Eurasia

Lindernia dubia (L.) Pennell var. dubia, 1988 J.K. Bissell et al.

Morton et al., Flora of Crawford County, Pennsylvania

Melampyrum lineare Desr. var. americanum, 1965 J.K. Bissell

and B.W. Danielson 1997:069

Mimulus alatus Ait., 1959 H.A. Wahl 19405

Mimulus ringens L. var. ringens, 2004 S.P. Grund and L. Miller

3503

Nuttallanthus canadensis (L.) D.A. Sutton, 1952 W.E. Buker s.n.

[Antirrhinum canadensis L.]

Pedicularis canadensis L. spp. canadensis, 2004 J.A. Isaac

17462

Pedicularis lanceolata Michx., 2004 J.A. Isaac 18462 G5:S1S2

Penstemon digitalis Nutt. ex Sims, 1981 R.C. Leberman s.n.

Penstemon laevigatus Aiton, 1972 R.C. Leberman s.n. G5:S3

Scrophularia lanceolata Pursh, 2004 J.A. Isaac 17608

Scrophularia marilandica L., 1970 W.E. Buker s.n.

Verbascum blattaria L., 2004 J.A. Isaac and R. Coxe 17581

Eurasia

Verbascum thapsus L., 1963 P.B. Monk s.n. Eurasia

Veronica americana Schwein. ex Benth., 2004 J.A. Isaac

Veronica arvensis L., 1919 O.E. Jennings s.n. Eurasia

Veronica chamaedrys L., 1996 J.K. Bissell et al.

Eurasia

Veronica officinalis L. var. officinalis, 2004 J.A. Isaac 17459

1990:09

Europe

Veronica peregrina L. ssp. xalapensis (Kunth) Pennell, 2004

J.A. Isaac 17353

Veronica persica Poir.,

Veronica scutellata L.,

1919 H.W. Mossman s.n. Eurasia

1997 J.K. Bissell and G. Buckley

Veronica serpyllifolia L. ssp. serpyllifolia, 1924 O.E. Jennings

Veronicastrum virginicum (L.) Farw., 1962 R.C. Leberman s.n.

Smilacaceae

Smilax herbacea L., 1995 R.C. Leberman s.n.

Smilax rotundifolia L., 1919 O.E. Jennings s.n.

Smilax tamnoides L., 1981 R.C. Leberman s.n.

Solanaceae

Physalis heterophylla Nees var. heterophylla, 1948 W.E. Buker

Sum

Solanum carolinense L. var. carolinense, 1998 R. Leberman s.n

Solanum dulcamara L. var. dulcamara, 2005 L. Speedy LSB726

Furasia

Sparganiaceae

Sparganium americanum Nutt., 2004 S.P. Grund and L. Miller

3485

Sparganium erectum L. ssp. stoloniferum (Graebn.) Hara, 2004

J.A. Isaac 18473

Sparganium eurycarpum Engelm. ex Gray, 2005 L. Speedy

LSB729

Staphyleaceae

Staphylea trifolia L., 1997 R.C. Leberman s.n.

Thymelaeaceae

Dirca palustris L., 2004 J.A. Isaac 18469

Tiliaceae

Tilia americana L. var. americana, 1997 R.C. Leberman s.n.

651

Typhaceae

Typha angustifolia L., 2005 J.K. Bissell 2005:043 Exotic

Typha latifolia L., 2005 L. Speedy LSB740

Typha x glauca Godr. (pro sp.) [angustifolia or domingensis],

1988 J.K. Bissell 1988:141

Ulmaceae

Ulmus americana L., 2004 S.P. Grund and L. Miller 3663

Ulmus rubra Muhl., 2004 S.P. Grund and L. Miller 3537

Urticaceae

Boehmeria cylindrica (L.) Sw., 2004 S.P. Grund and L. Miller

3492

lap rton rannrlonci

a canadensis (L) Wedd., 1909 O.E. Jennings s.n.

Pilea fontana (Lunell) Rydb., 2005 L. Speedy LSB1561

Pilea pumila (L.) Gray var. pumila, 2004 S.P. Grund and L.

Miller 3611

Urtica dioica L. ssp. gracilis (Ait.) Seland., 1944 O.E. Jennings

S.N.

Valerianaceae

Valeriana officinalis L., 2004 J.A. Isaac 17613 Eurasia

Verbenaceae

Phyla lanceolata (Michx.) Greene, 2000 J.K. Bissell and P.

Houghton 2000:086

Verbena hastata L. var. hastata, 2004 S.P. Grund and L. Miller

3491

Verbena urticifolia L. var. urticifolia, 1901 J.A. Shafer s.n.

Verbena x engelmannii Moldenke [hastata x urticifolia], 1901

Violaceae

Viola arvensis Murr., 1947 W.E. Buker s.n. Europe

Viola blanda Willd. var. blanda, 2004 J.A. Isaac 17493

Viola canadensis L. var. canadensis, 1930 W.R. VanDersal

1301

Viola cucullata Ait., 2005 L. Speedy LSB316

Viola hastata Michx., 2004 J.A. Isaac 17324

Viola labradorica Schrank, 2005 L. Speedy LSB317

Viola macloskeyi Llyod ssp. pallens (Banks ex Ging) M.S. Baker,

2004 J.A. Isaac 17322

Viola pubescens Ait. var. pubescens, 1942 O.E. Jennings s.n.

Viola pubescens Ait. var. scabriuscula Schwein. ex Torr. & A.

Gray, 1993 J.A. Isaac 4629

Viola rostrata Pursh, 2004 J.A. Isaac 17323

Viola rotundifolia Michx., 1952 L.K. Henry and W.E. Buker s.n.

Viola sororia Willd., 2005 L. Speedy LSB313

Viola striata Ait., 2004 J.A. Isaac 17347

Viola x bissellii House [cucullata x sororia], 1955 L.K. Henry

Sm:

Vitaceae

Parthenocissus quinquefolia (L.) Planch., 2005 L. Speedy

LSB728

Parthenocissus vitacea (Knerr) A.S. Hitchc., 1990 AW. Cusick

29027 [Ampelopsis quinquefolia var. vitacea Knerr]

Vitis aestivalis Michx. var. bicolor Deam, 1997 R.C. Leberman

s.n.

Vitis riparia Michx., 2005 L. Speedy LSB722

652

Zannichelliaceae

Zannichellia palustris L., 1881 J. Whitesides s.n.

GYMNOSPERMS

Pinaceae

Larix deciduas P. Mill, 2005 R. Thompson 05-1066 Europe

Larix laricina (Du Roi) K. Koch, 1997 J.K. Bissell and B. Gordon

1997:106

Pinus banksiana Lamb., 2005 L. Speedy LSB326

Pinus strobus L., 2005 L. Speedy LSB321

Pinus sylvestris L., 1975 W.E. Buker s.n. Eurasia

Tsuga canadensis (L.) Carr., 2005 L. Speedy LSB10832

Taxaceae

Taxus canadensis Marsh., 1999 J.K. Bissell 1999:091

LYCOPHYTES

Lycopodiaceae

Huperzia lucidula (Michx.) Trevisan, 2004 J.A. Isaac 17326

Lycopodium annotinum L., 1990 J.K. Bissell and B. Danielson

1990:196

Lycopodium clavatum L., 1984 J.K. Bissell 1984:106

Lycopodium dendroideum Michx., 1966 R.C. Leberman s.n.

Lycopodium digitatum Dill. ex A. Braun, 2005 L. Speedy

Lycopodium hickeyi W.H. Wagner, Beitel & Moran, 1965 DLL.

Pearth s.n.

Lycopodium obscurum L., 2005 L. Speedy LSB1090

Lycopodium tristachyum Pursh, 1962 R.C. Leberman s.n.

Lycopodium x habereri House [digitatum x tristachyum], 1997

J.K. Bissell and G. Buckley 1997:156

Lygodiaceae

Lygodium palmatum (Bernh.) Sw., 1968 R.C. Leberman s.n.

PTERIDOPHYTES

Aspleniaceae

Asplenium platyneuron (L) B.S.P. var. platyneuron, 1991 B.L.

Isaac and J.A. Isaac 3550

Blechnaceae

Woodwardia virginica (L.) 5m., 2004 J.A. Isaac 18476

Dennstaedtiace

Dennstaedtia ne (Michx.) T. Moore, 1997 R.C.

Leberman s.n.

Pteridium aquilinum (L.) Kuhn var. latiusculum (Desv.)

Underwood ex Heller, 2004 S.P. Grund and L. Miller

3666

Dryopteridaceae

Athyrium filix-femina (L.) Roth ssp. angustum (Willd.) Clausen,

1984 J.K. Bissell 1984:126

Cystopteris bulbifera (L.) Bernh., 1965 J. Stull and D. Stull s.n.

Cystopteris tenuis (Michx.) Desv., 1982 R.C. Leberman s.n.

Deparia acrostichoides (Sw.) M. Kato, 1984 J.K. Bissell

84:126

Diplazium pycnocarpon (Spreng) Broun, 2004 J.A. Isaac

18430

Journal of the Botanical R h Institute of Texas 1(1)

pteris carthusiana (Vill.) H.P. Fuchs, 2004 S.P. Grund and

Es Miller 3542

Dryopteris clintoniana (D.C. Eat.) Dowell, 2004 J.A. Isaac

18465 G5:52

Dryopteris cristata (L.) Gray, 2005 L. Speedy, M. Bowers and

M. Fodse LSB1621

Dryopteris goldiana (Hook. ex Goldie) Gray, 2004 J.A. Isaac

et al. 17718

Dryopteris intermedia (Muhl. ex Willd.) Gray, 2005 L. Speedy

LSB750

Dryopteris marginalis (L.) Gray, 1962 R.C. Leberman s.n.

Dryopteris x boottii (Tuck) Underwood (pro sp.) [cristata x,

2004 J.A. Isaac 18467

Dryopteris x triploidea Wherry |

J.K. Bissell 1984:120

Gymnocarpium dryopteris (L.) Newman, 2004 J.A. Isaac

17498

Matteuccia struthiopteris (L.) Todaro, 2004 J.A. Isaac 18423

Onoclea sensibilis L., 2005 L. Speedy LSB1131

Polypodium appalachianum Haufler & Windham, 1906 O.E.

Jennings s.n.

Polypodium virginianum L., 1914 C.A. Darling s.n.

Polystichum acrostichoides (Michx.) Schott var. acrostichoides,

2005 L. Speedy LSB746

husi intermedia], 1984

Equisetaceae

Equisetum arvense L., 1994 J.D. Wagner and J.K. Bissell 344

Equisetum fluviatile L., 2005 L. Speedy, J. Bissell, M. Bowers

and T. Pearson LSB1584

Equisetum hyemale L. var. affine (Engelm) A.A. Eat., 1970 W.E.

Buker s.n.

Equisetum sylvaticum L., 2005 L. Speedy LSB340

Isoetaceae

Isoétes tenella Léman, 1933 W.R. Van Dersal s.n. [/soétes braunii

Durieu]

Ophioglossaceae

Botrychium dissectum Spreng.,

Hoberecht 1985:048

Botrychium lanceolatum (Gmel.) Angstr. var. angustiseg-

mentum Pease & Moore, 1930 O.E. Jennings and G.K.

Jennings s.n

Botrychium matricariifolium (A. Braun ex Dowell) A. Braun ex

Koch, 1993 J.K. Bissell and B. Danielson 1993:091

Botrychium multifidum (Gmel.) Trev., 1984 A.W. Cusick

1985 J.K. Bissell and M.

Botrychium oneidense (Gilbert) House, 1985 J.K. Bissell and M.

Hoberecht 1985:048

Botrychium simplex E. Hitchc.,

Furope

NT virginianum (L.) Sw., 1984 J.K. Bissell 1984106

glossum pusillum Raf., 1939 J. Wurdach s.n.

o vulgatum L., 1970 R.C. Leberman s.n. G5:S4

1982 R.C. Leberman s.n.

Osmundaceae

Osmunda cinnamomea L. var. cinnamomea, 2005 L. Speedy

LSB304

Osmunda claytoniana L., 2004 J.A. Isaac 17470

Morton et al., Flora of Crawford County, Pennsylvania 653

Osmunda regalis L. var. spectabilis (Willd) Gray, 2005 L.Speedy Thelypteridaceae

Phegopteris connectilis (Michx.) Watt, 1955 L.K. Henry s.n.

preridscese Phegopteris hexagonoptera (Michx.) Fée, 1968 R.C. Leberman

Adiantum pedatum L., 1997 R.C. Leberman s.n. An E ,

Thelypteris noveboracensis (L.) Nieuwl., 2005 L. Speedy

Selaginellaceae LSB751

Selaginella apoda (L.) Spring, 2004 J.A. Isaac 17486 Thelypteris palustris Schott var. palustris, 1999 J.K. Bissell

1999:079

ACKNOWLEDGMENTS

Our thanks go to the Wild Resource Conservation Fund of the Pennsylvania Department of Conservation

and Natural Resources for partial funding of the field work. We would also like to thank Ann Rhoads for

reviewing the manuscript. We are also thankful to Kelly Tuite for data entry.

REFERENCES

Bates, S. P. 1885. History of Crawford County, Pennsylvania. Warner Beers and Co., Chicago, IL.

Brumm, R.K. and C.E. Powel. 1992. Authors of plant names. Royal Botanic Gardens, Kew.

Kartesz, J.T. 1999. A synonomized checklist and atlas with biological attributes for the vascular flora of the United

States, Canada and Greenland. First ed. In: Kartesz, J.T. and C.A. Meacham. Synthesis of the North American

Flora, Version 1.0. North Carolina Botanical Garden, Chapel Hill.

RHOADS, A.F. and W.M. KLEN, JR. 1993. The vascular flora of Pennsylvania: annotated checklist and atlas. American

Philosophical Society, Philadelphia, Pennsylvania.

RHOADS, A.F. and T.A. Block 2000. The plants of Pennsylvania. University of Pennsylvania Press, Philadelphia,

Pennsylvania.

THE NATURE CONSERVANCY (1996 version). Element Ranking List—Pennsylvania Natural Heritage Program. http://www.

dcnr.state.pa.us/forestry/pndi/rank.htm.

USDA. 1979. Soil Survey of Crawford County Pennsylvania. USDA Soil Conservation Service. US Government

Printing Office, Washington, D.C.

USDA, 2000. County Lookup. http://www.ers.usda.gov/Data/UrbanInfluenceCodes/2003/LookUpUIC.asp

654 Journal of the Botanical R h Institute of Texas 1(1)

DOOK NOTICES

RJ. CHinNock. 2007. Eremophila and Allied Genera: A Monograph of the Myoporaceae. (ISBN 978-1-

877-05816-5, hbk). Rosenberg Publishing Pty Ltd, PO Box 6125, Dural Delivery Centre, NSW 2158,

Australia. (Orders: www.isbs.com, 503-287-3093, 503-280-8832 fax). $99.95, 672 pp., 335 color

plates, 300 maps, 325 line drawings, 842" x 111^".

The seven genera of Myoporaceae are described up, down, sideways, and in yet intangible dimensions. Treatments include formal tax-

onomy, detailed descriptions and maps, beautiful and informative line drawings, many interspersed color photos of plants and flowers,

and notes on distribution, ecology, conservation status and relationships. Three new genera, 94 new species, and 37 new subspecies are

described, and revised infrageneric classifications of Eremophila and Myoporum are proposed. All this is presented in such an elegant

fashion that it invites being laid on the coffee table for recreational reading (not to belittle its academic value).

Contents: 1) Introduction; 2) History of the Myoporaceae; 3) Morphology and Anatomy; 4) Trichomes; 5) Floral and Fruit Mor-

phology; 6) Palynology; 7) Cytology and Cytogeography; 8) Reproductive Biology; 9) Distribution and Ecology; 10) Phytochemistry

and Toxicity; 11) Traditional and Economic Uses; 12) Horticultural Uses; 13) Phylogeny and Relationships of the Myoporaceae; 14)

Taxonomic Treatment of the Family Myoporaceae

Five of the seven genera of Myoporaceae occur in Australia, where all but one of these, Myoporum, are endemic. Bontia and Penta-

coelium each contain a single species and are restricted to the Caribbean (plants have recently been found in Florida, near Miami) and

to southern China, Japan, and northern Vietnam. Myoporum is largely Australian but species extend into the Pacific (including New

Zealand and Hawaii) and one species to the Indian Ocean.

The high salt tolerance and drought resistance of Myoporaceae combined with attractive iss has made this family, especially

Myoporum, à bs AN introduction in drier regions around the world. Most sq bs with a variety of flower colour

AS. 1

Forst. i alifornia.—Guy

Nesom, Botanical o mu of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U. s A

ALAN E. BESSETTE, WILLIAM C. Roopy, ARLEEN R. BESSETTE, AND Dai L. Dunaway. 2007. Mushrooms of the South-

eastern United States. (ISBN 978-0-8156-3112-5, hbk). Syracuse DESY Press, 621 Skytop Road,

Suite 110, Syracuse, NY, 13244-5290, U.S.A.. (Orders: www.sy I syr.edu, 315-443-

5534, 315-443-5545 fax). $95.00, 400 pp., 517 color images, 1 map, 74" x 1014".

More than 450 species are treated, with 517 color photographs (on 75 pages) and detailed descriptions, for an area that includes Texas,

Oklahoma, and Kansas east to West Virginia, Virginia, and southward to (and including) Florida. The authors note that the book provides

a “relatively comprehensive guide” and that “the number of species described and illustrated in color is substantially higher than has

previously appeared in any other single work devoted to the mushrooms of the southeastern United States. We provide cross-references

to additional species occurring in the region that are illustrated in Mus) jj f Nortl North America" (the companion volume, also

Syracuse Univ. Press). For use by a general audience, the bool hasizes identifi

based primarily on macroscopic field characters

but the authors note that it also provides necessary detail nr by advanced students and professional mycologists

There are lots of field guides and other books on mushrooms of eastern North America, some with large pictures and numer-

ous species, but this one apparently is the ticket. On the other hand, local field guides remain indispensable. For example, from Texas

mushrooms: a field guide (Metzler & Metzler 1992), there are 43 species (8 in Amanita) and 8 genera (Lentaria, Oudemansiella, Phlogiotis,

Sarcodon, Scutellina, Sphaerobolus, Steccherinum, Tremellodendron) not accounted for in the index of Bessette et al. It's not easily apparent

whether this is because E taxonomic/ nomenclatural issues or because the southern and western edges of Texas include the ranges of

primarily ti f both factors).— Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth,

TX 76102-4060, US. A.

J. Bot. Res. Inst. Texas 1(1): 654. 2007

VASCULAR FEORA OF DHEBOURCANYOINSPRESBERVE:

ELLIS COUNTY<OKLATOM:A

Bruce W. Hoagland Amy K. Buthod

Oklahoma Biological Survey and Oklahoma Biological Survey

Department of Geography University of Oklahoma

University of Oklahoma Norman, Oklahoma 73019, U.S.A.

Norman, Oklahoma 73019, U.S.A. amybuthod@ou.edu

bhoagland@ou.edu

ABSTRACT

This paper reports the results of an inventory of the vascular plants from The Nature Conservancy’s Four Canyons Preserve in Ellis

County, Oklahoma. A total of 371 taxa of vascular plants in 244 genera and 77 families were collected, with the most species occurring

in the families Asteraceae (69), Poaceae (64), and Fabaceae (38). One hundred-twelve species were annuals, four biennials, and 255

perennials. Fifty-one species of woody plants were present. Twenty-nine exotic species were collected representing 7.8% of the flora.

Six tracked by the Oklahoma Natural Heritage Inventory were found.

RESUMEN

En este articulo se presentan los resultados de un inventario de plantas vasculares de The Nature Conservancy’s Four Canyons Preserve

en Ellis County, Oklahoma. Se colectaron un total de 371 taxa de plantas vasculares de 244 géneros y 77 familias, siendo la mayoria

de las especies pertenecientes a las familias Asteraceae (69), Poaceae (64), y Fabaceae (38). Ciento doce especies fueron anuales, cuatro

bienales, y 255 perennes. Estan presentes cincuenta y una especies de plantas lefiosas. Se colectaron veintinueve especies exdticas que

representan el 7.8% de la flora. Se encontraron seis especies a las que hace un seguimiento el Oklahoma Natural Heritage Inventory.

INTRODUCTION

There are currently 501 taxa of vascular plants known to occur in Ellis County, Oklahoma (Hoagland et al.

2006). The first collections in the county were made in 1913 by G.W. Stevens, who deposited 69 species. In

the following year, 63 collections were deposited at the University of Oklahoma Herbarium (OKL) by R.L.

Clifton. No collections from Ellis County were deposited in state herbaria between 1915 and 1927. After

that time, there was steady collection in the county. The 1970s were an active decade for botanical study

of Ellis County, culminating in the deposit of 101 specimens at (OKL) collected by Barber, K. Pearce, and

R. Thompson in 1976. From 1985 to 1986, F.B. Erteeb deposited 405 specimens from Ellis County at the

Oklahoma State University (OKLA) herbarium as part of a floristic study of Northwest Oklahoma. In 1998,

N. McCarty and B. Hoagland deposited 226 specimens at OKL in conjunction with a study of wetland and

woody plants. Since that time there has been little collection effort in Ellis County. The objective of this

study was to provide a floristic inventory to aid Nature Conservancy personnel in management of the Four

Canyons Preserve (FCP) and remedy a gap in our knowledge of the flora of western Oklahoma.

STUDY AREA

The FCP (Fig. 1) was established by The Nature Conservancy in 2004 and protects 1,376 hectares. It is within

the Subtropical Humid (Cf) climate zone (Trewartha 1968). Summers are warm (mean July temperature =

26.6° C) and humid, and winters are relatively short and mild (mean January temperature = 0.3? C). Mean

annual precipitation is 60.4 cm (Oklahoma Climatological Survey 2006).

Physiographically, the study area is located in the Western Redbed Plains (Curtis & Ham 1979) of

Osage Plains Physiographic Province (Hunt 1974). The topography consists of gently rolling hills with deeply

eroded canyons. The surface geology is comprised of Permian age sandstones and shale in the uplands and

quaternary alluvial deposits on the Canadian River floodplain (Branson & Johnson 1979). The Quinlan-

Woodward soil association, a reddish loamy upland soil predominates at the site. The Lincoln-Spur Associa-

J. Bot. Res. Inst. Texas 1(1): 655 — 664. 2007

656 Journal of the Botanical R h Institute of Texas 1(1)

AAA

Arnett

Four Canyons

Nature Preserve

Cana dian

4 r r f D. * Ellie f£

Fic. 1. Location of The N y Four Canyons f ty, Oklahoma

tion, consisting of sandy to laomy soils, predominates on the Canadian River floodplain (Cole 1961). Duck

and Fletcher (1943) classified the study area as part of the mixedgrass eroded plains, defined as having a

“mixed grass composition and a definite ravine relief which is generally wooded. It is part of an extensive

area of similar vegetational conditions extending as an overlapping of tall grass species from the east, with

short grass species from the west northward across the United States." Vegetation along the Canadian River

was classified as bottomland, a category that was not clearly defined, but would include cottonwoods and

riparian shrubs and herbaceous species in the study area.

METHODS

Five collection sites were regularly visited for intensive floristic sampling. Additional collections were also

made randomly throughout the site. Collecting began on March through October 1999. Sites were selected

following a review of U. S. Geological Survey 1:24,000 topographic maps and field reconnaissance. The

predominant vegetation association at these sites was classified according to Hoagland (2000). Vouchers

for exotic species were made from naturalized populations only, thus excluding cultivated and ornamental

plants. Specimens were processed at the Robert Bebb Herbarium of the University of Oklahoma (OKL)

following standard procedures. Manuals used for specimen identification included Correll and Johnston

(1970), Great Plains Flora Association (1986), and Waterfall (1973). Origin, either native or introduced, was

determined using Taylor and Taylor (1991) and USDA-NRCS (2006). Nomenclature follows the United States

Department of Agriculture-Natural Resources Conservation Service (USDA-NRCS 2006). Voucher specimens

were deposited at the Robert Bebb Herbarium (OKL) at the University of Oklahoma.

RESULTS AND DISCUSSION

A total of 371 taxa of vascular plants in 77 families and 244 genera were collected (Appendix 1). Among

the angiosperms, 86 were monocots and 283 were dicots. In addition, there were two ferns and allies and

one gymnosperm. The Asteraceae (69), Poaceae (64), and Fabaceae (38) had the greatest number of species.

The genus Asclepias had the greatest number of species (10). One hundred-twelve species were annuals, four

Hoagland and Buthod, Flora of Four Canyons Preserve, Oklahoma 657

biennials, and 255 perennials. Fifty-one species of woody plants were collected. One hundred and eleven

species previously unreported from Ellis County were collected in this study.

Thirty taxa (8.196 of the flora) from 14 families were exotic. The families with the greatest number of

introduced species were Poaceae with twelve, Asteraceae with four, and Fabaceae with two. Genera with

the most exotic species were Bromus with three and Vulpia with two. The percentage of exotic species in

this flora is comparable with reports from other floras in Oklahoma (7-15% exotic) (Hoagland € Johnson

2001, 20042, 2004b; Hoagland € Buthod 2003, 2004; Hoagland & Wallick 2003; Hoagland et al. 2004;

Hoagland et al. 2004; Hoagland & Buthod 2005a; Hoagland & Buthod 2005b; Hoagland & Johnson 2005),

with the exception of two sites in McCurtain County, where 6.696 of the flora consisted of exotic species

(Hoagland € Johnson 20049). See Table 1 for a floristic summary of FCP.

No federally listed threatened or endangered species were encountered. However, five species tracked

by the ONHI (2005) were present: Argythamnia mercurialina (G5,S2S3), Echinocereus reichenbachii (G5, S2),

Escobaria vivipara (G5, $253), Muhlenbergia bushii (G5, S152), Vitis rupestris (G3, S?) and Zinnia grandiflora

(G4G5, S?). Species are ranked according to level of imperilment at the state (S) and global (G) levels on a

scale of 1—5; 1 representing a species that is imperiled and 5 one that it is secure. A “?” indicates a species

with rank influx (Groves et al. 1995).

The five collection sites at Four Canyons occurred within six vegetation associations. A brief description

of each follows:

1. Disturbed areas and old-fields were locations exhibiting signs of physical disruption, such as roadsides and home sites. This includes

a portion of the eee River E ns was ds tones n in dd prior to n by The Nature Conservancy.

Common pl gilo] icus, Cucurbita foetidissima, Eragrostis

A EC i lini Hord ill Mollug( Ta Oxalis stricta, Phytol i Polygonum aviculare, Portulaca

[9] [o]

halimoides, Quincula lobata, Rhus glabra, Er Tribulus terrestris.

2. Quercus muehlenbergii-Juniperus virginiana woodland association occupied deep sandstone canyons at the FCP. This vegetation type

does not appear in Hoagland (2000). Although Q. muehlenbergii has been reported from counties immediately south of Ellis, it does not

occur as a dominant species. The co-occurrence of J. virginiana is likely the result of fire suppression, but additional research is neces-

sary to character species composition and stand history. Associated species included Acalypha ostryifolia, Celastrus scandens, Cornus

drummondii, Elymus ei Galium o ame Juglans A ai Parietaria penslvanica, Pellaea atropurpurea, Ribes aureum, Sideroxylon

E i € 1

lanuginosum

, Toxicodendron radicans, Tridens flavus, and Ulmus rubra. Muhlenbergia bushii, a

species tracked by ONHI, "T in this UM

3. Quercus havardii-Sporobolus cryptandrus-Schizachyrium scoparium shrubland association was limited to a deposit of sandy soils on the

western edge ofthe pe Extensive occurrences M this Ed association are west of the FCP on deep sandy soils. Associated species

included Art i g igo Itiflora, Oenothera grandis, Prunus gracilis, and Yucca glauca.

DI +: TP PES) VE Been iation occurred on the well-drained soils and rockv

slopes. It was the most abundant vegetation type at FCP. Plant cover was sparse on sandstone outcrops Associated forb species included

Amorpha canescens, id AM Rd gracilis, — ao dd a ane: herbaceous, Comandra umbellata, Croton

texensis, Dalea aurea, D i deoma drummon longifl Lesquerella

ova ae ia, Krameria NM oca bc Pie dl r cuspidatum, 1 Ptelea nfl iata, med nutans, Sphaeralcea coccinea,

y the ONHI that occurred in this association

were ani ia Escobaria vivipara ad Zinnia grandiflora.

5. Schizachyrium scoparium— Castilleja purpurea var. citrina—Lesquerella gordonii herbaceous association occurred on shallow soils over

gypsum. The extent of escam cover varies with nue NA of TRUE exposing Associates include Bouteloua hirsuta, Erioneuron

pilosum, Chaet ides, Liatris punctata Pl and Ti ji Echinocereus reichenbachii,

which is e) byd the Oklahoma Natural Heritage inventory "es in ae habitat type.

6. Wetland and R an vegetation included t] g ll ponds at FCP and wet areas along the floodplain of the Canadian River.

Although P s land g i i the FCP, th hey are limited in aerial extent and readily intergrade. Classifying

Canadian River floodplain vee Ue is further con ere by land use/land conversion practices iR landowner a ior to The Nature

Conservancy. Possible ne g types present include Pol lvani Pol

Schoenoplectus americanus hari i is herbaceous association, and the a pd hrubland association

APPENDIX 1

Annotated species list for The Nature Conservancy's Four Canyon Preserve. The first entry indicates habitat

(DAOF = disturbed areas and old fields, QHSC = Quercus havardii-Sporobolus cryptandrus-Schizachyrium

658 Journal of the Botanical R h Institut

of Texas 1(1)

TABLE 1. Summary of floristic collections from the Four Canyons Preserve, Ellis County, Oklahoma.*

Taxonomic Group Species Native Exotic

Pteridophyta 2 2 0

Coniferophyta 1 1 0

Magnoliophyta 364 337 27

Magnoliopsdia 20 261 16

Liliopsida 87 76 11

Table follows the format of Palmer et al. 1995.

scoparium shrubland association, QMJV - Quercus muehlenbergii-Juniperus virginiana woodland association,

SSBC - Schizachyrium scoparium-Bouteloua curtipendula-Bouteloua gracilis herbaceous association, S$CP -

Schizachyrium scoparium-Castilleja purpurea var. citrina-Lesquerella gordonii herbaceous association, WETL =

wetland and riparian) followed by life history is designated as A=annual, B=biennial, or P=perennial, and

collection number. Introduced species are noted with an asterisk.

PTERIDOPHYTA

Equisetaceae

Equisetum laevigatum A. Braun—DAFL, WETL; P; 46103

Pteridaceae

Pellaea atropurpurea (L.) Link—QMJV; P; 44030

CONIFEROPHYTA

Cupressaceae

Juniperus virginiana L.—GMJV, DAUP, SSBC P; 4G019

MAGNOLIOPHYTA—MAGNOLIOPSIDA

Acanthaceae

Ruellia humilis Nutt.—SSBC; P; 4C330

Amaranthaceae

Amaranthus albus L.—DAUP; A; 4329

Anacardiaceae

Rhus aromatica Aiton—QHSC, SSBC; P; 4C027

Rhus glabra L.—SSBC; P; 4C214

Toxicodendron radicans (L.) Kuntze—QOMJV; P; 4C307

Apiacea

PURGE popei Torr. & A. Gray—SSBC; A; 4C075

Cymopterus acaulis (Pursh) Raf.—SSBC; P; 4C015

Eurytaenia texana Torr. & A. Gray—SSBC; A; 46288

Spermolepis echinata (Nutt. ex DC) A.Heller—DAUP, SSBC;

Apocynaceae

Apocynum cannabinum L.—DAFL, DAUP, SSBC; P; 4C187

Asclepiadaceae

Asclepias arenaria Torr.—SSBC; P; 4C280

Asclepias asperula (Dcne) Woods.—SSBC; P; 4C078

Asclepias engelmanniana Woods.—SSBC; P; 4C266

Asclepias latifolia (Torr.) Raf.—SSBC; P; 4C434

Asclepias pumila (A. Gray) Vail —OHSC; P; 46378

Asclepias stenophylla A. Gray—SSBC; P; 4239

Asclepias syriaca L.—DAFL; P; 4-432

Asclepias tuberosa L.—SSBC; P; 4C164

Asclepias verticillata L.—SSBC; P; 4C336

Asclepias viridiflora Raf—SSBC; P; 4345

Astera

ien em DC.—DAFL, DAUP, OMJV, SSBC; P;

Amphiachyris dracunculoides (DC.) Nutt.—DAUP, SSBC; A;

4C29]

Aphanostephus skirrhobasis (DC.) Trel—QHSC, SSBC; a;

-217

Artemisia dracunculus L.—SSBC; P; 4-293

Artemisia filifolia Torr.—QHSC, SSBC; P; 4C292

Artemisia ludoviciana Nutt.—DAUP, SSBC; P; 4C-037

Baccharis salicina Torr. & A. Gray—DAFL; P; 4C340

Brickellia eupatorioides (L.) Shinners—SSBC; P; 4-398

Centaurea americana Nutt.—DAUP, SSBC; A; 4C290

Chaetopappa ericoides (Torr.) G.L. Nesom

Chloracantha spinosa (Benth.) G.L. Nesom—DAFL; P; 46232

Cirsium ochrocentrum A. Gray—DAUP, SSBC; P; 4C180

Cirsium undulatum (Nutt.) Spreng.—DAUP, SSBC; P; 4274

Conyza canadensis (L) Cronquist—DAUP, QMJV, SSCP; A;

2205

Croptilon hookerianum (Torr. & A. Gray) House—SSBC; A;

4C-326

Echinacea angustifolia DC.—SSBC; P; 446264

Engelmannia peristenia (Raf) Goodman & C.A.Lawson—SSBC;

bellidiast.

H IUI

I astrum Nutt.—OHSC; A; 4C112

eee a & A. Gray—SSBC; P; 4-419

Erigeron strigosus Muhl. ex Willd.—SSBC; A; 4C165

Euthamia gymnospermoides Greene—SSBC; P; 4C384

Does Nutt. ex DC.—DAUP, SSBC; A; 4C110

pest eM tee DAUP; A; 4C377

Walter) H.Rock—QHSC, SSBC; P; 4C385

Gaillardia pulchella rue —QHSC, SSBP; A; 4052

Gaillardia suavis (AGray & Engelm.) Britt & Rusby—SSCP; P;

(5nillanrdin

4C-060

Grindelia lanceolata Nutt. SSBC; P; 4C366

Hoagland and Buthod, Flora of Four Canyons Preserve, Oklahoma

Grindelia nuda Wood var. nuda—DAUP; P; 4C:349

Grindelia squarrosa (Pursh) Dunal—DAUP; A; 4G372

Gutierrezia sarothrae (Pursh) Britt & Rusby

Haloesthes greggii A. Gray—SSCP; P; 44289

Helenium amarum (Raf) H.Rock—DAUP, SSBC; A; 4C043

Helianthus annuus L.—DAFL, DAUP, WETL; A; 4G312

Helianthus petiolaris Nutt.—DAFL; A; 4-352

Heterotheca stenophylla (A. Gray) Shinners var. stenophylla—

SSBC; P; 4C286

Heterotheca subaxillaris (Lam.) Britt € Rusby—SSBC; A; 4G

Hymneopappus flavescens A. Gray—SSBC; B; 4C170

Hymneopappus tenuifolius Pursh—QHSC, SSBC; P; 4C237

lva annua L.—DAFL; A; 4354

Lactuca serriola L*—DAUP; A; 4343

Liatris mucronata DC.—SSBC; P; 4C416

Liatris punctata Hook.—SSBC, SSCP; P; 4C371

Liatris squarrosa (L.) Michx.—SSBC; P; 46331

Lygodesmia juncea (Pursh) D.Don ex Hook.—SSBC; P; 4C271

Machaeranthera pinnatifida (Hook.) Shinners—SSBC; P; 4G

285

Packera plattensis (Nutt) W.A Weber & A. Love—SSBC; B;

Pluchea odorata (L.) Cass.—DAFL, WETL, QMJV; A; 4C-361

Psilostrophe tagetina (Nutt.) Greene var. cerifera (A.Nels.) B.L.

Turner—SSCP; B; 4C-031

Pyrrhopappus grandiflorus (Nutt.) Nutt.—DAUP, SSBC; P;

4C-094

Ratibida columnifera (Nutt. Wooten & Standl.—SSBC; P;

4C-303

Senecio riddellii Torr. & A. Gray—DAFL; P; 4C040

Solidago altissima L.—SSBC; P; 4415

Solidago canadensis L. var. canadensis—QMJV, SSBC; P; 4G

428

Solidago gigantea Aiton—WETL, SSBC; P; 46419

Solidago petiolaris Aiton—SSBC; P; 4C411

Sonchus asper (L.) Hill*—DAUP, QMJV; A; 4C186

Symphyotrichum ericoides (L.) G.L. Nesom var. ericoides—SSBC;

P; 4C394

Symphyotrichum fendleri (A. Gray) G.L. Nesom—SSBC; P;

Symphyotrichum oblongifolium (Nutt) G.L. Nesom—SSBC;

; 98

Symphyotrichum subulatum (Michx.) G.L. Nesom—DAFL,

WETL; A; 4C035

Taraxacum officinale G Weber ex AWigg.*—DAUP; P; 4C-029

Tetraneuris linearifolia (Hook) Greene—SSBC; A; 4C010

Tetraneuris scaposa (DC.) Greene—SSCP; P; 4C011

He megapotamicum (Spreng. Kuntze—SSBC; P;

rons Escapa (H.Richards.) om s P; 4C 003

Tragop ibius Scop.*—DAUP; A; 4C10

Venn baldwinii Torr.—DAUP, QMJV; P; na 319

Xanthium strumarium L.—DAFL, WETL; A; 4C356

Zinnia grandiflora Nutt.—SSCP; P; 4C-316

1LUUOIU

Boraginaceae

Cryptantha minima Rydb.—SSBC; A; 44-084

659

Lappula occidentalis (S. Watson) Greene var. occidenta-

lis—DAUP; A; 4C 141

Lithospermum caroliniense (Walter ex J.F.Gmel.) MacMil.—

SSBC; P; 4C 144

Lithospermu m incisum Lehm.—SSCP, SSBC; P; 4C-016

Brassicaceae

Camelina microcarpa DC.*—DAUP; A; 4C072

Descurainia pinnata (Walter) Britt—DAUP; A; 4G020

Draba cuneifolia Nutt. ex Torr. & A. Gray—DAUP, SSBC; A;

4C-022

Lepidium densiflorum Schrad.—DAUP, SSBC; A; 46-093

Lepidium oblongum Small —DAUP, SSBC; A; 4C017

a ud diei (A. Gray) S.

Le rella ovalifol a Rydb.ex Britt —SSBC; P; AGI 18

SD E hyacinthoides Hook.—SSBC; A; 4C258

Cactaceae

Echinocereus reichenbachii (Terscheck ex Walp.) Haage

f.—SSBC, SSCP; P; 4C136

Escobaria vivipara (Nutt.) Buxb.—SSBC; P; 4C174

Opuntia macrorhiza Engelm.—DAUP, SSBC; P; 4C-424

Campanulaceae

Triodanis holzingeri McVau

gh—DAUP, SSBC; A; 4C246

Triodanis perfoliata (L.) Nieuwl.—

DAUP, SSBC; A; 4C273

Capparaceae

Polanisia dodecandra (L.) DC.—OHSC, SSBC; A; 4379

Caprifoliaceae

Symphoricarpos

biculatus Moench—QMJV; P; 4C 147

Caryophyllaceae

Dianthus armeria L*—SSBC; A; 4C 088

Paronychia jamesii Torr. & A. Gray—SSCP; P; 4C247

Celastraceae

Celastrus scandens L.—OMJV; P; 4C 039

Chenopodiaceae

Atriplex canescens (Pursh) Nutt.—SSBC; P; 4C104

Chenopodium leptophyllum (Moq.) Nutt. ex S. Watson—DAUP;

E 68

PUE simplex (Torr.) Raf —QMJV; A; 4376

nolepis (Schult) Greene—WETL; A; 4C143

brin tragus L*—DAFL, DAUP; A; 4G341

PREIS d 5d

Convolvulaceae

Convolvulus arvensis L*—DAUP; P; 4C210

Fvolvulus li, Schult.—SSCP; P; 4C051

Ipomoea leptophylla Torr.—SSBC; P; 4C218

Cornaceae

Cornus drummondii C. A.Mey.—OMUV; P; 4C203

Cucurbitaceae

Cucurbita foetidissima Kunth—DAUP; P; 4C-178

Elaeagnaceae

Elaeagnus angustifolia L*—DAFL; P; 4374

Euphorbiaceae

Acalypha ostryifolia Riddell—DAUP, OMJV; A; 4185

Argythamnia mercurialina (Nutt.) Müll.Arg.—SSBC; P; 4C197

660

Chamaesyce fendleri (Torr. & A. Gray) Small —DAUP, OHSC;

P; 4C099

Chamaesyce glyptosperma (Engelm.) Small —DAUP, SSBC;

154-394.

Chamaesyce missurica (Raf) Shinners—SSBC; A; 4G359

Chamaesyce serpens (Kunth) Small—SSBC; A; 46335

Croton texensis (Klotzsch) Mull.Arg—QHSC; A; 4C227

Euphorbia marginata Pursh—DAUP, SSBC; A; 4C304

Euphorbia spathulata Lam.—SSCP; A; 4-085

Stillingia sylvatica Garden ex L.—SSBC; P; 4C163

Tragia ramosa Torr.—SSCP; P; 4355

Fabaceae

Amorpha canescens Pursh—SSBC; P; 4C150

Amorpha fruticosa L—DAFL; P; 4C160

Astragalus gracilis Nutt.—SSBC; P; 4C132

Astragalus lotiflorus Hook.—SSBC; P; 4C-007

stragalus missouriensis Nutt.—SSBC; P; 44-008

Astragalus mollissimus Torr.—SSBC; P; 4445

Astragalus plattensis Nutt.—SSBC; P; 4C133

Baptisia australis (L.) R.Br. ex Aiton—SSBC; P; 4C 044

Baptisia bracteata Muhl. ex Elliot —SSBC; P; 4C106

Se d (Torr. & A. Gray) Fisher—SSBC; P; 4323

Dalea aurea Nutt. ex Pursh—SSBC; P; 4C212

Dalea d Michx. ex Willd.—SSBC; P; 4C265

Dalea enneandra Nutt.—SSBC; P; 4220

Dalea lanata Spreng—DAFL; P; 44284

Dalea purpurea Vent.—SSBC; P; 4G213

Dalea villosa (Nutt.) Spreng.—DAFL; P; 4C283

esmanthus illinoensis (Michx.) MacMil.. ex B. L. Rob. &

Fernald—DAFL; P; 46313

Desmodium sessilifolium (Torr.) Torr. & A. Gray—SSBC; P;

-369

Glycyrrhiza lepidota Pursh—DAFL; P; 4C184

Gymnocladus dioicus (L.) K. Koch—OQOMJV; P; 4406

ud miniata Ortega—OHSC; P; 4C444

pedeza capitata Michx.—SSBC; P; 4C440

D stuevei Nutt.—SSBC; P; 4C381

Melilotus officinalis (L) Lam. *— DAFL, DAUP; A; 4G309

Mimosa borealis A. Gray—SSBC; P; 44161

Mimosa nuttallii (DC.) B.L. Turner—SSBC; P; 4-062

Oxytropis lambertii Pursh var. artciulata (Greene) Barneby—

SSBC; P; 4C-426

Oxytropis lambertii Pursh var. lambertii—SSBC; P; 4C134

Pediomelum dad. (Pursh) Rydb.—SSBC; P; 4C156

Pediomelum digitatum (Nutt. ex Torr. & A. Gray) Isely—SSBC;

Pediomelum linearifolium (Torr. & A. Gray) J. W. Grimes—SSBC;

Psoralidium tenuiflorum (Pursh) Rydb.—SSBC; P; 4C-263

Robinia pseudoacacia L.—OMJV; P; 4504

Senna marilandica (L) Link—QMUJV; P; 46342

Sophora nuttalliana B.L. Turner—SSBC; P; 4425

Strophostyles leiosperma (Torr. & A. Gray) Piper—SSBC; A;

Vicia americana Muhl. ex Willd.—SSBC; P; 44096

Vicia ludoviciana Nutt.—SSBC; A; 4C113

Fagaceae

fal, Dat M ID hi Pr

Journal of of Texas 1(1)

Quercus havardii Rydb.—OQOHSC; P, 4C046

Quercus muehlenbergii Engelm. -OMJV; P; 4C-034

Quercus stellata Wangenh.—QHSC; P; 46-446

Fumariaceae

Corydalis micrantha (Engelm. ex A. Gray) A. Gray—SSBC; A;

Gentianaceae

Eustoma exaltatum (L.) Salisb. ex G.Don—DAFL; P; 4297

Geraniaceae

Geranium carolinianum L.—DAUP; A; 4-087

Grossulariaceae

Ribes aureum Pursh var. villosum DC.—QMJV; P; 4C 014

Hydrophyllaceae

Nama stevensii C.L. Hitchc.—SSCP; A; 44142

Phacelia integrifolia Torr.—SSCP; A; 4G125

Juglandaceae

Juglans microcarpa Berl.—QMJV; P; 4C205

Krameriaceae

Krameria lanceolata Torr.—SSBC; P; 4C148

menia eae

ma drummondii Benth.—SSBC; P; 4C222

uL E hispida T DAUP, SSCP; A; 4C-090

Monarda clinopodioides A. Gray—SSCP; A; 4C27

onarda punctata L. var. occidentalis (Epling) Palmer &

Steyerm.—SSBC; A; 4C254

Salvia azurea Michx. ex Lam.—SSBC; P; 4C272

Scutellaria drummondii Benth.—SSBC; P; 4C100

Scutellaria resinosa Torr.—SSBC; P; 4C102

Teucrium canadense L—QMJV, WETL; P; 4C182

Linaceae

Linum lewisii Pursh var. lewisii—SSBC; A; 4C006

Linum rigidum Pursh—SSBC; A; 4C-120

Loasaceae

Mentzelia decapetala (Pursh ex Sims) Urb. & Gilg ex Gilg—

SSBC; P; 46370

Mentzelia multiflora (Nutt.) A. Gray—QHSC; A; 4370

Mentzelia nuda (Pursh) Torr. & A. Gray—SSBC; P; 4G324

Mentzelia oligosperma Nutt. ex Sims—SSCP; P; 4C193

Malvaceae

Callirhoe involucrata (Torr. & A. Gray) A. Gray—SSBC; P; 4G

Sphaeralcea coccinea (Nutt.) Rydb.—SSBC; P; 4-079

Molluginaceae

Mollugo verticillata L—DAUP, WETL; A; 4C:308

Moraceae

Maclura pomifera (Raf) Schneid.—QOMJV; P; 4C196

Morus alba L*—QOMJV; P; 4-259

Morus rubra L.—QMJV; P; 4C422

Nyctaginaceae

Mirabilis linearis (Pursh) Heimerl—SSBC; P; 4C257

Mirabilis nyctaginea (Michx.) MacMil..—QOMJV; P; 4C179

Hoagland and Buthod, Flora of Four Canyons Preserve, Oklahoma

Onagraceae

Calylophus berlandieri Soach—SSBC; P; 4-041

Calylophus hartwegii (Benth.) P.H.Raven—SSBC; P; 4C 071

Calylophus serrulatus (Nutt.) P.H.Raven—SSBC; P; 4097

Gaura coccinea Nutt. ex Pursh—SSBC; P; 4C126

Gaura longiflora Spach—DAUP; A; 4C241

Gaura villosa Torr—QHSC; P; 4C410

Oenothera grandis (Britt) Smyth —DAUP, OHSC; P; 4C 089

Oenothera jamesii Torr. & A. Gray—WETL; B; 4C387

Oenothera laciniata Hill—DAUP, SSBC; A; 4-059

Oenothera macrocarpa Nutt.—SSBC; P; 4C 050

Oenothera rhombipetala Nutt. ex Torr. & A. Gray—QHSC;

Stenosiphon linifolius (Nutt. ex James) Heynh.—SSBC; P;

Oxalidaceae

Oxalis stricta L—DAUP, SSBC; P; 4C 067

Oxalis violacea L.—SSBC; P; 4-058

Papaveraceae

Argemone polyanthemos (Fedde) G.B. Ownbey—SSBC; A;

C-061

Phytolaccaceae

Phytolacca americana L.—OMJV; P; 44198

Plantaginaceae

Plantago patagonica Jacq.—DAUP, SSBC; A; 4C111

Plantago rhodosperma Dcne.—DAUP, SSBC; A; 4C 114

Polemoniaceae

Ipomopsis longiflora (Torr.) V.E.Grant— OHSC, SSBC; A; 4G

395

Polygalaceae

Polygala alba Nutt.—SSBC, SSCP; P; 4C-055

Polygonaceae

Erigonum annuum Nutt.—SSBC; A; 4-306

Erigonum longifolium Nutt.—SSBC; P; 46-032

Polygonum amphibium L—WETL; P; 4C262

Polygonum aviculare L*—DAUP; A; 46-328

Polygonum lapathifolium L—WETL; A; 4209

sylvancium L.—WETL; A; 4C360

Polygonum ramosissimum Michx.—DAUP, WETL; A; 4322

Polygonum pen

rur d qe

Portulacaceae

Portulaca halimoides L.—DAUP; A; 4-436

| | L.—DAUP; A; 44358

D +

POPLUIUCU VICIULCO

Primulaceae

Androsace occidentalis Pursh—DAUP, SSBC; A; 46021

Samolus ebracteatus Kunth—DAFL; P; 4C318

natos

Anemon

ina Walter—SSBC; P; 4025

Delphinium ODIT Walter—SSBC; P; 4415

Ranunculus sceleratus L.—WETL; A; 4G091

Rhamnaceae

Ceanothus herbaceous Raf.—SSBC; P; 4C076

Rosaceae

Prunus angustifolia Marsh—QHSC, SSBC; P; 4C-005

661

Prunus gracilis Engelm. & A. Gray—QHSC; P; 4G 443

Pyrus communis L*—OMJV; P; 4C437

Rubiaceae

Cephalanthus occidentalis L.—WETL; P; 4C183

Galium aparine L.—DAUP, QMJV; A; 44190

Hedyotis nigricans ene Fosberg

Rutaceae

Ptelea trifoliata L.—SSBC; P; 4C047

Salicaceae

Populus deltoides Bartram ex Marsh—DAFL, WETL; P; 4G

Salix exigua Nutt.—DAFL, WETL; P; 4C108

Salix nigra Marsh—DAFL, WETL; P; 4C107

Santalaceae

Comandra umbellata (L.) Nutt.—SSBC; P; 4C109

Sapindaceae

Sapindus saponaria L. var. drummondii (Hook. & Arn.) L.D.

Benson—OMAJV; P; 4C317

Sapotaceae

Sideroxylon lanuginosum Michx.—QMJV; P; 4C200

Scrophulariaceae

Agalinis aspera (Douglas ex Benth.) Britt—SSBC; A; 4:386

Castilleja purpurea (Nutt.) G.Don var. citrina (Pennell)

Shinners—SSCP; P; 4C 081

Castilleja sessiliflora Pursh—SSBC; P; 46-023

Nuttallanthus texanus (Scheele) D.A.Sutton—SSBC; A; 4G

Penstemon albidus Nutt.—SSCP; P; 4C083

Penstemon buckleyi Pennell—SSBC; P; 4C115

Penstemon cobaea Nutt.—SSBC; P; 4C 049

Veronica arvensis L*—DAUP; A; 4C 080

Veronica peregrina L.—SSBC; A; 4C116

Solanaceae

Chamaesaracha conioides (Moric. ex Dunal) Britt—DAUP,

SSBC; P; 4C009

Datura stramonium L.—OMAUJV; A; 4C423

Physalis longifolia Nutt.—DAUP; P; 4C234

DAUP; P; 4C117

Quincula lobata (Torr.) Raf..—DAUP, SSBC; P; 4C234

Solanum elaeagnifolium Cav.—DAUP, OMJV, SSBC; P; 4G

045

Solanum dimidiatum Raf..—DAUP, SSBC; P; 4C269

Solanum rostratum Dunal—DAUP; A; 4C206

Tamaricaceae

Tamarix chinensis Lour *—DAFL; P; 4C 048

Ulmaceae

Celtis laevigata Willd.—OMJV; P; 4C215

Ulmus americana L.—OMAJV; P; 4C042

Ulmus pumila L*—QOMJV; P; 4442

Ulmus rubra Muhl.—QMJV; P; 44396

Urticaceae

Parietaria penslvanica Muhl. ex Willd.—OMJV; A; 46202

Verbenaceae

Glandularia pumila (Rydb.) Umber—DAUP, SSBC; A; 4C-086

662

Phyla nodiflora (L.) Greene—DAFL, WETL; P; 4C221

Verbena bracteata Lag. & Rodr.—DAUP; A; 4C140

Verbena stricta Vent.—DAUP; P; 4C235

Vitaceae

Cissus trifoliata (L.) L.—OMJV; P; 4065

Parthenocissus quinquefolia (L.) Planch.—QMJV; P; 4C057

Vitis acerifolia Raf —QMJV; P; 4C074

Vitis rupestris Scheele—OMJV; P; 4C074

Zygophyllaceae

Tribulus terrestris L.*—

DAUP, QHSC; A; 4C-367

MAGNOLIOPHYTA—LILIOPSIDA

Agavaceae

Yucca glauca Nutt.—DAUP, QHSC, SSBC; P; 4C146

Commelinaceae

Commelina erecta L.—DAUP, SSBC; P; 4C226

Tradescantia occidentalis (Britt) Smyth—SSBC; P; 4C137

Tradescantia ohiensis Raf —SSBC; P; 4C162

Cyperaceae

Carex gravida Bailey—WETL; P; 4231

Carex festucacea Schkuhr ex Willd.—WETL; P; 4287

Cyperus schweinitzii Torr.—SSBC; P; 4C204

Eleocharis erythropoda Steud.—DAFL, WETL; P; 4168

Eleocharis montevidensis Kunth—DAFL, WETL; P; 4C167

Eleocharis obtusa (Willd.) Schult.—DAFL, WETL; A; 4C223

Eleocharis tenuis (Willd.) Schult. var. verrucosa (Svenss.)

Svenss.—DAFL, WETL; P; 4C168

Schoenoplectus pungens (Vahl) Palla—DAFL, WETL; P; 4G

101

Iridaceae

Sisyrinchium angustifolium P.L.Mill—SSBC; P; 4C-082

Juncaceae

Juncus brachyphyllus Wiegmann—DAFL, WETL; P; 4G224

Juncus interior Wiegmann—DAFL, WETL; P; 4-169

Juncus torreyi Coville—DAFL, WETL; P; 4:344

Liliaceae

Allium drummondii Regel—SSBC; P; 46012

Allium perdulce SV.Fraser—SSBC; P; 4-077

Androstephium caeruleum (Scheele) Greene—SSBC; P; 4C

Najadaceae

Naja guadalupensis (Spreng.) Magnus—WETL; A; 4350

Poaceae

Aegilops cylindrica Host*—DAUP; A; 46053

Agrostis hyemalis (Walter) B.S.P.—WETL; P; 4-348

Andropogon glomeratus (Walter) B. DAFL; P; 4-300

Andropogon hallii Hack.—SSBC; G, P; 4C311

Add iE RD E L.—DAUP, OHSC; A; 4C-389

tida oli a Michx.—DAUP, OHSC, SSBC; A; 4:382

Wc m Nutt. var. longiseta (Steud.) Vasey—DAUP,

+P; 4

Aristida purpurea Nutt. var. purpurea—DAUP, SSBC; P; 4G

2/7

Bothriochloa laguroides (DC.) Herter—DAUP, SSBC; P; 4C278

fal, Dat M ID hi ee i

Journal of

of Texas 1(1)

Bothriochloa saccharoides (Sw.) Rydb.—DAUP, SSBC; P; 4G

Bouteloua curtipendula (Michx.) Torr. SSBC; P; 4C195

Bouteloua gracilis (Willd. ex Kunth) Lag. ex Griffiths—DAUP,

QHSC, SSBC; P; 4G333

Bouteloua hirsuta Lag.—SSCP; P; 44195

Bromus catharticus Vahl* —DAUL; A; 4C-068

Bromus japonicus Thunb. ex Murr*—DAUL; A; 4C261

Bromus tectorum L. *—DAUL, SSBC; A; 4-064

Buchloe dactyloides (Nutt.) Engelm. d

Calamovilfa gigantea (Nutt.) Scribn. € Merr—DALF; P; 4G

302

Cenchrus spinifex Cav.—DAUP, QHSC; P; 4C255

Chloris verticillata Nutt.—DAUP; P; 4C207

Cynodon dactylon (L.) Pers*—DAUP, DAFL; P; 4C194

Dichanthelium acuminatum (Sw.) Gould & C.A. Clark var.

fasiculatum (Torr.) Freckmann—SSBC; P; 4C233

Dichanthelium clandestinum (L.) Gould —OHSC, SSBC; P;

4C122

Dichanthelium villosissimum (Nash) Freckmann var. praecocius

(A.S. Hitchc. € Chase) Freckmann—SSBC; P; 46332

Distichlis spicata (L.) Greene—DAFL; P; 46151

Echinochloa muricata (P.Beauv.) Fernald—WETL; A; 4C253

Elymus canadensis L.—DAUP, QMJV, SSBC; P; 4C230

Elymus virginicus L—DAUP, QMJV, SSBC; P; 4C-405

Eragrostis cilianensis (All) Vignet ex Janch.*—DAUP; A; 4G

390

Eragrostis secundiflora J.Presl—DAUP, SSBC; P; 4C236

Eragrostis spectabilis (Pursh) Steud.—DAUP, SSBC; P; 4337

Eragrostis trichodes (Nutt.) Wood—DAUP, QHSC; P; 4G383

Erioneuron pilosum (Buckl.) Nash—QHSC, SSBC; P; 4C191

Hordeum jubatum L.—DAFL, DAUP; P; 4G175

Hordeum pusillum Nutt.—DAUP; A; 4C-069

Leptochloa fusca (L.) Kunth—WETL; A; 4C431

Lolium arundianceum (Schreb.) Darbysh.*—DAUP; P; 4C-427

Monroa squarrosa (Nutt.) Torr. —OHSC, SSBC; P; 4C211

Muhlenbergia asperifolia (Nees & Meyen ex Trin) Parodi—

DAFL; P; 4C408

Muhlenbergia bushii Pohl—OMJV; P; 4C202

Muhlenbergia racemosa (Michx.) B.S.P.—OMJV; P; 4C420

Panicum capillare L—DAUP, SSBC; A; 4C314

Panicum hallii Vasey—SSBC; P; 4-392

Panicum obtusum Kunth—DAFL, SSBC; P; 4C199

Panicum virgatum L—DAFL, SSBC; P; 4325

Pascopyrum smithii (Rydb.) A. Lóve—DAUP; P; 4C171

Paspalum setaceum Michx.—QHSC, SSBC; P; 4225

Phalaris caroliniana Walter—-WETL; A; 4-138

Phragmites australis (Cav.) Trin. ex Steud.—DAFL; P; 4-438

Poa arachnifera Torr.—DAUP, SSBC; P; 4C13

Polypogon monspeliensis (L.) Desf*—DAFL, WETL; G, A;

4C-139

Saccharum ravennae (L) L* DAFL, WETL; G, P; 4301

Schizachyrium scoparium (Michx.) Nash—QHSC, SSBC; P;

Setaria parviflora (Poir.) Kerguélen—DAFL, DAUP; P; 4-338

Sorghastrum nutans (L) Nash—SSBC; P; 46357

Sorghum halepense (L.) Pers *—DAUP; P; 4C177

Spartina pectinata Bosc ex Link—DAFL; P; 4C315

Hoagland and Buthod, Flora of Four Canyons Preserve, Oklahoma 663

Sphenopholis obtusata (Michx.) Scribn.—WETL; A; 4G153 Vulpia octoflora (Walter) Rydb.—DAUP; A; 4C130

Sporobolus cryptandrus (Torr) A. Gray—QHSC, SSBC; P; Vulpia sciurea (Nutt.) Henry—DAUP; G, A; 4C-131

4C391

Sporobolus neglectus Nash—DAUP, OHSC; A; 4-399

Tridens albescens (Vasey) Wooten & Standl.—SSBC, OHSC;

P: 4C380 Typhaceae

Tridens flavus (L.) A.S. Hitchc.—QMJV, SSBC; P; 4C310 Typha domingensis Pers.—DAFL, WETL; P; 4C249

Smilacaceae

Smilax tamnoides L.—OMJV; P; 4C201

ACKNOWLEDGMENTS

We thank Priscilla Crawford, Shannon Hall, and Lacy Brookshire for field assistance. This project was

funded by a grant from The Nature Conservancy. The comments of one anonymous reviewer are greatly

appreciated.

REFERENCES

Branson, C.C. and K.S. Johnson. 1979. Generalized geologic map of Oklahoma. In: K.S. Johnson et al., eds. Geology

and Earth resources of Oklahoma. Oklahoma Geological Survey, Norman.

Cote, E.L., AJ. CoNRADI, and C.E. RHOADS. 1961. Soil survey of Ellis County, Oklahoma. United States Department of

Agriculture, Washington, DC.

ConntLL, D. S. and M. C. Johnston. 1970. Manual of the vascular plants of Texas. Texas Research Foundation,

Renner.

Curtis, N.M. and W.E. Ham. 1979. Geomorphic provinces of Oklahoma. In: K.S. Johnson et al., eds. Geology and

Earth resources of Oklahoma. Oklahoma Geological Survey, Norman.

Duck, L.G., and J.B. FLETCHER. 1943. A game type map of Oklahoma. A survey of the game and furbearing animals

of Oklahoma. Oklahoma Department of Wildlife Conservation, Oklahoma City.

GREAT PLAINS FLORA ASSOCIATION. 1986. Flora of the Great Plains. University Press of Kansas, Lawrence.

Groves, C.R., M.L. Ke, and T.F. BreDEN. 1995. Natural Heritage Programs: public-private partnerships for biodiversity

conservation. Wildlife Soc. Bull. 23:784-790.

HOAGLAND, B.W. 2000. The vegetation of Oklahoma: a classification of landscape mapping and conservation

planning. SouthW. Naturalist 45:385-420.

HOAGLAND, B.W. and A.K. BuTHop. 2003. Vascular flora of the Keystone Wildlife Management Area, Creek, Pawnee,

and Osage Counties, Oklahoma. Oklahoma Native Pl. Rec. 3:23-37.

HOAGLAND, B.W. and A.K. ButHop. 2004. Vascular flora of Hugo Lake Wildlife Management Area, Choctaw County,

Oklahoma. SouthE. Naturalist 3:701-714.

HOAGLAND, B.W. and A.K. ButHop. 2005a. Vascular flora of a gypsum dominated site in Major County, Oklahoma.

Proc. Oklahoma Acad. Sci. 85:1-8.

HOAGLAND, B.W. and A.K. ButHop. 2005b. Vascular flora of a site along the Arkansas River, Pawnee County, Oklahoma.

Oklahoma Native Pl. Rec. 5:61-72.

HOAGLAND, B.W. and F.L. JoHNson. 2001. Vascular flora of the Chickasaw National Recreation Area, Murray County,

Oklahoma. Castanea 66:383-400.

HOAGLAND, B.W. and F.L. JoHNson. 2004a. Vascular flora of Chouteau Wildlife Management Area, Wagoner County,

Oklahoma. Oklahoma Native Pl. Rec. 4:30-39.

Hoagland, B.W. and F.L. Johnson. 2004b. Vascular flora of Love Valley Wildlife Management Area, Love County,

Oklahoma. Proc. Oklahoma Acad. Sci. 83:47-62.

HOAGLAND, B.W. and F.L. JoHnson. 2004c. Vascular flora of Red Slough and Grassy Slough Wildlife Management

Areas, Gulf Coastal Plain, McCurtain County, Oklahoma. Castanea 69:284-296.

HOAGLAND, B.W. and F.L. JoHNson. 2005. Vascular flora of the Deep Fork River in Okmulgee, Creek, and Okfuskee

Counties. Publ. Oklahoma Biol. Survey 6:15-29.

HOAGLAND, B.W. and K. Watuck. 2003. Vascular flora of Oologah Wildlife Management Area in Nowata County,

Oklahoma. Proc. Oklahoma Acad. Sci. 83:47-62.

664 Journal of the Botanical R h Institute of Texas 1(1)

HOAGLAND, B.W., A.K. Buruob, and W. Eustws. 2004. Vascular flora of Washita Battlefield National Historic Site, Roger

Mills County, Oklahoma. Sida 21:1187-1197.

HOAGLAND, B.W., P. CRAWFORD-CALLAHAN, P. CRAWFORD, and F.L. JOHNSON. 2004. Vascular flora of Hackberry Flat, Frederick

Lake, and Suttle Creek, Tillman County, Oklahoma. Sida 21:429-445.

HOAGLAND, B.W. and F.L. Johnson. 2001. Vascular flora of the Chickasaw National Recreation Area, Murray County,

Oklahoma. Castanea 66:383-400.

HOAGLAND, B.W., A. Burnop, |. BUTLER, P. CALLAHAN-CRAWFORD, W. ELisens, A. Upasi, and R. Tye. 2006. Oklahoma vascular

plants database. (www.biosurvey.ou.edu.). University of Oklahoma, Norman.

Hunt, C.B. 1974. Natural regions of the United States and Canada. W. H. Freeman, San Francisco.

OKLAHOMA CLIMATOLOGICAL SURVEY. 2006. Oklahoma climatological data. (www.ocs.ou.edu/). University of Oklahoma,

Norman.

OKLAHOMA NATURAL HERITAGE INVENTORY (ONHI). 2006. Oklahoma Natural Heritage inventory working list of rare Okla-

homa plants. (www.biosurvey.ou.edu/publicat.html). University of Oklahoma, Norman.

Parmer, M.W, G.L. Wac, and P. Neat. 1995. Standards for the writing of floras. Bioscience 45:339-345.

TayLor, RJ. and C.S. Taytor. 1991. An annotated list of the ferns, fern allies, gymnosperms, and flowering plants of

Oklahoma. Southeastern Oklahoma State University. Durant.

TREWARTHA, G.T. 1968. An introduction to Climate. McGraw-Hill, New York. USDA.

USDA-NRCS 2006. The PLANTS database. (plants.usda.gov/plants). National Plant Data Center, Baton Rouge,

LA.

WATERFALL, U.T. 1973. Keys to the flora of Oklahoma. Published by the author, Stillwater, OK.

DHEWVASCUDPARSSEORSOPIEHERBEGDANDONEBD RICE FIELD

GEORGETOWN SOU TE CAROLINA» A 30 YEAR COMPARISON

Richard Stalter John Baden Dwight Kincaid

Dept. of Biological Sciences U.S. Army Corp of Engineers Dept. of Biological Sciences

St. John's University Wilmington, North Carolina 28403, U.S.A. Lehman College CUNY

Jamaica, New York 11439, U.S.A. Bronx, New York 10468, U.S.A.

ABSTRACT

The vascular plant species at th land 1 rice fields, Alderly, Airport, and Thousand Acre Rice Field at the Belle W. Baruch Institute

for Marine Biology and Coastal Research, Georgetown, South Carolina were sampled and compared over a 39 year period, 1967-2006.

One hundred twenty-four species in 91 genera and 44 families were identified at the three marshes, including eight non-native spe-

cies. The Institute experienced a category 4 hurricane, Hurricane Hugo, on September 22, 1989. The number of vascular plant species

decreased at the two least saline marshes, by 32% at Alderly and by 30% at Airport, immediately after Hugo. The number of species at

"ep BI

Alderly increased when surveyed in 2002-2006, while there was no change in the number of species at the Airport marsh since Hugo.

There was little change in the number of species at Thousand Acre Rice Field, the most saline marsh, during the 39 year study. Phragmites

has recently invaded Airport Marsh and Alderly and may out compete and exclude native vascular plant species in the future.

Key Wonps: vascular flora, South Carolina, abandoned rice fields, brackish marshes, Hurricane Hugo

RESUMEN

Se muestrearon y compararon durante un periodo de 39 años, 1967-2006, las plantas ] de tr d bandonad

Alderly, Airport, y Thousand Acre Rice Field en el Belle W. Baruch Instituto de Biologia Marina e TC Costera, Georgetown,

Carolina del Sur. Se identificaron ciento veinticuatro especies de 91 géneros y 44 familias en los tres lugares, incluyendo ocho especies

no nativas. El Instituto sufrió un huracán de categoría 4, Huracán Hugo, el 22 de septiembre de 1989. El número de plantas vasculares

decreció en las dos últimas lagunas salinas, en un 32% en Alderly y en el 30% en Airport, inmediatamente después del Hugo. El nú-

mero de especies en Alderly aumentó cuando fue revisado en 2002-2006, mientras que no hubo cambio en el nümero de especies en

la laguna de Airport desde el Hugo. Hubo pocos cambios en el nümero de especies en Thousand Acre Rice Field, la laguna más salina

durante los 39 años de estudio. Phragmites ha invadido recientemente Airport Marsh y Alderly y puede que comita y excluya especies

vasculares nativas en el futuro.

INTRODUCTION

The objective of the present study was to compile a vascular flora of three abandoned rice fields and to de-

scribe changes in the flora of the three brackish marshes in which they exist, on the Belle W. Baruch Institute

for Marine Biology and Coastal Research Georgetown, South Carolina, over a 39 year period, 1967-1971,

1987-1991 and 2002-2006. The marshes, Alderly, Airport, and the 1000 Acre Rice Field, were selected in

1967 by John Baden for a masters degree project when he was a graduate student at the University of South

Carolina (Baden 1971). Airport marsh, 62.9 hectares, 33.3467 N, 79.2488 W, was the smallest study site.

Alderly, 33.3544 N, 79.2439 W and the 1000 Acre Rice Field, 33.3029 N, 79.2514 W, are 173.3 and 248.6

hectares, respectively (Fig. 1).

The vascular flora of these marshes was first catalogued by Barry (1968) in relation to soil types during

the growing season of 1967-1968. The brackish marsh soils are acid, continually saturated, and very poorly

drained. Baden (1971) examined the effect of tidal flooding, salinity, pH, soil texture and organic content on

the composition of vascular plants in these three brackish marshes. Baden et al. (1975) found that salinity,

pH, texture, and organic matter were not important edaphic factors influencing the zonation of vegetation

in Alderly and the Airport Marshes.

Stalter and Baden (1994) compared the vascular flora of Airport, Alderly, and Thousand Acre Rice Field

in 1987-91 with the vascular flora at the same three marshes in 1967-68, concluding that there was little

change during the 20 year interval. The Jaccard indices of similarity were high (> 0.8) in pairwise parisons

J. Bot. Res. Inst. Texas 1(1): 665 — 677. 2007

666 Journal of the Botanical Research Institute of Texas 1(1)

PEE DEE

RIYER

GEORGETOWN

e OAM DEBIDUE

ISLAND

Í ISLAND

i». cp

Fic. 1. Location of three study sites, Alderly (A), Airport (AP), and Thousan d Acre Rice Field (TRF) at the Baruch Institute, South Carolina

Stalter et al., Flora of abandoned rice fields, South Carolina 667

of vascular plant species among marshes from 1967-68 to 1987—91. Of the species observed in 1968 and

1991, 91% were native to the United States. By comparison, native vascular plant species comprised 72.5%

of the flora at Fort Sumter, South Carolina (Stalter & Lamont 1993); 65.696 of the flora at Cape May Point

State Park, New Jersey (Sutton et al. 1990) and 76.396 of the flora at Assateague Island, Virginia (Stalter &

Lamont 1990).

Stalter (1973) studied the factors influencing the distribution of vascular plant species in the Cooper

River estuary, South Carolina, in July 1971. The marsh vegetation was sampled by transects at 23 sites from

Sullivan Island at the mouth of the Cooper River to freshwater marshes well inland. The transects were laid

out perpendicular to the Cooper River and marked at the elevated end by a concrete post or, if available,

by a Coast and Geodetic Survey bench mark. Forty nine vascular plant species were identified in Stalter's

(1973) study. Zonation of vegetation was pronounced in salt marshes. Spartina alterniflora was the most flood

tolerant taxon and occupied the widest range in elevation. Zonation of vascular plant species in the brackish

marshes (salinity 1.0 ppt to 15 ppt) was less pronounced than in the salt marshes. Spartina cynosuroidies,

Juncus roemerianus and Scirpus validus occupied the most flood prone zone of the brackish marshes. Fresh-

water marshes (those with salinity values less than 1.0 ppt) were populated by Alternanthera philoxerodies,

Ludwigia spp., Pontederia cordata, Saururus cernuus and Zizaniopsis miliacea. All of the aformentioned species

at Cooper River were present in the abandoned rice field marshes at Baruch.

Eleuterius (1972) described the marshes of Mississippi based on extensive field work in 1968 and

1969. Vegetation was sampled by line transects and list-count quadrats at 19 stations in all the estuarine

marshes of Mississippi. He reported over 300 species of vascular plants in Mississippi marshes; no species

list was published (Eleuterius 1972). He reported that there was a greater diversity of vascular plant species

in freshwater marshes than in brackish or salt marshes; these results are similar to those in the abandoned

rice fields of the Baruch Institute. Distribution of species at the Baruch brackish marshes was also similar to

that reported by Stalter (1973) at the Cooper River. Eleuterius also noted the effect of salt water on vegetation

during a growing season. Crinum americanum and Iris virginica, were present in brackish marshes in April

when water salinity value were low. By June, these same species, “were dead or disappeared.” Crinum and

Iris virginica were growing vigorously and producing flowers in freshwater marshes.

Stalter (1972) conducted a survey of the summer and fall flora at Brookgreen Gardens from July to De-

cember 1970. He reported 639 species of vascular plants though he did not describe the plant communities

at Brookgreen. Many of the taxa reported by Stalter (1972) at the brackish marshes at Brookgreen, notably

Juncus and Spartina, were also present in brackish marshes at the Baruch Institute.

Stalter (1971) reported the summer and fall flora of Huntington Beach State Park, Georgetown County,

South Carolina. Included in the 321 taxa were 66 wetland species, though not all of these species were found

at the abandoned rice fields at the Baruch Institute. Common to both sites are species of Carex, Eupatorium,

Hydrocotyle, Juncus, Rhynchospora, Sagittaria, Spartina, Taxodium, and Typha.

A study at the Yawkey Center, a site adjacent to the Baruch Institute, was conducted by Nelson from

early spring 1990 to September 1992. Ten plant communities were described by Nelson, in an unpublished

inhouse document. One of his communities was old rice fields. Nelson stated that these sites, depending on

flood history and salinity may be highly complex and botanically diverse. Nelson cites 21 taxa specifically

found in the adandoned rice fields. Most of these taxa were also identified at the abandoned rice fields at

the Baruch Institute.

History

The marshes in the study are located on Waccamaw Neck, bordering Winyah Bay (Fig. 1). In 1718, George

II granted Lord Carteret 7500 hectares that composed the original Hobcaw Barony. Carteret sold his land to

a land speculator, John Roberts in 1735, who subdivided the property and sold it to developers. Ultimately,

the area was settled and the tidal wetlands bordering the Waccamaw River were diked and drained for rice

production. By the beginning of the Civil War (1861), 13 major rice plantations at Hobcaw were producing

approximately a half a million pounds of rice annually (Porcher 1976).

668 Journal of the Botanical R h Institute of Texas 1(1)

At the end of the Civil War (1865), plantation owners lost their slaves who provided the labor that

sustained the rice industry (Porcher 1976). The few rice growing plantations that survived were devastated

by a series of hurricanes in the 1890s and early 1900s (Baden 1971). The most severe hurricane, the storm

of 1893, rivaled Hurricane Hugo (1989) in intensity and destruction. The devastating hurricane of 1893 plus

additional hurricanes and competition from rice growers in the Gulf States terminated the rice industry in

South Carolina by the early 20th century.

Many took advantage of the demise of the coastal plantation system. South Carolinian Bernard Baruch

purchased 7,500 hectares along Waccamaw Neck between 1905 and1907, as a hunting preserve. Baruch

(1957), claimed the salt marshes (at Hobcaw Barony), “once offered the finest duck hunting in the United

States.”

After his daughter's demise, Bernard Baruch established the Belle Baruch foundation in his daughter's

honor as a natural laboratory for research and teaching. Today, two research institutes, one dealing with

marine science and affiliated with the University of South Carolina and the other with forestry and affiliated

with Clemson University, pursue scientific studies in the forest and marshes at Hobcaw and in the Atlantic

Ocean. The foundation was selected by NSF in the mid 1980s as a Long Term Ecological Research Site.

Climate

Coastal South Carolina has a subtropical climate. Average rainfall at Georgetown is 1,368 mm. August is

the wettest month averaging 180 mm while November is the driest month averaging 75.4 mm. Snowfall

is rare; 10 mm falls in an average year. July is the warmest month with an average temperature of 27.1 C.

January, the coldest month, has an average temperature of 8.7 C. The growing season averages 254 days

(Anonymous 1996).

Soils

Soils at Alderly are Levy silty clay loams. Levy soils are, “fine mixed superactive acid (pH 5.0—5.5) thermic

Typic Hydraquents. Organic content of the soils at Alderly ranged from 24-30% (Baden 1971). The average

mineral component at Alderly was sand (25%) silt (51.5%) and clay (23.5%). Table 3 reports the soil data

of Baden (1971).

Soils at the more saline Airport and 1000 Acre Rice Field are included in the Bohicket Series. These

soils are very poorly drained, very permeable soils that formed in marine sediments in tidal marshes. The

taxonomic class: Fine mixed Typic Sulfaquents. Soils at Airport and Thousand Acre Rice Field are acid in

reaction at the two sites with pH ranging from 5.0 to 6.0 and 5.0 to 5.5 at the two respective sites. The

organic content of soils at Airport marsh ranged from 25-2896; organic content of soils at Thousand Acre

Rice Field from 19-33%. Silt was the dominant soil fraction at both sites. Soil fractions at Airport were sand

(1896), silt (57.596) and clay (24.596). At the Thousand Acre Rice Field the fractions were 2496 sand, 41.296

silt and 34.896 clay (Table 3).

Elevation measurements within the marshes were made using a surveryors transit and stadia pole.

Transit measurements indicated that there was little difference in elevation across the abandoned rice fields

(Baden 1971).

Hurricane Hugo

Hurricanes have struck the South Carolina coast an average of once every 2.5 years in the 20th century

(Gentry 1971). The most notable during the course of this study was Hurricane Hugo in 1989 (Gardner et al.

1991), which was a “storm of the century” hurricane, causing catastrophic damage to upland forests at the

Baruch Institute (Blood et al. 1991; Gardner et al. 1991). Sustained winds of 122 km/m with gusts as high

as 150 km/hr were recorded in Charleston, 75 km south. Winds at Georgetown peaked at 112 km/hr. Many

trees were toppled and/or broken (Stalter & Baden 1994). Hugo's storm surge in the Thousand Acre Rice

Field, Airport and Alderly marshes was approximately four meters, depositing debris up to 0.5 meters at the

Thousand Acre and Airport marshes (Stalter and Baden 1994). The resulting raft of vegetation was colonized

by vascular plant species not normally associated with brackish marshes (Stalter and Baden 1994). These

Stalter et al., Flora of abandoned rice fields, South Carolina 669

were Cakile edentula, a dune species; Erectites hieracifolia, a disturbed site invader; Eupatorium capillifolium and

Glottidium vesicarum, a disturbed field associate; Panium amarum, an upper, rarely flooded salt marsh fringe

associate; Pinus taeda, an upland species; Triadica sebifera, an Asiatic tree that invades disturbed sites; Solidago

sempervirens, an occupant of coastal dunes and the upper, rarely flooded salt marsh fringe; and Strophostyles

helvola, a coastal dune associate (Stalter and Baden 1994). Thirteen years later, by 2002, the raft of “wrack”

had decomposed or washed away along with the aforementioned ephemeral vascular plants.

METHODS

The vegetation in three brackish marshes, Thousand Acre Rice Field, Airport and Alderly was sampled at

least once a month during the growing seasons beginning July 2002 and terminating in May 2006. The

flora was compared with the vascular flora reported by Baden et al. (1975) during the collecting seasons

of 1968-69 and Stalter and Baden (1994) during the growing seasons of 1987-91 (Appendix). Voucher

specimens collected in 1968-69, 1987-91 and 2002-06 were deposited in the herbarium at the University

of South Carolina, with the exception of taxa mailed to experts for verification. Accession numbers will be

assigned to the voucher specimens by Dr. John Nelson, Batson Herbarium, University of South Carolina.

The Appendix contains an inventory of the vascular plant species reported in 1968-69 by Baden et al.

(1975), the growing season of 1987-91 by Stalter and Baden (1994) and the present study 2002-06. Only

species that reproduce and persist longer than a single growing season without cultivation are tabulated in

Table 1. Non-native species are designated by an asterisk; non-native status follows Gleason and Cronquist

(1991) and Wunderlin (1998). Nomenclature presented in this flora agrees with Kartesz (1994). When dif-

ferences in nomenclature occur, the older name, listed in Radford et al. (1968) is listed as a synonym and

enclosed in brackets. The families, genera and species in the Appendix are listed alphabetically.

Salinity data presented by Baden et al. (1975) were combined with that recorded by Dr. Dennis Allen, Direc-

tor, Baruch Institute (2002 pers. comm.) and Conner and Inabinette (2005) over a twenty year period (Table 2).

Pairwise among the three sites, floristic similarity was measured by Jaccard coefficients (Kaufman and

Rousseeum 1990, Jaccard 1908), calculated from 2 x 2 contingency tables, and tested for significance by

approximate randomization (Noreen 1989). Each coefficient (test statistic) was compared to its sampling

distribution under the null hypothesis (Ho) of a random distribution of taxa across the three study sites.

These sampling distributions were achieved after 99,999 randomizations of the species labels in the JMP

(SAS) row-by-column data, with the observed data also considered as a randomization under Ho. P-values

were calculated in standard fashion as P = (NGE + D / (NS + 1) where NGE is the number of null coefficients

in the sampling distribution that were greater than or equal to the test statistic and with NS as the number

of randomizations performed (Manly 1997). For reference in Table 4, the mean of the null distribution is

given for each observed Jaccard coefficient.

RESULTS AND DISCUSSION

The vascular flora at the three brackish marshes of the Baruch Institute includes 124 species in 91 genera

within 44 families. Dicots (64 species) are more numerous than monocots (55 species) (Table 1). Eight spe-

cies, Alternanthera philoxeroides, Arundo donax, Chenopodium album, Echinochloa crus-galli, Murdania keisak,

Phragmites australis, Triadica sebifera and Verbena brasiliensis are non-native (Appendix). Seventy-one species

occurred exclusively at only one site, while only 26 species occurred at all three sites. Alderly, the least saline

marsh, contained the greatest number of species (110). Wass and Wright (1969) working in coastal wetlands

of Virginia, reported a greater species diversity in fresh water marshes than in salt water marshes. Airport

and Thousand Acre Rice Field had 54 and 39 species, respectively. Cyperus drummondii collected at Alderly

was the rarest species identified in this study. It has been collected only once before in South Carolina, in

the 1940s (Nelson, Director, Batson Herbarium, USC, 2003 pers. comm.). This is the first South Carolina

record of this taxon in nearly 60 years.

Feral pigs are a major source of soil disturbance at all marsh sites, especially at the intertidal zone and

670 Journal of the Botanical R h Institute of Texas 1(1)

TABLE 1. Summary of the vascular flora at three brackish marshes, Georgetown, South Carolina.

Ferns Conifers Dicots Monocots Total

Families 3 1 30 10 44

Genera S 1 59 cU 91

Species 4 ] 64 5D 124

Native Species 4 1 60 5] 116

Introduced species 0 0 4 4 8

TABLE 2. Water salinity at three brackish marsh sites. Data from Baden 1975, Stalter and Baden (1994), and Conner and Innabi-

nette (2005). Values at all sites were greater than 20 ppt during the Hurricane Hugo storm surge.

Study site Salinity, (ppt)

Thousand Acre Rice Field 0-23

Airport marsh 0-16

Alderly 0-18.5

TABLE 3. Organic content and soil fraction (96) at three abandoned rice fields. Data from Baden (1971).

Study Site Range of Organic 96 of Soil Fractions

Content (96) Sand Silt Clay

Thousand Acre 19-33 24.0 41.2 34.8

Rice Field

Airport 25-28 18.0 Ore 24.5

Alderly 24-30 25.0 SES 299

TABLE 4. Species similarity measured by Jaccard coefficients among three sites. Statistical significance determined by appoximate

randomization tests involving 99,999 randomizations of the species label in the row-by-column JMP (SAS) data file. The null,

mean Jaccard coefficients are from the sampling distribution of the Jaccard achieved after the randomizations.

Jaccard Species in

Site comparison coefficient Null mean P-value common/total

Thousand Acre Rice Field vs. Airport 0.388 0.124 0.00001 26/67

Thousand Acre Rice Field vs. Alderly 0.263 0:157 0.00019 21/118

Airport vs. Alderly 0.414 0.197 0.00001 48/116

upland border. Disturbance may enhance species diversity at the three marshes since disturbed areas may

be colonized by Chamaecrista (Cassia) and Chenopodium album as well as by true brackish marsh species,

e.g., Pluchea camphorata and P. foetida.

Ten woody species are reported at Alderly for the first time: Ilex cassine var. cassine, Ilex opaca, Itea vir-

ginica, Nyssa sylvatica var. biflora, Rosa laevigata, Rosa palustris, Salix caroliniana, Sambucus canadensis, Triadica

sebifera, and Viburnum nudum. These woody taxa occupy the upper wooded fringe of Alderly marsh that

experiences tidal flooding at the time of the new and full moon. While the aforementioned taxa are not true

marsh associates they all can be found in fresh water swamps (Radford et al. 1968).

Among the three study sites, there is great variability with respect to species composition, dominance

and diversity. The most saline marsh, Thousand Acre Rice Field, supports the fewest number of species,

was least affected by Hurricane Hugo, and is little changed in species composition over 39 years. Airport

marsh supports a large expanding population of Phragmites, a non-native species not observed here 39 years

Stalter et al., Flora of abandoned rice fields, South Carolina 671

ago. Alderly, the least saline marsh, contained the highest number of vascular taxa, 110. Alderly was most

impacted by Hurricane Hugo resulting in a loss of species diversity when sampled immediately after Hur-

ricane Hugo (Stalter and Baden 1994).

The two sites, Alderly and Airport, with 48 species in common, had the greatest floristic similarity

(0.414, P = 0.00001). Alderly, the least saline site, and the Thousand Acre site, the most saline site, had the

lowest floristic similarity (0.263, P = 0.00019, Table 4).

Species diversity was greatest at the least saline sites, Alderly and Airport, with 110 and 54 species,

respectively. Twenty six species occur at all sites while 71 occur at only one site. Seven species including salt

marsh associates Borrichia frutescens, Distichlis spicata, Limonium carolinianum, Spartina alterniflora and S. pat-

ens occur exclusively at Thousand Acre Rice Field, the most saline site. Ipomoea sagittata, a salt marsh fringe

species was also found exclusively at Thousand Acre Rice Field. The genera Borrichia, Distichlis, Ipomoea and

Limonium were found exclusively at Thousand Acre Rice Field. Baccharis halimifolia was the only woody species

occurring at all sites. Baccharis has been listed as a salt marsh associate (Gleason & Cronquist 1991; Radford et

al. 1968) yet it grows in profusion in North and South Carolina along 1-95, miles from salt marshes. Triadica

sebifera, an exotic, was observed in the upper intertidal Airport marsh. Iva frutescens and Borrichia frutescens

were the only woody salt marsh species found exclusively at the most saline Thousand Acre Marsh.

Alderly

The most notable change in species dominance at Alderly is the increase in cover of Phragmites australis,

a non-native species not present in 1968. Taxodium distichum was planted at this site in 1999 by Clemson

University researchers; seedling survival ranged from 0 to 7396 (Conner and Inabinette 2005). Several mature

T. distichum in the Waccamaw River, adjacent to the Alderly marshes, have died; these individuals were alive

in 1993 (Stalter & Baden 1994, Fig. 2).

Seedlings of Taxodium distichum from eight estuarine areas, from Mississippi to Virginia, were planted

at Alderly in 1999 (Conner and Inabinette 2005). Only seedlings from Louisiana, Alabama and Florida sur-

vived the severe drought of 2001 when salinity values peaked at 18.5 parts per thousand (ppt) November

30, 2001 (Table 2). Seedlings from Louisiana were the most salt tolerant as 7396 survived, while survival

rates for Florida and Alabama seedlings were approximately 2096 and 1496 respectively. Experimental data

from Conner and Inabinette (2005) suggest that there may be a wide range of salinity tolerance within

populations of Taxodium distichum.

Periodic wet and dry periods plus infrequent hurricane surges have affected salinity at the brackish

marshes during the 39 year study. The highest water salinity at Alderly occurred during Hurricane Hugo,

when salt water in excess of 20 ppt penetrated up to 5 km inland (Blood et al. 1991; Gardner et al. 1991).

“Normal” water salinity for the Alderly marshes is 1 ppt (Conner & Innabinette 2005). During a dry period

from May 2002 to August 2002 salinity values at Alderly were above 8 ppt peaking at 14.6 ppt and 16.3 ppt

in June and August, respectively. The high salinity values for Alderly followed the dry year of 2001, where

salinity values ranged from 4 to 6 ppt from April to October; salinity values peaked at 18.5 ppt on November

30, 2001. Salinity values were generally | ppt or lower from 2003 to the spring of 2004.

Common species at Alderly are Crinum americanum and Lilaeopsis chinensis which grow in profusion

around the edge of the marsh. Crinum is abundant around the boat dock today but nowhere else at Alderly.

Other common vascular plants are Alternanthera philoxeroides, Aster subulatus, Aster tenuifolias, Bidens laevis,

Phragmites australis, Pontederia cordata, Sagittaria falcata, Scirpus americanus, S. robustus, S. validus, Spartina

cynosuroides, Typha angustifolia and. T. domingensis.

Alderly may have been most affected by Hurricane Hugo’s storm surge that brought salt water and salinity

of up to 20 ppt to the marsh. In 1991, following Hurricane Hugo, 34 vascular plant species were identified.

Forty seven species were reported in 1968 (Baden 1971). The latest survey 2002-2006, was conducted over

a time of relatively high water salinity followed by a wet period when salinity values were less than 1 ppt.

High vascular plant diversity in 2002-06 (86 species) compared to the studies of 1968 and 1991 may be

a function of varying water salinity (Table 2). Fight woody taxa, those experiencing tidal flooding during

672 Journal of the Botanical R h Institute of Texas 1(1)

Fi) Y At Liege ah ttha TH NA Dian |

new and full moon tides, reported earlier in the paper, are reported at Alderly for the first time. If the eight

woody taxa were excluded from the list, the 78 vascular plant species at Alderly 2002-2006 would still be

significantly higher than the 47 species reported at Alderly in 1968. A total of 105 species have been identi-

fied at Alderly during the 39 year study.

Thousand Acre Rice Field

Water salinity is highest in the Thousand Acre Rice Field (Table 2). Common species observed here were

Spartina cynosuroides, Cladium jamaicense, Juncus roemerianus, Scirpus robustus, Aster tenuifolius, A. subulatus,

Lilaeopsis chinensis, Scirpus spp. and Typha spp. The Thousand Acre Rice Field also supports salt marsh spe-

cies, notably Spartina alterniflora, S. patens, Distichlis spicata and Aster tenuifolius. Spartina alterniflora borders

the creeks and ditches that meander through this marsh and which experience flooding during every tide.

Over half of the bald cypress, Taxodium distichum, at Thousand Acre Rice Field have died since the marsh

was inundated by Hurricane Hugo’s storm surge in 1989. The severe drought of 2001 and “high” salinity

may also have contributed to the demise of Taxodium (Fig. 2).

Thousand Acre Rice Field supports the fewest number of vascular plant species. Twenty three taxa were

reported in 1968, while 28 were identified here following Hurricane Hugo and 24 in 2006. The brackish plant

species at the Thousand Acre Rice Field were little affected by Hurricane Hugo's storm surge. Disturbance

by feeding pigs opening new habitats may account for the increase in number of plant species in 2006.

Prominent species in the Thousand Acre Rice Field were Juncus roemerianus, Scirpus robustus, S. ameri-

canus, Spartina cynosuroides, Typha angustifolia and Rhynchospora miliacea. Barry (1968) reported 12 vascular

plant species at Thousand Acre Rice Field. The smaller number of species noted by Barry (1968) compared

with 24 species collected during 2002-06 and 39 species over the 39 year study period, represents differ-

ences in sampling techniques. Barry sampled vegetation within small quadrats, placed along line transects;

in the present study, the vascular flora in the whole marsh was sampled.

Barry (1980) noted the preference of Juncus roemerianus in less saline portions of salt marsh where

salinity was less than 15 ppt. Where salinity exceeded 15 ppt, Spartina alterniflora replaced J. roemerianus.

Stalter et al., Flora of abandoned rice fields, South Carolina 673

Spartina alterniflora tolerated longer and deeper tidal flooding than J. roemerianus, and borders J. roemerianus

along the Waccamaw River.

The vegetation noted by Barry (1968) in the Airport marsh and abandoned Thousand Acre Rice Field

is similar to that observed today. Both Barry (1968) and Baden (1971) reported Spartina cynosuroides and

Scirpus robustus growing in profusion on the western border of the old rice field along with the more numer-

ous Typha angustifolia, T. latifolia with Juncus spp. Barry (1968) and Baden (1971) observed Pontederia cordata,

Sagittaria falcata, Peltandra virginica, Lilaeopsis chinensis and Hymenocallis crassifolia. Hymenocallis crassifolia

disappeared after Hurricane Hugo (Stalter and Baden 1994) and was not observed in 2002-2006.

Airport

The most significant change in vascular plant composition at the Airport Marsh is the increase of the non-

native, Phragmites australis. Phragmites was first reported in a marsh bordering the west side of the Waccamaw

River by Stalter (1975). Dikes and dredge spoils deposited on Baruch property in the 1970s may have created

habitat favorable for the invasion of Phragmites (Stalter @ Baden 1994).

The Airport Marsh experienced a decline in species composition over the 39 year study. Forty three

species were identified in 1968-1969 (Baden 1971) while 30 were identified in 2002-2006. Phragmites

has invaded Airport Marsh, outcompetes native species, and may be responsible for the decline in species

diversity and absence of some vascular plant species reported in 1968. High salinity < 15 ppt during the

drought of 2001 and 2002 may have killed salt intolerant taxa (Table 2).

Common taxa at Airport Marsh include Spartina cynosuroides, Zizania aquatica, Juncus spp., Typha

angustifolia, Cladium jamaicense, Scirpus robustus, Scirpus validus, Pontederia cordata, Aster subulatus, A. tenui-

folius, Pluchea foetida and Lythrum lineare. Lilaeopsis chinensis and Ptilimnium capillaceum are common at the

edge of the marsh as are Juncus biflorus, J. coriaceus, J. marginatus, J. nodosus, Hibiscus moscheutos and Samolus

parviflorus. Species diversity at Airport Marsh is greatest within the intertidal area.

APPENDIX

Species composition in three brackish marshes on the Belle W. Baruch Institute for Marine Biology and

Coastal Research, Georgetown, South Carolina. Species observed in 1967-1971 are compared with those

observed in 1987, 1991 and 2002-2006. Taxa in brackets are synonyms. Non-native taxa are scored with

an asterisk (*) preceding the scientific name. Collection dates are omitted for woody taxa occurring at the

upper marsh fringes flooded at the new and full moon.

POLYPODIOPHYTA

Blechnaceae

Woodwardia areolata (L.) Moore; occasional at marsh edge,

Alderly, 2002-06

Osmundaceae

Osmunda cinnamomea L.; occasional at marsh edge, Alderly,

2002-06

Osmunda regalis (L.) var. spectabilis (Willd.) A. Gray; rare at

marsh edge, 1968-06 Airport; occasional at Alderly,

2004-06

Thelypteridaceae

Thelypteris palustris Schott; occasional at marsh edge, Alderly,

PINOPHYTA

Cupressaceae

Taxodium distichum (L.) Rich; occasional at 1000 Acre Rice

field and Alderly, 1968—06; declining at both sites due to

Hugo 1989 and drought of 2003.

MAGNOLIPHYTA—MAGNOLIOPSIDA

Amaranthaceae

*Alternanthera philoxeroides (Mart.) Griseb.; common at

Alderly 1991—06

Amaranthus cannabinus (L) Sauer; occasional at marsh edge

Airport and Alderly, 1968-06

Apiaceae

Eryngium aquaticum L. rare at Airport 1991; occasional at

Alderly 1991; rare at Alderly 2004

Hydrocotyle sp.; common at marsh border at all three

marshes

Lilaeopsis chinensis (L.) Kuntze; common at marsh border at

all three marshes

Ptilimnium capillaceum (Michx.) Raf; common at marsh border

at all three marshes

Sium suave Walt.; occasional at Airport 1991-06; occasional

at Alderly 1968-06

Aquifoliaceae

lex cassine L. var. cassine; rare at marsh edge Alderly

llex opaca Ait.; occasional at marsh edge Alderly

674

Asteraceae

Aster carolinianus Walt.; occasional at Alderly, 2004-06

Aster subulatus Michx.; frequent at 1000 Acre Rice Field 1968

and 1991; frequent at marsh edge Airport 1968-06; oc-

casional at Alderly 1968 and 1991

Aster tenuifolius L.; occasional at marsh edge 1000 Acre Rice

Field 1968 and 1991; occasional at Airport 1968-06; oc-

casional at Alderly 1968-06

Baccharis halimifolia Michx.; occasional at marsh edge, Alderly,

2004-06

Bidens laevis (L.) BSP.; occasional at marsh edge Airport and

Alderly 1968 and 1991

Boltonia asteroides (L.) U Hér; rare at marsh edge Airport

Borrichia frutescens (L.) DC.; occasional at edge of marsh, 1000

Acre Rice Field, 1968-06

Eupatorium album L.; occasional at Airport and Alderly im-

mediately after Hurricane Hugo, 1991

Fupatorium coelestinum L.; occasional at marsh edge, Alderly,

2002-06

Eupatorium perfoliatum L; occasional at marsh edge, Alderly,

lva frutescens L; occasional, upper fringe of marsh at 1000

Acre Rice Field 1968-2006

Mikania scandens (L.) Willd.; occasional, upper fringe of marsh

at 1000 Acre Rice Field and Alderly 1968-2006

Pluchea camphorata (L.) DC; rare, 1000 Acre Rice Field, 1968

Pluchea foetida (L) DC; occasional at Airport, 1968-2006;

occasional at Alderly 1968 and 1991

Solidago sempervirens L.; occasional at edge of marsh, 1000

Acre Rice Field 1991; Airport 1968 and 1991; Alderly

1968-06

Verbesina occidentalis (L.) Walt.; frequent at marsh border of

Alderly, 1991-06

Betulaceae

Alnus serrulata (Ait.) Willd.; rare, marsh border of Alderly

Callitrichaceae

Callitriche peploides L.; occasional, marsh edge, Alderly,

Caprifoliaceae

Sambucus canadensis L.; rare at edge of Alderly

Viburnum nudum L, rare at edge of Alderly

Chenopodiaceae

*Chenopodium album L.; rare at edge of Airport and Alderly,

1968

Convolvulaceae

Calystegia sepium (L.) R. Br.; occasional, Alderly 2006

Dichondra carolinianus Michx.; frequent at marsh edge of

Alderly, 2004-06

Ipomoea sagittata Poir; occasional upper fringe of 1000 Acre

Rice Field 1991-2006

Cuscutaceae

Cuscuta pentagona Engelm. [Cuscuta campestris Yunker]; oc-

casional on assorted taxa, Alderly 1968 and 2004

fal, Dat o ID L

Journal of

titute of Texas 1(1)

Euphorbiaceae

*Triadica sebifera (L) Small [Sapium sebiferum (L.) Roxb.]; oc-

casional on marsh fringe of Airport, 2004-06

Fabaceae

Chamaecrista fasciculata (Michx) Greene [Cassia fasiculata

Michx.]; occasional upper marsh fringe of Airport and

Alderly, 1968

Gentianaceae

Sabatia calycina (Lam.) Heller; rare, 1000 Acre Rice Field

1968;Alderly 1968 and 1991

Sabatia stellaris Pursh.; frequent 1000 Acre Rice Field and

Alderly 1968-06; occasional Airport 1968

Grossulariaceae

Itea virginica L.; rare, upper marsh fringe of Alderly

Lamiaceae

Lycopus virginicus L.; occasional at marsh edge, Alderly,

2004-

Scutellaria integrifolia L; rare, Alderly, 2004-06

Stachys floridana Shuttlew. ex Benth.; abundant, Alderly

Lythraceae

Lythrum lineare L.; abundant at all marshes 1968-06

Malvaceae

Hibiscus moscheutos L.; occasional at edge of marsh, Airport

and Alderly 1968-06

Nymphaeaceae

Nuphar lutea (L.) Sm. ssp ifolia (Walt.) E.O. Beal; occasional

at small ponds at Alderly, 1968-06

Nymphaea odorata Ait; rare by culvert, Alderly 2004-06

Nyssaceae

Nyssa biflora Walt. [Nyssa sylvalica var. biflora (Walt.) Sarg.];

occasional at marsh fringe, Alderly

Plumbaginaceae

Limonium carolinianum (Walt.) Britt; rare, 1000 Acre Rice Field

1968 and 2004

Polygonaceae

Polygonum hydropiperoides Michx.; frequent at edge of

marsh1000 Acre Rice Field, 1991-06 and Airport 1968

and 2006, and Alderly, 1968-06

Polygonum sagittatum L.; rare at Alderly, 2004-06

Rumex verticillatus L.; rare at Airport, 1968; occasional at

Alderly 1968-06

Potamogetonaceae

Potamogeton diversifolius Raf, rare, Alderly at pool near culvert

Primulaceae

Samolus valerandi L. ssp. parviflorus (Raf) Hulten [S. parviflorus

Raf]; frequent at edge of marsh at all sites 1968-2006;

occasional upper marsh fringe of Alderly, 1968-06

Rosaceae

Rosa laevigata Michx.; rare, upper marsh fringe of Alderly

Rosa palustris Marsh.; occasional upper marsh fringe of

Alderly

Stalter et al., Flora of abandoned rice fields, South Carolina

Rubiaceae

Cephalanthus occidentalis L.; rare, upper fringe of Airport

1968-06; occasional, upper marsh fringe of Alderly

1968-06

Galium obtusum Bigel.; frequent, Alderly 2006

Salicaceae

Salix caroliniana Michx; occasional, upper marsh fringe of

Alderly

Saururaceae

Saururus cernuus L. frequent at Alderly 1968-06

Scrophulariaceae

Bacopa caroliniana (Walt.) B.L. Robins; occasional at Alderly,

Bacopa monnieri (L.) Pennell; rare at edge of marsh, Airport

Urticaceae

Boehmeria cylindrica (L.) Swartz.; occasional at Alderly,

1991-06

Verbenaceae

Phyla lanceolata (Michx.) Geene; occasional at marsh edge,

1000 Acre Rice Field 1968-06; abundant at marsh edge,

Airport 1968-06

*Verbena brasiliensis Vell.; rare, Alderly, 2006

MAGNOLIOPHYTA—LILIOPSIDA

Alismataceae

Sagittaria lancifolia L. [S. falcata Pursh]; occasional at Airport,

1991-06; abundant at Alderly 1968-06

Araceae

Orontium aquaticum L, rare at Alderly, 1968-06

Peltandra virgil uca (L.) Schott; freq uentat Airport and Alderly

1968-06

Commelinaceae

*Murdannia keisak (Hassk.) Hand.-Maz. [Aneilema keisak Hassk.];

abundant at Alderly, 2004-06

Cyperacaeae

Cladium jamaicense Crantz; abundant at all marshes

1968-06

Carex alata Torr; occasional at marsh edge Airport and Alderly

1968-

Carex atlantica L.H. Bailey; occasional, Alderly 2006

Carex lurida Wahleng.; occasional, Alderly 2006

Carex stipata Willd.; rare, Alderly 2006

Cyperus drummondii; rare at Alderly 2004

Cyperus refractus Engelm. ex Boeckl.; occasional at Aldery

2004-06

Cyperus strigosus L.; occasional at edge of marsh Alderly

Eleocharis engelmannii Steud.; occasional at all 3 marshes,

1968; and at 1000 Acre Rice Field and Alderly, 1991

Eleocharis flavescens (Poir.) Urb. occasional, Alderly, 2006

Eleocharis quadrangulata(Michx.) R.& S; rare at Airport and

Alderly 1968; rare at Alderly 2004-06

Fimbristylis thermalis S. Wats. [Fimbristylis spadicea auct. non

675

(L) Vahl]; frequent at 1000 Acre Rice Field and Airport

1968-06; rare at Alderly 68

Rhynchospora colorata (L.) H. Pfeiffer [Dichromena colorata (L.)

Hitchc.]; occasional at marsh edge at Alderly 1968-06

Rhynchospora miliacea (Lam.) Gray; rare at 1000 Acre Rice

Field, 2004—06; occasional to locally abundant at Adlerly,

1968-06

Scirpus cyperinus (L) Kunth; occasional at Alderly 1968-06

Scirpus pungens Vahl. [S. americanus Pers.]; abundant at 1000

Acre Rice Field and Airport 1968-06; occasional at Alderly

1968-91.

Scirpus tabernaemontani K.C. Gmel. [Scirpus validus Vahl];

abundant at 1000 Acre Rice Field 1968-2006 and at

Alderly 1968-91.

—_

Iridaceae

Iris virginica L.; frequent at Alderly 1968-06

Juncaceae

Juncus acuminatus Michx; rare at Alderly 1968

Juncus biflorus Ell; occasional at 1000 Acre Rice Field 1968 and

1991; rare at Airport and Alderly, 1968

Juncus coriaceus Mackenzie; occasional at marsh edge, 1000

Acre Rice Field 1968 and 1991; occasional at Airport and

Alderly 1968-06

Juncus dichotomus Ell; occasional at Alderly 2006

Juncus effusus L.; occasional at Alderly 2006

Juncus marginatus Rostk.; occasional at Airport 1968-06

Juncus nodosus L.; occasional at Airport 1991 and 2004

Juncus roemerianus Scheele; abundant at 1000 Acre Rice Field

1968-06 and Alderly at the border of the Waccamaw River

1968-06; occasional at Airport 1968

Liliaceae

Crinum americanum L; abundant in vicinity of Taylor boat

landing, Alderly at the edge of the marsh 1968-06

Hymenocallis floridana (Raf) Morton [Hymenocallis crassifolia

Herbert]; occasional at Alderly 1968, extirpated by Hur-

ricane Hugo 1989

Poaceae

*Arundo donax L; rare at marsh edge 1000 Acre Rice Field 1968

and 1991; rare at marsh edge Airport 1991; occasional at

Alderly near Waccamaw River 1968-06

Chasmanthium latifolium (Michx.) Yates [Uniola latifolia Mi-

chx.]; occasional at edge of marsh, Alderly, 1968-1991

Chasmanthium laxum (L.) Yates [Uniola laxa (L.) B.S.P]; occa-

sional at edge of marsh, Alderly 1968-06

Cinna arundinacea L, frequent at marsh border Airport and

Alderly 1968-06

Distichlis spicata (L.) Greene; abundant at 1000 Acre Rice

Field 1968-06

*Echinochloa crus-galli (L.) P. Beauv.; occasional at marsh

border Airport 1968-06; occasional at marsh border

Alderly 1968-06

Erianthus giganteus (Walt.) Muhl.; frequent at marsh edge

Alderly 1968-06

Leersia oryzoides (L) Sw. abundant at Alderly 2004-06

Panicum virgatum L.;frequent at marsh border 1000 Acre Rice

676

Field 1991 and 2006; frequent at marsh border Airport

and Alderly 1968-06

Panicum sp.; occasional at marsh border Alderly 2004-06

Paspalum distichum L.; occasional at marsh border, Airport

1968 and 2004-06

*Phragmites australis (Cav.) Trin.; abundant at all marshes

2004-06; see text for additional information

Setaria magna Griseb,; occasional at marsh border, all marshes

Spartina alterniflora Loisel; abundant along ditches, 1000 Acre

Rice Field 1968-06

Spartina cynosuroides (L.) Roth; abundant at all marshes

Spartina patens (Ait. Muhl.; abundant at 1000 Acre Rice Field

£sthka Dat o ID L

Journal of

titute of Texas 1(1)

Tripsacum dactyloides (L.) L; rare at 1000 Acre Rice Field edge

004-06; occasional at marsh edge Alderly 1968-06

Zizania aquatica L; occasional at 1000 Acre Rice Field 2004;

abundant at Airport and Alderly 1968-06

Zizaniopsis miliacea (Michx.) Doell & Aschers.; frequent at

Airport and Alderly 1968-06

Pontederiaceae

Pontederia cordata L.; rare at Alderly 2004-06

Typhaceae

Typha angustifolia L; frequent at all marshes 1968-06

Typha domingensis Pers.; frequent at 1000 Acre Rice Field

1991-06, occasional at Alderly 1991-06

Typha latifolia L; abundant at 1000 Acre Rice Field 1968-06;

occasional at Alderly 1968-06

ACKNOWLEDGMENTS

We wish to thank Dennis Allen, Assistant Director, Belle W. Baruch Institute for Marine Science and Coastal

Research, for granting collecting permits, for providing us with general information on water salinity and

providing a vehicle and a boat which were used in collecting forays at the Institute; Steve Hutchinson, Baruch

Institute, who provided us with the salinity data for the Winyah Bay Estuary; Eric Lamont for identifying the

Asteraceae; Steve Clemants, Brooklyn Botanical Garden for identifying Juncus; John Nelson, USC Herbarium

for identifying several taxa; Anthony Reznicek of the University of Michigan Herbarium for identifying Carex;

Gordon Tucker for identifying additional Cyperaceae; Georgia Angelopoulos and Sabrina Truc undergraduate

research students at St. John's University, and St. John's University for purchasing herbarium supplies.

REFERENCES

Anonymous. 1996. Weather America. Toucan Valley Publications, Inc. Milipita, CA.

BADEN, J. 1971. Tidal marsh vegetation on abandoned rice fields, Winyah Bay, Georgetown, South Carolina. M.S.

thesis. University of South Carolina, Columbia.

BADEN, J., W.T. BATSON, and R. STaLTER. 1975. Factors affecting the distribution of vegetation of abandoned rice fields,

Georgetown County, South Carolina. Castanea 40:171-184.

Barry, J.M. 1968. A survey of the native vascular plants of the Baruch Plantation. Georgetown, South Carolina.

M.S. thesis. University of South Carolina, Columbia.

Barry, J.M. 1980. Natural Vegetation of South Carolina. Columbia: University of South Carolina Press.

BanucH, B.M. 1957. Baruch: My own story. New York. Henry Holt and Co.

BLOOD, E.R., P. ANDERSON, P.A. Smith, C. Nveno, and K.A. GINSBERG. 1991. Effects of Hurricane Hugo on coastal soil solu-

tion chemistry in South Carolina. Biotropica 23:348-55.

Conner, W.H. and L.W. INAgiNETTE. 2005. Identification of salt tolerant baldcypress (Taxodium distichum (L.) Rich) for

planting in coastal areas. New Forests 29:305-312.

ELEUTERIUS, L. 1972. The mashes of Mississippi. Castanea 37:153-168.

GARDNER, L.R., W.K. MICHENER, E.R. BLoop, T.M. WiLLIAMs, D.J. Liescome, and W. Jerrerson. 1991. Ecological impact of Hur-

ricane Hugo- salinization of a coastal forest. J. Coastal Res. 8:301-317.

Gentry, R.C. 1971. Hurricanes, one of the major features of air-sea interaction in the Carribean Sea. Symposium

on investigation and resources of the Carribean and adjacent regions. Paris: UNESCO.

GLEASON, H.A. and A. Cronauist. 1991. Manual of vascular plants of the Northeastern United States, Canada, and

Greenland. 2nd Edition. The New York Botanical Garden, Bronx.

JACCARD, P. 1908. Nouvelles recherches sur la distribution florale. Bull. Soc. Vaud. Sci. Nat. 44:223-270.

KARTESZ, J.T. 1994. A synonymized checklist of the vascular flora of the United States, Canada and Greenland. 2nd

ed. Volume 1-Checklist. Timber Press, Inc. Portland, OR.

Stalter et al., Flora of abandoned rice fields, South Carolina 677

KAUFMAN, L. and PJ. Rousseeum. 1990. Finding groups in data: An introduction to cluster analysis. John Wiley €

Sons, Inc., New York.

ManLv, B.F.J. 1997. Randomization, bootstrap and Monte Carlo methods in biology, 2nd ed. Chapman & Hall,

New York.

Noreen, E.W. 1989. Computer-intensive methods for testing hypotheses: An introduction. John-Wiley & Sons,

New York.

PALMER, M.W, G.L. Wabe, and P. NEAL. 1995. Standards for the writing of floras. Bioscience 45:339-345.

PorcHer, R.D. 1976. A history of the land use of Hobcaw Barony. Report to the Belle W. Baruch Forest Science

Institute of Clemson University, Georgetown, S.C.

RADFORD, A.E., H.E. Ares, and C.R. Bett. 1968. Manual of the vascular flora of the Carolinas, 2nd ed. The University

of North Carolina Press, Chapel Hill.

SrALTER, R. 1968. An ecological study of a South Carolina salt marsh. Ph. D. dissertation. University of South

Carolina. Columbia.

STALTER, R. 1971. The summer and fall flora of Huntington Beach State Park, Georgetown County, South Carolina.

Castanea 36:167-1 74.

STALTER, R. 1972. The summer and fall flora of Brookgreen Gardens, Georgetown Co., South Carolina. Castanea

37:214-226.

STALTER, R. 1973. Factors influencing the distribution of vegetation of the Cooper River Estuary. Castanea

38:18-24.

STALTER, R. 1975. Phragmites communis in South Carolina. Rhodora 77:159.

STALTER, R. and J. BADEN 1994. A twenty year comparison of vegetation of three abandonded Rice Fields, George-

town County, South Carolina. Castanea 59:69-77.

STALTER, R. and E.E. Lamont. 1990. The vascular flora of Assateague Island, Virginia. Bull. Torrey Bot. Club 117:

48—56.

STALTER, R. and E.E. Lamont. 1993. The vascular flora of Fort Sumter and Fort Moultrie, South Carolina, one year

after Hurricane Hugo. Castanea 58:141-152.

SUTTON, P, R. Meyer, and R. Stater. 1990. The vascular plants of Cape May Point State Park, Cape May County, New

Jersey. Bull. Torrey Bot. Club 117:294-300.

Wass, M.L. and T.D. Wnicur. 1969. Coastal wetlands of Virginia. Virginia Institute of Marine Science, Glouster

Point.

WUNDERLIN, R.P. 1998. Guide to the vascular plants of Florida. University Press of Florida, Gainesville.

678 Journal of the Botanical R h Institute of Texas 1(1)

BOOK NOTICES

GERALDINE Etuis Watson. 2006. Big Thicket Plant Ecology: An Introduction (3rd Ed.). (ISBN 978-1-

57441-214-7, pbk). University of North Texas Press, PO Box 311336, Denton, TX 76203-1336, U.S.A.

(Orders: www.unt.edu/untpress, 940-565-2142, 940-565-4590 fax, 1-800-826-8911). $14.95, 144

pp., 20 illustrations, 15 maps, 8 figures, 6" x 9".

This small volume is the starting point for anyone interested in the biology of the Big Thicket -- an expanded and updated edition of the

original from 1975. Included are discussions of the definition of the “Big Thicket,” geological history, plant communities and succession,

and descriptions of the subdivisions (*units") of the Big Thicket National Preserve. Good maps, diagrams, and photos.

Other useful books on Big Thicket biology:

—Ajilvsgi, G. 1979. Wildflowers of the Big Thicket, East Texas, and Western Louisiana. Texas A&M Univ. Press, College Station

—Gunter, P. 1971. The Big Thicket: A Challenge for Conservation. Viking Press, New York.

—Gunter, PA.Y. 1993. Big Thicket: An Ecological Reevaluation. Univ. North Texas Press, Denton

—Peacock, H. 1994. Nature Lovers Guide to the Big Thicket. Texas ASM Univ. Press, College Station.

An All-Taxa-Biological-Inventory (ATBI) is underway for the Big Thicket National Preserve. For detailed information on this project,

contact Linda C. Brindle, Executive Director, Big Thicket Association «di @bigthicl rg>.—Guy Nesom, Botanical Research Institute

of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A.

Kris M. Havstap, Laura E HUENNEKE, AND WILLIAM H. SCHLESINGER (eds.). 2006. Structure and Function of

a Chihuahuan Desert Ecosystem: The Jornada Basin Long-Term Ecological Research Site.

(ISBN 978-0-195-11776-9, hbk). Oxford University Press, 2001 Evans Road, Cary, NC 27513, U.S.A.

(Orders: www.oup.com, 1-800-451-7556, 919-677-1303 fax). $74.50, 492 pp., 98 b/w halftones,

maps, line drawings, 6" x 914",

The Jornada Basin Long-Term Ecological Research Site is located in south-central New Mexico (25 km northeast of Las Cruces). It in-

cludes the 78,000 ha Jornada Experimental Range operated by the USDA Agricultural Research Service and the 22,000 ha Chihuahuan

Desert Rangeland Research Center (CDRRC) DOM by New Mexico State University. Research toward understanding the causes and

consequences of desertification began in this basin in 1912, and the site was established in 1982 as one of NSF's Long-Term Ecologi-

cal Research (LTER) sites. Research reported and summarized here is collaborative across a number of disciplines. The book has 18

chapters, written by 36 contributors.

Contents.—1) Introduction (Havstad & Schlesinger); 2) Regional Setting of the JB; 3) Climate and Climatological Variations in

the JB; 4) Soil Development in the JB; 5) Patterns and Controls of Soil Water in the JB; 6) Nutrient Cycling within an Arid Ecosystem; 7)

Biogeochemical Fluxes across Piedmont Slopes of the JB; 8) Water and Energy Balances within the JB; 9) Eolian Processes on the JB; 10)

Plant Communities in the JB: The Dynamic Landscape; 11) Patterns of Net Primary Production in Chihuahuan Desert Ecosystems; 12)

Chihuahuan Desert Fauna: Effects on Ecosystem Properties and Processes; 13) Grazing Livestock Management in an Arid Ecosystem;

14) Remediation Research in the JB: Past and Future; 15) Applications of Remotely Sensed Data From the JB; 16) Modeling the Unique

Attributes of Arid Ecosystems: Lessons from the JB; 17) A Holistic View of an Arid Ecosystem: A Synthesis of Research and Its Applica-

tions; 18) Future Directions in Jornada Research: Applying an Interactive Landscape Model to Solve Problems.

Two websites provide information on Jornada Basin research ce ar bit and active data sets, current projects, research

dia access authorization forms, program descriptions, and an i

ofthe history of publications in the Jornada

:<http://usda-ars.nmsu.edu M «http: Titties -www.nmsu.edu>.—Guy ee Botanical Research Institute of Texas, 509 Pecan

Street, Fort Worth, TX 76102-4060, U.S.A.

J. Bot. Res. Inst. Texas 1(1): 678. 2007

THBVASCUDEARDEORAOPINASELPRAIRIE:

AUOAXASDIALBRAIRIE.REMDNABNTOIBCODRAZORDS-GOUNTY, TEXAS

David J. Rosen

U. S. Fish and Wildlife Service

17629 El Camino Real, Suite 211

Houston, Texas 77058-3051, U.S.A.

ABSTRACT

An intensive survey of the vascular flora of Nash Prairie, a ca. 120 ha Coastal Prairie remnant in Brazoria County, Texas, resulted in a

checklist of 311 species of vascular plants representing 63 families and 197 genera. The seven families containing the most species are

Poaceae (70), Cyperaceae (37), Asteraceae (46), Fabaceae (17), Euphorbiaceae (10), Apiaceae (7), and Scrophulariaceae (7). Rich native

genera include Carex (11 spp.), Cyperus (9 spp.), Juncus (7 sppJ, Panicum (7 spp.), and Paspalum (7 spp.). Non-native species account

for only 7% of the total, 50% (11) of which are grasses. The native flora comprises 289 species distributed in 63 families. The native

grass flora includes 59 species in five subfamilies, 71% of which were C, species, with most of these belonging to the Panicoideae (35

spp.). Noteworthy collections of globally and regionally rare species and species with temperate amphitropical distributions from Nash

Prairie are discussed, and an annotated checklist of vascular plants is provided. This research suggests previous estimates of species

richness for climax Coastal Prairie in Texas are low and that historic and potential losses of botanical diversity are greater than previ-

ously thought.

RESUMEN

Un seguimiento intensivo de la flora vascular de Nash Prairie, de unas 120 ha de restos de pradera costera en Brazoria County, Texas,

dio como resultado un listado de 311 especies de plantas vasculares de 63 familias y 197 géneros. Las siete familias con mayor número

de especies son Poaceae (70), Cyperaceae (37), Asteraceae (46), Fabaceae (17), Euphorbiaceae (10), Apiaceae (7), y Scrophulariaceae (7).

Los géneros nativos ricos en especies incluyen Carex (11 spp.), Cyperus (9 spp.), Juncus (7 spp.), Panicum (7 sppJ, y Paspalum n spp.).

Las especies exóticas son sólo el 7% del total, siendo el 50% (11) gramíneas. La flora nativa comprende 289 especies distrib

familias. Las gramíneas nativas incluyen 59 especies de cinco subfamilias, de las que el 71% son especies C,, perteneciendo la mayoría

1 Ta] ps |

de ellas a las Panicoideae (35 spp.). Se discuten | nivel UA como global y las especies

con distribución templada y amfitropical de Nash Prairie, y se aporta un XM de las plantas vasculares. Esta investigación sugiere

que las estimaciones previas de riqueza de especies de la climax de la Coastal Prairie de Texas son bajas y que las pérdidas históricas y

potenciales de diversidad botánica son mayores de lo que se había pensado antes.

INTRODUCTION

The original extent of the Coastal Prairie region encompassed ca. 3,800,000 ha extending from southcentral

Louisiana to south Texas along the northwestern Gulf of Mexico, and contributed about 196 to the total of all

major grassland types of the contiguous United States (Sims & Risser 2000). Most of the original vegetation

of the Coastal Prairie has been altered by overgrazing, conversion to various agricultural uses, encroach-

ment of woody vegetation due to fire protection, or destroyed by industrial, commercial, and residential

development (Diamond & Smiens 1984; Smeins et al. 1991; Sims & Risser 2000). The Coastal Prairie was

characterized in detail by Smeins et al. (1991), and divided in Texas into an upper and lower section based

on variation in soil and climate (Diamond & Smiens 1984). A full account of the flora of the Coastal Prairie

would be difficult to compile since 9996 of the original vegetation has been destroyed and the best and larg-

est remaining remnants occur on private property. Several years ago, I was presented with the opportunity

to survey a relatively large and intact example of this rare plant community.

METHODS

Nash Prairie is a ca. 120 ha remnant Coastal Prairie on the Kittie Nash Groce (KNG) Ranch. Nash Prairie

is managed as a native hay meadow and has been maintained for decades by frequent mowing, haying,

and burning. The entire prairie is seldom dry enough to burn or hay all at once, which results in a random

J. Bot. Res. Inst. Texas 1(1): 679 — 692. 2007

680 Journal of the Botanical R h Institute of Texas 1(1)

pattern of disturbance across the landscape. The topography of Nash Prairie is intact and includes wetland

depressions and abandoned stream meanders, numerous pimple mounds, and inter-mound flats (Fig. 1).

Discussions with KNG Ranch personnel revealed that the prairie has probably never been subjected to

overgrazing, and the native sod has never been broken. Access for an intensive floristic survey was arranged

through cooperation between the KNG Ranch and the Nature Conservancy of Texas.

Nash Prairie is located in southwest Brazoria County, Texas, ca. 35 km south of the Houston met-

ropolitan area (Fig. 2). The prairie is bounded on all sides by agricultural land subject to various uses,

including rice and row crop farming, improved pasture, and grazing. A barbed-wire fence along the south,

west, and north boundaries accounts for some of the woody species that occur in the annotated checklist.

Nash Prairie occurs roughly in the geographic center of the Upper Coastal Prairie region, which lies within

the Coastal Plain Province at the northern limit of the subtropical vegetation zone (Fenneman 1928; Good

1953). The regional climate is moist subhumid mesothermal characterized by long hot summers and mild

winters (Thornthwaite 1948). Average annual rainfall is 132 cm, with 6096 occurring from April through

September (Crenwelge et al. 1981). The average daily summer temperature is 27°C, and average daily winter

temperature is 13°C (Crenwelge et al. 1981).

Soils mapped in Nash Prairie are Edna fine sandy loam and Edna-Aris complex (Crenwelge et al. 1981).

The Edna-Aris complex is generally associated with older stream meanders and is distinguished by the

distinctive pimple mounds, with Aris fine sandy loam soils occupying the mounds, and Edna clayey soils

occurring on the flats in between (Crenwelge et al. 1981). Pimple mounds are characteristic of remnant

Coastal Prairie sites which are topographically intact, and they are a reliable indicator that the native sod

is unbroken. Pimple mounds, like prairie wetlands (Fig. 3), also provide a microhabitat differing in soil

texture, slope, soil moisture and elevation from the surrounding landscape.

Collecting trips were made to the prairie from August 2003 through September 2006. The prairie was

visited on average three times a month from March to October throughout the study with the exception of

a single visit in December, 2005. A complete set of voucher specimens are housed at the University of Texas

at Austin Plant Resources Center Herbarium (TEX). Some duplicate specimens can also be found at BRCH,

BRIT, MICH, SBSC, TAES, VSC and US (acronyms follow Holmgren et al. 1995). Plant identifications were

made using the Flora of North America (2000; 2002a; 2002b; 2003) and various regional manuals, including

Correll and Johnston (1970), Gould (1975), Isely (1990), and Smith (1994).

RESULTS AND DISCUSSION

This research resulted in collections of 311 species of vascular plants representing 63 families and 197

genera (Table 1; Appendix I). The seven families containing the most species are Poaceae (70), Cyperaceae

(37), Asteraceae (46), Fabaceae (17), Euphorbiaceae (10), Apiaceae (7), and Scrophulariaceae (7). Rich native

genera include Carex (11 spp.), Cyperus (9 sppJ, Juncus (7 spp.), Panicum (7 spp.), and Paspalum (7 spp.). Non

native species accounted for 7% of the total species, 50% (11) of which were grasses. A subjective estimate

of the dominant and sub-dominant species of climax plant assemblages occurring at various landscape

positions is presented in Table 2. This is meant to aid managers and restorationists by providing lists of

target species for different cover types from a relatively intact Coastal Prairie remnant. During the study,

species occurrence and dominance varied seasonally, and dominance roles appeared to shift with seasonal

and annual precipitation patterns and disturbance from mowing and haying. Because mowing and haying

occurred sporadically across the site during the study years, it is difficult to know what role disturbance

played in the distribution and abundance of species in each cover type. Long-term quantitative vegetation

studies at Nash Prairie are greatly needed.

Infrafamilial Diversity in Poaceae

The native grass flora of Nash Prairie includes 59 species in five subfamilies and comprises 71% C, species

(Table 3). As indicated by Smeins et al. (1991) for the Coastal Prairie, the native grass flora of Nash Prairie

Rosen, Flora of Nash Prairie, Brazoria County, Texas 681

is dominated by the Panicoideae. The greatest number of C, taxa also belong to the Panicoideae (33 spp.),

followed by the Chloridoideae (14 spp.; Table 3). Sixty-four percent of the introduced grass species found use

the C, photosynthetic pathway, including potentially aggressive species such as Cynodon dactylon, Paspalum

dilatatum, P. notatum, P. urvillei, and Sorghum halepense. The dominance of C, species in the Coastal Prairie

is expected given their competitive advantage over C, species in a subtropical climate regime (Diamond &

Smeins 1988).

Endemic and Rare Taxa

This survey yielded collections of several rare or otherwise noteworthy taxa. Species with distributions

limited to the Coastal Prairie or with the greatest extent of their range occurring therein include Amsonia

repens, Asclepias linearis, Cooperia traubii, Euphorbia texana, Liatris bracteata, Rudbechia texana, and Thalictrum

texanum (Correll & Johnston 1970). Species that are regionally rare and previously unreported from the

Coastal Prairie (including some significant range extensions from collections previously mapped by Turner

et al. 2003a; Turner et al. 2003b) include the Great Plains Eleocharis compressa var. acutisquamata and the

eastern species E. wolfti, Scleria muhlenbergii, Juncus elliottii. var. elliottii, and Sporobolus silveanus. Diamond

and Smeins (1985) described a novel S. silveanus- Carex meadii grassland type from the northern end of the

Blackland Prairie. The occurrence of members of this same assemblage at Nash Prairie suggests this com-

munity type might have been more widespread prior to European settlement.

Several species with temperate amphitropical distribution are known from the Coastal Prairie region

f4L,D

682 Journal of tani h Institute of Texas 1(1)

xp A PDA y y.

wt f- Ii 27 T iN

po a To vin AREA

OPS FAR

ara AS "

UM i;

at!

-r

ES | .

a Mei UM SS f k,

A ALAA IRE ;

I Dant £

Pal £1 f T 7 24 RON NA

Fic 2 Sd id sa ICAQ» (UGSTICU

and are considered by most botanists to be native taxa with disjunct distributions. Raven (1963) suggested

a late Pliocene or Pleistocene introduction by long range dispersal for temperate amphitropical disjuncts in

North America and considered coastal, seasonally wet, and/or open weedy grassland habitats particularly

suitable for disjuncts to establish. Several amphitropical disjuncts listed by Raven (1963) occur at Nash Prai-

rie, including Soliva sessilis [as S. pterosperma (Juss.) Less.], Hordeum pusillum, Phalaris angusta, Lepuropetalon

spathulatum, and Proserpinaca palustris. The following discussion provides details of other amphitropical

elements of the Coastal Prairie.

Bothriochloa exaristata is a relatively rare species known from sites with heavy soils in the Coastal

Prairie of Texas and Louisiana, with a disjunct distribution in coastal areas of southern Brazil and adjacent

Argentina, inland to Paraguay (Gould 1975; Allred 2003). Tucker et al. (2003) suggested that Cyperus cepha-

lanthus might be naturalized in the United States. However, more detailed accounts support C. cephalanthus

as a temperate amphitropical disjunct and a reliable indicator of undisturbed Coastal Prairie remnants in

Texas and Louisiana (Carter @ McInnis 1993; Grace et al. 2000). Collections of C. cephalanthus from Nash

Prairie are only the second reported (and the only extant population known) in Texas since the type speci-

men was collected near Galveston Bay (Rosen & Christoffersen 2004). Cyperus drummondii is another reli-

able indicator of Coastal Prairie remnants in Texas and Louisiana, as well as other undisturbed habitats in

the southeast United States (Carter et al. 1999; Rosen 2004). Panicum bergii is distributed in eastern South

America and southeastern Texas. Although Gould (1975) considered P. bergii to be introduced, Correll and

Rosen, Flora of Nash Prairie, Brazoria County, Texas 683

Fic. 3. Examples of wetland habitats at Nash Prairie, Brazoria County, Texas. Foreground is an abandoned stream meander dominated by Panicum

HCHIRUPHIUII Ji l J y Ww y MAar viet WOUJ}.

TABLE 1. Taxonomic summary of vascular plants of Nash Prairie, Brazoria County, Texas.

Species

Group Families Genera Native Non-native Total

Monocots 13 59 UE 14 154]

Eudicots 50 139 172 8 180

Totals 63 197 289 22 311

Johnston (1970) suggested it might be bicentric in distribution. Panicum bergii has been established in the

flora of the Coastal Prairie for some time, has a similar distribution to other amphitropical species, and does

not appear to be aggressive. Paspalum wrightii occurs in Cuba, Mexico (Campeche), Bolivia, Paraguay, and

Argentina (Allen & Hall 2003). Along the upper Texas coast P. wrightii occurs in prairie wetlands, where it

has probably frequently been confused with Paspalum plicatulum. Although Cuphea glutinosa is considered

native to South America, my experiences with this species have been restricted to late successional vegeta-

tion in Coastal Prairie remnants in Louisiana and Texas, suggesting it is also native to the flora. Temperate

amphitropical distributions have been reported for other grassland species, indicating floristic disjunctions

might have played a role in the development of the Coastal Prairie flora, in addition to other origins reported

by Smeins et al. (1991).

Invasive Species

Most introduced species at Nash Prairie were encountered along a paved public road that bounds the prairie

to the East and unimproved roads that provide access for farm equipment into the prairie (Fig. 1). Several

684 Journal of the Botanical R h Institute of Texas 1(1)

TABLE 2. Subjective estimate of dominant and sub-dominant species of climax plant assemblages occurring at various land-

scape positions in Nash Prairie. Species occurrence and dominance varied seasonally and across study years. Species are

listed in alphabetical order.

WETLANDS

Semi-permanently flooded depressions

Eleocharis quadrangulata Polygonum hydropiperoides

Hydrolea ovata Pontederia cordata var. cordata

Hymenocallis liriosme Proserpinaca palustris var. amblyogona

Iris brevicaulis Rhynchospora corniculata

Juncus effusus var. solutus Sagittaria graminea subsp. graminea

Juncus nodatus Sagittaria papillosa

Panicum hemitomon Sesbania drummondii

Paspalidium geminatum Thalia dealbata

Paspalum wrightii

Seasonally wet lowland prairie

Amsonia repens Juncus elliottii var. elliottii

Arnoglossum plantagineum Lobelia puberula

Axonopus fissifolius Neptunia lutea

Boltonia diffusa var. diffusa Panicum rigidulum subsp. rigidulum

Carex festucacea Panicum tenerum

Carex longii Panicum virgatum

Carex tetrastachya Paspalum floridanum

Carex triangularis Paspalum praecox

Cyperus drummondii Rhynchospora caduca

Cyperus virens var. virens Rhynchospora globularis

Eleocharis compressa var. acutisquamata Rudbeckia texana

Euthamia gymnospermoides Tridens strictus

Helianthus angustifolius Tripsacum dactyloides var. dactyloides

Juncus brachycarpus Vernonia missurica

UPLANDS

Upland prairie (incl. pimple mounds)

Acacia angustissima var. hirta Liatris pycnostachya

Andropogon gerardii subsp. gerardii Mimosa hystricina

Andropogon ternarius var. ternarius Muhlenbergia capillaris

Aristida purpurascens var. purpurascens Paspalum plicatulum var. plicatulum

Arnoglossum plantagineum Polytaenia nuttallii

Baptisia bracteata var. leucophaea Schizachyrium scoparium var. scoparium

Baptisia sphaerocarpa Scleria ciliata var. elliottii

Carex bushii Scleria pauciflora var. pauciflora

Carex meadii Silphium gracile

Croton glandulosus var. lindheimeri Solidago stricta

Cyperus echinatus Solidago tortifolia

Dichanthelium aciculare subsp. angustifolium Sorghastrum nutans

Eragrostis spectabilis Symphyotrichum dumosum

Eryngium yuccifolium Symphyotrichum ericoides var. ericoides

Fimbristylis puberula var. puberula lephrosia onobrychoides

Liatris acidota

species observed to disperse from disturbed sites into existing prairie vegetation include Cyperus entrerianus,

Paspalum urvillei, Sorghum halepense, and Triadica sebifera. A good deal of research has been published on the

invasive potential of T. sebifera in the Coastal Prairie of Texas and Louisiana (e.g., Bruce et al. 1997; Barril-

leaux & Grace 2000). However, little attention has been given to other species that appear to be invasive in

established vegetation in the Coastal Prairie region.

Rosen, Flora of Nash Prairie, Brazoria County, Texas 685

Taste 3. Distribution of grass taxa and C, photosynthetic pathway by Subfamily for Nash Prairie, Brazoria County, Texas. Sub-

familial classification follows the Catalogue of New World Grasses (Judziewicz et al. 2000; Peterson et al. 2001; Soreng et al.

2003; Zuloaga et al. 2003).

Subfamily Genera Species %C, Species

Introduced Native Introduced Native

Aristidoideae 1 0 1 - 100

Chloridoideae 7 2 12 100 100

Ehrhartoideae ] 0 ] - 100

Panicoideae 17 5 35 100 80

Pooideae 11 4 10 0 0

CONCLUSIONS

This research reports numerous taxa not previously cataloged in early descriptions of the Coastal Prairie

(e.g., Bray 1906; Tharp 1926) or more recent accounts of the flora of the Upper Coastal Prairie (e.g., Butler

1979; Smeins et al. 1991). Indeed, the flora of the Coastal Prairie probably includes many more species not

reported herein or previously by other authors. Given the overall geological and environmental diversity of

the region (Smeins et al. 1991), the flora might approach ca. 600 species (Allain & Johnson 1997; Allen et

al. 2001; D. J. Rosen, unpublished data & personal observation). Only continued intensive floristic research

in remnants of Coastal Prairie will provide an accurate estimate. I believe that both historic and potential

losses of botanical diversity in the Coastal Prairie are greater than previously thought. As this research has

indicated, numerous plant taxa with their distributions centered in other habitat types extend into the Coastal

Prairie. It has been suggested that the conservation of potentially genetically distinct populations is at least

as important as the conservation of an entire species (Ehrlich 1988). Expanding populations in the Upper

Coastal Prairie region, rapid urbanization of rural areas, a continued poor understanding of the flora, and

the lack of an aggressive strategy for Coastal Prairie conservation could result in the loss of the remaining

large remnants. The approach of ex-situ preservation of a few endangered Coastal Prairie plant and animal

species will have little environmental value in the absence of large areas of suitable habitat (Jordan 1988).

APPENDIX 1

ANNOTATED CHECKLIST OF SPECIES

Families are arranged alphabetically, beginning with monocots, and followed by eudicots, following the

classification presented in APGII (2003). Genera, species, and infraspecific names are arranged alphabetically

under families and their classification generally follows Jones et al. (1997) with a few exceptions. Recent

molecular data supports the recognition of Dichanthelium as a genus distinct from Panicum (Freckmann

& Lelong 2003). I also follow APGII (2003) in considering Apocynaceae to include Asclepiadaceae. Some

species names are preceded by special symbols to indicate nativity and conservation interest as follows:

(1) non-native species are indicated by an asterisk (*), based on review of Hatch et al. (1990) and Correll

and Johnston (1970); (2) endemic, rare, or regionally rare species are indicated by a superscript dagger (+),

based on review of Correll and Johnston (1970), Turner et al. (20032; 2003b), or personal experience; and

(3) amphitropical species are indicated by a superscript bold capital AT. Synonyms, if considered useful,

are provided in brackets following the species name. Following each name is an abbreviation from Palmer

et al. (1995), representing one of the following subjective estimates of the relative abundance of that species

in the particular habitat(s) where it was collected: r = rare (very difficult to find and limited to one or very

few locations or uncommon habitats); i = infrequent (difficult to find with few individuals or colonies but

found in several locations); o = occasional (widely scattered but not difficult to find); f = frequent (easily

seen or found in one or more common habitats but not dominant in any common habitat); and a = abundant

(dominant or codominant in one or more common habitats). Following the relative abundance, the habitat(s)

686 Journal of the Botanical R h Institute of Texas 1(1)

where that species is typically found is indicated by the following general categories: Prairie = Infrequently

flooded to upland elevation grassland throughout the study area; Wetlands - All wetland sites, including

deep, seasonally flooded depressions and abandoned stream meanders and temporarily flooded inter-mound

flats and lowland prairie; Pimple mounds - Distinctive circular mounds of various elevations and diameters

that occur throughout the site; Disturbed - Sites where the native sod has been disturbed, including dirt

roads, roadside ditches, and fence-lines with encroaching woody vegetation.

The abundances and habitat preferences indicated are based on my observations during the duration

of this research. They may reflect the response of each species to management of Nash Prairie and recent

rainfall conditions, but they are not meant to indicate the expected dominance or habitat preference for each

species throughout the Coastal Prairie. Collection numbers are mine with the exceptions of a few specimens

collected by William R. Carr (WRC). Annotations for grasses are followed with their designation as either

C, or C, following Waller and Lewis (1979). Global and state conservation ranks following Carr (2004) are

provided for some rare taxa.

MONOCOTS

Agavaceae

Manfreda virginica (L.) Salisb. ex Rose, prairie, i, 3004

Alismataceae

Sagittaria graminea Michx. subsp. graminea, wetlands, f,

2707

Sagittaria papillosa Buchenau, wetlands, i, 2859

lliaceae

Nothoscordum bivalve (L.) Britton, prairie, f, 2690

Amaryllidaceae

*Cooperia traubii W. Hayw., prairie, i, 3155, G3QS3

Hymenocallis liriosme (Raf) Shinners, wetlands, o, 2713

Commelinaceae

Commelina erecta L. var. deamiana Fernald, disturbed, i,

923

Tradescantia ohiensis Raf., disturbed, pimple mounds, i,

Cyperaceae

Carex bushii Mack., prairie, i, 2875

Carex cherokeénsis Schwein., prairie, o, 2717

Carex complanata Torr. & Hook, prairie, i, 2746

Carex festucacea Schkuhr ex Willd., wetlands, i, 2763

Carex flaccosperma Dewey, prairie, i, 2765

Carex leavenworthii Dewey, wetlands, r, 3659

Carex longii Mack., wetlands, i, 2848

Carex meadii Dewey, prairie, pimple mounds, a, 2688

Carex microdonta Torr. & Hook., prairie, i, 4048

Carex tetrastachya Scheele, wetlands, o, 2766

Carex triangularis Boeck., wetlands, o, 2764

*Cyperus articulatus L., wetlands, i, 2899

*ATCyperus cephalanthus Torr. & Hook., wetlands, r, 2950,

G2QS1

Cyperus croceus Vahl, disturbed, pimple mounds, i, 2949

*ATCyperus drummondii Torr. 8 Hook., wetlands, o, 2631

Cyperus echinatus (L.) Aloh. Wood, prairie, o, 3013

*Cyperus entrerianus Boeck., disturbed, i, 2960

Cyperus fraternus Kunth, prairie, r, 2896

*Cyperus haspan L., wetlands, 0, 2676

Cyperus pseudovegetus Steud. var. pseudovegetus, wetlands,

j 8

Cyperus reflexus Vahl, prairie, r, 2865

Cyperus retrorsus Chapm. var. retrorsus, pimple mounds, i,

3586

Cyperus virens Michx. var. virens, wetlands, o, 2900

*Eleocharis compressa Sull. var. acutisquamata (Buckley) S.G.

Sm., wetlands, f, 2911

Eleocharis microcarpa Torr. var. microcarpa, wetlands, i, 2832

Eleocharis montana (Kunth) Roem. & Schult., wetlands, i,

2846

Eleocharis palustris (L.) Roem. & Schult., wetlands, r, 2922

Eleocharis quadrangulata (Michx.) Roem. & Schult., wetlands,

i, 298

*Eleocharis wolfii (A. Gray) A. Gray ex Britton, wetlands, r,

2910

Fimbristylis puberula (Michx.) Vahl var. puberula, prairie, a,

2815

Rhynchospora caduca Elliott, wetlands, f, 2907

Rhynchospora corniculata (Lam.) A. Gray, wetlands, o, 2839

Rhynchospora globularis (Chapm.) Small, wetlands, f, 2816

Rhynchospora recognita (Gale) Kral, prairie, o, 2957

Scleria ciliata Michx. var. elliottii (Chapm.) Fernald, wetlands,

229/10

*Scleria muhlenbergii Steud., wetlands, r, 3122

Scleria oligantha Michx., prairie, i, 2840

Scleria pauciflora Muhl. ex Willd. var. pauciflora, prairie, o,

Hypoxidaceae

Hypoxis hirsuta (L.) Coville, pimple mounds, prairie, i, 2698

Iridaceae

Herbertia lahue (Molina) Goldblatt, prairie, i, 2755

Iris brevicaulis Raf., wetlands, r, 2852

Sisyrinchium angustifolium Mill., disturbed, i, 2825

Sisyrinchium minus Engelm. & A. Gray, disturbed, i, 2749

Juncaceae

Juncus acuminatus Michx., wetlands, o, 2871

Juncus brachycarpus Engelm., wetlands, o, 2818

Juncus effusus L. var. solutus Fernald & Wiegand, wetlands,

i, 2847

Rosen, Flora of Nash Prairie, Brazoria County, Texas

* uncus elliottii Chapm. var. elliottii, wetlands, i, 2817

Juncus marginatus Rostk., wetlands, o, 2814

Juncus nodatus Coville, wetlands, o, 2918

Juncus tenuis Willd. var. dichotomus (Elliott) Alph. Wood,

wetlands, i, 2854

Marantaceae

Thalia dealbata Fraser ex Roscoe, wetlands, i, 3372

Orchidaceae

Spiranthes vernalis Engelm. & A. Gray, prairie, i, 2945

Poaceae

Agrostis elliottiana Schult., pimple mounds, o, 2743, C,

Agrostis hyemalis (Walter) Britton, Sterns & Poggenb. var.

hyemalis, pimple mounds, f, 2738, C

Andropogon gerardii Vitman subsp. gerardii, prairie, f, 3120

Me EI glomeratus (Walter) Britton, Sterns & Poggenb.

var. glomeratus, wetlands, i, 3148, C,

Andropogon ternarius Michx. var. ternarius, prairie, f, 2662, C,

Aristida purpurascens Poir. var. purpurascens, prairie, o, 2673,

arenes fissifolius (Raddi) Kuhlm., prairie, pimple mounds,

wetlands, f, 2944, C

Axonopus furcatus (Fluggé) Hitchc., wetlands, i, 3152, C,

*Bothriochloa exaristata (Nash) Henrard, prairie, o, 3151,

353, E

Bothriochloa longipaniculata (Gould) Allred & Gould, dis-

turbed, prairie, i, 3012, C

Bouteloua curtipendula (Michx.) Torr. var. curtipendula, prairie,

[2128€

*Briza minor " POlsuurbeG: i, 2770, C;

*Chloris canterae Arechav. var. a disturbed, i, 2758,

*Cynodon dactylon (L.) Pers. var. dactylon, disturbed, i, 3149,

Dichanthehum aciculare (Desv. ex Poir) Gould & C.A. Clark

subsp. aciculare, prairie, i, 2711, C

Dichanthelium aciculare (Desv. ex Poir) Gould & C.A. Clark

subsp. angustifolium (Elliot) Freckmann & Lelong, prairie,

2741€.

Dichanthelium acuminatum (Sw.) Gould & C.A. Clark subsp.

acuminatum, prairie, f, 2740, C

Dichanthelium dichotomum (L) Gould subsp. dichotomum,

prairie, i, 2912,

Dichanthelium E COEO DOR (Elliott) Gould, prairie, f, 2739,

*Digitaria ciliaris (Retz.) Koeler var. ciliaris, disturbed, i, 2988, C,

Digitaria cognata (Schult.) Pilg., pimple mounds, o, 3583, C,

Elymus virginicus L. var. virginicus, disturbed, r, 3658, C.

Fragrostis elliottii S. Watson, pimple mounds, i, 3582, C

Eragrostis intermedia Hitchc. var. intermedia , prairie, o, 2952,

nos lugens Nees, pimple mounds, o, 3584, C

Eragrostis refracta (Muhl. ex Elliott) Scribn., prairie, i, 3124, C,

Eragrostis spectabilis (Pursh) Steud., prairie, o, 3583, €,

Eriochloa contracta Hitchc, disturbed, i, 3008, C,

*Hordeum pusillum Nutt., disturbed, i, 2775, C.

687

Leersia hexandra Sw., wetlands, r, 3042, C.

Limnodea arkansana (Nutt.) L.H. Dewey, disturbed, i, 3376,

*l olium arundinaceum (Schreb.) Darbysh., disturbed, i, 3348,

*Lolium perenne L. var. perenne, disturbed, 027056:

Mnesithea cylindrica (Michx.) de Koning & Sosef [Sy = Coelora-

chis cylindrica (Michx.) Nash], wetlands, 129786,

Muhlenbergia capillaris (Lam.) Trin., prairie, o, 3110, €,

Nassella leucotricha (Trin. € Rupr.) RW. Pohl, disturbed, i,

Panicum anceps Michx. var. anceps, wetlands, i, 3471, C,

ATPanicum bergii Arechav. [Sy = Panicum pilcomayense Hack.],

pimple mounds, i, 3464, C

Panicum dichotomiflorum Michx. subsp. dichotomiflorum,

disturbed, o, 3102, C,

Panicum hemitomon Schult„wetlands, a, 2908, C,

Panicum rigidulum Bosc ex Nees subsp. cua wetlands,

0; 26072, C

Panicum tenerum Beyr. ex Trin., wetlands, o, 2872, C,

Panicum virgatum L., wetlands, a, 3118, C

Paspalidium geminatum (Forssk.) Stapf var. geminatum, we-

tlands, i, 2951, C

Paspalum denticulatum Trin. [Sy = P. lividum Trin.], wetlands,

f, 3423,C,

*Paspalum dilatatum Poir., disturbed, i, 2895, C,

Paspalum floridanum Michx., wetlands, o, 3014, C,

*Paspalum notatum Fluggé, disturbed, i, 2989, C,

Paspalum plicatulum Michx. var. plicatulum, prairie, a, 2905,

PREDA praecox Walter, wetlands, r, 3119, C,

Paspalum setaceum Michx. var. muhlenbergii (Nash) D.J. Banks,

prairie, i, 2954, C,

*Paspalum urvillei ao disturbed, i, 2894, C,

A'Paspalum wrightii Hitchc. & Chase [Sy =P. texanum Swallen],

wetlands, o, 2990, C,

^'Phalaris angusta Nees ex Trin., disturbed, i, 2774, C,

Phalaris caroliniana Walter, disturbed, o, 2769, C.

*Poa annua L, disturbed, i, 2694, C,

Schizachyrium scoparium (Michx.) Nash var. scoparium, prairie,

a 3109, €.

Setaria parviflora (Poir.) Kerguélen, disturbed, prairie, i, 2948,

asia nutans (L.) Nash, prairie, a, 3121, C,

*Sorghum halepense (L.) Pers., disturbed, i, 2893, C

Sphenopholis obtusata (Michx.) Scribn. var. obtusata, dis-

turbed, i2855, C

Sporobolus compositus (Poir. Merr. var. compositus, prairie,

i, 3877,

Sporobolus pompous (Poir.) Merr. var. macer (Trin.) Kartesz &

Gandhi, pimple mounds, i, 3153, C,

Sporobolus indicus (L.) R. Br. var. e disturbed, o, 2706,

li silveanus Swallen, prairie, r, 360

Steinchisma hians (Elliott) Nash, wetlands, o, 2946, C,

Tridens strictus (Nutt.) Nash, wetlands, f, 3541, C,

Tripsacum dactyloides (L.) L. var. dactyloides, wetlands, r,

688

Urochloa platyphylla (Munro ex C. Wright) R.D. Webster,

disturbed, i, 3879, C

Vulpia octoflora (Walter) Rydb. var. octoflora, pimple mounds,

o, 2744, C,

Pontederiaceae

Pontederia cordata L. var. cordata, wetlands, i, 2868

EUDICOTS

Acanthaceae

Hygrophila lacustris (Cham. €: Schltdl.) Nees, wetlands, r,

3040

Justicia ovata (Walter) Lindau var. lanceolata (Chapm.) R.W.

Long, wetlands, r, 311

Ruellia humilis Nutt. var. depauperata Tharp & F.A. Barkley,

prairie, o, 3043

Ruellia nudiflora (Engelm. & A. Gray) Urb. var. nudiflora, dis-

turbed, o, 3006

Anacardiaceae

Toxicodendron radicans (L.) Kuntze, disturbed, i, 2877

Apiaceae

Chaerophyllum tainturieri Hook. var. dasycarpum (Nutt) S.

Watson, disturbed, r, 3603

Daucus pusillus Michx., mounds, r, 4090

Eryngium yuccifolium Michx., prairie, f, 2982

Hydrocotyle umbellata L., wetlands, i, 3375

Limnosciadium pinnatum (DC.) Mathias & Constance, prairie,

i 2915

Polytaenia nuttallii DC., prairie, o, 2936

Sanicula canadensis L. var. canadensis, disturbed, r, 2856

Apocynaceae (incl. Asclepiadaceae)

tAmsonia repens Shinners, wetlands, o, 2703

tAsclepias linearis Scheele, wetlands, r, 3115

Asclepias longifolia Michx. subsp. longifolia, wetlands, i, 2943

Asclepias verticillata L., prairie, o, 2937

Asclepias viridis Walter, disturbed, i, 2833

Asteraceae

Acmella oppositifolia (Lam.) R.K. Jansen var. repens (Walter) R.

K. Jansen, wetlands, i, 2861

Ambrosia psilostachya DC., prairie, o, 2666

Ambrosia trifida L. var. texana Scheele, disturbed, r, 3596

Arnoglossum plantagineum Raf, prairie, o, 281

Astranthium integrifolium (Michx.) Nutt. subsp. ciliatum (Raf)

eJong, pimple mounds, r, 3607

Boltonia diffusa Elliott var. diffusa, wetlands, f, 3106

Calyptocarpus vialis Less., disturbed, r, 3422

Cirsium horridulum Michx. var. elliottii Torr. & A. Gray, prairie,

r, 2786

Conoclinum coelestinum (L.) DC., wetlands, o, 3129

Conyza canadensis (L.) Cronquist var. glabrata (A. Gray) Cron-

quist, pimple mounds, i, 3924

Coreopsis tinctoria Nutt. var. tinctoria, dsturbed, o, 2892

Erigeron philadelphicus L., disturbed, i, 3347

Erigeron tenuis Torr. & A. Gray, prairie, f, 2708

Eupatorium lancifolium (Torr. & A. Gray) Small, prairie, r, 3925

Eupatorium serotinum Michx., disturbed, i, 3472

Euthamia gymnospermoides Greene, wetlands, f, 2663

(09)

£sthka Dat o ID L

Journal of

titute of Texas 1(1)

Euthamia leptocephala (Torr. & A. Gray) Greene ex Porter &

Britton, prairie, i, 3543

Gamochaeta purpurea (L.) Cabrera [Sy 2 Gnaphalium purpu-

reum L.], disturbed, o, 2751

Gutierrezia texana (DC.) Torr. & A. Gray var. texana, prairie, i,

Helenium amarum (Raf) H. Rock var. amarum, disturbed, i,

2669

Helenium flexuosum Raf., wetlands, o, 2897

Helianthus angustifolius L., wetlands, a, 3469

Helianthus maximiliani Schrad., wetlands, r, 3592

*Hypochaeris microcephala (Sch. Bip.) Cabrera var. albiflora

(Kuntze) Cabrera , disturbed, i, 2748

lva annua L., prairie, r, 3931

Krigia cespitosa (Raf) K.L. Chambers, pimple mounds, r, 3609

Krigia dandelion (L.) Nutt., pimple mounds, i, 2696

Liatris acidota Engelm. & A. Gray, prairie, f, 3111

*Liatris bracteata Gaiser, prairie, r, 3101, G2G 35253

Liatris pycnostachya Michx. , prairie, f, 2904

Mikania scandens (L.) Willd. , wetlands, i, 3117

Packera glabella (Poir.) C. Jeffrey, prairie, i, 2716

Pityopsis graminifolia (Michx.) Nutt., pimple mounds, r, 3660

Pluchea baccharis (Mill) Pruski, wetlands, i, 3928

Pyrrhopappus pauciflorus (D. Don) DC., disturbed, i, 2845

Ratibida columnifera (Nutt.) Wooton & Standl., disturbed, r,

Rudbeckia hirta L. var. angustifolia (T.V. Moore) Perdue, pimple

mounds, i, 2834

tRudbeckia texana (Perdue) P.B. Cox & Urbatsch, wetlands,

f, 2983

Silphium gracile A. Gray, prairie, pimple mounds, f, 2822

Solidago canadensis L. var. scabra (Muhl. ex Willd.) Torr. & A.

Gray, prairie, i, 3544

Solidago stricta Aiton, prairie, pimple mounds, f, 2671

Solidago tortifolia Elliott, prairie, pimple mounds, f, 3610

ATSoliva sessilis Ruiz & Pav., disturbed, r, 2867

Symphyotrichum dumosum (L.) G. L. Nesom, prairie, pimple

mounds, i, 2664

Symphyotrichum ericoides (L.) G. L. Nesom var. ericoides, prairie,

pimple mounds, i, 2665

Symphyotrichum lanceolatum (Willd.) G. L. Nesom var. lanceo-

latum, wetlands, o, 3589

Symphyotrichum patens (Aiton) G.L. Nesom var. gracile (Hook.)

G. L. Nesom, prairie, r, 3597

Symphyotrichum pratense (Raf) G. L. Nesom, prairie, pimple

mounds, r, 3104

Vernonia missurica Raf., wetlands, f, 3927

Boraginaceae

Lithospermum incisum Lehm., prairie, r, 2715

Myosotis macrosperma Engelm., wetlands, r, 3656

Brassicaceae

Lepidium densiflorum Schrad. var. densiflorum, disturbed, i,

3349

Lepidium virginicum L. var. medium (Greene) C.L. Hitchc.,

disturbed, o, 2823

Lepidium virginicum L. var. virginicum, disturbed, o, 2783

Rosen, Flora of Nash Prairie, Brazoria County, Texas

Callitrichaceae

Callitriche peploides Nutt., wetlands, i, 2761

Campanulaceae

Lobelia puberula Michx., prairie, i, 2667

Triodanis lamprosperma McVaugh, disturbed, i, 2874

dos due ) rie var. biflora (Ruiz & Pav.) Bradley,

e, pimple mounds, o

me perfoliata (L.) EU var. perfoliata, prairie, pimple

mounds, o, 2736

Caryophyllaceae

Cerastium glomeratum Thuill., prairie, i, 2709

Cistaceae

Lechea mucronata Raf., prairie, i, 2986

Clusiacea

Hypericum D (L.) Crantz , prairie, r, 3712

Convolvulaceae

Dichondra carolinensis Michx., prairie, pimple mounds, i

2750

Evolvulus sericeus Sw. var. sericeus, prairie, i, 3011

Ipomoea cordatotriloba Dennst., disturbed, i, 3007

Commaccae

us drummondii C.A. Mey., disturbed, i, 2880

Cuscutaceae

Cuscuta pentagona Engelm., wetlands, r, 3041

Droseraceae

Drosera brevifolia Pursh, pimple mounds, i, 2734

Ebenaceae

Diospyros virginiana L., disturbed, r, 2853

Euphorbiaceae

Acalypha gracilens A. Gray var. gracilens, prairie, r, 22178

(WRC)

*Caperonia palustris (L.) A. St.-Hil., wetlands, r, 3930

Chamaesyce maculata (L.) Small, disturbed, o, 3127

Croton capitatus Michx. var. bod (Engelm. & A. Gray)

Müll. Arg., disturbed, o

Croton glandulosus L. var. eu MT Arg. prairie, pimple

mounds, o, 3045

Euphorbia bicolor Engelm. & A. Gray, disturbed, o, 3046

Euphorbia spathulata Lam., prairie, pimple mounds, o, 2757

*Fuphorbia texana Boiss., prairie, pimple mounds, i, 2737

Tragia betonicifolia Nutt., pimple mounds, o, 2941

*Triadica sebifera (L.) Small (2Sapium sebiferum (L.) Roxb.),

wetlands, f, 2921

Fabaceae

Acacia ang (Mill) Kuntze var. hirta (Nutt. ex Torr. & A.

Gray) B.L. Rob., prairie, pimple mounds, o, 2891

Baptisia bracteata Muhl. ex Elliott var. leucophaea (Nutt)

Kartesz & Gandhi, prairie, i, 2902

Baptisia sphaerocarpa Nutt., prairie, f, 2837

Centrosema virginianum (L.) Benth., disturbed, i, 3015

Chamaecrista fasciculata (Michx.) Greene, disturbed, 0, 2981

Dalea candida Willd. var. candida, prairie, i, 2959

Desmodium ciliare (Muhl. ex Willd.) DC. var. ciliare, pimple

mounds, 0, 3105

689

Galactia marginalis Benth., pimple mounds, r, 3112

Lathyrus pusillus Elliott, disturbed, i, 2753

*Medicago lupulina L., disturbed, o, 2693

*Medicago minima (L.) L., disturbed, i, 2844

*Melilotus indicus (L.) All., disturbed, i, 2843

Mimosa hystricina (Small ex Britton & Rose) B.L. Turner, prairie,

0,2835

Mimosa nuttallii (DC.) B.L. Turner, prairie, o, 2938

Neptunia lutea (Leavenw.) Benth., wetlands, o, 2939

Neptunia pubescens Benth. var. pubescens, wetlands, r, 22175

(WRC

Sesbania drummondii (Rydb.) Cory, wetlands, r, 3044

Stylosanthes biflora (L.) Britton, Sterns € Poggenb, prairie,

12959

Tephrosia onobrychoides Nutt., pimple mounds, o, 2942

*Trifolium repens L. var. repens, disturbed, i, 2890

Vicia ludoviciana Nutt. ex Torr. & A. Gray subsp. ludoviciana,

disturbed, i, 3604

Fagaceae

Quercus virginiana Mill. var. virginiana, disturbed, r, 2987

Gentianaceae

Sabatia campestris Nutt., pimple mounds, prairie, o, 2830

Geraniaceae

Geranium carolinianum L. var. carolinianum, disturbed, o,

Haloragaceae

ATProserpinaca palustris L. var. amblyogona Fernald, wetlands,

Hydrophyllaceae

Hydrolea ovata Nutt. ex Choisy, wetlands, f, 2999

Krameriaceae

Krameria lanceolata Torr., prairie, r, 3420

Lamiaceae

Hedeoma hispidum Pursh, prairie mounds, i, 2756

Physostegia intermedia (Nutt.) Engelm. & A. Gray, wetlands,

o, 2850

Salvia lyrata L., disturbed, i, 3346

Scutellaria parvula Michx. var. parvula, pimple mounds, o,

Linaceae

Linum medium (Planch.) Britton var. texanum (Planch.) Fernald,

prairie, f, 2903

Lythraceae

A? Cuphea glutinosa Cham. & Schltdl., prairie, r, 3714

Lythrum alatum Pursh var. lanceolatum (Elliott) Rothr., wet-

lands, i, 2947

Malvaceae

Callirhoë involucrata (Torr. & A. Gray) A. Gray var. lineariloba

(Torr. & A. Gray) A. Gray, prairie, o, 2836

Modiola caroliniana (L.) G. Don, disturbed, i, 2776

Sida ciliaris L. , disturbed, i, 3010

Sida rhombifolia L. , disturbed, i, 3009

690

Melastomataceae

Rhexia mariana L. var. mariana, wetlands, i, 2984

Myricaceae

Morella cerifera (L) Small, prairie, i, 3264

Onagraceae

Gaura longiflora Spach, disturbed, o, 3039

Ludwigia glandulosa Walter, wetlands, o, 2901

Ludwigia linearis Walter, wetlands, i, 2985

Oenothera laciniata Hill, pimple mounds, i, 2735

Oenothera linifolia Nutt., prairie, r, 2785

Oenothera speciosa Nutt., disturbed, o, 2782

Oxalidaceae

Oxalis dillenii Jacq., disturbed, o, 2781

Oxalis violacea L., prairie, r, 3154

Passifloraceae

Passiflora incarnata L., disturbed, o, 2914

Plantaginaceae

Plantago aristata Michx., disturbed, o, 2841

Plantago virginica L., disturbed, i, 2826

Polemoniaceae

Phlox cuspidata Scheele, prairie, r, 2787

Polygalaceae

Polygala incarnata L., pimple mounds, prairie, f, 2828

Polygonaceae

Polygonum hydropiperoides Michx., wetlands, f, 2674

Rumex chrysocarpus Moris, disturbed, i, 2883

Primulaceae

Anagallis minima (L.) E.H.L. Krause, pimple mounds, f, 2714

Ranunculaceae

Anemone berlandieri Pritz., pimple mounds, i, 2699

Delphinium carolinianum Walter, pimple mounds, i, 2821

Ranunculus laxicaulis (Torr. & Gray) Darby, wetlands, r, 3605

Ranunculus pusillus Poir., wetlands, i, 2760

*Thalictrum texanum (A. Gray) Small, pimple mounds, r, 2701,

fal, Dat o ID L

Journal of

titute of Texas 1(1)

Rubus argutus Link, disturbed, i, 2876

RUBIACEAE

Cephalanthus occidentalis L. var. californicus Benth., wetlands,

r, 3933

Diodia virginiana L. var. virginiana , wetlands, i, 2920

Galium obtusum Bigelow subsp. obtusum, wetlands, i, 2759

Houstonia pusilla Schoepf, pimple mounds, f, 2689

Rutaceae

Zanthoxylum clava-herculis L., disturbed, i, 3371

Salicaceae

Salix nigra Marshall, wetlands, r, 2869

Saxifragaceae

AT! epuropetalon spathulatum Elliott, pimple mounds, r, 2702

Scrophulariaceae

Agalinis heterophylla (Nutt.) Small ex Britton, prairie, f, 3107

Agalinis viridis (Small) Pennell, prairie, o, 3103

Buchnera americana L., pimple mounds, prairie, o, 2831

Castilleja indivisa Engelm., pimple mounds, prairie, f, 2692

Gratiola virginiana L. var. virginiana, wetlands, i, 2762

Mecardonia acuminata (Walter) Small var. acuminata, we-

tlands, r, 3123

Nuttallanthus canadensis (L.) D.A. Sutton [Sy = Linaria canaden-

sis (L) Dum. Cours], disturbed, i, 2704

Solanaceae

Physalis cinerascens (Dunal) Hitchc. var. cinerascens, pimple

mounds, i, 2771

Ulmaceae

Ulmus americana L., wetlands, r, 3373

Valerianaceae

Valerianella woodsiana (Torr. & A. Gray) Walp., disturbed, o,

2780

Verbenaceae

Phyla nodiflora (L.) Greene , disturbed, i, 2913

Verbena halei Small , prairie, o, 2754

*Verbena brasiliensis Vell., disturbed, i, 2863

Verbena xutha Lehm. prairie, r, 3419

saceae

Prunus angustifolia Marsh. var. angustifolia, disturbed, i,

2879

ACKNOWLEDGMENTS

I am grateful to the KNG Ranch for granting access to Nash Prairie. Thanks to Larry Brown, J. Wipff, Guy

Nesom, and Tony Reznicek for annotating difficult specimens. Thanks to Larry Allain and Bill Carr for com-

ments on an earlier version of this manuscript. Monique Reed and Michael MacRoberts provided helpful

comments and suggestions that improved the manuscript. I offer special thanks to Bill Carr for inviting me

on the inaugural trip to Nash Prairie, and to Peter and Susan Conaty for their support, interest, enthusiasm

and companionship in the field, and for their commitment to conservation.

REFERENCES

ALLAIN, L. and S. JOHNSON. 1997. The prairies of coastal Texas and Acadiana. Wildflower 13:42-45.

ALLEN, C.M., M. VipniNe, B. Borsarl, and L. Attain. 2001. Vascular flora of the Cajun prairie of southwestern Louisiana.

Proc. 17th N. A. Prairie Conference. Pp. 35-41.

Rosen, Flora of Nash Prairie, Brazoria County, Texas 691

ALLEN, C.M. and DW. Hau. 2003. Paspalum L. In: M. A. Barkworth, K.M. Capels, S. Long, and M.B. Piep, eds. Flora of

North America North of Mexico. Vol. 25. Oxford University Press, NY. Pp. 566-599.

ALLRED, KW. 2003. Bothriochloa Kuntze. In: M.A. Barkworth, K.M. Capels, S. Long, and M.B. Piep, eds. Flora of North

America North of Mexico. Vol. 25. Oxford University Press, NY. Pp. 639-647.

APGII. 2003. An update of the Angiosperm Phylogeny Group classification for the orders and families of flower-

ing plants: APG Il. Bot. J. Linn. Soc. 141:399-436.

BARRILLEAUX, T.C. and J.B. Grace. 2000. Growth and invasive potential of Sapium sebiferum (Euphorbiaceae) within

the Coastal Prairie Region: The effects of soil and moisture regime. Amer. J. Bot. 87:1099-1106.

Bray, W.L. 1906. Distribution and adaptation of the vegetation of Texas. Univ. Texas Bull. 82(10).

Bruce, K.A., G.N. Cameron, PA. HARCOMBE, and G. JugiNskv. 1997. Introduction, impact on native habitats, and manage-

ment of a woody invader, the Chinese tallow tree, Sapium sebiferum (L.) Roxb. Nat. Areas J. 17:255-260.

BurLER A.C. 1979. Mima mound grasslands of the upper coastal prairie of Texas. M.S. Thesis. Texas A&M University.

College Station.

Cann, W.R. 2004. An Annotated List of the G3/T3 and Rarer Plant Taxa of Texas. Texas Conservation Data Center.

The Nature Conservancy of Texas. San Antonio, TX.

Carter, R., D.K. ALEXANDER, C.T. Bryson, and A. Lazari. 1999. The taxonomy of Cyperus virens and Cyperus drummondii

in the Southeastern United States. Sida 18:1049-1063.

Carter, R. and N. Mclnnis. 1993. Status report on Cyperus cephalanthus. Unpublished report submitted to U.S. Fish €:

Wildlife Service, Jackson, MS.

Correll, D.S. and M.C. Johnston. 1970. Manual of the vascular plants of Texas. Texas Research Foundation,

Renner.

CRENWELGE, G.W., J.D. Crour, E.L. GRIFFIN, M.L. GOLDEN, and J.K. Baker. 1981. Soil Survey of Brazoria County, Texas. United

States Dept. of Agriculture, Soil Conservation Service, Washington, D.C.

DiAMOND, D.D. and FE. Smeins. 1984. Remnant grassland vegetation and ecological affinities of the upper coastal

prairie of Texas. SouthW. Naturalist 29:32 1-334.

Diamonb, D.D. and F.E. Smeins. 1985. Composition, classification, and species response patterns of remnant tall

grass prairies in Texas. Amer. Midl. Naturalist 113:294—308.

DIAMOND, D.D. and F.E. Smeins. 1988. Gradient analysis of remnant true and upper coastal prairie grasslands of

North America. Canad. J. Bot. 66:2152-2161.

EHRLICH, PR. 1988. The loss of diversity; causes and consequences. In: E.O. Wilson, ed. Biodiversity. National Acad-

emy Press, Washington D.C. Pp. 21-27.

FENNEMAN, N.M. 1928. Physiographic divisions of the United States. Annals Association American Geographers,

18:261-353.

FLORA OF NORTH AMERICA EDITORIAL Committee (eds.). 2000. Flora of North America North of Mexico. Vol. 22. Magno-

liophyta: Commelinidae (in part), and Zingiberidae. Oxford Univ. Press, NY.

FLORA OF NORTH AMERICA EDITORIAL Committee (eds.). 2002a. Flora of North America North of Mexico. Vol. 23. Magno-

liophyta: Commelinidae (in part), Cyperaceae. Oxford Univ. Press, NY.

FLORA OF NORTH AMERICA EDITORIAL Committee (eds.). 2002b. Flora of North America North of Mexico. Vol. 26. Magno-

liophyta: Lilidae: Liliales and Orchidales. Oxford Univ. Press, NY.

FLORA OF NORTH AMERICA EDITORIAL Committee (eds.). 2003. Flora of North America North of Mexico. Vol. 25. Magno-

liophyta: Commelinidae (in part), Poaceae, part 2. Oxford Univ. Press, NY..

FRECKMANN, R.W. and M.G. LeLONG. 2003. Dichanthelium (Hitchc. € Chase) Gould. In: M.A. Barkworth, K.M. Capels,

S. Long, and M.B. Piep, eds. Flora of North America North of Mexico. Vol. 25. Oxford University Press, NY. Pp.

406—450.

Goop, R. 1953. The geography of flowering plants. Longmans, Green and Co., London.

GouLb, F.W. 1975. The grasses of Texas. Texas A&M Univ. Press, College Station, TX.

Grace, J.B., L. ALLAIN, and C. ALLEN. 2000. Vegetation associations in a rare community type - coastal tallgrass prairie.

Pl. Ecol. 147:105-115.

692 Journal of the Botanical R h Institute of Texas 1(1)

Hatch, S.L., K.N. GANDHI and L.E. Brown. 1990. Checklist of the vascular plants of Texas. Texas Agric. Exp. Sta. Bull.

MP-1655, TX.

HouLMcGREN, PK., N.H. HOLMGREN, and L.C. BarnerT. 1990. Index herbariorum Part I: the herbaria of the world. New

York Botanical Garden, Bronx.

IseLy, D. 1990. Vascular flora of the southeastern United States, Volume 3, Part 2, Leguminosae (Fabaceae). Univ.

of North Carolina Press, Chapel Hill.

Jones, S.D., J.K. Wierr, and PM. Montcomery. 1997. Vascular plants of Texas: a comprehensive checklist including

synonymy, bibliography, and index. University of Texas Press, Austin.

JoRDAN, W.R. 1988. Ecological restoration: Reflections on a half-century of experiences at the University of Wis-

consin-Madison Arboretum. In: E.O. Wilson, ed. Biodiversity. National Academy Press, Washington D.C. Pp.

311-316.

JupziEWICZ, E.J., RJ. SORENG, G. DaviDsE, P.M. PETERSON, T.S. FILGUERIAS, and F.O. ZULOAGA. 2000. Catalogue of New World

grasses (Poaceae): |. Subfamilies Anomochlooideae, Bambusoideae, Ehrhartoideae, and Pharoideae. Contr.

U.S. Natl. Herb. 39:1-128.

PALMER, M.W, G.L. Wac, and P. Neat. 1995. Standards for the writing of floras. Bioscience 45:339-345.

PETERSON, P.M., R.J. SORENG, G. DaviDsE, T.S. FILGUERIAS, F.O. ZULOAGA, and EJ. Jubziewicz. 2001. Catalogue of New World

grasses (Poaceae): Il. Subfamily Chloridoideae. Contr. U.S. Natl. Herb. 41:1-255.

Raven, PH. 1963. Amphitropical relationships in the floras of North and South America. Quart. Rev. Biol. 38:151-

Rosen, D.J. 2004. Noteworthy collections of Cyperus drummondii (Cyperaceae) from Texas. Sida 21:495-497,

Rosen, D.J. and B.J. CHRISTOFFERSEN. 2004. Rediscovery of Cyperus cephalanthus (Cyperaceae) in Texas. Phytologia

86:107-109.

Smelns, F.E., D.D. DIAMOND, and C.W. HaNsELkA. 1991. Ch. 13, Coastal Prairie. In: R.T. Coupland, ed. Ecosystems of the

World 8A; Natural grasslands, introduction and Western Hemisphere. Elsevier, Amsterdam. Pp. 269-290.

Sims, P.L. and PG. Risser. 2000. Ch. 9, Grasslands. In: M.G. Barbour and W.D. Billings, eds. North American Terrestrial

Vegetation, 2nd ed. Cambridge University Press, NY.. Pp. 323-356.

Smit, E.B. 1994. Keys to the flora of Arkansas. The University of Arkansas Press, Fayetteville.

SORENG, R.J., PM. Peterson, G. Davidse, E.J. JubziEwicz, F.O. ZULOAGA, T.S. FILGUERIAS, and O. Morrone. 2003. Catalogue of

New World grasses (Poaceae): IV. Subfamily Pooideae. Contr. U.S. Natl. Herb. 48:1—730.

Tuan, B.J. 1926. Structure of Texas vegetation east of the 98th meridian. Univ. Texas Bull. 2606.

THORNTHWAITE, C.W. 1948. An approach toward a rational classification of climate. Geogr. Rev. 38:55-94.

Tucker, G.C., B.G. Marcks, and J.R. Carter. 2003. Cyperus Linnaeus. In: Flora of North America Editorial Committee,

eds. Flora of North America North of Mexico. Vol. 23. Oxford University Press, New York. Pp. 141-191.

TURNER, B.L., H. NicHoLs, G. Denny, and O. Doron. 2003a. Atlas of the vascular plants of Texas. Vol. 1. Sida, Bot. Misc.

24:1-648.

TURNER, B.L., H. NicHoLs, G. Denny, and O. Doron. 2003b. Atlas of the vascular plants of Texas. Vol. 2. Sida, Bot. Misc.

24:649-888.

WALLER S.S. and J.K. Lewis. 1978. Occurrence of C, and C, photosynthetic pathways in North American grasses.

J. Range Managem. 32:12-28.

ZULOAGA, F.O., O. Morrone, G. Davipse, T.S. FILGUERIAS, P.M. Peterson, R.J. SORENG, and E.J. Jupziewicz. 2003. Catalogue of

New World grasses (Poaceae): Ill. Subfamilies Panicoideae, Aristidoideae, Arundinoideae, and Danthonioideae.

Contr. U.S. Natl. Herb. 46:1-662.

THE VASGULAK FLORA OF A AWOODEAND PARK SDE UN

PSL rans COUN ITY. DER

PA. Harcombe, I.S. Elsik, W.W. Pruess L.E. Brown

Department of Ecology and Evolutionary Biology Houston Community College

Rice University, PO Box 1892 1300 Holman, Houston, Texas 77004, USA

Houston, Texas 77251-1892, USA. Herbarium, Spring Branch Science Center

8856 Westview Drive, Houston, Texas 77005, USA.

ABSTRACT

An annotated, vouchered checklist is provided of the vascular plant taxa of a 50 ha segment of Herman Brown Park, Houston, Harris

County, Texas. Four hundred forty eight are documen ted for the site, of which 361 are native and 87 are non native. Weedy species

constitute 5696 of the native species and 6896 of the non natives, but only a few are highly invasive. The native and non native taxa are

predominantly from the southeastern or eastern USA. The non native taxa are predominantly Asian or European in origin, and approxi-

mately 2596 are from Central or South America.

RESUMEN

€ + +41 i E aH 1 1 1 pa

to de 50 ha del Herman Brown Park, Houston,

d nd eo jes En o

Harris County, Texas. Se documentan cuatrocientas cuarenta y ocho taxa para el lugar, de las que 361 son nativas y 87 no nativas. Las

especies de malas hierbas constituyen el 5696 de las especies nativas y el 6896 de las no nativas, pero sólo unas pocas son invasoras. Los

taxa nativos son predominantemente del sureste o del este de Estados Unidos. Los taxa no nativos son principalmente de origen asiático

0 europeo, y aproximadamente el 2596 son de América Central o Sur América.

INTRODUCTION

This inventory was conducted as part of a biological assessment of a potential site for the Botanic Garden

of Houston, TX (BGH). The main goal was to document the flora of an urban natural area and to consider

factors influencing its floristic composition. An additional goal was to provide a baseline for assessment of

biodiversity changes wrought by development as a botanical garden.

Floristic richness of a site should be a function of the size, habitat diversity, and disturbance history of

the site. The number of species should increase with site size and habitat diversity. Disturbance could elimi-

nate some species while providing new habitat for others, so its influence is unclear. The BGH site is small

(50 ha) and site conditions are relatively homogeneous. Negative impacts to the site include timber cutting,

grazing, oilfield activity and pipeline construction, as well as dredging for Hunting Bayou flood control,

and urbanization of the surrounding area. Furthermore, the site appears to have undergone considerable

ecological change, from prairie and riparian woodland to closed forest following cessation of grazing and

burning.

Site location should also affect the relative size of the flora, considering that the pool of potential non

native invaders would be higher in an urban setting. For example, rural nature preserves typically have a

small fraction of non natives. In nearby Texas nature preserves, the values are in the range of 5-696 (Fleming

et al. 2002; Singhurst et al. 2003; MacRoberts et al. 2004; Brown et al. 2005). Few studies have been reported

for urban natural areas in the US, but values range from 2696 (Middlesex Fells, metropolitan Boston; Drayton

and Primack 1996) to 4096 (Pelham Bay Park, New York City: DeCandido et al. 2004). In regional floras the

non native fraction is as high as 4396 (Great Britain: Crawley et al. 1997) or even 6096 for some islands (Wu

et al. 2004). In Central Europe non natives comprise about 4096 of the floras of 54 cities, and the fraction

of non natives increases with city size and decreases with increasing latitude (Pysek 1998). More studies

from American cities are necessary to determine whether similar trends exist.

In addition to being in an urban setting, the site under study is of interest because it contains a remnant

J. Bot. Res. Inst. Texas 1(1): 693 — 711. 2007

694 Journal of the Botanical R h Institute of Texas 1(1)

of a Quercus similis (bottomland post oak) woodland, a vegetation type that is poorly known ecologically

and floristically. Also, it is located near the western boundary of the eastern deciduous forest biome where

forests rapidly give way to woodlands, savannas, prairies and marshes moving west and south. In the

northeast quadrant of Harris County, the forest communities resemble those of the outer coastal plain of the

southeastern US, with an upland matrix of mixed pine-hardwood forest dissected by oak-gum bottomland

hardwood forest. In the southwest quadrant, uplands are predominantly prairie while riparian zones are

often dominated by Quercus virginiana (live-oak), Q. nigra (water oak), Populus deltoides (cottonwood), and

Celtis laevigata (hackberry). The question addressed in this paper is whether an urban natural area near this

ecotone might have elevated floristic richness since sources of post-disturbance colonists are more varied

than they might be fora site within a uniform regional vegetation matrix. In addition, a substantial admixture

of species with Western or Midwestern distributions may also occur.

To summarize, this paper focuses on the question of how the local flora reflects biogeography and human

activity including whether there is a mix of species from different habitats and if the non native species ratio

is high. The paper also establishes a baseline for investigating floristic change in an urban natural area.

SITE AND METHODS

The collection site was the section of Herman Brown Park (City of Houston, Harris County, TX) west of

Hunting Bayou, herein called the Botanic Garden of Houston site. Herman Brown Park was acquired by the

City of Houston in 1980. Land use at the BGH site has been varied. Deed records indicate timber sales in

1869. The site was later acquired by a local rancher and presumably grazed. Ages of canopy trees indicate

that grazing ceased about 50 years ago. There are several abandoned oil wells on the site which appear to

have been in operation from about 1937, when the Clinton Field was discovered, through the 1950s. Hunting

Bayou was dredged for flood control purposes in the late 1940s, and a deep drainage ditch was constructed

running east-west through the site. A residential subdivision was established on the south boundary of the

site in 1952. The site is bordered on the north and west by pasture and industrial lands. The BGH is now

predominantly wooded with two small mowed fields and several open drainage ditches.

The climate of the area is characterized by long hot summers and mild winters. Yearly average precipi-

tation at Hobby Airport (3 km SW of the site) is 134.6 cm, well distributed throughout the year. Average

annual temperature is approximately 21°C. January is the coldest month with mean daily temperatures of

12°C. July is the warmest month with highs averaging 34°C. The growing season is about 270 days (NCDC

2004).

Harris County lies within the Gulf Coastal Plain Physiographic Province (Fenneman 1938), which

is a broad region of low relief stretching from the coast inland 100-150 miles and from Texas to western

Georgia. Streams and rivers of the Gulf Coastal Plain drain into the Gulf of Mexico. The BGH site is situated

on the Beaumont Geologic Formation (Shelby et al. 1993), a broad, flat, poorly drained sedimentary surface

of sands and clays that appeared in the late Pleistocene when sea level was high during a warm interval

(approximately 30-40,000 years ago) within the Wisconsin glacial period (Moore and Aronow 1994). The

surface is cut, sometimes deeply, by shallow, short bayous draining into the Gulf of Mexico.

Houston is located on the edge of the Eastern Deciduous Forest Biome. The northern portion of the city

is part of the Upper West Gulf Coastal Plain ecoregion (Weakley et al. 1998) often referred to as the Piney-

woods (Gould 1975; Correll & Johnston 1970). In this ecoregion, Quercus-Pinus (oak-pine) forests dominate

the landscape; stream and river bottoms are typically bottomland hardwood forest dominated by Quercus

and Liquidambar. The southern portion of the city is part of the Gulf Prairies and Marshes ecoregion. In this

ecoregion the original vegetation of the area was likely to have been mostly prairie, with bands of Quercus-

Pinus forest on bluffs along bayous (Gould 1975; Correll & Johnston 1970) and Ulmus americana, Fraxinus

pennsylvanica, and Celtis laevigata forest in the bottoms. Today, where native trees persist along drainages,

Quercus similis Ashe (bottomland post oak) is sometimes one of the dominant oaks.

Three types of forest vegetation were identified at the site. On the flat uplands away from the bayou,

Harcombe et al., Flora of a iland park site, Harris County, Texas 695

Ulmus americana, Fraxinus pennsylvanica, and Celtis laevigata dominate [National Vegetation Classification

System (NVCS) type Ulmus americana- Celtis laevigata-Fraxinus pennsylvanica Forest (NatureServe 2005)].

Ligustrum spp. and Ilex decidua are important shrubs. In the ground layer, coverage by graminoids is high,

especially Carex spp. The second vegetation type occurs on flats and bluffs nearer the bayou. Q. similis and

Pinus taeda are dominant but bands of Quercus phellos follow shallow drains. Ilex decidua and Ilex vomitoria

form dense patches, and non native Ligustrum species are abundant. Coverage of graminoids in the ground

layer is low to moderate. The only NVCS type containing Q. similis as a dominant is the Quercus phellos-Q.

similis/Crataegus marshallii-C. spathulata/Chasmanthium laxum Forest (NatureServe 2005). All of these spe-

cies except C. spathulata occur on the site. However, Pinus taeda is a canopy dominant, and C. laxum is of

only modest importance in the ground layer. The likely interpretation is that the site vegetation represents

a seral stage of this NVCS type. The third vegetation type on the site is a narrow riparian strip in the deeply

incised ravine of Hunting Bayou dominated by Platanus occidentalis, Salix nigra, and Populus deltoides.

Soils of the site are clays, clay loam, and fine sandy loams of several series with Lake Charles Clay and

Aldine Very Fine Sandy Loam being the most widespread (SCS 1976). The soils are poorly drained and of

low agricultural productivity.

Plant collections were made during parts of three growing seasons, from Oct 2004 to Dec 2006. The

site was visited sporadically Oct-Dec 2004, at approximately two-week intervals during Feb-Dec 2005, and

occasionally Feb—June 2006. Approximately 700 specimens were collected and deposited at Rice University.

In most cases, the nomenclature follows Jones et al. (1997) and Kartesz and Meacham (1999). Some recent

literature sources were followed, especially some nomenclatural adjustments published in Flora of North

America volumes (FNA 1993+).

Nativity was defined as presence in North America north of Mexico and was determined using Kartesz

and Meacham (2001). Origins of non natives were determined using a variety of web sources, including Flora

North America Online (www.efloras.org), Purdue University Dept of Horticulture (www.hort.purdue.edu/

newcrop), University of California Jepson Herbarium (www.ucjeps.berkeley.edu/interchange.html), Global

Compendium of Weeds (www.hear.org/gcw), Plants for a Future (www.pfaf.org/database), and Floridata

(http://www.floridata.com)

The non native species were divided into four categories, ruderals (weedy herbs of waste places, disturbed

areas, and dry,sandy soils), garden weeds, ornamentals (plants that appeared to have arrived at the site by

dumping or planting), and invasives (species naturalized in native forests, wetlands or prairies). Native spe-

cies were classed as ruderals, forest and woodland species, species of natural open habitats (savannas and

prairies), or wetland species. Assignment to categories was based on the primary habitat as given by Correll

and Johnston (1970) and modified by our experience. Classification of weedy species followed Kartesz and

Meacham (2001).

Three major sources were used to check for possible county or state records: Turner et al. (2003) Diggs

et al. (2006), and the collection at SBSC.

RESULTS

The vascular flora of the BGH site consists of 448 taxa in 259 genera in 97 families. The list may be found in

the Appendix which contains information on life form, nativity, distribution, introduction, original source,

habitat, and weediness.

All of the families are Angiosperms except five: two Gymnosperms (Pinaceae, Cycadaceae) and three

Pteridophytes (Lygodiaceae, Polypodiaceae, Thelypteridaceae). Nearly half the families are represented by

a single species, with only 18 containing more than five species. The top four familes contain more than

one-third of the taxa. They are Asteraceae (59 taxa; 13.5%), Poaceae (57; 13.1 %), Cyperaceae (35; 7.6%),

and Fabaceae (22; 5.0%). The largest genera are Carex (15 species), Juncus (9), Cyperus (9), Euphorbia (8),

Quercus (7), and Paspalum (6).

More than half (54%) of the species are broad-leafed herbs (Monocots and Dicots). Graminoids are the

696 Journal of the Botanical R h Institute of Texas 1(1)

second most important element (2396), with the remainder being split relatively equally between trees (996),

shrubs (696), and vines (896).

The species are mostly of broad distribution with more than half being from the southern US (Table 1).

Only 696 are found primarily in Texas; one, (Rhynchospora indianolensis) is endemic to the state and another

(Zephyranthes traubii) is known only from coastal Texas and NE Mexico (FNA 1993+ vol 19:302; Kartesz

and Meacham 1999; Diggs et al. 2006). None of the species is known to be threatened or endangered (TOES

1995), though R. indianolensis and Z. traubii are poorly collected (Kartesz and Meacham 2001). Species of

predominantly drier regions (central and southwest US) make up a very small fraction of the flora (ca 696).

About one-third of the species are from highly disturbed habitats, being either ruderals, garden weeds or

ornamentals (Table 2). Open-habitat species (prairie, field and savanna) are slightly more common than

closed-habitat species (forest or woodland). Approximately 5896 of the taxa are classified as weeds according

to Kartesz and Meacham (2001). None are on the Federal list of noxious weeds (USDA 2006). Two species

(Alternanthera philoxeroides and Triadica sebifera) are listed as noxious in Texas (Texas Administrative Code

We found one new state record, Syngonium podophyllum (per Kartesz & Meacham 2001; Jones & Wipff

2003). This is a horticultural species introduced from Mexico which we have observed growing wild in several

areas but apparently has not yet been formally documented as an escape from cultivation. Five new county

records were found. Cestrum nocturnum is listed in Jones & Wipff (2003) as cultivated only. The specimen

reported here is the first documentation of an escaped Texas population of this species. Eleocharis acicularis,

Rhynchospora indianolensis, Triodanis lamprosperma, and Zephyranthes pulchella are indicated as absent from

Harris County by Turner et al. (2003). Another species, Juncus elliottii, is known for Harris County only

from a specimen at New York Botanical Garden (Diggs et al. 2006) and from the specimen collected in this

study.

Species noteworthy for their showiness include one orchid (Spiranthes vernalis), spider lily (Hymenocallis

liriosme), a Crinum cultivar, and two species of rain lily (Zephyranthes pulchella, Z. traubii).

Most of the species (8196) are native to the United States. Of non native species, about 7696 are of Old

World origin, 2496 are from Mexico or South America, and 496 are of undetermined origin. The major Old

World locations are Asia (2996), Europe (1496), and Eurasia (1096). Only 3 of the species are African in origin

(Cynodon dactylon, Crinum bulbispermum and Dioscorea bulbifera).

DISCUSSION

The taxon count of 448 for the BGH site is somewhat lower than counts for most other sites in the region

(401—930 taxa; Table 3). However, considering the relative uniformity of the site, as well as the absence of

native prairie or savanna with their characteristically rich ground layer, the BGH site is surprisingly rich.

This very small site («50 ha) contains nearly 10% of all the species found in Texas. Species richness surpasses

that of the 125 ha Hickory Creek Unit of Big Thicket National Preserve, which is approximately 200 km to

the east (MacRoberts et al. 2002).

One contributor to the relative species richness of the site is non natives. They comprise 1996 of the flora

(87 spp), a substantially larger fraction than the 4—1396 for natural areas in the region (Table 3). The largest

number of non native species are ruderals (43 spp.; e.g., Duchesnea indica, Stellaria media, Euphorbia nutans,

Lamium amplexicaule) or garden weeds (Dioscorea bulbifera, Fatoua villosa, Ipomoea quamoclit, Oxalis debilis var.

corymbosa, Phyllanthus urinaria) that are adapted to highly disturbed sites and are commonly found at BGH

in mowed areas of fields or along trails, pipelines or ditches. A second group of non natives (n=15), mostly

ornamentals, appears to owe its presence at the site to more direct human agency, probably dumping from

nearby homes (Canna xgeneralis, Cestrum nocturnum, Citrus aurantium, Crinum bulbispermum, Cycas revoluta,

Iris sp., Kalanchoe sp., Lycoris radiata, Mirabilis jalapa, Musa sp., Philodendron hederaceum var. oxycardium,

Tradescantia pallida, Yucca sp.). Cyrtomium falcatum (holly fern) appears to have arrived via floodwaters from

upstream and Pyracantha koidzumi apparently germinated from bird-transported seed. The final group of

non native species (n222) appear to be naturalized components of native forests and prairies. This group

Harcombe et al., Flora of a

697

TABLE 1. Geographic distributions within USA of taxa at the BGH site (generalized from Kartesz & Meacham 2001).

Number Fraction

Central 16 0.04

East 80 0.18

South 82 0.18

Southeast Iss 0.35

Southwest 8 0.02

TX 29 0.06

USA 23 0.16

Ornamental 5 0.01

Grand Total 448 1.00

Taste 2. Numbers of taxa for which each listed habitat was considered to be the primary habitat (after Correll & Johnston

197

Number Fraction

Forest 119 0.27

Prairie, field, savanna 140 0.31

Wetland 50 0.11

Ruderal 114 0.25

Ornamental 15 0.03

Garden weed 6 0.01

Unknown 4 0.01

Total 448 1.00

TABLE 3. Species richness, total area and fraction non native for selected natural areas.

Site Species Area Fraction Source

(ha) Non native

Botanic Garden of Houston, Harris Co., TX 446 50 0.19 This study

Hickory Creek Unit, BTNP, Tyler Co., TX 401 284 0.04 MacRoberts et al. 2002

Turkey Creek Unit, BTNP, Polk and Tyler Co., TX 738 3150 0.06 Brown et al. 2004

Big Branch NWR, St Tammany Parish, LA 553 5870 0.13 Rosen et al. 2003

Big Lake Bottom WMA, Anderson Co., TX 459 1685 0.05 Fleming et al. 2002

Gus Engeling WMA, Anderson Co., TX 930 4465 0.06 Singhurst et al. 2002

Middlesex Fells,Boston, MA 400 69 0.26 Drayton & Primack 1998

Pelham Bay Pk, New York City, NY 956 0.34 DeCandido & Lamont 2004

Highland Hts Community Pk, Cuyahoga County, OH 403 141 0.35 Jog et al. 2005

Shaker Median Park, Beachwood, Cuyahoga Co. OH 298 140 0.47 Delong et al. 2005

includes both herbaceous and woody plants (including shrubs and vines), some of which appear to be highly

invasive locally. The most aggressive invaders at the BGH site are Paederia, Lygodium, Ligustrum spp, and

Triadica. Paederia is very dense in parts of the BGH site, and it has been reported to be extremely aggressive

at another nearby site (Brown 1992).

Ecologically, the non native flora differs from the native flora of the site in several ways. A slightly larger

percentage of the non-natives is widely distributed in the USA (2696) than natives (1496) and more of the

non native species are weedy and/or ruderal (6896) than natives (5696). Habitat designations are strongly

skewed to disturbed sites and waste areas for non natives (5696) as opposed to natives (2096). It is also worth

noting that the contribution of non native species from Latin America is substantially higher at BGH (21%)

698 Journal of the Botanical R h Institute of Texas 1(1)

than has been reported farther north (Central New York; 396), as would be expected, considering location

and climate.

Although the non natives are important at the BGH site, the non native percentage (1996) is somewhat

lower than reported in other urban preserves (26-47%). Perhaps the percentage is lower because most of

the site is either closed forest or frequently-mowed parkland. Areas of early successional vegetation are quite

small. Also, urbanization has been more recent around the BGH site than around the other urban preserves

listed in Table 3, so non native species may still be accumulating at the site. If insufficient time has passed

for the full complement of non natives to have colonized the site, then it may explain lack of a southward

increase in the fraction of non natives such as the trend identified by Pysek (1998) for central European cit-

ies. Clearly, more published data on the floras of urban preserves are required to identify important trends

and their causes.

Generally speaking, a large fraction of the non native flora of the USA arrived via purposeful introduction

for ornamentals, gardens, crops, forage or soil stabilization (Mack and Lonsdale 2001; Reichard and White

2001). For example, 4996 of the non native flora of the Cayuga region of NY was planted at some time in that

region or elsewhere in the USA (Marks et al. 2006). This appears true of the BGH site, as well. A substantial

fraction (4196) of the non native species appear to have been introduced for some specific human use. The

most notorious Texas example of escape from cultivation is Triadica sebifera (Chinese tallow tree), originally

introduced as an ornamental by Benjamin Franklin, and subsequently as a potential oil-seed crop by the

US Bureau of Plant Industry (Bruce et al. 1997).

Aside from non native plants, another important cause of high richness at the BGH site is the large

percentage of native weeds (4596 of the site flora). While weediness generally has a negative connotation, it

must be emphasized that native weeds are a heterogeneous group, including not only native ruderals that

are mostly restricted to disturbed and waste areas such as Erigeron philadelphicus, Helianthus annuus, and

Euphorbia nutans (1296 of the flora), but also many forest, prairie or marsh species that, because of broad

ecological tolerances, high fecundity, and wide dispersal, are good colonizers of disturbed areas (3296 of

the flora), and

canopy dominant trees at the site, including Ulmus, Celtis, Pinus, and Fraxinus.

juently are present in many forest types. This latter group includes several important

The importance of weediness is illustrated by the fact that, combining natives and non natives, 5896 of

flora is classed as weeds according to Kartesz and Meacham (2001). The weedy nature of the flora highlights,

in turn, the long history of human use for grazing and logging and its role in determining species richness.

Following cessation of burning and grazing nearly 50 years ago, there was ample opportunity for vagile

members of native communities to colonize the site.

The final class of species that constitute the flora is non-weedy species that are commonly associated

with less-disturbed habitats, and by definition are less fecund, vagile, or tolerant of a range of conditions

than those species classified as weedy. They amount to 3696 of the flora, and include species of both forested

and non-forested habitats. The most notable forest species in this class is Quercus similis, one of the original

site dominants which remains important. Many species of prairies and marshes (4396 of the flora) are still

present, but the prairie imprint has undoubtedly been substantially diluted by succession of prairie to forest

on the western part of the site.

It is interesting to note that few species in the flora of the site have predominantly Western or Mid-

western distributions. We found only one woody species that is more typical of drier regions to the west

(Maclura pomifera) than to the humid region to the east. Other species such as Symphoricarpos orbiculatus,

Sapindus drummondii, and Acacia farnesiana, which are present in the Houston area, were absent at this site.

The prominence of eastern and southeastern taxa was previously identified for another protected area ap-

proximately 200 km North (MacRoberts and MacRoberts 2004). Together the results support the finding

of MacRoberts and MacRoberts (2003) that the broad transition from eastern to western floras begins just

west of Houston. Consequently, in spite of marked differences in appearance of the plant communities and

the landscape across the transition from forest to prairies and woodlands that occurs in Houston, the flora

Harcombe et al., Flora of a ite, Harris County, Texas 699

remains predominantly Eastern. The richness of the flora of the site does not appear to be a result of mixing

of species from different habitats or different regions.

With respect to disturbance, our results show that the list of habitat generalists is large and site het-

erogeneity is low, and so the effect of disturbance is to offer opportunity for weedy species to colonize and

persist. Thus, richness is high because of the abundance of native and non native weedy species that tend

to be habitat generalists of broad distribution.

These observations on the BGH site clearly illustrate the importance of weeds and non natives in modern

urban floras. Because of this and the low abundance of habitat specialists, there seems little risk of harm

through escape of additional non natives. Indeed, the opportunity exists to enhance diversity by control-

ling woody invasives and by reintroducing elements of the prairie vegetation that once occupied part of the

site.

APPENDIX

Ecological characteristics of taxa at Herman Brown Park, Houston, Harris County, Texas. Column heads are as follows: F=life

form; O=origin; D=Distribution in US; I=reason for introduction; H=habitat; W=weed status; C=specimen collection numbers

(I.S.Elsik & W.W. Pruess unless otherwise noted). See below for key to column entries.

FAMILY F (0) D l H W C

Acanthaceae

Dicliptera brachiata (Pursh) Spreng. H N SE F N 4602

Hygrophila lacustris (Schlecht. & Cham.) Nees H N SE E W 4432

Justicia ovata (Walter) Lindau var. lanceolata H N SE F N 4161

(Chapm.) RW, Long

Ruellia caroliniensis (J.F. Gmel.) Steud. H N E P W 4282, 4434

Ruellia nudiflora (Engelm. & A. Gray) Urb. H N TX P N 4445

Aceraceae

Acer negundo L. i, N US f W 4187

Agavaceae

Yucca aloifolia L. S N SE O Or N 4490, 4660

Yucca sp. L. 5 Or 4665

Alismataceae

Sagittaria platyphylla (Engelm.) J.G. Sm. H N SE W W 4429, 4304

Amaranthaceae

Alternanthera philoxeroides (Mart.) Griseb. H SA S U R W 4165

Anacardiaceae

Toxicodendron radicans (L.) Kuntze V N E E W 4146, 4561

Apiaceae

Bowlesia incana Ruiz & Pav. H N SW R W 4094

Centella erecta (L. £) Fernald H N SE R W 4276

Chaerophyllum tainturieri Hook. H N TX P W 4426

var. dasycarpum Hook. ex S. Wats

Chaerophyllum tainturieri Hook. var. tainturieri H N 3 P W 4064

Cyclospermum leptophyllum (Pers.) Sprague H SA S A W 4266, 4202

ex Britton & Wilson

Cynosciadium digitatum DC. H N SE P N 4308

Eryngium hookeri Walp. H N TX P N 4450

Hydrocotyle verticillata Thunb. H N S W N 4265

var. triradiata (A. Rich.) Fernald

Hydrocotyle verticillata Thunb. var. verticillata H N S W N 4170

Limnosciadium pumilum (Engelm. & A. Gray) H N vis O F W 4193

Mathias & Constance

Ptilimnium capillaceum (Michx.) Raf. H N E W W 4310

Sanicula canadensis L. H N E F N 4159

Torilis arvensis (Huds.) Link H EA S A R W 4209

Trepocarpus aethusae Nutt. ex DC. H N SE P W 4377

700

fal, Dat

Journal of titute of Texas 1(1)

FAMILY F (0) D l H W C

Apocynaceae

Trachelospermum difforme (Walter) A. Gray V N E F W 4531

Aquifoliaceae

llex decidua Walter S N SE [5 W 4103

llex opaca Aiton ii N SE F N 4685

llex vomitoria Aiton S N SE F W 4132

Araceae

Colocasia esculenta (L.) Schott H P SE O F W 4466

Philodendron hederaceum (Jacq.) Schott V SA @ O Or N 4644

var. oxycardium (Schott)

Syngonium podophyllum Schott V SA SE A F N 4538

Xanthosoma sagittifolium (L.) Schott H SA TX O E N 4444

Araliaceae

Hedera helix L. V EA SE O F W 4333

Arecaceae

Sabal minor (Jacq.) Pers. S N SE F W 4301

Asclepiadaceae

Asclepias viridis Walter H N SE R W 4376

Matelea gonocarpos (Walter) Shinners V N SẸ E N 4381, 4307

Asteraceae

Acmella oppositifolia (Lam.) Jansen H N SE W N 4535

var. repens (Walter) Jansen

Ageratina altissima (L) King & H. Rob. H N E P W 4648

Ambrosia usi DE H N US P W 4585

Ambrosia trifida À H N US R W 4428

Arnoglossum ovatum (Walter) H. Rob. H N SE P N 4449

Baccharis halimifolia L. S N E È W 4591, 4590

Bidens bipinnata L H N E P W 4575

Calyptocarpus vialis Less. H N 5 W W 4070

Cirsium horridulum Michx. H N S P W 4183

Cirsium texanum Buckle H N C R N 4321

Conoclinium coelestinum (L.) DC. H N E F W 4052

Conyza canadensis (L.) Cronquist var. canadensis H N US R W 4424, 4417

Coreopsis tinctoria Nutt. H N US P W 4395

Eclipta prostrata (L.) L. H N D W W 4481, 4245

Elephantopus carolinianus Raeusch. H N SE F N 4031

Erigeron philadelphicu H N US R W 4068

Erigeron tenuis Torr. E H N S R N 4081, 4812

Eupatorium capillifolium (Lam.) Small H N SB R W 4635

Eupatorium semiserratum DC. H N SE P N 4472

Eupatorium serotinum Michx. H N E R W 4027

Eupatorium X pinnatifidum Elliott H N SE R W 4634

Euthamia leptocephala (Torr. & A. Gray) H N SE P N 4586

Greene ex Porter & Britton

Gaillardia aestivalis (Walter) H. Rock var. aestivalis H N SE P N 4494

Gamochaeta coarctata (Willd.) Kerguelen H N D P N 4230, 4200

Gamochaeta purpurea (L.) Cabrera H N E P W 4092, 4198

Helenium flexuosum Ra H N E R W 4271

Helianthus annuus L. H N US R W 4309

Helianthus simulans E.E. Wat H N TX P N 4589

Heterotheca subaxillaris ae ) Britton & Rusby H N S P W 4447, 4129

Hypochaeris microcephala (Sch. Bip.) Cabrera H SA TX A R N 4084, 4199

var. albiflora (Kuntze) Cabrera

lva angustifolia Nutt. ex DC. H N 5 P N 4636

Iva annua L H N C W W 4584

Krigia cespitosa (Raf) K.L. Chambers forma cespitosa H N S P W 4190, 4126

Harcombe et al., Flora of a ite, Harris County, Texas

701

FAMILY F (0) D l H W C

Krigia cespitosa (Raf) K.L. Chambers H N S P W 4172

forma gracilis (DC) KJ. Kim

Lactuca floridana (L.) Gaertn. H N E P W 4506, 4587

Mikania scandens (L.) Willd. V N E P W 4402

Packera tampicana (DC.) C. Jeffrey H N SE P W 4156

Pluchea camphorata (L.) DC. H N SE P W 4452, 4599

Pluchea foetida (L.) DC. H N SE P N 4608

Pluchea odorata (L.) Cass. H N S P W 4034

Pterocaulon virgatum (L.) DC. H N TX P N 4632

Pyrrhopappus pauciflorus DC. H N TX P N 4087

Rudbecki plexicaulis Vahl H N SE P N 4264

Rudbeckia hirta L. H N US R W 367

Rudbeckia texana (Perdue) P.B. Cox & Urbatsch H N SE P N 4407, 4505

Solidago canadensis L. H N US R W 4656, 4013

Solidago gigantea Aiton H N US P W 4597

Solidago stricta Aiton H N SE P N 4631

Soliva sessilis Ruiz & Pav. H SA SB A R W 4201

Sonchus asper (L.) Hill H E US A R W 4086

Symphyotrict divaricatum (Nutt.) G.L. Nesom H N E R W 4629

symphyotricl umosum (L.) G.L. Nesom H N SE P N 4315

Symphyotrichum lanceolatum (Willd.) H N C P N 4658

G.L. Nesom var. lanceolatum

symphyotrichum racemosum (Elliott) G.L. Nesom H N E P N 4058

Symphyotrichum subulatum (Michx.) G.L. Nesom H N E R W 4037

Taraxacum erythrospermum Andrz. ex Besser H C US A R W 4102

Ver besir IU virgit "ica A H N S P W 4543

Vernonia missurica Raf. H N 5 P N 015

Youngia japonica (L.) DC. H AS S A R W 4083, 4077

Betulaceae

Carpinus liniana Walter ale N E F N 4389

Bignoniaceae

Campsis radicans (L.) Seem. V N B F W 4366

Catalpa bignonioides Walter i N E E N 4387

Boraginaceae

Heliotropium procumbens Mill. H N SE R W 4461

Myosotis macrosperma Engelm. H N E F N 4082

Brassicaceae

Brassica juncea (L.) Czern. H AS US A R W 4150

ardamine debilis D. Don H P S A R N 4096, 4409

Lepidium virginicum L. H N E R W

Bromeliaceae

Tillandsia usneoides (L.) L. H N SE F N 4262, 4113

Cactaceae

untia sp. (L.) Mill. S TX P 4489

Callitrichaceae

Callitriche heterophylla Pursh H N US W N 4176, 4242

Campanulaceae

Lobelia appendiculata A. DC. var. appendiculata H N E F W 4673

Lobelia puberula Michx. H N E P N 4568

Triodanis lamprosperma McVaugh H N S. P N 4224

Triodanis perfoliata (L.) Nieuwl. var. biflora H N US R W 4167

(Ruiz & Pav.) T.R. Bradley

Cannaceae

Canna x generalis L.H. Bailey H E O Or N 4596

Caprifoliaceae

Lonicera japonica Thunb. V AS E O F W 4144

702

Journal o

fal, Dat

| titute of Texas 1(1)

FAMILY F O D I H WwW C

Sambucus nigra L. subsp. canadensis (L.) R. Bolli B N US n N 4356

Viburnum dentatum L. S N E F N 4340

Caryophyllaceae

Cerastium glomeratum Thuill. H EA US A R W 4088

Sagina decumbens (Ell.) Torr. & A. Gray H N SB R W 4243

Stellaria media (L.) Vill. H C US A R W 4069

Clusiaceae

Hypericum hypericoides Crantz 5 N E P W 4471,4035

Commelinaceae

mmelina caroliniana Walter H N SE E N 4628

Commelina diffusa Burm.f. var. diffusa H N S F N 4600

Commelina erecta L. H N E R W 4662

Gibasis pellucida (M. Martens & Galeotti) D.R. Hunt H SA SE O N 4443

Tradescantia pallida (Rose) D.R. Hunt H SA SE O Or N 4642

Convolvulaceae

Dichondra carolinensis Michx. H N 5 R W 4136

Ipomoea cordatotriloba Dennst. var. cordatotriloba V N SE R W 4504

Ipomoea quamoclit L. V SA S O G W

Cornaceae

Cornus drummondii C.A. Mey. > N E P N 4155

Cornus foemina Mill. S N S W N 4311

Nyssa sylvatica Marsh. i N E F N 4360

Crassulaceae

Kalanchoe cv. Adans. H O Or N 4643

Cycadaceae

Cycas revoluta Thunb. 5 As SE O Or N 4580

Cyperaceae

Carex caroliniana Schwein. G N S F N 4229

Carex cherokeensis Schwein. G N SE F W 4101

Carex complanata Torr. & Hook. G N SE F N 4305

Carex corrugata Fernald G N SE F N 4285

Carex flaccosperma Dewey G N SE F N 4147, 4272

Carex frankii Kunth G N S P W 4415, 4292

Carex intumescens Rudge G N E P N 4100, 4353

Carex leavenworthii Dewey G N E P. N 4223

Carex louisianica L.H.Bailey G N SE P W 4249

Carex lupulina Muhl. ex Willd. G N E P W 4687

Carex oxylepis Torr. & Hook. G N SE k W 4231

arex reniformis Small G N SE P N 4284

Carex retroflexa Muhl. ex Willd. G N E F W 4226

Carex triangularis Boeckeler G N SE R W 4268, 4287

Carex tribuloides Wahlenb. G N E W N

Cyperus croceus Vahl G N SE P W 4363, 4550

Cyperus entrerianus Boeckeler G SA SE A F W 4051

Cyperus esculentus L. G N US R W 4624

Cyperus ochraceus Vahl G N SE W N 4612, 4059

Cyperus odoratus L. G N US P W 4577

Cyperus pseudovegetus Steud. G N SE P W 4253

Cyperus retrorsus Chapm. G N SE P W 4574

Cyperus thyrsiflorus Schltdl. G N SE F N 4355, 4548

Cyperus virens Boeckeler G N SE P W 4234

Eleocharis acicularis (L.) Roem. & Schult. G N US W N 4263, 4293

Eleocharis montana Roem. & Schult. G N SW W N 4252

Eleocharis montevidensis Kunth G N SW P N 4236, 4253

Fimbristylis dichotoma Vahl G N SE P W 4578

Harcombe et al., Flora of a

703

FAMILY F (0) D l H W C

Fimbristylis miliacea Vahl G N SE E N 4462

Kyllinga brevifolia Rottb. G N 5 R W 4238, 4482

Kyllinga odorata Vahl G N SE P N 4579

Rhynchospora caduca Ell. G N SE P N 4278

Rhynchospora corniculata (Lam.) A. Gray G N SE P W 4288

Rhynchospora indianolensis Small G N TX P N 4283, 4519

Scleria oligantha Michx. G N S p N 365

Dioscoreaceae

Dioscorea bulbifera L. V Af S O G W 4351

Ebenaceae

Diospyros virginiana L. i N C E W 4259, 4194

Epod ae

alypha gracilens A. Gray H N E P W 4459

Acalypha rhomboidea Raf. H N E P W 4416

Croton capitatus Michx. var. lindheimeri H N SE R W 4394, 4457

(Engelm. & A. Gray) Muell.-Arg.

Croton monanthogynus Michx. H N E E W 4397

Euphorbia bicolor Engelm. & A. Gray H N S P N 4475

Euphorbia dentata Michx. H N E P W 4148

Euphorbia heterophylla L. H N S P W 4540

Euphorbia maculata L. H N US P W 463

Euphorbia nutans Lag. H N E R W 4458, 4507

Euphorbia prostrata Aiton H N S R W 4241

Euphorbia serpens Kunth H N US R W 4559

Euphorbia spathulata Lam. H N Ws R W 4130

Phyllanthus pudens L.C. Wheeler H N S R N 4562

Phyllanthus urinaria L. H AS SE A G W 4547

Ricinus communis L. H M US O F N 556

Triadica sebifera (L.) Small if Ch SE G F W 4258, 4588

Fabaceae

Albizia julibrissin Durazz. T Ch 5 O P W 4373

Cercis canadensis L. var. canadensis Ji N E F N 4419

Chamaecrista fasciculata (Michx.) Greene H N E R W 4024, 4440

Desmanthus virgatus(L.) Willd. H N TX P W

E glabellum DC. H N SE F N 4645, 4570

odium paniculatum (L.) DC. H N E F N 4510, 4647

mu herbacea L. 5 N SE F N 4649

Gleditsia triacanthos L JE N US F N 4653, 4542

Glottidium vesicarium R.M. Harper H N SE R W 4512

Kummerowia striata (Thunb. ex Murray) Schindl. H AS > F R W 4565

Lathyrus pusillus Ell. V N > P W 4151

Medicago polymorpha L. H EA US A R W 4079

Mimosa strigillosa Torr. & A. Gray H N SE E W 4341

Neptunia pubescens Benth. H N SE P N 4396

Rhynchosia minima (L) DC. V N SE P N 4573

Sesbania drummondii (Rydb.) Cory > N SE P W 4385

Sesbania herbacea (Mill) McVaugh H N 5 P W 4621

Strophostyles helvola (L.) Elliott V N E P W 4508, 4509

Trifolium incarnatum L. H E [E E R W 4207

Trifolium repens L. H AS US E R W 116

unknown V 4652, 4557, 4661

Vicia minutiflora D. Dietr. H N SE E N 4143

Vicia tetrasperma (L.) Schreb. H E E F R W 4080

Fagaceae

Quercus alba L. i N E F N 4553

704

Journal o

fal, Dat

f titute of Texas 1(1)

FAMILY F O D l H W C

Quercus falcata Michx. T N E F N 4532

Quercus laurifolia Michx. Jj N SE F N 4674

Quercus nigra L. F N SE F N 4479

Quercus phellos L. 2 N SE F N 4133, 4464

Quercus similis Ashe F N SE ls N 4609, 4603

Quercus virginiana Mill. el N BE F N 4500

Gentianaceae

Centaurium pulchellum (Sw.) Druce H E E A R N 4260

Sabatia campestris Nutt. H N TX P N 4298

Geraniaceae

Geranium carolinianum L. H N US R W 4063

Haloragaceae

Myriophyllum pinnatum (Walter) H N S W W 4289

Britton, Sterns & Poggenb.

Proserpinaca palustris L. var. amblyogona Fernald H N E W W 4303

Hamamelidaceae

Liquidambar styraciflua L. Ji N E P N 4124

Hydrophyllaceae

Hydrolea ovata Nutt. H N SE W W 4412

Hydrolea uniflora Raf. H N SE W W 4403

Iridaceae

Herbertia lahue (Molina) Goldblatt H N TX F N 4164

HIS Spec H 4484

Sisyrinchium langloisii Greene H N TX R N 4131

Sisyrinchium minus Engelm. & A. Gray H N TX P N 4180

Sisyrinchium rosulatum E.P. Bicknell H N SE R N 4163, 4141

Juglandaceae

Carya illinoensis K. Koch d N E ls N 4386

Juncaceae

Juncus acuminatus Michx. G N US W W 4233

Juncus bufonius L. G N Us W W 4244

Juncus diffusissimus Buckley G N SE P N 4290

Juncus effusus L. G N E P W 4185

Juncus elliottii Chapm. G N Sie P N 4257

Juncus marginatus Rostk. G N SE P N 4392, 4227

Juncus repens Michx. G N SẸ P N 4313

Juncus tenuis Willd. G N US F W 4306, 4221

Juncus validus Coville G N SE P W 4404

Lamiaceae

Hedeoma hispida Pursh H N US P W 4273

Lamium amplexicaule L. H EA US A R W 4122

Lycopus rubellus Moench. H N SE F N 4607

Lycopus virginicus L. H N E F N 4025

Micromeria brownei Benth. H N SE [5 N 4317

Monarda citriodora Cerv. ex Lag. H N US P W 4339

Prunella vulgaris E H N E R W 419]

Salvia coccinea Juss. Ex Murr. H N SE P N 4676

Salvia lyrata L. H N E F W 4076

Scutellaria drummondii Benth. H N SW R N 4378

Scutellaria parvula Michx. H N E R N 4177,4127

tachys crenata Phil. H N TX F N 4089

Teucrium canadense L. H N US W W 4026, 4354

Lemnaceae

Lemna aeq tialis Welw. H N S W W 4483, 4473

Spirodela punctata (G. Mey.) C.H. Thomps. H N SE W W 4299

Harcombe et al., Flora of a ite, Harris County, Texas

705

FAMILY F (0) D H W C

Lentibulariaceae

Utricularia gibba L. H N E W W 4314

Liliaceae

Allium canadense L. var. canadense H N E P N 4075

Allium canadense L. var. mobilense (Regel) Ownbey H N SE E N 4186

Crinum bulbispermum (Burm. H Af SE Or N 4686, 4641

Milne-Redh. & Schweick.

Hymenocallis liriosme (Raf) Shinners H N SE P N 4312

Lycoris radiata Herb. H Ch SE Or N 4536

Nothoscordum bivalve (L.) Britton H N S R W 4074

Zephyranthes pulchella J.G. Sm. H N 5 P N 4534, LB30762

Zephyranthes traubii (Hayward) Moldenke H N TX P N

Linaceae

Linum medium (Planch.) Britton H N US W N 4295, 4493

var. texanum (Planch.) Fernald

Loganiaceae

Gelsemium sempervirens (L.) Aiton V N SE P W 4119

Mitreola petiolata (Gmel.) Torr. & A. Gray H N SE P N 4522

Polypremum procumbens L. H N 5 R W 4640

Lygodiaceae

Lygodium japonicum (Thunb.) Sw. V AS SE F W 4323

Lythraceae

Ammannia coccinea Rottb. H N S W W 4142

Lagerstroemia indica L. S AS SE P W 4425

Lythrum alatum Pursh var. lanceolatum H N SE W W 4393

(EIL) Torr. & A. Gray ex Rothrock

Magnoliaceae

Magnolia grandiflora L. y N SE E N 4324

Malvaceae

Malvaviscus arboreus Cav. var. drummondii E N SE P N 4350

(Torr. & A. Gray) Schery

Modiola caroliniana (L.) G. Don H N S R W 4152

Sida rhombifolia L. H N S P W 4071, 4296

Sida spinosa L. H N E R W 572

Meliaceae

Melia azedarach L. it As S E W 4418

Menispermaceae

Cocculus carolinus DC. V N SE [: W 4357

Moraceae

Fatoua villosa Nakai H AS S G W 4485

Maclura pomifera (Raf.) C.K. Schneid. T N US P W 4604

Morus alba L E Ch US F W 4567

Morus rubra L. Jj N g F W 4106

Musaceae

Musa sp. L. H Or N 4488

Myricaceae

Myrica cerifera L. 5 N SẸ P W 4406

Nyctaginaceae

Mirabilis jalapa L. H SA SW Or W 4486

Oleaceae

Chionanthus virginicus E T N SE F N 4437, 4362

Forestiera ligustrina Poir. 3 N SE F N 4539, 4325

Fraxinus americana L. JE N E E W 4216

Fraxinus berlandieriana DC. iy N SW F W 4560, 4382

Fraxinus pennsylvanica Marshall ie N C F W 4173, 4411

706 Journal of the Botanical R hl te of Texas 1(1)

FAMILY F (0) D l H W C

Ligustrum lucidum Aiton S AS SE O W W 4623

Ligustrum sinense Lour. S As SE O n W 4192

Onagraceae

Gaura lindheimeri Engelm. & A. Gray H N TX P N 4503, 4492

aura parvi flora Dougl. ex Lehm. H N US R W 4427

ee decurrens Cus) H N E W W 4521

Ludwigia glandulosa Walter H N SE W N 4281

Ludwigia octovalvis Tum PH. Raven H N SE W W 4018

Ludwigia palustris (L.) Elliott H N US W W 4212, 4430

Oenothera speciosa Nutt. H N S P W 4162

Orchidaceae

Spiranthes vernalis Engelm. & A. Gray H N E R N 4294

Oxalidaceae

Oxalis corniculata L. var. wrightii (A. Gray) B.L. Turner H N C R N 4014

Oxalis debilis Kunth var. corymbosa (DC.) Lourteig H SA SE O G W 4062

Oxalis violacea L. H N BS P W 4555

Passifloraceae

Passiflora incarnata L. V N SE E W 4516

Passiflora lutea L. V N SE F N 4474

Phytolaccaceae

Phytolacca americana L. H N E R W 4442

Pinaceae

Pinus taeda L. i N SE [s W 4491

Plantaginaceae

Plantago rhodosperma Dcne. H N S P N 4091, 4134

Plantago virginica L. H N Us R W 4160

Platanaceae

Platanus occidentalis L. AN N E F W 4361

Poaceae

(Walter) Britton, Sterns & Poggenb. G N E W 4181, 4197

Andropogon glomeratus (Walter) Britton, Sterns G N S P W 4593, 4657

& Poggenb. var. pumilus (Vasey) L.H. Dewey

Andropogon virginicus L. G N S R W 4619

Aristida purpurascens Poir. var. virgata (Trin.) Allred G N Bs P W 4620

Avena sativa L. G E US E R W 4208

Bothriochloa ischaemum (L.) Keng var. songarica G As SW F R W 4639

(Rupr. ex Fisch. & C.A. Mey.) Celarier & Harlan

Bothriochloa longipaniculata (Gould) Allred & Gould G N TX P N 4371

Briza minor L. G E S A F W 4178

Bromus catharticus Vahl G SA US F R W 4120

Chasmanthium latifolium (Michx.) H.O. Yates G N S F N 4528

Chasmanthium laxum (L.) H.O. Yates var. laxum G N SE F N 4438

Chasmanthium laxum (L.) H.O. Yates G N SE F N 4467

var. sessiliflorum (Poir.) Wipff & S.D. Jones

Chloris canterai Arech. G SA x U R N 4368

See dactylon (L.) Pers. G Af US F R W 4267

thelium acuminatum (Sw.) Gould & G N E N LB30549

"e Clark var. densiflorum (E.L. Rand & Redfield)

Gould & C.A. Clark

C.A. Clark var. lindheimeri (Nash) Gould & C.A. Clark

Dichanthelium commutatum (Schult.) G N 5 P N 4109

Gould subsp. commutatum

Dichanthelium acuminatum (Sw.) Gould & G N E R N 4157, 4274

Dichanthelium dichotomum (L.) Gould subsp. G N E P N 4246, 4614

microcarpon (Muhl. ex Ell.) Freckmann & Lelong

Harcombe et al., Flora of a ite, Harris County, Texas 707

FAMILY F O D l H W C

Dichanthelium dichotomum (L.) Gould G N E P N LB30556

subsp roanokKense (Ashe) Freckmann & Lelong

Dichanthelium laxiflorum (Lam.) Gould G N SE P N 4256, 4364

Dichanthium aristatum (Poir.) C.E. Hubb. G | S k R N 4622

Digitaria ciliaris (Retz.) Koeler G N S R N 4514

Echinochloa colonum (L.) Link G | S A R W 4423

Echinochloa muricata (P.Beauv.) Fernald G N US R W 4204, 4627

Echinochloa walteri (Pursh) in A. Heller G N E P W 4530

Eleusine indica (L.) Gaertn. G EA US A R W 4626

Elymus virginicus L. G N G E W 4149

Eragrostis intermedia Hitchc. G N S P N 4513

Hordeum pusillum Nutt. G N US R W 4093

Leersia virginica Willd. G N E F N 4280

Lolium perenne L. G EA US G R W 4179

Melica mutica Walter G N SE F N 4108

Oplismenus hirtellus (L.) Beauv. subsp. G N SE F N 4551

setarius (Lam.) Mez

Panicum anceps Michx. G N SE F W 4615

Panicum dichotomiflorum Michx. G N US R W 457]

Panicum gymnocarpon Elliott G N SE W N 4520

Panicum rigidulum Bosc ex Nees G N S W W 4480, 4446

Paspalum dilatatum Poir. G SA S F R W 4145

Paspalum langei (E. Fourn.) Nash G N SE F N 4254, 4222

Paspalum lividum Trin. Ex Schltdl. G N TX W N

Paspalum notatum Flueggé G SA 3 F R W 4239

Paspalum pubiflorum Rupr. G N S R N 4237

Paspalum urvillei Steud. G SA SE E R W 4320, 4659

Phalaris lini Walter G N E R W 4300

Poa annua L G E US A R W 4073

Polypogon monspeliensis (L.) Desf. G E US A R W 4211

Saccharum giganteum (Walter) Pers. G N SE W W 4633

Schizachyrium scoparium (Michx.) Nash G N SE P N 4616

var. divergens (Hack) Goul

Setaria parviflora (Poir.) Kerguélen G N S R W 4331, 4433

Sorghum halepense (L.) Pers. G M US F R W 4196

Sphenopholis obtusata (Michx.) Scribn. G N US R W 4203

Sporobolus compositus Merr. var. drummondii G N TX R N 4569

(Trin.) J. Kartesz & Gandhi

Sporobolus indicus (L.) R. Br. G SA Sie A R N 4286

Steinchisma hians (Elliott) Nash G N SE W N 4189, 4401

Stenotaphrum secundatum (Walter) Kuntze G N SE G W 4335

Tridens strictus Nash G N SE F W 4617

Urochloa reptans (L.) Stapf G N TX R W 4455

Polygalaceae

Polygala verticillata L. H N C P N 4277

Polygonaceae

Brunnichia ovata (Walter) Shinners V N SE F W 4527, 4653, 4672

Polygonum densiflorum Meisn. H N Sb W N 4554

Polygonum hydropiperoides Michx. H N US W W 4175, 4020

Rumex crispus L H E US A R W 4319

Polypodiaceae

Cyrtomium falcatum (L. f.) C. Presl F As US O Or N 4663

Polygonum punctatum Elliott H N US W 4630

Pleopeltis polypodioides (L.) E.G. Andrews & F N JE $ N 4670

Windham subsp. michauxiana (Weatherby)

E.G. Andrews & Windham

708

fal, Dat

Journal of titute of Texas 1(1)

FAMILY F (0) D l H W C

Pontederiaceae

Heteranthera limosa Vahl H N E W W 4408

Portulacaceae

Portulaca oleracea L. H N US P W 4460

Primulaceae

Anagallis arvensis L. H E US A R W 4090

Anagallis minima (L.) K. Krause H N US P W 4302, 4228

Samolus valerandi L. subsp. parviflorus (Raf.) Hultén H N US W N 4316

Ranunculaceae

Anemone berlandieri Pritz. H N SE P W 4135

Clematis crispa L. H N S P N 4110, 4469

Clematis terniflora DC. V J E O R N 4439

Ranunculus muricatus L. H Ẹ 5 A R W 4078

Ranunculus pusillus Poir. H N 5 R W 4125

Ranunculus sardous Crantz H E SE A P W 4112

Rhamnaceae

Berchemia scandens (Hill) K. Koch V N SE F W 4085, 4174

Rosaceae

Crataegus marshallii Eggl. ER N SE F W 4104

Crataequs viridis L. da N SE F N 4105, 4275

Duchesnea indica (Andrews) Focke H AS E O R W 4114

Prunus caroliniana (P. Mill.) Ait. T N SE E N Jes

Pyracantha ea (Hayata) Rehd. S AS SE O Or N 4558

Rubus argutus Lin V N S R W 4061

Rubus trivialis Michx. V N Sie R W 4552

Rubiaceae

Diodia virgil var IU lt H N SE P W 4332

Galium aparine L. H N us p W 4065

Galium tinctorium L. H N E W N 4072, 4138

Houstonia pusilla Schoepf H N S R N 4107

Mitchella repensL. H N E F N HE

Paederia foetida L. V AS TX O F W 4405, 4345, 4655

herardia arvensis L. H M E A R W

Rutaceae

Citrus aurantium L. i AS SE € Or N 4526

Zanthoxylum clava-herculis L. Ji N SE E W 4217, 4400

Salicaceae

Populus deltoides subsp. deltoides Bartram ex Marshall T N US F W 4206

Salix nigra Marshall 2 N E F W 4115,4117

Sapotaceae

Sideroxylon eee Michx. i N S p N 4487, 4359, 4370

Scrophulariac

Agalinis ol (Nutt.) Small H N SẸ P N 4592, 4646

Bacopa monnieri (L.) Pennell H N S W W 4166

Gratiola virginiana L. H N S W N 4169

anagallidea (Michx.) Cooperr.

Mazus pumilus (Burm. f.) Steenis H EA S A P W 4261

Mecardonia procumbens Small H N SW W N 4171

Penstemon tenuis Small H N TX P N 4153

Veronica arvensis L. H EA Us A P W 4095, 4128

Micranthemum umbrosum (J.F. Gmel.) S.F. Blake H N SE W W 4210

Veronica peregrina L. subsp. peregrina H N US R N 4139

Veronica persica Poir. H AS US A R W 4137

Smilacaceae

Smilax bona-nox L. V N SE F W 4399, 4601

Harcombe et al., Flora of a ite, Harris County, Texas 709

FAMILY F (0) D l H W C

Smilax rotundifolia L. V N E F W 4523

Smilax smallii Morong V N SE E W 411]

Solanaceae

Calibrachoa parviflora Jussieu) D'Arcy H N S P W 4240

Cestrum nocturnum L. 5 SA TX O Or N 4689, 4598

Solanum capsicastrum Link ex Schauer $ U S O R N 4010, 4067

Solanum carolinense L. H N US R W 4184

Solanum ptychanthum Dun. H N US R W 4012

Sterculiaceae

Melochia pyramidata L. H N TX R N 4625

Thelypteridaceae

Thelypteris kunthii (Desv.) CV. Morton E N SE E N 4420, 4650

Typhaceae

Typha domingensis Pers. H N US W W 4338

Typha latifolia L. H N US W W 4337

Ulmaceae

Celtis laevigata Willd. T N US F W 4501, 4097

Ulmus alata Michx. ji N E F W

Ulmus americana L. iF N E F W 4098, 4517

Ulmus crassifolia Nutt. JR N S E N 4654, 4390

Urticaceae

Boehmeria cylindrica (L.) Sw. H N E E W 4476

Valerianaceae

Valerianella woodsiana Walp. H N SE F W 4066

Verbenaceae

Callicarpa americana L. S N SẸ P W 4478, 4336

Lantana camara L. $ N SE R W 4441, 4391

Phyla nodiflora (L.) Greene H N S R W 4372

Verbena brasiliensis Vell. H SA S A R W 4195

Verbena halei Small H N S E N 4297

Verbena xutha Lehm. H N S R N 4495

Violaceae

Viola sororia Willd. var. missouriensis H N a F W 4318

(Greene) McKinney

Viscaceae

Phoradendron serotinum (Raf) M.C. Johnston H N SE P N 4322

subsp. serotinum

Vitaceae

Ampelopsis arborea Koehne V N Sie R W 4422, 4344

Ampelopsis cordata Michx. V N SE F W 4421, 4270

Parthenocissus quinquefolia (L.) Planch V N E F W 4380

Vitis cinerea (Engelm.) Millardet var. cinerea V N S F N 4497, 4269

Vitis mustangensis Buckley V N TX R W 4021

Vitis rotundifolia Michx. V N SE F W 4398

Life form. Tree (T); shrub or subshrub (S); vine (V); broad-leaved herb (Monocot or Dicot) (H); graminoid (G); fern (F).

Origin. United States (N); Asia (As); Africa (Af); cosmopolitan (C); China (Ch); Europe (E); Eurasia (Ea); India (I); Japan (J);

Mediterranean (M); South America (SA).

Distribution. Throughout US (US); mostly limited to Texas (TX); Southern (S); Southeastern (SE); Southwestern (SW); Eastern

(E); Central (C).

Introduction reason. Accidental arrival (A); Crop (C); forage (F); ornamental (O); ; unknown (U).

Habitat. Forest or woodland (F); prairie, field or savanna (P); wetland (W); ruderal (R); garden weed (G); ornamental (Or).

Weed status. Weedy (W); non-weedy (N) per Kartesz & Meacham (2001).

Collection Number. All specimens were collected by I.S. Elsik & W.W. Pruess, except for a few collected by L. Brown denoted

by (LB) or Jeremy Caves (JC). *One species, Ir quamociit, is listed as sight record only.

710 Journal of the Botanical R h Institute of Texas 1(1)

ACKNOWLEDGMENTS

This study was funded by a gift to Rice University from the Botanic Garden of Houston. We thank many

volunteers for their assistance, including Katy Emde, Walt Hesson, Ray Jordan, Barbara Kertz, TJ Marks,

Irene Pendergrast, Lelia Rodgers, and Lan Shen. We also thank student assistants Niki von Hedemann and

Chandrika Achar. We are most grateful to Merrill O'Neal for his enthusiasm and encouragement.

REFERENCES

BROWN, L.E. 1992. Cayratia japonica (Vitaceae) and Paederia foetida (Rubiaceae) adventive in Texas. Phytologia.

/2:45-47.

BROWN, L.E., B.R. MAcRoserts, M.H. MacRoserts, P.A. HARCOMBE, W.W. Pruess, I.S. ELsik, and D. Johnson. 2005. Annotated

checklist of the vascular flora of the Turkey Creek Unit of the Big Thicket National Preserve, Tyler and Hardin

Counties, TX. Sida 20:781-795.

Bruce, K. G. CAMERON, PA. HARCOMBE, and G. JugiNskv. 1997. Introduction, impact on native habitats, and management

of the weedy invader, Chinese tallow tree (Sapium sebiferum (L.) Roxb.). Nat. Areas J. 17:255- 260.

ConntLL, DJ. and M.C. Johnston. 1970. Manual of the vascular plants of Texas. Texas Research Foundation,

Renner.

Crawley, M.J., PH. Harvey, and A. Purvis. 1997. Comparative ecology of the native and alien floras of the British

Isles. In: J. Silvertown, M. France and J.L. Harper, eds. Plant life histories: ecology, phylogeny and evolution.

Cambridge Univ. Press, Cambridge, UK. Pp 36-56.

DeCAanpibo, R. and E.E. Lamont. 2004. The historical and extant vascular flora of Pelham Bay Park, Bronx County,

New York 1947-1998. J. Torrey Bot. Soc. 131:368-386.

DECANDIDO, R., A.A. Muir, and M.B. GaraiuLLo. 2004. A first approximation of the historical and extant vascular flora

of New York City: Implications for native plant species conservation. J. Torrey Bot. Soc. 131:243-251.

DeLong, M.K., K.K. Joc, J.R. JOHANSEN, and G.J. WiLoeR. 2005. Floristic survey of a highly disturbed wetland within

Shaker Median Park, Beachwood (Cuyahoga County), Ohio. Ohio J. Sci. 105:102-105.

Dicas, Jr., G.M., B.L. LirscowB, M.D. Reep, and R.J. O'Kennon. 2006. Illustrated flora of East Texas. Vol. 1. Sida, Bot. Misc.

No. 26. BRIT Press. Ft Worth, TX.

DRAYTON, B. and R.B. Primack. 1996. Plant species lost in an isolated conservation area in Metropolitan Boston from

1894 to 1993. Conserv. Biol.10:30-39.

FENNEMAN, N.M. 1938. Physiography of the eastern United States. McGraw-Hill. New York and London.

FLEMING, K.M., J. SincHurst, and W.C. HoLmes. 2002. Vascular flora of Big Lake Bottom Wildlife Management Area,

Anderson County, TX. Sida 20:355-372.

FNA. 1993+, Flora of North America north of Mexico. 12+ vols. Flora of North America Editorial Committee, eds.

New York and Oxford.

GouLb, F.W. 1975. Texas plants. A checklist and ecological summary. Texas Agric. Exp. Stat., Texas A&M University,

College Station, TX. MP-585/Revised.

Joc, S.K., J.T. KARTESZ, J.R. JOHANSEN, and G.J. Wiper. 2005. Floristic Study of Highland Heights Community Park,

Cuyahoga County, Ohio. Castanea 70:136-145.

Jones, S., J.K. Wierr, and PM. Montcomery. 1997. Vascular plants of Texas: a comprehensive checklist including

synonymy, bibliography, and index. University of Texas Press, Austin.

Jones, S. and J.K. Wierr. 2003. Updated checklist of the vascular plants of Texas. (unpublished CD)

KARTESZ, J.R. and C. Meacham. 1999. Synthesis of the North America Flora. North Carolina Botanical Garden, Chapel

Hill.

Mack, R.N. and W.N. Lonspate. 2001. Humans as global plant dispersers: getting more than we bargained for.

BioScience 51:95-102.

Marks, PL., F.R. Westey, and S. Ganpescu. 2006. The flora of the Finger Lakers Region of central New York: changes

in the late 1800s and 1900s. MS in review

Harcombe et al., Flora of a iland park site, Harris County, Texas 711

MacRostRrs, B.R., M.H. MacRoserts, and L.E. Brown. 2002. Annotated checklist of the vascular flora of the Hickory

Creek Unit of the Big Thicket National Preserve, Tyler County, TX. Sida 20:781-795.

MacRoserts, M.H. and B.R. MacRoserts. 2003. The east-west transition of flora in Texas, a biogeographical analysis.

Sida 20:1693-1700.

MacRoseets, M.H. and B.R. MacRoserts. 2004. The Post Oak Savanna: a floristic assessment of its uniqueness. Sida

21:399-407.

Moore, R.C. and S. Aronow. 1994. A preliminary geomorphological and archaeological assessment of the City of

Houston Herman Brown Park, Harris County, Texas. In: Herman Brown Park site analysis C.I.P. No. F-0365-01-2.

Background Data. Document prepared by Llewelyn-Davies Sahni, Inc. for the City of Houston. Pp 34-70.

NatureServe. 2005. NatureServe Explorer: An online encyclopedia of life [web application]. Version 4.4. NatureServe,

Arlington, Virginia. http://www.natureserve.org/explorer. Accessed 3 Aug 2006)

NCDC 2004. Climatography of the United States No. 20. 1971-2000. pp 146A-E. National Climatic Data Center.

http://www.ncdc.noaa.gov/oa/normals/usnormals.html. Issue date February 2004. Accessed 3 Aug 2006

Pysek, P. 1998. Alien and native species in Central European urban floras: a quantitative comparison. J. Biogeog.

25:144-163.

Rosen, D.J., S.D. Jones, and V.E. Rettic. 2003. A floristic survey of Big Branch National Wildlife Refuge, St Tammany

Parish, Louisiana. Sida 20:1189-1216.

REICHERD, S.H. and P. Waite. 2001. Horticulture as a pathway of invasive plant introductions in the US. BioScience

51:103-113.

SES 1976. Soils of Harris County, TX. Soil Conservation Service (now Natural Resources Conservation Service).

Washington, D.C.

SHELBY, C.A., M,K. McGowan, S. Aronow, W.L. Fisher, L.F. Brown, JR, J.H. McGowen, CG Groat, and V.E. Barnes. 1968

(Revised 1992). Geologic atlas of Texas: Beaumont Sheet. Bureau of Economic Geology, University of Texas,

Austin, TX.

SINGHURST, J.R., J.C. CATHY, D. PRocHASKA, H. HAUCKE, G.C. KroH, and W.C. HoLmes, 2003. The vascular flora of Gus Engeling

Wildlife Management Area, Anderson Co., TX. Southeastern Nat. 2:347-368.

TOES. 1993. Endangered, threatened, and watchlists of Texas Plants. Texas Organization for Endangered Spe-

cies. Publication 9. 34 Revision. August 1993, with updates. http://www.csdl.tamu.edu/FLORA/toesplants.

htm. Accessed 2 Jan 2007.

TURNER, B.L., H. NicHoLs, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. Sida Bot. Misc., No. 24.

BRIT Press, Fort Worth, TX.

Texas ADMINISTRATIVE Cope. 2005. Quarantines and Noxious Plants, Chapter 19. State of Texas.

USDA. 2006. Federal noxious weed List (as of June 30, 2006). US Department of Agriculture http://www.USDA.

gov/ppq/weeds. Accessed 2 Jan 2007.

WEAKLEY, A.S., K.D. PATTERSON, S. LANDAAL, and M. Pyne. 1998. International classification of ecological communities:

terrestrial vegetation of the southeastern United States. Working Draft. The Nature Conservancy, Chapel Hill,

Wu, S-H., EF. Hse, S-M. Chaw, and M. Remanek. 2004. Plant invasions in Taiwan: Insights from the flora of casual

and naturalized alien species. Diversity Distrib. 10:394—362.

712 Journal of the Botanical R h Institute of Texas 1(1)

BOOR NOTICES

Wu ZneNGYI and Peter H. RAVEN (EDITORIAL Co-Cuairs). 1994. Flora of China Illustrations, Vol. 14: Apiaceae

through Ericaceae. (ISBN 1-930723-51-2, hbk.). Missouri Botanical Garden Press, PO. Box 299, Saint

Louis, MO 63166-0299, U.S.A. (Orders: www.mbgpress.org, orders@mbgpress.org, 314-577-9547,

314-577-9594 fax). $125.00, 726 pp., line drawings, 8%" x 111A".

“This publication is the 11th in a series of 25 volumes of the Flora of China Illustrations. It accompanies the Flora of China (FOC) Volume

14, which was published in 2005 and treated 132 Benes Dun 1498 species in nine familes. This volume of illustrations includes 697

figures representing a total of 1013 species in the ), Aucubaceae (7 species), Clethraceae (7 species), Cornaceae (23

species), Diapensiaceae (5 species), Ericaceae (484 species), TERNI (4 species), Mastixiaceae (2 species), and Toricelliaceae (2

species).” The FOC Illustrations include line drawings previously published in the Chinese language Flora Reipublicae Popularis Sinicae

(FRPS) and additional ones for taxa not or insufficiently illustrated in FRPS.—Guy Nesom, Botanical Research Institute of Texas, 509 Pecan

Street, Fort Worth, TX 76102-4060, U.S.A.

Mary E. BARKWORTH, KATHLEEN M. CAPELS, SANDY LONG, LAUREL K. ANDERTON, and MichHarL B. Prep (eds.). 2007.

Flora of North America North of Mexico, Vol. 24: Magnoliophyta: Commelinidae (in part):

Poaceae, Part 1. (ISBN 978-0-19-531071-9, hbk.). Oxford University Press, 2001 Evans Road, Cary,

NC 27513, U.S.A. (Orders: www.oup.com, 1-800-451-7556, 919-677-1303 fax). $95.00, 944 pp.,

line drawings, 8%" x 1144",

This volume is, both logically and numerically, the first of the two volumes in the Flora of North America devoted to the Poaceae (Grass

Family). [Volume 25 was published in 2003]. It contains the family description and a key to all the tribes recognized in the two volumes.

In addition, it includes treatments for genera in four subfamilies: Pharoideae [Phareae], Bambusoideae [Bambuseae, Olyreae], Eh-

rhartoideae [Ehrharteae, Oryzeae], and Podideae [Brachyelytreae, Nardeae, Diarrheneae, Meliceae, Stipeae, Brachypodieae, Bromeae,

Triticeae, Poeae].” “For the most part, the authors of the treatments determined the limits of the genera, species, and infraspecific taxa.

The most controversial generic decisions are those in the Triticeae. At the species level, changes from existing treatments will be found

in several genera n Poa, Elymus, Calamagrostis, Deschampsia, and Puccinellia." In all, 236 genera and 1373 species are treated in the

two FNA grass volumes. Native represent 6596 of the total.

The county-level distribution g feature of the FNA grass volumes. The maps are prepared from a geographic

database maintained at the Intermountain Herbarium. “Larger, more informative versions of the maps are available on the Web at http://

herbarium.usu.ed

^webmanual/. They are updated as well-documented information is submitted." Combined [literature] citations for

the two volumes are available at http://herbarium.usu.edu/grassbib.htm.”—Guy Nesom, Botanical Research Institute of Texas, 509 Pecan

Street, Fort Worth, TX 76102-4060, U.S.A.

FLORA or NORTH AMERICA EDITORIAL COMMITTEE (eds.). 2007. Flora of North America North of Mexico, Vol.

27, Bryophyta, Part 1. (ISBN 978-0-19-531823-4, hbk.). Oxford University Press, 2001 Evans Road,

Cary, NC 27513, U.S.A. (Orders: www.oup.com, 1-800-451-7556, 919-677-1303 fax). $95.00, 944

pp., line drawings, 8%" x 1144".

Press Release from Oxford University Press.—"Flora of North America Volume 27 is the result of the collaboration of respected U.S. and

Canadian botanical institutions. It treats the first half of the mosses, including 32 of the acrocarpous families, with introductory chapters

on bryophyte morphology, the history of North America fl and economic uses. The 84 species of sphagnum, of consider-

able economic importance, are examined fully, as well as the 37 genera and 621 species that make up the taxa."

Features include: 1) full descriptions of the taxa; 2) 136 panels of detailed line drawings; 3) geographic distributions at the prov-

ince and state level; 4) distribution maps for native and established species, and 5) a list of synonyms currently in use for the accepted

names.—Barney Lipscomb, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A.

J. Bot. Res. Inst. Texas 1(1): 712. 2007

HYPERICUM ADPRESSUM (CLUSIACEAE) NEW TO

ARKANSAS AND THE OUACHITA MOUNTAINS, U.S.A

C. Theo Witsell

Arkansas Natural H leritage Commission

1500 Tower Building

323 Center St.

Little Rock, Arkansas 72201, U.S.A

theo@arkansasheritage.org

ABSTRACT

Hypericum adpressum Bart. (Clusiaceae) is reported as new to the state of Arkansas. It was collected from a complex of natural depression

wetlands in the eastern Ouachita Mountains of Saline County. A description of the habitat and list of associated species is provided.

RESUMEN

Se cita Hypericum adpressum Bart. (Clusiaceae) como nuevo del estado de Arkansas, en un mplejo de

en las montafias Ouachita del este del condado de E Se aporta una descripción del hábitat y una dd de especies asociadas.

INTRODUCTION

Creeping St. John's wort, Hypericum adpressum Bart., (Fig. 1) is endemic to the eastern United States and

is considered to be a species of conservation concern in all 19 states from which it is known (NatureServe

2006). Based on data from state natural heritage programs, this species is considered to be extant in 14 states,

with a state conservation status rank of either S1 (critically imperiled) or S2 (imperiled), and is considered

to be extirpated or possibly extirpated in 5 states. It has a global conservation status rank of G3 (vulner-

able) and has the following ranks in each state: CT (SH), DE (S2), GA (S2?), IL (S1), IN (SD), KY (SH), MD

(S1), MA (S2), MI (S1), MO (S1), NJ (S2), NY (S2), NC (SH) PA (SX) RI (92) SC (S1), TN (S1), VA (S1), and

WV (SH), where S1 = critically imperiled, S2 = imperiled, SH = possibly extirpated, and SX = presumed

extirpated (NatureServe 2006). Habitats listed for the species include marshes, shores, marshy shores, wet

meadows, bogs, swales, ditches, moist depressions in sand prairies, and along the shores and in shallow

water zones of freshwater ponds (Gleason & Cronquist 1991; Godfrey & Wooten 1981; Radford et al. 1968;

Yatskievych 2006; Enser 2001).

Hypericum adpressum was listed (as a woody plant) for Arkansas without comment or citation of a voucher

by Moore (1941) and was also listed without comment or citation by Demaree (1943). Tucker (1976) excluded

it from his treatment of the woody flora of Arkansas, noting its inclusion by Moore (1941) in a checklist of the

ligneous flora of Arkansas despite the species being “wholly herbaceous.” Smith (1988; 1994) excluded the

species altogether from the flora of Arkansas noting that while Moore (1941) and Demaree (1943) included

it, Adams (1973) did not include Arkansas in the range for H. adpressum. No specimens of the species have

been reported in the recent inventory of herbaria by the Arkansas Vascular Flora Committee, who excluded

it from the 2006 Checklist of the Vascular Plants of Arkansas (Arkansas Vascular Flora Committee 2006).

On 12 April 2006, while surveying a complex of natural depression wetlands on a terrace of the Alum

Fork Saline River in northern Saline County, I observed several large colonies of young sterile stems and the

previous year’s dead stems of a rhizomatous, perennial Hypericum growing on the margins of two ponds (Fig.

2). Based on the habitat and rhizomatous character of the plants, I thought the plants might be H. adpressum,

which I had seen in southeastern Missouri the previous year. Two rammets were removed from the Alum

Fork population and maintained in cultivation until mid June 2006, when the plants began to flower. Based

on reproductive features I confirmed that the plants were H. adpressum. I returned to the site on 21 June

2006 to collect voucher specimens, take photographs, and collect data for the Arkansas Natural Heritage

J. Bot. Res. Inst. Texas 1(1): 713 — 716. 2007

714 Journal of the Botanical R h Institute of Texas 1(1)

Fic. 1. Hyperi dj Saline County, Arkansas. Not luteleaf ^ Fic. 2. Habitat for H. adp p AE T T

margins. 20 June 2006. on niceties of Alum iu Saline jd Saline County, Arkansas. Emergent

I. adpressum. 20 June 2006

Program. I visited the ponds again on 17 August 2006 to collect fruiting specimens and conduct additional

inventory. Tens of thousands of stems of Hypericum adpressum were found along the margins of these ponds

within the zone of seasonal water fluctuation. The ponds were Peau) dry on 17 M 2006.

Voucher Specimens: ARKANSAS. Saline Co.: Ouachita Mountains, locally c f rfl i d l

natural ponds on terrace above Alum Fork Saline River, 0.4 mi (0.6 km) SW JofL Lake Winona Dam, N34.79247, W92.85592 (NAD 83,

Zone 15), Paron 7.5’ quad. (specimen in flower), 20 Jun 2006, T. Witsell & J. Pelton 06-273 (anhc [herbarium of the Arkansas Natural

Heritage Commission], MO, UARK); same locality (specimen in fruit), 17 Aug 2006, T. Witsell 06-397 (anhc [herbarium of the Arkansas

Natural Heritage Commission], MO, UARK).

DISCUSSION

Natural depression wetlands are found along several of the larger streams in the Ouachita Mountains and

are known to support disjunct or relict populations of plant species more typical of the Gulf Coastal Plain

and Mississippi Alluvial Plain Ecoregions (e.g. Pinus taeda L., Quercus lyrata Walter, Myrica cerifera L., Carex

hyalinolepis Steud., C. gigantea Rudge., C. lupuliformis Sartwell ex L.H. Dewey, Eleocharis microcarpa Torr.,

Panicum verrucosum Muhl., Eupatorium album L. var. glandulosum (Michx.) DC., Acer rubrum L. var. drum-

mondii (Hook. & Arn. ex Nutt.) Sarg., Lyonia mariana (L.) D. Don., Planera aquatica J. F. Gmel., and Taxodium

distichum (L.) Rich). Geomorphologically, these wetlands occur in abandoned stream channel scars on older

stream terraces above present day floodplains. Along the Alum Fork Saline River, Middle Fork Saline River,

and North Fork Saline River in the eastern Ouachita Mountains, these wetlands typically occur as small 0.1

Witsell, Hypericum adpressum new to Arkansas and the Quachita Mountains 715

to 1.6 ha (0.25 to 4 acre) forested depressions similar in appearance and species composition to the hydric

flatwoods of the Mississippi Alluvial Plain and Gulf Coastal Plain. These wetlands are poorly drained, have

clay soils, and typically have standing water up to 10 cm deep in the winter and spring but dry out on the

surface in the summer. They are typically dominated by Quercus phellos L., Quercus nigra L., Liquidambar

styraciflua L., Fraxinus pennsylvanica Marshall (and in rare cases support Planera aquatica, Quercus lyrata and

Taxodium distichum) with scattered shrubs and a graminoid understory rich in sedges (Carex and Eleocharis

spp.) and rushes (Juncus spp.) and often with extensive bryophyte cover (particularly Sphagnum spp.).

In at least three sites on the Alum Fork and North Fork in northern Saline County, these wetlands

occur with deeper, open water ponds fringed by concentric zones of emergent herbaceous vegetation and

shrubs. Water levels fluctuate throughout the growing season as water is lost to evaporation, exposing a ring

of mudflats colonized primarily by annual species. The ponds at the Alum Fork site are dominated by open

water with a fringe of scattered Cephalanthus occidentalis and dense stands of emergent vegetation dominated

by perennial species including Juncus effusus L., Juncus nodatus Coville, Carex lurida Wahlenb., Carex joorii

L.H. Bailey, Carex ozarkana P. Rothr. & Reznicek, Carex gigantea Rudge*, Dulichium arundinaceum (L.) Brit-

ton*, Rhynchospora glomerata (L.) Vahl, Scirpus cyperinus (L.) Kunth, Rhexia virginica L., Xyris jupicai Rich.,

Viola lanceolata L., Eleocharis quadrangulata (Michx.) Roem. & Schult. in Roem. et al, Sagittaria platyphylla

(Engelm.)J.G. Sm., Hypericum lobocarpum Gatt., Triadenum walteri (J.F. Gmel.) Gleason, and Panicum rigidulum

Bosc ex Nees. As the water recedes over the course of the summer, Hypericum mutilum L., Eleocharis obtusa

(Willd.) Schult., Eleocharis microcarpa Torr.*, Oldenlandia boscii (DC.) Chapm., Lindernia dubia (L.) Pennell,

and Gratiola brevifolia Raf.* dominate the exposed soil of the pond margins along with the perennial Juncus

repens Michx. [* = species tracked as elements of conservation concern by the Arkansas Natural Heritage

Commission].

The presence of small spoil piles with trees growing on them around several of the ponds at the Alum

Fork site indicate that they were partially excavated in the past in an attempt to deepen them, perhaps as a

source of water for cattle. However, it is clear from the size of the ponds relative to the size of the spoil piles

as well as the physical, geomorphological, and floristic similarity of these ponds to undisturbed terrace

wetlands nearby that they are natural wetlands.

Aside from H. adpressum, there are 15 taxa of Hypericum known from Arkansas (Arkansas Vascular Flora

Committee 2006). Of these, H. sphaerocarpum Michx., which is also often rhizomatous and herbaceous, is the

most likely to be confused with H. adpressum. It can be distinguished from H. adpressum by its flat leaf margins

(vs. revolute in H. adpressum), its broadly ovoid to globose seed capsules with 4 to 8 seeds (vs. ellipsoid to

ovoid capsules with numerous seeds in H. adpressum), and 2.0 to 2.7 mm long seeds (vs. 0.6 to 0.8 mm in H.

adpressum) (Yatskievych 2006). H. sphaerocarpum often also has suffrutescent lower stems and rootstock (vs.

entirely herbaceous in H. adpressum) (Yatskievych 2006). In Arkansas H. sphaerocarpum is typically found

in calcareous glades and woodlands, shale barrens, and prairies, but is reported also from banks of streams

and rivers, fens, and margins of ponds and lakes in Missouri by Yatskievych (2006). H. punctatum Lam.,

H. perforatum L., and H. pseudomaculatum Bush ex Britton are occasionally rhizomatous but differ from H.

adpressum by having characteristic black glands on the sepals and/or petals.

The nearest known extant site to the Saline County population for H. adpressum is approximately 395

km (245 miles) to the northeast in Scott County, Missouri where it occurs in a moist depression in a sand

prairie near Blodgett in the Mississippi Alluvial Plain (Tim Smith pers. comm.; NatureServe 2006). There

are also two historical collections from Mississippi and Scott Counties, Missouri, both from 1933. (Tim

Smith pers. comm., Yatskievych 2006); Aside from these records, H. adpressum is known only from east of

the Mississippi River (NatureServe 2006). It is possible that the inclusion of H. adpressum for Arkansas by

Moore (1941) and Demaree (1943) was based on a misidentified specimen that has since been annotated

as another Hypericum species or that it was based on a properly-identified sight record or a specimen now

lost.

When garden-grown plants of H. adpressum were compared to in situ plants of their parent population

716 Journal of the Botanical R h Institute of Texas 1(1)

it was apparent that the cultivated plants, which grew in pots with well-drained soil that was allowed to

dry out on occasion, branched profusely from the axils of the lower leaves (a condition absent in the wild

plants). Wild plants also frequently had a shiny upper leaf surface compared to dull upper leaf surfaces in

the cultivated plants. Both wild and cultivated plants grew in full sun. Wild plants in the wettest zones of

the pond margins had a spongy, thickened lower stem consistent with plants called variety spongiosum by

Robinson (1902) which was later downgraded to forma spongiosum by Fernald (1949). Both this form and

the more typical form without this character were present in the population and were obviously correlated

to the amount of water present where they grew.

ACKNOWLEDGMENTS

Thanks to Paul McKenzie (US Fish and Wildlife Service), Tim Smith, and Bob Gillespie (both of the Missouri

Department of Conservation) for showing me plants of H. adpressum in the field in southeastern Missouri

in 2005. This certainly led to my finding it in Arkansas. Thanks also to George Yatskievych for additional

information on the species in Missouri and to John Pelton for the photographs of the plant and habitat.

Tanya Miller-Witsell provided a Spanish translation of the abstract. Jennifer Akin, Staria Vanderpool and an

anonymous reviewer provided comments which strengthened an earlier draft of this manuscript. Thanks

also to Karen Smith and Bill Holimon of the Arkansas Natural Heritage Commission.

REFERENCES

ApAMSs, P. 1973. Clusiaceae of the southeastern United States. J. Elisha Mitchell Sci. Soc. 89:62—71.

ARKANSAS VASCULAR FLORA Committee. 2006. Checklist of the vascular plants of Arkansas. University of Arkansas Her-

barium. Fayetteville.

Demart, D. 1943. A catalogue of the vascular plants of Arkansas. Taxodium 1:1—88.

Enser, R.W. 2001. Hypericum adpressum (creeping St. John's-wort) conservation and research plan. New England

Wild Flower Society. Framingham, MA, USA (http://www.newfs.org).

FERNALD, M.L. 1949. Hypericum adpressum Bart., forma spongiosum (Robinson), stat. nov. Rhodora 51:112.

GLEASON, H.A. and A. Cronauist. 1991. Manual of the vascular plants of northeastern United States and adjacent

Canada. New York Botanical Garden Press. Bronx.

Goorrey, R.K. and J.W. Wooten. 1981. Aquatic and wetland plants of the southeastern United States, dicotyledons.

University of Georgia Press. Athens.

Moore, D.M. 1941. Checklist of the ligneous flora of Arkansas. Proc. Ark. Acad. Sci. 1:41—55.

NatureServe. 2006. NatureServe Explorer: an online encyclopedia of life [web application]. Version 6.1. NatureServe,

Arlington, VA. availablehttp://www.natureserve.org/explorer. (Accessed January 25, 2007).

RADFORO, A.E., H.E. Ahles, and CR. BELL. 1968. Manual of the vascular flora of the Carolinas. University of North

Carolina Press. Chapel Hill.

Rosinson, B.L. 1902. Two new Hypericums of the adpressum group. Rhodora 4:135— 137.

Smith, E.B. 1988. An atlas and annotated list of the vascular plants of Arkansas, 2"? ed. Privately printed. Fayette-

ville, AR.

StH, E.B. 1994. Keys to the flora of Arkansas. University of Arkansas Press. Fayetteville.

Tucker, G.E. 1976. A guide to the woody flora of Arkansas. Ph.D. dissertation. University of Arkansas at Fayette-

ville., AR.

YATSKIEVYCH, G. 2006. Steyermark's flora of Missouri. Volume 2. Missouri Botanical Garden Press. St. Louis, MO.

MONARDA LINDHEIMERI (LAMIACEAE): NEW TO ARKANSAS

Walter C. Holmes Jason R. Singhurst

Department of Biology Wildlife Diversity P! ogram

Baylor University Texas Parks and Wildlife Department

Waco, Texas 76798-7388 U.S.A. 3000 South IH-35, Suite 100

walter_holmes@baylor.edu Austin, Texas 78704 U.S.A.

jason.singhurstetpwd.state.tx.us

ABSTRACT

Monarda lindheimeri is reported as new to Arkansas. The overall distribution of the species in Louisiana and Texas and a comparison

with a similar species are also discussed.

RESUMEN

Se cita como nueva para Arkansas Monarda lindheimeri. También se comenta la distribución total de la especie en Louisiana y Texas, y

una comparación con una especie semejante.

Monarda lindheimeri Engelm. & A. Gray ex A. Gray occurs mainly in east Texas, where Turner et al. (2003)

mapped it in 15 counties (Anderson, Brazos, Cass, Fannin, Galveston, Gonzales, Grayson, Grimes, Harris,

Harrison, Montgomery, Morris, Walker, Waller, and Upshur). Additional Texas records include Colorado

(Brown 7472, SBSC), Hardin (Singhurst 2161, BAYLU), Liberty (Brown 15414, SBSC), Madison (Neill 1732,

TAMU), Marion (Holmes 9967 & Singhurst, BAYLU), and San Jacinto (Brown 7566, SBSC) counties. The few

scattered records from Louisiana include Calcasieu Parish (Correll & Correll 1941), St. Landry Parish (Mc-

Clintock & Epling 1942), Acadia and Jefferson Davis parishes (Thomas & Allen 1998), and Rapides Parish

(Camp Beauregard, Leonard 1560, US!, determined and annotated by R. Scora).

The species is characterized by its single, terminal flower cluster (glomerule) composed of white flow-

ers. The stamens are exserted beyond the upper lip of the corolla. Plants are more or less unbranched above

and have midstem petioles 3-7 mm long or longer (~ 9 mm) if spreading pilose hairs are present. Although

closely related to Monarda fistulosa L. var. mollis (L.) Benth., M. lindheimeri has been maintained as distinct

by McClintock and Eppling (1942), Scora (1967), and Turner (1994, 2003). The cited variety of M. fistulosa

may be distinguished from M. lindheimeri by being branched above, thus having several glomerules in a

loose corymb, has flowers that are more pinkish to lavender, but occasionally white with a pinkish tint, and

its midstem petioles are 8 mm or greater in length and are never spreading pilose.

Based upon the specimens cited below, we report the occurrence of Monarda lindheimeri in Arkansas.

Specimens: ARKANSAS. Miller Co.: County Road 32, 0.1 mi E of jct. with County Road 107 near Brightstar, 24 May 2006, Holmes

13620 & Singhurst (BAYLU); County Road 107, 0.7 mi S of jct. with AR Hwy 160 near Brightstar, 24 May 2006, Holmes 13628 & Singhurst

(BAYLU).

Three populations of Monarda lindheimeri were found growing in red clay soils on exposed iron ore ridges on

the Weches Geologic Formation (University of Texas Bureau of Economic Geology 1979). The first population

mentioned above consisted of 55 genets, while the second population included 20 genets. Specimens were

not collected from the third population, which was located about 200 m west from the first collection men-

tioned above. This population was comprised of 37 genets. Characteristic vegetation included Pinus elliottii,

Quercus marilandica, Q. stellata, Liquidambar styraciflua, Diospyros virginiana, Juniperus virginiana, Viburnum

rufidulum, Vaccinium arboreum, Rhus copallina, Aesculus pavia, Berchemia scandens, Toxicodendron radicans,

Schizachyrium scoparium, Panicum sphaerocarpon, Symphyotrichum patens, Hypericum punctatum, Helianthus

hirsuta, Liatris elegans, Vernonia missurica, Stylosanthes biflora, Hypochaeris hirsuta, Lechea tenuifolia, Ascyrum

hypericoides, Tephrosia virginiana, Echinacea angustifolia, Plantago aristata, and Ruellia humilis.

J. Bot. Res. Inst. Texas 1(1): 717 — 718. 2007

718 Journal of the Botanical R h Institute of Texas 1(1)

ACKNOWLEDGMENTS

We thank the herbaria (DUKE, SBSC, TAES, TAMU, TEX/LL, and US) whose specimens, in part, made

this study possible. Special thanks are due Gina Gollub, a student and herbarium worker at BAYLU, who

assisted with preparation of the manuscript. Staria Vanderpool and an anonymous reviewer provided help-

ful comments.

REFERENCES

Correll, D.S. and H.B. CorreLt. 1941. A collection of plants from Louisiana. Amer. Midland Naturalist 26:30-64.

McCuntock, E. and C. EPuiNG. 1942. A review of the genus Monarda. Univ. Calif. Publ. Bot. 20:147-194.

Scora, R. 1967. Interspecific relationships in the genus of Monarda (Labiatae). Univ. Calif. Publ. Bot. 41:1—71.

Thomas, R.D. and C.M. ALLEN. 1998. Atlas ofthe vascular flora of Louisiana. Vol. 3: dicotyledons. Fabaceae-Zygophyl-

laceae. Louisiana Department of Wildlife & Fisheries, Baton Rouge.

Turner, B.L. 1993. Taxonomic treatment of Monarda (Lamiaceae) for Texas and Mexico. Phytologia 77:56-79.

Turner, B.L., H. NicHoLs, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. Vol. 1. Sida Bot. Misc. 24.

Botanical Research Institute of Texas, Fort Worth.

UNIVERSITY OF TEXAS BUREAU or ECONOMIC GEOLOGY. 1979. Texarkana map. University of Texas, Austin.

NEW RECORDS OF WETLAND AND RIPARIAN PLANTS IN

SOUTHERN CALIFORNIA, WITH RECOMMENDATIONS AND ADDITIONS

TOCDEENATIONALPEISTOLTDTPEANT SPEGIES THAT OCCUR DINSWVETEAINIDS

Richard E. Riefner, Jr. Steve Boyd

Research Associate Herbarium (RSA-POM)

Rancho Santa Ana Botanic Garden Rancho Santa Ana Botanic Garden

1500 North College Avenue 1500 North College Avenue

rriefner@earthlink.net steve.boyd@cgu.edu

ABSTRACT

Panicum coloratum, Panicum virgatum, and Setaria megaphylla are exotic species reported for the first time for California. The following

new or noteworthy records of native and non-native vascular plants are reported from wetland and riparian habitats in coastal southern

California: Amaranthus blitum subsp. emarginatus, Atriplex polycarpa, Bacopa monnieri, Beta vulgaris subsp. maritima, Chloris truncata,

Ehrharta erecta, Elytrigia rey Epilobium brachycarpum, Eriochloa aristata, Glinus radiatus, Limonium ramosissimum, Limonium indet., On-

cosiphon piluliferum, M dichotomiflorum subsp. dichotomiflorum, Paspalum urvillei, Psilocarphus chilensis, Salvinia molesta, and Setaria

adhaerens. Of these, Amaranthus blitum subsp. emarginatus, Beta vulgaris subsp. maritima, Chloris truncata, Ehrharta erecta, Limonium

ramosissimum, Oncosiphon piluliferum, Panicum coloratum, Panicum virgatum, Psilocarphus chilensis, Salvinia molesta, Setaria adhaerens, and

Setaria megaphylla are facultative wetland plants in California herein proposed for addition to the National List of Plants that Occur in

Wetlands. Supplemental information is also provided to revise the current wetland indicator status for Epilobium brachycarpum, Glinus

radiatus, and Paspalum urvillei. The new records presented herein suggest that focused floristic studies are needed to thoroughly docu-

ment the flora of ae and ud communities in the South Coast ju In NEAN watershed urbanization and exotic species

ge Adlara ] ] ft)

in the v Wlid!ana modified the cl character or the native vegeta

tion and integrity of many wetland and riparian ecosystems in coastal southern CA

Key Wonps: exotic species, hydrophytes, riparian, Salvinia molesta, urban runoff, wetlands, wetland indicator status, wetland plant lists,

wildland-urban interface

RESUMEN

Panicum coloratum, Panicum virgatum, y Setaria megaphylla son especies exóticas citadas por vez primera para California. Se citan nuevos

o importantes registros de plantas vasculares nativas y no nativas de humedales y zonas ribereñas de la costa sureña de California:

Amaranthus blitum subsp. emarginatus, Atriplex polycarpa, Bacopa monnieri, Beta vulgaris subsp. maritima, Chloris truncata, Ehrharta

erecta, Elytrigia repens, Epilobium brachycarpum, Eriochloa aristata, Glinus radiatus, Limonium ramosissimum, Limonium indet., Oncosiphon

uliferum, Panicum dichotomiflorum subsp. dichotomiflorum, Paspalum urvillei, Psilocarphus chilensis, Salvinia molesta, y Setaria adhaerens.

Las siguientes especies: Amaranthus blitum subsp. emarginatus, Beta vulgaris subsp. maritima, Chloris truncata, Ehrharta erecta, Limonium

SS Mb dod um dei, Panicum coloratum, Panicum E Psilocarphus chilensis, Salvinia molesta, Setaria adhaerens, y

rd 1 del ess]

laLista Nacional de Plantas

que Hann en m. ales. Se proporciona también información MN para revisar el actual status indicador en humedales de

Epilobium brachycarpum, Glinus radiatus, y Paspalum urvillei. Los nuevos registros presentados aquí sugieren que son n ri

Bs pene

florísticos para documentar la flora de los humedales y las ribereñas de la región de la Costa Sur, además demue traqu la

urbanización de la cuenca hidrográfica y la presencia de especies exóticas en comunidades suburbanas y en el punto de contacto entre

tierras y urbanas han modificado considerablemente el aspecto de la vegetación nativa y la integridad de muchos humedales

y ecosistemas riberefios in la costa surefia de California.

INTRODUCTION

Wetlands are lands periodically covered by shallow water or where saturation is the dominant factor that

determines the nature of soil development and the types of plants and animals living in the soil and on its

surface (Tiner 1999; USFWS 2004). Thus, flooding or soil saturation occurs at a duration and frequency that

excludes many organisms not tolerant of the wetland environment (Brinson & Malvárez 2002). Wetlands

are generally characterized by one or more of the following three attributes: (1) at least periodically, the land

supports predominantly hydrophytes; (2) the substrate is predominantly undrained hydric soil; and (3) the

J. Bot. Res. Inst. Texas 1(1): 719 — 740. 2007

720 Journal of the Botanical R h Institute of Texas 1(1)

substrate is non-soil and is saturated with water or covered by shallow water at some time during the grow-

ing season of each year (Cowardin et al. 1979). The attributes of hydrology, hydric soils, and hydrophytic

vegetation implicit to wetland definitions, were introduced by the U.S. Fish and Wildlife Service (USFWS)

and remain an important and recurring theme in wetland regulatory policy in the United States (Shaw &

Fredine 1956; Mitsch & Gosselink 2000; Somerville & Pruitt 2006).

As wetland policy evolved, the U.S. Army Corps of Engineers (USACE), the primary agency responsible

for regulating wetlands, adopted a slightly different, but more restrictive definition than Cowardin et al.

(1979). The term wetlands is defined as those areas that are inundated or saturated by surface or ground

water at a frequency and duration sufficient to support, and that under normal circumstances do support,

a prevalence of vegetation typically adapted for life in saturated soil conditions (USACE 1987, 2006). This

definition requires that each of the three attributes of hydrology, hydric soils, and hydrophytic vegetation

typically be present to identify legal (or jurisdictional) wetlands. Although it is implicit that all wetlands

have wetland hydrology, the Cowardin et al. (1979) and other definitions indicate that only one of the three

parameters needs to be present (Batzer & Sharitz 2006). The USACE Wetland Delineation Manual, and the

Arid West Regional Supplement for the western states, provide the technical guidance and procedures neces-

sary to identify and delineate wetlands that may be subject to regulatory jurisdiction under Section 404 of

the Clean Water Act (USACE 1987). Wetland delineations are based on a three-parameter approach involving

regional indicators of hydrophytic vegetation, hydric soil, and wetland hydrology (USACE 2009).

Riparian habitats on the other hand are *mesic islands" of vegetation associated with narrow corridors

of land that border lakes, creeks, rivers or other waters (Gregory et al. 1991; Malanson 1993). Riparian eco-

systems are highly variable. They can support hydrophytic vegetation, upland vegetation, unvegetated areas,

or a mosaic of these types, but usually exhibit distinctive geomorphic features and vegetation that develops

in response to periodic flooding or exchange of surface or ground waters between rivers and streams and

adjacent habitats (USFWS 2004; USACE 2006). In addition, riparian habitats support many facultative

species adapted to and/or are tolerant of high soil moisture conditions that are not usually present elsewhere

in western arid landscapes, including many species of deciduous trees that are often restricted to riverine or

floodplain communities (Brinson et al. 1981; Holstein 1984; Smith et al. 1989; Pattern 1998). The indicators

of hydrophytic vegetation, hydrology, and hydric soils, are therefore important attributes used to classify

and characterize riparian habitats, which are also typically used to delineate and separate them from juris-

dictional wetlands (Bennett et al. 1989; USACE 2006).

Wetlands of the California Floristic Province are extremely varied and complex (Ferren & Fiedler 1993).

Rapid urbanization, poor historical d tation, and uncoordinated conservation efforts have often resulted

in the widespread and unmitigated loss of wetland resources in southern California (Ferren & Fiedler 1993).

Approximately 91% of California’s historic wetlands no longer exist, or the remaining habitats are often highly

degraded (USFWS 1977; Dahl 1990, 2000). In addition, many wetland types have been underestimated in their

diversity, richness, and unique functions and values. Since these losses occurred prior to detailed study, biolo-

gists and the general public may never fully understand or appreciate the composition, diversity, and functional

values of many unique and regionally rare wetland communities in southern California (Ferren et al. 1996).

Riparian ecosystems have also suffered great losses. Estimates of 90-9896 of the historic riparian habitats

of central and southern California have been lost or seriously degraded by agricultural development, expanding

urbanization, flood control projects, and other human-caused impacts (Katibah 1984; Swift 1984; Faber et al.

1989). Long-term human activities have also physically modified or permanently altered riparian ecosystems,

which have frequently resulted in a decline of native species diversity (Bowler 1989; Brinson 1990). Thus,

structurally complex riparian communities that support a diverse assemblage of native species are threatened

in southern California (Bowler 1989; Ferren & Fiedler 1993).

Paradoxically, as a result of expanding urbanization, the extent of wetland and riparian vegetation

communities has significantly increased in many suburban and urban watersheds. Expanding wetland and

riparian habitat in urban settings can be attributed to new water sources generated by over-irrigation of

Riefner and Boyd, Wetland and riparian plants in southern California 721

lawns and landscaping in commercial and residential development projects, discharge of municipal water

treatment plant effluent into drainages, and increased dry-season stream flows associated with decreased

precipitation infiltration and increased hard-surface runoff following storm events (Arnold & Gibbons 1996;

Paul & Meyer 2001; Greer & Stow 2003; Burkhart & Kelly 2005; Burkhart 2006; White & Greer 2006). In a

San Diego County case study, the acreage of riparian vegetation in the lower Los Pefiasquitos Creek watershed

increased by 56-7496 between 1969 and 1982, and increased by 118—12996 between 1988 and 2000, which

was accompanied by a shift from an historic oak-dominated riparian community, characterized by intermit-

tent stream flows, to a willow-dominated community associated with urban-induced perennial stream flows

(White & Greer 2006). With ongoing development, more buildings and paved areas mean greater quanti-

ties of storm runoff, and growth in population and industry will also generate a proportional increase in

wastewater discharges into stream and river channels. In Orange County, it is predicted that base flows for

the Santa Ana River may increase by 100,000 acre-feet over the next 20 years due to urban development in

upstream areas (OCWD 2006). This phenomenon is especially important since almost no perennial streams

existed in southern California prior to urban development (Ellis & Lee 1919; Troxell 1948; Poland 1959;

Bader 1969). In addition, escalating urban development in the upper reaches of coastal watersheds is also

the likely source of new exotic plant propagules that are actively being dispersed downstream via urban

drainage systems into native plant communities. Similar circumstances have been documented in expand-

ing desert urban communities. In Las Vegas, Nevada, urban growth has generated increased discharge of

municipal waters, hard-surface runoff, and storm water flows into ephemeral washes that has promoted

development of extensive native wetland communities, which, over time, have been degraded by erosion

and the invasion of exotic species as population pressures increase (Shanahan & Crear 2004).

Although species richness may be high, many urban wetlands are floristically degraded, often with 1596

to more than 5096 of the plants present being introduced species (Magee et al. 1999; Burkhart & Kelly 2005;

Burkhart 2006). However, there has been little research published on the influence of urbanization-induced

hydrologic changes on wetland and riparian vegetation communities, and the distributions of native and

exotic plants in multi-use urban watershed environments (Poff et al. 1997; Dwire et al. 2000; Paul & Meyer

2001; Burkhart 2006). Other examples of large intermittent streams in the urban environments of southern

California that are now characterized by increased dry-season base flows and other altered hydrologic regimes,

and/or that support perennial flows across large segments of their drainage, include the lower watersheds

of the Los Angeles River and Arroyo Seco Creek in Los Angeles County, the Santa Ana River, San Gabriel

River, San Diego Creek, and San Juan Creek in Orange County, and Temescal Wash in Riverside County.

The composition and areal extant of the pre- and post-urban development of wetland and riparian vegetation

communities relative to new inputs of municipal water supplies has not been critically studied in many of

these and other watersheds in southern California.

The natural hydrologic regimes of many of our coastal waters have now become altered owing to the advent

of urbanization. Because identification of individual hydrophytes and hydrophytic vegetation communities is

a major determinant of federally regulated wetlands, it has become increasingly important to identify native

and exotic plants as wetland indicators in urban and urban fringe environments, which can further assist

biologists during wetland delineations (Tiner 1991, 1993, 2006). During wetland delineations, hydrophytic

vegetation decisions are based primarily on the wetland indicator status assigned to individual plant species

compiled on the National List of Plant Species that Occur in Wetlands (Reed 1988; USACE 2009).

The purpose of this paper is to identify overlooked native or previously unreported exotic hydrophytes

that would: 1) improve the accuracy and efficiency of wetland delineation procedures; 2) identify native

and exotic elements of hydrophytic vegetation communities important to the conservation and restoration

of wetland and riparian habitats in coastal southern California; and 3) demonstrate the need to carefully

inventory unique seasonal wetland habitats prior to development, and to establish ongoing floristic monitor-

ing of anthropogenic changes to wetland and riparian vegetation in southern California's rapidly urbanizing

communities

722 Journal of the Botanical R h Institute of Texas 1(1)

THE NATIONAL LIST OF PLANT SPECIES THAT OCCUR IN WETLANDS

The distribution and abundance of hydrophytes in the flora and vegetation of southern California often define

or characterize wetland and riparian ecosystems. The USFWS developed lists of wetland plants in order to as-

sist in the field identification and classification of wetland communities according to the Cowardin et al. (1979)

system (Reed 1988). Plant species that occur in wetlands (hydrophytes) are defined as species that demonstrate

the ability to achieve maturity and reproduce in an environment where all or portions of the soil within the

root zone is periodically saturated, continuously saturated, or inundated with water, and/or the substrate is at

least periodically deficient in oxygen due to excessive wetness during the growing season (Reed 1988; Mitsch

& Gosselink 2000; Tiner 1991, 2006). Roughly one third of the nation's vascular plants have some potential

for being hydrophytes (Tiner 2006).

Two editions of the National List of Plant Species that Occur in Wetlands (National List) are currently

published and available for use, an approved list compiled by Reed (1988) and a revised draft list prepared by

the USFWS (1996); hereafter referred to as the 1988 list and the 1996 draft list (respectively). The National

List was generated from 13 regional lists, including a list for California, which is Region 0 (Reed 1988; Tiner

2006). A given plant species may be a better indicator of wetlands in one region than it is in another; the

regional wetland plant lists have been used to help identify wetland communities that can be recognized

entirely on their vegetation components (Tiner 1991, 2006). Accordingly, wetland indicator categories of

individual plant species on the regional lists have been created in order to further assist biologists with the

delineation of wetland and riparian habitats throughout the United States (Tiner 1993, 2006; Mitsch & Gos-

selink 2000; USACE 2006). Each native or non-native species on these lists is assigned an indicator status

reflecting its frequency of occurrence (not degree of wetness) in wetlands (Reed 1988), which is described in

Table 1.

METHODOLOGY

Voucher specimens and other data were collected during wetland delineations, focused botanical surveys,

and casual floristic surveys of wetland and riparian habitats associated with urbanized watersheds in south-

ern California. Primary areas of study included the coastal bays and estuaries of Los Angeles, Orange, and

San Diego counties, Arroyo Seco Creek and the San Gabriel River drainage in Los Angeles County, Aliso

Creek, San Diego Creek, San Juan Creek, and the Santa Ana River drainage in Orange County, and the Lake

Elsinore basin and San Jacinto River-Salt Creek drainage in western Riverside County.

Our approach to identifying previously unreported hydrophytes follows the “individualistic concept" of

a hydrophyte defined by Tiner (1988). The individualistic concept recognizes that plant species may exhibit

considerable plasticity or ecological amplitude in their adaptations to wet environments, which may represent

the entire population of a species or only a subset of individuals (Tiner 1988, 1991). The potential adaptation

and behavior of a species that functions as a hydrophyte in new environments is particularly important in

coastal southern California where new sources of urban water has effected the spread and establishment of

several exotic plant introductions that would not likely occur in this otherwise arid environment.

Field experiences in southern California, a review of the literature, a review of habitats and species data

available from local herbaria (RSA-POM, UCR) and online from the Consortium of California Herbaria (2006),

and well-known wetland species with which it is commonly associated were used to assign an indicator

status for each of the unreported hydrophytes evaluated for addition to the National List. Identification of

associated hydrophytic vegetation, wetland hydrology, and/or hydric soils of the plant community in which

the suspected hydrophyte occurs follows the methodology described in the USACE Wetland Delineation

Manual and the Arid West Regional Supplement (USACE 1987, 2006).

In addition, evaluation of landform position and plant community type in which the suspected hy-

drophyte is typically found (i.e., streamside riparian scrub), widespread wetland types (i.e., coastal bay salt

marsh), aquatic sites (i.e., shallow ponds and slow-moving streams), location of its microhabitat community

along moisture gradients (i.e., depressions in vernal alkali flats), man-made or man-modified habitats that

Riefner and Boyd, Wetland and riparian plants in southern California 723

TaBLe 1. Wetland Indicator Status for the National List of Wetland Plant Species that Occur in Wetlands.

Definitions for Wetland Indicator Status

Indicator Category Estimated Probability of Occurrence in Wetlands

Obligate Wetland (osL) Plants that almost always occur in wetlands; >99% of the time.

Facultative Wetland (FAcw) Plants that usually occur in wetlands; 67-99% of the time.

Facultative (FAC) Plants that are just as likely to occur in wetland and in non-wetland areas;

34-66% of the time.

Facultative Upland (Facu) Plants that occasionally occur in wetlands; 1-33% of the time.

Upland (uet) Plants that almost always occur in uplands; 99% of the time.

No Indicator (Ni) Assigned to plants that lack sufficient information to base an indicator status.

No Agreement (NA) Assigned where reviewers could not achieve a unanimous agreement for an indicator.

Asterisk (*) Assigned to indicators derived with limited ecological information, which indicates a

tentative assignment that requires further review.

In addition, a plus (+) or minus (-) designation, respectively, indicates the higher or lower range for a particular indicator.

exhibit minimal hydric characteristics (i.e., ditches with seasonal flowing water or seasonal saturation that

support a predominance of facultative weedy species), and/or easily recognized morphological adaptations to

wet areas (e.g., shallow root systems developed on or near the soil surface) were observed during this study

and further indicate these plants are functioning as hydrophytes in southern California.

NEW RECORDS OF WETLAND AND RIPARIAN PLANTS

Voucher specimens, generalized distribution, habitat association, taxonomic notes (where appropriate), a

literature review, and the wetland indicator status for California are cited for each taxon discussed in this pa-

per. References to biogeographic provinces follow Hickman (1993). Common names provided in the species

discussions follow the treatments of generic experts, or if not available, we use the common name provided by

Roberts (1998), DiTomaso & Healy (2003), Roberts et al. 2004), and Rebman & Simpson (2006). Facultative

plants proposed for addition to the California regional list, and revisions and/or recommendations to the

wetland indicator status for other species currently on the list are provided. Where appropriate, additional

representative voucher specimens with notes on habitat and associated facultative plants are also provided

to further establish the facultative behavior of each taxon discussed, which follows:

Amaranthus blitum L. subsp. emarginatus (Uline & Bray) Carretero (Amaranthaceae)

CALIFORNIA. Los Angeles Co.: City of Pico Rivera, San Gabriel River, Thienens Rd. near confluence with San Jose Creek, UTM (NAD

83) 11S 0405107E 3766943N, elev. 74 m (243 ft), uncommon on river banks and in wet sand, 28 Aug 2004, Riefner 04-406 (RSA). Or-

ange Co.: City of Anaheim, Santa Ana River bottom near 57-Freeway between Orangewood St. and Chapman Ave., UTM (NAD 83) 11S

0418311E 3739098N, elev. 41 m (135 ft), locally abundant in disturbed wetlands and on sandbars, 8 Aug 2004, Riefner 04-370 (RSA);

City of Yorba Linda, N floodplain of Santa Ana River near Yorba Linda Regional Park, UTM (NAD 83) 11S 0430355E 3748269N, elev. 98

m (321 ft), common in wet river wash sand, 25 Sep 2004, Riefner 04-442 (RSA); City of Huntington Beach, Huntington Central Park at

Goldenwest St., UTM (NAD 83) 11S 0406898E 3730128N, elev. 1.5 m (5 ft), locally common and highly invasive in wet ditches, drying

streambeds, and margins of riparian woodlands, 9 Jul 2005, Riefner 05-546 (RSA); City of Irvine, San Diego Creek at Alton Pkwy., UTM

(NAD 83) 11S 0429107E 3724401N, elev. 45 m (148 ft), locally common in wet sand along slow-moving waters, 30 Sep 2006, Riefner

06-458 (RSA).

Previous knowledge.—Amaranthus blitum L. (purple amaranth, livid amaranth), of tropical origin, is well

established in many disturbed habitats in the eastern United States and Canada (Mosyakin & Robertson 2003).

The distribution of the infraspecific taxa of the A. blitum complex is poorly known in North America however, and

requires additional study (Mosyakin & Robertson 2003). Amaranthus blitum subsp. emarginatus was not treated

in The Jepson Manual (Henrickson 1993). It was first reported in California from waste ground and potted plant

containers in Los Angeles and Riverside counties (Hrusa et al. 2002). Additional localities for A. blitum subsp.

724 Journal of the Botanical R h Institute of Texas 1(1)

emarginatus, including populations from cultivated ground, have been reported for western Riverside County,

but not from Orange County (Roberts 1998; Roberts et al. 2004). Amaranthus blitum (infraspecific taxon not

cited) has been reported from a yard in San Diego County (Consortium of California Herbaria 2006; Rebman

& Simpson 2006).

Wetland Indicator Status —Amaranthus blitum subsp. emarginatus was not assigned a wetland indicator status

for any region on the 1988 list or the 1996 draft list, but A. blitum has been assigned a FAC wetland indicator

status for Hawaii on the 1996 draft list.

Significance and Recommended Indicator Status.—First report of A. blitum subsp. emarginatus documented

for Orange County, and second report for Los Angeles County. Adapted to tropical and subtropical climates, A.

blitum subsp. emarginatus was expected to be only a minor weed of greenhouse or garden situations in California

(Hrusa et al. 2002). However, it has naturalized and is spreading from yards and fields to disturbed vernally moist

soils, seasonal wetlands, and riparian habitats in the mild Mediterranean climate of coastal southern California.

In Huntington Beach, this taxon is highly invasive in disturbed riparian scrub and wetland habitats. These new

habitat records indicate A. blitum subsp. emarginatus is undergoing a range expansion and will likely colonize

numerous disturbed wetlands and other moist sites associated with urbanized watershed habitats.

Based on field observations, and information available in the literature and the Consortium of California

Herbaria (2006), A. blitum subsp. emarginatus should be added to the California regional list. We propose a

FAC* wetland indicator status. Additional regional review is needed to specifically define its frequency of oc-

currence in wetlands as the species undergoes further expansions of range and invasion of new habitats.

Atriplex polycarpa (Torr.) S. Watson (Chenopodiaceae)

CALIFORNIA. Riverside Co.: City of Lake Elsinore, back basin of Lake Elsinore, W ca. 0.5 mi off Pete Lehr Dr. at Diamond Stadium

in Wildlife Viewing Area, UTM (NAD83) 11S 0471224E 3723561N, elev. 402 m (1320 ft), locally common in Distichlis grassland, 7 Jul

2004, Riefner 04-306 (RSA, UCR).

Previous knowledge.—Atriplex polycarpa (desert saltbush) inhabits fine-textured saline soils of warm desert

shrub and saltgrass communities from 60-1500 m elevation in Arizona, California, Nevada, Utah, and northern

Mexico (Welsh 2003). In California, it occupies alkaline flats and dry lakes in the San Joaquin Valley, Transverse

and Peninsular Ranges, and the region lying east of the Sierra Nevada south through the California deserts

(Taylor & Wilken 1993). It has not been recorded from Orange County or western Riverside County (Roberts

1998; Roberts et al. 2004; Consortium of California Herbaria 2006). With the exception of the alkali plains of

the Hemet and San Jacinto River Valleys, and the coastal salt marshes, little attention has been paid to the study

of alkaline soil plant communities in southern California (Riefner & Boyd 20053).

Wetland Indicator Status.—Atriplex polycarpa has been assigned a FACU wetland indicator status for California

on the 1988 list and the 1996 draft list.

Ç: ifi

County. This is one of a suite of seasonal wetland species, including Eleocharis obtusa (Willd.) Schultes var.

led Indicator Status.—First report of A. polycarpa documented for western Riverside

engelmannii (Steud.) Gilly and Psilocarphus chilensis (P. tenellus Nutt. var. globiferus [DC.] Morefield), that is

more typical of California’s Central Valley (Boyd & Ross 1996; Riefner et al. 2002). No changes to the wetland

indicator status are proposed at this time.

Bacopa monnieri (L.) Wettst. (Scr ophular iaceae)

CALIFORNIA. Orange Co.: City of Yorba Linda, Santa Ana River bottom near Yorba Linda Regional Park, UTM (NAD 83) 11S

0429795E 3747941N, elev. 96 m (315 ft), locally common in wet river wash sand, 3 Sep 2004, Riefner 04-419 (RSA)

Previous knowledge.—Bacopa monnierii (Monnier water hyssop), native to tropical and subtropical regions

nearly worldwide, grows in wet soil or in shallow water in the Sonoran Desert (eastern Riverside County) to the

southern United States (Strother 1993; DiTomaso & Healy 2003). It is apparently a recent alien, just becoming

naturalized in our area, including San Diego County (Strother 1993; Consortium of California Herbaria 2006;

Rebman & Simpson 2006). Bacopa monnierii, however, has not been recorded from western Riverside County

or Orange County (Roberts 1998; Roberts et al. 2004).

Wetland Indicator Status —Bacopa monnierii was not assigned a wetland indicator status for California on

the 1988 list, but has been assigned an OBL wetland indicator status on the 1996 draft list.

Riefner and Boyd, Wetland and riparian plants in southern California 725

Significance and Recommended Indicator Status.—First record of B. monnierii documented for Orange County.

Bacopa monnierii is one of many subtropical and tropical weeds that are adapting well to disturbed urban

wetlands in the South Coast region. We concur with the revised OBL indicator status assigned on the 1996

draft list.

Beta vulgaris L. subsp. maritima (L.) Arcangeli (Chenopodiaceae)

CALIFORNIA. Los Angeles Co.: Torrance, Victoria Park, Dominguez Channel, vicinity of Del Amo Blvd. and Carson Plaza Dr.,

UTM (NAD 83) 11S 0382502E 3746087N, elev. m (28 ft), uncommon, growing with Distichlis and Atriplex lentiformis in seasonally

saturated alkali grasslands, 29 Jul 2006, Riefner 06-329 (RSA). Riverside Co.: City of Wildomar, near Lake Elsinore, N ca. 0.25 mi from

the intersection of Union St. and Corydon Rd., UTM (NAD 83) 11S 0471584E 3720254N, elev. 367 m nee ft), uncommon, growing

with Distichlis, Plantago elongata, Nitrophila occidentalis, and C

in alkali meadows, 22 Apr 1998, Riefner 98-266 (RSA).

Previous knowledge —Beta vulgaris subsp. maritima (sea beet), native to southern Europe, inhabits waste areas

1 soils of depressions

E (e)

and moist, sandy places near the coast in New Jersey and southern California (Shultz 2003). In California, it has

been collected from low-lying and upland habitats in Los Angeles, San Diego, and Santa Barbara counties, but

not Orange or Riverside counties (Roberts 1998; Roberts et al. 2004; Consortium of California Herbaria 2006;

Rebman & Simpson 2006).

Wetland Indicator Status.—Beta vulgaris subsp. maritima was not assigned a wetland indicator status for

California or any other region on the 1988 list or the 1996 draft list.

Significance and Recommended Indicator Status. —First report of B. vulgaris subsp. maritima documented for

Riverside County. Based on field observations, and information available in the literature and the California

Consortium of Herbaria (2006), B. vulgaris subsp. maritima should be added to the California regional list. We

propose a FAC* wetland indicator status and recommend additional regional review to specifically define

its frequency of occurrence in wetlands as the species undergoes further expansions of range in southern

California. Beta vulgaris subsp. vulgaris (cultivated beet) is sporadic in waste areas, roadsides, and fields (Shultz

2003); it has been assigned a FACU wetland indicator status on the 1988 list and on the 1996 draft list.

Chloris truncata R. Br. (Poaceae)

CALIFORNIA. Riverside Co.: Menifee Valley, along Newport Rd. 1 mi E of Goetz Rd. intersection, UTM (NAD 83) 0479208E,

3727278N, elev. 442 m (1450 ft), locally common in wet ditch with Chloris virgata, 21 Oct 2003, Riefner 03-461 (RSA); Perris Valley,

Rider Street near Perris Valley Storm Drain, UTM (NAD 83) 11S 0480399E 3742338N, elev. 483m (1585 ft), common in irrigated alfalfa

fields, in disturbed seasonal wetlands with Polypogon and Cyperus, and disturbed margin of Salix scrub, 27 May 2003, Riefner 03-254

(RSA); Homeland, State Hwy. 74 near 1* St., UTM (NAD 83) 11S 0483321E 3734325N, elev. 439m (1439 ft), locally common in roadside

swale with Cynodon dactylon, Cyp alternifolius, and Echinochloa colona, 3 Aug 2006, Riefner 06-354 (RSA). San Bernardino Co.: City

of Chino, along Edison Ave. at veo Ave., UTM (NAD 83) 11S 0437972E 3762021N, elev. 230 m (755 ft), uncommon, agricultural

ditch with Cyperus and Eleusine in fallow farmland, 31 Jul 2006, Riefner 06-343 (RSA).

Chloris truncata (black wind-mill grass), native to Australia, is known as a weed of alfalfa fields, orchards,

irrigated turf grass nurseries, and roadsides in Imperial, Merced, and Riverside counties, but not from Los An-

geles, Orange or San Bernardino counties (Hrusa et al. 2002; Barkworth 2003; Roberts et al. 2004; Consortium

of California Herbaria 2006; Rebman & Simpson 2006). It has also been collected in South Carolina (Barkworth

2003).

Wetland Indicator Status —Chloris truncata was not assigned a wetland indicator status for California or any

other region on the 1988 list or the 1996 draft list.

Significance and Recommended Indicator Status.—First report of C. truncata documented for San Bernardino

County. Chloris truncata is spreading from orchards and fields to disturbed vernally moist soils, seasonal wet-

lands, and riparian habitats. Based on field observations, information available in the literature, and review of

the Consortium of California Herbaria (2006), C. truncata should be added to the California regional list. We

propose a NI* wetland indicator status and recommend additional regional review to specifically define its

frequency of occurrence in wetlands as the species undergoes further expansions of range and invasion of

new habitats.

Ehrharta erecta Lam. (Poaceae)

CALIFORNIA. Imperial Co.: El Centro, Imperial Ave. at Ocotillo Rd., UTM (NAD 83) 11S 0633931E 3628013N, elev. -9 m (31

£s+haD o ID L

726 Journal of t titute of Texas 1(1)

ft), few plants growing in a moist gutter and in an irrigated commercial landscape, 9 Oct 2006, Riefner 06-488 (RSA). Los Angeles

Co.: Verdugo Mountains near Sunland, La Tuna Canyon, N of La Tuna Canyon Rd. along La Tuna Creek, S of 210-Fwy. and ca. 1.5 mi

W of 210-Fwy. intersection with La Tuna Canyon Rd. exit, UTM (NAD 27) 0379653E 3788712N, elev. 411 m (1350 ft), common, grow-

ing with Cyperus eragrostis in damp sand along stream banks and with Carex spissa in shaded willow-oak riparian woodlands, 29 Aug

2000, Riefner 00-764 (RSA); City of Pasadena, Arroyo Seco Creek, S of Arroyo Blvd., UTM (NAD 83) 11S 0392637E 3779048N, elev. 226

m (740 ft), common in wet sand along creek in oak-willow riparian woodland, 22 Aug 2004, Riefner 04-381 (RSA); City of Pasadena, San

Rafael Hills, canyon in vicinity of Candeo Pl. and Wierfield Dr., UTM (NAD 83) 11S 0391056E 3779872N, elev. 320 m (1049 ft), com-

mon in oak riparian woodland, growing with Cyperus on damp stream banks, 29 Apr 2006, Riefner 06-159 (RSA). Orange Co.: City of

Seal Beach, San Gabriel River at River's End Café, First St. and Ocean Blvd., UTM (NAD 83) 11S 0396779E 3734148N, elev. 3 m (10 ft),

common in irrigated landscapes, 23 May 1994, Riefner 94-387 (RSA); City of Buena Park, Lincoln Ave. at Magnolia Ave., UTM (NAD

83) 11S 0409756E 3743996N, elev. 36 m (117 ft), common in roadside gutter, 18 Jun 1994, Riefner 94-441 (RSA); City of Laguna Beach,

N of Laguna Beach Country Club along Aliso Creek, UTM (NAD 83) 11S 0431343E 3708831N, elev. 20 m (65 ft), locally common on

stream banks with Salix, 4 Mar 2002, Riefner 02-103 (RSA); City of Lake Forest, Serrano Creek, SE of Bake Pkwy., UTM (NAD 83) 11S

0434618E 3723575N, elev. 107 m (352 feet), locally abundant in mule fat scrub along margins of ephemeral streambed, 31 May 2004,

Riefner 04-194 (RSA, UCR); City of Rancho Santa Margarita, Los Flores Rd. near Santa Margarita Pkwy., UTM (NAD 83) 11S 0444330E

3722423N, elev. 292 m (958 ft), common in irrigated landscape, 5 Jul 2004, Riefner 04-286 (RSA, UCR); City of Aliso Viejo, Aliso Creek

near Entidad Rd. at Los Alisos Blvd., UTM (NAD 83) 11S 0439164E 3723107N, elev. 220 m (721 ft), common, growing with Cyperus on

shaded stream banks in oak riparian woodland, 5 Oct 2005, Riefner 05-709 (RSA). Ventura Co.: City of Simi Valley, Santa Susana Park

UTM (NAD 83) 11S 0347007E 3792286N, elev. 331 m (1087 ft), common on stream bank with Epilobium ciliatum in oak-willow riparian

woodland, 9 Sep 2005, Riefner 05-656 (RSA).

Previous knowledge.—Populations of Ehrharta erecta (panic veldt grass), a native of South Africa, have been

reported from the San Francisco Bay Area, Santa Barbara, Ventura, Los Angeles, western Riverside, and San

Diego counties (Bossard et al. 2000; Roberts et al. 2004; Rebman & Simpson 2006). Although it is a common

urban weed in southern California, it has not been reported from Orange County (Roberts 1998; Bossard et al.

2000). However, it has been recently collected from roadside habitats in Orange and San Bernardino counties

(Consortium of California Herbaria 2006). Ehrharta erecta infests wildlands in most or all of the North and

Central Coast, but there are relatively few reports of it in southern California native habitats (Sigg 1996, 2003;

Roberts et al. 2004). Its preference for moist environments suggests it might become a threat to wetlands and

riparian habitats (Sigg 1996, 2003). Ehrharta erecta is also recognized as a species with the potential to spread

explosively in California (CalEPPC 1999).

Wetland Indicator Status.—Although E. erecta inhabits moist environments in urban and wildland situations,

it has not been assigned a wetland indicator status for California or any other region on the 1988 list or the

1996 draft list.

Significance and Recommended Indicator Status.—First report of E. erecta documented for Imperial County,

where it has likely been introduced with landscape plantings for urban development projects; verification of a

herbarium record for Orange County. Additional records of F. erecta are documented from native riparian and

stream course habitats in Los Angeles, Orange, and Ventura counties. In Orange County, E. erecta is vigorously

expanding its range from the coast into the foothills of the Santa Ana Mountains in many types of urban envi-

ronments owing to landscape gardening practices associated with expanding urbanization. Based on field

observations, information available in the literature, and review of the Consortium of California Herbaria (2006),

E. erecta should be added to the California regional list and assigned a FAC wetland indicator status.

Elytrigia repens (L.) Nevski (Poaceae)

CALIFORNIA. Orange Co.: City of Costa Mesa, South Coast Dr. at Susan Rd., UTM (NAD 83) 115 0415261E 3728412N, elev. 16

m (52 ft), swale and margin of mule fat scrub, 1 Oct 2005, Riefner 05-704 (RSA).

Previous knowledge.—Elytrigia repens (Agropyron repens [L.] Beauv.; quackgrass), native to Eurasia, is

a weed of cultivated ground and other disturbed sites throughout California (except the deserts and high

elevation montane habitats) to the eastern United States (Jarvie & Barkworth 1993). However, it has not

been reported from Orange, Riverside, or San Diego counties (Roberts 1998; Roberts et al. 2004; Rebman

& Simpson 2006). Most records from the South Coast region are from Santa Barbara County (Consortium

of California Herbaria 2006).

Wetland Indicator Status.—Elytrigia repens has been assigned a NI designation for California on the 1988

list (as Agropyron repens) and a FAC* wetland indicator status on the 1996 draft list.

VAl asl J | mee. if I T +h

Riefner and Boyd, California 727

Significance and Recommended Indicator Status. —First report of E. repens documented from Orange County.

We concur with the revised FAC* indicator status as the species requires further review as it spreads in the

South Coast region.

ipic brachycarpum C. Presl (Onagraceae)

IFORNIA. Butte Co.: along flood control slough, ca. 4 mi N of Chico on Hwy 99, 8 Sep 1978, Taylor 1894 (CHSC). Humboldt

Co.: Pur Coast region, near Hydesville, gravel bar on Yager Creek, 9 Sep 1900, Tracy 946 (UC). Lake Co.: below the high water line

of Indian Valley Reservoir, W of the causeway in the NW part of the reservoir, W side of Bartlett Springs Rd., 4 Oct 2004, Ahart 11446

(CHSC); Mendocino National Forest, Stonyford Quadrangle, sag pond 1 mi N of Sheep Corral, 19 Sep 1936, Schreiber 2334 (UC). Los

Angeles Co.: Transverse Ranges, Liebre Mountains, Cow Spring Pond, sag pond on the San Andreas Fault at the N foot of Liebre Mtn.

on the S side of West Oakdale Canyon Rd., ca 100 m W of the Cow Spring Canyon drainage, 34?44'12"N 118?38'45"W, 21 Sep 1994,

Ross & Boyd 8327 (RSA); Liebre Mountains, SW end of Quail Lake, seasonally flooded depression on S side of Hwy 138, 34?46'13.6"N

118?45'16.5"W, 14 Oct 1996, Boyd & Raz 9086 (RSA). Mendocino Co.: Northern California Coast Range Preserve, Elder Creek, margins

of creek about 0.5 to 1 mi upstream above confluence with South Fork Eel River, 23 Jun 1961, Sharsmith 4905 (UC). Mono Co.: Leavitt

Meadows, West Walker River, gravel bench along river, 1 Sep 1944, Alexander & Kellogg 4174 (UC). Orange Co.: City of Yorba Linda,

Santa Ana River, Horseshoe Bend in Santa Ana Canyon, UTM (NAD 83) 11S 0431466E 3749276N, elev. 99 m (323 ft), floodplain in Bac-

charis salicifolia scrub, 5 Oct 2005, Riefner 05-705 (RSA). Plumas Co.: W side of Little Last Chance Creek, strand of Frenchman Lake, 29

Aug 2001, Ahart 9254 (CHSC). Riverside Co.: Peninsular Ranges, Perris-Aguanga Basin region, floodplain E of dike along San Jacinto River,

W of Sedco Hills community, 29 Sep 1994, Boyd & Banks 837 (RSA); City of Hemet, vicinity of Warren Rd. at Florida Ave., UTM (NAD 83) 11S

0496489E 3724636N, 460 m (1510 ft), scattered in vernal alkali grassland with Hordeum intercedens, Cressa truxillensis, Veronica peregrina

a occidentalis, 31 Jul 2004, Riefner 04-347 (RSA); SE of Temecula and

E of Pechanga Indian Reservation, Cleveland National Forest, Agua Tibia Wilderness, N slope of Agua Tibia Mountain, ca. 1.5 mi WNW of

Woodchuck Rd., UTM (NAD 83) 11S 0497415E 3699899N, 665 m (2180 ft), scattered along bank and bed of unnamed blueline stream with

Baccharis salicifolia, 31 Jul 2005, Riefner 05-57 (RSA); San Jacinto Mountains, McCall Park, McCall Park Rd. near Hwy 74, UTM (NAD 83)

11S 0524882E 3729641N, 1360 m (4462 ft), growing with Muhlenbergia rigens along ephemeral stream, 7 Aug 2005, Riefner 05-591 (RSA);

San Jacinto Mountains, near McMullen Flat on SR 243, Valley Hi County Park, UTM (NAD 83) 11S 0516223E 3747936N, 1139 m (3737 ft),

common, growing with Crypsis and Amaranthus californicus along edge of drying pond and ephemeral creek, 12 uns i Riefner & Sanders

subsp. xalapensis, Centromadia pungens subsp. laevis, and Chamomi

05-601 (RSA). San Bernardino Co.: Transverse Ranges, San Bernardino Mountains region, NW shore, alkali g Baldwin Lake, s.d

Thorne & Wisura 53689 (RSA). San Diego Co.: bank of stream 12 mi E on hwy to Santa Ysabel- Ramona, 29 Aug 1927, Wiggins 2607 (UC);

Agua Tibia Mtns., Rainbow Heights Rd. East, ca. 0.7 mi N from Rainbow Heights Rd. West, UTM (NAD 83) 115 0490270E 3698344N,

elev. 491 m (1611 ft), perennial seep with Juncus and Cyperus, 7 May 2006, Riefner 06-182 (RSA). Trinity Co.: Van Duzen River, gravel bar,

river 3 mi above Low Gap, 4 Sep 1939, Tracy 16483 (UC). Yuba Co.: along a small stream, below the high water line of Sly Creek Reservoir,

about 1/8 mi SW of Day Camp, Sly Creek Reservoir, 9 Oct 2002, Ahart 9956 (CHSC).

Previous knowledge.—Epilobium brachycarpum (E. paniculatum Torr. & A. Gray; summer cotton weed) is

common in dry open woodlands, grasslands, and roadsides below 3300 m elevation in the California Flo-

ristic Province (except the Channel Islands) and the Modoc Plateau north to British Columbia, and east to

South Dakota, New Mexico, and eastern Canada (Hoch 1993). In the South Coast region, E. brachycarpum

has been documented from Los Angeles, Riverside, and San Diego counties, but not from Orange County

(Roberts 1998; Roberts et al. 2004; Consortium of California Herbaria 2006; Rebman & Simpson 2006).

Wetland Indicator Status.—Epilobium brachycarpum was not assigned a wetland indicator status on the

1988 list, but was included on the 1996 draft list with a UPL designation.

Significance and Recommended Indicator Status.—First report of E. brachycarpum documented from Orange

County. In California, E. brachycarpum is found on open, usually dry disturbed ground in many plant com-

munities (Munz & Keck 1959; Munz 1974; Hoch 1993). However, recent field work, review of local floras,

and a search of local herbaria records indicates E. brachycarpum frequently inhabits floodplains, drying pond

and lake margins, vernal alkali plains, gravel bars, banks and drying beds of ephemeral and intermittent

streams, sloughs, ditches, sag ponds, seasonally flooded depressions, and perennial seeps (Yoder 1996; Banks

1999; Boyd 1999; Roberts et al. 2004; Consortium of California Herbaria 2006). Based on field observations,

and information available in the literature and the Consortium of California Herbaria (2006), the status of

E. brachycarpum should be revised and assigned a FACU wetland indicator status for California.

Eriochloa aristata Vasey (Poaceae)

CALIFORNIA. Riverside Co.: City of Hemet, Florida Ave. near Grant Ave., UTM (NAD 83) 0511109E 3734246N, elev. 547 m

(1796 ft), locally common in wet ditch and roadside swales, 31 Jul 2004, Riefner 04-352 (CS, RSA).

f4L,D o ID L

728 Journal of t titute of Texas 1(1)

Previous knowledge. —Eriochloa aristata (bearded cupgrass) occupies seasonal streams and riverbanks in

the San Francisco Bay region, Riverside, and Imperial counties in the Sonora Desert region of California, and in

southern Arizona south through Mexico and Central America to Colombia (Munz 1974; Webster 1993a; Shaw

et al. 2003). Eriochloa aristata has not, however, been reported from Orange, western Riverside, and San Diego

counties (Roberts 1998; Roberts et al. 2004; Consortium of California Herbaria 2006; Rebman & Simpson

2006).

Wetland Indicator Status.—Eriochloa aristata has been assigned a FACW wetland indicator status on the 1988

list and the 1996 draft list.

Çi Wf AD

led Indicator Status. —First record of E. aristata documented from western Riverside

On’.

County. Rarely collected in southern California (Webster 1993a), E. aristata is expected to expand its range and

occupy moist roadside swale habitats in other inland valleys of western Riverside County and in eastern San

Diego County. No changes to the wetland indicator status are proposed at this time.

Glinus radiatus (Ruiz Lopez & Pavon) Rohrb. (Molluginaceae)

CALIFORNIA. Orange Co.: City of Lake Forest, Upper Oso Reservoir, UTM (NAD 83) 11S 0441970E 3724638N, elev. 293 m (960 ft),

drying margin of fl ing lake, 24 Sep 2005, Riefner 05-688 (RSA). Riverside Co.: City of Murrieta, ca. 1.5 mi E of I-15 Fwy. on Clinton

Keith Rd., vicinity of Smith Ranch Rd., UTM (NAD 83) 11S 0479562E 3717088N, elev. ca. 427 m (1400 ft), uncommon in drying stock

pond, 26 Sep 2003, Riefner 03-379 (RSA, UCR).

Previous knowledge.—Glinus radiatus (radiate sweetjuice), native to tropical America, is known in California

from the Sacramento Valley and the Peninsular Range Province (Ferren 1993). Glinus radiatus is local, but well

documented from drainage courses, drying vernal pools and ponds, and receding shores of reservoirs and

lakes in San Diego County (Beauchamp 1986; Consortium of California Herbaria 2006). It has not however,

been reported from Orange or Riverside counties (Roberts 1998; Roberts et al. 2004; Consortium of California

Herbaria 2006).

Wetland Indicator Status.—Glinus radiatus was not assigned a wetland indicator status on the 1988 list, but

was included on the 1996 draft list with a NI designation.

Significance and Recommended Indicator Status.—First records of G. radiatus documented for Orange and

Riverside counties. Glinus radiatus is easily confused with G. lotoides L., and it is likely more widespread than

herbarium records indicate. Glinus radiatus is expected to occur at other large seasonal pond and drying lakeshore

habitats in southern California. Based on field observations, and information available in the literature and the

Consortium of California Herbaria (2006), the status of G. radiatus should be revised and assigned an OBL

wetland indicator status for California. Glinus lotoides, which also occupies moist or seasonally dry margins of

wetlands, has been assigned an OBL wetland indicator status on the 1988 and 1996 draft lists.

Limonium ramosissimum (Poir.) Maire (Plumbaginaceae)

CALIFORNIA. Los Angeles Co.: City of Long Beach, Belmont Shores, Colorado Lagoon in vicinity of Park Ave. and Colorado St.,

UTM (NAD 83) 11S 0394757E 3737423N, elev. 6 m (20 ft), uncommon in disturbed salt marsh, 21 Jan 2006, Riefner 06-8 (RSA, UC).

Orange Co.: City of Lake Forest, Aliso Creek near Portola Pkwy. at El Torro Rd., UTM (NAD 83) 11S 0439205E 3724613N, elev. 225 m

(739 ft), drying streambed in alkaline soils, 9 Sep 2005, Riefner 05-654 (RSA, UC); City of Newport Beach, E of Newport Bay, E of Bayside

Dr. at Big Canyon Creek drainage, UTM (NAD 83) 11S 0418191E 3721588N, elev. 6 m (20 ft), common in salt marsh, 16 Sep 2005, Riefner

05-669 (RSA); City of Irvine, Bonita Creek, along La Salud St. near Milano Dr., UTM (NAD 83) 11S 0420070E 3723153N, elev. 9 m Q9

ft), common along bike trail, disturbed scrub, and on roadsides, 5 Oct 2005, Riefner 05-707 (RSA, UC); City of Newport Beach, Newport

Bay, Bay View Dr. E of Jamboree Rd., UTM (NAD 83) 11S 0419864E 3723987N, elev. 6 m (20 ft), uncommon, edge of salt marsh, 23 Nov

2005, Riefner 05-772 (RSA, UC); City of Newport Beach, San Diego Creek near confluence with Newport Bay near SR 73 overpass, UTM

(NAD 83) 11S 0419867E 3723886N, elev. 6 m (20 ft), common, edge of disturbed riparian woodland with Saliconia, Euthamia occidentalis,

and Pulicaria paludosa, 12 Nov 2006, Riefner 06-670 (RSA). San Diego Co.: San Onofre State Beach, near Echo Arch Campground, UTM

(NAD 83) 11S 0449452E 3691583N, elev. 7 m (22 ft), uncommon in wet sand along edge of Distichlis seep, 31 Aug 2004, Riefner 04-409

(RSA, UC); City of Carlsbad, N side of San Marcos Creek at Carlsbad Blvd., 0.2 mi S of Avenida Encinas, UTM (NAD 83) 11S 0470849E

3661229N, elev. 10 m (32 ft), uncommon on edge of mule fat-willow scrub, 7 Jan 2005, Riefner 05-3 (RSA, UC).

Previous knowledge.—Limonium ramosissimum (Poir.) Maire (sea-lavender) is a European species not treated

in The Jepson Manual, but recently reported from the Carpenteria salt marsh in Santa Barbara County as the

subsp. provinciale (Pignatti) Pignatti, where it is abundantly naturalized (McClintock 1993; Hrusa et al. 2002;

Smith 2005). Its identification is somewhat problematic, and due to the complexity of the genus, it is prema-

Riefner and Boyd, Wetland and riparian plants in southern California 729

ture to assign a subspecies to this taxon without detailed comparison with European material (Smith 2005).

Limonium ramosissimum has not been reported from Los Angeles, Orange, or San Diego counties (Roberts 1998;

DiTomaso & Healy 2003; Consortium of California Herbaria 2006; Rebman & Simpson 2006). It has been

cited by Roberts et al. (2004) from moist, low-lying alkaline habitats in western Riverside County, but this speci-

men (Lake Elsinore, Riefner 03-241, UCR) is referable to Limonium indet., which is discussed below. Limonium

ramosissimum and Limonium indet. are superficially similar, often co-occur in disturbed wetland or moist ruderal

habitats, are frequently used in the nursery trade, and are therefore easily confused.

Wetland Indicator Status.—Limonium ramosissimum has not been assigned a wetland indicator status for

California or any other region on the 1988 list or the 1996 revised list.

Significance and Recommended Indicator Status.—First annotated reports of L. ramosissimum documented

for Los Angeles, Orange, Riverside, and San Diego counties. Based on field observations and information avail-

able in the literature, L. ramosissimum should be added to the California regional list and assigned a FACW

wetland indicator status. Limonium ramosissimum may be a threat to the endangered Cordylanthus maritimus

Nutt. in salt marsh habitats at Carpenteria, Santa Barbara County (Smith 2005). Limonium ramosissimum is

spreading rapidly, especially in salt marsh habitats, where it may also threaten sensitive species in Orange

and San Diego counties. The California Invasive Plant Council should monitor it closely.

Limonium indet. (Plumbaginaceae)

CALIFORNIA. Orange Co.: City of Newport Beach, E of Newport Bay and E of Bayside Dr., Big Canyon Creek drainage, UTM (NAD

83) 11S 0418191E 3721588N, elev. 6 m (20 ft), common in salt marsh, 16 Sep 2005, Riefner 05-667 (RSA, UC); City of Irvine, Bonita Creek,

along La Salud St. near Milano Dr., UTM (NAD 83) 11S 0420070E 3723153N, elev. 9 m (29 ft), common along bike trail, disturbed scrub,

and on roadsides, 5 Oct 2005, Riefner 05-708 (RSA); City of Rancho Santa Margarita, along Robinson Ranch Rd. near North Peak Rd., UTM

(NAD 83) 11S 0448529E 3724126N, elev. 500 m (1639 ft), common on disturbed roadside slopes, in ditches, and banks of ephemeral

streambed, 4 Nov 2005, Riefner 05-739 (RSA); City of Newport Beach, Upper Newport Bay, Bay View Dr. E of Jamboree Rd. at San Diego

Creek, UTM (NAD 83) 11S 0419864E 3723987N, elev. 6 m (20 ft), uncommon, edge of salt marsh, 23 Nov 2005, Riefner 05-774 (RSA),

City of Newport Beach, E of Newport Bay, E of Back Bay Dr., flats along Big Canyon Creek, UTM (NAD 83) 11S 0418069E 3721609N, 6

m (18 ft), salt flats and scrub with Salicornia and Atriplex glauca, 11 Aug 2006, Riefner 06-386 (RSA, UC); City of Newport Beach, Upper

Newport Bay E of Jamboree Rd., UTM (NAD 83) 11S 0419547E 3723812N, elev. 6 m (20 ft), common, salt marsh with Salicornia, 13

Aug 2006, Riefner 06-391 (RSA, UC); City of Newport Beach, San Diego Creek near confluence with Newport Bay near SR 73 overpass,

UTM (NAD 83) 11S 0419867E 3723886N, elev. 6 m (20 ft), uncommon, edge of disturbed riparian woodland with Salicornia, Euthamia

icaria paludosa, 12 Nov 2006, Riefner 06-671 (RSA). Riverside Co.: Temescal Valley, W side of I-15 Freeway, ca. 1.1

mi N of Temescal Canyon Rd., UTM (NAD 83) 11S 0453861E 3739203N, 289 m (949 ft), common, planted in residential landscapes

and spreading to roadsides, 2 Aug 2006, Riefner 06-356 (RSA, UC); Corona-La Sierra area, vicinity of Green River Rd. and Palisades Dr.,

UTM (NAD 83) 11S 0440802E 3749217N, 141 m (462 ft), uncommon, spreading from nearby ial landscape pl g

to edge of mule fat scrub, roadside ditch, and dirt lot, 22 Aug 2006, Riefner 06-398 (RSA, UC).

—

occidentalis, and Pu

Previous knowledge —Another introduced and naturalized perennial Limonium, as yet unidentified to

species, but probably originating from the Mediterranean region of Eurasia, has been recently collected from

salt marshes in San Diego County (Smith 2005; Rebman & Simpson 2006). This unidentified taxon has not

been reported from Orange or Riverside counties (Roberts 1998; Roberts et al. 2004). This species is actively

being planted in tract home development projects in Orange and Riverside counties, and is spreading rapidly to

native riparian and wetland habitats.

Wetland Indicator Status.—An indicator status has not been assigned to this currently unidentified species

of Limonium.

Significance and Recommended Indicator Status.—First annotated records of the Limonium indet. reported for

Orange and Riverside counties. This exotic perennial is more robust and aggressive than L. ramosissimum, and is

abundantly naturalized in and around sensitive salt marsh ecological reserves. It is rapidly displacing native salt

marsh species, and could also become a threat to the endangered Cordylanthus maritimus in the South Coast

region. The California Invasive Plant Council should monitor it closely. After taxonomic problems have been

resolved, it should be added to the California regional list and assigned a FACW wetland indicator status.

Oncosiphon piluliferum (L.f.) Kállersjó (Asteraceae)

CALIFORNIA. Imperial Co.: E of El Centro, Ross Rd. ca. 0.2 mi E of Dogwood St., UTM (NAD 83) 11S 0637549E 3628051N, elev.

-9 m C28 ft), two plants on disturbed, vernally moist alkaline flats with Atriplex and Suaeda, 20 Aug 2005, Riefner 05-644 (RSA). Los

730 Journal of the Botanical R h Institute of Texas 1(1)

Angeles Co.: Transverse Ranges, San Gabriel Mountains region, alluvial fan below San Antonio Canyon at southern base of the range, just

west of Los Angeles/San Bernardino County line along north margin of the west-bound 210-Fwy. at Monte Vista/Baseline Ave. off-ramp, UTM

(NAD 83) 11S 0436039E 3776202N, elev. 488 m (1600 ft), single plant along edge of road shoulder in hydroseeded scrub, 11 May 2006,

Boyd 11659 (RSA), City of Long Beach, Studebaker Rd. at 22-Fwy., UTM (NAD 83) 11S 0397935E 3737465N, elev. 6 m (20 ft), uncommon,

ruderal alkaline plant community on dirt lot, 8 Jul 2006, Riefner 06-294 (RSA); City of Long Beach, Shopkeeper Rd. at Westminster Blvd.,

UTM (NAD 83) 115 0397433E 3735926N, elev. 3 m (9 ft), uncommon, edge of salt marsh and vernally moist alkaline flats, 8 Jul 2006,

Riefner 06-296 (RSA). Riverside Co.: Romoland, Sherman Rd. near Ethanac Rd., UTM (NAD 83) 115 0483282E 3733376N, elev. 440

m (1443 ft), common in dry basin of seasonal pool with Crypsis vaginiflora and adjacent upland ruderal habitats, 30 May 2006, Riefner

06-213 (RSA).

Previous knowledge.—Oncosiphon piluliferum (Matricaria globifera [Thunb.] Fenzl; stink-net or globe

chamomile), a native to the Cape region of South Africa, was first reported in California from Orange and

San Diego counties (Sanders 1996). It is now locally common and spreading rapidly across western River-

side County (Hrusa et al. 2002; Roberts et al. 2004; Riefner & Boyd 20053). It has also been reported from

Arizona, where it has been documented from roadside, desert scrub, and wash and floodplain riparian

habitats (Landrum et al. 2005).

Wetland Indicator Status.—Oncosiphon piluliferum has not been assigned a wetland indicator status for

California on the 1988 list or the 1996 draft list in any region of the United States.

Significance and Recommended Indicator Status.—First records of O. piluliferum documented for Imperial

and Los Angeles counties. In western Riverside County, this species is well documented from alkali playa and

vernal alkali plain habitats dominated by facultative wetland species including Plagiobothrys leptocladus, Cressa

truxillensis, Crypsis schoenoides, Rumex crispus, and Atriplex argentea (Hrusa et al. 2002). It is spreading to low-ly-

ing vernal alkaline habitats in Imperial and Los Angeles counties. However, O. piluliferum is invading a variety

of disturbed upland and seasonal wetlands habitats, including roadside, field, floodplain, seasonal pool, and

scrub habitats in southern California and Arizona (Roberts et al. 2004; Riefner & Boyd 2005a; Landrum et

al. 2005). Oncosiphon piluliferum is expected to become a widespread invasive species in Arizona, which serves

as an example of the potential impacts to native vegetation when a non-native ornamental species becomes

naturalized (Landrum et al. 2005). It is also spreading explosively in southern California, and the California

Invasive Plant Council should monitor O. piluliferum closely. Based on field observations, information available

in the literature, and review of the Consortium of California Herbaria (2006), O. piluliferum should be added to

the California regional list. We propose a FACU* wetland indicator status and recommend additional regional

review to specifically define its frequency of occurrence in wetlands as the species undergoes further expan-

sions of range and invasion of new habitat types in California.

Panicum coloratum L. (Poaceae)

CALIFORNIA. Imperial Co.: S of El Centro, Bowker Rd. near McCabe Rd., UTM (NAD 83) 11S 0643807E 3624673N, elev. 2 m (6

ft), uncommon in wet irrigation ditch, 9 Oct 2006, Riefner 06-497 (RSA); S of El Centro, Heber Rd. near Hwy. 86, UTM (NAD 83) 11S

0635522E 3622345N, elev. 2 m (6 ft), uncommon in ditch with Typha and Chloracantha spinosa, 9 Oct 2006, Riefner 06-500 (RSA); E

of El Centro, along Ross Rd. near Bass Cove Rd., N of Hwy 8., UTM (NAD 83) 11S 0638008E 3628013N, elev. -9 m (-30 ft), common and

widespread in ditches, along irrigation canals, and in low-lying wet fields, 9 Oct 2006, Riefner 06-512 (RSA, UWSP). Los Angeles Co.:

City of Altadena, along Calveras St. near Hollister Ave., UTM (NAD 83) 11S 0396508E 3783148N, 390 m (1280 ft), uncommon, street gutter

in moist soil with Eleusine, 22 Aug 2004, Riefner 04-378 (RSA). Orange Co.: City of Los Alamitos, Oak Middle School, vicinity of Oak St. at

Catalina St., UTM (NAD 83) 11S 0400172E 3741242N, 14 m (45 ft), very rare, moist depressions in ball field turf grass with Eleusine, 3 Sep

2004, Riefner 04-425 (RSA).

Previous knowledge.—Panicum coloratum (kleingrass) is an African species occasionally cultivated for forage

that has been introduced into subtropical and tropical regions worldwide (Freckmann & Lelong 2003). In

the United States, it is currently known only from New Mexico and Texas, where it grows in open, usually

wet ground (Freckmann & Lelong 2003).

Wetland Indicator Status.—Panicum coloratum has not been assigned a wetland indicator status for Cali-

fornia on the 1988 list or the 1996 draft list.

Significance and Recommended Indicator Status.—First record of P. coloratum documented for California,

which likely represents a recent introduction. As with many other African grasses now established in south-

ern California, this weedy species could spread rapidly to other disturbed, moist-soil habitats in the south

Riefner and Boyd, Wetland and riparian plants in southern California 731

coast region. Based on initial field observations and information available in the literature, P. coloratum should

be added to the California regional list. We propose a FACW* wetland indicator status. Additional regional

review is needed to refine its frequency of occurrence in wetlands, as the species will likely undergo further

expansions of range and colonization of different habitat types in southern California.

Panicum dichotomiflorum Michx. subsp. dichotomiflorum (Poaceae)

CALIFORNIA. Orange Co.: City of Huntington Beach, Huntington Central Park at Goldenwest St., UTM (NAD 83) 11S 0406884E

3729853N, elev. 1 m G ft), common in and along the edge of shallow water ponds, 20 Sep 2005, Riefner 05-674 (RSA, UWSP), City of

Yorba Pon Santa Ana River near Yorba Linda Regional Park, UTM (NAD 83) 115 0430104E 3748155N, elev. 98 m (320 ft), common on

1 vith perennial waters, 27 Oct 2005, Riefner 05-735 (RSA, UWSP). Riverside Co.: City of Corona, E of Santa Ana

Canyon along Santa Ana River, ca. 0.5 mi S of 71 Freeway, UTM (NAD 83) 11S 0439689E 3749438N, elev. 135 m (442 ft), uncommon

in wet river wash sand, 18 Sep 2004, Riefner 04-435 (RSA).

Previous knowledge.—Panicum dichotomiflorum subsp. dichotomiflorum (fall panic grass), native to the eastern

United States, is found in the San Joaquin Valley and the South Coast region of California (Webster 1993b). It

grows in open, often wet disturbed areas in a wide variety of habitats, including cultivated and fallow fields,

roadsides, ditches, stream banks, along receding shores of lakes, clearings in floodplains, and in shallow water

(Freckmann & Lelong 2003). In southern California, P. dichotomiflorum has been collected from San Bernardino,

San Diego, and Santa Barbara counties, but not from Orange or Riverside counties (Munz 1974; Roberts 1998;

Roberts et al. 2004; Consortium of California Herbaria 2006, Rebman & Simpson 2006). This weedy species

was not included in the treatment of aquatic and riparian weeds by DiTomaso & Healy (2003).

Wetland Indicator Status.—Panicum dichotomiflorum subsp. dichotomiflorum has been assigned a FACW

wetland indicator status for California on the 1988 list and the 1996 draft list.

Significance and Recommended Indicator Status.—First records of P. dichotomiflorum subsp. dichotomiflorum

documented for Orange and Riverside counties. This grass will likely become a common weed of disturbed

wetland and riparian habitats throughout coastal southern California.

The P. dichotomiflorum complex shows great plasticity of growth forms in response to environmental

conditions, which is the case of the extremely robust plants collected from the Santa Ana River in Orange and

Riverside counties. These specimens have spikelets similar to P. dichotomiflorum subsp. puritanorum (Svenson)

Freckmann & Lelong, but no other characters. Given the great variation in the P. dichotomiflorum complex and

especially the effect of growing conditions on robustness, the Santa Ana River specimens are best treated as the

subsp. dichotomiflorum (pers. com., RW. Freckmann, October 2006). No changes to the wetland indicator status

are proposed at this time.

Panicum virgatum L. (Poaceae)

CALIFORNIA. Orange Co.: near City of Anaheim, Santa Ana Canyon, N side of Santa Ana River channel, Santa Ana Canyon, W

(downstream) ca. 1.25 mi from Green River Golf Course, UTM (NAD 83) 11S 0434148E 3748318N, elev. 120 m (387 ft), uncommon in

moist swales, on river sandbars, and in Baccharis salicifolia scrub, 19 Sep 2006, Riefner 06-439 (RSA, UWSP); City of Irvine, San Diego

Creek near 405-Freeway, UTM (NAD 83) 115 0429336E 3724184N, elev. 49 m (162 ft), uncommon on moist sandy banks and margin of

riparian woodland, 8 Oct 2006, Riefner 06-482 (RSA).

Previous knowledge.—Panicum virgatum (including P. havardii Vasey; switchgrass, panic raide) has not been

reported previously for California (Webster 1993b; Freckmann & Lelong 2003). It grows in mesic to wet tallgrass

prairies, and on dry slopes, sand, open oak or pine woodlands, shores, riverbanks, and brackish marshes. Pani-

cum virgatum occurs primarily on the eastern side of the Rocky Mountains, extending from southern Canada to

Mexico, Cuba, Bermuda, and Costa Rica, but is also found in Nevada, Utah, and Arizona (Freckmann & Lelong

2003). Panicum virgatum is planted for range and wildlife habitat improvement, and has been introduced as a

forage grass to other parts of the world (Freckmann & Lelong 2003).

Wetland Indicator Status.—Not previously known from California, P. virgatum was not assigned a wetland

indicator status on the 1988 California list, but it was included on the 1996 draft list with a FAC wetland indica-

tor status for most other regions.

Significance and R led Indicator Status.—First records of P. virgatum documented for California, which

likely represents an escape from cultivation or habitat enhancement plantings. Based on initial field observa-

tions and information available in the literature, P. virgatum should be added to the California regional list. We

732 Journal of the Botanical R h Institute of Texas 1(1)

propose a FAC* wetland indicator status and recommend additional regional review to refine its frequency of

occurrence in wetlands, as the species will likely undergo further expansions of range and colonization of

different habitat types in southern California.

Paspalum urvillei Steudel (Poaceae)

CALIFORNIA. Orange Co.: City of Lake Forest, Alton Pkwy. at Bertea Rd., UTM (NAD 83) 115 0434065E 3724402N, elev. 123m

(402 ft), common, growing with Baccharis salicifolia and Picris in ditches, swales, and fields, 4 Nov 2005, Riefner 05-742 (RSA, UCR).

Previous knowledge.—Paspalum urvillei (vaseygrass), a native of South America, occupies disturbed, moist

to wet places in the Sacramento Valley and the South Coast region (Webster 1993c; DiTomaso & Healy 2003).

However, Allen & Hall (2003) did not report it from southern California. Local herbaria have specimens from

San Bernardino and San Diego counties, where it occupies streambeds, margins of lakes and meadows, and

roadside habitats, but it has not been collected from Riverside and Orange counties (Roberts 1998; Roberts et al.

2004; Consortium of California Herbaria 2006; Rebman & Simpson 2006). Paspalum urvillei is more common

in the southern United States, but is expected to expand its range in California (DiTomaso & Healy 2003).

Wetland Indicator Status.—Paspalum urvillei has been assigned a NI* designation for California on the 1988

list and the 1996 draft list.

Significance and Recommended Indicator Status.—First record of P. urvillei documented for Orange County,

and additional populations will likely be discovered in the South Coast region. Field observations, a literature

review, and a review of herbarium data indicate this species frequently inhabits moist soil habitats in California

(DiTomaso & Healy 2003; Consortium of California Herbaria 2006). A revised FACU wetland indicator status

is proposed for the California regional list.

Psilocarphus chilensis A. Gray (Asteraceae)

LIFORNIA. Riverside Co.: Winchester, Patton Rd. at Patterson Ave., UTM (NAD 83) 11S 0492961E 3728074N, elev. 452 m (1483

ft), common in shallow roadside depressions with Plagiobothrys leptocladus, 22 Apr 2005, Riefner 05-239 (RSA, UCR); City of Lake Elsinore,

back basin of Lake Elsinore, W ca. 0.5 mi off Pete Lehr Drive at Diamond Stadium in Wildlife Viewing Area, UTM (NAD 83) 11S 0471224E

3723561N, elev. 402 m (1320 ft), uncommon in shallow depressions with Plagiobothrys acanthocarpus and Plantago elongata in saltgrass

meadows, 8 Apr 2006, Riefner 06-306 (RSA, UCR); Romoland, Sherman Rd. near Ethanac Rd., UTM (NAD 83) 11S 0483293E 3733281N,

elev. 440 m (1443 ft), uncommon in dry basin of detention pond with Crypsis vaginiflora, 30 May 2006, Riefner 06-215 (RSA).

Previous knowledge. —Psilocarphus chilensis (P. tenellus Nutt. var. globiferus [DC.] Morefield; round woolly

marbles) is an amphitropical disjunct, known in North America only from California, and from Chile in

South America (Morefield 2006). Psilocarphus chilensis occupies saturated to drying vernal pools, seasonally

inundated sites, and coastal interdune areas of the Sierra Nevada foothills to the Central Coast region and

south to the San Joaquin Valley (Morefield 1993). In southern California, P. chilensis is known only from the

Winchester and Domenigoni Valley region of western Riverside County (Boyd & Ross 1996; Roberts et al.

2004; Morefield 2006).

Wetland Indicator Status.—Psilocarphus chilensis (P. tenellus var. globiferus) has not been assigned a wetland

indicator status for California on the 1988 list or the 1996 draft list.

Significance and Recommended Indicator Status.—First report of P. chilensis documented for the Romoland

and Elsinore Valley regions of western Riverside County. This species is tolerant of disturbed conditions, occurs

in man-made or modified seasonal pool habitats, and is expected to occur elsewhere in cismontane southern

California, especially interior valleys in Riverside and San Diego counties. Based on field observations, informa-

tion available in the literature, and review of the Consortium of California Herbaria (2006), P. chilensis should

be added to the California regional list and assigned an OBL wetland indicator status.

Salvinia molesta D.S. Mitch. (Salviniaceae)

CALIFORNIA. Orange Co.: City of Huntington Beach, Huntington Central Park at Goldenwest St., UTM (NAD 83) 11S 0407062E

3730088N, elev. 1 m (3 ft), very rare in still-water pond with Lemna and Eichhornia ipes in willow riparian woodland, 30 Sep 2005,

Riefner 05-700 (RSA), City of Irvine, along Jeffrey Rd. N of Barranca Rd., UTM (NAD 83) 11S 0427803E 3726479N, elev. 47 m (155 fo,

uncommon, growing with Eichhornia crassipes in irrigation pond in agricultural field, 5 May 2005, Riefner 05-304 (RSA).

Previous E molesta (giant salvinia) is an invasive, free-floating aquatic fern native to

South America that has proven to be a troublesome weed in many warm freshwaters of the world (Barrett

Riefner and Boyd, Wetland and riparian plants in southern California 733

1989). In the United States, this federally listed noxious weed occurs across the southern states from Florida

to California (Barrett 1989; DiTomaso & Healy 2003). In California, S. molesta is well known from the lower

Colorado River drainage in Imperial and Riverside counties, but small populations have also been reported

along the coast in Orange, San Diego, and San Luis Obispo counties (Hrusa et al. 2002; Riefner & Boyd

2005b).

Salvinia molesta appears to only reproduce vegetatively in the United States, but is capable of rapid

growth in warm, slow-moving waters where it can be devastating to aquatic native species by covering large

areas (Mitchell et al. 1980; DiTomaso & Healy 2003). In most cases, the range of this plant is temperature

limited. Salvinia molesta survives extremes of 3'C and 43°C, however optimal growth occurs at 24°C to 28°C

(Whiteman & Room 1991). Salvinia molesta has been recognized as a species with the potential to spread

explosively in California (CalEPPC 1999).

Wetland Indicator Status.—Salvinia molesta has not been assigned a wetland indicator status for Califor-

nia on the 1988 list or the 1996 draft list. Salvinia auriculata Aubl. is included on the 1996 draft list with

an OBL wetland indicator status for the Caribbean and North Central regions, and a NI indicator status

for the Northeast, Southeast, and South Plains regions, but not for California. Salvinia auriculata has been

misapplied to Salvinia species in the United States, including S. minima Baker (S. auriculata auct. non Aubl.),

which occurs in New Mexico and other southern states (Nauman 1993; DiTomaso & Healy 2003).

Si led Indicator Status.—New locations of S. molesta documented for Orange County.

This EOM is spreading, most likely by waterfowl, in the mild Mediterranean climate of southern California.

Salvinia molesta may be expected to naturalize in the same areas where water hyacinth (Eichhornia crassipes

[Mort.] Solms) is known to grow, including slow-moving freshwaters in regions that may experience frost but

not the formation of ice (Whiteman & Room 1991; DiTomaso & Healy 2003; Riefner & Boyd 2005b). Based

on field observations and information available in the literature, S. molesta should be added to the California

regional list and assigned an OBL wetland indicator status.

Setaria adhaerens (Forssk.) Chiov. (Poaceae)

CALIFORNIA. Los Angeles Co.: City of Pasadena, Arroyo Seco Creek, S of Arroyo Blvd., UTM (NAD 83) 11S 0392637E 3779048N,

226 m (740 ft), uncommon along creek in wet sand and on sand bars with Eleusine, Ehrharta erecta, Leptochloa fusca subsp. uninervia,

and Xanthium strumarium, 22 Aug 2004, Riefner 04-383 (RSA). Orange Co.: City of Anaheim, Santa Ana River bottom, E of 57-Freeway

between Orangewood St. and Chapman Ave., UTM (NAD 83) 11S 0418311E 3739098N, 41 m (135 ft), locally abundant, disturbed

wetlands, margins of active channels, and on sandbars with Echinochloa crus-galli, Leptochloa fusca subsp. uninervia, and Xanthium stru-

marium, 8 Aug 2004, Riefner 04-373a (RSA); City of Irvine, San Diego Creek at Woodbridge High School, S of intersection of Barranca Rd

and West Yale Loop Rd., UTM (NAD 83) 11S 0425279E 3727007N, elev. 27 m (89 ft), uncommon, growing with Cynodon dacty

Xanthium strumarium on disturbed creek terrace, 27 Aug 2004, Riefner 04-389 (RSA); City of Irvine, San Diego Creek near 405-Freeway,

UTM (NAD 83) 11S 0429336E 3724184N, elev. 51 m (166 ft), uncommon on moist sandy banks and wet sand along creek channel with

Echinochloa crus-galli, Leptochloa fusca subsp. uninervia, and Ludwigia repens, 8 Oct 2006, Riefner 06-480 (RSA); City of Santa Ana, 17th

St. at 55-Fwy., UTM (NAD 83) 11S 0422908E 3735773N, elev. 49 m (160 ft), common along roadside in ruderal vegetation, 23 Jul 2006,

Riefner 06-310 (RSA); City of Huntington Beach, Atlanta St. at Surge Ln., UTM (NAD 83) 11S 0411693E 3724638N, elev. 11 m (35 ft),

sidewalk weed, 6 Aug 2006, Riefner 06-373 (RSA). Riverside Co.: City of Riverside, residential lots and roadside along Elderwood Ct. at

Tequesquite Ave., UTM (NAD 83) 11S 0463408E 3759616N, elev. 237 m (777 ft), common in irrigated landscape and roadside seepage

areas, 7 Oct 2005, Riefner 05-723 (ASC, RSA); City of La Sierra; Gramercy Pl. near La Sierra Ave., UTM (NAD 83) 11S 0454476E 3754271N,

elev. 227 m (746 ft), sidewalk and gutter weed, 30 Dec 2005, Riefner 05-793 (RSA). San Bernardino Co.: City of Fontana, Merrill St. at

Olive St., UTM (NAD 83) 11S 0459593E 3772444N, elev. 381 m (1250 ft), uncommon in drainage ditch with Cyperus in vacant dirt lot,

5 Oct 2005, Riefner 05-722 (RSA). San Diego Co.: Kearny Mesa, Clairemont Mesa Rd. near Ruffner Rd., UTM (NAD 83) 11S 0485359E

3632725N, elev. 122 m (401 ft), irrigated landscape with Agrostis, 19 Nov 2006, Riefner 06-679 (RSA).

Previous knowledge.—Setaria adhaerens (burr bristlegrass) grows in subtropical and tropical regions

—

on and

throughout the world (Rominger 2003). In North America, it occurs in the southern United States, northeast-

ern Mexico, Cuba, and the Bahamas (Rominger 2003). In California, it is known only from the Kern County

region, which may represent a recent introduction (Rominger 2003). Setaria adhaerens has not been collected

from Los Angeles, Orange, Riverside, San Bernardino, or San Diego counties (Webster 1993d; Roberts 1998;

Rominger 2003; Roberts et al. 2004; Consortium of California Herbaria 2006; Rebman & Simpson 2000).

Wetland Indicator Status.—Setaria adhaerans was not assigned a wetland indicator status on the 1988 list or

734 Journal of the Botanical R h Institute of Texas 1(1)

the 1996 draft list for any region in the United States. Setaria verticillata (L.) P. Beauv., native to Europe, resembles

S. adhaerens and also occupies disturbed moist-soil habitats in southern California. Setaria verticillata has been

assigned a NI* wetland indicator status for California on the 1988 list and a FACU wetland indicator status on

the 1996 draft list.

Significance and Recommended Indicator Status.—First documented report of S. adhaerens for Los Angeles,

Orange, Riverside, San Bernardino, and San Diego counties. Based on initial field observations and information

available in the literature, S. adhaerens should be added to the California regional list. We propose a FACU*

wetland indicator status. This species will likely undergo further expansions of range and colonization of

different habitat types in southern California, which will require further review to refine its frequency of

occurrence in wetlands.

Setaria megaphylla (Steud.) T. Durand & Schinz (Poaceae)

CALIFORNIA. Orange Co.: City of Huntington Beach, Huntington Central Park near Goldenwest St., UTM (NAD 83) 11$ 0406845E

3730055N, elev. 1 m (3 fD, uncommon in disturbed Salix-dominated riparian woodland with Echinochloa crus-galli and Paspalum dilatatum,

27 Oct 2005, Riefner 05-731 (RSA); City of Newport Beach, urban creek draining into San Diego Creek near confluence with Newport Bay

at SR 73 overpass, UTM (NAD 83) 115 0419895E 3723901N, elev. 6 m (19 ft), locally common in moist disturbed riparian woodland with

Euthamia occidentalis and Pulicaria paludosa, 12 Nov 2006, Riefner 06-667 (ASC, RSA); City of San Juan Capistrano, San Juan Creek at

La Novia Ave., UTM (NAD 83) 11S 0439744E 3707154N, elev. 22 m (72 ft), uncommon and widely scattered in riparian scrub, 19 Nov

2006; Riefner 06-683 (RSA).

Previous knowledge.—Setaria megaphylla (bigleaf bristlegrass), native to tropical Africa and tropical America,

has not been reported previously for California (Webster 1993d; Rominger 2003). In North America, it is ap-

parently established only in Florida, although it is occasionally cultivated for its ornamental, plicate leaves

(Rominger 2003).

Wetland Indicator Status.—Setaria megaphylla was not assigned a wetland indicator status on the 1988 list

or the 1996 draft list for any region in the United States. Setaria palmifolia J. Konig) Stapf, a species similar to

S. megaphylla, has been assigned a NI wetland indicator status for the Southeast region and a FACU wetland

indicator status for Hawaii.

Significance and Recommended Indicator Status.—First report of S. megaphylla documented for California,

which likely represents an escape from cultivation. Based on initial field observations and information avail-

able in the literature, S. megaphylla should be added to the California regional list. We propose a NI wetland

indicator status. This species may undergo further expansions of range and colonization of different habitat

types in southern California, but may be restricted to moist soils and mild climate along the immediate

coast. Additional monitoring of new populations is necessary to determine its frequency of occurrence in

wetlands.

DISCUSSION

Wetlands, streams and their associated riparian vegetation communities are important natural resources in

urban and suburban southern California (White & Greer 2006). The increased area of impervious surfaces

and imported municipal water supplies associated with the urbanization of coastal watersheds have driven

significant and often undocumented changes in many of coastal southern California's wetland and riparian

vegetation communities by altering stream flow characteristics, channel geomorphology, and historic plant

community composition (Greer & Stow 2003; Burkhart 2006). In southern California, and many other urban

watersheds, these changes also include dramatic shifts from historic episodic stream flows following major

storm events to perennial base flows, which is often accompanied by an expansion of wetland and riparian

vegetation, including human-adaptive or disturbance-tolerant native species, and an overall decline in na-

tive species richness (Magee et al. 1999; Dwire et al. 2000; Shanahan & Crear 2004; Burkhart & Kelly 2005;

Burkhart 2006; White & Greer 2006). Also, the direct effects of the introduction of exotic species in suburban

and urban environments have significantly modified the character of the native vegetation and integrity of

wetland and riparian ecosystems associated with many urban environments, which is directly related to avail-

able moisture, and secondarily to disturbance (Suarez et al. 1998; Magee et al. 1999; McKinney 2002; Radeloff

et al. 2005). The effects of urban development on ecological communities have only recently become a topic

Riefner and Boyd, Wetland

tsi th California 735

TABLE 2. Summary of distributional records for southern California, and proposed revisions and additions to the National List

of Plant Species that Occur in Wetlands.

Species

(*=exotic taxon)

Significance in California

and Recommended Wetland Indicator Status

*Amaranthus blitum subsp.

emarginatus

Atriplex polycarpa

*Bacopa monnierii

*Beta vulgaris subsp. maritima

*Chloris truncata

*Fhrharta erecta

*Elytrigia repens

Epilobium brachycarpum

Eriochloa aristata

*Glinus radiatus

*limonium ramosissimum

*Limonium indet.

*Oncosiphon piluliferum

*Panicum coloratum

*Panicum dichotomiflorum

Sl Jbsp dic hi JO! Jf JUTTI

*Panicum virgatum

*Paspalum urvillei

Psilocarphus chilensis

*Salvinia molesta

*Setaria adhaerens

*Setaria megaphylla

New to Orange County; spreading to urban wetlands and stream course habitats;

propose FAC* wetland indicator status.

New to western Riverside County; alkaline soil habitats in southern California

are in need of focused study; concur with current FACU indicator status.

New to Orange County; one of many subtropical and tropical weeds invading

coastal southern California; concur with revised OBL indicator status.

New to Riverside County; propose FAC* wetland indicator status.

New to San Bernardino County; propose NI* wetland indicator status.

New to Imperial County and verification of an unreported Orange Count

herbarium record; spreading rapidly from urban to native habitats; new records

documented from native riparian and stream course habitats in Los Angeles, Orange,

and Ventura counties; assign FAC wetland indicator status.

New to Orange County; concur with revised FAC* indicator status.

New to Orange County; revise from UPL and assign FACU wetland indicator status.

New to western Riverside County; expected elsewhere in disturbed wetland

habitats in southern California; concur with current FACW indicator status.

New to Orange and western Riverside counties; likely more widespread than

current records indicate; revise from NI and assign OBL wetland indicator status.

New to Los Angeles, Orange, Riverside, and San Diego counties; spreading rapidly;

threat to the endangered Cordylanthus maritimus in salt marsh habitats;

assign FACW wetland indicator status.

New to Orange and Riverside counties; spreading rapidly from cultivation;

potential threat to the endangered Cordylanthus maritimus in salt marsh habitats;

assign FACW wetland indicator status following identification to species.

New to Imperial and Los Angeles counties; potential to spread explosively; propose

FACU* wetland indicator status.

New to California; African species with potential to spread explosively; propose

FACW* wetland indicator status.

New to Orange and Riverside counties; expected to invade disturbed wetland

and riparian habitats throughout the South Coast region; concur with current

FACW indicator status.

New to California; propose FAC* wetland indicator status.

New to Orange County; revise from NI* and propose FACU wetland indicator status.

New to Romoland and Elsinore Valley regions of western Riverside County;

assign OBL wetland indicator status.

New populations reported for Orange County; federally listed noxious weed

with potential to spread explosively; assign OBL wetland indicator status.

New to Los Angeles, Orange, Riverside, San Bernardino, and San Diego

counties; subtropical and tropical | spreading rapidly in coastal southern

California; propose FACU* wetland indicator status.

New to California; recent introduction of another tropical species; propose NI

wetland indicator status.

of study, however, and few comprehensive surveys have been carried out in coastal southern California, or

in other regions, to document this relatively recent phenomenon (Dwire et al. 2000; Hansen et al. 2005;

Burkhart 2006; White & Greer 2006). Botanists and ecologists have traditionally focused their research on

wildland ecosystems, but the new records presented herein, and summarized in Table 2, suggest that detailed

floristic studies are needed to thoroughly document the flora of wetland and riparian communities in urban

ecosystems and in surrounding native habitats degraded by exotic species throughout the South Coast region.

Expected declines in native plant biodiversity of wetland and riparian ecosystems in or near wildland

736 Journal of the Botanical R h Institute of Texas 1(1)

habitats, and in urban nature preserves and parks, pose major environmental problems to resource conservation

planning, especially in California, which has the highest number of wildland-urban interface (WUD housing

units in the United States (McKinney 2002; Pysek et al. 2002; Hansen et al. 2005; Radeloff et al. 2005). As long

as the amiable climate and the strong economy of coastal southern California continues to attract urban-

ization and other forms of high density human activity, and without coordinated land-use planning and

ecologically-based conservation efforts, it will not be likely that substantial gains in area, native floristic

diversity, and unique wetland- and riparian-related functions and values will occur (Brinson & Malvárez

2002; Broberg 2003).

ACKNOWLEDGMENTS

J. Travis Columbus and LeRoy Gross of the Rancho Santa Ana Botanic Garden (RSA), and Andrew Sanders,

University of California at Riverside (UCR), provided annotations and/or helpful discussion regarding the range

and habitat affinities of several taxa. We also thank Robert W. Freckmann, University of Wisconsin at Stevens

Point (UWSP), who annotated Panicum and provided helpful discussion of the P. dichotomiflorum complex,

James W. Rominger, Northern Arizona University (ASC), who annotated Setaria, Robert Shaw, Colorado State

University (CS), who annotated Eriochloa, and Alan R. Smith, University of California Berkeley (UC), who an-

notated Limonium and provided helpful discussion regarding the invasive behavior of exotic Limonium species.

Vanessa Ashworth, Rancho Santa Ana Botanic Garden, Roy Shlemon, Roy J. Shlemon & Associates, Inc., and

two anonymous reviewers provided helpful comments that greatly improved this manuscript. Nancy Refulio-

Rodriguez, Rancho Santa Ana Botanic Garden, prepared the Spanish resumen, and Kurt Campbell, Jones &

Stokes Temecula Office, provided helpful discussion regarding riparian plant communities. This research was

funded, in part, by a grant to the senior author from the International Foundation for Applied Research in the

Natural Sciences (IFARNS).

REFERENCES

ALLEN, C.M. and D.W. Hatt. 2003. Paspalum. In: Flora of North America Editorial Committee, eds. Flora of North

America north of Mexico, Vol. 25, Magnoliophyta: Commelinidae (in part): Poaceae, part 2. Oxford University

Press, New York, NY. Pp. 566-599.

ARNOLD, C.L. and C.J. GIBBONS. 1996. Impervious surface coverage: the emergence of a key environmental indica-

tor. J. Am. Plan. Assoc. 62:243-258.

Baber, J.S. 1969. Ground-water data as of 1967, south coastal subregion, California. U.S. Geol. Survey, Water Re-

sources Division, Open-File Report 30, Melano Park, CA.

Banks, D.L. 1999. A vascular flora of the Agua Tibia Mountains, southern California. Rancho Santa Ana Botanic

Garden Occasional Publications, No. 4, Claremont, CA.

BARKWORTH, M.E. 2003. Chloris. In: Flora of North America Editorial Committee, eds. Flora of North America north

of Mexico, Vol. 25, Magnoliophyta: Commelinidae (in part): Poaceae, part 2. Oxford University Press, New

York, NY. Pp. 204-218.

Barrett, S.C.H. 1989. Waterweed invasions. Scientific American 261:90-97.

Batzer, D.P. and R.R. SHARITZ. 2006. Ecology of fresh water and estuarine wetlands: an introduction. In: D.P. Batzer

and R.R. Sharitz, eds. Ecology of fresh water and estuarine wetlands. University of California Press, Berkeley,

CA. Pp. 1-6.

BEAUCHAMP, R.M. 1986. A flora of San Diego County, California. Sweetwater River Press, National City, CA.

Bennett, PS., M.R. KUNZMANN, and R.R. JOHNSON. 1989. Relative nature of wetlands: riparian and vegetational consid-

erations. In: D.L. Abell, ed. Proceedings of the California riparian systems conference. U.S. Dept. of Agriculture,

Forest Service, Pacific Southwest Forest and Range Experiment Station, Berkeley, CA. Pp. 140-142.

BossarD, C.C., J.M. RANDALL, and M.C. HosHovesky. 2000. Invasive plants of Californias wildlands. University of

California Press, Berkeley, CA.

Bower, PA. 1989. Riparian woodlands: an endangered habitat in southern California. In: A.A. Schoenherr, ed.

Endangered plant communities of southern California. Southern California Bot. Spec. Publ. No. 3. Pp. 80-97.

Riefner and Boyd, Wetland and riparian plants in southern California 737

Bovp, S. and T. Ross. 1996. Noteworthy collections-California: Cyperus acuminatus, Psilocarphus tenellus var. glo-

biferus, and Scirpus supinus var. saximontanus. Madrono 43:33.

Bovp, S. 1999. Vascular flora of the Liebre Mountains, Western Transverse Ranges, California. Aliso 18:93-139.

Brinson, M.M., B. L. Swiet, R.C. PLantico, and J.S. Barclay. 1981. Riparian ecosystems: their ecology and status. U.S.

Fish and Wildlife Service Biological Rep. 81/17.

Brinson, M.M. 1990. Riverine forests. In: A.E. Lugo, M.M. Brinson, and S. Brown, eds. Ecosystems of the world, Vol.

14: forested ecosystems. Elsevier, Amsterdam, The Netherlands. Pp. 87-141.

Brinson, M.M. and A.I. Malvárez. 2002. Temperate freshwater wetlands: types, status, and threats. Environm. Conserv.

29:115-133.

BROBERG, L. 2003. Conserving ecosystems locally: a role for ecologists in land-use planning. BioScience 53:

670-673.

BunkHART, B. and M. KeLLY. 2005. Mapping and planning: key first steps to eradicating the weedy plants that dominate

riparian systems in urban California. Ecological Restoration 23:40-45.

BurKHART, B. 2006. What weeds dominate southern California urban wetlands and why? Fremontia 34:14-19.

CALIFORNIA Exotic Pest PLANT Council (CalEPPC). 1999. Exotic plants of greatest ecological concern in California,

California Exotic Pest Plant Council. CalEPPC Newsletter, October:1-12

CONSORTIUM OF CALIFORNIA HERBARIA. 2006. Smasch Accession Results: Amaranthus blitum subsp. emarginatus, Atriplex

polycarpa, Bacopa monnieri, Beta vulgaris subsp. maritima, Chloris truncata, Ehrharta erecta, Elytrigia repens

(Agropyron repens), Epilobium brachycarpum, Eriochloa aristata, Glinus radiatus, Limonium ramosissimum, On-

cosiphon piluliferum, Panicum dichotomiflorum, Paspalum urvillei, Psilocarphus chilensis ME tenellus

Val globiferus), Salvinia molesta, Setaria adhaerens ,and Setaria gad]

bin/get_chc.pl). Accessed September 2 to December 18, 2006.

COWARDIN, L.M., V. Carter, F.C. Gotet, and E.T. Laroe. 1979. Classification of wetlands and deepwater habitats of the

United States. Department of the Interior, U.S. Fish and Wildlife Service, Washington, D.C.

Dan, T.E. 1990. Wetlands losses in the United States 1780s to 1980s. Department of the Interior, U.S. Fish and

Wildlife Service, Washington, D.C.

Dat, T.E. 2000. Status and trends of wetlands in conterminous United States 1986 to 1997. Department of the

Interior, U.S. Fish and Wildlife Service, Washington, D.C.

DiTomaso, J.M., and E.A. HEALv. 2003. Aquatic and riparian weeds of the West. U.C. Agriculture and Natural Resources

Publication 3421, Oakland, CA.

Dw, K.A., J.B. KAUFFMAN, and J. Baham. 2000. Relations among redox potentials, water levels, and riparian vegetation.

In: PJ. Wigington, Jr. and R.L. Beschta, eds. Riparian ecology and management in multi-land use watersheds.

American Water Resources Association, TPS-00-2, Middleburg, VA. Pp. 23-28.

Elus, AJ. and C.H. Lee. 1919. Geology and groundwaters of the western part of San Diego County, California. U.S.

Geol. Survey Water-Supply Paper 446.

Faber, P.M., E. KELLER, A. Sanps, and B.M. Massey. 1989. The ecology of riparian habitats of the southern California

coastal region: a community profile. U.S. Fish and Wildlife Service, National Wetlands Research Center, Wash-

ington, D.C. Biological Rep. 85 (7.27).

FERREN, W.R. 1993. Glinus. In: J.C. Hickman, ed. The Jepson manual: higher plants of California. University of Cali-

fornia Press, Berkeley, CA. P. 764.

FERREN, W.R. and P. FiebLER 1993. Rare and threatened wetlands of central and southern California. In: J. Keeley, ed.

Interface between ecology and land development in California. Southern California Academy of Sciences, Los

Angeles, CA. Pp. 119-131.

FERREN, W.R., P. FIEDLER and R.A. Ley. 1996. Wetlands of California, part I: history of wetland habitat classification. Ma-

drono 43:105-124.

FRECKMANN, R.W. and M.G. LeLone. 2003. Panicum. In: Flora of North America Editorial Committee, eds. Flora of North

America north of Mexico, Vol. 25, Magnoliophyta: Commelinidae (in part): Poaceae, part 2. Oxford University

Press, New York, NY. Pp. 450-488.

$ "i f I^ | | l

(http//ucje| du/cgi-

738 Journal of the Botanical R h Institute of Texas 1(1)

Greer, K.A. and D.A. Stow. 2003. Vegetation type conversion in Los Peñasquitos Lagoon: an examination of the

role of watershed urbanization. Environm. Manag. 31: 89-503.

GREGORY, S.V., F.J. SWANSON, W.A. McKee, and K.W. Cummins. 1991. An ecosystem perspective of riparian zones. BioSci-

ence 41:540-551.

HANSEN, A.J., R.L. KNIGHT, J.M. MarzLure, S. PoweLL, K. Brown, P.H. GUDE, AND K. Jones. 2005. Effects of exurban develop-

ment on biodiversity: patterns, mechanisms, and research needs. Ecological Applications 15:1893-1905.

HENRICKSON, J. 1993. Amaranthus. In: J.C. Hickman, ed. The Jepson manual: higher plants of California. University

California Press, Berkeley, CA. Pp. 131-134.

Hickman, J.C. 1993. Geographic subdivisions of California. In: J.C. Hickman, ed. The Jepson manual: higher plants of

California. University California Press, Berkeley, CA. Pp. 37-48,

Hoc, PC. 1993. Epilobium. In: J.C. Hickman, ed. The Jepson manual: higher plants of California. University California

Press, Berkeley. Pp. 793-798.

Horsten, G. 1984. California riparian forests: deciduous islands in an evergreen sea. In: R.E. Warner and K.E. Hendrix,

eds. California riparian systems: ecology, conservation, and productive management. University of California

Press, Berkeley, CA Pp. 2-22.

HRUSA, F., B. ERTTER, A. SANDERS, G. Leppic, and E. Dean. 2002. Catalogue of non-native vascular plants occurring spon-

taneously in California beyond those addressed in The Jepson Manual-Part |. Madroño 46:61-98.

Jarvie, J.K. and M.E. BAnkwoRrH. 1993. Elytrigia. In: J.C. Hickman, ed. The Jepson manual: higher plants of California.

University of California Press, Berkeley, CA. Pp. 1256-1257.

Katean, E.F. 1984. A brief history of riparian forests in the Central Valley of California. In: R.E. Warner and K.E. Hendrix,

eds. California riparian systems: ecology, conservation, and productive management. University of California

Press, Berkeley, CA. Pp. 23-29.

LANDRUM, L.R., L. Ducan, S. Whrrcome, J. ANDERSON, D. Dameet, and F.E. Taylor. 2005. Noteworthy collection-Arizona:

Oncosiphon piluliferum. Madrono 52:270-271.

Macte, T., T. Ernst, M.E. KeNrULA, and K. A. Dwire. 1999. Floristic comparison of freshwater wetlands in an urbanizing

environment. Wetlands 19:517-534.

MALANSON, G.P. 1993. Riparian landscapes. Cambridge University Press, Cambridge, England.

McCuntock, E. 1993. Limonium. In: J.C. Hickman, ed. The Jepson manual: higher plants of California. University

California Press, Berkeley, CA. Pp. 822-824.

McKinney, M.L. 2002. Urbanization, biodiversity, and conservation. BioScience 52:883-890.

MircHELL, D.S., T. Petr, and A.B. Vine. 1980. The water fern Salvinia molesta in the Sepik River, Papua New Guinea.

Environm. Conserv. 7:115-122.

MirscH, W.J., and J. G. GosseLiNk, 2000. Wetlands. 3* Ed. John Wiley & Sons, Inc., New York, NY.

MontriELD, J.D. 1993. Psilocarphus. In: J.C. Hickman, ed. The Jepson manual: higher plants of California. University

California Press, Berkeley, CA. P. 329.

MontriELD, J.D. 2006. Psilocarphus. In: Flora of North America Editorial Committee, eds. Flora of North America

north of Mexico, Vol. 19, Magnoliophyta: Asteridae (in part): Asteraceae, part 1. Oxford University Press, New

York, NY. Pp. 456-460.

MoSYAKIN, S.L. and K.R. Rogertson. 2003. Amaranthus. In: Flora of North America Editorial Committee, eds. Flora of

North America north of Mexico, Vol. 4, Magnoliophyta: Caryophyllidae, part 1. Oxford University Press, New

York, NY. Pp. 410-435.

Munz, PA. 1974. A flora of southern California. University of California Press, Berkeley, CA.

Munz, PA. and D.D. Keck. 1959. A California flora and supplement. University of California Press, Berkeley, CA.

NAUMAN, C.E. 1993. Salvinia. In: Flora of North America Editorial Committee, eds. Flora of North America north of

Mexico, Vol. 2, Pteridophytes and Gymnosperms. Oxford University Press, New York, NY. Pp. 336-338.

ORANGE County Water District (OCWD). 2006. Groundwater recharge operation (http//www.ocwd.com/ html/recharge.

htm). Accessed October 15, 2006.

PATTERN, D.T. 1998. Riparian ecosystems of semi-arid North America: diversity and human impacts. Wetlands

18:498-512.

Riefner and Boyd, Wetland and riparian plants in southern California 739

Paul, M.J. and J.L. Mever. 2001. Streams in the urban landscape. Ann. Rev. Ecol. Syst. 32:333-365.

Porr, N.L., J.D. ALLAN, M.B. Bain, J.R. Kann, K.L. PREsTEGAARD, B.D. RicHTer, R.E. Sparks, and J.C. STROMBERG. 1997. The natural

flow regime: a paradigm for river conservation and restoration. BioScience 47:769-784.

PoLAND, J.F. 1959. Hydrology of the Long Beach-Santa Ana area, California, with special reference to the water-

tightness of the Newport-Inglewood structural zone. U.S. Geol. Survey Water-Supply Paper 1471.

Pysek, P, V. Jarosik, and T. Kucera. 2002. Patterns of invasion in temperate nature reserves. Biological Conserv.

104:13-24.

RADELOFF, V.C., R.B. HAMMER, S.I. STEWART, J.S. FRIED, S.S. HoLcome, and J.F. McKeerry. 2005. The wildland-urban interface

in the United States. Ecological Applications 15:799-805.

REBMAN, J.P. and M.G. Simpson. 2006. Checklist of the vascular plants of San Diego County. 4^ Ed. San Diego Natural

History Museum, San Diego, CA.

REED, PB., Jr. 1988. National list of plant species that occur in wetlands: California (Region 0), Biological Rep. 88 [26.10].

U.S. Department of the Interior, U.S. Fish and Wildlife Service, Washington, D.C.

RIEFNER, R.E., JR, S. Boyd, and RJ. SHLEMON. 2002. Eleocharis obtusa var. engelmannii new to southern California. Cros-

sosoma 27:52-54.

RieFNER, R.E., JR. and S. Bovo. 2005a. Noteworthy collections from western Riverside County, southern California.

Crossosoma 31:28-32.

RiEFNER, R.E., JR. and S. Bovo. 2005b. Noteworthy collection-California: Salvinia molesta. Madroño 52:272-273.

RoserTs, F.M., JR. 1998. A checklist of the vascular plants of Orange County, California. 2"? Ed. F.M. Roberts Publica-

tions, Encinitas, CA.

Roserts, F.M., Jr., S.D. Wurre, A. C. SaNpERs, D.E. BramLer, and S. Bovo. 2004. The vascular plants of western Riverside

County, California: an annotated checklist. EM. Roberts Publications, San Luis Rey, CA.

ROMINGER, J.M. 2003. Setaria. In: Flora of North America Editorial Committee, eds. Flora of North America north of

Mexico, Vol. 25, Magnoliophyta: Commelinidae (in part): Poaceae, part 2. Oxford University Press, New York,

NY. Pp. 539-558.

SANDERS, A.C. 1996. Noteworthy collections-California: Certonia siliqua, Chloris truncata, Cynanchum louiseae,

Eragrostis curvula, Fatoua villosa, Matricaria globifera, Panicum maximum, Pistacia atlantica, Schismus polygamus,

Solanum mauritianum. Madroño 43:523-532.

SHANAHAN, S.A. and K.C. Crear. 2004. The Las Vegas Wash riparian ecosystem: development, degradation, and

active rehabilitation. Poster presented to the Mojave Desert Science Symposium, November 12-14, 2004,

Redlands, CA.

SHAW, S.P. and C.C. FreDINE. 1956. Wetlands of the United States. Circular 39. U.S. Fish and Wildlife Service, Wash-

ington, D.C.

SHAW, R.B., R.D. Wesster, and C.M. Bern. 2003. Eriochloa. In: Flora of North America Editorial Committee, eds. Flora

of North America north of Mexico, Vol. 25, Magnoliophyta: Commelinidae (in part): Poaceae, part 2. Oxford

University Press, New York, NY. Pp. 507-515.

SHULTz, L.M. 2003. Beta. In: Flora of North America Editorial Committee, eds. Flora of North America north of

Mexico, Vol. 4, Magnoliophyta: Caryophyllidae, part 1. Oxford University Press, New York, NY. Pp. 266-267.

Sica, J. 1996. Ehrharta erecta: sneak attack in the making? CalEPPC News, Summer/Fall:8—9.

Sica, J. 2003. Triple threat from South Africa. Fremontia 31:21-28.

SMITH, S.D., J.L. NACHLINGER, A.B. WELLINGTON, and C.A. Fox. 1989. Water relations of obligate riparian plants as a func-

tion of streamflow diversion on the Bishop Creek watershed. Pp. 360-365. In: D.L. Abell, ed. Proceedings of

the California riparian systems conference. U.S. Dept. of Agriculture, Forest Service, Pacific Southwest Forest

and Range Experiment Station, Berkeley, CA.

SurrH, A.R. 2005. Limonium. In: Flora of North America Editorial Committee, eds. Flora of North America north of

Mexico, Vol. 5, Magnoliophyta: Caryophyllidae, part 2. Oxford University Press, New York, NY. Pp. 606-611.

Somerville, D.E. and B.A. Pruitt. 2006. United States wetland regulation and policy. In: D.P. Batzer and R.R. Sharitz,

eds. Ecology of fresh water and estuarine wetlands. University of California Press, Berkeley, CA. Pp. 313-347.

740 Journal of the Botanical R h Institute of Texas 1(1)

STROTHER, J.L. 1993. Bacopa. In: J.C. Hickman, ed. The Jepson manual: higher plants of California. University of

California Press, Berkeley, CA. Pp. 1015-1016.

Suarez, J.G., D.T. BoLGER, and T.J. Case. 1998. Effects of fragmentation and invasion on native ant communities in

coastal southern California. Ecology 79:2041-2056.

Swit, B. 1984. Status of riparian ecosystems in the United States. Water Res. Bull. 20:223-227.

Tavlor, D. and D.A. Wilken. 1993. Atriplex. In: J.C. Hickman, ed. The Jepson manual: higher plants of California.

University of California Press, Berkeley, CA. Pp. 501-505.

Tiner, RW. 1988. Field guide to nontidal wetland identification. Maryland Department of Natural Resources, Annapolis,

and US. Fish and Wildlife Service, Newton Corner, MA.

Tiner, RW. 1991. The concept of a hydrophyte for wetland identification. BioScience 41:236-247.

Tiner, RW, 1993. The primary indicators method-a practical approach to wetland recognition and delineation in the

United States. Wetlands 13:50-64.

Tiner, RW, 1999. Wetland Indicators: a guide to wetland identification, delineation, classification, and mapping.

Lewis Publishers, CRC Press, Boca Raton, FL.

Tiner, RW. 2006. Lists of potential hydrophytes for the United States: a regional review and their use in wetland

identification. Wetlands 26: 624—634.

TROXELL, H.C. 1948. Hydrology of western Riverside County, California. Riverside County Flood Control and Water

Conserv. Dist. Rep., Riverside, CA.

U.S. Army Corps or ENGINEERS (USACE). 1987. Environmental laboratory, wetlands delineation manual. Technical

Report Y-87-1, U.S. Army Engineer Waterways Experimental Station, Vicksburg, MS.

U.S. Army Corps or ENGINEERS (USACE). 2006. Interim regional supplement to the Corps of Engineers wetland delin-

eation manual: arid west region. J.S. Wakeley, RW. Lichvar, and C.V. Noble, eds. U.S. Army Engineer Research

and Development Center, ERDC/EL TR-06-16, Vicksburg, MS.

U.S. Fish and Wildlife Service (USFWS). 1977. Concept plan for waterfowl wintering habitat preservation. Central

Valley California. Region 1, Portland, OR.

U.S. FisH AND WILDLIFE Service (USFWS). 1996. National list of vascular plant species that occur in wetlands: 1996

national summary. Washington D.C. (http://wetlands.fws.gov/bha/download/1996/national.pdf). Accessed

November 2005 through December 2006.

U.S. FisH AND WILDLIFE Service (USFWS). 2004. Technical procedures for wetlands status and trends. Department of the

Interior, U.S. Fish and Wildlife Service, Branch of Habitat Assessment, Arlington, VA.

Weester, R. 1993a. Eriochloa. In: J.C. Hickman, ed. The Jepson manual: higher plants of California. University of

California Press, Berkeley, CA. Pp. 1258-1259.

Weester, R. 1993b. Panicum. In: J.C. Hickman, ed. The Jepson Manual: higher plants of California. University of

California Press, Berkeley, CA. Pp. 1277-1278.

WEBSTER, R. 1993c. Paspalum. In: J.C. Hickman, ed. The Jepson manual: higher plants of California. University of

California Press, Berkeley, CA. P. 1280.

Wesster, R. 1993d. Setaria. In: J.C. Hickman, ed. The Jepson manual: higher plants of California. University of

California Press, Berkeley, CA. Pp. 1294-1296.

WELSH, S.L. 2003. Atriplex. In: Flora of North America Editorial Committee, eds. Flora of North America north of

Mexico, Vol. 4, Magnoliophyta: Caryophyllidae, part 1. Oxford University Press, New York, NY. Pp. 322-381.

Wuite, M. D. and K.A. Greer. 2006. The effects of watershed urbanization on the stream hydrology and riparian

vegetation of Los Peñasquitos Creek, California. Landscape and Urban Planning 74:125-138.

WHITEMAN, J.B. and P.M. Room. 1991. Temperatures lethal to Salvinia molesta Mitchell. Aquatic Bot. 40:27-35.

Yooer, V. 1996. An Alabama Hills plant list. California Native Plant Society, Bristlecone Chapter, Independence,

CA:

NOTEWORTHY PLANTS FROM NORTH FLORIDA. VIII

Loran C. Anderson

Professor Emeritus, Department of Biological Science

Florida State University

Tallahassee, Florida 32306-4370 U.S.A.

ABSTRACT

Additions to the flora of north Florida are documented. Bradburia pilosa (Nutt.) Semple, Hydrocotyle bowlesioides Mathias & Constance,

Ottelia alismoides (L.) Pers., Ranunculus laxicaulis (Torr. & A. Gray) Darby, and Solidago gigantea Ait. are new to the state; several species

new to the panhandle are listed. Additionally, many first reports for counties within the Florida panhandle are given.

RESUMEN

T3 4] H sg WT

Sed as adiciones a la flora del norte de Florida

i ) Semple, Hydrocotyle bowlesioides Mathias & Constance,

Ottelia alismoides (L.) Pers., Ranunculus laxicaulis (Torr. & A. Gray) Darby, y S

1:3

Ait. son Į lestado; se listan varias

especies nuevas para el *panhandle." Aditionalmente, se dan muchas primeras citas para condados del *panhandle" de Florida.

In most previous installments of this series (Anderson 1984, 1986, 1988, 1989, 1991, 1995), “north Florida"

was defined as the Florida panhandle west of the Suwannee River. In Anderson 2000 the area of coverage

was expanded to include selected counties of northeastern Florida. This installment returns to the original,

more restricted area of coverage, i.e., the area covered by Clewell (1985). Nomenclature generally follows

Wunderlin and Hansen (2003). Austin's study (1980) was used to identify Cuscuta species, and Flora of North

America treatments were used for Cyperaceae. Herbarium specimens are at FSU unless noted otherwise.

These findings update species distributions as mapped by Wunderlin and Hansen (2004).

TAXA NEW TO THE AREA

Arachis prostrata Benth.—Calhoun Co.: Rte 20 on E edge of Blountstown, 22 Jul 2002, Anderson 20353,

26 Jun 2003, Anderson 20736. Florida Department of Transportation workers (pers. comm.) planted this

here in 1999 as “Arachis glabrata cultivar. Ecoturf,” and it is naturalized and spreading. New to the Florida

panhandle.

Bradburia pilosa (Nutt.) Semple—Leon Co.: Capitol Circle NW at railroad crossing, Tallahassee, 23 May

2003, Anderson 20728, 22 May 2004, Anderson 20931. Population persisting but presently showing no signs

of becoming invasive. New to Florida; Semple (2006) mentioned possible introduction in Leon County,

based on my collections, but did not put a dot for Florida on the map.

Cichorium intybus L.—Escambia Co.: W of Cantonment, 26 May 2001, J. Burkhalter 17811 (FSU, UWFP).

New to the Florida panhandle.

Coccinia grandis (L.) Voigt.—Escambia Co.: vacant lot on Cervantes St, Pensacola, 16 Sep 2005, J. Burkhalter

20502 (UWFP), 16 Oct 2005, J. Burkhalter 20541 (FSU, UWFP). New to Florida panhandle..

Cuscuta obtusiflora HBK var. glandulosa Engelm.—Jackson Co.: limestone glade (Williams no. 2), NW

of Marianna, 21 Jul 2005, Anderson 21197. New to the Florida panhandle.

Emilia sonchifolia (L.) DC.—Leon Co.: bordering Call St on FSU campus, Tallahassee, 17 Nov 2006,

Anderson 22642; by Kosha Lab on campus, 27 Dec 2006, Anderson 22708. New to north Florida. Flowers of

these population were exserted 2 mm from the involucre, thus the plants were not distinguishable from E.

fosbergii Nicolson (which is also on campus) when using the Wunderlin and Hansen (2003) key. Distinguishing

features (lyrate leaves and much shorter corolla lobes) as listed by Nicolson (1975 ) assured proper identifica-

tion; also this species has lilac corollas, whereas E. fosbergii corollas are salmon to red-orange in color.

J. Bot. Res. Inst. Texas 1(1): 741 — 751. 2007

742 Journal of the Botanical R h Institute of Texas 1(1)

Euphorbia graminea Jacq.—Escambia Co.: near Pensacola (6709 Pensacola Blvd), 18 Dec 2004, J. Burkhalter

20223 (FSU, UWFP); Leon Co.: 3606 Maclay Blvd, Tallahassee, 17 Nov 2006, Anderson 22643. New to the

Florida panhandle.

Hydrocotyle bowlesioides Mathias & Constance.—Leon Co.: 3606 Maclay Blvd, Tallahassee, 17 Nov

2006, Anderson 22645. New to Florida and second report for the United States. The first known occurrence

of this species in the United States was from southern Georgia (Anderson 1983). I monitored the Georgia

population; it gradually died out by the early 1990's. Erection of the building and its associated landscaping

for the Florida site occurred in 2003, so the Georgia population was not the source for this introduction

into Florida.

Hyptis verticillata Jacq.— Bay Co.: S of Hwy 98 beside Liddon St., Panama City, 14 Sep 2005; J.R. Burkhalter

20490 (FSU, UWFP). New to the Florida panhandle.

Kalanchoe daigremontiana Raym.-Hamet & H. Perrier.—Wakulla Co.: several plants at Rte 365 bridge

over Wakulla River, 5 Jan 2006, Anderson 21525. New to Florida panhandle.

Momordica charantia L.—Leon Co.: St. Augustine Rd, ESE of Tallahassee, 25 Oct 2005, Anderson 21467.

This population dies back each winter but increases in area coverage every following year. New to the

Florida panhandle.

Ottelia alismoides (L.) Pers.—Escambia Co.: 11 Aug 2005, J. Burkhalter 20479 (FSU, UWFP). First observed

here in 2003 by J. Van Dyke of Florida Dept. Environmental Protection; Burkhalter (letter of 15 Dec 05)

reported this invasive species occurs in *uncountable thousands" and has taken over Thompson's Bayou

and Ferry Pass Bayou of the Escambia River delta. New to Florida; previously known only in the United

States in Louisiana and California.

Ranunculus laxicaulis (Torr. & A. Gray) Darby.—Leon Co.: locally abundant in meadow N side of Hwy

90 (2.5 mi ENE of Baum community), 19 Apr 2006, Anderson 21680, 22 Apr 2006, Anderson 21711. New to

Florida. Wunderlin and Hansen (2003) list the species as excluded from the state (specimens of R. pusillus

misapplied with this name). My specimens key to R. flaxicaulis; the flowers have 5-6 petals which are 5-5.5

mm long, whereas petals are fewer and much smaller in flowers of R. pusillus Poir. Whittemore 1997).

Ruellia ciliatiflora Hooker.—Leon Co.: North Monroe St (Hwy 27) at Old Bainbridge Rd, Tallahassee, 20

Nov 2004, Anderson 20957. New to the Florida panhandle.

Solidago gigantea Ait —Liberty Co.: mesic woodland bordering Apalachicola River at Estiffanulga, 13 Sep

2001, Anderson 19945. New to Florida; Semple and Cook (2006) noted this collection was the only one they

saw from Florida—all other Florida collections so named were S. leavenworthii Torrey & A. Gray.

Symphyotrichum kralii Nesom.—Santa Rosa Co.: edge of lake at Camp Paquette, 27 Oct 1995, Anderson

16036. This population plus some from Karick Lake and Krull Lake had been identified as Aster laeve L. var.

concinnum Willd. by A. G. Jones and were the basis of reporting that species in Florida (Anderson 2000), but

I was uncomfortable with that determination. Guy Nesom recently identified a duplicate of Anderson 16036

(BRIT) as S. kralii. Semple (pers. comm.) suggests S. kralii may be included in S. dumosum (L.) Nesom or S.

simmondsii (Small) Nesom, and Wunderlin and Hansen (2003) include it in the latter, but these specimens

look like neither of those species as I understand them and are possibly new to Florida. Obviously, the

taxonomy of the asters is still a work in progress.

Symphyotrichum laeve (L.) Love & Love var. concinnum (Willd.) Nesom.—Jackson Co.: edge of lime-

stone glade, 30 Sep and 20 Oct 2005, Anderson 21385 and Anderson 21461. Duplicates of these specimens

were identified by J. Semple (WAT), so this taxon remains in the flora of Florida, but its presence is based

on these more recently discovered, different plants.

Tradescantia pallida (Rose) D.R. Hunt—Leon Co.: naturalized in woodland remnant in N part of Tal-

lahassee, 31 Jan 2006, Anderson 21530. New to the Florida panhandle.

Tridax procumbens L.—Leon Co.: edges of lawn adjacent to Richards Building, F.S.U. campus, Tallahas-

see, 18 Nov 2005, Anderson 21514, many plants in cracks of sidewalk and parking area at Nuclear Research

Building, F.S.U. campus, Tallahassee, 13 May 2006, Anderson 21803. New to the Florida panhandle.

Anderson, Noteworthy plants from north Florida 743

Zinnia violacea Cav.—Jackson Co.: frequent along Bumpnose Rd bordering open pasture, 2.2 mi N of Hwy

90 (Marianna), 20 Oct 2005, Anderson 21465. New to the Florida panhandle.

ADDITIONAL RANGE EXTENSIONS

Some of the following are county records that “fill in” gaps (and thus were expected occurrences); they are

listed without additional notation. Other collections bear significance worthy of further comment.

Acer saccharinum L.— Calhoun Co.: disturbed mesic woodland along Apalachicola River, SE of Blount-

stown, 20 Mar 2006, Anderson 21575.

Aesculus pavia L.—Calhoun Co.: wooded calcareous slopes bordering Chipola River WSW of Altha, 12

Apr 2006, Anderson 21639.

Agalinis linifolia (Nuttall) Britton.—Leon Co.: moist depression beside Old Plank Rd, SE of Tallahassee,

25 Oct 2005, Anderson 21474.

Agrostis perennans (Walter) Tuck.—Calhoun Co.: wooded calcareous slopes bordering Chipola River

WSW of Altha, 12 Apr 2006, Anderson 21647.

Albizia julibrissin Durazz.— Calhoun Co.: roadside ditch, Blountstown, 24 May 2006, T. MacClendon 460.

Ambrosia artemisiifolia |..—Calhoun Co.: Myers Dairy Rd, SW side of Blountstown, 27 Sep 2006, T.

MacClendon 524.

Amsonia taberaemontana Walter.—Calhoun Co.: wooded calcareous slopes bordering Chipola River

WSW of Altha, 12 Apr 2006, Anderson 21646.

Arenaria serpyllifolia | —Calhoun Co.: disturbed roadside near landing at Apalachicola River, SE of

Blountstown, 20 Mar 2006, Anderson 21580.

Aeschynomene americana L.—Gulf Co.: under Overstreet bridge, 11 Nov 2006, Anderson 22625.

Aureolaria flava (L.) Farw.—Gadsden Co.: edge of Brickyard Glade, 23 May 2006 (atypical phenology),

Anderson 21869.

Brickellia eupatorioides (L.) Shinners.—Gadsden Co.: hillside glade, 0.8 mi N of Dolan Rd, 8 Nov 2005,

Anderson 21504.

Bromus commutatus Schrad.—Bay Co.: East Panama City Beach, 14 May 1996, Keppner s. n.; Jackson

Co.: roadside depression along Rte 167 near Dry Creek, 12 May 2006, Anderson 21781; Leon Co.: Tallahas-

see, 23 Apr 1984, Anderson 7075; N of Tallahassee, 14 May 1984, Godfrey 81299; Hwy 90 roadside 0.8 mi

E of Magnolia Rd, 19 Apr 2006, Anderson 21686 (the two 1984 collections were originally identified as B.

japonicus Thunb, but later annotated by L. E. Pavlich as B. commutatus).

Calyptocarpus vialis Less.—Jefferson Co.: moist shaded loam N side of Hwy 90 at Lake Miccosukee boat

landing, 19 Apr 2006, Anderson 21665.

Campsis radicans (L.) Seem. ex Bureau—Calhoun Co.: Ocheesee Landing on Apalachicola River, 9

Nov 2006, MacClendon 542.

Carex alata Torr—Wakulla Co.: hydric hammock at McBride Slough near Rte 267, 26 May 2006, Anderson

21871.

Carex aureolensis Mack.—Jefferson Co.: Lake Miccosukee boat landing by Hwy 90, 19 Apr 2006, Anderson

21671, 2 Aug 2006, Anderson 22160. Wunderlin and Hansen (2003) do not list this species, but may have

included it under C. frankii Kunth. Ford and Reznicek (2002) consider the two species distinct with only C.

aureolensis occurring in Florida—either way this is a new county record.

Carex chapmanii Steudel.—Gadsden Co.: Flat Creek, W of Rte 269, 20 Apr 2006, Anderson 21741; Jackson

Co.: mesic woodland near boat ramp, Three Rivers State Park, 20 May 2006, Anderson 21833.

Carex cherokeensis Schwein.— Calhoun Co.: calcareous mesic woodland along Apalachicola River, SE of

Blountstown, 20 Mar 2006, Anderson 21553.

Carex dasycarpa Muhl.—Calhoun Co.: sandy, wooded calcareous slopes bordering Chipola River WSW

of Altha, 12 Apr 2006, Anderson 21640.

744 Journal of the Botanical R h Institute of Texas 1(1)

Carex festucacea Schkuhr ex Willd.—Wakulla Co.: hydric hammock, St. Marks Natl Wildlife Refuge

headquarters, 26 May 2006, Anderson 21879.

Carex fissa Mack. var. aristata FJ. Herm.—Calhoun Co.: 0.5 mi E of Chipola River beside Rte 20, 5 May

2003, Anderson 20698.

Carex gigantea Rudge.—Jackson Co.: shallow waters of Dry Creek at Rte 167 bridge, 12 May 2006, An-

derson 21783.

Carex kraliana Naczi & Bryson.—Jackson Co.: wooded edge of limestone glade NW of Marianna, 25 Mar

2006, Anderson 21594. This rare species is also found in Gadsden and Leon counties in Florida.

Carex leavenworthii Dewey.—Calhoun Co.: disturbed mesic woodland along Apalachicola River, SE

of Blountstown, 20 Mar 2006, Anderson 21576. Ball in Flora of North America (2002) lists this species as

distinct from C. cephalophora Muhl. ex Willd., but Wunderlin and Hansen (2004) list the two species under

the latter in their atlas. Either way, it is a new county record.

Carex leptalea Wahlenb.—Jackson Co.: hydric hammock W of Rte 73 along N side of Dry Creek, 12 May

2006, Anderson 21790.

Carex muhlenbergii Schkuhr.—Gadsden Co.: Booth Hill, W end of Dolan Rd, 16 Apr 2003, Anderson

20679.

Carex oxylepis Torr. @ Hook.—glade like opening (Brooks # 3) in woods NW of Marianna, 2 Jun 2006,

Anderson 21926.

Carex thornei Naczi.—Leon Co.: Lafayette Heritage Trail Park, Tallahassee, 2 Apr 2003, Anderson 20615.

This rare species is also found in Gadsden and Liberty counties in Florida.

Carex typhina Michx.—Calhoun Co.: mesic hardwoods on low bluff along Apalachicola River, SE of

Blountstown, 8 Jun 2006, Anderson 2966.

Carex venusta Dewy.— Gadsden Co.: seepage below Powerline Glade, 23 May 2006, Anderson 21843.

Centella erecta (L. f.) Fern.—Calhoun Co.: open firelane beside ti-ti thicket, ca. 7.8 air mi WSW of Altha,

5 Jun 2006, Anderson 21956. Wunderlin and Hansen (2003) list this species as C. asiatica (L.) Urb. Years

ago, Lincoln Constance told me C. asiatica (a diploid) does not occur in the continental United States and

that ours is C. erecta (a tetraploid). Either way, the species is new to the county.

Cercis canadensis L.—Calhoun Co.: disturbed mesic woodland along Apalachicola River, SE of Blount-

stown, 20 Mar 2006, Anderson 21583.

Chaerophyllum tainturieri Hook.—Calhoun Co.: disturbed mesic woodland along Apalachicola River,

SE of Blountstown, 20 Mar 2006, Anderson 21574.

Cinnamomum camphora (L.) Presl.— Calhoun Co.: disturbed mesic woodland along Apalachicola River,

SE of Blountstown, 16 Feb 2006, Anderson 21532.

Cocculus carolinus (L.) DC.—Calhoun Co.: disturbed mesic woodland along Apalachicola River, SE of

Blountstown, 20 Mar 2006, Anderson 21557.

Commelina virginica L.— Calhoun Co.: mesic woodland along Apalachicola River, SE of Blountstown, 4

Apr 2006, Anderson 21620.

Conyza bonariensis (L.) Cronquist.—Calhoun Co.: Myers Dairy Rd, SW side of Blountstown, 17 Aug

2006, T. & K. MacClendon 521.

Croton glandulosus L.—Calhoun Co.: Myers Dairy Rd, SW side of Blountstown, 19 Aug 2006, T. & K.

MacClendon 522.

Cuscuta gronovii Willd. ex Schult. in Roem. & Schult.—Jackson Co.: limestone glade (Williams no. 2),

NW of Marianna, 30 Sep 2005, Anderson 21372.

Cuscuta pentagona Engelm.—Jackson Co.: limestone glade (Williams no. 2), NW of Marianna, 30 Sep

2005, Anderson 21380.

Desmodium ochroleucum M.A. Curtis ex Canby—Jackson Co.: woods at edge of fairway, Marianna

Caverns Golf Course, 30 Sep 2005, Anderson 21360 (this doubles the known sites for this taxon in Florida).

This species is listed as endangered in Florida (Coile & Garland 2003).

Anderson, Noteworthy plants from north Florida 745

Dichanthelium leucothrix (Nash) Freckmann.—Calhoun Co.: ephemeral pond bottom, ca. 5.8 air mi

WSW of Altha, 5 Jun 2006, Anderson 21945.

Dichanthelium oligosanthes (Schult.) Gould.—Jackson Co.: Williams glade # 2 , NW of Marianna, 2 Jun

20056, Anderson 21913.

Dichanthelium strigosum (Muhl. ex Elliott) Freckmann.— Calhoun Co.: fallow field (cutover flatwoods)

along River St in Blountstown, 8 Jun 2006, Anderson 21971; Gadsden Co.: edge of Humphrey Glade between

1-10 and Dolan Rd, 8 Nov 2005, Anderson 21491.

Dioscorea alata L.—Wakulla Co.: locally common on N side Hwy 98 at Newport, 30 Nov 2006, Anderson

22684

Dioscorea bulbifera |..—1 eon Co.: still in my yard in Tallahassee after 30 years of trying to eradicate, 2

Dec 2006, Anderson 22692; Wakulla Co.: frequent on N side Hwy 98 at Newport, 30 Nov 2006, Anderson

220095.

Eleocharis baldwinii (Torr. Chapm.—Calhoun Co.: ephemeral pond bottom, ca. 5.8 air mi WSW of

Altha, 5 Jun 2006, Anderson 21944.

Eleocharis tortilis (Link) Schultes.— Gadsden Co.: shaded floodplain of Flat Creek, 29 Apr 2006, Ander-

son 21728. This species is infrequently collected in the state; three 19th century Florida collections exist at

NY. Two are labeled “Chapman s.n., Florida" and one is “Chapman s.n., Aspalaga, Florida.” Aspalaga is a

historic river landing in Gadsden County, so the current collection confirms the species' continued pres-

ence in the county.

Eragrostis cilianensis (All. Vignolo ex Janch.—Leon Co.: near loading zone behind Conradi Bldg on

Florida State University campus, Tallahassee, 18 Sep 2006, Anderson 22388. This is the third county of

record for the state.

Erigeron annuus (L.) Pres.—Calhoun Co.: W side of Blountstown (3.7 mi E of Rte 71 on Rte 20), 1 Jun

2006, Anderson 21898.

Eryngium prostratum Nutt. ex DC.— Calhoun Co.: W side of Blountstown (3.7 mi E of Rte 71 on Rte 20),

] Jun 2006, Anderson 21897.

Fimbristylis autumnalis (L.) Roem. € Schult.—Calhoun Co.: ephemeral pond bottom, ca. 5.8 air mi

WSW of Altha, 5 Jun 2006, Anderson 21943.

Fragaria virginiana Duchesne.—Jackson Co.: rich calcareous woods, Three Rivers State Park, 4 Apr 1979,

A. K. Gholson 7587; edge of limestone glade (Brooks no. 1), NW of Marianna, 21 Jul 2005, Anderson 21206;

edge of limestone glade (Williams no. 2); 30 Sep 2005, Anderson 21383. This species was previously known

only from Leon County from collections made by R. M. Harper in 1927 (NY), but it has apparently not been

collected in that county since then.

Galium aparinae L.—Calhoun Co.: disturbed mesic woodland along Apalachicola River, SE of Blountstown,

20 Mar 2006, Anderson 21573.

Galium circaezans Michx.—Gadsden Co.: wooded slopes above Apalachicola River near Aspalaga, 29

Apr 2006, Anderson 21745.

Gelsemium sempervirens (L.) Ait. f.—Calhoun Co.: woodland along River St near Apalachicola River, SE

of Blountstown, 20 Mar 2006, Anderson 21581.

Gentiana pennelliana Fern.—L eon Co.: wet savanna in Gully Bay extension of Grand Bay, Apalachicola

National Forest, 11 Dec 2002, Anderson 20552. This species is listed as endangered in Florida (Coile and

Garland 2003).

Glyceria striata (Lam.) Hitchc.—Leon Co.: wet depression in powerline corridor on S side Hwy 90, 0.5

mi W of Lake Miccosukee boat landing, 19 Apr 2006, Anderson 21674.

Gymnopogon ambiguus (Michx.) BSP.—Jackson Co.: edge of limestone glade (Williams no. 2), NW of

Marianna, 30 Sep 2005, Anderson 21382.

Gymnopogon brevifolius Trin.—Gulf Co.: wet flatwoods W of Depot Creek, S of Port St. Joe, 13 Oct

2005, T. Mitchell s.n.

746 Journal of the Botanical R h Institute of Texas 1(1)

Gymnostyles anthemifolia Juss.—Escambia Co.: E. Pensacola Beach, 8 Nov 2005, J. Burkhalter 20341

(FSU, UWFP). New to western panhandle and second record for north Florida.

Hexaletris spicata (Walt.) Barnh.—Gadsden Co.: large population at hunt camp near Aspalaga, 27 Jul

2005, Anderson 21247. This species is listed as endangered in Florida (Coile and Garland 2003).

Hydrocotyle verticillata Thunb. var. triradiata (A. Rich.) Fern.—Jefferson Co.: shaded mud at Lake Mic-

cosukee Hwy 90 boat landing, 2 Aug 2006, Anderson 22162. New county record for this variety.

Hypochaeris glabra L.—Jackson Co.: E of Rte 71 near Rocky Creek, 20 Apr 2006, Anderson 21707, picnic

area, Three Rivers State Park, 20 May 2006, Anderson 21834.

Hypoxis rigida Chapm.—Gadsden Co.: E.B. Glade (recently burned), 23 May 2006, Anderson 21854.

Hypoxis wrightii (Baker) Brackett —Gadsden Co.: E.B. Glade, 23 May 2006, Anderson 21853; Jackson Co.:

Williams #2 Glade, 2 Jun 2006, Anderson 21907; Liberty Co.: N of Rte 20 opposite Rte 2224, 4 Aug 2006,

Anderson 22169.

Ipomoea pandurata (L.) G. Mey.— Calhoun Co.: Abe Springs, SW of Blountstown, 15 Jun 2006, T. & K.

MacClendon 504.

Ipomoea triloba L.—Liberty Co.: Apalachicola River floodplain under Rte 20 bridge, 19 Oct 2005, Anderson

21428. Listed as a Federal Noxious Weed.

Iva angustifolia Nutt. ex DC.—Calhoun Co.: Myers Dairy Rd, Blountstown, 21 Jun 2006, T. & K. Mac-

Clendon 509.

Iva imbricata Walter.—Gulf Co.: sand dunes N end of St. Joseph Peninsula, 17 Sep 2005, Anderson

21320.

Juncus repens Michx.— Calhoun Co.: ephemeral pond bottom, ca. 5.8 air mi WSW of Altha, 5 Jun 2006,

Anderson 21946.

Lamium amplexicaule |..—Calhoun Co.: disturbed mesic woodland along Apalachicola River, SE of

Blountstown, 20 Mar 2006, Anderson 21545; Gadsden Co.: Rte 12 roadside SW of Greensboro, 20 Mar 2006,

Anderson 21535.

Landoltia punctata (G. Mey.) Les € DJ. Crawford.—Jefferson Co.: with Wolffiella in shallow water at Lake

Miccosukee Hwy 90 boat landing, 2 Aug 2006, Anderson 22176.

Lemna valdiviana Phil.— Calhoun Co.: abundant along shaded pond margin, ca. 5.7 air mi WSW of Altha,

5 Jun 2006, Anderson 21957.

Lespedeza procumbens Michx.—Gadsden Co.: N edge of Humphrey Glade between I-10 and Dolan Rd,

8 Nov 2005, Anderson 21480.

Lespedeza repens (L.) Bart.— Calhoun Co.: oak-pine sandhill, ca. 5.8 air mi WSW of Altha, 5 Jun 2006,

Anderson 21940.

Lindernia crustacea (L.) F. Muell.—Liberty Co.: Apalachicola River floodplain under Rte 20 bridge, 28

Sep 2005, Anderson 21353.

Lolium arundinaceum (Schreb.) Darbysh.—Jackson Co.: beside Rte 73 just NW of Marianna, 2 Jun 2006,

Anderson 21899.

Lonicera sempervirens L.—Calhoun Co.: woods bordering Chipola River WSW of Altha, 12 Apr 2006,

Anderson 21664.

Ludwigia glandulosa Walt.— Calhoun Co.: fallow field (cutover flatwoods) along River St in Blountstown,

8 Jun 2006, Anderson 21973.

Ludwigia leptocarpa (Nutt.) Hara.— Calhoun Co.: Ocheesee Landing on Apalachicola River, 9 Nov 2006,

MacClendon 541.

Malvastrum coromandelianum (L.) Garcke.— Calhoun Co.: disturbed edge of mesic woodland along

Apalachicola River, SE of Blountstown, 8 Jun 2006, Anderson 21962.

Matelea floridana (Vail) Woodson.—Jackson Co.: both flowering and fruiting, near E entrance of Florida

Caverns State Park, 27 Jul 2005, Anderson 21218; W side of Florida Caverns State Park, 27 Jul 2005, Anderson

21227; both flowering and fruiting, Williams 43 Glade, 2 Jun 2006, Anderson 21921. This species is listed

as endangered in Florida (Coile and Garland 2003).

Anderson, Noteworthy plants from north Florida 747

Medicago arabica (L.) Huds.—Calhoun Co.: disturbed mesic woodland along Apalachicola River, SE of

Blountstown, 20 Mar 2006, Anderson 21570.

Medicago lupulina I..— Calhoun Co.: disturbed mesic woodland along Apalachicola River, SE of Blount-

stown, 8 Jun 2006, Anderson 21963.

Medicago polymorpha L.—Calhoun Co.: disturbed mesic woodland along Apalachicola River, SE of

Blountstown, 20 Mar 2006, Anderson 21540; Jefferson Co.: Lake Miccosukee boat landing, 19 Apr 2006,

Anderson 21670.

Melia azedarach L.—Calhoun Co.: disturbed mesic woodland along Apalachicola River, SE of Blountstown,

20 Mar 2006, Anderson 21559. This species is listed as a Category II invasive exotic (FLEPPC 2005).

Melilotus albus Medik.—Calhoun Co.: roadside ditch, Blountstown, 31 May 2006, T. MacClendon 467;

beside Bear Head Rd., ca. 7.8 mi WSW of Altha, 5 Jun 2006, Anderson 21958.

Micromeria brownei (Sw.) Benth. var. pilosiuscula A. Gray.—Wakulla Co.: Rte 267 roadside by McBride

Slough. 26 May 2006, Anderson 21874.

Mitchella repens L.—Calhoun Co.: woods bordering Chipola River WSW of Altha, 12 Apr 2006, Anderson

21658.

Modiola caroliniana (L.) G. Don.— Calhoun Co.: disturbed mesic woodland along Apalachicola River, SE

of Blountstown, 20 Mar 2006, Anderson 21550.

Muhlenbergia schreberi J. F. Gmel.—Gadsden Co.: wooded edge of E.B. Glade, 23 May 2006, Anderson 21849.

Murdannia keisak (Hask.) Handel-Mazz.—Gadsden Co.: shaded wet shoals in Flat Creek, 29 Apr 2006,

Anderson 21740.

Murdannia nudiflora (L.) Brenan.— Calhoun Co.: roadside near Apalachicola River boat landing, SE side

of Blountstown, 19 Oct 2005, Anderson 21436.

Myosotis macrosperma Engelm.—Calhoun Co.: disturbed mesic woodland along Apalachicola River, SE

of Blountstown, 20 Mar 2006, Anderson 21543.

Nemophila aphylla (L.) Brummitt.— Calhoun Co.: disturbed mesic woodland along Apalachicola River,

SE of Blountstown, 16 Feb 2006, Anderson 21533, 20 Mar 2006, Anderson 21576; fourth county of record

and southernmost population in the state.

Oxalis debilis Kunth var. corymbosa (DC.) Lourteig.— Calhoun Co.: wooded floodplain of Apalachicola

River, SE side of Blountstown, 1 Feb 2005, Anderson 20958.

Oxycaryum cubense (Poeppig & Kunth) Lye.—Jackson Co.: inflorescences monocephalous, shore of Lake

Seminole at boat ramp, Three Rivers State Park, 20 May 2006, Anderson 21830. This species is “morpho-

logically plastic” (Bruhl 2002); the typical form is polycephalous, but most plants in the panhandle have

compactly monocephalous inflorescences and can be referred to as O. cubense forma paraguayense (Maury)

T. M. Pedersen. Wunderlin and Hansen (2003) list this species as Scirpus cubensis Poeppig & Kunth. (either

way new to the county).

Packera glabella (Poir.) C. Jeffrey.— Calhoun Co.: disturbed mesic woodland along Apalachicola River, SE

of Blountstown, 20 Mar 2006, Anderson 21548.

Parthenium hysterophorus L.— Calhoun Co.: disturbed mesic woodland along Apalachicola River, SE of

Blountstown, 20 Mar 2006, Anderson 21541.

Pediomelum canescens (Michx.) Rydb.—Calhoun Co.: oak-pine sandhill, ca. 5.8 air mi WSW of Altha,

5 Jun 2006, Anderson 21941.

Phalaris angusta Nees ex Trin.—Jackson Co.: roadside ditch, W of Chipola River, 1.3 mi S of Rte 278 on

Rte 73, 12 May 2006, Anderson 21800; Leon Co.: Conner Creek in Lafayette Heritage Trail Park, Tallahas-

see, 1 Jun 2002, Anderson 20254, wet depression along powerline S of Hwy 90 near Jefferson County line,

19 Apr 2006, Anderson 21679.

Phoradendron leucarpum (Raf.) Reveal & M. C. Johnst.— Calhoun Co.: in trees along River St. in Blount-

stown, 4 Apr 2006, Anderson 21627.

Physalis angulata L —Calhoun Co.: with P. heterophylla and P. pubescens in fallow field (cutover flatwoods)

along River St. in Blountstown, 8 Jun 2006, Anderson 21972.

748 Journal of the Botanical R h Institute of Texas 1(1)

Physalis pubescens L.—Calhoun Co.: fallow field (cutover flatwoods) along River St. in Blountstown, 8

Jun 2006, Anderson 21969.

Piriqueta cistoides (L.) Griseb. subsp. caroliniana (Walt.) Arbo.—Gadsden Co.: E.B. Glade, 23 May

2006, Anderson 21851.

Pityopsis flexuosa (Nash) Small.—Jefferson Co.: ca. 2.1 mi ESE of Wacissa, 27 Oct 05, Nee

endemic species is listed as endangered in Florida (Coile and Garland 2003).

0502. This

Plantago rugelii Decne.—Jefferson Co.: shaded mud at Lake Miccosukee Hwy 90 boat landing, 2 Aug 2006,

Anderson 22159. Third county of record in the state for this species (and best looking population).

Plantago wrightiana Decne.—Jackson Co.: drying shoulder of Rte 71 at Rocky Creek bridge, 12 May

2006, Anderson 21806.

Platanus occidentalis L.—Calhoun Co.: disturbed mesic woodland along Apalachicola River, SE of

Blountstown, 20 Mar 2006, Anderson 21543.

Pluchea baccharis (Millet) Pruski.—Calhoun Co.: roadside ditch, Blountstown, 12 Apr 2006, T. Mac-

Clendon 466. This species is listed as P. rosea Godfrey by Wunderlin and Hansen (2003), but Nesom (2006)

listed that species as a synonym for the earlier name.

Poa annua L.—Calhoun Co.: disturbed mesic woodland along Apalachicola River, SE of Blountstown, 20

Mar 2006, Anderson 21576.

Polygonum hirsutum Walt.— Calhoun Co.: ephemeral pond bottom, ca. 5.8 air mi WSW of Altha, 5 Jun

2006, Anderson 21949.

Prunus caroliniana (Mill.) Ait.—Calhoun Co.: disturbed mesic woodland along Apalachicola River, SE of

Blountstown, 16 Feb 2006, Anderson 21584.

Prunus persica (L.) Batsch—Calhoun Co.: disturbed mesic woodland along Apalachicola River, SE of

Blountstown, 16 Feb 2006, Anderson 21531.

Pteris multifida Poir—Gadsden Co.: limestone outcrop in maple, beech, magnolia, oak woodland above

Apalachicola River floodplain near Aspalaga, 27 Jul 2005, Anderson 21256.

Pycnanthemum albescens T. & G. ex A. Gray.—Jefferson Co.: 4.5 mi W of Wacissa, 31 Oct 2005, Neel 0504.

Pycnanthemum setosum Nuttall.—]Jackson Co.: limestone glade (Williams no. 3), NW of Marianna, 21 Jul

2005, Anderson 21176; limestone glade (Williams no. 2), NW of Marianna, 2 Jun 2006, Anderson 21912.

Ranunculus abortivus 1.—Calhoun Co.: disturbed mesic woodland along Apalachicola River, SE of

Blountstown, 20 Mar 2006, Anderson 21547.

Ranunculus sardous Crantz.—Jackson Co.: wet roadside depression of Rte 73 at edge of hydric hammock

along Dry Creek, 12 May 2006, Anderson 21788.

Rhynchosia tomentosa Hook. & Arn. var. tomentosa.— Calhoun Co.: oak-pine sandhill, ca. 5.8 air mi

WSW of Altha, 5 Jun 2006, Anderson 21938.

Rhynchospora compressa J. Caery ex Chapm.—Gadsden Co.: Powerline Glade, 23 May 2006, Anderson

21902.

Richardia humistrata (Cham. & Schltdl.) Schult. & Schult. f —]Jackson Co.: edge of fairway, Marianna

Caverns Golf Course, 27 Jul 2005, Anderson 21222. New for county and significant range extension eastward

within the state.

Sagittaria filiformis J. G. Sm.—Calhoun Co.: ephemeral pond bottom, ca. 5.8 air mi WSW of Altha, 5

jun 2006, Anderson 21948.

Sanicula odorata (Raf) Pryer & Phillippe.—Calhoun Co.: shaded mesic woodland along Apalachicola

River, SE of Blountstown, 4 Apr 2006, Anderson 21619.

Sassafras albidum (Nutt.) Nees.—Calhoun Co.: woodland along River St near Apalachicola River, SE of

Blountstown, 20 Mar 2006, Anderson 21582.

Setaria pumila (Poir.) Roem. & Schult.—Calhoun Co.: Abe Springs, Blountstown, 26 Jul 2006, T. & K.

MacClendon 519.

Sida elliottii Torr. & A. Gray.—Jackson Co.: limestone glade (Williams no. 3), NW of Marianna, 21 Jul

Anderson, Noteworthy plants from north Florida 749

2005, Anderson 21179; limestone glade (Williams no. 2), 30 Sep 2005, Anderson 21373; limestone glade W

of Rte 73, NW of Marianna, 20 Oct 2005, Anderson 21453.

Silene regia Sims.—Jackson Co.: both flowering and fruiting plants observed, W side of Florida Caverns

State Park, 27 Jul 2005, Anderson 21225. King (1981) noted this species is well established in Missouri and

“elsewhere, the species is rare and widely dispersed, if not extirpated.” Its presence in Florida (Jackson

County) was documented in 1940 by Henry 2312 (ISC, MIN); this new report confirms the rare species is

not extirpated in Florida (certainly a candidate for listing as endangered in Florida).

Smilax ecirrhata Wats.—Calhoun Co.: wooded calcareous slopes bordering Chipola River WSW of Altha,

12 Apr 2006, Anderson 21644.

Smilax lasioneuron Hook.—Calhoun Co.: wooded calcareous slopes bordering Chipola River WSW of

Altha, 12 Apr 2006, Anderson 21648.

Sonchus asper (L.) Hill.—Calhoun Co.: roadside weed beside Rte 20 on E side of Blountstown, 20 Mar

2006, Anderson 21585.

Spermacoce densiflora (DC.) Alain.—Calhoun Co.: abundant along roadsides near Apalachicola River

boat landing, SE side of Blountstown, 19 Oct 2005, Anderson 21431.

Spiranthes odorata (Nuttall) Lindley. —Gadsden Co.: partially shaded edge of Humphrey Glade between I-10

and Dolan Rd, 8 Nov 2005, Anderson 21485; Jefferson Co.: 7 air mi SE of Wacissa, 31 Oct 2005, H. Neel 0503.

Sporobolus vaginiflorus (Torr. ex A. Gray) A. W. Wood.—Jackson Co.: limestone glade (Williams no.

2), NW of Marianna, 30 Sep 2005, Anderson 21390; limestone glades (Brooks no. 1 and 4), 20 Oct 2005,

Anderson 21445, Anderson 21449; Bumpnose glade, N of Marianna, 20 Oct 2005, Anderson 21463. Previously

known only to Gadsden County in Florida.

Symphyotrichum fontinale (Alexander) G.L. Nesom.—Gadsden Co.: between Humphrey Glade and Dolan

Rd, 8 Nov 2005, Anderson 21495.

Symphyotrichum shortii (Lindl.) G.L. Nesom.—Jackson Co.: limestone glade (Brooks no. 1), NW of Mari-

anna, 20 Oct 2005, Anderson 21440. This aster was previously known in Florida only in Gadsden County .

Symphyotrichum simmondsii (Small) G.L. Nesom.— Calhoun Co.: Ocheesee Landing on

Apalachicola River, 9 Nov 2006, MacClendon 538; Franklin Co.: Bloody Bluff, Apalachicola River, 15 Nov

1984, Anderson 7776, Hickory Landing, Owl Creek, 4 Dec 1984, Anderson 7816, St. Vincent Island, 25 Oct

1985, Anderson 8993, Van Horn Landing, Apalachicola River, 24 Jan 1986, Anderson 9091; Jefferson Co.:

Wacissa Springs, 31 Oct 2002, Anderson 20505; Leon Co.: wet edge of stormwater pond N of Tallahassee, 30

Nov 2005, Anderson 21521; Madison Co.: lakeside in Greenville, 19 Dec 1986, Anderson 10271; Wakulla Co.:

McBride Slough, 1 Nov 1984, Anderson 7765, ditch near St. Marks post office, 5 Nov 1985, Godfrey 81961.

Tilia americana L. var. caroliniana (Mill.) Castigl.—Calhoun Co.: wooded calcareous slopes bordering

Chipola River WSW of Altha, 12 Apr 2006, Anderson 21652.

Tillandsia recurvata (L.) L—Leon Co.: epiphytic on Quercus geminata Small and Ulmus parvifolia Jacq. along

edge of commercial parking lot, Raymond Diehl Rd, Tallahassee, 16 Nov 2005, Anderson 21513. Naturalized;

county record and new to central panhandle. These relatively young trees apparently came from nursery

stock in south Florida; the bromeliads are healthy and laden with fruiting branches.

Tradescantia ohiensis Raf.—Gadsden Co.: maple, beech, magnolia, oak woodland near Aspalaga, 27 Jul

2005, Anderson 21245.

Tridens flavus (L.) Hitchc. var. flavus.— Calhoun Co.: roadsides near boat landing, SE side of Blountstown,

19 Oct 2005, Anderson 21432.

Trifolium campestre Schreber.— Calhoun Co.: disturbed mesic woodland along Apalachicola River, SE of

Blountstown, 20 Mar 2006, Anderson 21542.

Trifolium carolinianum Michx.—Calhoun Co.: disturbed mesic woodland along Apalachicola River, SE

of Blountstown, 20 Mar 2006, Anderson 21578.

Trifolium incarnatum 1.—Calhoun Co.: disturbed mesic woodland along Apalachicola River, SE of

Blountstown, 20 Mar 2006, Anderson 21571.

750 Journal of the Botanical R h Institute of Texas 1(1)

Trifolium nigrescens Viv.— Calhoun Co.: disturbed roadside just S of Rte 274 Chipola River bridge SW

of Altha, 12 Apr 2006, Anderson 21660.

Urena lobata L.—Leon Co.: street side in Tallahassee, 8 Oct 2005, Anderson 21427. County record and

new to central panhandle. This species is listed as a Category II exotic invasive species in Florida (FLEPPC

2005).

Verbena officinalis L. subsp. halei (Small) Barber.—Calhoun Co.: disturbed roadside just S of Rte 274

Chipola River bridge SW of Altha, 12 Apr 2006, Anderson 21661.

Veronica arvensis L.—Calhoun Co.: disturbed mesic woodland along Apalachicola River, SE of Blount-

stown, 16 Feb 2006, Anderson 21534.

Veronica peregrina L.—Calhoun Co.: disturbed mesic woodland along Apalachicola River, SE of Blount-

stown, 20 Mar 2006, Anderson 21567.

Veronica persica Poir.—Gadsden Co.: Rte 12 roadside SW of Greensboro, 20 Mar 2006, Anderson 21537.

Viburnum obovatum Walter.— Calhoun Co.: disturbed mesic woodland along Apalachicola River, SE of

Blountstown, 20 Mar 2006, Anderson 21562.

Vicia minutiflora Dietr.— Calhoun Co.: disturbed mesic woodland along Apalachicola River, SE of Blount-

stown, 20 Mar 2006, Anderson 21560.

Vicia sativa L.—Calhoun Co.: disturbed mesic woodland along Apalachicola River, SE of Blountstown,

20 Mar 2006, Anderson 21571.

Vicia tetrasperma (L.) Schreb.—Jackson Co.: calcareous loam of disturbed site, W edge of Florida Caverns

State Park, N of Marianna, 25 Mar 2006, Anderson 21614.

Wisteria sinensis (Sims) Sweet.—Calhoun Co.: woodland along River St near Apalachicola River, SE of

Blountstown, 20 Mar 2006, Anderson 21579.

Wolffiella gladiata (Hegelm.) Hegelm.—Jefferson Co.: shallow water at Lake Miccosukee Hwy 90 boat

landing, 2 Aug 2006, Anderson 22163.

Xanthium strumarium L. var. glabratum (DC.) Cronquist.—Calhoun Co.: roadside near boat landing,

SE side of Blountstown, 19 Oct 2005, Anderson 21434.

Youngia japonica L.— Calhoun Co.: disturbed mesic woodland along Apalachicola River, SE of Blount-

stown, 20 Mar 2006, Anderson 21577.

Yucca aloifolia (L.) DC.—Jackson Co.: Williams #2 Glade, 2 Jun 2006, Anderson 21910.

Zizia aptera (A. Gray) Fernald.— Calhoun Co.: wooded calcareous slopes bordering Chipola River WSW

of Altha, 12 Apr 2006, Anderson 21642. Second known site for the state of Florida.

Knowledge on the composition and distribution of the state’s flora is still incomplete. The large number

of reports given here for Calhoun County of relatively common species (plus some rare ones) reinforces

the observation (see Moerman & Estabrook 2006) that generally the further a county is from a university

in the state, the less well-known is its flora [e.g., Leon County (home of Florida State University) had 1664

species listed by Wunderlin and Hansen (2004) prior to this paper; Alachua County (home of University of

Florida) had 1535; Calhoun County had 911 prior to this paper; and Lafayette County (equidistant between

FSU and UF) had the fewest with 423 species listed].

Emphasis on Calhoun County has been twofold. Anderson made frequent visits to his dentist in Blount-

stown and then collected while in the county. Travis and Karen MacClendon live in the county and have begun

a “Calhoun County Herbarium" with support from the county officials who donated herbarium supplies. This

grassroots effort will provide a valuable plant identification resource for interested local citizenry.

The high number of first reports for Gadsden and Jackson counties is one result of the ongoing extensive

survey of the limestone glades (unique within the state) by Wilson Baker and Loran Anderson.

ACKNOWLEDGMENTS

Bill and Pam Anderson, Wilson Baker, James Burkhalter, the MacClendons, and Harry Neel contributed

collections or made localites known to the author. Charles Bryson provided help with identifications with

some sedges; Paul Berry and Michael Huft did so with the somewhat atypical spurge from Leon County;

Anderson, Noteworthy plants from north Florida 751

and Guy Nesom and John Semple did so with some asters. NY provided a virtual herbarium loan. Elizabeth

A. Jensen and Elaine M. Norman provided valuable reviews of the manuscript.

REFERENCES

ANDERSON, L. C. 1983. Hydrocotyle bowlesioides in Georgia-new to United States. Castanea 48:317.

ANDERSON, L. C. 1984. Noteworthy plants from north Florida. Sida 10:295-297.

ANDERSON, L. C. 1986. Noteworthy plants from north Florida. Il. Sida 11:379-384.

ANDERSON, L. C. 1988. Noteworthy plants from north Florida. Ill. Sida 13:93-100.

ANDERSON, L. C. 1989. Noteworthy plants from north Florida. IV. Sida 13:497-504,

ANDERSON, L. C. 1991. Noteworthy plants from north Florida. V. Sida 14:467-474.

ANDERSON, L. C. 1995. Noteworthy plants from north Florida. VI. Sida 16:581—587.

ANDERSON, L. C. 2000. Noteworthy plants from north Florida. VII. Sida 19:211-216.

Austin. D. F. 1980. Studies of the Florida Convolvulaceae. ll. Cuscuta. Florida Scientist 43:294-302.

BALL, PW. 2002. Carex sect. Phaestoglochin. Flora N. Amer. 23:285-297.

Brun, J.J. 2002. The genus Oxycaryum. Flora N. Amer. 23:140.

CiEwtLL, A.F. 1985. Guide to the vascular plants of the Florida panhandle. Univ. Presses of Florida, Tallahassee.

Core, N.C. and M.A. GanLAND. 2003. Notes on Florida s endangered and threatened plants. Florida Department of Ag-

riculture & Consumer Serv., Division of Plant Industry-Botany Section Contrib. No. 38, 4th ed., Gainesville.

Cronauist, A. 1980. Vascular Flora of the Southeastern United States. |. Asteraceae. Univ. North Carolina Press,

Chapel Hill.

FLEPPC. 2005. Florida's most invasive plants list. Florida Exotic Pest Plant Council. http://www/fleppc.org.

Fonp, B.A. and A.A. REzNicEk. 2002. Carex sect. Squarrosae. Flora of N. Am. 23:518-519.

Kine, C.C. 1981. Distribution of royal catchfly (Silene regia) with special reference to Ohio populations. Ohio Biol.

Surv. Biol. Notes 15:131-141.

Moerman, D.E. and G.F. EsragRook. 2006. The botanist effect: counties with maximal species richness tend to be

home to universities and botanists. J. Biogeogr. 33:1969-1974.

NESOM, G.L. 2006. The genus Pluchea. Flora N. Amer. 19:478-484.

Nicotson, D.H. 1975. Emilia fosbergii, a new species. Phytologia 32:33-34.

SEMPLE, J.C. 2006. The genus Bradburia. Flora N. Amer. 20:211-212.

SEMPLE, J.C and R.E. Cook. 2006. The genus Solidago. Flora N. Amer. 20:107-166.

WHITTEMORE, A. T. 1997. The genus Ranunculus. Flora N. Amer. 3:88-135.

WUNDERLIN, R.P. and B.F. Hansen. 2003. Guide to the vascular plants of Florida, 2nd ed. Univ. Press of Florida,

Gainesville.

WUNDERLIN, R.P. and B.F. HANSEN. 2004. Atlas of Florida vascular plants. http://www.usf.edu/~isb/projects/atlas/atlas.

html.

752 Journal of the Botanical R h Institute of Texas 1(1)

BOOK NOTICES

PAMELA A. Matson, ASHOK GADGIL, and Dante. M. Kammen (eds.). 2006. Annual Review of Environment and

Resources, Vol. 31, 2006. (ISBN 0-8243-2331-9, hbk.). Annual Reviews, 4139 El Camino Way, PO

Box 10139, Palo Alto, CA 94303-0139, U.S.A. (Orders: www.annualreviews.org, service@annualre-

views.org, 650-493-4400, 650-424-0910 or 650-855-9815 fax, 1-800-523-8635). $85.00, 494 pp.,

color figures, 614" x 914".

This volume *will be a useful resource for researches and practitioners working on nature-society interactions who want and ought to

know the current state of affairs on the topics reviewed, but who do not have the time to cover the individual articles in each of the dozen

or so high-impact journals that would need to be read to keep up to date." The individual reviews are divided into four categories.

I. Earth's Life Support Systems

1) Abrupt change in earth's climate system. 2) Earth's cryosphere: current state and recent changes. 3) Integrated regional changes in

arctic climate feedbacks: implications for the global climate system. 4) Global marine biodiversity trends. 5) Biodiversity conservation

planning tools: present status and challenges for the future.

II. Human Use of Environment and Resources

6) Energy efficiency policies: a retrospectiveg examination. 7) Energy-technology innovation. 8) Water markets and trading. 9)

Bio technology in agriculture.

III. Management, Guidance, and Governance of Resources and Environment

10) Environmental governance. 11) Neoliberalism and the environment in Latin America. 12) Assessing the vulnerability of social-

environmental systems. 13) Environment and security. 14) Sustainability values, attitudes, and behaviors: a review of multinational

and global trends.

IV. Integrative Themes

15) Linking knowledge and action for sustainable development.

— Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A.

DIANTHA STATES and Jack States. 2004. Wildflowers of Wyoming. (ISBN 0-87842-496-2, pbk.). Mountain

Press Publishing Company, Box 2399, Missoula, MT 59806, U.S.A. $20.00. (Orders: www.mountain-

press.com, info@mtnpress.com, 406-728-1900, 406-728-1635 fax). $19.00, 254 pp., 5!^" x 81^".

Both of the authors of this handy field guide are experienced botanists. They begin the book with a concise description of the varied

biomes of Wyoming: alpine, subalpine, montane, foothills, steppe, and plains, noting the dominant plants and common wildflowers in

each. Following a key to plant families, there are plates of postage stamp- size photographs of the flowers arranged by color, by corolla

form, and other distinguishing features. The page number leads the user to the text. Arrangement is by plant family. Each species is

described with blooming period, growth habitat, and general | | 1. A photograph of each species is on the opposite page. For

the most part, the photographs are adequate for assured identification.—Joann Karges, (TCU Library, retired), Botanical Research Institute

of Texas, Fort Worth, TX, 76102-4060, U.S.A.

DONALD ANTHONY SCHIEMANN. 2005. Wildflowers of Montana. (ISBN 0-87842-504-7, pbk.). Mountain Press

Publishing Company, Box 2399, Missoula, MT 59806, U.S.A. $20.00. (Orders: www.mountain-press.

com, info@mtnpress.com, 406-728-1900, 406-728-1635 fax). $22.00, 306 pp., 52" x 81^".

Following the introduction describing the general landscape of the state, its climate, and vegetation zones, the author provides hints for

flower identification, floral forms, leaf forms, and terms that are useful in consulting the text. Arranged by plant family, the text clearly

describes each species and any with which it might be confused.

Habitat, range, and significance of the plant are noted. The color photographs are excellent. A “thumb-nail” sketch of photographs

of the flowers by color and form lead even a beginner to the appropriate textual description. An added feature is the appended guide

to flowers of the Bitteroot Region and of Glacier National Park which should be most welcome to visitors of those areas. This is a well-

conceived and executed field guide

Dr. Schiemann earned his doctoral degree in i l Science and Microbiology at the University of North Carolina, Chapel

Hill.—Joann Karges, (TCU Library, retired), REN Research Institute of Texas, Fort Worth, TX, 76102-4060, U.S.A.

J. Bot. Res. Inst. Texas 1(1): 752. 2007

NEW, CORRECTED, AND INTERESTING RECORDS FOR THE

KANSAS VASCULAR FLORA

Caleb A. Morse Craig C.Freeman Ronald L. McGregor

R. L. McGregor Herbarium, Division of Botany

Bi Jiversit Resedi ch Center

University of Kansas, 2045 Constant Avenue

Lawrence, KS 66047, U.S.A.

cmorseaku.edu ccfree@ku.edu

ABSTRACT

Field and herbarium studies have produced dd for 25 vascular plants previously undocumented in Kansas, or known only from

historic records. Three taxa are considered t i d 2? id 1 the results of recent introductions. Non-native

taxa represent 21.7% of the vascular flora of the state.

RESUMEN

Los estudios de campo y de herbario han dado citas de 25 plantas vasculares previamente sin documentar en Kansas, o conocidas

ünicamente de citas históricas. Tres taxa se consideran nativos y 22 son considerados introducciones recientes. Los taxa no nativos

representan 21.796 de la flora vascular del estado.

Continuing field and herbarium studies in the grassland biome of central North American have produced

records for 24 vascular plants previously undocumented in Kansas, and one species known only from a

single, historic collection. Only three taxa are considered to be native occurrences in the state; the remaining

22 (marked with an asterisk below), including 21 species and two interspecific hybrids, are considered the

results of recent introductions. Following the concepts of Richardson et al. (2000), we consider 11 of these

introduced taxa to be casual aliens in Kansas, seven to be naturalized, and four to be invasive in the state.

In reports of floristic novelties for the state since 1998 (Freeman et al. 1998; Freeman 2000; Freeman et al.

2003; Singhurst & Holmes 2005; Barnard 2006) new non-native taxa (52) have outpaced native taxa (33),

which illustrates a trend in our knowledge of the Kansas flora. While the number of vascular taxa known for

the state rose from 1,872 taxa in 1940 (Stuckey & Barkley 1993) to 2,366 in 2007 (Freeman, unpublished

data), the percentage of introduced taxa has increased from 17.496 to 21.796. A similar change in the relative

abundance of non-native taxa has been documented for the Missouri flora by Yatskievych and Raveill (2001),

who found that introduced taxa had increased from 22.896 to 27.796 between 1965 and 2001. We anticipate

that the introduced component of the Kansas flora will continue to increase as a percentage of all taxa known

for the state. These reports update the distribution data contained in Atlas of the Flora of the Great Plains

(Great Plains Flora Association 1977) and Flora of the Great Plains (Great Plains Flora Association 1991). AII

voucher specimens are deposited in the R. L. McGregor Herbarium (KANU), University of Kansas.

*Aethusa cynapium L. (Apiaceae). Though reportedly poisonous, this European native is cultivated in North

America for use as a homeopathic remedy. Fool’s-parsley has been documented from most of the northeast

U.S. and from Alabama (USDA, NRCS 2004). In the Midwest, itis known from Illinois (Mohlenbrock 1986),

Minnesota (Ownbey & Morley 1991) and Wisconsin (Harriman 1972). Early reports of fool's-parsely in Kansas

(Carruth 1873; Carruth 1877; Smyth 1890; Smyth 1892) were not substantiated with voucher specimens,

however, and this species was not treated for the Great Plains by McGregor (1991). Upon reexamination,

a specimen from Miami County reported by Freeman et al. (2003) as Carum carvi L. has proven to be A.

cynapium, where it is considered to be a casual alien.

Carex bulbostylis Mack. (Cyperaceae). Thick-style sedge is native to the south-central U.S. and, in the

Great Plains, has been reported as far north as Oklahoma (Naczi & Bryson 2003). Recent examination of

J. Bot. Res. Inst. Texas 1(1): 753 —761. 2007

754 Journal of the Botanical R h Institute of Texas 1(1)

specimens at KANU confirms the presence of C. bulbostylis in the Chautauqua Hills, in the southeast part

of the state.

Voucher specimen: KANSAS. Woodson Co.: 10 mi N Yates Center, wooded area along Turkey Creek, 18 Jun 1975, Stephens 85266.

Carex corrugata Fernald (Cyperaceae). Native to the southeast U.S., as far north as Illinois, corrugate sedge

was documented from several of the western two tiers of counties in Missouri (Yatskievych 2006) and was

mapped for Kansas by Naczi and Bryson (2003), based on a single, historical specimen at NY (R.F.C. Naczi,

in litt.). However, no vouchers in Kansas herbaria had been verified as belonging to C. corrugata and this

species was not treated for the Great Plains by Kolstad (1991). Examination of historic specimens at KANU

and several recent collections confirm the presence of corrugate sedge in two counties along the eastern

border of the state.

Voucher specimens: KANSAS. Cherokee Co.: 1 mi E, 1 mi S Chetopas, 0.5 mi E Neosho River, roadside ditch, 3 Jun 1964, Harms 2230

[Additional specimen at NY, fide Naczi]; 0.25 mi N, 3 mi E jct of US Hwy 69 & KS Hwy 96 at Crestline, Spring River Wildlife Area,

T33S, R25E, sec 10, NEY4, SE & sec 11, Wi, SW, near 37?10'53", 94*38'59"W, elev 245—265 m, sandstone glade, post-blackjack

oak forest on E-facing slope, and floodplain forest along Spring River, 17 Jun 2003, Morse 9653 [Additional voucher at DOV]. Linn Co.:

3.5 mi S, 3.5 mi E Trading Post, Marais des Cygnes National Wildlife Refuge, T21S, R25E, sec 24, S!5, SW, sec 25, NZ, NW'!ÁA, & sec

26, NE, NE!4, 38.1944—38.1961?N, 94.6131—94.6254°W, elev 780—850 ft, periodically inundated swamp white-pin oak-sycamore

bottomland forest and wet sedge-cordgrass prairie along tributary to Mine Creek, 14 May 2004, Morse 10423 et al. [Additional voucher

at DOV]; 1.75 mi S, 4 mi E Trading Post, Marais des Cygnes National Wildlife Refuge, T21S, R25E, sec 13, NE, NE, 38.2259°N,

94.6124°W, elev 780 ft, bottomland pin oak-hickory-pecan forest remnant along Marais des Cygnes River, 20 May 2004, Morse 10471

& Morse [Additional voucher at DOV].

Tess A

*Cerastium dubium (Bast.) Guépin (Caryophyllaceae). First reported in North America from W in

1973 and from eastern North America in 1986 (Shildneck & Jones 1986), doubtful chickweed is now BOUE

from a handful of states in the Pacific Northwest, Midwest, and southern U.S. (Chester 2000; Belden et al.

2004; USDA, NRCS 2004). Though it has been documented from several of the western tier of counties in

Missouri (Yatskievych 2006), a single collection confirms the presence of this European species in Kansas,

where it is considered to be naturalized.

Voucher specimen: KANSAS. Labette Co.: Parsons, E-central part of town, Marvel Park, T31S, R20E, sec 20, NW 1⁄4. 37.3387°N,

95.2481°W, elev 870—880 ft, mowed, weedy bank of Labette Creek and disturbed ground near parking lot in Marvel Park, 15 Apr 2003,

Freeman 19660.

*Cymbalaria muralis Gaertn. et al. (Scrophulariaceae). Native to the Mediterranean, Kenilworth ivy has

been introduced as an ornamental throughout the northeast U.S. and in several western states (USDA, NRCS

2004). In the Great Plains, C. muralis has been documented in Nebraska and South Dakota. A recent collec-

tion confirms the presence of this species in Kansas. It is considered to be a casual alien in the state.

Voucher specimen: KANSAS. Chase Co.: Cottonwood Falls, N side of town, N end of Main St, on W side of street, T19S, R8E, sec 29,

NE, NEM, 38.3730?N 96.5418? W, elev 1200 ft, cracks in foundations of buildings and in sidewalk, 1 Apr 2006, Freeman 21456.

*Cyperus fuscus L. (Cyperaceae). Introduced to North America from Eurasia, brown galingale is known

from scattered localities throughout the U.S. and southeastern Canada (Gillett 1971; Tucker et al. 2003).

This species has been reported along the Platte River in Douglas and Lincoln counties, Nebraska (Rolfsmeier

1995), and from several counties along the Missouri River in central Missouri (Yatskievych 1999). A large

population was documented in 2003 and 2004 on dry bars along the Kansas side of the Missouri River,

where it is considered to be naturalized.

Voucher specimens: KANSAS. Leavenworth Co.: Ft. Leavenworth, NE part, N side of Sherman Army Airfield, just S of Weston Bend

of the Missouri River, 39°22'48"—23'28"N, 94?54'01"—11"W, elev ca 770 ft, mud flat along Missouri River, 30 Sep 2003, Morse 9815;

Ft. Leavenworth, NE part, N side of Sherman Army Airfield, just S of Weston Bend of the Missouri River, 39°22'39"—41"N, 94°53'26"—

52'50"W, elev 760—770 ft, muddy bar along river, 20 Nov 2003, Morse 9911 et al.; Ft. Leavenworth, E-central part, E side of Sherman

Army Airfield, just S of Weston Bend of the Missouri River, 39.3645—39.3663°N, 94.8966—94.8902°W, elev ca 770 ft, sandy mudflats

and muddy bank of river, 15 Oct 2004, Morse 10871; Ft. Leavenworth, NE part, N side of Sherman Army Airfield, just S of Weston

Bend of the Missouri River, 39.3835°—39.3837°N, 94.9206—94.9290°W, elev 760 ft, sandy mud flats and muddy bank along river, 12

Morse et al., New and ted Is for the K fl 755

Oct 2004, Morse 10858; Ft. Leavenworth; S-central part, Area ca 0.5 mi NE main entrance to Fort and due N of water treatment plant,

39.3421—39.3348°N, 94.9111—94.9172°W, elev 760 ft, mudflats and muddy bank of Missouri River, 21 Oct 2004, Morse 10898.

Dichanthelium laxiflorum (Lam.) Gould (Poaceae). Soft-tufted panicgrass is known from throughout

the southeast U.S. (Freckmann & Lelong 2003) and has been collected along the Kansas-Missouri border

in Newton County, Missouri (Yatskievych 1999). A large population recently was discovered in extreme

southeast Kansas, growing on sandy soil in a post oak-blackjack oak forest.

Voucher specimen: KANSAS. Cherokee Co.: 0.25 mi N, 3 mi E jct of US Hwy 69 & KS Hwy 96 at Crestline, Spring River Wildlife

Area, T33S, R25E, sec 10, NE 1⁄4, SE 1⁄4 and sec 11, WZ, SW, near 37?10'53"N, 94°38'59"W, elev 245-265 m, sandstone glade and

post-blackjack oak forest on E-facing slope, 17 Jun 2003, Morse 9665.

*Euonymus alata (Thunb.) Siebold (Celastraceae). Winged burningbush is locally established throughout

the northeastern U.S. and in Montana (Gleason & Cronquist 1991; USDA, NRCS 2004) and has been re-

ported as *sporadic but becoming more widespread" in Missouri (Yatskievych 2006). In Kansas, this species

is known from scattered stations in the eastern two tiers of counties, where it is considered to be invasive.

It appears to be especially common in white oak-shagbark hickory upland forests in the northeasternmost

part of the state.

Voucher specimens: KANSAS. Cherokee Co.: 1.5 mi E, 1.75 mi S Baxter Springs, Woods S of small stream, old home site in area, 2

Jun 1980, Brooks 14724. Douglas Co.: Lawrence, E side of ri of Kansas Main E T12S, R19E, sec 36, SEM, 38.0576—

38.9620?N, 95.2430—95.2424?W, elev 980—1010 ft, weedy 1] Battenfeld Scholarship Hall & Lilac Ln, 17 Apr 2006, Morse

12672. Leavenworth Co.: Ft. Leavenworth, W-central part, area to N of water reservoir and E of T Hill, 39°21'57"—22'03"N,

94°56'06"—55'47"W, elev 850-1060 ft, mesic oak-hickory-basswood forest on NE-facing slope and hackberry-mulberry floodplain for-

est along tributary to Quarry Creek, 11 Jun 2003, Morse 9497; Ft. Leavenworth; W-central part, area to N and W of National Cemetery,

39°20'59"—21'01"N, 94°56'00"—14"W, elev 920-1010 ft, disturbed oak-hickory-basswood forest on SE-facing slope, 23 Jun 2003,

Morse 9695; Ft. Leavenworth, W-central part, ravine 0.25 mi E of Bell Point, between reservoir and radio tower, 39°21'46"—43'N,

94°55'58"—56'14"W, elev 880-1000 ft, disturbed oak-hickory-basswood forest on moderate E-facing slopes with deep, steep-sided

draws along tributary to Quarry Creek, 18 Jul 2003, Morse 9761; Ft. Leavenworth, W-central part, area to E of Wagner Point and NW of

National Cemetery, 39?21'10"—16"N, 94°56'21"—55'49"W, elev 850-1030 ft, disturbed oak-hickory-basswood forest on steep slopes

above tributary to Quarry Creek and floodplain forest along creek, 31 Oct 2003, Morse 9877. Neosho Co.: 1 mi S of Erie, Centerville

Township, in woods just above Neosho River, near old river bridge, 17 Nov 2004, Holland 10709. Wyandotte Co.: 0.75 mi S, 0.75 mi E

northern jct KS Hwy 5 & 1-435, NW side of Wyandotte Co Lake, T10S, R24E, sec 19, NW14, NW!A, 39.1702°N 94.7867°W, 850-900

ft, disturbed oak-hickory forest on moderate E-facing slope, 9 May 2004, Morse 10384.

*Eriochloa villosa (Thunb.) Kunth (Poaceae). Native to eastern Asia, woolly cup grass is known from

scattered records throughout the U.S. (Shaw et al. 2003; Belden et al. 2004). In the Great Plains, this spe-

cies has been documented in Nebraska (Rolfsmeier et al. 1991) and Iowa, where it has reportedly become a

serious weed of agricultural crops (Stubbendieck et al. 1994; Darbyshire et al. 2003). Though reported for

the state (USDA, NRCS 2004), woolly cup grass was not mapped from Kansas by Shaw et al. (2003) and to

our knowledge a single, recent specimen confirms the presence of E. villosa in northeast part of the state.

Based on its behavior in other states in the Great Plains, it is considered to be invasive.

Voucher specimen: KANSAS. Douglas Co.: 0.5 mi S, 2 mi E Big Springs, T12S, R17E, sec 18, SY, 39.0051?N, 95.4553?W, elev 1100

ft, Conservation Reserve Program field with high graminoid cover and low forb cover; terraced site with weedy, low-lying patches, 14

Jul 2005, Fritts & Castle s.n.

*Fatoua villosa (Thunb.) Nakai (Moraceae). Native to China, mulberry-weed was first reported in North

America from Louisiana in 1964, where it had already been known for about 15 years (Vincent 2004). The

species has subsequently spread through the continental U.S., and is now reported from 30 states and the

District of Columbia (Vincent 2004). It often is reported as a weed of greenhouses and cultivated ground.

In the Great Plains, F. villosa is known from eastern Iowa (Cusick 2002), eastern Missouri (Yatskievych &

Raveill 2001), and Oklahoma (Taylor & Taylor 1981; Taylor et al. 1996), where it has reportedly become

a nuisance. Mulberry-weed is added to the flora of Kansas based on a small population, introduced as a

seed contaminant that has persisted for several years in a flower bed in the eastern part of the state. It is

considered to be a casual alien.

756 Journal of the Botanical R h Institute of Texas 1(1)

Voucher specimen: KANSAS. Douglas Co.: Lawrence, SW side of town at 3410 W 24th Place, T13S, R19E, sec 10, NE, NE%, 38°56'23"N

95?16'49"W, elev 870 ft, flower bed in front of house, 2 Sep 2004, Freeman 20106.

*Ligustrum obtusifolium Sieb. & Zucc. (Oleaceae). Native to Japan, obtuse-leaf privet is naturalized

throughout the northeastern U.S. (USDA, NRCS 2004). Recent fieldwork and examination of specimens

at KANU misidentified as L. vulgare L. have confirmed that this species is naturalized in forested areas of

eastern Kansas, where it is considered to be invasive. Ligustrum vulgare, for its part, is frequently planted in

hedges in eastern Kansas, but has not been verified as an escape.

Voucher specimens: KANSAS. Cherokee Co.: E side of Baxter Springs, Kiwanis Park, along N side of US Hwy 166-400, T35S, R24E,

sec 1, SEY4, NE, 37?01'35"N, 94°43'30"W, elev 790 ft, disturbed floodplain forest edge on W side of Spring River, 13 Apr 2004, Morse

10343 & Roth. Douglas Co.: University of Kansas Campus West, brushy wooded area SW of Bridwell Lab, area undisturbed for 45 years,

6 Nov 1995, McGregor 41206, 11 Jun 1996, McGregor 41211; Baldwin Woods, ca 2 mi N Baldwin City, Kansas Ecological Reserves, Ray

and Eleanor Wall Woods, T14S, R20E, sec 28, S!'ó, NW, elev 990-1100 ft, oak-hickory forest and floodplain forest along Coal Creek,

disturbed right of way of abandoned RR tracks, 10 Aug 1991, Freeman 4028. Leavenworth Co.: 2 mi S, 3 mi W Leavenworth, steep,

wooded bank of small creek, 19 Oct 1967, Stephens 19459; Ft. Leavenworth, W-central part, T8S, R22E, sec 15, SEY4, elev 900-1050

ft, mesic, upland, oak-hickory-maple-basswood forest on slopes SE of Wagner Point, steep draws with few limestone outcrops, 18 May

1995, Freeman 7133; Ft. Leavenworth, E of Wagner Point, T8S, R22E, sec 15, NE!4, SE, elev 950-1050 ft, mesic edge of oak-hickory

forest clearing, 15 Jun 1995, Elliott & Morse 634; Ft. Leavenworth, SE corner along Corral Creek, E and W along Grant Ave, T8S, R22E,

sec 23, SE%, elev 770—800 ft, open banks of creek and wooded riparian area immediately W of Grant Ave, 12 Oct 1995, Freeman 7863;

Ft. Leavenworth, SE corner, T8S, R22E, sec 23, SE%. 39.337 2?N, 94.9247? W, elev 800 ft, disturbed riparian forest along N side of Cor-

ral Creek, 27 May 2003, Freeman 19863; Ft. Leavenworth, NW part, Fort de Cavagnial Picnic Area, 39°22'19"N, 94°55'55"W, elev 1020

ft, weedy edge of oak-hickory-basswood forest and mowed parkland, 11 Jun 2003, Morse 9519; Ft. Leavenworth, W-central part, area

to E of water reservoir, below Hancock Hill, 39?21'57"—22'00"N, 94°56'06"—55'41"W, elev 850—950 ft, mesic oak-hickory-basswood

forest on NE-facing slope and hackberry floodplain forest along tributary to Quarry Creek, 8 Aug 2003, Morse 9792; Ft. Leavenworth,

S-central part, area ca 0.5 mi NE main entrance to Fort and due N of water treatment plant, 39.3421—39.3348°N,94.9111—94.9172°W,

elev 760—850 ft, weedy, early successional woodlands and forests on N and E-facing slopes along Corral Creek, dominated by Carya-

J abel

Quercus and Celtis-Gleditsia-Maclura assocations and exotic understory and mid-successional floodplain forest on E side of RR

tracks, dominated by Acer negundo-A. saccharinum-Salix and Platanus-Populus associations with Carex abundant in understory, 21 Oct

2004, Morse 10888. Neosho Co.: 2 mi W Erie, low woods above Neosho River, 24 Nov 1994, Holland 8228.

*Lonicera xbella Zabel (L. morrowii A. Gray x L. tatarica L., Caprifoliaceae). Pretty honeysuckle is a culti-

vated shrub that has become established in the northeast U.S., New Mexico, and Wyoming (USDA, NRCS

2004). Recent examination of specimens at KANU revealed that plants from extreme northeastern Kansas,

previously misidentified as L. tatarica, represent the first verified records of this fertile hybrid from the state,

where it is considered to be invasive. Interestingly, in examining specimens of the Lonicera tatarica complex

sensu Green (1966) for his treatment in the Flora of Missouri, Whittemore (2006) did not find material refer-

able to either parent species occurring outside cultivation in that state. However, L. xbella is known from

scattered localities, primarily through the eastern half of Missouri. Our re-examination of Kansas material

leaves L. tatarica represented by a single collection from Cloud County. Lonicera morrowii has not been

verified from the state, although, using characters employed by Green (1966) in distinguishing hybrids

of this complex, two specimens here referred to L. xbella (Morse 12673b, Morse 12673c) appear somewhat

intermediate between L. xbella and L. morrowii.

Voucher specimens: KANSAS. Douglas Co.: Lawrence, just N of 19th St at S end of alley between the 1800 blocks of Indiana and Mis-

sissippi Sts, T13S, R19E, sec 1, SE!4 , NE%. 38.9503°N, 95.2437°W, elev 880 ft, weedy, brushy right of way in alley, 11 Apr 2006, Morse

12671; Lawrence, E side SD of Kansas Main Campus, T12S, R19E, sec 36, SE%. 38.9576—38.9620°N, 95.2430—95.2424°W,

elev 980—1010 ft, weedy 11 Battenfeld Scholarship Hall & Lilac Ln, and on S side of 12th St, just E section with Oread

Ave, 17 Apr 2006, Morse 12673a, Morse 12673b, Morse 12673c, 25 Apr 2006, Morse 12673b-A; Lawrence, E side of ence of Kansas

Main Campus, T12S, R19E, sec 36, SE%. 38.9620?N 95.2424°W, elev 980 ft, weedy woodlot on S side of 12th St, just E inters with Oread

Ave, 5 Jun 2006, Morse s.n; Lawrence, N side of University of Kansas Main Campus, T12S, RIĐE, sec 36, NE%, SW, near 38.9640°N,

95.2493°W, elev 940—990 ft, weedy woodlot on moderate, N-facing slope along 11th St, just N of Phi Kappa Theta fraternity, 25 Apr

2006, Morse s.n, 5 Jun 2006, Morse s.n. Leavenworth Co.: Ft. Leavenworth, W-central part, Hills ESE of Wagner Point, T8S, R22E, sec

15, SEM, elev 900-1050 ft, mesic, upland, oak-hickory-basswood-maple forest on hills ESE of Wagner Point, 3 May 1995, Freeman 7087,

Ft. Leavenworth, S-central part, area ca 0.5 mi NE main entrance to Fort and due N of water treatment plant, 39.3421—39.3348°N,

Morse et al., New and ted ls for the K fl 757

94.9111—94.9172°W, elev 760 ft, weedy, early successional woodlands and forests on N and E-facing slopes along Corral Creek, domi-

nated by Carya-Quercus and Celtis-Gleditsia-Maclura assocations and exotic shrubby understory, 21 Oct 2004, Morse 10879.

Malus L. (Rosaceae). Three species of Eurasian crab-apple, which are frequently cultivated in the U.S. and

reported as occasional escapes throughout the northeastern states (Gleason & Cronquist 1991; Rhoads &

Block 2000; USDA, NRCS 2004), have been observed growing spontaneously in eastern Kansas, where they

sometimes occurred with congener M. floribunda Sieb. ex Van Houtte. This latter species was first reported

for the state by Freeman et al. (1998).

*Malus baccata Borkh. (Rosaceae). Siberian crabapple is consdered to be naturalized in the state.

Voucher specimens: KANSAS. Douglas Co.: Lawrence, University of Kansas Campus West, just S of Foley Hall, T13S, R19E, sec 2, S15,

NE, SE. 38.9480°N, 95.2638°W, elev 910 ft, weedy tree line along chain link fence, 11 Apr 2005, Morse s.n, 25 May 2005, Morse s.n;

Lawrence, University of Kansas Campus West, T13S, R19E, sec 2, S!ó, NE!A, SEM. 38.9477°N, 95.2632°W, elev 920 ft, weedy lots near

Bridwell Lab, 7 Apr 2006, Morse 12665, Morse 12666; 25 Apr 2006, Morse 12665-A, Morse 12666-A; 2 Aug 2006, Morse 12665-B, Morse

12666-B, 11 Oct 2006, Morse 12665-C, Morse 12666-C; Lawrence, SW side of town, T13S, R19E, sec 20, NE. 38.9131?N, 95.3189°W,

elev 840 ft, low, wooded, disturbed ground S of the radio-control airfield, 17 Apr 2006, Freeman 21459 & Morse.

*Malus prunifolia (Willd.) Borkh. Plumleaf crabapple is considered to be a casual alien in the state.

Voucher specimens: KANSAS. Douglas Co.: Lawrence, near inters of 31st and Louisiana Sts, T13S, R19E, sec 12, SE 1⁄4, SE 1⁄4, near

38.9293°N, 95.2435°W, elev 820 ft, weedy bank of Naismith Creek, 6 Apr 2006, Morse 12661, Morse 12662; 24 Apr 2006, Morse 12661-A,

Morse 12662-A; 2 Aug 2006, Morse 12661-B, Morse 12662-B; 11 Sep 2006, Morse 12662-C.

*Malus sieboldii Regel. Toringo crabapple is considered to be a casual alien in the state.

Voucher specimens: KANSAS. Douglas Co.: Lawrence, University of Kansas Campus West, brushy woodland SW of greenhouse; area

originally an open wooded pasture, undisturbed for 45 yr, 7 Apr 1995, 26 Apr 1996, McGregor 41119; Lawrence, just N of intersection

of Iowa St and 21st St, T13S, R19E, sec 1, W edge NW!A, SW. 38.9480?N, 95.2607°W, elev 920 ft, weedy drainage ditch along E side

of Iowa St, 7 April 2006, Morse 12664, 24 Apr 2006, Morse 12664-A; 2 Aug 2006, Morse 12664-B.

*Najas minor Allioni (Hydrocharitaceae). Native to Europe, brittle waternymph has steadily expanded its

range westward in North America since first documented in New York in 1934 (Clausen 1936; Meriláinen

1968; Haynes 1979). The species has been reported from southeast Missouri (USDA, NRCS 2004) and south-

central Oklahoma (Nelson & Couch 1981, Haynes 2000). A recent collection confirms the presence of N.

minor in Kansas, where the collector reports this species has been present for several years. It is considered

to be naturalized in the state.

Voucher specimen: KANSAS. Coffey Co.: ca2 mi N, 2.5 mi E Burlington, Wolf Creek Lake: vicinity of Wolf Creek Generating Station,

T21S, R15E or R16E, near 38.2280°N, 95.6842°W, elev ca 1070 ft, station intake and elsewhere in lake, growing in mixed population

with Najas guadalupensis subsp. guadalupensis, 09 Aug 2006, Haines s.n.

*Oxalis corniculata L. (Oxalidaceae). Introduced to the U.S. from tropical America, creeping woodsorrel

is a common weed of greenhouses (Cusick 2002). It previously has been reported from 43 states and the

District of Columbia in the continental U.S. (USDA, NRCS 2004) and from the majority of the states in

the Great Plains. In Kansas, this species was first noticed in greenhouses around the University of Kansas

in 1982; it has been collected from disturbed sites throughout the eastern part of the state since then. It is

considered to be naturalized in the state.

Voucher specimens: KANSAS. Chautauqua Co.: 4 mi N, 0.5 mi E Peru, Riley Memorial Cemetery, T33S, R12E, sec 3, SE, SE!4, SW.

37?08'28"N, 96%04'56"W, elev 260 m, mowed, weedy, upland tallgrass prairie in cemetery, 23 Apr 2001, Morse 5419 & Michener; 0.75 mi

W Peru, Peru Cemetery, T34S, R12E, sec 20, W14, NE, 37°04'45"N, 96%06'41"W, elev 285 m, mowed, weedy, upland tallgrass prairie

in cemetery, scattered oaks and junipers, 23 Apr 2001, Morse 5435 & Michener. Cherokee Co.: 4.75 mi S, 2.75 mi E jct of US Hwy 69 «€

KS Hwy 96 at Crestline, S side of SE Lostline Rd, T34S, R25E, sec 3, S15, SEY4. 37.107 7?N, 94.6550°W, elev 810—900 ft, open post oak-

blackjack oak-Texas hickory woodland on moderate to steep, predominately W- and S-facing slopes above Spring River, 28 Sep 2005,

Morse 11803 et al. Douglas Co.: Lawrence, University of Kansas West Campus, in greenhouse beds and pots, 8 Mar 1982, McGregor 32859;

Lawrence, 521 Lawrence Ave, in garden and lawn, 14 oct 1997, McGregor 41335; Lawrence, yard of private residence at 1733 Mississippi

St, T13S, R20E, sec 1, SE%4, NE, 38.9518°N, 95.2446°W, elev 890 ft, cultivated garden, 25 Oct 2005, Morse s.n.

f4L,D o ID L

758 Journal of t titute of Texas 1(1)

Philadelphus L. (Hydrangeaceae). Three taxa of mock-orange are reported here for the first time from

Kansas. Two occurrences may represent instances in which individuals have merely persisted from prior

cultivation. However, both observations were of large plants, several meters in diameter, occurring well away

from any recent human habitation. A fourth taxon, P. pubescens Loisel (hoary mock orange), is occasionally

planted as an ornamental in eastern Kansas, but has not been observed to escape.

*Philadelphus coronarius L. Native to Eurasia, sweet mock-orange has been reported from the eastern

U.S., as far west as Minnesota and Missouri (Mohlenbrock 1986; Rhoads & Block 2000, USDA, NRCS 2004).

A single large individual inside the edge of an oak-hickory forest confirms the presence of this species in

Kansas, where it is considered to be a casual alien.

Voucher specimens: KANSAS. Wyandotte Co.: 0.5 mi N, 1.75 mi E jct of KS Hwy 32 & Loring Rd on E side of Bonner Springs, Theo-

dore Naish Boy Scout Reservation, T11S, R23E, sec 22, S14, SW14 and sec 27, NW. 39.0689 —39.0735?N, 94.8507—94.8481°W, elev

900—980 ft, mesic, upland, white oak-white ash-hickory forest on generally N-facing slopes of ridge above N side of Kansas River, 18

April 2004, Morse 11053; 12 May 2005, Morse 11216 et al.

*Philadelphus inodorus L. Scentless mock-orange is native to southeastern North America, but is fre-

quently cultivated and has escaped throughout the northeast U.S. (Mohlenbrock 1986; Rhoads & Block

2000; USDA, NRCS 2004). A single individual found in extreme northeast Kansas confirms the presence

of this species in the state, where it is considered to be a casual alien.

Voucher specimen: KANSAS. Leavenworth Co.: Ft. Leavenworth, E of Sherman Army Airfield along Missouri River, T8S, R23E, sec

12, S !5, elev 750—800 ft, open disturbed area along Missouri River, 28 Jun 1998, Elliott & Freeman 834.

*Philadelphus xnivalis Jacques (P. coronarius L. x P. pubescens Loisel). Though not reported as an escape

in North America, this hybrid mock-orange was noted by Hu (1954-1956) as “one of the most commonly

cultivated Philadelphus in the gardens of Boston.” A single large individual growing 30 m inside the edge of an

oak-hickory forest confirms the presence of this taxon in Kansas, where it is considered to be a casual alien.

Voucher specimens: KANSAS. Leavenworth Co.: Ft. Leavenworth, W-central part, ravine 0.25 mi E of Bell Point, between reservoir

(to N) and radio tower (to S), 39?21'46"N 94°55'58"W, elev 880-1000 ft, disturbed oak-hickory-basswood forest on moderate E-facing

slopes with deep, steep-sided draws along tributary to Quarry Creek, 18 Jul 2003, Morse 9759; 3 Jun 2004, Morse 10537 & Rossow.

*Poncirus trifoliata (L.) Raf. (Rutaceae). Native to Asia, hardy-orange has been naturalized throughout the

southeastern U.S. and as far north as Pennsylvania (Rhoads & Block 2000; USDA, NRCS 2004). In the Great

Plains, this species has been reported from Oklahoma and eastern Texas (USDA, NRCS 2004). A single col-

lection confirms the presence of hardy-orange in Kansas. It is considered to be a casual alien in the state.

Voucher specimen: KANSAS. Cowley Co.: 1.5 mi W Arkansas City, Chaplin Nature Center, T34S, R3E, sec 16, Sí. 37°05'32"N,

97°06'17"W, elev 1080-1170 ft, forested uplands and floodplain, and open, sandy floodplain along W side of the Arkansas River, 25

Sep 2004, Freeman 20199.

*Pyracantha coccinea M. Roem. (Rosaceae). Native to Eurasia, scarlet firethorn is reported from scattered

states through the southern U.S., a handful of northeastern states, and Oregon (USDA, NRCS 2004). The

species is reported by Welsh et al. (2003) as “persisting, and escaping rarely” in Utah. A single occurrence

confirms the presence of this species in Kansas, where it is considered to be a casual alien.

Voucher specimen: KANSAS. Douglas Co.: Lawrence, ca 2 mi S, Haskell Ave, 0.5 mi S of Wakarusa River, T13S, R20E, sec 20, W Y.

38.9043°N, 95.2225°W, elev 860 ft, grazed, upland pasture E of blacktop, 17 Apr 2006, Freeman 21461 & Morse.

*Rhodotypos scandens (Thunb.) Makino (Rosaceae). A native of Japan, jetbead is widely cultivated in

eastern North America as an ornamental shrub and has been reported as an occasional escape, or perhap

a more aggressive invader, in the northeast U.S. and from Alabama and Georgia (Rhoads & Block 2000;

Lamont & Young 2002; USDA, NRCS 2004). In the Midwest, jetbead is known from Illinois (Mohlenbrock

1986) and Minnesota (Ownbey & Morley 1991), but the species has not been reported in the Great Plains.

Two populations were discovered recently in forested areas of northeast Kansas, where it is considered to

be naturalized.

Morse et al., New and ted Is for the K fl 759

Li los T

Voucher specimens: KANSAS. Leavenworth Co.: Ft. Leavenworth, W-central part, area to N of water reservoir and E of Hancock Hill,

39°21'57"—22'03"N, 94?56'06"—55'47"W, elev 850-1060 ft, mesic oak-hickory-basswood forest on NE-facing slope and hackberry-

mulberry floodplain forest along tributary to Quarry Creek, 11 Jun 2003, Morse 9507; Ft. Leavenworth, NW part, Bluffs W of N end of

Sherman Army Airfield and ca 0.75 mi NE Hancock Hill, 39°22'04"—29"N, 94°55'59"_40"W, elev 770—850 ft, maple-basswood-oak

forest on steep NE- and W-facing bluffs above Missouri River, and marshy floodplain forest at base of bluffs, 10 Nov 2003, Morse 9896

& Loring. Shawnee Co.: SE side of Topeka, Dornwood Park, T12S, R16E, sec 9, E%, near 39?01'18"N, 95?38'18"W, elev 900—960 ft,

wooded hillside, 9 May 2002, Hansen s.n.

*Viola striata Aiton (Violaceae). Native to the eastern U.S. as far west as Oklahoma (USDA, NRCS 2004),

striped cream violet was mapped by Steyermark (1963) from throughout the Ozarks and in Clay Co., Mis-

souri. Though it is occasionally cultivated in shaded gardens and lawns, the species was excluded from

the Great Plains by Brooks & McGregor (1991) for lack of specimen evidence. A recent observation of a

persistent population confirms the presence of V. striata in eastern Kansas. It is considered to be a casual

alien in the state

Voucher specimen: KANSAS. Lyon Co.: 0.5 mi S Hartford, T20S R13E, sec 22, SEY4, SEA, elev 350 m, mowed upland cemetery with

remnant prairie, surrounded by mesic to dry-mesic tallgrass prairie in Osage Questas, 23 Apr 1997, Morse et al. 1160.

*Zoysia japonica Steud. (Poaceae). Korean lawngrass is naturalized sq throughout the eastern U.S.

and in California (Catling et al. 1977; Anderson 2003; USDA, NRCS 2004). Though commonly cultivated in

lawns in Kansas, the species has only recently been confirmed as naturalized in the state.

Voucher specimen: KANSAS. Neosho Co.: 1 mi S Galesburg, New Mount Hope Cemetery, T30S, R19E, sec 5, SW'4, SEM. 37?27'33"N,

95?21'01"W, elev 970 ft, mowed, grassy, E side of cemetery, scattered moist depressions with abundant Juncus and Cyperus, 16 Jun 2005,

Freeman 20505.

ACKNOWLEDGMENTS

We thank R.F.C. Naczi (DOV) for identification of specimens of Carex corrugata; the curators of FHKSC,

KSC, KSP, KSTC, and WASH for access to, and information about, holdings in their collections; and Robert

Kaul, Stephen L. Timme, and an anonymous reviewer for helpful comments on the manuscript.

REFERENCES

ANDERSON, S.J. 2003. Zoysia. In: M.E. Barkworth, K.M. Capels, S. Long, and M.B. Piep, eds. Flora of North America

north of Mexico. Volume 25: Magnoliophyta: Commelinidae (in part): Poaceae, part 2. Oxford University Press.

New York, NY. Pp. 281-284.

BARNARD, l. 2006. Exotic plant introduction in Kansas, two new species. Sida 22:777-779.

BELDEN, A., JR, G.P. FLEMING, J.C. Lubwig, J.F. TowNsEND, N.E. Van Acstine, C.E. Stevens, and T.F. Wiegorr. 2004. Noteworthy

collections: Virginia. Castanea 69:144-153.

Brooks, R.E. and R.L. McGrecor. 1991. Violaceae. In: Great Plains Flora Association. Flora of the Great Plains. University

Press of Kansas, Lawrence. Pp. 255-264.

CARRUTH, J.H. 1873. Report on the botany of Kansas for the year 1873. Trans. Kansas Acad. Sci. 2:9-14.

CARRUTH, J.H. 1877. Centennial catalog of the plants of Kansas. Trans. Kansas Acad. Sci. 5:40-59,

CATUNG, PM., A.A. Rezwicek, and J.L. Riev. 1977. Some new and interesting grass records from soutern Ontario.

Canad. Field-Naturalist 91:350-359.

CHESTER, EW. 2000. Noteworthy collections: Tennessee. Castanea 65:168- 170.

CLAUSEN, R.T. 1936. Studies in the genus Najas in the northern United States. Rhodora 38:333-345 + 1 plate.

Cusick, A.W. 2002. Six non-native species newly discovered in the lowa vascular flora. Sida 20:405-407.

DAR8YSHIRE, S.J., C.E. WiLsoN, and K. ALuison. 2003. The biology of invasive alien plants in Canada. 1. Eriochloa villosa

(Thunb. Kunth. Canad. J. Plant Sci. 83:987-999.

FRECKMANN, R.W. and M.G. LeLoNG. 2003. Dichanthelium. In: M.E. Barkworth, K.M. Capels, S. Long, and M.B. Piep, eds.

Flora of North America north of Mexico. Volume 25: Magnoliophyta: Commelinidae (in part): Poaceae, part

2. Oxford University Press. New York, NY. Pp. 406-450.

760 Journal of the Botanical R h Institute of Texas 1(1)

FREEMAN, C.C. 2000. Vascular plants new to three states in the central United States. Trans. Kansas Acad. Sci.

103:51-54.

FREEMAN, C.C., R.L. MCGREGOR, and C.A. Morse. 1998. Vascular plants new to Kansas. Sida 18:593-604.

FREEMAN, C.C., C.A. Morse, and R.L. McGrecor. 2003. New vascular plant records for the grassland biome of central

North America. Sida 20:1289-1297,

Gittett, J.M. 1971. Cyperus fuscus L. new to Canada. Canad. Field-Naturalist 85:190.

GLEASON, H.A. and A. Cronauist. 1991. Manual of the vascular plants of northeastern United States and adjacent

Canada. (second edition). New York Bot. Gard. Bronx, NY.

GREAT PLAINS FLoRA ASSOCIATION. 1977. Atlas of the flora of the Great Plains. lowa State University Press, Ames.

GREAT PLAINS FLoRA ASSOCIATION. 1991. Flora of the Great Plains. University Press of Kansas, Lawrence.

Green, PS. 1966. Identification of the species and hybrids in the Lonicera tatarica complex. J. Arnold Arbor.

47:75-88.

HARRIMAN, N.A. 1972. Records on the flora of Wisconsin. Rhodora 74:156-157.

Haynes, R.R. 1979. Revision of North and Central American Najas (Najadaceae). Sida 8:34-56.

Haynes, R.R. 2000. Najadaceae. In: Flora of North America Editorial Committee, eds. Flora of North America north of

Mexico. Volume 22: Magnoliophyta: Alismatidae, Arecidae, Commelinidae (in part), and Zingiberidae. Oxford

University Press. New York. NY. Pp. 77-83.

Hu, S.-Y. 1954-1956. A monograph of the genus Philadelphus. J. Arnold Arbor. 35:275-333, 36:53-109, 36:325-368,

37:15-90.

KoLsTAD, O.A. 1991. Cyperaceae. In: Great Plains Flora Association. Flora of the Great Plains. University Press of

Kansas, Lawrence. Pp. 1059-1113.

Lamont, E.E. and S.M. Young. 2002. Noteworthy plants reported from the Torrey Range-2001. J. Torrey Bot. Soc.

129:363-371.

McGrecor, R.L. 1991. Apiaceae. In: Great Plains Flora Association. Flora of the Great Plains. University Press of

Kansas, Lawrence. Pp. 584-604.

MERILAINEN, J. 1968. Najas minor All. in North America. Rhodora 70:161-175.

MonHLENBROCK, R.H. 1986. Guide to the vascular plants of Illinois. Southern Illinois University Press, Carbondale.

Naczi, R.F.C. and C.T. Bryson. 2003. Carex sect. Griseae. In: Flora of North America Editorial Committee, eds. Flora

of North America north of Mexico. Volume 23: Magnoliophyta: Commelinidae (in part): Cyperaceae. Oxford

University Press. New York, NY. Pp. 448-461.

NELSON, E.N. and R.W. Couch. 1981. Occurrence of Najas minor and Najas marina (Najadaceae) in Oklahoma. Proc.

Oklahoma Acad. Sci. 61:78.

Ownsey, G.B. and T. Mortey. 1991. Vascular plants of Minnesota: a checklist and atlas. University of Minnesota

Press, Minneapolis.

RICHARDSON, D.M, P. Pyšek, M. REjuANEK, M.G. Bangoun, F.D. PANErTA, and C.J. West. 2000. Naturalization and invasion of

alien plants: concepts and definitions. Diversity & Distrib. 6:93-107.

RHOADS, A.F. and T.A. Block. 2000. The plants of Pennsylvania: an illustrated manual. University of Pennsylvania

Press, Philadelphia.

ROLFSMEIER, S.B. 1995. Keys and distributional maps for Nebraska Cyperaceae, part 1: Bulbostylis, Cyperus, Dulichium,

Eleocharis, Eriophorum, Fimbristylis, Fuirena, Lipocarpha, and Scirpus. Trans. Nebraska Acad. Sci. 22:27-42.

ROLFSMEIER, S.B., R. KauL, and D.M. SUTHERLAND. 1991. New and corrected floristic records for Nebraska — 4. Trans.

Nebraska Acad. Sci. 18:141-150.

SHAW, R.B., R.D. Wester, and C.M. Bern. 2003. Eriochloa. In: Flora of North America Editorial Committee, eds. Flora of

North America north of Mexico. Volume 25: Magnoliophyta: Commelinidae (in part): Poaceae, part 2. Oxford

University Press. New York, NY. Pp. 505-515.

SHILDNECK, P. and A.G. Jones. 1986. Cerastium dubium (Caryophyllaceae) new for the eastern half of North America

(comparison with sympatric Cerastium species, including cytological data). Castanea 51:49-55.

SINGHURST, J.R. and W.C. Homes. 2005. Uvularia sessilifolia L. (Liliaceae): new to Kansas. Trans. Kansas Acad. Sci.

108:57-58.

Morse et al., New and ted Is for the K fl 761

SMYTH, B.B. 1890. Catalog of the flowering plants and ferns of Kansas. Bull. Washburn Coll. Lab. Nat. Hist.

2:43-61.

SMYTH, B.B. 1892. Check list of the plants of Kansas. Published by the author. Topeka, Kansas.

STEYERMARK, J.A. 1963. Flora of Missouri. lowa State University Press, Ames.

STUBBENDIECK, J., G.Y. Frisoe, and M.R. Bouck. 1994. Weeds of Nebraska and the Great Plains. Nebraska Dept. of

Agriculture, Lincoln.

Stuckey, R.L. and T.M. Barkley. 1993. Weeds. In: Flora of North America Editorial Committee, eds. Flora of North

America north of Mexico. Volume 1: Introduction. Oxford University Press. New York, NY. Pp. 193-198.

TavLoR, R.J. and C.E. Tayor. 1981. Plants new to Arkansas, Oklahoma and Texas. Sida 9:25-28.

TAYLOR, C.E., L.K. MacnarH, P. FoLtey, P. Buck, and S. Carpenter. 1996. Oklahoma vascular plants: additions and distri-

butional comments. Proc. Oklahoma Acad. Sci. 76:31-34.

Tucker, G.C., B.G. Marcks, and J.R. Carter. 2003. Cyperus. In: Flora of North America Editorial Committee, eds. Flora

of North America north of Mexico. Volume 23: Magnoliophyta: Commelinidae (in part): Cyperaceae. Oxford

University Press. New York, NY. Pp. 141-191.

USDA, NRCS. 2004. The PLANTS database, Version 3.5 (http://plants.usda.gov). National Plant Data Center, Baton

Rouge.

ViNcENT, M.A. 2004. Spread of Fatoua villosa (Mulberry Weed; Moraceae) in North America. J. Kentucky Acad. Sci.

65:67-74.

WELSH, S.L. 2003. Rosaceae. In: S.L. Welsh, N.D. Atwood, S. Goodrich, and L.C. Higgins, eds. A Utah Flora. Monte L.

Bean Life Science Museum, Brigham Young University. Provo, UT. Pp. 550-577.

Wuittemore, A. 2006. Caprifoliaceae. In: G. Yatskievych Steyermark's flora of Missouri. Vol. 2. Missouri Botanical

Garden, St. Louis. Pp. 765-794.

YATSKIEVYCH, G. 1999. Steyermark's flora of Missouri. Vol. 1. Missouri Dept. of Conservation, Jefferson City.

YATSKIEVYCH, G. 2006. Steyermark's flora of Missouri. Vol. 2. Missouri Botanical Garden, St. Louis.

YATSKIEVYCH, G. and J.A. RAVEILL. 2001. Notes on the increasing proportion of non-native angiosperms in the Mis-

souri flora, with reports of three new genera for the state. Sida 19:701-709.

762 Journal of the Botanical R h Institute of Texas 1(1)

BOOK NOTICES

DouaLas J. Furuyma, H. BRADLEY SHAFFER, AND DANIEL SIMBERLOFF (EDs.). 2006. Annual Review of Ecology,

Evolution, and Systematics, Vol. 37, 2006. (ISBN 0-8243-1437-9, hbk.). Annual Reviews, 4139

El Camino Way, PO Box 10139, Palo Alto, CA 94303-0139, U.S.A. (Orders: www.annualreviews.org,

service@annualreviews.org, 650-493-4400, 650-424-0910 or 650-855-9815 fax, 1-800-523-8635).

$80.00, 682 pp., color figures, 742" x 942".

Collection of new reviews in easy-reading format, covering a wide array of topics. Abstracts and author information are free at <http://

arjournals.annualreviews.org> and full text is available with subscription

Contents.—1) Birth-death models in mE 2) The posterior and the prior in Bayesian phylogenetics. 3) Unifying and

testing models of sexual selection. 4) Genetic inl the age of genomics. 5) Ecological effects

of invasive arthropod generalist predators. 6) The QUIS of Bento architecture. 7) The major histocompatibility complex, sexual

selection, and mate choice. 8) Some evolutionary consequences of being a tree. 9) Late Quaternary extinctions: state of the debate.

10) Innate immunity, environmental drivers, and disease ecology of marine and freshwater invertebrates. 11) Experimental methods

for measuring gene interactions. 12) Corridors for conservation: integrating pattern and process. 13) The population biology of large

brown seaweeds: ecological consequences of multiphasic life histories in dynamic coastal environments. 14) Living on the edge of two

changing worlds: forecasting the responses of rocky intertidal ecosystems to climate change. 15) Has vicariance or dispersal been the

predominant biogeographic force in Madagascar? 16) Limits to the adaptive potential of small populations. 17) Resource exchange in

the rhizosphere: molecular tools and the microbial perspective. 18) The role of hybridization in the evolution of reef corals. 19) The

new bioinformatics: integrating ecological data from the gene to the biosphere. 20) Incorporating molecular evolution into phylogenetic

analysis, and a new compilation of conserved pol hain reaction primers for animal mitochondrial DNA. 21) The developmen-

tal, physiological, neural, and genetical causes and consequences of frequency-dependent selection in the wild. 22) Carbon-nitrogen

interactions in terrestrial ecosystems in response to rising atmospheric carbon dioxide. 23) Ecological and evolutionary responses to

recent climate change.—Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A.

WINSOME SHEPHERD. 2000. Wellington's Heritage: Plants, Gardens, and Landscape. (ISBN 0-909010-73-0,

pbk.). Te Papa Press, Museum of New Zealand Te Papa Tongarewa, Cable Street, PO Box 467, Wel-

lington, New Zealand. (Orders: www.tepapa.govt.nz/TePapa/English/TePapaPress, +64 (0)4 381-7470,

+64 (0)4 381-7280 fax). ~$34.29 ($49.99 NZD), 256 pp., color photos and illustrations, b/w photos,

OA x ll

“This book tells the story of the Wellington [New Zealand] landscape and its private and public gardens, from colonial times to the

present. It looks in detail at key pan un ee gardeners in the nineteenth century in Wellington and the Hutt Valley, gives a history of

] t of the Botani d d the Town Belt, as well as other parks. Wellington's

aiken lille are EM well illustrated." The author notes that “the book's origins go back to 1979 when [he] was asked by the

New Zealand Historic Places Trust to research plant introductions to New Zealand after the Missionary period.

It is an amazing chronicle of the evolution of the botanical landscape of this beautiful city, its “eco-history.” Paintings, maps, and

hundreds of photos of structures and associated landscape provide documentation of such detail and care that this account surely is a

strong academic contribution. But the text is fascinating and non-technical and many interested in similar aspects of their own city, in

whatever part of the world, will be interested to have this book.— Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort

Worth, TX 76102-4060, U.S.A.

James LL. Morison AND MICHAEL D. Morecrort (Eps.). 2006. Plant Growth and Climate Change. (ISBN

978-1-4051-3192-6, hbk.). Blackwell Publishing, 2121 State Avenue, Ames, IA 50014-8300, U.S.A.

(Orders: www.blackwellpublishing.com, 515-292-0140, 515-292-3348 fax, 1-800-862-6657). $199.99,

232 pp., illustrations, m x oy".

T al £f se ] £, 1 ph

The book examines the major aspects of how , temperature, water Md

and the interactions between these factors. Topics are MED Seed and cover the field suh saca but easily

Plenty of good illustrations and charts. “It is directed at advanced-level university students, researchers and professionals across the

range of plant science disciplines, including plant physiology, plant ecology and crop science.”

Contents. —1) Recent and future climate change and its implications for plant growth; 2) Plant responses to rising atmospheric

carbon dioxide; 3) The significance of temperature in plant life; 4) Temperature and plant development: phenology and seasonality; 5)

ue of pen do oud dip: to changes in water P in a changing climate; 6) Water availability and productivity; 7)

Effects of temi plant ) Issues in modelling plant ecosystem rest to elevated CO2:

interactions with soil Soie 9) M the effect of climate change on global plant productivity and the carbon cycle; References.

—Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A.

J. Bot. Res. Inst. Texas 1(1): 762. 2007

MINUARTIA DRUMMONDII (CARYOPHYLLACEAE) AND GRATIOLA FLAVA

(PLANTAGINACEAE) REDISCOVERED IN LOUISIANA AND GRATIOLA FLAVA

HISTORICALLY IN ARKANSAS

Michael H. MacRoberts Christopher S. Reid

Barbara R. MacRoberts Patricia L. Faulkner

Bog Research, 740 Columbia Louisiana Natural Heritage Program

Shreveport, Louisiana 71104, U.S.A. Louisiana Department of Wildlife and Fisheries

Herbarium, Museum of Life Sciences 2000 Quail Drive

Louisiana State Univeristy in Shreveport Baton Rouge, Louisiana 70898-9000, U.S.A.

Shreveport, Louisiana 71115, U.S.A.

Dwayne Estes

University of Tennessee Herbarium

Department of Ecology and Evolutionary Biology

University of Tennessee

Knoxville, Tennessee 37996-1100, U.S.A.

ABSTRACT

Collected in Louisiana in the mid-nineteenth century and not found since, Minuartia drummondii and Gratiola flava have recently been

discovered in saline prairies in Caddo and De Soto parishes. Two herbarium sheets of M. drummondii collected in the mid-nineteenth

century in Louisiana were located as well as an herbarium specimen of M. drummondii, misidentified as M. muriculata (M. muscorum),

collected in a saline prairie in Red River Parish in 1990. Herbarium sheets from the mid-nineteenth century containing G. flava specimens

from Texas, Louisiana, and possibly Arkansas were also located.

RESUMEN

Minuartia drummondii y Gratiola flava colectadas en Louisiana a mediados del siglo diez y nueve, y que no habian vuelto a encontrarse

desde entonces, se han descubierto recientemente en paraderas salinas en las parróquias de Caddo y De Soto. Fueron localizados dos

m de e de M. drummondii colectados a mu del siglo e y nueve en Louisiana, así como un pliego de M. drummondii,

M. muriculata (M. muscorum), colect ia de Red River en 1990. Se localizaron

E L

también pliegos de herbario de la mitad de la mitad del siglo diez y nueve que contienen especímenes de G. flava de Texas, Louisiana,

y posiblemente Arkansas

Minuartia drummondii (Shinners) McNeill [Arenaria drummondii Shinners] was first reported in Louisiana

by Riddell (1852) under the name Stellaria nuttallii Torr. & A. Gray. Schutz (1979) reported it under the

same name for Rapides Parish on the basis of a specimen collected in the mid-nineteenth century by Josiah

Hale (s.n. NO). Hale's specimen is presumably the source of Riddell's (1852) inclusion since he and Hale

collaborated. MacRoberts (1984) included the taxon as M. drummondii in his checklist of the Louisiana flora

on the basis of the Schutz report. MacRoberts (1989) later reported it for Louisiana citing the Hale (s.n. NO)

specimen. Thomas and Allen (1996) excluded M. drummondii from the Louisiana flora apparently because

they were unable to locate an herbarium specimen. It is currently reported from Texas, Oklahoma, and

Arkansas by Kartesz and Meacham (1999), Turner et al. (2003), NatureServe (2006), and USDA (2006) and

in Texas, Oklahoma, Arkansas, and Louisiana by the Flora of North America (2006).

We located two Josiah Hale (s.n. NO) specimens of M. drummondii collected in Louisiana in the mid-

nineteenth century. One label reads: *Stellaria Nuttallii, Prairies on Red River, J. Hale, M.D.” and the other,

“Stellaria Nuttallii, Moist Prairies, Caddo Par. La., April, Collected by Dr Josiah Hale.” Neither of these

specimens has a specific date.

Gratiola flava Leavenw. ex Pennell, a West Gulf Coastal Plain endemic (MacRoberts et al. 2002), is cur-

rently reported from Texas and Louisiana (Kartesz @ Meacham 1999; Turner et al. 2003; NatureServe 2006;

J. Bot. Res. Inst. Texas 1(1): 763 — 767. 2007

£+sha D o ID

764 Journal of t h Institute of Texas 1(1)

USDA 2006). In Louisiana, G. flava is currently considered to be an historical species (Louisiana Natural

Heritage Program 2006) because it has not been observed or collected since Josiah Hale’s collections in the

mid-nineteenth century (Vincent 1982; Pennell 1935; Thomas & Allen 1998).

Specimens of G. flava collected by Hale exist at GH, NO, and NY. The specimen at GH is part of a com-

posite sheet containing additional specimens of G. flava and a single individual of Gratiola neglecta Torr. The

G. flava specimens are separated on the sheet into six different groups, each of which is associated with its

own typed label, packet, or handwritten inscription. The Hale specimen is located in the lower left corner of

the sheet above a label bearing the following information: “Torr. & Gray; Flora, N. Amer. Louisiana. Hale.”

Three of the other specimens on the sheet were collected from Texas. Two of these are Charles Wright (s.n.)

collections, one from “Texas” and the other from “Texas. On the Colorado [River] below Lagrange [Fayette

County] The third Texas specimen is a collection by Elihu Hall (414) from Hempstead, Waller County. One

packet containing three individuals, located in the lower right corner, bears the note “Texas & Arkansas,

Leavenworth (Hb. Torr.).” Just above this packet in the right-central portion of the sheet are two individuals

and to the lower right of these specimens there is an annotation "Arkansas, Leavenworth."

Whether "Arkansas" here refers to modern Oklahoma or Arkansas is unknown. Between 1819 and 1835

the Arkansas Territory incorporated both and Melines Conkling Leavenworth was collecting in the upper

West Gulf Coastal Plain when Oklahoma and Arkansas were divided in 1835. To possibly further confuse

matters, in territorial times there was a town of Leavenworth on the Arkansas River in Oklahoma. But from

the sheet as a whole, it appears unlikely that the reference is to the town of Leavenworth.

At NO, there is a Hale specimen that is a collection consisting of a single individual of G. flava and two

stems of Gratiola brevifolia Raf. The specimen has no original label data but has been annotated by Francis

W. Pennell (in 1921), Karl A. Vincent (in 1980-81), and Dwayne Estes (in 2005). It is unclear from the

available data whether this specimen was actually collected in Louisiana.

At NY, there are two Hale specimens. One has a form label that bears the information *Gratiola flava.

Josiah Hale, M.D. Alexandria, Louisiana.” This specimen was deposited at NY in 1983 and was formerly

part of the Sartwell Collection at Hamilton College in Clinton, New York. It was annotated by Karl Vincent

in 1988 but has not been annotated by anyone else. The second specimen at NY is of a single individual

of G. flava collected by Hale from “Moist Prairies. April" and was given the collection number “12”; it was

identified only as “Gratiola.” Someone later wrote the specific epithet “pusilla Torr.” on the specimen follow-

ing the genus name. This specimen was also annotated by F.W. Pennell in 1920 (as G. tenella Pennell ined.)

and again in 1930 (as G. flava Leavenworth). Vincent verified the specimen in 1980—81 and provided the

following annotation: *The specimen of Gratiola flava is not a type specimen. This Hale collection is merely

cited by Pennell as from Louisiana in his 1935 monograph (pg 83)"

Examination of all available Hale material of G. flava revealed that these specimens lack specific locality

data. Therefore, it is quite interesting that Thomas and Allen (1998) and the USDA Plants Database (2006)

map the species from Rapides Parish, Louisiana. Thomas and Allen (1998, p. 159) cite the Hale specimen

at NO as the basis for the Rapides Parish record in spite of the lack of explicit label data on the specimen.

Therefore, the attribution of G. flava to Rapides Parish is likely based on the fact that Hale primarily col-

lected in the vicinity of Alexandria where he lived for a period of time (Ewan 2005; Anne Bradburn, Tulane

University, pers. comm .). It is also possible that the species was attributed to Rapides Parish based on one

of the specimens at NY that has a form label with "Alexandria, Louisiana" printed on it. Whether or not the

specimen actually came from the Alexandria area cannot be known, and it is possible that this is merely

the address of Hale and not the origin of the specimen. However, it is not unreasonable to believe that G.

flava could have been collected from Rapides Parish for three reasons. First, the species is known from

within a few kilometers of the Louisiana state line in northern Newton County, Texas, only about 55 km

west of Rapides Parish. Second, the geologic formation with which populations of G. flava are associated in

Newton County, the Catahoula Formation, extends eastward into northern Rapides Parish. Last, since the

Hale specimen at NO was mixed with G. brevifolia, a species that within Louisiana is found only in Allen,

MacRoherts et al ; Arl i Louisi floristic records 765

Beauregard, Calcasieu, Cameron, Jefferson Davis, Rapides, and Vernon parishes (Thomas & Allen 1998;

Knapp & Estes 2006) mostly in pineland seeps, it seems quite plausible that G. flava could have been col-

lected by Hale from central or southwestern Louisiana, possibly in Rapides Parish.

Alternatively, it is also possible that Hale collected the specimens of Gratiola flava from northwestern

Louisiana in prairies along the Red River. It is interesting to note that one of the specimens at NY bears

the label data “Moist Prairies. April" and that similar data are found on one of the specimens of Minuartia

drummondii collected by Hale from Caddo Parish in northwestern Louisiana (this paper).

The presence of specimens of Gratiola flava labeled as having been collected from *Arkansas" (Leavenworth

s.n. GH) prompted us to investigate the literature in search of additional notes concerning the distribution of

this species. Small (1903) reported G. flava (as Gratiola pusilla Torr.) from *prairies" in Arkansas and Texas

without citing any vouchers, although he most likely attributed the species to Arkansas based on a specimen

at NY bearing the label data "Arkansas. Dr. Leavenworth." Later, Pennell (1921) gave the distribution of G.

flava as *sandy prairies in and near the pinelands of southern Arkansas, western Louisiana, and eastern

Texas." Pennell based his citation of the species from Arkansas on the same Leavenworth specimen at NY

following Small, and his attribution of the species to Louisiana was based on one of the Hale specimens at

NY (Pennell 1921, p. 473). Later, Pennell (1935, p. 83) maintained G. flava as a component of the Louisiana

flora based again on the Hale specimens (at both GH and NY); however, he did not attribute the species to

Arkansas. In the list of synonyms for G. flava, Pennell (1935, p. 83) dismisses the occurrence of the species

in Arkansas, noting “As no specimen was found marked as from ‘Arkansas, the plant must have originally

been attributed to that territory because of Leavenworth's residence there or his form-labels so printed."

Smith (1988:424) also excluded G. flava from Arkansas and provided the following note: *reported (as G.

pusilla) for Arkansas by Small (1913) [Smith probably meant 1903], but apparently not in the state.”

Extant populations of Minuartia drummondii and Gratiola flava in saline prairies in Louisiana —lIn March and

April 2006, we found thousands of flowering M. drummondii and G. flava scattered across the 9.7 ha (23.4

acres) Barron Road Saline Prairie in southern Caddo Parish. On 13 April 2006, we found thousands of M.

drummondii and a small number of G. flava on the 10.9 ha (26.8 acres) Dickson Saline Prairie in northern De

Soto Parish. Subsequently, these two species have been found at two more saline prairies in De Soto Parish.

The discovery of M. drummondii in these prairies was initially thought to be the first discovery in Louisiana

since Hale's collections. However, subsequently while visiting NLU we found a specimen of M. drummondii

(Thomas 115,384) misidentified as M. muriculata (Maguire) McNeill (M. muscorum (Fassett) Rabeler), collected

in 1990 from a saline prairie in Red River Parish.

Saline prairies have been described by McInnis et al. (1993), Keith et al. (2004), Lester et al. (2005),

and Arkansas Natural Heritage Commission (2006) in Louisiana, Texas, and Arkansas. Those in Caddo

and De Soto parishes resemble other saline prairies, being open grassy expanses with treed mima mounds

scattered over them. They are a mosaic of dense to sparse herbaceous vegetation with interspersed bare soils

or “slicks.” The soils, which are typically cryptogamic, are poorly drained with slow permeability and high

sodium content. The soil series for the saline prairies in Caddo, De Soto, and Red River parishes is Bonn

silt loam (Edwards et al. 1980, 1991). Brimstone and Lafe soils are also known to support saline prairies in

Louisiana. In addition to M. drummondii and G. flava, species found in Louisiana saline prairies include Ana-

gallis minima (L.) Krause, Aristida longespica Poir., Aristida oligantha Michx., Astragalus distortus Torr. & Gray,

Cooperia drummondii Herbert, Coreopsis tinctoria Nutt., Crassula aquatica (L.) Schoenl., Croton michauxii G.L.

Webster, Evolvulus sericeus Sw., Fimbristylis puberula (Michx.) Vahl, Geocarpon minimum Mackenzie, Habranthus

tubispathus (LHer.) Traub, Houstonia micrantha (Shinners) Terrell, Houstonia pusilla Schoepf, Houstonia rosea

(Raf.) Terrell, Isolepis carinata Hook. & Arn. ex Torr., Iva angustifolia Nutt. ex DC., Krigia occidentalis Nutt.,

Lepuropetalon spathulatum EIL, Lotus unifoliolatus (Hook.) Benth., Marshallia caespitosa Nutt. ex DC., Mimosa

strigillosa Torr. & Gray, Minuartia muscorum (Fassett) Rabeler, Mirabilis albida (Walt.) Heimerl, Neptunia lutea

(Leavenworth) Benth., Nothoscordum bivalve (L.) Britt., Opuntia humifusa (Raf.) Raf., Phacelia glabra Nutt., Pha-

laris caroliniana Walt., Plantago pusilla Nutt., Rumex hastatulus Baldw., Sabatia campestris Nutt., Schedonnardus

f4L,D o ID

766 Journal of t h Institute of Texas 1(1)

paniculatus (Nutt.) Trel., Schoenolirion wrightii Sherman, Sporobolus pyramidatus (Lam.) Hitchc., Sporobolus

vaginiflorus (Torr. & Gray) Wood., Talinum parviflorum Nutt., Tradescantia hirsutiflora Bush, Tradescantia oc-

cidentalis (Britt.) Smyth, and Valerianella radiata (L.) Dufr.

Since we collected M. drummondii and G. flava in saline prairies, it is possible that Hale also did so. If

that is the case and if the label information for his M. drummondii is correct, it would follow that at least some

of the prairies on the Red River floodplain that existed in Caddo Parish in the mid-nineteenth century were

saline prairies (MacRoberts et al. 1997; MacRoberts & MacRoberts 2005), and perhaps it was from these

that Hale collected G. flava. However, it is also possible that G. flava was collected by Hale farther to the

south in western or west-central Louisiana quite possibly from Rapides Parish. Given the close proximity

of the newly discovered Louisiana populations to Arkansas, it is quite possible that Leavenworth collected

G. flava from southwestern Arkansas in saline prairies or similar habitats.

Voucher for Gratiola flava. LOUISIANA. Caddo Parish: B.R. & M.H. MacRoberts 7296 (LSUS), B.R. & M.H. MacRoberts 7310 (TENN).

De Soto Parish: B.R. & M.H. MacRoberts 7584 (LSUS), Reid 5724 LSU).

Vouchers for Minuartia drummondii. LOUISIANA. Caddo Parish: B.R. & M.H. MacRoberts 7314 (LSUS). De Soto Parish: B.R. & M.H.

MacRoberts 7586 (LSUS), Reid 5721 (LSU). Red River Parish: Thomas 115384 (NLU).

ACKNOWLEDGMENTS

We appreciate the cooperation of Sidney Evans and James Taylor who gave permission to survey the Bar-

ron Road site. We thank George Dickson and Ralph Dalton for allowing our survey of Dickson and Dalton

prairies and for their interest in the sites. Anne Bradburn, NO, provided photocopies of some of the Hale

specimens. Thanks are also extended to the curators of GH and NY for assistance during visits to these

herbaria and for loan of specimens.

REFERENCES

ARKANSAS NATURAL HERITAGE COMMISSION. 2006. Warren Prairie Natural Area. www.naturalheritage.com

EDWARDS, J.P, PG. MARTIN, JW. Macoun, W.W. Kitpatrick, and C. Henry. 1980. Soil survey of Caddo Parish, Louisiana.

U.S.D.A. Soil Service, Washington, D.C.

EpwARDS, J.P, M. Cooley, and C.L. Gobrrey. 1991. Soil survey of De Soto Parish, Louisiana. U.S.D.A. Soil Service,

Washington, D.C.

Ewan, J.A. 2005. Notes on Louisiana botany and botanists, 1718-1975. Sida 21:2275-2296.

FLORA OF NorTH America. 2006. www.eFlora.org.

KARTESZ, J.T. and C.M. Meacham. 1999. Synthesis of North American flora. Version 1.0. North Carolina Botanical

Garden, Chapel Hill.

KerrH, E.L., J.R. SINGHURST, and S. Cook. 2004. Geocarpon minimum (Caryophyllaceae), new to Texas. Sida 21:1165-

1169.

Knapp, W.M. and D. Estes. 2006. Gratiola brevifolia (Plantaginaceae) new to the flora of Delaware, the Delmarva

Pensinsula, and the mid-Atlantic. Sida 22: 825-829.

Lester, G.D., S.G. SORENSEN, P.L. FAULKNER, C.S. Reip, and I.E. Maxit. 2005. Louisiana comprehensive wildlife conservation

strategy. Louisiana Department of Wildlife and Fisheries. Baton Rouge.

Louisiana NATURAL HERITAGE PROGRAM. 2006. Rare plant species of Louisiana. Louisiana Department of Wildlife and

Fisheries. Baton Rouge.

MacRostRrs, D.T. 1984. The vascular plants of Louisiana: an annotated checklist and bibliography of the vascular

plants reported to grow without cultivation in Louisiana. Bull. Mus. Life Sci., Louisiana State University-Shreve-

port 6:1-165.

MacRogerTs, D.T. 1989. A documented checklist and atlas of the vascular flora of Louisiana. Bull. Mus. Life Sci.

Louisiana State Univ. 8:257-536.

MacRostRrs, D.T., B.R. MacRoserts, and M.H. MacRostnrs. 1997. A floristic and ecological interpretation of the Free-

man and Custis Red River expedition of 1806. Bull. Mus. Life Sci. Louisiana State Univ. 12:1-26.

MacRoherts et al ; Arl i Louisi floristic records 767

MacRoserts, H.H., B.R. MacRoserts, B.A. SorRIE, and R.E. Evans. 2002. Endemism in the West Gulf Coastal Plain: im-

portance of xeric habitats. Sida 20:767-780.

MacRoseets, M.H. and B.R. MacRoserts. 2005. Reference conditions of the Red River floodplain and upland, Caddo

Parish, Louisiana. Sida 21:1793-1806.

McInnis, N.C., L.M. Smith, and A.B. Pitman. 1993. Geocarpon minimum (Caryophyllaceae), new to Louisiana. Phy-

tologia 75:159-162.

NatureServe 2006. NatureServe Explorer. An online encyclopedia of life [web application]. Version 4.7. NatureServe,

Arlington, Virginia. Available http://www.natureserve.org/explorer.

PENNELL, FW. 1921. Scrophulariaceae of the west gulf states. Proc. Acad. Nat. Sci. Philadelphia 73:459-536.

PENNELL, EW, 1935. The Scrophulariaceae of eastern temperate North America. The Academy of Natural Sciences

of Philadelphia, Philadelphia, PA.

RipDELt, J.L. 1852. Catalogus florae Ludovicianae. New Orleans Med. Surg. J. 8:734—754.

Schurz, S.E. 1979. A preliminary survey of the vascular flora of Rapides Parish, Louisiana. M.S. Thesis, Northeast

Louisiana University, Monroe.

SMALL, J.K. 1903. Flora of the southeastern United States. Published by the author, New York.

SmrrH, E.B. 1988. Atlas and annotated list of the vascular plants of Arkansas. Privately Printed, Fayetteville. (Avail-

able online at: http://www.csdl.tamu.edu/FLORA/arkansas/arkindex.htm)

THomas, R.D. and C.M. ALLEN. 1996-1998. Atlas of the vascular flora of Louisiana. Vols. 2 and 3. Louisiana Depart-

ment of Wildlife and Fisheries, Baton Rouge.

Turner, B.L., H. NicHots, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. Sida, Bot. Misc. 24:1-

USDA NRCS. 2006. The PLANTS Database (http://plants.usda.gov, 5 July 2006). National Plant Data Center, Baton

Rouge, LA 70874-4490.

VINCENT, K.A. 1982. Scrophulariaceae of Louisiana. M.S. Thesis, University of Southwestern Louisiana, Lafayette.

768 Journal of the Botanical R h Institute of Texas 1(1)

BOOK NOTICES

Jim L. Bowyer, RUBIN SHMULSKY, and JOHN G. HAYGREEN. 2007. Forest Products and Wood Science: An Intro-

duction (5th Ed.). (ISBN 978-0-8138-2036-1, hbk.). Blackwell Publishing, 2121 State Avenue, Ames,

IA 50014-8300, U.S.A. (Orders: www.blackwellpublishing.com, 515-292-0140, 515-292-3348 fax,

1-800-862-6657). $89.99, 568 pp., illustrations, 7!4" x 1014".

A completely revised and updated edition of this textbook, providing students, wood scientists, and wood product professionals with an

introduction to the anatomical and physical nature of wood and the relationship of these characteristics to use of wood as an industrial

raw material. Chapters 1-7 introduce processes of growth and structure and chemical and structural characteristics; chapters 8-12

discuss physical properties of wood; chapters 13-17 discuss major wood-based products, the basic manufacturing processes associated

with each, and how raw material selection affects product properties; chapters 18 and 19 discusses wood as a source of energy and

chemicals and environmental implications of wood use.

Contents.—Introduction. 1) Tree Growth and Production of Woody Tissue. 2) Macroscopic Character of Wood. 3) Composition

and Structure of Wood Cells. 4) Softwood Structure. 5) Hardwood Structure. 6) Juvenile Wood, Reaction Wood, and Wood of Branches

and Roots. 7) Bark. 8) Wood and Water. 9) Density and Specific Gravity. 10) Strength and Mechanics. 11) Wood Durability and Protec-

tion. 12) Silvicul | Practices and Wood Quality. 13) Lumber. 14) Structural Panels. 15) Nonstructural Panels. 16) Composite Lumber

Products. 17) Pulp and Paper. 18) Energy and Chemical Products. 19) Wood in the Global Raw Materials Picture.— Guy Nesom, Botanical

Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A.

ANTHONY R. Yeo and Timotuy J. Flowers (eds). 2007. Plant Solute Transport. (ISBN 978-1-4051-3995-

3, hbk.). Blackwell Publishing, 2121 State Avenue, Ames, IA 50014-8300, U.S.A. (Orders: www.

blackwellpublishing.com, 515-292-0140, 515-292-3348 fax, 1-800-862-6657). $199.99, 424 pp.,

illustrations, 644" x 915".

A technical consideration of solute transport, with easy-to-reach information owing to the detailed and highly organized subtopical

structure of the discussions. Some of the topics include ERA and roles of solutes, physical bases of ion and water movement, solute

adaptation of solute use to more extreme environmen ts, and internally-controlled dehydration concomi-

tant with seed formation. “The book is directed at postgraduates, researchers, and professionals in plant physiology, biochemistry, and

molecular biology."

Contents.—1) General Introduction. 2) Solutes, what are they, where are they and what do they do? 3) The driving forces for wa-

ter and solute movement. 4) Membrane structure and the study of solute transport across plant membranes. 5) Transport across plant

membranes. 6) Regulation of ion transporters. 7) Intracellular transport: solute transport in chloroplasts, mitochondria, peroxisomes

and vacuoles, and between organelles. 8) lon uptake by plant roots. 9) Transport from root to shoot. 10) Solute transport in the phloem.

11) Factors limiting the rate of supply of solutes to the root surface. 12) Mineral deficiency and toxicity. 13) Water-limited conditions.

14) Salinity. 15) Desiccation tolerance —Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060,

Ron Russo. 2007. Field Guide to Plant Galls of California and Other Western States. (ISBN 978-0-

520-24885-4, pbk.). California Natural History Guide series no. 91. The University of California Press,

Berkeley, CA 94704, U.S.A. (Orders: California Princeton Fulfillment Services, 1445 Lower Ferry Road,

Ewing, NJ 08618, U.S.A., www.ucpress.edu, 609-883-1759, 609-883-7413 fax). $24.95, 400 pp., 338

color illustrations, 89 line illustrations, 15 tables, 412" x 714".

Plant gall fascinati ] pli 1 to understand. Some may appear to be no more than deformities on the plant; others

; 1 : £T 1 1 TRAC qas

in e texture, and color. Alli fthe gall inducer to the plant, whether it be bacterium,

TENE us, insect, or mistletoe. In the clear, concise, and ] ivei ] to cecidology, the author explains the various inducers

and the differences between such gall responses and T of abnormal plant growth such as cankers, burls, and root nodules

As a field guide, the book is arranged by plant types—trees, shrubs and then by Ee d as ash Led, oak galls, with indication

E COE fe) E

of the parts affected, which may be stem, leaf, inflorescence, root, etc. Color the descriptions

of the galls, their distribution, and approximate dates of appearance.

While the book concentrates on euis of the western states, many of the galls or similar ones, especially those of the cynipid wasps

that constitute 7596 of gall makers, are found t. This book will be useful to horticulturists, particularly perhaps those

lt gl

engaged i , but it will also be useful t

d others who interpret the natural world.—Joann Karges, (TCU Library,

retired), Botanical RE. Institute of Texas, Fort Worth, TX, 76102-4060, U.S.A.

J. Bot. Res. Inst. Texas 1(1): 768. 2007

NOTEWORITMY COLLECTIONS BERONDSEHEB

XAZOOSMISSISSIPPEDBELASSEGIOUN ODMISSISSIPPI

Daniel A. Skojac, Jr. Charles T. Bryson

U.S. Forest Service, Southern Research Station USDA-ARS

Southern Hardwoods Laboratory Southern Weed Science Laboratory

PO. Box 227 Stoneville, Mississippi 38776, U.S.A. PO. Box 350 Stoneville, Mississippi 38776, U.S.A.

dskojacefs.fed.us charles.bryson@ars.usda.gov

Charles H. Walker, Il

U.S. Forest Service, Southern Research Station

Southern Hardwoods Laboratory

PO. Box 227 Stoneville. Mississippi 38776, U.S.A.

ABSTRACT

The flora of the Yazoo-Mississippi Delta Region is the least represented in the checklist of Mississippi plants currently being compiled

for the state. This paper reports 20 noteworthy collections from the region and discusses their distributions within the state. Typha

angustifolia is reported new to Mississippi and Bowlesia incana, Caperonia palustris, Carya cordiformis, Carya glabra, Chenopodium stand-

leyanum, Eichhornia crassipes, Hottonia inflata, Oplismenus setarius, Podophyllum peltatum, Polystichum acrostichoides, Proserpinaca palustris,

Scirpus atrovirens, Senecio vulgaris, Sparganium americanum, and Spigelia marilandica are reported new to the Yazoo-Mississippi Delta

Region. Range extensions are reported for Callicarpa americana, Carex intumescens, Carya laciniosa, and Menispermum canadense within

the Yazoo- ii d Delta Region. Of the 20 species, Bowlesia incana, rad palustris, Eichhornia crassipes, and Senecio ud are

weed f agri ultural and humankind disturbed areas. Carya laciniosa, H infl , and Menispermum canadense are plant p ]

concern within the state and are on the state’s special plant tracking list.

RESUMEN

La flora de la región del delta del Yazoo- -o es la menos representada en el catálogo de plantas de Mississippi qu stá compi

lando actualmente para el estado. En est itan 20 colecciones notables de la región y se discute su distribución en el estado.

Typha angustifolia se cita como nueva para Mississippi y Bowlesia incana, Caperonia palustris, Carya cordiformis, Carya glabra, Chenopodium

standleyanum, Eichhornia crassipes, Hottonia inflata, Oplismenus setarius, Podophyllum peltatum, Polystichum acrostichoides, Proserpinaca

palustris, Scirpus atrovirens, Senecio vulgaris, Sparganium americanum, y Spigelia marilandica se citan como unevas para la región del delta

de Yazoo-Mississippi. Se incrementa el área de extensión de Callicarpa americana, Carex intumescens, Carya laciniosa, and Menispermum

canadense dentro de la región Yazoo-Mississippi. De las 20 especies, Bowlesia incana, Caperonia palustris, Eichhornia crassipes, y Senecio

vulgaris son malas hierbas para la agricultura y de áreas alteradas por el hombre. Carya laciniosa, Hottonia inflata, and Menispermum

canadense son plantas de especial interés en el estado y están en la lista estatal del plan especial de seguimiento.

INTRODUCTION

The Yazoo-Mississippi Delta Region is not a delta in the geomorphic sense, but rather is the floodplain of

the Mississippi and Yazoo rivers. This vast alluvial plain occupies approximately 18,389 km? and covers 10

counties entirely including Bolivar, Coahoma, Humphreys, Issaquena, Leflore, Quitman, Sharkey, Sunflower,

Tunica, and Washington counties and parts of 8 other counties including Carroll, DeSoto, Grenada, Holmes,

Panola, Tate, Warren, and Yazoo in northwest Mississippi. The Yazoo-Mississippi Delta Region is bordered to

the west by the Mississippi River and to the east by the Loess Bluff Region (Lowe 1921; Morris 1989). From

its northern terminus near Memphis, Tennessee, the Yazoo-Mississippi Delta Region extends southward to

the confluence of the Yazoo and Mississippi rivers near Vicksburg, Mississippi. Topography of the region is

flat and drainage is slow. Extant ridges (natural levees) rise slightly above adjacent areas of lower elevation

along ancient and current stream-channels, and represent the highest elevations in the region. Soils range

from poorly drained, fine clays on sites of lower relief to well drained, silt-loams and sandy soils on natural

levees. Construction of artificial levee systems during the last century has altered historical flooding regimes

and minimized the severity of seasonal flooding events across the region.

J. Bot. Res. Inst. Texas 1(1): 769 — 775. 2007

£+sha D o ID L

770 Journal of t titute of Texas 1(1)

The Yazoo-Mississippi Delta Region remains the least understood floristically and most under-collected

region in the state of Mississippi (Carter et al. 1980). Poor geographic distribution data also characterize the

region's flora. Significant collections from the region, however, are provided in Carter (1978), Gunn et al.

(1980), and Wiseman (1982). The mass conversion of the native bottomland hardwood forests to agricul-

ture production has hindered efforts to accurately define the area's historical floristic composition. Native

forest vegetation is now mostly restricted to sites of lower relief not suited to agricultural production due to

poor drainage. Species of these sites comprise the majority of taxa presented in the three works referenced

earlier. Because of the desirability for row crop production, higher elevation, well-drained sites remaining

in native vegetation are rare in the region. These specialized sites have yielded significant additions to the

Yazoo-Mississippi Delta Region flora in recent decades (Bryson and Jones 1990; Carter et al. 1990; Rabler

and Bryson 1990; Bryson and Elmore 1991; Bryson and Carter 1992; Bryson et al. 1992; Rosen et al. 2006),

and provided some historical perspective on the original floristic composition of the area. Additionally,

state and nation-wide geographic distribution data often preclude the Yazoo-Mississippi Delta Region in

the native range of many plant species. Thus, the area apparently interrupts geographic distributions of

species, whose ranges continue in areas of differing physiography adjacent to the Yazoo-Mississippi Delta

Region. This is attributed partially to the edaphic characteristics of the region, but anthropogenic activi-

ties, particularly widespread conversion of natural ecosystems to agriculture, are also relevant. Collections

presented by Carter et al. (1990) and those presented here from the isolated ridge sites remaining in native

vegetation have bridged the native ranges of many species otherwise extirpated or omitted from the Yazoo-

Mississippi Delta Region flora. Continued exploration of these sites will be required to attain an accurate

floral description for the Yazoo-Mississippi Delta Region of Mississippi.

NOTEWORTHY COLLECTIONS

Listed below are noteworthy collections from the Yazoo-Mississippi Delta Region of Mississippi. Many rep-

resent significant new discoveries to the region. Others are range extensions of species considered rare or

weedy within the Yazoo-Mississippi Delta first reported by Carter et al. (1990), or otherwise documented in

the region by McCook and Kartesz's (2000) preliminary checklist of Mississippi plants. Distributional data

are taken from Little (1971), McCook and Kartesz's (2000) checklist, and The PLANTS database (USDA,

NRCS 2006). Herbarium abbreviations follow Holmgren et al. (1990) and Holmgren and Holmgren (1998),

except the Southern Hardwoods Laboratory, Stoneville, MS (shl).

Bowlesia incana Ruiz & Pavon (Apiaceae). Hoary bowlesia (USDA, NRCS 2006), a non-native invasive weed,

continues to spread northward in Mississippi in container plants supplied at nurseries and as contaminates

of sod, mowing, and lawn maintenance equipment. Previously, this plant was only reported from a single

collection in Harrison County in Mississippi (McCook & Kartesz 2000). Our collections are important

because they document the northward range extension of B. incana in Mississippi and report it as a weed of

lawns, landscape beds, woodland edges, nurseries, and no- and reduced-tillage row crop production systems

in the Yazoo-Mississippi Delta Region. This weed is often misidentified in the seedling stage because of its

similarity to Ranunculus parviflorus L., also a common weed of gardens, lawns, landscape beds, and woodland

edges. In B. incana seedlings, the deeply lobed leaves are not toothed compared to the deeply lobed toothed

leaves of R. parviflorus. Our vouchers represent significant range extensions northward and the first records

of this species in the Yazoo-Mississippi Delta Region.

Voucher specimens: U.S.A. MISSISSIPPI. Bolivar Co.: Cleveland, Delta State University campus, 29 Apr 1993, Stewart 4709 (SWSL);

29 Feb 1996, Stewart 5379 (SWSL). Issaquena Co.: Grace, N of Hwy MS 14 in mowed lawn, 27 Apr 2001, Bryson 18365 & Goodlett

(SWSL). Washington Co.: Leland, along Deer Creek Drive, 14 Apr 1991, Bryson 10715 & Bryson (SWSL); Leland NE jct. of Broad and

Willeroy streets, 13 Mar 1994, Bryson 13309 & Bryson (SWSL); Leland, 2 blocks SE of Broad St. at 306 Garrison St. (private residence),

in flower beds and lawn, T18N R7W S14, 20 Mar 2004, Skojac 785 (shD; Stoneville, 16 Mar 2006, Bryson 20609 (DOV, MISS, MISSA,

MMNS, SWSL, USMS, VSC).

Callicarpa americana L. (Verbenaceae). American beautyberry (USDA, NRCS 2006) is a common woodland

Skojac et al., Yazoo-Mississippi Delta Region floristic records 771

rr wd

shrub of hardwood and mixed pine-hardwood forests throughout the state. It was most likely a common

understory shrub on sites with higher elevation than surrounding areas throughout the Delta prior to the

conversion of theses sites to row crop production. This species was first reported in the region by Carter et

al. (1990) from ridge-bottom sites in two Yazoo-Mississippi Delta counties. Our vouchers more than double

the number of documented sites within the Yazoo-Mississippi Delta Region where this otherwise common

shrub occurs.

Voucher o U.S.A. MISSISSIPPI. Humphreys Co.: SW of Belzoni and Gunn Bayou near Townson Lake, T15N R2W S16,

well-drained ridg ing bottomland hardwood forest, 2 Sep 2005, Skojac 1142 (shl). Leflore Co.: NW of Sidon, between Sidon

cut-off and old Yazoo a run, TIBN RIE S19, 1 Jul 1996, Bryson 15400 & MacDonald (SWSL); 16 Jun 1999, Bryson 17254 (SWSL).

Sharkey Co.: Delta National Forest, approx. 4 mi S of Holly Bluff, F.S. compartment 39, T11N R5W S34, bottomland hardwood forest

ridge-bottom site with notably higher relief than surrounding terrain, 26 May 2005, Skojac 1108 (shl).

Caperonia palustris (L.) St.-Hil. (Euphorbiaceae). Sacatrapo (USDA, NRCS 2006) or Texasweed (WSSA

1989) is reported in four southern states, including neighboring Arkansas and Louisiana, but is not listed

in Mississippi (USDA, NRCS 2006). McCook and Kartesz (2000) do not report vouchers of C. palustris from

Mississippi either, but do indicate that the species has likely been introduced into the state. Our vouchers from

Washington County are the first published accounts of this species from the Yazoo-Mississippi Delta Region,

where it is a weed of rice and soybeans, two of the principal agricultural row crops of the region. During the

past decade, C. palustris has become a major weed of rice production in Washington County, Mississippi.

Voucher specimens: U.S.A. MISSISSIPPI. Washington Co.: ca. 4 mi W of Arcola in rice field, 30 Aug 1982, McDaniel (SWSL); 5 mi W of

Arcola in soybean field, 11 Jul 1983, Elmore 208 (SWSL); Leland, weed in flowerbed along Cotton Drive, 28 Jul 1997, Bryson 16076 (SWSL);

Leroy Percy State Park, along S side of dirt road near ball field NE of alligator pond, T13W R5W S9, 6 Nov 2005, Walker MS24 (shl).

Carex intumescens Rudge (Cyperaceae). Greater bladder sedge (USDA, NRCS 2006) was first reported new

to the region in Carter et al. (1990). It is locally common in many bottomland hardwood forests throughout

the rest of Mississippi. Our voucher represents a range extension and county record of this seemingly rare

sedge in the Yazoo-Mississippi Delta Region.

Voucher specimen: U.S.A. MISSISSIPPI. Bolivar Co.: appox. 3.7 mi NW of Shelby, MS and ca. 2.0 mi W of US 61 in an isolated tract

of bottomland hardwood forest, 33°59'N 90°48'W, T25N ROW S34, 3 Jun 2004, Skojac 871 & Bryson (shl, SWSL).

Carya cordiformis (Wangenh.) K. Koch Juglandaceae). Bitternut hickory (USDA, NRCS 2006) has wide

distribution across much of the eastern U.S., but was excluded from the Yazoo-Mississippi Delta Region

by Little (1971). In Mississippi, it occurs in the northern half of the state on mesic to dry upland sites with

other hardwoods and it is somewhat common on well-drained soils of minor stream-bottom systems as

well. Our voucher is the first record of this hickory in the Yazoo-Mississippi Delta Region. Interestingly,

this collection was made from a river-bordering county rather than from one bordering the Loess Bluffs,

where the species is quite abundant.

Voucher specimen: U.S.A. MISSISSIPPI. Washington Co.: Yazoo National Wildlife Refuge, wooded tract S of Alligator Pond along

extant ridge bordering slough, 17 Aug 2004, Skojac 982 (shl).

Carya glabra (Mill.) Sweet (Juglandaceae). According to Little (1971), the native range of pignut hickory

(USDA, NRCS 2006) extends across all of Mississippi with the exception of the Yazoo-Mississippi Delta

Region. Our collections extend the known native range of this hickory into the Yazoo-Mississippi Delta,

where it appears to be restricted to the higher elevation ridge sites in the region. It is possible the material

we collected is of the C. leiodermis Sarg. form of pignut hickory, which is of special concern in the state.

Voucher specimens: U.S.A. MISSISSIPPI. Holmes Co.: Hillside National Wildlife Refuge, ca. 3 mi N of Eden in woods near the Alligator

Slough Nature Trail on ridge-bottom site, 29 Jul 2004, Skojac 937 (shl). Humphreys Co.: SW of Belzoni and Gunn Bayou near Townson

Lake, T15N R2W S16, well-drained ridge site supporting bottomland hardwood forest, 2 Sep 2005, Skojac 1143 (shl).

Carya laciniosa (Michx. f.) Loud. (Juglandaceae). Shellbark hickory (USDA, NRCS 2006) is a scattered

tree of alluvial sites north of Mississippi. According to Little (1971), its native range extends southward to

£+sha D o ID L

772 Journal of t titute of Texas 1(1)

southwestern Tennessee and northeastern Arkansas along the Mississippi river. A single record of this species

from DeSoto County in extreme northwest MS is reported in McCook and Kartesz's 2000) checklist of Mis-

sissippi plants. Our vouchers represent a significant range extension southward in the state. Shellbark hickory

is listed on the Mississippi special plant tracking list (Mississippi Natural Heritage Program 2002).

Voucher specimens: U.S.A. MISSISSIPPI. Bolivar Co.: ca. 3.7 mi NW of Shelby, MS and ca. 2.0 mi W of US 61 in an isolated tract

of bottomland hardwood forest, 33°59'N 90°48'W, T25N R6W S34, 3 Jun 2004, Skojac 865 & Bryson (SWSL, shl). Tunica Co.: ca. 4

mi S of Tunica; 1 mi S jct. Hwy US 61 and MS 4, then 1 mi E of Hwy US 61 on Hurt Rd., S of Hurt Rd., T5S R11W S28, N34°37.574' W

090°22.189', bottomland hardwood forest, 3 May 2006, Skojac 1257 & Bryson (shl).

Chenopodium standleyanum Aellen (Chenopodiaceae). Standley's goosefoot (USDA, NRCS 2006) has

limited distribution in Mississippi and is known only from the Yazoo-Mississippi Delta counties of Sharkey

and Leflore (McCook & Kartesz 2000; USDA, NRCS 2006). Our specimens were taken within and along

the edges of bottomland hardwood forests. This habitat differs from that of the more common C. album

L., which occurs most frequently in open fields and along row crop field margins in the Yazoo-Mississippi

Delta Region, and elsewhere in the state. Both McCook and Kartesz (2000) and USDA, NRCS (2006) cite our

Leflore County voucher based on duplicate material provided by the authors. Our vouchers document the

Leflore County station and provide an additional county record within the Yazoo-Mississippi Delta Region,

and represent the only other documented records of this species in the state.

Voucher specimens: U.S.A. MISSISSIPPI. Bolivar Co.: Approx. 3.7 mi NW of Shelby, MS and ca. 2.0 mi W of US 61 inan isolated tract

of bottomland hardwood forest, along southern edge of forest growing on spoil material from adjacent ditch, 33°59'N 90°48'W, T25N

ROW S34, 12 Aug 2004, Skojac 971 (SWSL, shl). Leflore Co.: NW of Sidon, between Sidon cut-off and old Yazoo River run, T18N RIE

$19, 31 Aug 1998, Bryson 16654 et al. (MISS, SWSL).

Eichhornia crassipes (Mart.) Solms (Pontederiaceae). Common water hyacinth (USDA, NRCS 2006) is

an exotic aquatic weed native to S. America. It has invaded most southern states, where it is a problem in

waterway canals, ditches, and ponds. McCook and Kartesz (2000) report this invasive aquatic weed in seven

counties in southern Mississippi and NRCS (2006) reports it from an additional two counties. Our vouchers

are a significant range extension northward in the state, and represent the first records of this species in the

Yazoo-Mississippi Delta Region.

Voucher specimen: U.S.A. MISSISSIPPI. Issaquena Co.: ca. 5 air mi NE of Mayersville at jct. of Grace Road and Steele Bayou, 30 Oct

2006, Bryson 21940 & Hoagland (DOV, SWSL, VSC). Leflore Co.: ca. 6 mi SW of Greenwood on S end of Lake Roebuck and just N of

Leflore County Road 512, 31 Oct 2006, Bryson 21943 (DOV, MISS, MISSA, MMNS, SWSL, VSC). Washington Co.: SW of Greenville

at Warfield Point Park, sandbar along Mississippi river, TIBN ROW S13, 10 Oct 2005, Walker MS11 (shl); ca. 2.2 air mi E of Glen Allan

at jct. of Hwy MS 1 and Steele Bayou, 30 Oct 2006, Bryson 21938 & Hoagland (DOV, MISS, MISSA, SWSL, VSC); ca. 5.8 mi NE of Glen

Allan at jct. of Bear Garden Road and Steele Bayou, 30 Oct 2006, Bryson 21942 & Hoagland (MISSA, SWSL).

Hottonia inflata Ell. (Primulaceae). American featherfoil (USDA, NRCS 2006) is a submersed aquatic

native to the eastern half of the United States. Where observed, it is known to be sporadic in appearance

from year to year. In Mississippi, the species is of special concern and is listed on the state’s special plant

tracking list (Mississippi Natural Heritage Program 2002). McCook and Kartesz (2000) report this species

from neighboring Quitman County in the Delta, and a second voucher is reported from the eastern part of

the state in Oktibbeha County by USDA, NRCS (2006). Our voucher represents only the third published

account of this species in the state, and doubles the number of sites within the Yazoo-Mississippi Delta

Region where this species occurs.

Voucher specimen: U.S.A. MISSISSIPPI. Tunica Co.: ca. 5.5 mi S of Tunica; lake parallel to Margie Rd., T5S R11W S33, growing in

lake, 3 May 2006, Bryson 21414 & Skojac (ALA, DOV, FLAS, FSU, GH, LSU, MICH, MISS, MISSA, MMNS, MO, NY, SWSL, TENN, UARK,

US, USCH, USMS, VDB, VPI, VSC, WIN), Skojac 1266 & Bryson (shl).

Menispermum canadense L. (Menispermaceae). Common moonseed (USDA, NRCS 2006) is considered

rare in Mississippi (Morris et al. 1993) and is listed on the state’s special plant tracking list (Mississippi

Natural Heritage Program 2002). It was first reported from the Yazoo-Mississippi Delta Region by Carter et

AA: H : «Male D

rr wd

Skojac et al., Yazoo floristic records 773

al. (1990) from Bolivar, Coahoma, and Sunflower counties. Two other stations in Bolivar County are listed

in McCook and Kartesz’s (2000) checklist, representing the only other published accounts of this species in

the Yazoo-Mississippi Delta Region. Our voucher from Washington County provides an additional county

record for this woody vine rare to Mississippi.

Voucher specimen: U.S.A. MISSISSIPPI. Washington Co.: Delta Experimental Forest, along eastern side of woods directly N of W

entrance onto Forest via Feather Farms Rd., T19W R7W S33, bottomland hardwood forest, 6 Nov 2004, Skojac 1091 & M. Skojac (shl).

Oplismenus setarius (L am.) Roem. & Schult. (Poaceae). Bristle basketgrass (USDA, NRCS 2006) is reported

in several southern counties within the state by McCook and Kartesz (2000). It was not, however, reported

in Carter’s (1978) flora of Delta National Forest, most likely due to the rather isolated population from which

the Sharkey County material was collected. Our vouchers represent a significant range extension northward

in the state and are the first records of this species in the Yazoo-Mississippi Delta Region, where it appears

restricted to well-drained ridge sites.

Voucher specimens: U.S.A. MISSISSIPPI. Sharkey Co.: Delta National Forest, Sweetgum Research Natural Area, T12N R5W S9, old-

growth bottomland hardwood forest, 19 Sep 2003, Skojac 775 (shl). Washington Co.: Yazoo National Wildlife Refuge, wooded tract S

of Alligator Pond along extant ridge bordering slough, 17 Aug 2004, Skojac 980 (shl).

Podophyllum peltatum L. (Berberidaceae). This is the first published record of mayapple (USDA, NRCS

2006) from the Yazoo-Mississippi Delta Region. Its principle distribution within the state lies east of the

Delta in the rich deciduous woods of the Loess Bluffs (McCook & Kartesz 2000; USDA, NRCS 2006). It is

common elsewhere in the state as well, except in the extreme lower coastal plain, where it occurs on mesic

sites in upland hardwood forests and occasionally on ridge sites within minor stream-bottom forests.

Voucher specimen: U.S.A. MISSISSIPPI. Tunica Co.: ca. 6 mi S of Tunica; along Margie Rd., T5S R11W S33, on elevated ridge-bottom

parallel to lake, 3 May 2006, Bryson 21417 & Skojac (DOV, SWSL, VSC); Skojac 1272 & Bryson (shl).

Polystichum acrostichoides (Michx.) Schott (Dryopteridaceae). Specimens of Christmas fern (USDA, NRCS

2006) were collected from a small group of plants growing on an elevated ridge site with notably higher relief

than the surrounding terrain. This single isolated station was not reported in Carter’s (1978) flora of Delta

National Forest. This voucher represents the first record of Christmas fern from the Yazoo-Mississippi Delta

Region. It is common elsewhere in the state, except in the extreme lower coastal plain, usually occurring in

upland hardwood and mixed pine-hardwood forests.

Voucher specimen: U.S.A. MISSISSIPPI. Sharkey Co.: Delta National Forest, approx. 4 mi S of Holly Bluff, F.S. compartment 39, T11N

R5W S34, 7 Nov 2005, Skojac 1198 (shl).

Proserpinaca palustris L. (Haloragaceae). Marsh mermaidweed (USDA, NRCS 2006) is a polymorphic,

aquatic to semi-aquatic herb of ponds, sloughs, and seasonally inundated depressions. It has a sporadic

distribution within Mississippi, occurring in seven counties ranging from the extreme northern county of

Alcorn to the coastal counties of Hancock and Jackson (McCook & Kartesz 2000; USDA, NRCS 2006). Our

voucher is the first collection of the species from a county within the Yazoo-Mississippi Delta Region, where

it was found in a depression at the edge of a beaver damaged area of bottomland hardwoods.

Voucher specimen: U.S.A. MISSISSIPPI. Washington Co.: ca. 5.0 mi W of Hollandale on Hwy MS 12 at Leroy Percy State Park; ca. 0.3

mi W of Park entrance in woods S of Hwy 12 and E of West Park Rd., T15N R7W S17, 12 Jun 2006, Skojac 1342 (SWSL, shl).

Scirpus atrovirens Willd. (Cyperaceae). Green bulrush (USDA, NRCS 2006) is locally common in the

northern third of Mississippi, excluding the Yazoo-Mississippi Delta Region. McCook and Kartesz (2000)

and USDA, NRCS (2006) list S. atrovirens from six north-central Mississippi counties outside of the Yazoo-

Mississippi Delta Region. This voucher represents the first record of this species in the Yazoo-Mississippi

Delta, where it is apparently restricted to wet open areas adjacent to aswamp predominated with Taxodium

distichum (L.) Richard and Salix nigra Marshall.

Voucher specimen: Bolivar Co.: SE of Rosedale, ca. 1.5 mi SE jct. Hwy MS 1 and MS 8, 17 Jun 1996, Bryson 15343 & Carter (DOV, MISS,

MISSA, MMNS, MO, SWSL, VSC).

£s+haD H ID L

774 Journal of t titute of Texas 1(1)

Senecio vulgaris L. (Asteraceae). Common groundsel (WSSA 1989) or old-man-in-the-spring (USDA,

NRCS 2006) is a common weed of agricultural and non-agricultural areas throughout most of the United

States; however, it was undetected in the Yazoo-Mississippi Delta Region by Gunn et al. (1980) and others.

McCook and Kartesz (2000) and USDA, NRCS (2006) report S. vulgaris from Mississippi but do not pro-

vide data. This voucher represents the first collection of this species in the Yazoo-Mississippi Delta Region,

where it was found locally on clay soils in no- and reduced-tillage cotton and soybean fields in association

with Alopecurus carolinianus Walt., Bowlesia incana, Conyza canadensis (L.) Cronq., Coronopus didymus (L.)

Sm., Lamium amplexicaule L., Myosurus minimus L., Oenothera laciniata Hill, and other early season weeds

commonly found in reduced-tillage row crop production systems.

Voucher specimen: U.S.A. MISSISSIPPI. Washington Co.: 2.5 mi NE of Stoneville, W side of Napanee Rd., T19N R7W sect. 36; lo-

cally common weed on edge of no-till soybean field, 6 Apr 2006, Bryson 20617 (ALA, DOV, MICH, MISS, MISSA, MMNS, MO, SWSL,

UARK, USMS, VDB, VSC, shl).

Sparganium americanum Nutt. (Sparganiaceae). Threesquare burreed (WSSA 1989) or American bur-

weed (USDA, NRCS 2006) is reported from 9 Mississippi counties (McCook & Kartesz 2000; USDA, NRCS

2006) excluding the Yazoo-Mississippi Delta Region. This voucher represents the first collection of this

species to the Yazoo-Mississippi Delta Region, where it was found locally in a ditch and small oxbow lake

in association with Hottonia inflata and Hydrocotyle ranunculoides L. and surrounded by Taxodium distichum,

Salix nigra and Styrax americana Lamarck.

Voucher specimen: U.S.A. MISSISSIPPI. Tunica Co.: 5.5 mi S of Tunica; lake parallel to Margie Rd., T5S R11W S33, growing in lake,

3 May 2006, Bryson 21416 & Skojac (DOV, SWSL, VSC), Skojac 1267 & Bryson (shl); 3 Oct 2006, Bryson 21928 (MISS, MISSA, SWSL).

Spigelia marilandica L. (Loganiaceae). Indian pink or woodland pinkroot (USDA, NRCS 2006) isa common

herb of rich woodland sites. In Mississippi, it occurs on mesic upland sites and on elevated ridges within

minor stream-bottom systems. This species has wide distribution throughout the state but is not reported

from the Yazoo-Mississippi Delta Region (McCook & Kartesz 2000). Our voucher is the first record of this

species from the region, where it appears restricted to well-drained ridge bottom sites.

Voucher specimen: U.S.A. MISSISSIPPI. Tunica Co.: ca. 6 mi S of Tunica along Margie Rd., T5S R11W S33, elevated ridge-bottom

parallel to lake, 3 May 2006, Skojac 1274 & Bryson (shl).

Typha angustifolia L. (Typhaceae). Narrowleaf cattail (USDA, NRCS 2006) has limited distribution in the

southeastern U.S. The species is reported from the neighboring states of Arkansas, Louisiana and Tennessee

(USDA, NRCS 2006), but is not included in the preliminary checklist of Mississippi plants being compiled

by McCook and Kartesz (2000). Our voucher from the Yazoo-Mississippi Delta Region is the first record of

the species in the state of Mississippi.

Voucher specimen: U.S.A. MISSISSIPPI. Washington Co.: 0.5 mi S of southern entrance onto Delta Experimental Forest, in ditch on

E side of Experiment Station Rd., 28 May 2003, Skojac 724 (shl).

ACKNOWLEDGMENTS

We thank Richard Carter, J. Paige Goodlett, Robert Hoagland, John R. MacDonald, Sidney McDaniel, Robert

Stewart, and Randy Warren for help with field work. The authors thank Samuel W. Rosso, Lucile M. McCook

and an anonymous reviewer for helpful comments on the manuscript.

REFERENCES

Bryson, C.T. and C.D. Eumore. 1991. Two weedy species, Ammoselinum butleri (Umbellifereae) and Lepidium aus-

trinum (Crucifera), new to Mississippi. Sida 14:506-508.

Bryson, C.T. and S.D. Jones.1990. Carex comosa (Cyperaceae), new to Mississippi. Sida 14:311-312.

Bryson, C.T. and R. Carter. 1992. Notes on Cyperus and Kyllinga (Cyperaceae) in Mississippi with records of six

species new to the state. Sida 15:119-124.

* rh n D

Skojac et al., Yazoo-Mississi floristic records 775

Bryson, C.T., REC. Naczi, and S. McDanteL. 1992. Notes on noteworthy records of Carex (Cyperaceae) from the

southeastern United States. Sida 15:125-135.

Carter, J.R. 1978. A floristic study of the Delta National Forest and adjacent areas. M.S. thesis, Department of

Botany, Mississippi State University, Mississippi State. 79 p.

CARTER, J.R., M.W. Morris, and C.T. Bryson. 1990. Some rare or otherwise interesting vascular plants from the Mis-

sissippi Delta Region of Mississippi. Castanea 55:40—55.

GUNN, C.R., T.M. PULLEN, E.A. STADELBACHER, J.M. CHANDLER, and J. Barnes. 1980. Vascular flora of Washington County,

Mississippi, and environs. Science and Education Administration, U.S. Department of Agriculture, New

Orleans, LA.

HoLMGREN, P.K., N.H. Hot waREN, and L.C. Barnertr (eds.). 1990. Index herbariourm. Part 1: The Herbaria of the world,

8^ ed. New York Botanical Garden, Bronx.

HOLMGREN, PK. and N. H. HoLmGrEn. 1998 onwards (continuously updated). Index herbariorum. New York Botanical

Garden. Web site: http://sciweb.nybg.org/science2/IndexHerbariorum.asp

Lite, E.L. 1971. Atlas of United States Trees. Vol. I. Conifers and important hardwoods. U.S. Department of Agri-

culture, Forest Service Misc. Publ. No. 1146. U.S. Government Printing Offices, Washington, D.C.

Lowe, E. N. 1921. Plants of Mississippi. Mississippi State Geological Survey. Bulletin No.17. 292p.

McCook, L.M., and J. Kartesz. 2000. A preliminary checklist of the plants of Mississippi. Website: herbarium.olemiss

edu/checklist.html.

Mississippi NATURAL HERITAGE PROGRAM. 2002. Special Plants Tracking List. Museum of Natural Science, Mississippi Dept.

of Wildlife, Fisheries, and Parks, Jackson, MS. 8 pp. Website: mdwfp.com/museum/downloads/plant. tracking.

Morris, M. W. 1989. Spiranthes (Orchidaceae) in Mississippi. Selbyana 11:39-48.

Morris, M.W., C. T. Bryson, and R.C. Warren. 1993. Rare vascular plants and associate pl'ant communities from Sand

Creek chalk bluffs, Oktibbeha County, Mississippi. Castanea 58:250-259,

RaaLER, R.K. and C.T. Bryson. 1990. Montia linearis (Portulaceae), new to Mississippi. Sida 14:310-311.

Rosen, D.J., R. CARTER, and C.T. Bryson. 2006. The spread of Cyperus entrerianus (Cyperaceae) in the southeastern Unit-

ed States and its invasive potential in bottomland hardwood forests. Southeastern Naturalist 5:333-344.

USDA, NARCS. 2006. Plants database. http://plants.usda.gov. Accessed October 2006.

Wiseman, J.B., Jr. 1982. A study of the composition, successional relationships, and floristics of Mississippi River

floodplain forests in parts of Washington, Bolivar, and Sharkey counties, Mississippi. PhD dissertation, Depart-

ment of Biological Sciences, Mississippi State University, Mississippi State.

WSSA. 1989. Composite list of weeds. Weed Sci. Soc. Amer. Champaign, IL.

776 Journal of the Botanical R h Institute of Texas 1(1)

ANNOUNCEMENTS

THE 2007 DELZIE DEMAREE TRAVEL AWARD

Applications for the 2007 Delzie Demaree Travel Award should include a letter from the applicant telling how symposium attendance

will benefit his/her graduate work and letter of recommendation sent by the major professor. Please send letters of application to: Dr.

Donna M.E. Ware, P.O. Box 8795, Herbarium, Biology Department, The College of William and Mary, Williamsburg, VA 23185-8795,

U.S.A. 1-757-221-2799; Email: ddmware@wm.edu. The period for receiving applications will end three weeks prior to the date of the

symposium if a sufficient number of appli in hand at that time. Anyone wishing to apply after that date should inquire whether

applications are still being accepted before applying. The Systematics Symposium dates for 2006 are 12-13 October.

he Delzie Demaree Travel Award was established in 1988 honoring Delzie Demaree who attended 35 out of a possible 36 sym-

posia before he died in 1987. Delzie Demaree was a frontier botanist, explorer, discoverer, and teacher. His teaching career as a botanist

began in Arkansas at Hendrix College in 1922. He also taught botany at the University of Arkansas, Navajo Indian School, Yale School of

Forestry, Arkansas A&M, and Arkansas State University at Jonesboro where he retired as professor emeritus in 1953. One of the things

he enjoyed most as a botanist was assisting students with their field botany research.

THE RUPERT BARNEBY AWARD

The New York Botanical Garden is pleased to announce that Valquiria Ferreira Dutra, of the Universidade Federal de Vicosais the recipient

of the Rupert Barneby Award for the year 2007. She will be studying the genus Mimosa in the “campos rupestres" in Eastern Brazil.

The New York Botanical Garden now invites applications for the Rupert Barneby Award for the vee 2008. The award of US$

1,000.00 is to assist researchers to visit The New York Botanical Garden to study the rich collection of I (Fab ). Anyone

interested in applying for the award should submit their curriculum vitae, a detailed letter describing the project for which the award is

sought, how a visit to the NYBG would help mplish the goals of the project, and the names of 3 referees. Travel to the NYBG should

be planned for sometime in the year 2008. The HE should be addressed to Dr. Fabián A. Michelangeli, (e-mail: fabianOnybg.

org). Institute of Systematic Botany, The New York Botanical Garden, 200th Street and Kazimiroff Blvd., Bronx, NY 10458-5126 USA,

and received no later than December 1, 2007. Electronic applications are preferred. Announcement of the recipient will be made by

December 15th. Anyone interested in making a contribution to THE RUPERT BARNEBY FUND IN LEGUME SYSTEMATICS, which

supports this award, may send their check, payable to The New York Botanical Garden, to Dr. Michelangeli.

ENHANCED SPELLEX BOTANICAL DICTIONARY

NOW COMPATIBLE WITH 2007 MICROSOFT OFFICE

Spellex C | he rel f Spellex Botanical 2007, their popular botanical ae software. The new release includes

more than 50 ,000 oO bs m the botanical industry and new 2007 Mg Office Suite compatibility.

Benefits: The bot lli ff 1l l pap ical dictionaries and word books. By adding

a Botanical 2007 to 2007 Microsoft Office's English language spell checker, mos will verify correctly spelled botanical words and

de correct spelling choices for incorrectly spelled terms. Both the standard English language dictionary and the Spellex specialty

alone, are E simultaneously with one mouse click.

h fe

an now be combined with the new features of 2007 Microsoft Office, which allow you to:

—Find EE flag annoying ocn eei) spelling errors.

—Check all 2007 applications including Word, Excel, Access, Outlook, and more.

—Share custom dictionaries with others, which allow you to add your own words to the spell checker.

— Create exclusion dictionaries, which allow you to flag words that you wish to avoid using.

—Perform global changes to your spelling options. Your changes will now be saved across the entire Microsoft Office suite.

Features: Spellex Botanical is a comprehensive botanical terminology spell checker which includes the correct spelling of tens of thou-

sands of vascular plants, mosses, liverworts, hornworts, and lichens grown around the world.

Spellex Botanical includes the spelling of thousands of common plant names, scientific names and botanical names as well as plant

families, B m n symbols. This comprehensive speller even includes the correct spelling of a wide ce! i peu) d

common medicinal plants and plants used for habitat creation and |

includes common names and scientific names of hundreds of plant diseases including bacterial deum fungal FUE nematodal

and parasitic diseases, genetic diseases, phytoplasmal diseases and much more!

Spellex Botanical dictionaries start at $64.95. Multiuser licenses and educational discounts are available. Spellex dictionaries are

compatible with Microsoft, most Windows programs, developer tools, custom applications, Web browsers, and are available in both US

English and UK English.

To request product information or to place an order, contact Spellex Corporation at 800-442-WORD (9673) or 813-792-7000.

You can also visit www.spellex.com/products/biotech.htm.—Spellex Corporation, Attn: Melissa Rossi, 10820 Sheldon Road, Tampa,

FL 33626, U.S.A. mrossi@spellex.com.

J. Bot. Res. Inst. Texas 1(1): 776. 2007

TALINUM RUGOSPERMUM (PORTULACACEAE) NEW TO OKLAHOMA

Christopher S. Reid Barbara R. MacRoberts

Patricia L. Faulkner Michael H. MacRoberts

Louisiana Natural Heritage Program Bog Research, 740 Columbia

Louisiana Department of Wildlife and Fisheries Shreveport, Louisiana 71104 U.S.A. and

000 Herbarium, Museum of Life Sciences

Baton Rouge, Louisiana 70898-9000 U.S.A. Louisiana State University-Shreveport

Shreveport, Louisiana 71115 U.S.A.

ABSTRACT

Talinum rugospermum Holz. (Portulacaceae) is reported new to Oklahoma. It was discovered at The Nature Conservancy’s Boehler Seeps

and Sandhills Preserve in Atoka County.

RESUMEN

Se cita cono nuevo para Oklahoma Talinum rugospermum Holz. (Portulacaceae). Se ha descubierto en la reserva Nature Conservancy's

Boehler Seeps and Sandhills en el condado de Atoka.

Talinum rugospermum Holz. has not been reported from Oklahoma although it occurs in Texas, Arkansas,

Kansas, and Louisiana (Cochrane 1993; Tayor & Taylor 1994; Singhurst 1996; MacRoberts & MacRoberts

1997; Kartesz & Meacham 1999; Kiger 2003; USDA, NRCS 2006; NatureServe 2006; Oklahoma Natural

Heritage Inventory 2003; Hoagland et al. 2004, Bruce Hoagland pers. comm., Jason Singhurst, pers. comm.).

On May 18 and 19, 2006, we found over one hundred T. rugospermum plants in flower in xeric sandylands at

The Nature Conservancy’s Boehler Seeps and Sandhills Preserve, Atoka County, Oklahoma. Also known as a

bluejack oak sandhill community or Quercus incana-Quercus stellata woodland association, xeric sandylands

occur in eastern Texas, southern Oklahoma, southwestern Arkansas, and western Louisiana, and are rare

in all but Texas (Jones 1993; Hoagland 2000; MacRoberts et al. 2002a; Diggs et al. 2006). Xeric sandylands

are characterized by deep sands, an open understory, sparse vegetation, and a distinct assemblage of plants,

many of which are endemic to the West Gulf Coastal Plain (MacRoberts et al. 2002a, 2002b). In Oklahoma,

Boehler Seeps and Sandhill Preserve is the best remaining site for this community (Jones & Carpenter 1995;

Gatti Clark 1997). The global conservation status rank for T. rugospermum assigned by NatureServe (2006)

is G3G4, meaning that it is either vulnerable across its range (G3) or apparently secure (G4). Associate spe-

cies at the site included Cnidoscolus texanus, Collinsia violacea, Commelina erecta, Eriogonum longifolium, Evax

prolifera, Loeflingia squarrosa, Opuntia humifusa, Paronychia drummondii, Phacelia strictiflora, Quercus incana,

Selaginella arenicola ssp. riddellii, and Stylisma pickeringii.

Voucher specimen: U.S.A. Oklahoma. Atoka Co.: Boehler Seeps and Sandhills Preserve, just W of Boehler, SE1/4 SE1/4 $25 and NE1/4

NE1/4 S36, T4S R13E. 18 May 2006, Reid, Faulkner, MacRoberts, MacRoberts 5754 (LSU, OKL)

ACKNOWLEDGMENTS

We thank Amy Buthod and Lawrence K. Magrath for reviewing the manuscript and providing helpful feed-

back. Bruce Hoagland, Oklahoma Biological Survey, and Jason Singhurst, Texas Parks and Wildlife Depart-

ment, provided information on the distribution of T. rugospermum. We appreciate The Nature Conservancy

of Oklahoma for allowing access to the property.

REFERENCES

COCHRANE, T.S. 1993. Status and distribution of Talinum rugospermum Holzinger (Portulacaceae). Nat. Areas J.

13:33-41.

J. Bot. Res. Inst. Texas 1(1): 777 — 778. 2007

778 Journal of the Botanical R h Institute of Texas 1(1)

Dices, G.M., B.L. Lirscome, M.D. REED, and R.J. O’KENNON. 2006. Illustrated flora of east Texas. Sida, Bot. Misc. 26:1-1594.

Garm CLark, L.C. 1997. Floristic and biosystematic investigations in plant taxonomy. Ph.D. Dissertation. Okla-

homa State University, Stillwater.

HOAGLAND, B. 2000. The vegetation of Oklahoma: classification for landscape mapping and conservation planning.

SouthW. Naturalist 45:385-420.

HOAGLAND, B.W., A.K. ButHob, I.H. Butler, PH.C. CRAwroRD, A.H. Upasi, W.J. Euisens, and RJ. TvrL. 2004. Oklahoma Vas-

cular Plants Database (http://geo.ou.edu/botanical), Oklahoma Biological Survey, University of Oklahoma,

Norman.

Jones, N. 1993. A baseline study of the water quality, vegetative gradients, and hydrology of Boehler Seeps and

Sandhills Preserve. Unpublished report. The Nature Conservancy, Tulsa, OK.

Jones, N. and B. Carpenter. 1995. Boehler Seeps and Sandhills site conservation plan. Unpublished report. The

Nature Conservancy, Tulsa, OK.

KARTESZ, J.T. and C.A. MEAcHAM. 1999. Synthesis of North American flora. Version 1.0. North Carolina Botanical

Garden, Chapel Hill.

KiGER, R.W. 2003. Phemeranthus. In: Flora of North America Editorial Committee, eds. Flora of North America North

of Mexico. Vol. 4. New York. Pp. 488-495.

MacRoserts, M.H. and B.R. MacRoberts. 1997. Talinum rugospermum Holz., new to Louisiana with notes on terete-

leaved Talinum in Louisiana. Phytologia 82:86-93.

MacRostRrs, B.R., M.H. MacRoserts, and J.C. CatHey. 2002a. Floristics of xeric sandylands in the post oak savanna

region of east Texas. Sida 20:373-386.

MacRogerTs, M.H., B.R. MAcRoserts, B.A. Sonne, and R.E. Evans. 2002b. Endemism in the West Gulf Coastal Plain:

importance of xeric habitats. Sida 20:767-780.

NatureServe. 2006. NatureServe Explorer: An online encyclopedia of life [web application]. Version 4.7. NatureServe,

Arlington, Virginia. Available http://www.natureserve.org/explorer. (Accessed: July 3, 2006).

OKLAHOMA NATURAL HERITAGE INVENTORY. 2003. Working list of rare Oklahoma plants. Oklahoma Biological Survey,

University of Oklahoma, Norman.

SINGHURST, J.R. 1996. The status of nine endangered plant species of east Texas: historical, ecological, and phyto-

geographical notes. M.S. Thesis, Stephen F. Austin State University, Nacogdoches, TX.

TAYLOR, R.J. and C.S.E. TavLor. 1994. An annotated list of the ferns, fern allies, Gymnosperms and flowering plants

of Oklahoma. Southeastern Oklahoma State University, Durant.

USDA, NRCS. 2006. The PLANTS Database (http://plants.usda.gov, 3 July 2006). National Plant Data Center, Baton

Rouge, LA 70874-4490 USA.

ADDITIONAL NOTEWORTHY COLLECTIONS OF CYPERUS DRUMMONDIL

(CYPERACEAE) PROM TEXAS AND FIRST REPORT FROM MEXICO

David J. Rosen Richard Carter

S. M. Tracy Herbarium

Department of Ecosystem Science & Management Herbarium, Department of Biology

Texas A&M University Valdosta State University

College Station, Texas 77843-2138, U.S.A. Valdosta, Georgia 31698, U.S.A.

ABSTRACT

Recent noteworthy collections of Cyperus drummondii from the Texas Gulf Prairies and Marshes, Piney Woods Natural Regions, and a

first record from Mexico are reported.

RESUMEN

Se citan colecciones recientes notables de of Cyperus drummondii de las paraderas y charcas del Golfo de Texas, Piney Woods Natural

Regions, y una primera cita de México.

Collections of Cyperus drummondii Torr. & Hook. from the Gulf Coast Prairies and Marshes as defined by

Gould (1975) from Texas counties not previously reported by Rosen (2004) were made during field work

in 2004-2007 as follows:

TEXAS. Fort Bend Co.: Buffalo Creek Unit of the San Bernard National Wildlife Refuge, S of FM 442, 7.2 km W of its intersection with

Hwy. 36, between the towns of Boling and Needville, 10 May 2007, Rosen 4114 (BRIT, TAES, TEX, VSC).

At this location, Cyperus drummondii occurred frequently in prairie wetlands on clayey-loam soils with Carex

aureolensis Steud., C. festucacea Schkuhr ex Willd., C. triangularis Boeck., Cyperus reflexus Vahl, Eleocharis sp.,

Juncus acuminatus Michx., J. brachycarpus Engelm., J. marginatus Rostk., Polygonum sp., Sesbania drummondii

(Rydb.) Cory., and Steinchisma hians (Elliott) Nash.

Victoria Co.: McFaddin Ranch, about 2.8 mi E of the jct. of U.S. Hwy. 77 and FM 445 in the town of McFaddin, 11 Sep 2004, Rosen

3056 & Carter (MICH, TEX); Carter 15438 & Rosen (VDB, VSC, others to be distributed).

At this location, Cyperus drummondii was locally abundant in a large prairie wetland on clayey-loam soils of

the Lissie Formation with Acacia farnesiana (L.) Willd., Andropogon gerardii Vitman, Cyperus spp., Eleocharis

ravenelii Britton, Leersia hexandra Sw., Paspalum spp., Polygonum sp., Prosopis glandulosa Torr., and Rhyncho-

spora spp.

Reports of Cyperus drummondii from the Piney Woods in the Illustrated Flora of East Texas, Vol. 1. (Diggs et al.

2006) are based on two very old collections from the southeast margin of East Texas (Rosen 2004). Recent

field work in Hardin County has resulted in collections of C. drummondii from a hillside seepage bog and

the margins of a flat-woods pond as cited here:

TEXAS. Hardin Co.: Roy E. Larsen Sandyland Sanctuary, N of State Hwy. 327, between the towns of Silsbee and Kountze, 08 Oct 2006,

Rosen 3949 & Brown and Boensch (BRIT, MICH, TAES, TEX, VSC).

The label data from the historic collections of Cyperus drummondii from eastern Texas provide no details

on its habitat (Rosen 2004). We expand the habitat description of this species in Texas to include hillside

seepage bogs and flat-woods ponds, in addition to the relict Coastal Prairie wetlands noted by Rosen (2004).

At the Roy E. Larsen Sandyland Sanctuary, C. drummondii was occasional to frequent on sandy soils in

shallow water on the fringes of a flatwoods pond and wet sandy soils of a hillside seep with Dichanthelium

sp., Eleocharis spp., Fuirena sp., Morella cerifera (L.) Small, Panicum hemitomon Schult., Saccharum giganteum

(Walter) Pers., and Scirpus cyperinus (L.) Kunth. This is very similar to the habitat of this species in the Gulf

Coastal Plain of Georgia and Mississippi (Carter et al. 1999).

J. Bot. Res. Inst. Texas 1(1): 779 — 780. 2007

£+sha D o ID L

780 Journal of t titute of Texas 1(1)

Denton (1978) reported Cyperus drummondii [as C. virens Michx. var. drummondii (Torr. & Hook.) Kük.] as

occurring in North America (Texas and Louisiana), Nicaragua, Jamaica, the Galapagos Islands, Surinam,

and Brazil. Tucker (1994) and Espejo Serna and López-Ferrari (1997) included C. drummondii under C.

virens in Mexico. During field work near the city of Durango following the Second Botanical Symposium of

Northern Mexico, we collected specimens of C. drummondii, confirming its occurrence in Mexico. Review of

specimens at CIIDIR also resulted in the discovery of other collections of C. drummondii from the Mexican

state of Durango previously identified as C. virens.

Specimens examined. MÉXICO. Durango: proximidad a Canelas, terrenos de la UAF Topia, Mpio. Canelas, montemojino, transición

entre el bosque tropical caducifolio y el bosque de pino, 1414 m snm, suelo profundo, café muy pedregoso, con pendientes hasta de 45°,

27 Sep 1990, Benítez 2409 (CIIDIR); Carretera Durango-Nombre de Dios, Km 246.5, Mpio. Nombre de Dios, Aug1997, matorral xerófilo,

González 354 (CIIDIR); 17 km al E de Durango por la carretera a Fresnillo, lugar inundado cerca de la carretera, localmente abundante,

1960 m, 18 Oct 1983, González & Acevedo 2747 (CIIDIR); Río Tunal, bajo puente en libramiento de carretera Zacatecas—Torreón (cerca

del Balneario San Juan), Mpio. Durango, 19 Jan 1999, Pinedo 24 (CIIDIR); 31 km de La Guajolota, por el camino a Los Charcos, Mpio.

El Mezquital, estanque en medio de bosque de pino-encino, escaso, dentro del agua, 2000 m, 8 Oct 1983, González 2701 con González

y Acevedo (CIIDIR); Carretera 45 al E de la Ciudad de Durango (Puente Gavilanes), Mpio. Durango, 24°0'51" N, 104°29'25" W, 1850 m,

vegetación riparia (Salix bonplandiana), común, hierba, 08 Oct 1999, González 3195 (CIIDIR); Mpio. Durango, 17.7 km al SSE de Du-

rango, por la carretera a El Mezquital, al S de Felipe Ángeles, 23°55'21" N, 104°32'40" W, 1850 m, vegetación subacuática, abundante

a orilla de canal, 16 Sep 2005, González 7091 con Guaglianone, Torres, Rosen, Carter y Peterson (CIIDIR, ANSM, ENCB, IEB, MEXU, SD;

Carter 16149, 16150 & González, Guaglianone, Torres, Rosen, & Peterson (VDB, VSC, others to be distributed); Rosen 3493, 3494 & Carter,

González, Guaglianone, Torres, & Peterson (TAES).

The following key modified from Denton (1978) and Carter et al. (1999) will separate Cyperus drummondii

from other members of the C. virens complex in Mexico.

1. Primary peduncles 3-5; floral scales 1.0-1.5(-1.8) mm long; spikelets at least 2.25 mm wide__________ C.drummondii

1. Primary peduncles 6-12(-14); floral scales 1.5-2.4 mm long; spikelets (2-)2.2-3.3 mm wide.

2. Achenes (2.5-)3-5 times longer than wide, (1-)1.2-1.5 mm long; spikelets (5-)7-15 mmlong ^ | | Q. virens

var. virens

2. Achenes 2-2.5 times longer than wide, 1-1.2 mm long; spikelets 5-6.5 mm long C. virens

var. minarum

ACKNOWLEDGMENTS

We are grateful to Wade Harrell with the Nature Conservancy's Refugio-Goliad Prairie Project and Kirk

Feuerbacher with the McFaddin Ranch for arranging access for field work in Victoria County, and to Bob

Boensch and Wendy Ledbetter with the Nature Conservancy for access to the Roy E. Larsen Sandyland

Sanctuary. We would like to thank Guy Nesom for reviewing this manuscript and Socorro Gonzalez-Eli-

zondo for reviewing this manuscript and the kind hospitality and her indispensable assistance, without

which the field work resulting in discovery of this species in México would not have been possible. The

Valdosta State University Foundation and Faculty Internationalization Fund are gratefully acknowledged

for financial support.

REFERENCES

CARTER, R., D.K. ALEXANDER, C. T. Bryson, and A. Lazari. 1999. The taxonomy of Cyperus virens and Cyperus drummondii

in the southeastern United States. Sida 18:1049-1063.

Denton, M.F. 1978. A taxonomic treatment of the Luzulae group of Cyperus. Contr. Univ. Michigan Herb. 11:

197-271.

Dices, G.M. Jr, B.L. Liescome, M.D. Reep, and R.J. O'KeNNoN. 2006. Illustrated flora of East Texas, Vol. 1. Sida, Bot. Misc.

26.

ESPEJO SERNA, A. and A.R. Lopez-Ferrari. 1997. Las monocotiledóneas mexicanas Una sinópsis floristica. 1. Lista de

Referencia. Parte V Cyperaceae. Consejo Nacional de la Flora de México, A.C., México, D.F.

GouLb, F.W. 1975. Texas plants—a checklist and ecological summary. MP-585 Revised, Texas Agricultural Experi-

ment Station, College Station.

Rosen, D.J. 2004. Noteworthy collections of Cyperus drummondii (Cyperaceae) from Texas. Sida 21:495-497.

Tucker, G.C. 1994. Revision of the Mexican species of Cyperus (Cyperaceae). Syst. Bot. Monogr. 43:1-213.

LAWRENCE K. MAGRATH

1943-2007

Barney Lipscomb

Botani ical Research Institute of Texas

509 Pecan Street

Fort Worth, Texas 76102-4060, U.S.A.

It is with great sadness to report the loss of a friend and colleague Lawrence K. Magrath, biologist, orchi-

dologist and curator, who died Saturday, 24 February 2007, after a long illness. Magrath, a 35-year member

of the University of Science and Arts of Oklahoma (USAO) science faculty, previously served as professor

of biology, director of interdisciplinary studies and curator of the OCLA Herbarium (herbarium acronym).

Though he officially resigned March 2005, he continued teaching on a limited schedule.

Through the 1990s he led or supported one campus beautification project after another, adding flowers,

trees, and shrubbery to create the String of Pearls Garden surrounding Austin Hall, the Water Garden to

the north, the Sculpture Garden east of Davis Hall, and more. As a result of his leadership, the college won

two Keep Oklahoma Beautiful awards in the 1990s, and the USAO Board of Regents presented a special

commendation to him.

As a long-time leader in USAO’s unique Interdisciplinary Studies Program, Magrath was recognized

for classroom teaching, his dedication to liberal arts education, mentoring young scientists, and encourag-

ing learning both inside and outside the classroom. He advised several student organizations and academic

honor societies at USAO.

In 1994, Magrath was presented a plaque by the Oklahoma Junior Academy of Science (OJAS) for “Out-

standing Contributions and Long-time Service.” Magrath welcomed the OJAS conference to the USAO campus

on several occasions. Also during 1994, Magrath published three poems in the “Journal of Evolutionary

Psychology.” As one of Oklahoma’s authorities on Orchids in this region, he attended the American Orchid

Society Trustees Meeting in New York City to serve as vice chair for the AOS Conservation Committee. He

was a member of more than a dozen national and international societies in science, wildlife preservation,

and teaching. For decades, he worked with the Oklahoma Academy of Science, the American Orchid Society,

the Oklahoma Native Plant Society, the Flora of Oklahoma Project, and the Flora of North America Project,

authored or co-authored some 50 articles, read at least 30 papers at professional conferences.

Larry was born 28 March 1943 in Garnett, Kansas. He was salutatorian at Westphalia Rural High School

in Coffey County, Kansas. His bachelor’s and master’s degrees were from Kansas State Teachers College

(now Emporia State University). He received an undergraduate fellowship to Argonne National Laboratory

in Illinois. Larry was awarded his Ph.D. in botany in 1973 at the University of Kansas; his dissertation was

entitled “The native orchids of the prairies and plains region of North America.” Orchids remained the primary

focus of Larry’s research throughout his career. He developed the OCLA herbarium (ca. 22,000 specimens)

with noteworthy collections from southeastern Oklahoma and the Orchidaceae of the central and south-

eastern United States.

ACKNOWLEDGMENTS

Thanks to the USAO News Bureau (University of Science and Arts of Oklahoma) for permission to extract

information from the USAO website (http://www.usao.edu/news/spring-07/LarryMagrath.htm). Randy Talley,

Michael Bendure, and Charles Mather (USAO) provided the photograph. The Oklahoma Flora committee,

particularly Susan Barber and Wayne Elisens, gave encouragement and support. Judy MacKenzie kindly

read and suggested improvements to the manuscript.

J. Bot. Res. Inst. Texas 1(1): 781 — 784. 2007

782 Journal of the Botanical R h Institute of Texas 1(1)

SCIENTIFIC PUBLICATIONS AND REPORTS

MagRATH, L.K. 1969. The vascular flora of Coffey County, Kansas. Master's Thesis; Kansas State Teachers College,

Emporia.

MacRATH, L.K. 1970. Spiranthes tuberosa, new for Kansas. Rhodora 72:141.

MacRATH, L.K. 1971. Vascular plants new for Oklahoma, Kansas, and Nebraska. Rhodora 73:300-304.

MacRath, L.K. 1971. Opposing the construction of Hillsdale Reservoir. Hearings, House of Representatives, Public

Works Appropriations for 1972, 92nd congress, Ist Session, Part 5.

MacRATH, L.K. 1971. Native orchids of Kansas. Trans. Kan. Acad. Sci. 74(384):287-309.

Maarat, L.K. 1971. Identification of Smilax herbacea (Greenbriar). The Anderson Countian. Sept. 16, 1971, Sec-

tion 2, p. 7.

MacRATH, L.K. and K.L. Johnson. 1971. The genus Rhynchospora (Cyperaceae) in Kansas. SouthW. Naturalist

15389.

MacnATH, L.K. 1972. The Nelson Tract botanical survey, 1971. In: Barr, B.G., J.R. Eagleman, R.J. Eastmond, and S.A.

Morain. Atmospheric pollution and its effect in the Kansas City-Topeka Corridor. The University of Kansas.

NSF Grant #Gl-29760. The Center for Research, Inc.

MAGRATH, L.K. and R.R. Weepon. 1972. A report of fall-fruiting plants of Ophioglossum engelmannii Prantl in eastern

Kansas. Amer. Fern J. 62:22-23.

McGntGOR, R.L., L.K. MAGRATH, and R.R. Weebon. 1972. New and interesting plants from the Great Plains. Rhodora

74:378-388.

Weepon, R.R. and L.K. MacratH. 1973. New and interesting plants from the Great Plains. SouthW. Naturalist

18:341-342.

MacnATH, L.K. 1973. The native orchids of the Prairies and Plains Region of North America. Ph.D. Dissertation;

University of Kansas, Lawrence.

MacRath, L.K. 1974. The native orchids of the prairies and plains regions of North America. Diss. Abstr. Int., B 34.

(12):5838.

MacRath, L.K. and R.R. Weebon. 1974. New and interesting plants from the central plains States. Rhodora

76:489-490.

MAGRrATH, L.K. and J. Tavor. 1978. Orchids and other new and interesting plants from Oklahoma. Publ. Herb. S.E.

Oklahoma State Univ. 2:1-16.

MacRath, L.K. 1980. Listera australis Lindley (Orchidaceae) new for Oklahoma. SouthW. Naturalist 25:275.

MacRATH, L.K. 1983. Orchidaceae native to Oklahoma: distribution and ecology. Amer. J. Bot. 70:122.

MacRATH, L.K. 1984. New native orchids of Oklahoma and Kansas. Proc. Oklahoma Acad. Sci. 64:43-44.

MacRath, L.K. 1985. The native orchids of Oklahoma. Tulsa Garden Center Annual & Directory. Official Publication

of the Tulsa Garden Center. Pp. 21-26.

MacRath, L.K. 1987. Glaucium flavum Crantz (Papaveraceae) and Silybum marianum (L.) Gaertn. (Asteraceae) in

Oklahoma 12:243-244.

MacRath, L.K. and J.L. Norman. 1989. Nomenclatural notes on Calopogon, Corallorhiza, and Cypripedium (Orchida-

ceae) in the Great Plains Region. Sida 13:371—372.

MAGRATH, L.K., G. BuLmer, and |. SHarer. 1989, Dendrochilum javieriense, a new species in section Acoridium from

Luzon, the Philippines. Lindleyana 4:135-138.

MacnATH, L.K. 2001. Native orchids of Oklahoma. Crosstimbers 2001 (Spring):18-25.

MAGRATH, L.K. 2001. Native orchids of Oklahoma. Oklahoma Native Pl. Rec. 1:39-66.

MacraTH, L.K. 2001. Galium parisiense var. leiocarpum Tausch, new for Oklahoma. Oklahoma Native Pl. Rec.

TOIT.

MacRath, L.K. 2001. Schoenoplectus hallii var. S. saximontanus: 2000 Wichita Mountain Wildlife Refuge Survey.

Oklahoma Native Pl. Rec. 2(1):54-6467.

MacRATH, L.K. and R.J. COLEMAN. 2002. Listera. In: Flora of North America Editorial Committee, eds. Fl. North Amer.

26:586-592. Oxford Univ. Press., New York and Oxford.

Lawrence K. Magrath 783

Lawrence K. Magrath (1943-2007).

784 Journal of the Botanical R h Institute of Texas 1(1)

MagRATH, L.K., R.L. DRESSLER, J.T. Atwooo, and J. Beckner. 2002. Key to the genera of Orchidaceae. In: Flora of North

America Editorial Committee, eds. Fl. North Amer. 26:494-499. Oxford Univ. Press., New York and Oxford.

MacRath, L.K. and J.V. FREUDENSTEIN. 2002. Corallorhiza. In: Flora of North America Editorial Committee, eds. Fl. North

Amer. 26:633-638. Oxford Univ. Press., New York and Oxford.

MacRath, L.K. 2003. Triphora trianthophora and Tipularia discolor in Oklahoma. Oklahoma Native Pl. Rec. 3(1):

68-72.

SCIENTIFIC PRESENTATIONS/LECTURES

MacRath, L.K. 1970. The native orchids of Kansas. Paper read before Kansas Academy of Science.

MAGRATH, L.K. 1972. A checklist of native orchids occurring in Oklahoma, Kansas, Nebraska, South Dakota, and

North Dakota. Paper read before the Southwestern Association of Naturalists.

MacRath, L.K. 1973. Native orchids of Kansas. Paper read before the Kansas Orchid Society.

MacRath, L.K. 1976. The native orchids of Oklahoma. Paper read before the Oklahoma Academy of Science.

MAGRATH, L.K. and J. TayLor. 1977. Oklahoma orchids: new species and new distributions. Paper read before the

Oklahoma Academy of Science.

MacRath, L.K. 1978. Corallorhiza striata Lindley complex. Paper read before the Oklahoma Academy of Science.

MAGRATH, L.K. 1979. Native orchids of Oklahoma and adjacent regions. Banquet Address, 35th Annual Meeting

of the Southwestern Regional Orchid Growers Association.

MacRATH, L.K. 1980. Spiranthes praecox (Orchidaceae), new for Oklahoma. Paper read before the Oklahoma

Academy of Science.

MAGRATH, L.K. 1983. Orchidaceae native to Oklahoma: distribution and ecology. Paper presented at American

Society of Plant Taxonomists annual meeting.

MacnATH, L.K. 1983. A morphological comparison of two taxa of Calopogon occurring in Oklahoma. Paper read

before the Oklahoma Academy of Science.

MacRath, L.K. 1984. Orchids and other spring wildflowers. Presentation at 7th Annual Oklahoma Wildflower

Meeting.

MacRath, L.K. 1984. Cypripedium kentuckiense (Rafinesque's ladyslipper) a previously unrecognized ladyslipper in

southeastern Oklahoma. Paper read before the Oklahoma Academy of Science.

MacRath, L.K. 1985. Latest Royal Horticultural Society Awards in Cymbidium (1978-1984). Presentation made at

Great Plains Regional Judging Center of the American Orchid Society Meeting.

MaGRATH, L.K. 1986. Spaceship earth and the geopolitical implications of resource allocation and overpopulation.

Banquet Address, 1986 Model United Nations at University of Science & Arts of Oklahoma.

MAGRATH, L.K., W.W. WiLson, and J.D. Hurr. 1986. A macromorphological comparison of Cypripedium parviflorum

and C. kentuckiense in Oklahoma. Paper read before the Oklahoma Academy of Science.

NEW NAMES

Aster oblongifolius f. albaliqulatus Magrath—Rhodora 73:303. 1971.

Commelina erecta f. alba Magrath—Rhodora 73:300. 1971.

Cypripedium parviflorum f. albolabium Magrath & J.L.Norman—Sida 13:372. 1989.

Sida spinosa f. albiflora Magrath—Rhodora 74:383. 1972.

BOOK REVIEW

GERHARD GOTTSBERGER and ILSE SILBERBAUER-GOTTSBERGER. 2006. Life in the Cerrado: a South American Tropical

Seasonal Ecosystem. Vol. I. Origin, Structure, Dynamics and Plant Use. (ISBN 3-00-017928-

3, hbk.). Abteilung Systematische Botanikund Okologie und Botanischer Garden, Universitat, Ulm,

GERMANY (Orders: http//www.cerrado.eu/avail.html). €49.00 ($64.60), 277 pp., numerous color

figures, drawings, and tables, 8%" x 015".

GERHARD GOTTSBERGER and [LSE SILBERBAUER-GOTTSBERGER. 2006. Life in the Cerrado: a South American Tropi-

cal Seasonal Ecosystem. Vol. II. Pollination and Seed Dispersal. (ISBN 3-00-017929-1, hbk.).

Abteilung Systematische Botanikund Okologie und Botanischer Garden, Universitat, Ulm, Germany.

(Orders: http://www.cerrado.eu/avail.html). €59.00 ($77.78), 385 pp., numerous color figures, draw-

ings, and tables, 85%" x 9⁄2".

yi TT

This authoritative monograph on one of the most ecosystems of the Neotropics is based on more than 35 years of field work

and research by the two authors. It gives not only a survey of the main results of their own multidisciplinary studies, but offers an up-

to-date and competent synthesis of the relevant and voluminous literature, often not easily accessible and partly written in Portuguese.

The text is well organized and very clearly written, copiously illustrated by many excellent colour p nd drawings, and sup-

ported by numerous schemata and tables.

Volume I covers the more general aspects of the cerrado biome and starts with chapters characterizing the eco-geographical

and floristic position of the Central Brazilian cerrado. Relationships are discussed with the Amazonian, Guiana and Central American

savannas and the caatinga to the North and Northeast, as well as the Pantanal and Beni savannas, and the chaco and campo rupestre

ecosystems towards the West and South. The cerrado and other South American savanna pues ee asa CP DNE of NÉ

changes from the Miocene (about 25 mya bp) onward. During the Pleistocene they had th

tropical rain forests. An important difference with ble African is that large and mostly ungulate herbivore

Is E

mammals had died out in South America by the early Postglacial. For many flowering plant families and genera (e.g., palms, Annona,

Jacaranda, etc.) an origin of cerrado taxa from tropical rain forest ancestors can be demonstrated. There is impressive fossil evidence for

an increase and final dominance in the cerrado grass flora of better adapted C, over less specialized C, taxa from 10 to 3mya bp. Species

diversity is remarkably high in the cerrado flora: For one hectare pect 350—400 vascular plant species are recorded, including 50-90

shrubs and trees. For the whol d bout 10.000 species. Annual changes of pronounced dry and rainy seasons,

and regular fires dominate the cerrado ecosystem. This influences its vegetation rhythm, physiognomy and life form spectrum, and

explains the frequent occurrence of excessive bark formations and subterranean A o Thus, cerrado has an excellent regeneration

capacity. Literature reports and personal contacts with Kayapó and Xavante l the many ways in which cerrado plants

are used as fire wood, food, medicine, game attractant, fertilizer and for cultivation by the natives. Remarkable is their management of

artificial forest islands in which they concentrate useful plants (also from outside of the cerrado). This contrasts with the destruction

of cerrado areas by modern developments which had reached 3796 already in 1990 and is sadly accelerating since. Thus, immediate

measures are necessary, to preserve at least parts of this unique ecosystem

Volume II concentrates on the reproductive nnd of the NAM plants of the cerrado biome. After an introduction, a first

major block of chapters deals with pollination and | , another with seed dispersal. For not less than 625

of flowering EM more or less detailed descriptions HS partly illustrations of pollination modes are presented, including EC

to many hundreds of pollinating animals. Data are arranged according to the following principles: flowers of generalist versus specialist

nature; pollinators either collecting pollen, nectar or oil; small to large bees, beetles, butterflies, moths and flies, as well as hummingbirds

and bats as pollinators; wind pollination. —Flowering plant spectra of different pollination modes for the single Botucatu plot versus

the whole of the cerrado area demonstrate: The majority of species are melittophilous and dependent on small (22/2696) and large bees

(16/2496); generalist taxa come second (37/2296), followed by anemophilous (13/496) and ornithophilous (2/7,596) taxa, whereas all oth-

ers remain below 596. This spectrum differs from other biomes, e.g. from the tropical rain forest where more ornithophilous and less

anemophilous species occur. Of particular interest are chapters concerned with families and genera which exhibit evolutionary radiation

Jls

in response to different p and different flowering periods. A good example is the Vochysiaceae which have differentiated from

large bee to small bee, hawkmoth and hummingbird pollination. The Bignoniaceae are represented by a group of 6 genera with 10 species

in the cerrado which produce pollen and nectar and are pollinated in different proportions by 74 bee species. In addition Jacaranda has

specialized in the production of perfumes from glands at the large staminode, attracting Euglossine bees. Among 19 cerrado species of

Annonaceae 13 belong to the genus Annona and are pollinated by large scarab beetles, whereas 6 species from other genera have smaller

beetles and thrips as pollinators. The Annona species attract a similar spectrum of scarab species by floral heating and strong odor emis-

sion in the evening, but are differentiated with respect to their flowering seasons throughout the year. The 9, mostly short stemmed

cerrado palms (Arecaceae) studied, are pollinated partly by bees, partly by beetles, but only rarely by wind. Comparative analyses of the

J. Bot. Res. Inst. Texas 1(1): 785. 2007

786 Journal of the Botanical R h Institute of Texas 1(1)

breeding system are another important approach, relevant for evolutionary differentiation: 6-15% of the cerrado taxa studied are dioe-

cious, among the hermaphrodite and monoecious taxa 40-50% are reported as self-compatible and 6-8,6% as apomictic A remarkable

set of data is presented on different seed dispersal modes among 301 flowering plant species in the cerrado area of Botucatu. Generally,

there is more zoochory (6496, mainly endozoochory) and anemochory (3496), but less autochory (296) in the tree layer as compared with

the ground layer. Endozoochory is even more dominant in trees of the tropical rain forest. Epizoochory greatly declines from the more

pen cerrado sesu stricto (21%) to the quite dense cerradáo (196). Most of the larger animals involved in seed dispersal (birds, mammals

and bats) live in the gallery forests adjacent to the cerrado. Further chapters deal with interactions of flowering plants with herbivores

and fungi with as well as with ants and termites.

In retrospect: The two volumes on the cerrado ecosystem by Gerhard Gottsberger and Ilse Silberbauer-Gottsberger are a major

contribution to our understanding of the biological problems of South America and a must for all interested in this field. Beyond that

these volumes are an important step forward in current efforts to better evaluate the links between species diversity, ecological inter-

dependences and evolutionary aspects in the different nd bn our ci —Prof. Dr. F. Ehrendorfer, Institute of Botany, University of

ie.ac.at

Vienna, A-1030, Rennweg 14, Vienna, Austria, friedrich.el univie.ac.at.

J. Bot. Res. Inst. Texas 1(1): 786. 2007

BOOKS RECEIVED

BLACKWELL PUBLISHING: ANNUAL PLANT REVIEWS

Jeremy ROBERTS and ZINNIA GONZALEZ-CARRANZA (eds.). 2007. Annual Plant Reviews, Volume 25: Plant Cell

Separation and Adhesion. (ISBN-13 978-14051-3892-5, hbk.). Blackwell Publishing. 2121 State

Ave., Ames, IA 50014-8300, U.S.A. and 9600 Garsington Road, Oxford, OX4 2DQ, UK. (Orders: www.

blackwellprofessional.com, orders@ blackwellpublishing.com, 515-292-0140, 515-292-3348 fax,

1-800-862-6657). $199.99, 232 pp., 6" x 9".

Contents.—1) Cell separation and adhesion o cad in eae z Cell wall structure, biosynthesis and assembly; 3) Vascular cell dif-

ferentiation; 4) Cell adhesion, set idan l eproduction; 5) Cell separation in roots; 6) Organ abscission;

7) Dehiscence; 8) Fruit ripening; 9) the role Bi ates Cross- Swan in m IE adhesion. References, Index.

Kraus D. Grasser (ed.). 2006. Annual Plant Reviews, Volume 29: Regulation of Transcription in Plants.

(ISBN-13 978-14051-4528-2, hbk.). Blackwell Publishing. 2121 State Ave., Ames, IA 50014-

8300, U.S.A. and 9600 Coe) ode i OX4 200 UK. Ordas: www.blackwell-

professional.com, orders@ ll hing.com, 515-292-0140, 515-292-3348 fax,

1-800-862-6657). $199.99, 350 pp., 6" x o"

Contents.—1) General transcription factors and the core promoter: ancient roots; 2) Transcription factors of Arabidopsis and rice: a

on PREE 3) Chromatin-associated architectural HMGA and HMGB pon assist transcription factor function; 4) Histone

in maintaining

] transct ipt ionin plants 5) Chromatin remodeling and histone

DNA methylation; 6) Matrix attachment regions and transcriptional gene silencing; 7) Dues I M 8) Transcription of

plastid genes; 9) Control of flowering time; 10) Combinatorial control of floral organ identity by MADS-domain transcription factors;

11) Networks of transcriptional regulation underlying plant defense responses toward phytopathogens; 12) Temperature-regulated

gene expression; 13) Application of inducible transcription in plant research and biotechnology; 14) Modulation of transcriptional

networks in crop plants.

Garry C. WnrreLam and Karen J. Haruma (eds.). 2007. Annual Plant Reviews, Volume 30: Light and

Plant Development. (ISBN-13 978-14051-4538-1, hbk.). Blackwell Publishing. 2121 State Ave.,

Ames, IA 50014-8300, U.S.A. and 9600 Garsington Road, Oxford, OX4 2DQ, UK. (Orders: www.

blackwellprofessional.com, orders? blackwellpublishing.com, 515-292-0140, 515-292-3348 fax,

1-800-862-6657). $199.99, 325 pp., 6" x 9".

Contents.—1) Phytochromes; 2) Cryptochromes; 3) Phototropins and other LOV-containing proteins; 4) Phytochrome-interacting fac-

tors; 5) Phosphorylation/dephosphorylation in photoreceptor signalling; 6) The role of ubiquitin/proteasome-mediated proteolysis in

photoreceptor action; 7) UV-B perception and signal transduction; 8) Photocontrol of flowering; 9) Red:far-red ration perception and

shade avoidance; 10) Photoreceptor interactions with other signals; 11) Photoreceptor biotechnology; 12) Light-quality manipulation

by horticulture industry. References, Index.

Davip C. Locan (ed.). 2007. Annual Plant Reviews, Volume 31: Plant Mitochondria. (ISBN-13 978-

14051-4939-6, hbk.). Blackwell Publishing. 2121 State Ave., Ames, IA 50014-8300, U.S.A. and 9600

Garsington Road, Oxford, OX4 2DQ, UK. (Orders: www.blackwellprofessional.com, orders@ames.

blackwellpublishing.com, 515-292-0140, 515-292-3348 fax, 1-800-862-6657). $199.99, 342 pp.,

"x 9",

Contents.—1) Mitochondrial dynamics: the control of mitochondrial shape, size, number, motility, and cellular inheritance; 2) The

unique biology of mitochondrial genome instability in plants; 3) Expression of the plant mitochondrial genome; 4) Import of nuclear-

encoded mitochondrial proteins; 5) Mitochondrial respiratory complex biogenesis: communication, gene expression and assembly; 6)

Supramolecular structure of the oxidative phosphorylation system in plants; 7) Mitochondrial m transport and oxidative stress;

8) Mitochondrial metabolism; 9) Cytoplasmic male sterilities and mitochondrial gene mutations in plants; 10) The mitochondrion and

plant programmed cell death. References, Index.

J. Bot. Res. Inst. Texas 1(1): 787. 2007

788 Journal of the Botanical R h Institute of Texas 1(1)

PauL Martin Brown (Text) and STAN Forsom (Original Artwork). 2007. Wild Orchids of the Northeast:

New England, New York, Pennsylvania. (ISBN 978-0-8130-3034-0, flexibind). University Press

of Florida, 15 Northwest 15th Street, Gainesville, FL 32611-2079, U.S.A. (Orders: www.upf.com,

352-392-1351, 352-392-7302 fax, 1-800-226-3822). $29.95, 376 pp., 316 color plates, 94 b/w il-

lustrations, 81 maps, 6" x 9".

ANDRUS Vorrk and Maria Vorrk. 2006. Orchids on the Rock: The Wild Orchids of Newfoundland. (ISBN

0-9699509-3-4, pbk.). Gros Morne Co-operating Association, Publicity Dept., Rocky Harbour, New-

foundland AOK 4NO (Orders: http://www.grosmornetravel.com/intro_details.asp?Id=15). $12.95, 96

pp., color photographs, 5/4” x 612".

Luis M. Botana (ed.). 2007. Phycotoxins: Chemistry and Biochemistry. (ISBN 13 978-0-8138-2700-1,

hbk.). Blackwell Publishing. 2121 State Ave., Ames, IA 50014-8300, U.S.A. and 9600 Garsing-

ton Road, Oxford, OX4 2DQ, UK. (Orders: www.blackwellprofessional.com, orders@ames.

blackwellpublishing.com, 515-292-0140, 515-292-3348 fax, 1-800-862-6657). $199.99,

345 pp., 7” x 10”.

Contents.—1) Gambierol; 2) Brevetoxins: Structure, Toxicology, and Origin; 3) Chemistry of Maitotoxin; 4) Biochemistry of Maitotoxin;

5) Chemistry of Palytoxins and Ostreocins; 6) Biochemistry of Palytoxins and Ostreocins; 7) Chemistry of Cyanobacterial Neurotox-

ins—Anatoxini-a: Synthetic Approaches; 8) Anatoxin-a and Analogues: Discovery, Distribution, and Toxicology; 9) Pectenotoxins; 10)

Chemistry, Origins, and Distribution of Yessotoxin and It's analogues; 11) Pharmacology of Yessotoxin; 12) Chemistry of Diarrhetic

Shellfish Poisoning Toxins; 13) The Molecular and Integrative Basis to DomoicAcid Toxicity; 14) Hepatotoxic Cyanobacteria; 15) Poly-

cavernosides; 16) Structural Assignment and Total Synthesis of Azaspiracid-1; 17) Biochemistry of Azaspiracid Poisoning Toxins; 18)

Cyclin Imines: An Insight into this Emerging Group of Bioactive Marine Toxins. Index.

J. Bot. Res. Inst. Texas 1(1): 788. 2007

Illustrated Flora of East Texas

George Diggs, Barney Lipscomb, ES Reed, Bob O'Kennon

From Big “D” to is Big Thicket th fl

tely nes hardwoo ds,

wisteria, ie as ‘ilies, ae rcp azaleas, and Texas Pride!

Volume 1 of the Illustrated Flora of East Texas is the first fully illustrated flora for East

Texas, a species rich area that contains roughly two-thirds of all the plant species known

for Texas. The book covers all the native and naturalized ferns and similar plants

gymnosperms, and monocotyledons (1,060 species) known to occur in East Texas

Publication date: 26 hy 2006 Botanical Research Institute of Texas (SBM 26)

712" x 101", hbk, 1,614 pp., 1200+ ae figures, 1000+ county distribution maps

ISSN 0833-1475; E 1-889878-12-X

$89.05. Texas residents add 8.2596 tax ($7.42); P&P $7.50 domestic (outside U.S.A.,

please inquire). Available from: Botanical Research Institute of Texas, 509 Pecan Street,

Fort Worth, D. 76102-4060, U.S.A.; jmackenzie@brit.org; Fax 817-332-4112;

www.brit.org/sida/sbm/

BRIT ata Cotte e

PRESS -

Steven P. McLaughlin

Floristic elements and floristic areas for North America

Tundra to Tropics: The Floristic Plant Geography of North America

nalyses were conducted: 1) a PCA onam

ircumscribed using principle

components e (PCA) of a sample of 245 local floras from Canada, the United States, and

Mexico. Three

matrix of Otsuka similarity indices based

provinces; and 3) a PCA on a matrix of Pears

formed nested hierarchical

levels of the hierarchy. When compared with ea aie bio

by Dice, Ud

Publication date: 20 April 2007 Botanical Research Institute of Texas

on shared species, which identified 27 floristic subprovinces; 2) a PCA on a matrix of Pearson

correlations on the log number of species Per a in each flora, which identified 12 floristic

lation g number of species per

family in each flora, which identified 4 osc Rd on eight percent of the 245 floras

grout ll three analyses; 98% formed nested groups over two

geographic treatm

vardy, and Cronquist, McLaughlin's results supported different aspects of each one

but also showed that none completely captured the major floristic patterns on the continent.

ents of North America

(SBM 30)

7" x 10", pbk, 58 pp, 50 b/w maps * ISBN-10-889878-17-0; ISBN-13-978-1889878-17-1

$20.00 Texas residents add 8.2596 ($1.82 tax for one book; $1.77 for each additional copy);

P&P $2.00 domestic (outside U.S.A., please inquire). Available from: Botanical Research Institute

of Texas, 509 Pecan S

Fax 817-332-4112; www.brit.org/sida/sbm/

treet, Fort Worth, Texas 76102-4060, U.S.A.; jmackenzie@brit.org;

Trees in the Life of the Maya World

Regina Aguirre de Riojas and Elfriede de Póll

Trees in the Life of the Maya World brings together the knowledge of the shaman and the scientist

the myths and arts of ancient civilizations and the practices of modern people and treats it all

with wisdom and clarity of vision. It is an example of how we should respect and treat not just

trees but our entire natural surroundings. Chapters include: 1) Introduction; 2) Trees in the Life

of the Maya World; 3) Trees in the Popol Vuh; 4) Trees as Nourishment; 5)

Trees Used in Build-

ing and Carpentry; 6) Tr

ees as Firewood; 7) Trees as Medicine; 8) Other Uses; 9) The Special

Place of Trees; Glossary, Bibliography, and Index.

Publication date: 23 April 2007 Botanical Research Institute of Texas (BRIT Book)

91?" x 12", hbk, French Folded Jacket 208 pp, 48 b/w drawings, 94 color images

ISBN-10-889878-18-9; ISBN-13-978-1889878-18-8

$ $50.00 Texas residents add 8.25% tax ($4.54 tax for one book; $4.41 for each additional

y copy); P&P $5.00 domestic (outside U.S.A., please inquire). Available from: Botanical

Research Institute of Texas, 509 Pecan Street, Fort Worth, Texas 76102-4060, U.S.A.;

BRIT jmackenzie@brit.org; Fax 817-332-4112; www.brit.org/sida/sbm/

PRESS

[9]

ISSN 1934-5259

JOURNAL OF THE BOTANICAL RESEARCH INSTITUTE OF TEXAS

The Journal of the Botanical Research Institute of Texas, Formerly

called Sida, Contributions to Botany, publishes research in classical

and modern systematic botany—including studies of anatomy,

biogeography, chemotaxonomy, ecology, evolution, floristics,

genetics, paleobotany, palynology, and phylogenetic systematics.

Geographic coverage is global. Articles are published in either

English or Spanish; an abstract is provided in two languages. AII

contributions are peer reviewed and frequently illustrated with

maps, line drawings, and full color photographs.

BRIT The journal is published twice yearly in a multiple topic format

and provides information to scientists, professionals, and the general public for research,

education, and cultural enhancement. All articles are indexed and abstracted in print

and/or electronic form by the following: AGRICOLA Database (National Agricultural

Library); Applied Botany Abstracts; Biosciences Information Service of Biological

Sciences (BIOSIS); Current Awareness in Biological Sciences (CABS); Excerpta

Botanica; The Kew Record of Taxonomic Literature; Natural Products Alert

(NAPRALERT); and Referativnyi Zhurnal (Abstracts Journal of the Institute of Scientific

Information of the Republic of Russia).

Lloyd H. Shinners, whose herbarium and library were the basis of the Southern

Methodist University (SMU) legacy that became BRIT in 1987, founded the journal in

1962. William F. Mahler succeeded Shinners as SMU herbarium director in 1971 and

inherited both ownership and editorship of the journal. Since 1993, it has been published

by the Botanical Research Institute of Texas.

The current issue of the Journal of the Botanical Research Institute of Texas is available online

at http://www britpress Org