JOURNAL OF THE BOTANICAL RESEARCH INSTITUTE OF TEXAS

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

VOLUME 3 NUMBER 1 15 JULY 2009

Botanical Research Institute of Texas (BRIT)

500 East 4th Street

Fort Worth, Texas 76102-4025, USA

EDITOR: Barney Lipscomb

ASSISTANT EDITOR: Brooke Byerley

Botanical Research Institute of Texas

500 East 4th Street

Fort Worth, Texas 76102-4025, USA

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

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HISTORY AND DEDICATION

1962— Lloyd H. Shinners

(left), à 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)

CONTRIBUTING SPANISH EDITOR

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

botany at SMU and director emeritus of BRIT,

inherited editorship and copyright.

1993—BRIT becomes publisher/copyright holder.

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

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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. Dot. 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) as one volume

by the Botanical Research Institute of Texas.

Félix Llamas

Dpto. de Botánica, Facultad de Biologia

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PRINTING

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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.

BRIT

PRESS

TABLE OF CONTENTS

COMMENTARY

Learning about other species: an updated component of a liberal arts education

Guy L. NESOM AND ÁLAN S. WEAKLEY

SYSTEMATICS

Orobanche riparia (Orobanchaceae), a new species from the American Midwest

L. TURNER COLLINS, ALISON E.L. COLWELL, AND GEORGE YATSKIEVYCH

Bristlecone pine dwarf mistletoe: Arceuthobium microcarpum subsp. aristatae (Viscaceae),

a new subspecies of western spruce dwarf mistletoe from northern Arizona

JARED M. SCOTT AND ROBERT L. MATHIASEN

A new cliff-dwelling species of Zamia (Zamiaceae) from Belize

MICHAEL CALONJE

A new species of Zamia (Zamiaceae) from the Maya Mountains of Belize

MICHAEL CALONJE, JAN MEERMAN, PATRICK GRIFFITH, AND GEOFFREY HOESE

What is Zamia prasina (Zamiaceae: Cycadales)?

MICHAEL CALONJE AND JAN MEERMAN

Festuca aloha and F. molokaiensis (Poaceae: Loliinae), two new species from Hawai'i

PILAR CATALÁN, ROBERT J. SORENG, AND PAUL M. PETERSON

A new genus, Mexotis, for five Mexican species of Hedyotideae (Rubiaceae)

EDWARD E. TERRELL AND HAROLD ROBINSON

Capsicophysalis: a new genus of Solanaceae (Physaleae) from Mexico and Central America

JOHN E. AvERETT AND MAHINDA MARTÍNEZ

Synopsis of Gonolobus s.s. (Apocynaceae: Asclepiadoideae) in Trinidad and Tobago

ALEXANDER KRINGS

An overview of the Diospyros campechiana complex (Ebenaceae) and description of three

new species

MITCHELL C. PROVANCE AND ÁNDREW C. SANDERS

Novedades y notas misceláneas en las Bromeliaceae de Mesoamérica

J. FRANCISCO MORALES

Una nueva especie y notas misceláneas en el género Oreopanax (Araliaceae) en Centro América

J. FRANCISCO MORALES AND ALVARO IDARRAGA

Five new species of Myrtaceae from Ecuador

MARIA Lúcia KAWASAKI AND BRUCE K. HOLst

Folia taxonomica 12. Paradrymonia (Gesneriaceae: Episcieae) from the Guiana Shield: P. magu-

irei, a new species from Amazonas, and distribution and floral morphology of P. maculata

CHRISTIAN FEUILLET

New species of Baccharis (Asteraceae: Astereae) from Rio de Janeiro state, southeastern Brazil

G. HEIDEN, J.F.A. BAUMGRATZ, AND R.L. ESTEVES

Diplycosia indica (Ericaceae): a new species and a new generic record for India

M.R. DEBTA AND H.J. CHOWDHERY

Amorpha confusa, a new name for an old Amorpha (Fabaceae: Amorpheae)

SHANNON C.K. STRAUB, BRUCE A. SORRIE, AND ALAN S. WEAKLEY

A new combination in North American Tephrosia (Fabaceae)

Guy L. NESOM AND JAMES L. ZARUCCHI

113

117

123

133

139

147

151

157

A recombination for varieties of Anticlea elegans (Melanthiaceae)

WENDy B. ZOMLEFER AND WALTER S. JUDD

Taxonomic overview of Eurybia sect. Herrickia (Asteraceae: Astereae)

Guy L. Nesom

The phylogeny of Selenia (Brassicaceae) inferred from chloroplast and nuclear sequence data

JAMES B. BECK :

Preliminary insights into the phylogeny and speciation of Scalesia (Asteraceae), Galápagos Islands

JEREMY D. BLASCHKE AND RoGER W. SANDERS

Introduced lesser celandine (Ranunculus ficaria, Ranunculaceae) and its putative subspecies in

the United States: a morphometric analysis

ANGELA R. Post, ALEXANDER KRINGS, WADE A. WALL, AND JOSEPH C. NEAL

The types of Astragalus section Diphysi (Fabaceae), a complex endemic to Western North America,

Part I: lectotypifications, epitypifications, and new combinations of several taxa

J. ANDREW ÁLEXANDER

The typification of Crotalaria rotundifolia and Crotalaria maritima (Fabaceae)

DaNiEL B. WARD

Lectotypification of Cardamine flexuosa (Brassicaceae)

ANGELA R. Post, ALEXANDER KRINGS, QUI-YUN (JENNY) XIANG, BRYON R. SOSINSKI, AND JOSEPH C. NEAL

An investigation of morphological evidence supports the resurrection of Pyrrocoma scaberula

(Asteraceae: Astereae)

Curtis R. BJORK AND MARK DARRACH

Miscellaneous typifications, one new series, and one new varietal combination in Crataegus

(Rosaceae)

J.B. Pipers

What is Ranunculus gelidus (Ranunculaceae)?

ALAN T. WHITTEMORE

Infrageneric taxonomy of Astrophytum (Cactaceae), with remarks on the status of Digitostigma

RICHARD R. MONTANUCCI

Mycological literature on Texas fleshy Basidiomycota, two new combinations, and new fungal

records for Texas

Davip P. Lewis AND CLARK L. OVREBO

Twenty-eight new three-letter family acronyms for vascular plants (with comprehensive listings

on-line)

NEIL SNOW

CHROMOSOME NUMBERS

Chromosome numbers of miscellaneous angiosperm taxa

Marc BAKER, Jon REBMAN, BRUCE PARFITT, DONALD PINKAVA, CHARLOTTE CHRISTY, ANDREW SALYWON,

RAUL PUENTE-MARTINEZ, ALLAN ZIMMERMAN, AND J. Huso COTA

ANATOMY AND MORPHOLOGY

Anatomia de la epidermis foliar en las especies Mexicanas del género Muhlenbergia (Poaceae)

OCTAVIO ROSALES CARRILLO AND YOLANDA HERRERA ARRIFTA

FLORISTICS, ECOLOGY, AND CONSERVATION

Primer reporte de Cystopteris reevesiana (Aspleniaceae) para México

José A. VILLARREAL-Q. AND CELESTINO FLORES-L.

159

161

169

177

193

211

219

227

231

239

245

251

257

273

279

285

307

Floristic composition, abundance, and distribution pattern of woody plants in a tropical savanna

in northern Ghana

ALEX ASASE, PATRICK K. EKPE, AND JOHN Y. AMPONSAH

New state records of vascular plants for the Southern Rocky Mountain region

PAMELA F. SMITH, DONALD L. HAZLETT, AND NEIL SNOW

Pyracantha koidzumii (Rosaceae) new to the Arkansas flora

BRETT E. SERVISS

Medicago rigidula and M. truncatula (Fabaceae): new to the California flora

RICHARD E. RIEFNER, JR. AND ERNEST SMALL

New and noteworthy plants from Florida

Jonn M. KUNZER, RICHARD P. WUNDERLIN, LORAN C. ANDERSON, AND JAMES R. BURKHALTER

Vascular flora of Churchill Ranch, Sarasota County, Florida

ALAN R. FRANCK AND RICHARD P. WUNDERLIN

Vascular flora of a saline prairie in Winn Parish, Louisiana

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

Vascular flora of Morse Clay prairies in northwestern Louisiana

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

Floristics of upland shortleaf pine/oak-hickory forest in northwestern Louisiana

BARBARA R. MACROBERTS AND MICHAEL H. MACROBERTS

A revision to the vascular flora of Calcasieu Parish, Louisiana

Ray NEYLAND

Additions and emendations to the vascular flora of Caddo Parish, Louisiana

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

Mentha suaveolens and M. xrotundifolia in North Carolina: a clarification of distribution

and taxonomic identity

MICHAEL W. DENSLOW AND DERICK B. POINDEXTER

Castilleja angustifolia var. dubia (Scrophulariaceae), a new record for South Dakota

GRACE KOSTEL AND LYNN HETLET

Glyceria declinata (Poaceae) new to the flora of Texas

CHARLES M. ALLEN, PATRICIA LEWIS, AND Davip P. Lewis

Vegetation areas of Texas: concept and commentary

Ray C. TELFAIR II

Vascular plant species richness of the Big Thicket, East Texas

MicHAEL H. MACROBERTS AND BARBARA R. MACROBERTS

A classification of geographic elements and analysis of the flora of Big Bend region of Texas

ANNA SAGHATELYAN

Annotated checklist of the vascular flora of the Menard Creek Corridor Unit of the Big Thicket

National Preserve, Polk, Liberty, and Hardin counties, Texas

Larry E. Brown, BARBARA R. MACROBERTS, MICHAEL H. MACROBERTS, AND WARREN W. PRUESS

Cyperus pilosus (Cyperaceae) new to the flora of Texas

RICHARD CARTER, CHARLES M. ALLEN, PATRICIA LEWIS, AND Davip P. Lewis

Cryptostegia grandiflora (Apocynaceae: Asclepiadoideae), a new non-native weed for Texas

THOMAS F. PATTERSON AND Guy L. NESOM

Tournefortia hirsutissima (Boraginaceae) new to the flora of Texas

ALFRED RICHARDSON AND W. KEN KING

309

317

319

323

331

339

349

355

367

375

379

383

391

393

395

443

457

461

Book Reviews and Notices 12, 22, 30, 42, 50, 76, 84, 122, 146, 156, 168, 192, 210, 226, 244, 272, 278,

284,308, 322,330, 0239197804390, 400, 406, 442, 456, +00: 404

Announcement 468

INDEX to new names and new combinations in J. Bot. Res. Inst. Texas 3(1), 2009

Amorpha confusa (Wilbur) S.C.K. Straub, Sorrie @ Weakley, comb. et stat. nov.—154

Anticlea elegans var. glaucus (Nutt.) Zomlefer € Judd, comb. nov.—159

Arceuthobium microcarpum subsp. aristatae J.M. Scott & Mathiasen, subsp. nov.—14

Astragalus lentiginosus var. bryantii (Barneby) J.A. Alexander, comb. nov.—215

Astragalus lentiginosus var. iodanthus (S. Watson) J.A. Alexander, comb. nov.—215

Astragalus lentiginosus var. pseudiodanthus (Barneby) J.A. Alexander, comb. nov.—216

Baccharis altimontana Heiden, Baumgratz & Esteves, sp. nov.—140

Baccharis friburgensis Heiden, Baumgratz & Esteves, sp. nov.—143

Calyptranthes compactiflora M.L. Kawasaki & B.K. Holst, sp. nov.—123

Calyptranthes fusca M.L. Kawasaki & B.K. Holst, sp. nov.—125

Calyptranthes sparsiflora M.L. Kawasaki & B.K. Holst, sp. nov.—125

Capsicophysalis (Bitter) Averett & M. Martínez, stat. nov.—72

Capsicophysalis potosina (B.L. Rob. & Greenm.) Averett & M. Martínez, comb. nov.—72

Crataegus series Altaicae J.B. Phipps, ser. nov.—243

Crataegus chrysocarpa var. subrotundifolia (Sarg.) J.B. Phipps, comb. et stat. nov.—240

Diospyros camposii M.C. Provance & A.C. Sanders, sp. nov.—93

Diospyros crotalaria M.C. Provance & A.C. Sanders, sp. nov.—95

Diospyros haberi M.C. Provance & A.C. Sanders, sp. nov.—98

Diplycosia indica M.R. Debta @ H.J. Chowdhery, sp. nov. —147

Eurybia aberrans (A. Nels.) G.L. Nesom, comb. nov.—166

Eurybia kingii (D.C. Eaton) G.L. Nesom, comb. nov.—166

Eurybia kingii var. barnebyana (Welsh & Goodrich) G.L. Nesom, comb. nov.—166

Festuca aloha Catalán, Soreng & PM. Peterson, sp. nov.—51

Festuca molokaiensis Soreng, PM. Peterson & Catalán, sp. nov.—54

Hygrocybe chamaeleon (Cibula) Lewis & Ovrebo, comb. nov.—264

Hygrocybe mississippiensis (Cibula) Lewis & Ovrebo, comb. nov.—264

Mexotis Terrell & H. Rob., gen. nov.—60

Mexotis galeottii (M. Martens) Terrell & H. Rob., comb. nov.—61

Mexotis kingii (Terrell) Terrell & H. Rob., comb. nov.— 64

Mexotis latifolia (M. Martens & Galeotti) Terrell & H. Rob., comb. nov.—61

Mexotis lorencei Terrell & H. Rob., sp. nov.—67

Mexotis terrellii (Lorence) Terrell & H. Rob., comb. nov.—64

Myrcia aequatoriensis M.L. Kawasaki € B.K. Holst, sp. nov. —128

Myrcia verticillata M.L. Kawasaki € B.K. Holst, sp. nov. —130

Oreopanax paramicolus J.F Morales & A. Idárraga, sp. nov.—118

Orobanche riparia L.T. Collins, sp. nov. —7

Paradrymonia maguirei Feuillet, sp. nov.—134

Paradrymonia subg. Pagothyra (Leeuwenb.) Feuillet, comb. et stat. nov. —134

Tephrosia xintermedia (Small) G.L. Nesom & Zarucchi, comb. nov.—157

Werauhia luctuosa J.F Morales, sp. nov. —114

Zamia decumbens Calonje, Meerman, M.P Griff. & Hoese, sp. nov —31

Zamia meermanii Calonje, sp. nov.—23

COMMENTARY

LEARNING ABOUT OTHER SPECIES:

AN UPDATED COMPONENT OF A LIBERAL ARTS EDUCATION

Guy L. Nesom Alan S. Mid

2925 Hartwood Drive Herbarium/NC B

Fort Worth, Texas 76109, U.S.A. Univ North Carolina, Coker Hall, E 2

guynesom@sbcglobal.net Chapel Hill, North Carolina 27599-3280, U.S.A.

weakley@unc.edu

Some aspects of a liberal arts education—critical thinking, respect for truth and clarity of expression—are

unchanging in value. Knowledge of humans and human culture remains central. But if the goal of the cur-

riculum is to prepare students for broad engagement in the world, an understanding of the Earth and all of

life calls for equal time.

Humanity is, for now at least, bound to Earth. Here, unbridled growth of the human population and

profligate use of natural resources threaten our own existence and put at risk the well-being and continuance

of all life and the environment. An appreciation of the planet and preparation toward committed stewardship

should be among the primary goals of ae E arts education can not afford to be self-centered, in

the sense of study of human cult l context and without attention to the thousands

of species that are our evolutionary associates.

We propose a course that guides students towards an appreciation of the Earth as home to many spe-

cies. At the end of the essay, the course is integrated within an updated liberal arts curriculum.

Learn to know other species. Perhaps the most effective path toward an appreciation of life on Earth is

through first-hand knowledge of species other than our own. The primary goal of the proposed course is

to learn the identities of commonly encountered species. The course need not be heavily laden with deep

academic underpinning—the value lies in learning to recognize species by sight, as they are encountered

in everyday life, and to identify them by name.

A know-the-species course provides foundation for one of the most basic lessons in humility—humans

are one species among hundreds of thousands. So many of us are hardly aware that others exist, except in

an abstract way. Daily acquaintance with other species on a "first-name" basis greatly broadens one's view

of the natural world and engenders a closeness, a sense of unity, with it. Even if names of species eventually

are forgotten, at least the memory remains that each is unique, distinguished by features that most have

previously been unaware of.

One can quickly learn to identify a half-dozen species (hollies, oaks, maples, etc.) and to see that some

of the species are more similar among themselves while others are more unique. Such an appreciation of

variation and diversity, even in a simple system, is the quickest way possible toward real insight into the

process of evolution— patterns of similarity among species result from common ancestry. Each species on

Earth is the product of long evolutionary lineage, interrelated among the others, each unique and beauti-

ful.

“Lam strongly induced to beli hat, as in music, tl ho understands every note will, if he also possesses a proper taste,

more thoroughly enjoy the whole, so ene que examines a part of a aoe view, n c puo iia the full

d be a botanist, for in all I "Charles

Darwin, 1836, Voyage i the E.

Nature of the course. Plants are the most conspicuous and abundant expression of non-human life, even

in urban settings, and we believe they are the best focus for the proposed course. It's joyful to walk along a

J. Bot. Res. Inst. Texas 3(1): 1 — 2. 2009

sidewalk, through a park, or through the woods and recognize species of oaks, elms, and maples by name.

Even in a cityscape, dozens of species are commonly planted and usually many more are scrabbling out

their existence, unassisted, seeding themselves in unlikely places.

Awareness of other species also could be gained from an introduction to birds, beetles, fish, or fungi, or

practically any other kind of organism, but none of these is as easily accessible as plants. Do other courses in

biology (e.g., physiology, genetics, molecular biology, cell biology) offer the same potential insights? Simply,

no—most are based on study of a single species or of no species in particular. Some courses in ecology re-

quire a basic knowledge of species diversity, but the focus is different. Studies of geology, meteorology, and

astronomy may be doorways to recognition of the diversity of everyday extra-human experience and could

be analogous to identification of plants, but diversity, or at least every-day accessibility, is much lower.

Residents of the United States and other highly urbanized areas are so removed from other species that

it is easy to be unaware that plants underpin our existence. Do undergraduates recognize that our diet is

mostly of roots, stems, leaves, fruits, and seeds? Do typical students know that most of the energy expended

by human individuals comes directly from starch in corn, rice, cassava, wheat, and potatoes? Beans, ba-

nanas, and sorghum are the world's dietary staples. Chocolate, coffee, sugar—the plant foods that tempt us.

Cotton, flax, hemp, mulberry—the plants of fashion. Even the meat in our diet is but one step away from

plant nutrition. Plants are responsible for nearly all our oxygen, our clothes (directly or indirectly), and most

of our non-food energy (fossil fuels —oil, natural gas, coal). These topics deserve a place in a liberal arts

education.

Since the suggested course is not an abstruse undertaking, basic knowledge of other species should be

taught not only as part of a liberal arts collegiate education but in elementary through high school as well.

Of course, this is one among many basic topics, but lack of knowledgeable teachers is a constraint and the

responsibility usually comes to rest at college level.

Suggested course outline. (A) identification of 100 species, concentrating on first-hand recognition of

those most easily at hand and abundant; (B) a brief overview of plant vegetative features and reproduction

(flower and fruit structure usually are critical in identification) and common families encountered among

the species; (C) perspectives on evolution, the interrelatedness of life, and global patterns of diversity; and

(D) a brief introduction to ecological and economic values of plants tered in everyday life (e.g., urban

forests, invasive species, food, clothes, construction, fuel). Simply learning to know the species is the first

priority.

Goals of a liberal arts education. In the broad view, we believe that a liberal arts curriculum should be

aimed (I) toward continued existence of our own species and all others and (2) toward maximizing the

potential for freedom and happiness (including creativity, appreciation of beauty, productivity, service, and

increasing understanding of the world).

Overview of an updated liberal arts curriculum. We divide a suggested curriculum into four major, overlap-

ping categories.

1. The Earth and all things non-human. Life and life processes (humans as animals, other species, ecology,

evolution), geology, weather and climatology. Our proposed course would figure as a central focus in

this category.

2. Intellectual outlook. Reasoning and critical thinking (including scientifi hod), respect for truth, clarity

of expression, philosophy and ethics.

3. Humans and human culture. Beauty (visual art, music, perception), cultural diversity and history (history,

language, philosophy, ethics, and religion).

4. The Universe and universal principles. Astronomy and cosmology, physics and chemistry, mathematics.

OROBANCHE RIPARIA (OROBANCHACEAB),

A NEW SPECIES FROM THE AMERICAN MIDWEST

L. Turner Collins Alison E.L. Colwell

Science and Technology Department Resources Management and Science

vangel University Yosemite National Par

111 N Glenstone PO. Box 700, 5083 Foresta Road

Springfield, Missouri 65802, U.S.A. El Portal, California 95318, U.S.A.

collinst@evangel.edu alison_colwell@nps.gov

George Yatskievych

Missouri Botanical Garden

PO. Box

St. Louis, Missouri 63166, U.S.A.

george.yatskievychemobot.org

ABSTRACT

Orobanche dedi mE oe in the n Plains, is tl idely distributed species i t. Nothaphyllon. It was first

described by T in 181 ted at Ft. Mandan, North Dakota. During the foll and a half,

most botanists had HQ field Ee "ih living plants of the g ] very few herbarium peci ith e to a ke

newly collected eae! bl fusion al he ci ion of thi i lits g

N hyllon ( M hiza) by B (1890, 1930) aud 1930) 1 led | i l species

name too broadly to par whose mn was s discordant with Nuttall's original protologue, a the dinde ofa an Bn. broad

led hi O. ludoviciana

pn (Collins 1973) 1d his problem, including f l gnition of logical itia O. ludoviciana sensu stricto

with different h inhabi land d tl I fined to I land iated with major river systems. Recent

additional investigations have revealed fu further evid pporting gnition of tl listi ic entiti

: oe logical distincti i 1 : p i f the upland f O. ludoviciana

] gnition the | land f distinct species, O. riparia, sp. nov.

RESUMEN

Orobanche ludoviciana, cuya extensión abarca mayormente la zona de Great Plains, es la especie de mayor distribución en la Sección

Nothaphyllon. El o que n una ee fue pas Nuttall en 1818, que se K en una muestra NAME en Ft. Mandan,

Dakota del N D te el g Į i p g y con

ejemplares de a con las due comparar los e obtenidos igui hubo! fusión en torno a la circun-

scripción de esta especie y su distribución geográfica. Los estudios ROM con la Sección Nothaphyllon (sec. Myzorrhiza) hechos

por i ie pid y Munz Coa hacen que los botánicos apliquen de modo demasiado amplio los nombres de las plantas cuya

uttall. El problema que permanece es el de un concepto demasiado amplio de

E 8

la E Estudios doctoral ducid por el autor pip durante los primeros años de la década de los setenta resultaron en

TRE d 1 I O. ludoviciana (Collins 1973), incluso un reconocimiento formal de dos

tro de O. ludovici j distribución algo dif éspedes. Una de las plantas era s tierras

altas y es s retenida aquí como O. ludoviciana y l finada a ti baj iadas principal i

Jecerincion: Reci ] lido a 1a] asd dich t a. En r

trabajo se comentan las diferencias morea entre estos dos ecotipos y se propone reconocer les adams de las tierras bajas

ta O. riparia, sp. nov.

COLL p

Key Wonps: Orobanche, Orobanchaceae, parasitic plants, broomrape, Flora of North America

Orobanche L. sect. Nothaphyllon (A. Gray) Heckard (sect. Myzorrhiza Phil.) comprises of an array of ca. 13

species that account for most of the taxonomic diversity within New World holoparasitic Orobanchaceae. It

has been considered a monophyletic group by most botanists based on morphological and molecular data

(Olmstead et al. 2001; Schneeweiss et al 2004; Bennett & Matthews 2006). Within sect. Nothaphyllon, two

J. Bot. Res. Inst. Texas 3(1): 3 — 11. 2009

major clades exist whose relationships have not yet been fully resolved: the O. californica Cham. & Schltdl.

complex (Heckard 1973), which has diversified mainly in the far-western United States; and the O. ludoviciana

Nutt. complex (Collins 1973), which is widespread in the remainder of the United States and Canada, and

southward into Mexico (also disjunct in southern South America).

Orobanche ludoviciana Nutt., with a range primarily in the Great Plains, is the most widely distributed

species in sect. Nothaphyllon. It was first acude. by Thomas Nuttall in 1818 from a specimen collected at

Ft. Mandan, North Dakota. B ists during the following century had limited field experience

with living plants of the genus and had very few herbarium specimens with which to compare newly collected

materials, the species name was applied too broadly to include plants whose morphology is discordant with

Nuttall's original protologue. The result has been considerable confusion about the circumscription of this

species and its geographic distribution. All those who have treated Section Nothaphyllon (sect. Myzorrhiza)

(Beck 1890, 1930; Munz 1930; Collins 1973; Heckard 1973), have at times recognized various infraspecific

taxa, some of which are no longer accepted and therefore excluded from this discussion. Nevertheless, the

lingering problem has been that of an overly broad species concept. Even Philip Munz (1930), who provided

good insights into taxonomic variation within O. ludoviciana in his otherwise excellent taxonomic revision

of sect. Myzorrhiza, overlooked significant morphological variation. Subsequent authors of floristic works

who relied on his revision to determine specimens continued to some degree to overlook the taxonomic

heterogeneity of plants treated by him as O. ludoviciana var. genuina Beck.

Doctoral studies by the senior author led him to suggest a number of

ludoviciana complex (Collins 1973), one of which was formal recognition of two ecological races within O.

ludoviciana sensu stricto with differing host preferences: one inhabiting only uplands and the other confined

to bottomlands associated with major river systems. Recent discoveries of new populations and examina-

tion of additional existing herbarium specimens of the bottomland plants have added evidence in support

of this proposal. The present paper therefore discusses the morphological distinctions between these two

ecotypes, and we propose to retain the upland form as O. ludoviciana and to recognize the bottomland form

as a distinct species, O. riparia. It should be noted that, based on Collins's dissertation, the epithet O. riparia

was discussed far in advance of the present publication by Musselman (1982, on Orobanchaceae in Virginia)

and by Kartesz (1994, in a North American plant checklist), but neither of these usages affects the validity

of the present description.

Bringing this new taxon to the botanical literature requires a brief review of the existing nomenclature

and morphology of O. ludoviciana sensu stricto (O. ludoviciana var. genuina, sensu Beck). Several species of

Orobanche in North America are commonly misidentified as O. ludoviciana, but are currently recognized

as distinct species. They include O. multiflora Nutt., O. cooperi (A. Gray) A. Heller, O. multicaulis Brandegee,

and O. valida Jepson. Of these, the two with the most morphological similarity and name association are O.

ludoviciana and O. cooperi, and must therefore be contrasted with O. riparia.

Collins (1973) found that Munz's 1930 description and illustration of the corolla lobes of O. ludoviciana

did not match the morphology of Nuttall's type specimen. The corolla lobes of the type specimen are rounded

on both the upper and lower lips, but Munz's illustration, based on a specimen from White County, Illinois,

shows a corolla with triangular-pointed lobes. Because Munz used material that matched his illustration in

assembling his description of O. ludoviciana, subsequent regional and state floras continued to overlook the

mu gical eub pee Mp i to i euer

ated

within the O.

range has revealed that the character of founded e ibe is consistent throughout for UN PR

(Fig. 1). In contrast, the corolla lobes of bottomland plants, hereafter referred to as O. riparia, are consistently

triangular and pointed (Fig. 1). The populations of O. riparia share several additional characteristics (Table

1): D They are found almost exclusively in river bank and sandbar habitats in floodplains; 2) The native

hosts are annual members of Ast tribe Heliantheae, subtribe Ambrosiinae, primarily Ambrosia trifida

L. and Xanthium strumarium L., and occasionally Ambrosia artemisiifolia L. and Dicoria canescens A. Gray 3)

Collins et al., Orobanche riparia, a new species

y as

karl Ae,

7 VL x2 D^

re Af ncn A

à - "FE

Fic. 1. Comp

ludoviciana.

6 Journal of tt IR h Institute of Texas 3(1)

Tape 1. C | id f Orot | ludoviciana, and cooperi

Character Orobanche riparia Orobanche ludoviciana Orobanche cooperi

Calyx len 7-11 mm 8-14 mm 7-12 mm

Corolla length 15-22 mm 14-20 mm 15-32 mm

ape and size Triangular-acute, 4-5mm, Rounded-obtuse, 4-8mm, Triangular-acute, 5-8 mm,

of corolla erect or slightly reflexed erect or ascending apiculate apex, ascending

lips/lobes or reflexed

Corolla tube Strongly curved, tube Slightly curved, tube ascending, Strongly curved, tube

Corolla color

horizontal slightly flared distally

Pallid, or upper lobe apex

purple/lavender

slightly flared distally

Purple, lavender, rose, or pallid,

tube exterior whitish

ae nalig: (alee ay

Dar | half with

T eeii and throat

Pubescence pubescent with glandularand Pubescent with "uad Pubescent with glandular

eglandular trichomes or trichomes except in trichomes on tube, eglandular

glabrate lobe surface on inner surface of lobes

Flowering August-October April-August December-April

Inflorescence Open raceme, flowers in Compact raceme, flowers in Open or compact raceme with

arrangemen loose, regular spiral on dense, often irregular spiral on flowers in loose, regular spiral

Primary hosts

Habitat and

floral axis

Annuals; Ambrosia, Xanthium,

icoria

River banks, sandbars eastern

floral axis

Perennials; Grindelia, Artemisia,

Heterotheca

Prairie, desert, dunes; central

and western U.S.

on floral axis

Paranniale Amhrocin

Sonoran Desert, ravines, dry

stream beds, dunes;

main range U.S. and prairie provinces

southwestern U.S,

The flowering period is from mid-August to early October; 4) Inflorescence color is typically pallid with

only the interior tips of the corolla lobes tinted purple; 5) The corolla tube is strongly curved with the distal

portion of the tube horizontal, slightly flared distally, and palatal folds that are either glabrous or densely

pubescent.

Compared with O. riparia, O. ludoviciana has slightly shorter corollas with a flared throat and rounded

corolla lobes (Table 1). It occurs mainly through the Great Plains (and eastward along the Prairie Peninsula)

in various habitats, including sandy prairies, sand dunes, dry washes, calcareous (caliche) soils, and lower

slopes of the Rocky Mountains. Its hosts are almost exclusively perennial members of Asteraceae tribes As-

tereae and Anthemideae, including Grindelia squarrosa (Pursh) Dunal, Heterotheca spp., Artemisia spp., and

occasionally other genera. The principal flowering period is from late April in the southern plains through

mid-August in the northern plains, although sporadic flowering has been observed rarely at other times in

marginal habitats. The corolla tube is slightly curved at the constriction, ascending, and flared distally, with

the palatal folds sparsely pubescent. Floral pigmentation varies from purple to pale rose and occasionally

yellow, usually with dark purple upper lobes (Table 1).

Additionally, populations of O. riparia are ecologically and somewhat geographically distinct, and the

habitat distinction is maintained in the western populations, where the two ecological races are geographi-

cally somewhat sympatric. Populations east of the Mississippi River (but including ell are allopatric

with O. ludoviciana. West of the Mississippi River, sympatry of m eastern Nebraska

and Kansas to eastern Colorado, southward into New Mexico and the Texas Panhandle. A small overlap in

flowering times in early August could provide an opportunity for hybridization between the two species.

Although no suspected cases dix uen have been addas based on morphology or field observa-

tic st ded to answer this question. Molecular

+} + ors

tions of populations, data from g

studies by Colwell are currently underway.

Munz (1930) treated O. cooperi as a variety of O. ludoviciana, but subsequent authors mostly have ac-

cepted it as a separate, but closely related species (Shreve & Wiggins 1964; Munz 1974). Orobanche cooperi is

Collins et al., Orobanche riparia, a new species 7

distinguished by its dark purple color, larger, di lla tube, and erect or strongly reflexed, triangular

lobes with apiculate apices. The corolla tube has a whie exterior and is slightly curved and flared distally,

with palatal folds pubescent at the mouth and becoming glabrous in the throat. The habitat is primarily

Sonoran Desert scrub, often associated with washes that remain dry except in monsoon season. Its hosts

are shrubs in various tribes of Asteraceae, principally species of Ambrosia and Encelia (all Heliantheae, Am-

brosiinae), but also reported on Gutierrezia (Astereae). The flowering period is mainly December through

April (Table 1). Collins (1973) found that the cross-sectional stem anatomy of O. cooperi and O. riparia are

very similar and that O. ludoviciana differs significantly from both of the other taxa.

Several characters suggest that O. riparia is more closely related to O. cooperi than to O. ludoviciana.

Orobanche riparia and O. cooperi share an affinity for riparian or water-eroded habitats and hosts in the sub-

tribe Ambrosiinae. They differ in that O. cooperi occurs only in seasonally arid habitats, whereas O. riparia

occurs in both mesic and semi-arid habitats, and that the former parasitizes shrubby hosts, whereas the

host taxa of the latter are annuals. The pubescence of the corolla lobes in both species is eglandular on the

inner surface with vestiture considerably longer than the short glandular trichomes typical of the exterior

surface. Likewise, the palatal folds are invested with eglandular trichomes, differing between the species

only in their distribution on the folds. This shared pattern of trichomes is more evident in the western

populations of O. riparia than in the eastern populations. Similarly, the eastern populations have reduced

indumentum as compared with the western populations. One can speculate that O. riparia and O. cooperi

share a recent common ancestor. A possible scenario is that a progenitor’s range became bisected, with

populations that became O. riparia adapting and dispersing into new habitats eastward across the American

Midwest and populations now referable to O. cooperi attaining a distribution in the southwestern U.S. and

adjacent Mexico. If this is the case, O. riparia adapted quickly to annual hosts and a late summer to early

fall flowering period, in contrast with the shrubby hosts and late winter to early spring flowering period of

O. cooperi. Of this character set, O. ludoviciana shares only the semi-arid habitat and perennial hosts with

O. cooperi, although the host taxa of O. ludoviciana often are herbaceous.

Orobanche riparia L.T. Collins, sp. nov. (Fig. 1). ter: UNITED STATES: Inpiana: Gibson Co.: common on bank of slough

.5 mi N of Griffin, 16 Aug 1931, Deam 50941 (noLorvee: IND; isorvees: A, E GH, IND, MINN, WIS).

Ab O. ludoviciana differt lobis corollae apice acutis, puberulis vel cs eure horizontali, tricto flexoq per ovarium,

inflorescentia laxe racemosa. Ab O. cooperi differt corolla 15-22 mm longa, lo! I piculati flexis, inflorescentia

pallide purpure

Annual root parasite, lacking chlorophyll. Coralloid roots few, rudimentary, stout, fleshy, usually forming

a single host attachment. Stem stout or slender, fleshy, 5—35 cm long, mostly subterranean, solitary or clus-

tered, Aoi or more ee ae from near base with a few to rarely numerous branches,

each beari larly enlarged or not, 5-20 mm in diameter; glabrous,

creamy siue: yellow, or tan, drying Piraeus: dark bone. or black. Leaves reduced to scales, 7-10 mm

long, cauline, spirally alternate, appressed, broadly ovate to ovate triangular, becoming narrower distally.

Inflorescence an open spike-like raceme, glandular pubescent; bracts mostly ca. 1 cm long, the distal ones

shortened, narrowly oblong-lanceolate, acute at the tip, pale lavender to dark purple. Flowers sessile or the

proximal ones on short pedicels, evenly and spirally inserted on the axis, subtended by 2 small, linear-subulate

bracteoles; calyces 7-11-13) mm long, purple, deeply 5-lobed, the lobes lance-linear to linear-subulate,

about equal in length, densely glandular pubescent; corollas (13—)15-22 mm long, constricted above the

ovary, strongly curved at the constriction, the tube nearly horizontal, exterior surface glandular puberulent

or pubescent, persistent, the tube narrow or slightly flared distally, cream-white sometimes tinted with

purple, the throat with yellow palatal folds, glabrous or pubescent on folds and in grooves with eglandular

trichomes; upper lip ca. 5 mm long, erect, divided about half its length, the 2 lobes triangular-acute, lavender

or purple, puberulent or densely pubescent with eglandular trichomes; lower lip 3-4 mm long, divided to

its base into 3 linear lobes with an acute apex, ca. 1 mm wide, pubescent with eglandular trichomes, pale,

often with 1-3 purple veins. Stamens 4, inserted on the corolla tube at its constriction, generally glabrous

or with a few scattered hairs, anthers ca. 1 mm long, white, glabrous or sparsely pubescent, equalling or

shorter than style. Ovary somewhat asymmetrically ovoid, 2-carpellate, nectary not evident; placentation

parietal, with 4 lateral placentae; style filiform, equaling the corolla tube or shorter, stigma peltate, discoid-

crateriform or rarely somewhat 2-parted Fruits 2-valved capsules, 0.7-1 cm long; seeds numerous, 0.3-0.5

mm long, ovoid to broadly ellipsoid, reticulate-pitted, light tan to dark brown. 2n = 48 (Collins 1973).

Flowering mid-August to early October.

The species ranges (Fig. 2) from the Potomac and Shenandoah River valleys of Virginia to southern Ohio

and Indiana along the Ohio River to Central Illinois along the Wabash and Sangamon Rivers, to the Meramec

River in eastern Missouri (near St. Louis), southward to a tributary of the Mississippi River in western Ten-

nessee, and westward along the Platte River in Nebraska and Eastern Colorado. Some apparently isolated

populations occur in the Kansas, Oklahoma, Texas, and New Mexico along the Canadian, Cimarron, and

Rio Grande rivers and their tributaries. Only historical collections are currently known from the District

of Columbia, Virginia, and West Virginia. The species is found entirely on mixed sand and silt deposits on

stream banks of eastern rivers or on sandbars or sandy banks of western rivers. Hosts are Ambrosia trifida

and Xanthium strumarium, and occasionally Ambrosia artemisiifolia and Dicoria canescens. Rarely reported

alternate hosts are Nicotiana tabacum L. and N. glauca Graham.

Representative specimens: UNITED STATES. COLORADO: Crowley Co.: cultiv: m ae near nee 22 Je 1948, ld: s.n.

(CS). Jackson Co.: N sandhills E of Cow uA 1 Oct Mine Wi pd 13922 is ).I ] near Ster-

== anonymous, 8 Sep 1965 (CS). P ,6mi E of Two Buttes Peak, 24 Aug 1964, Martin s.n. (COLO,

CS). Weld Co.: sandhills 3 mi NE of Roggen, 1 Sep 197, pud s.n. He CS). DISTRICT OF COLUMBIA: E end of New Long

Bridge, 2 Sep 1905, Ward s.n. (US). ILLINOIS: nim: ~ s of Embarras River, 2 mi E of Greenup, 18 Sep 1980, Collins

1618 (MO). Mason Co.: bank of Salt Creek and edg field, 5 mi N per cati - nis in os 1615 (MO). Menard Co.:

bottoms, Athens, 1860, 1861, 1866, 1867, E. Hall s.n. (F, GH, IL, MO, POM, WIS). V mel along banks of Wabash

River and Greathouse Creek, 18 Sep 1980, Collins 1619 (MO). INDIANA: Harrison Co.: W of ig at ER 25 Aug 1945,

Deam 63566 (IND). Jefferson Co.: tob field 0.5 mi E of Brooksburg, 15 Sep 1935, Banta s.n. (IND). Knox Co.: along ditch 2 mi N

x ud 9 Oct 1938, Deam 59255 Ends vieni Co.: E bank of Wabash River at New Harmony, 28 AE 1971, Collins 1555 (MO, WIS).

k, 6.5 mi SW of Carlisle, 1 Sep 1939, Deam 59424 (IND). Vandenburgh Co: river slough, 17

Oct 1941 Zeiner s.n. (IND), Warrick Co.: banks of Ohio River at Newburg, 28 Aug 1971, Collins 1543 (UWM). KANSAS: Allen Co.:

Iola, Sep 1925, anonymous (KSC). Greenwood Co.: potato patch near Eureka, 24 Sep 1935, Farmer s.n. (KS). Morton Co.: Cimarron

River bed N of Wilburton, Aug 1929, Gates 16205 (F, KSC, MO). KENTUCKY: Pendleton Co.: Ohio River bank at Ivor, 27 Nov 1942,

Chicoine 4745 (MO, US). MISSOURI: St. Louis Co.: a pes a bar on S bank of Meramec River, 2 mi N of Crescent, 1 Sep 1991,

Yatskievych & Colwell 91-195 (MO). NEBRASKA: B Platte River, 2 mi S of Odessa, 8 Oct 1999, Collins 1620 (MO,

s Co.: sandbar in Platte River, 2 mi NE of Louisville, 15 Sep D. Morrison 1342 (NEB). Dawson Co.: sandbar in Platte

River, S of Lexington, 8 Oct 1999, Collins 1621 (MO). Kearney Co.: sandbars of Platte River, Minden, 15 Sep 1922, Hapeman s.n. (PH,

POM/RSA, WTU). Lincoln Co.: sandbar in Platte River, Sutherland, 13 in 1936, anonymous (NEB). Merrick Co.: sandbar in Platte

River, near Central City, 22 Aug 1938, I. Mueller s.n. (NEB). NEW MEXICO: Doña Ana Co.: sand, od e 25 s 1906, Wooten

& Stanley s.n. (US). Sandoval Co.: sandy soil, Arroyo Chico, 22 Sep 1973, ee 1758 (ARIZ). San

River, 30 m S of Farmington, pipeline ee 7 on BIA Road 2, 4 sep 2007, Collins & Heil 2032 (MO). Socorro Co.: sand along Rio

Cani San Marcial, 1 Aug 1897, H k 82 ). Union Co.: idle field, 17.5 mi SW of Clayton, 1 Sep 1936, Cogdell 125 (GH). OHIO:

5 mi E of Chilo, 20 Oct 1949, Bardy s.n. (OS, US). Hamilton Co.: moni a ae Miami River, 25 pace wee Haller s.n.

(US). Scioto Co.: Nile Bae a Ohio River, 29 Aug 1949, zn s.n. An f Canadian

River, 10 mi NW of N , 8 Sep 1946, Goodman 4078 (MO). T dale C hofLower Forked

Deer Creek, ca.2 m NE of in 16 Oct 1972, Piehl s.n. (TENN). pois Brewster Co.: »: Boquillas Given. Chisos Mts., sand near

Rio Grande River bank, 5 Aug 1937, Warnock 838 (ARIZ, TEX, US). D dy soil, Rita Blanca Creek N of Dalhart, Aug 1949,

York & Rogers 233 (OSU, TTC). El Paso Co.: vicinity of El Paso, 1911, Stearns 40 (MO, US). Hartley Co.: deep sand on creek bank,

Punta del Agua Creek 30 mi SW of Dalhart, 26 Sep 1965, Collins 821 (OSU, TTC). Wheeler Co.: 5 mi 5 & 7 mi E of Miami, Jul 1967,

Baggerman s.n. (TTC, WIS, UWM). VIRGINIA: Campbell Co.: 8 mi from Lynchburg, 13 Oct 1933, Freer s.n. (US). Page Co.: banks of

Shenandoah River, Luray, 22 Aug 1912, Holtzman s.n. (US). Warren Co.: sandy field, Front Royal, 18 Sep 1897, Miller s.n. (US). WEST

RGINIA: Summers Co: New River, above Shanklin's Ferry, 23 Aug 1946, McNeill s.n. (WVA)

Eastern and western populations have an apparent disjunction in distribution between eastern Missouri

and east central Nebraska, a distance of ca. 650 km. A full list of specimens examined is available upon

request to the authors. It is uncertain whether this is a true disjunction or simply the lack of intensive field

work in this region to locate obscure populations (many botanists avoid botanizing in the midst of dense

Collins et al., Orobanche riparia, a new species

11000 100*0'0"w 90*go"W

80'00"W

TO*OO"W

50”0'0"N

1 20*00W

EAS

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EL ul

40"0'0"NH LFFEPEED ATE H

LEN

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Fic. 2. Distribution of Orobanche riparia.

e O. riparia Sems e

a E T

NE S

Q 125250 500 750 1,000

Kilometers

110"0'0W 400*00"W 9o*oo"W 80*0'0"W

g allergy season). The eastern and western populations of O. riparia

stands of ragweed and cockleburs d

exhibit slightly different morphologies. Plants of the eastern populations (including the Missouri plants)

are generally pallid with very pale lavender or purple pigmentation in the corollas. Pubescence is reduced

and sometimes almost absent, especially on the corolla lobes. The primary host is Ambrosia trifida. The

habitat of these populations is floodplains with mixed sand and heavy silty loam soils of the midwestern

rivers cited above.

The western populations (west of eastern Missouri) have more deeply purple-tinted corolla lobes,

calyx lobes, and floral bracts. The upper stems, where exposed to sunlight, may be infused with a rosy

purple. Pubescence is rather dense on the inflorescence, corolla lobes, and palatal folds. The primary host

is Xanthium strumarium. Populations of this variant are concentrated along sand bars of the Platte River in

Nebraska, with a few records of populations widely scattered in Kansas, Oklahoma, Texas, Colorado, and

New Mexico. Specimens of this species collected along the Rio Grande River from central New Mexico to

the Big Bend of Texas have proven problematic for identification throughout the study of Section Nothaphyl-

lon. Only recently, when they were compared with the Nebraska plants, did it become apparent that they

represent populations of O. riparia.

It is tempting to name these two variants formally as subspecies within O. riparia. However, the life

history and morphological distinctions are not sufficient to warrant taxonomic status at this time. This

problem requires more intensive study of both morphological and genetic variation within and between

populations before a conclusion can be reached as to the merits of segregating infrataxa within O. riparia.

Host information for Orobanche is generally difficult to assess from herbarium labels. Collectors sometimes

list as a host any adjacent plant without confirming a physical connection or list no host at all. However,

host data for O. riparia reported here have been observed by all three authors. Only two non-Asteraceous

MO Joumalof tani Insti Texas 3(

host reports have been confirmed. Some populations in southern Ohio occasionally parasitize cultivated

tobacco (Nicotiana tabacum). Likewise, some populations along the Rio Grande in western Texas have been

reported on the weedy, introduced N. glauca (the only perennial host with which the species has been associ-

ated). It is not possible at this time to judge He a of ak exceptions, if any. Anecdotal evidence

suggests that other species of Orobanche lternate hosts at the margins of their

ranges.

The populations east of the Appalachian Mountains are of particular interest because they represent

the extreme eastern portion of the range and may possibly be extirpated. Virginia records are represented

by four herbarium specimens from four counties (on the James, Shenandoah and Potomac rivers). The most

recent collection we have seen was made in 1933. A single specimen was seen from the New River, Summers

County, West Virginia. The present study could not confirm extant populations in the District of Columbia,

Virginia, or West Virginia. The species seems reasonably secure elsewhere in its range, although current

herbarium vouchers document a highly fragmented range. For example, the single Tennessee record rep-

resents a southward disjunction of ca. 325 km from the St. Louis County, Missouri station. However, field

experience of the authors suggests that this species is probably more common than the herbarium records

indicate. The senior author discovered several previously unreported populations in Nebraska, Illinois, and

Indiana. Additionally, a few populations were observed in Illinois, Ohio, and Texas that were located on

minor tributaries (creeks) of the major river systems. Some of the historical populations were determined

to be extant. Thus, it is likely that concentrated field work will reveal additional populations of O. riparia.

The status of the historical populations in the District of Columbia, Virginia, and West Virginia especially

needs to be updated by further field work.

The habitat of O. riparia is disturbed annually by levents and h tivity. Although it is widely

distributed, populations seem to be locally isolated and therefore fragile. The senior author has observed the

obliteration of a large population in Posey County, Indiana, as a result of riverbank development. Because of

its unique distribution of widely separated populations in regularly disturbed habitats, the species is likely

to be threatened on a local or state basis.

ACKNOWLEDGMENTS

The authors wish to express their appreciation to the staff and curators of the following herbaria for the

loan of specimens and/or for providing images and label data from specimens in the Orobanche riparia alli-

ance: A, ARIZ, COLO, CS, F, GH, ILL, IND, JEPS, KS, MICH, MIN, MO, NEB, NY, OSC, OKLA, PH, POM,

RSA, TENN, TTC, UC, US, UWM, WIS, WTU, and WVA. Collins is grateful to Ken Heil (San Juan College)

for accompanying him during field work in the southwestern U.S. Financial support for some of the field

work of Collins was provided by a grant from the Evangel University Faculty Summer Research Program.

Colwell's initial contributions to this project during her doctoral studies in St. Louis were supported by the

Department of Biology at Washington University. The Flora of North America Project additionally provided

funding for one of Colwell's trips to the Missouri Botanical Garden.

REFERENCES

Beck von MANNAGETTA, G. 1890. Monographie der Gattung Orobanche. Biblio. Bot. 4(19):78-85.

Beck von MANNAGETTA, G. 1930, Orobanchaceae. In: A. Engler, ed. Das Pflanzenreich: Regni vegetabilis conspectus

im Auftrage der Kóngl. preuss. Akademie der Wissenschaften vol. IV (261), issue 96. Wilhelm Engelmann,

Leipzig. Pp. 1-348.

BENNETT, J.R., AND S. MarraEws. 2006. Phylogeny of the parasitic family Orobanchaceae inferred from phytochrome

A. Amer. J. Bot. 93:1039-1051.

Cotuns, LT. 1973. Systematics of Orobanche section Myzorrhiza. Ph.D. dissertation, University of Wisconsin,

Milwaukee.

HeckarD, L.R. 1973. A taxonomic reinterpretation of the Orobanche californica complex. Madroño 22:41-104.

Collins et al., Orobanche riparia, a new species 11

KARTESZ, J.T. 1994. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland, 2

vols. Timber Press, Portland, Oregon.

Munz, PA. 1930. The North American species of Orobanche, section Myzorrhiza. Bull. Torrey Bot. Club 57:

611-624.

Munz, PA. 1974. A flora of southern California. Univ. of California Press, Berkeley.

MussLeman, L.J. 1982. The Orobanchaceae of Virginia. Castanea 47: 266-275.

NurraLL, T. 1818. The genera of North America plants, and a catalogue of the species, to the year 1817, 2 vols.

Published by the author (printed by D. Heartt), Philadelphia, Pennsylvania.

OLMSTEAD, R.G., C.W. DePampnitis, A. Wotre, N.D. Young, W.J. ELison, AND P. Reeves. 2001. Disintegration of the Scrophu-

lariaceae. Amer. J. Bot. 88:348-361.

ScHNEEWEISS, G.M., A. CouweLL, J.-M. Park, C.-G. JANG, AND T.F. Steussy, 2004. Phylogeny of holoparasitic Orobanche

(Orobanchaceae) inferred from nuclear ITS sequences. Molec. Phylogen. Evol. 30:465-478.

Shreve, F. AND ILL. Wicains. 1964. Vegetation and flora of the Sonoran Desert, 2 vols. Stanford University Press,

Stanford, California.

SrRAUsBAUGH, P.D. AND E.L. Core. 1978. Flora of West Virginia, ed. 2. Seneca Books, Grantsville, West Virginia.

I Al S£ Das anli D L De if

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ied Bent ud den San more uA He 000 d on e to Pe at the Le DRS at the TRE time to take

ng ml

Nevada

r O E

1 1 ET * [gens | 1 p A ful E Ee Lae 1 + 1

r r Eu ao, e ? r

Basin Desert, Mojave Desert, and Colorado Desert (part of the Sonoran Desert). The intent of tl horis to span tl dership f

] public, naturalist, a t li ] iati f nati l h tion, and he deserves five

o r T: rr Pul

stars for surpassing the highest standards.

This book is divided into five chapters: (1) Evolution, Classification, and Botanical Characteristics; (2) Ecology and Habitats of

Cacti, Agaves, and Yuccas; (3) HK Profiles; (4) Conservation and d and (5) Exploring the Cactus Country of California

and Nevada. The eight appendices a lly informative and increase the value of the book: Cactus, Agave, and Yucca Species of

California and Nevada; Men Vegetation Types and ppt for Cacti, Dee dy Yuccas; senec Names ie Non- ates Species;

Cholla Species C C

Species List for ME Public tended in Califonia and Nev ada

The species profiles for each entry includ name, the scientific name, a phonetic pronunciation guide, recent syn-

elie oth common dé a commentary al he plant, a landscape, habit, and flower photograph pue a range map. Ad ing

the species. The chapt

ipti f the habit, stems, spines or leaves, flowers, and f pprot

on oo ation 2 Cultivation”: pecially y ] ion i Siena aves cultivation

A E 3111 1 1 1 1 1

? li ] ific si 1 lected i be found on

bs section on on "Exploring n Cactus Country of calina ae Ney ada

ate parks, national parks, nati I sildeenéss areas, wildlife ee and recreation areas.

c : 1 wa. : 1 ] id i y common, in igenous rare I new species are

th llent | th dagl hat defi ] inol lin the book. The 15 genera of Cactaceae due cli

that occur in California. Ru Nevada are a by d ios ao paintings by Eugene O. Murman and th M dd Susan

text. For cad “Packrat Middens me Vegetation Histor

*Yuccas and Yucca Moths" in both dd to tl iosi iated plants, especially the yucca moth-flower o

biology often featured in biology textbooks as obligate un.

tho push the Pus are not M in ue text, the pees has more n 200 citati ly published in the last ten

Bazell. T

*t*r.l

ears. J ing

ted afr a : 1 decr ] A px

by g genus and species li

s book must ls on Hee E a did WB. botanist, a: on o landscaper, conservationist,

dde]

E E

E 1 L JE 1 1 L th

and national parks E in Western states. Cachuma Press, the editors

will be cherished by all bibliophilesi—Harold W. Keller, Ph.D., Research Associate Botanical Research Institute of dois Fort Worth, Texas

76102-4025, U.S.A

J. Bot. Res. Inst. Texas 3(1): 12. 2009

BRISTLECONE PINE DWARF MISTLETOE:

ARCEUTHOBIUM MICROCARPUM SUBSP. ARISTATAE (VISCACEAE),

A NEW SUBSPECIES OF WESTERN SPRUCE DWARF MISTLETOE

FROM NORTHERN ARIZONA

Jared M. Scott Robert L. Mathiasen

School of Forestry School of Forestry

Northern Arizona University Northern Arizona University

Flagstaff Arizona 86011, U.S.A. Flagstaff, Arizona 8601 1, U.S.A.

Jared Scott@nau.edu Robert. Mathiasen@nau.edu

ABSTRACT

We describe Arceuthobium microcarpum subsp. aristatae (Viscaceae), a parasite of bristlecone pine (Pinus aristata Engelm.) and

Engelmann spruce (Picea engelmannii Parry ex Engelm.) in northern Arizona. oe and aer Me data were used to com-

pare Á. microcarpum populations parasitizing bristlecone pine on the San Francisco Peaks, A a, with populations on Engelmann

ME and a e (Picea pungens a from nto Pid in a and New Mexico. diit ieu e phenological, and

Francisco Peaks and Kendrick

rr r

Peak in northern veneer as a new als

RESUMEN

Se describe A thobi i p subsp. aristatae (Vi ,un e de Pinus aristata NEN y de Picea engeimannii

Parry e Engelm en el norte de Arizona. S 1 fol 5gi y g I p de A. microcarpum que

D. 1

r 1 F

Shee [PES f cee wees Aéxic Dis fala f, J hili Jad Anal

parasitan a Pi istata de San F i , Arizona, blaci i Pi l ti APEA EN Pod

mentan la clasificación de las SNAM de A A. microcarpum de San Fand Peaks y de eig Peak en el norte de Arizona como

una subespecie nueva

Key Wonps: Arceuthobium, Picea engelmannii, Picea pungens, Pinus aristata, Arizona

Western spruce dwarf mistletoe (Arceuthobium microcarpum (Engelm.) Hawksw. & Wiens) is an important

parasite of blue spruce (Picea pungens Engelm.) and Engelmann spruce (Picea engelmannii Parry ex Engelm.)

in the southwestern United States (Mathiasen et al. 1986; Hawksworth & Wiens 1996). It occurs as far

south as the Pinaleño Mountains, Arizona and Sacramento Mountains, New Mexico and as far north as the

Kaibab Plateau, Arizona. However, western spruce dwarf mistletoe is most common in the White Moun-

tains, Arizona and Mogollon Mountains, New Mexico (Mathiasen et al. 1986; Hawksworth & Wiens 1996).

On the San Francisco Peaks, Arizona, it also parasitizes Rocky Mountain bristlecone pine (Pinus aristata

Engelm.) (Mathiasen & Hawksworth 1980). Initially, the dwarf mistletoe infecting bristlecone pine on the

San Francisco Peaks was classified as limber pine dwarf mistletoe (Arceuthobium cyanocarpum (A. Nelson

ex Rydb.) Coulter & Nelson) by Hawksworth and Wiens (1972). It was later classified as A. microcarpum

based on its host affinities and chemical characters (flavonols) (Crawford & Hawksworth 1979; Mathiasen

& Hawksworth 1980; Hawksworth & Wiens 1996). Although Mathiasen and Hawksworth (1980) reported

morphological, phenological, and host susceptibility differences between the A. microcarpum populations

on the San Francisco Peaks and other A. microcarpum populations in the Southwest, they did not recom-

mend giving the San Francisco Peaks populations taxonomic status at that time. In 2006, we began a more

detailed analysis of the morphological characteristics of the A. microcarpum populations in Arizona. Our

data further substantiated the differences between the A. microcarpum populations on the San Francisco

Peaks and those in other mountain ranges of Arizona first reported by Mathiasen and Hawksworth (1980).

Because of the differences in plant size, plant color, phenology, host affinities, and geographic isolation of

J. Bot. Res. Inst. Texas 3(1): 13 — 21. 2009

14 J t ical Insti Texas 3(

the dwarf mistletoe populations on the San Francisco Peaks, and nearby Kendrick Peak, there is sufficient

evidence to describe these populations as a new subspecies of A. microcarpum.

Arceuthobium microcarpum (Engelm.) Hawksw. & Wiens subsp. aristatae J.M. Scott & Mathiasen, subsp.

nov. (Figs. 1-2). Tre: U.S.A. Arizona. Coconino Co.: near summit of Schultz Peak, a southeast subsidiary ridge of the San

Francisco Peaks, Coconino National Forest, elev. 3,060 m, Lat. 35? 18' 43" N, Long. 111? 37' 52" W, parasitic on Pinus aristata, 8

Aug 2006, J.M. Scott 2006-6 ( : ASC; IsoTYPES: JEPS, UNM, US).

Plantae (1237) lt li principales basi (121.8(—3) mm diam; fructus maturi 3.3 x 2.1 mm; anthesis mense Julio-Augusto;

antde i=;

; F F

fructus maturitas Augusto—Septembri; in Pinus aristata et Picea engelmannii parasiticae.

Male plants (0.8-)2.7(-7.0) cm in height, basal diameter of dominant plants (1.0-)1.8(3.0) mm; female

plants (1.4—)3.6(-7.0) cm in height, basal diameter of dominant plants (0.6-)1.8(3.0) mm; male and female

plant plants primarily light green, but some brown-green or purple; male flower diameter 2.5 mm, perianth

lobe dimensions 1.2 x 1.0 mm, anther diameter 0.5 mm, distance from anther to perianth tip 0.5 mm; fruit

dimensions 3.3 x 2.1 mm; seed dimensions 2.4 x 1.1 mm.

Common name.—bristlecone pine dwarf mistletoe.

Etymology.—We selected aristatae, a name that indicates the principal host of this dwarf mistletoe is

Pinus aristata.

Phenology.—Anthesis from early July to late-August with peaks in late-July to early-August; seed dis-

persal from mid-August to late-September with peaks in mid- to late-August to early-September.

Habit.—Parasitic principally on Pinus aristata and occasionally on Picea engelmannii on Schultz Peak,

parasitic principally on Picea engelmannii in the Inner Basin of the San Francisco Peaks and on nearby Ken-

drick Peak. Not known on Picea pungens. Rarely infects Pinus flexilis James and Abies lasiocarpa (Hook.) Nutt.

on Schultz Peak.

Distribution.—Bristlecone pine dwarf mistletoe only occurs in three small populations on the San

Francisco Peaks (Schultz Peak, Weatherford Trail, and Inner Basin, Fig. 3). It also occurs near the summit

of Kendrick Peak (Fig. 4). It ranges in elevation from 2890 to 3130 m on Schultz Peak.

METHODS

Collection Locations.—1n 2006 and 2007, plants of A. microcarpum were collected from 18 populations

in Arizona (Appendix A; Appendices A—F available online at http://www.for.nau.edu/SOFArchive/MSF-

Students/JMScott/20090325.pdf). Four populations were sampled near Flagstaff, Arizona (host plants):

Schultz Peak (bristlecone pine and Engelmann spruce), Weatherford Trail (bristlecone pine), Inner Basin

(bristlecone pine and Engelmann spruce) (Fig. 3), and Kendrick Peak (Engelmann spruce) (Fig. 4). A total

of 12 populations of A. microcarpum were sampled in the White Mountains (host plants): Cache Cienega

(blue and Engelmann spruce), Coleman Creek (blue spruce), Hannagan Creek (blue spruce), Johns Canyon

(blue spruce), Lee Valley (blue spruce), Williams Valley (blue spruce), Forest Road 249 (blue spruce), For-

est Road 402 (blue and Engelmann spruce), and Forest Road 72 (blue and Engelmann spruce) (Fig. 4). In

addition, two populations were sampled from the North Rim of the Grand Canyon: Point Royal Road (blue

spruce) and Route 67 (blue spruce).

In 1975 and 1976, plants of A. microcarpum were collected by the junior author from the same loca-

tions and hosts on the San Francisco Peaks that were sampled in 2006-07. Plants were also collected from

the North Rim of the Grand Canyon (blue spruce), the White Mountains (blue and Engelmann spruce), the

Mogollon Mountains (blue spruce), and the Pinaleño Mountains (Engelmann spruce) (Fig. 4).

Morphological Measurements.—The following morphological characters of male plants were measured:

dominant plant height, dominant plant basal diameter, flower diameter, anther diameter, perianth lobe

length, perianth lobe width, and distance from the outer edge of the anther to the tip of the perianth lobe.

The following morphological characters were measured for female plants: dominant plant height, dominant

plant basal diameter, length and width of both fruits and seeds. The color of plants, fruits, and seeds were

recorded. Plant heights and basal diameters were measured with digital calipers to the nearest 0.1 cm. All

Cents his ; A thah: : H aricfataa 15

Plants of Arizona

VISAS

tatae

Arceuthobium microcarpam

sap. aris da M; "ems

Mathiassen

rona Tesoro C6 Cocenmo NF Servite

Eevu tOnoze

Habitat: On birstiecone pine

Jared Scot 2006.8

23 August 2005

Northern Arizona University (NAU)

Fic. 1. Holotype of Arceuthobium microcarpum subsp. aristatae J.M. Scott & Mathiasen, subsp. nov., J.M. Scott 2006-6, 8 Aug 2006 (ASC). Photo by R.L.

Mathiasen.

; Arizona. Photo

F f

7,714 2008 hv 1 M Scott (Rott 1 Mala A

taken 30

May 2008 by J.M. Scott.

Schultz Peak, Arizona. Photo

Scott and Mathiasen, Arceuthobium microcarpum subsp. aristatae

San Francisco Peaks

Arizona

Fic. 3. The four infestations of western spruce dwarf mistletoe on the San Francisco Peaks, AZ. Dark circles indicate an infestation on bristlecone pine,

white circles indicate an infestation on Engelmann spruce, half-dark and white circles indicate an infestation on both bristlecone pine and Engelmann

Spruce.

A LL ÉEc

North

San

D . Francisco

Kendrick Peaks

Peak @ a

New

. White Mexico

Arizona „Mountains

ss”

e Mogollon

Mountains

inaleño

Mountains

PI te locations for plant collections of western spruce dwarf mistletoe in 1975 and 1976 (gray squares) and in 2006 and 2007 (closed

circles). Plant lj 1f, : ER nara dd Sec a ap liqus 2007 (dosi

of the Grand Canyon.

other morphological characters were measured using a 10x hand lenses with a micrometer to the nearest

0.1 mm. Male plants were collected during peak anthesis and female plants were uda NE fruits were

mature. Over E male or female ps were collected for each population and

were comp

characters that were measured in 2006—07 were measured lou the 1970s ae Characters EXER for both

datasets included male and female plant height and basal diameter, flower diameter, perianth lobe length

and width, fruit length and width, seed length and width, and plant color.

Phenology Observations.—Anthesis and seed dispersal data were recorded when plants were collected

for morphological measurements in 1975, 2006, and 2007. Data for the last two years were primarily from

the San Francisco Peaks and Kendrick Peak due to frequent visits to these areas. During these years, the

White Mountains were only visited twice each year: once during anthesis and again during seed dispersal.

Therefore, the precise times of peak anthesis and seed dispersal were estimated for the White Mountains

in 2006-07. In 1975, phenology data was collected from the San Francisco Peaks, Kendrick Peak, White

Mountains, and Pinaleño Mountains, Arizona.

Host Susceptibility.—In 2008, temporary 0.012 ha (radius 6 m) plots tablished around 26 severely

infected bristlecone pines on Schultz Peak and 13 severely infected Engelmann spruces in the Inner Basin.

In each plot, all trees > 1.4 m in height were examined for dwarf incas dud For each tree, species,

diameter at breast height (dbh; to the nearest cm), and a dwarf mistl g R, Hawksworth 1977) were

] qos 1

] using ten randomly selecrea 1 plants f IOT each led Not el of

c 4 I Mash f A hah’ ` | | I aristatae 19

recorded. These data provided information on the relative susceptibility of bristlecone pine and Engelmann

spruce to dwarf mistletoe infection on Schultz Peak and for Engelmann spruce in the Inner Basin.

Data Analysis. Morphological data were combined from hosts on the San Francisco Peaks (bristlecone

pine and Engelmann spruce) and Kendrick Peak (Engelmann spruce) and also for the White Mountains

and North Rim (blue and Engelmann spruce) for analyses. Morphological data from 1975 and 1976 were

included within these two data sets, but the 1970s data also included morphological measurements from

the Mogollon and Pinaleño Mountains. A one-way analysis of variance (ANOVA, P-value < 0.05) was used

to test for significant differences between the means of each morphological character measured.

RESULTS

Morphological Data.—On average, male and female plants of western spruce dwarf mistletoe (WSDM, subsp.

microcarpum) were larger than those of bristlecone pine dwarf mistletoe (BPDM, subsp. aristatae) and the

differences in size were significantly different (Table 1, Appendices B-E, online). Although the largest plants

we measured in 2006 and 2007 were collected from blue spruce in the White Mountains (Appendix B), the

largest plants measured in the 1970s were from Engelmann spruce in the Pinaleño Mountains (Appendix

E). Plant heights for the 1970s data were slightly larger than those for the 2006-07 data for BPDM, but both

sets of data demonstrated that plants of WSDM were larger than BPDM. The mean basal diameter of male

plants was similar, but the mean basal diameter of female WSDM plants was larger on average than female

plants of BPDM and significantly different. The means for flower diameters, fruit lengths, and seed widths

were significantly different, even though the differences were only about 0.1 mm or less (Table 1).

The color of male and female plants of BPDM was commonly light green or green-brown. However,

many plants on the San Francisco Peaks appeared purple (Fig. 2). The color of male and female plants of

WSDM was similar to BPDM, except that some plants in the White Mountains were green-blue.

Phenology.—Peak anthesis for BPDM occurred one to two weeks earlier on the San Francisco Peaks

than for WSDM in the White Mountains in 2006 and 2007. Seed dispersal of BPDM also starts and ends

one to two weeks earlier on the San Francisco Peaks than seed dispersal of WSDM in the White Mountains.

The 1970s observations also found that BPDM flowers and disperses seed earlier than WSDM (Mathiasen

& Hawksworth 1980)

Host Susceptibility.—Infection of bristlecone pine on Schultz Peak was 9696 (n = 111) indicating it is a

principal host of bristlecone pine dwarf mistletoe there. However, infection of Engelmann spruce was only

3496 (n = 224) on Schultz Peak placing it in the occasional host class of Hawksworth and Wiens (1996).

Engelmann spruce is clearly much less susceptible than bristlecone pine on Schultz Peak. However, infec-

tion of Engelmann spruce in the Inner Basin, just 5 km north of Schultz Peak, was 94% (n = 101). All of

the bristlecone pines growing near severely infected Engelmann spruces in the Inner Basin were infected,

but there are only about 10 bristlecone pines in the area. We also observed rare infection of limber pine (2

trees) and subalpine fir (3 trees) by BPDM on Schultz Peak.

Table 2 summarizes the principal morphological and physiological differences between BPDM and

WSDM used to distinguish the two subspecies. The shoot heights, colors, and phenology, of BPDM on the

San Francisco Peaks and Kendrick Peak were similar enough to group them as one population. Morpho-

logical data from Mathiasen and Hawksworth (1980) and our data demonstrated that BPDM plant heights

were smaller than for WSDM populations in southern Arizona and central New Mexico. The differences in

mean plant height between BPDM and WSDM were statistically significant. Additionally, color differences

were found between BPDM (purple plants) and WSDM (blue-green plants). Furthermore, the periods of

flowering and seed dispersal for BPDM on the San Francisco Peaks occurred one to two weeks earlier than

for populations of WSDM in the White Mountains in 1975-76 and 2006-07.

Another important set of A ecc characteristics that separates BPDM from WSDM are

their host affinities. Bristlecone pine pal host of BPDM on Schultz Peak (9696 infection), but we

found that only 3496 of the Engelmann Soria were infected there. Mathiasen and Hawksworth (1980) also

TABLE 1. por : morphological characters of Arceuthobium microcarpum subsp. aristatae and A. microcarpum subsp.

microcarpum. Data combine measurements from and 2006—07 for all hosts and are presented as means (ranges)

[n]. Characters ul an asterisk had significantly different ica Using ANOVA (P < 0.05). The means for seed width were

rounded to the nearest 0.1 mm, but actual values g different.

Character A. microcarpum subsp. aristatae A. microcarpum subsp. microcarpum

Plant Height (cm)

Male* 2.7(0.8-7.0) [152] 5.6(1.8-14.9) [283]

Female*

Basal Diameter (mm)

Male

Female*

Flower Diameter SUBE

3.6(1.4-7.0) [177]

1.8(1.0-3.0) [121]

1.8(0.6-3.0) [167]

2.5(1.8-4.0) [287]

3:3(3.5-5.1) [281]

2.1(1.7-2.9) [281]

6.4(2.0-15.7) [353]

1.9(0.8-3.4) [157]

2.0(0.8-3.8) [313]

2.4(1.6-3.1) [266]

34(3.4-5.2) [440]

2.2(1.9-3.1) [440]

Seed Length (mm) 24(1.5-34) [107] 24(1.3-3.4) [224]

Seed Width (mm)* 1.1(0.8-1.4) [107] 1.1(0.7-1.5) [244]

Tagle 2. Summary of the principal E and physiological differences between Arceuthobi j 1 ubsp.

aristatae and A. microcarpum subsp. microcarp

Character A. microcarpum subsp. aristatae A. microcarpum subsp. microcarpum

Mean Plant Height (cm)

Male Plants 27 5.6

Female Piants 36 64

Plant Color Light green; green-brown; purple Light green; green-brown; blue-green

Peak Flowering Period One to two weeks earlier than One to two weeks later than

subsp. microcarpum subsp. aristatae

Host ad

Bristlec ne Principal Unknown

ey spruce Principal/Occasional? Principal

Blue spruce nknown Principal

Limber pine Rare Unknown

Subalpine fir Rare Rare

Southwestern white pine Unknown Immune

'Host susceptibility classificati j described in Hawl th and Wiens (1996). The host susceptibility designa-

tions used for A, nin un micorcarpum are based on information in Hawksworth and Wiens (1996) also.

25 | | fiar

| host for Subsp: aristatae in the Inner Basin of the San Francisco Peaks and on

Kendrick Peak, Arizona, but it was an sal host of | n Schultz Peak.

reported that bristlecone pine was the principal host of BPDM on Schultz Peak, and they reported a similar

level of infection for Engelmann spruce (32%), but they used a different method of collecting infection data;

a systematic sampling design using 27 rectangular plots. Although they classified Engelmann spruce as a

secondary host of BPDM on Schultz Peak, the low incidence of infection we found indicates Engelmann spruce

should be classified as an occasional host there. Based on our observations and those of Lynch (2004), blue

spruce is more susceptible to infection by WSDM than Engelmann spruce, but both are principal hosts of

WSDM. Peas blue spruce has not been reported on Kendrick Peak or the San Francisco Peaks, we were

unable to infection data for BPDM on this host. There is also a report of dwarf mistletoe on Chihuahua

spruce (Picea chihuahuana Martinez) in northern Mexico by Ledig et al. (2000) which we assumed would be

WSDM. However, our examination of several Chihuahua spruce populations in Chihuahua and Durango

Scott and Mathi , A thobi i | bsp. aristatae 21

reported to be infested with dwarf mistletoe, indicated that the spruces were infected with spruce broom

rust (Chrysomyxa arctostaphyli Dietel) and not dwarf mistletoe. Spruce broom rust induces the formation of

witches' brooms on Chihuahua spruce that are similar to those caused by dwarf mistletoes (Cibrián et al.

2007), which was undoubtedly the reason for the report of dwarf mistletoe on Chihuahua spruce in Mexico.

Therefore, none of the Mexican species of Picea have been reported as the host of a dwarf mistletoe thus far

(Hawksworth et al. 2002).

Hawksworth and Wiens (1972, 1996) defined a subspecies as geographically restricted populations

having small, but consistent variations. Variations in dwarf mistletoe plant size, color, host range, and phe-

nology have been used as the primary characteristics to identify subspecies of dwarf mistletoe (Hawksworth

& Wiens 1965; Hawksworth & Wiens 1972, 1977, 1996; Hawksworth et » T Wass & Mathiasen 2003;

Mathiasen 2007; Mathiasen & Daugherty 2007). Furthermore, the g lation of the San Francisco

Peak/Kendrick Peak BPDM populations is also consistent with Hw and Wiens (1972, 1996) defini-

tion that a subspecies should be a "geographically restricted population."

ACKNOWLEDGMENTS

We thank M. Socorro González Elizondo for the Spanish translation for the Resumen. We also appreciate the

field assistance of James White and Laine Smith in northern Arizona and Brian Howell in northern Mexico.

Gregory M. Filip and Del Wiens provided helpful suggestions to an earlier version of the manuscript.

REFERENCES

CIBRIAN, T.D., D. ALVARADO, AND S.E. GARCIA. (EDS.). 2007. Forest diseases in Mexico. Universidad Autonoma Chapingo,

Mexico.

CRAWFORD, D.J. AND F.G. HawksworTH. 1979. Flavonoid chemistry of Arceuthobium (Viscaceae). Brittonia 31:212-216.

HawkswoRTH, F.G. 1977. The 6-class dwarf mistletoe rating system. Gen. Techn. Rep. R.M. U.S. Forest Serv. 48:1-7.

Hawksworth, F.G. AND D. Wiens. 1965. Arceuthobium in Mexico. Brittonia 17:213-238.

HawksworTH, F.G. AND D. Wiens. 1972. Biology and classification of dwarf mistletoes (Arceuthobium). Agriculture

Handbook 401, USDA Forest Service, Washington, D.C.

Hawksworth, F.G. AND D. Wiens. 1977. Arceuthobium in Mexico: Additions and range extensions. Brittonia 29:

411-418

HAWKSWORTH, F.G. AND D. Wiens. 1996. Dwarf mist! biology, pathology, and systematics. Agriculture Handbook

709, USDA Forest Service, Washington, D.C.

HAwKSWORTH, F.G., D. Wiens, AND B.W. Gas. 2002. Arceuthobium in North America. In: Mistletoes of North American

conifers. BW. Giels and T.D. Cibrian, tech. coords. Gen. Techn. Rep. R.M.RS. U.S. Forest Serv. 98:29-56,

HAWKSWORTH, F.G., D. Wiens, AND D.L. NickRENT. 1992, New western North American taxa of Arceuthobium (Viscaceae).

Novon 2:204-211.

LEDIG, Eis M. ebay el B. diia V. HERNANDEZ Reyes, C. FLORES LOPEZ, AND me bin ARTEAGA. 2000.

Mexico and the d | f Pi Madroño

Locations of g | por graphy of!

47:71-88.

LyncH, A.M. 2004. Fate and characteristics of Picea damaged by Elatobium abietinum (Walker) (Homoptera:

Aphididae) in the White Mountains of Arizona. W. N. Amer. Naturalist 64:7-17.

Mathiasen, R.L. 2007. A new combination for Hawksworth's dwarf mistletoe (Viscaceae). Novon 17:217-221.

MATHIASEN, R.L. AND C.M. DaucHerry. 2007. Arceuthobium tsugense subsp. amabilae, a new subspecies of hemlock

dwarf mistletoe (Viscaceae) from Oregon. Novon 17:222-227.

MATHIASEN, R.L. AND F.G. HAwkswonrH. 1980. Taxonomy and effects of dwarf mistletoe on bristlecone pine on the

San Francisco Peaks, Arizona. Res. Pap. R.M. 224:1-10

MATHIASEN, R.L., FG. HAw«swonrH, AND C.B. EDMINSTER. 1986. Effects of dwarf mistletoe on spruce in the White Moun-

tains, Arizona. Great Basin Naturalist 46:685-689.

Wass, E.F. AND R.L. MATHIASEN. 2003. A new subspecies of A g (Viscaceae) from British Columbia

and Washington. Novon 13:268-276.

BOOK REVIEW

Arnos Farjon. 2008. A Natural History of Conifers. (ISBN 978-0-88192-869-3, hbk.). Timber Press, Inc.,

The Haseltine Building, 133 S.W. Second Avenue, Suite 450, Portland, Oregon 97204, U.S.A. (Orders:

www.timberpress.com, 1-800-827-5622). $34.95, 304 pp., 157 color photos, 39 line drawings, 7 3/8"

x 10 3/8".

1 A 3M» 3115 E ‘A J E J 11 x x 1 A x zt

plant group y g y This is what we have in

this volume. Mr. Farjon, retired head of tł ti f the Kew Herbarium, is an obviously dedicated botanist and always had

a working knowledge of the conifers. | i d decidi ite a book on th j d porn) them all over again.

io we are dus an extensive T of the conifers - enlightening naper on corrected classificati y methods

t y o DNA studies, g ore accurate ob of the group.

T} 1 fé if, JL 1 es D 1 AR th lah

their viia is Hong and involved. DH course w are e also a pano of the pope Since some are fast-growing and quite dense. their

connecti is very important

b 2 2 E

ff, ] ¡lod inf, ; hath th

1 ae ne |

TI iting should be of i I ienti ientist alike. The photographs by tl I ;

some of which depict trees you would have to travel long, difficult ways to see for yourself, make this a wonderful book.—Elaine Bell,

Volunteer, Botanical Research Institute of Texas, Fort Worth, Texas 76102-4025, U.S.A.

J. Bot. Res. Inst. Texas 3(1): 22. 2009

A NEW CLIFF-DWELLING SPECIES OF ZAMIA (ZAMIACEAE) FROM BELIZE

Michael Calonje

Montgomery Botanical Center

11901 Old Cutler Road

iami, Florida 33156, U.S.A

michaelcamontgomerybotanical.org

ABSTRACT

Zamia meermanii (Zamiaceae), a new cliff-dwelli f l Belize is described. It is distinguished by having 1 to 3 pendent

iid EE coriaceous ue that are entire or r erenulately nee on e uis un MM on A gda surface, and

ich it most closely resembles, as well as to

E d

M : Jiff_q 1 š 7 ; hha} Gach Cchut & Deh

or r£ O

RESUMEN

z> do Roli Cantal Pee € PD po :

Se describe Zamia meermanii oe j tral q precip p guir porque

tiene de l a 3 f T lad ] ee ee eee 2m

t T Y

en la superficie adii. y persi la superficie abaxial. Esta especi I Zamia furfi L.f., la especie más

his 1 T : des "e E Z3

semejante, yt Y A 1 4 A t

Vovides, Schut & Dehgan y Z lovalii Nelson

INTRODUCTION

Zamia is a New World genus ranging from the southeastern USA to Bolivia. It currently comprises 57 species

(Hill et al. 2007) and is considered to be the most ecologically and morphologically diverse cycad genus

(Norstog & Nicholls 1997). It includes species growing in areas with extremely high rainfall, such as Z.

roezlii Linden from the Colombian Chocó region, and adjacent coastal Ecuador to species growing in full sun

in extremely dry conditions, such as Z. encephalartoides Stevenson from the Santander region of Colombia.

The genus also includes the only known obligately epiphytic cycad, Z. pseudoparasitica Yates, as well as two

cliff-dwelling species: Z. cremnophila Vovides, Schutzman & Dehgan, and Z. sandovalii Nelson.

Jan Meerman, a Dutch ecologist who resides in Belize, li a third cun dwelling species of Zamia

in February of 1999 together with Martin Meadows of the Belize Botanic G during a Rapid Ecological

Assessment of a private protected area. Since discovering the plant, Meerman has peen collecting ecologi-

cal and distribution data for this species, some of which is used in this description. He was instrumental

in bringing a team from Montgomery Botanical Center to Belize to study this taxon and other Belizean

Zamiaceae in August and September of 2008.

DESCRIPTION

Zamia meermanii Calonje, sp. nov. (Figs. 1-2). Tr: BELIZE. Beuze D : 60 m, 28 Aug 2008, M. Calonje, J. Meerman &

P Griffith BZ08-152 (moLoryre: BRH; isotypes: FTG, MO, NY, XAL).

Species insignis habitui pulicol dex brevis, ape pendulis y litus; foliola coriacea ginil integris vel crenulatis

triente apicali, is distinctis adaxialil pagini 3 : pro]

Stems globose to cylindrical, to 27 cm long and 18 cm wide, MM or occasionally branching on older

plants. Cataphylls chartaceous, stipulate, triangular with lanceolate apex. Leaves 1—3 per stem, pendent,

2-146.5 cm long. Petiole 26-60 cm long with abruptly swollen base 2-3 cm wide, variously unarmed or

carrying numerous prickles (50+) up to 3 mm tall. Rachis 50 to 109 cm long, unarmed or bearing a few

prickles (« 20). Leaflets 7—23 pairs per leaf on adult plants, jam 3- > cm c from each other with the

point of attachment to the rachis 8.5-13.3 mm wide, obovate t narrowly oblong, coriaceous,

12.5-32 cm long, 3.4-7.5 cm wide, with 31-56 veins prominently raised on adaxial surface, visible but not

J. Bot. Res. Inst. Texas 3(1): 23 — 29, 2009

Fic.1 (A) Habit of plant 1 M Inge ME fal . ight (Di E ey ee | : gl £4 le, E, " L hs y (0 Detail

of trunk and petioles. (D) Variation in leaflet sh (E) à f daxial surface of leaflet showi ised vei d late t

flat L £: " lnc el : ETT RAT fev ct £ al. :4l £. laaflat L

duit leaf. A and C rep t Calonje et al. BZ08-152; B and F represent Calonje et al. BZ08-156; D, E and G represent Calonje

et al. BZ08-125.

Calonje, Zamia meermanii, a new cliff-dwelling species from Belize

[M 4 : : vn hl IDVA Emi Ave ile 2 IL 1 f^ id lu £ mat vll

Fig. 2

f J t F 3 pi F t ve) b LÁ

(PI Ad 1 £ hall /EY RA : tun? a PA (E\ Ra H trnhil $ 1 dehicri (G) Mature

V F PM SAT ort J f 3 > 3 1

J uir nne "m el hull Dhatanranhe A E£ m 1 : r RUD + al fall E ire

2 rJ q“ t d 2"

kil hab: lew RA f RA J u tc Calanie et al. R708-

y j 152.

26 J t ical i Texas 3(

protuberant on abaxial surface; margins entire or crenulately notched in the distal third; surface on emergent

leaves reddish-brown, but densely white-tomentose, turning cream-colored with saffron-orange tomentum,

and gradually maturing to green and glabrous adaxially, with some persistent brown tomentum abaxially

along leaflet margins and near the po of Su to the rachis. Eophylls typically carrying a single

pair of ovate leaflets. } te strobilus conica l-cylindrical, erect, at pollen release 13-16 cm long,

2.2—2.5 cm diam. pu individually or in groups of 2—5 or more, cream to brown colored; peduncle

9-10 cm long, 0.9-1 cm diam., covered by ferrugineous pubescence. Microsporophylls arranged in 12-18

columns, 17-30 rows; obtrullate, distal face hexagonal to oblong-hexagonal, 5.5-6 mm wide, 2.5-3 mm

tall, and 1-2 mm thick ; microsporangia spheroidal, 1-1.6 mm diam., present on abaxial surface, absent

on adaxial surface, 22-28 per microsporophyll arranged in two separate groups along margins. Megaspo-

rangiate strobilus cylindrical, solitary, erect at maturity, 12-20 cm long, 6-7 cm diam., tan to light-brown

tomentose from emergence to maturity; peduncle 8-12 cm long, 1.5-3 cm wide, tan to brown tomentose.

Megasporophylls arranged in 6-10 columns, 7-15 or more rows; distal face oblong-hexagonal, 1.5-2.2 cm

wide, 1.0—1.5 cm tall, 1-1.2 cm thick, extruded to a narrow, depressed terminal facet. Seeds with sarcotesta

ovoid-pyramidal, red at maturity, 2.0-2.5 cm long, 1-1.2 cm wide; sclerotesta ovoid to ovate, 1.5-1.8 cm

long, 0.9-1.1 cm wide.

ined: BELIZE. Belize District: 40-70m, 25-26 Aug 2008, M. Calonje & J. Meerman BZ08-120 (FTG), BZ08-122

(FTG), BZ08-124 (FTG), BZ08-125 (BRH, FTG, MO, NY, XAL), BZ08-126 (FTG), BZ08-140 (FTG), BZ08-146 (FTG); 60 m, 28 Aug 2008,

M. Calonje, J. Meerman & P. Griffith B208-154 (FTG); 70m, 27 Aug 2008, M. Calonje, J. Meerman & P. Griffith BZ08-156 (BRH).

Etymology.—The specific epithet honors Jan Meerman, who discovered the species.

Distribution and habitat Endemic to Cayo and Belize Districts in Belize, it occurs in seasonally dry

tropical evergreen broad-leaved lowland forests on steep karstic hills as defined by Meerman and Sabido

(2001), reaching elevations of up to 200 m. The arboreal vegetation is characterized by species such as Co-

mocladia guatemalensis Donn. Sm., Metopium brownei (Jacq.) Urb., Plumeria rubra f. acutifolia (Poir.) Woodson,

Bursera simaruba (L.) Sarg., Forchhammeria trifoliata Radlk. var. trifoliata , Coussapoa oligocephala Donn.Sm.,

Caesalpinia gaumeri Greenm., Erythrina standleyana Krukoff, Pseudobombax ellipticoideum A. Robyns, and

Thouinia paucidentata Radlk. Disturbed open spots are characterized by the endemic Louteridium donnell-

smithii S. Watson. These forests are semi-deciduous, with more than half of the trees being leafless for at

least one month during the dry season, and with several species being leafless for four months per year.

This semi-deciduous cl implies that the amount of light reaching the forest floor (and under-canopy

cliffs) is dramatically different between the dry and rainy seasons.

Scattered throughout these hills are vertical limestone cliffs where this species is found. It appears to

be an obligate cliff-dwelling species, as it is typically found growing in cracks and crevices on sheer vertical

walls and absent from th ling forest floor. The cliff faces are largely bare but depending on the level

of hering of the rock and the amount of shading they receive, the accompanying cliff vegetation consists

largely of herbaceous plants, hemi-epiphytes and vines such as Adiantum tenerum Sw., Agave angustifolia Haw.,

Anthurium schlechtendalii Kunth subsp. schlechtendalii, Anthurium verapazense Engl., Philodendron radiatum

Schott var. radiatum, Clusia sp., Begonia sericoneura Liebm., Pitcairnia recurvata (Scheidw.) K. Koch, Trades-

cantia spathacea Sw., Passiflora cobanensis Killip, and Passiflora xiikzodz J.M. MacDougal subsp. xiikzodz.

Climate.—The average annual precipitation within this species’ range is estimated at 2000-2500 mm,

with a distinct dry season occurring from February through May. The wettest month is July with an average

monthly range of 300-440 mm, and the driest month is April with a range of 50-70 mm. The temperature

ranges from 18°C to 31°C, with an annual mean temperature of 25°C. The coldest month is January and the

warmest month is May. (Data derived from GIS analysis using Worldclim 1.4 climate layers as described by

Hijmans et al. (2005)).

Conservation status. —The extent of for thi is estimated to be 750 sq. kmas delimited

by the eee range occupied by suitable karst hills in Belize and Cayo Unas The area of occupancy,

which incl only these karst formations, is 300 sq. km. Based on an g population density

Calonje, Zamia meermanii, a new cliff-dwelling species from Belize 27

of 30 adult plants per km?, the estimated total population size for this species is 9,000 plants (Meerman,

unpub. data). The karst hills where this species occurs are often isolated and separated by unsuitable habi-

tats such as lowland forest, lowland savannas, and agricultural areas. In addition, this species occurs in

discrete populations of only a few individuals, and does not appear to occur on all suitable habitats within

its geographic range. As a result of the isolation between karst mountains and the clumped distribution of

this species, populations are considerably fragmented. Visits to 12 different localities where this species

Occurs appear to indicate that reproduction is occurring and seedling regeneration is healthy.

The primary threat to this species appears to be fire, utilized in adjoining areas for slash and burn

agriculture. These fires have been observed spreading up surrounding karst hills and decimating native

vegetation. Another important threat to this species is mining for construction aggregates and dolomite,

with several active and planned quarries in the region. Recent fieldwork in Belize suggests that illegal wild

collection of cycad plants and seeds has occurred in the past and may present an additional threat to this

species. Hopefully, the difficult terrain and inaccessibility of this cliff dwelling plant, combined with the

fact that approximately 5096 of its area of occupancy lies within protected areas will help minimize the

risk of illegal harvesting. Specific locality information has been purposefully withheld in order to further

minimize this risk.

Due to its limited extent of occurrence and area of occupancy, the fragmented nature of its populations,

and the threats caused by fire, mining, and illegal harvesting, this species should be listed as Endangered

(EN) based on IUCN Red List criteria ab(i-iv) and 2ab(i-iv) (IUCN, 2001).

Reproductive phenology.—Elongating microsporangiate strobili have been observed in August and

September in habitat, and pollen releasing strobili have been observed in od and OVID: under

cultivation at Green Hills Botanical Collections near San Ignacio. A si

with about a third of the seeds already dehisced, was collected in September of 2006, ud o en tócally

dehisced with most seeds cleaned of sarcotesta, was collected in January 19 of 2006 by Martin Meadows

of Belize Botanic Gardens. During the course of fieldwork with this species in August of 2008 our team

found several distinct groupings of seedlings with newly-emerged eophylls, indicating that

strobili probably disintegrated two to three months earlier. Further work is required to peier A

the reproductive phenology of this species.

Ecology.—A large number of young seedlings observed during the course of fieldwork indicate that

reproduction is healthy and the pollinating agent is active, although it has not yet been observed. Seedlings

germinating on the forest floor suffer 100% mortality within a year or two, suggesting the forest floor habitat

is unsuitable for this species (J. Meerman, pers. comm.). In August of 2008 our team observed a female plant

with its peduncle attached and several seedlings growing directly underneath on the forest floor. Several

of these seedlings from this same strobilus had also germinated in a fissure on the cliff directly above the

mother plant, indicating that some unknown dispersal agent had moved individual seeds or a piece of the

strobilus to this location. Larvae of Eumaeus toxea Godart butterflies were observed feeding on emergent

leaves.

DISCUSSION

Zamia meermanii is most similar in appearance to Zamia furfuracea L.f. from southeastern Veracruz, Mexico.

Both species have obovate to oblanceloate or narrowly oblong coriaceous leaflets with persistent tomentum

on the abaxial side of leaflets, distinct veins on the adaxial side of leaflets, and a similar number of maxi-

mum leaflet pairs per leaf (ca. 25). Zamia meermanii differs from Zamia furfuracea in having usually solitary

stems as opposed to freely branching stems; holding 1-3 pendent leaves per crown, rather than 5-7 erect

or slightly arching leaves per crown; eophylls with a single pair of leaflets rather than eophylls with two

pairs of leaflets, leaves cream-colored and covered with saffron-orange tomentum just prior to maturing

as opposed to light green with predominantly white tomentum prior to maturing; coriaceous leaflets to

0.8 mm thick compared to coriaceous or extremely coriaceous leaflets to 1 mm thick; leaflet length to 28

28 1 intel ID t + f Texas 3(1)

Taste 1. D lucti | for Za ia furfi (d cliff-dwelli g Zamia species

Zamia Zamia Zamía Zamia

meermanii cremnophila sandovalii furfuracea

Microsporangia per microsporophyll 22-28 14-18 14-18 40-42

Microsporangia diameter (mm) 1-1.6 0.65-0.78 0.6 0.8-1.2

Microsporangiate strobilus 9-10 2.5-3 6.7 8-10

peduncle length (cm)

Megasporangiate strobilus length (cm) 12-20 8.5-14 9-17 10-24

Megasporophyll rows 7-15 4-8 4-7 4-15

Megasporophyll width (cm) 1.5-22 1.6-2.6 2.5-3.5 1.5-1.8

Megasporantiate strobilus 8-12 4—5 3.6-6 10-20

eduncle length (cm)

Sclerotesta length (mm) 1.5-1.8 1.5-1.7 1.8-2.2 1.1-1.5

Data derived from measurements by author, Whitelock 2002, Nelson 2006, and Schutzman et. al. 1988.

cm compared to 18 cm; protuberantly raised veins on adaxial surface to 1.5 mm wide rather than slightly

raised, narrower veins to 0.8 mm wide; and 22-28 microsporangia on abaxial side of microsporophylls as

opposed to 40 or more on Z. furfuracea

Zamia meermanii shares this unusual cliff-dwelling habit with two other Mesoamerican Zamia species:

Zamia cremnophila from Mexico and Zamia sandovalii from Honduras. All three species typically carry 1-3

pendent leaves. Zamia meermanii is easily differentiated from the two other species by leaflet features alone

(see key below).

1. Leaflets with distincly raised veins on adaxial surface and persistent tomentum on abaxial surface

Z. meermanii

1. A | | I 1 : jore QE | f. lue . 1 "sl £.

Z. cremnophila

Z. sandovalii

2. Adaxial surface of petiole with d

2. Adaxial surface of petiole tod ded without d qd groove

It has distinctly raised veins on the adaxial surface, persistent tomentum on the abaxial surface, and mar-

gins that are entire or crenulately notched in the upper third, whereas Z. sandovalii and Z. cremnophila both

have plane leaflets with no distinct raised veins on the adaxial surface, are glabrous on the adaxial surface,

and have margins that are distinctly toothed on the distal half. Zamia sandovalii median leaflets are strongly

falcate as opposed to those of Z. meermanii and Z. cremnophila, which are typically straight. Zamia meermanii

and Z. cremnophila eophylls typically possess two leaflets, whereas Z. sandovalii eophylls typically possess

four leaflets.

In addition to the vegetative characters that easily distinguish these three species, reproductive char-

acters are also useful (Table 1). Zamia meermanii microstrobili have longer peduncles and possess more and

hyll than Z. cremnophila and Z. sandovalii. Megastrobili of Z. meermanii

larger i I gi pet

can be longer and with more sporophyll rows and have longer peduncles than either Z. cremnophila or Z.

sandovalii. Seeds of Z. sandovalii are the largest of the group, and those of Z. meermanii and Z. cremnophila are

of a similar smaller size. Until a well-resolved phylogeny of Zamia is available, we have no way to determine

if the cliff-dwelling habit evolved more than once in the genus.

ACKNOWLEDGMENTS

Funding in support of the 2008 Belize expedition which resulted in the description of this species was

generously provided by the Association of Zoological Horticulture (AZH), Tim Gregory, and Montgomery

Botanical Center (MBC). Jan Meerman of Biodiversity & Environmental Resource Data Systems of Belize

introduced me to this species and participated in all fieldwork. Patrick Griffith (MBC) assisted in data and

Calonje, Zamia meermanii, a new cliff-dwelling species from Belize 29

specimen collection, and editing the manuscript. Green Hills Botanical Collections donated significant in-

kind support by providing lodging, equipment and work infrastructure in Belize. Belize Botanic Gardens

provided megasporangiate strobilus images and phenological data. Alan Meerow and Bart Schutzman pro-

vided assistance with the Latin diagnosis and comments on the manuscript. Alberto Taylor (PMA), Anders

Lindstrom of Nong Nooch Tropical Botanic Garden, Miguel Angel Pérez-Farrera (HEM), and Chad Husby

(MBC) provided comments on the manuscript. The Belize Forest Department granted the scientific research

and collecting permit (CD/60/3/08(45)) supporting research on Belizean Zamiaceae, as well as herbarium

infrastructure for preservation of voucher specimens.

REFERENCES

HUMANS, R.J., S.E. CAMERON, J.L. PARRA, P.G. JONES, AND A. Jarvis, 2005. Very high resolution interpolated climate surfaces

for global land areas. International J. Climatology 25:1965-1978.

Hitt, K.D., D.W. STEVENSON, and R. OsgorNE, 2007. The world list of cycads. Proc. 7th Int. Conf. on Cycad Biol. (CYCAD

2005), Xalapa, Mexico, January 2005. Mem. New York Bot. Gard. 97:454—483.

IUCN. 2001. IUCN Categories & Criteria, ver. 3.1. IUCN Species Survival Commission, Gland, Switzerland. Website:

<http://www.rediist.org/info/categories_ criteria2001.html>. Accessed 16 October 2008.

Meerman, J. AND W. Sagipo. 2001. Central American ecosystems: Belize. Programme for Belize, Belize City. 2

volumes.

Netson, C.H. 2006. Dos plantas del genero Zamia (Gimnosperma) nuevas de Honduras. Ceiba 46:41-44.

Norstoc K.J AND T.J. NicHoLLs. 1997. The biology of cycads. Cornell University Press, Ithaca, New York.

ScHuTZMAN, B., A.P VoviDEs, AND B. DEHGAN. 1988. Two new species of Zamia (Zamiaceae, Cycadales) from southern

Mexico. Bot. Gaz. 149:347-360.

Wuitetock, L.M. 2002. The cycads. Timber Press, Portland, Oregon.

BOOK REVIEW

Lynne CHAPMAN, NOELENE DRAGE, Di DursTON, JENNY JONES, HILLARY MERRIFIELD, AND BILLY West. 2008. Tea Roses:

Old Roses for a Warm Climate. (ISBN 9781877058677, hbk.). Rosenberg Publishing Pty Ltd, PO.

Box 6125, Dural Delivery Centre, New South Wales 2158, Australia. (Orders: www.rosenbergpub.com.

au, 61-2-9654-1502, 61-2-9654-1338 fax). $59.95, 240 pp., color photographs, glossary, appendices,

bibliography, index, 9 1/4" x 11 1/4”.

This beautiful book is presented by six ladies who love roses with a passi ] realized the lack of hi £d

tralia. The tea rose or P Scented China rose hidd China f. ies. They were BUB to Europe in about 1820 and later

to Australia. They | d, studied, and hybridized tt hth losing an g over and over again.

bia over two hundred pages covering every De of the tea rose, this book is a sioe work and just fascinating for a rose

lover. T d in detail, from the horticultural and botanical facts to bs po history. ue teas are presented

in all, each with a list of distinguishing features, cultivation background, location, The pepe is

quite fantastic as is rud polo by the AE as as ems um close- NUS T aen MS views.

and other p

Of additio note is the authors’ f di listinguisl ] ] f family ions. Tea rose

“imposters” are discussed and a “What not to grow” list from 1922 is given.

ook for the beauty of the rose and a book for the factual study of the rose, Tea Roses should strike a chord with a variety of

readers.— Elaine Bell, Volunteer, Botanical Research Institute of Texas, Fort Worth, Texas 76102-4025, U.S.A.

J. Bot. Res. Inst, Texas 3(1): 30. 2009

A NEW SPECIES OF ZAMIA (ZAMIACEAE) FROM THE MAYA MOUNTAINS OF BELIZE

Michael Calonje Jan Meerman

Montgomery Botanical Center Green Hills Butterfly Ranch

11901 Old Cutler Road and Botanical Collections

Miami Florida 33156, U.S.A Cayo District, BELIZE

michaelcamontgomerybotanical.org

Patrick Griffith and Geoffrey Hoese

Montgomery Botanical Center

11901 Old Cutler Road

Miami, Florida 33156, U.S.A

ABSTRACT

Zamia decumbens (Zamiaceae), a new species from the Maya Mountains of Belize, is described and illustrated. It has affinity with

Zamia tuerckheimii, Zamia bussellii/onan-reyesii, and Z. monticola, to which it is compared.

RESUMEN

mu acce = - q 1 (7 : A le] nac M de Belize M t finidad Zamia

tuerckheimii, Zamia bussellii/onan-reyesii, y Z. monticola, con las que se compara.

INTRODUCTION

In December of 1997, while lucting research on wild nutmeg trees, John Janovec and Amanda Neill were

led to the bottom of a large sinkhole in the Maya Mountains of Belize by naturalist guide Valentino Tzub

to look at an interesting group of plants he called “corn palms.” At the bottom of the sinkhole, they found

an old Mayan kiln and many pottery shards, and a dense population of Zamia (Janovec & Neill 2003). The

plants did not match the description of any Zamia known from Belize at the time, but shortly after their

return, Dr. Dennis Stevenson of the New York Botanical Garden informed them that the holotype of Zamia

prasina Bull, a species described in 1881 (Bull 1881), had recently been re-discovered at the Kew herbarium.

Janovec and Neill (2003) wrote that they connected these specimens to the sinkhole plants, and the name

Zamia prasina has since been applied to this plant in the horticultural trade and several publications (see

Balick 2000; Whitelock 2002).

In August, 2008, Montgomery Botanical Center (MBC) sponsored an expedition to Belize to shed some

light on this poorly-known species and other Belizean Zamiaceae. During the course of the expedition, it

became clear that the leaflets of these sinkhole-dwelling plants, which were chartaceous, gradually acumi-

nate, with a distinctly raised longitudinal fold, and margins that are entire or with a few minute teeth at

the apex, were very different from the leaflets of the Z. prasina holotype, which were coriaceous, abruptly

acuminate, strongly serrulate and lacking a longitudinal crease. We determined that Z. prasina was in fact a

prior valid name for Z. polymorpha DW. Stev., A. Moretti & Vazq. Torres (see Calonje et al., pages 43 — 49,

this volume), and that the cycads found in the sinkhole belonged to an undescribed species.

During the cycad research expedition, the authors studied four separate populations of this species in

Belize, gathering enough data to formally describe it.

SPECIES DESCRIPTION

Zamia d I Calonje, Meerman, M.P Griff. & Hoese, sp. nov. (Figs. 1-3). Tree: BELIZE. Torno District: bottom

of sinkhole, 350—400 m, 2 Sep 2008, M. Calonje, J. Meerman & P Griffith BZ08-201 (HoLorYrE: BRH; isotypes: FTG, MO, NY, XAL).

Species Z. tuerckheimii Donn.Sm. affinis sed caule d bente et pedunculo pollinis longissimo.

J. Bot. Res. Inst, Texas 3(1): 31 — 41. 2009

Journal of the Botanical Research Institute of Texas 3(1)

Fic. 1. Habit and habitat of Z. decumbens. A. Adult female plant with naturalist guide Valentino Tzub. B. View from sinkhole bottom at type locality of

Z. decumbens.

tains, Belize

Calonie etal.. Z.

J} ,

16 mm

Fic. 2. Rey j f £7 d b All pl g pl j jfi yl | lity | I lation. A. C f I g bilus, adaxial

side. B. C cti f | g bilus, abaxial side. C Rhopalotria sp., put i I ll fZ decumbens. D. Pharaxonotha SP., putative

pollinator of Z. decumbens. E Microsporophyll abaxial side. F. Mi I iate strobilus. G. M g gi ti bil | Il | ivi y stag H

M gas gi trobilus. |. Seed witt test 1.J. M I l yll f KM poro Í yll [ 1, abaxial side

34 i i Texas 3(

Stems globose to cylindrical, to 80 cm long and 6.7-11 cm wide, decumbent and occasionally branch-

ing on older plants, in habitat often rotting at base and rooting adventitiously from along the horizontal

stems. Vegetative cataphylls tan-tomentose, chartaceous, stipulate, shallowly triangular to triangular with

short, brown apex; strobilar cataphylls tan-tomentose, chartaceous, narrowly triangular with elongated

light-tan lanceolate apex. Leaves on adult plants 5-17 per crown, erect or arching, 90—175 cm long, light

green when emerging, turning dark green when mature. Petiole 30—55 cm long with abruptly swollen base

2.2-3.2 cm wide, moderately to heavily armed with prickles up to 5.3 mm long. Rachis 60-120 cm long,

unarmed or sparsely armed in the proximal third. Leaflets to 28 pairs per leaf, 2.8-4.1 cm wide, 17.5-29

cm long, spaced 2—5.5 cm apart from each other with the point of attachment to the rachis 4.5-6 mm wide,

chartaceous, lanceolate with gradually tapering apex and distinct longitudinal furrow, straight or slightly

falcate, margins entire or with a few teeth restricted to apex. Eophylls typically with a single pair of ovate

leaflets. Microsporangiate strobilus conical-cylindrical, emerging erect with peduncles progressively lean-

ing outward horizontally, at pollen release 10—16.5 cm long, 2.1—2.5 cm diam., occurring individually or

in groups of 2-6 or more, cream to brown; peduncle 12-23 cm long, 0.9-1 cm diam., green with brown

pubescence. Microsporophylls arranged in 16-18 columns, 26-36 rows; obtrullate, distal face hexagonal to

oblong-hexagonal, 3.0-3.5 mm wide, 4.0—4.5 mm tall; microsporangia spheroidal, 1-1.2 mm diam., present

on abaxial nae absent on ue surface, 12-18 per microsporophyll arranged in two separate groups

along margins. Meg bilus cylindrical, solitary, erect at maturity, 12-20 cm long, 5.7-6.4 cm

diam. , emerging cream- color tomentose and maturing to glabrous light or dark green with patches of tan

tomentum ing on inner facet and margins of megasporophylls; peduncle 7-13 cm long, 1.5-2 cm wide,

green to brown tomentose. Megasporophylls arranged in 8-12 columns, 4—16 rows; distal face hexagonal

to oblong-hexagonal, 2.25-2.9 cm wide, 1.6-1.8 cm tall, 2.5-4 mm thick, with a depressed terminal facet

9.5-12mm wide, 3.5—4.55 tall. Seeds with sarcotesta ovoid to ovoid-pyramidal, red at maturity, 1.7-2.0 cm

long, 1.1-1.45 cm wide; sclerotesta ovoid to ovate, 1.45-1.65 cm long, 1.05-1.43 cm wide.

Other vouchers examined: BELIZE. Cayo District: 170—200 m, 19 Aug 2008, M. edd J. Meerman, M.A. Perez-Farrera, B. Arevalo

BZ08-040 (FTG), BZ08-041 eg BZ08-053 (BRH). Toledo District: 100—200 m, 9,11,12 Mar 1987, Davidse & Brant 32232 (MO); bottom

of sinkhole, 700 m, 3 Oct 1999, Janovec & Neill 1185, 1186 (FTG). bottom of sinkhole, 300—320m, 1 Sep 2008, M. Calonje & P. Griffith

BZ08-180 (FTG), BZ08-189 (FTG), BZ08-194 (FTG); bottom of sinkhole, Toledo District, 700 m, 2 Sep 2008, M. Calonje, P. Griffith, J.

Meerman, & V. Tzub BZ08-222 (FTG), BZ08-224 (FTG), BZ08-225 (FTG), BZ08-227 (FTG), BZ08-231(BRH), BZ08-232 (FTG); 350-400m,

2 Sep 2008, M. Calonje, J. Meerman & P. Griffith BZ08-180 (FTG), BZ08-189 (FTG), BZ08-194 (FTG), BZ08-202 (FTG).

Etymology.—The specific epithet refers to the decumbent habit of the stems.

Distribution and habitat. —Known from several locations in the Maya Mountains of Belize in Toledo,

Cayo, and Stann Creek districts at elevations of 150-700 m. Currently considered a Belizean endemic, but

some populations occur near the border with Guatemala, and since the Maya Mountains extend into South-

eastern Petén Province, it is likely to occur here as well. These locations were within Tropical Evergreen

Broadleaf Lowland Forest variants as defined by Meerman and Sabido (2001) and over a number of geologi-

cal formations, most notably late Cretaceous limestones, but also older Triassic metamorphic bedrock to a

lesser extent (Cornec d

The authors h rved seven different f thi i f 1 throughout the Maya Moun-

tains of Belize, encompassing an area of approximately 2600 km". OR each of these populations was

extremely small and restricted to rocky mountaintops and ridges or on the bottom of sinkholes. The largest

populations found have been inside two steep-walled sinkholes in Toledo district, each approximately 50-80

m wide and 30-60 m in depth, with approximately 150 large plants of reproductive age. The mountaintop

and ridge populations observed to date have been much smaller, with a maximum of 20 plants observed,

all much smaller in size than those observed in the sinkholes. The factor limiting the distribution of this

species appears to be reduced soil moisture. In the sinkholes, the plants are found growing near the sinkhole

walls within the sinkhole overhang drip line, where there is less light and they are protected from direct

rainfall. Few other plant species are present in these areas, in low densities, with large areas of unoccupied

soil (Table 1). The mountaintop populations may benefit from the quick-draining substrate provided by

Calonie et al., Z

J ,

Tage 1. À g | wit! t locality for Za d b

Taxon Family

Justicia sp. Acanthaceae

Louteridium donnell-smithii Acanthaceae

Adiantum macrophyllum Adiantaceae

Syngonium podophyllum Araceae

Chamaedorea geonomiformis Arecaceae

visa veracruzana Aristolochiaceae

Forchhammeria trifoliata var. trifoliata Capparidaceae

lis spathacea Commelinaceae

Dracaena americana Dracaenaceae

ponen Ho URN Fuphorbiaceae

Episcia pun Gesneriaceae

Calatola d Icacinaceae

Dorstenia lindeniana Moraceae

icus Sp. oraceae

Passiflora lancetillensis Passifloraceae

Peperomia sp. Piperaceae

Piper spp. Piperaceae

Deherainia smaragdina var. smaragdina oo

Myriocarpa heterostachya Urticac

Myriocarpa longipes i

Urera Ho Urticaceae

the rocky environment. In Toledo district, other lithophytic species such as Chamaedorea adscendens and

Chamaedorea schippii were also observed growing in the same environment.

Climate—The average annual precipitation within this species’ range is estimated at 1800-2800 mm,

with a distinct dry season occurring from February through May. The wettest month is July, with an aver-

age monthly range of 205-580 mm; the driest month is April, with a range of 50-70 mm. The temperature

ranges from 16-31°C, with an annual mean temperature of 22-25?C. The coldest month is January and the

warmest month is May (data derived from GIS analysis using Worldclim 1.4 climate layers as described by

Hijmans et al. (2005).

Reproductive phenology.—The reproductive phenology of this little-studied species is not well known and

no distinct periodicity is evident based on the authors' visits in September of 2008, as well as the examina-

tion of photographic evidence archived at Montgomery Botanical Center from John Janovec and Amanda

Neill's visits to the sinkhole populations in August of 1999, and June of 2001. Immature, pollen-releasing,

and old microsporangiate strobili were observed during all three visits. Receptive megasporangiate strobili

were observed in August and September, and near-mature strobili were observed in June and September.

Ecology.—Several microsporangiate strobili at pollen-release stage observed at both sinkhole localities

harbored numerous snout weevils of an unknown Rhopalotria species, as well as clavicorn beetles of an

unknown Pharaxonotha species (Fig. 2c, d). Both genera are believed to be pollinators of Zamia and have

previously been found in mi giate strobili of Zamia furfuracea L.f. (Vovides 1991) and Z. pumila L.

(Tang 1987), as well as entering nt exiting both microsporangiate and megasporangiate strobili in many

natural Zamia populations in Panama (Alberto Taylor, pers. comm.). A preliminary examination of the

Rhopalotria insects suggests that they are either conspecific or close relatives to Rhopalotria mollis, one of the

known pollinators of Zamia furfuracea L.f. (William Tang, pers. comm.).

Ceratozamia robusta Miq. occurs throughout the range of Z. decumbens and both species were observed

growing sympatrically at one location in Northern Cayo district. Zamia decumbens was also observed grow-

ing together with Zamia variegata Warsz. at one location in southern Toledo district, with no evidence of

hybridization. These two species are unlikely to overlap much geographically, as most observed populations

of Z. decumbens occur above 300 m, which is the maximum of the altitudinal range of Z. variegata.

—

JOU OF

Notice

f Z. decumbens.

c. 3. Di

D. Eophylls

Calonie et al., Zamia d b ies from the Maya Mountains, Belize 37

J 7 7 f P 7

DISCUSSION

Zamia decumbens appears to be most closely related to Zamia tuerckheimii Donn.Sm. from Guatemala as well

as a new species described from Honduras. The taxonomic priority for this Honduran species, described

separately as Zamia bussellii Schutzman, R.S. Adams, J.L. Haynes & Whitelock and Z. onan-reyesii C. Nelson

& Sandoval is currently in dispute and will poo be aed here as Z. bussellii/onan-reyesii.

Zamia decumbens differs from both species in 1 tems to 80 cm long that often rot at the

base and re-root horizontally. Zamia tuerckheimii has erect or leaning stems to 3 m tall, and Z. bussellii/onan-

reyesii has erect or leaning stems to 2 meters tall. All three species have leaflets with a marked longitudinal

crease most prominent near the point of attachment to the rachis, but the leaflets of Z. bussellii/onan-reyesii

are characteristic enough to distinguish it vegetatively from the two other species (Fig. 4). Zamia bussellii/

onan-reyesii leaflets are papyraceous and with prominent teeth on the lower margin compared to those of

the other two species, which are chartaceous with margins entire or minutely toothed at the apex. Leaflets

of Z. decumbens, measuring 2.8—4.1 cm in width, are Mia s narrower than those of Z. tuerckheimii, which

typically measure 4—9 cm in width. Another useful veg tinction between the two species is that Z.

decumbens has moderately to heavily armed petioles wl | f Z. tuerckheimii are unarmed or sparsely

armed.

Mature microsporangiate strobili of Zamia decumbens are easily distinguished from those of the two

other species as they have long peduncles 11-23 cm in length causing them to lean outward, whereas Z.

tuerckheimii has erect to slightly leaning strobili with peduncles to 6.5 cm long and Z. bussellii/onan-reyesii

has erect to slightly leaning strobili with peduncles to 8.5 cm long. Zamia bussellii/onan-reyesii has the larg-

est microsporangiate strobili, measuring up to 27.5 cm long and 3.8 cm in diameter, whereas those of Z.

tuerckheimii measure 14-17 cm in length and 2.5 to 4 cm in diameter, and those of Z. decumbens measure

10-16.5 cm. Zamia bussellii/onan-reyesii is further differentiated from the other two species because it has

more than 40 microsporangia arranged in a single group on the abaxial surface of the microsporophyll

whereas 2. decumbens has 12-18 ón arranged in two separate groups and Z. tuerckheimii has

18-26 I gia arranged int The microsporophylls of Z. tuerckheimii are elongate

triangular, approximately 1. 8-2. 5 times as one as they are wide, whereas those of Z. decumbens and Z. bus-

sellii/onan-reyesii are roughly triangular and n a 1.3 to 1.5 times as long as they are wide.

TI bili of Z. bussellii/o the largest of the group, measuring 22.4—25.2

cm tall and 10- 11. 2 cm mi wide with sporophylls faces 4. 5-5 cm tall "m 4.3-4.6 cm wide, compared to those

of Z. tuerckheimii which are 16.3—22 cm tall and 8.3-10 cm wide with sporophyll faces 1.8-2.2 cm tall and

2.3-3.8 cm wide, and to those of Z. decumbens which are 12-20 cm tall and 5.7-6.4 cm wide with sporophyll

faces 1.6-1.8 cm tall and 2.25-2.9 cm wide. See the following dichotomous key and Table 2 for summary

of diagnostic characters. The known geographical ranges of the three species do not overlap (Fig. 5).

KEY TO Z. BUSSELLII/ONAN-REYESII, Z. DECUMBENS, Z. TUERCKHEIMII

. Leaflets papyraceous with prominent teeth on lower margin Z. bussellii/onan-reyesii

. Leaflets chartaceous with entire margins Or us toothed at the apex.

2. Stem decumbent | g 11-23 cm long Z. decumbens

2. Stem erect or leaning, ¡ gi bili peduncle to 6.5 cm long Z. tuerckheimii

One other species, Zamia monticola Chamb. appears to share some similarities with Z. decumbens in leaf

length, leaflet shape and size, and microsporangiate strobilus size. It was described by Charles J. Cham-

berlain based on a single male plant cultivated from seed reportedly collected near Naolinco Crater in the

vicinity of Xalapa, Mexico. Chamberlain (1926) contends that the single plant sprouted from a batch of

seeds thought to be Ceratozamia mexicana. The species has never been found again in the area where it was

reportedly collected, and it is thought that it is either extinct, or that Chamberlain's seeds became mixed

in his glasshouse and this species was acquired elsewhere (Hill 2004). Material collected in southern Alta

Verapaz in Guatemala is currently being identified as this species, but this Guatemalan material is poorly

understood at this time. Considering the fact that the species was described from a single cultivated indi-

vidual obtained under unusual circumstances, that its present geographic distribution remains unclear, and

that its megasporangiate strobili are unknown, a detailed comparison cannot currently be made between

this species and Z. decumbens. However, based on Chamberlain's description, this species appears to have

an erect stem 14 cm in diameter compared to the stems of Z. decumbens, which are decumbent and up to

11 cm across. In addition, Z. monticola is described as having 20-32 microsporangia per microsporophyll,

whereas Z. decumbens has 12-18.

Conservation status.—Although we observed seven different populations in the Maya Mountains add-

ing up to an extent of occurrence of 2600 km”, the populations were all small, occupying a small area, and

limited to the very specialized mountain top and sinkhole habitats. Despite visiting seven populations, only

the two sinkhole populations had numerous large plants of reproductive size (about 150 each). The rest of

the populations, located on rocky mountaintops, all had less than 15 individual plants of a much smaller

size than those observed in the sinkholes. In total, less than 350 plants were observed within a combined

area of occupancy of less than a square kilometer. Discussion with local residents throughout the range of

Z. decumbens confirms that plants of this species have been extracted commercially in the past and seeds

continue to be extracted for commercial purposes. During our fieldwork in the sinkholes, we found evi-

dence of past mechanical removal of megasporangiate strobili, and a seemingly low number of seedling and

young plants, perhaps suggesting that continued harvesting of seeds may be affecting the health of these

populations. Specific locality information has been purposefully withheld from this paper in order to further

minimize the risk of illegal harvesting.

Considering how fragmented the distribution of this species is, most likely resulting from its specific

habitat requirements, the small number of healthy, reproductively active populations observed, the small

combined area of occupancy for this species, and the evidence of past and present commercial exploitation,

Calonje etal. Z

Belize

TABLE 2. $ y of di icd

Enr 7

tuerckheimii, Z. bussellii/onan-reyesii, and Z. decumbens.

Z. tuerckheimii

Z. bussellii/onan-reyesii

Z. decumbens

Microstrobilus length

Microstrobilus width

Microstrobilar peduncle

eng

Microsporophyll shape

Microsporangia

Megastrobilus length

Megastrobilus width

Petiole armature

Leaflet length

Leaflet texture

Leaflet margins

Eophyll leaflet pairs

Stem height

Stem habit

14-17 cm

2.5-4 cm

to 6.5 cm

Elongate triangular

18-26 Arranged in

two rows

Sparse or unarmed

19-30 cm

to 14

Chartaceous

Entire or with a few

27.5 cm

38cm

8.5 cm

Broadly triangular

ed in

single row

22.4-25.2 cm

Light to moderate

15.5-36 cm

to 23

Papyraceous

rominent teeth on

minute teeth at distal end the lower margin

to3m

Erect or leaning

to2m

Erect or leaning

rotting at base and

11-16.5 cm

2.1-2.5 cm

11-23 cm

Broadly triangular

12-18 arranged

in two rows

Moderate to heavy

17.5-29 cm

to 28

Chartaceous

Entire or with a few

minute teeth at distal end

]

to 80 cm long.

Decumbent, often

rooting horizontally

Zamia bussellii/onan-reyesii measurements derived from Schutzman et al., 2008. Zamia tuerckheimii measurements derived

from Donnell Smith (1903), Standley & Steyermark (1958), Vannini 2008, and measurements taken by primary author. Zamia

decumbens measurements taken in-situ by authors. All measurements based on mature plants.

we consider this species critically endangered and therefore recommend a Red List Category of CR for this

species based on criteria B2ab (i, ii, iii, iv, v) (IUCN 2001). It ue be noted that although the criteria have

changed somewhat based on new information, ntinuation of the Red List category

currently listed for this species under the misapplied name “Zamia prisa

ACKNOWLEDGMENTS

The Association of Zoological Horticulture (AZH) funded our research and conservation work with this

species through the grant "Zamia prasina in Belize." Tim Gregory and Montgomery Botanical Center sup-

ported additional work in Belize.

John Janovec and Amanda Neill were the first to scientifically document and collect the sinkhole popu-

lations of Z. decumbens, providing the inspiration for our return to Belize on a research expedition. Green

Hills Butterfly Ranch and Botanical Collections donated significant in-kind support by providing lodging,

equipment and work infrastructure in Belize. The Belize Forest Department granted the scientific research

and collecting permit (CD/60/3/08(45)) supporting research on Belizean Zamiaceae, as well as herbarium

infrastructure for preservation of voucher specimens. Valentino Tzub and Boris Arevalo provided field as-

sistance. Willie Tang offered to identify the possible pollinators of Z. decumbens and provided the images of

them. Jay Vannini pru information on Z. tuerckheimii and comments on the manuscript. Bart Schutzman

helped in the Latin diag 1 provided comments on the manuscript. Anders Lindstromm, Chad Husby,

and Alberto Sidney Taylor provided comments on the manuscript. Nancy Korber provided assistance in

locating hard-to-find references. Dennis Stevenson provided herbarium specimen images and references.

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Fic. 5. Geographical distribution of Z. bussellii/onan-reyesii, Z. decumbens, and Z. tuerckheimii.

REFERENCES

BaLick, MJ, M.H. Nee, AND D.E. Atha. 2000. Checklist of the vascular plants of Belize, with common names and uses

Mem. New York Bot. Gard. 85:246.

Butt, W. 1881. Retail List 20.

CHAMBERLAIN, C.J. 1926. Two new species of Zamia. Bot. Gaz. 81:218-227

Cornec, J. 2003. Geology Map of Belize. Private Publication.

Donnett SMITH, J. 1903. Undescribed plants from Guatemala and other Central American republics XXIV. (Zamia

tuerckheimii). Bot. Gaz. 35:8-10.

IUCN 2001. IUCN Red List Categories and Criteria: Version 3.1. IUCN Species Survival Commission. IUCN, Gland

Switzerland and Cambridge, U.K.

HUMANS, R.J., S.E. CAMERON, J.L. PARRA, PG. JONES, AND A. Jarvis, 2005.Very high resolution interpolated climate surfaces

for global land areas V. Int. J. Climat. 25:1965- :

Hitt, K. 2004. The Cycad Pages. Zamia monticola. Sydney: Royal Botanic Gardens. http//plantnet.rbgsyd.nsw

8].

gov.au/cgi-bin/cycadpg?taxname=Zamiat+monticola [Accessed 27 December 200

Janovec, J.P. ANO A.K. Neti. 2003. Exploring the palms and cycads of the Maya Mountains of Belize: reflections on

MBC-sponsored expeditions to Belize, 1999-2001. The Montgomery News 11(1):5-6.

MEERMAN, J. AND W. Sasipo. 2001. Central American ecosystems: Belize. Programme d Belize) Belize City.

7 ZU A A L

aceae) aer

NELSON-SUTHERLAND, C.H. AND G.G. SANDOVAL-GONZALEZ. 2008. Una especie nueva de

Ceiba 49:135-136.

aadlil

]

Calonie et al., Z

J 7

la decumbens, pecies from the Maya Mountains, Belize "

SCHUTZMAN, B., R. ADAMS, J.L. HAYNES, AND L.M. WurreLock. 2008. A new endemic Zamia from Honduras (Cycadales:

Zamiaceae). Cycad Newsl. 31(2/3):22-26.

STANDLEY, P.C. AND J.A. STEYERMARK. 1958. Zamiaceae. In: Flora of Guatemala, Part |. Field Mus. Nat. Hist., Bot, Ser.

24:19-20.

TANG, W. 1987a. Insect pollination in the cycad Zamia pumila (Zamiaceae). Amer. J. Bot. 74:90-99.

VANINI, J.P. 2008. Notes on the Guatemalan cycad Zamia tuerckheimii Donn. Sm. Cycad Newsl. 30(3):4-7.

Vovipes, A.P. 1991. Insect symbionts of some Mexican cycads in their natural habitat. Biotropica 23:102-104.

WhrreLock, L.W. 2002. The cycads. Timber Press, Portland, Oregon.

BOOK REVIEW

Bonnie J. GiseL, with images by STEPHEN J. Joseph. 2008. Nature's Beloved Son: Rediscovering John Muir's

Botanical Legacy. (ISBN 978-1-59714-106-2, hbk.). Heyday Books, PO.Box 9145, Berkeley, California,

94709 U.S.A. (Orders: www.heydaybooks.com, 1-510-549-3564, 1-510-549-1889 fax). $45.00, 247

pp., 9 1/2" x 12 1/4".

A 1 1 13.21 14 1 ža mi } des fTIAl € ees - C TON, | Es J Lt. P NEA

O J 2 I

GE and yes, his Eu as s explored and found p f t ifers, all of which he considered friends

as well as expressions of divinity. The auth 1 photographs follows Muir's ventures into wilderness with

assages from the explorer's own works, drawings from hisi journals and EUM d i d i ross the United

p P Į p

States. In a way the book is a botanical Bormo of Muir—not definitive, as that would take vol , but llent sampling of his

botanical endeavors.

Tl 1 1 1 +1 f. iH 1 4l ] ce | fil $4] f, 1 in th «

p y I I I Plant Gallery

Citations," with luced pi , a herbarium legend, and a quotation from Muir on his discovery of each plant.

Di 1 1 T Jis 1 1 lod farh: : 1 i ; [1 1 : ; 1

e r d I I cC

adding contrast, and eli 1 f he label d d 1 ] 1 d t

Lo e a r Y Y

"T 1 iE 1 ben bear es 1| N34 11 3111 e 1 11 1 J CNA.

4 i=)

Joann Karges, Texas Christian University Library (retired), Box 298400, Fort Worth, Texas 76129, U.S.A.

J. Bot. Res. Inst. Texas 3(1): 42, 2009

WHAT IS ZAMIA PRASINA (ZAMIACEAE: CYCADALES)?

Michael Calonje Jan Meerman

Montgomery Botanical Center Green Hills Butterfly Ranch

11901 Old Cutler Road and Botanical Collections

Miami, Florida 33156, U.S.A. Cayo District, BELIZE

ABSTRACT

Maya Mountains: in Belize. An examination of the 1 ical ial of Zami ina, herbarium specimens, and wild Zamia popula-

tions in Pipe ] ] d t l y to o this taxon but instead is a valid prior name for Zamia

] l 1 A lectotype for Z

4L ra

I» Is E

7 : 1 1

Key Worps: Nomenclature, Priority, Belize, Zamia prasina 1 I

RESUMEN

El nombre Zamia prasina se está aplicando en horticultura y literatura sistemática a un taxó ido de las Montañas Maya de

Belize. U del material histórico de Zamia prasina, pecí de herbario, y poblaciones e de Zamia en imd nos

1] 1 1 A 1 1 1 z

X: r L

yporl i prioridad 1 1. Se designa un lectotipo [ Z 1

INTRODUCTION

Zamia prasina Bull was described in 1881 in English nurseryman William Bull's horticultural catalog.

Historically, this species has received little mention in the literature, but in recent years the name has been

applied in the horticultural industry and systematic literature to a relatively unknown taxon from the Maya

Mountains of Belize. We re-examined the historical material related to Zamia prasina as well as herbarium

specimens and wild Zamia populations in Belize in order to uncover the true identity of this species, which

is presented in the following analysis.

HISTORICAL MATERIAL AND EARLY USAGE OF THE NAME ZAMIA PRASINA

William Bull's 1881 horticultural catalog pa the ne ias for Zamia prasina:

“A hand d distinct-looking Cycad, i luced from H f g h Honduras, now Belize]. The stems in the young

plants roundish or roundish-ovate, clothed with the imbri les left by the falling of the 1 The leaves are equally pinnate, the

dark Eigen ae terete, with an indistinct PME in genu and Enea with afew small Men prickles. The leaf-blade is oblong-ovate,

pinnate, I I entire, and tar base, where they are set on by a oy

swollen articulation, denticul 1s tł he upp f f a bright grass-green colour, whence the name. 1 Y? guinea.”

Bull also provided a leaf of Zamia prasina, which was subsequently mounted as an herbarium specimen at

Kew. In a letter attached to this specimen, dated March 22, 1881, he invites William Turner Thiselton-Dyer,

at the time Assistant Director at Kew, to stop by his nursery and collect a leaf of Z. prasina. Although there

was no notation on the specimen indicating when it was collected, it was probably shortly after Thiselton-

Dyer received the invitation.

The specimen at Kew tated ible hol fZ. prasina by Dennis Stevenson of the New

York Botanical Garden on September of 1999. However, it has not been designated as a type for the species

in any of the printed matter we consulted. This specimen, presumably provided by Bull the same year as he

described the species in the horticultural catalog, undoubtedly represents the original material on which the

species was based. However, because no reference was made to this specimen in the protologue, it cannot be

considered a holotype under article 9.1 of the 2005 International Code of Botanical Nomenclature ("Vienna

Code”). We remedy this situation by here designating this specimen as the lectotype for Z. prasina.

J. Bot. Res. Inst. Texas 3(1): 43 — 49, 2009

4 tani i Texas 3(

Zamia prasina Bull, Hort. Cat. 176:20. 1881. Ter: [BRITISH] HONDURAS [BELIZE]: cultivated from Belize, William Bull s.n.,

ar 1881 (Lecrorvre, here designated: K!).

Zamia polymorpha D.W. Stev., A Moretti & Vázq.Torres, Delpinoa 37—38:3-8. 1998. Tyre: BELIZE. Caro: 22 Jan 1989, D. Stevenson et al.

1119 (uoLorvpe: NY; isotypes: BRH, FTG, MO, NY, U.

In addition to this specimen, there is an illustration of a leaf and caudex of Zamia prasina at Kew (dated

1881). A letter attached to the illustration (dated June 15, 1881) provides some background on Bull’s original

importation of Zamia prasina. The letter, addressed to “J. Smith Esq.” (presumably Kew curator John Smith),

mentions that a “Mr. Watson” (presumably Kew assistant curator William Watson) “talked him out of two

plants that he did not want to give up.” He also mentions receipt of two separate shipments of Zamia from

Belize and that each of the plants taken by Mr. Watson came from a separate shipment. The first plant,

described as having “widely separate” leaflets was the only surviving plant from one of the shipments. The

second plant, which Bull called Z. prasina, was one of the larger plants from the second shipment of which

only a few plants survived. Mr. Bull clearly did not want to let go of the two plants and was asking Mr. Smith

to return them. In return, he promised to provide “a couple” of the smaller surviving plants of Z. prasina for

the collection at Kew. The illustration appears to be traced from the original Z. prasina plant provided by

Bull before it was returned, as it is on a herbarium-specimen-sized sheet and bears the hand-written words

“returned Zamia prasina.” Another inscription on the illustration reads “cf. Z. latifolia Lodd.,” and an inscription

on the herbarium specimen reads “Z. latifolia Lodd.” These notes, in the same handwriting, were most likely

added by Thiselton-Dyer, as he included Z. prasina in his treatment of Mexican and Central American cycads

(Thiselton-Dyer 1884) as a synonym of Z. latifolia Lodd. ex A.DC (see de Candolle 1868). Thiselton-Dyer

added a question mark next to his listing of Zamia prasina indicating he was uncertain about its synonymy

with Z. latifolia. In fact, today it is not entirely clear what Z. latifolia is, as it was a horticultural name from

Loddiges’ catalog for which no type specimen or illustration was provided. Miquel (1843) considered Zamia

latifolia a synonym of Z. muricata var. obtusifolia, later bringing it to species rank (1849). In 1847, when he still

considered Z. latifolia a variety of Z. muricata, he published an illustration of a leaflet labeled Zamia latifoliae

(Linnaea 19(4): Tab. VII, fig. a. 1847), which Stevenson and Sabato (1986) selected as the neotype for Z.

muricata var. obtusifolia (Fig. 1a). Since so little is known about the true identity of what was originally called

Z. latifolia in Loddiges’s catalog, it must be considered a nomen dubium as well as its synonym, Z. muricata

var. obtusifolia. Furthermore, the leaflet illustrated in Miquel's publication is unlikely to have been derived

from a Belizean species, as none of the known species have leaflets with strongly serrated rounded leatlet

tips. The leaflet shape and serrations in Miquel's illustration match closely those of Z. furfuracea plants at

Montgomery Botanical Center grown from seed eed near Alvarado, Veracruz, Mexico (Fig. 1b).

Thiselton-Dyer clearly considered Bull's specimen to be di than Z. muricata Willd., as both species

are compared on the Zamia prasina illustration in the same handwriting as the "Z. latifolia" annotations. The

comparison indicates that Z. prasina has leaflets that are coriaceous, abruptly acuminate, and serrulate at

the apex, whereas Z. muricata has leaflets that are chartaceous, gradually acuminate, and spinulose-serrate

towards apex. Zamia muricata was originally described from Venezuela, and as currently circumscribed, is

a species occurring only in Venezuela and Colombia.

In his 1932 cycad treatment, Schuster listed Z. muricata var. obtusifolia as well as Zamia prasina as

synonyms of Z. loddigesii var. latifolia (Lodd.) J. Schust., incorrectly citing the publication date for Bull's

horticultural catalog as 1822 rather than 1881 (Schuster 1932).

MODERN USAGE OF THE NAME ZAMIA PRASINA

Since Schuster's synonymization of Zamia prasina, the species name received little use until recent years, as

apparently the original description and Bull's specimen were misplaced and not re-discovered until 1998

(Whitelock 2002). Since then, the name has been misapplied to a rare and little known taxon from isolated

localities in the Maya Mountains of Belize (see Balick et al. 2000; Whitelock 2002; Janovec & Neill 2005;

Hill 2004). This species, formally described in this issue (Calonje et al., page 31), differs from Zamia prasina

in having an epigeous stem, leaflets that are chartaceous, gradually acuminate, with a raised longitudinal

Lenta I Jin RAS 1(184N RI flat nf 7, DEF AP £ I 4 Marta]

Fic. 1. A. Leaflet of Zamia muricata var. obtusi ifolia (= Ze Ji ] ) I

near Alvarado, Veracruz, Mexico (MBC Accession # 20011305*A).

crease, and margins that are entire or with few minute teeth at the apex, rather than a hypogeous stem and

leaflets that are coriaceous, abruptly acuminate, without a raised longitudinal crease, and strongly serrulate

on the upper half (Fig. 2).

Since it is clear that the plant currently being called Z. prasina in horticult d recent literat

not match the original description, lectotype or illustration of this species at Kew, then the logical ensuing

question to ask is: What is Zamia prasina?

The only other described Zamia species currently known from Belize are Z. variegata Warsz. and Z.

polymorpha DW. Stev., A. Moretti & Vázq.Torres. While Zamia variegata has strongly serrulate leaflets as

described in the protologue for Z. aiio. ee are pola rather than coriaceous, and are variegated

with yellow flecks, a unique and horti that Bull undoubtedly would have men-

tioned in his horticultural catalog. However, e coriaceous, strongly serrulate leaflets of Zamia polymorpha

exactly match the description of Zamia prasina.

SYNONYMIZATION OF ZAMIA POLYMORPHA

Zamia pol } tl ibed ranges from the Yucatan Peninsula in Mexico to Belize (Steven-

son et al 1998). It was arene considered to be a form of Z. loddigesii Miq. with a highly variable karyo-

Vee (see Mog & Olivares 1996). Stevenson et al. (1998) noted the karyotype variability as an important

f this species in addition to its high level of pol; phism in leaf and leaflet morphology. This

variability was ascribed to phenotypic plasticity due to dii censes in sun exposure, with plants exposed to

full sun having shorter and narrower leaves with lanceolate leaflets, those growing in Mdb shade nas

longer, broader leaves with ellit ] late leaflets, and those g gini

displaying leaf morphologies that are intermediate between the two extremes.

During a cycad research expedition to Belize in August of 2008 sponsored by Montgomery Botanical

Center, an extensive survey of populations matching Stevenson et als description of Z. polymorpha with

strongly serrulate leaflets uncovered some P ao with remarkably consistent vegetative morphology,

and others that were highly variable. Tl ] 11

y consistent populations included 1 narrow-leafleted

[| | £ pl D Hi Hi fF

46 Journal of Texas 3(

laaflat af 7, a n £ kal fal 1 Mantas £ > A I £ I+ 7 nracina (helow

b UL F 7 F

Fic. 2. Same

Michael Calonje et al. BZ08-201).

plants growing in savannas closely resembling Zamia loddigesii, and also other populations growing in

tropical evergreen seasonal broad-leaved forests with larger leaves and wider leaflets. In addition to these

morphologically consistent populations, others were found to be highly variable, containing the two forms

mentioned above as well as intermediate plants.

The variability in these mixed populations appeared to be at least partially genetically determined, as

plants with wide leaflets were sometimes found growing in full sun, and some narrow-leaflet plants were

found growing in more shaded areas. Plants with narrow leaflets collected in an open savanna by the second

author have retained their morphological characteristics despite years of growing in a shaded environment

at Green Hills Botanical Collections.

A survey of one highly variable population halfway between Belmopan and Belize City uncovered in-

dividual plants closely matching Bull's holotype (Fig. 3) of Zamia prasina and the drawing of this species at

Kew (Fig. 4), as well as the holotype for Z. polymorpha (Fig. 5), indicating that Z. prasina and Z. polymorpha

are the same species, and therefore Z. prasina takes nomenclatural precedence over Z. polymorpha.

In addition, there is also some circumstantial evidence suggesting that Z. prasina is a prior valid name

for the species now considered Z. polymorpha. William Bull did not provide a locality or specific habitat

information for where in Belize Z. prasina may have been collected, but two years after its description, Morris

(1883) identified plants common on ridges and banks near Point Ycacos as belonging to this species. Morris

did not provide a description or illustration of the plants he saw, but the cycad that is common in this area,

visited by the second author, matches the description of Zamia polymorpha. Furthermore, this species is the

most common cycad throughout Belize and would very likely be the first species encountered by collectors

for William Bull.

Zamia prasina (now including Z. polymorpha) appears to belong to a species complex that includes Z.

loddigesii and other related species such as Z. paucijuga Wieland, and Z. spartea A. DC (Schutzman 1987).

Members of this a a display high levels of morphological variation (Gonzalez-Astorga et al.

2006) and may easily hybridize (Schut 1987). Although Schuster’s (1932) taxonomic work with cycads

is not highly regarded (see Stevenson & Sabato 1986; De Luca 1990, Taylor et al. 2008), his inclusion of

Z. prasina as a synonym of Z. loddigesii suggests that the name at the time may have been applied to plants

within this P ne oe

l variability observed within populations of Zamia prasina, coupled with

the high Ec RI vatisbility between p lations and the la ce a by Stevenson

“1.1 1

populations i d coriaceous,

and Sabato (1986), indicate that while it i

serrulate leaflets represent one highly poyini species, further research including genetic sampling

£ 7, . A [RP f ote r] . " IL Dal 1 DAI Cs late Michnal falania nt

X Y

Fic. 3. Same-scal

al. BZ08-086).

across a wide range of populations will be needed to better clarify the relationship between the different

forms and their placement within the iia piod d iil complex. Until species nn are PEN we

cognition of Z I ith Z. polymorpha included within it

ACKNOWLEDGMENTS

Funding in support of the 2008 Belize expedition was g ly provided by the Association of Zoological

Horticulture (AZH), Tim Gregory, and Montgomery Botanical Center (MBC). Green Hills Botanical Col-

lections donated significant in-kind support by providing lodging, equipment and work infrastructure in

Belize. Patrick Griffith, Chad Husby, Anders Lindstrom, Walter S. Judd, and Bart Schutzman provided

comments on draft versions of the manuscript. The Belize Forest Department granted the scientific research

and collecting permit (CD/60/3/08(45)) supporting research on Belizean Zamiaceae, as well as herbarium

infrastructure for preservation of voucher specimens. Dennis Stevenson and Andrew Vovides provided im-

ages of historical material of Zamia prasina. Nancy Korber of Fairchild Tropical Botanical Garden graciously

assisted me in the search of references.

48 I I af sl Das : ID KI FEES {T

F AS 1 . £7 5 n J . V. l^ f lant kad hu WR Dull f/eiaht

IG. p g I Pr y (right)

Dal ] DAI Fitu (late Mic I falanio ot nl D7n0. 72

y Li y i

REFERENCES

BaLick, M.J., M.H. Nee, AND D.E. ATH. 2000. Checklist of the vascular plants of Belize, with common names and uses.

Mem. New York Bot. Gard. 85:246. The New York Botanical Garden Press, Bronx.

Butt, W. 1881. Zamia prasina. Bull horticultural catalog. ÉS E

De Luca, P. 1990. A historical perspective on cycads fi tl t. Mem. New York Bot. Gard. 57:1—7.

DE CANDOLLE, A.P. 1868. Zamia latifolia. In: Prodromus systernati naturalis regni vegetabilis sive enumeratio con-

tracta ordinum, generum specierumque plantarum huc usque cognitarum, juxta methodi naturalis normas

digesta. Paris. P. 533.

GONZÁLES-ASTORGA, J., A.P. VOVIDES, P. OCTAVIO-AGUILAR, D. AGUIRRE-FEY, F. NICOLALDE-MOREJÓN, AND C. IGLESIAS. 2006. Genetic

diversity and structure of the cycad Zamia loddigesii Miq. (Zamiaceae): implications for evolution and con-

servation. Bot. J. Linnaean Soc. 152:533-544.

Hitt, K. 2004. The ean pages Zamia pana Sydney: Royal Botanic Gardens. http://plantnet.rbgsyd.nsw.gov.

au/cgi-bin/cycadpg Zamia+prasina [Accessed 27 December 2008].

Janovec, J.P. AND A.K. Nei. 2003. Exploring the palms and cycads of the Maya Mountains of Belize: Reflections on

MBC-sponsored Expeditions to Belize, 1999-2001. The Montgomery News 11(1):5-6.

McNett, J, ER. Barrie, H.M. Bunper, V. Demoutin, D.L. Hawksworth, K. MARHOLD, D.H. NicoLson, J., PRADO, P.C. Siva, J.E.

Skoc, J.H. WIERSEMA, AND N.J. TuRLAND, eds. 2006. International code of botanical nomenclature (Vienna Code)

AE YORE PRA TAR

Fi. 5. $ | pari f Zamia polymorpha holotype (right) | bet Imo; d Beli y (left, Michael

Calonje et. al BZ08-085). Image of Zamia polymorpha holotyt | with permission from The C.V. Starr Virtual Herbarium of the New York Botanical

adopted by the Seventeenth International Botanical Congress Vienna, Austria, July 2005. Gantner Verlag,

Ruggell, Liechtenstein.

Miquel, FAW. 1843. De cycadeis loddigesianis epistola. Tijdschrift Nat. Ges. Phys. 10(1):68-74.

Moue, FAW. 1847. Collectanea nova ad Cycadearum cognitionem. Linnaea 19:411—430.

Miquel, FA.W. 1849. Epicrisis systematis cycadearum. Tijdschr. Wis-Natuurk. Wetensch. Eerste Kl. Kon. Ned. Inst.

Wetensch. 2:280-302.

Morass, D. 1883. The colony of British Honduras, its resources and prospects; with particular reference to its

indigenous plants and economic productions. Edward Stanford, 55, Charing Cross, London.

SCHUSTER, J. 1932. Cycadaceae. In A. Engler, ed. Das Pflanzenreich 99(4):1-168.

SCHUTZMAN, B. 1987. Mesoamerican Zamias. Fairchild Trop. Gard. Bull. 42:16-19.

STEVENSON, D.W, A. Moretti, AND L. Gaupio. 1998. A new species of Zamia (Zamiaceae) from Belize and the Yucatan

Peninsula of Mexico. Delpinoa 37-38:3-8.

STEVENSON, D.W. AND S. SABATO. 1986. Typification of names in Zamia L. and Aulacophyllum Regel (Zamiaceae).

Taxon 35:134-144.

TAYLOR, A.S. J.L. HAYNES, AND G. HoLzMAN. 2008. Taxonomical, nomenclatural and biogeographical revelations in the

Zamia skinneri complex of Central America (Cycadales: Zamiaceae). Bot. J. Linnaean Soc. 158:399-429.

THIsELTON-Dyer, WT. 1884. Cycadaceae. In: Helmsley, W.B. Biologia Centrali-Americana, Botany 3(16):190-195.

Vovipes, A.P. AND M. Otivares. 1996. Karyotype polymorphism in the cycad Zamia loddigesii (Zamiaceae) of the

Yucatan Peninsula, Mexico. Bot. J. Linnaean Soc. 120:77-83.

WhrreLock, L.W. 2002. The cycads. Timber Press, Portland, Oregon.

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Botante Research institute of Texas, Fort Worth, Texas 76102-4025, U.S.A.

J. Bot. Res. Inst. Texas 3(1): 50. 2009

FESTUCA ALOHA AND F. MOLOKAIENSIS (POACEAE: LOLIINAE),

TWO NEW SPECIES FROM HAWATT

Pilar Catalán Robert J. Soreng and Paul M. Peterson

Departamento de Agricultura (Botanica) Department of Bota

T Satie Superior de el esca National Museum of Naural “a

ersidad de Zaragoz mithsonian Instit

Ctra. ree um 1, 22071 freu SPAIN Washington, DC oa USA,

pcatalangunizar.es sorengr@si.edu, peterson@si.edu

ABSTRACT

Festuca aloha, sp. nov. of Kaua'i and F. molokaiensis, sp. nov. from Molokz'i are described and illustrated. The two species are en-

demic to the Hawaiian archipelago and grow on steep mountain slopes and volcanic cliffs with other native mesic flora. A key to the

species of Festuca found in the Hawaiian Islands is given.

Key Words: Festuca, new species, Hawaii, Loliinae, key

RESUMEN

Se describen e ilustran Festuca aloha, sp. nov. de Kaua'i y F. molokaiensis, sp. nov. de Moloka'i. Las d peci démicas del

sataj Torr ti 31,1 o lad = J; a f1 A R +

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mésicos € H 1 da 3 if L da] i Jo Eu Lakit 1 icl Ap] heel

r L K e

Is E

While revising specimens of Festuca deposited at the United States National Herbarium (US) the first author

found two specimens of fine-leaved individuals from Kaua'i that were identified as F. rubra L. However,

these individuals exhibit several characters that separate them from holarctic red fescues of the F. rubra

complex, including; leaf blades flat with involute margins, sheaths open, relatively long ligules, and ovaries

with densely hairy apices. After reviewing specimens of Festuca from Molokai'i deposited at the Bishop

Museum (BISH) we noticed an additional taxon. Previously, the only species of Festuca and relatives re-

corded from Hawai'i and the Pacific include the endemic, F. hawaiiensis Hitchc. (Hitchcock 1922), a robust

broad-leaved species placed in F. subg. Drymanthele V.1. Krecz. & Bobrov sect. Banksia E.B. Alexeev (Alexeev

1980), and three Eurasian species: the fine-leaved F. rubra L. (F. subg. Festuca sect. Aulaxyper Dumort.); and

the broad-leaved F. arundinacea Schreb. and F. pratensis Huds. [F. subg. Schedonorus (P. Beauv.) Peterm. sect.

Schedonorus (P. Beauv.) Endl]. The taxonomy of Festuca s.l., the largest genus of monophyletic subtribe

Loliinae Dumort., is i tate of flux. Much additional research is needed to properly align the taxa, althoug

preliminary data from DNA sequencing (Catalán et al. 2007; Inda et al. 2008) suggest that two major clades

may exist. As elements of the "broad-leaved" clade, the latter two species have been classified within the

separate genus Schedonorus P. Beauv., as Schedonorus arundinaceus (Schreb.) Dumort and S. pratensis (Huds.)

P. Beauv. (Soreng et al. 2003; Snow 2008). In addition, Festuca subg. Drymanthele is sometimes recognized

as the genus Drymochloa Holub. The “fine-leaved” clade of fescues, including F. rubra, are placed in Festuca

s.s. (Soreng et al. 2003). However, results are still equivocal concerning the resolution of the division at the

base of the two major clades.

Specimens ined from the Bishop Museum (BISH), National Tropical Botanic Garden (PTBG),

Missouri Botanical Garden (MO), and the United States National Herbarium (US).

Festuca aloha Catalán, Soreng & PM. Peterson, sp. nov. (Figs. 1A-J, 2). Tye: U.S.A. Hawar: Kava‘, Hanalei district,

Kalalau Rim, Kalala, side bendi and W iun "i d neo. lookout, isolated hanging side-valley of iind diverse mesic forest

Peucedanum, Poa mannii, Hedyotis, 1 hia el Melicope

surrounded by F

pallida, NA dtm. Dubautia. Dodd: POSER Li ta, Lobelia niil is, Lipidium, Neti Hibiscus kokio,

Eragrostis, a Myrsine, Acacia, and Psychotris, 790 n m, 13 Mar 1992, K.R. Wood 1701 & S. Verba (HoLOTYPE: PTBG-17679;

Iso TYPE: US-32522

J. Bot. Res. Inst. Texas 3(1): 51 — 58. 2009

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Fic. 1. Illustration of Festuca aloha Catalán, Soreng & P.M. Peterson (isotype, US-3252239). A. Habit. B.

with rachilla. E. Lemma, ventral view. F. Pa

Sheath, ligule, and blade. C. Spikelet. D. Floret

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A Festuca hawaiiensis Hitchc. paniculis 8-11 cm longis, angustibus, ramis erectis lentibus; pedicellis (11.5—3.5(-5) mm longis;

lemmatibus 5.8-8 mm longis; laminis 1.5-2.4 mm longis, recedit.

Plants perennial, cespitose, with extravaginal innovations; cataphylls conspicuous, brownish. Culms

45-70 cm tall, erect, slender (ca. 1 mm diam.), nodes 23, smooth and glabrous. Leaf sheath margins fused

for 1-2 mm at base, overlapping more than !^ the length below, sparsely villose abaxially when young,

becoming glabrous at maturity, purple-brownish at the base, greenish above, becoming fibrous in age at

base; auricles absent; collars glabrous; ligules 1-1.5 mm long, scarious, glabrous, brownish, apex obtuse,

dentate-erose; leaf blades of vegetative shoots 22-33 cm x (12)1.5-1.7 mm, erect to pendant, flat with

involute margins, glabrous and smooth abaxially, hirsute along protruding ribs adaxially, the hairs up to

0.1 mm long, cross section (Fig. 2) about 1 mm bearing 13-16(-20) larger vascular traces or veins, each

with prominent ribs adaxially alternating with smaller ribs on the secondary and tertiary veins, scleren-

chyma strands forming trabeculae or girders on most veins; leaf blades of culms about 22 cm x 2.4 mm,

erect to pendant. Panicles 8-11 x 2.5—4 cm, erect, axis and peduncle smooth; loosely contracted with

34-66 spikelets, spikelets mostly at branch tips; panicle branches 2(3) per node, basal branches 4-6 cm

and ca. 1/2 the length of panicle, angled, erect or ascending, smooth proximally to scabrous, angled dis-

tally. Spikelets 8.2-13 mm x 3.5-6.0 mm, 4-6-flowered, broadly lanceolate, pale green, imes tinged

with purple; pedicels (1-21.5-3.5(-5) mm long; rachilla internodes 0.8-1.2 mm long, scabrous; glumes

lanceolate, green, margins very narrowly scarious («0.5 mm), apex acute; lower glumes (3.6-)3.9-4.2(-4.5)

mm long, 1-veined; upper glumes 5.2—5.5(-6.0) mm long, 3-veined; lemmas 5.8-8.0 mm long, 5-veined,

glabrous below, scabrous near the apex, green or tinged with purple at the apex, mucronate to awned, the

mucro or awn 0.5-1.1 mm long, scabrous; callus rounded, obliquely angled (to 45°), smooth to dorsally

minutely scabrous at the apex; paleas longer or shorter than lemma, scabrous on and between keels; stamens

3, anthers 3.1-4.2 mm long; ovary densely hairy on upper 1/3. Caryopsis not seen.

Additional material examined. U.S.A. Hawaiʻi. Kaua‘i: Hanalei district, Kalalau Rim, N of Kahuamaa flat, 990-1020 m, 3 Mar 1991,

K.R. Wood 631, M. Query, S. Montgomery (PTBG-9498); north aspect, 1060—1190 m, 5 Apr 1991, K.R. Wood 901 & M. Query (PTBG-13876);

Kalalau side below and W of the first una lookout, 13 Mar a K.R. Wood 1704 & S. Perlman (PTBG-17678, topotype?; Honopu

Rim, undisturbed cliffs, r ith old i seen with Panicum lineale, 2800 ft, 6 Nov 1993, K.R. Wood 2832

(PTBG-29690; MO- pened Kalalau valley, Gumping on N facing al cliffs, seen with Poa mannii, Panicum lineale, Eragrostis vari-

abilis, Carex Mn ca. 3000 ft, 17 Jun 1994, K.R. Wood 3245 (PTBG-20438); Pohakuao, hanging valley between Kalalau and Hanakoa,

and Kaaalahine Ridge, 2000 ft, 4 Jan 1992, K.R. Wood 1761, S. Perlman & J. Lau (BISH-621133, PTBG-12875, US-3250261);

Waimea D. with Panicum lineale pn P. FPE 2000 ft, 30 Mar 1993, K.R. Wood 2470, & S. Perlman (PTBG-18788), K.R. Wood 2471

& S. Perlman (PTBG-18779); A alley, N facing slopes above stream, 0.5 mi along trail, growing with Wilkesia gymoxiphium

and Lipidium serra, 3300—3500 ft, 18 May 1994, K.R. Wood 3193, S. Perlman & M. Hartley (PTBG-15270); Upper Kawaiiki: Kaluahaulu

Ridge PP dh W of ] i iiis into Hager Acacia n Metrosideros iss morpha mixed mesic for-

est with D 11 1 Poa ie eee S nd Cl

1149 m, 14 Nov 1996, K.R. Wood 7605 (PTBG-42661).

Comments.—Festuca aloha has panicles 8-11 cm long, narrow, branches erect and ascending; lemmas

5.8-8 mm long; and leaf blades 1.5-2.4 mm wide; in contrast F. hawaiiensis has panicles 30-40 cm long,

widely open, branches patent or patent-erect; lemmas 9 mm long; and leaf blades 2-3 mm wide.

The illustrator Alice Tangerini noticed that all florets with mature anthers of F. aloha contained

small, undeveloped ovaries (Fig. 1H). Mature ovaries with well-developed styles were found only in florets

that had already shed their anthers (filaments were still present) [Fig. 11]. Therefore, F. aloha is apparently

protandrous.

Etymology.—The specific epithet F. aloha name derives from the Kalalau mountains of Kauai.

Conservation status —Festuca aloha has been found in at least six different localities of Kauaii, covering

a distribution area of approximately 700 km?. The number of individuals varies among populations but

usually there are less than 1000 individuals per population. Based on these preliminary data, the new spe-

cies clearly falls within Vulnerable (VU) category as defined by the IUCN (2001). The major threats to in-

dividuals of F. aloha are the allochthonous plants, such as: Cyperus meyenianus Kunth, Erigeron sp., Kalan-

choe pinnata (Lam.) Pers., Lantana camara L., Melia sp., Passiflora mollissima (Kunth) L.H. Bailey, Rubus

Ec? l £hlad na £ tati L £ Fact Inha Catalán Sorena & PM DP 1110 2353320 Y ). Scale bar = 0.1 mm

Li 3 A

rosifolius Sm. ex Baker, Setaria gracilis Kunth, Triumfetta sp.; and goats and pigs (K.R. Wood, on herbarium

label).

Festuca molokaiensis Soreng, PM. Peterson & Catalán, sp. nov. (Fig. 1K). tw U.S.A. Hawari. Moroka‘: Kupaia

Gulch, on steep slopes in mesic forest, occasional, associated vegetation: Metrosideros, Coprosma, Leptecophylla, Dodonaea, Alyxia,

Viola, Melicope, Myrsine, Diospyros, Dryopteris, Phyllanthus, Luzula, Selaginella, A Carex, 21.119564 N 156.936999 W, 880 m, 3 Apr

2007, H.L. Oppenheimer H40704, S. Perlman & N. Tangalin (noLorvre: BISH-7287

li b tib i bris, arista lon-

A Festuca aloha Catalán, Soreng & P.M. Peterson, p

giori 1.5-2.8 mm, antheris brevioribus 2-3 mm, shins inferis (s. 0-)5.2- 5. sc 5.7) mm, EET superis (6.0-)6.2-6.8(-7.0) mm, oe

Plants perennial, cespitose, with extravaginal innovations; cataphylls conspicuous, brownish. Culms up

to 60-70 cm tall, scabrous below panicle. Leaf sheaths with margins fused for 1-2 mm at base, overlapping

more than Y the length below, purple-brownish at the base, greenish above, becoming fibrous at base in

age; auricles absent; collars glabrous; ligules 1.5—2.5 mm long, scarious, glabrous, brownish, apex obtuse,

dentate-erose; leaf blades of vegetative shoots approximately 34 cm x 2 mm, more or less pendant, flat

with involute margins, glabrous and smooth abaxially, hirsute along protruding ribs adaxially and on mar-

gins and apex, dull green, cross section luu to that of F. aloha, about 2 mm wide bearing 25 larger vas-

cular traces or veins, each with prominent ril ially alternating with smaller ribs on the secondary and

tertiary veins, bulliform cells between adaxial ribs, sclerenchyma strands present forming trabeculae or

girders on most veins, sclerenchyma strands with thin cell walls (adaxially) and thick cell walls (abaxially);

leaf blades of culms about 21 cm x 2 mm, flat with involute margins, densely scabrous adaxially. Panicles

8.5-13 x 4 cm, loosely contracted with approximately 75 spikelets, spikelets located at the ends of the

branches, 3(4) branches on basal node, 2(3) branches in other nodes, axis and branches strongly scabrous;

basal branches as long as 7 cm, more than 1/2 the length of the panicle, angled, suberect to ascending.

Spikelets 7-9 x 3-4 mm, 5-6-flowered, broadly lanceolate, pale green; pedicels Q— 4(-9) mm long; ra-

chilla internodes scabrous; glumes lanceolate, green, margins very narrow, scarious, apex acute; lower

glumes (5.025.2—5.5(5.7) mm long, 1-veined; upper glumes (6.0—)6.2-6.8(-7.0) mm long, 3-veined;

lemmas (5.0-)5.5—6.0 mm long, 5-veined, scabrous dorsally and near apex, green, awn 1.5-2.5 mm long,

scabrous; callus rounded, obliquely angled (to 459), scabrous; paleas slightly shorter than lemma, bidentate

at apex, keels scabrous, between keels scabrous; stamens 3, anthers 2.8-3.0 mm long; ovary densely hairy

on upper 1/3. Caryopsis not seen.

Comments.—Festuca molokaiensis differs from F. aloha by having more delicate and thinner, more nu-

merous and lax branches; scabrous panicles and spikelets; lemmas scabrous throughout, awns longer

1.5-2.8 mm; anthers shorter 2-3 mm; lower glumes (5.0—)5.2—5.5(-5.7) mm; and upper glumes (6.0—6.2—

6.87.0) mm. In comparison to F. aloha, culms are scabrous on the nodes; panicles 8.5-13 cm long, more

delicate, axis and peduncles scabrous, branches densely scabrous, angled; spikelets more delicate; rachilla

more slender; lower glumes (5.0-)5.2—5.5(-5.7) mm long; upper glumes (6.0—)6.2—6.8(-7.0) mm long; lem-

mas 5-6 mm long, evenly scabrous across the back from the base to the apex, 3-veined or infrequently

faintly 5-veined (lateral veins obsolete or obscure), awned, the awn 1.5-2.8 mm long; and anthers 2-3 mm

long.

Catalán et al ; T

Etymology.—The specific epithet of F. molokaiensis derives from the island of Moloka'i.

Conservation status.—Festuca molokaiensis is only known from the type locality where it has been re-

corded as occasional (Oppenheimer, on herbarium label). Due to the scarcity of available data on the po-

tential distribution, demography, and population status of F. molokaiensis, we consider to be a Data Deficient

(DD) taxon (IUCN 2001).

TAXONOMY AND PHYLOGENY

Festuca aloha and F. molokaiensi logically similar to the Hawaiian endemic, F. hawaiiensis in ligule

size and shape, ovary hairiness, and leal hee y (Hitchcock 1922). However, F. hawaiiensis is a more

robust plant up to 150 cm tall with longer (30-40 cm) and wider panicles that are open with 3—5 spreading

and drooping branches at the nodes, and larger lemmas up to 9 mm long. Festuca hawaiiensis grows at

higher elevations, at ca. 2000 m, in rich soil on moist wooded hills on the island of Hawaii.

Festuca aloha and F. molokaiensis resemble the western North American, F. californica ssp. hitchcockiana

(E.B. Alexeev) Darbysh. [Darbyshire & Pavlick 2007] in leaf blade cross section, panicle features, and hairy

ovary apices. However, F. californica ssp. hitchcockiana differs from the new species by having a more robust

and cespitose habit, possession of a continuous sclerenchyma layer on the abaxial side of the leaf blade, a

partially hairy collar, and longer ligules with ciliate apices.

The new Festuca taxa also show similarities with Macaronesian volcanic cliff dwellers: F. agustinii Lind-

ing., F. jubata Lowe, and F. francoi Fern.Prieto, e Aguiar, E. Días & M.I. Gut. (Saint-Yves 1922; Fernández

Prieto et al. 2008) because all sh d flat to inrolled (or conduplicate) leaf-blades

with some complete sclerenchyma cele However, the latter species all differ from the new species by

having glabrous ovary apices and short (« 1.5 mm) ligules.

We analyzed the ITS and trnL-F sequences of two samples of Festuca aloha from different populations

from Kauai [F. aloha 1: Kalalau, US-3252239 (isotype), GenBank GQ162205 (ITS) and GQ162208 (trnLF)

accessions; F. aloha 2: Pohakuao, US-3250261, GenBank GQ162206 (ITS) and GQ162209 (trnLF) accessions]

and one sample of F. molokaiensis from the only known population from Molokai [F. molokaiensis 1: Kupai

Gulch, BISH-728771 (holotype), GenBank GQ162207 (ITS) and GQ162210 (trnLF) accessions]. We then

conducted heuristic parsimony analyses for the combined ITS and trnL-F dataset (10,000 random entry

trees, TBR, mulpars off, saving no more than 10 trees of length «10 per replicate; 1000 bootstrap replicates

with the same parameters as in the original search) using these three samples and combined them with our

previous data for Festuca on a worldwide level (Catalán et al. 2007; Inda et al. 2008).

All three samples grouped together within the broad-leaved Festuca clade (Fig.3), nested within the

well supported clade Subulatae + Leucopoa p.p. [92% bootstrap(BS)], which were separated from F. califor-

nica, F. rubra, and the M ian Festuca sect. Aulaxyper s.l. species. Festuca aloha was monophyletic (9996

BS) and sister to F. molokaiensis (98% BS).

The two F. aloha specimens from dn ode on Kauai (F. aloha 1 & F. aloha 2) had similar

sequences (9996 BS; 1 trnL-F nucleotid titution difference); however, the sample of F. molokaiensis from

Molokai showed several nucleotide diff in both the ITS (five substitutions) and trnL-F (two substitu-

tions) regions with respect to F. aloha. These molecular differences also support the taxonomic separation

of the two species. Despite the limited infraspecific sampling, the ITS variation found between F. aloha and

F. molokaiensis, from the same Hawaiian archipelago, is remarkable as these species show more nucleotide

differentiation than that found between other closely related species [e.g. the Pyrenean F. eskia Ramond ex

DC., and F. gautieri (Hack.) K. Richt. with only two ITS nucleotide substitutions], and between species from

different Macaronesian archipelagos (e.g., Madeiran F. jubata and Azorean F. petraea Guthn. ex Seub. with

three ITS nucleotide substitutions). Unfortunately, F. hawaiiensis could not be included in the molecular

analysis because it is extremely rare and we have no material other than the type.

The closest relative of F. aloha and F. molokaiensis in our analyses (Fig. 3) was the eastern Asian F.

parvigluma Steud. (8896 BS), followed by the ] North American F. subulata Trin. (7796 BS), both

belonging to F. subg. Subulatae, and then by the Siberian-North American F. altaica Trin. (9296 BS), placed

Festuca luciarum

Festuca multinodis

Vulpia membranacea

Vulpia unilateralis

Cienopsis mE

Vulpia ci

Fesucrm

Festuca violac

Vulpía octo Hora

gautieri

Festuca quadriflora

Festuca rieulmis

estuca thurbert

Festuca arundinacea

Festuca atlantigena

1 Festuca leo urnexiana

Festuca gincescens

Festuca mai

Festuca fen

Festuca bačtica

foe einig

pad

Tes

Festuca aloha2

Festuca moloksiensis

s parvigrum

[| l af pl

JUuiriidi OF

Drymanthele s.s. gr.

Exaratae p.p./

American Vulpia

Leucopoa p.p. +

pee S.S. +

Aulaxyper s.l./

Vulpia 2x

Festuca/

Wangenheimia

Micropyrum

Narduroides

Loretia gr. +

Ctenopsis

Psilurus/

Vulpia (4x-6x)

Exaratae p.p.

American II

American ]

Eskia-

Dimorpha gr.

Leucopoa p.p.

Neozeylandic I./

American

Schedonorus gr.

Subulbosae

Subulatae +

Asian-American

Leucopoa s.s.

Lojaconoa

Castellia

pa nig

Cynosurinae/

Fine-leaved

Loliinae

Broad-leaved

Loliinae

conc

oly

Fic. 3. Strict t (

r. ini

fatali

, Soreng & PM. Peterson and

retention index 0.762) I

Catalán et al T iac nf Factura fi T sg T

within F. subg. Leucopoa sect. Breviaristatae. In a separate ITS strict consensus tree the F. aloha/F. molohai-

ensis clade fell within the broad-leaved clade, and was closely related to the Asian F. parvigluma and F.

modesta Steud. [sometimes treated as Drymochloa modesta (Nees ex Steud.) Holub]. In contrast to the ITS

data, a trnL-F strict consensus tree placed F. aloha/F. molokaiensis in a polytomy at the base of the fine-leaved

clade, among fine-leaved taxa and taxa intermediate between fine- and broad-leaved forms of Festuca.

Given the different resolutions between separate nuclear and plastid DNA sequence topologies for our

species, it is possible that F. aloha and F. molokaiensis are of allopolyploid origin. On morphological grounds

F. aloha, F. molohaiensis, and F. hawaiiensis are all part of the same complex, representing a common pattern

of morphological speciation in upland habitats of different islands along the Hawaiian archipelago.

KEY TO THE SPECIES OF FESTUCA IN HAWAI

1. Lower leaf blades with auricles; leaf blades usually 5-12 mm wide, flat.

2. Auricles ciliate on margin; lemmas awned (awn 2—3 mm), more or less scabrous distally Festuca arundinacea

2. Auricles not ciliate on margin, lemmas muticous, smooth or only sparsely scabrous distally — baba pratensis

1. All le af bl f blades 0.3- 5m m wide, i involute to flat but when flat

3. Ligules 0.1-0.5 mm long; c ovary apices glabrous; leaf blades conduplicate; leaf sheaths usually closed

below (open « Y their length); leaf blades of vegetative shoots in cross section without sclerenchyma

girders. estuca rubra

. Ligules 1-2.5 mm long; ovary apices densely hairy; leaf blades uu d Tel margina; m Aa

generally open Y their length [they are fused only for 1-2 mm att fi

in cross d with M dps Sues

4. Panicles 30-40 cm long, open, t | ling or drooping; leaf blades 3-5 mm wide ; known

only dd M. wall Festuca hawaiiensis

4. Panicles 3.5-25 cm long, usually narrow, the branches patent, erect or ascending; leaf blades 0.3-2.4

mm wide.

5, qui scabrous only in the upper third, i liate veins distinct, awns 0.5-1 mm long; anthers

3.1-42 mm long; ligules 1-1.5 mm long; lower glumes (3.6-)3.9—4.2(-4.5) mm long; upper glumes

eh 5.5(- is mm long; aca only from pale! i Festuca aloha

veins absent or obscure, awns 1.5-2.8 mm; ee

2-3 mm long: ligules 1.5— 25m mm long; lower glumes (5.0-)5.2—5.5(-5.7) mm long; RES gium

(6.0-)6.2-6.8(-7.0) mm long; known only from Moloka'i Festuca end:

ACKNOWLEDGMENTS

This work has been subsidized by the Spanish Ministry of Science and Technology Grant Project CGL2006-

00319/BOS and Sabbatical Research Stay Grant PR2008-0020 to Pilar Catalán. We thank Stephen Darbyshire

and Neil Snow for their critical review of an earlier version of the manuscript; Napua Harbottle, Tim Flynn,

and Gerrit Davidse, for facilitating for us the study of Hawaiian Festuca materials deposited at BISH, PTBG

and MO, respectively; Hank Oppenheimer, Neil Snow, and Cliff Morden for valuable information on the

ecology and distribution of F. aloha and F. molokaiensis in Kauai'i and Molokai'i; Sasha Savytskyy, Dai

Tsuchiya, and Juan Viruel for helping us with figure preparation; Alain Touwaide for correcting the Latin

diagnoses; and Alice R. Tangerini for providing the illustration.

REFERENCES

ALEXEEV, E.B. 1980. Novye podrody i sekzii ovsjaniz (Festuca L.) severnoj ameriki | meksiki. (Festuca L. subgenera et

sectiones novae ex America Boreali et Mexica). Novosti Sist. Vyssh. Rast. 17:42—53.[in Russian].

CATALÁN, P, P. TORRECILLA, J.A. Lopez-Ropricuez, J. MULLER, AND C.A. Stace. 2007. A systematic approach to subtribe

Loliinae (Poaceae: Pooideae) based on phylogenetic evidence. Aliso 23:380—405.

DaRBYSHIRE, S.J. AND L.E. PAvuck. 2007. 14.01 Festuca L. In: Barkworth, M.E., K.M. Capels, S. Long, L.K. Anderton, and

M.B. Piep, eds. Magnoliophyta: Commelinidae (in part): Poaceae, part 1. Flora of North America north of

Mexico, volume 24:389—444. Oxford University Press, New York.

FERNÁNDEZ PRIETO, J.A., C. AGUIAR, E. Días, AND M.I. Gutiérrez ViLLARÍAS. 2008. On the identity of Festuca jubata Lowe

L D H In l.l PA fT,

58 Journal of t| Texas 3(

(Poaceae) and the description of a new Festuca species in the Azores Islands. Bot. J. Linnean Soc.

157:493—499.

HitcHcock, A.S. 1922. The grasses of Hawaii. Mem. Bernice Pauahi Bishop Mus. 8:100-230.

INDA, L.A., J.G. SEGARRA-MORAGUES, J. MULLER, PM. Mine AND P. CATALÁN. e Pains iS biogeography of the

temperate Loliinae (Poaceae, Pooideae) grasses in t tl h Molec. Phylogen.

Evol. 46:932—957.

IUCN. 2001. IUCN Red List Categories and Criteria: Version 3.1. IUCN Species Survival Commission. IUCN, Gland,

Switzerland and Cambridge, UK.

Saint-Yves, A. 1922. Les Festuca (subgen. Eu-Festuca) de l'Afrique du Nord et de les Isles Atlantiques. Candollea

Snow, N. 2008. Notes on grasses (Poaceae) in Hawaii. Bishop Mus. Occas. Pap. 100:38-43.

Soreng, RJ, PM. Peterson, G. Davipse, E.J. Jubziewicz, FO. ZULOAGA, AND O. Morrone 2003. Catalogue of the New World

grasses (Poaceae): IV Subfamily Pooideae. Contr. U.S. Natl. Herb. 48:1—730.

A NEW GENUS, MEXOTIS, FOR FIVE MEXICAN SPECIES OF

HEDYOTIDEAE (RUBIACEAE)

Edward E. Terrell Harold Robinson

Research diy Department of Botany, MRC 166 Department of Botany, MRC 166,

National Museum of Natural History National Museum of Natural History

20. PO. Box 37012

Smithsonian Institution Smithsonian Institution

Washington, DC 20013-7012, U.S.A. Washington, DC 20013-7012, U.S.A.

terr60@ msn.com robinsoh@si.edu

ABSTRACT

In furtl icti f th t of the g Hedyotis in Mexico, four | pecies, H. galeottii, H. kingii, H. latifolia, and H. terrel-

lii, are m inanew eee Mexotis. A new species, M. lorencei, i gni d described. Morphological and distributional dat

RESUMEN

Enl icción adicional 1 pto del gé Hedyotis en México, lel peci idas, H. galeottii, H. kingii, H. latifolia

ie. M. lorencei

r i i

y H. terrellii,

y de distribución y se citan las PA representativas.

In early 20* century literature on the tribe Hedyotideae of the Rubiaceae the name Hedyotis L. was applied

to many members of the tribe in both hemispheres. The main recent proponent of the name Hedyotis was

Fosberg (1943) in his treatment of 24 Polynesian species of Hedyotis. He stated that he was adopting Hedyo-

tis in the broadest sense because “the criteria to segregate the species involve not very fundamental differ-

ences in structure" (Terrell 1996:2—5). Under Hedyotis he recognized five subgenera and commented that

many botanists would consider these as genera. Despite his broad view of genera, he favored extreme split-

ting for subdivisions of species, e.g., he described 39 M forms of Dou acuminata a & Schltdl.)

Steud.(now Kadua acuminata Cham. & Schltdl.). He als idered seed cl because

“they vary so much that no two are alike in a single capsule". In subsequent papers rockers continued to

use the name Hedyotis.

A recent study of Asian Hedyotis shows that many species related to the type, Hedyotis fruticosa L., of

Sri Lanka and India, have unique capsule and seed morphology (Terrell & Robinson 2003). These species,

in the genus Hedyotis, so far as known, are restricted to Asia and the western Pacific. Excluded from this

concept are the mid-Pacific elements more recently treated as Kadua Cham. & Schltdl. (Terrell et al. 2005)

and the New World species that have currently been placed in Hedyotis.

Many of the American Hedyotideae that were once placed in Hedyotis are now recognized as separate

genera on the basis of structural features, particularly seeds and habit. Some of the segregate genera were

described as new, including Carterella, with one Mexican species (Terrell 1987), Stenaria with five U.S. and

Mexican species (Terrell 2001a), and Stenotis with seven Baja California and one Arizona species (Terrell

2001b). In addition, Houstonia and North American species of Oldenlandia each have five Mexican or

Mexican-U.S. species (Terrell 1996; Terrell & Robinson 2006). In recent years further study has included

about 12 additional hedyotoid genera. None of these genera were found to have seeds and other characters

similar to those of the five species studied here.

We provide a further needed reclassification for Mexican Hedyotis species. A new genus, Mexotis, is

described, including Hedyotis galeottii, H. kingii, H. latifolia, H. terrellii, and a newly described species, M.

lorencei.

The removal of Western Hemisphere Hedyotideae from Hedyotis is supported by two recent studies

J. Bot. Res. Inst. Texas 3(1): 59 — 70. 2009

60 Journal of tani i Texas 3(

using DNA sequencing. In the DNA study of Andersson and Rova (1999), such species once placed in

Hedyotis are not closely related to the Asian and eastern Pacific Hedyotis, the latter represented by H. mac-

rostegia Stapf or the Central Pacific H. hillebrandii (Fosberg) Wagner & Herbst. (now Kadua axillaris (Wahra)

Wagner & Lorence). In a more complete survey, Kárehed et al. (2008) show the division more clearly, and

they call for the narrower interpretation of Hedyotis. Thus, the morphological and DNA evidence agree in

the need to exclude American species of Hedyotideae from the genus Hedyotis. The Kárehed et al. study

subdivides the Hedyotidae (as Spermacoceae) into many discrete clades that are not only distinct from

Hedyotis, but distinct from each other. Two such groups are the Arcytophyllum/Houstonia/Stenaria Clade, to

which many of the previously studied American Hedyotideae listed above belong and the Manettia/Bou-

vardia branch, the latter being close to the Spermacoce Clade. No DNA sequences are available for any of the

species placed here in Mexotis, and conclusions regarding which group is closest to Mexotis are based on

structural features.

A paper by Church (2003) contributed an important molecular study on Houstonia and related genera,

but did not include any of the five Mexican species that we investigate here.

Mexotís includes four species of large herbs or subshrubs with woody stems, long internodes, large

elliptic or ovate leaves, long petioles, filiform pedicels, and generally flattened seeds with central punctiform

hila. These species are native to Oaxaca and Veracruz. A fifth smaller species, M. latifolia, has herbaceous

or woody-based stems and smaller leaves. It appears closely related to the four more robust species and

ranges from Veracruz to Guatemala. This species is the only one of the five with a known chromosome

number, n - 17, a number not known to occur in any other species of Hedyotideae.

The five species (Table 1; Figs. 1—5) have significant reproductive and vegetative characters. The seed

types fall into two groups, the kingii group and the galeottii group. The kingii group is represented by M.

kingii, the type species, which has somewhat thicker seeds, sometimes polygonal in outline and being often

obtusely angulate or obtusely ridged. The compression is moderate or slight. These exact characters have

not appeared in other Hedyotideae species. Mexotis terrellii is also in this group because of its apparent

similarity to M. kingii; however, unfortunately the collections have immature seeds. The second seed type

group includes M. galeottii, M. latifolia, and M. lorencei. These species have thin, flat or strongly compressed,

concavo-convex seeds. In M. galeottii and M. latifolia the seeds are slightly to moderately concavo-convex.

In M. lorencei the seeds become strongly concavo-convex and may appear cupulate. In M. galeottii and M.

latifolia the seed areoles (cells) are radially elongated, and M. galeottii is unique in having narrow, usually

partial, sometimes complete, wings. These same two characters also occur in the genus Bouvardia, which,

however, carries the winged character to extremes, as many Bouvardia species have broad, conspicuous

wings (and large capsules) (Blackwell (1968).

Mexotis Terrell & H. Rob., gen. nov. Tee species: Mexotis kingii (Terrell) Terrell & H. Rob.

Plantae UNS saepe Luis b i a herbaceae vel suffrutescentes. Caules ad 1.5 m alti; laminae foliorum ad 11 cm longae

vel ellipticae; planatae aut leniter compressae interdum alatae, hilis centralis punctiformibus,

ovatae

He d PA Dal 1 1: 1

Perennial herbs or subshrubs typically with thick, woody, erect to decumbent, stems ca. 0.4-1.5 m long, in

M. latifolia stems 15—55 cm tall, woody only at base. Leaves opposite, blades ovate to elliptic, acute to

slightly acuminate, glabrous to sparsely or minutely puberulent or pubescent. Stipules interpetiolate. Inflo-

rescences with slender or filiform pedicels, branching alternate or partially opposite. Calyx lobes lancolate.

Corollas white or purplish to rarely rose, 3-13 mm long, funnelform, valvate, 4-lobed, not appendaged,

glabrous to densely pubescent inside; heterostylous, with pin and thrum forms. Capsules subglobose to

broader than long, where stages are first loculicidally then septicidally dehiscent. Seeds numerous, concavo-

convex or obtusely ridged, one species with narrow wing, hilum central, punctiform, areoles isodiametric

or radially elongated.

The genus name, Mexotis, is derived from Mex- as in Mexico and —otis as in Hedyotis, referring to an

, Mexotis, g f Rubi 61

KEY TO SPECIES OF MEXOTIS

1. Plants decumbent or erect, slender and only slightly woody at base, height 15-55 cm; leaf blades 4-32(-45)

x 3-24 mm; calyx lobes 0.4-1.5 mm long 2. M. latifolia

1. Plants erect, with woody stems and stout woody bases, height 45-150 cm; leaf blades 25-110 x 8-40 mm;

calyx lobes 1-6 mm lon

2. Stems and infl i ly hirtell g ls oft bvi ly wil iged 1. M. galeottii

2. Stems and infl lab t winged.

3. Stipules to 10 mm long; calyx lobes 5s 6 mm long; capsules 5.0-5.5 x 6-7 mm 5. M. lorencei

3. Stipules 2-4 mm long; calyx lobes 1-4.7 mm long; capsules 1-4.5 mm in diam.

4. Inflorescences 3-15 x 6-18 cm; pedicels 7-32 long; calyx lobes 1-2 mm long; corollas 5-6.5 mm

long; stipule appendages 1-2 mm long 4. M. terrellii

4. Inflorescences to 8 x 11 cm; pedicels 3-13 mm long; calyx lobes 1.5-4.7 mm long; corollas 7-13 mm

long; stipule appendages only 0.3-0.5 mm long 3. M. kingii

1. a galcote s ban. Terrell & H. Rob., comb. nov. (Fig. 1A—D). Declieuxia galeottii M. Martens, in

d. Roy: Sci. Brux. 11:231. 1844. Hedyotis Minas (M. Martens) Terrell & Lorence, Phytologia 66:1—4.

1989. Tyre: MEXICO. Oaxaca: Llano Verde, 3-7000 ft, Apr 1840, H. Galeotti 2603 BR!; isorvres: BR!, US!)

Manettia liebmannii Standley, J. Washington Acad. Sci. 17:337—338. 1927. Tere: MEXICO. Oaxaca: Pelado, Aug 1842, Liebmann 11485

(HOLOTYPE: C; ISOTYPE fragment: US 13157721). Paratype: MEXICO. Oaxaca: Cuesta de San Juan del Estado, Liebmann 11487 (C; frag-

ment US 1315771!)

Perennial herbs or small shrubs. Stems 60—150 cm tall, to ca. 2.5 mm thick and woody toward base, stout,

subterete, erect, glabrous or minutely hirtellous on upper stem and in inflorescence, few-several branched.

Leaves with slender petioles 2-7 mm long, blades 40-85 x12—38 mm, ovate or ovate-lanceolate, acuminate

glabrous above, glabrous or hirtellous on midrib beneath. Stipules 1-3 mm, broadly deltate, cuspidate with

tip 1-2 mm long, glabrous or with few thick marginal teeth. Inflorescence with terminal, few-flowered

cymes, branches densely and minutely hirtellous, flowers heterostylous, pedicels 2-7 mm long, very slender

or filiform. Hypanthium glabrous; calyx lobes 1-2 x 0,3-0.7 mm, lanceolate, acute or submucronate, gla-

brous. Corollas 4.5-9.7 mm long, funnelform or broadly funnelform, white, glabrous externally; tubes

2.8-4 mm long, 1-2.5 mm wide at base, 2.2-4.5 mm wide at throat, densely puberulent within; lobes

2.2—5.2 x1-2.5 mm, ovate, densely downy on adaxial face with white gland-tipped hairs. Pin flowers with

stigma lobes 0.6-1.5 mm long, oblong or linear, exserted 1-2 mm beyond throat, anthers located at 3/4-point

from base of corolla tube. Thrum flowers with anthers 1.0—1.5 mm long, linear or narrowly oblong, sessile

or on filaments 0.5 mm long, attached at sinuses, stigmas located at midpoint of corolla tube. Capsules

2.2—4.0 x 2.2-4.5 mm, subglobose or somewhat compressed, 3/4—4/5 inferior, rather thick-walled, glabrous,

loculicidally dehiscent. Seeds 0.8—1.2 x 0.8—1.2 mm, black, strongly compressed, very thin, flat, slightly to

moderately concavo-convex, in outline orbicular or suborbicular, margins thin, wings present or absent,

complete or partial, very narrow, 0.1-0.2 mm wide, fragile, hilum punctiform, testa finely reticulate, areoles

radially elongated (Fig. 1). The pollen was found by Joan Nowicke (Terrell et al. 1986) to be similar to that

of Hedyotis kingii. Chromosome number unknown.

Distribution and Habitats.—Forests and rocky slopes. México: Oaxaca. Llano Verde area, Sierra de

Juarez, north of the city of Oaxaca; Veracruz, in Sierra Madre Oriental northeast and northwest of Xalapa.

Notes.—Additional nomenclatural and morphological notes were included in the Terrell and Lorence

(1989) description of Hedyotis galeottii. The minute pubescence on the stems and inflorescence branches

a the species.

Addi i Labelled Hedyotis liebmannii ( l bove) and cited ! don data from D. H. Lorence. MEXICO

Veracruz: Mpio. Alto Lucero, El Cerro la Cima, entre Plan de las Hayas y Tierra Blanca, 1600m, Castillo C. & Narave 2158 (XAL); Mpio.

Atzalan, Cerro del Aguila, 850 m, 10 Sep 1982, Ventura A. 19740 (ENCB); Mpio. Atzalan, Alseseca, 950 m, 24 Mar 1975, Ventura A. 11118

(ENCB); Mpio. ii La Calavera, 1000m, 27 Apr 1978, Ventura A. 15232 (ENCB); Mpio. Naolinco, Naranjillo, 1250 m, 13 Nov 1976,

Ventura A. 13605 (ENCB); Mpio. Yecuatla, Santa Rita, 1450 m, 12 pu 1973, Ventura A. 7824 (ENCB); Mpio. Yecuatla, Loma Santa Rita,

1480 m, 12 Jan hn Ventura A. 4764 (ENCB). Oaxaca: Yolox R de, al oesta de Cuasimulca, Miranda 1094 (MEXU).

2. Mexotis latifolia (M. Martens & Galeotti) Terrell & H. Rob., comb. nov. (Fig. 1E, F). Oldenlandia latifolia M.

£ 41 D A ID Li PS £T

62 Journal of Texas 3(

£-4- Ot hue

latifolia. Purpus 2951 ( NY), Veracruz. A, B, E. Seeds, ventral views, flat, gs. C

D, F. Areoles enlarged.

Fic. 1. Seeds examined by SEM. A-D. Mexotis galeottii. A. Castillo C. & M 2158 (XAL) V B-D. Ventura A. 11118 (ENCB), Veracruz. E, F. Mexotis

E A Il he I Init I tad ch .

owina hilum

T Hu JI DAR: , Mexotis, g £ DAR: 63

Martens & Galeotti, Bull. Acad. Roy. Sci. Brux. 11:235. 1844. Hedyotis latifolia (M. Martens & Galeotti) Walp., Rep. 6:55. 1846.

Type: MEXICO. Veracruz: Mirador, at 3000 ft, near Jalapa, May 1840, Galeotti 2556 (holotype: BR; isotypes: F! GH! K!).

tune:

Hedyotis dichotoma Sessé & Moc., Fl. Mexic. ed.2. 20.1893, non H. dichotoma Cav., Icon. 6:53. 1801. Type: Sessé, Mociño, Castillo,

<= Mala 2 aang y

Houstonia gracilis Brandegee, Zoe 5:238. 1907, non Hedyotis gracilis DC., Prodr. 4:419. 1830. Type: MÉXICO. Veracruz: on rocks near

Orizaba, May 1905, C.A. Purpus 1251 (lectotype: UC! isolectotypes: F! MON.

Hedyotis exigula WH. Lewis, Rhodora 63:221. 1961. New name.

Perennial herbs with roots sometimes woody. Stems 15—55 cm tall, occasionally rather woody at base, tet-

ragonal or rounded-tetragonal, slender, erect, decumbent or spreading, glabrous or rarely pubescent,

sparsely to much branched. Leaves with petioles to ca.10 mm long, blades 4—32(-45) x 3-14(-24) mm,

median blades ovate to broadly elliptic or ovate-lanceolate, upper blades becoming lanceolate or linear-

lanceolate, glabrous or puberulent above, glabrous beneath, base rounded to cuneate, apex obtuse or

rather acute. Stipules to 2 mm long and wide, ovate or deltate, glabrous or short-pubescent, margin with

several glandular teeth to ca. 1.5 mm long, apices truncate to lacerate. Inflorescence with terminal, few-

flowered cymes, flowers heterostylous, closed at night, pedicels to 5(-10) mm long, very slender or filiform.

Hypanthium glabrous or puberulent; calyx lobes 0.4—1.5 x 0.2-0.7 mm, scarcely to somewhat longer than

capsules, erect, deltate to lanceolate, apices acute or obtuse, margins and sinuses glabrous or with white

hairs to ca. 0.1 mm long. Corollas 3-7.3 mm long, funnelform or broadly funnelform, white, purplish, or

rarely rose, glabrous externally; tube (222.3-3.3(-4) mm long, usually 1-2 times longer than lobes, some-

what widened distally, 0.6-1.4 mm wide at base, 1.8-3 mm wide at throat, glabrous or puberulent within;

lobes 13.3 x 0.6-2 mm, ovate, glabrous or puberulent within. Pin flowers with stigma lobes 0.4-1.3 mm

long, linear, exserted 0.5—2 mm beyond corolla throat, style 2.4-4.3 mm long, filiform, anthers 0.5-0.8 mm

long, 0.2 mm wide, narrowly oblong or elliptic, yellow or whitish, included near midpoint of corolla tube,

sessile or subsessile. Thrum flowers with anthers 0.7—1.2 x 0.2-0.3 mm, narrowly elliptic or narrowly oblong,

yellow or whitish, exserted at throat on filaments 0.2-0.8 mm long, stigma lobes 0.4-0.6 mm long, in-

cluded near midpoint of corolla tube on style 1-1.6 mm long, filiform. Capsules 1-2.7 x 1-3.5 mm, usually

slightly wider than long, or subequal, usually 3/4 to 9/10 inferior, moderately compressed, thin-walled,

glabrous or with few hairs on calyx margins, cuneate or rounded toward base, apex retuse, dehiscing

widely loculicidally and then septicidally. Seeds 8-24 per capsule, 0.5-1 x 0.4-0.9 mm, black, strongly

compressed, very thin, slightly to moderately concavo-convex, in outline broadly elliptic to suborbicular,

margins thin or slightly rounded, hilum punctiform, testa finely reticulate, areoles radially elongated (Fig.

1). Chromosome number: n = 17 (Lewis in Terrell et al. 1986, as Houstonia gracilis).

Phenology.—Flowering throughout year.

Distribution and Habitats.—Moist soil, shaded places, banks, slopes, among rocks, sides and bases of

cliffs, montane rain forest; altitudes recorded for a few collections vary from 330-1430 m (1000-4300 ft).

Mexico: Veracruz (mainly from Cofre de Perote and Jalapa south to Orizaba), northern Oaxaca, Chiapas,

and west and central Guatemala.

Notes.—Stephen Koch reported (pers. comm. 1980) that this species was for sale in Coyoacan, Mexico

City, as a house plant. It grows rapidly and continually produces numerous small flowers.

Standley (1918) listed Oldenlandia latifolia as a synonym of O. microtheca DC., but the type of O.

latifolia (see above) has flat seeds and reticulate testas, as opposed to O. microtheca (Schltdl. & Cham.) DC.

which has angular seeds with the areole walls coalescent. The basionym O. latifolia thus supplants the

previously accepted names, Hedyotis exigula and Houstonia gracilis.

A paper on Oldenlandia (Terrell & Robinson 2006) provided a key to Oldenlandia microtheca and

Hedyotis latifolia (Mexotis latifolia), two superficially similar species differing by seed characters and chro-

mosome numbers (the former n = 11. latter n = 17).

Mexiotis latifolia is a variable species. Its relationships to the other four species in Mexiotis remain to

be determined. It resembles Houstonia purpurea L. of eastern U.S. in aspect and leaf shape and size, but

differs in seeds and reproductive characters.

Das A ID h l titt. ET

1 I £ al 5f

64 Journal of Texas 3(

Representative specimens: MEXICO. Chiapas: Lagos de Montebello, 42 km NE of La Triniteria, 23 Oct 1971, Breedlove & Thorne 21112

(DS, MICH, MO). Veracruz: mpio. Ixtaczoquitlan, Parque de Cerveceria Moctezuma, 1 km SE of Fortin, 970 m, 9 Nov 1978, Koch &

Fryxell 78185 (CHAPA, ENCB, XAL); Cerro de Chicahuaxtla-Cuauhtlapan, 22 Jan 1968, Marino Rosas R. 1034 (CAS, MO); Mt. Orizaba,

Seaton 101 (F, GH, K, NY, US); near Rio Blanco and Orizaba, 4300 ft, 15 Sep 1944, Sharp 44889 (GH, MEXU, TENN, US); side road

between Orizaba and Fortin, 3200 ft, 23 May 1973 Terrell & King 4450, (US); Tepejilotla, mpio. Chocaman, 29 Jun 1979, Ventura A. 16282

(ARIZ, CHAPA, ENCB, MO). GUATEMALA. Huehuetenango: 5 mi SE of Barillas, Sierra de los Cuchumatanes, 1150 m, Steyermark

49537 (GH)

3. Mexotis kingii (Terrell) Terrell & H. Rob., comb. nov. (Figs. 2, 3). Houstonia kingii Terrell, Brittonia 32:491. 1980.

Hedyotis kingii (Terrell) G.L. Nesom, Syst. Bot. 13:434. 1988. Tre: MEXICO. Oaxaca: E-facing slopes, mountains along rt. 175, 28

km by road S of Valle Nacional on way to Ixtlan de Juarez; rain forest abounding in ferns, bromeliads, orchids, and aroids; elev. ca.

2000 m; not common; full sun; gravelly soil, 30 Jul 1959, R.M. King 2107 (HoLorvre: MICHI, 2 sheets, one flowering, one fruiting).

Perennial herbs with thick, woody roots. Stems to 45 cm tall, to ca. 5 mm thick and somewhat woody toward

base, tan or straw-colored, subterete, rather stout, erect, glabrous, branched. Leaves with petioles to 10 mm

long, blades 40-85 x 8-30 mm, elliptic, ovate, or lanceolate-ovate, acuminate, glabrous, slightly paler be-

neath. Stipules to 2 mm long, broadly rounded, lobed, the central lobe with recurved glandular hairs. In-

florescence in terminal, few-flowered cymes, widely branching, to 8 x 11 cm, flowers heterostylous, pedicels

to 13 mm long, very slender or filiform. Hypanthium glabrous; calyx lobes 1.5-4.7 x 0.3-0.8 mm, 1/2-3/4

as long as corolla tube, linear or narrowly lanceolate, acute or acicular, sometimes divaricately spreading.

Corollas 7-13 mm long, broadly funnelform, white or light purple, glabrous externally; tube 3-9 mm, 3-7

mm wide at throat, glabrous or puberulent within; lobes 2.5-7.0 x 1.5-3.3 mm wide, shorter than to

slightly longer than the tube, ovate, glabrous or puberulent within. Pin flowers with stigma lobes 0.5—1.3

mm long, narrowly oblong or linear, near corolla throat or slightly exserted, anthers located at 1/3-1/4-point

from base of corolla tube. Thrum flowers with anthers 1.3-2.0 mm long, narrowly oblong, purple, attached

at corolla throat, subsessile or filaments to 1.5 mm long, stigmas located at 1/4—point from base to midpoint

of corolla tube. Capsules (only previous year's dehisced capsules seen) 3-4 x 3-4 mm, subglobose, thin-

walled, glabrous, 3/4—7/8 inferior. Seeds 0.6—1.1 x 0.5-0.9 mm, dark brown, moderately to slightly com-

pressed, often polygonal in outline, obtusely angulate or obtusely ridged, margins rounded, hilum puncti-

form, testa finely reticulate, areoles small, isodiametric. Pollen illustrated in Terrell et al. 1986, figs 21, 22.

Plant and seeds illustrated here in Figures 2, 3. Chromosome number unknown.

Phenology.—Flowering December to July.

Distribution.—México: Oaxaca. Original collection in a cloud forest of the Sierra Juarez range along or

near highway 175 from Ixtlán de Juarez to Valle Nacional, 28 km south of Valle Nacional. Additional occur-

rences in Oaxaca cited below are mostly along hwy. 175 between Tuxtepec and Oaxaca city; the location of

Tarabunde is unknown. On 22 July 1987 Stephen Koch and Terrell, somewhat hampered by a torrential

rain, searched unsuccessfully for the species along Highway 175.

Additional specimens examined. MEXICO. Oaxaca: Distr. de Ixtlan, roadside, trop. v itl ferns, 24 mi S of Valle Nacio-

nal, Hwy. 175, 22 Mar 1978, Poole et al. 1283 (LL); bosque mesofilo perturbado, 1750 m, 38 km S of Valle Nacional, 11 Apr 1976, Rze-

dowshi 34092 (ENCB); km 128 entre Llano de Las lores y Tuxtepec, 26 Dec 1965, Delgadillo 121 (MEXU), reported by Lorence; Mpio. de

Comaltepec, Hwy. 175, “Vista hermosa,” selva secundaria, km 137, 1500 m, 11 May 1966, Martinez Calderon 801 (ENCB, MEXU, XAL);

Tarabunde, 7 Feb 1966, MacDougal 17 (US); 22-26 mi S from Valle Nacional on Hwy. 175, cloud forest, 7 Jan 1989, Todzia et al. 2839

(TEX)

4. Mexotis terrellii (Lorence) Terrell & H. Rob., comb. nov. Hedyotis terrellii Lorence, Novon 7:51.1997. Tyre: MÉXICO.

Oaxaca: Distrito de Mixe, Municipio de Totontepec, 3 km al SO de Totontepec, carretera a Mitla, 1910 m, bosque mesófilo, 6 Sep

1986, R. Torres C. & C. Martinez 9067 (HoLorvre: PTBG 12314; isotype: MEXU)

The following description is modified from that of David Lorence (1997). Perennial herbs. Stems 45-80 cm

tall, branching from the base with several erect stems 2.0-3.5 mm diam. and somewhat woody toward base,

terete, pee adi with petioles 3. on mm long, blades (25—)45-100 x 10-40 mm, elliptic to ovate-

elliptic, the b decurrent, te, glabrous, chartaceous, margins sometimes with few small

teeth. Stipule en to 4 mm ine: MR or deltate, with apical and lateral linear gland-tipped teeth 1-3

, Mexotis, g f Rubi 65

Fe? € de afhalat £ M. n ae A Joy CEA) A R Seeds polygonal, flat, IE ele PIPER] I CD A E el I l,

not radially elongated.

mm long. Inflorescences terminal, compound cymes 3-15 x 6-18 cm, subtended by reduced leaves, branch-

ing dichasially to 2° or 3°, the primary branches 2-5 cm long, the axes slender, glabrous, subtended by

stipule-like bracts, flowers on filiform pedicels 7-32 mm long. Hypanthium broadly obconic, glabrous.

Calyx lobes 1-2 x 1-2 mm, deltate, glabrous. Corolla in bud quadrangular, rounded at apex, at anthesis

5-6.5 mm long; tube 2-2.5 mm long glabrous, lobes 3-4 mm long, erect or slightly spreading, elliptic, acute

at apex, externally glabrous, internally villosulous with crinkled, white trichomes. Stamens attached near

middle of tube below sinuses, the filaments 0.5 mm long, anthers 1 mm long, linear-elliptic, style 2-2.5

mm. Stigma lobes 1 mm long, linear. Capsules 2-3 x 3-4 mm, slightly wider than long, 1/2 inferior, thin-

walled. Seeds immature. Plant illustrated in Lorence 1997.

Phenology.—Collected in flower in March, and in flower and fruit in June and September.

Distribution. —Known only from the area around the type locality in cloud forest at ca. 1900 m. near

Totontepec in the mountainous Mixe District of Oaxaca.

Notes.—Lorence (1997) compared certain characters of M. terrellii with those of M. galeottii and M. kingii.

Mexotis terrellii mede resembles M. hingii, as summarized in Table 1

d. P MEXICO. Oaxaca: Distrito Ixtlán, Llano Verde, collector unknown, Herb. Reichenbach fils

III-396(W); Dio Mixe, mpio. ded COE Totontepec, bosque mesófilo de montaña, 1900 m, 17?15'N, 9602"W, 11 Mar 1990, Rivera R. &

66 Journal of tl

fo il d y Pz

alt ce 9 72

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AA AS y 1

ae Y 7 KET

A "dA iN » 4 A /À E Y

ey AQ PLA a A

" h á 4 TÁ j 44 2 y x »

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X AND

X y»: ye

PV Any

u Ny A

J f ay

equine Ce ee ae 2.

Fic. 3. Mexotis kingii, holotype. A. Habit. B. Flower. C. Corolla with st style, and stigma D Ovary sectioned, ith calyx lol i sti E. In-

£1 agt L L L £. + > An^ 19

F. Mature capsules. D i late Regina O. H

3 i

DE, 2 2

Terrell and Robinson, Mexotis, a new genus of Rubiaceae 67

gag on : : Jhy CEM A D I £. R Ventral f ith ina-like margin. €

3 3

Fic. typ y

Areoles, dorsal surface, isodiametric. D. Areoles, ventral surface, somewhat radially elongated.

Martin 1403 (PTBG, UC); Distrito Mixe, mpio-Totontopec, 2 km al SO de Totontepec, carretera a Oaxaca, 1900 m, 17 Jun. 1986, Torres C & Téllez

8643 (MEXU, PTBG).

5. Mexotis jorence! Tere & H. Rob., sp. nov. (Figs. ^ 3» Terr: XEM ea id de pino-encino perturbado,

Distr. S Mpio. San Martin Peras, 1 | del t 24 carretera San Sebastián

oia Lat. 17.17 N. Long. 98.09 W., Alt. 2655 m, 29 Nov 1994, IL Calzada 19568 ee MEXU; isorvre: PTBG).

Ad M. tida bi ot M t sio: 11 di i e ABS >] IH s il dicti t

This description is based only on one E branch in ae condition. Perennial herb to 80 cm tall. Stems

8 mm or more thick, widely branched. Leaves with slender petioles 10-25 mm long, blades 70-110 x 13-30

mm, elliptic or ovate-lanceolate, acuminate. Stipule body to ca. 5 mm long, with several linear apical and

lateral gland-tipped teeth to 10 mm long. Inflorescence in fruiting condition only, pedicels to ca. 15 mm

long, slender. Calyx lobes ca. 5-6 mm long, linear or narrowly lanceolate. Corollas lacking. Capsules mature,

5-5.5 x 6-7 mm, distinctly wider than long. Seeds (only a few available) 1.4-1.8 x 1.4—1.8 mm, dark brown,

orbicular or broadly elliptic in outline, thin, strongly (cupulate) to moderately concave-convex, hilum punc-

tiform, testa reticulate, areoles not clearly visible. Fig. 5.

Distribution and habitats —The collection label stated “Negro con calizas,” which we suppose referred

to black soil with limestone or limey soil. The locality was in a disturbed pine-oak forest on the west side

of Oaxaca not far from the Guerrero border. This area is roughly 100 miles or 160 km from the other species

and we have not seen any related collections from this part of Oaxaca.

5 Mp! is florence; lesdl[ » MBA,

Hs ¡Upa

Edvard Terex de 2009

Mesas

Ss de Kn rd correo MNA, NY NEN eee

Vany UR

Fic. 5. Mexotis lorencei, J.I. Calzada 19568, isotype (PTBG)

Sp.

w David is Lene

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‘rita

T, T i Dah: , Mexotis, £Duhk: 69

Taste 1. Morphological ct

M. galeottii M. latifolia M. kingii M. terrellii M. lorencei

Stems height cm 60-150 15-55 to 45 45-80 to 80

Leaf blades L xW mm | 40-85 x 12-38 4-32(-45) x 3-24 40-85 x 8-30 25-100x10-40 . 70-110x 13-30

Inflor. L x W cm 3.5 x 2.5-5 no data to8x 11 3-15 x 6-18 no data

pedicel Lmm 2-7 to 5(-10) 3-13 7-32 to 15

vesture hirtellous glabrous glabrous glabrous glabrous

Calyx lobes L mm 1-2 0.4-1.5 1.5-4.7 1-2 5-6

Corollas L mm 4.5-9.7 3-7.3 7-13 5-6.5 no data

tubes L 2.8-4 2-4 3-9 2-2.

inner vesture villous glab/puberulent ^ glab/puberulent glabrous

lobes L 22-52 1-3.3 2.5-7.0 -4

Capsules L x W mm 2.02-40 X 22-45 | 1-27 x 1-3.5 3-4 x 3-4 2-3x3-4 5-55 x 6-7

Seeds L x W mm 0.8-1.2 x 0.8-1.2 0.5-1.0 x 0.4-0.9 0.6-1.1x0.5-0.9 immature; no data 1.4-1.8 x 1.4-1.8

Notes.—We name this species for David Lorence, who called our attention to the Calzada collection

and who has contributed much to knowledge of Rubiaceae in Mexico.

This species resembles the other three species with stout woody stems and large leaves. Although only

one collection is known, there are capsule, seed and stipule characters for consideration. The capsules and

seeds are distinctly larger than in other species. The seeds have a thin, strongly (cupulate) to moderately

concavo-convex body. Other species in this genus have flat or slightly concavo-convex seeds.

ACKNOWLEDGMENTS

David Lorence provided loans and important data on Mexotis galeottii, M. terrellii, and M. lorencei. In earlier

times Stephen Koch provided very helpful field assistance. We thank Scott Whittaker, Electron Microscope

Laboratory, Smithsonian Institution, for the SEM illustrations, and Marjorie Knowles and Alice Tangerini

for preparing the figures. David Lorence and Warren Wagner are thanked for providing in-depth reviews

of the manuscript. The curators of the cited herbaria helpfully provided Terrell with loans during a number

of years long past.

REFERENCES

ANDERSSON, L. AND J.H.E. Rova. 1999. The rps16 intron and the phylogeny of the Rubioidae (Rubiaceae). Pl. Syst. Evol.

214:161-186.

BLackweLL, W.M. 1968. Revision of Bouvardia (Rubiaceae). Ann. Missouri Bot. Gard. 55:1—30.

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. Phylogenet. Evol. 27:223-38.

FosBERG, F.R. 1943. The Polynesian species of Hedyotis (Rubiaceae). Bernice P Bishop Mus. Bull. 174:1-102.

KAREHED, J., |. GROENINCLX, S. Dessein, TJ. MoTLEY, AND B. Bremer. 2008. The phylogenetic utility of chloroplast and nu-

clear DNA markers and the phylogeny of the Rubiaceae tribe Spermacoceae. Molec. Phylogenet. Evol.

49:843-866.

Lorence, D.H. 1997. New taxa, a new name, and a new combination in Rubiaceae from southern Mexico and

Mesamerica. Novon 7:46-58.

STANDLEY, P.C.1918. Rubiaceae, Oidenlandieae. In: N. Amer. Fl. 32(1):17-39.

TERRELL, E.E. 1987. Carterella (Rubiaceae), a 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. 2001a. Taxonomy of Stenaria (Rubiaceae: Hedyotideae), a new genus including Hedyotis nigricans.

Sida 19:591-614,

I afl D : ID hi M. £ T,

70 J t Texas 3(

TerreLL, E.E. 20015. Stenotis (Rubiaceae), a new segregate genus from Baja California, Mexico. Sida 19:899-911.

TerRELL, EE. AND D.H. Lorence. 1989. Hedyotis galeottii (Rubiaceae), new combination for a Mexican species. Phyto-

logia 66:1—4.

TerreLt, E.E. AND H. Rosson. 2003. Survey of Asian and Pacific species of Hedyotis and Exallage (Rubiaceae) with

nomenclatural notes on Hedyotis types. Taxon 52:775-782.

Tengen, E.E. AND H. Rosinson. 2004. Seed and capsule characters in Arcytophyllum, Bouvardia, and Manettia (Rubi-

aceae), with notes on A. serpyllaceum. Sida 21:911-927.

TerreLt, E.E. AND H. Rosinson. 2006. Taxonomy of North American species of Oldenlandia (Rubiaceae). Sida 22:

TERRELL, E.E, W.H. Lewis, H. RoBinson, AND J.W.Nowicke. 1986. Phylogenetic implications of diverse seed types, chromo-

some numbers, and pollen morphology in Houstonia (Rubiaceae). Amer. J. Bot.73:103-115.

Tengen, E.E., H. Rosson, W.L. Wacner, AND D.H. Lorence. 2005. Resurrection of genus Kadua for Hawaiian Hedyo-

tidinae (Rubiaceae), with emphasis on seed and fruit characters and notes on South Pacific species. Syst. Bot.

30:818-833.

CAPSICOPHYSALIS: ANEW GENUS OF SOLANACEAE (PHYSALEAE)

FROM MEXICO AND CENTRAL AMERICA

John E. Averett ee Martinez

sic ati Ol P uela de Biologia

PO, Box a Autonoma de Querétaro

Georgia So es UTER Querétaro, 76010 MEXICO

pee Georgia 30460, U.S.A. Mahindaguaq.mx

averett@georgiasouthern.edu

ABSTRACT

B new o from the Moe 2 eastern and southern Mexico and Central America is na da is a m

Luis Potosi, Mexico. Áthenaea cernua

£1 1 f. E y * A +41 1 1

onymy. The genus hasan condo 5-lobed corolla, red fruit, 5-lobed g cal; y

testa, features unknown in related genera of Solanaceae. A p giving the distinguishing psicoy

Cha eee is provided.

RESUMEN

Se descril distrib ] fias del le México, así como en Centro América. Capsicophysalis

está basado en iaa potosina, descri de San Luis on México. Athenaea cernua, descrita originalmente de Guatemala y

que ha sido transferida a tres géneros, qu nero tiene una corola aaa 5-lobulada, fruto aN

un i p D M al pa y eae con una testa tubercalada, T d g

3: f, ] A O a A ]

ES r t r T pal

Key Wonps: Solanaceae, Athenaea, Chamaesaracha, Physalis, Physaleae, Physalineae, Guatemala, Mexico

Hunziker (2001) recognized section Capsicophysalis within Chamaesaracha, a genus of about 10 species

distributed in the arid regions of the southwestern United States and adjacent northern Mexico. Within the

section he included three species: C. cernua, C. potosina, and C. rzedowskiana. Hunziker listed C. cernua as

the type species of the section. All of the species are relatively rare. Hunziker (1980) noted that only seven

collections of C. cernua were known. We add an additional 7 for a total of 14. Until this study C. potosina

has been known only from the type collection, all other specimens having been referred to C. cernua (=

Athenaea cernua). Only two collections of C. rzedowshiana are known to us, both of which are from San Luis

Potosi, Mexico.

Chamaesaracha cernua also has been included in Physalis and Athenaea and is treated as Athenaea cernua

in most recent literature. Waterfall (1967) treated the taxon as a variety of Physalis melanocystis (Robins.) Bit-

ter. More recently, Hunziker (1980) agreed with Gentry (1973) that the treatment of this species in Physalis

was unacceptable and, in addition, made a very strong argument for its exclusion from Athenaea, which, as

now conceived, is a small Brazilian genus of about ten woody species.

Careful morphological comparisons indicate that C. cernua and C. potosina are conspecific with po-

tosina the oldest specific epithet. The species differs in a number of critical features from Chamaesaracha,

Physalis, and Athenaea, and is best treated within a new and separate genus, Capsicophysalis. Chamaesaracha

rzedowshiana differs from Capsicophysalis in distribution, flowers, fruit and fruiting calyx and, for now, is

retained in Chamaesaracha.

Chamaesaracha and Capsicophysalis (as Chamaesaracha cernua) were included by Estrada and Martínez

(1999) in their morphology based cladistic analysis of Physalis and related genera. They concluded that C, po-

tosina was not closely related to either Physalis or Chamaesaracha. In the strict consensus tree, C. potosina forms

a clade with Leucophysalis viscosa which Averett (20092) now recognizes as a distinct genus, Schraderanthus.

C. potosina further differs from Chamaesaracha in that it is an annual herb of mesic riparian forests (Table 1).

J. Bot. Res. Inst. Texas 3(1): 71 — 75. 2009

Capsicophysalis potosina is relatively rare and is represented by only a few collections in herbaria. Un-

fortunately, mature fruiting calyces are not always seen and the feature has not been noted in the literature.

However, it clearly is present on a number of specimens, including type material of both C. potosina and C.

cernua. The very distinctive irregularly lobed corolla, once seen, is easily observed on herbarium sheets but

has not been noted in the literature. The mature calyx also appears to be a dark red on herbarium sheets.

More information on these interesting characters would be welcome.

Capsicophysalis (Bitter) Averett & M. Martínez, stat. nov. Physalis sect. Capsicophysalis Bitter. Repert. Spec. Nov. Regni

70. 1924. Chamaesaracha sect. Capsicophysalis (Bitter) Hunz., Genera Solanacearum 230. 2001. Tyre spectes, Capsicophysalis

potosina (B.L. Rob. & Greenm.) Averett & M. Martínez

Annual or weak perennial herbs to 1 m high; herbage glandular pubescent, mixed with longer hairs; leaves

petiolate, thin to membranous; flowers 1-2 in axils, campanulate—subrotate, 1.5-3 cm wide, corolla white,

yellowish, or yellow-green with villous pads in the throat, unequally 5-lobed and slightly irregular, aestiva-

tion plicate; calyx campanulate, 5-lobed, accrescent in fruit, at first loosely investing the berry, then splitting

and becoming reflexed below the berry, red, the lobes thickened along the margins; fruit a berry, shiny red

or orange-red; seeds discoid 1-1.5 mm long, testa tuberculate.

The striking features of this distinctive genus include the irregular corolla, red or orange-red fruit, the

red reflexed, deeply lobed structure of the mature calyx, and rod-like projections on the seed testa (Fig. 1).

All of these features are uncommon among related genera, and the irregular corolla is completely novel.

The latter character is evident in the types of both C. potosina and C. cernua and present in all of the cited

specimens with flowers. The red fruit is largely unknown among potential relatives except in Brachistus and

Schraderanthus viscosus, both of which have 6—8(-10) flowers arising in fascicles from the axils and other

distinguishing features.

Capsicophysalis potosina (B.L. Rob. & Greenm.) Averett E M. Da comb. nov. (Figs. 1-2). Basionym,

B.L. Rob. & Greenm. Amer. J. Sci. 50:161. 189 (B.L. Rob. € Greenm.) Averett, Ann. Missouri

Bot. Gard. 57: 380. 1971. Ere: MEXICO. San Luis Porosr: Tamasopo Puis M 1880, Pringle 3654 (HoLorvrE: VT!; isorvee: GHI).

Athenaea cernua Donnell Smith, Bot. Gaz. 48:297. 1909. Physalis melanocystis (Robins.) Bitter var. cernua (Donnell Smith) Waterfall,

Rhodora 69:99. 1967. Chamaesaracha cernua (Donn.-Sm.) Hunz., Contr. Gray Herb. 210:25-26. 1980. Tree: GUATEMALA. Derr.

in Verapaz: Sasia, 900 m, May 1908, Tuerckheim II 2245 (hoLoTvee: US!; isotype: CORD

pl id Repert. Spec. Nov. Regni Veg. 20:371. 1924. Tyr: aaa VERACRUZ: : Papantla, pue 1829, ans Note.—

Hunger (1980) terial of thi g

thet i tion. We also have not seen ister of this collection.

Plants herbaceous annuals or weak perennials to 1 m high; herbage largely glabrous except for a few hairs

along the stems and leaf margins; leaves petiolate, blades 2—4 cm long and 1-2 cm wide (about Y as wide as

long), narrowly ovate-lanceolate, margins entire, acute-acuminate at the tip, lamina thin, on short petioles;

flowers 1-2 from axils on pedicels 3 cm long, flowering calyx ca 15 mm long and 12 mm wide, campanulate

and rounded at the base, divided 1/2-3/4 its length, lobes acute to slightly acuminate; corolla ca. 1 cm long,

yellow-white, rotate-campanulate; anthers white-yellow, 2 mm long, filaments ca 3 mm long, connected

to the base of the anthers; fruiting calyx 8-12 mm long and wide, campanulate, exceeding the berry and

becoming reflexed at maturity, berry orange-red to bright red at maturity; seeds dark brown, testa tuber-

culate with rod-like projections.

Distribution —Chamaesaracha potosina is distributed from Guerrero and Tamaulipas in Mexico south

to Guatemala and Honduras (Fig. 2).

Additional material examined: BELIZE. a Arenal-Valentine road, Jun-Aug 1936, Lundell 6181 (US); Vaca Plateau, 8 Mar 1980,

Whitefoord 2023 (MO). GUATEMALA. Peten: Dolores, 22 Aug 1961, Contreras 2746 (LL); Dolores, 5 km E of village, 30 Aug 1961,

Contreras 2830A (TEX); Tikal, 18 Jan 1962, ee 17186 (LL); Tikal Nat'l. Park, 14 Feb 1959, Lundell 15548 (LL); Tikal Nat'l. Park, 9

Mar 1959, Lundell 15805 (LL); Ciebal, Sayaxche, 17 Mar 1970, Contreras 9764 (LL). HONDURAS. Dist. Toledo: Edwards road beyond

Columbia, 15 May 1948, Gentle 6535 (LL). MEXICO. chisves: 9km S E oe 6 Apr 1985, E & Cabrera 8168 (TEX); Mpio.

Ococingo, 14 Sep 1985 Martínez 13823 (TEX). G le O ril 1938, Hinton 14034 (GH, LL); Dist. Galena, Carrizo-El

Río, 20 Oct 1939, Hinton 14689 (GH). Oaxaca: Mpio. Sta. Maria a. 26 July 1985, H. Hernandez G. 1375 (TEX). Tamaulipas:

Averett and Martínez, Capsicophysalis, a new genus of Solanacaeae

GSU0079

Fis. 1. Capsicophysalis potosina: A. flower; B. cleared flower showing unequal lobes; C. SEM of seed; D. immature fruit with calyx enclosing the berry; E.

mature fruit with red berry and reflexed calyx.

na,

Mi pr

Fic. 2. Distribution of Capsicophysalis potosina.

H CG 41

Mpio. Victoria, Cañon de La Libertad, May 1994, Martínez s.n. (UAT); Mpio. de San Nicolás, a 5 km del inicio de la b

28 Nov 1998, Martínez 5329 (QMEX).

GENERIC RELATIONSHIPS

Capsicophysalis probably is most closely related to Schraderanthus viscosus (Schrad.) Averett which Averett

(2009a) recognized as a distinct genus. Hunziker (1991) had included S. viscosus in Leucophysalis and later

(1995) in Chamaesaracha. He returned the species to Leucophysalis in his Genera Solanacerum (2001). Nei-

ther Capsicophysalis nor Schraderanthus seems to be especially close to Chamaesaracha and certainly not

congeneric.

Capsicophysalis has a distribution similar to that of Schraderanthus, Brachistus and Tzeltalia, but C. poto-

sina extends farther to the north in the Mexican states of San Luis Potosi and Tamaulipas. Morphologically,

Capsicophysalis is similar to Schraderanthus and Brachistus which also have orange or red berries, but the

flowers are not in fascicles and, at maturity, the fruiting calyx is reflexed under the berry. Table 1 compares

Capsicophysalis to Schraderanthus and Chamaesaraccha. Averett (2009a, 2009b) provides further discussion

of the history and taxonomy of Schraderanthus and its relationship to Leucophysalis and Brachistus, including

supporting molecular data from Olmstead et al. (2008) and Whitson and Manos (2005).

Hunziker (2001) placed Brachistus in Tribe Solaneae, subtribe Witheringinae while Olmstead et al., (1999,

2008) place Brachistus in Tribe Physaleae, subtribe Physalineae. Capsicophysalis has a strongly accrescent fruit-

ing calyx characteristic of the Physaleae as described by D'Arcy and Averett (1996). We therefore include

Capsicophysalis in Physaleae, subtribe Physalineae, near Schraderanthus and Brachistus. All three have bright

red or orange-red fruit which is relatively uncommon in the Physalineae.

Taste 1. Comparative characters of Capsicophysalis, Schrad hus, and Chamaesaracha.

Capsicophysalis Schraderanthus Chamaesaracha

Habit Erect, herbaceous, annual Erect, herbaceous to woody, Ascending or spreading perennial

annual or perennial herbs

Habitat and Mesic forest, southern Mexico, Mesic forest, southern Mexico, Arid regions of southwestern U.S.

distribution Guatemala and Honduras Guatemala and northern Mexico

Inflorescence

1-2 flowers from axils

6-8 flowers from axils in

1-2 flowers from axils

fascicles

Corolla less than 3 cm wide, w/o 4-5 cm wide, with green broken 2-4 cm wide, with white

maculations in the throat maculations in the throa tomentose pads in the throat

Fruit Red, fleshy berry Red, fleshy berry Green, dry berr

Fruiting calyx Accrescent, reflexing under Accrescent, rotate to slightly Accrescent, appressed to and

the berry at maturity, red reflexed under the berry at partially enclosing the

maturity, red berry, green

Seeds Testa tuburculate with rod- Testa rugose-reticulate, Testa rugose-reticulate, honey-

shaped projections honeycombe combed

ACKNOWLEDGMENTS

Marshall Crosby provided helpful comments about the nomenclature and we appreciate the assistance of

those herbaria which were visited and those from which specimens were borrowed, GH, LL, TEX, QMEX,

US, UAT, and VT. We thank Alan Harvey for assistance with the photographs and distribution map and 3

anonymous reviewers for their corrections and helpful comments on the manuscript.

REFERENCES

Averett, J.E. 2009a. Schraderanthus, a new genus of Solanaceae. Phytologia 91:54—61.

AvtRETT, J.E. 2009b. Taxonomy of Leucophysalis (Solanaceae, Tribe Physaleae). Rhodora (in press).

D'Arcy, W.G. AND J.E. Averett. 1996. Recognition of tribes Capsiceae and Physaleae, subfamily Solanoideae, Solan-

aceae. Phytologia 80:273-275.

ESTRADA, E. AND M. Martinez. 1999. Physalis L. (Solanoideae: Solaneae) and allied genera: a morphology-based

cladistic analysis. In: M. Nee, D.E. Symon, R.N. Lester, L. Jessop, and J.P. Jessop, eds. Solanaceae 4:139-159,

Royal Botanic Gardens, Kew.

Gentry, J.L. 1973. Studies in Mexican and Central American Solanaceae. Phytologia 26:255-278.

Hunziker, A.T. 1980. Studies on Solanaceae XII. Additions to the genus Chamaesaracha. Contr. Gray Herb.

Hunziker, A.T. 2001. Genera Solanacearum. Koeltz Scientific Books, Kónigstein, Germany.

OLMSTEAD, R.G, J.A. Sweere, R.F. SPANGLER, L. Bons, AND J.D. Pamer. 1999. Phylogeny and provisional classification of

the Solanaceae based on chloroplast DNA. In: M. Nee, D.E. Symon, R.N. Lester, L. Jessop, and J.P. Jessop, eds.

Solanaceae 4:111-137. Royal Botanic Gardens, Kew.

OuusreAD, R.G., L. Bons, H.A. Micip, E. SANTIAGO-VALENTIN, V.F. GARCIA, AND S.M. Cottier. 2008. A molecular phylogeny of

the Solanaceae. Taxon 57:1159-1181.

WATERFALL, U.T. 1967. Physalis in Mexico, Central America, and the West Indies. Rhodora 69:84- 120.

Wuitson, M. AND P. Manos. 2005. Untangling Physalis from the physaloids: a two-gene phylogeny of the Physa-

lineae. Syst. Bot. 30:216-230,

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J. Bot. Res. Inst, Texas 3(1): 76. 2009

SYNOPSIS OF GONOLOBUS S.S. (APOCYNACEAE: ASCLEPIADOIDEAE)

IN TRINIDAD AND TOBAGO

Alexander Krings

Herbarium, Department of Plant Biology

North Carolina State University

Raleigh, North Carolina 27695-7612, U.S.A.

ABSTRACT

A synopsis of Mauda S.S. rios OR: "e in d and D Nine; a ee descriptions, and

illustrations, i I g I [ tobagensis The

E 1 $2 1 1; s A a) +h aA 1 2 EI A

RESUMEN

S i isd lol (Apocynaceae, Asclepiadoid ción en ue y ai Aue incluye una clave, des-

ples e codes para f l imi y G. tobagensis.

Se discute 1 ] ] la bibliografía y en los espiclinenes. Sirainens

About fifty Species of subtribe G lobi (A l loid ) occur on the islands comprised

by the Greater and Lesser Antilles, the T Trinidad and Tobago, and Aruba and the Netherland

Antilles. Evidence from the chloroplast (Rapini et al. 2003; Liede-Schumann et al. 2005; Rapini et al. 2006;

Krings et al. 2008) and nuclear genomes (Krings et al. 2008) supports the monophyly of the subtribe, as

well as of Gonolobus s.s. Krings (2008a) recently provided a revision of Gonolobus s.s. in the West Indies.

However, Trinidad and Tobago were excluded from that treatment due their South American affinity. The

objective of this note is to provide a key to and descriptions of the species of Gonolobus s.s. from Trinidad

and Tobago to facilitate their recognition and conservation.

This treatment is derived from a larger critical study of ca. 250 specimens of the fifty some known

species in West Indian Gonolobinae, obtained in part through: (1) loan requests from ninety herbaria—of

which sixty-five responded with either loans, digital images, or negative search results (see Acknowledge-

ments), (2) visits to BM, BSC, DUKE, HAC, HAJB, IJ, K, UCWI, UPRRP, US, and P, and (3) forty-eight days

of field work by the author in the Bahamas (Long Island), Cuba, Dominica, Jamaica, Puerto Rico, St. Lucia,

and St. Vincent. The treatment is also informed by analyses of sequences of portions of the chloroplast

(trnL-F, rps16) and nuclear genomes (LEAFY) of selected accessions (see Krings et al. 2008).

TAXONOMIC TREATMENT

Two species of Gonolobus s.s. are recognized from Trinidad and Tobago. Descriptions of the species are

provided following a key to the species. Corona morphological terminology follows Liede and Kunze (1993)

and Kunze (1995): Ca = faucal annulus (corolline corolla); Cd = dorsal anther appendage; Ci = interstaminal

gynostegial corona; Cs) = fused staminal and interstaminal gynostegial corona; Cs = staminal gynostegial

corona. Species are arranged alphabetically. Following Franz et al. (2008), taxon concept mapping is pro-

vided to facilitate databasing. The operator =is used to indicate whether a given taxon concept is equal to a

previously published concept. The symbol + is used to indicate the misapplication of a name to a concept.

KEY TO GONOLOBUS S.S. IN TRINIDAD AND TOBAGO

1. Calyx lobes ovate, 4.1-8.4 mm x 1.9-5.5 mm, btuse t ded; lla lobes glal laxially .G.

rostratus

1. Calyx lobes linear-lanceolate, 8.5-13 mm x 1.7-2.6 mm, apices acute to narrowly obtuse; corolla lobes

pubescent adaxially . G. tobagensis

J. Bot. Res. Inst. Texas 3(1): 77 — 83. 2009

78 tanical h Insti f Texas 3(

1. Gonolobus rostratus (Vahl) Schult. in Roemer & Schultes, Syst. Veg. 6:61. 1820. (Fig. 1). Cynanchum ros-

tratum Vahl, Symb. Bot. 3:45. 1794. Tyee: Taina: (von Rohr?) Hb. Liebmann (nororvee: C [IDC microfiche photo: Vahl herbarium

nr. 17: III, 5!]; ISOTYPE: B

Gonolobus broadwayae Schltr. in Urban, Symb. Antill. 7(3):340. 1912. Tyre: TeiniDap: Lopez 2419 ( , designated by Krings 2008b:

Herbaceous perennial vines. Latex unknown, presumably white. Stems pubescent, both short, capitate-

glandular, and longer, sharp, eglandular trichomes present, the latter throughout, spreading, or retrorse

internodally, and antrorse-appressed along an indistinct, horizontal ridge between two opposing petioles

and/or just above the node, to 1.07 mm long; nodes pubescent, gland field apparently absent. Leaf blades

ovate to oblong-ovate, 1.9-10.2 cm x 0.9-6.6 cm, apices gradually acuminate with the acumen narrowly

obtuse, to 1.2 cm long, bases deeply cordate, margins entire, evenly strigillose on both surfaces, trichomes

sharp, eglandular, to 1 mm long, glandular emergences from the surface apparently absent, colleters 23,

0.9-1 mm long; petioles 0.9—6.8 cm long, pubescent on all sides, capitate trichomes to 0.16 mm long, sharp,

eglandular trichomes antrorsely-ascending or —appressed, ca. 0.7 mm long; stipular colleters 2, ca. 0.3 mm

long, one borne on each side of the petiole base (rarely on the stem), ca. 0.3 mm long. Inflorescences race-

miform, peduncles 0.4-2.2 cm long, capitate, as well as sharp, eglandular trichomes present, the latter

mostly antrorsely-appressed or -ascending, distributed throughout, ca. 0.4 mm long; pedicels 1.2-4 cm

long, evenly pue da apex to base, capitate-glandular trichomes ca 0.1 mm long, longer, sharp, eg-

landular tricl ly-ascending, ca. 0.8 mm long, bracts linear-lanceolate, ca. 5.1-5.5 mm x 0.8-1.1

mm long, caducous, KAN surface glabrous, abaxial surface coarsely pubescent, trichomes sharp, eglan-

dular, 0.38-0.4 mm long, antrorse. Calyx lobes 5, ovate, 4.1-8.4 mm x 1.9-5.5 mm, apices obtuse to

rounded, margins sparsely glabrous or ciliate, abaxial surface sparsely pubescent at the base and glabrous

toward the apex, trichomes antrorsely appressed or -ascending, to 0.3 mm long; colleters 1 per sinus. Co-

rolla lobes 5, linear-lanceolate or lanceolate, 12-22 mm x 4.3-6.7 mm, slightly lobed at the base, a glandu-

lar swelling frequently present in the sinus, adaxial surface glabrous, abaxial surface sparsely pubescent

with coarse, sharp, eglandular, antrorsely-appressed or -ascending trichomes, trichomes ca. 0.25 mm long;

faucal annulus (corolline corona or Ca) interrupted, a raised bump or indistinct ridge opposite eac h corolla

lobe sinus, pubescent or glabrous; gynostegial corona of iuen staminal (Cs) and ea (Ci) parts,

prostrate-undulating, single, secondary nectaries in i l position absent or at least not formed into

distinct bumps or mounds; anther guiderails without ME laminar dorsal anther appendages (Cd)

1.3-1.5 mm wide, truncate to rounded; style-head ca. 5.4 mm diam, stipe « 0.8 mm long, edentate. Polli-

naria: corpuscula ca. 0.23 mm long, pollinia borne horizontally, narrowly ovate, ca. 1.1 mm x 0.4 mm.

Follicles appareritly known only from a photo taken by P. and Y.S. Comeau, but this was not obtained on

loan.

Phenology.—Flowering in March, April, May. Fruiting in March.

Distribution and habitat. —1In the Caribbean region, G. rostratus is known only from Trinidad, where it

has been found on road banks. Its distribution in South America needs to be re-examined (see discussion

below).

Etymology.—The specific epithet means “beaked.”

Taxon concept mapping.— + G. rostratus sensu Schlechter (1899); = G. broadwayae sensu Schlechter

(1912); = G. broadwayi sensu Cheesman (1947).

Notes.—Schlechter (1899) mistakenly listed “Gonolobus rostratus Griseb. Flor. (1861'[1864]) p. 420 (nec

R.Br)” in synonymy with G. ciliatus Schltr. The lectotype of the latter, i.e., Eggers 5561 (P!; designated by

Krings 2008b), is referable to acd ae ata (Vahl) Fontella & E.A. Schwarz (= Gonolobus denticulatus

(Vahl) W.D. Stevens, if lato; holotype: von Rohr120, C [IDC microfiche photo: Vahl

herbarium nr. 17: III, 11); js. 2008b). For two reasons it seems clear that Grisebach (1864) did in fact

mean G. rostratus as based on Cynanchum rostratum Vahl, rather than *Matelea" denticulata, when he penned

his description. First, he noted the corolla lobes of “his” G. rostratus to be lanceolate-linear. Corolla lobes of

Krinas, $

wm 7

Fic. 1. Ink Al Jig Bi. 0} A

f th ll i , and style- head. E. Detail of f;

| view). Bii. 0 fl ( baxial view). C. Calvcine colleter. D. Faucal

i T i ,

£ J nt A Life 1: . I

im segments (style- head removed). Side view vof gynostegial c corona, b Styl I

. K. Pollinarium. A, C-K based on Britton & Hazen 1601 (GH). Bi-ii

bed on ede 67 faucal annulus of corolla; Ci = interst

corpusculum; C V» y) y y PVE.

80 Journa

“Matelea” denticulata are broadly elliptic to ovate, whereas those of G. rostratus are linear-lanceolate to lan-

ceolate. Second, Grisebach (1864) cited a plate (i.e., t. 7) in Vahl's Icones illustrationi plantarum America-

narum (1798), which is so strikingly similar to the holotype of C. rostratum at C, as to leave little doubt that

it must have been based on it. The ovate calyx lobes of the mature flower are clearly visible in the illustra-

tion. A contrasting illustration of C. denticulatum (basionym of “M” denticulata) is found on plate 8 of the

Icones (Vahl 1798).

Perhaps following Schlechter (1899), Cheesman (1947) also mistakenly considered “Gonolobus rostratus

R.Br. sens. Griseb Fl. 420" synonymous with the entity today known as “Matelea” denticulata, by submerging

the former name under a synonym of the latter: Matelea viridiflora (G.FW. Meyer) Woods. Based on Chees-

man's description (e.g., *cal[yx] lobes lanceolate,” "corolla [...] lobes ovate,” “anthers without dorsal append-

ages”), his concept of Matelea viridiflora (G.FW. Meyer) Woods undoubtedly corresponds to “M.” denticulata

and not Gonolobus rostratus sensu Grisebach (1864) and (Vahl) Schult. as discussed above.

The confusion regarding the application of names appears to extend beyond the Caribbean region and

affects our understanding of the ranges of G. rostratus, G. tobagensis, and potentially other species. Fontella

& Schwarz (1981) proposed that the following names (and combinations based on them) be added to the

synonymy of G. rostratus: Cynanchum rotatum Vell. (lectotype: Fl. Flum. Icones t. 79!, designated by Fon-

tella & Schwarz 1981), Gonolobus patens Decne. (holotype: Martius s.n., M, n.v. [apparently seen by Fontella

and Schwarz, but not relocated by M in searches in Fall 2008]), and G. oxyanthus Turcz. (holotype: Funck 2,

G!, PD). As a consequence of this decision, the name G. rostratus, has now been applied widely to plants from

Tobago to Brazil, Peru, and Argentina (e.g., Fontella & Schwarz 1981; Morellato & Leitáo-Filho 1999; Hechem

& Ezcurra 2006). Plants from West Africa— considered recent introductions (D. Goyder, pers. comm.)—

have been referred to G. rostratus, as well. However, based on a preliminary survey of specimen images

available to me, including three African collections, it appears that G. rostratus may exhibit a much nar-

rower range than presently considered, if not be endemic to Trinidad. The majority! of northern South

American and African material available to me bearing the name G. rostratus, G. patens, or G. oxyanthus

exhibits calyx lobes with acute to acuminate apices, whether linear-lanceolate or ovate, rather than with

obtuse to rounded apices as in G. rostratus (e.g., Argentina: Cabrera et al. 32105, SI; Argentina: Vanni et al.

83, K; Argentina: Venturi 9677, K, SI; Bolivia: Wood €- Goyder 16744, K; Ghana: Merello et al. 1662, K; Ivory

Coast: Fosberg 40488, US; Paraguay: Hassler 227, SI; Venezuela: Aristeguieta 6505, US; Venezuela: Funck 2, G,

P; Venezuela: Pittier 11787, US). In general appearance this material seems much closer to G. tobagensis, al-

though I am not convinced that the latter is the correct name nor that a single taxon is represented. Based

on the protologue, Gonolobus patens, for instance, also exhibits similar linear-lanceolate to lanceolate calyx

lobes (Decaisne 1844). The holotype of G. patens (Brazil, Martius s.n.) resided at M according to Decaisne's

(1844) account, but was not recently relocated (F. Schuhwerk, pers. comm.). Fournier (1885) apparently

saw this specimen—or at least cited it. He described the calyx lobes of this species as ovate and provided

an illustration (t. 94) depicting calyx lobes ovate to lanceolate, but with acute apices, not obtuse as in G.

rostratus (Fournier 1885). Neither Decaisne (1844) nor Fournier (1885) recognized G. rostratus from Brazil.

Fontella and Schwarz (1981) were apparently the first to associate G. patens and G. rotatus with G. rostratus.

Unfortunately, a complete survey of South American and African entities to which the name G. rostratus has

been applied is beyond the scope of the present study and limited by the lack of availability of the type of

G. patens. It appears that either G. rostratus displays considerable morphological heterogeneity or that mul-

tiple taxa are represented. It is hoped that the key and descriptions provided here will contribute to this

much needed survey.

Additional specimens examined: Hab. in Ins. Carib., Anonymous s.n. (L). Trinidad. Apr [18?]74, O. Kuntze 881 (E); Morne Cocoa Road,

bank, 9 Apr 1920, N.L. Britton and T.E. Hazen 1601 (GH, K); Gaura Old Rd., 19 May 1937, Cheesman 13168 (TRIN); 1877-80, A. Fendler

637 (BM, E); Heights of Aripo Rd., close to small ravine or close to Rhapsey's Estate, Mar 1987, F. Moreau s.n. (TRIN); Mar [18?]85, H.

Prestoe s.n. (K).

m . £ D. Quipuscoa 378 (Kl) hac! £f, " I n J r f " ypi 1 mn : " " a

Krings, Sy a er Lal *a IC | I TAL g 81

Fic. 2. Gonolob b g is. A.l l infl Bi 0j fl ( laxial vi ) Bii 0} fl ( baxial vi ) Biii. Flower bud. C Calycine

lleter. D. lla, gynost i , and style-head E Style head and lami | l antl pi | g ( pvi ). F. Detail

lami G. Pollinariu m. Based on Sandwith 1840 (K) f ial

P tvla-head

SQUIRT, NU — Iaimndt Ural GEILE! apperid ys, vuU — CUI puotuluti, S5 — Staal yyliU»tegidi LUTUNI. A »ecymie, rU — pummiit, 311 = style nied’.

82 tani i Texas 3(1)

2. Gonolobus tobagensis Urb., Repert. Spec. Nov. Regni Veg. 16:37. 1919. (Fig. 2). Tye: Tosaco: The Widow, a

climbing plant, 28 Apr 1913, WE. Broadway 4467 (vecromvre, designated by Krings 2008b: BM!).

Herbaceous perennial vines. Latex unknown, presumably white. Stems pubescent, both short, capitate-

glandular, and longer, sharp, eglandular trichomes present, the latter throughout, spreading, or retrorse

internodally, and antrorse-appressed along an indistinct, horizontal ridge between two opposing petioles

and/or just above the node, to 1.3 mm long; nodes pubescent, gland field apparently absent. Leaf blades

ovate to oblong-ovate, 3-10.8 cm x 1—5.7 cm, apices gradually or abruptly acuminate with the acumen nar-

rowly obtuse, 0.9—1.9 cm long, bases deeply cordate, margins entire, evenly pubescent on both surfaces,

trichomes sharp, eglandular, glandular emergences from the surface apparently absent, colleters 2, ca. 0.9

mm long; petioles 1—4.5 cm long, spreading pubescent on all sides, some trichomes capitate, to 0.2 mm

long, but mostly sharp, eglandular, to 1.4 mm long; stipular colleters 2, ca. 0.3 mm long, one borne on each

side of the petiole base (rarely on the stem). Inflorescences racemiform, peduncles 0.2-4.7 cm long, capitate,

as well as sharp, eglandular trichomes present, the latter mostly spreading to somewhat retrorse, distrib-

uted throughout, ca. 1.5 mm long; pedicels ca. 3.6 cm long, relatively evenly pubescent from apex to base,

capitate-glandular trichomes ca 0.16 mm long, longer, sharp, eglandular trichomes antrorsely-ascending,

ca. 1.5 mm long, bracts linear to linear-oblong, 7.6-16 mm x 1.2-2.4 mm, persistent, adaxial surface gla-

brous, abaxial surface coarsely pubescent, trichomes sharp, eglandular, ca. 0.6 mm long, antrorse. Calyx

lobes 5, linear-lanceolate, 8.5-13 mm x 1.7-2.6 mm, apices acute to narrowly obtuse, margins sparsely

ciliate or glabrous, abaxial surface densely and coarsely pubescent at the base and variously toward the apex,

trichomes antrorsely appressed or -ascending, to 0.8 mm long; colleters 1 per sinus. Corolla lobes 5, nar-

rowly lanceolate to elongate triangular, 13.8-18.4 mm x 4. n 3 mm, slightly lobed at the base, a [deed

swelling frequently present in the sinus, adaxial surf te-pubescent on the right half, ab

pubescent with coarse, sharp. eglandular, UN, or -ascending trichomes, particularly at the

base and variously toward the apex, ca. 0.38 mm long; faucal annulus (corolline corona or Ca) interrupted,

a raised bump or indistinct ridge opposite each corolla lobe sinus, pubescent; gynostegial corona of fused

staminal (Cs) and interstaminal (Ci) parts, prostrate-undulating, single, secondary nectaries in interstami-

nal position unknown (could not be determined from specimens); anther guiderails without appendages,

laminar dorsal anther appendages (Cd) ca. 2.9 mm wide, truncate; style-head ca. 4.3 mm diam, stipe ca. 1

mm long, edentate. Pollinaria: corpuscula ca. 0.3 mm long, pollinia borne horizontally, narrowly ovate, ca.

1.3 mm x 0.29 mm. Follicles unknown.

Phenology. Flowering in April, October, and November.

Distribution.—Gonolobus tobagensis has been previously considered endemic to Tobago and Trinidad,

but was recently cited from Guyana, though not the rest of the Guianan Shield (i.e., not in French Guiana,

Surinam, Venezuela: Amazonas, Bolivar, Delta Amacuro; Morillo 2007). In light of the issues presented

above, a re-evaluation of the application of this name to South American material is warranted. In Trinidad

and Tobago, G. tobagensis is known from only four collections (including the type). Its habitat is unclear, but

likely mid-elevation rainforests as for other West Indian congenerics.

Etymology.—The specific epithet means “from Tobago."

Taxon concept mapping.— = G. tobagensis sensu Cheesman (1947).

lditi ined. Tobago: Arima (sp?)-Blanchi Rd. 8^ m. ee oo pls ME Mount E GRO:

Castara Rd (sp? i f f Main Ridge, 18 Oct 1937, N.Y. Sandwith 1840 (K). Ti

1 Apr 1928, W.E. Broadway 6876 (MO).

ACKNOWLEDGMENTS

Ithank the curators and staff of the following herbaria for searching, or providing access to or loans of their

collections: B, BG, BH, BKL, BM, BOLO, BR, BREM, BSC, BUF, C, CGE, COLO, CR, DUKE, E, F, FI, FLAS,

FR, FTG, G, GH, GOET, H, HAC, HAJB, HBG, IA, IJ, ISC, JBSD, JE, K, L, LD, LE, LINN (Linnean and

Smithean herbaria), M, MICH, MIN, MO, MSC, NCU, NEU, NSW, NY, O, OXF, P, PH, RSA, 5, U, UBT, UC,

Krings, S . er ILL . o me sJ J J'TAL 83

UCWI, UPRRP, UPS, US, USF, TUR, WILLI, WU, Z. I also thank the following persons for graciously pro-

viding images of specimens in their collection cited herein: N. Deginani (SD, L. Gautier (G), D. Goyder (K),

A. Lehmann (G), Carolina Loup (P), Marc Pignal (P). The Interlibrary Loan Office of North Carolina State

University was instrumental in obtaining titles not in the collection.

LITERATURE CITED

Cheesman, EE. 1947. Asclepiadaceae. In: EE. Cheesman and RO. Williams, eds. Flora of Trinidad and Tobago.

Department of Agriculture, Port-of-Spain. Pp. 162-175.

Decaisne, J. 1844. Gonolobae. In: A.P. de Candolle, ed. Prodromus Sy tis Naturalis Regni Vegetabilis 8. Treuttel

& Würts, Paris. Pp. 591—605.

FowrELLA P, J. Schwarz, AND E.A. Schwarz. 1981. Estudos em Asclepiadaceae, XIII. Novos sinónimos e novas com-

binacóes. Bol. Mus. Mun. Curitiba 46:1-10.

Fournier, E.PN. 1885. Exolobus. In: CEP. von Martius, ed. Flora Brasiliensis 6(4). F. Fleischer, Monaco. Pp. 318-319.

Franz, N.M, R.K. Peet, AND A.S. WeakLev. 2008. On the use of taxonomic concepts in support of biodiversity research

and taxonomy. New Taxonomy Proceedings of the 5^ Biennial Meeting of the Systematics Association,

Cardiff.

GnisEBACH, A.H.R. 1864. Flora of the British West Indian Islands. Reeve & Co., London.

Hechem, V. AND C. Ezcurra. 2006. Asclepiadaceae. In: LJ. Novara, ed. Flora del Valle de Lerma. Serie Flora 7 (13).

Facultad de Ciencias Naturales, Universidad Nacional de Salta, Salta. Pp. 1—64.

Houmcren, PK. AND N.H. HoLmren. 1998-present (continuously updated). Index Herbariorum. New York Botanical

Garden. [http//sciweb.nybG.org/science2/IndexHerbariorum.asp]

KRINGS, A. 2008a. Revision of Gonolobus s.s. (Apocynaceae, Asclepiadoideae) in the West Indies. J. Bot. Res. Inst.

Texas 2:95-138.

KRINGS, A. 2008b. Index of names and types in West Indian Gonolobinae (Apocynaceae, Asclepiadoideae), includ-

ing fourteen new lectotypifications, one weotyoification: and a new combination. J. Bot. Res. Inst. Texas

2:139-163.

KRINGS, A., D.T, Thomas, AND Q.- Y. XIANG. 2008. On the generic circumscription of Gonolobus (Apocynaceae: Ascle-

piadoideae): Evidence from molecules and morphology. Syst. Bot. 33:403-415.

Kunze, H. 1995. Floral morphology of some Gonolobeae (Asclepiadaceae). Bot. Jahrb. Syst. 117:211—238.

Lieve, S. AND H. Kunze. 1993. A descriptive system for c analysis in Asclepiadaceae and Periplocaceae. Pl. Syst.

Evol. 185:275-284.

LIEDE-SCHUMANN, S., A. Rapini, D.J. Goyder, AND M.W. Chase. 2005. Phylogenetics of the New World subtribes of Ascle-

piadeae (Apocynaceae-Asclepiadoideae): Metastelmatinae, Oxypetalinae, and Gonolobinae. Syst. Bot.

30:184-195

MontLLATO, PC. AND H.F. Lerráo-FiHo. 1999. Reproductive phenology of climbers in a southeastern Brazilian forest.

Biotropica 28:180-191.

Morio, G. 2007. Asclepiadaceae. In: V. Funk, T. Hollowell, P. Berry, C. Kelloff, and S.N. Alexander, eds. Checklist of

the plants of the Guiana Shield. Contr. US Natl. Herb. 55:208-212.

Rapini, A., M.W. CHASE, AND T.U.P. Konno. 2006. Phylogenetics of South American Asclepiadoideae (Apocynaceae).

Taxon 55:119-124.

Rapin, A, M.W. Chase, DJ. GOYDER, AND J. GrirritHs. 2003. Asclepiadeae classification: evaluating the phylogenetic

relationships of New World Asclepiadoideae (Apocynaceae). Taxon 52:33-50.

Schlechter, R. 1899. Asclepiadaceae. In: |. Urban, ed. Symbolae Antillanae. Gebrüder Borntraeger, Berlin. Pp.

ScHLEcHrER, R. 1912. Asclepiadaceae. In: |. Urban, ed. Symbolae Antillanae. Gebrüder Borntraeger, Berlin. Pp.

Var, M. 1798. Icones illustrationi plantarum Americanarum. Hauniae, Copenhagen.

BOOK REVIEWS

PuiriP V. Werts. 2000. The Manzanitas of California: also of Mexico and the World. (ISBN: 0-933994-

22-2, hbk.). Originally published by the author, Department of Botany, University of Kansas, Lawrence,

Kansas 66045, U.S.A. (Orders: check or money orders by mail to Dr. Anke M. Wells, 4638 Bluebird

Lane, McLouth, Kansas 66054-3109). $60.00 incl. ph (plus sales tax for Kansas residents), 151 pp.,

8 1⁄2" x 11".

Published by the author in 2000, this li d edition book is based han 40 f field and hert ji

by the late Philip V. Wells (1928-2004, Professor Emeritus of Ecol 1 Evolutionary Biol he University of Kansas in Lawrence

Kansas. His lifelong interest in manzanitas started in the late 1950s while Mods at the neu of California in Santa Barbara

and culminated in this 151-page work, the most up-to-date and complete treatise of the genus. The book i dee d synonyms,

oe and peas for 61 Audios bd is 150 o 146 are pere for the first tim ludi ital scans of

und-breaking work. n attractive PUE book

di fu xq rir i DER

measures 8.5 x 11 inches and is d on acid- AR paper and bound E red e 7 and gold lettering. Approximately 200 copies

are available.— Craig C. Freeman, Curator (R.L. McGregor Herbarium), Associate Scientist (Kansas Biological Survey), University of Kansas,

Lawrence, Kansas 66047, U.S.A.

Davip Yerman. 2007. The Great Cacti: Ethnobotany & Biogeography. (Arizona-Sonora Desert Museum

Studies in Natural History) (ISBN 978-0816524310, hbk.). University of Arizona Press, 355 S. Euclid

Ave., Suite 103, Tucson, Arizona 85719, U.S.A. (Orders: www.uapress.arizona.edu, ordersQuapress.

arizona.edu, 1-800-426-3797). $59.95, 297 pp., 8 3/4" x 11 1/4".

The Great Cacti reviews the more than one hundred species of columnar cacti, with detailed discussions of those that have been the

most beneficial to humans or are the most spectacular. The book is divided into three Ae parts. The first covers the great cacti

in general and looks at their uses, Scotos esito their origins ev p and tax y. The second, and largest of the three

Eos ies of the genus. Some genera are

parts, presents the plants by genera, b

grouped by geographical region, Stier; are not. x. The third part anli where the ho spots for

he has not personally visited but discusses what he considers to be the most impressive, accessible locations where one can see them

has not listed sites where

in their native lands

tman focuses particularly on northwestern Mexico and the southwestern United States and South America to some degree.

He examines the role of each a in uud society, oo ae cacti have provided ae ae e even Tehsious y

significant hallucinogens. great i "elo t

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y ielding their pr duct He draws upon!

importance of these TE to the native MCA in these regions. The oral uses are listed with each species, generally in a

small portion of the first section.

TH 11 2n 1

y I 1 col ps, The Great Cacti i lous book, one which I enjoyed just paging through.

The use of color d helm th Ith of sci h he displays throughout his narration. Recommended for public

libraries, int d student d ] y levels.

David Y a 1 001 he Gaul pea pe eke. reas J] frhe televici ies The

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d f North N , also published e the ais of Arizona Press.—Gary L. Jennings, Librarian, Botanical Research

Institute of Texas, Fort Worth, Texas 76102-4025, U.S.A

J. Bot. Res. Inst. Texas 3(1): 84. 2009

AN OVERVIEW OF THE DIOSPYROS CAMPECHIANA COMPLEX (EBENACEAE)

AND DESCRIPTION OF THREE NEW SPECIES

Mitchell C. Provance and Andrew C. Sanders

UCR Herbarium

m of Botany and Plant Sciences

Uni y of California, Riverside

uic ‘aloo 9252 i -0124, U. SA.

ABSTRACT

The Di hi lex (El ) is described in detail. Six speci gnized for the complex in Mesoamerica, and

one species in South America. Expanded d ipti ided for the three M i ies, and three new species are de-

P I p p E P

scribed. O ii,i finedt 11 he Pacific Sl f the Si Madre del Sur, Oaxaca,

Mexico, RS two ee are om n only hon rum Rica, including Diospyros crotalaria, en is restricted to the Osa Peninsula,

and Diospyros haberi, which is widespread in mountainous regions. Members of the complex are remarkable for having fruiting

pedicels that tend to hold fast to the calyx, despite being articulated to the receptacle. The fruit itself is weakly attached to the calyx.

Photographs, comparison tables, and a key to the species of the complex are provided.

RESUMEN

Sed ik detalle el lejo Di } El JS i i M ica, y una en Sur América.

Se aportan descripciones en extenso para las tres Ao O y se describen tr peci Una de las especies

la Si Madre del Sur, Oaxaca, México,

nuevas, Dio pyr p :

t d sólo de Costa Rica, iicluyendo Diospyros iota lista: que está restringida a la Península de Osa,

y Diospyros aver de = oo montañosas. Los eee de d son Mrd hor tener OB a a pesar de estar

Las semillas se

dispersan pobbi por pájaros. Se aportan fotografías, tablas ans y una clave de especies del complejo.

While ining material during continuing studies of Mesoamerican Diospyros (e.g., Provance & Sanders

2005; Provance & Sanders 2006; Provance et al. 2008), building toward production of a comprehensive

treatment of the Mexican species, three new taxa belonging to the Diospyros campechiana complex were

discovered. Because this complex has only been discussed briefly (Provance et al. 2008), here we provide a

more detailed account. The addition of three new species brings the number of included species to seven.

One of the new species, D. camposii, is described from Oaxaca, Mexico, based on collections from a restricted

area of the Pacific Slope of the Sierra Madre del Sur, and appears to be a local endemic. Two other species,

including D. crotalaria and D. haberi are described for Costa Rica. Diospyros crotalaria appears to be a local

endemic restricted to the Osa Peninsula, while D. haberi is fairly widespread in the central mountains. Previ-

ously described species include D. campechiana Lundell, D. panamense S. Knapp, D. hartmanniana S. Knapp,

and D. juruensis A.C. Smith. The distribution of the complex extends from Oaxaca and Veracruz, Mexico

southward to Panama, and likely enters Columbia, since D. panamense has been collected very close to its

border. The complex definitely occurs in South America because D. juruensis occurs in the Jurua River Basin

of the Amazon watershed in western Brazil, where the only known collection was made.

We are unsure about the complete status of this complex in South America because most South Ameri-

can specimens are unavailable for examination. In addition, the protologs of twenty-five recently described

species of South American Diospyros (Wallnófer 1999, 2000, 2003, 2005) completely lack photos or other

illustrations, and are devoid of comparisons to allied, or even similar-looking taxa. This makes many of

them difficult or impossible to place with confidence. Identification keys were also not provided, and no

attempt was made to fit them into existing keys (e.g., MacBride 1959; Cavalcante 1963a; White 1981; So-

thers & Berry 1998). While not violations of the International Code of Botanical Nomenclature (Greuter

J. Bot. Res. inst. Texas 3(1): 85 —112. 2009

86 Jour tani i Texas 3(

et al. 1994 2000), the figure omissions are inconsistent with Recommendation 32B, and the omission of

comparisons to allied taxa is contrary to Recommendation 32C of the Code. Comparisons among taxa are

made in virtually all of the other relevant literature (e.g., Miquel 1856; Hiern 1872; Milbraed 1927; Gleason

& Smith 1933; Sandwith 1931, 1949, 1963; MacBride 1959; Cavalcante 1963a, 1963b, 1966, 1977; White

1981; Sothers & Berry 1998; Sothers 2003), and the illustration of types has been standard procedure for

most modern taxonomists. Consequently, a more thorough examination of South American specimens may

reveal that additional members of the D. campechiana complex are present there. However, our examination

of the literature and the limited South American Diospyros specimens to which we have had access did not

reveal any other such species.

The species in this complex are so poorly represented in herbaria that from over fifty herbaria that

provided specimens of Diospyros, we received only seventy-three specimens (representing fifty collection

events) that were referable to species in it. Little or nothing has been recorded concerning bark texture and

color, color of the slash, wood characteristics, fruit color, texture and taste, flower odor, and color of the seeds

and guard cells in life for most species. Flowers are completely unknown in D. camposii and D. crotalaria,

and male flowers are known from only one or a few collections in the other species. Specimens with female

flowers are ially rare: we have seen open female flowers only in D. campechiana, and only a single female

flower bud in D. haberi. Regardless of sex, flowering specimens typically have few open flowers. Fruits seem

to be unknown in D. juruensis.

METHODS

All of the relevant literature and specimens, including those types we could obtain, were evaluated. Collec-

tions were sorted into preliminary groups based on ue and LAB RUE morphological characters,

and these eodd were considered taxonomic hypotheses nvestigation. Comparisons

were then made | I igned to each group and ee specimens, specimens determined to be

consistent wit! i tol ( ially if from near the type locality) and paratypes determined to be

consistent with the al description: We E examined specimens in light of floristic and monographic

treatments, and the annotations of previous workers. Expanded descriptions have been constructed for pre-

viously described species based on the studied herbarium specimens. In this treatment we use ‘lanceolate’

in the sense of Jackson (1916), being broadest near the lower third of the leaf, not at mid-leaf in the sense of

Stearn (2000). Descriptions apply to herbarium material except when indicated, or when obviously referring

to fresh material. Map coordinates geo-referenced by the authors are given in brackets. Distribution maps

were constructed using The Generic Mapping Tools 4.1.4 (Wessel and Smith 2006) accessed through OMC

(Weinelt 2006) and amended using Adobe Photoshop (Ver. CS 8.0, Adobe Systems Incorporated, San Jose,

CA). Illustrations are by the first author.

TAXONOMIC TREATMENT

Diospyros campechiana complex: Members of the Diospyros campechiana complex are evergreen trees

with medium to large, more or less evenly spaced leaves. Petioles and midribs are occasionally transversely

fissured, and frequently have a bloom of minute white crystals (scintillae) that are probably derived from

cuticular wax, though their origin has not been studied. Stems and leaves are glabrous to sparsely hairy,

and members of the complex have been variously referred to as “black strigillose,” in reference to D. campe-

chiana (Standley & Williams 1967), or “minutely strigillose” in reference to D. hartmanniana (White 1978).

At low magnification the vestiture appears to be comprised of minute, mostly black, appressed, fusiform

hairs. At higher magnification it becomes evident that most hairs are dark red and 2-armed, with a short

basiscopic arm, and a longer acroscopic arm. Less often hairs are simple, and in some speci there may be a

high frequency of similar, but paler, yellowish, or reddish brown hairs. In addition, some taxa are minutely

hirtellous or minutely pubescent, with hairs much smaller than those of the aforementioned strigillose

vestiture. Stems sometimes have localized patches of atypical hairs (e.g., upright, aspergilliform, etc.) which

D Ic J E n [ hi [| 87

we think represent endophytic fungi. In this paper the term “strigillose” refers to the common vestiture of

dark red to black, simple and 2-armed hairs, with the addition of modifiers for special cases (e.g., “reddish

brown strigillose”).

The abaxial lamina surfaces typically have some conspicuously darkened guard cells (Fig. 1), and in all

species except D. campechiana, the stomatal apparatus is recessed, giving the epidermis a black-puncticulate

appearance at 20x magnification. The black dots on the abaxial lamina surface of D. hartmanniana that were

interpreted by S. Knapp (1997) to be the bases of broken hairs may have been these pigmented guard cells.

We suspect that darkening of guard cells is a result of secondary compound degradation in non-functioning

cells of older leaves.

Male inflorescences are (1—)2—23(-29)-flowered glomerules, compact cymes, paniculiform cymes,

or occasionally fasciculate cymes, in leaf axils of young stems. Their axes are 4-angled, and more densely

hairy than leaves and stems, though the hairs are otherwise similar. Female inflorescences are solitary or

2—5(-10)-flowered cymes in leaf axils of young stems, and often more or less 4-angled.

Flowers have 4—5 calyx and corolla lobes, though rarely a calyx or corolla will have only 3 lobes. Co-

rollas are urceolate and white in life, though they turn dark brown upon drying. Male flowers have 10—28

stamens with their filaments fused into 5-14 pairs (comprised of an inner and outer member). Filaments

are adnate to the base of the corolla or inserted on the receptacle. Filaments and/or anther connectives are

sericeous. The apex of the anther is tapered to rostrate, and consists mostly of minutely muricate connective

tissue. The pistillode in male flowers is markedly lobed, with the number of lobes being equal to, or slightly

less than, the number of stamen pairs surrounding it. In D. campechiana the filaments of adjacent stamen

pairs are separated by these lobes, and in D. panamense and D. hartmanniana the indentations between the

lobes resemble impressions of the filaments, suggesting that pressure from filaments against the pistillode

during floral development may be responsible for the lobe pattern on the pistillode.

Female flowers in D. campechiana poses eat unpalted. staminodes, with several short hairs on the

baxial surf; fth tive, and l 1 les with slits. Open female flowers and their internal

structures have only been ea in this complex in this one species, though a single mature flower bud

has been observed in D. haberi. Despite female flowers being largely unknown in the complex, styles often

persist on fruits, and their number (1-2) and morphology are useful in species identification.

The female calyx accresces during enlargement of the developing fruit, though not as markedly as in

members of the D. rosei complex. The fruiting calyx tube is patent (“saucer-shaped” sensu White 1978; Knapp

1997) with a short basal protuberance. The apex of the fruiting pedicel is 4-angled, + pyramidal in shape,

and is jointed to the receptacle within the protuberance at the base of the calyx. Although Diospyros is typi-

cally described as having fruits with a persistent calyx (e.g., Wallnófer 2001), members of the campechiana

complex have a fruit that is weakly attached to the calyx. Despite having a pedicel with an articulated apex

(as in other Diospyros species), in members of this group the fruit often detaches from the calyx, while the

connection of the pedicel apex to the base of the calyx holds firm.

The fruits of members of the complex are small to medium-sized, ovoid-ellipsoid or depressed-globose

to subglobose, and reportedly green, yellow, orange, or red when mature. Comparing fruit shape between

species is not straightforward, since it is highly influenced by the number of seeds that develop. Fruits are

usually asymmetric (e.g., Figs. 4b, 6b) when seeds only develop on one side. The fruits have only a small

amount of flesh, which is reddish and vitreous when dry, though possibly viscous and clear in life. There

are 1-4 locules separated by complete septa in the taxa in which fruit anatomy was studied. Based on fruit

lobes, this also seems to be the case in the remaining taxa. Each locule contains a lone seed which lacks

adherent pericarp. The large locules include considerable space around the seeds, and when dry fruits are

shaken they produce a rattling sound.

The combination of brightly colored small to medium-sized fruits, that readily separate from the calyx,

and contrasting black or very dark inflorescence branches, suggests avian seed dispersal in the complex.

We suspect that dispersal in the campechiana complex is effected by removal of the fruit from the calyx,

Journal of the Botanical Research Institute of Texas 3(1)

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Fic. 1. Epidermis details in species of the Diospyros campechiana complex. A. Abaxial leaf surface of D. camposii (scale = 0.5 mm) from A. Campos V. & R.

Torres C. 825 (MO). B. Abaxial leaf surface of D. crotalaria (scale = 0.5 mm) from R. Aguilar 2877 (K). C. Pigmented and non-pigmented guard cells on the

abaxial leaf surface of D. hartmanniana (scale = 0.125 mm) from G. McPherson & N. Hensold 15310 (B-100152291). D. Pigmented and non-pigmented

guard cells on the abaxial leaf surface of D. crotalaria (scale — 0.25 mm) from R. Aguilar 2877 (K).

D Ic J , Di hs I 89

Dei r T

and swallowing of the fruit whole, unlike in other groups (e.g., the salicifolia complex), where fruits are torn

open and seed llowed incidentally along with adherent pericarp (White 1978; Provance pers. obs.).

Alternatively, the combination of detachment of the fruit from the calyx and air space surrounding seeds in

mature fruit may suggest a hydrochorous dispersal syndrome (Kubitzki & Ziburski 1994).

Diospyros campechiana Lundell, Contr. Univ. Michigan Herb. 4:23. 1940. (Fig. 2, 3, 15b). Ber: MEXICO.

CAMPECHE: Palizada, [18° 15'N, 92? 05'W 3 m], 25-28 Jul 1939, E. Matuda 3843 (HoLoTYPE: MICH; isotypes: E K!)

Trees 6—20 m tall, evergreen; trunk up to 45 cm dbh, bark unknown, latex reportedly yellow to orange;

young stems smooth to shallowly furrowed, brownish, usually dark, viscid, sparsely to moderately strigil-

lose, also densely hirtellous, the hairs minute, light gray to + colorless, shorter than the appressed hairs;

mature stems lenticellate, glabrate. Leaves alternate, simple, entire; PEOS E B mm long, minutely

hirtellous, sometimes with a few transverse fissures, rounded below, the ey linally wrinkled,

often scintillant, + flat above, somewhat canaliculate distally; lamina subcoriaceous, EOM M to

oblong-elliptic, sometimes oblong-oblanceolate, 100—285 mm long, 35-90 mm wide, length to width ratio

ca. 2.1-3.5 : 1, base acute to attenuate, sometimes Epes: eee decurrent on the petiole, margin revolute,

apex acuminate with a rounded tip; 1 ll brown pure to strigillose, guard cells

sometimes conspicuously darkened, put not ee pper |! 11 brown to somewhat glossy,

usually papillose. Venation ] ; midrib + rounded below, longitudinally

wrinkled, depressed above, d and minutely firtellous ad strigillose near the base of the lamina;

lateral veins 7—12 per side, prominent below, slightly raised above; 3? veins slightly raised below, more so

above. Laminar extrafloral nectaries on the abaxial surface only, common, round to elliptic, 0.3-0.5 mm

long, minutely rimmed, drying dark green to black. Male inflorescences a single (3)6-23(29)-flowered

compound cyme in the leaf axil of young stems (sometimes bordering on being a fascicle of 23) cymes),

5-17(31) mm long, initially paniculiform, but the terminal and often the penultimate units dichasia, axes

4-angled, densely strigillose, also minutely hirtellous, sulcate, subtended by ovate to triangular, concave

bracts + 1 mm long; peduncles 1.5-5.5 mm long; pedicels 0-1.5 mm long. Male flowering calyx sub-

crateriform (urceolate prior to anthesis), with a short basal protuberance, exterior drying dark brown to

black, strigillose, papillate, interior glabrate to locally strigillose, 272.4 mm wide, tube + 0.7 mm long, lobes

4—5, depressed-orbicular to widely ovate, 0.8-1 mm long, 0.8-1.4 mm wide, sometimes emarginate, often

slightly asymmetric, suberect; male corolla white in life, drying brown, glabrous except for some minute

papillae, tube urceolate, 1.2-1.7 mm long, + 2.3 mm wide, lobes (324(—5), 1-1.6 mm long, 1.4-1.9 mm

wide, depressed-orbicular to widely ovate, sometimes emarginate, often slightly asymmetric; stamens

(12-)16(-20?), with Whitefoord and Knapp (1998—onward) reporting 12-16 stamens and Pacheco (1981)

reporting 16 stamens, these inserted on the receptacle, fused near the base of their filaments into (6—)8(-10?)

pairs, the outer stamen larger, bowing inward at the middle, anthers yellow, sometimes with minute red

spots, lanceolate, outer anthers 1-1.2 mm long (including the connective), sericeous on the abaxial surface

along the connective and sometimes on the adaxial surface, hairs yellow to reddish black, inner anthers

similar except smaller, less hairy, and the hairs mostly on the adaxial surface, the apex (connective) tapered,

minutely muricate, filaments 0.2-0.8 mm long, sericeous near the base of the anther; pistillode rotate,

markedly 8(-10)-lobed, 0.7-1 mm wide, glabrous, with 2(-1) broadly rostrate apical structures. Female

inflorescences 1-3 compound cymes in leaf axils on young stems, paniculiform, but the terminal units

dichasia, (123-5(-10)-flowered, though no more than three fruit seen on a single inflorescence, 5-8 mm

long, axes 4-angled, strigillose, also minutely hirtellous, sulcate, subtended by persistent bracts similar to

those of male inflorescences; flowering peduncles 0.5-2 mm long; flowering pedicels 0.5-2 mm long;

fruiting peduncles 4-6 mm long; fruiting pedicels 2-5 mm long, with a 0.4—0.5 mm long, 4-angled,

pyramidal apex, only the distal 0.2-0.3 mm forming a joint within the basal protuberance of the calyx.

Female flowering calyx campanulate-infundibuliform to crateriform (urceolate prior to anthesis), with a

short basal protuberance, exterior drying dark brown to black, moderately strigillose, minutely papillate, ca.

2.5 mm wide, tube 1.2-1.5 mm long, lobes 4-5, depressed orbicular, (1.121.7-1.9 mm long, 1.5-2.5 mm

Fic. 3. Diospyros campechiana. A. Male flower (C.L. Lundell & E. Contreras 20739, MO). B. Pistillode (C.L. Lundell & E. Contreras 20739, MO). C. Female flower

(L. Pacheco & J.J. Calzada 20, XAL). D. Female flower (L. Pacheco € J.I. Calzada 20, XAL).

L Das H ID El A p, ET,

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92 Journal of Texas 3(

wide, erect; female flowering corolla white in life, drying brown, tube 0.9 mm long, 1.3-1.5 mm wide,

slightly constricted distally, throat 1.2 mm long, semiglobose to urceolate, 2.3-2.5 mm wide, lobes 4—5,

widely ovate, 1.7—1.9 mm long, ca. 1.6 mm wide, erect (L. Pacheco & J.I. Calzada 20) or spreading slightly

(W.E. Harmon & J.A. Fuentes 5790); pistil + globose, glabrous; style 12.1 mm long, glabrous, the basal

portion 4-angled, divided from the basal 1/3 to 1/2 into two ascending branches (style and stigma [see

below] details based on those seen in flowering material and persisting on fruit); stigmas minute, undulat-

ing irregularly along the abaxial perimeter of the style branches; staminodes 8, with several short hairs

on the abaxial surface of the connective, antherodes lanceolate, with slits. Fruiting calyx not markedly

accrescent, relatively thin (not fleshy), drying very dark, exterior glabrate to strigillose, minutely glandular

punctate (punctae developing from the minute papillae of flowers), tube patent, with a basal protuberance

0.8-2 mm long and 1.5-2 mm wide, the tube 3.5-6 mm wide between opposing sinuses, interior glabrate

to moderately strigillose, sometimes minutely glandular punctate, scattered clavate glandular hairs some-

times present, a circular band of short reddish black hairs sometimes pnus at the base, lobes spreading,

(222.5-4 mm long, (324—5(-6) mm wide, depressed orbicular, ly revolute,

ciliate, adaxial surface glabrate to moderately strigillose. Fruit glabrous, (10213- 17 mm long. 10-17 mm

wide, ovoid-ellipsoid, often asymmetric, locules 1—4, separated by complete septa; flesh sparse to negligible,

red and vitreous when dry, unknown in life, though likely viscous and clear; epidermis probably smooth

in life, usually wrinkled when dry, color progressing from green to yellow to orange with maturity (rarely

reported to be red), orange to dark brown when dry; seeds 1—4, reddish brown to brown, loosely contained

in the fruit, ovoid to ellipsoid, the shape depending on the number of seeds that develop, circular in x.s

(when one seed develops), or with 1—2 flat surfaces (when 2-4 seeds develop), 9.5-12 mm long, 5.7-8.4

mm wide, the dorsal surface with a prominent vascular strand, texture rugulose-foveolate, hilum apical on

a minute protuberance.

Diospyros hiana is probably most closely related to D. camposii. Separation of these taxa is discussed

under the description of the latter. Generally, D. campechiana is an easy species to identify. In addition to

the geographical separation of this species from other members of the complex, the narrow oblong leaves

are distinct, being reminiscent only of D. crotalaria. However, the minutely hirtellous vestiture of the stems,

petioles, midrib, and inflorescence seems to be diagnostic.

Fruits in fragment folders that have retained their calyx often have the broken tip of the pedicel apex

lodged inside the protuberance of the calyx. Thus, in many cases, fruits and calyces that broke away as a

unit during the collection process, did not separate from the mother plant at the point of articulation of the

pedicel and receptacle/calyx, but rather the pedicel broke near the opening of the cavity in the protuberance

at the base of the calyx.

It seems likely that pressure from filaments during floral development creates the lobed pattern on the

pistillodes of species in this complex. Because we observed mostly 8-lobed pistillodes in this species, we

surmise that the stamen number is id is 16.

Distribution and ecology.—Di ana occurs in lowland tropical forests associated with wet-

lands and riparian areas of major river m and their tributaries between sea-level and 270 m elevation

in the states of Tabasco, Campeche, Chiapas, and Veracruz, Mexico, and Guatemala (Fig. 5).

Specimens po MEXICO. Camrecne. Mpio. Palizada: 20 km de dade i e Au a A Pil 07' 19"N, 92? 07'

15"W, [0-4 mi, Dec 2000, P. Sima et al 2555 (MO). CHIAPAS Mpio Ocosingo: a3 Corozal, I Rio Usumacinta,

selva alta Rd [near 16? 49'N, 90? 53'W], 120 m, 17 Aug 1984, E. Martinez S. 7326 did Tarasco: “Habitat D margins fluvii

Gonzalez,” [18° 15'N, 92° 55'W, 10 m], 8 May 1889, J.N. Rovirosa 482 (K). Mpio Jalapa: 41 pa-Tacotalpa,

potrero, [near 17? 50'N, 92? 48'W], 10 m, 22 Nov 1983, F. Ventura A. 20785 (MO, XAL); Mpio. Comalcalco, San Cayetano, potrero, [18°

22'N, 93? 13'W], © m, F. Ventura A. 20796 (MO, XAL). Mpio. Jalapa-Tacotalpa: en el cerco de un potrero, 10 m, 23 Nov 1983, R. Curiel

A. & M.A. Guadarrama O. 108 (CHAPA, XAL). Mpio. Macuspana: 8 km de la entrada de Macuspana hacia Escarcega y 1.5 km al N,

s FREE asociado con Pachira, Bactris, [near 17° 41'N, 92° 34'W, 45 a 25 Aug 1981, M.A. ue Á. & S. Zamudio 358 (XAL).

M cajuca: Tucta, a 2 km de Incunac median caducifolia, 2°, gado, [18° 12'N, 92? 59'W], 26

m, 6 Oct 1978, J.I. Calzada 4906 (XAL). Veracruz. Mpio. Ignacio de la Llave: [probably the town of Ignacio de la Llave], acahual, [18°

43'N, 95? 58’W], 50 m, 13 Nov 1967, G. Martínez C. 1555 (USF); 500 m de d pieza la desviacion Villa Nueva-Zacate Colorado,

n AC [| : n: hi I 93

en potrero, 18° 43'N, 95° 59'W, 50 m, 29 Apr 1981, L. Pacheco & J.I. Calzada 20, 21 (XAL). Mpio. Las Choapas: Las Choapas, orillas

del Rio Playas, cerca de Abasal (one sheet indicates “Rio Playas, arriba de Abasal”), [near 17° 45'N, 93° 57'W], 30 m, 4 Aug 1984, Miguel

Chazaro & Luis Robles 3081 (XAL-2 sheets). GUATEMALA. ALTA VERAPAZ: Sebol, along Rio Sebol, in high forest, [15° 48'N, 89? 57', 153

m], Aug 1964, Elias Contreras 3354 (F, MO); Sebol and vicinity, [15° 48'N, 89? 57', 153 m], Aug 1964, Elias Contreras 5355 (1J, F digital

d Chahal, bordering Rio Chiyú, 1km W, in low forest, [near 15? 50'N, 89° 34'N], 29 Oct 1968, Elias Contreras 8032 (F, MO). PETEN

Cumbre, is 142 of Cadenas Road, in low forest on wet land, [16? 05'N, 89? 21'W, 270 m], 24 Sep 1966, Elias C 220 (F); Rio

Pasion, Al ifi in high forest on bank of river Pd 30'N, 90? 32'W, + 380 ft], 8 Feb 1964, - L. Lundell e (F, If, MO); El

Rosario, the FAO-FYDEP Camp at S h li laguna, [16? 31'N, 90? 10'W], 1 19 Jun 1971, W.E. Harmon

eLA.F 5790 (UMO); La Cumbre, caserio "Sapurul", bordering a “Sapurul”, in zapotal x a forest, Ade 16? 05'N, 89?

21'W, 270 m], 7 April 1977, C.L. Lundell & Elias Contreras 20739 (WIS, MO); La Cumbre, Pusila Road, 5 km N, in acahual, [16° 07'N,

89? 21"W, 250 ml, 19 Aug 1976, C.L. Lundell & Elias C 20224 e UCR); Brecha Chinajá, 12 km from Laguna Petexbatun, high

forest, [16° 18'N, 90° 08'W, 540 ft], 20 May 1965, Elias Contreras 5401 (MO).

Diospyros camposii M.C. Provance & A.C. Sanders, sp. nov. (Figs. la, 4, 15c). Tre MEXICO. Oaxaca. Mpio. San

Jerónimo Coatlán: 41.5 km al SW de San Jerónimo C.[Coatlán], brecha a Progreso, bosque de pino-encino alterado, suelo amarillo

arcilloso, 16? 10'N, 96? 59'W, 1550 m, 29 Nov 1990, A. Campos V. 3452 (HoLotyre: CHAPA!; isotype: MEXU)

ae D. n anm oL similis, sed yen IBS, jum ce eee crassiore, intervallo grandiore inter sinos oppositos, lobis

fructibus, et st ibus, caulibus, petiolis, costa inflorescentibusque non hirtellis,

TUM breviore et marginis minute porcatis, sedi non revolute. ten:

Trees 8-25 m tall, pr obably evergreen, t | ] ; stems brownish, shallowly furrowed when young,

sometimes smooth, sparsely strigillose, becoming lenticellate and scaly in older stems. ET alternate,

simple, entire; petioles 8—14 mm long, with a short marginal wing distally, y fissured,

sparsely strigillose, epidermis scintillant; rounded below, epidermis longitudinally wrinkled, + flat above,

somewhat canaliculate distally; lamina chartaceous to subcoriaceous, widely ovate to elliptic, 55-152

mm long, 33-69 mm wide, length to SUE ratio ca. 1.5-2.5 : 1, base acute to shortly-attenuate, sometimes

oblique, decurrent on the petiole inutely ridged, apex acuminate with a rounded tip; lower lamina

surface dull brown, sparsely ls the fiae ] near the leaf base, many guard cells darkened and

often recessed; upper lamina surface dull brown, slightly paler than the lower surface, smooth, glabrous.

Venation camptodromous to eucamptodromous; midrib subprominent below, + rounded, sparsely strigil-

lose, the epidermis longitudinally wrinkled, somewhat raised above with a narrow central canal, glabrous;

lateral veins (526—9(-11) per side, fine, but prominently raised below, flush or slightly raised above; 3°

veins fine, conspicuously raised below, barely raised above. Laminar extrafloral nectaries on abaxial

surface only, uncommon, mostly in the proximal half of the lamina, + round, ca. 0.25 mm wide, green to

black. Male inflorescences unknown. Female inflorescences not seen with flowers, apparently cymes,

1-3 in leaf axils of young stems, each 13-flowered, though no more than a single fruit seen on any inflores-

cence, axes subterete, moderately to densely strigillose, fruiting peduncle 0.5-2 mm long; fruiting pedicel

1.5-5 mm long, with a 4-angled pyramidal apex jointed to the receptacle within the basal protuberance of

the calyx, subtended by 23 lanceolate bracts, ca. 2mm long. Female flowers unknown, except the style

at least 2.4 mm long and 0.5 mm wide, hairy, the hairs off-white to reddish, up to 0.7 mm long, + straight,

+ terete, divided into two appressed, ascending branches at least 0.7 mm long (style details based on those

persisting on fruit). Fruiting calyx thick, but not fleshy, exterior moderately strigillose, the epidermis viscid,

tube patent, with a basal protuberance 1.3-2 mm long, 6-7.5 mm wide between opposing sinuses, interior

with numerous minute, reddish hairs, lobes 4—5, reflexed, roundish, the margins weakly revolute, 4—6 mm

long, 4-8 mm wide, interior glabrescent, viscid. Fruit ovoid-ellipsoid, often asymmetric, (15-)18-24 mm

long, (11213-15(-17) mm in diameter; hairy near the apex, sometimes with hairs similar to those of the

style scattered over the distal third, locules unknown, probably 1-4; flesh unknown; epidermis + smooth,

yellow to orange in life when mature, brownish yellow in herbarium material; seeds not observed, loosely

contained (audibly so) in the locules of dried specimens.

Two of the paratypes (A. Campos V. 1017, MO and A. Campos V. & R. Torres C. 825, MO) were originally

identified as D. campechiana, but annotated by R. Durán “No es Diospyros campechiana” in 1995. These

specimens were later identified as D. campechiana by B. Wallnófer in 1999. While this species is clearly a

n IC J , Di he I 95

close relative of D. campechiana, that taxon is found primarily in lowland tropical forests associated with

wetlands and riparian areas of major river systems between sea-level and 270 m elevation in the states of

Tabasco, Campeche, Chiapas, and Veracruz, Mexico, and Guatemala, while D. camposii has been found only

in Oaxaca in cloud forest between 1200 and 1550 m elevation. There is approximately a 300 km disjunc-

tion from the nearest populations of D. campechiana in the coastal lowlands of Veracruz to the mountains

in Oaxaca where D. camposii occurs.

Diospyros camposii can be separated from D. campechiana by its shorter, widely ovate to elliptic lamina

with a minutely ridged, but not at all revolute, margin. Although sometimes elliptic, the lamina of Diospyros

campechiana is typically longer and more oblong in outline, and the margin is distinctly revolute. While both

species are strigillose, D. camposii lacks the additional hirtellous i always present in D. campechiana.

Diospyros camposii also has a larger, thicker, fruiting calyx, with reflexed sepal lobes, in contrast to the thinner

calyx with spreading lobes of D. campechiana. Finally, the fruits of D. camposii are longer and have a hairy

apex, with the pubescence extending onto the style. The fruits and style of D. campechiana are glabrous.

Distribution and ecology.—This species is currently known only from a small area on the Pacific Slope

of the Sierra Madre del Sur (Fig. 5), where it is reported to occur in cloud forest (bosque mesófilo) in asso-

ciation with Alnus acuminata, Arbutus xalapensis, Chiranothodendron pentadactylon, Clethra mexicana, Cornus

disciflora, Dendropanax arboreus, Ficus pertusa, Inga eriocarpa, Ocotea helicterifolia, Oreopanax xalapensis, Or-

mosia oaxacana, Parathesis brevipes, Picramnia lindeniana, Quercus candicans, Siparuna andina, Ulmus mexicana,

Viburnum elatum and Zinowiewia integerrima (Campos-Villanueva & Villaseñor 1995). The area has a large

number of canyons with permanent streams, and soils consisting of a deep layer of organic matter over red

clay (Campos-Villanueva and Villaseñor 1995). The holotype was collected in pine-oak forest.

Etymology.—This species is named in honor of the Mexican botanist Alvaro Campos-Villanueva, who

collected the holotype, and participated in all known collections of the species. He is also an author of a

flora of the Municipio de San Jerónimo Coatlán in the Sierra Madre del Sur.

Specimens examined. MEXICO. Oaxaca. Mpio. San Jerónimo Coatlán: 41 km al SW de San Jerónimo Coatlán, cañada de bosque

dp suelo amarillo dea 16? 10'N, 96? 58'W, pid m, 17 Jan 1988, A. Campos V. 1017 (MO); 12.3 km al N de Piedra Larga,

E de Piedral arga, carr. a Miahuatlán, | 5fil bosque de

pino, en cañada, 16? 09! 00'N, 97° 01' 00"W, 1200 m, 16 Dec 1987, R. Torres C. & A. Campos 10895 (MO). Mpio. Piedra Larga: 12.5

km al NE de Piedra Larga, sobre el camino a El Progreso, bosque mesófilo, suelo negro, 16? 10'N, 97? 01'W, 1300 m, 15 Dec 1987, A.

Campos V. & R. Torres C. 825 (MO).

Diospyros crotalaria M.C. Provance & A. E nae qur nov. (Figs: 1b, 1d, 6, 15a). Tree: COSTA RICA. Puntar-

ENAS. Cantón de Osa: R.E Golfo Dulce, Pení uemado, Sector Oeste, 08°, 41' 10"N, 83° 35' 10"W, 400 m, 9

Sep 1992, J. Marín 512 (HoLorYrE: MO-5316690!; ISOTYPES: “CR K!).

ous D. d S. Knapp a sed e ind bip MD d e oen LERNEN stylibus glabratis, gracilioribus,

Trees 10-20 m tall, eats evergreen; trunk recorded up to 15 cm dbh, bark reportedly thick; young

stems reddish brown, somewhat compressed, shallowly channeled, or with some minor furrows, glabres-

cent to sparsely strigillose, the hairs pale yellow to dark red or nearly black; mature stems terete, rugose,

densely lenticellate. Leaves alternate, simple, entire; petioles 4—9(-10) mm long, subterete, epidermis often

scintillant, atropurpureous, rugulose, the wrinkles often transverse, sometimes developing into fissures;

glabrous to sparsely strigillose, the hairs slightly flexuose, ascending, and pale yellow, or straight, appressed,

and dark red to nearly black; shallowly V-grooved above, the groove sometimes with an additional narrow

channel down its center, glabrous below, sometimes with a few small longitudinally oriented lenticels;

lamina subcoriaceous, oblong to oblong-obovate, sometimes narrowly ovate, 74-150 mm long, 26-49

mm wide, length to width ratio ca. 2.53 : 1, base acute to subacute, margin flat, apex acuminate with a

rounded tip; lower lamina surface reddish brown, slightly darker than above, dull, glabrescent or with

scattered minute, 0.2-0.5 mm long, straight, appressed, dark red to nearly black, sometimes pale yellow

hairs, epidermis sometimes with aggregates of opaque light-colored excrescences, guard cells often very

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dark and recessed, upper i f; ish green, not glossy, though slightly more shiny than below,

glabrous, epidermis densely papillose, the EM angular. Venation eucamptodromous; midrib narrow

but prominently raised below, rounded, longitudinally wrinkled, glabrate to sparsely strigillose; shallowly

V-grooved above, glabrous; lateral veins 6—9(-10) per side, fine, raised below, forming inconspicuous loops

t laterals, flush or slightly depressed above, papillae absent; 3? veins fine, inconspicuously

raised Deu Avperceotible above, or barely so, and then depressed. Laminar extrafloral nectaries on the

abaxial surface only, relatively common, round to elliptic, 0.2-0.5 mm wide, black. Male inflorescences

unknown. Female inflorescences cymes, 1 per leaf axil on young stems, 1-2-flowered, producing 1-2

fruit, axes « terete, densely strigillose, the hairs flexuose, pale yellow to dark reddish brown, subtended by

2-3 opposite pairs of strigillose, persistent, deltoid bracts; fruiting peduncle 0.5-1.5 mm long; fruiting

pedicel 1-2 mm long, with a 1 mm long 4-angled apex that forms a joint within the basal protuberance of

the calyx. Female flowers unknown, except styles (2) erect, divided + to the base, 4-angled, at least 0.9 mm

long, 0.2 mm wide, essentially glabrous, though a long solitary hair was seen in the style branch fork on two

of the known fruit (style details based on remnants from fruit). Fruiting calyx relatively thin (not fleshy),

reportedly green in life, drying very dark, exterior strigillose, the hairs pale yellow to dark red, epidermis

Vee tube Pa with a ca. 1.5 mm long basal protuberance, 5-6 mm wide between opposing sinuses,

with inute, reddish hairs, lobes 4-5, usually reflexed, ovate, 2.4-3 mm long,

3-4 mm wide, the margins flat to weakly tE with a few cilia near the sinuses, moderately pubescent

inside, the hairs minute, appressed, pale yellow to dark red. Fruit glabrous, 13-19 mm long, 15-20 mm

wide, depressed globose to subglobose, often appearing 2-4 lobed (when dry) due to conformation of the

fruit wall to the shape of the (1-4) locules with developing ovules; flesh exiguous; epidermis smooth, yellow

in life when nearly mature, drying orangish-brown, red when mature, drying dark reddish brown; seeds

1-4, brown, loosely contained in the fruit, + ellipsoid, 9-11.5 mm long, with 1 or 2 flat surfaces, ca. 6 mm

wide, dorsal surface broadly convex, with a prominent vascular strand, texture rugulose-foveolate.

Specimens examined. COSTA RICA. PUNTARENAS. Cantónde Osa: R.F. Golfo Dulce, Península de Osa, Cerro Chocuaco, camino el

98 tani i Texas 3(

faro, 08? 43' 20"N, dod e oe m, 28 Dec 1993, R. i eal i P Ki, A Cantón de Golfito Jiménez: Dos Brazos de Río

Tigre, C N erro Rincón bradi followi g 1 I hl I 11

creeks), 08? 30' 35"N, 83? 28' 15"W, 782 m, 25 Nov 1990, G. Herrera 4647 (CR, K!, MOD.

The specimens of D. crotalaria we examined had not previously been determined to species. Diospyros crota-

laria is similar to D. hartmanniana, but can be separated (Table 1) by its oblong lamina, glabrous fruits, and

more slender, glabrous, erect styles, which are divided nearly to the base. Furthermore, D. crotalaria has a

thinner calyx, and smaller, reflexed calyx lobes.

Distribution and ecology.—Apparently endemic to hills and canyons with wet tropical evergreen forest

between 400 and 800 m elevation on the Osa Peninsula on the coast of southwestern Costa Rica (Fig. 14).

Reports of D. hartmanniana from the Osa Peninsula (e.g., Zamora et al. 2004) may have been in reference to

this species. The closest population of D. hartmanniana to the Osa Peninsula, that we know of, is about 75

km southeast in Panama.

Etymology.—This species i 1 for the rattling noise of the seeds in the locules of fruit on herbarium

specimens.

Diospyros haberi M.C. Provance & A.C. MR sp. nov. (Figs. X deis Tere: COSTA RICA. AtAjuErA. Cantón de

Upala: P. N. Rincón de la Vieja, Cordillera d 7 km de la casa de Administración, 10? 47' 50"N,

85? 18' 19"W, 1500 m, 6 Jul 1991, G. Rivera 1422 (HoLoTYPE: MO}; ISOTYPES: im

Arbor D. hartmanniana S IDE siu 1 lobis caly daxialles prope glabratis, 1.5— mm aces ae oe aia oe

ES TA LU E Js

t T

BLISEO

Trees 8-28 m tall, BN evergreen; trunk up to 60 cm dbh, the bark reportedly black with white

patches; young stems 4—5 angled, somewhat compressed and channeled, sulcate, shiny, minutely black

gland-dotted, sparsely strigillose; mature stems terete, glabrate, the epidermis light to dark green, smooth

to half-netted, eventually lenticellate. Leaves alternate, simple, entire; petioles 3.5-9 mm long, + terete,

epidermis light green to castaneous or atropurpureous, shiny, sometimes transversely fissured, rugulose

below, glabrous to sparsely strigillose, rounded ee idees and sparsely strigillose, channeled, often

deeply, glabrous within or with scattered black hairs; i , elliptic, 30-96 mm long, 17-40

mm wide, length to width ratio ca. 1.8—2.7 : 1, base acute to attenuate, margin flat, though thickened, apex

acute to obtuse, rarely slightly acuminate; lower lamina surface greenish to grayish brown, glabrate to

sparsely strigillose, guard cells often dark, the stomatal apparatus often recessed; upper lamina surface

greenish brown, shinier and slightly lighter than the lower surface, glabrate, epidermis markedly papillose,

the papillae angular. Venation Doe ATEAN to aid midrib narrow below, subpromi-

nent, glabrate to sparsely strigillose, epid ften darkened, 1 lly wrinkled, shallowly V-grooved

above, glabrous or with some scattered dark appressed hairs; lateral veins 6-9 per side, fine, raised below,

barely raised above; 3? veins slightly raised below, inconspicuous above. Laminar extrafloral nectaries

on the abaxial surface only, usually near the midrib and towards the base, round to elliptic, 0.1-0.3 mm

wide, dark green to dark brown. Male inflorescences solitary compact cymes in leaf axils of young stems,

(1323 -flowered, 6 mm long, axes 4-angled, strigillose, subtended by 1-2 pairs of deltate, 0.7 mm long, 0.6

mm wide bracts; peduncles 0.3 mm long; pedicels 0.3-0.8 mm long. Male flowering calyx crateriform

to subrotate, exterior drying dark greenish brown, 3 mm wide, tube 1.5 mm long, exterior sparsely black

strigillose, interior unknown, lobes 4, widely ovate, 0.5 mm long, 1-1.3 mm wide, erect, exterior glabrate

to sparsely black strigillose, margins densely ciliate, interior glabrous; male corolla white in life, drying

dark brown, glabrous, tube urceolate, 2 mm long, 2.6 mm wide, lobes 4, quadrate, 1.4 mm long, ca. 1.4

mm wide; only three flowers were seen, none were dissected, thus stamen morphology is unknown, except

that anthers have a slender, tapering connective. Female inflorescences solitary, subsessile flowers in

leaf axils of young stems, or rarely 2-flowered compact cymes, but no more than a single fruit seen in an

inflorescence; axes 4-angled, sparsely strigillose, with 1—2 pairs of persistent, ovate bracts, 0.8 mm long,

1 mm wide; fruiting peduncles 0—0.5 mm long; fruiting pedicels ca. 0.5 mm long, with a 0.3 mm long

URI

BOTANICAL GARDEN

PESBARIL :

MECO MENE

NO 5600

Fic. 7 Diospyros haberi. A. 1

£. H. L

flower bud (G. Rivera 1422, K). D.

(W. Haber & S. Daniel 9895, MO). B. B

(W. Haber & S. Daniel 9895, MO)

). C. Female

100 Journal of the Botanical R h Institute of Texas 3(1)

TABLE 1 MA pt | E | E y d £ my L L ] f, n J n.

D. panamense D. hartmanniana D. crotalaria D. haberi

Fruiting calyx thick thick thin thick

Fruiting calyx tube width 7-10 mm 5-6 mm 5-6 mm 5.1-6.6 mm

(distance between

opposing sinuses)

Fruiting calyx lobe length 2./-5 mm 2./-3.8 mm 24-3 mm 1.5-2 mm

Fruiting calyx lobe width 4.5-9 mm (4-)4.4-5.5 mm 3-4mm 3.8-5.3 mm

Fruiting calyx lobes spreading to t spreading reflexed + spreading

slightly reflexed

Fruit length up to 33 mm (14-)17-20 mm 13-19 mm 14-18 mm

Fruit vestiture glabrous when usually hairy, glabrous glabrous when

mature, hairy at

apex and base

when immature

especially at the

base and apex

mature, hairy at

apex immature

Style 1, hairy, minutely 1, hairy, divided 2, + glabrous, divided 1(-2), glabrous,

deltoid-bifid nearly to the base, to the base, parallel shortly deltoid-bifid,

spreading in the and erect n cleft on

form of a 'V' e side

Stem, petiole and midrib minutely curved- not curved-acicular not curved-acicular not curved-acicular

acicular pubescent pubescent pubescent pubescent

Lamina shape oblong-elliptic to elliptic, apex oblong to oblong- elliptic, apex

elliptic, apex minate obovate, sometimes rounded

acuminate narrowly ovate,

apex acuminate

Lamina size (96-)127-240 mm (50-)64-163 mm 74-150 mm long, 30-96 mm long

long, 45-107 long, 25-70 26-49 mm wide 17-40 mm wide

wide mm wide

4-angled pyramidal forming a joint within the basal protuberance of the calyx. Female flowers known

from a single flower na and style remnants on immature fruit. Female flowering calyx subcampanulate

prior to anthesis, sparsely strigillose, 6.3 mm wide, tube 2.5 mm long, lobes (3-34, thick (ca. 0.8 mm at

1/4 length), + evenly tapered to the apex, depressed-orbicular, 1.2-1.8 mm long, 3.9-4.1 mm wide, some-

times emarginate, asymmetric (hooking left viewed abaxially), ciliate on the left margin; female flowering

corolla drying dark brown, tube ca. 2.2 mm long, with some scattered dark two-armed hairs, lobes 4, ca.

1.3 mm long, shape and width indiscernible, with some indistinct papillae; pistil unknown; styles 122),

0.8-1.2 mm long, 0.7-1 mm wide, angular, bifid, the branches short, 0.2-0.3 mm long, deltoid, style often

grooved on one side, rarely notched apically or grooved on both sides, glabrous, though strigillose near

the base on the apex of the fruit (style and stigma [see below] details based on those persisting on fruit);

stigmas depressed orbicular, 0.3 mm long, 0.7 mm wide; staminodes unknown. Fruiting calyx barely

accrescent, thick, probably fleshy and green in life, drying brown to black, exterior sparsely strigillose, tube

patent, with a basal protuberance 1.2-1.5 mm long, 5.1-6.6 mm wide between opposing sinuses, interior

glabrous, or with some scattered reddish hairs, lobes + spreading, depressed orbicular, 1.5-2 mm long,

3.8—5.3 mm wide, margins flat to slightly reflexed, ciliate, glabrous inside. Fruit 14-18 mm long, 10-20

mm wide, depressed globose to subglobose, often appearing 2-4 lobed when dry due to conformation of

the fruit wall to the (1-4) locules with developing ovules, glabrous except for some dark hairs at the apex

in immature fruit; flesh sparse, reddish, translucent and vitreous when dry; epidermis green in life when

immature, becoming orange to red when mature, delicate, golden brown to dark reddish brown and loosely

wrinkled when dry; seeds 1—4 (based on fruit lobes), reddish brown, loosely contained in the fruit, probably

n AC A , Di h; l 101

wedge-shaped, 14-18 mm long (width indeterminable, and shape uncertain, as the only seed observed had

been sectioned longitudinally), surface texture rugulose in a maze-like pattern.

Distribution and ecology.—This species is associated with moist and windswept forests between 1100 and

1700 m elevation on the Pacific slopes of the Cordilleras de Guanacaste, Tilarán, and Talamanca, in Costa

Rica (Fig. 14). Populations occur near, but as far as known do not overlap, populations of D. hartmanniana

in the former two mountain ranges. All of the specimens examined had been recently identified as D. hart-

manniana, to which it is doubtless closely related. The most obvious differences between the species (Table

1.) involve leaf apex shape, and style number and morphology. In addition to these characters, the interior

of the sepal lobes in male and female flowers of D. haberi are glabrous, while in D. hartmanniana they are

densely dark strigillose. The color and brilliance of the lamina is also different in these species. In D. haberi

the greenish brown upper surface of the lamina is shinier and slightly lighter than the greenish to grayish

brown lower surface. Brilliance is reversed in D. hartmanniana, where the grayish brown upper surface is

duller than the reddish brown lower surface.

Etymology.—This species is named in honor of William A. Haber, a biologist who has

to our knowledge of the biota of Costa Rica and who has documented the presence of this new species at

Monteverde.

tributed greatly

O ya

Specimens examined. COSTA RICA. GuANAcAsTE. Cantón de Tilarán: Sa Río Cañas, indes Pacifico, 10? ut 84? d 1200 m,

25 Aug 1989, E. Bello 1124 (MO). PUNTARENAS. C g 217

40"N, 83? 11' 50"W, 1700 m, 24 Sep 1989, G. Herrera 3542 (MO). Cantón de Osa: San Luis, Monteverde Buen Amigo, 10? 16' 00'N, 84?

49' 00"W, 1100 m, 10 Nov 1993, Z. Fuentes & E. Fuentes 565 (MO). Cantón de Puntarenas: Monteverde, Bajo Tigre Reserve, 10? 18'N,

84° 48'W, 1200-1300 m, 3 Apr 1991, W. Haber & W. Zuchowski 10600 (MO); Monteverde, above Quebrada Máquina, along Fonseca,

Hotel de Montaña and Savage Farms, 10? 18'N, 84? 48'W, 1100-1300 m, 3 Jun 1990, W. Haber & S. Daniel 9895 (MO).

Diospyros hartmanniana S. Knapp, Novon 7:256. 1997. (Figs. 8, 9, 15e). Ter: PANAMA. Chiriquí: near Coast

Rican border, ca. 13 road-km from Río Sereno, Finca Hartmann, 1550-1750 m, 08? 50'N, 82? 45'W, 23 Oct 1992, G. McPherson &

PM. Richardson 15959 (HoLotyee: BM; isotypes: MO, P

Trees 7—20 m tall, rarely a 1.5 m tall shrub, apparently evergreen; trunk up to 72 cm dbh, bark smooth and

black, wood oxidizing bright yellow; young stems angular, often compressed, sulcate to channeled, shiny,

often minutely black gland-dotted, glabrous to sparsely strigillose; mature stems dark, terete, smooth,

glabrate to sparsely hairy, later half-netted, scaly and lenticellate. Leaves alternate, simple, entire; peti-

oles 4—10 mm long, subterete, epidermis castaneous to atropurpureous, sometimes transversely fissured,

sometimes scintillant, longitudinally wrinkled to colliculate below, shiny, glabrous to strigillose; glabrous

above, markedly channeled; lamina subcoriaceous, elliptic, (50-)64-163 mm long, 25-70 mm wide,

length to width ratio ca. 2-3 : 1, base acute to attenuate, margin flat, though slightly thickened, apex acute

to subacute, rarely obtuse, acuminate, the tip rounded; lower lamina surface reddish brown, glabrate to

sparsely strigillose, aggregates of opaque off-white excrescences sometimes present, guard cells often dark

and the stomatal apparatus often recessed; upper lamina surface grayish brown, duller than the lower

surface, glabrous, epidermis conspicuously papillose, the papillae angular. Venation brochidodromous to

eucamptodromous; midrib narrow below, though prominently raised, rounded, longitudinally wrinkled,

glabrous to strigillose, the hairs pale or dark; V-grooved above, glabrate; lateral veins 6—11 per side, fine,

raised below, slightly raised to barely depressed above, lacking papillae; 3° veins slightly raised below,

indiscernible above. Laminar extrafloral nectaries on the abaxial surface only, common near the midrib

and the base of the lamina, 0.1-0.5 mm wide, castaneous. Male inflorescences solitary compact cymes in

the leaf axils of young stems, or several near the base of young stems, (12)3—7-flowered, 7-11 mm long, axes

4-angled, densely reddish brown to black strigillose, subtended by several lanceolate to widely ovate, 1-2 mm

long, ca. 1 mm wide bracts; peduncles 0.5-1.5 mm long; pedicels 0.5-1.5 mm long. Male flowers known

only from buds. Male flowering calyx subrotate, with a basal protuberance ca. 1 mm long, exterior drying

dark greenish brown, moderately strigillose, interior densely strigillose, 3.5-4.1 mm in widest dimension,

tube 1-1.2 mm long, lobes 4-5, ovate, 0.9—1.2 mm long, 1.3-1.6 mm wide, suberect to spreading, margins

ciliate; male corolla white to cream in life, drying dark brown, thick, glabrous, papillate, tube urceolate,

102

J I| £ 4.

JOUTM al OF

TERR. HORT, KEW,

Fic. 8. D

(G. Herrera 5669, K) B. Fruiti

(E Bello 1767, MO). C. Im-

mature fruits (£. Bello 2442, MO).

103

Fic. 9. Di D)

IR Ecninnza etal 224 MM A Male fl

T ,

bud. C, Stamens

£ el Dat tai D khi Fr £ T.

104 Journal of Texas 3(

1.6-2.8 mm long, 3.2-4.5 mm wide, lobes 4—5, widely ovate, 2.1-3.1 mm long, ca. 3 mm wide; stamens

ca. 27 (based on R. Espinoza et al. 884 (MO) from Costa Rica) adnate to the base of the corolla, the filaments

fused most of their length into ca. 13 pairs, comprised of inner and outer members (Knapp [1997] reports

10-12(20) stamens, and illustrates 14 stamens in pairs based on McPherson & Hensold 15303 (MO) from

Panama), anthers cream, the inner anthers often larger than the outer, lanceolate, (2.5)3—3.2 mm long (in-

cluding connective), golden to reddish sericeous along the connective on both sides, the apex (connective)

rostrate, slightly constricted basally, minutely muricate, filaments 1.2-1.5 mm long (fused portion of the

filament pair), glabrate, free portion of the filaments 0.5—0.7 mm long, densely hairy, the hairs minute and

curly; pistillode obturbinate, 1.4 mm high, 1.2 mm wide, ca. 8-lobed/grooved, the ‘lobes’ narrower than the

‘grooves’, presence of style-like structures indeterminable due to a dense vestiture of 0.5 mm long, red, flat,

twisting hairs in the upper half. Female inflorescences unknown with flowers, apparently a 1-3-flowered

compact cyme in leaf axils of young stems, with 1-2 fruit developing per inflorescence, axes + 4-angled,

strigillose, with ca. 2 pairs of persistent, widely ovate bracts, 1.2-1.8 mm long, ca. 1.5 mm wide; fruiting

peduncles 0—1.8 mm long; fruiting pedicels 0.5-2 mm long, with a 1-1.5 mm long 4-angled pyramidal apex,

the distal 1/2 forming a joint within the basal protuberance of the calyx. Female flowers unknown, except

style divided nearly to the base, branches angled away from one another in the form of a ‘V’, each 0.7-1.3

mm long, ca. 0.5 mm wide, densely reddish brown strigillose basally and where the style branches diverge

(style and stigma [see below] details based on those persisting on fruit), stigmas minute, deltoid. Fruiting

calyx thick, probably fleshy, reportedly brown in life, drying dark brown to black, exterior strigillose, the

hairs pale yellow to dark, sometimes glabrescent, epidermis viscid, interior Oe strigillose, tube patent,

with a basal protuberance ca. 1.5 mm long, 5-6 mm wide between opt , lobes 4—5, spreading,

ovate, 2.73.8 mm long, (424.4—5.5 mm wide, the margins flat, n ciliate. Fruit (1417-20 mm

long, (12-)17-20 mm wide, depressed globose to subglobose, often appearing 2-4 lobed when dry due to

conformation of the fruit wall to the (1—4) locules with developing ovules, usually strigillose basally and at

the apex; flesh sparse, reddish; epidermis smooth to wrinkled, in life orange to orangish red when mature,

light brown to dark reddish brown when dry; seeds 1-4, loosely contained in the fruit, + wedge-shaped,

reddish brown and rugulose-foveolate textured, though only immature seeds were seen.

Distribution and ecology—This species is associated with cloud and moist forests between 500 and

2200 m elevation, on slopes of the Cordilleras de Guanacaste and Tilarán in Costa Rica, and Talamanca (La

Amistad) in Panama (Fig. 14). Populations occur near, but as far as known do not overlap, populations of

D. haberi in the former two mountain ranges.

Collections examined. COSTA RICA. ALAJUELA. Cantón de San Carlos: La Fortuna, Finca El Jilguero, 10? 26' 35"N, 84? 41' 25"W, 1140

m, 27 Nov 1992, G. Herrera 5669 (K). Cantón de San R R iol. Monteverde, Valle del Río Peñas Blancas, Fila de Toro, Sendero

Pipilacha y Capo Tres, [acca. W3 Tropicos 10? 18' 00"N, 84° 43' mw 900—1100 m, 21 Jan 1991, W. Haber (ex E. Cruz) 10632 (MO);

Res. Biol. iuri a Eladio’s, 10? 19'N, 84? 43'W, 820 m, 2 Oct 1990, E. Bello 2442 (MO). Cantón de Upala: P.N. Rincón

de la Vieja dill te, sendero a la Quebrada Mora, camino a la Colonia Blanca, 10° 46' 32"N, 85° 15' 10"W, 840 m, 30

Nov 1990, G Rivera 920 (K, MO); P.N. Guanacaste, Cordill Estación, San Ramón, Dos Ríos, Sector La Campana, 10?

52! 50"N, 85? 24! 05"W, 550 m, 1 May 1993, R. Espinoza et al. 884 (K, MO); GUANACASTE. Cantón de La Cruz: De Bahia Salinas a Santa

Cecilia Hda. El Oro, 10? 59' 26"N, 85° 25' 40"W, 500 m, 5 Sep 1922, R. Espinoza 531 (K, MO); Puntarenas. Cantón de Coto Brus: Zona

Protectora Las Tablas, Cuenca Térraba-Sierpe, 08? 58' abe 82° 50' 14"W, 1500-2000 m, 1 Nov 1996, A. Rojas & E. Navarro & E. Alfaro

1757 (M m de Osa: Reserva Biológica M brada Veracruz, Finca Pablo Morales, 10° 15'N, 84° 48'W, 1600 m, 11

Jan 1990, E. ay 1767 (MO); San Luis, Monteverde, Cerro nude camino a Surtubal, 10? 15' 25"N, 84° 47' 20"W, 1100-1200 m, 22

Mar 1994, Z. Fuentes 704 (MO). PANAMA. Chiriquí. Near border with Costa Rica, ca. 13 road km from Río Sereno, Finca Hartmann,

08? 50'N, 82? 45'W, 1400—1800 m, 12 May 1991, G. McPherson & N. Hensold 15310 (B, 2 sheets, CAS); ridges leading to Cerro Pelota,

Parque Nacional Amistad, 08° 52-53'N, 82° 44'W, 1700-2200 m, 21 Aug 2000, 5. Knapp & A. Monro 9254 (MO)

€

Diospyros juruensis A.C. Smith, Brittonia. 2:163. 1936. (Figs. 10, 11). ‘Tere: BRAZIL. Amazonas: on varzea land,

r mouth of Rio Embira (tributary of Rio Tarauaca), 7? 30'S, 70? 15'W, 26 Jun 1933, B.A. Krukoff 5003 (HoLotYrE: NY; ISOTYPES:

E MICH, MOD.

A lengthy description here of the species based on the single sheet with male flowers we have seen seems

premature to us. The characters that lead us to believe that D. juruensis belongs to the D. campechiana complex

105

1 Al ype Specimen

QJ iJ / HERA, MLC

P e =—

P. A. EEUKOTT'S Bh EXPEEFTION Te AAN Ad

BAKIN GF MM: JURMUA

ud P due. “E

amen p Á É t Loe

7: E ato av fd.

Embira (tribulary of

: near mouth af Rio

T^ 2908 ;Jongs te th’ W.

as York aries! Gardin

varo

Ris caca,

Fic. 10. The isot

is (B.A, Krukoff 5003, MO). A. Fruiti

106 J i i Texas 3(

Fic. 11. The isotype of Diospyros juruensis (B.A. Krukoff 5003, MO). A. Pistillodes. B. Stamens.

include the large oblong-oblanceolate leaves with frequent darkened guard cells and somewhat recessed

stomatal apparatus; male inflorescence axes that are markedly 4-angled; depressed orbicular male calyx

lobes; a lobed pistillode (Fig. 11a) very similar to that seen in D. campechiana (Fig. 3b); stamens adnate to

the base of the corolla, fused near the base of their filaments into ca. 16 pairs, outer and inner stamens +

same length, densely golden sericeous on the filament and connective, anthers reddish orange, lanceolate,

the apex (connective) tapered and minutely muricate.

Diospyros panamense S. Knapp, Novon 9:524. 1999. (Figs. 12, 13, 15f). Ter: PANAMA. San Bras: Rio Diablo y

vecinidad de Duque Sui, a unos 10 km de la costa frente a la Isla de Nargana, ruta hacia Cerro Ibedón, 80-110 m, 09° 22'N, 78°

35'W, [1 Jul 1992 acc. to W3Tropicos], H. Herrera et al. 1175 (noLorvrE: BM; isorvees: MO, PMA) Replaced name: Dios

Knapp, Novon 7:258. 1997. Non Diospyros whitei Dows.-Lem. & Pannell, Bull. Jard. Bot. Nat. Belg. 65:399. 1966.

t holl OW,

Trees 5-20 m tall, evergreen; trunk up to 40 cm dbh, bark black, wood reportedly soft; y

distally angular, + compressed and shallowly channeled, terete and smooth proximally, dark. green, brown

or gray, occasionally sparsely strigillose, the hairs dark, or sometimes yellowish, minutely pubescent, the

hairs colorless, curved-acicular, ca. 0.03 mm long; mature stems with thick, scaly, gray to dark brown bark,

lenticels prominent, stemwood reddish brown in stems ca. 10 mm wide. Leaves alternate, simple, entire;

petioles 7-14 mm long, subterete, minutely curved-acicular pubescent, epidermis green or atropurpureous,

sometimes glaucous and scintillant, sometimes with a few transverse fissures, rounded below, glabrate to

sparsely strigillose, + flattened above, sparsely strigillose, with a densely glandular narrow central channel;

lamina subcoriaceous to coriaceous, oblong-elliptic to elliptic, though slightly wider in the upper half,

(96-)127-240 mm long, 45-107 mm wide, length to width ratio ca. 22.6 : 1, base acute to obtuse, sometimes

abruptly decurrent on the petiole, margin flat to revolute, apex acuminate with a rounded tip; lower lamina

surface mu pou PR to sparsely strigillose, faintly papillose, guard cells sometimes dark and

recessed; upp ish to greenish brown, markedly duller than the | , glabrous

to use TE near pb epidermis M a bind papillose, the papillae angular. Venation

(sensu Knapp 1997), minutely curved-acicular pubescent below; midrib

pionnen below, rande, glabrate to sparsely strigillose, longitudinally wrinkled, concave above, sparsely

to moderately strigillose and glandular; lateral veins + evenly spaced, 6-9 per side, fine below, though

prominently raised, barely depressed above, the surface smooth, and so, conspicuous adjacent to the papil-

late surface of the lamina; 3° veins fine below, though prominently raised, scarcely, if at all apparent above.

Laminar extrafloral nectaries on the abaxial surface only, occasional, more frequent near the midrib,

dark brown to black, ca. 0.6 mm wide, minutely rimmed. Male inflorescences glomerules, or compact

cymes, (3-)10-20-flowered, a few at the base of young stems, or solitary in leaf axils, axes 4-angled, dark

reddish brown strigillose, subtended by + 2 strigillose triangular bracts, ca. 0.9 mm long; peduncles 0.1-1

#ISSOURI

BOTANICAL GARDEN

HERBARIU?

il NIE de UE

BAIRD

) d

E Bii

Fic. A. Immature fruit (E. Alfaro 291, MO). B. Fruiting t hlets (R. Aguilar et al. 2727, MO). C. Male infl (R. Aguilar

2749, K). D. ‘Fruits (R. Aguilar et al, 2727, MO).

108 I Ll pnfthn DC 12a n MEER PR

JOUTMal OF

Fic. 13. Diospyros panamense (R. Aguilar 2749, MO). A. Detail of male infl ith fl buds from fi t folder. B. Interior of male cal

with pistillode. C. Stamens.

mm long; pedicels 0—1.4 mm long. Male flowers known only from buds; male fl i I brotat

with a short basal 1 , exterior drying dark brown to black, strigillose, interior acy strigillose,

3-4.2 mm in widest dimension, tube 1-1.5 mm long, lobes 4-5, depressed orbicular to ovate, 1-1.6 mm

long, 1.8-2.4 mm wide, suberect nearing anthesis; male corolla white in life, drying dark brown, thick,

glabrous, papillate, tube urceolate, 0.8-1.9 mm long, 2.6-3.2 mm wide, lobes 4-5, ovate, 1.4-2.2 mm long

1.6-1.7 mm wide; stamens 24-28, adnate to the base of the corolla, fused near the base of their filaments

into 12-14 pairs, outer stamens bowing inwards, outer and inner stamens + same length, the outer stamen

golden seri abaxially on the filament and connective, the hairs long and curved, also golden sericeous

D ne J , Di hs [| 109

on the connective adaxially and on both surfaces of the inner stamen, but the hairs shorter, anthers reddish

orange, lanceolate, ca. 2 mm long, the apex (connective) tapered and minutely muricate, filaments 0.2-0.4

mm long; pistillode obturbinate, markedly 12-14-lobed, + 1 mm wide, with 2-3 style-like apical struc-

tures, each with a few long straight golden hairs. Female inflorescences not seen with flowers, apparently

a solitary 1-3-flowered compact fasciculate cyme in the leaf axil of young stems, though no more than one

fruit per inflorescence observed, axes 4-angled, sparsely pale strigillose, subtended by several thin, deltoid,

strigillose, persistent bracts, ca. 1.5 mm long; fruiting peduncles 0-1 mm long; fruiting pedicels 1-4 mm

long, with a 1-1.2 mm long 4-angled apex, the distal 0.5 mm depressed-pyramidal, forming a joint with the

receptacle within the basal protuberance of the calyx. Female flowers unknown, except style 1.3-1.6 mm

long, 0.9-1.3 mm wide, hairy, the hairs pale, yellowish, slightly wavy, terete to slightly 4-angled, bifid, the

branches short, 0.2-0.3 mm long, deltoid (style and stigma [see below] details based on those persisting on

fruit), stigmas depressed orbicular, 0.3 mm long, 0.7 mm wide. Based on the holotype (which we have not

seen), Knapp (1997) reported, “styles 5, 1-1.5 mm, fused at the base” and “stigmas irregularly lobed.” Fruit-

ing calyx thick, possibly fleshy in life, drying dark brown to black, exterior sparsely to densely strigillose,

the hairs mostly pale, tube patent, y a short protuberance in immature fruit, this apparently absent at

maturity, 7-10 mm wide! j , interior densely golden strigillose, lobes 4-5, spreading

to slightly reflexed, depressed- M 2. 7—5 mm long, 4.5-9 mm wide, the margins sometimes ciliate,

interior sparsely golden strigillose. Fruit up to 33 mm long, 33 mm wide, globose to subglobose, immature

fruit densely hairy near the base and the apex, the remainder sparsely hairy, the hairs golden to dark red,

mature fruit glabrous, locules probably 1-4, small damaged fruits with + 2 locules observed, and Whitefoord

and Knapp (1998—onward) reporting “lóculos 4?”; flesh unknown in mature fruit; epidermis mature fruit

loosely wrinkled, cracked and scaly near the base and apically, the largest fruits seen (R. Aguilar 2727 et al.,

MO) reportedly brilliant green; seeds unknown. Fruits are suspected to contain up to 4 seeds, though fruits

in E. Alfaro 291 (MO), are probably 1 and 2-seeded based on the number of enlarged locules.

Distribution and ecology.—Lowland rain forests of Costa Rica, Panama, and possibly Columbia, between

80 and 1200 m elevation (Fig. 14). There are reports (Zamora et al. 2004) of this species from both near the

highest peaks (617—745 m) and in the low forests (e.g., 0-300 m) on the Osa peninsula. However, we have

not seen collections from either of these regions.

Specimens examined. COSTA RICA. PUNTARENAS. Canton de Osa: Fila Costeña, Río Piedras Blancas, junto a la casa, faldas Cerro An-

guciana, Fila Cruces. 08? 49' 02"N, 83° 11' 23"W, 900 m, 9 Dec 1993, R. Aguilar et al. 2727 (MO, K); Fila Costeña, Rio Piedras Blancas,

cerca de la casa, Cerro Anguciana, Fila Cruces, 08? 49! 02"N, 83° 11' 23"W, 900 m, 10 Dec 1993, R. Aguilar 2749 (MO, K). SAN Jose.

Cantón de Perez Zeledon: Cordillera de Talamanca, La Nubes, Santa Elena, 09? 23' 30"N, 83? 35' 50"W, 1210 m, 3 Aug 1995, E. Alfaro

291 (MO). PANAMA. Darien: Cerro Pirre, 4 Aug 1967, N. Briston 1229 (MO-2 sheets).

KEY TO THE MESOAMERICAN SPECIES OF THE DIOSPYROS CAMPECHIANA COMPLEX

1. Fruits mostly globose t | ts of Panama and Costa Rica.

2. Styles 2 or 1 style divided nearly to the base (Fig. 15).

3. Lamina oblong to oblong obovate; fruiting calyx thin (not fleshy), the lobes 3-4 mm wide, usually

reflexed; TUE apa aud. eam mm s erect, glabrous D. crotalaria

3. | ti | e lobes 3.8-5.5 mm wide, spreading; fruit apex dons styles

0.5-1 mm thick, diverging from near the base i in the form of a'V'to + erect, hairy or glabrou

4. Lamina apex acuminate, upper lamina surface duller and slightly darker than reddish EUM lower

P fruiting calyx lobes 2.7-3.8 mm lang ~ dark strigillose inside; style hairy, branches

diverging from near the base i i the iudi ofa D. hartmanniana

Lamina apex rounded, upper hinier and slightly lighter than the g hto ae

brown lower surface; unig calyx lobes 1.5-2 mm long, glabrous inside; style branches glabr

A DAT ideo)

+ erect ( p ypically has one style with some apical notches, or a groove down

D. haberi

2. Style 1 (Fig. 15)

5. Fruiting calyx lobe 2.7-5 mm long, 4.5-9 mm wide; fruiting calyx tube 7-10 mm from sinus to opposite

sinus; lamina 96-250 mm long, 45-107 mm wide, the apex acuminate; young stems, petioles, and

D. hartmanniana = O

D. panamense = © | |

D. crotalaria — e | |

D. haberi = x po

pum ppp

O 50 100

fet ia 2008 Sep 1223:52:38 OHC - Martin Weinatt

Fic. 14. Distribution of Di talaria, D. hartmanniana, D. haberi and D.

midribs with a sparse, minute, curved-acicular pubescence; fruits up to 32 mm long; style undivided,

i 5- | .6 mm long, 1-1.5 mm thic D. panamense

alyx lobe 1.5-2 mm long, 3.8-5.3 mm wide; fruiting calyx tube 5.1-6.6 mm from sinus to op-

im sinus; lamina 30-96 mm long, 17-40 mm wide, the apex rounded; young stems and midribs

without a minute curved-acicular pubescence; fruits 14-18 mm long D. haberi

. Fruits mostly ovoid-ellipsoid; plants of Mexico and Guatemala.

Stems, petioles, midribs (in addition to being dark api fruit

13-17 mm long, the apex glabrous; style 1-2.1 mm long, glabrous; i calyx thin, the tube 3

from sinus to opposite sinus, the lobes 2-4 mm long, not reflexed; lamina 100-285 mm long, 2n ong to

oblong-lanceolate to elliptic, ti oblong- -oblanceolate, the margins ome o D. campechiana

6. Stems, petioles, midribs i minutely ¡ it (15-)18-24

mm long, the apex hairy; style ca. 2.4 mm long, hairy; fruiting x thick, the tube 6- 7. 5 mm from sinus

to opposite sinus, the lobes 4-6 mm long, reflexed; lamina 55- 152 mm long, widely ovate to elliptic, the

margins minutely ridged . camposii

ACKNOWLEDGMENTS

We would like to express our appreciation to Martha L. Orozco-Cardenas at the UCR Plant Transformation

Research Center who kindly allowed access to microscopes a did equipment. We thank Lia

Mansfield for providing Latin translations. Ed Plummer g h appreciated technical support. We would

especially like to extend our gratitude to the following ebat for specimen loans: B, CAS, CHAPA, F, IJ,

K, MO, UCR, UMO, USF, WIS, XAL. Paulo T. Sano and an anonymous reviewer provided helpful comments

for improving the manuscript. Finally, we appreciate Giles Waines' continuing support and encouragement

of our taxonomic work on the Ebenaceae at the UCR Herbarium.

REFERENCES

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nc/omc. intro.html

NOVEDADES Y NOTAS MISCELÁNEAS EN LAS BROMELIACEAE DE MESOAMÉRICA

J. Francisco Morales

Instituto Nacional de Biodiversidad (INBio)

pto 22-3100

Santo Domingo de Heredia, COSTA RICA

RESUMEN

1,0 1 q [eae S |

Se describe Werauhia luctuosa (Bromeliaceae: Tillandsoideae)

con W. pittieri. Adicionalmente, se reporta la presencia de W. brunei y W. tanrtiddda en Panamá. B

de herbario, material EDS y taba de O se ne que el material de Mesoamérica previamente identificado como qr.

penduliflora y g América) tualmente A. angustifolia y G. blasii respectivamente.

n el ectiidi

PALABRAS CLAVE: Bromeliaceae, Honduras, Costa Rica, Panama, Aechmea, Werauhia

ABSTRACT

Werauhia luctuosa Sime peus NA a new species from Honduras is described and its ic affinities with W.

pittieri are discussed. Werauhia brunei and W. pua eie are pps for the ieu time in iran is on ME n of cui

Speed DB NA m field work, i i 1 ij

America), are actually A angustifolia and G. }

Key Wonps: Bromeliaceae, Honduras, Costa Rica, Panama, Aechmea, Werauhia

Como resultado de la binación del trabajo de campo y el estudio de colecciones de herbario tendientes al

desarrollo de una Guía de Campo para las Bromeliaceae del Sur de Centroamérica, se proponen la siguiente

novedad y notas taxonómicas misceláneas.

Aechmea penduliflora André, Enum. Bromél. 3. 1888. Tiro: COLOMBIA. SantanveR: Isla Brava, Río Magdalena, entre

ondo y Chucuri, 8 dic 1875, André 378 (noLoripo: K, GH-fotografía).

Distribución. —Colombia y Venezuela a Ecuador y Perú, donde crece en bosques muy húmedos entre los

50 y 700 m.

Aechmea penduliflora ha sido reportada para Nicaragua, Costa Rica y Panamá en diversas ocasiones

(Correa et al. 2004; Morales 2003; Utley & Burt-Utley 1994; Utley et al. 2001). Sin embargo, el estudio de

su tipo, así como de todos los especimenes citados como testigos en los tratamiento antes mencionados,

han revelado que las colecciones (e.g., Espinoza 94, INB, MO; McPherson 8533, MO; Rueda & Mendoza 17152,

MO) representan típicos especimenes de A. angustifolia Poepp. & Endl. (concordantes en todo sentido con

el tipo de este taxón) y que por lo tanto, A. penduliflora debe considerarse como una especie no presente aún

en Mesoamérica. Esta última especie algunas veces puede tener inflorescencias reducidas, en cuyo caso los

ejemplares pueden parecer diferentes de la típica variedad con inflorescencias relativamente largas y elon-

gadas, pero diferentes grados de variación pueden ser observados en el campo, así como en los especimenes

de herbario. Esta misma plasticidad en la morfología y tamaño de la inflorescencia está presente en otras

especies del genero en Mesoamerica (e.g., A. tillandsioides (Schult. f. ex Mart.) Baker).

Guzmania mitis L.B. Sm., Contr. Gray Herb. 98:31, t. 6, f. 4-5. 1932. Tiro: COLOMBIA. Norte DE SANTANDER: valle de

Pica-Pica, sobre Tapata, al N de Toledo, 1-5 mar 1927 (fl), Killip & Smith 20195 ( : GH; isoriro: US, INB-fotografía).

Distribución. —Colombia y Venezuela, donde crece en bosques muy húmedos y bosques nubosos entre los

1800 y 2800 m

En su revisión monográfica de las Bromeliaceae, Smith y Downs (1977) reportó esta especie para Costa

Rica, basado en un especimen recolectado por Foster en la región de Cartago, lo cual ha sido seguido en

forma subsecuente por diversos trabajos florísticos (e.g., Luther 1995; Morales 2003; Utley & Burtley-Utley

J. Bot. Res. Inst. Texas 3(1): 113 —116. 2009

114 tani i Texas 3(

1994). Ese especimen corresponde a una ejemplar relativamente pobre, con frutos maduros y carece de flores

o incluso brácteas florales completas. El pobre ue » ese especimen fue mencionado anteriormente por

Utley y Burtley-Utley (1994). Sin embargo, como resultad Be pd Ee oe especimen (Foster 2686, US),

asi como del ejemplar tipo de Guzmania mitis y otr de esa especie en herbarios de

Estados Unidos, Colombia y Venezuela, es claro que la eos colectada por Foster en Costa Rica, fue mal

identificada por Smith y Downs (1977) y que en realidad, corresponde a un ejemplar de Guzmania blassii

Rauh, un taxón similar, pero endémico a Costa Rica. Esta última especie se puede reconocer y separar de G.

mitis por sus hojas con las láminas con líneas moradas o rojizas longitudinales (vs. concoloras y sin líneas),

flores con pedicelos de 4—7 mm de largo (vs. sésiles a subsésiles), pétalos amarillos (vs. blancos) y rangos

altitudinales distintos, ya que usualmente G. blasii crece en elevaciones entre 700 y 1700 m, mientras que

G. mitis se encuentra entre los 1800 y 2800 m. Por lo tanto, G. mitis debe excluirse como un representante

de la flora Mesoamericana y considerarse con un taxón restringido al N de Suramérica.

Werauhia brunei (Mez & Wercklé) J. - Grant, D o pue 91:31. 1995. vriesea brunei Mez &

Wercklé, Bull. Herb. Boissier, ser. 2, 4(9):865. 1904. Tiro: C Alajuela): Candelaria,

oct 1902, Brune in Werckle Brom. Costar 45 ee B, M

Distribución.— Costa Rica y el O de Panamá, donde crece en bosques nubosos y robledales en elevaciones

de 1700-2750 m

Werauhia brunei pertenece a un complejo de especies que se caracterizan por sus inflorescencias

simples, las flores siempre con una bráctea adyacente, que Imente cubre gran parte de los sépalos y con

las brácteas variadamente imbricadas (Morales, 2003). Este complejo incluye entre otras a W. ampla (L.B.

Sm.) J.R. Grant, W. bicolor (L.B. Sm.) J.R. Grant, W. burgeri (L.B. Sm.) J.R. Grant, W. gladioliflora (H. Wendl.)

J.R. Grant, W. macrantha (Mez & Werckle) J.F. Morales, W. macrochlamys (Mez & Wercklé) J.F. Morales, W.

osaensis (J.F. Morales) J.F. Morales, W. tiquirensis (J.F. Morales) J.F. Morales y W. tonduziana (L.B. Sm.) J.R.

Grant, las cuales en general, son superficialmente similares y de difícil distinción (Morales, 1999, 2003).

Werauhia brunei se puede separar por sus brácteas florales usualmente dísticas antes y después de la antesis

(incluso con frutos maduros), lisas, verdes a verde-canela y de (4.9—)5.2-6 cm de largo. Para mayor infor-

mación, véase el trabajo de Morales (2003).

Especímenes examinados. PANAMÁ. Chiriquí: cerro Punta, falda NO, camino desde Finca Drácula hacia el último lodge dentro del

Parque, 10 ago 2006 (fD, Morales & Santamaría 14625 (INB

Werauhia camptoclada (Mez & Wercklé) J.E Morales, Monogr. Syst. Bot. Missouri Bot. Gard. 92:360. 2003.

Vriesea camptoclada Mez & Wercklé, Repert. Spec. Nov. Regni Veg. 14(400-404):247. 1916. Tipo: COSTA RICA. San José: en las

cercanías de La Palma, oct 1908, Wercklé in Inst. Costaric. 17292 (HoLotiro: B, INB-fotografía; isoro: GH).

Distribución. —Costa Rica y el O de Panamá, creciendo en bosques nubosos en elevaciones de 1500-2100

Werauhia camptoclada es una especie distintiva que se puede confundir con W. moralesii Luther, W.

kupperiana (Suess.) J.R. Grant y W. werckleana (Mez) J.R. Grant, pero que se puede reconocer por su roseta

densa y pequeña (en relación al tamaño de la inflorescencia), hojas que no exceden los 35 cm de largo e

inflorescencias con las ramas secundarias que raramente exceden los 12 cm de largo. Esta especie ha sido

considerada endémica a Costa Rica (e.g., Morales 2003) pero ahora se conoce en el O. de Panamá en las

estribaciones de la Cordillera de Talamanca.

PANAMÁ. eh cerro Punta, falda NO, camino desde Finca Drácula hacia el ültimo lodge dentro del

Pardue 10 ago 2006 (fl), Morales & S 14626 (INB).

Werauhia luctuosa J.F Morales, sp. nov. (Fig. 1). Tiro: HONDURAS. Santa BArBara: lago Yojoa, El Sauce, 9 abr 1951 (fl), L.

Williams & A. Molina 17704 (moLotipo: EAP, INB-fotografias).

A Werauhia pittieri (Mez) J.R. Grant, cui similis, scapo 36-37 cm longo (vs. 45-55 cm), sepalis 3.2-3.5 cm longis (vs. 1.7-2.8 cm) et

coma alba (vs. ferruginea) differt.

Plantas acaulescentes, con una altura en floración de 50 a 60 cm. Hojas rosuladas, más o menos erectas,

Morales, Nueva especie en Bromeliaceae

2cm

Ps -

4 "Pe

HONDURAS

Hentcme al

17704 Facurla Agri Canaria

: į >

Vriesia A jd ` T ra. V.

AN Gau

Rain forest area above

olot,

Bl Sauce

Departamento

Lows O. Wüll&ms

Sta. -arbara

Antonio Mola P

att 1000

April 9, 1951 na:

115

116 tani Insti Texas 3(

41-46 cm de longitud, densamente punteado-lepidotas en la superficie abaxial, el indumento escaso y más

esparcido en la superficie adaxial, Mich verdes p ambas superficies; bases de la lámina anchamente

elípticas, 9-11 x 4.5—6 cm, pálidas, usualment las láminas, densamente punteado-lepidotas

en la superficie abaxial; lámina de las hojas e abruptamente corto-acuminadas, 31-34.5 x 3.5-4.1

cm. Escapo erecto, 36-37 x 0.5-0.6 cm, diminutamente papilado; brácteas del escapo erectas, 4.3-4.6

cm de longitud, anchamente ovadas, mucho más largas que los internudos, densamente imbricadas, café

oscuro basalmente, algo más claras distalmente, sobretodo en la región del ápice, lisas. Inflorescencias

erectas, simples, 12-13 cm de longitud; brácteas florales anchamente ovadas, agudas, 3.5-3.8 x 2.9-3.3 cm,

relativamente delgadas, ligeramente imbricadas igualando los sépalos en longitud o ligeramente más cortas,

café oscuro, los bordes más claros, ecarinadas, lisas. Flores 5 a 6, con los pedicelos de 4-5 mm de longitud,

gruesos, erectos, esparcidamente papilados; sépalos 3.2-3.5 cm de longitud, erectos, elípticos, coriáceos,

café oscuro. Corola y cápsula desconocidas.

Distribución & habitat. —Endémica a Honduras, donde se conoce solo de la localidad tipo, en bosques

hümedos en elevaciones de 900 a 1000 m.

Werauhia luctuosa se encuentra cercanamente relacionada a W. pittieri (Mez) J.R. Grant, ya que ambas

especies comparten inflorescencias simples, con brácteas florales y flores unilaterales, café oscuro, densa-

mente imbricadas y de 2.6—3.9 cm de largo, pero la primera especie se puede separar por su escapo más

corto (36-37 cm vs. 45-55 cm), sépalos más largos (3.2-3.5 cm vs. 1.7-2.8 cm) y frutos con coma blanca

(vs. ferrugínea). Adicionalmente, W. pittieri es una especie predominante en bosques nubosos y robledales

sobre los 2000 m (raramente a 1850 m), mientras que W. luctuosa crece en bosques húmedos entre 900 y

1000 m de elevación. Werauhia luctuosa solo es conocida por la colección tipo y a pesar de la búsqueda en

otros herbarios no fue posible localizar colecciones adicionales. El epíteto de esta especie está dedicado a la

memoria de Rosenda Morales y María Ester Padilla, familiares (tías) fallecidos en forma reciente.

AGRADECIMIENTOS

Quiero agradecer a los siguientes herbarios por permitir el estudio de sus colecciones: BR, CR, EAP, F, K,

LAGU, MHES, MO, Y, P, U, US. Barry Hammel (MO) amablemente realizó el resumen en inglés. Asimismo,

la visita de algunas instituciones fue financiada por el proyecto “Compartiendo tecnología y desarrollando

capacidades para la gestión de la biodiversidad en Centroamérica" financiada por el Gobierno de Noruega

y ejecutado en forma parcial por el Instituto Nacional de Biodiversidad (INBio)

REFERENCIAS

CORREA A, M.D., C. GALDAMES Y M.S. DE STAPF, 2004, Catálogo de las plantas vasculares de Panamá. Smithsonian

Tropical Research Institute, Ciudad de Panamá, Panamá.

LurHER H.E. 1995. An annotated checklist of the Bromeliaceae of Costa Rica. Selbyana 16:230-234.

Morales, J.F. 1999, Seis nuevas especies de Vriesea section Xiphion (Bromeliaceae: Tillandsoideae) para Costa

Rica. Novon 9:401—406.

Moraes, J.F. 2003. Bromeliaceae. En: B. Hammel, M. Grayum, C. Herrera y N. Zamora, eds. Manual de plantas de

Costa Rica. Monogr. Syst. Bot. Missouri Bot. Gard. 92:297-375.

SmrrH, L.B. v RJ. Downs. 1977. Tillandsioideae (Bromeliaceae). Fl. Neotrop. 14(2):663-1492.

Urey, J. v K. Bunr.ev-Uriev. 1994. Bromeliaceae. En, G. Davidse, M. Sousa y A. Chater, eds. Fl. Mesoamer. 6:89-156.

1994

UrtEy, J., K. Bunr-Uriev v M.J. Hurr. 2001. Bromeliaceae. En: W.D. Stevens, C. Ulloa Ulloa, A. Pool y O.M. Montiel, eds.

Flora de Nicaragua Vol. 1. Mongr. Syst. Bot. Missouri Bot. Gard. 85:460-495.

UNA NUEVA ESPECIE Y NOTAS MISCELÁNEAS EN EL GÉNERO

OREOPANAX (ARALIACEAE) EN CENTRO AMÉRICA

J. Francisco Morales Alvaro Idárraga

Instituto Nacional de Biodiversidad (INBio) Herbario, Centro de Investigaciones

pto 22-3100 Facultad de Ciencias Exactas y Naturales nro

Santo Domingo de Heredia, COSTA RICA Universidad de Antioquia, Apartado Postal 122

Medellín, Antioquia, COLOMBIA

RESUMEN

aa ee de la revisión de las especies del género Oreopanax (Araliaceae) en Costa Rica, Nicaragua y Panamá, se proponen una

yen la descripción de una nueva especie, O. paramicolus, la redefinición del concepto de otras On nica-

raguensis), hectificación del rango de distribución geográfica para dos taxones (O. geminatus, O. peltatus) y el reporte d p

Panamá (O. nubigenus).

PALABRAS CLAVE: Centro América, Costa Rica, Panama, Araliaceae, Oreopanax

ABSTRACT

A lt of th isi f species of the g O Arali ) in Costa Rica, Nicaragua and Panama, Idi ies |

come ght. TI i d ,0 Pe here described. tl O. nicaraguensis and the need

oli

to adjust the geographical range b three taxa, O. geminatus, O. peltatus, and O. nubigenus, the latter ay reported for Panama.

Key Wonps: Central America, Costa Rica, Panama, Araliaceae, Oreopanax

Araliaceae es una familia pantropical de plantas, con alrededor de 41 géneros y ca. 1350 especies, distribuidas

principalmente en los trópicos (Plunkett et al. 2004). No existen tratamientos monográficos recientes de los

géneros del Nuevo Mundo y a nivel de Centroamérica, lo único disponible son sinopsis (Cannon & Can-

non 1989) o tratamientos florísticos (e.g., Cannon & Cannon 2001; Standley 1938). La ausencia de trabajos

monográficos y la alta variabilidad foliar en algunos géneros (e.g., Dendropanax, Oreopanax) ha provocado

una taxonomía confusa, sobretodo en lo que la correcta aplicación de nombres se refiere. Oreopanax, el cual

abarca cerca de 80 especies, 24 de ellas presentes en México y Centroamérica (Cannon & Cannon 1989,

2001), tampoco ha escapado de la situación descrita anteriormente, lo que ha traido como consecuencia la

incorrecta aplicación del concepto de algunos taxones y datos de distribución incorrectos para otros.

Como resultado de la elaboración del tratamiento de Araliaceae para el Manual de Plantas de Costa

Rica, se realizó un detallado estudio taxonómico de las especies de Oreopanax presentes en Costa Rica y

países aledaños, que incluyó la revisión de especimenes tipo y material en los principales herbarios de

Centroamérica, Estados Unidos y Europa. Como resultado, una serie de novedades fueron encontradas, las

cuales son descritas o comentadas a continuación. Estas incluyen la descripción de una nueva especie, así

como notas misceláneas sobre la incorrecta aplicación del concepto de algunos taxones y la rectificación

del rango de distribución geográfica de otros. Algunos datos citados en el manuscrito han sido tomados del

tratamiento de Araliaceae para el Manual de Plantas de Costa Rica (Morales et al., datos sin publ.), por lo

que pueden diferir de los de otros tratamientos previamente publicados.

Milos geminatus Marchal, Bull. Acad. Roy. Sci. Belgique, ser. 2, 47:91. 1879. Tiro: NICARAGUA. Nueva

GovIa: "America Centralis, ad Sajonia,” sin fecha (fl), A. Oersted 7 (HoLotIPo: C).

Distribución.—México, Belice, Honduras y Nicaragua, en elevaciones de 900—1750(-2000) m.

Oreopanax geminatus fue descrita con base en una colección hecha por Oersted en un sitio inespecífico

de Centroamérica, con una localidad vaga “Sejovia” (Marchal 1879). Desde entonces, se ha considerado que

esta especie está presente en Costa Rica, pues se asume que la localidad tipo corresponde a una localidad en

ese país. En general, muchas de las localidades donde recolectó Oersted en Costa Rica son zonas cercanas

J. Bot. Res. Inst. Texas 3(1): 117 —121. 2009

118 t tani i Texas 3(

al Valle Central, donde se ha conoce relativamente bien la flora. Sin embargo, aunque dicho colector realizó

colecciones principalmente en Costa Rica, también recolectó varios centenares de colecciones en Nicaragua

entre 1846 y 1848 (Stevens & Montiel 2001). Luego del estudio del especimen tipo de O. geminatus, así como

del material de esa especie presente en diferentes herbarios de Europa, Estados Unidos y Centroamérica,

hemos llegado a la conclusión de que la localidad "Sejonia" pone es análoga con el departamento

de Nueva eu en dd N de Nicaragua, donde E geminatus es te común (R. Abarca,

com, pers.). A , este taxón ] els EN Nicaragua ni en ninguna otra región

de Costa Rica. Por lo tanto, debemos delimitar que O. geminatus tiene un rango geográfico que se extiende

del S de México al N de Nicaragua, cuya colección tipo, fue recolectada en algún sitio del departamento de

Nueva Segovia en Nicaragua y que por el momento, basado en la evidencia suminstrada por especimenes

de herbario, no se conoce en Costa Rica.

Oreopanax nicaraguensis M.F Cannon & Cannon, Ann. Missouri Bot. Gard. 73:482, f. 1-2. 1986. Two:

NICARAGUA. Jinoreca: camino a Aranjuez, Santa Elena, 1-30 nov 1983 (fr), S. Vega & J. Quesada 197 (HOLOTIPO: BM; isoripos:

HNMN, MO)

Distribución —Endémica a Nicaragua, en elevaciones de 100—1300(-1500) m.

En la descripción de Oreopanax nicaraguensis Cannon & Cannon (1986) citaron seis especimenes de

Nicaragua, cuatro de Costa Rica y uno de Panamá. Desde entonces, se ha manejado un concepto de esta

especie que incluye especímenes que crecen casi a nivel del mar (100 m), hasta las zonas más altas de las

Cordilleras en Costa Rica y Panamá (3400 m), lo cual ha sido ampliamente seguido en colecciones de her-

barios y tratamientos florísticos subsecuentes (e.g., Cannon & Cannon 1989; Cannon & Cannon 2001).

Sin embargo, durante el estudio de material para el tratamiento de Araliaceae para el Manual de Plantas de

Costa Rica (Morales et al., datos sin publ, llegamos a la conclusión de que en la descripción original tres

especies distintas fueron citadas en los paratipos: O. nicaraguensis, a la cual pertenecen todos los especimenes

citados de Nicaragua, O. donnell-smithii Standl., a la que corresponde la colección de la cordillera de Tilarán

en Costa Rica (Dryer 1360, F, MO) y la última de ellas, una especie sin describir y restringida a las zonas

más altas de la cordillera de Talamanca en Costa Rica y Panamá (descrita en este trabajo). De esta forma, O.

nicaraguensis debe considerarse un taxón endémico al N de Nicaragua, el cual se encuentra algo relacionado

con O. donnell-smithii, O. paramicolus y O. striatus MJ. Cannon & Cannon, pero que se diferencia de este

grupo por sus hojas simples, enteras, con la base obtusa redondeada, i

o glabrescentes, las hermafroditas con cabezuelas eat pedunculadas, con grupos de 5a 10 flores

por cabezuela y frutos globosos o subglobosos al madurar, de 6—7 mm de largo y con los estilos con más de

la mitad inserta dentro de la depresión apical del fruto.

Oreopanax nubigenus Standl., J. Wash. Acad. Sci. 17:315. 1927. Two: COSTA RICA. San José: Las Nubes, 21 mar 1924

(£D, P Standley 38806 (noLoriro: US).

Distribución. —Costa Rica y el O de Panamá, en elevaciones de 155026002850) m.

Esta especie era considerada endémica a la Cordillera Central en Costa Rica (Cannon & Cannon 1989)

pero es reportada por primera vez para Panamá, donde se conoce en las estribaciones de la Cordillera de

Talamanca, en la provincia de Chiriquí. Oreopanax nubigenus puede ser confundida con O. standleyi A. C.

Sm., aún no reportada para Panamá (Correa et al. 2004), pero se diferencia por sus láminas foliares usual-

mente más pequeñas (6—10(-11.5) x 4-9.5(-11) cm vs. 12-25.5 x 5.5-20 cm), ramitas jóvenes glabras o

glabrescentes con el indumento muy esparcido (vs. tomentosas o tomentulosas) e inflorescencias con las

flores hermafroditas con 6 a 8 estilos (vs. 10—12)

Especímenes examinados. PANAMA. Chiriquí: Cerro Pate Macho, 31 dic 1985 (fr), de Nevers & Chanley 6689 (MO).

i oed o. E Morales & A. Idárraga, sp. nov. (Fig. 1). Two: COSTA RICA. San José: cantón de Pérez

Cerro de la Muerte, páramo Buena Vista, 19 oct 1993 (fr), J.E Morales, N. Zamora, E. Lépiz & V. Ramírez 1890 (uoLormo:

m. isoripo: CR).

RA J 1144 ga, N ia daf 119

Fic. 1. 0 icolus. A. Rama con frutos. B. Detalle de los frut

120

A Oreopanax nicaraguensis MJ. Cannon & Cannon, cui similis, inflorescentia hermaphrodita 15—30-floris, fructus 8-10 mm longus et

stylis exsertis, non inmersis differt.

Árbol o arbusto epífito de 2-10 m de altura, las ramitas glabras o glabrescentes y con el indumento incon-

spicuo. Hojas simples, enteras; lámina 7-22 x 9-21 cm, na a a ae ae de forma menos

común elíptica, el ápice acuminado o agudo, la base l comúnmente

redondeada a obtusa, glabra, pecíolo 4-20 cm de largo. Inflorescencias usualmente paniculadas, con las

ramificaciones evidentes, 12-17 cm de e ee o PO y con el qua inconspicuamente pu-

berulento justo antes de las cabezuela (González et al. 1592,

INB); bea de las a masculinas 8— 12 mm de diámetro, con grupos de 15 a 30 flores,

las flores con un estilo solitario; cabezuelas de las inflorescencias hermafroditas 5-7 mm de diámetro, con

grupos de 4 a 5(6) flores, las flores con 6-8 estilos. Frutos en grupos de 4-6 por cabezuela, ovoides, con un

diámetro de 8-10 mm, los estilos casi totalmente exsertos fuera de la depresión apical del fruto.

Distribución, habitat, y ecología.—Restringida a la Cordillera de Talamanca en Costa Rica y el O de

Panamá, donde crece en formaciones de robledales (Quercus spp., Fagaceae) y páramos, en elevaciones de

2500-3400 m. Floración se produce entre enero y febrero. Fructificación ocurre de enero a marzo, mayo, y

de julio a noviembre.

Oreopanax Duomo ha uu confundida con Oreopanax nicaraguensis M.J. Cannon & Cannon, pero

se diferencia por sus infl tas con mucho más flores (15 a 30 vs. 5-10), frutos de forma

diferente y más grandes (ovoides y de 8-10 mm de largo vs. globosos o subglobosos y de 6-7 mm de largo)

y estilos casi totalmente exsertos fuera de la depresión apical en el fruto (vs. estilos con más de la mitad in-

cluida). Adicionalmente, O. paramicolus crece en robledales y páramos (i de transición entre

ambos ecosistemas), entre 2500 y 3400 m, mientras que O. nicaraguensis crece en bosques muy húmedos,

entre los 100 y 1300(-1500) m.

Especímenes examinados. COSTA RICA. Cartago: Cerro de la Muerte, La Georgina, 2 ago 1965 (fr), Croat 261 (MO); reserva forestal

Rio Macho, estación Ojo de Agua, 12 ene 1996 (fr), Gamboa & Picado 950 (INB, MO); reserva forestal Río Macho, Cerro de la Muerte,

6 sep 1996 (fr), Rodríguez et al. 1470 (INB). Limón: parque nacional Chirripó, 15 feb 1983 (fl), Garwood et al. 1273 (BM, MO); parque

internacional La Amistad, Tararia, 12 abr 2002 (fr), González et al. 1592 (INB). Puntarenas: Cordillera de Talamanca, Cerro Echandi, 23

ago 1983 (fr), Davidse et al. 23876 (MO); parque internacional La Amistad, Cerro Echandi, 13 ago 1997 (fr), Quesada et al. 1994 (INB). San

José: entre Canaán ny cheapo via pu Angeles, ue El Río a 19-22 enel970 s Burger & Liesner 7381 (F, MO), 24 ago 1971 (fr),

d 8328 (F, ^ I hirripó, Indios, 1 may 1997 (fr), Gamboa & Aguilar 1271 (INB) reserva forestal

s Santos, camino a Pu 19 jul 1997 (fr), Gamboa 1620 (INB); Pérez eil albergue Cuerecí, 8 jul 2000 (fr), Miller 38 (INB,

m, cis: 10 mar 1994 (fr), Ramírez & Hammel 287 (INB, MO). San José-Cartago: Cordillera de Talamanca, Cerros Cuerecí, 15

set 1983 (fr), Davidse 24705 (MO); Copey, cerro Las Vueltas, 28 Mar 1994 (fr), Vargas et al. 1748 (CR). P. MA. Bocas del Toro: cerro

Fábrega entre cerros Itamut y Bine, Mar 1984 (fr), Gómez et al. 22461 (MO). Chiriquí: Boquete, camino a Volcán Barú, 18 may 1976 (fr),

Croat 34885 (MO); Potero Muleto, Volcán Barú, 27 nov 1975 (st), Volcán de Chiriquí, 20 nov 1975 (fr), Davidse & D'Arcy 10273 (MO); El

Volcán, SE de la Nivera, sin fecha (fr), D'Arcy 12505 (MO); Volcán Barú, Potrero Muleto, 6 nov 1978 (fr), Hammel 5642 (MO); Volcán Barú,

24 jul 1975 (fr), Mori & Bolten 7440 (MO); parque nacional Volcán Barú, 15 ene1992 (fl), Vega & Rincón (MO, PMA

Oreopanax peltatus Linden, Gartenflora 11:170, t. 363. 1862. Tiro: MÉXICO: Datos perdidos, J. Linden s.n. (notorio:

Distribución. —México, Guatemala y El Salvador, en elevaciones de 700—1900(—2400) m.

Algunos especimenes de Costa Rica y Panamá, han sido identificados como esta especie (e.g., Liesner

et al. 15518, MO), pero estas colecciones en realidad corresponden a especimenes de Dendropanax estériles,

género en el cual la presencia de hojas peltadas e irregularmente lobuladas es algo comün en los brotes

inmaduros de muchas especies, las cuales se tornan no peltadas con la edad. En Centroamérica Oreopanax

peltatus se puede reconocer fácilmente por sus hojas simples, 5—7-lobuladas, peltadas y con la base redon-

deada o subcordada a no peltatas y cordadas e inflorescencias con las flores estaminadas de 3-6 mm de

diámetro.

AGRADECIMIENTOS

Deseamos reconocer a los siguientes herbarios por permitir el ingreso a sus colecciones: BM, BR, CGE, COL,

, A k An ft 121

CR, CUVC, F, G, HUA, HUQ, INB, JAUM, K, MEDEL, MA, MO, NY, P, Q, QCA, QCNE, TULV, U, W, WAG.

Esta investigación fue posible además gracias al convenio de cooperación entre el Ministerio de Ambiente

y Energía (MINAE) y el Instituto Nacional de Biodiversidad (INBio) con el fin de completar el Inventario

Nacional de Biodiversidad en Costa Rica.

REFERENCIAS

Cannon, M.J. 8: J.F.M. Cannon. 1986. Studies in the Araliaceae of Nicaragua, and a new widespread species of

Oreopanax. Ann. Missouri Bot. Gard. 73:481—485.

Cannon, M.J. & J.F.M. CANNON.1989. Central American Araliaceae - A precursory study for the Flora M icana.

Bull. Brit. Mus. (Nat. Hist), Bot.19:5-61.

Cannon, M.J. & J.F.M. Cannon. 2001. Araliaceae. En: W.D. Stevens, C. Ulloa Ulloa, A. Pool & O.M. Montiel, eds. Flora

de Nicaragua Vol. |. Monogr. Syst. Bot. Missouri Bot. Gard. 85:188-192,

CORREA, M., C. GALDAMES & M. DE STAPF. 2004. Catálogo de las plantas vasculares de Panamá. Universidad de Panmá,

Instituto Smithsonian de Investigaciones Tropicales, Ciudad de Panamá, Panamá.

Marcha, E. 1879. Révision des Hédéracées américaines. - Description de dix-huit espéces nouvelles et d'un

genre inédit. Bull. Acad. Roy. Sci. Belgique, ser. 2, 47:70-96.

PLUNKETT, G.M., J. WEN, AND PP. Lowey II. 2004. Infrafamilial relationships in Araliaceae: insights from plastid (trni-trnF)

and nuclear (ITS) sequence data. Pl. Syst. Evol. 245:1-39,

STANDLEY, P.C. 1938. Araliaceae. En: P. C. Standley, ed. Flora of Costa Rica. Publ. Field Mus. Nat. Hist., Bot. Ser.

18:851-858.

STEVENS, W. & O. Montiet. 2001. Reseña de la exploración botánica. En: W.D. Stevens, C. Ulloa Ulloa, A. Pool & O.M.

Montiel, eds. Flora de Nicaragua Vol. |. Monogr. Syst. Bot. Missouri Bot. Gard. 85.

122 [| rnal of tl Das H ID khi HH £ T,

BOOK REVIEW

JuLes G. Evans. 2008. California Natural History Guides: Natural History of the Point Reyes Peninsula,

Second Edition. (ISBN 978-0-520-25467-1, pbk.). University of California Press, 2120 Berkeley Way,

Berkeley, California 94704-1012, U.S.A. (Orders: The University of California Press c/o California/

Princeton Fulfillment Services, 1445 Lower Ferry Road, Ewing, New Jersey 08618, U.S.A.; www.ucpress.

edu, email ordersOcpfsinc.com, 1-800-777-4726, 1-800-999-1958 fax). $24. i 366 pp, numerous

figures, Es images, maps, tables, checklists, index, and references, 5 1/4" x

Natural Hi t fl T». T D. L9 | I 11 J 31 1 Tz , geology. | various fauna

(e.g. birds, insects marsupials, carnivores pinnipeds, bats whales, M reptiles, hibians) 1fl ities that inhabi

this Calif p fthis book (1989, revised 1993), at f hers! jucted studi

on Point Reyes, and the inf ion they accumulated ini nint conan this new ie

Nw. J ze J D T DP. 1

4, J [94 E +

à MES 1 "E Hu Xt hofthereseión'ehabitare- (tr trial, riparian, freshwater

r £ É O > Eu

and usd wi I I 1 prinklesi hi ical asid fa parti lari I 1 d , and definitions greatly

enhancing the overall reader experience.

I found this book to be very informative and a good resource for any ici or pum ns is dl to learn more about the

diversity of biota found on Point Reyes doa M QUEM The anh size and weig I companion ad any hiker or

outdoorsman. The author used easy-to ] der witl d sci

The inclusion of a comprehensive checklist of the fauna and ofa with information on whether Pus species is listed as endangered.

vulnerable, rare, or threatened further enhanced this book.—Keri McNew, MS Biology, Prog ger, Botanical Research Institute of

Texas, 500 E 4" Street, Fort Worth, TX 76102, U.S.A.

J. Bot. Res. Inst. Texas 3(1): 122. 2009

FIVE NEW SPECIES OF MYRTACEAE FROM ECUADOR

Maria Lúcia Kawasaki

Bruce K. Holst

Department of Botany Marie Selby Botanical Gardens

Field Museum of Natural History 811 South Palm Avenue

00 South Lake Shore Drive

Sarasota, Florida 34236-7726, U.S.A.

b lb

Chicago, Illinois 60605-2496, U.S.A. holst@selby.org

lkawasaki@fieldmuseum.org

ABSTRACT

Five new species of M from Ecuad described and ill d: Calyptranthes compactiflora, C. fusca, C. sparsiflora,

Myrcia aequatoriensis, and M. verticillata.

RESUMEN

Se J } E Y do NM t EE 1

: Calyptranthes compactiflora, C. fusca, C. sparsiflora,

Myrcia cea aioe y M. verdeillata:

INTRODUCTION

Among the Myrtaceae from Ecuador, there are approximately 30 species of Calyptranthes and ca. 40 spe-

cies of Myrcia, including many new taxa (Holst 1999; Kawasaki & Holst 2005; Holst & Kawasaki 2006,

2008). In preparation of the Myrtaceae treatment for the Flora of Ecuador, five additional new species of

these genera are herein described: Calyptranthes compactiflora, C. fusca, C. sparsiflora, Myrcia aequatoriensis,

and M. verticillata.

Calyptranthes compactiflora M.L. Kawasaki & B.K. Holst, sp. nov. (Fig. 1). Tre: ECUADOR. Sucumsios (Napo on

label): downstream Río Cuyabeno from Laguna de Cuyabeno to Puerto Bolívar, 00°03'S, 76°10'W, 250 m, 14 Apr 1980 (bud), J.

Brandbyge, E. Asanza & L. Reib 30543 (noLotYPE: AAU; isotypes: E, QCA, SEL).

Arbor, ind to rufo-brunneo , ferrugineo vel flavo-brunneo; ramulis teretis; foliis anguste ellipticis vel ellipticis, chartaceis, 5-8 x

s x Te lA TL CEST: if] H lah = 1 E

p 3 m longis

1.8-3 cm, abrupto-acuminatis, si | I

Tree ca. 4 m tall, the trichomes dibrachiate, reddish-brown, ferrugineous to yellowish-brown; branchlets

terete, appressed-pubescent. Leaf blades narrowly elliptic to elliptic, chartaceous, 5-8 x 1.8-3 cm, the up-

per surface drying dark-brown, glabrous and lustrous, indistinctly or impressed-punctate, the lower surface

reddish-brown in drying, sparsely appressed-pubescent, distinctly dark-punctate; midvein impressed above,

convex below; lateral veins 35—40 pairs, parallel, scarcely raised on both surfaces; marginal vein 1, ca. 1 mm

from the margin, equaling the lateral veins in prominence; apex abruptly acuminate, the acumen to ca. 1.3

cm long; base cuneate; petioles 5-9 mm long, channeled, puberulous, blackish. Inflorescences of paired,

very abbreviate panicles appearing glomerulate by the reduction of the main axes, axillary, with up to ca. 6

flowers, the peduncle 1-2(-10) mm long, pubescent; bracts lanceolate, 3-5 mm long, appressed-pubescent,

early deciduous; bracteoles ovate, ca. 2 mm long, appressed-pubescent, deciduous. Flowers: buds closed,

obovoid, 3-4 mm long, sessile, appressed-pubescent, glabrescent, the e furfuraceous, yellowish-

t; disk ca. 2 mm diam., glabrous;

white; calyptra obtuse to shortly apiculate, s parsely pubescent; p

stamens numerous, ca. 2-3 mm long, the anthers ca. 0.5 mm long; style ca. 2 mm long; ovary 2-locular;

ovules 2 per locule. Fruits not seen.

Distribution.—Known only from the type collection from northeastern Ecuador in the Río Cuyabeno

region, in riverside forests at 250 m elevation.

Calyptranthes compactiflora is readily recognized by the greatly reduced inflorescences, also seen in C.

smithii McVaugh, a species known only by the type collection from Guyana. These two species may be dis-

tinguished by leaf characters: in C. compactiflora, the leaves are chartaceous, 5-8 x 1.83 cm, the midvein

J. Bot. Res. Inst. Texas 3(1): 123 -132. 2009

124

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FLORA OF ECUADOR

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Weed collection in ÁAU.

Fic. 1. Calyp h | ctiflora M.L. Kawasaki 8. B.K. Holst (Brandby tal. 30543: Holotype, AAU; i f

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is clearly impressed above, and the petioles are 5-9 mm long; in C. smithii, the leaves are coriaceous, 9-12

x 3.5-4.5 cm, the midvein is plane to impressed above, and the petioles are 10-13 mm lon

Calyptranthes fusca M.L. Kawasaki € B.K. Holst, sp. nov. (Fig. 2). Type: ECUADOR. CARCHI: Tulcán Canton,

Parroquia Tobar Donoso, Sector Sabalera, Reserva Indígena Awa, 01°00'N, 78°24'W, 650-1000 m, 19-28 Jun 1992 (fI), G. Tipaz, J.

Zuleta & N. Guanga 1438 (HoLoTiPE: QCNE; isotypes: E MO, SEL)

Arbor, ind fl i li is; folii llipticis, ellipticis vel oblanceolatis, chartaceis, 5.5-14 x 3-5 cm, abrupto-

acuminatis, siccati | i iculi ifloris; alal is 1-3 mm longis, petalis nullis; baccis globosis, atropurpureis.

Tree 10-20 m tall, 15-40 cm dbh, the trichomes dibrachiate, yellowish; branchlets terete, puberulous.

Leaf blades narrowly elliptic to elliptic, or oblanceolate, chartaceous, 5.5-14 x 3-5 cm, drying dark-brown

to brownish above, paler below, glabrous on the upper surface, sparsely appressed-pubescent to glabrous

on the lower surface, the trichomes located especially along venation; glands distinctly dark-brown, plane

above, salient below, also on venation, branchlets, and inflorescences; midvein impressed above, convex

below; lateral veins 10—15 pairs, impressed above, salient below; marginal veins 2, the innermost arched,

3-5 mm from the margin, similar to the lateral veins in prominence, the outermost less prominent, paral-

lel to the margin, ca. 1 mm from it; apex abruptly acuminate, the acumen 1-1.5 cm long; base cuneate to

obtuse; petioles 5-10 mm long, channeled, sparsely pubescent, blackish. Inflorescences of paired panicles,

subterminal or axillary, with up to ca. 20 flowers, 2-8 cm long, the branches densely appressed-pubescent,

gland-dotted; bracts ovate, ca. 2 mm long, puberulous, early deciduous; bracteoles lanceolate, ca. 1 mm

long, puberulous, deciduous. Flowers: buds closed, obovoid to subglobose, of very different sizes, 1-3 mm

long, sessile, the lateral flower buds of the terminal triads apparently abortive; hypanthium prolonged ca.

1 mm beyond the ovary, glabrous; calyptra obtuse to minutely apiculate, glabrous; petals absent; disk ca.

2 mm diam., glabrous; stamens numerous, ca. 4—5 mm long, the anthers ca. 0.3 mm long; style ca. 5 mm

long; ovary 2-locular; ovules 2 per locule. Fruits berries 1-2 cm diam., globose, crowned by the hypan-

thium scar, dark-purple, glabrous, prominently gland-dotted, the pericarp thick and hard; seed 1, ca. 9 x

7 mm, the seed coat membranous; embryo myrcioid, the cotyledons leafy and folded, the radicle elongate,

equaling the cotyledons in length.

Distribution.—Known from several collections from northwestern Ecuador of the same area (Reserva

Etnica Awá) in the provinces of Carchi and Esmeraldas, in humid, lowland to premontane forests at 200-1600

m elevation. The wood is utilized for ax handles and other tools (Tipaz et al. 1290) and the edible fruits to

cure diarrhea (Tipaz et al. 1233). Common names are recorded as “ungal nusma” (Tipaz et al. 1290), “unga

lusma” (Tipaz et al. 1438), and “pishuilde” (Tipaz et al. 1126).

The leaves of Calyptranthes fusca and C. cuspidata DC., a species from Amazonian Brazil and Peru, are

morphologically similiar and dark-brown in drying. Calyptranthes fusca is distinguished from C. cuspidata

by having the leaves with impressed (vs. convex) midveins above, conspicuous (vs. inconspicuous) and dark

glands on both surfaces, along the venation and on the inflorescence axes, and relatively few-flowered (vs.

multiflorous) panicles. The varying size of the flower buds is unusual in the family.

Additional coliecti ined: ECUADOR. Carchi: Maldonado, Parroquia Tobar Donoso, Reserva Etnica Awá, Sabalera, 00?55'N,

78°32'W, 900 m, 22 Nov 1992 (fr), C. Aulestia et al. 658 (F, MO, QCNE, SEL). Tulcán Cantón, Parroquia Tobar Donoso, Sector El Baboso,

Reserva Indígena Awa, 00°53'N, 78°00'W, 1600 m, 3 Oct 1991 (fr), G. Tipaz et al. 310 (F, MO, QCNE, SEL). Tulcán Cantón, Parroquia

Chical, Sector Gualpi medio, Reserva Indígena Awá, 01°02'N, 78°16'W, aD 23-27 May 1992 (fr), G. Tipaz et al. 1126 (F, MO, QCNE,

SEL). Tulcán Cantón, Parroquia Tobar Donoso, Sector Sabalera, R igena Awá, 01°00'N, 78°24'W, 650-1000 m, 19-28 Jun 1992

(fr), G. Tipaz et al. 1233 (F, MO, QCNE, SEL). Tulcán Cantón, Parroquia Tobar Donoso, Sector Sabalera, Reserva Indígena Awá, 01?00'N,

78°24'W, 650-1000 m, 19-28 Jun 1992 (fl), G. Tipaz et al. 1290 (F, MO, QCNE, SEL). Esmeraldas: San Lorenzo Cantón, Parroquia

Mataje, Reserva Etnica Awa, Centro Mataje, on the banks of Rio Mataje, 01°08'N, 78?33'W, 200 m, 21 Sep 1992 (fr), C. Aulestia et al. 551

F, MO, QCNE, SEL). San Lorenzo Cantón, Parroquia Mataje, Reserva Etnica Awd, Centro Mataje, on the banks of Río Mataje, 01°08'N,

78*33'W, 200 m, 21 Sep 1992 (fr), C. Aulestia et al. 568 (F, MO, QCNE, SEL).

Calyptranthes sparsiflora M.L. Kawasaki & B.K. Holst, sp. nov. (Fig. 3). Type. ECUADOR. SUCUMBÍOS: Lago

Agrio Cantón, Reserva Cuyabeno, Laguna Canangueno, 00%02'S, 76°13'W, 230 m, 18 Nov 1991 (ID, W. Palacios, G. Tipaz, D. Rubio,

E. Gudiño & C. Aulestia 9126 (HoLotYrE: QCNE; IsorYPE: MO, SEL)

126

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128 Journal of t t Texas 3(

Frutex vel arbor, indumen to rufo-l li inatis; folii llipticis, chartaceis, 4—6.5 x 1.5-2.5 cm, caudato-acuminatis;

> o

inflorescentiis 1 vel 3 floris, dense appresso-pubescentibus; baccis globosis.

Shrub or tree 1.5—4 m tall, the trichomes dibrachiate, reddish-brown, to 0.5 mm long; branchlets bicari-

nate, thinly appressed-pubescent to glabrous. Leaf blades narrowly elliptic, chartaceous, 4-6.5 x 1.5-2.5

cm, the upper surface drying olive-green to dark-brown, glabrous, indistinctly or impressed-punctate, the

lower surface light-brown in drying, sparsely appressed-pubescent, minutely punctate; midvein impressed

above, convex below; lateral veins 30—40 pairs, parallel, scarcely raised on both surfaces; marginal vein 1,

0.5-1 mm from the margin, equaling the lateral veins in prominence; apex caudate-acuminate, the acumen

to ca. 1.5 cm long; base obtuse to cuneate; petioles 1-3 mm long, channeled, puberulous. Inflorescences

paired, subterminal, with 1 or 3 subsessile flowers, borne at the end of a 1.3-2.8-mm long peduncle, densely

appressed-pubescent; bracts and bracteoles early deciduous, not seen. Flowers: buds not seen, the hypan-

thium 1.5-2 mm long, densely appressed-pubescent; calyptra ca. 1 mm long, shortly apiculate, densely to

loosely appressed-pubescent; petals not seen; disk ca. 2 mm diam., glabrous; stamens not seen expanded,

the anthers ca. 0.3 mm long; style ca. 7 mm long. Fruits berries, immature, globose, 8-9 mm diam., yel-

low, appressed-puberulous, the trichomes located especially at the base and by the hypanthium scar; seed

1, ca. 8 x 7 mm, the seed coat membranous; embryo myrcioid, the cotyledons leafy and folded, the radicle

elongate, equaling the cotyledons in length.

Distribution.—Known from eastern Ecuador in Sucumbíos and Pastaza provinces, in primary rain

forests at 200—230 m elevation.

Calyptranthes sparsiflora is characterized by the very reduced, 1- or 3-flowered inflorescences, that are

densely appressed-pubescent with reddish-brown mne ns the species from Ecuador, it resembles

C. bipennis O. Berg on branchlet, leaf, and infl y. These two species may be distinguished

by the presence of indumentum on young vegetative MN [e inflorescences, flowers, and fruits in

C. sparsiflora (ys. mostly glabrous in C. bipennis).

Additional collection examined: ECUADOR. Pastaza: Lorocachi, SW of the military camp, 01?38'S, 75°58’W, 200 m, 26 May 1980

(fr), J. Jaramillo et al. 31106 (AAU, F, QCA).

Myrcia A id L. Kamarak & B.K. Holst, sp. nov. (Fig. 4). Tre: ECUADOR. Sucuwsios: Reserva Faunistica

Cuyabeno, gs, including Río Cuyabeno from Puerto Bolívar to above Laguna Canangueno, 00°00'S,

76°10'W, 2651 m, 11 Mar 1990 (fl), H. Balslev, C.C. Berg, M. Gavilanes, A. Thygesen, D.E. Christensen, L. Ellemann & R. Brucculeri 97496

(HOLOTYPE: AAU; sorres: ASU, E QCA, SE

Frutex vel arbor, indumento flavescenti vel cano-flavescenti; ramulis teretis; foliis lanceolatis, chartaceis, 5.6-14 x 1.6—3 cm, longi

acuminatis, basi obtusis; paniculis axibus gracilibus; baccis ellipsoideis.

Shrub or tree 2—10 m tall, the trichomes simple, yellowish to yellowish-white; branchlets terete, densely

pubescent when young, peeling in thin strips when slightly older. Leaf blades lanceolate, 5.6—14 x 1.6-3

cm, membranous to chartaceous, the upper surface glabrous except for the midvein, drying olive-green to

brownish, the lower surface paler, sparsely appressed-pubescent, the trichomes present especially along

the venation; apex long-acuminate; base obtuse; midvein impressed above, pubescent, convex below; lat-

eral veins 20—30 pairs, impressed above, sharply convex below; marginal vein 1, to ca. 1 mm from blade

margin, indistinct above, slightly salient below; glands numerous, indistinct or minutely punctiform on

both surfaces; petiole 1-3 mm long, channeled, densely pubescent. Inflorescences paniculate, axillary or

mostly subterminal, with to ca. 20 flowers, 3—6.5 cm long, the axes appressed-pubescent, filiform; bracts

and bracteoles early deciduous, not seen. Flowers 5-merous; buds obovoid to io d mm 1 long;

calyx-lobes triangular, to 1 mm long, appressed-pubescent without, glabrous within; p , Cà

2 mm diam., sericeous without; stamens numerous, the filaments ca. 3-4 mm long, the anthers ca. 0.3 mm

long; style ca. 4 mm long, the stigma punctiform; hypanthium not prolonged beyond the ovary, sericeous

without; disk ca. 2 mm diam., densely pubescent; ovary 2-locular; ovules 2 per locule. Fruits (immature)

berries, ellipsoid, ca. 10 x 8 mm, crowned by the persistent calyx lobes, yellow, gland-dotted, puberulous

Hire TOR W^

FLORA or. E ECUAD: OR

Collected by H. Balsiov, M. Gavilanes,

A, Thygesen, D. E. Christ solacia mian BR: Brucculeri

97496

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is M.L. Kawasaki & B.K. Holst (Balsi |, 97496: Holotype, AAU; inset from isotype at SEL).

Fic. 4. Myrcia aeq

£ 4L Das s.. 1D Llar ET

130 Journal of Texas 3(

to glabrous; seed 1, ca. 8 x 7 mm, the seed coat membranous; embryo myrcioid, the cotyledons leafy and

folded, the radicle elongate, equaling the cotyledons in length.

Distribution.—This species is known from a few collections from Sucumbíos (Reserva Faunística Cuy-

abeno) in northeastern Ecuador, and one collection from Imbabura in northwestern Ecuador; it occurs in

lowland tropical rainforests at 220-500 m elevation.

Myrcia aequatoriensis belongs to sect. Myrcia, among the group of species related to the M. fallax (Rich.)

DC. — M. splendens (Sw.) DC. complex. These species are separated by characters of the indumentum, leaf

morphology, including venation and glands, inflorescences, and size of flower buds (McVaugh 1969). From

this group, M. aequatoriensis is distinguished by the combination of these characters: lanceolate, thin, sub-

sessile leaves, long-acuminate at the apex, obtuse at the base, with impressed lateral veins on the upper

surface, and by delicate panicles with filiform axes.

Addi l collecti ined: ECUADOR. Imbabura: Lita, 501 m, 28 Apr 1949 (fr), M. Acosta Solís 12296 (F). Sucumbios: Reserva

Faunística Cuyabeno, Laguna Grande and surroundings, including Río Cuyabeno from Puerto Bolívar to above Tapuna E m

00°00'S, 76°10'W, 265 m, 11 Mar 1990 (fl), H. Balslev et u 97071 (AAU, QCA). Reserva Faunística Cuyabeno, I

roundings, including Río Cuyabeno from Puerto Bolívar to above Laguna Canangueno, 00°00'S, 76°10'W, 265i m, 11 Mar 1990 (f),

H. Balslev et al. 97217 (AAU, ASU, F, QCA, SEL). Estación Científica Cuyabeno, 220 m, 21 Jul 1992 (fr), J. Jaramillo 14913 (NY, QCA).

Reserva Faunística Cuyabeno, 1 ha plot ca. 1 km N of Laguna Grande and surroundings, 00°00'S, 76°12'W, 265 m, 11 Apr-10 Jun 1988

(st), A.D. Poulsen 78348 (AAU-2 sheets, QCA).

Myrcia verticillata M.L. Kawasaki € B.K. Holst, sp. nov. (Fig. 5). Tere: ECUADOR. ORELLANA (Napo on label): Estación

Científica Yasuní, Río Tiputini, NW of confluence with Río Tivacuno, 6 km E of Maxus road, Km 44, detour to Tivacuno well, parcela 50

ha, 00938"S, 76930'W, 200—300 m, 21 Mar 1996 (fr), K. Romoleroux & R. Foster 2147 (HoLoTveE: QCA; isotypes: F-2 sheets, SEL).

Frutex vel arbor, indumento branie vel O Sa ramulis quadrangulatis; foliis verticillatis, RN MEN coriaceis, 19-29 x

lis 5-13 cm longis; baccis globosis,

E Is E

Shrub or tree qud m tall, to ca. 20 cm dbh, the trichomes simple, brown to yellowish-brown; bark red-

dish, the stems lar in cross section with rounded angles, densely pubescent. Leaves subsessile to

short-petiolate, verte biet in 4-merous whorls; blades oblanceolate, 19-29 x 4—7.5 cm, coriaceous, the upper

surface drying dark brown to brownish, puberulous, the lower surface paler, sparsely hirsute-pubescent,

the trichomes present especially on the venation; apex abruptly acuminate; base obtuse; midvein hirsute,

impressed above, convex below; lateral veins 15—20 pairs, these and major interconnecting tertiary veins

impressed above, convex below; marginal veins 2, the innermost 2-4 mm from blade margin, similar in

prominence to the lateral veins, the outermost less than 1 mm from margin; glands numerous, punctiform,

indistinct above, evident below; petiole 3-9 mm long, stout, channeled, densely pubescent. Inflores-

cences paniculate, axillary, 5-13 cm long, the axes hirsutulous; bracts and bracteoles early deciduous, not

seen. Flowers 5-merous; buds not seen; calyx-lobes ca. 1 x 2 mm, truncate to broadly rounded at apex,

appressed-pubescent to puberulous without, glabrous within; petals, stamens, and style not seen; hypan-

thium not prolonged beyond the ovary; disk ca. 2 mm diam., hirsutulous. Fruits berries, globose, 1-1.7

cm diam., turning from green to yellow-orange to red, and finally purple-black (Acevedo-Rdgz. & Cedeño

7628), puberulous, crowned by the slightly overlapping calyx lobes; seeds 1 or 2, ca. 8-10 x 7-8 mm, the

seed coat membranous; embryo myrcioid, the cotyledons leafy and folded, the radicle elongate, equaling

the cotyledons in length.

Distribution.—Ihis species is known only from northeastern Ecuador in Orellana province, in the region

of Yasuní National Park; it occurs in primary, lowland wet forests, at 200—300 m elevation.

From all the other species of Myrcia in Ecuador (Holst 1999; Holst & Kawasaki 2008), M. verticillata

is promptly recognized, even in sterile condition, by the large, oblanceolate, dark brown leaves (when dry),

that are subsessile and verticillate, in tetramerous whorls, and by the quadrangular stems. Species of Myr-

cia with both opposite and verticillate leaves were described by Berg (1857) in Martius' Flora Brasiliensis,

but at least in northern South America, M. verticillata is the only known species with leaves consistently in

tetramerous whorls. Myrcia verticillata may be allied with the Gomidesia group of Myrcia; flowers are needed

to confirm the relationship.

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FELD MUSEUM

NATURAL HISTORY

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Neil et al. 8233, SEL)

132 Journal of the Botanical R h Institute of Texas 3(1)

Additional collections examined: ECUADOR. Orellana: Yasuní Forest Reserve, along road between Km 70 and 100, E of PUCE Scien-

tific Station, 0°50.014’S, 76220.518"W — 0%54.730'S, 76?13.304"W, 200 m, 2 Jul 1995 (fr), P. Acevedo-Rodriguez & J.A. Cedeño 7628 (SEL,

US); Yasuní, Parque Nacional Yasuní, oil well Conoco-Amo 2, 00?57S, 76?13"W, 230 m, 9-19 Jan 1988 (fr), D. Neill et al. 8233 (F, MO,

SEL); Aguarico, Yasuní, Reserva Etnica Huaorani, Maxus road and pipeline construction project, Kms 98-99, 00°55’S, 76*13'W, 250

m, 18 Jun 1994 (fr), N. Pitman & G. Romero 299 (F, MO, QCNE, SEL); Estación Científica Yasuni, Río Tiputini, NW of confluence with

Río Tivacuno, E of Repsol-YPF road, Km 7, detour to Tivacuno well, parcela 50 ha, column 29-02(2,3), 00°38'S, 76°30"W, 200-300 m,

17 Feb 2001 (fr), G. Villa & L. Velez 912 (F, QCA).

ACKNOWLEDGMENTS

We are grateful to the curators and staff of the herbaria cited in the manuscript for providing the collections

used in this study, especially to Henrik Balslev and Benjamin Dilgaard from AAU, and Ron Liesner and

Jim Solomon from MO. Fred Barrie, Les Landrum, and Jim Luteyn offered helpful suggestions to improve

this manuscript.

REFERENCES

Bera, O. 1857-1859. Myrtaceae. In: C.P. von Martius, ed. Fl. Bras. 14(1+suppl.):1-655.

Ho st, B.K. 1999. Myrtaceae. In: P.M. Jørg d S. León-Yánez, eds. Catalogue of the vascular plants of Ecuador.

Monogr. Syst. Bot. Missouri Bot. Gard. 75:618-622.

Hoist, B.K. AND M.L. Kawasaki. 2006. New species of Myrtaceae from Ecuador and Peru. Sida 22:931-934,

Ho st, B.K. and M.L. Kawasaki. 2008. New species of Myrtaceae from Ecuador. J. Bot. Res. Inst. Texas 2:297-303.

Kawasaki, M.L. AND B.K. Hoist. 2005. Two new species of Calyptranthes (Myrtaceae) from Ecuador. Sida 21:1955-

1960.

McVaucn, R. 1969. The botany of the Guayana Highland—Part VIII. Myrtaceae. Mem. New York Bot. Gard.

18(2):55-286.

FOLIA TAXONOMICA 12. PARADRYMONIA (GESNERIACEAE: EPISCIEAE) FROM

THE GUIANA SHIELD: P. MAGUIREI, A NEW SPECIES FROM AMAZONAS, AND

DISTRIBUTION AND FLORAL MORPHOLOGY OF P. MACULATA

Christian Feuillet

Department of Botany, MRC-166

Smithsonian Institution, PO. Box 37012

Washington, DC 20013-7012, U.S.A.

feuillec@si.edu

ABSTRACT

Parad i irei is described f I fA Venezuela. The distribution of Paradrymonia maculata in Venezuela

and the Guiya is enel: as well as its unusual corolla morphology.

RÉSUMÉ

Paradry i irei est décrit de l'état d'A Venezuela. La répartition de Parad i lata au Venezuela et dans les

i M 11

eid ta th i 1 11 Ap f,

RESUMEN

Se describe P be puli del estada de a Msn bord se documenta la distribución de Paradrymonia

maculata en Venezuela y las

Hanstein (1854: 206) described the genus Paradrymonia Hanst. with only one species, P. glabra (Benth.) Hanst.

(1854, p. 207, fig. 43), which is currently a synonym of P. ciliosa (Mart.) Wiehler. The name Paradrymonia

was in use only for a short time as Hanstein (1864) reduced it to a synonym of Episcia Mart. When Wiehler

(1973) re-established Paradrymonia, he transferred species from other genera. From Drymonia Mart. he took

species that did not have the typical Drymonia anther dehiscence through a basal pore, and from Episcia

Mart. he moved species that lacked stolons and did not otherwise belong to Nautilocalyx Linden ex Hanst.

Currently Paradrymonia includes 38 species.

Recent molecular studies (Clark et al. 2006) have shown that it is likely that the species of Paradrymonia

will separate into two natural groups of species, mostly with large leaves (20—50 cm long) and small axillary

inflorescences, and a few isolated species. The larger group, true Paradrymonia, would mostly include species

with petioles longer than the inflorescences and either with stems 5-10 mm thick and rooting at nodes on

the substrate, and the smaller group has thinner and shorter stems and a “rosette-like” habit (Wiehler 1978).

Those two groups are likely to stay in Paradrymonia. They are present in continental America from Mexico

(Oaxaca) to Bolivia (Cochabamba) and eastward to Brazil (Amapá).

Afew Species in eee have tain affinities. P. anisophylla Feuillet & L.E. Skog is an epiphyte

with hang gly unequal in a pair; the molecular data (Clark et al. 2006) suggest that

it may not be a Paradrymonia. The same data set places outside Paradrymonia an epiphyte with erect thick

stems, P. longifolia (Poepp.) Wiehler. Paradrymonia campostyla (Leeuwenb.) Wiehler and P. barbata Feuillet

& L.E. Skog from the Guianas are climbers with smaller leaves (3-15 cm long) and 1(-3) axillary flowers.

Here two other hard-to-place species are dealt with: P. maguirei, a new species from Amazonas (Venezuela),

which is vegetatively unlike other Paradrymonia species with a rosette-like habit, short petioles, and sharply

biserrate paper-thin leaves, when dry; and P. maculata (Hook. f.) Wiehler with large condensed inflorescences

and large bracts (see below) that is endemic to the Guiana Shield.

J. Bot. Res. Inst. Texas 3(1): 133 -138. 2009

134 i itute of Texas 3(

A.—Paradrymonia maguirei Feuillet, sp. nov.

Although it seems obvious that the genus Paradrymonia will prove to be polyphyletic (Clark et al. 2006), in

the absence of a comprehensive molecular study of Paradrymonia, along with Nautilocalyx and Chrysothemis

Decne., the only reasonable option at the moment is to place this new species in Paradrymonia as P. magu-

irei. It does not belong in any of the other genera of the Episcieae (Weber 2004; Skog & Boggan 2006) and

anticipating the split of Paradrymonia by describing a new genus without the proper data would be taking

a high risk of creating a generic synonym.

Paradrymonia maguirei Feuillet, sp. nov. (Fig. 1). Tree: VENEZUELA. AMAZONAS. Depto. aan od al adc d

forest, 1000 m, 3 May 1949, B. Maguire & B. Maguire, Jr. 29185 (Hototyre: NY, pro p I

= [P marahuacana Wiehler] invalid: in sched.

iin d maguirei ab aliis speciebus a chatactenbus angen o "dus pro parte manie praeter infra foliis intervenia,

10-11 x 7-8 cm, basi asym-

vestita, petiolo 2-2.5 cm longo, 1 velutina PI

metrica, apice obtuso rotundo, margine bi- serrata, in sicco FRANE

Epiphyte or saxicolous. Stem creeping, 0.5 cm thick or more, 5 cm long (in the type collection), with a

dense brown-red indumentum, apical few internodes with leaves, about 3 mm long, forming some kind

of a loose pauci-leaved rosette. Leaves opposite, strongly unequal in a pair, the smaller about 2 cm long

including petiole, ligulate, 2-3 mm wide; the larger with petiole 2-2.5 cm long, thick, covered with dense,

long, brown-red trichomes; blade membranous when dry, elliptic, 10—11 x 7-8 cm, asymmetrically acute to

obtuse at base, widely rounded at apex, margin sharply biserrate, above velutinous or appressed-pubescent,

beneath appressed-pubescent or hirsute on veins. Inflorescence axillary, fasciculate; pedicels up to 2 cm

long, with a dense, long, brown-red indumentum. Flowers with sepals lanceolate, long acuminate, 0.8-1.3

x 0.2 cm, with a dense, long, brown-red indumentum; corolla oblique in the calyx, with red trichomes

outside, basal gibbosity 1-1.5 x 2 mm, tube cylindric, 1.8-2 cm long, lobes suborbicular, 0.8 x 0.6 mm,

undulate at margin. Fruit not seen.

Distribution.—Paradrymonia maguirei is known only from the type collection from the area North of La

Esmeralda in the Duida-Marahuaca National Park, on a forested slope of the Cerro Marahuaca (Amazonas,

Venezuela), 65?24"W 3°40'N according to maps, at 1000 m elevation. It was blooming in May.

The color of the corolla is not known. There are two specimens that I marked A and B on the herbarium

sheet in the New York herbarium. Specimen A, including the fragments in the pocket, is the type of the new

species; it is on the middle left of the sheet and the pocket on the lower right above the label. Specimen B

(sterile), on the upper right, is probably Nautilocalyx cordatus (Gleason) L.E. Skog. The only open corolla,

preserved in the pocket, is very unlike the corolla of N. cordatus and is the one described here. The affinities

of Paradrymonia maguirei in the genus are not clear. This species shows a unique combination of characters:

rosette-like habit, petioles short, with dense, long, appressed pubescence, leaf blade drying paper-thin,

asymmetric at the base, broadly rounded at the apex, and sharply biserrate at the margin. Other species

with short petioles have leaf blades long-decurrent or are long stemmed epiphytic climbers. This species

was Paradrymonia “sp. E" in the text and the key (Feuillet & Steyermark 1999).

Etymology.—The epithet maguirei refers to the senior collector, a great botanist, collector, and student

of the flora of the Guiana Shield.

B.—Subgenus Pagothyra

Like other infrageneric taxa, Episcia sect. Pagothyra Leeuwenb. coined for E. maculata Hook. f. (Leeuwen-

berg 1958: 312) was never transferred to Paradrymonia, although P. maculata stands alone there as well as

in Episcia.

Paradrymonia subg. Pagothyra (Leeuwenb.) Feuillet, comb. et stat. nov. Basioww: Episcia sect. Pagothyra Leeuwenb.,

Blumea 7:312. 1958

Feuillet, New species of Paradrymonia

The Sow Virk ülantcal Gordon

Planie al

THE RUNUARDT VESEZEELAN FAPEDIMOY gsis gs

Verri Xacahboaza, Teitilnoiia (ee) cman

^a 29155

f Kd wA 0o, Sa d

ee LA d

Siape forest, alt. 1000 meters.

WBasaedi Ya

salia

lassi Maguire, Ji

Way 5,

—— dai NT E IST

Fic. 1. Parad,

B. Maguire & B. Maguire, Jr. 29185 (NY)

j photograph by € F

136 Journal of i i Texas 3(

Paradrymonia maculata (Hook. f.) Wiehler, Selbyana 5:57. 1978. Episcia maculata Hook. f., Bot. Mag. 116: pl. 7131.

1890. [Nautilocalyx maculatus] Wiehler 1970, invalid: in Varie Tere: ee Origin GUYANA: Cult. Hort. Kew.. 2 Sep 1889

(fl), collector unknown s.n. (HOLOTYPE: K; soTyPE: K; pl type BH, NY, U, US, WAG).

Paradrymonia maculata climbs by way of short roots at the nodes and along internodes, similar to Hedera

helix L. The stems are tightly applied to the bark and have been reported to grow 1-3 meters high. The leaves

are opposite and equal or subequal in a pair with a long petiole; the blade is large and broadly elliptic, with

serrate margins. The pedunculate inflorescences are axillary, unilateral cymes, with bracts that are large,

greenish yellow with purplish or reddish veins. The corolla is creamy yellow with reddish dots and the

ventral lobe acts as a cover closing the tube. Longer descriptions can be found in Leeuwenberg (1958) and

Skog & Feuillet (2008).

Distribution.—Paradrymonia maculata is known from the forests of French Guiana, Guyana, and Ven-

ezuela (Delta Amacuro) at 0—500 m. It has been collected in bloom every month of the year and in fruit in

March, June, August, and December.

Corolla.—As noted and illustrated in Hooker (1890), the ventral lobe of the corolla of Paradrymonia

maculata closes the tube (Fig. 2). The nectary gland is in dorsal position at the base of the ovary, next to the

basal gibbosity forming a nectar chamber at the base of the 2.7-3.5 cm long corolla tube. The ventral corolla

lobe is effectively a barrier between the nectar produced at the base of the tube and most pollinators. Exert-

ing pressure on both sides on the apical third of the tube flips the ventral lobe from its position of convex

lid closing the throat to a more classic concave corolla lobe by moving it more than 90°. It allows access to

the nectar to strong pollinators, possibly carpenter bees. The bracts and sepals are pale yellow with red or

purple veins. In the Guianas similar corolla morphology is found in a Solanaceae, Marhea formicarum Dam-

mer, where the lower lobe closes the throat. That species is lacking bracts, but the large calyx is cream- or

straw-colored with purple veins, showing a similar color pattern as the bracts and calyces of P. maculata. It

might be of interest to note that an Asian Gesneriaceae, Agalmyla chorisepala (C.B. Clarke) Hilliard & B.L.

Burtt, has orange corollas with the ventral lobe closing the tube, but in this example the lobe flips at anthesis

and opens the access to the tube without further obstacle to pollination.

The type of Paradrymonia maculata (Hook. f.) Wiehler from an unknown collector, comes from a plant

cultivated at the Royal Botanic Gardens, Kew. It was blooming in September 1889. The plant was grown

from material collected in British Guiana, now Guyana. Subsequent collections all came from Guyana and

this limited distribution was acknowledged by Leeuwenberg (1958) and Wiehler (1978). During the comple-

tion of the treatment of the Gesneriaceae for Flora of the Guianas (Skog & Feuillet 2008), collections of P.

maculata from Venezuela and French Guiana came to my attention. That species was not mentioned in the

treatment for Flora of the Venezuelan Guayana (Feuillet & Steyermark 1999) but is present in the Antonio

Díaz Department, the part of the state of Delta Amacuro neighboring Guyana. No collections from Surinam

have been made, but it is likeiy that it is, or has been, present there.

Material studied: VENEZUELA. Delta Amacuro. Depto. Antonio Díaz: low forest, 12 km S of San José de Amacuro, 8?28'N 60°27'W,

sea level, Feb 1987 (fD, A. Fernandez 3903 (MO, MYF, NY, PORT, US); primary rain forest, Río Grande, 60 km NE of El Palmar, about

8°25'N 61°45'W, 120 m, 15 Mar 1987 W, e ana 5411 (PORT).

GUYANA. Arawai Creek, ight! Riv., Feb 1952 (fD, C.A. Persaud 140 = F.D. (Forestry Sd 6897 (K, NY, U);

Essequibo River, cain IE Groete Creek, 14 Ap: 1944 (f), B. Maguire & D.B. Fans) 22826 (F, GH, K, NY, U, US); Morawhanna,

Oct 1905 (fD, A.W. B 00 (K); Rockstone, 15 July — 1 Aug 1921 (fl), H.A. Gleason 664 (NY); Unabaruka p Aug 1930 (fD, E.B.

a 225 (BRG). Barima. Waini: Barima Riv., 15 mi E of Arakaka, 7°37'N 59°54'W, 38 m, 26 July 1986 (fD), J.J. Pipoly 8059 (BRG, CAY,

Y, US); Barima Riv., May 1907 (fl), R.D. Ward s.n. (K); Baramita airstrip — Millionaire trail, 7°22'N 60°28'W, 91 m, 3 Apr 1991 (fl), T.D.

o et al. 4182 (NY, US); Matthews Ridge, Barima Riv., 23 Jan 1955 (fl), R.S. Cowan 39337 (NY, US); T between Aruau Riv. &

Yarikita Riv., 8°00'N 59°55'W, 17 Jan 1920 (fT), A.S. Hitchcock 17601 (GH, K, NY, S, US); Sebai Riv., + 5 km of Sebai Village, 15-20

m, 7°49'N 59°57'W, 16 Dec 1991 (fr), B. Hoffman et al. 615 (US); Upper Aruau Riv., Áruka Riv., Apr-May 1929 > E.B. Martyn 53 (K);

Upper Kaituma R.; 3 km w of Port Kaituma, 7°42'N 59°54°W, 0—5 m, 8 Dec Mid (fD, B. Hoffman & H. Benjamin 525 (US); Waini Riv.,

July 1906 (fn, J. E. Beckett s.n (K, U). Cuyuni Mazaruni: Aurora. heli 1; 6°47'N 59?44"W, 4 Oct 1989 AD, LJ. Gillespie 2084

(US; Bartica, 12-15 mi f t 28 Aug 1935 (fr), D. Potter 5358 (GHD: Essequibo county, near Mazaruni Forest Station, 9 Aug 19

(fD, W.A. Archer 2432 (BRG, K, US); id., July-Sep 1942 (fD, D.B. Fanshawe 785 = F.D. 3521 (K). Essequibo Islands. West Demerara:

Macouria Riv., right bank of Lower Essequibo Riv., Nov 1886, G.S. Jenman 2419 (K); Upper White Creek, near Blue Mountain, 6°35'N

Feuillet, New species of Paradrymonia 137

Fc? Pp Ir ; lata, phot ph by Chris D

58°43'W, 5-20 m, 14 Apr 1993 (fD, T.W. Henkel et al. 1879 (NY, US). Pomeroon. Supenaam: Pomeroon District, Mt. Russell, Mar 1886

(fl), G.S. Jenman 2097 (K); Pomeroon Riv., 20 Aug 1959 (fI), V. Graham 352 (K); Pomeroon Riv., Yawiami Creek, Aug 1882 (fI), G.S. Jen-

man 1939 (K); Pomeroon Riv., Pomeroon apis 17-24 Dec 1922 (fl), J.S. de la Cruz 3124 (F, GH, MO, NY, PH, UC, US); id., 14-20 Jan

1923 (fl&fr), J.S. de la Cruz 3022 (GH H, US); id., Mar 1884 (fD, G.S. Jenman 2002 (K, NY); Abrahms Creek, Mar 1904 (st), "G.S.

Jenman” 7808 ER 3km Pu of AM Mission Mig sa 7?15'N 58°45'W, 0-10, 25 Sep 1992 (fl bud), B. Hoffman & L. Roberts 2839

(NY, US). Potaro. S : Garraway stream, 102,5 mi on Bartica — Potaro rd., 5°22'25"N 59°7'20"W, 38 m, 12 Mar 2004 (fl), K.M.

Redden 2231 (US); ieee inf R pukari — Annai Road, 4°28'14"N 58?47'16"W, 400—500 m, 21 Mar 1997 (fD, H.D.

Clarke, S.A. Mori €: S. Heald 4181 (US); Kaieteur a '5°10'N 59°29'W, 23 Oct - 3 Nov 1923 (fD, J.S. de la Cruz 4395 (F, GH, NY, PH,

US, VEN); Potaro, 10 mi S of Potaro landing, 5°10'N 59°00'W, 7-8 Jan 1920 (fl), A.S. Hitchcock 17397 (GH, K, NY, S, US); North Fork

Riv., 0.5-1.5 km N of Konawark Riv., 5°9'N 59°8'W, 137 m, 18 May 1991 (fl), T.D. McDowell, C.L. Kelloff & A. Stobey 4819 (US). Upper

Demerara. Berbice: Haiowa Falls, Essequibo Riv. basin, 5°7'N 58°49'W, 27 Sep 1937 (fD, A.C. Smith 2123 (F, G, GH, K, NY, S, U, US);

Mabura as W inn: compartment, 5°01'95"N 58?37'73"W, 12 Oct 1993 (fI), R.C. Ek, P.].M. Maas, H. Mass & C. Górts 942 (U, US);

Mabura region, Holder Falls, 5°20'N 58°10'W, 21 Aug 1993 (fI), R.C. Eh, R. Zagt, L. Brouwer & N. Eernisse 896 (US).

Upper Takutu. Upper Esequibo: Maparri R., S bank, 3°20'N 59°15'W, 3 June 1996 (st), H.D. Clarke & T. McPherson 1928 (US); Upper

Rupununi Riv., near Dadanawa, 2°45'N 59°31'W, 13 June 1922 (fl&fr), J.S. dela Cruz 1518 (CM, F, MO, NY, PH, US); id., 13 June 1922

(fl), J.S. de la Cruz 1535 (CM, GH, F, MO, NY, PH, UC, US)

138 Journal of the Botani Insti Texas 3(1)

FRENCH GUIANA. Approuague Riv. Basin: Crique Cascade, 390 m, 15 Mar 2002 (fl&fr), J.F. Smith, E. Teppe & C. Davidson

4134 (CAY). Matoury: Plateau de Nancibo, 4°40'N 520°30'W, 24 Oct 1983 (fI), F. Billiet & B. Jadin 1857 (BR, CAY). Oyapock Riv. Basin:

Crique Gabaret, 3°55'42"N 51?48'7"W, 15 Apr 1988 (fD, G. Cremers 9951 (CAY, NY, P, U, US); Roche Touatou, 130 m, 20 May 1995 (fD,

J.-J. de Granville & G. Cremers 13009 (CAY).

ACKNOWLEDGMENTS

I thank Alain Chautems and an anonymous reviewer for their careful reading of the manuscript. I am grate-

ful to Eduardo Garcia-Milagros who translated the abstract into Spanish. This work could not have been

completed without the help of the curators of the herbaria CAY, K, MO, NY, P, and VEN who made available

to me the material in their care. This is number 144 in the Smithsonian's Biological Diversity of the Guiana

Shield Program publication series.

REFERENCES

CLARK, J.L., PS. HERENDEEN, L.E. SKOG, AND E.A. Zimmer. 2006. Phylogenetic relationships and generic boundaries in the

Episcieae (Gesneriaceae) inferred from nuclear, chloroplast, and morphological data. Taxon 55:313-336.

FEUILLET, C. AND J.A. STEYERMARK. 1999. Gesneriaceae. In: Steyermark, J.A., PE. Berry, K. Yatskievych, and B.K. Holst, Flora

of the Venezuelan Guayana, vol. 5. Missouri Botanical Garden Press, St. Louis. Pp. 542-573.

HANSTEN, J. 1854. Die Gesneraceen des Kóniglichen Herbariums und der Gárten zu Berlin, nebst Beobachtungen

über die Familie im Ganzen I. Abschnitt. Linnaea 26:145-216; fig. 1-68.

HANSTEIN, J. 1864. Gesneraceae. In: Martius, Flora Brasiliensis 8(1):341—428; pl. 58-68. Fleischer, Leipzig.

Hooker, J.D. 1890. Paradrymonia maculata. Bot. Mag. 116: pl. 7131.

LEEUWENBERG, A.J.M. 1958. The Gesneriaceae of Guiana. Acta Bot. Neerland. 7:291-444.

SkoG, L.E. AND J.K. BOGGAN. 2006. A new classification of the Western Hemisphere Gesneriaceae. Gesneriads

56(3):12-17.

Skoc, L.E. and C. FEuuLEr. 2008. Gesneriaceae. In M.J. Jansen-Jacobs, ed. Flora of the Guianas ser. A, 26. 136 pages.

Royal Botanic Gardens, Kew.

WEBER, A. 2004. Gesneriaceae. Pp. 63-158 in K. Kubitzki and J.W. Kadereit, eds. The families and genera of vascular

plants, Dicotyledons. Lamiales (except Acanthaceae including Avicenniaceae) vol. 7. Berlin: Springer.

Wiener, H. 1973. Seven transfers from Episcia species in cultivation (Gesneriaceae). Phytologia 27:307-308.

WienLer, H. 1978. The genera Episcia, Alsobia, Nautilocalyx, and Paradrymonia (G iaceae). Selbyana 5:1 1-60.

NEW SPECIES OF BACCHARIS (ASTERACEAE: ASTEREAE) FROM

RIO DE JANEIRO STATE, SOUTHEASTERN BRAZIL

G. Heiden J.F.A. Baumgratz

. Sc, CNPq-PROTAX Fellow Diretoria de Pesquisa Científica

Escola pisa de Botánica Tropical Instituto de Pesquisas

uto de Pesquisas Jardim Botánico do Rio de Janeiro

Jardim as do Rio de Janeiro Rua Pacheco Leão 915

Pacheco Leáo, 915, Jardim Botánico 22460-030, Rio de Janeiro, RJ, BRAZIL

Rio de Janeiro, RJ 22460-030, BRAZIL

gustavo.heidenggmail.com

R.L. Esteves

Universidade do Estado do Rio de Janeiro

Cen dico

Departamento de Biologia o e ~

Rua São Francisco X

Maracanã, 20550-900, Rio de A T BRAZIL

ABSTRACT

m i f NE. 1 pi. det . n +7 4 du A +11 IR 1 sc alti

Heiden, Baumgratz & Esteves, assigned to sect - Caulopterae DC., is characterized by th ] hes, female capitulum with

5 "x | je] Tod 2 n ] i fth

urceolate or E glcally P

ki ii sgeniseades CORDE * pu friburgensis, assigned to sect. Oblongifoliae DC., is cl ized by tl i l

J 1 Af 1 n 1 À f, 1 f1

E > o

odas dor 1] d tl } istics diffi iate it f B hylla D Additionally. keys to identify the spe-

cies of sect. Caulopterae Anda sect. Oblongifoliae occurring in the state of Rio de Jairo are provided.

RESUMO

Duas novas espécies de Baccharis do estado do Rio de Janeiro, sudeste do Brasil, sáo descritas e ilustrada RI A

ee A & Esteves, pos á sect. E E DC., é ee Bod ramos jid M

[Al

cer

proximas do "(c I 1 R x 11.53 Dp 1 = f...21. g RLW PP | D g e A Oblongifoliae

DC., é esa a pelas folhas cc coriáceas, com 2 agudo ou obtuso, gem intei lut panicula een capitulos

I ápice d la denticulado, isti ] B. macrophylla

Dus é fol imilar. Adicionalmente, são f id } d inação d écies d có

e Pr. ocorrentes no estado do Rio de Janeiro.

INTRODUCTION

Baccharis was never revised taxonomically as a whole. The last worldwide compilation of accepted scientific

names and synonyms of the genus was published by Malagarriga (1977). Following Müller (2006), infrage-

neric classification of Baccharis is still not well resolved, and, additionally due to the high species number,

revisionary work in the genus is only possible with a geographical approach. For Brazil, the last revisionary

work of the genus was provided by Barroso (1976), comprising 125 species. Later, Oliveira et al. (2006)

published a compilation of 146 names, without citing vouchers to assure some occurrences of species not

listed by Barroso (1976) in the country. The genus occurs in all Brazilian biomes and most of its species are

concentrated in the central and eastern regions of the country, growing mainly in savannas (cerrado) and

grasslands (campos de altitude, campos rupestres and campos sulinos). For the state of Rio de Janeiro information

concerning the genus is available only in geographically restricted florulas (Barroso 1957, 1959; Esteves &

Barroso 1996) or in the taxonomic revision for Brazil (Barroso 1976).

J. Bot. Res. inst. Texas 3(1): 139 —145. 2009

140 i i Texas 3(

During the taxonomic revision of Baccharis for the flora of the state of Rio de Janeiro two new species

belonging to the sect. Caulopterae DC. and sect. Oblongifoliae DC. were recognized. These new species are

described and illustrated, and keys to identify the species of these sections occurring in Rio de Janeiro state

are presented.

MATERIAL AND METHODS

The study has been based on literature revision and the examination of material or photos from herbaria

GUA, HAS, HB, HBR, HPNI, HRJ, M, P, R, RB, RBR, RFFP, RUSU and SP. Specimens of the related taxa

contrasted to the new taxa are cited just after the description and before the keys. The terminology used

in the morphological descriptions is based on Radford et al. (1974) and Muller (2006). The measures were

taken by the mean of a digital calliper in the wider portion of the structures. All colours mentioned are of

dried mature structures. The illustrations were prepared by stereomicroscopy using a Carl Zeiss Stemi SV6.

The conservation status is presented following the IUCN (2001) guidelines.

TAXONOMY

Baccharis alti t Heiden, B tz & Esteves, sp. nov. (Fig. 1). Tyre: BRAZIL: Rio DE Janeiro: Teresópolis, Serra

dos Órgãos, 27 Nov 1933, 9 Brade 12510 E RD.

+ollnid qr J s ti-natentibi s (non

Ad Baccharis sect. acia DC. pa B. iade (Lam.) Pers. subsp. geni p us (non

erectis) munitis j 9-4 mm ee s 4— of mm Hiep dde Pad foemineorum 2.5-3.4 mm longis

f] f.

(non 3.5-6.8 mm longis),

Dioecious subshrubs 0.2—0.4 m tall, erect-patent or prostrate. Stems and branches 3-winged, wings 0.5—6.5

cm long, 0.3-1.6 cm wide, plane to undulate, strongly resinous, seemingly glabrous, indument tufted, tufts

appearing under magnification as small resinous dots. Leaves 0.3-2 mm long, 0.2-0.5 mm wide, bract-

like. Panicles 18-24 cm long, of pseudospikes 3-18 cm long, capitula solitary or 23, sparsely along the

axis, the main pseudospike longer than the axillary ones. Male capitula 4—5.5 mm long, involucre 4.4-6.5

mm long, 4-6.8 mm wide, urceolate to campanulate; pyllaries in 5-8 series, abaxial surface pale yellow to

dark, outer and median phyllaries ovate, inner elliptical, apex entire or denticulate, margins light yellow,

denticulate, scarious; clinanthium obconical, plane, alveolate, densely covered by biseriate trichomes. Male

florets 35-63; corolla 2.9-4 mm long, 5-laciniate, tube 1.8-2.5 mm long, throat 0.2-0.4 mm long, cup-

shaped, lobes 1-1.4 mm long, externally with sparse biseriate trichomes on the throat and tube; styles 3.2-4

mm long., apex 1/2-bifidous; pappus 3.2-4.2 mm long, uniseriate, apex slightly broadened, terminal cell

ends slightly protruding. Female capitula 6.5-8 mm long; involucre 4.4-6.5 mm long, 4-6.8 mm wide,

urceolate to campanulate; phyllaries in 6—9 series, abaxial surface pale yellow to dark brown or olive green,

outer ovate, median ovate to elliptical, internal elliptical, apex entire or denticulate, margins light yellow,

denticulate, scarious; clinanthium obconical, alveolate, densely covered by biseriate glandular trichomes,

paleae absent. Female florets 34-42; corolla 2.5-3.4 mm long, filiform, apex shortly ligulate, 3-5 teeth of

unequal size; styles 3.1—5.1 mm long, branches 0.5-0.8 mm long. Cypselae 1.2-1.5 mm long, cylindrical,

cal papillose, A pappus 4—4.7 mm long, uniseriate, not-accrescent.

S ined. BRAZIL. Rio de J iaia, 4 Nov 1965, Eiten 6612 (RB); 12 Sep 2007, Heiden & Baumgratz 823 (RB); 17 Dec

2007, Heiden 932, 933 (RB); 13 Apr 2008, Heiden 994 (RB); 18 Oct 1977, Landrum 2103 (RB); 5 Dec 1964, Vianna 207 (GUA, RB). Nova

Friburgo: 22 Mar 2008, Heiden & Bcumgratz 711 (K, RB). Teresópolis: 27 Jun 2007, Heiden 791, 792 (RB); 26 Sep 2007, Heiden 879, 880

(RB); 21 Mar 2007, Nadruz 1772 (RB); 6 Sep 1981, Ribeiro 138 (GUA); Jan 1952, Vidal 11-173, 11-175 (R); 12 Feb 1952, Vidal 11-551 (R) Feb

1952, Vidal 11-662 (R), 15-20 Dec 1952, Vidal 11-5768 (R); Feb 1953, Vidal 1478 (R).

Distribution & ecology.—Baccharis altimontana is restricted to southeastern Brazil, known up to now only

from the state of Rio de Janeiro. It occurs in the Atlantic Rain Forest biome, above 1900 m, in the massifs

of Itatiaia (Serra da Mantiqueira), Serra dos Órgãos, Pico da Caledônia and Pico do Desengano (all of them

belonging to the Serra do Man). It grows in rather small populations in rock grooves on bare rock outcrops

along the high altitude grasslands, preferably in places with constant wetness. Flowering and fruiting time

is between September and December.

Haidan ef al A : £D L ?- 4 a + D A |

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rot

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Fic. 1 D hon DI. m Uni Duce te 0 Estoune: A Hahi f € I lant) D. Mal

pit | C. Male fl (I pl 1); D. Female

capitulum; E. Phyllaries from female capitulum; F. Femal lla and style; G. Cypsela. Scale I A: 3 cm; B, D: 2 mm; C, E, F, G: 1 mm. (A: Brade

12510; B, C Heiden 879; D, E, F, G Heiden 823). Drawn by Joao Iganci.

Das H ID Ll titt £T

142 Journal of ti Texas 3(

Vernacular—Carqueja (Ribeiro 138; Heiden 711, 791, 792, 823, 879, 880, 932, 933, 994).

Conservation.— Common in high altitude grasslands, the species is considered near threatened (NT)

due to the loss of quality of the mountainous environments caused by anthropogenic pressures.

Etymology.—The name ban to the habitat along the summits of mountain ranges.

Notes.—Baccharis alti igned to the sect. Caulopterae DC., mainly due to presence of winged

stems, epaleaceous clinanthia densely red with biseriate glandular hairs, papillose glabrous achenes,

and pappus bristles of female flowers enlarged basally and Med into a ring. The species belongs to the

“Baccharis genistelloides Complex” ing at least eight } lly close taxa [B. crispa Spreng., B.

genistelloides (Lam.) Pers. subsp. ES B. genistelloides subsp. lorentzii Joch. Müll., B. jocheniana Heiden

& Macias, B. myriocephala DC., B. opuntioides Mart. ex Baker and B. riograndensis Malag. & J. E. Vidal]. The

new taxon is closer related to Baccharis genistelloides subsp. genistelloides, which occurs along the Andes from

Colombia to northern Chile and Bolivia (Müller 2006). The studied specimens of B. altimontana were found

in iota aa as B. ena P aio: B. myriocephala, B. puntada or B. trimera (= B. crispa).

1 of B. genistelloides subsp. genistelloides by male corollas 2.9—4

mm don female corollas 2. 5 4 mm long and female florets with uniseriate pappus (vs. male corollas 4-6

mm long, female corollas 3.5-6.8 cm long and female florets with multiseriate pappus).

Specimens of B. altimontana are commonly identified as B. opuntioides, since Barroso (1976) merged

both species in a broad circumscription of B. opuntioides. However, we consider B. opuntioides as endemic to

the Serra do Caparaó at the boundary between the states of Espírito Santo and Minas Gerais. This species

may be recognized by erect branches with short wings, 0.31.8 cm long and 0.2-0.8 cm wide, pseudospikes

with apically crowded capitula (in the apical 1-2.5 cm), with the secondary branches longer than the main

axis of the capitulescence, male corolla 4—5.2 mm long, female corolla 4.2—5 mm long and cypselae 0.8-1

mm long.

Baccharis may be distinguished from B. crispa by erectopatent or prostrate stems and shortly

ligulate corollas of female florets with teeth of irregular size (vs. erect stems, corolla of female florets trun-

cate or denticulate), and from B. myriocephala by the height (0.2—0.4 m tall), panicles 18-24 cm long with

pseudospikes 3-18 cm long, involucre of female capitulum urceolate to campanulate (vs. height 100—250

cm, panicles 25-80 cm long, pseudospikes 10-35 cm long and involucre of female capitulum cylindrical).

Ted £

Selected action specimens:

Baccharis cri Spreng., Syst. veg. 3:466. 1826. BRAZIL. Rio de Janeiro: Itatiaia, 13 Apr 2008, Heiden 978, 993 (RB); Mar 1894, Ule

177 (R); ene 12 Nov 2007, Heiden & Baumgratz 815 (RB); 13 Sep 2007, Heiden €- Baumgratz 833, 834, 850, 851 (RB). URUGUAY.

Montevideo: 1821/1822, Sellow d397 GsoTYPE: R).

Baccharis genistelloides (Lam.) Pers., Syn. pl. 2:425. 1807. BOLIVIA. Santa Cruz: 13 Nov 1959, Maguire & Maguire 44496 (RB)

QUADOR. Cotopaxi: “province de Tacunga,” Jussieau s.n. (P-Lam, photo of holotype). Pichincha: San Juan, 28 Nov 1952, Fagerlind

& Wibom 1556 (R)

Baccharis myriocephala DC., Prodr. 5:426. 1836. BRAZIL. Minas Gerais: Vauthier 265 (G-DC, photo of lectotype). Rio de Janeiro:

Miguel Pereira, 18 Apr 2007, Heiden & Baumgratz 727 (RB). Resende, 26 Jul 1966, Eiten 7447 (RB); 12 Nov 2007, Heiden & Baumgratz 802,

803 (RB); Rio de Janeiro, Jun 2000, Esteves & Esteves 462 (RB); Santa Maria Madalena, 14 May 2007, Heiden 734, 735 (RB); Teresópolis,

27 Jun 2007, Heiden 779, 780 (RB); 26 Oct 2007, Heiden 878 (RB).

B h M ex Baker in Martius, Eichler & Urban, Fl. bras. 6(3):39. 1836. BRAZIL. Espírito S /Mi G

Caparaó, 15 Naw 1960, Flaster 101 (GUA); Nov 1922, Lobo s.n. (R 37944); 8-12 Mar 1917, Lutz 1197 (R); Manse n. (M, photo of holotype);

21 Oct 1947, Moreira 44 (R); 11 Jul 1998, Oliveira 1468 (GUA); 29 Jun 1950, Santos s.n. (R 52182); 2 Jul 1888, Schwacke s.n. (R 37949

KEY TO BACCHARIS SECT. CAULOPTERAE IN RIO DE JANEIRO STATE, BRAZIL

. Stems aptero

2: cm blades c coriaceous, margins enre, m. 3 d Muy basal veins; panicles 1.5-6 cm long B. organensis

2. Leaf b nately veined or with 3 acrodromous 1-5 mm distant from

ES base suprabasal veins; panicles 4-16c cm ant B. regnellii

inged

1. Stems w

3. Leaves well-developed.

4. Subshrubs (0.5-1 m tall); | lospike branches 5-20 cm long B. junciformis

Heiden etal., N peci f Baccharis f ti tern Brazil 143

4. Shrubs (0.5-3 m tall); pseudospike branches 0.5-5 cm lon

5. Wings 1-3 mm wide; leaf blades coriaceous, with attenuate base B. burchellii

5. Wings 5-13 mm wide; leaf blades papiraceous, with cuneate or rounded base B. glaziovii

3. Leaves bract-like.

DEGREES EIS ende corolla aN or metan Ada B. crispa

E , Scande | | ligulate.

7. Subshrub 02- 04 m tall; S 3-18 cm m een solitary or 2-3; involucre of female

capitulum urceolate to campanulate B. altimontana

7. Subshrub 1-2.5 m tall; pseudospikes 10-35 cm long; capitula solitary or gl les with 2-8 zs

involucre of female capitulum cylindrical B. myriocephala

Baccharis friburgensis Heiden, Baumgratz & Esteves, sp. nov. (Fig. 2). Tre: BRAZIL: ih DE JANEIRO: Nova Friburgo,

Macaé de Cima, 16 Aug 1989, 3, Vieira 48 (HoLorYPE: RBI; isotypes: FCAB!, GUA!, NY, RI, RB!, RBR!, SP

Ad Baccharis sect. Oblongifoliae DC. pertinens, B. dll ndi Papi Md coriaceo, dbi ai es in d cuin contracta

similis, sed capitulis foemineis 2 ad 3 floribus (non 11 ad 21 ;

et capitulo masculo 8 ad 12 floribus (non 30 ad 45 floribus) munito differt.

Dioecious shrubs 1.5-2 m tall. Stems erect, branching dichotomous, shoots glabrescent, with uniseriate

and pedestal trichomes. Leaves spirally alternate, crowded at the apex of the branches, with petioles 0.2-0.8

cm long; blades 30-68 mm long, 7-21 mm wide, coriaceous, narrowly elliptic or oblanceolate to obovate,

base attenuate, apex acute to obtuse, margins entire, revolute; pinnately veined; both surfaces seemingly

glabrous, abaxial surface with tufted indument appearing as resinous dots, scarce biseriate, uniseriate and

pedestal trichomes among the tufts. Panicles 1.9-4.4 cm long, 1.8-5.5 cm wide, corymbose, terminal;

peduncles 1-1.8 cm long. Male capitula 2.9-4.8 mm long, involucre 2.9-4.8 mm long, cup-shaped, 2.75

mm wide; phyllaries in 3-5 series, brown, outer and median phyllaries ovate, inner linear-ovate, apex fimbri-

ate, margins light-brown, shortly dentate; clinanthium convex, glabrous; paleae absent. Male florets 8-12,

apex 5-laciniate; corolla 2.3-2.5 mm long., tube 0.9-1.1 mm long, externally densely covered by biseriate

trichomes near the apex, throat 0.5-0.6 mm long, cup-shaped, lobes 0.7-0.8 mm long, apex not revolute;

anthers included; styles 2.4—2.6 mm long, apex capitate by sweeping hairs of unequal size, slightly exserted;

sterile ovary 0.5-0.7 mm long; pappus 2-2.3 mm long, uniseriate, apex broadened, cell ends erectopat-

ent, shortly protruding. Female capitula 5.3-5.8 mm long, involucre 4.3-5 mm long, 2.2-3.1 mm wide,

cylindrical; phyllaries in 4—6 series, brown, outer phyllaries ovate, median linear-ovate, inner linear, apex

long-fimbriate, margins light-brown, shortly dentate; clinanthium convex, paleae linear, acute, deciduous.

Female florets 2-3, filiform; corolla 2.9-3.2 mm long, apex 5-denticulate, distal half with subapical scat-

tered biseriate trichomes; styles 3-3.5 mm long., branches 0.3-0.6 mm long. Cypselae 2-2.6 mm long,

0.4-0.6 mm wide, light brown, cylindric, narrowed at both extremities, with 5-6 longitudinal ribs, with

scattered biseriate trichomes near the apex; pappus 2.5-3 mm long, slightly shorter to slightly longer than

the style, biseriate, persistent, not accrescent at maturity.

Additional specimens examined. BRAZIL. Rio de Janeiro: Nova Friburgo, 9, 24 Aug 1986, Leitman 196 (K n.v.; RB-2x); d, 19 Aug

1990, Moraes 414 (RB-2x).

Distribution & ecology.—Restricted to southeast Brazil (state of Rio de Janeiro). It occurs in the Atlantic Rain

Forest biome, from 1000 to 1500 m above sea level, and is endemic to the summits of the Macaé de Cima

Mountains, a part of the Organ Mountains/Serra do Mar Ranges. It grows directly on rock outcrops, along

river headwaters or in edges of nebular forest. Flowering and fruiting in August

Conservation.—Taking into account the few collections available, the restricted area of occurrence and

the presence of anthropogenic pressure in surrounding areas, represented by cattle grazing, agriculture,

tourism and urban expansion, the species is considered endangered (EN B2abliiil).

Etymology.—The name refers to the type locality in the municipality of Nova Friburgo.

Notes. —Baccharis friburgensis is assigned to sect. Oblongifoliae DC. for the pinnately veined leaves,

capitate style apex in male flowers due to sweeping hairs of unequal size, paleate female capitula and female

flowers with the pappus not elongated at cypsela maturity. The species is remarkable for the few-flowered

Fc? R n 25 £pl A] D £5 0 Cop AD le aff, I lene. D AL allan £. f" Mal sel m Mala £l

dct

AE E I nul

t papi

Phyllari itul G. Palea: H. Fi ji d style: 1. Cypsela. Scale bar near

Ld * £1

A: 2 cm; B, D, F, G, H, I: 1 mm; C, E: 2 mm (A, B, E, F, G, H, I: Leitman 196; C, D: Vieira 48). Drawn by Joao Iganci.

Heiden et al., N pecies of Baccharis fi theastern Brazil 145

male (8-12) and female 2-3) capitula, which results in seemingly epaleaceous female clinanthia, because

all paleae are, due to the low flower number, more distal then the distalmost flowers and hidden by the

innermost phyllary series. Some further distinguishable characters are the coriaceous leaves with acute to

obtuse apex, entire and revolute margins, and the corymbose panicles. Baccharis friburgensis is vegeta-

tive morphologically similar to the allopatric B. macrophylla Dusén, although this similarity seems to be a

convergence to the habitat in sunny habitats on summits of mountains. The later one has leaf margin more

commonly with 1—5 subapical teeth, seldom entire, male capitula with 30—45 flowers, female capitulum

with 11-21 flowers, and female corollas ligulate.

d ane o

la Dusén, Arq. Mus. Nac. Rio de Janeiro 13:14. 1905. BRAZIL. Rio de Janeiro: Itatiaia, Dusén 4844 (HOLOTYPE:

R); 23 Jul 1960, Handro 941 (SP); 17 Dec 2007, Heiden 930 (RB); 16 Aug 1969, Sucre 5775 (CEPEC); s.d., Tamandaré & Brade 6562 (SP);

Resende, 12 Sep 2007, Heiden & Baumgrat. 7, 828 (RB); Jul 1902, Moreira & Teixeira s.n. (R 38027

KEY TO BACCHARIS SECT. OBLONGIFOLIAE IN RIO DE JANEIRO STATE, BRAZIL

1. Panicles pyramidate.

2. Foliar indument ferruginous B. rufidula

2. Foliar indument not ferruginous B. oblongifolia

1. Panicles corymbose.

3. Leaf blade papyraceous, margins not revolute B. grandimucronata

3. Leaf blade mE margins revolute

4. Male capitula 30-45 florets; female capitula 11-21 florets, female corollas with ligulate apex —— B. macrophylla

4. Male capitula 8- E 20 florets, female capitula 3- d florets, female corollas with denticulate apex. B. friburgensis

ACKNOWLEDGMENTS

The authors acknowledge the consulted herbaria, André Scarambone Zaú, Cecilia Cronemberger de Faria,

Denise Pinheiro da Costa, Izar Aximoff, Marcus Nadruz and Marina Wolowski Torres for providing field

support, Joào Iganci for the illustrations, Jochen Müller for the valuable comments and an anonymous

reviewer.

REFERENCES

Barroso, G.M. 1957. Flora do Itatiaia—Compositae. Rodriguésia 32:175-241.

Barroso, G.M. 1959. Flora da cidade do Rio de Janeiro—Compositae. Rodriguésia 33-34:69-155.

Barroso, G.M. 1976. Compositae—Subtribo Baccharidinae Hoffman. Estudo das espécies ocorrentes no Brasil.

Rodriguésia 28(40):3-273.

Esteves, R.L. AND G.M. Barroso, 1996. Compositae. In: Lima, M.PM. & Gutpes-BRuNI, R.R. Reserva Ecológica de Macaé

de Cima: Nova Friburgo—RJ: aspectos florísticos das espécies vasculares. Rio de Janeiro: Jardim Botánico do

Rio de Janeiro, 1996, 2:189-245

INTERNATIONAL UNION FOR CONSERVATION OF NATURE AND NATURAL Resources (IUCN). 2001. IUCN Red List Categories and

Criteria, Version 3.1. IUCN, Gland, Switzerland and Cambridge, United Kingdom. http://www.iucn.org.

MALAGARRIGA Heras, R.P. 1977. Nomenclator baccharidinarum omnium. Mem. Soc. Ci. Nat. La Salle 37:129-224.

MÜLLER J. 2006. Systematics of Baccharis (Compositae-Astereae) in Bolivia, including an overview of the genus.

Syst. Bot. Monogr. 76:1—341.

Ouv&RRA, A.S., L.P. Desle, A.A. SCHNEIDER, AND J.N.C. Mancuion. 2006. Checklist do género Baccharis L. para o Brasil

(Asteraceae-Astereae). Balduinia 9:17-27.

Raproro, A.E., W.C. Dickison, J.R. Massey, AND CR. BeLL. 1974. Vascular plant systematics. Harper & Row, New York.

146 Journal of the Botanical R h Institute of T

BOOK REVIEW

Bruce M. Pavuk. 2008 The California Deserts: An Ecological Rediscovery. (ISBN 978-0-520-25145-8,

pbk.). University of California Press, 2120 Berkeley Way, Berkeley, California 94704-1012, U.S.A.

(Orders: The University of California Press c/o California/Princeton Fulfillment Services, 1445 Lower

Ferry Road, Ewing, New Jersey 08618, U.S.A.; www.ucpress.edu, email orders@cpfsinc.com, 1-800-

777-4726, 1-800-999-1958 fax). $27.50, 365 pp, numerous figures, graphs, images, maps, tables,

references, index, and art credits, 7" x 10".

The California Deserts: An Ecological Rediscovery, is divided i i 1 ifically d ibe tl

p y gins, discovery, climate,

geology, resources, fauna 1 fl he 25 milli hat f he tł d ts of California: the S Mojave, and

Great Basin ac nd Bruce Pavlik thread ies of various early explorati hrougl he fi ion of this book, provid-

: 1 1 ST Ta fi £, TT s J 1 Lt ra D. y Ry pe | ca

ing the rea Į y

: 1 NT v 1 na 1 A ECT 1; E el 1 len f, r 1

mo day oe and how most o are ca between 1 10° and a latitude ae and south of the Go afar M. three dm

are in fact very ,andth : ge, precipitation, temperature

that are present within

Pavlik ao the “remarkable id that can js a Ww unn the California desert region. A pri ple is the

Tat ^ f. m Ll TT "m

y its abilit y to procure wat

book by listing tl 1 (e.g. fragmentation, i i ies) of thi i 1 id ideas f ing tl

DOOK DY B 5 5 ? L e F o d

deserts

12] 1 3 o, ; 1 acta] f. list, educator, I ]

Pi Pi r

da. t th J f +1 + California Ib J; th 1 1 e 1 1 1;

including A Vaot number

f pl f the flora and fauna that are found w a ee regions. Ted Tees that were bus throughout ce PUDE ld the

reader WIA a

California deserts. en McNew, MS Biol) Prog Manager, Botanical R h Institute of Texas, 500 E 4th Street, Fort Worth, TX 76102,

U.S.A

J. Bot. Res. Inst. Texas 3(1): 146. 2009

DIPLYCOSIA INDICA (ERICACEAE):

A NEW SPECIES AND A NEW GENERIC RECORD FOR INDIA

MR. Debta' and HJ. Chowdhery

Botanical Survey of India (Northern Circle)

192, Kaulagarh Road, Dehradun- 248 195, INDIA

ABSTRACT

Diplycosia Bl (Eri )—a new generi land Dipl ia indica M.R. Debta & HJ. Chowdhery—a new species from India

is described and illustrated.

Key Worbs: Diplycosia, India, new species, new generic record

RESUMEN

+ Dy] Gyi A 4 e 4 +7 E Tin? iaindica MR Debta @HJ Chowdhery—una nueva

o E

especie de la India.

INIRODUCIION

The genus Diplycosia consisting of about 99 species (Mabberley 1997), is widely distributed in the Ma-

laysian region. Sleumer (1967) revised the genus for Flora Malesiana treating 97 species. He reported that

the genus has its highest diversity in Borneo, while Mt. Kinabalu has the highest concentration of species.

Powell and Kron (2001), based on molecular studies, have suggested treating Diplycosia under Gaultheria

L. Argent 2002) —Uwhile studying Diplycosia from Borneo and peninsular Malaysia— described seven new

species, one new form, and made two new combinations. Diplycosia can be distinguished from Gaultheria

with the following key.

KEY TO THE GENERA DIPLYCOSIA AND GAULTHERIA

1. Flowers in fascicles; anther cells unawned, not aristate, with short or long apical tubules at apex; tubules

connate or free Diplycosia

1. Flowers in racemes; anther cells awned or aristate, not having apical tubules at apex Gaultheria

punc a20 Ms plant couecuing = to Singalila National Park e Bengal), in the eastern Himalayan

, some i 1 unusual plant speci ing to the family Ericaceae were

collected. "Initially ty were iden nea as species of Gaultheria, but dene. ] that some of the

specimens belonged to the closely allied genus Diplycosia, which is hitherto unknown from India. Further

critical examinations of the specimens have shown that though one specimen had some resemblance to a

Malaysian species, Diplycosia aperta JJ. Sm., but differs widely from it in habit, leaf venations, number of

flowers, bracteoles, and calyx character. The new collection is therefore described as a new species, Diplyco-

sia indica, which is also a new generic record for India. Table 1 shows the differences with its allied species

Diplycosia aperta. An illustration of the new species is provided for easy identification.

1; 1

Diplycosia indica M.R. Debta & H.J. Chowdhery, sp. nov. (Fig. 1). Treus: INDIA. West Bencar: Darjeeling district, Singalila

National Park, from Kainyakata to Kalapokhri, ca. 2950 m, 02 Jun 2006, M.R. Debta 40813 (HoLorrrus: CAL; isorvrus: BSD)

Lan nova b e eae Sm. arte affinis, sed habitu i; infl ii oe floribus multum brevioribus, albis;

: 1

[e] E

Terrestrial, prostrate shrubs, up to 1.5 m high. Stems terete, us to densely brown-hirsute or setose, reddish.

Leaves subcoriaceous, 13-36 mm long x 7-18 mm wide, elliptic to ovate-elliptic, mucronate or apiculate,

‘Corresponding author's e-mail: manas_debta@rediffmail.com

J. Bot. Res. Inst. Texas 3(1): 147 —150. 2009

148

£ sl Das

Journal of

0.5 mm

Soc

327:

W, LJ

X 2 Y

ORE

ASE

Fis. 1. Diplycosia indica. A. Habit. B. Leaf, upper surface. C. leaf, lower surface. D. Flower. E. Bract. F. Bracteole. G. Calyx, dorsal surface. H. Calyx, ventral

surface. |. Corolla split open. J. Stamen. K. Pistil (Scale bars: A = 3 cm; B-C = 1 cm; D - land K = 1 mm; J = 0.5 mm).

Taste 1. Distinguishing d f Diplycosia indi | D. aperta

Diplycosia indica Diplycosia aperta

Habit Terrestrial, prostrate shrubs Epiphytic, sometimes scandent shrubs

Leaves Margin with short cilia; lateral veins in Margin with long cilia; lateral veins in 2 pairs,

3-4 pairs, distinct beneath obscure beneat

Flowers (3-)4-5; pedicel 0.5 mm, glabrous 1 or 2, rarely 3; pedicel 0.8-1.1cm, with bristles

Bracteoles Acute to shortly acuminate, 3-3.5 mm long, Obtuse, ca. 1.5 mm long, glabrous throughout

very sparsely puberulous in the apical the dorsal surface

half along midrib on dorsal surface

Calyx Minutely puberulous on both surfaces Glabrous to some fine glandular warts on

in apical portion dorsal surface

Corolla ca. 4 mm long, white, lobes 0.75 mm 8-9 mm long, red or rose, lobes 1.5 mm

rounded to broadly cuneate at base, margin serrulate-ciliate with short cilia, cilia 0.5-1 mm long, laxly softly

pilose at the base along mid-vein, dark green above, setulose, light green beneath; veins impressed above,

distinct beneath, lateral veins in 3-4 pairs; petiole 1.5-3 mm long, setulose, red. Inflorescence a fascicle, up

to 7 mm long, (32)4—6-flowered, glabrous; bracts 1.5-2 mm long x 1.8-2.2 mm wide, basal, ovate, acute,

glabrous, margin membranous, ciliate, reddish-green; bracteoles 2, 33.5 mm long x 1.3-1.8 mm wide,

ovate, acute to shortly acuminate, opposite, concave, margin membranous, ciliate, very sparsely puberulous

in the apical half along the middle on dorsal surface. Flowers 4—5 mm long, urceolate; pedicel ca. 0.5 mm

long, inconspicuous, glabrous. Sepals 5-6, 1.5-3 mm long x 1-1.5 mm wide, ovate-triangular, acuminate,

minutely puberulous in the apical portion on both the surfaces, margin membranous, thickly puberulous,

adnate to the ovary, rosy-pink. Corolla ca. 4 mm long, urceolate, white, 6-lobed; lobes ca. 0.7 mm long,

ovate, apex obtuse to rounded, margin obscurely crenulate, glabrous. Stamens 10, ca. 1.5 mm long, loosely

epipetalous, caducous, filament ca. 0.8 mm long, oblong, dilated at middle, papillose; anther with tubules

0.6 mm long, very small, 2-lobed, oblong. Pistil 2.5-3.5 mm long; ovary superior, 1 mm long x 1.5 mm wide,

subglobose, subglabrous or scattered puberulous, light green; style ca. 2 mm long, slender, sparsely minutely

hairy in upper half, disk cupular, 8-10 lobed, pressed against the ovary, papillose. Fruit not seen.

Diplycosia indica is closely allied to Diplycosia aperta J.J.Sm. but can be distinguished from it by its ter-

restrial habit; inflorescence with 4—6, glabrous pedicelled, much smaller, white flowers; acute to acuminate,

EE pd mob and poto poc

] , in association with Viburnum erubescens,

Thamnocalamus M etc. and i at an altitude of about 3000 m.

Flowering.—June.

Distribution —India: Eastern Himalaya (West Bengal). Endemic.

Etymology.—Diplycosia indica is named after India, the country of its origin.

Conservation status.—The new species could not be traced from anywhere inside the Singalila Na-

tional Park except for a small population. Since Singalila National Park—especially from Manebhanjeng to

Sandakphu—is a famous trekking route, it attracts a large number of tourists and trekkers throughout the

year. The increasing pressure from the tourism industry and large scale cattle grazing in and around park

areas, are posing severe threats to the natural habitat of numerous species aaa paa indica.

Additional specimen examined: INDIA. West Bengal: Darjeeling district, 32 km N of Manebhanjeng, from K Kalapokhri

ca. 2950 m, 02 Jun 2006, M.R. Debta 40813 (CAL, BSD).

ACKNOWLEDGMENTS

The authors are indebted to the director, Botanical Survey of India for providing facilities. Grateful thanks

are due to G.C.G. Argent and DJ. Middleton, Royal Botanic Garden, Edinburgh for their expert suggestions

3 af ob Dat H ID hi bitit. £f T,

150

and helps, to VJ. Nair, ex-joint director, Botanical Survey of India for providing Latin diagnosis of the new

species. Thanks are also acknowledged to the officials of the Forest Department, Government of West Bengal

for permission to carry out research work inside the Singalila National Park and to Brijesh Kumar, Botanical

survey of India, Dehradun for illustrating the new species. We greatly appreciate the helpful review of one

anonymous reviewer.

REFERENCES

Arcent, G.C.G. 2002. New taxa and new combinations in the genus Diplycosia (Ericaceae) of Borneo and Peninsular

Malaysia. Gard. Bull. Singapore 54:217-238.

Mapsertey, DJ. 1997. The plant book: a portable dictionary of the vascular plants. Cambridge University Press.

Pp. 234

PowtLL, E.A. and K.A. Kron. 2001. An analysis of the phy ic relationships in the winterg group (Diplycosia,

Gaultheria, Pernettya, Tepuia; Ericaceae). Syst. Bot. 26: :808- 817.

SiEuMER H. 1967. Diplycosia Blume. In: van Steenis, C.G.G.S, ed. Fl. Males. 6, ser. 1:696—740.

AMORPHA CONFUSA, A NEW NAME FOR

AN OLD AMORPHA (FABACEAE: AMORPHEAE)

Shannon C.K. Straub Bruce A. Sorrie

Cornell University North Carolina ped Heritage Program

LH. Bailey Hortorium Office of Natural Resources and Conservation Planning

Department of Plant Biology 3076 o Pos thage Roa

412 Mann Library Whispering Pines, North Carolina 28327, U.S.A.

Ithaca, New York 14853, U.S.A.

ss463@cornell.edu

Alan S. Weakley

University of North Carolina at Chapel Hill

ios of North Carolina Herbarium i U)

Nor rolina Botanica den

rudi m North Carolina 275 s Ud

ABSTRACT

bination, Amorph fusa (Fab : Amorpheae), based on A. georgiana var. confusa, is established. The variety is raised

in sank due to differences in morphology, distribution, habitat preference, phenology, ploidy, and genetic variation when compared to

A. georgiana.

RESUMEN

Se ao una A nueva, Amorpha cantas a Amorpheae), parana en A. PCM var. confusa. La variedad se

, distribución f t, fenología, ploidía, y variación

genética en toiipaba ción. con Á. georgiana.

For many years, plants currently known as Amorpha d aid UT var. confusa Wilbur (Fabaceae Juss.:

Amorpheae Boriss.) have been a source of tural confusion. Wilbur (1964) elegantly

explored the intricacies of the situation in his revision of the dwarf species of Amorpha L. Various names often

had been applied incorrectly to this species (e.g., A. caroliniana H. B. Croom, A. cyanostachya M.A. Curtis)

or were unavailable (e.g., A. glabra Desf. ex Beadle, nom. illeg.). This left Wilbur no option but recognize it

with a new name, which he did at the rank of variety due to its similarity to A. georgiana. At the time, this

was the most conservative course of action due to the paucity of collections of both var. georgiana and var.

confusa, even though Wilbur recognized that the two differed morphologically. Later, Wilbur (1975 p. 367)

commented in his monograph of the genus that he remained "skeptical" that his treatment would prove

"satisfactory when more is learned about them" and acknowledged that "future investigation may well dem-

onstrate that the two taxa are specifically distinct." Most recent floristic treatments and species checklists

(e.g., Isely 1990, 1998; Kartesz 1999) have followed Wilbur's treatments in recognizing two varieties. These

varieties differ morphologically in many characters with var. confusa having larger leaflets [((10-)15-25(35)

mm long and (7-)9-15(-18) mm wide versus (36-10(-15) mm long and (2—)3-5(-8) mm wide], longer

petioles [(6-)8-15(-20) mm versus 1-3(-5) mm] and racemes [10-20(30) cm versus (2-)3-5(6) cm], less

numerous leaflets, clustered, panicle-like inflorescences rather than generally solitary racemes, and bright

blue rather than reddish-violet vexilla (Fig. 1; Wilbur 1964; Sorrie 1995; Weakley 1995).

Recent investigations into the current distribution, conservation status, and genetics of the two varieties

have revealed additional differences and supported Wilbur's (1975) notion that they should each be recog-

nized at the specific level. The distributions of the varieties do not overlap currently, nor did they histori-

cally based on herbarium records (Fig. 2). Amorpha georgiana var. confusa is an endemic of a few counties in

J. Bot. Res. Inst. Texas 3(1): 151 —155. 2009

Journal of the Botanical Research Institute of Texas 3(1)

Fis. 1. a. Amorpha confusa at Green Swamp Preserve, North Carolina. (Photograph by Andrew Walker). b. Amorpha georgiana at Fort Bragg, North

Carolina (Photograph by Bruce Sorrie).

Straub et al., A new name for an old Amorpha 153

Amorpha confusa

BÉ Current Distribution

[2] Historical Distribution

Reported Distribution

a Amorpha georgiana

eS lI Current Distribution

[5] Historical Distribution

[s] Reported Distribution

Atlantic

Ocean

Fic. 2. Current, historical,

extreme southeastern North Carolina and immediately adjacent South Carolina, though it is now believed

to be extant in only Brunswick and Columbus counties of North Carolina (Weakley 1995). Amorpha geor-

giana var. georgiana is found in the middle and inner Coastal Plain of North Carolina, South Carolina and

Georgia (Sorrie 1995). Both varieties are associated with the longleaf pine savanna ecosystem but differ in

the details of their habitat preferences with var. confusa occurring in flat, moist to rather dry outer Coastal

Plain savannas with loamy soils, especially of the Foreston series (Weakley 1995); whereas var. georgiana

occupies more moist to occasionally inundated areas, chiefly sandy river terraces and river banks above

blackwater rivers traversing the sandhills of the middle and inner Coastal Plain, and more infrequently the

edges of swampy floodplains (Sorrie 1995). Both varieties have suffered habitat loss, fragmentation, and

degradation in recent years due to human activities, including fire suppression, agriculture, and land devel-

opment, causing them to be of conservation concern (Sorrie 1995; Weakley 1995). In North Carolina, var.

confusa is currently considered to be threatened (North Carolina Department of Agriculture & Consumer

Services 2008) because its population numbers are estimated to be less than 14,000 individuals, and only

those populations occurring in The Nature Conservancy's Green Swamp Preserve are likely to receive long-

term conservation-oriented management (Weakley 1995).

The two varieties also differ in phenology, with Amorpha georgiana var. confusa flowering from late

May to mid-July and var. georgiana flowering from late April to late May (Sorrie 1995). Additionally, recent

genetic work has indicated that the genome of var. confusa is likely tetraploid, while that of var. georgiana

is diploid (Straub et al. 2009). A comparison of the microsatellite variation observed for the varieties at the

population level indicates that they are quite well differentiated genetically and preliminary phylogenetic

analyses of chloroplast spacer region and low-copy nuclear gene DNA sequence data indicate that they are

likely not each other's closest relative among Amorpha species (S. Straub & J. Doyle, unpublished data).

Since the time of Wilbur's (1964) original publications, additional studies of the morphology, distribu-

tions, habitat preferences, phenology, and genetics of the two varieties have shown the extent to which they

are distinct from one another. These differences warrant the neon an var. confusa at the rank of species

(Sorrie 1995; Weakley 1995). Recognition at this rank further empl tion importance and

need of both of these imperiled species.

154 t tani i Texas 3(

Amorpha confusa (Wilbur) S.C.K. Straub, Sorrie & Weakley, comb. et stat. nov. Amorpha georgiana var. confusa

Wilbur, J. Elisha Mitchell Sci. Soc. 80: 58. 1964. Tyre: UNITED STATES. NORTH CAROLINA. Brunswick Co.: Savannah, 7 mi SW of

Wilmington, 31 May 1938, R.K. Godfrey & IV. Shunk 4122 (Lectory, designated by Wilbur 1975: GH, digital image!; DUPLICATE OF

THE LECTOTYPE: US, digital image!).

Amorpha glabra Beadle, Bot. Gaz. 25:279. 1898; EE. Boynton in Smalls Fl. S.E. U.S. 626. 1903, non Poir., Encycl. (Lamarck)

Suppl. 1:330. 1810

Amorpha caroliniana sensu Torr. & A. Gray, Fl. N. ea 1:305. 1838 in part; C.K. Schneider, Ill. Handb. Laubholzk. 2:74. Mar 1907 &

Bot. Gaz. 43:302. Jun 1907; Rydberg, Fl. N. Amer. 24:29. 1919, non H.B. Croom, Amer. J. Sci. Arts 25:74. 1834

Amorpha cyanostachya sensu E.J. Palmer, J. Arnold pum 12:169. 1931; Small, Man. S.E. Fl. 639. 1933, non M.A. Curtis, Boston J. Nat.

Hist. 1:140. 1835

Wilbur (1964) did not choose E Pen two acl listed in the protologue as the holotype, although he

later indicated that he considered P GH to be the holotype (Wilbur 1975: 366), here corrected

to lectotype pursuant to Article 9.8. of the ICBN ene et al. 2006).

Additional collections. NORTH CAROLINA. Bladen C Bil Herbarium 5767B (NY). B k Co.: pineland near

Wilmington, 28 Aug 1938, R.K. Godfrey 6233 (GH, US, NCSC- 4939); gs woods along route 17 about 3.5 mi NE of Bolivia, 22 Jun 1947,

C.E. Wood, Jr. an D. elle ih (GH, NCU-175175); dry sandy pineland, 20 mi S of Wilmington on route 17, 27 Jun 1950, F.H. Sargent

10008 (GH); 1 Shall d Ash, 6 Jul 1951, R.K. Godfrey & H.L. Blomquist 51177 (FLAS, NCSC); dry, sandy

ie: savanna, 2.4 mi W of Shallotte, Highway 130, 10 Jul 1951, H.L. P ds R.K. a ds il R. E Wilbur15225 (PURE, ee in

h about 1 mi W of Shallotte on NC 130; 6 Jun 1957, R.L. Wilbur 5991 (

M : f] 4 1 4 ad e

bod 6 mi NW of Southport on Route 87, 12 Jun 1957; R.L. Wilbur 6169 (DUKE-14 41699) p

route 87-133, 10 Jul 1963, R.L. mee 6954 (DUKE, FSU); open piney woods about 11 mi NW of Supply on Route 211, 18 Aug 1967, R.L.

Wilbur 9466 (DUKE-186242); powerline right-of-way, s E state roads 1518 and 1521, N of Funston,14 Jun 1979, DJ. Sieren 1835

i of borrow pit on east side of NC 211, 9.4 km N of US 17, 18 Jul 1986,

J.B. Taggart 224 (NCU-557630); Green Swart Preserve, 5.5 mi N of Supply, 22 May 2003, A.S. Weakley & G.T. Chandler 7242 (NCU-

569103, NCU-569104, NCU-569106); Green pie Preserve, W of Big Island Savanna, 22 in 2003, A.S. Weakley & G.T. ene 7244

6-11

Lad

bord

(NCU-569107, NCU-569109, NCU-569111) g Lake Preserve, C d, 11 Aug 2006, J.C. Morris 0

(NCU-584945, WILM). Columbus Co.: ing DE aki 25 TAE dd PO. “Shallert s.n. aye ileal dan near Route 76, 1

mi SE of Delco, 31 Jul 1949, W.B. Fox & R.K. pa o (NCSC-3 : 3.7 mi NW of Old Dock on NC 130, 2 Jul 1968, S.W.

Leonard & K. Moore 1720 (FLAS, FSU, GH, NCU sand il along CR 1928 to Shulkins, 7 Jul 1989, J.A. Churchill 89-685

(VDB). New Hanover Co.: Wilmington, 1 Jul 1904, bim DRM 1391-L (NCU): Wilmington, 6 Oct 1908, E.A. Bartram s.n. (PH);

Wilmington, 11 Jun 1917, T.G. Harbison 3415 (NCU-3469); low swampy ground, Wilmington, 11 Jun 1917, T.G. Harbison 16 (A). SOUTH

CAROLINA. Horry Co Sea, 16 Jun 1941, H.R. Totten s.n. (NCU-22550, NCU-577773).

ACKNOWLEDGMENTS

The authors thank the curators of A, DUKE, FLAS, FSU, GH, HUH, NCSC, NCU, US, VDB and WILM for

access to specimens and digital images, the North Carolina Natural Heritage Program for access to locality

data, James L. Reveal (BH) for helpful discussions related to nomenclature issues, oe Tu = Reveal and

Guy Nesom for valuable comments which improved the manuscript. Th li

the the U.S. Fish and Wildlife Service for status surveys, a Garden Club of America ein H. Beattie

Fellowship to S.C.K.S., and National Science Foundation grant DEB-0709960.

g from

REFERENCES

IseLy, D. 1990. Amorpha. In: Vascular flora of the southeastern United States, vol. 3, part 2: Legumi (Fabaceae).

University of North Carolina Press, Chapel Hill. Pp. 71-76.

IseLy, D. 1998. Amorpha. In: Native and naturalized Leguminosae (Fabaceae) of the United States (exclusive of

Alaska and Hawaii). Brigham Young University, Provo, UT. Pp. 132-144.

KARTESZ, J.T. 1999. A synonymized checklist and atlas with biological attributes for the vascular flora of the United

States, Canada, and Greenland. First Edition. In: Kartesz, J.T. & C.A. Meacham. Synthesis of the North American

Flora, Version 1.0. North Carolina Botanical Garden, Chapel Hill, NC.

McNELL, J., ER. Barrie, H.M. Bunper, V. Demouun, D.L. HAwKsworTH, K. MARHOLD, D.H. NicoLson, J. PRADO, PC. Sit va, 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.

Straub et al., A new name for an old Amorpha 155

NORTH CAROLINA DEPARTMENT OF AGRICULTURE & CONSUMER Services. 2008. Plant Industry Division — Plant Protection Section

- Plant Conservation Program: Protected plant list [online data]. NCDA&CS, Raleigh, NC. Available at http://

www.ncagr.com/plantindustry/ plant/plantconserve/plist.htm, accessed 3 January, 2008.

SorRIE, B.A. 1995. Status survey of Amorpha georgiana var. georgiana. Submitted to the US Fish and Wildlife Service,

Office of Endangered Species, Asheville, NC, and to the North Carolina Natural Heritage Program, Division of

Parks and Recreation, Raleigh, NC.

STRAUB, S.C. K,, S.M. BOGDANOWICZ, AND J.J. Dove. 2009. Characterization of twel | hic mic lli |

for Georgia false indigo (Amorpha georgiana Wilbur var. georgiana), an endangered species, and their utility

in other dwarf Amorpha L. species. Molec. Ecol. Resour. 9:225-228.

Weak.ey, A. 1995. Status survey for the savanna Indigo-bush, Amorpha georgiana Wilbur var. confusa Wilbur.

Submitted to the US Fish and Wildlife Service, Office of Endangered Species, Asheville, NC, and to the North

Carolina Natural Heritage Program, Division of Parks and Recreation, Raleigh, NC.

WiLgur, R.L. 1964. A revision of the dwarf species of Amorpha (Leguminosae). J. Elisha Mitchell Sci. Soc. 80:

51-65.

WiLBUR, R.L. 1975. A revision of the North American genus Amorpha (Leguminosae-Psoraleae). Rhodora

77:337-409,

156 [] Ingel D saal D L

BOOK REVIEW

MICHAEL WINK AND BEN-Enik VAN Wyk. 2008. Mind-Altering and Poisonous Plants of the World. (ISBN-13

978-0-88192-952-2, hbk.). Timber m Inc., 133 S. bid eae Avenue, Suite 450, Portland, Oregon

97204-3527, U.S.A. (Orders: j com, com, 800-327-5680, 503-227-

3070 fax). $49.95, 464 pp., ose throughout black/white a ans 6 5/8" x 9 1/2".

Mind-Altering and P Pl of the World i k. Its subtitle—A scientifically te guide to 1200 Loss cinia

ing plants i fect d inti frbied lightf ] A ful book. I ; 2; NE id

this book to be. a valuable aud liabl finf ti

on poisonous Slate Not only is this Book for the botanist, but a variety

ranchers, teachers, students, homeowners, gardeners, horticulturists, florists,

botanists, chemists, medical and ing enforcement personnel, health care professionals, parents, and anyone with a curiosity in the

power of green plants

of audiences would find ful E

It has been said, “Poisons tl he] J 251 I p 1] FA y : g lif y ly ly A <4] iA

have ever had a peculiar fascination for mankind." Poi 1 poi pl inly fasci 1 tl thors, Wink and van Wyk;

they have done a spectacular job in assimilating a vast t of inf tion int tical and user friendly format. Beautiful but

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iuis ] ins derived f I Mind-Alteri 1 Poi Plants of the World tal P

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In the authors’ words, “About 200

d by concise monographs.

Essential facts about another 1000 e are provided in the ‘Quick guide to poisonous and mind-altering plants’ at the end of the

book.” Eacl l |

12 sections: 1) Name, 2) plants with similar properties, 3) plant characters, 4)

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and 12) systematics. Beautiful color photograpl ] gray

ake note, accidental eal happen and therefore I was O to see a list of suggesti f d to avoid pete. or

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8 rZ. : Dar Hey Lip

Botanical Research Institute of Texas, Fort Worth, Texas 76102-4025, U.S.A.

J. Bot. Res. Inst. Texas 3(1): 156. 2009

A NEW COMBINATION IN NORTH AMERICAN TEPHROSIA (FABACEAE)

Guy L. Nesom James L. Zarucchi

2925 Hartwood Drive Missouri Botanica! Garden

Fort Worth, Texas 76109, U.S.A

guynesom@sbcglobal.net

O. Box 299

St. Louis, Missouri 63166-0299, U.S.A.

james.zarucchigmobot.org

ABSTRACT

Tephrosia xintermedia (Small) G.L. Nesom & Zarucchi, comb. n

is the correct name for plants regarded as hybrids between

Tephrosia espia and T. florida, replacing T. xfloridana (Vail) s sb of typification are given for Cracca intermedia Small and

Cracca floridana Vai

RESUMEN

t iderad mo hí-

Tephrosia xintermedia (Small) G.L. Nesom & Zarucchi, comb. nov., es el nombre correcto a a

bridos entre Tephrosia chrysophylla y 1. florida, que reemplaza a T. pcne (Vail) Isely. Se aportan det tir

intermedia Small y Cracca floridana Vail.

Cracca intermedia Small was described (Small 1894) as a new species intermediate between C. chrysophylla

(Pursh) Kuntze (= Tephrosia chrysophylla Pursh) and C. ambigua (M.A. Curtis) Kuntze (= Tephrosia florida

(F. Dietr.) C.E. Wood). Vail (1895) curiously provided C. intermedia with a new name, Cracca smallii Vail,

because she apparently regarded C. intermedia as invalidated by the earlier Tephrosia intermedia Graham in

Wall. In the same publication, Vail described C. floridana Vail, noting that it was *very close" to C. smallii.

Isely (1982, 1990, 1998) provided the combination Tephrosia xfloridana (Vail) Isely and used it in reference

to the putative T. chrysophylla—T. florida hybrids, even though he correctly recognized Graham’s T. intermedia

to be a nomen nudum.

Tephrosia intermedia Graham was indeed published without a description and is

nudum, but it does not block the transfer of Cracca intermedia Small to Tephrosia. Many of the nomina nuda

in Wallich's list were validated by description in Don (1831-1834), but Tephrosia intermedia is not among

them. *T. intermedia Grah. in Wall. Cat. 5632" was listed at the end of the paragraph for Tephrosia tinctoria

Pers. var. intermedia [W & A.] in J.D. Hooker, Fl. Brit. India 2:112. 1879, so the epithet "intermedia" at the

rank of species under “Tephrosia” was not there validated.

illegiti nomen

Tephrosia xintermedia (Smail) al L. Nesom & o comb. nov. Cracca intermedia Small, a d Bot. Club

(non Tepl ud.], Numer. List [Wallich] n 2. Between 1831 and 1832). Cracca smallii

Vail, Bull. Torrey Bot. Club 22:33. 1895 [nom. nov. e C. intermedia Small]. phim smallii ci. B.L. Rob., Bot. Gaz. 28:198. 1899.

Tyre: U.S.A. FLORIDA. [Duval Co.]: near Jacksonville, Florida, 31 May, Fr. 11 Jul 1893, A.H. ahs 4231 M Rn NY AA

digital image!; isorEcrorvees: NY 00006578, xd image!; US, US-digital image!) I

de "by Mr. A.H. Curtiss in 1893, flowering on do 3lst ne in mature Pie on July 1 Ith” and that it was lee a Sd Y

» Alc

poor ‘Blackjack thickets’ about Jacksonville,

Isely (1982) indicated “Curtiss 4231” at NY to be ihe “holotype” but he did not ere which of o Ao was e Due. Sheet

00006579 has a cie pun “type” bn 00006578 has a printed label “isotype,”

bel of 00006578 has “C. smallii A.M. Vail” handwritten below the typed “Cracca intermedia Small. sp. nov.’

designations. T

Cracca dicun Vail, Bull. Torrey Bot. Club 22:35. 1895. Tephrosia xfloridana (Vail) Isely, Brittonia 34:340. 1982. Type: U.S.A. FLORIDA.

o.: high pineland, vicinity of Eustis, Jul 1894, G.V Nash 1198 (1ecrorvrE, designated by Isely 1982: NY, digital ni Vail

me ke Florida, G.V. Nash Nos. 49415, 1198, 1263, 1334, 1552, 1615. Louisiana, New Orleans Ingalls. Tj rb

Columbia College.” Photos of all syntypes except Nash 1334 are displayed on the NY web site. As oda " = (1982), e xe

collection does not appear to represent a different taxon.

The original hypothesis of hybrid origin of Tephrosia intermedia has been accepted by those who have further

studied the genus (Vail 1895; Robinson 1899; Wood 1949). Isely (1998, p. 856—857) noted that “As alleged

J. Bot. Res. Inst. Texas 3(1); 157 —158. 2009

El D H ID LI Ht f T,

158 Journal of t Texas 3(

by Wood (1949), Tephrosia xfloridana probably is a derivative of hybridization between T. chrysophylla and

T. florida. Whatever its origin, it is sufficiently widely distributed and consistent in characters to deserve

binomial listing.” It has been collected from Alabama, Florida, Georgia, and Mississippi, but the distribution

is sporadic and its origin as a recurrent hybrid seems a reasonable hypothesis. It occurs in association with

the two putative parents and is intermediate in morphology.

ACKNOWLEDGMENTS

We are grateful to Kanchi Gandhi (GH) and Richard P. Wunderlin (USF) for comments and analysis.

REFERENCES

Don, G. 1831-1834. A general history of the dichlamydeous plants, vols. 1—4.

sev, D. 1982. New combinations and one new variety among the genera Indigofera, Robinia, and Tephrosia

(Leguminosae). Brittonia 34:339-341.

Isety, D. 1990. Vascular flora of the southeastern United States, Vol. 3, Part 2: Leguminosae (Fabaceae). Univ. of

North Carolina Press, Chapel Hill.

sey D. 1998. Native and naturalized Leguminosae (Fabaceae) of the United States (exclusive of Alaska and

Hawaii). M.L. Bean Life Science Museum. Univ. of Utah, Provo.

Rosinson, B.L. 1899. Revision of the North American species of Tephrosia. Bot. Gaz. 28:193-202.

Smal, J.K. 1894. Studies in the botany of the southeastern United States.—ll. Bull. Torrey Bot. Club 21:300-307.

Van, A.M. 1895. A revision of the North American species of the genus Cracca. Bull. Torrey Bot. Club 22:25-36.

Woon, C.E. 1949. The American barbistyled species of Tephrosia (Leguminosae). Rhodora 51:193-231, 233-302,

305-364, 369—384.

A RECOMBINATION FOR VARIETIES OF ANTICLEA ELEGANS (MELANTHIACEAE)

Wendy B. Zomlefer Walter S. Judd

Department of Plant Biology Department of Biology

University of Georgia University of Florida

2502 Plant Sciences PO. Box 118526

Athens, Georgia 30602-7271, U.S.A. Gainesville Florida 32611-8526, U.S.A.

ABSTRACT

The new bination is 1 le for Anticlea eleg . glaucus, formerly within the polyphyletic Zigad pl Zigad

elegans var. glaucus.

RESUMEN

Se | binació Anticlea elegans var. glaucus, previamente incluida en el complejo polifilético Zigadem

Zigadenus elegans var. glaucus.

Based on molecular and morphological data (Zomlefer et al. 2001), the authors made recombinations at

the generic and species-level for taxa of the former Zigadenus complex (Zomlefer & Judd 2002). While as-

sisting in the update of these taxa for the Plants Database (USDA-NRCS 2009) for the National Resources

Conservation Service (Cooperative Agreement 68-3H75-3-122 Mod 14; PI Craig C. Freeman, KANU), the

first author realized the necessity of making the following infraspecific transfer to Anticlea elegans (formerly

Zigadenus elegans), presented below.

Anticlea elegans (Pursh) Rydb. var. glaucus (Nutt.) Zomlefer & Judd, comb. nov. Basionvm: Melanthium glaucum

Nutt., Gen. 1:232. 1818

Zigadenus elegans Pursh var. glaucus (Nutt.) P Gl & Cronq. Man. Vasc. Pl. NorthE. U.S. Canad., ed. 2, 864. 1991.

The wide-ranging Anticlea elegans (Alaska- Canada south to northern Mexico), occurs in various habitats:

generally bogs, beaches, and calcareous wetlands in eastern North America, and prairies, coniferous forests,

and alpine meadows in the west (Zomlefer 1997). The differences between the eastern element, A. elegans

var. glaucus (plants glaucous; leaves blunt or subacute, coriaceous; inflorescence usually paniculate; bracts

herbaceous, subulate; tepals intensely colored; capsule ovoid-conic), and the western A. elegans var. elegans

(plants green; leaves pointed, more herbaceous; inflorescence usually racemose; bracts scarious margined;

tepals pale; capsule lance-conic), as outlined by Fernald (1935), are most evident in the geographical extremes

(Preece 1956), and these two taxa require further study in their area of morphological integradation in the

eastern Dakotas—western Minnesota region (Gleason & Cronquist 1991; Schwartz 2002). Varietal recogni-

tion is appropriate given our current understanding of the pattern of variation.

ACKNOWLEDGMENTS

We thank Fred Barrie and Guy Nesom for their careful check of this nomenclatural note.

REFERENCES

FERNALD, M.L. 1935. Critical plants of the upper Great Lakes region of Ontario and Michigan. Rhodora 37:238-262.

GLEASON, A. AND A. CRONQUIST. 1991. Manual of vascular plants of northeastern United States and adjacent Canada.

The New York Botanical Garden, Bronx.

Preece, S.J. 1956. A cytotaxonomic study of the genus Zigadenus (Liliaceae). Ph.D. dissertation. State College of

Washington, Pullman.

J. Bot. Res. Inst. Texas 3(1): 159 —160. 2009

160

Schwartz, F.C. 2002. 13. Zigadenus Michaux. In: Flora of North America Editorial Committee, eds. Flora of North

America north of Mexico, Vol. 26, Magnoliophyta: Liliidae: Liliales and Orchidales. Oxford University Press,

New York and Oxford. Pp. 81-8

USDA-NRCS. 2009. The PLANTS Database, National Plant Data Center, Baton Rouge, Louisiana. http://plants.

usda.gov. Accessed 23 April 2009,

ZomLerer, W.B. 1997. The genera of Melanthiaceae in the southeastern United States. Harvard Pap. Bot. 2:133-177.

Zomuerer, W.B. AND W.S. Jupp. 2002. Resurrection of segregates of the polyphyletic genus Zigadenus s.l. (Liliales:

Melanthiaceae) and resulting new combinations. Novon 12:299-308.

ZomLerer, W.B., N.H. WiLuAMs, W.M. WHITTEN, AND W.S. Jupp. 2001. Generic circumscription and

Melanthieae (Liliales, Melanthiaceae), with emphasis on Zigadenus: evidence from ITS and trnl- -F sequence

data. Amer. J. Bot. 88:1657-1669.

in the tribe

TAXONOMIC OVERVIEW OF EURYBIA SECT. HERRICKIA

(ASTERACEAE: ASTEREAE)

Guy L. Nesom

2925 Hartwood Drive

Fort Worth, Texas 76109, U.S.A.

www.guynesom.com

ABSTRACT

hia horrid five closely related j included I ithin Eurybi Horrida. These species are E. glauca, E. horrida, E.

pulchra, and E. wasatchensis, as a ipao treated by DE Vs two e cede m nn T o oe as close

relatives: a kingii din G.L. Nesom, comb. n g y g y Welsl L. Nesom,

b (A. Nels.) G.L. Nesom, comb. nov. The first fi fu peci 1 by Brouill 1 1 1

IT DEN dE 1 1 Tel lin D +11 eee Cu De lan NAAA 1 1 EE 1

isle Herrickia and Trini } 3 g he} frhe Marl 1 lado? hological d

of Herrickia and Triniteurybia f Eurybi ivial or non itl p f Eurybia at g AS ,

1 N 1 1 : ] hi distribu 1 for the si i f Eurybi Herrickia

RESUMEN

Herrickia horrida y cinco especies muy relacionadas con ella se incluyen aqui en Eurybia sect. Horrida. Estas especies son E. glauca, E.

horrida, E. pulchra, y E. wasatchensis, tal como habían sido tratadas previamente por Nesom, y otras dos especies reconocidas reciente-

nte mediante pue E adu como LOMA próximos: Perla King (D.C. Eat.) Nesom, comb. nov. (que inclu e Furybla

kingii var. ] N comb. nov.) y E (A. Nels.) Nesom, comb. nov. I

especies fueron TN por Brouillet l gé A pli Heridas la última fue bn como hig o tinotibico Trini-

teurybia Brouillet, Urbatsch & Roberts. Los d lecul indican que Eurybia, Herrichia, y Tri un grado de

tres taxa en la base del clado i las dif i g Herrickia y able in di con n Eurybia son triviales

1 Syi tak d

MV LIMLAlSLELLILES y F o o i

ibuci Afi ] i ies de Eurybia sect. Herrickia

J 5-5 E E

Prior to 1994, Herrickia horrida Woot. & Standl. either had been maintained as a mo gens or treated

as Aster horridus (Woot. & Standl.) Blake. Nesom (1994) united it with tl lly similar species

in the western U.S.A. and treated the group as sect. Herrickia (Woot. & Standl.) son of the genus Eurybia

(Cassini) S.F. Gray: Eurybia glauca, E EM E. pulchra, and E. wasatchensis. Eurybia in the sense of the

present overview includes a total of 28 species distributed over eastern and western North America.

Based on molecular study, Brouillet et al. (2004) reinstated Herrickia Woot. & Standl. at generic rank,

including the species noted above but reducing Eurybia pulchra to varietal rank within Herrickia glauca and

expanding the group to include Aster/Tonestus hingii. They also discovered Haplopappus/Tonestus aberrans to

be a close relative of Herrickia and Eurybia and segregated it as the ions Bi genus Triniteurybia Brouillet,

Urbatsch & Roberts (Brouillet et al. 2004). Molecular evidence indicates that this group of “herrickioid”

taxa forms a grade at the base of subtribe Machaerantherinae: Oreostemma (Herrickia (Eurybia (Triniteurybia

(Machaerantherinae)))) (Brouillet et al. 2004, from ITS and 3'ETS nrDNA; Selliah and Brouillet 2007, from

the nuclear CNGC4 gene). The boundaries of Eurybia were further adjusted with the transfer of Eurybia

(Aster) pygmaea (Lindl.) Nesom and Eurybia (Aster) chapmanii (Torrey & A. Gray) Nesom to Symphyotrichum

(Brouillet & Selliah 2005; Brouillet et al. 2006).

The taxonomic repositioning of Symphyotrichum chapmanii and S. pygmaea is justified, based on mo-

lecular as well as morphological criteria, and those two species are outside of the immediate relationship

of the species of Herrickia/Eurybia/Triniteurybia. The inclusion of Aster kingii and Haplopappus aberrans in

this group of species also is a valuable and morphologically justified insight. The implicit rationale of strict

monophyly, however, for recognizing three separate genera among these few species of the eurybioid grade

J. Bot. Res. Inst. Texas 3(1): 161 —167. 2009

W200 oumalof t tani i Texas 3(

is based on a philosophical position not universally shared by systematists (e.g., Nordal & Stedje 2005),

whether or not further molecular data may confirm the pattern of cladistic relationship among them. There

is no consistent, diagnostic morphological difference to separate Herrickia (sensu Brouillet) from Eurybia,

and Triniteurybia is distinguished from Herrickia and Eurybia by a single feature (lack of ray florets), which

is sometimes variable within species and at most no greater than sometimes differentiating species among

genera of Machaerantherinae sensu stricto. Of course, as among species, there are no consistent or objective

criteria specifying the number or kinds of differences that justify the recognition of closely related genera

(or see McVaugh's set of recommended criteria [1945]). Similarly, however, there is no constraint that the

pattern of descent be mirrored in lat that does not account for phenotypic modification (or lack of

it). Were Herrickia, Eurybia, and Triniteurybia distinguished by significant morphological features, combined

with a preliminary cladistic hypothesis, their treatment as separate genera would certainly be justified.

The present overview examines the generic concepts of Herrichia, Eurybia, and Triniteurybia and shows

geographical distributions of species included here within Eurybia sect. Horrida. Map data are taken from

specimens at ASC, BRIT-SMU, MO, SJC, and TEX-LL.

Morphological distinction of Herrickia Hom abad

Eurybia (sensu Nesom 1994, 2000) is di

rounded phyllaries with a green, often basally truncate apical patch, leararen aie disc style appendages,

cylindric and multinerved cypselae, 2-seriate pappus of equal-length, apically thickened bristles, and base

chromosome number of x = 9. Brouillet et al. (2004, 2006) did not provide morphological criteria for the

distinction of Herrickia from Eurybia, but for the FNANM treatment, the key to genera (FNANM Editorial

Committee 2006, p. 19) used the following contrast.

1 Coe A a |

, ciliate-fringed,

1, Stems and leaves usually stipitate-glandular, sometimes eglandular and glaucous; leaves mostly cauline,

entire or spinulose-serrate, glabrous or scabrellous; phyllaries sometimes rounded, usually keeled, apices

acute to long-acuminate; rays 8-27; disc corolla tubes shorter than limbs (w Cordilleras) Herrickia

Stems and leaves usually eglandular, sometimes stipitate-glandular (e North Arnerica only), not glaucous;

leaves basal and/or cauline, serrate (teeth sometimes spinulose or spinose, blades then linear, grass-

like, se North America) or entire, hairy or glabrous; phyllaries usually rounded, sometimes keeled, apices

obtuse to acute; rays 5-60; disc corolla tubes shorter or longer than limbs Eurybia

—

Each of the comparisons in the key shows broadly overlapping characters and there is no consistent, diag-

nostic morphological difference to separate Herrickia (sensu Brouillet) from Eurybia.

My rationale for the recognition of sect. Herrickia (Nesom 1994) was similarly lacking in diagnostic

cohesiveness, reflecting more a perception of the geographical coherence of the species. Eurybia horrida and

E. wasatchensis are similar in their subshrubby habit, sessile and subclasping leaves, subequal phyllaries, and

tendency to produce foliaceous bracts immediately subtending the involucre. Eurybia glauca and E. pulchra

are similar to the former two in habit and subclasping leaves but differ in graduate phyllaries and lack of

foliaceous bracts. Only the subshrubby habit is distinct among other species of the genus and E. glauca and

E. pulchra sometimes are more herbaceous than subshrubby. Tonestus aberrans and Aster kingii are similar

between themselves (see comments below) and both show features of Eurybia, but they are distinct from

other “herrickioids” in their herbaceous habit and reduced infl ; both produce non-clasping leaves

and d o pid and od lack foliaceous bracts. Thus, there apparently is no ace morphological

1 group, but tl j glyi interrelated species lust ] their geographic

coherence (Figs. 1 and 2) suggest that they are closely related within the larger Eurybia, and molecular data

tentatively confirm this.

Morphological distinction of Eurybia and Triniteurybia

In the original description of Triniteurybia as a new genus (Brouillet et al. 2004), its diagnostic features were

noted (in the Latin diagnosis only, as differences from Eurybia) as eradiate heads and stipitate-glandular

leaves. The western U.S.A. species Eurybia integrifolia and E. conspicua, however, have stipitate-glandular

foliage, as does Herrickia (Eurybia) horrida. Herrickia glauca var. pulchra (= Eurybia pulchra), Herrickia (Eu-

164

sita af T, a7)

T€AdS 311)

—

O Eurybia glauca

A Eurybia pulchra `

O Eurybia horrida

Fic. 2. Distributi

f Eurybia gl. | E. pulchra, and E. horrida. The record for E, pulchra in Iron Co., Utah, is based on the citation by Welsh (2003).

T i i f Eurvbi t. Herrickia 165

Nesom,

rybia) wasatchensis, and the recently joined Herrickia kingii have stipitate-glandular stems, peduncles, and

phyllaries. Distal leaves of H. kingii sometimes are minutely stipitate-glandular (e.g., Smith 3508, TEX,

from Box Elder Co., Utah). Some species of Eurybia in the eastern U.S.A. also produce stipitate-glandular

vestiture: e.g., E. macrophylla, E. schreberi, E. spectabilis. In summary, the vestiture of Triniteurybia aberrans

is not distinct among its close relatives. Brouillet later observed (2006, p. 364) that stipitate-glandularity

“is most likely a shared primitive feature within [Herrickial and is therefore not indicative of a particular

relationship within the group."

In the FNANM treatment of Triniteurybia, Brouillet (2006, p. 382) observed that "the cylindro-cam-

panulate heads with imbricate phyllaries and a wide green area, and the coarse, dentate foliage, are similar

to those of Eurybia. The lack of ray florets clearly distinguishes Triniteurybia.” In fact, lack of rays has been

the only morphological feature noted in any discussion that is diagnostic of the new genus. Triniteurybia

aberrans is the only rayless species in the eurybioid grade, but within subtribe Machaerantherinae sensu

stricto, Dieteria is is variable in ray production, Arida is rayless, and Xanthisma grindelioides is

rayless. Ray producti ble among species of Grindelia. Lack of ray florets may iue. recognition

of a taxon at specific rank, but there is no other example in the Astereae where it is the sole morphological

character for distinction of a genus.

Eurybia kingii and Furybia aberrans

In the positioning of Aster kingii within Tonestus (Nesom 1991), the species was noted (p. 124) to be most

closely similar to Tonestus aberrans: “Indeed, as a pair the two are somewhat set apart from the rest of the

genus in the toothed leaves often pais ponies teeth, narrowly lanceolate-attenuate and apically spreading

or reflexed phyllaries, styl with more widely arranged sweeping hairs, and purplish disc corolla

lobes.” And (p. 125) “in mie a its white rays, it fits more securely in [Tonestus] than in Aster, particularly

when placed next to T. aberrans.” Molecular evidence confirms that the two are closely related, and their

morphological and geographical similarity (Fig. 1) support this hypothesis. Both are treated here within

Eurybia.

Taxonomic status of Aster glaucodes subsp. pulcher Blake

Nesom (1994) treated Aster glaucodes subsp. pulcher at specific rank within Eurybia, as E. pulchra, distinct

from E. glauca. Eurybia pulchra “differs from E. glauca in its smaller leaves, apically acute phyllaries, and well-

developed glandularity. ... Eurybia pulchra has a restricted geographic range, and in the specimens I have

examined, there appears to be no intermediacy between it and E. glauca” (p. 194). The present study shows

that E. pulchra (northern Arizona and southern Utah) is sympatric with the more widespread E. glauca (Fig.

2) and confirms a paucity of intermediates between them. Plants of E. pulchra have distal stems, peduncles,

and phyllaries densely stipitate-glandular, while those of E. glauca are completely glabrous.

Numerous collections of both taxa have been made in the Grand Canyon, in the area of Havasupai

Canyon and roughly between Havasupai Canyon and Bright Angel Canyon, e.g.: Eurybia glauca: Clover

4513, 5136, 7018 (LL), Deaver 1524, 2135, 2677, 2944, 3053 (ASC), Hodgson 16026 (ASC); Eurybia pulchra:

Clover 7189 (LL), Deaver 3042 (ASC), Rink 4455, 4477a (ASC), Stevens s.n. [14 Aug 1992], s.n. [27 Sep 1992]

(ASC). Among all plants I have studied, the only three that might suggest the possibility of gene flow are

from this area: Watters 26 (ASC) and Deaver 4412 (ASC) are typical E. glauca, except that the peduncles of

each are minutely and sparsely glandular for about one centimeter immediately below the heads; Stevens

s.n. [10 Apr 1991] (MO) is glabrous except for sparsely glandular distal peduncles and glandular phyllary

margins.

A putative distinction noted by Brouillet et al. (2006) in number of disc florets (12-32 in Eurybia glauca,

29-40 in E. pulchra) is not confirmed here, as E. pulchra has florets evenly distributed in number down to

at least 19. Leaf and phyllary morphology also overlap, though E. pulchra does tend to have slightly smaller

leaves and more acute phyllaries. Within the area of sympatry, differences in habitat and phenology are not

apparent.

laftha Dat H ID hl bip, £T,

166 Journal of Texas 3(

In his decision to treat Eurybia pulchra at varietal rank within E. glauca, Brouillet (2003, p. 1561) noted

that “the ranges of the two taxa appear to be parapatric in southern Utah-northern Arizona with E. pulchra

restricted to the vicinity of the Grand Canyon while E. glauca ranges widely to the north and east into the

Southern Rocky Mountains; it must be noted however, that populations of the latter are found in Arizona

south and east of the Grand Canyon. Nonetheless, no mixed population of the two taxa has been reported.

The restricted range and distinct glandularity justify that E. pulchra be recognized as a taxon. Nevertheless,

glandularity alone does not seem to justify recognizing E. pulchra as a distinct species.” Brouillet's observation

that the lack of glands in E. glauca represents a derived condition is reasonable and probably correct but in

itself not pertinent to the decision of rank. The contrasting observation Bre that the ranges are sympatric

suggests that the distinct polarity in vestiture i ] through g lation, in which case specific

rank for each entity is justified. It is possible that in the southern part of the range (the range of E. pulchra),

some kind of genetic switching turns on or off the expression of glands, but similar variation does not occur

within other taxa of sect. Herrickia or any other Eurybia species. If E. pulchra is to be recognized at any rank

above “forma” the behavior of these two taxa as distinct species is the simplest hypothesis as the basis for

assignment of rank. Field study clearly is needed.

Welsh (2003) recognized the distinctiveness of Eurybia puchra and treated it as a variety of E. wasatch-

ensis rather than a closer relative of E. glauca. The implication by Brouillet et al. (2006) that E. pulchra and

E. glauca are sister taxa is better supported by morphology.

TAXONOMIC SUMMARY OF EURYBIA SECT HERRICKIA

Eurybia sect. Herrickia (Woot. & Standl.) Nesom, Phytologia 77:258. 1994. Herrickia (Woot. & Standl.). Triniteurybia

Brouillet, Urbatsch & Roberts, Sida 21:898.

Lack of a clear diagnosis is the herrickioid group, apart f I ingly i lated species cl d the geographic col

of the species, rend of sect. Herrickia weakly justified. Nevertheless, it i gory in reference to

the group.

l. reo aberrans (A. Nels.) G.L. Nesom, comb. nov. Basionym: Macronema aberrans A. Nels.; Haplopappus aberrans (A.

1s.) H.M. Hall; ra pti (A. Nels.) Rydb.; Tonestus aberrans (A. Nels.) Nesom & Morgan; Triniteurybia aberrans (A.

v ) Brouillet, Urbat

2. Eurybia glauca (Nutt.) Nesom, Phytologia 11:260. dd Hiephahs glaucus Nutt.; ickia gl (Nutt.) Brouillet;

Aster glaucus (Nutt.) Torrey & A. Gray 1841 (non Nees 1818); A od

3. Eurybia horrida (Woot. & Standl.) Nesom, Phytologia 77:260. 1994. Herrickia horrida Woot. & Standi.; Aster hor-

ridus (Woot. & Standl.) Blake.

4. Eurybia kingii (D.C. Eaton) G.L. Nesom, comb. nov. Basionvm: Aster kingii D.C. Eaton in S. Watson, Botany (Fortieth

Parallel) 141, plate 16, figs. 1-6. 1871. Mni ee (D.C. Eaton) Cronq. & Keck; Tonestus kingii (D.C. Eaton) Nesom;

Herrickia kingii (D.C. Eaton) Brouillet, Urbats

4a. ee kingii (D.C. Eaton) G.L. Nesom var. barnebyana (Welsh & Goodrich) G.L. Nesom, comb.

NOV. Basionym: Machaeranthera kingii var. barnebyana Welsh & Goodrich, Brittonia 33:299, fig. 5. 1981. Aster kingii var. barnebyana

(Welsh & Goodrich) — e iin var. barnebyana (Welsh & Goodrich) Nesom; Herrickia hingii var. barnebyana (Welsh &

Goodrich) Brouillet, Urbat

4b. Eurybia kingii (D.C. Eaton) G.L. Nesom var. kingii

5. Eurybia pulchra (Blake) Nesom, Phytologia 77:261. 1994. Aster glaucodes subsp. pulcher Blake; Aster glaucodes var.

pulcher (Blake) Blake; Aster wasatchensis var. pulcher (Blake) Welsh; Eurybia glauca var. pulchra (Blake) Brouillet; Herrickia glauca var.

pulchra (Blake) Brouillet.

6. Eurybia wasatchensis (M.E. Jones) Nesom, Phytologia 77:262. 1994. Aster glaucus var. wasatchensis M.E. Jones;

Aster wasatchensis (M.E. Jones) Blake; Eucephalus wasatchensis (M.E. Jones) Rydb.; Herrichia wasatchensis (M.E. Jones) Brouillet.

CONCLUSION

While the overall understanding of the herrickioid group is essentially the same as that of Brouillet et al.

Nesom, T. i i fE yl i t. Herrickia 167

(2004) and Brouillet (2006), the taxonomic alternative proposed here conveys an equally or more useful

picture of the evolutionary standing of these species. Treating them within Eurybia, while explicitly point-

ing out evolutionary relationships suggested by the cladistic hypothesis, emphasizes their morphological

similarity and geographic coherence. The Brouillet et al. taxonomy emphasizes the evolutionary discrete-

ness of species that are morphologically inseparable by characters used elsewhere in the tribe and family

to distinguish genera. The formal recognition of a paraphyletic group at generic rank, compared to the

cladistically-derived alternative, has merit in this extreme example where morphology plays essentially no

role at all in the classification. Further research may resolve the issue with less ambiguity.

ACKNOWLEDGEMENTS

I am grateful for loans from ASC and SJC, assistance of staff during study at BRIT, MO, and TEX-LL, speci-

men data from NMC for Eurybia horrida, and comments on the manuscript from John Pruski and three

anonymous reviewers.

REFERENCES

BrouiLLEr, L. 2003. New combination in Eurybia (Asteraceae: Astereae) from North America. Sida 20:1561-1563.

BrouiLter, L. 2006. Eurybia, Herrickia, Triniteurybia. |n: Flora of North America Editorial Committee, eds. Flora of North

America north of Mexico. Oxford University Press, New York and Oxford. 20: 365-382; 361-365; 382-383

BrOuILLeT, L, L.E. URBATSCH, AND R.P. Roserts. 2004. Tonestus kingii and T. aberrans are related to Eurybia and the

Machaerantherinae (Asteraceae: Astereae) based on nrDNA (ITS and ETS) data: Reinstatement of Herrickia

and a new genus, Triniteurybia. Sida 21:889—900.

BROUILLET, L. AND S. SELLIAH. 2005. Symphyotrichum pygmaeum: transfer of Eurybia pygmaea from the eurybioid

grade to the subtribe Symphyotrichinae (Asteraceae: Astereae). Sida 21:1633-1635.

BRouiLLET, L., J.C. SEMPLE, G.A. ALLEN, K.L. CHAMBERS, AND S.D. SUNDBERG. 2006. Symphyotrict In: Flora of North America

Editorial Committee, eds. Flora of North America north of Mexico. Oxford University Press, New York and

Oxford. 20:465-539.

FNANM EbrroriaL Comittee. 2006. Magnoliophyta: Asteridae, part 7: Asteraceae, part 2. Vol. 20. Flora of North

America North of Mexico. Oxford University Press, New York and Oxford.

Hatt, H.M. 1928. The genus Haplopappus, a phylog ic study in the Compositae. Carnegie Inst. of Washington

Publ. No. 389, Washington, D.C.

McVauch, R. 1945. The genus Triodanis Rafinesque, and its relationships to Specularia and Campanula. Wrightia

1:13-52.

Nesom, G.L. 1991. Transfer of Aster kingii to Tonestus (Asteraceae: Astereae). Phytologia 71:122-127.

Nesom, G.L. 1994. Review of the taxonomy of Aster sensu lato (Asteraceae: Astereae), emphasizing the New World

species. Phytologia 77:141-297.

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. 2009 (in press). Eurybia (Asteraceae). In Heil, K.D., S. O'Kane, and L. Reeves, eds. Flora of the Four Cor-

ners Region: vascular plants of the San Juan River Drainage - Arizona, Colorado, New Mexico, Utah. Missouri

Bot. Garden Press, St. Louis.

Nesom, G.L. ano D.R. Morcan. 1990. Reinstatement of Tonestus (Asteraceae: Astereae). Phytologia 68:174-180.

NORDAL, |. AND B. STeDJE [and 148 signatories]. 2005. Paraphyletic taxa should be accepted. Taxon 54:5-8.

SELLIAH, S. AND L. BrouiLtet. 2007. Molecular phylogeny of the North American eurybioid asters, Oreostemma, Her-

rickia, Eurybia, and Triniteurybia (Asteraceae, Astereae) using a low copy nuclear gene, CNGCA, a. Abstract

1229, Botany & Plant Biology 2007, Joint Congress (Chicago). «www.2007.botanyconference.org» Accessed

Oct 2007.

SHELLY, J. S. AND M. MANTAS. 1993. Noteworthy collections, Montana. Madrono 40:271-273.

WELSH, S.L. 2003. A Utah flora (ed. 3, rev). Brigham Young Univ. Press, Provo.

168 Journal of the Botanical R h Institute of Texas 3(

BOOK REVIEW

Tuomas G. Barnes, DEBORAH WHITE, AND Marc Evans. 2008. Rare Wildflowers of Kentucky. (ISBN 978-0-

8131-2496-4; hbk.). The University Press of Kentucky, 663 South Limestone, Lexington, Kentucky

40508-4008, U.S.A. (Orders: www.kentuckypress.com, Hopkins Fulfillment Service, RO. Box 50370,

Baltimore, Maryland 21211-4370, 1-800-537-5487, 410-516-6998 fax). $39.95, 220 pp., 220 full-color

ee 10" x 8 1/2".

1 1 1 11 : 1 : JST d LJ 1 "e ds 1 ls]

y 5 y losing aU P P

l. These activi-

1 J > : J; MES 1 Y 1 fl o iy co Hot H 1 El 1 A A th t d

wildflowers. T

an additional 52 of special concern, and 60 species (historic) have not Mu seen in 20 or more a a color i images were ño with

film Men s transition to po cameras. C

TS » | J E 1

he text Photographs were by Thomas G. Barnes, mmn,

The first sections of the book are divided into a discussion of native species: Reasons for Decline of Native Pro Reasons for

Protecting Native Plants, Natural ne of Kentucky ene a map that shows the nae ceps of the pate, Rarity, and

Extinction and the Preservation Bt | EN ti

any one wl of our native

(hh

floras. An all too familia

I ted: human lifestyl d ] d ) lt in Habitat Loss, Invasive

Exotic oe Pollution, bn ci e There is a thoughtful and compelling discourse on reasons for protecting native plants.

d by pean a and habitat color images.

e section on TI is divided i 96), Prairies and pare as In and Hone Rivers n 13D,

Wetlands (133-160), and Cliff Lines and Rock! (161-166). The fl

against a a dark or black background.

of plant names (both common and SCIRE d as well as rare un isu dd a ma a

(E= EL T=threatened, S Icon H-histo

A list of References and E follows

Tl ] 21115 n 3 1 FinlA TH : 1 > PAR | if, 1 1

1 [1 12€ anal

but in many cases

1 : Y;

ide. The threatened biodiversity, esthetic beauty of flowers, and the appeal for more

f£] ] f, f, I qu 1 d: 1 1 1 d `} E 1 tat

conservation efforts to save Hablas and larger

flora of Kentucky.—Harold W. Keller, Ph.D., Research Associate, Botanical Research Institute of Texas, Fort Worth, Texas 76102-4025, U.S.A.

J. Bot. Res. Inst. Texas 3(1): 168. 2009

THE PHYLOGENY OF SELENIA (BRASSICACEAE) INFERRED FROM

CHLOROPLAST AND NUCLEAR SEQUENCE DATA

James B. Beck

E of buo

Duke University

T Not CU 27708, U.S.A.

es.be

ck@duke.e

ABSTRACT

Selenia Nutt. (Brassi ) is a North American g f five species distril lf ] land ] U.S.A. to o

Moa 17131 hal : y els g 1 1 11 calliehad , Very elsi] 1 ha hin] gy find.

or ic relationsl I In this study, DNA sequence iation fi I I i 1 ibed spacer and

haul Tl Š dom f £1; 1 1 " J

four non- coding a regions he v trnL intron; pan the n NOR Miu de n MR ON Lm quus was used

A Yd 2l : a 1 1 1 ];

I ] resolved intra pnylogeny

hl 1 1 indi 1 sonal hybridization Denda Selenia grandi d Seleni i The phylogenetic distinctiveness

of S. mexicana, “knows om none few collections in Nuevo León, combined with the relative lack of collections from Mexi ggested

hat add in this group.

RESUMEN

Selenia Nutt. (Brassicaceae) es un gé i cinco especies que se distribuyen desde el centro y sudoeste de Estados

Unides hasta] te de México. Aunque ! ía bási grup tá bien establecida, se sabe muy poco e la pene de

1 ; 1 Toss f Ati 1 g

e E

en las secuencias del espaciador de e interna “específico del ADN ribosomal (ADND; dificantes del

ADN pue (el intrón trnL 2 I g petå-psbJ, trnQ- eee y srr) Los analisis de máxima p

1 £5 As] Á

monia yl o r E r T e

completamente resuelta El conflict las topologías obtenid ADN nuclear y cloroplástico indican hibrid histérica entre

elenia grandis y listinción filogenética de S. mexicana, ida d pocas localidades en Nuevo León,

junto con las p g | pera descubrir más diversidad en este M

INTRODUCTION

Selenia Nutt. (Brassicaceae) is a distinctive genus of five species distributed from the central and southwest-

ern U.S.A. to northern Mexico (Fig. 1). Selenia species are small («50 cm tall), spring flowering, herbaceous

annuals found on a wide range of often seasonally wet habitats from sandstone glades (S. aurea Nutt.) to

limestone hills (S. dissecta Torr. & A. Gray) to alluvial soils (S. grandis R.F. Martin) (Rollins 1993). Selenia

can be easily distinguished from all other genera of the tribe Cardamineae by a combination of an annual

habit, fully bracteate inflorescences, yellow flowers, silicles with distinct style, and biseriate, broadly winged

seeds. Although S. aurea and S. dissecta can be found in multiple states, known from 57 and 14 counties,

EE L S. a as S. jonesii Cory are endemic respectively to southern and western Texas in the

L is known only from the Mexican states of Coahuila and Nuevo León, although

the individual ranges of this species and of $ dieu, S. jonesii, and S. grandis will surely expand following

additional fieldwork in northern Mexico. This | graphic uncertainty is rey ive of a basic lack of

knowledge regarding Selenia, and little to no alora oa exists concerning the reproductive biology, ecol-

ogy, and phylogenetic relationships within this distinctive group (Al-Shehbaz 1988).

Although both morphological and biogeographic patterns within Selenia suggest certain null phyloge-

netic hypotheses, no study has addressed these evolutionary relationships. Selenia aurea is morphologically

divergent from its congeners, with unappendaged sepals, pinnate (vs. bipinnate) leaves, and relatively long

(>5 mm) styles. These features and its disjunct range (Fig. 1) suggest an isolated phylogenetic position for

J. Bot. Res. Inst. Texas 3(1): 169 —176. 2009

170 Journal of the Botanical R h Institute of Texas 3(1)

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. N E y 5n

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this species. Within the remaining species, several characters, including possession of a horn-like (vs.

pouch-like) sepal appendage and relatively long (>8 mm) sepals and anthers (22.5 mm) suggest that S.

dissecta, S. grandis, and S. mexicana form a natural group. These patterns of shared character variation, the

morphological cohesiveness of individual species (Al-Shehbaz 1988), and the small size of the genus sug-

gest that reconstructing the evolutionary relationships within this distinctive North American taxon will

be tractable. This study aims to resolve the phylogenetic relationships between the five recognized species

of Selenia using both chloroplast and nuclear DNA sequence variation.

MATERIALS AND METHODS

Taxon Sampling

Cols 1 ]

Sample information appears in Appendix 1. Ten d, including two samples from

each of the five species recognized in perhaps the most focused examination of the genus (Martin 1940).

Three taxa that have been recognized by certain authors were not included, S. jonesii var. obovata Rollins, S.

Beck, Phylogeny of Selenia 171

aperta (S. Watson) Small, and 5. ipd Steyerm. pos first taxon is known only from the type collection

(Rollins 1993), and the di te fruits) has been considered to be an artifact of

pressing the inflated fruits (Al-Shehbaz in ms.). Martin (1940) provided a detailed discussion of the lack of

distinctiveness of both S. aperta and S. oinosepala, although both taxa warrant additional study (see discus-

sion). Leavenworthia Torr. has been shown to be sister to Selenia, and the monotypic genus Planodes Greene

has been shown, along with Barbarea R. Br., to be sister to the Selenia/Leavenworthia clade (Beilstein et al.

2006). Two Leavenworthia samples [L. uniflora (Michx.) Britton and L. alabamica Rollins] and a P. virginicum

(L.) Greene sample were therefore used as outgroups. Nine of ten Selenia samples were obtained from her-

barium material, and collections made as early as 1958 (see Appendix 1) yielded successful amplifications

and sequences.

Molecular Methods

Extractions were performed with either a Qiagen (Qiagen, Valencia, CA) DNeasy Plant Mini Kit or a Viogene

(Viogene U.S.A., Sunnyvale, CA) extraction kit. The nuclear internal transcribed spacer (ITS) region was

amplified using the primers "ITS 1" (White et al. 1990) and either "ITS 4" (White et al. 1990) or "ITS2-

265.4” (Rauscher 2002). The trnL"^^ intron was amplified using the primers “C” and *D" (Taberlet et al.

1991). A portion of the chloroplast trnSSU-trnGUY intergenic spacer was amplified with the primers “1F”

and “1R” (Säll et al. 2003). A portion of the chloroplast petA-psbJ intergenic spacer was amplified with the

primers “5F” and “5R” (Säll et al. 2003). A portion of the chloroplast trnQ""9—rps16 intergenic spacer was

amplified with the primers *trnQU"*" and *rpS16x1" (Shaw et al. 2007). All reactions were performed under

standard conditions. Products were visualized and purified via agarose gel electrophoresis with a Viogene

gel extraction kit. Products were dye-labeled using a Big Dye Terminator Kit (Applied Biosystems, Foster

City, CA), and analyzed on either a MJ Research BaseStation (MJ Research, Waltham, MA) or an Applied

Biosystems 3130xl Genetic Analyzer. All sequences have been deposited in the EMBL nucleotide sequence

database (Appendix 1).

Phylogenetic Analyses

The ITS and combined chloroplast (trnL, trnS-trnG, petA-psbJ, and trnQ—rps16) datasets were analyzed sepa-

rately. Sequences were manually aligned in Se-Al (Rambaut 2002) and the aligned matrix was exported as

a NEXUS file. All insertion/deletion (indel) events, both autapomorphic and synapomorphic, were scored

except in the case of nucleotide rep resulting in more than two indel character states (which were viewed

as likely homoplasious), or in regions of uncertain alignment. In the case of overlapping indel events, the

"simple gap coding" method of Simmons and Ochoterena (2000) was used. All positions involved in indels,

or situated in regions of uncertain alignment were deleted prior to analysis, and indel events were coded

as pagus i dd and added to the end of the NEXUS file. Positions exhibiting poor sequence or

additivity (multi ks presumably due to the presence of divergent ITS sequences in a single individual)

were coded as bo For each dataset a heuristic maximum parsimony search with 100 random addi-

tion replicates was performed using PAUP* 4.0b10 (Swofford 2002) with the following parameters: starting

trees obtained by stepwise addition, TBR branch swapping, “MulTrees” turned on, and steepest descent not

in effect. Ten thousand bootstrap replicates were cond ] with PAUP* 4.0b10 in order to obtain bootstrap

Suppor e s uas. model of sequence evolution for each DNA region (indels and poorly aligned

1 using the Akaike Information Criterion in Modeltest 3.06 (Posada & Crandall

1998), and a Bayesian Markov Chain Monte Carlo analysis was performed on each dataset in MrBayes 3.1.2

(Huelsenbeck & Ronquist 2001). The combined chloroplast data were analyzed as a partitioned dataset,

with the best-fitting model of sequence evolution for each separate region assigned to the corresponding

partition (see Table 1). For both the ITS and combined chloroplast analyses, the indel characters were as-

signed the binary model of character evolution (Nst=1, Coding=Variable) as recommended in the MrBayes

documentation. All Bayesian analyses comprised four independent runs, with four chains (one cold and

three heated). Flat priors were used, with the exception of the rate prior that was set to allow rates to vary

among partitions. Chains were run for 5 million generations, and trees were sampled every 1000 genera-

tions. Stationarity was evaluated by examining the standard deviation of split frequencies among runs and

172 Journal of tl tanical h Instit Texas 3(

Taste 1. Relative phylogenetic information and model of seq lution chosen in each ofthe five DNA regi lyzed. ‘includes one inversion event. "Only a subset of the

Sequence characteristic ITS trL petA-psbJ trnS-trnG trnQ-rps16

Aligned length (bp) 520 578 397 301 523

Analyzed characters, including indels 516 327 286 470

Variable characters, including indels (%) 107 (21%) 23 (5%) 31 (9%) 13 (5%) 35 (7%)

Parsimony informative characters, 92 (18%) 9 (2%) 8 (2%) 3 (1%) 12 (3%)

including indels (%

Number of indels (parsimony informative) 3 (2) 7 (4) 8 (3) 3 (0) 9? (43)

substitution model selected by Modeltest TVMef-H K8luf+l — K8luf K81uf+l K81uf

substitution model implemented in MrBayes? — GTR+1 GTR-H GTR GTR-H GTR

by plotting the log likelihood values from each run using Tracer 1.4 (Rambaut & Drummond 2007). These

diagnostics indicated that runs reached stationarity quickly (within 100,000 generations) and I conserva-

tively excluded the first 500,000 generations before obtaining a consensus phylogeny and clade posterior

probabilities (PP).

RESULTS

Details regarding the length, variability, and model of sequence evolution chosen for each gene region are

presented in Table 1. The analyzed ITS matrix of 516 characters yielded 107 (2196) variable and 92 (1896)

parsimony-informative characters. The matrix contained 10 (0.196) cells coded as either missing or ambigu-

ous. Additivity, indicated by multiple peaks at a single nucleotide position, was limited to three samples.

Two samples (S. aurea sample 1 and S. mexicana sample 2) exhibited multiple peaks at one position each,

while S. jonesii sample 2 exhibited multiple peaks at four positions. At each of these four 5. jonesii positions

one of the two inferred nucleotides matched that from the other S. jonesii sample, with the other nucleotide

a E CÓ typically observed in several congeners and the outgroup. Two of the three insertion/

were parsimony-informative. Each of the 100 random addition replicate parsimony searches

using the ITS dataset usi the same island of five most parsimonious trees (MPTS (length = 141, con-

sistency index = 0.86, retention index = 0.91). One of the five MPTs, along with bootstrap percentages and

Bayesian posterior probabilities, is shown in Figure 2a. The ITS data provided low support for the monophyly

of Selenia (0.60 PP, 38% BS), but strong support (1.0 PP, 100% BS) for a “core Selenia" clade comprising S.

jonesii, S. grandis, S. dissecta, and S. mexicana. The ITS data also provided strong support (0.97-1.0 PP) for the

sister relationship of each pair of conspecific samples. Certain chloroplast regions failed to amplify in four

samples (the trnQU"6— rps16 intergenic spacer in S. aurea sample 2, S. mexicana sample 2, and the L. alabamica

sample and the trnSGC"-trnG""€ intergenic spacer in S. dissecta sample 2). These samples were excluded from

the combined chloroplast analysis. The analyzed chloroplast matrix of 1576 characters yielded 102 (6%)

variable and 32 (296) parsimony-informative characters. The chloroplast data matrix contained 29 (0.2%)

cells coded as either missing or ambiguous. Eleven of the 27 insertion/deletion events were parsimony-

informative. Each of the 100 random addition replicate parsimony searches using the combined chloroplast

dataset recovered the same MPT (length = 112, CI = 0.95, RI = 0.88). The MPT, along with bootstrap per-

centages and Bayesian posterior probabilities, is shown in Figure 2b. Unlike the ITS data, which provided

minimal support for the monophyly of Selenia, the combined chloroplast dataset strongly indicated such a

relationship (1.0 PP, 99% BS). Strong support was also provided for the monophyly of “core Selenia” (1.0 PP,

100% BS), and two additional clades nested within this group. Although the four-locus chloroplast dataset

was only able to assess the sister relationships of each pair of S. jonesii (1.0 PP, 100% BS) and S. grandis (1.0

PP, 95% BS) samples, a dataset including only trnL intron and petA-psbJ intergenic spacer sequences for all

10 Selenia samples provided strong support for the sister relationship of each pair of conspecific samples:

173

Beck, Phylogeny of Selenia

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S. aurea (1.0 PP, 100% BS), S. dissecta (1.0 PP, 90% BS), and S. mexicana (1.0 PP, 86% BS). The only conflict

between the chloroplast and nuclear topologies involved the placement of S. mexicana, which was sister to

S. dissecta in the nuclear topology and sister to S. grandis in the chloroplast topology.

DISCUSSION

Selenia phylogeny

In general the nuclear and chloroplast datasets provide well-resolved and strongly supported phylogenetic

reconstructions that are not only congruent with each other (Fig. 2), but with patterns of shared morpho-

logical character states. The core Selenia clade comprising S. jonesii, S. grandis, S. dissecta, and S. mexicana is

distinguished by sepals bearing a dorsal appendage, bipinnate (vs. pinnate) leaves, and relatively short («6

mm) styles. A more exclusive clade comprising S. grandis, S. dissecta, and S. mexicana is distinguished by

possession of a horn-like (vs. pouch-like in S. jonesii) sepal appendage and relatively long (>8 mm) sepals

and anthers (>2.5 mm). Interestingly, even though the Selenia key of Martin (1940) is artificial and there-

fore doesn’t necessarily imply relatedness, if it is viewed as a bifurcating tree it perfectly matches the ITS

topology presented in Figure 2a. The data clearly support the recognition of S. mexicana as distinct from S.

dissecta (see below), suggesting that additional diversity is yet to be documented in this group. The three

taxa not analyzed in this study (S. jonesii var. obovata, S. aperta, S. oinosepala) should therefore be subject to

future molecular and morphological evaluation. Selenia aperta is a particularly intriguing case. This taxon

was originally described as a variety of S. aurea by Watson (1895) based on material from San Augustine

County, Texas, which exhibited broadly inflated silicles, a reduced septum, and a relatively long style. The

variety was later given species status by Small (1903). Martin (1940) failed to locate the type material but

examined both potential types and other specimens exhibiting these character states. He observed the

variation described by Watson but found both variation among individuals from a single collection and no

specimens that exhibited the full complement of characters. This variation, the potential disjunct range of

S. aperta (noted in Fig. 1), and the chromosome number variation noted by Rollins and Rüdenberg (1977)

all suggest that additional lineages remain to be identified within S. aurea.

Potential Hybridization and Chloroplast Capture

The only incongruence between the two topologies is the placement of S. mexicana, and morphological and

biogeographical evidence suggest that the nuclear placement (as sister to S. dissecta) is correct and that the

anomalous placement of S. mexicana by the chloroplast data is due to historical hybridization between S.

mexicana and S. grandis followed by chloroplast capture. Chloroplast capture has been well documented em-

pirically (Rieseberg & Soltis 1991), and appears to be possible under a range of biologically realistic situations

(Tsitrone et al. 2003; Chan & Levin 2005). From a morphological standpoint, S. dissecta and S. mexicana are

difficult to distinguish, with the latter exhibiting shorter («2.5 vs. »3.5 mm) styles and spongy (vs. winged)

seed margins, and recent workers (Al-Shehbaz 1988; Rollins 1993) have considered it a synonym of S. dis-

secta. Although the ranges of most Selenia taxa are poorly known, the existing biogeographic data also lend

support to the proposed hybridization scenario, as known populations of S. mexicana are approximately

250 km closer to known populations of S. grandis than they are to populations of S. dissecta (Fig. 1). This

evidence notwithstanding, S. mexicana is by far the most poorly known Selenia species, and additional cyto-

logical, genetic, and field studies are needed to thoroughly test this hypothesized gene flow. Selenia dissecta

has been reported to be 2n - 14, while S. grandis is known to be 2n = 24 (Warwick & Al-Shehbaz 2006).

The currently unknown chromosome number of S. mexicana will therefore reveal if the proposed hybridiza-

tion event was via a polyploid or homoploid pathway. Evidence of additivity in the S. grandis or 5. mexicana

ITS sequences was limited to a single position in S. mexicana sample 2, indicating that any heterospecific

ITS repeats have been largely eliminated due to backcrossing to conspecifics (the homoploid scenario) or

concerted evolution (Franzke & Mummenhoff 1999). Particularly in a homoploid hybridization scenario,

sequencing both individual cloned ITS sequences and other nuclear loci in an expanded sample set from

across both species’ ranges will reveal the extent of proposed introgression. Three individuals exhibiting

Beck, Phylogeny of Selenia 175

S. mexicana morphology (short styles in particular) were discovered in a recent survey of specimens from

six major herbaria (BRIT, GH, MO, NY, TEX-LL, and US), all from within 100 km of the type locality of

Galeana, Nuevo León (Standley 1937). Unfortunately, these collections contain few specimens from Mexico,

and additional fieldwork is clearly needed. As noted in the introduction, the range of S. mexicana is but one

of many aspects of Selenia species that are poorly documented. Future work, including expanded sampling

of all proposed Selenia taxa, is therefore needed to understand the biology of this genus, information that

can now be placed in an evolutionary context.

APPENDIX 1

Sample information. Taxon (sample number), voucher, year of collection (if herbarium material) try, state, county

(if applicable), ITS EMBL, trn£ EMBL, trnS-trnG EMBL, petA-psbJ EMBL, trnQ-rps16 EMBL.

oe alabamica Rollins, Beck 486 (MO)—U.S.A. ALaBAMA. Franklin Co.: FM957596, FM957609, FM986404, FM986416,

non

menso po pal Britton, Beck 516 (MO) —U.S.A. ALasama. Morgan Co.: FM957595, FM957608, FM986403,

nive e n pm Al-Shehbaz s.n. (MO)—U.S.A. Missouri: FM957594, FM957607, FM986402, FM986414,

428

Selenia aurea Nutt. (1), Beck 774 (MO)—U.S.A. Missouri. St. Clair Co.: FM957598, FM957611, FM986406, FM986418,

Selenia aurea Nutt. (2), Stephens 29996 (GH) 1969—U.S.A. OxtAHoMA. Nowata Co.: FM957597, FM957610, FM986405,

FM9 ,none

Selenia dissecta Torr. & A. Gray (1), Worthington 11630 (NY) 1984—MEXICO. Chihuahua, FM957600, FM957613, FM986407,

F 20, FM986431

Selenia dissecta Torr. & A. Gray (2), Correll 38395 (TEX-LL) 1970—U.S.A. Texas. Culberson Co.: FM957599, FM957612, none,

FM986419, FM986430

Selenia Pas E F. Martin (1), Turner 4323 (TEX-LL) 1958—U.S.A. Texas. Nueces Co.: FM957603, FM957616, FM986410,

FM 986435

Selenia ards E F. Mei (2), Correll 36762 (TEX-LL) 1969—U.S.A. Texas. Hidalgo Co.: FM957604, FM957617, FM986411,

a jonesii oie (1 pum & Lundell 16958 (GH) 1961—U.S.A. Texas. Dawson Co.: FM957601, FM957614, FM986408,

FM986421, FM986432

Selenia jonesii Cory (2), Mahler 8846 (GH) 1981—U.S.A. Texas. Reagan Co.: FM957602, FM957615, FM986409, FM986422,

F 643

Selenia mexicana Standl. (1), Hinton 27036 (TEX-LL) 1997—MEXICO. Nuevo León: FM957605, FM957618, FM986412,

Eden Eee piso

(2), Crutchfieid & Jot (GH) 1960—MEXICO. Nuevo León: FM957606, FM957619, FM986413,

FM986426, none

ACKNOWLEDGMENTS

The author would like to thank the curators of BRIT, GH, MO, NY, TEX-LL, and US for herbarium loans.

Special thanks go to Ihsan Al-Shehbaz for critical discussions, to George Yatskievych for help with locating

Selenia aurea, to Eric Schuettpelz for help with Bayesian analyses, to Lisa Pokorny for providing the Span-

ish abstract, and to Ray Collier for laboratory assistance. Manuel B. Crespo (ABH) and two anonymous

reviewers provided helpful comments on the manuscript. This study was funded by the Graduate Studies

Program at the Missouri Botanical Garden and the Division of Biology and Biomedical Sciences at Wash-

ington University in St. Louis.

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Á 2r

PRELIMINARY INSIGHTS INTO THE PHYLOGENY AND SPECIATION OF

SCALESIA (ASTERACEAE), GALÁPAGOS ISLANDS

Jeremy D. Blaschke Roger W. Sanders

uu Pane #7071 Bryan College # 7802

721 Bryan Dr.

Dayton, Rae m U.S.A. Dayton, lennessee 37321, U.S.A.

jblaschke0998@bryan.edu rsanders@bryan.edu

ABSTRACT

Scalesia Arn. (A Heliantl )i lyg f fifi peci demi I lapagos Island ee distribution

J : ; C pb RE 3 £. 1 1; 1 1 1 All species

of Scalesia Pappo lus SE Blake and Sim sia Pers., along with selected species of Viguiera Kunth, were ejected to phy logenetic analysis

(63 characters in 78 taxa). Homoplasy and i g g lting pud resolutio d comparison

of speciation events dd: to its eds du dd phologically, species throug! g ked primarily by homoplastic

apomorp ies. Base fhabit ph 1 gi lai g f, 1 f, 1 y Lab; pr 1

poorly correlated. cup for f dies should b ded to include other groups in the derived Helianthinae

RESUMEN

a (A Heli I ) gé lef d I i p ies, dcus t tasta láp g 1 1 5 lógi

Lito 1 x T J aie [ad 1 1 Li J Tye Oe P p fs ] * ah

seleccionados. Todas | ies de Scalesi TP S.F. Blake, y Simsia Pers., junto con algunas E de Wiewers Kunth, fueron

eae 1 un análisis Ei inodo iid caracteres en ids nni La ee e Ud M. entre zs árboles epe idw la

E 4 E r E -

E ; : fine] leet Bacadecen | pli ización de los hábitat q divergencia

1 f, di Lb

INTRODUCTION

1: L: y . Ef, species, all J a trat? land láp

Scalesia Arn. (Asteraceae: Heliantheae: E

Islands. Howell (1941) accepted 18 species in four Secun and related it to the Ecliptinae Lessing. Based

on extensive field study, Eliasson (1974) recognized only 14 species, avoided the use of sections, discussed

aspects of character evolution, and placed the genus in the Helianthinae Dumort. Hamann and Wium-

Andersen (1986) described an additional species.

Recent studies on Scalesia have focused on intergeneric relationships (Schilling et al. 1994; Spring et

al. 1999; Panero 2007), chemical diversity (Adsersen € Baerheim Svendsen 1986; Spring et al. 1997, 1999;

Petersen et al. unpubl.), anatomy (Carlquist 1982), autecology (Itow 1995; Kitayama & Itow 1999; Hamann

2001), adaptive reproductive strategies (McMullen & Naranjo 1994; Nielsen et al. 2002, 2007), and popula-

tion structure (Nielsen et al. 2003; Nielsen 2004). Chloroplast DNA restriction site analysis suggests that

Scalesia belongs to a group of specialized genera, the “derived Helianthinae,” that are embedded within a

derived clade of Viguiera Kunth (Schilling et al. 1994). Viguiera, whose taxonomy has been unresolvable on

morphological grounds, appears as a paraphyletic assemblage basal to all other genera in the Helianthinae

on the basis of cpDNA restriction sites and internal transcribed spacer (ITS) sequences of nuclear ribosomal

DNA (Schilling & Jansen 1989; Schilling & Panero 2002). In an analysis in which S. pedunculata Hook.f. and

several species of Pappobolus S.F. Blake were sampled (Schilling et al. 1994), Scalesia and Pappobolus were

sister groups, and the next closest clade consisted of Simsia Pers. plus Viguiera ser. Pinnatilobatae S.F. Blake.

However, the authors noted problems with interpretation of the three restriction sites synapomorphic for

Scalesia and Pappobolus and concluded, "Thus, the relative relationships among Scalesia, Pappobolus, Simsia

J. Bot. Res. Inst. Texas 3(1): 177 —191. 2009

178 Journal of t tani Insti Texas 3(

and Viguiera ser. Pinnatilobatae are not well resolved by these data." Indeed, subsequent work has shown that

chloroplast restriction sites, chloroplast genes, and ITS regions lack sufficient variation to resolve relation-

ships in the derived Helianthinae (Schilling & Panero 1996, 2002; Petersen et al. unpubl.). However, recent

analyses using external transcribed spacer (ETS) regions did find sufficient diversity to resolve species of

Helianthus (Timme et al. 2007), a member of the derived Helianthinae, as well as species in other genera of

the Heliantheae, such as Montanoa (Plovanich & Panero 2004).

Our interest is primarily in patterns of diversification, homoplasy, speciation rates, and degree of ad-

aptation using Scalesia because it is the most speciose endemic angiosperm genus in the Galápagos. It is of

interest that homoplasy among morphological characters of Simsia species prevented Spooner (1990) from

publishing a cladistic analysis in his monograph. Likewise, Panero (1992) chose not to include phylogeny

in his monograph of Pappobolus, instead recognizing only phenetic groupings. Thus, notable amounts of

unanalyzed data are available in the literature for addressing the issue of homoplasy across Scalesia and

relations. While it is our hope in the future to sample ETS regions in Scalesia species to determine their ap-

plicability in phylogenetic analysis, as well as use pr Mp: in investigating issues of interest to

us, our goal here is to mine the existing pertinent nd ecological data that are available in the

literature to provide a comparative context for later molecular ce That is, we seek to provide insights

into: 1) sister-group and intrageneric relationships of Scalesia, 2) homoplastic traits, 3) relative amounts

of speciation per clade, and 4) directions for future molecular sampling. We anticipate that phylogenetic

analysis of morphology may not yield consistent assessments of relationships or be congruent with mo-

lecular phylogenies (for example, see Plovanich and Panero [2004] concerning homoplasy in morphological

taxonomic criteria in the Heliantheae). However, that result is not certain, for the hand-calculated Wagner

parsimony networks of morphological data of Dendroseris and Robinsonia (Sanders et al. 1987), two other

island endemics of the Asteraceae (Lactuceae and Senecioneae, respectively), did prove to be cong with

later molecular phylogenies (Crawford et al. 1992; Sang et al. 1995).

METHODS

Data.—Taxon sampling is based on the sister-group conclusions and Figure 1 of Schilling et al. (1994)

and availability of supplemental specimens at the Botanical Research Institute of Texas and Bryan College.

Morphological traits, coded as binary and multistate unordered characters, were extracted from published

monographs of Pappobolus (Panero 1992), Scalesia (Eliasson 1974; Hamann & Wium-Andersen1986), Simsia

(Spooner 1990), and species representing Helianthus L. (Schilling 2006), Viguiera sect. Maculatae (S.F. Blake)

Panero & Schilling (Panero & Schilling 1988), and the outgroup Bahiopsis Kellogg (Schilling 1990). Selected

dried specimens were consulted to verify codings obtained from the literature, supply missing data, and

score representative species from Viguiera ser. Grammatoglossae S.F. Blake and ser. Pinnatilobatae (Table 1).

Characters were chosen to maximize distinctions within Scalesia, Pappobolus, and Simsia and scored accord-

ingly in the remaining taxa (Table 2), resulting in a number of characters being coded as polymorphisms.

The compiled data constitute 63 characters in 78 species (Appendix).

Phylogenetic Analysis —Parsimony analysis was conducted using PAUP* 4.0b10 (Swofford 1998). Heu-

ristic searches were made with character optimization set to both accelerated and delayed transformation

and with the following options: character weighting equal, 10 rounds of random addition sequence with

100 trees held at each addition, branch swapping by tree-bisection, MulTrees in effect, MaxTrees=100,000.

Bootstrap analysis (10,000 replicates) was conducted using accelerated transformation by heuristic search

with 10 trees held at each addition step. A final heuristic search, in which the majority-rule consensus tree

from the bootstrap analysis was input for branch swapping only, was conducted using accelerated transfor-

mation with options as above. Based on the strict consensus tree from the first heuristic search, a reduced

matrix of only the ancestral nodes of the outgroup, Helianthus, Pappobolus, Simsia, and Viguiera grammatoglossa

+ V. stenophylla; the remaining Viguiera species; and the species of Scalesia was generated. This matrix was

subjected to a branch-and-bound search (options: accelerated transformation, equal weighting, MulTrees

in effect, furthest addition sequence) and bootstrap analysis as above.

Tagle 1. Hert ited to suppl ify y y recognized by p y to

Taxon Specimen Locality Herbarium

hiopsi

B. deltoidea A. Gray S. White Mexico: Baja, California. La Paz SMU

B. parishii Greene Mahler & ar USA: Arizona: Maricopa Co: Sagauro Lake SMU

Helianthus

W. L. Henning Acc. No.B802 USA: Missouri: Boone Co.: W of Columbia BRYAN

G. Varga Acc. No. B 1794 USA; Tennessee: Rhea Co, BRYAN

H. tuberosus L. W. L. Henning Acc. No. B 804 USA: Missouri: Boone Co: S of Columbia BRYAN

Pappobolus

P. acutifolius (S.F. Blake) Panero ac 1399 Perd: Ancash: Caráz BRIT

P. matthewsii (Hochr) Panero J. Panero 135. P : Pedro Ruiz BRIT

P robinsonii Panero Pa cM 1225 Perú: Cajamarca: Celendin BRIT

P steubelii (Hieron.) Panero Panero et al. 932 Perú: Cajamarca: Chalhuayaco BRIT

Scalesia

S. affinis Hook.f. Mears 5296 Ecuador: Galápagos: Floreana BRIT

S. helleri B.L. Rob. Mears 5494 Ecuador: Galápagos: Santa Fe BRIT

S. stewartii L. Riley Mears 5556 Ecuador: Galápagos: Bartolomé BRIT

S. villosa A. Stewart Mears 5226 Ecuador: Galápagos: Gardner BRIT

Simsia

S. amplexicaulis (Cav.) Pers. A. Cronquist 9611 Mexico: Michoacán: La Piedad SMU

S. calva A. Gray J. Rodriguez Mexico: Nuevo León: Vallecillo SMU

U. Waterfall 16660 Mexico: Coahuila: Sabinas SMU

S. eurylepsis S.F. Blake U. Waterfall 14300 Mexico: San Luis Potosí: Ciudad de Valles SMU

S. foetida (Cav.) S.F. Blake Yen & Estrada 6479 Mexico: Chihuahua: Presa Chihuahua BRIT

S. fruticulosa (Spreng) S.F.Blake — King € Guevara 5817 Colombia: Cundinamarca. Chipaque MU

S. holwayi S.F. Blake R. M. King 7337a Guatemala: Alta Verapaz: San BRIT

Cristóbal Verapaz

S. sanguinea A. Gray C.G. Pringle 11513 Mexico: Jalisco: Guadalajara SMU

Viguiera ser. Grammatoglossae

V. cordifolia A. Gray J. Cornelius 227 USA: Texas: Brewster Co.: Black Gap WMA SMU

D. S. Correll 15006 USA: Texas: Jeff Davis Co.: Davis Mts. SMU

V. grammatoglossa DC. J. Rzedowski 34497 Mexico: Oaxaca. Chilapa de Díaz VDB

Viguiera sect. Maculatae

V. adenophylla S.F. Blake E. Estrada 1889 Mexico: Nuevo León. Iturbide BRIT

Viguiera ser. Pinnatilobatae

V. stenoloba S.F. Blake A. Krings 288 USA: Texas: Pind Co.: Big Bend BRIT

Ranch State Park

Nee & Diggs 25354 Mexico: Edo. pom Mun. San Pedro BRIT

A. Treverse 2215 USA: Texas: Brewster Co; Big Bend BRIT

National Park

Bayesian analysis was conducted using MrBayes v3.1.2 (Huelsenbeck & Ronquist 2001) on both the full

and the reduced matrices using the default settings of the standard discrete evolutionary model. Analysis

of the full matrix was run for 200,000,000 generations and sampled once every 100,000 generations; the

reduced matrix was run for 400,000 generations and sampled every 100 generations.

Habitat Characterization.—Geographic distributions and habitat features were estimated from Cron-

quist (1971), Eliasson (1976), Hamann and Wium-Andersen (1986), and personal observation of one of us

(RWS)

180

Taste 2. Characters and cl

states used in data matrix (Appendix). Character states are unordered.

1. Habit 0: shrub, 1:tree, 2: suffrutescent perennial, 3: peren-

nial herb, 4: annual

2. Hai 0

to PAN 2: scabrous or strigose

3. Glandular trichomes 0: absent, 1: present

4. hod pubescence color 0: white to gray, 1: yellow or

, 1: villous

> Leaf Phyllotaxy o: aoe 1: peo

is 1:lvs. partially or gradually ERO. 2 pu

much reducing into inflor.

7. lla 0: ovate, 1: lanceolate, 2: cordate, 3: triangular,

4: elliptic, 5: linear-oblon

8. Leaf margin lobing 0: unlobed, 1: lobed 1/4 to midrib,

2: lobed 1/2 to midrib, 3: lobed 3/4 to midrib, 4: regularly

deeply lobed nearly to midrib

9, Leaf margin serration 0: ai era, 1: crenate or

serrulate, indistinctly toothe

10. Leaf margin orientation 0: flat, 1: revolute

11. Leaf adaxially strigose 0: not strigose, scabrous, or

sericeous, 1: moderately strigose, scabrous, or sericeous,

2: densely strigose, scabrous, or sericeous

12. Leaf abaxially strigose 0: not strigose, scabrous, or

co 1: moderately strigose, scabrous, or sericeous,

sely strigose, scabrous, or sericeous

13. Leaf abaxially lanate 0: not villous or lanate, 1: moder-

ately villous or lanate, 2: densely villous or lanat

14. Leaf surface reflectance, adaxially 0: dull, 1: A

15. Leaf texture 0: P ous/chartaceous, 1: leathery, 2:

thinly m

a Leaf venation 0 tripinerved, M PERS

1 a 2: distinctly ICIAL

1: sunken below surface

18. Petiole shape 0: unwinged, 1: wing tapering in apex, 2

wing tapering above 3: wing broad to basal inser-

tion, 4: a at base only

19, Petiole length 0: 0-9 mm, 1: >

20. inflorescence reiteration t monochasal, 1: dichasial

2 litary, 1: open panicle,

2: no aggregate ibus

2.1 large c large 15-30

mm, 2: moderate 7- 157 s z lee

23.1 p en

lat dea

24. Phyllary series 0: 3-4, 1: 2, 2: (4-)5-6

25. Phyllary shape 0: oblong to obtrullate, oo elliptic,

2: lanceolate, 3: o Suena i

idric, 1: urceo-

26. Phyllary, o outer serie t 1:spatulate

27. Phyll florets 0: sul Ji gf , 1: overtop-

pin ng flore

28. Phyllary size ratio, outer/inner 0: outer + inner, 1: outer

inner

29. Phyllary pes A oe green, 1: stramineous + with

green strip , 3: purple

30. Phyllary o scale- like, 1: foliaceous

31. Phyllary pubescence density 0: revealing surface, 1:

obscuring surface

32. Phyllary margin, cilia 0: without cilia, 1: ciliate

33. Phyllary tip shape 0: blunt or abruptly acute, 1: acumi-

nate, long acute

4. Phyllary tip ori

1, 1: reflexed

nyral y up MP

Crapredelid

5. Phyllary thicl 0: untl

base slightly thickened indurate, 2: base conspicuously

thickened indur

36. la jm 1: present in full complement, 2:

present in pa

37. is ae ish 0: spreading, 1: strongly reflexed or

| |, herbaceous, 1:

ved

38. a ligule length 0: «1.5 cm, 1: 1.5-3.0 cm, 2: > 3 cm

39. Ray apex fusion 0: shallowly 2-3 toothed, 1: deeply 2-3

dede 2: Jua. barely fused aa

1: lin

. Ray MP WEG

41. Palea length 0: : about equalling dor 1: a

above phyllaries, 2: shorter than phyllari

42. Palea apex pubescence 0: Pede 1: E

43. Palea segmentation 0: lacking, 1: shallow, 2: deep

44. Palea segments, shape 0: elliptic, 1: triangular, 2: ovate-

rounded, 3: subulate, 4: oblong-ligulate

45. de piel orientation 0: erect, 1: diverging or

xed, 2: strongly Xena no! g, 3: a or iud

t uch

longer than laterals

47. Disk corolla color 0: yellow to orange, 1: brown, 2: pale

yellow, 3: white, 4: pink, 5: deep purple

48. Disk corolla tube to throat length ratio 0: -3-4, 1:

~5—10, 2:

49. m corolla tube pubescence 0: glabrous, 1: puberu-

len

50 raul n I L f gl |

51 Dial n i L Th ri “al

with black pigment, 2: with purple pigm

52. eae yellow, 1: black, 2: galo nine distally,

, 1: puberulent

dark pigment, 1:

53. Anther appendix coloro. stramineous, 1: all or part black,

white

54. ud branch Red i e 0: without black pigment,

1: with black pigm

55. Style branch apex n deltate, 1: attenuate

56. Style branch appendage 0: absent, 1: present

57. Achene length 0: « 3 mm, 1: 3-5 mm, 2: » 5 mm

58. Achene pubescence 0: glabrous, 1: sericeous

59, Achene compression 0: biconvex-lenticular, 1: laterally

flat but e ly bi 2: strongly lat. flattened, 3: terete

or trigon

60. de developmen 0: absent, 1: callous ring only, 2:

ns and/or scales

61. Ba appus, no. awns 0: 0, 1: 1 (often small), 2: 2, 3: multiple

62. Pappus, intervening scales 0: absent, 1: present

63. Pappus persistence 0: persistent, 1: caducous

RESULTS

Sister-group relationships.—The first two heuristic searches (random-addition with accelerated vs. de-

layed character transformations) resulted in 100,000 shortest trees each (442 steps). These and their strict

consensus trees were partially incongruent with the majority-rule tree of the bootstrap analysis. In the

delayed transformation search, Viguiera adenophylla was sister to all other ingroup taxa, and Pappobolus was

paraphyletic with P. ecuadoriensis sister to all remaining taxa. Of these, one clade contained P. sagasteguii, a

subclade of V. stenoloba + Scalesia, and a subclade e the remaining Mrs species, Helianthus, and

Simsia as monophyletic genera. The other clad I ppobolus. The accelerated

transformation search resulted in V. adenophylla as above but the aida o taxa constituted five

clades in an unresolved polytomy: V. cordifolia, Helianthus, Scalesia, Simsia, and one having a monophyletic

Pappobolus sister to V. grammatoglossa + V. stenoloba

The third heuristic search (bootstrap majority-rule tree input and branches swapped) resulted in all

100,000 trees being congruent with the bootstrap analysis, though one step longer (443) than the trees from

the first two searches. In the strict consensus tree of this analysis (Fig. 1), the ingroup formed three major

clades. A monophyletic Pappobolus was sister to the remaining ingroup taxa. Of these, one clade consisted

of V. adenophylla and Scalesia as sister groups. The other clade contained a tetrachotomy: Simsia, Helianthus,

V. cordifolia, and V. grammatoglossa + V. stenoloba.

The Bayesian majority-rule consensus tree (analysis final average standard deviation 0.0078) added yet

another possible arrangement. Of the ingroup taxa, Scalesia + V. adenophylla were sister to the remainder,

which formed a polytomy: V. grammatoglossa, V. stenoloba, nine species of Pappobolus, a clade with all the

remaining Pappobolus, and a clade consisting of V. cordifolia, Helianthus, and Simsia.

In all of the consensus trees, Simsia was completely unresolved or nearly so, and Pappobolus contained

two to three large sets of unresolved species. Scalesia was reasonably well resolved but its topology differed

among trees. All heuristic searches found the arboreous species as a resolved clade (S. cordata A. Stewart,

S. microcephala B.L. Rob., S. peduculata basal), the lobe-leaved species (S. baurii Robinson & Greenman, S.

helleri Robinson, S. incisa Hook.f., S. retroflexa Hemsl.) as a partially or fully resolved clade, the three spe-

cies with elongate phyllaries (S. atractyloides Arn., S. stewartii L. Riley, S. villosa A. Stewart basal) as a grade

or clade, and a clade of S. divisa Andersson + S. gordilloi OJ. Hamann & Wium-And. In two searches the

arboreous clade was sister to the remainder with the elongate-bracted clade deeply imbedded; in the third

the elongate-bracted group was a basal grade with the arboreous clade deeply imbedded. In the Bayesian

majority rule tree, Scalesia was an eight-way polytomy of the arboreous, elongate-bracted, and lobe-leaved

clades, S. affinis Hook.f., S. aspera Andersson, S. crocheri J.T. Howell, S. divisa, and S. gordilloi.

Branch-and-bound analysis of the reduced matrix produced 13 trees of equal length (163 steps). In

the strict consensus tree (Fig. 2), V. adenophylla was sister to the other ingroup taxa, which formed a tetra-

chotomy: Scalesia, Helianthus, Simsia + V. cordifolia, and Pappobolus + the V. grammatoglossa-stenophylla ances-

tor. In Scalesia, the arboreous clade (unresolved) was sister to the remainder which formed a polytomy of S.

affinis, S. aspera, S. crocheri, S. divisa, S. gordilloi, a partially resolved clade of the lobe-leaved species, and a

resolved elongate-bracted clade. However, the Bayesian majority rule tree (analysis final average standard

deviation 0.0070) of the reduced matrix differed by being nearly identical to one of the most parsimonious

branch-and-bound trees (Fig. 2) except that 1) the arl ies formed a basal grade with S. cordata + S.

microcephala sister to the remaining species, 2) there was no PUDE among S. affinis, S. aspera, S. crocheri

and s Masc. pa dno 3) S. retroflexa was basal to the other BOO. of the lobe-leaved clade.

In one ofthe 100,000 dl lly 1 from the third heuristic

Seni the Composite CURRO index (CD was 0. 24 ( (excluding two unif tive ch Dis rescaled CI

(RC) was 0.17, and the retention index (RI) was 0.71. In this tree, the onl ies of S with a con

nd M over 0. y were el florets absent and anthers black. b p. comparable DE for Pap-

[ 1 for Simsia, phyllaries not thickened at laterally flattened

pobolus

The RE Cl of the branch-and-bound trees (Fig. 2) was 0.49 (including only 51 informative char-

182

rir

B deltoidea

B parishii

/ adenophylla

Sc pedunculata

c corda

Sc microcephala

c crockeri

c aspera

c villosa

Sc atractyloides

tewartii

c helleri

c retroflexa

urii

c incisa

cordifolia

| annuus

1 tuberosus

eurylelpis

lagasciformis

calva

sanguinea

setosa

villasenorii

Fic. 1. Strict

` rr » F F [d 4

Generic abbreviatio

ns: B=Bahiopsis, H=Helianthus, P=Pappobolus, Sc=Scalesia, Si=Simsia, V=Viguiera.

Bahiopsis

V adenophylla

V gramm steno

Pappobolus

Helianthus

V cordifolia

Simsia

Sc microcephala

"M ie

100

» Ep]

a (p

Sc cordata

Sc pedunculata

Sc affinis

Sc villosa

Sc atractyloides

Sc stewartii

Doa

[po Sc crockeri

121

Ee Sc aspera

Sc divisa

Sc gordilloi

Sc retroflexa

Sc helleri

Sc baurii

Sc incisa

| 3543] J I

Anek IE

probabilities

183

184 J t tani i Texas 3(

acters), the RI 0.66, and RC 0.37. In this tree, the synapomorphies for Scalesia supported by a consistency

index of 0.4 or greater included: capitula 15-30 cm diameter; involucre hemispheric to urceolate; phyllaries

oblong to obtrullate and erect/appressed; ray absent but, when present deeper within the clade, reflexed

with irregularly fused lobes; paleae deeply segmented; corollas white; anther appendices white; and achenes

glabrous. Black anthers, instead, appeared to be synapomorphic for the ingroup minus V. adenophylla.

The third heuristic search of the full matrix resulted in only six characters that were completely con-

sistent: four por autapomorphies (or e for species pairs), whereas only three involved

gnificant clades. Elev ters had consistency indices of 0.5 or higher.

Bout a iliése a base thickening, ray Bm fusion of ray lobes, and shape of palea segments) were

pcne or reversals within Scalesia; only two (growth habit, orientation of palea segments) were paral-

leli t species of Scalesia and other genera. Forty-six characters had consistency indices lower than

0.5, of which 30 aipsdredd in both Scalesia and other genera, 15 in only other genera, and only one (ratio of

corolla tube to limb lengths) just in Scalesia.

The branch-and-bound matrix had only 55 variant characters. Sixteen were consistent, and, of these,

eleven involved a of significant clades. Twenty-five cl ters were homoplastic with a con-

sistency index of 0.5 or higher including ni ing within Scalesia and six in Scalesia and related genera.

Only 14 characters were below the 0.5 eorsisieney index level with only one restricted to species of Scalesia

(as above), only one Guilde of Scalesia, and the remaining 12 appearing in both Scalesia and other genera.

aphic di and ecology.—All species except Scalesia affinis, which is sympatric with S.

aspera, S. ae - helleri, S. retroflexa, ide eue are narrowly ad or AO (Fig. 3). Some

have disj ions occurring All th (S. pedunculata, S. cordata,

and S. cU a) are found in the uos forest zone in mid to upper E and are geographically

isolated from each other. Scalesia affinis, the only species with consistently radiate capitula, has the widest

distribution and occupies the widest range of habitats; occurring most commonly in the arid zone, it ranges

from coastal to lower parts of the moist forest zone. All remaining species are found in the low elevation

(littoral, arid, and dry forest zones) (Table 2). Due to overlap of preferences, there appears to be little habitat

differentiation among these species. Only the rare species S. crocheri and S. retroflexa are known only from

littoral sites. Other species (e.g., S. atractyloides, Mauchamp et al. 1998) are restricted to cliffs due to grazing

by feral goats but historically ranged over more littoral and arid sites.

DISCUSSION

Phylogenetic relationships.—The present ttl hyly of Scalesia, Simsia, and Helianthus.

Although molecular data (Schillling et al. 1994) ap ait Panpovoliis as monophyletic, the present data are

equivocal in that regard, in some cases placing Scalesia, Simsia, Helianthus, and associated Viguiera species

within a paraphyletic Pappobolus.

Morphological data do not resolve the sister-group relationships of Scalesia, leaving open the possibility

that Scalesia is sister to a group of Viguiera species or that Scalesia arose more or less simultaneously with

Simsia, Helianthus, and Pappobolus (with any associated Viguiera species). As Viguiera sect. Maculatae is basal

among the derived Helianthinae based on DNA restriction site and ITS data (Schilling & Jansen 1989;

Schilling et al. 1994; Schilling & Panero, 1994, 1996, 2002), the sister-group placement of Scalesia and V.

adenophylla in some results is due possibly to experimental error in coding or identification. Furthermore,

Schilling and Panero’s (1996) molecular analysis suggests that Tithonia Desf. ex Gmelin, Viguiera subg.

Amphilepis S.F.Blake, and V. sect. Paradosa S.F Blake are closer to Pappobolus than is Simsia and should be

considered as potential sister groups of Scalesia.

Within Scalesia there is general support for the lobe-leaved, arboreous, elongate-bracted, and divisa-

gordilloi clades. Because the arboreous clade did not receive support from a minority of analyses, it is inter-

esting that Eliasson (1974) considered S. pedunculata to have developed arborescence convergently with S.

cordata and S. microcephala. If all variant trees based on the various analyses preformed are considered, the

only clades receiving total support are the lobe-leaved clade and a terminal clade of S. atractyloides and S.

50 km

Galápagos Islands 50 mi

baurii m Marchena

Genovesa

atractyloides

LE He

90°

Bartolomé |.

stewartii

Faá

Española

Floreana E)

(Santa María)

Fic. 3. Distributi f Scalesi pecies, esti if literat including histori lly | Fine stippli Scalesia affinis, course stippling

lnha.l d rlad hatehi h lad tical Ii l " a sta Gd dart Ai HHA clad Ed Inlet

lobe g , Vertical lines — elon gate , g ; g

gray = remaining species.

stewartii, more in line with Eliasson's conclusions. Within the lobe-leaved clade, there is total support for S.

baurii and S. incisa as a clade, but only partial support for S. helleri + S. retroflexa. In this case, these clades

and all remaining species would radiate from a basal polytomy. If, indeed, Scalesia is an example of radia-

tion by the rapid dispersal of founder populations from an initial colonizer, an unresolved basal polytomy

may portray more accurately the history of Scalesia than any of the less-supported but more-resolved trees.

Homoplasy.—Obviously, the degree of resolution of the particular cladogram examined will affect

the level of homoplasy among taxa. Because the branch-and-bound analysis resulted in 13 equally par-

simonious well-resolved trees and the comparison of all analyses suggest a minimally resolved polytomy

within Scalesia, the level of homoplasy discussed is based on the branch-and-bound consensus tree,

which is intermediate in resolution (Fig. 2). Scalesia itself is delimited by five synapomorphies and 11

homoplastic apomorphies (two among Scalesia species, five with external taxa, and four occurring both

inside and outside Scalesia). Of the 15 species and 6 clades in Scalesia, only 9 are delimited by unique

apomorphies. Of the 90 total character-states apomorphic for clades and species, 14 (16%) are unique, 42

(46%) are homoplastic only within Scalesia, 9 (10%) are homoplastic only between Scalesia and external

taxa, and 25 (28%) are homoplastic between Scalesia taxa as well as with external taxa. It will be of inter-

est to see the level of morphological homoplasy on DNA-sequence trees when these become available.

186 tani i Texas 3(

Tase 3. CI ization of habi f Scalesia species, |f hel

Moist Forest Littoral Arid Dry Forest Volcanic Lava Fissured

e Zone Soil Gravel lava

cordata x x

microcephala — x x x

pedunculata x x x

affinis x X X x x x x

villosa X X x x

atractyloides X X ? X

tewartii X x x x

incisa ? x X

bautii ? x x ? x

retroflexa x x

helleri x x x x

gordilloi x X X X

divisa X X x X

aspera X X x x x

crockeri X x x

The characters that (at least some states of which) are not homoplastic in Scalesia include tree habit,

leaf outline, leaf marginal lobing, dense villous hairs on abaxial leaf and phyllary surfaces, petiole shape,

phyllary shape, ray orientation, palea segment shape and orientation, disk corolla color, disk corolla shape,

anther appendage color, achene pubescence, and pappus development. However, all other characters and

some states of the above are homoplastic. Some interesting examples include: 1) the presence of villous hairs

in S. villosa and Pappobolus; 2) more or less solitary capitula of most species of Scalesia and Viguiera gram-

matoglossa and V. stenoloba; 3) multiple changes in size of capitula in Scalesia; 4) phyllary shape in S. crockeri

and Helianthus; 5) palea length in S. microcephala, the elongate bracted clade, and Pappobolus; 6) glabrous

paleae in S. aspera, S. baurii, S. incisa, S. microcephala, and Pappobolus; 7) length of the central lobe of the

paleae in S. affinis, S. crocheri, S. microcephala, S. villosa; 8) disk corolla tube length in S. affinis, S. baurii, S.

stewartii, the lobe-leaved clade, and Viguiera adenophylla; and 9) glabrous disk corolla tubes in S. affinis, S.

aspera, S. villosa, the arboreous clade, and Simsia. Many of these characters are associated with the palea and

corolla structure. cu to Plovanich and a idi us CODI associated with reproductive

in the Helianth tion pressures. Whether this will be true

in Scalesia remains to be TENT using molecular data BON

In regard to the presence of rays in certain species of Scalesia, Eliasson (1974) concluded that rays were

lost in the ancestor of Scalesia, regained as scattered bilabiate disk corollas in the lobed-leaved species, and

regained as nearly typical rays in S. affinis. His hypothesis is supported by the results presented here. If

the affinis-type rays are the end of a character transformation involving the bilabiate disk corollas or are a

reversal to true rays, then this constitutes an additional homoplastic trait. Presumably rays increase insect

pollination and should be selected for on islands as the insect fauna diversifies, as suggested by the wider

distribution of S. affinis. However, the addition of artificial rays to S. pedunculata did not increase its fitness

(Nielsen et al. 2002). Therefore it is not clear that this character has high adaptive value in Scalesia.

Despite the species and clades of Scalesia being delimited primarily by unique combinations of ho-

moplastic character states as opposed to unique sud shee E species all jaan to be distinct. Moreover,

the full data set suggests that there is a real lack of morpl phy within the

continental genera because many species groups and es are ase defined 2i by unique combina-

tions of homoplastic characters states, not unique states.

Tucteibution i relation to nbl 4 th 11

1 R th 14

islands in the ar-

chipelago are in the southeast and the youngest in the west and northwest, correlation of geology with the cla-

DI Ll le J Dhul " J 224i £Cralaci 187

dograms is not straightforward. Ifthe po diverged from east to west, the basal split should produce a group

of eastern species with the west td E However, the main split is between lowland and

upland species. This may suggest that the lowland species diverged after the older islands from San Cristóbal

west to Santiago were already in puce and the lineage ancestors were able to disperse among islands easily.

Among the upland species, Scalesi hala and S ona (basal in some ie occur on pue youngest

islands. Presumeably, Scalesi 1 lata was already distinct and dispersed on t d founded

populations on the new ean that later formed Isabela to originate the two other arboreous species.

Species of Scalesia are characterized by nearly allopatric distributions in similar habitats (12 spp. in

arid communities, 3 spp. in upland moist communities) within the archipelago. The only synapomorphy

correlated with the origin of the upland-habitat lineage is the tree growth habit, though four homoplastic

characters also accompany the habitat (loss of leaf adaxial strigosity, moderately sized capitula, glabrous

paleae, and corolla tube glabrous). The development of arborescence in a moist habitat under reduced

competition is easy to understand (see Itow 1995; Hamann 2001), but further study is needed to determine

if the other apomorphies are correlated with reproductive ecology. Eight homoplastic o but no

as are aa ith origin of the li in the lowland habitat (pul

gins, solitary P Blum phyllaries, pe deeply divided into elliptic es

en lacking awns). Un] dditi l factors, such idity, ion content, pollinators, or

dispersers, etc. significantly differentiate among both lowland and upland habitats, the species within these

two elevational zones appear to occupy nearly the same range of habitats. For example, Scalesia villosa is the

only Scalesia species having a dense covering of villous hairs, which presumably functions as a protection

from high solar radiation. But several species occupying such habitats are not villous even though villous

hairs occur in the related genera. Although S. helleri bears pinnatifid leaves as an autapomorphy, the nearly

parapatric and perhaps sister species is distinguished only by two homoplastic apomorphies. Only a single

autapomorphy (fully winged petioles) and three homoplastic traits distinguish S. crockeri; nearly parapatric

with it is S. aspera, which is differentiated by only four | plastic traits. Furthermore, diversification among

the lowland species has resulted in some sharing apomorphies with some or all of the upland species and

vice versa. Scalesia microcephala of mesic forests shares two palea dud states (see above) with S. villosa

but not with its close congeners in the mesic zone. Thus, g adaptation of distinguishing features

of these species may prove to be challenging.

Speciation Patterns.—Because the sister-group to Scalesia remains obscure, comparison of speciation

amounts among clades is not possible. It is apparent that this situation will not change until multiple con-

gruent lines of molecular evidence resolve the relationships of the infrageneric groups of Viguiera and other

genera in the derived Helianthi However, if a DNA sequence in which there is variation among species of

Scalesia can be found and analyzed, then, at least speciation rates within Scalesia should be ipai d

Sampling recommendations.— Given the above situation, it is clear that sampling for f

] hould include, in addition to the present taxa, at least e. of Tithonia; Viguiera sube. AD

sect Macias, and sect. Parad d other seg g of the derived Helianthinae. When congruent

lines of molecular evidence point t to one of these lineages as sister to a a up ene of species

should be attempted t to determine whether the wl g E ter to Scalesia.

CONCLUSION

The present study provides a large morphological data set for comparison with molecular phylogenies of

Scalesia and close relatives when the molecular data become available. The results confirm that additional

taxa and DNA sequences must be sampled to resolve the interg d internal relationships of Scalesia.

Furthermore, divergence of Scalesia from its origin to terminal speciations is characterized by combinations

of homoplastic apomorphies. Likewise divergence and EDS geo graphy e to Me poorly correlated.

The seeming uniformity within habitat zones, though, appear: plasy associated

1 1 . 1 Lc

with divergence in Scalesia. Determining the degree to "iud } logical a Í

are adaptive should clarify the process of speciation in this and other mana ae

{23} K={24}

{14} 1={15} J

{12} G={13} H

={03} D={04} E-(05) F=

123456789012345678901234567890123456789012345678901234567890123

00100104001100010010011000000000001210211A24N030110120011000007?

f(111 RNA O

Se een

APPENDIX

(2345) Y2(01234].

024) T=(1 24} U={234} V={01 23} X

{023}S

{35} N={01 2} P={013} Q={015} R

Data matrix. Missing, unknown, or inapplicable=?

Sc_helleri

Sc atractyloides

Taxa

Egl

20001100F01100100001F2002001B001100100A1014A10101102A0011FA22200

Si_fruticulosa

Si_santoros.

1000004A10F00200AB0011A0100A101A101EA102A0011FA2NB0O0

U01011CA1000000004A1F2002001C0011001004A101A1010A102A0011FA2000?

Silagasciformis 48 1011CYF0OAA00000411FK0A2001C001100100A101AG010A10220011FA2BB00

i i 00 ?? ? |

villasenorii

sanguinea

. eurylelpis

Si chaseae

1:2 1:0 2:0:0:010 001 8.970 1:0:09A.1-0-1-& 1:0 41 2:91 92 500 11 2 A9 $2 X8

100000411

1 CNFO 1

mbeyana

Si. do

4B101

— —

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1000000A1022A1

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1390.9 1.0 0-9. 1.0:1.0-0:1. 10-090 OO OO OO 22:01

001200B0001000002

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00010101010001 0AC0001A02211

P_hypargyreus

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00041 1

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3B101

H_tuberosus

190 Journal of ti tanical Texas 3(

ACKNOWLEDGEMENTS

We thank Todd Wood for encouragement and comments and Stephanie Mace for assistance with graph-

ics. The herbaria BRIT, SMU, and VDB are thanked for the loan and use of specimens as is Amanda Neill,

Botanical Research Institute of Texas, for expediting those loans to Bryan College.

REFERENCES

+ l cc / : a (Act

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CanLOussT, S. 1982. Wood and bark anatomy of Scalesia (Asteraceae). Aliso 10:301-312.

Craweoro, D., T.F. Stuessy, M.B. Cosner, D.W. Haines, M. Siva, AND M. BAEZA. 1992. Evolution of the genus Dendroseris

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Euasson, U. 1974. Studies in Galápagos plants XIV. The genus Scalesia Arn. Opera Bot. 36:1-117.

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Sci. 49:17-30.

Kitayama, K. AND S. row. 1999. Aboveground biomass and soil nutrient pools of a Scalesia pedunculata montane

forest on Santa Cruz, Galápagos. Ecol. Res. (Tokyo). 14:405-408.

MAUCHAMP, A, |. ALpAz, E. Ortiz, AND H. VALDEBENITO. 1998. Threatened species, a re-evaluation of the status of eight

endemic plants of the Galápagos. Biodivers. & Conservation 7:97-107.

McMutten, C.K. AND S.J. NARANJO. 1994. Pollination of Scalesia baurii ssp. hopkinsii (Asteraceae) on Pinta Island. Not.

Galápagos 53:25-28.

NueLsen, LR. 2004. Molecular differentiation within and among island populations of the endemic plant Scalesia

affinis (Asteraceae) from the Galápagos Islands. Heredity. 93:434-442.

Nietsen, L.R., R.S. Cowan, H.R. SiecismunD, H. ADSERSEN, M. PHILIPP, AND M. i Fay. 2003. T el and pete

microsatellite variation in populations of Scalesia divisa and S. incisa (Asteraceae) from t pag

Bot. J. Linn. Soc. 143:243-254.

Nievsen, LR, M. PHILIPP, AND H.R. SleGismUND. 2002. Selective advantage of ray florets in Scalesia affinis and S. pedun-

culata (Asteraceae), two endemic species from the Galápagos. Evol. Ecol. 16:139-153,

NieLsen, LR, H.R. SIEGISMUND, AND T. HANSEN. 2007. Inbreeding depression in the partially self-i |

plant species Scalesia affinis (Asteraceae) from Galápagos Islands. Evol. Ecol. 21:1-12.

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Panero, J.L. 2007. Compositae: Tribe Heliantheae. In: JW. Kadereit and C. Jeffrey, eds. Families and genera of vascular

plants, vol. VIII, Flowering plants, Eudicots, Asterales. Springer-Verlag, Berlin, Heidelberg. Pp. 440-477.

Panero, J.L. ano E.E. ScHILLING. 1988. Revision of Viguiera sect. Maculatae (Asteraceae: Heliantheae). Syst. Bot.

13:371-400.

Petersen, G., O. SEBERG, AND T. JORGENSEN. Unpubl. OO ce a plana Paice. National Centers for Biotechnology

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PLOVANICH, A.E. AND J.L. Panero. 2004. A phylogeny of the ITS and ETS for Montanoa (Asteraceae: Heliantheae).

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SANDERS, R.W., T.F. Stuessy, C. MARTICORENA, AND M. SitvA. 1987. Phytogeography and evolution of Dendroseris and

Robinsonia, tree-Compositae of the Juan Fernandez Islands. Opera Bot. 92:195-215.

SANG, T., D.J. CRAwFORD, T.F. STUESSY, AND M. SiL vA. 1995. ITS sequences and the phylogeny of the genus Robinsonia

(Asteraceae). Syst. Bot. 20:55-64.

ScHILLING, E.E. 1990. Taxonomic revision of Viguiera subg. Bahiopsis. Madroño. 37:149-170.

SCHILUNG, E.E. 2006. Helianthus. In: Flora of North America Editorial Committee, eds. Flora of North America north

of Mexico. Oxford University Press, New York and Oxford. 21:141.

SCHILLING, E.E. AND R.K, JANSEN. 1989. Restriction fragment analysis of chloroplast DNA and the systematics of Viguiera

and related genera (Asteraceae: Helianthinae). Amer. J. Bot. 76:1769-1778.

SCHILLING, E.E. AND J.L. Panero. 1996, Phylogenetic reticulation in subtribe Helianthinae. Amer. J. Bot. 83:939-948.

SCHILLING, E.E. AND J.L. Panero. 2002. A revised classification of subtribe Helianthinae (Asteraceae: Heliantheae). |.

Basal lineages. Bot. J. Linn. Soc. 140:65-76.

SCHILLING, E.E., F.B. Da Costa, N.P. Lores, AND PJ. Heise. 2000. Brazilian species of Viguiera (Asteraceae) exhibit low levels

of ITS sequence variation. Edinburgh J. Bot. 57:323-332.

SCHILLING, E.E., J.E. Panero, AND U.H. ELtasson. 1994. Evidence from Chloroplast DNA restriction site analysis on the

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SPOONER, D.M. 1990. Systematics of Simsia (Compositae-Heliantheae). Syst. Bot. Monogr. 30.

SPRING, O, N. He AND U. Eliasson. 1999. Chemosystematic studies on the genus Scalesia (Asteraceae). Biochem.

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SWOFFORD, D.L. 1998. PAUP*. Phylogenetic lysis using parsi y (“and Other Methods). Version 4.0610. Sinauer

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185-265 ribosomal DNA external transcribed spacer. Amer. J. Bot. 94:1837-1852.

L H Ip hl botit, ET

| || £ 4

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192

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Chamberlain, ane Research Institute of Texas, Fort Worth, Texas 76102- 4025, US. A.

J. Bot. Res. Inst. Texas 3(1): 192. 2009

INTRODUCED LESSER CELANDINE (RANUNCULUS FICARIA,

RANUNCULACEAE) AND ITS PUTATIVE SUBSPECIES

IN THE UNITED STATES: A MORPHOMETRIC ANALYSIS

Angela R. Post' Alexander Krings

go of Horticultural Science Herbarium, Department of de ui

North Carolina State University North Carolina State University

Raleigh, men Carolina 27695-7609, U.S.A. Raleigh, North Carolina rosca 12, U.S.A.

Alexander Kringsncsu.edu

Wade A. Wall Joseph C. Neal

ie of Plant Biology Eo of Horticultural Science

North Carolina State University North Carolina State University

Raleigh, M. Carolina 27695-7612, U.S.A. Raleigh, o Carolina 27695-7609, U.S.A.

ABSTRACT

Ranunculus ficaria is native to Europe, but was introduced to the United States by at least the 19* century as a garden ornamental.

Followi ing) x een the m epea un aber Because, in contrast to European floristic treatments, previous North

Ámerican I B , Our O for this ÓN to men if and nii

1 1 E) 11 i P 2i P Ls R a z n T IT n

ds e ES e J I

to th b if foll 1 in Europe, and lyze the distributi habitats d rat f d of each entity. To meet our

objectives, we conducted a Borphameue pn based on id of 319 specimiens from forty-seven herbaria, The ecompined dina

= I Y e

diverged T ld limited f overlap of operational ic units (OTUs) | Principal aoi

nates Analysis and Principal Components AROS results, llas i pletel 1OTU i lust d cl

tree analyses. Based on our current all fi b i in the United States. They are best adapted to moist sites,

y y P

exhibit overlapping distributions, and appear to be spreading at DUM rates.

RESUMEN

Ranunculus ficaria es nativa de re pero fue introducida en los Estados Unidos como ornamental en el siglo XIX. Después de la

ultivo. Forau en ee con los tratamientos floristicos europeos, los norteamericanos no

b if stros objetivos en este estudio fueron determinar si existen y cuantas enti-

la esneci

introducción, p p

imient

habían enfatizado en el recon

dades O E de R. caña se dan en los E Unidos, evaluar en que medida

Europa E analizar las distribuciones, hábitats, y tasas de ode de cada a Para

1 n epto P ME O

lograr t bjetivos, realizamos un análi basado en E estudio de 19 esp ta y siete herbarios.

binados indi ] ia de ci idad l I i ] Europa. Si

E bespeci lí diverg i pl dria est una aa ae de esolapamiemto en m Aa

t 5 OTUSs) n Análisis de C J

r I Eu

OTU l denad álisis de cl y árboles de clasificació "Pene en nuestro

conocimiento [ ] i 1 peci án en los Estados Unidos. Están j 1 I d l g hú muestran distri

d pa | 1 a AN 4

buciones que se Solar yI 1 j

INTRODUCTION

Ranunculus ficaria L. (Ranunculaceae) is native to Europe (Tutin 1964; Taylor & Markham 1978; Sell 1994;

Whittemore 1997), but was introduced to the United States (U.S.) through the garden ornamental trade for

its showy flowers (Bailey 1935). It was collected with certainty in the U.S. in 1867 (Philadelphia County,

Pennsylvania, Burke s.n., PH) and required 141 years to “spread” from Philadelphia, Pennsylvania to Fort

Worth, Texas (Nesom FW08-1, BRIT, MO, NCSC, NCU, TEX)—the southernmost extent of the current

known distribution (Nesom 2008).

"Present address: Der f Plant Biology, Cornell University, Ithaca, NY 14853, arp232@cornell.edu

J. Bot. Res. Inst. Texas 3(1): 193 — 209. 2009

194 Journal of tl tanical Insti Texas 3(

In Europe, five subspecies of Ranunculus ficaria are recognized (Tutin 1964; Sell 1994), whose “ecol-

ogy and [...] distribution appear to overlap, but tend to be different" (Sell 1994). Flora Europaea currently

recognizes: (1) Ranunculus ficaria subsp. ficaria from western Europe, eastward to southern Italy, (2) subsp.

bulbilifer Lambinon from northern and central Europe, extending to Spain, Albania and east-central Russia,

(3) subsp. calthifolius (Reichenb.) Arcangeli from south-central and eastern Europe, (4) subsp. chrysocephalus

PD. Sell from Greece and Crete, and (5) subsp. ficariiformis (FW. Schwartz) Rouy & Fouc from southern

Europe. Plants are known to be diploid (2n=16), triploid (2n=24), or tetraploid (2n=32). Diploids have been

referred to subspecies calthifolius and ficaria, and tetraploids to bulbilifer, chrysocephalus, and ficariiformis

(Greilhuber 1974; Sell 1994). Triploids may represent putative hybrids and have apparently been collected

from widely separated localities in Europe (Marchant & Brighton 1974; Sell 1994). Sell (1994) suggested

that a large proportion of the pollen of triploids, as well as the tetraploid subsp. bulbilifer, is non-viable and

few t. Pollen from diploids and the large-flowered tetraploids (i.e., chrysocephalus and ficariiformis)

is apparently viable and many achenes are produced (Sell 1994). Two subspecies are capable of producing

bulbils in their leaf axils: bulbilifer and ficariiformis. Subspecies bulbilifer tends to exhibit globose bulbils and

subsp. ficariiformis produces ellipsoid bulbils (Sell 1994.

In contrast to European accounts, North American floristic treatments either recognized no subspe-

cific taxa in R. ficaria (Fernald 1950; Gleason 1952; Gleason & Cronquist 1963; Whittemore 1997) or only

variety bulbifera Marsden-Jones (=subsp. bulbilifer Lambinon; Magee & Ahles 1999). However, the recent

P of entities poop referable to subsp. ficariiformis in North Carolina (Krings et al. 2005) and

Texas (I 2008; bsp. bulbilifer, but with ellipsoid bulbils and flower dimensions within the

range of ficariiformis) caused us to question whether additional subspecies may be present in North America

that have not been previously recorded and if so, whether these differed in their distributions, habitats, and

rates of spread. Because prior North American treatments did not emphasize subspecific recognition and

as subspecies are not uniformly accepted, our objectives were to: (1) determine if and how many morpho-

logically recognizable entities within R. ficaria occur in the United States, (2) evaluate to what extent such

entities correspond to the subspecific concepts followed in Europe (based on the work of Sell 1994), and

(3) analyze the distributions, habitats, and rates of spread of each entity.

METHODS

Distribution and habitat information led from 319 herbari j , requested from the follow-

ing forty-seven herbaria based on previous literature reports (Benson 1942; Bell 1945; Gleason & Cronquist

1991; Whittemore 1997): A, AUA, BALT, BH, BKL, BRIT, CONN, CU, DOV, F, FLAS, GA, GH, HNH, ILLS,

KE, LGO, LSU, MARY, MASS, MICH, MISS, MO, MOR, MSC, MT, MU, NA, NCSC, NCU, NHA, NY, OS,

OSC, PH, POM, TENN, TEX, UNA, US, USF, USCH, VDB, VPI, WTU, WVA, Y (Appendix A). Herbarium

label data recorded for each specimen included collector name, collector number, date, habitat, and county

and state of collection. Specimens Hone cando were excluded from the study.

Collection localities were classified into g nine habitat classes: (1) adjacent to a water source,

(2) disturbed areas, (3) dry woods, (4) fields, (5) Bostieuib ss (6) lawns, (7) lowlands, (8) moist areas, and

(9) roadsides. A specimen was classified as adjacent to a water source if it was collected along the banks ofa

river, stream, or pond. Moist areas were defined as moist or alluvial woods, swamp or bog areas, and other

moist shade. The horticultural class was defined as being collected in a nursery or garden under cultivation.

Lowlands were defined as low or depressed areas where moisture level was not mentioned on the label.

Fields were defined as any open grassy area not maintained as a lawn and in full sun such as pastures and

meadows.

To determine if and how many morphologically recognizable entities within R. ficaria occur in the

United States, each herbarium sheet was treated as an operational taxonomic unit (OTU) for data capture

and pertinent subsequent analyses. To evaluate to what extent such entities correspond to the subspecific

concepts followed in Europe, each specimen was determined to subspecies following the key constructed

by Sell (1994):

Post et al., A pl tri lysis of R lus ficari 195

1. Leaf blades to 8 x 9 cm; petioles io 28 cm; MEMES to ne mm diam; achenes 5.0 x 3.5 mm.

2. Stems rather robust, but st bulbil eaf axils after flowering subsp. ficariiformis

2. Stems robust and erect; without bulbils i in leaf axils P flowering subsp. chrysocephalus

1. Leaf blades to4 x : cm; petioles to 15 cm; flowers to 40 mm diam; achenes to 3.5 x 2.2 mm.

. Leaves | the base with few on short stems subsp. calthifolius

3. Leaves less crowded at base and more numerous an the ae stems.

4. Buibils not present in leaf axils atte) fl g; V veloped subsp. ficaria

4. Bulbils present in leaf axils after f ing; acl p rly d developed subsp. bulbilifer

Sell (1994) recognized the difficulty in identifying Ranunculus ficaria to the subspecific level, noting that

specimens should be examined throughout the growing season for positive identifications. We agree with

him and others (Whittemore 1997; Nesom 2008) that identification can be challenging and recognize the

impact identifications have on analysis results. However, we feel reasonably confident in our subspecies

assignments due in part to the quality of specimens, which facilitated taxon assignment, as well as our own

field observations and phenological analyses. Of the 319 herbarium specimens examined 232 had at least a

month and year date on the label and 9096 of these were collected mid-April through June. Another 596 were

collected in the last week of March and the remaining 596 were collected in January through mid-March.

Based on date of collection, the majority of specimens examined in this study were collected late enough in

the spring that they would exhibit bulbils if uA were i nad capable of producing them.

Using a digital caliper, the following mort were taken from each OTU: (1) leaf

length from up to ten leaves, Q) leaf width from s to ten leaves, (3) petiole length from up to ten leaves

(using same leaves measured for length and width), (4) petal length from up to ten flowers, (5) petal width

from up to ten flowers, (6) achene length of all achenes present, (7) achene width for all achenes present.

The presence or absence of bulbils was also recorded.

Quantitative and qualitative data were studied jointly and separately. Statistic analyses, including

ANOVAs and post-hoc tests (Tukey's HSD), were carried out in the statistics package R (Ihaka & Gentleman

1996; R Foundation for Statistical Computing 2008). Prior to multivariate analysis, we tested all quantita-

tive univariate variables using the Shapiro-Willks normality test and subsequently log, transformed them

to minimize the influence of allometry on the results (Dufréne et al. 1991; Almeida-Pinheiro de Carvalho

et al. 2004; Pimentel et al. 2007). Gower's dissimilarity coefficient for mixed data was used to quantify

resemblances between OTUs (Gower 1971). The relationships between OTUs were subsequently explored

with both hierarchical agglomerative cluster analyses and principal coordinates analyses (PCoA) using the

complete set of characters. Three different sorting algorithms were used to help distinguish between data-

dependent and potential OR Rd MEN ct in results, following Dickinson & Phipps (1985) and

Pimentel et al. (2007): single linkage, g „and average linkage (UPGMA; Sneath & Sokal 1973).

Quantitative characters were also MA separately using Principal Components Analysis (PCA).

PCA is an objective, correlation-based technique that allows the variance in quantitative characters to be

considered simultaneously and the subsequent visualization of dispersion patterns in a number of dimen-

sions that explain the greatest amount of variance (Sargent et al. 2004; Joly & Bruneau 2007; Pimentel et

al. 2007). A Kaiser-Meyer-Olkin (KMO) test was performed prior to the PCA to assess the suitability of the

data for multivariate analysis (see also Almeida-Pinheiro B ee et a dil Pimentel et al. 2007).

Classification trees were acci to help identify sy I g hat could reliably

separate the OTU group o the fi tati ] j Sell (1994). Classification trees

divide datasets vit pre- assigned aie membership into Weee homogeneous subsets in tree-like

fashion based on the included hological traits. Recovery proceeds until ps obtained are pure or

until a dividing threshold is ed (Joly & Bruneau 2007). For the n tree, all morphological

characters were included and quantitative data was not log transformed prior to analysis.

The distribution of R. ficaria was mapped based on herbarium specimens and using ArcGIS 9.2 (ESRI

2004). The rate of spread for each subspecies was determined based on the number of counties each was

present in during each decade from 1860 to the present. These data were analyzed using proc mixed in SAS

196 Journal of ical Institute of Texas 3(

9.1.3 with a critical value of 0.05 (SAS Institute 2002-2005). Note that we use “spread” in a broad sense, as

we suspect that not all new county records are the result of physical movement of propagules from estab-

lished parental plants, but that many may reflect novel introduction events.

RESULTS

Descriptive statistics

Table 1 provides non-transformed means and standard deviations for

sured for each group of OTUs assignable to one of five putative subspecies sensu Sell (1994) and summarizes

the results of the one-way ANOVAs and subsequent post-hoc tests (Tukey's HSD) on log,, transformed data

(see superscripts). Figure 1 exhibits box-plots showing the distribution of non-transformed quantitative

measurements taken by OTU group. Significant differences in the means for each of the five groups of OTUs

assigned to the subspecies sensu Sell (1994) for all seven characters were found (Table 1). Post-hoc tests

(Tukey's HSD) to determine which sample means differed from which others showed that means of leaf

length and petal width of the OTU group assignable to subsp. chrysocephalus differed significantly (p«0.05)

from the respective means of the groups of OTUs assignable to the four other subspecies (Table 1). OTUs

assignable to the diploid subsp. calthifolius and ficaria differed significantly (p«0.05) in mean leaf length,

mean leaf width, and mean petiole length (Table 1). They did not differ significantly in mean petal length,

mean petal width, mean achene length, or mean achene width. OTUS assignable to the tetraploid subsp.

bulbilifer, chrysocephalus, and ficariiformis differed significantly (p«0.05) from one another in mean leaf length

and mean petal width. Subspecies bulbilifer differed significantly (p«0.05) from both subsp. chrysocephalus

and ficariiformis in mean leaf width, mean petiole length, and mean petal length. Subspecies chrysocephalus

differed significantly (p<0.05) from subsp. bulbilifer in mean achene length and width, but subsp. ficariiformis

differed neither from subsp. chrysocephalus nor bulbilifer in these characters (Table 1).

1 titati 1 f

Cluster analyses

In all three cluster analyses—average, complete, and single linkage—OTUs were resolved into two large

divisions, these corresponding to (1) the bulbil bearing taxa: subsp. bulbilifer and subsp. ficariiformis sensu

Sell (1994) and, (2) the non-bulbil bearing taxa: subsp. calthifolius, subsp. chrysocephalus, and subsp. ficaria

sensu Sell (1994; Fig. 2). Within these two divisions, the topologies resulting from the three different

algorithms differed notably only for those resulting from single linkage. Within the bulbilifer/ficariiformis

division, average linkage recovered a cluster predominantly composed of OTUs referable to subsp. ficarii-

formis sister to a larger cluster of OTUs predominantly e to denn bulbilifer Both recovered clusters

contained OTUs referable to either subspecies. Th 1 three clusters within

the bulbilifer/ficariiformis division—one of OTUs predominan ble to ficariiformis, nested within two

composed predominantly of OTUs referable to subsp. bulbilifer. Single linkage similarly recovered a cluster

of OTUs referable to subsp. ficariiformis nested within OTUs referable to subsp. bulbilifer. In the calthifolius/

chrysocephalus/ficaria division, both average and complete linkage analyses recovered a cluster of OTUs

predominantly referable to subsp. ficaría nested within clusters of OTUs predominantly referable to subsp.

calthifolius. OTUs referable to subsp. ficaria did not emerge in a distinct cluster in the single linkage analy-

sis, but rather were interspersed throughout those referable to subsp. calthifolius. OTUs referable to subsp.

chrysocephalus emerged interspersed in grades of OTUs referable to subsp. calthifolius and subsp. ficaria in a

sister position to the rest of the division in all three analyses.

PCoA and PCA

Consistent with cluster analysis results, two non-overlapping clusters of OTUs were recovered in the PCoA

corresponding to (1) the bulbil bearing taxa: subsp. bulbilifer and subsp. ficariiformis sensu Sell (1994) and

Q) the non-bulbil bearing taxa: subsp. calthifolius, subsp. chrysocephalus, and subsp. ficaria sensu Sell (1994;

Fig. 3A). Within both of these clusters, cohesiveness was exhibited by each group of OTUs referable to one

of the five subspecies sensu Sell (1994), although each group overlapped with another to some degree.

Consistent with expectations for infraspecific entities, distinct but overlapping clusters of OTUs were

Post et al., A pl tri lysis of R lus ficari 197

Toate 1.1 No £ J ral Tei | n . i : £ 1 e E.

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V t 310 7 2f

bulbilifer calthifolius chrysocephalus ficaria ficariiformis F p-value

Leaf length 2.18 24 (0.41; 82) 1.93?(0.46; 96) 3.545(1.03;6) | 242? (0.39; 37) 2.97 * (0.49; 12) 9.79 «0.001

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petole ln 10.41 ? (2.79; 82) 6.55*(1.62; 97) 16.21 (1.44;6) 12.198% (2.16; 37) 14.75 %4 (3.85; 12) 2844 <0.001

mo no 10,23 9(1.67; 75) 11.11 #(1.91;97) 17.43 ©4 (3.79; 6) 12.11 © (1.95;37) 13.91% (1.95,11) 16.89 «0.001

(mm)

ous. p 11) 23.62 «0.001

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(mm

Achene length 2.95 ? (0.65; 53) 3.04? (0.65;28) 4.33 £ (0.69;5) 3.27 = (0.41; 6) 3.50 ** (0.90; 7) 530 «0.001

Achene width 4.35 è (0.90; 53) 4.262(1.00;28) 622°°(0.77;5) 458**(1.12;6) 4.70 ** (1.31; 7) 383 «001

DN £11

exhibited in the PCA comprising all OTUs when symbol coded for a priori

the infraspecific concepts of Sell (1994) (Fig. 3B). Among these, OTUs eee to subsp. bulbil ifer exhibited

the most cohesive and least diffuse cluster. In this analysis, 81% of the variation is explained by the first two

axes. PCI is positively correlated most strongly with petiole length, leaf width, and leaf length, whereas PC2

is positively correlated most strongly with petal width and petal length (Table 2).

A separate analysis of only OTUs referred to the two diploid taxa subsp. calthifolius and ficaria, resulted

in two very well-defined clusters with minor overlap (Fig. 3C; Table 3). In this analysis, 73 % of variation is

explained by the first two axes. PC] is positively correlated most strongly with petiole length, leaf length, and

leaf width, whereas PC2 is positively correlated most strongly with petal width and petal length (Table 3).

An analysis of only OTUs referred to the tetraploid taxa—subsp. bulbilifer, chrysocephalus, and ficariiformis—

showed evident clustering, but with greater overlap among the three a priori defined subspecies (Fig. 3D;

Table 4). In this analysis, 76% of the variation is explained by the first two axes. OTUs defined a priori as

subsp. ficariiformis occupied a central coordinate space in the tetraploid analysis, flanked along the primary

axis by subsp. chrysocephalus to the left and bulbilifer to the right. PC1 is negatively correlated most strongly

with petal width, leaf length, and leaf width, whereas PC2 is positively correlated most strongly with petiole

length (Table 4).

Among the five subspecies of is ee eee i Sell (1994), only the tetraploid subsp. bulbilifer and

ficariiformis are known to prod A y f only OTUs with bulbils, showed two rather

cohesive clusters with imed overlap on to a priori assignment to these two subspecies sensu

Sell (1994; Fig. 3E; Table 5). In this analysis, 69% of the variation is explained by the first two axes. PC1

is negatively correlated most strongly with petal width, whereas PC2 is positively correlated most strongly

with petal width and petal length (Table 5). A separate analysis of OTUs without bulbils (Fig. 3f; Table 6),

showed three rather cohesive clusters with limited overlap, corresponding to a priori assigned subspecies.

In this analysis, 7996 of the variation is explained by the first two axes. PCL is positively correlated most

strongly with petiole length, leaf width, and leaf length, whereas PC2 is positively correlated most strongly

with petal width and petal length (Table 6).

Classification tree

Classification tree analysis showed that 9596 or greater of the OTUs we aa to subsp. calthifolius, subsp.

ficaria, and subsp. bulbilifer using Sell (1994) could be placed i Į g g groups (Fig. 4)

198

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PCI PC PG

Leaf length 71 0.06007699 -0.5807531

Leaf width 0.4659988 0.01441147 -0.4945180

Petiole length 0.6942403 -0.45147042 0.5562742

Petal length 0.2116039 0.44013398 0.1484652

Petal width 0.2415118 0.77371804 0,2944399

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PCI PQ PG

Leaf length 0.4393161 0.1933816 -0.5527152

Leaf width 0.4473662 0.1682932 -0.4790156

Petiole length 0.7371736 -0.4948622 0.4585902

Petal length 0.1785614 0.4539478 0.2102479

Petal width 0.1776393 0.6952149 0.4588473

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Leaf width -0.4139542 0.2858958 -0.4349598

Petiole length -0.4282971 0.5777908 0.6924294

Petal length -0.3705260 -0.3867720 0.1113257

Petal width -0.5797091 -0.5918072 0.1221624

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PC1 PQ PG

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Leaf width -0.4395120 -0.2045104 0.4630089

Petiole length -0.4585246 0.6345035 -0.6193181

Petal length -0,3303427 0.3747980 -0.1413127

Petal width -0.5602160 0.6109399 -0.2063936

Taste 6. C] | (PC) in the PCA of th bulbiliferous OTUs.

PC1 PQ PG

Leaf length 0.4474063 0.1388268 -0.5661383

Leaf width 0.452295] 1139208 -0.4887299

Petiole length 0.6954902 -0.5497825 0.4608629

Petal length 0.2280402 0.4510623 0.2216459

Petal width 0.2440242 0.6797288 0.4232127

Seventy-five percent (N26) and 87.5% (N=7) of OTUs we referred respectively to subsp. chrysocephalus and

subsp. ficariiformis using Sell (1994) could be placed into corresponding homogeneous groups. Á quarter

of the OTUs (N22) assigned to subsp. chrysocephalus in the analysis were specimens we referred to subsp.

ficaria using Sell (1994). The presence of bulbils separated the calthifolius/chrysocephalus/ficaria group from

202 Jour nal of t tani f Texas 3(

Bulbils Present

No Yes

Petiole Petiole Petal Length | Petal Length

29.9 cm «13.5 mm

Petal Width | Petal Width

>6.9 m

«6.9 mm

calthifolius bulbilifer ficariiformis

73 (94.8%) bulbilifer 7 (87.5%) ficariiformis

PHP A QE 1 (1.3%) calthifolius 1 (12.5%) bulbilifer

> 3 (3.9%) ficariiformis

ficaria chrysocephalus

31 (96.9%) ficaria 6 (75%) chrysocephalus

1 (3.1%) calthifolius 2 (25%) ficaria

is 4. Classification t ly is. Numi gi t node t inals indicat iori classificati Sell (1994) ). For example, the model as-

cial to ñ Icaria by us.

the dl ied Within the ud iole l liscrimi ] best between OTUs referable

to subsp d those referable t onda and EM (oa Petal width discriminated

best between euism ficaria a subsp. ola Petal length d best bet bsp. bulbilifer

and subsp. ficariiformis. The shape of bulbils was not scored for the classification tree analysis as we were

interested in seeing what additional vegetative character distinguished these putative taxa.

DISCUSSION

Subspecies recognition.—The combined results indicate the presence of five entities that can be reason-

ably referred to the subspecies accepted by Sell (1994). If one accepts subspecies as incompletely diverged

lineages, one would expect a limited amount of overlap of OTUs as seen in our PCoA and PCA results, as

well as incompletely sorted OTUs as seen in our cluster and classification tree analyses (Rosen et al. 2007).

The ANOVA results are also informative on this issue, particularly because the assignment of each OTU to

a putative subspecies was based exclusively on the key by Sell (1994; see above). In this key, quantitative

measurements were used only to distinguish two groups of subspecies (i.e., chrysocephalus/ficariiformis and

bulbilifer/calthifolius/ficaria). Qualitative characters are used in Sell's (1994) key to distinguish individual

subspecies within these two groups. Thus, contributing evidence of the morphological cohesiveness of the

subspecies concept of Sell (1994) is the extent to which differences in quantitative characters are found be-

tween all subspecies pairs. Of course, had we found that our OTU groups assigned to the subspecies sensu

Sell (1994) did not differ significantly in quantitative characters, it would not necessarily have challenged

Sell's concepts, as the taxa may truly differ only in qualitative characters. However, the finding that the

OTU groups corresponding to the subspecies sensu Sell (1994) do in fact differ in various combinations of

the quantitative characters we examined provides some additional evidence of distinctness. Although the

groups differed primarily in the means of quantitative characters and showed overlap in maximum dimen-

sions, if one accepts a subspecies as an incompletely diverged lineage, overlap in character states cannot be

Post et al., A pt tri lysi fR Jus ficari 203

unexpected. In addition, distinct means in quantitative characters could be viewed as a reflection of partial

isolation and potentially emerging distinct evolutionary trajectories, possibly leading to speciation. In rec-

ognizing that our analysis is limited to plants introduced to the United States, our results could be biased

if our data sets largely contained “non-controversial” individuals (e.g., individuals from subspecific centers

of distribution in Europe, rather than regions of overlap). There is no way to know this, except through

a broader study. However, we did not explicitly seek to test the subspecies concept sensu Sell (1994), but

rather whether plants introduced to the United States could be reasonably referred to that concept—which

we believe they can. The recognition of subspecies of R. ficaria has obvious practical consequences in weed

management, as not all subspecies may behave in the same manner.

Summary of taxon distribution, habitar, and rate eof spread.—Based on our current understanding,

the subspecies of R. ficaria exhibit in the United States (Fig. 5). Subspecies calthifo-

lius occurs in a states and the District of Columbia. It was apparently first collected in the United

States in 1867 (Pennsylvania: Burke s.n., PH). Collections of this subspecies account for 35.5% of specimens

examined. Subspecies bulbilifer currently occurs in sixteen states and the District of Columbia. It was appar-

ently first collected in the United States in 1891 (New York: Hollick s.n, LGO) and accounts for 31.5% of the

specimens examined. Subspecies ficaria occurs in ten states and the District of Columbia. It was apparently

first collected in the United States in 1876 (New York: Schrenck s.n., LGO) and accounts for 15% of the col-

lections examined. Subspecies ficariiformis currently occurs in Missouri, North Carolina, New York, Ohio,

Pennsylvania, and Texas. It was aj ly first collected in the United States in 1910 (Pennsylvania: St. John

111, GH) and accounts for only 8% of specimens examined. Subspecies chrysocephalus currently is known

only from Maryland, New York, Oregon, and Washington. It was apparently first collected in the United

States in 1975 (Oregon: Hatch s.n., NY, OSC). The narrower range documented for subsp. chrysocephalus

vis-a-vis the other subspecies is attributed to the recency of introduction—collections of this subspecies in

the United States are unknown prior to 1975. The disjunct populations in the east and west likely resulted

from secondary introductions through the horticultural trade.

All subspecies of Ranunculus ficaria are best adapted to moist sites (Taylor & Markham 1978). All

perform well in irrigated landscapes, such as lawns and horticultural plantings, but occur in a variety of

habitats from moist woods to roadsides and lawns (Fig. 6). The majority of collections of all subspecies

(45.9%) were made adjacent to a water source such as a river, stream, or pond. An additional 15.5% came

from other moist areas. Collections from lawns and horticultural plantings were equal at 8.13% each. A few

specimens have been collected in other habitats such as disturbed sites, lowlands, and fields. Twenty-three

percent of specimens of subsp. ficariiformis were collected from dry woods suggesting that this subspecies

may tolerate more xeric environments than the other four. Habitat distributions of subsp. chrysocephalus

and subsp. ficariiformis likely represent only a limited percent of the habitat range of these taxa due to the

limited number of collections they are based upon (N = 6 and N = 13, respectively).

Vegetative spread occurs through tuberous roots, although subsp. bulbilifer and subsp. ficariiformis also

produce axillary bulbils for reproduction (Taylor & Markham 1978; Sell 1994). All subspecies except subsp.

bulbilifer produce viable seed which frequently fall adjacent to parent plants (Marsden-Jones 1937). Disper-

sal over long distances likely occurs anthropogenically. The subspecies multiply easily along riverbanks,

forming dense mats where there is seasonal flooding (Taylor & Markham 1978). Short distance dispersal is

effected by seasonal flood waters which may transport tubers or bulbils downstream. This dispersal pattern

was confirmed by primary observation on subsp. ficariiformis in Wake Co., North Carolina, in the spring

of 2006. The subspecies was distributed along a drainage ditch, through a culvert under the road, and into

a local waterway where it colonized banks downstream from the source. Persistence in the landscape is

exacerbated by continued use in the nursery trade as a garden plant. Plants may slowly escape from cultiva-

tion and spread when tuberous roots, bulbils, or small plants are discarded in yard waste.

Figure 7 shows the relative rate of spread of each subspecies by the number of counties in which it was

collected by decade. It does not appear that any single subspecies of R. ficaria is more invasive than another

204 Journal

Distribution of R lus ficaria and its five subspecies in the United Stat f2008. O = subsp. bulbilifer; 4 = subsp. calthifolius; + = subsp.

chrysocephalus; * = subsp. ficaria; Y = subsp. ficariiformis.

in the United States. There is no significant difference among the expansion slopes of the five subspecies

during the first forty years after each introduction (p=0.0769), suggesting that each pecies behaves simi

larly, at least in the early phases of expansion. It may be expected, therefore, that more recently introduced

taxa, such as subsp. chrysocephalus, subsp. ficaria, and subsp. ficariiformis, will follow a similar pattern to

that of the earlier introductions, subsp. bulbilifer and subsp. calthifolius, and extend their ranges at similar

exponential growth rates in the next hundred years.

All subspecies of R. ficaria should be expected to persist where introduced throughout most of Canada,

New England to Iowa, possibly northern California, and as far south as Texas. We have not seen collections

from California, Iowa, Indiana, Maine, Rhode Island, or Vermont, although expect that subspecies will

persist there, as well. The Midwestern states of Arizona, Colorado, Idaho, Kansas, North Dakota, Nebraska,

Nevada, Oklahoma, South Dakota, Utah, and Wyoming are likely too arid for widespread naturalization.

However, subspecies may establish locally in irrigated areas or wetlands following introduction.

APPENDIX A

List of exsiccatae of Ranunculus ficaria in the United States. Arranged alphabetically by subspecies. * 2 handwriting difficult

to decipher; d» = double flowers.

Ranunculus ficaria L. subsp. bulbilifer Lambinon

U.S.A. CONNECTICUT. New Haven Co.: 12 May 1992, Mehrhoff 15469 (CONN, Y); 4 Jun 1997, Morehead Il! 3561 (CONN

DELAWARE. New Castle Co.: 24 Apr 2004, Clancy 5905 (DOV); Apr 1978, Lindtner 109 (DOV); Schuyler 7210 (PH). DISTRICT OF

COLUMBIA: 27 Apr 1993, Redman 6651 (BALT). ILLINOIS. Cook Co.: 6 May 1987, Evert 11832 (MOR); 6 May 1987, Evert 11833

(MOR); 17 May 1989, Evert 16279 (MOR, NA); 8 May 1997, Hick 514 (MOR); 26 May 1978, Kamin 945-3000 (MOR); 2 May 1982,

Lace s.n. (MOR); 5 May 1998, Masi, Epting, & ! ich 817 (ILLS); 1 May 1960, Venrick 122 (MO). Dupage Co.: 27 Apr 2003, Kobal

FPD03-02 (MOR); 5 May 1995, Lampa 95-03 (MOR). Lake Co.: 27 May 1998, Fiest 24 (LLS). MARYLAND. Baltimore Co.: 26 Apr

1975, Beach 0107 (MARY); 17 Apr 1984, Hill 13552 (BRIT, GH, MARY, MO, MSC, NY-2 sheets, POM); 22 Mar 1989, Redman 6320

(BALT); 30 Apr 1993, Redman 6492 (BALT); 14 Apr 1974, Romeo 41 (MARY). Howard Co.: 30 Apr 1965, Engh s.n. (BKL, MARY).

—

205

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Montgomery Co.: 25 Apr 1937, Benedict Jr. 3679 (NA); 18 Apr 1976, Morris 1 1 (MARY). Prince Georges Co.: 19 Apr 1974, Bunn 37

(MARY). MASSACHUSETTS. Suffolk Co.: 24 May 2004, Mehrhoff 21169 (CONN). MICHIGAN. Clinton Co.: 9 May 1982, Gereau

966 (MICH, MSC). Eaton Co.: 22 May 1984, Blouch s.n. (MSC); 31 May 1984, Blouch s.n. (MSC). lonia Co.: 21 May 2003, Reznicek &

Kogge 11470 (MICH). MISSOURI. Saint Louis Co.: 21 Apr 1985, Brant 55 1 (MO); 26 Apr 1972, Brown s.n. (MO); 12 Apr 1992, Ochs

19 (MO). NEW HAMPSHIRE. Hillsborough Co.: 13 May 1962, Stiff s.n. (NHA). NEW JERSEY. Essex Co.: 25 Apr 2003, Glenn 8181

(BKL). Hunterdon Co.: 26 Apr 2001, Glenn 5425 (BKL). Mercer Co.: 22 Apr 2002, Glenn 8988 (CONN, BKL). Passaic Co.: 17 Ma

1939, Clausen s.n. (BH); 6 May 1941, Langmuir & Lawrence s.n. (BH). Somerset Co.: 23 Apr 2003, Glenn 8149 (BKL). NEW YORK.

Bronx Co.: 4 May 1964, Bennett s.n. (NY); 28 Apr 1939, Swift R432/37 (NY); 4 May 1996, Walker 1868 (NY); 19 Apr 1989, Yost

355 (DOV). Cayuga Co.: 23 May 1932, Hazard 17883 (CU). Nassau Co.: 27 Apr 2001, Steward 416 (BKL). Richmond Co.: 3 May

1937, Guiler s.n. (CU); 19 Apr 1891, Hollick s.n. (LGO). Tompkins Co.: 10 May 1935, Burnham 18847 (CU); 22 May 1935, Clausen

s.n. (Bm), 22 May ud 7, Gershoy 8104 (CU). OHIO. Franklin Co.: 17 Apr 1977, Carr 68 (OS); 27 Apr 1980, Carr 2645 (OS); 28 Apr

1987, 4 (OS); 16 Apr 1992, Lowden 4908 (OS). Hamilton Co.: 21 Apr 1982, Cusick 21469 (OS); 12 Apr 1989, Cusick

27953 (OS).M Co.: 6 May 1978, Cusick 18074 (KE, OS); 23 Apr 1998, McCormac 6484 (MICH). OREGON. Multnomah

Co.: 26 Apr 1962, Sib (OSC). PENNSYLVANIA. Berks Co.: 5 May 1972, Brumbach 7910 (BH, NA-2 sheets, NY). Blair Co.:

1 May 1987, k 8564 (PH). Bucks Co.: 3 May 1959, Forman s.n. (PH); 18 Apr 1998, Mehrhoff 20039 (CONN). Chester Co.:

18 Apr 1959, Webb & Wherry s.n. (PH). Delaware Co.: 6 Apr 1937, Blaser s.n. (CU); 7 May 1944, Carter 5084 (DOV); Apr 1908,

Painter s.n. (MO, NA); 16 Apr 1942, Wheeler 5600 (POM). LeHigh Co.: 26 Apr 1959, Schaeffer Jr. 58388 (PH). Montgomery Co.:

22 Apr 1922, Dreisbach 868 (F, MICH); 24 Apr 1954, Wherry s.n. (PH). Philadelphia Co.: 26 Apr 1911, Eckfelds s.n. (PH-2 sheets);

1 May 1912, Fackenthall s.n. (NA); 20 Apr 1911, St. John s.n. (GH); 12 Apr 1908, Van Pelt s.n. (MICH, PH). VIRGINIA. Fairfax Co.:

13 Apr 1976, Bradley & Frederickson 9954 (WVA). WASHINGTON. King Co.: 17 Mar 2002, Zika & Jacobson 16885 (WTU). WEST

VIRGINIA. Ritchie Co.: 15 May 1970, Elliott s.n. (WVA). Wood Co.: 8 Apr 2003, Grafton s.n. (WVA).

m a on. | subsp el: iD L hy Arcan nge

U.S.A. CONNECTICUT. Fairfield Co.: s Apr 1985, Mehrhoff 11192 (CONN). New Haven Co.: 20 Apr 2001, Murray 01-001

(CONN). pilin co 29 Apr 2 í 0744 (CONN). DELAWARE. New Castle Co.: 4 Apr 1985, Meyer & Mazzeo 20693

(NA). DISTRICT :20 Mar 1983, Flemming s.n. (MARY); 14 Apr 1983, Fleming s.n. (NA); 3 Apr 1986, Fleming 4 (NA);

13 Apr 1899, qs 72 (NA); de 1874, McCarthy s.n. (NA); 26 Apr 1884, McCarthy s.n. (NA); 18 Apr 1897, Topping s.n. (NA). ILLI-

NOIS. Cook Co.: 2 May 1996, Antonio & Masi 7577 (ILLS); 24 Apr 1966, Argent M. D. s.n. (ILLS). Lake Co.: 24 Apr 1988, Snydacker

lysis of R lus ficari 207

Post et al., A | [

561 (F). KENTUCKY. Campbell Co.: 12 Apr 1981, Buddell I| 108 (NY). MARYLAND. Anne Arundel Co.: 4 Apr 1991, Longbottom

1460 (MARY); 15 Apr 1993, Longbottom 3473 (MARY). Baltimore Co.: 23 Apr 1971, Chanoski 043 (BALT); 3 Apr 1980, Critikos 9

(BALT); 4 Apr 1977, Lears s.n. (MARY); 13 Apr 1974, Ness 33 Mp 1 Mod idi Redman 4029 (BALT). Caroll Co.: 13 Apr 1963,

Burroughs 31 (MARY). Howard Co.: 17 Apr 1965, Stolze 386 (F-2 sheets). A y Co.: 25 Jan 1950, Cross s.n. (NA); 27 Mar

1976, Mora 22 (MARY); 1 ii e Sappington s.n. (MARY); 6 Apr en Schlossberg 0002 (MARY); 19 Mar 1983, Zastrow 1 (OSC).

Prince Georges Co.: 9 Apr 1987, Bowman 377 (MARY); 13 Apr 1970, Thompson Jr. s.n. (MARY). MASSACHUSETTS. Middlesex

Co.: 8 May 1982, Wood 4690 m MICHIGAN. lonia Co.: 23 Apr 1989, Penskar 1085 (MICH); 24 Apr 1989, Penskar 1086 (MICH).

MISSOURI. Saint Louis Co.: 4 Apr 1989, Yatskievych, Yatskievych, iod 89-05 (MO); 1 Apr 1999, a ied

Harris, Harris, & S 99-04 (MO). NEW JERSEY. Burlington Co.: 16 Apr 1932, Stokes M.D. s.n. a :

1898, Saunders s.n. (PH). Middlesex Co.: 12 Apr 2003, P 354 (CONN). Somerset Co.: 7 Apr ian. dd pu

(BKL). Union Co.: 3 Apr 2002, Glenn & Steward 6197 (BKL). NEW YORK. Bronx Co.: 11 Apr 1976, n Ne (BKL). Dutchess

Co.: May 1941, Van Melle s.n. (BH). Queens Co.: Apr 1876, Schrenk s.n. (LGOJ; 1876, Schrenk s.n. (LGO); 25 Apr 1877, Schrenk s.n.

(BKL, MOJ; Apr 1877, Schrenk s.n. (PH); 11 Apr 1878, Schrenk s.n. (MICH); Apr 1878, Schrenk s.n. (BKL, F-2 sheets, HNH, NA, NY-2

sheets, POM); Apr 1878, Schrenk s.n. (LGO-2 sheets, NA); May 1882, Bisky s.n. (BKL). Richmond Co.: 13 Apr 1898, Coheu* s.n.

ini 2 sheets). Tompkins Co.: 7 May 1937, Anderson s.n. (MASS); e 19 Apr 1959, Dress 5984 (BH); 1 May 1996, Dress 19996 (BH).

OHIO. Clark Co.: 31 Mar 1992, Cusick 30056 (OS). Clermont Co.: 2 Apr 1996, Cusick 32886 (MO, OS). Clinton Co.: 3 Apr 1989,

inr 27946 (OS). Franklin Co.: 5 Apr 1986, Cooperband 5 (OS); 29 Mar bu Cusick 26207 (NY). Greene Co.: 31 Mar 1992,

gu es (OS). Lake Co.: 2 Jun 1901, Hacker s.n. (OS). Van Wert Co.: 22 Apr 1946, Brooks 1393 (OS); 28 Apr 1947, Brooks

REGON. Multnomah Co.: 27 Mar 1991, Zika 11064 (OSC). IAN Bucks Co.: 18 Apr 1962, Wherry s.n.

(Ph Delaware Co.: 13 Apr 1934, Fogg Jr. 6316 (PH); 12 Apr 1938, Fogg Jr. 14021 (GH); 7 Apr 1894, MacElwee Jr. s.n. (PH); 12 Apr

1920, Meredith M.D. s.n. (NY); 15 Apr e Schaeffer Jr. 16953 (PH-2 sheets); 19 Apr 1936, Thompson Jr. 17 (PH); Greene Co.: 4

Apr 1953, Buker s.n. (PH). M y Co.: e 19 Apr 1963, Fogg Jr. 22220 (A); 3 Apr 1921, Long 23784 (PH); 20 Apr 1937, Long

49720 (PH); 18 Feb 1954, Long 77721 o 1 May 1985, Weaver s.n. (PH). Philadelphia Co.: 1867, Burke s.n. (PH); e10 Apr

1954, Fogg Jr. 21460 (PH-2 sheets); e 18 Apr 1954, Fogg Jr. 21474 (PH); 3 Apr 1933, Hermann 3953 (NA); 18 Apr 1974, Jers* (PH);

27 Apr 1924, Lang 112 (GH); %1 May 1920, Meredith M.D. (NY); 30 Mar 1909, St. John 110 (GH). TENNESSEE. Knox Co.: 17

Mar 1977, BA (TENN); 25 Mar 1966, Thomas s.n. (BRIT); 31 Mar 1966, Thomas & Rogers s.n. (TENN). VIRGINIA. Albemarle

Co.: 25 Mar 19 25629 (VPI). Chester Co.: 3 Apr 1999, Huber 1 (ILLS, OS). Fairfax Co.: 13 Apr 1947, Sargent s.n. (NCSC);

23 Mar 1974, rd (MARY). WASHINGTON. Whatcom Co.: 5 Apr 1968, Sundquist 1550 (POM). WEST VIRGINIA. Ritchie

Co.: 15 Apr 1963, Stonestreet s.n. (WVA). WISCONSIN. ohh Co.: May 1970, Larkin s.n. (MOR).

R ficaria L. subsp. cl halus PD. Sell

U.S.A. MARYLAND. diss Georges Co.: 27 Mar 1977, Wirick 05 (MARY). NEW YORK. Tompkins e MAR Dress d

(CU). OREGON. Benton Co.: 31 Mar 1991, Zika 11065 (OSC). Lane Co.: 22 Feb 1975, Hatch s.n. (NY,

Co.: 17 May 2002, pois & Jacobson 13733 (WTU).

Ranunculus beris L. subsp. ficaria

» S.A. DISTRICT OF COLUMBIA: e 29 Apr 1896, Pollard 776 (NY); #29 Apr 1896, Pollard 776 (MSC); %29 Apr e Pollard

n. (POM); 417 Apr 1898, Pollard s.n. (NY); 417 Apr 1898, Steele s.n. (MSC); #18 Apr 1897, Steele s.n. (GH); de 18 Apr 1897,

med s.n. (NA); e 18 Apr 1897, Steele s.n. (NY). ILLINOIS. Lake Co.: 24 Apr 1988, Snydacker 560 (MOR). aÑo Baltimore

Co.: 13 Apr 1980, Caruso 22 (BALT); 5 Apr 1980, King 10 (MARY). Howard Co.: 25 Mar 1989, Redman 6321 (BALT). Montgomery

Co.: 19 Apr 1975, Hollenberg 8 pea 19 Apr 1979, Trumball 4 (BRIT); 13 Apr 1975, Yinger 17 (MOR). Prince Georges Co.: 10

Apr 1977, Dochtermann 29 (MARY); 16 Apr 1994, Hedge 13 (MARY); 2 Apr 1982, Kunowsky 10 (MARY); 4 Apr 1964, Weigel Jr. s.n.

ane MASSACHUSETTS. Hampshire Co.: 24 Apr 2002, Mehrhoff 21611 (CONN, MASS). MISSOURI. Saint rs Co.: 1

990, Christ s.n. (MO); NEW JERSEY. Mercer Co.: 18 Apr 2001, Glenn 5386 (BKL). Somerset Co.: 19 Apr a

Ps 5401 (BKL). NEW YORK. Cayuga Co.: 7 Jun 1935, Petry 18846 (CU). Dutchess Co.: 1932, Van ma s.n. (BH).

Queens Co.: 22 Apr 1990, sel s.n. (BKL); 1921, Martin s.n. (PH); Apr 1877, Redfield 10896 (MO); Apr 1876, Schrenk

s.n. (LGO). OHIO. Butler C Apr 1997, Turner 31 (MU). PENNSYLVANIA. Chester Co.: 4922 Apr 1941, Terrell Jr.

375 (PH); %6 May 1954, idus n. (PH). Delaware Co.: 30 Apr 1892, Brinton M. D. (PH-3 sheets); Apr 1903, Conard

s.n. (PH); 5 Apr 1935, Fogg Jr. 7996 (PH); 1 May 1904, Jahn s.n. (PH); 4 Apr 1946, Proctor 1716 (NHA). Northampton

Co.: 2 May 1969, Tucker s.n. (DOV). Philadelphia Co.: %20 Apr 1921, Hur s.n. (PH); 23 Apr 1932, Hermann 2752

(NA); e May 1878, Martindale s.n. (NA, LGO); %12 May 1904, Van Pelt s.n. (PH). WASHINGTON. King Co.: 8 Mar

2000, Zika 6 Jacobson 14827 (WTU). WEST VIRGINIA. Wood Co.: 12 Apr 2003, Grafton s.n. (WVA)

R ficariiformis (FW. Schwartz) Rouy & Fouc

U.S.A. MISSOURI. Sui Du Co.: 19 May 1994, Ladd 18515 (MO). NEW YORK. Bronx Co.: 17 Apr 1988, Mori & Gracie "o

(GH, MO, NY). Nassau Co.: 8 May 1950, Abbott s.n. (CU). Suffolk Co.: 1 May 2003, Glenn 8215 (BKL). Westchester Co.: 2 May

1994, Walker 684 (NY). NORTH CAROLINA. Wake Co.: 11 Apr 2005, Krings 1271 (AUA, F, FLAS, GA, LSU, MISS, NCSC-2 rub

NCU, TEX, UNA, US, USF, USCH, VDB). OHIO. Butler Co.: 5 Apr 1988, Cusick 27224 (OS). PENNSYLVANIA. Delaware Co.: 16 May

£ 4L D H ID hl EP. FT,

208 Journal of Texas 3(

1915, Cramfondi* s.n. (PH); 26 May 1920, Long 23006 (PH). Philadelphia Co.: 7 May 1910, St. John 111 (GH). TEXAS. Tarrant

Co.: 8 Mar 2008, Nesom FW08-1 (NCSC)

Ranunculus ficaria L. (subspecific determination not feasible)

U.S.A. CONNETICUT. Middlesex Co.: 27 Apr 1991, Swan s.n. (Y). New Haven Co.: 18 Apr 1995, Brown 1 (Y); 10 May 1993,

Souther s.n. (Y). MARYLAND. Baltimore Co.: 25 Apr 1948, Moudry s.n. (MARY); 21 Apr 1968, Redman s.n. (BALT). Howard Co.:

17 Apr 1966, Engh s.n. (MARY). Prince Georges Co.: 19 Apr 1979, Mills Jr. 14 (BRIT). MASSACHUSETTS. Barnstable Co.: 27

Apr 2001, Mehrhoff 21129 (CONN). MICHIGAN. Ingham Co.: 10 Apr 1991, Stephenson s.n. (MSC). NEW JERSEY. Union Co.: 3

Apr 1977, Moldenke & Moldenke 31285 (NY); 10 May 1978, Moldenke & Moldenke 31642 (NY). NEW YORK. Nassau Co.: 4 May

2004, Bennett 030031 (BKL). Queens Co.: 27 Apr 1918, Ferguson 7 (NY); 17 Apr 1921, Martin s.n. (NY); 30 Apr 1876, Schrenck

s.n. (CU). OHIO. MONTGOMERY Co.: 18 Mar 1990, McCormac 1977 (MICH, OS). OREGON. Multnomah Co.: 4 Apr 1991, Zika

11066 (OSC). PENNSYLVANIA. Chester Co.: 15 Apr 1957, Wilkens 9129 (PH). Delaware Co.: 7 May 1977, Brown 199 (Y); 24 Apr

1894, MacElwee Jr. s.n. (PH). Philadelphia Co.: 11 Apr 1954, Fogg Jr. 21461 (PH). WEST VIRGINIA. Monongalia Co.: 12 Apr

1995, Baer s.n. (WVA). Ritchie Co.: 25 Apr 1971, Elliott s.n. (WVA); 27 Apr 1991, Grafton s.n. (WVA).

—

ACKNOWLEDGMENTS

We thank the following herbaria and their staff for ilable to us for study: A, AUA, BALT,

BH, BKL, BRIT, CONN, CU, DOV, F, FLAS, GA, GH, HNH, ILLS, KE, LGO, LSU, MARY, MASS, MICH,

MISS, MO, MOR, MSC, MT, MU, NA, NCSC, NCU, NHA, NY, OS, OSC, PH, POM, TENN, TEX, UNA, US,

USF, USCH, VDB, VPI, WTU, WVA, Y. We also thank David Schuman and Steve Stanislav for statistical

support. Three anonymous reviewers offered constructive comments on an earlier draft of the ms.

REFERENCES

ALMEIDA-PINHEIRO DE CARVALHO, M.A., C.C. WiLcock, T.M. Marques Dos SANTOS, IC. VatE-Lucas, J.F. TEIXEIRA-GANANGA, E. FRANCO,

D. THANGADURAI, D. MURALIDHARA-RAO, AND D. FrEITAS-SOUSA. 2004. A review of the genus Semele (Ruscaceae) sys-

tematics in Madeira. Bot. J. Linn. Soc. 146:483-497.

BAiLEY, L.H. 1935. Cyclopedia of horticulture. MacMillan Co., New York.

Bett, FH.1945. The genus Ranunculus in West Virginia. Amer. Midland Naturalist 34:735—743.

Benson, L. 1942. North American Ranunculi-V. Bull. Torrey Bot. Club 69:373-386.

Breiman, L., J.H. FRIEDMAN, R.A. OLSHEN, AND C.J. STONE, 1984. Classification and regression trees. Wadsworth & Brooks/

Cole, Monterrey.

Dickinson, T.A. AND J.B. Phipps. 1985. Studies in Crataegus L. (Rosaceae: Maloideae).XIll. Degree and pattern of phe-

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DurRENE, M, J.L. GATHOYE, AND D. TYTECA. 1991 | Dactylorhiza (Orchidaceae)—

the D. maculata group. PI. Syst. Evol. 175:55-72.

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GLEASON, H.A. AND A. CronquisT.1991. Manual of vascular plants of northeastern United States and adjacent Canada.

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Gower, J.C. 1971. A general coefficient of similarity and some of its properties. Biometrics 27:857-871.

GREILHUBER, J. 1974. Ein chromosomensatz von Ranunculus ficaria subsp. calthifolius. Mitt. Bot. Arbeitsgem.

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IHAKA, R. AND R. GENTLEMAN. 1996. R: A language for data analysis and graphics. J. Computat. Graphical Statistics

5(3):299-314.

Jovy, S. AND A. Bruneau. 2007. Delimiting species boundaries in Rosa Sect. Cinnamomeae (Rosaceae) in Eastern

North America. Syst. Bot. 32:819-836.

KRINGS, A, A.S. WEAKLEY, J.C. NEAL, AND E.C. Swag. 2005. Ranunculus ficaria (Ranunculaceae) new to North Carolina

and an updated key to Carolina congeners. Sida 21:2429-2437.

Marchant, CJ. AND C.A. BnicHTON. 1974. Cytological diversity and triploid frequency in a complex population of

Ranunculus ficaria L. Ann. Bot.38:7-15.

Marosen—Jones, E.M.1937. Ranunculus ficaria Linn.: life-history and pollination. J. Linn. Soc. Bot. 50:39-55.

Nesom, G.L. 2008. Ranunculus ficaria (Ranunculaceae), naturalized in Texas. J. Bot. Res. Inst. Texas 2:/41—742.

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SARGENT, D.J., M. Gege, J.A. HAWKINS, MJ. WILKINSON, N.H. Batey, AND D.W. Simpson. 2004. Quantitative and qualitative

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210

BOOK REVIEW

WILLIAM A. WEBER AND RONALD C. WITTMANN. 2007. Bryophytes of Colorado: Mosses, Liverworts, and

Hornworts. (ISBN 978-0979090912, pbk.). Pilgrims Process, Inc., PO. Box 32597, Santa Fe, New

Mexico 87594-2597, U.S.A. (Orders: www.pilgrimsprocess.com, 1-720-937-6641). $29.95, 231 pp.,

8 1/2" x 11".

Almost too small to bud NE PAPER en masse, De are not pe When viewed with a hand lens or microscope some of the

unusual, lare revealed

e r D

The genesis for the cur 1 1 with a preliminary led by Weber in 1973. He presented a joi del i keys B us

mosses, at an time e identifying shou 290 pe itl i g d hornworts. The current

I is th prehensive study of ‘Colorado mosses and liverworts to date. It en

b ful in adj f hl d intl i gi ftl western e States.

Bernd: and students will find this bool luable ref though it d t Keys to the

genera and pe i mbined s notes on E in the dae th trate on n distinguishing characteristics of

idi 1 for the identificati but the authors

J g I

have been undis to produce a useful ke to he eae They hat tl family | ilabl the internet, but that for the

seasoned MISCERE family id d that for the beginner tl |

11 d D pa | $, A 1 id f, de 1 11 f A

Only a few e&t e E

1] D 13 1 ET A t

y and anatomy.

Recommended for 1 1 ] ig 11 1 El J 1 Should h 215411 7 j 11 RI +L, f. 1

x E XA7AL e LEN. f, E. E 1 Td E fr^ J n 14 J +l f, fel a PONE x Ir

William A. y olorado

Museum Herbarium. He is tl I f bool fl ing plants, ferns, and lichens as well as bryophytes, concentrating

on Colorado m E d M E Pon in scope as well

Wi plished amateur botanist, and is the co-autl f the l versions of

Ronald

the Colorado Flora.—Gary L. prem Librarian Botanical Resear Institute of Texas, Fort Worth, Texas 76102-4025, U.S.A.

J. Bot. Res. Inst. Texas 3(1): 210. 2009

THE TYPES OF ASTRAGALUS SECTION DIPHYSI (FABACEAE), A COMPLEX

ENDEMIC TO WESTERN NORTH AMERICA, PART I: LECTOTYPIFICATIONS,

EPITYPIFICATIONS, AND NEW COMBINATIONS OF SEVERAL TAXA

J. Andrew Alexander

Curator, Utah Valley University Herbarium (UVSC) and Research Associate

Pu A as Utah Valley University Wesley E. Niles Herbarium (UNLV)

st University Parkway as of Nevada, Las Vegas

prn Utah, 84058, U.S.A. 4505 Maryland Parkwa

alexanja@uvu.edu Las Vegas, o 4004 U.S.A.

RESUMEN

This is the first of several Pipe discussing typification issues Pipes JURE snis leading to a monograph of Astragalus L. Section

Diphysi A. Gray. À lectot designat Hook. Original material designated here as the

nol ype of Ass diphysus A. Gray var. s albiflorus- A. nid [sAsirzgalus lentiginiosus var. albiflorus (A. Gray) Schoener] was rediscovered

at GH over y used name, A. lentiginosus var. diphysus (A. oy M.E. Jones, is discussed.

ds are deere for. lennon var. micans Barneby and A. lentiginosus var. oropedii Barneby. Three new combinations, previ-

are proposed: A g ar. bryantii (Barneby) J.A. Alexander, A. lentiginosus var. iodanthus (S.

Watson) J A. Alexander and A. lentiginosus var. pseudiodanthus (Barneby) J.A. Alexander.

RESUMEN

T 1 - > : 1 dz +1 de tin Es A ey J 1 As 43

1 F I £

T ph ZA Gra 1 Al x um T* I1 T. a |

A sidus L.S

e como Rd de Astragalus dps A idi var. t albiflorus 5 Gray [Astragalus lennan Si var. albiflorus (A. Gray) Sdhóeed fue

i lentiginosus var. diphysus (A. Gray) M.E. ones Son

deste d epitipos de A. lentiginosus y var. micans Barneby y A. lentiginosus var

lentiginosus var. bryantii (Barneby) J. A. aa A. lentiginosus var. iodanthus (S.

watsen) A. Alexander, y A. nia var. pseudiodanthus (Barneby) J.A. Alexander.

INTRODUCTION

Astragalus L. Section Diphysi A. Gray, a section native to Western North America, is a problematic species

complex composed of taxa that were originally described as or have been split as species in the past 150

years. Astragalus lentiginosus Douglas ex Hook. (in Hooker 1831) and A. diphysus A. Gray (in Gray 1849)

were the core species of Section Diphysi in the first comprehensive monograph of Astragalus by Asa Gray

(1863). Sereno Watson (1871) expanded Gray's concept of Section Diphysi to include A. coulteri Benth. and

A. platytropis A. Gray, both of which have inflated pods similar to A. lentiginosus. Gray's monograph (1863)

and Watson's (1871) revision were the primary references for Astragalus taxonomy until Marcus E. Jones

began publication of treatments in the genus in the late 1890s.

In his 1898 publication, Jones proposed that species from Gray's (1863) Section Diphysi should be com-

bined, as varieties, into a greatly expanded concept of A. lentiginosus. His full treatment of these species was

not widely known until he published his Revision of North-American Species of Astragalus in 1923 (Barneby

1964). Jones was a field botanist of unmatched experience. As a taxonomist, he had a disdain for rules of

priority and nomenclature, and was known for his often frustrating brevity. Barneby (1945) found “that many

of his names covered unreasonable extremes of variation, or that his descriptions and indications of range

were either inaccurate or actually misleading" (p.65). Barneby was able to decipher Jones' species only after

extensive study of his annotations and vouchers at POM and duplicates at other herbaria (see discussion

of A lancearius A. Gray and A. episcopus S. Watson, Barneby 1964, p. 267; compare descriptions of varieties

of A. lentiginosus of Jones 1923, p.124-125 and Barneby 1945, p. 65-152; and discussion of the taxonomic

J. Bot. Res. Inst. Texas 3(1): 211 — 218. 2009

BR oummalof ical I i Texas 3(

confusion between Jones’ species and hybrids which are now synonyms of A. argophyllus Nutt. var. martinii

M.E. Jones, A. marianus Barneby, and A. desereticus M.E. Jones, Barneby 1964 p.629—635).

Concurrently, Rydberg (1929) delimited all members of Section Diphysi as separate species of the

genus Cystium Steven. Rydberg appeared to have an idealized concept of each of his species. He regularly

composed his morphological descriptions and keys from features exclusively on a single type. As a result,

his keys and descriptions sometimes did not match the morphology on all specimens he annotated for his

monograph. Despite this, Rydberg for his time was more precise than all previous workers in the genus

(Barneby 1964).

Barneby (1945) was the first to comprehensively evaluate Jones’ and Rydberg’s hs and found

that only one overall classification scheme fit the taxa related to A. lentiginosus well: he don of

all former species into varieties. Barneby's (1945) treatment and his later Monograph (1964) still serve as a

precise baseline from which all taxonomic treatments in this group are based.

Different taxonomic interpretations of the degree of morphological differentiation among the many

widespread, sympatric, and geographically isolated taxa within this complex have been the major source of

disagreement between the revisions of Rydberg (1929), Jones (1923), Barneby (1964, 1989), Isely (1998), and

Welsh (2007). Ultimately Ene major B in varietal delimitations poe these workers stem from

their individual i of ty] d selection of Į | i g the

nomenclaturally ‘ ‘typical” variant. Despite Barneby’s decades of work in sorring out the enon issues

leading up to his Monograph, it is still difficult to determine which variant is nomenclaturally “typical” due

to the often poor quality of the type material. Barneby M Qs for is adr nature of many ee by

examining specimens from the vicinity of the type locality. Th

the informal basis for his concept of the nomenclaturally typical variant and ds sia morphological

descriptions.

The most recent monographs, Isely (1998) and Welsh (2007), have made significant additions to the

knowledge of taxonomic boundaries in this section, but each has added their own entangled varietal de-

limitations. However, both still use Barneby (1964) as a nomenclatural foundation. For a monograph of the

section that is more comprehensive and thorough than Barneby's, nomenclatural issues that have resulted

in conflicting varietal delimitations between the major monographs must be resolved. Fortunately, there

are more formal processes in use today that seek refine the delimitation of taxa with problematic types. The

Linnaean Plant Name Typification Project (<http://www.nhm.ac.uk/research-curation/research /projects/

linnaean- preto has been the forerunner in using modern additions to the lectotypification, epitypifi-

cation and ion articles in the I ional Code (McNeill et al. 2006). Using the procedures and

philosophy of Vander Kloet (1989), Turland and Jarvis (1997), and Krings (2008) as models, this treatment

is the first a series of papers proposing new combinations, lectotypifications and epitypifications within

Section Diphysi. Due to the varietal and nomenclatural complexity of Section Diphysi, a series of papers

instead of a single, expansive treatment is necessary. The paali is to stabilize and refine the nomenclature in

this group, which will enable the creation of a j d less problematic taxonomic revision

of this morphologically and taxonomically difficult TE

METHODS

Herbarium specimens were examined at UC in December of 1999, GH in August of 2002, and NY in October

of 2003. Additional herbarium specimens were obtained on loan from CAS, DS, K, POM, RM, and RSA.

In addition to loans, research was conducted from 2002 to 2008 using the following online type specimen

databases: Consortium of California Herbaria Specimen Databases (CAS, DS, JEPS, RSA, SD, UC, UCR,

UCSB), University and Jepson Herbaria (2008b); Index of Botanical Specimens (GH), Harvard University

Herbaria (2008); Type specimens at the Herbaria (JEPS, UC), University and Jepson Herbaria (2008b); Kew

Herbarium Catalogue (K), Royal Botanic Gardens, Kew (2008); Tropicos (MO), Missouri Botanical Garden

(2008); The C V. Starr Virtual Herbarium (NY), New York Botanical Garden (2008); Type Specimen Register

Alexander, Types of Astragalus sect. Diphysi 213

(US), United States National Herbarium (2008). Though the International Code of Botanical Nomenclature

(McNeill et al. 2006) was the basis for all ] | decisions, many papers resulting from The Linnaean

Plant Name Typification Project (<http://www.nhm.ac.uk/research-curation/research /projects/linnaean-

typification/>) were examined for examples of specific applications of the Code. Other typification papers,

especially those resulting from the various tropical flora projects, were also examined. Turland and Jarvis

(1997) and Krings (2008) were most frequently consulted for example typifications. Vander Kloet (1989)

was consulted to determine how other Douglas taxa described by Hooker were lectotypified (i.e., Vaccinium

membranaceum Douglas ex Hook., p.133).

RESULTS

ps m lentiginosus us ex Hook Fl Bor. bend 1:151. 1831. Tragacantha cie (Douglas ex Hook.)

e, Revisio Gen. Pl. 2:946. 1891. P} (D Hook.) Piper, Contr. U.S. Natl. Herb. 11:368. 1906. d

n (Douglas ex mee ae Bull. Torte Bot. Club 40: 50. 1913. PROTOLOGUE: = Ee ranges a des E Mountains

North West America. Douglas." Tyre: [U.S.A. OrEGON]: in the valley of the Bl awallah us

Walla] and Utala [Umatilla] Rivers, [no date], David Douglas s.n. (LecromwE [first step] designated by xis 1964: K; LECTOTYPE

ee ae undas mid K diga d smaller, fruiting, Di wd ao on the oe left hand side of the sheet

ight); [U.S.A. OREGON] thern branches of the

Goiaba oey collected at ihe same pie Ad as a lectoypel, ED cate David Douglas s.n. (zPrrYre, designated here: K 264945},

the mostly flowering,

1 1 3; f£, > 1 £] 1 SA 1 1 d us. P: 11 f .L E sel

MM p yI ig p

TNR Ine Tri : hiato] ich f fi i ftheD As with

MR Douglas opener described as new qe ~ Hooker (1831 El none of the pe spectes have a fone Ga matches the

From an examination of the annotations on the three type sheets, Asa Gray in 1868 was the first to annotate

the A. lentiginosus specimens at K. Gray did not indicate which of the sheets was the type nor did he annotate

every sheet, but he was the first to recognize that the central element (K264945) on one of the types was A.

lentiginosus. However, he did not recognize that the other two elements on this sheet were type material for

A. diaphanus Douglas ex Hook. After Gray, the delimitation of typical Astragalus lentiginosus has largely been

based on the morphology of specimens found in the vicinity of the Blue Mountains in Oregon. Since both

A. lentiginosus var. salinus (Howell) Barneby and A. lentiginosus var. platyphyllidius (Rydb.) Peck can be found

in southern Blue Mountains of Oregon, some populations of these taxa have been misinterpreted as typical

A. lentiginosus. Barneby (1945) was the first since Gray to critically analyze the types of the A. lentiginosus

complex and refine typical A. lentiginosus morphologically. The only Douglas specimen of A. lentiginosus he

saw for this revision was the fragmentary type at GH. Barneby (1964, p.917) indicated a specimen a K found

“near the source of the Wallahwallah and Utala rivers” was the holotype. This is the first step lectotypifica-

tion of Astragalus lentiginosus (McNeill et al. 2006, Article 9.8). Unquestionably, Barneby selected a fruiting

specimen at K as the lectotype of A. lentiginosus since he knew that Hooker (1831, p.151) indicated that "the

flowers of this do not appear to have been seen by Mr. Douglas” and that "floribus - ?” was the only reference

to flowers in the diagnosis. However, there are two sheets in fruit from the same individual at K: one from

the Herbarium Benthamianum (K 264012) and one from Herbarium Hookerianum (K 264017). It is likely

that Barneby chose the specimen from the Hooker Herbarium, however no specimens at K are annotated

by him. The individual, K264017, is mounted with a specimen of A. lentiginosus var. lentiginosus collected by

Dr. Lyall (K 264018). After an examination of the diagnosis and all the types of A. lentiginosus, the second

step lectotypification was made herein to unambiguously specify which sheet, Kew accession number, and

elements mounted on the sheet correspond to the lectotype in accordance with McNeill et al. (2006) Article

9.15. The second specimen, K 264012, is an isolectoype of A. lentiginosus.

Since the fruit of typical Astragalus lentiginosus is not by itself agree precise taxon delimitation

based on the lectotype is not possible. Both fruit type and flower si ic features that distinguish

typical A. lentiginosus from other sympatric varieties. Astragalus lentiginosus var. salinus has small whitish

flowers (keel «9 mm) and thin walled, bladdery inflated pods. Astragalus lentiginosus var. platyphyllidius has

£L Dag H ID ll II P, £T,

214 Journal of Texas 3(

larger whitish flowers (keel 11-15 mm long) and thick walled, curved pods inflated only towards the base.

Throughout its range, the fruit of A. lentiginosus var. lentiginosus can take both of these forms. The lectotype

is a late season specimen with only mature, thick walled, curved pods inflated only at the base. The size of

the flowers is unknown.

For a precise interpretation of nomenclaturally typical Astragalus lentiginosus, a flowering specimen

was chosen from the original Douglas specimens. The epitype of Astragalus lentiginosus is designated herein

(K264945; McNeill et al. 2006, Article 9.7) as the larger individual (keels 8-9 mm long) in the center of the

sheet. It is mounted with two other fragmentary elements. Element 2 (K264015) is a fruiting specimen of

A. diaphanus. Element three is mounted in two different places on the sheet, has received different accession

numbers (K264016 and K264014), and is a flowering specimen of A. diaphanus. This entire sheet was from the

Herbarium Hookerianum. Elements two and three are potential lectotypes or isolectotypes of A. diaphanus

and will be discussed in a future publication (Alexander, in prep). The Douglas label is associated with a

portion of element three and indicates it was collected “on the banks of streams on the southern branches

of the Columbia." Presumably, the flowering A. lentiginosus element was collected at the same locality as the

A. do elements. Alternatively, the epitype and the lectotype could have been collected from the same

l vicinity since the Walla Walla and Umatilla Rivers are two of several southern branches of the

Columbia River in northeastern Oregon.

desa braces var. albiflorus A. Gray, Pl. Fendler. Novo-Mexicanae, Mem. Amer. Acad. Arts II. 4:34.

1849 bib em lentiginosus var. albiflorus (A. Gray) Schoener, Great Basin Naturalist 34:180. 1974. ProroLoGue: “with the preced-

ing [Fend r 146 from “around Santa Fe,” New Mexico]. No. 147.” Tyre: U.S.A. Plantae Novo-Mexicanae [New Mexico]: [locality not

a on label], 1847, A. Fendler 147 (novorvee: GH 112365! H 587151, K 2640231, MO).

Notes.—When Barneby (1945) ined the Fendl fA dipl t GH, he only found one type specimen of Astragalus

diphysus var. albiflorus A. Gray (Fendler 147 GH58715) This sheet was nn of a collection transferred from the herbarium of the

Boston Society of Natural History to GH in 1941. pod pr aa P. i noted mu iioi pene “was received long after

Gray’s death and not annotated by him,” and sel YE d, Barneby may

have chosen the MO sheet because GH58715 has an uncertain history and was mounted wi i b p galus dipl vat.

diphysus (Fendler 146 GH58716).

In this study: e of Du Bes aus ee rues dl pu to uate institutions revealed mal d SUD.

E ¿E

Novo-Mexicanae, ] he other instituti ] inati heir dupli based his manuscript. The Fendler types

i] LI! 1 1 : : $ : 2 | J 1 n 1 Mi na 1 ab 1

In 2002, a thorough search was conducted at GH of all the Monk: American fold 1 types of Astragalus. The origina ELA

of A. diphysus var. albiflorus (Fendler 147 GH112365) with th j ] d the d

rediscovered. This type specimen was recently divided from another ipic collection and remounted on a new shee which

is the likely reason that Barneby was not able to find the original type. Currently, GH112365 is mounted with an unaccessioned,

non-type specimen of A. iiie Sn by pm in 1883 in mn Abuqurqe New Mexico. The emia of the holotype of

A. diphysus var. albifl bys (1 leill et al. 2006, Article 9.17.

aa pane var. diphysus (A. Gray) M.E. Jones, Proc. Calif. Acad. Sci. H 5:673. 1895. Astragalus

physus A. Gray, Pl. Fendler. Novo-Mexicanae, Mem. Amer. Acad. Arts IL 4:34. 1849. Gona diphysa (A. Gray) Kuntze, Revisio

Gen. Pl. 2:944. 1891. Cystium diphysum (A. Gray) Rydb. Bull. Torrey Bot. Club 32:659. 1905. ProroLoGUE: “around Santa Fe [New

Mexico]...No. 146." rr: U.S.A. Plantae Novo-Mexicanae [NEw js ei! not ud on label], 1847, A. Fendler 146

(noLorvee: GH 587141; isotypes: BM, E GH 587161, K! [3 Sheets], MO, NY

Notes.—The ar name iid what dini been nina i = as id. cate A var. diphysus (A. Gray) M.E. Jones has been con-

Barneby (1964, p. 941) stated “strict adherence

Rules of N ] ire that tl lier i varietal rank [in his opinion A. diphysus var. albiflorus] takes precedence

q. A. diphysus var. diphysus]." He rejected making a new Po A. ene var. albiflorus, since it would result in “an

absurdity and runs counter to common sense” of using th Based on her interpretation of the Article

11 of the 1972 edition of the International Code, Scheer (1974) concisa: “the name, var. diphysus, is anteceded by the name

A. diphysus var. albiflorus...[and] the older name in the same rank has poene. (p. 180). Schoener (1974) then made the new

combination, A. lentiginosus var. albiflorus (A. Gray) Schoener. S (Welsh 1978; Isely odd. have iini pd Schoeners

combination, while the most recent (Welsh 2007) has used Jones’. The most recent I j Il et al. 2006, Article

11.6 and 26.3) resolves this controversy. Two taxa were described by Gray (1849): first, A. diphysus (based on ae 146); and

Alexander, Types of Astragalus sect. Diphysi 215

second, A. diphysus var. albiflorus ind on "iid 147). The HE taxon, Á. iain var. albiflorus ically creates the priorable

tion, A. lentiginosus var. diphysus, that M.E.

autonym, A. diphysus var. diphysus.

Jones made in 1895 has priority over Schoeners 1974 Eom BA: A. Top var. albiflorus.

Astragalus lentiginosus var. bryantii (Barneby) J.A. Alexander, comb. nov. Astragalus bryantii Barneby, Proc. Calif.

Acad. Sci. 4, 25:156. 1944. PROTOLOGUE: “ARIZONA: at the head of Phantom Canyon in the Grand Canyon of the Colorado River,

Coconino Co., 15 Dec. 1939. Collected by Dr. H.C. Bryant...” Tyre: U.S.A. Arizona: [Coconino Co.]: head of Phantom Canyon, in

Grand Canyon, 15 Dec 1939, H.C. Bryant s.n. (HOLOTYPE: CAS 2939401; sorvre: US 1769041, internet image.

5 fn by) LA AI EI 1 1 A Pee | A itl A lentiginosus var. palans

Notes.—Astragalus lentiginosus var. }

(M.E. Jones) M.E. Jones or A. lentiginosus var. mkd (A. Gray) M.E. joes in hist (Barneby 1944,

Barneby ae any ie Isely 1998, Welsh 2007). Welsh et al. (2003) and mes e cra that Pies in Utah

prev iously ryantii Barneby are A. piles laa var. palans All full 1 I

been published, } 1 by Barneby (1964) and Wi Ish (2007) ines 828, Gaines 1005,

O E 4 JM 4

and Gaines 1009, all fom NAU. Although t! pecific speci ined, Al 1 (2008) ae not BER ay en

from Utah previously d d as A. bryantii Barneby or A. lentiginosus var. pal hat has tl found in

1 1 $ e 1 Dl D 1 E $ f. A Á Jaji * 11 AT 1 K^ rA QN

por y G Canyon y " non var. boat is

s » Fa 1 = 1 -1 c fl 333 J D e | f£ Dh t

p Ranch, Coconino

Co., Arizona. As a as N these voptladions are long-dist disjuncts from that of tl population of A. lentiginosus

var. palans. In addition, the Lin variation d in i specimens of A. up var. TUM is n e that of A

A. lentiginosus

lentiginosus var. palans. However.

var. wilsonii (Greene) Barneby and A. bici var. ursinus de Gray) inia Bameby ea theorized u that oui taxon was closely

related to A. lentiginosus var. mokiacensis, a rel y , Isely 1998,

Welsh et al. 2003, Welsh 2007). Further favestination of habitat in the canyons of the Colorado River eastward ni the nearest

populations of A. lentiginosus var. mokiacensis at Emory Falls (Clover 6079, CAS!) and Quartermaster Canyon (Goodding 15-41, RM!)

may reveal populations of A. lentiginosus var. bryantii intermediate to A. lentiginosus var. mokiacensis.

Astragalus lentiginosus var. iodanthus (S. Watson) J.A. Alexander, comb. nov. Astragalus iodanthus S. Watson, Bot.

g. 70. 1871. ProTOLOGUE: “in the foothills of Western Nevada from the Virginia to the West H ; 4500-6000 feet

altitude... [S. Watson] 269.” Tyre: U.S.A. Nevapa: [Pershing Co.]: West Humboldt Mountains [Humboldt Range, NE of Lovelock],

Jun 1868, S. Watson 269 [in part] (LecrorvrE, designated by Barneby 1964:962: US 469091; isoLecroryres: GH!, NYi, YU).

Astragalus iodanthus var. diaphanoides Barneby, Leafl. W. Bot. 4:50. 1944. Tire: U.S.A. Nevapa: Washoe Co.: between Reno and Dewey, 10

: RSA)

J 3, H.D. Ripley & R.C. Barneby 5659

Astragalus iodanthus var. vipereus Barneby, T4 New York Bot. Gard. 13:963. 1964. Tree: U.S.A. IDAHO: e Co.: on cobblestone

bluffs near Bruneau, 31 May 1945, H.D. ecd a R.C. ao Tum eee is ISOTYPES: n RSA).

Notes.—In his taxonomic revision, Al gh deg F E

ferentiation between A. ingre var salis. A. iodanthus and A. pseudiodanth 1 continued separation of th I

species is not supported. The new lin Alexander (2008) are made formal herein.

Traditionally, differences in pod morphology have been the basis for the separation of Astragalus iodanthus

S. Watson and A. pseudiodanthus Barneby from A. lentiginosus. Both have a deciduous, mostly uniloculate to

partially biloculate pod with a septum less than half the width of the locule. This feature does differentiate

these two taxa from the varieties of A. lentiginosus with bladdery inflated, completely biloculate pods. The

contrast appears so great that in other sections of Astragalus, these differences have been considered species-

level indicators. When these two taxa are merged into the complex folds of A. lentiginosus, along with the

widespread and morphologically diverse taxa allied with A. lentiginosus var. palans (some of which were also

originally recognized as a species), they form a continuum of variation—morphologically and geographi-

cally. In the Mojave and Sonoran Deserts, A. lentiginosus var. mokiacensis and A. lentiginosus var. maricopae

Barneby form the southern end of the continuum with persistent, mostly straight, tubular, scarcely inflated

pods and a septum extending from one-half to slightly over three quarters the width of the locule. In the

Colorado Plateau vicinity, A. lentiginosus var. palans, forms the central and eastern axis of the continuum

with deciduous, mostly straight to nearly 180° curved, tubular to triquetrous, scarcely inflated pods and a

septum from one-half to slightly over three quarters the width of the locule. In the Great Basin, A. lentiginosus

var. iodanthus (S. Watson) J.A. Alexander and A. lentiginosus var. pseudiodanthus (Barneby) J.A. Alexander

form the western axis of the continuum with deciduous, mostly 180° curved, triquetrous, scarcely inflated

pods and a septum from one-quarter to less than one-half the width of the locule. Barneby (1964) was the

216 J i i fTexas 3(1)

first to recognize this similarity. In his uniquely succinct style, he stated "this pair of species [A. iodanthus

and A. pseudiodanthus] posses no character which cannot be matched somewhere in A. lentiginosus” (p.

911) and *it is often difficult or nearly impossible to separate flowering material of [A. iodanthus] from the

polymorphic A. lentiginosus, from which it differs principally in a tendency to dorsiventral and triquetrous

compression of the fruit, which is never inflated and commonly very strongly incurved... however in A.

lentiginosus var. palans, the lace-ellipsoid, little inflated pod varies from erect to decurved and its section

varies from round to triangular, so that sometimes the only technical differential character that remains is

the broader septum" (p. 959-960).

Other varieties of A. iodanthus have been recognized in Barnbey (1964), Isely (1998) and Welsh (2007),

but they are treated herein and in Alexander (2008) as synonyms. Results of population level morphological

analyses in progress may warrant recognition of A. iodanthus var. diaphanoides Barneby and A. iodanthus var.

vipereus Barneby at the varietal-level within A. lentiginosus.

Astragalus lentiginosus var. micans Barneby, Leafl. W. Bot. 8:22. 1956. ProroLocur: “CALIFORNIA: lower slopes of

sand dunes at southeast end of Eureka Valley, east of Inyo Mts., Inyo County, elevation 3050 ft. May 13, 1955 (fr.) John C. Roos 6354, and at

the same place, ae ae i ape 9, Te = ), Munz & Roos 20851. o Rancho Santa Ana Bot. Gard.” Type: U.S.A. CALIFORNIA:

13 May 1955, J.C. R 354 ( , designated here: RSA

Invo C

of Inyo

1001791; isoLecToTYPES: GHI, Kl, NY! [2 sheets], OSC!, RSA 1146941, SD, ya ip sheets], US 2483071, internet image!); U.S.A. CALIFORNIA:

Inyo Co.: on sand dunes at S end of Eureka Valley, E of Inyo Mts., 9 Apr 1955, P Munz & J.C. Roos 20581 (errrvre, designated here: RSA

100180!; isoEprrvees: NY!, OSC!, RSA 1099741, RSA 115975!, UCD).

Notes.—Barneby (1956) eed two Ones (as p in the protelogue: one dios dim as C. Roos oi RSA) e one with

e e & Roos 2058 I rneby

t as es dAsd 7" lectotype Maleh (9007) al 1 R hire f, 1 u 4 hoth

* r

KDCAIT/An 1 L 1 (Me Nlei3ll et al 1

as npe. ap mene ee Roos a 1 79) is designated yp 2006, Article 9.2 and

0] 1 s 1 1 1 f Leal c i f, J

ES fq 4E.

on the T syntype. To keep all of tl ial used in the original type description (and the original i rol theanthon

intact, the flowering specimen Mite & Roos 20581 (RSA100180) is selected h the epitype (McNeill et al. 2006, Article 9.7)

It should be noted that in his original type publication, Barneby (1956) mistakenly cited “Munz & Roos 20851”

as the syntype at RSA. Also Welsh (2007) mistakenly stated that the RSA syntype was “Munz & Roos 20815”.

All syntype specimens at RSA bear the correct collection number, Munz & Roos 20581. Also all isosyntypes

at various institutions listed above are labeled with the collection number, Munz & Roos 20581. There do not

appear to be any types with labels bearing the typographical errors published by Barneby or Welsh.

Astragalus lentiginosus var. oropedii Barneby, Leafl. W. Bot. 4:135. 1945. Prorotocue: “Kaibab Trail to Roaring

Springs, Grand Canyon National Park, Coconino County, Arizona, 22 September 1938, fruct., Eastwood & Howell No. 7064 (Herb.

Calif. Acad. Sci. No. 262056). Also ibid., 23 June 1933, flor., Eastwood & Howell No. 1054 (Herb. Calif. Acad. Sci. No. 211208, co-

TyPE).” Type: U.S.A. Arizona: Coconino Co.: Kaibab Trail to Roaring Springs, 22 Sep 1938, A. Eastwood & J.T. Howell 7064 (LECTOTYPE,

designated by Barneby 1989:158: CAS 2620561); U.S.A. Arizona: Coconino Co.: Kaibab Trail to Roaring Springs, 23 Jun 1933, A.

Eastwood & J.T. Howell 1054 (errrveE, designated here: CAS 211208!)

Notes. mr (1945) designated two syntypes (as cotypes) in the protologue: one us i ee & ee oe baie. and

one with flowers (Eastwood & Howell 1054, CAS). In his monograph, pao (19 p

pet For the Intermountain Flora, Barneby (1989) listed E & Ho oe 7064 as the iE for A. lentiginosus var. oedi

bini winch rami isa latory pintaron (McNeill et al 2006, Article 9.8). The flowers of A. lentiginosus var. oropedii are the

À pityr led for precise taxonomic application of this rua cn D of A. lentiginosus var.

d herein as the floweri designated by Barneby as a sy twood & Howell 1054 (CAS; McNeill

1 1 ; 1 2a] Jin R hw ioinal d rintion

fa A

E C»

et al. 2006, Article 9.7), a d

Astragalus lentiginosus var. pseudiodanthus (Barneby) J.A. Alexander, comb. nov. Astragalus pseudiodanthus

TT iur W. Bot. H a m s iodanthus var. jum die enr Isely, Syst. Bot. 8:422. 1983. PRorOLOGUE:

"NEVA f Cactus Peak outheast of Tonopah, Nye Co.: alt. 5500 ft, 3 Jun

1941, Ripley & Barneby N 3725." Tre: U.S.A. Nevapa: Nye Co.: plateau N vet Cactus Peak, about 20 mi SE of Tonopah, 3 Jun 1941,

H.D. Ripley, R.C. Barneby 3725 (notorvee: CAS 2904051; isotype: RSA, PO

fd l } t I d A. lentiginosus the al t of A. lentigi-

F Agent ast m

Notes. discussion

nosus var. iodanthus.

Alexander, Types of Astragalus sect. Diphysi 217

Barneby (1964) was the first to describe in detail of the gas 2 nen of A. io den var. pseudio-

danthus and some populations of A. lentiginosus var. iodanthus. T por

but spread throughout its range. It is likely that A. lentiginosus var. pseudiodanthus recently diff iated from

several of these relictual, intermediate populations and became adapted to a stabilized sand dune habitat.

Similar adaptations have occurred in populations of A. lentiginosus var. variabilis Barneby, A. lentiginosus var.

fremontii (A. Gray ex Torr.) S. Watson and A. lentiginosus var. stramineus (Rydb.) Barneby (which may itself

be a sand dune derivative of A. lentiginosus var. fremontii or A. lentiginosus var. vitreus Barneby).

Population level morphological analyses in progress, leading to an overall monograph of Section Diphysi,

may provide more details on the taxonomic status of these sand dune variants.

ACKNOWLEDGMENTS

Numerous individuals provided advice, funding and encouragement during various stages of this project

including: Wesley E. Niles; Aaron Liston; Richard Halse; Kathryn Birgy of the UNLV Herbarium; Arnold

Tiehm and the Northern Nevada Native Plant Society (and the NNNPS Small Grants Program); Lisa Decesare

of the Library of the Gray Herbarium, Harvard University; and the Moldenke Fund for Plant Systematics

and the Hardman Fund at Oregon State University. Additional thanks go to the late R.C. Barneby and the

ond dun for SUM in Legume Systematics, which funded a month of research at NY. The following

lted for this project: CAS, DS, GH, JEPS, K, MO, NY, OSC, ORE, RSA,

SD, UC, UCR, UCSB, UNIV, US, WILLU. I thank the curators and staff of the following herbaria for access

to their collections or loans of their specimens: BRY, CAS, DS, GH, JEPS, K, NY, OSC, ORE, RENO, RM,

RSA, UC, UNLV, US, WILLU. Arnold Tiehm and one anonymous reviewer provided useful reviews.

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nosus complex of Section Diphysi. Ph.D. dissertation. Oregon State University, Corvallis, Oregon.

Barnesy, R.C. 1944. Pugillus Astragalorum alter. Proc. Calif. Acad. Sci. 4:147-167

Barnesy, R.C. 1945. Pugillus Astragalorum - IV: the section Diplocystium. Leafl. W. Bot. 4:65-147.

Barney, R.C. 1956. Pugillus Astragalorum - XVII: four new species and one variety. Leafl. W. Bot. 8:14-23.

Barnesy, R.C. 1964. Atlas of North American Astragalus. (2 volumes). Mem. New York Bot, Gard. 13:1—1188.

Barney, R.C. 1989. Intermountain flora. Volume III Part B. A. Cronquist, A.H. Holmgren, N.H. Holmgren, J.L. Reveal,

PK. Holmgren, eds. The New York Botanical Garden. Bronx, New York.

Gray, A. 1849. Plantae Fendlerianae Novi-Mexicanae. Mem. Amer. Acad. Arts, Ser. 2, 1:1-116.

Gray, A. 1863. A Revision and arrangement (mainly by the fruit) of the North American species of Astragalus and

Oxytropis. Proc. Amer. Acad. Arts 6:188-237

HARVARD University HErBARIA. 2008. Index of botanical specimens. <http://asaweb.huh.harvard. edu:8080/databases/

specimen, index.html». Accessed April 2002, July 2007, August 2008.

Hooker, WJ. 1831. Flora boreali-americana or the botany of the northern parts of British America. 2(12):97-160.

IseLv, D. 1998. Native and naturalized Leguminosae (Fabaceae) of the United States. Monte L. Bean Life Science

Museum. Provo, Utah.

Jones, M.F. 1898. Contributions to western botany. Contr. W. Bot. 8:1-43.

Jones, M.E. 1923. Revision of North American species of Astragalus. Text distributed Feb. 15, 1923; Plates, June

20, 1923. Salt Lake City, Utah.

KRINGS, A. 2008. Index of names and types in West Indian Gonolobinae (Apocynaceae: Asclepiadoideae), includ-

ing fourteen new lectotypifications, one neotypification, a new name, and a new combination. J. Bot. Res.

Inst. Texas 2:139-163.

McNent, J., ER. Barrie, H.M. Buroet, V. DemouLin, D.L. HawksworTH, K. MARHOLD, D.H. NicoLsoN, J. PRADO, P.C. SiLva, J.E. Skoc,

J.H. Wiersema, AND N.J. TURLAND, eds. 2006. International code of botanical nomenclature (Vienna Code) adopted

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by the Seventeenth International Botanical Congress Vienna, Austria, July 2005. Gantner Verlag, Ruggell,

Liechtenstein. - Regnum Veg. 146:1-568.

MissOURI BOTANICAL GARDEN. 2002. Tropicos. <www.tropicos.org>. Accessed Apr 2002, Jul 2007.

New York BOTANICAL GARDEN. 2008. The CV. Starr Virtual Herbarium. <http://sciweb.nybg.org/ science2/VirtualHer-

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ScHOENER, C.S. 1974, Notes on three varieties of Astragalus lentiginosus (Leguminosae). Great Basin Naturalist

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2002, July 2007, August 2008.

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Accessed August 2008.

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html». Accessed August 2008.

VANDER Kloer, S.P. 1989. Typification of some North American Vaccinium species names. Taxon 38:129-135.

Watson S. 1871. Botany. In: C. King. Report of the geological exploration of the fortieth parallel, Vol. 5. Govern-

ment Printing Office, Washington, DC.

WeLsH, S.L. 1978. Utah flora: Fabaceae (Leguminosae). Great Basin Naturalist 38:225—-367.

WeLsH, S.L. 2007. North American species of Astragalus (Leguminosae): a taxonomic revision. Monte L. Bean Life

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Museum, Provo, Utah.

THE TYPIFICATION OF CROTALARIA ROTUNDIFOLIA

AND CROTALARIA MARITIMA (FABACEAE)

Daniel B. Ward

d of Botany

University of Florida

nia ae 32611, USA.

ABSTRACT

T Fal he American S [ fused i 1 ftl prop l

of C. rotundifolia bou ex J.F. ad and dece el a type ino iti hapm.. A John F ly tł yp

he plant | eau Wier justifying io Fraser specimen

be upereaded by Į from South Carolina. A speci f he typ lity of C. maritima in southern Florida is

here selected as Leno D that Species:

RESUMEN

n xul Tarin (Gal ^ : DEP ene PE. a rS | Jahid la timifi SAGT de (C

rotundifolia Walter ex J.F. Gmel. y al ia d tipo para la C. maritima Chapm io espécimen: de John EHE que es actualmente

] ipop tundifoli ] quí q; ó mas Walter, justificando

jue el espéci le F plazado p ipo apropiado de Carolina del Sur Un T del pes tipo de C. maritima

en el Sur de Florid lecci ] ipo para esta especie.

(Fal ), section Alatae, is represented in t theastern United States by six species (Windler

1974; Ward 2009). Two taxa in this complex—C. s ia Walter ex J.F. Gmel. (1792), and C. maritima

Chapm. (1878)—have been variously interpreted as one undivided species (Isely 1990; Wunderlin & Hansen

2003), one species of two varieties (Windler 1974; Duncan & Kartesz 1981), or two species (Small 1933;

Senn 1939; Ward 2009). Though the variability and occasional intermediacy of individuals of these taxa is

obvious, recognition of their populations as worthy of specific status has been hindered by the absence of

type specimens that adequately represent their morphology.

The cornerstone of stability in the linkage between a plant and its scientific name is the existence of a

single specimen to which each name is permanently fixed. This single specimen—the type—permits later

investigators to know with exactness the form of the plant on which the original author based the new

name. But an author does not always select a type, or the type may become lost through misfortune, or the

type may in one way or another be found to be unrepresentative of the author's original concept. Wherever

ambiguity is caused by a missing or defective type, it is helpful to future studies of the characteristics and

classification of the plant that a type be determined or a defective type be corrected. The International Code

of poe ene Masa et al. 2006) dictates the rules that govern such selection or correction.

C familiar name but a poorly und 1 species. It has been mapped by Windler

(1974, as his var. vu lgaris) as extending from southeastern Virginia, south to central peninsular Florida, and

west to southeastern Louisiana. Crotalaria maritima was mapped by Windler (1974, as his var. rotundifolia)

as ranging from southeastern Georgia (one undocumented dot in South Carolina), to southernmost Florida,

and sparingly west to southeastern Louisiana. Though these ranges greatly overlap and need adjustment

where herbarium materials poorly represent the two entities, Windler's maps well demonstrate that on bal-

ance C. rotundifolia is northern, while C. maritima is significantly more southern

The present task is to establish types that will assist in separation of the two taxa. No effort is made

here to justify this separation; that analysis must await greater understanding of the gross morphology,

chromosome counts, DNA morphotypes, and differences in habitat preferences. Though the more southern

J. Bot. Res. Inst. Texas 3(1): 219 — 225. 2009

rf el D * 4l D hi [PPS

220 J tit f Texas 3(

population, Crotalaria maritima, requires for its typification only selection of a neotype, the more northern

population, C. rotundifolia, has a record of misinterpretation of its type that can be resolved only by a full

review of its taxonomic and nomenclatural history.

REPLACEMENT NEOTYPE FOR CROTALARIA ROTUNDIFOLIA

Crotalaria rotundifolia, the more northern of the two taxa, was the second species (to C. sagittalis L., 1753)

of this group to be recognized. The plant was first described by Thomas Walter (1788), using the name

Anonymous rotundifolia, an impermissible combination (Ward 1962, 2007a; Wilbur 1962). [Walter's use of

Anonymos has been proscribed (Art. 20.4).] Four years later J.F. Gmelin (1792) validated Walter's epithet,

as Crotalaria rotundifolia. Gmelin was merely assigning Walter's plant to a familiar Linnaean genus, Cro-

talaria, and his specific description was a restructured and shortened version of the description provided

by Walter. Crotalaria rotundifolia J.F. Gmel. was thus based on Anonymos rotundifolia Walter and whatever

materials Walter may have seen and used. [To reflect this origin, authorship of the legitimate name is often

cited ^Walter ex Gmelin."]

Walter, a plantation owner in rural South Carolina, with no means of long-term dried-plant preserva-

tion and surrounded by an abundance of fresh materials, kept no specimens that can now be termed types

(Ward 2007b). A folio herbarium at the Natural History Museum, London (BM), gathered by John Fraser,

the Scottish plant explorer, contains a specimen of a small Crotalaria. The folio was photographed in 1946

by B. G. Schubert and the specimen was believed by M. L. Fernald to be Walter's "type" of Anonymos rotun-

difolia (Fernald & Schubert 1948). Later authors (Ward 1962; Wilbur 1962; Windler 1974) accepted this

judgment. Though Fernald used only the term "type," his usage suggests the more precise term “holotype.”

But since (as described below) Walter could not have based his description on this Fraser specimen, the

type designation has been corrected to “neotype,” as authorized by the Code, Art. 9.8 (Ward 20073).

The origin and content of the Fraser folio (best termed the Fraser/Walter herbarium) has been studied

in detail (Ward 2006). Fraser was in the American Southeast only briefly, with opportunity to collect only

during 1787. He gathered plant materials with enthusiasm but with little proficiency; Andre Michaux, who

met and briefly traveled with him, spoke disparagingly of Fraser's botanical skills (Ward 2007b). Even so,

Fraser returned to England with a collection of 690 mostly small, sometimes fragmentary, specimens from

South Carolina and Georgia. Fraser also carried with him Walter’ ipt of the pioneer Flora Caroliniana

(1788), for publication in London.

Before Fraser returned to England, Walter had opportunity to see and perhaps study Fraser's speci-

mens. Attached slips indicate the specimens had not been named when seen by him (though many bore

three-digit numbers assigned by Fraser). Walter's handwriting is now present on 368 of the surviving slips,

with 345 representing his identifications and the remainder being his various comments and observations

(Ward 2007b). [Fraser's hand is borne by only 230 slips, many merely his assignment of a Walter epithet to

a specimen identified by Walter only to genus.]

The specimen that has been believed to be Walter's “type” is now numbered 67-D (Ward 2006). Its

photograph has been published (Fernald & Schubert 1948, plate 1107 - an image described as "quite horrible"

by John Lewis). A similarly poor image is available on commercial microfiches of the entire herbarium, and a

digital image of better quality has been made available by the Museum. [None of these images is sufficient to

show pubescence, a critical feature.] The specimen was examined in the 1890s by Britten & Baker (1897), in

August 1962 by John Lewis (BM) at the request of the present writer, again in the 1970s by Norman Robson

(BM) at the request of Windler (1974: 189), and yet again in July 1984 by the present writer.

Specimen 67-D (BM) is small, but not unrepresentative either of the Fraser specimens or the species

itself, consisting of a stem with six branches, perhaps 25 leaves, two crumpled flowers, and a single pod.

Its leaves are broadly elliptic to ovate or obovate, apically rounded, the larger ones +8 mm. long, +5 mm.

broad.

The label of specimen 67-D is unusually ample, with the plant attached near one end by its stem hav-

Ward, Typification of two Crotalaria species 221

ing been slipped through parallel cuts in the paper (Fraser's common method of affixing labels). The label

is sufficient in size to have received a series of notations extending from (probably) 1787 into the 1960s.

Almost hidden by the stem is a partially obscured (and thus readable only as ??6) three-digit number in

Fraser's hand, believed to be his collection number. The label also reads: “Lupinus affinis" in Walter's well-

formed script; this is one of the 23 specimens in the Fraser collection whose label bears Walter's hand but

was not identified by him. [Specimens 67-A, 67-B, and 67-C, mounted across top of the same folio page, were

labeled in Fraser's hand as Lupinus pilosus (2 spms.) and L. perennis.] Alongside the specimen and writings

by Fraser and by Walter is a more extended notation: “confer / Anonymos sagittalis p. 181 / = A. rotundifolia

Walt. / = Rhynchosia,” in the hand of Asa Gray, who examined the Fraser folio in 1839. Below the Gray nota-

tion a further entry reads: “Crotalaria maritima Chapm. / det. John Lewis 1962." Still another reads: “Anonymos

rotundifolia Walt. / Fl. Carol.: 181 (1788) / SYNTYPE / of / Crotalaria rotundifolia J. F. Gmel. / in Linn., Syst. Nat.,

ed 13 2:1095 (1792)" in an undetermined hand (below and thus subsequent to the 1962 notation of Lewis).

The source of specimen 67-D is unknown. Fraser, unlike Pehr Kalm in the American Northeast and

Andre Michaux throughout eastern America, left no account of his travels. But from occasional remarks on

labels and from specimens obtainable only in clearly defined locations, it is known Fraser reached (among

other more northern and western stations) the Altamaha River in southeastern Georgia (Ward 2006), well

within the distribution of the more southern taxon, Crotalaria maritima.

Pubescence of the stems and leaves has been long recognized to differ between northern populations

typical of Crotalaria rotundifolia and more southern populations assignable to C. maritima. Lewis (pers.

comm., Aug. 1962) directed his analysis primarily to this feature, making comparison with other col-

lections of the two taxa as idertified by Senn (1939) and from locations appropriate to the northern and

southern populations, respectively. In his words: “The critical difference is that the general level of the top

of the indumentum in [C. rotundifolia; Lewis used C. angulata] is equal in height (from the substratum) to

the thickness of the stem, while in C. maritima even the few spreading hairs do not reach this height." He

found the indumentum of the Fraser specimen to be very worn on the more-mature parts of the stem, but

even here "the sparse remnant is wholly appressed." Further, he found the leaf shape *not inconsistent" with

that of C. maritima. Lewis' conciusion was that Senn would have placed the Fraser specimen in C. maritima

if he had seen it. The present writer, with opportunity in 1984 to study the Fraser specimen directly, fully

agreed with Lewis' conclusions.

Windler (1971, 1974) addressed the judgment of Fernald & Schubert (1948), as well as information

given him by Robson and his own examination of a better-quality photo from the British Museum, and his

understanding of the approximate ranges of these two taxa. In consideration of the variable and intergrading

morphologies of the two populations, he chose to recognize the taxa at varietal level. He was compelled to

conclude the “Walter” type was of the more southern population, which obligated him to base C. rotundifolia

var. rotundifolia on Fraser's specimen. He then took the next logical step, by selecting a specimen from the

northern population to represent the non-typical variant. He proposed the new name C. rotundifolia var.

vulgaris Windler, and typified it with a specimen from Hampton County, South Carolina.

Windler’s acceptance of the Fraser specimen as typical of the southern population and a specimen

from the northern population as representing the non-typical variant had the unfortunate consequence of

putting in place as type of C. rotundifolia a specimen that Walter, the author of that name, would not have

recognized. Indeed, Walter (though he died in 1789) has himself had opportunity to speak to this issue. He

annotated specimen 67-D as “Lupinus affinis,” or “allied to Lupinus,” a comment similar to his many Latin-

based remarks throughout the folio (Ward 2007b). This notation well indicates he did not recognize the

species, though by later nomenclatural fiat he has now been claimed its author!

Windler's use of varietal status has suppressed the incongruity of a Walter name being applied to a

population of which its author had no direct knowledge. Thus persons who interpret the complex as undi-

a we see no issue, and those who use only varieties will scarcely be alerted. But, in the belief the two

it specific rank (Ward 2009), the present writer has long been intrigued of a proper solution.

I as

222 tani i Texas 3(

The Code (McNeill et al. 2006) permits any name to be retained by the process of "conservation" (Art.

14). Though there are several subtleties, the usual process would involve a petition to change the type by

which a name was determined, thereby changing the name itself. This petition must be of a form acceptable

for publication in a specific international journal (Taxon). The petition would then be reviewed by a special

committee for the appropriate taxonomic group, then by a senior committee, then by a future botanical

congress. Disapproval at any level would negate the proposal. Here, a proposal might ask that the type of

Crotalaria rotundifolia Walter ex J.F. Gmel. be replaced by a specimen more appropriate than the one provided

by John Fraser.

A less complex and uncertain pathway exists by which the issue may be resolved. The Code, Art. 9.16,

states: ^A neotype selected under Art. 9.15 [the article permitting selection of a neotype] may be superceded

if it can be shown to differ taxonomically from the holotype or lectotype that it replaced." This power does

not exist if a holotype or lectotype is involved, for in each of those categories a specimen seen and used by

the author would have priority. But a neotype, a replacement selected at a later date by another person, can

possibly be an inappropriate choice as a replacement for the missing holotype.

Rejection of the Fraser specimen as the type of Crotalaria rotundifolia quite exactly conforms to the

requirements of Art. 9.16. All investigators who have examined specimen 67-D or its images, even though

in part working independently and without full knowledge of the work of others, have agreed this specimen

matches the thern population. Even its author, Thomas Walter, by his annotation has left testimony

of his endorsement. Though it is permissible for an author to consider the variability within the group too

slight to permit naming, most authors have recognized the two populations at some taxonomic rank. And

there is no dispute that the one taxon is common in the area known to Walter, and the other taxon to be es-

sentially absent. It thus follows that the specimen previously treated as the neotype of Crotalaria rotundifolia

is taxonomically different from the specimen that Walter must have once held and used as the basis for his

new name.

Since the conditions for invoking Art. 9.16 are fully met, it also follows that the Fraser specimen treated

as neotype of Crotalaria rotundifolia may be superceded by a more appropriate specimen.

Crotalaria rotundifolia J.F Gmel., Syst. Nat. 2:1095. 1792. Tez: U.S.A. South CaroLiNa. Hampton Co.: ca. 3 mi NW of

Yemassee on SC Hwy 68, sandhill, 23 Jul 1967, D.R. & B.K. Windler 2769 (neorvee, designated here: NCU). This selection supersedes

the previous selection of J. Fraser 67-D, 1787 (BM) as neotype, as authorized by the Code, Art. 9.16.

The selected specimen is also the holotype chosen by Windler (1974: 193) for Crotalaria rotundifolia var.

vulgaris. Windler's judgment is accepted in his selection of a specimen appropriate for the northern popula-

tion. Although typification does not apply outside of rank, the use here of the same specimen as the type of

C. rotundifolia obviates all uncertainty as to application of the name without regard for the rank employed.

NEOTYPE FOR CROTALARIA MARITIMA

Crotalaria maritima, the more southern of the two species, is somewhat more variable than its northern

analogue, in that the leaves of some plants may range from ovate to linear. This variability has troubled

investigators (Windler 1974; Isely 1990), but has then been put aside as a poorly understood property of

the southern population. Other workers have separated the more clear-cut linear-leaved plants as C. linaria

Small (1933) or C. maritima var. linaria (Small) Senn (1939). Though there is unquestioned intergradation,

this linear-leaved population is also separated edaphically along the South Florida coastal dunes, and merits

specific rank (Ward 2009).

With Crotalaria linaria removed, the remaining plants of the southern population are appreciably more

uniform. A rare, recently described endemic, C. avonensis Delaney & Wunderlin (1989), with succulent

leaves and loosely appressed pubescence, from near the southern end of the Lake Wales Ridge in central

peninsular Florida, appears to be a local derivative. An erect species with leaves glabrous above, C. purshii

DC., is known to hybridize with the southern population (Windler 1974: 202). An erect, usually annual

Ward, Typification of two Crotalaria species

berbacuam ot rae Vins

Agnovitwai xp:

joy, Cane

s DOMI ..

3558

iversity of Flor la

sime Station

Plants of Florida

L FL

CIN 4 £ I

F Ff. H TH

of Chapman 1878, 1883), Cape Sable, E glad Park, Monroe County, Florida. Bulls y le 23 cm diameter.

Bot. Gaz. 3:4. 1878.F. { ly swale behind | tal shell ridg

223

224 tani i Texas 3(

species, C. sagittalis L., may hybridize with the northern plants here called C. rotundifolia (Windler 1974:

202), but appears not to overlap in range with the southern population.

The earliest name for the southern population is Crotalaria maritima Chapm. (1878, 1883). Chapman

described his plant with elements unique to the southern population: “Stem low, much branched, pubes-

cent with short appressed hairs; leaves simple, small, oblong, sub-sessile, very thick and succulent; stipules

minute and narrowly decurrent, or none; raceme 2-flowered; legume smooth; ... Stem 6 in. high. Leaves 1

in. long. Flowers not seen" (slightly rephrased in 1883). He gave the place of collection, perhaps well known

to Florida travelers in the 19th century, though now far from obvious, as "Sandy beach at Palm Cape, South

Florida." His notation of "Flowers not seen" suggests the plant was not overly familiar to him. Windler (1974)

did not trace this station; he noted Chapman’s type only as “not located."

Chapman's home was in Apalachicola, in the central Florida panhandle. At least once he is known to

have visited another early botanist, Dr. John Blodgett of Key West (who died in 1853). He made this journey

by taking a small boat along the western Florida coast, stopping and apparently collecting along the way.

He may have done so more than once, at least for part of the distance, in that along with the new Crotalaria

he reported many other plants newly discovered in South Florida (Chapman 1878). The careful account of

Florida botanical exploration by R.P. Wunderlin, B.F. Hansen & J. Beckner (in Wunderlin & Hansen 2000)

reported no documentation of these journeys.

Old late-19th century maps of Florida, however, do record *Palm Cape" as the middle point of Cape

Sable, on the southwestern tip of peninsular Florida. The Palm Cape beach is an obvious stopping-point for

shallow-draft vessels moving along the southern shore. This location is now within the Everglades National

Park, Monroe County, though it is still quite as remote—and perhaps less visited—as in Chapman's day.

In Sd 1964 "e writer, pee by a graduate student and two rangers, was able to reach the Cape.

was immediately obvious, broadly spreading in a dry sandy depression behind the front

beach. Specimens were collected, and one has been selected as typical of C. maritima Chapm..

Crotalaria maritima Chapm., Bot. Gaz. 3:4. 1878. (Fig. 1). Tre: U.S.A. Fiona. Monroe Co.: Middle Cape (“Palm Cape"

hapman 1878), a Sable, Everglades National Park, open dry sandy swale behind low coastal shell ridge, 18 Apr 1964, D.B.

Ward 3939 ( potype), designated here: FLAS)

With recognition of the types selected here, Crotalaria rotundifolia Walter ex J.F. Gmel. again becomes the

round-leaved, non-succulent, spreading-pubescent plant of the Carolina coastal plain and northern Florida,

and Crotalaria maritima Chapm. again is recognized as the ovate-leaved, succulent, appressed-pubescent

plant of peninsular Florida. These points of nomenclatural stability will serve as clearly understood data-

points for further work addressing the morphological subtleties of these and related taxa.

ACKNOWLEDGMENTS

My involvement with these little “rattleboxes” has extended intermittently over many years. I am grateful to

Jacqueline Patman (Mrs. Malchie Broughton), University of South Florida, who briefly joined me in study of

these plants, terminated too quickly by her movement westward; to Donald Windler, Towson State University,

Maryland, for keeping me abreast of his more detailed studies and remaining a friend to the present day; to

Derek Burch, Plantation, once my student and field companion and permanently my wise advisor; to John

Lewis, British Museum of Natural History, for answering in such wonderful detail (most not reported here)

my queries on the weary specimens of John Fraser; to Robert L. Wilbur, Duke University, for counsel on

matters of nomenclature; to Christine M. Housel for the Spanish translation of the Abstract; to Kent Perkins,

University of Florida, for the plate of Crotalaria maritima; and to Thomas G. Lammers and James L. Reveal

for meticulous and insightful suggestions in review of my manuscript.

REFERENCES

BRITTEN, J. AND E. Baker. 1897. Houstoun's Central American Leguminosae. J. Bot. 35:241-425.

CHAPMAN, A.W. 1878. An enumeration of some plants—chiefly from the semi-tropical regions of Florida—which

Ward, Typification of two Crotalaria species 225

are either new, or which have not hitherto been recorded been recorded as belonging to the flora of the

Southern States, Bot. Gaz. 3:2-6, 9-12, 17-21.

CHAPMAN, A.W. 1883. Flora of the southern United States, ed. 2. New York.

DeLaney, K.R. AND R.P WUNDERLIN. 1989. A new species of Crotalaria (Fabaceae) from the Florida central ridge. Sida

13:315-324.

Duncan, W.H. AND J.T. Kanresz. 1981. Vascular flora of Georgia: an annotated checklist. Univ. of Georgia Press,

Athens.

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.

GMEUN, J.F. 1792. Systema naturae. Leipzig.

Isei v, D. 1990. Vascular flora of the southeastern United States. Vol. 3, part 2: Leguminosae (Fabaceae). Univ. N.

Carolina Press, Chapel Hill.

LINNAEUS, C. 1753. Species plantarum. Stockholm.

McNett, J., ER. Barrie, H.M. Buroer, V. Demoutin, D.L. HAwkswonrH, K. MARHOLD, D.H. NicoLsow, J. PRADO, PC. Sit vA, J.E. SkOG,

NJ. TURLAND, AND J. Wiersema, eds. 2006. The International code of botanical nomenclature (Vienna Code), July

2005. Regnum Veg. 146:1—568.

Senn, H.A. 1939. The North American species of Crotalaria. Rhodora 41:317-366.

Smal, J.K. 1933. Manual of the southeastern flora. New York.

WALTER, T. 1788. Flora Caroliniana. London.

Warp, D.B. 1962. The genus Anonymos and its nomenclatural survivors. Rhodora 64:87-92.

Warp, D.B. 2006. Thomas Walter typification project, |. Observations on the John Fraser folio. Sida 22:

1111-1118.

Warp, D.B. 2007a. Thomas Walter typification project, Il. The known Walter types. J. Bot. Res. Inst. Texas 1:

407-423.

Waro, D.B. 2007b. The Thomas Walter Herbarium is not the herbarium of Thomas Walter. Taxon 56:917-926.

Warp, D.B. 2009. Keys to the Flora of Florida—24, Crotalaria (Leguminosae). Phytologia (in press).

WILBUR, R.L. 1962. The identity of Walter's species of Anonymos. J. Elisha Mitchell Sci. Soc. 78: 125-132.

WiNDLER, D.R. 1971. New North American unifoliolate Crotalaria taxa. Phytologia 21:257—266.

WiNpLER, DR. 1974. A systematic treatment of the native unifoliolate Crotalarias of Noth America (Leguminosae).

Rhodora 76:151—204.

WUNDERLIN, R.P. AND B.F. HANSEN. 2000. Flora of Florida. Vol. 1, Pteridophytes and Gymnosperms. Univ. Press of

Florida, Gainesville.

WUNDERLIN, R.P. AND B.F. Hansen. 2003. Guide to the vascular plants of Florida, 2nd ed. Univ. Press of Florida, Gainesville.

] | af al D , ID hi PO £ T.

226 t Texas 3(

BOOK REVIEW

LAWRENCE E. STEVENS AND Vicky J. Meretsky (eds). 2008. Aridland Springs in North America: Ecology and

Conservation. (Arizona-Sonora Desert Museum Studies in Natural History). (ISBN 978-0816526451,

hbk.). University of Arizona Press, 355 S. Euclid Ave., Suite 103, Tucson, Arizona 85719, U.S.A. (Orders:

www.uapress.arizona.edu, orders@uapress.arizona.edu, 1-800-426-3797). $75.00, 432 pp., 6" x 9".

t oft ia, one at the Arizona-Sonora Desert Museum in 2000 and the other at the Ecological Society of

e 3 1 i A 1 T f. 1 d

This volume grew posia,

erica meetings in 2002. The book rep d levelop ym[

i that followed.

The book begins with a series of S ies cda e issues of springs ecosystems, bad ie Hers): a Ene

springs. This is ao by

classification system

D and 8 tati ] 8 SE i g; i a ce Mexico: and p alm d frheQq +) P RN

1 i i lso di tl th Colorado Plateau, Zi d Grand Canyon

s e i e * e i O

A sai 3 iem f ; ` ; nel J; 1 Je LT ivit : ; us :

National Parks I y of spring I lj Į yand Springe been fae

3 £f : 1 € RE 1 Aricana TY 1

o rr oOo

this secti as they ] li d imp | f spring y i ity “This i is followed by légal options for

1 1 f Tal] 1 1 1 1 zum | A ic ].

p pring; g y g water. The conc ss

1 1 7 1 th SE Ta f Iç, A ER £ A TA

Esa) 111 ULLE F -F X 5 PE

conservation d restorat f 11 he chall f 1 bl i 1 d restorat f

r O O O O jo] Y Eu o Es e

ecosystems.

MN hook will ar to a wid li including the i 1 publi ] ] Il levels. Recommended for general

lil h level collections

library

Lawrence E. Stevens is curator of ecology and conservation at the Museum of Northern Arizona and the senior science advisor

for the Grand Canyon Wildlands Council, Inc., in Flagstaff. Vicky J. Meretsky is an associate professor of conservation biology at the

School of Public and Environmental Affairs at Indiana University, Bloomington.—Gary L. Jennings, Librarian, Botanical Research Institute

of Texas, Fort Worth, Texas 76102-4025, U.S.A.

Contents;

Prologue, by Gary P. Nabhan

1. Springs Ecosystem Ecology and Conservation, by Lawrence E. Stevens and Vicky J. Meretsky.

2. The Demise of Desert pups by Peter J. Unmach and W. L. Minch

3. nii dici fD prings in North America, by ur Kreamer and si Viet od

1 wpe l, and Ecological Criteria, by Abraham

E. pias Laven E Stevens ee pees Roderic A. Parnelll, David K. NN Lisa Levin, and Stephen P. Flora.

5. Edo amis Cauld gical Archives Ds Vance So.

P3 O

i be E G LE NE | TN E x NT XA7ATII ry

g Well, Arizona,

F + P

i Dean W. Blinn.

7. Combining Ecological Research and Conservation: A Case Study in Cuatro Ciénegas, Coahuila, México, by Dean A. Hendrickson,

Jane C. Marhs, Angie Moline, Eric Dinger, and Adam Cohen.

8. The Desert Fan Palm Casis, by James W. Corn

9. Spring-Supported "OE along the un River, Colorado Plateau: Floristics, Vegetation Structure and Environment, by John

R. did

10. ^ f Change in S Pl iti he Soutl Colorado Plateau, by Vicky J. Meretsky.

11. Email and Productivity at an Undisnibed Spring in Comparison with Adjacent Grazed Riparian and Upland Habitats, by

nca S. Perla and Lawrence E. Ste

12. aa Diversity Influenced by noe Management of Freshwater Springs: Flora of Quitovac, Sonora, Mexico, by Gary Paul

Nabhan.

13. Historic and Prehistoric Ethnobiology of D S , by Amadeo M. Rea

14. Hong Dynamics of Great Basin Springs: Potential Effects of Groundwater Withdrawal, by Duncan Patten, Leigh Rouse, Juliet

mberg.

15. i Knowles Canyon e Garden, Glen Canyon National Recreation Area, Eight Years after Burning: Changes in Vegetation

and Soil Biota, by Tim B

16. Between the Cracks: Water Law em Spring Conservation in Arizona, by Nancy Nelson.

17. Epilogue: After the Last Drop, by Lawrence E. Stevens

J. Bot. Res. Inst. Texas 3(1): 226. 2009

LECTOTYPIFICATION OF CARDAMINE FLEXUOSA (BRASSICACEAE)

Angela R. Post' Alexander Krings

Department of Horticultural Science Herbarium, Department of Plant Biology

North Carolina State University North Carolina State University

Raleigh North Carolina 27695-7609, U.S.A. Raleigh North Carolina 27695-7612, U.S.A.

Qui-Yun (Jenny) Xiang Bryon R. Sosinski

Department of Plant Biology Department of Horticultural Science

North Carolina State University North Carolina State University

Raleigh North Carolina 27695-7612, U.S.A. Raleigh North Carolina 27695-7609, U.S.A.

Joseph C. Neal

Department of Horticultural Science

North Carolina State University

Raleigh North Carolina 27695-7609, U.S.A.

In 1796, William Withering (1741-1799) described a new species of Brassicaceae from England as Cardamine

flexuosa With. The primary set of Withering’s collections is held at BM (Stafleu & Cowan 1976), but upon

inspection we found no specimens at BM for any taxa of Cardamine L. bearing his name or hand. There are

also none of his own Cardamine specimens or those collected by others that he may have studied in other

herbaria also known to house his material: BR, K, LINN, LIV, UPS-Thunb (Stafleu & Cowan 1976). Loss

of type material for C. flexuosa With. has been previously reported (Marhold 1995, Lihova et al. 2006);

however, a lectotype for C. flexuosa With. remains undesignated. In the absence of the holotype we refer to

illustrations cited in the protologue to lectotypify Cardamine flexuosa. Withering cited the following three

illustrations which serve as syntypes in the absence of other original material: t. 277 (Curtis 1777); t. 735

(Oeder 1770); C. hirsuta (Walcott 1778).

Here we designate one of these illustrations as the lectotype for Cardamine flexuosa With.:

Cardamine flexuosa With., Arr. Brit. Pl. ed. 3:578. 1796. (Fig. 1). Tree: “Rookery at Edgebaston” (Lecroryre, designated

here: Curtis t. 277, 1777).

The lectotype (Fig. 1) is a historical illustration from Curtis’ Flora Londonensis (1777). This work is a collection

of loose or bound plates depending on the copy, and some copies have been hand colored. There are both

colored and black and white examples of plate 277. The plate is accompanied by a brief but thorough Latin

description and an English translation (Fig. 2). Plate 277 is labeled as the closely related species C. hirsuta,

but clearly illustrates the siliques angled away from the inflorescence, flexing nodes, and a densely hairy

stem more typical of Cardamine flexuosa With. This plate undoubtedly refers to the taxon in Withering's

original description of the species. Basal leaflets in the illustration are not totally congruent with what we

have observed in the field and on herbarium specimens for the taxon, but the description accompanying the

plate reads: “Lobes of the radical leaves vary much in shape and are frequently much rounder than the figure

represents.” The only other caveat to our determination here is that a detailed illustration of the androecium

in the bottom right corner of the plate (labeled #3) cannot refer to C. flexuosa because it illustrates only

four stamens when C. flexuosa always has six. Information given in the description accompanying the plate

sheds some light on this problem. The main morphological difference between C. hirsuta and C. flexuosa is

stamen number, C. hirsuta having four and C. flexuosa six. Curtis’ illustration shows four stamens and gives

the following information in the description: “The same plant, early in the spring, when the weather is cold,

has only four stamina; as the advances, it has constantly six.” We believe that Curtis was encounter-

P ldress: D f Plant Biology, Cornell University, Ithaca, NY 14853, arp232@cornell.edu

J. Bot. Res. Inst. Texas 3(1): 227 — 230. 2009

228 Journal of the Botanical R h Institute of Texas 3(1)

Ec 1 Diaitali £ika lartat £o» fest H fi fa AI f. "PI I I: . Di44

iG Curtis

£1 4 t 377 IC 4 y £ M: ? D nl N P:

Post et al l

CARDAMINE HinsuTA. Harry LADIES-SMOCK.

CARDAMINE Lin, Gen. PL Terranrranta Siurerosa.

Siligua claflice diflliens valvulis revolutis- Siigme integrum. Cel,

fübhians.

Rati Syn. Got, 22. HERBE TETRAPETALA gatans Z ET SILICULOS 5.

CARDAMINE oe puse floribus tetrandris. Lin. Syf. Vegelab. p. 497. Sp. Pl. p- pis FI.

tele tie 587.

CARDAMINE foliis pinnatis hirluuix, pinnis fubrotundis, ftaminibus quaternis, Moder Mil. 472.

CARDAMINE firfute, Scopoli FI. Cora, n. 817. 1. 98.

NASTURTIUM uquaticum minus, Baud. Pía, 104.

CARDAMINE ime altera hirfutior. Rati Syn. p. 300. Tho leflor Hairy, impatient Cuckow-

or Ladics-Smock.

venum ULIS i Pit "rg ge e

RADIX annua, fibrofa, fibris albidis. ROOT annual and fibrous, the fibres whkifh.

CAULIS fj pie et ultra pro rationc loci, in 3 STALK about a d high, or more, according to

ho flis h midis reperitur etiam fefquipedalis, : e fituation ch it grows; in wet ditches

: m j

ofiffimus c

FOLIA radicalia plurima, in orbem polita, pinnata, |

bucal pri vn Hw ndato angulatis, ple-

hirfutis, pats $ round yet angular, ni div idea

emet E tada lobi; er equae into des lobes, hirfate, roug

x Cu nentis, cauli prominent points, the lobes une

gis profundo incila, lobis i times Pa im fomctimes pointed ; mE rol

des p e inis à and more deeply ig-

FLORES parvi, albi, primo vore tantum. tetrandri, FLOWERS pi ge white, eariy in the {pring

CALYX: Peaianrn 19M ee ies n | CAREER bi Po Rian TRUN of four leaves, of an

ees obt vis, dociduis, pilis 3 objeng fanpe, 9 , Obtufe, hollow, decida-

i p a few white hairs, He AF TS.

corona: Peraza quatuor, "alba, calyce dopio COROLLA: per i PsTALS, ais

ha ongiora, patentia, integerrimo, ob» length T the calyx, fpreading,

V5 obtufe, fig. s.

STAMINA: E senta plerumque fox, quoram SP: a molt part fix DEFINEN of

m bi ora, alba, Antasns minima, $ two are fhorter than the refi, of a

joek, As. 3 vw e E ke ÁNTHERE Ea ftoall and

ERREEN GRANEN peage, ea sTM Ganten oblong, flender, a little

ulo brevior, S A capitatum, fis. 4. one sé a Sriora forming

a sad, fig.

PERICARPIUM : Sigua erca, uncialis, fubcom- ¿ SEED-VESSEL : on upright Pod, about an inch in

refla, jer lattice diffilicns, valvulis length, fomcwhat flattened, of two valve»,

A revolutis, Az. 5. which burft with an elaflic force, end roll

: back, fr.

SEMINA duodecim circiter, fuborbiculata, comprefla, Y SEEDS rud A. i in number, nearly round and

glabra, e flavo fufca, fc. 6. RU meee Ímooth, and of a ycllowifh-browa

colour, fiz

‘We were inclined to believe with our ingenious friend Mr. Licmrr oor, that the Cardamine hirfute and

parviflora were difint poe but repeated obforvation and culture have convinced us, p they are both

the fame, varying only in Sze, in hairinels, and in the number of the pad

, In wet htuations, where E foil is luxuriant, it a foot or two in t, and lofcs in a great degree

its hairinele ; in shin gel. lere it feldom reaches iHa then fix or cight dud p is ally uch s hp

= airy: and, w Capiro nee more branched. The aa E early i r dep {pring

weather is Js iH Hid four ma os as the fummer advances, it has conftantly fi x, The bur of pol

radical leaves vary much in fhape, and are frequently m neni as the figure tc epee

This fpecics is b es ire spar ag ounds in particular places ; as by Cf water-

works, g wet ditches about Mamplicad, Highgate, and clícwhere, 5 V ga

Tt flowers in Apri and May. In the garden, y, the fituation in which it is fowa be deci and the fafon

not pacem dry, it continucs flowering and feeding during the whole of the fu

According to Mr. LiogrrooT, the leaves arc a good ingredient a and may eafily be

obtained in the fpring, when Muftard and Greh are not to be had, j oar

229

Fic. 2 Digit |

230 J | ical h Insti Texas 3(

ing two different taxa during the course of the growing season. Our personal observations suggest that C.

hirsuta has a winter annual life cycle and is only encountered in flower during the cooler months of spring.

Cardamine flexuosa, on the other hand, does not have such a set life cycle in the landscape and can be seen

flowering later in the year. We thus suggest that in the early Spring, Curtis was likely observing C. hirsuta

with four stamens and later, when he observed six stamens, he almost certainly was observing C. flexuosa

which is only subtly cd in eRe morphology compared with C. hirsuta.

The two remaining il t ited in the protologue (Walc. C. hirsuta and Fl. Dan.735) are poor rep-

resentations of the taxon in question. The Walcott illustration labeled C. hirsuta (1778) depicts a plant with

six stamens in the flower which could refer to C. flexuosa; however, a very straight stem and four stamens in

the detailed drawing of the androecium suggest C. hirsuta rather than C. flexuosa. Table 735 (Oeder 1770)

labeled C. impatiens exhibits a flexing stem, but the stem and rachis are very thick compared to C. flexuosa

and there are numerous cauline leaves with leaflet shapes more typical of C. impatiens, though Kuéera et al.

(2006) excluded this as an accurate depiction of C. impatiens citing the lack of auriculate leaf bases. Com-

pared to this illustration, Cardamine flexuosa has sparser cauline leaves which exhibit fewer leaflets per leaf

than the basal leaves.

ACKNOWLEDGMENTS

We would like to thank the curators of BM, K, and LINN for the courtesies extended during herbarium

visits, the curators at BR, LIV, and UPS-Thunb. for their assistance in searching for holotype material of this

taxon, and those at MO for providing high resolution images of the designated lectotype. Reviews of Jeffery

M. Saarela and Karol Marhold are greatly appreciated.

REFERENCES

Curtis, W. 1777. Flora Londinensis or, plates and descriptions of such plants as grow wild in the environs of

London. t. 277. Britain.

Liniova, J., K. MarHoLp, H. Kupon, AND M. Koch. 2006. Worldwide phylogeny and biogeography of Cardamine flexuosa

(Brassicaceae) and its relatives. Amer. J. Bot. 93:1206-1221.

Kucera, J., J. Linová, AND K. MARHOLD. 2006: Taxonomy and phylogeography of Cardamine impatiens and C. pectinata

(Brassicaceae). Bot. J. Linn. Soc. 152:169-195.

MarHoLD, K. 1995. Taxonomy of the genus Cardamine L. (Cruciferae) in the Carpathians and Pannonia. III. Folia

Geobot. Phytotax. 30:397—434.

Ocper, G.C. 1770. Enumeratio plantarum florae Danicae. t. 735, Hafniae, Sumptibus Heineck & Faber.

STAFLEU, F.A. AND R.S.CowAN. 1976. Taxonomic literature: a selective guide to botanical publications and collections

with dates commentaries and types Vol. Il. Bohn, Scheltema, and Holkema, Utrecht.

Watcotr, J. 1778. Flora Brittanica Indigena: or plates of the indigenous plants of Great Britain. S. Hazard, Bath.

WirHeRING, W. 1796. Arrangement of British plants. Ed. 3 Swinney and Walker, Birmingham. P. 578

AN INVESTIGATION OF MORPHOLOGICAL EVIDENCE SUPPORTS

THE RESURRECTION OF PYRROCOMA SCABERULA

(ASTERACEAE: ASTEREAE)

Curtis R. Bjórk Mark Darrach

Stillinger Herbarium Corydalis Consulting

University of Idaho 20762 Hemlock St. NE

Moscow, Idaho, U.S.A. 83854, U.S.A. Indianola, Washington 98342, U.S.A.

crbjorkegmail.com corydalis_mark@earthlink.net

ABSTRACT

Dou data were pro > ild populati [P liatriformi lato (A Astereae). These are measures of degree

tomentosit y gland y. f1 linfl ] hes, number, length and width of flower heads, width of phyllar-

ies, and y zd wp Tolo 1 f fl 2 £1 Tis T 1 a 1 e Lg T GO fih 1 7

A in e bera d 5 P. Te sensu lana: ] lentifiabl pholog p gto geographical

A Sener 8 hand and g lands of the Snake River Canyons/Camas Prairi gi the oth

T pl fiha] geog pl rcd g 1 y 1 yp Fp y la. hitl y ized ler P. liatriformis

ih igi 1 I Li: ; EE in 1909. Tk greg xs fr I lati 1 ted P. L lal P. liatriformis

trict 1 d ] d i C i i li i f thi aiv discussed.

RESUMEN

Cc 1 P A 31 I 41 1 j 1 D- 1. if 1 Astereae) E ms J I 1:4

pa] lg "i J a EXAMS | glandulosidad, 1 1 1 Ja] fl a J dal 1 la He Fl

J 1 £51 1 J 1 1 = = J T f1 1 A 1 1 1

revelaron falta d if idad en la morfología de P. liatriformi lato, y también d fologias identificabl li

a ue ara división Beegiance Los pastizales de Palouse y los pastizales del cañón Snake River Pradera de Camas. Las olaia s de

I o Pyrrocoma scaberula, previamente bajo el sinónimo de P. liatriformis desde

la publicación S cad de est 1909. La segregación de esta a con e resurgida P. n EH a la ue P.

iscuten |

Key Wonps: Pyrrocoma, Asteraceae, Astereae, Palouse, Snake River Canyon, Camas Prairie, principle components analysis

INTRODUCTION

Members of the genus Pyrrocoma Hooker (Asteraceae: Astereae) are taprooted, herbaceous perennials with

basal rosettes and leafy stems topped by one to several flower heads with yellow ray florets. The genus is

endemic to western North America, and includes 26 taxa as circumscribed in the most recent treatment

(Bogler 2006). Between Hall's monograph of the genus in 1928 and Mayes' monograph in 1976, Pyrrocoma

was generally merged along with numerous other genera under Haplopappus. Recent morphological and

molecular work suggests that such a broad use of Haplopappus results in a polyphyletic assemblage (Clark

et al. 1980; Morgan 1992).

Edward L. Greene presented the names Pyrrocoma liatriformis and Pyrrocoma scaberula in a single

publication in 1909, and subsequently, the latter has been treated as a synonym of the former (Hall 1928,

Mayes 1976, Hitchcock & Cronquist 1973). Pyrrocoma scaberula was represented in its type collection by

poor material, and until the 1990's, further collections from populations in the region of the type locality

were unknown. Upon the availability of these specimens, it became apparent that plants from the region of

P. scaberula consistently share many of the morphological traits described for and apparent in the type of P.

scaberula. These traits appear to make them morphologically and geographically separable from typical P.

liatriformis.

J. Bot. Res. Inst. Texas 3(1): 231 — 238. 2009

232 i i Texas 3(

Pyrrocoma liatriformis sensu lato is currently placed on the United States Fish and Wildlife Service's

Species of Concern list. At the time of its first listing as a conservation priority, the species was thought to

be found primarily in the Palouse grasslands, but in recent years, numerous large populations of the species

in its broad sense have been located in the canyon puce in ind vicinity the pa Clearwater and

Salmon Rivers (Mancuso 1997). These newly found pop istics with

the type of P. scaberula, which was also collected in this more southerly on (Hear the town of Forest, Nez

Perce County, Idaho).

The present study aims to test 1) the morphological coherence of populations of Pyrrocoma liatriformis

sensu lato, 2) whether any distinct morphologies correlate to geographical range, 3) whether all populations

can be unambiguously assigned to either of the hypothesized morphological entities, and 4) whether a por-

tion of the populations are comparable to the type of P. scaberula and so can be assigned that name.

METHODS

Plants of both of the P mn S were studied in the field in 31 populations. Seventeen

populations in Whitman County, Washington, and Latah and northern

Nez Perce Counties, a Fourteen popalio were sampled in grasslands of the Snake River Canyon

system and adjacent Camas Prairie in Asotin County, Washington, southern Nez Perce, Idaho and Lewis

Counties, Idaho. These 31 populations constitute more than one third of the 91 known extant occurrences

(K. Gray, pers. com., Kemper 2005) from throughout most of the known range of Pyrrocoma liatriformis

sensu lato. In total, 322 plants were studied in the field, 201 of them from the Palouse grasslands, and 124

from the E d Prairie a

] populations were studied based on herbari i y. Both of these populations

were newly discovered in 2005 by other i and were not seen by the antoje until after the conclu-

sion of field work for the present study. These sp were included in the study because they represent

populations in areas where Pyrrocoma liatriformis was previously not known to grow (one specimen from

Wallowa County, Oregon, and two specimens from Washington County, Idaho, both located far south of

the Palouse grasslands).

Plants in the field surveys were chosen randomly on intervals along walkabouts following contours

of the topology so as to sample plants on any occupied slopes, aspects and soil/moisture conditions. Eight

variable morphological characters d based on the hypothesis that they are inf tive of genetic

lineage and hence ic placement. Two of these characters are qualitative: 1) pubescence (tomentose,

non-tomentose or intermediate), 2) glandularity (strongly, weakly or intermediate). Six characters used

are quantitative: 3) number of lateral inflorescence branches longer than 1 cm, 4) number of flower heads,

5) length of flower heads (from the base of the lowest closely ensheathing phyllaries up to the tips of the

highest-positioned phyllaries; three heads, the lowest highest and a middle one were measured), 6) width of

flower heads (non-pressed; same heads measured for length), 7) width of the phyllaries (measured among

the second or third ranks of phyllaries on the same heads as measured for head length and width), and 8)

width of the lowest leaf of the inflorescence. All quantitative characters appear not to be interdependent,

with no correlation between them having an r? value higher than 0.463 (head width with phyllary width).

From large populations, voucher specimens (one fertile stem each) were gathered for every sampled

plant. Plants from small populations were vouchered nondestructively by use of digital photographs, taken

with a ruler for scale. Photos will be archived in both digital and printed format at the Stillinger Herbarium,

University of Idaho, where voucher specimens will be deposited as well.

Quantitative morphological characters were entered as continuous variables in principle components

analysis (PCA) run on PCORD (MjM Software Design). Characters 3 and 4 were discarded from the PCA

owing to the large number of aborted branches and heads, which was perhaps due to the severe drought

that occurred throughout the study area in the year of the field season. Correlations of characters with the

first and second axes were sought to determine which characters contribute most to any clustering pat-

Bjórk and Darrach, R tian af P basil 233

terns. Points in the PCA graphs were matched to their geographical range, pubescence type, and degree of

glandularity to visually seek any correlations between the discrete and quantitative variables.

Means and standard errors of morphological characters and hypothesis tests were generated using

JMP Version 3.2.6 for Windows (SAS Institute). Correlated discontinuities were sought using the ranges

and means of both individual plant data and population means arrayed by geographical range, pubescence

type and degree of glandularity.

High-resolution images were examined of the type specimens of Pyrrocoma liatriformis, Pyrrocoma scab-

erula, and Pyrrocoma suksdorfti E. Greene, which is the third name synonymized under P. liatriformis in all

treatments since the publication of all three names by E. Greene. All three are housed at the U.S. National

Herbarium, and may be viewed at the type specimen regi ster at http://ravenel.si.edu/botany/types.

RESULTS

All eight morphological characters reveal non-uniformity within Pyrrocoma liatriformis sensu lato (Table

1). Within- and among-population variation is seen in all quantitative variables, but pubescence type and

degree of glandularity were strongly uniform E populations.

The principle components analysi clusters of populations, separated along axis one, which ac-

counted for 81.88% of the variation. Axes two through four had broken-stick Figenvalues higher than their

actual Eigenvalues, and are thus not further considered. Correlation coefficients were greatest with axis one

for phyllary width (-0.5147) and head length (-0.5046).

For pubescence type, all populations scored for the tomentose and intermediate types overlap in the

right cluster in the PCA graph, along with two populations scored as non-tomentose (populations KS and

GR, Fig. 1). All other non-tomentose populations appear in the left cluster. Similarly, populations scored

as weakly and intermediately glandular overlap within the right cluster, along with two of the populations

scored for strong glandularity (populations KS and GR). The left cluster correlates to populations of the

canyon/Camas Prairie grasslands, while all plants of the Palouse grasslands cluster on the right.

When plants are assigned to geographical range (Palouse grassland versus canyon/Camas Prairie

grassland) without considering population means, the ranges of the quantitative variables are overlapping

to varying degrees, but are all significantly different (Table 1). Population means assigned to geographical

range reveals far less overlap in head length, head width, phyllary width, and leaf width, but no appreciable

separation in lowest and highest values is seen in number of lateral branches and number of flower heads

(Table 2). Two outlier low values for population means of the canyon/Camas Prairie plants occur for head

length and phyllary width; when these outliers are disregarded, there is no overlap in the ranges for popula-

tion means in phyllary width, and very little for head length.

Plants of the Palouse grasslands in all but two populations were strongly or intermediately tomentose,

with the tomentum being lanate and distributed most densely on the phyllaries, but often throughout the

entire plant. Plants of the canyon grasslands and two proximal populations on the Palouse (Kramer Prairie

and Gross Road) often have harsh, curled hairs on the stems and inflorescence branches, but never on the

phyllaries, and the hairs are never lanate. Degree of glandularity was distributed among populations almost

perfectly concommitantly with the occurrence of tomentum, the exceptions being that some Palouse popu-

lations could be strongly tomentose but Honc ee or nas tomentose and weakly

glandular. All plants of the canyon/Camas Prairi were strongly glandular, with resinous-punctate

glands being distributed most strongly on the oi lares and usually also on the cauline leaves. Any glands

among Palouse plants were limited to the phyllaries. Among the quantitative variables, the Palouse plants

had a greater number of lateral inflorescence branches, more numerous flower heads, shorter and narrower

heads, narrower phyllaries and narrower cauline leaves (Table 1).

The Kramer Prairie (KS) and Gross Road (GR) populations, though located on the Palouse grasslands,

share the lack of tomentum and strong glandularity of the canyon/Camas Prairie plants. However, they are

attributable to the Palouse-type morphology based on (means and standard errors, values for KS preceding

D „eal D h Inetti ET.

234 Journal of tl

Tag 1. Means, standard error,

Geographical range

Character Palouse Canyon/Camas Prairie

Number of lateral branches* 2.39 + 0.16(0-17) 1.78 + 0.19(0-7)

Number of heads** 757 + 0.34(1-57) 5.67 + 0.39(1-33)

Head length (mm)*** 11.79 + 0.12(8-17(-18.5)) 1422 + 0.14(9-)11-21)

Head width (mm)*** 6.56 + 0.10(4-10(-11)) 8.71 + 0.11(6-14)

Phyllary width (mm)*** 1.90 + 0.03(1-3(-3.5)) 2.83 + 0.04(1.5-)3-5)

Leaf width (mm)*** 7.23 + 0.21 (4-14) 10.45 + 0.21(4-) 6-19)

Taste 2. R f lati f iteti holoaical ct for nl ianed hical Vales i | innle-ahservatian outliers

pl Fr I 1 T UICE YGUVI VANT 9.

3 F 3 is Asics hind J i; J

Geographical range

Character Palouse Canyon/Camas Prairie

Number of lateral branches 0.5-6.4 0.0-3.17

Number of heads 1.6-14.0 3.0-15.0

Head length (mm) 10.3-13.9 (8.5-)13.8-15.1

Head width (mm) 5.8-8.7 74-11.

Phyllary width (mm) 14-2.3 (1.4-)2.3-3.3

Leaf width (mm) 5.0-9.4 93-154

in each case): head length (12.63 + 0.35; 12.53 + 0.35), head width (6.58 + 0.23; 6.66 + 0.18), phyllary

width (2.29 + 0.73; 2.27 + 0.12), and leaf width (7.44 + 0.57; 7.55 + 0.59), though their mean number of

lateral branches (0.53 + 0.29; 1.36 + 0.43), and number of flower heads (6.82 + 0.78; 5.64 + 0.71) fell near

the means of the canyon/Camas Prairie plants.

The type specimen of Pyrrocoma liatriformis (CV. Piper s.n. 29 August, 1903, Pullman, Washington,

US) shows a tomentose plant lacking conspicuous glandularity. The number of lateral branches average 6.5

between the two stems, and the number of flower heads averages 9.0. Head length averages 12.03 mm (6

heads), phyllary width ranges 1.5-2.0 mm, and width of the lowest leaves of the inflorescence is 7.5 mm.

The type of Pyrrocoma scaberula (A.A. Heller 3469, 29 July, 1896, Forest, Nez Perce County, Idaho, US) is a

non-tomentose plant with conspicuous glands on the phyllaries. Each of the 4 non-damaged stems bears no

lateral branches and a single flower head. The heads are 12.5 to 14.5 mm long, and the phyllaries are 2.2 mm

wide or greater. The width of the cauline leaves is narrow (mean of 4.2) compared to other plants from the

canyon/Camas Prairie examined in the present study. The type of Pyrrocoma suksdorfii is a tomentose plant

with no conspicuous glandularity. The number of lateral branches average 2.0, and the number of heads

per stem is 4.7. Head length ranges 10—11.5 mm, the phyllaries are under 2.0 mm long, and the cauline leaf

width averages 6.0 mm.

DISCUSSION

Qualitative and quantitative characters reveal that Pyrrocoma liatriformis sensu lato is morphologically

non-uniform, both within and among populations. Sorting plants and populations into geographical range

between the Palouse grasslands on the one hand and the canyon/Camas Prairie grasslands on the other reveal

that this variation is nonrandom and is correlated to geographical range. While geographical range does

not account for all variation, there are clearly two arrays of morphology that are each coherent within their

own geographical range. Further, narrow discontinuities in the arrays of morphological variation for half of

the quantitative characters (Head length, Phyllary width and Leaf width) are itant when population

Bjórk and Darrach, Resurrection of Pyrrocoma scaberula

Pyrrocoma Populations

235

PL GH

A A

N p

e PB E SB PR

E ge AN 5 div d

F à

RR IS Ab i RT A

^ de GR A

WR & A, ra

TS A A

FE e

Axis 1

Pyrrocoma Populations

iactat

AP : Region

TS WR ^. i :

à i A "

FB a

å KS

uc GE Ap

é "ad ^

FE l

A. A bd wie hi

m h LR

E CH AR

T N A

Ls A srÁ

Á fe

Axis 1

Fic. 1. PCA graph showing axes 1 versus 2, and 1 versus 3, respectively. The solid triangles ref populations of tl C

The open triangles represent the Palouse region.

Dat M ID hI de dg rh £T.

236 Journal of tl Texas 3(1)

means are sorted by geography. While there is considerable overlap in tl ining quantitative character

and two of the Palouse populations break with their co-distributed populations int

the distinctions among the most strongly diagnostic quantitative characters e width, Head leno.

Head width and Leaf width) allows their placement within the Palouse morphology.

In the initial herbarium-based investigation comparing the Palouse and canyon/Camas Prairie mor-

phologies within Pyrrocoma liatriformis sensu lato, a greater spread in means was found for number of lateral

branches (3.86 + 0.73 versus 1.54 x 0.49, respectively) and number of flower heads (11.89 + 1.86 versus 5.38

+ 1.07). Extreme drought occurred throughout the study area in the year of the field work, with very little

precipitation falling in the preceding winter and spring, when normally the bulk of annual precipitation falls

in both the Palouse and canyon/Camas Prairie regions. This suggests that under normal conditions, these

two quantitative characters would be more diagnostic than is suggested by the data gathered from the field

in 2005. Plants in all populations visited had a portion of their lateral branches and flower heads aborted,

as evidenced by the number of small, distorted branches and heads that senesced without maturing flowers

or fruits.

Examination of the type specimens of Pyrrocoma liatriformis, Pyrrocoma scaberula, and Pyrrocoma

suksdorfii reveal that these plants are comparable to others collected from the same geographical regions in

the present study. The observed morphologies of the Palouse grassland populations are clearly represented

by the types of Pyrrocoma liatriformis and Pyrrocoma suksdorfii. The latter shows no appreciable distinction

from the former, and its morphological characters fit within the ranges observed in the field for the Palouse

populations in general, thus the name P. suksdorfti appears to be correctly synonymized under P. liatriformis.

The type of P. scaberula fits well within the observed range of morphologies of the canyon/Camas Prairie

populations (except in the width of the cauline leaves, which are within the range of the Palouse populations,

and the anomalous single flower heads per stem, which was rarely observed in both geographical ranges).

Thus, the type of Pyrrocoma scaberula appears to be representative of all populations of the canyon/Camas

Prairie region.

KEY TO PYRROCOMA LIATRIFORMIS AND PYRROCOMA SCABERULA

. Plants mostly heavily lanate-tomentose throughout, sometimes only on and near the flower heads;

resinous-punctate glands usually absent or inconspicuous; population means of flower head length

10-13.8 mm; preria 1.4-2.3 mm wide; population means for width of the lowest leaf of the inflo-

scence up to 9.3 mm; plants occurring in the Palouse grasslands Pyrrocoma liatriformis

f Plants hispid, any longer hairs harsh and firm, never lanate; resinous-punctate glands conspicuous,

especially on the phyllaries; population means of flower head length 13.8-15.1, very rarely less; phyl-

laries 2.3-3.3 mm (very rarely less); population means for width of the lowest leaf of the inflorescence

more than 9.3 mm; plants occurring south of the Palouse grasslands Pyrrocoma scaberula

—

The degree of variation within Pyrrocoma liatriformis and Pyrrocoma scaberula, the overlap in certain char-

acters, and the narrow discontinuities in both the geographical ranges and among the most diagnostic

quantitative characters suggest a close relationship between the two. While the coherence of morphologies

within the Palouse and canyon/Camas Prairie geographical ranges suggests that it is highly unlikely that

the two species are randomly polyphyletic to each other, it is possible that one species is derived from the

other (one monophyletic and the other paraphyletic), rather than both being derived from a now extinct

common ancestral race (both monophyletic). Further, hybridization may have occurred giving rise to inter-

mediate populations. The two Palouse populations here assigned to Pyrrocoma liatriformis which have the

non-tomentose and strongly glandular condition of Pyrrocoma scaberula suggest either hybridization, or that

these populations might actually be closer related to P. scaberula. It is also possible that specific taxonomic

rank may be unmerited for P. scaberula, such that it would be best recombined as a variety of P. liatriformis.

An ongoing molecular study will address these questions, assaying AFLP loci from all plants measured in

the field in the present study.

Population means of head length, phyllary width and leaf width, along with the strong toward

237

r ty, Idaho, NH L

238 J ical Insti Texas 3(

character states in the other morphological variables allow the unambiguous assignment of populations to

either Pyrrocoma liatriformis or P. scaberula. Thus, the segregation of the relatively common and widespread P.

scaberula leaves P. liatriformi stricto as a very rare and highly threatened species. Several populations

of P. scaberula are over 1,000 individuals (K. Gray, pers. comm.) and occupy large tracts of natural grassland,

and the newly discovered populations in Wallowa County, Oregon and Washington County, Idaho suggest

that additional populations may await discovery in the poorly explored grasslands of Hells Canyon in the

intervening area. In contrast, nearly all of the populations of P. liatriformis are less than 500 individuals, and

its potential habitats are all much smaller tracts surrounded by agricultural fields (Fig. 2). Intensive surveys

in 2005 for additional populations in the Palouse grasslands by the present authors, as well as by the Idaho

Conservation Data Center (K. Gray, pers. comm.) and Washington Natural Heritage Program (T. Kemper,

pers. com.) in the same year make it highly unlikely that additional large populations will be discovered.

Thus, P. liatriformis sensu stricto is vulnerable to extinction and should be protected accordingly.

ACKNOWLEDGEMENTS

This research was supported by funding from the U.S. Fish and Wildlife Service and Bureau of Land Man-

agement. We are very grateful for the enthusiastic assistance from the Idaho Conservation Data Center,

the Stillinger Herbarium, University of Idaho, the Ownbey Herbarium, Washington State University, and

especially to Karen Gray for the seed idea, and we thank Federico Osorio for the Spanish translation. David

J. Bogler and an anonymous reviewer provided helpful comments.

REFERENCES

Boater, DJ. 2006. Pyrrocoma. In Flora of North America Editorial Committee. Flora of North America, Vol. 20.

Oxford Univ. Pr. New York.

Clark, W.D, et al. 1980. Systematic implications of flavonoid patterns in Haplopapt gregates. Biochem. Syst

Ecol. 8:257-259.

Greene, E. 1909. Pyrrocoma liatriformis. Leafl. Bot. Observ. Crit. 2:17.

Greene, E. 1909. Pyrrocoma scaberula. Leafl. Bot. Observ. Crit. 2:19.

Greene, E. 1909. Pyrrocoma suksdorfii. Leafl. Bot. Observ. Crit. 2:18.

Hau, H.M. 1928. The genus Haplopappus: A phylogenetic study in the Compositae. Publ. Carnegie Inst. Wash.

389.

HitcHcock, C.L. AND A. CronquisT. 1973. Flora of the Pacific Northwest. University of Wash. Pr. Seattle.

Kemper, T. 2005. Report on the status of Haplopappus liatriformis (Greene) St. John in Washington. Washington

Natural Heritage Program, Department of Natural Resources. Olympia.

MaNcuso, M. 1997. Palouse Goldenweed (Haplopappus liatriformis) Monitoring at Craig Mountain, Idaho—1996

Results. Unpublished report for the Idaho Department of Fish and Game, Boise.

Maves, R.A. 1976. A cytotaxonomic and chemosystematic study of the genus Pyrrocoma (Asteraceae: Astereae).

PhD dissertation. University of Texas, Austin.

MorGan, D.R. 1992. A systematic study of Machaeranthera (Asteraceae) and related groups using restriction site

analysis of chloroplast DNA. Syst. Bot. 17:511-531.

MISCELLANEOUS TYPIFICATIONS, ONE NEW SERIES, AND

ONE NEW VARIETAL COMBINATION IN CRATAEGUS (ROSACEAE)

J.B. Phipps

The University of Western Ontario

Department of Biology

1151 Richmond St. N.

London, Ontario, a 5B7, CANADA

jphippsauwo.ca

ABSTRACT

Fifteen specific epithets and one varietal epithet in Crataegus Mari are typified. These are: C. brainerdii Sarg., C. subrotundifolia

Sarg., C. brumalis Ashe, C. gravis Ashe, C. populnea Ashe, C. acutiloba Sarg., C. demissa Sarg., C. macrosperma, C. matura Sarg., C. pastorum

Sarg., C. dinis la C. roanensis Ashe, C. laurentiana Sarg., C pas Sarg., C. brunetiana Sarg., and C. laurentiana var. dissimilifolia

(the last ial hybrids). A combination, Crataegus chrysocarpa Ashe var. subrotundifolia and a new series of

central Asian: Crataegus, Altaicae, are also proposed.

Key Wonps: Crataegus (Rosaceae), typifications, Crataegus ch pa Ashe var. subrotundifolia (Sarg.) J.B. Phipps, comb. et stat. nov.,

North American, Crataegus ser. Altaicae J.B. Phipps, ser. nov.

RESUMEN

Se tipifican qui it y uno varietal en Crat (Rosaceae). ). Estos son: C. brainerdii Sarg., C. subrotundifolia Sarg., C.

brumalis Ashe, C. Eu Ashe, c. populnea Ashe, C. acutiloba n C. demissa RS C. macrosperma, C. matura Sarg., C. pastorum Sarg.,

e peana Sarg, i roanensis Ashe, C. Mua Sarg., C. Aa Sarg., C. brunetiana Sarg., y C. laurentiana var. dissimilifolia (híbridos

, Crataegus chrysocarpa Ashe var. subrotundifolia y una nueva

e de Crataegus, Altaicae, dé Asia central.

INIRODUCTION

Until recently, most of the names used in the forthcoming treatment of North American Crataegus for Flora

of North America, vol. 9, were not or not precisely, typified. This is gradually being rectified by the author,

in part with various colleagues, by the typification of hundreds of names, for the most part those that will

appear in the flora. Several important names from classical authors of the late eighteenth up to the later

nineteenth century have now been typified, including e.g., C. viridis L., C. crus-galli L., C. coccinea L., C.

intricata Lange, C. flava Aiton, C. pruinosa (H.L. Wendl.) K. Koch, and C. rivularis Nutt. However, by far the

majority of names recently typified come from the “explosion period” of Crataegus taxonomy, a time when

the type concept was beginning to take hold, so only a few of these names prove to have holotypes. Of

the major authors of this period, C.D. Beadle either cited cotypes (usually matching flowering and fruiting

specimens from the same tree) in the protologue or designated them on the sheets and his taxa are gener-

ally straightforward to lectotypify. Beadle's types are mainly at US and NY. Charles S. Sargent normally

indicated syntypes, either explicitly in his protologues or by annotating relevant specimens as “type” or

simply ‘n. sp.. There is virtually always adequate surviving material for lectotypification at A or GH, and

sometimes duplicates elsewhere, the difficulties instead lying in selecting an appropriate lectotype from the

putative types when slight discrepancies from the protologues are found, or if, sometimes, there are mixed

gatherings. Further, there may be complications as a consequence of the tree numbering system in which

various collaborators used the same number for a particular tree even when collecting at different dates.

Sargent's names also present a challenge by their sheer number (832) but several hundred have now been

typified. WW. Ashe is the major author of this period whose names present the most difficult problems as

his indications in the protologue of type, collector and location, are often poor, ambiguous, or even lacking

J. Bot. Res. Inst. Texas 3(1): 239 — 243. 2009

240 Journal of t i itute of Texas 3(

so that only the existence of handwriting by Ashe or one of his collectors remains t documen t authenticity.

The dispersal of authentic Ashe material during his active period was to a vide range of herbaria so it can be

difficult to track down, while his private collection, often poorly labeled and seemingly incomplete, had to

await curation at NCU until after his death by T.G. Harbison, this a labor of love never completed. Added to

all this, his protologues tend to have the least detail of these three authors, sometimes omitting characters

considered essential today. This is unfortunate for Ashe created many potentially earliest d certainly

many important species names bear his authorship, e.g., C. chrysocarpa, C. dodgei and C. margaretta (since

typified), C. macrosperma, C. roanensis, C. holmesiana (still untypified). Consequently, Ashe names usually

require neotypification, and not surprisingly, there is little enthusiasm for entering this minefield.

This paper concentrates on several Sargent and Ashe examples to provide valid names for clearing up

some taxonomic problems in ser. Tenuifoliae, Brainerdianae and Rotundifoliae and some putative interserial

hybrids probably between ser. Rotundifoliae and Macracanthae. The text is also the vehicle for introducing

the segregate central Asian series Altaicae.

—Crataegus series Brainerdianae

1. Crataegus brainerdii Sarg., Rhodora 3:27.1901. Te: U.S.A. Vermont. Addison Co.: SW Middlebury, “Garrett House,” 22

Sep 1900, E. Brainerd 6b ( , designated here: A 2536, duplicate at A).

ee la ee ue ne ther two syntypes, has both flowering and fruiting material. The lectotype shows beautifully the

dant, + ellipsoid fruit

B—Crataegus series Rotundifoliae

1. Crataegus chrysocarpa Ashe var. subrotundifolia (Sarg.) J.B. Phipps, comb. et stat. nov. Basionym: Crataegus

subrotundifolia Sarg., Bot. Gaz. 35:394. 1903. Tee: U.S.A. ILLINOIS. Lake Co.: shores of Lake Zurich on bluff bank, 13 May 1901, EJ.

Hill 31a (LecrorwrE, designated here: A).

Comment. a pone mO same label data dn 5 T ro dj Sa En qu suns a leaves on gu

However, as I

y y

to e ibis consequently wid d form, I he thie AN specimen for the type.

P q y P E g Sp YP

This variety is erected to accommodate those variants of Crataegus chrysocarpa with glabrous hypanthia and

variably hairy inflorescences.

C—Crataegus series Silvicolae

1. Crataegus brumalis Ashe, Ann. Carnegie Mus. 1:393.1902. Tre: U.S.A. Pennsyvania. Allegheny Co.: Pittsburgh, 18^

Ward, Stanton Ave., corner e Morine le hoe p ae 19a, 20 e aes: ee here: PH 648488).

Comment. pe. The stamens on the lectotype

s eo

are said to be puis

This taxon, often made a variety of C. iracunda Beadle, has + glabrous adaxial leaf surfaces at anthesis and

much broader truncate leaf-bases. Moreover, it is out of range for C. iracunda so it is best assigned to ser.

Silvicolae as a scarce presumptive interserial hybrid. Most records of it are probable misidentifications of a

form of C. macrosperma.

2. Crataegus gravis Ashe, J. Elisha Mitchell Sci. Soc. 20:49. 1904. Twe: U.S.A. Micucan. St. Clair Co.: Port Huron, CK

Dodge Type Tree 28, 26 May1902 (Lecrorvee, designated here: MICH). Errrype: U.S.A. Michigan: St. Clair Co.: Port Huron, C.K. Dodge

Type Tree 28, 7 Oct 1902 (1ecrorvre, designated here: MICH, duplicate at MICH J:

Comment. —This tree was apparently quite frequently collected and Kruschke (1965) mentions specimens at MSC and MIN. Un-

fortunately, this flowering syntype, the only one located, is of indifferent quality, so I am also epitypifying with a superior fruiting

specimen from the type tree.

This species was said to be ‘common’ by Ashe but it has not been seen in the vicinity of Port Huron, in-

cluding the very well-collected Sarnia area across the river in Ontario, in recent years. It is somewhat like

Crataegus pruinosa var. parvula in general appearance but has slightly hairy adaxial leaf surfaces, a propor-

tionately broader and slightly larger leaf, a larger and narrower fruit type, and lacks the elevated calyx of

C. pruinosa.

TEN 241

Phipps, Miscell Ur g

3. ge populnea Ashe, Ann. Carnegie Mus. 1:395.1902. Type: U.S.A. Penusvivania. Berks Co.: 2.5 mi NW of

town, beyond ee Hill, C.A. Gruber 31, 15 May 1902 (LecrorwrE, designated here: PH).

PH dated 20 Sep 1903, have otherwise a label data.

Comment. g SI

eiusd ien ion of this fairly common northeastern species,

TI + L| ££

suh; is followed in FNA vol. 9. The large leaves, n at anthesis, and squarish lobe-tips of many of

the leaves are characteristic.

D—Crataegus series Tenuifoliae

Forms of C. macrosperma that have been made into varieties have been selected for typification.

1. aa acutiloba E Rhodora 3:23. 1901. Tye: U.S.A. Mame. Hancock Co.: Mt Desert, 4 Sep 1899, B. Jones s.n.

OTYPE, designated here

E nt.—A beautiful 2s pes annotated i du n. Vd is nod Pi ue aid It à the m large,

cuneate-based, somewhat elongate (5-8

characteristic large-leaved type of the coast from Massasuchetts Bay to Nova Scotia but ated neher specimens nor any precise

localities.

An A specimen annotated ‘type’ by Kruschke is not supported in Kruschke (1965).

2. Crataegus demissa Sarg., Rhodora 5:139. 1903. Tre: U.S.A. Vermont. Chippenden Co.: Charlotte, 26 May1902, EH.

Horsford s.n. iu a. bonds here:

Commen mall shrub' and has all the short-shoot 1 + cordate, exactly fitting the

protologue. A fruiting syniype collected by per from Charlotte, Vt., is a good match. However, all the syntypes from Lennox,

Mass., have mostly a leaf bases, with only a few + truncate.

This is a very distinctive, apparently dwarf, form of C. macrosperma but there appears to be little material

from anywhere else closely matching the type.

3. Crataegus macrosperma Ashe, J. Elisha Mitchell Sci. Soc. 16:73. ad Tee: U.S.A. ALABAMA. DeKalb Co.: Desoto

State Park, pines 25 Sep 2001, R. I 170 , designated here: UWO

f1 the type locality: I

Com

omment

and M matching thé protologue, except for their apices (see below).

Crataegus macrosperma is the most important taxon dealt with in this paper. Ashe gives the distribution as

northern Alabama, northwestern Georgia and the adjacent portions of Tennessee, being frequent on Lookout

Mountain in the last state (type locality) and in the g mountains. Lacking authentic material and

working with an inadequate protologue, my interpretation is affected by attempting to create, if possible, an

entity in the Tenuifoliae that satisfies the protologue sufficiently and also differs from C. roanensis, the other

common regional member of this series, which Ashe held to be different. There is little meaningful differ-

entiation in the protologues except for range, C. macrosperma having ‘round’ versus ‘oblong’ fruit, a more

southwesterly, lower altitude distribution, leaf blades ‘deltoid’ (perhaps extension shoots, this not specified)

or ‘oval’, a rather generic term of that period. Crataegus macrosperma is also said to have leaves ‘obtuse’ at the

apex, but nothing like that has been seen in series Tenuifoliae, to which the protologue fully applies in other

respects. Thus I am assuming that this is a defect in the description. There are two extreme leaf types in the

general region of the type areas of C. macrosperma. and C. roanensis, southwestern Appalachia, that broadly

correlate with the different fruit types so I am typifying C. macrosperma on a specimen lacking cuneate leaf

bases but having orbicular fruit. This, or a similar entity, appears to be a widespread form extending to

the north of the range of the species though whether it merits varietal separation from C. roanensis requires

further work. Certainly, apparent intermediates with C. roanensis occur.

4. Crataegus matura Sarg., Rhodora 3:24. 1901. Tree: U.S.A. Vermont. Addison Co.: N of Bristol village, 19 Sep 1900, E.

Brainerd 10h (LecrorvrE, designated here: A).

Comment.—A fine fruiti i is selected for the 1 It has very large leaves to 9 cm long, broad ovoid fruit and

a broader angle at the leaf-base than C. acutiloba.

242 J i i Texas 3(

Crataegus matura and C. acutiloba appear to represent poles of variation of large-leaved northeastern C.

macrosperma. An A specimen annotated ‘type’ by Kruschke is not supported in publication.

5. eei pastorum Sarg., Rhodora 3:24. 1901. Tree: U.S.A. Massacuusetts. Worcester Co.: West Boylston, 6 Oct 1900,

G. Jack 11 E n cmd jus Ma

Comment. ith K i l di fi f C. macrosperma

from the Claro olt valley (Vermont) aud Berkshire Co. (ase ), to al and southern esaidak but cited no specimens nor

any precise localities. There are, however, plenty of specimens labeled ^n. sp. ' by Sargent from the stated area.

4 f

6. Crataegus pentandra Sarg., Rhodora 3:25. 1901. Tree: U.S.A. Vermont. Rutland Co.: West Rutland, 17 Sep 1899, WW.

Eggleston 1135 (1eczorrrE, designated here: A).

Comment.—A fruiting syntype specimen is selected for the lectotype. It has quite large leaves to 6 cm long, large subglobose fruit

and is somewhat similar to C. matura and C. acutiloba.

7. Crataegus roanensis Ashe, Bull. North Carolina Exp. Sta. 175:114.1900. Tyee: U.S.A. NORTH CAROLINA.

Henderson Co.: Bearwallow Mountain, Sep 1994, R.Lance 205 (NEOTYPE, designated here: NCU).

Palmer in 1946 made this a variety of Crataegus macrosperma eventually differentiating it by the depth of

lobing of the leaves while Kruschke raised it to the rank of species. This attention was given even though

the species had no formal type. Ashe gives the distribution as Yancey and Mitchell counties, North Carolina,

especially about the base of Roan Mountain (on the Tennessee border of Mitchell Co.), saying that it is not

common below 4000 ft but between 4000 and 6000 ft it is one of the most common thorns. Although,

presumably, Ashe held this to be different from C. macrosperma, there is little differentiation in the proto-

logues, this taxon having ‘oblong’ versus ‘round’ fruit. I find two extreme leaf types in the general region of

the type areas for C. roanensis and C. macrosperma that broadly correlate with the different fruit types so I

am typifying C. roanensis on a specimen with cuneate leaf bases and oblong fruit. Forms of the macrosperma

complex with narrow fruit are widespread, extending to Wisconsin, Ontario and New England and warrant

further study. Interestingly, the quite numerous specimens from Yancey and Mitchell counties at NCU better

match C. macrosperma than C. roanensis as interpreted here.

E—Putative interserial hybrids

Taxonomic discussion of the following may be found in Flora of North America, vol. 9, which it is anticipated

will be published later this year.

1. Crataegus laurentiana Sarg., Rhodora 3:77. n 901. Tres: CANADA. Quzszc. La Prairie Co.: La Tortue (outskirts of Delson),

8 Oct 1899, J.G. Jack 40 (Lectoryre, designated here:

Comment.—The type has eroded nutlets and in E 40 Gwan the petioles are virtually eglandular. The latter is a particularly

fine specimen from the type locality. Flowering syntypes have pink anthers. Further syntypes, also Jack collections, come from La

Tortue and Caughnawaga (same county).

2. Crataegus laurentiana Sarg. var. dissimilifolia Kruschke ex J.B. Phipps. Tree: U.S.A. Wisconsin. Ashland Co.:

TrA Island, 1.5 mi S of La Pointe, 11 Sep 1950, E.P Kruschke K-49-145 (LectoTYPE, designated here: MIDW, duplicate of lecto-

common — flowering co-type, same label data except 3 Jun 1949, would make an excellent epitype. Anthers of the latter are

Crataegus laurentiana Sarg. var. dissimilifolia Kruschke, Milwaukee Public Mus. Publ. Bot. 3:35. 1965

Although Kruschke’s protologue has the word type associated with the second, fruiting specimen of a

matching pair, it is clear both from the consistency of his usage as well as from the labeling of the speci-

mens, that he understands the pair as co-types, the first being the flowering specimen of the same collection

number. Kruschke’s only exception to this system is found in a few cases where he only collected a type at

one season.

3. Crataegus fernaldii Sarg., Rhodora 5:166. 1903. Tre: U.S.A. Maine. Aroostock Co.: U.S.A.: Maine: Aroostock Co.: Fort

Fairfield, river thicket, 27 Sep 1901, M.L. Fernald Cr. 21 (LECTOTYPE, designated here: A

The sharp and deep lobing of the leaves and relatively broad breadth: length ratio is reminiscent of C. chryso-

Phipps, AA: TT 4 se a2 tn unt 243

carpa but the near-glabrous and near-eglandular petioles are more like C. macracantha. Pink anthers in a

syntype of th ber (presumed type tree) could well originate from C. A but would be very

unusual in C. chrysocarpa while the somewhat eroded nutlets of the lectot liacy between

the species discussed. In sum, this seems to be a broader-leaved, en le: MM C. laurentiana.

4. Crataegus brunetiana Sarg., Rhodora 5: "a 1903. Tree: CANADA. Quesec. Quebec Co.: M y Falls, 30 May

1901, J.G. Jack 129 (Lectotype, designated here:

This is rather similar to C. laurentiana but the leaves are proportionately wider, even more sharply lobed

in flower and less hairy (rather as in C. fernaldii), the petioles at anthesis more glandular, the inflorescence

branches of a less silky villosity and the anthers cream. The flowering syntype, Jack 129, has pitted nutlets

like C. laurentiana while those of another syntype, Jack 120, do not.

F—Crataegus series Altaicae

Crataegus series Altaicae J.B. Phipps, ser. nov. Te: Crat ltaica 1 (Crat iana Hemsley & Lace).

Similes ad ser. parquitieaes ser. MU iem cum NUN luteis vel fuscis, non rubris vel atris, inflorescentiis glabris, non pilosis vel

subpilosis. Distributio: A il Species 1 vel 23

Series Altaicae is a small (1-2 or 3) species series widespread in Central Asia to the west of the central Asian

massif which ranges from Baluchistan (Pakistan) to the northern slopes of the Altai in south-central Sibe-

ria. It is named for a species widely known as C. altaica Lange, perhaps synonymous with C. wattiana. The

new series is segregated from ser. Sanguineae, with which it shares important characters of pitted nutlets

and falcate basal bracteoles. However, it has glabrous inflorescences, a very unusual yellowish to tan fruit

color and occurs, uniquely for sect. Sanguineae, in a semi-xeric upland climatic zone. Differently from sect.

Sanguineae as here emended, ser. Altaicae may also have an unusual but here not uncommon form in the

species complex C. wattiana/altaica which has very deeply lobed leaves and veins to the sinuses.

ACKNOWLEDGMENTS

The curators and staff of HUH and PH are thanked for their valuable cooperation, particularly Melinda

Peters at Harvard who imaged many syntypes for me. James L. Reveal and Kanchi Gandhi provided valu-

able review comments.

REFERENCES

KruscHKE, E.P. 1965. Contributions to the taxonomy of Crataegus. Milwaukee Public Mus. Publ. Bot. 3:1-273.

BOOK REVIEW

GRAHAM CHARLES. 2003. Cacti and Succulents. (ISBN: 978-1-1861268726, pbk.). The Crowood Press Inc.,

The Stable Block, Crowood Lane, Ramsbury, Marlborough, Wiltshire SN8 2HR, England. (Orders: www.

od E co.uk, in puse 01-672-520320). $35.00, 192 pp., 8 3/8" x 10 3/4".

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Texas 76102-4025, U.S.A.

J. Bot. Res. Inst. Texas 3(1): 244. 2009

WHAT IS RANUNCULUS GELIDUS (RANUNCULACEAE)?

Alan T. Whittemore

U.S. National Arboretum

3501 New York Ave. NE

Washington, DC 20002-1958 U.S.A.

ABSTRACT

Le 5 1 EP. 1 Ain Marth A : H r id ] i 1 but some recent

e P I A E

fe I lled this species R. karelinii, while Asi f I R. gelidus and R. karelinii fı lemic st

cies of central Asia 1 I R fi ] f Siberi d t North America. R 1 Iba]

r o

lemic in the Tian Shan and Jungar SI f central Asia, and R. karelinii is illegiti perf] l f

R. gelidus). Populati fi beri d North America. which differ f R. gelidus in thei EE EE Part lool

smaller meee and less deeply divided leaves, are correctly known as R. grayi. The section in which this species is d is correctly

called Ranunculus sect. Epirotes (Prantl) Benson— "Ranunculus sect. Auricomus Spach,” used in some recent suos is not valid.

RESUMEN

Ei hre R lus gelid 4 te en Norte Amé áncul á la zona ártico-montana, pero algunas

referencias recientes americanas han llamad ta especie R karelint ] ] fe los nombres R

li r dial y a Ae Aca central y AD 1 Jal d TL es 1 TES

Ey r 4

Norte América. R lus gelid ] | p demi de Tian Shan y Jungar Mrd en a a Y R ice es

ilegiti fluo de R. gelidus). I blaci del Je Sil

o I r o E P

401 2 INE EE ES 1 5 a hoias dividid fund leben ll R. grayi EUM

M ^r E 1 af J Is

lebe inclui t ie del ] R I t. Epi (Prantl) Benson—“Ranunculus sect. Auricomus Spach,”

t E

usada en algunas publicaciones recientes, no es válida.

Treatments of Ranunculus in North America have generally used the name R. gelidus Karelin & Kirilov for a

small buttercup that is widespread in arctic and alpine regions of western North America and eastern and

central Asia (Benson 1948; Hitchcock & Cronquist 1964; Scoggan 1978; Whittemore 1997). However, a

number of recent North American publications (Qian & Klinka 1998; Kartesz & Meacham 1999; Panjabi &

Anderson 2006; Kratz 2007; NRCS 2008) treat this species under the name R. karelinii Czerep. This usage is

questionable on several grounds. First, R. karelinii was not published until 1981, and three older names are

listed in the synonymy of R. gelidus in North American references (R. grayi Britton 1891, R. verecundus B.L.

Robinson ex Piper 1906, and R. ramulosus M.E. Jones 1912; Benson 1948; Hulten 19682; Whittemore 1997).

Second, recent Asian references (Ovchinnikov 1937; Baitenov et al. 1961; Voroshilov 1982; Tumokhina 1992;

Cherepanov 1995; Borodina-Grabovskaya et al. 2001; Wang & Gilbert 2001) use the name R. harelinii for

a species endemic to Central Asia, and treat specimens from eastern Siberia and North America as R. grayi

Britton. Third, the name R. karelinii is superfluous and illegitimate according to IPNI 2008).

This widespread buttercup is currently going under three names (R. gelidus, R. grayi, and R. karelinii)

in recent floristic literature, an unsatisfactory situation. The taxonomy and nomenclature of the species was

investigated, as it relates to both the North American species and the type material from central Asia, to

determine the correct name for the species.

NOMENCLATURE

Cherepanov (1981) concluded that R idus Karelin & Kirilov 1842 is an invalid homonym because

he felt that R. gelidus Hoffm. was validated a by Reichenbach (1830-33, p. 720). However, examination

of this publication makes it clear that R. gelidus Hoffm. was not validly published. Reichenbach listed "R.

gelidus Hffgg.” under R. glacialis forma al, formatted like synonyms elsewhere in the book, not like names of

other formae in the species (which have a single Latin epithet, not preceded by “R.”; Fig. 1). Hoffmansegg's

J. Bot. Res. Inst. Texas 3(1): 245 — 250. 2009

246

4584, R. el ACI iali is IL. "

ternatia, foliolis tripartito = Eg <A

caulinis sessilibus, caule paucifloro)

calycibus ferrugineo - hirsutis , petalis

rotundatis, squama nectar brevissima

bifida. — Tres inprimis formas dis.

tinguo: @. R, gelidus A, '8 8. 7X al-

pestri - glacialis? foliis R. alpestris

cordato - renilormibus 3 — 5$- partitis,

partitionibus trifidis obtusis, — . gla-

cialis genuinus ¢ foliis | circumscriptione B

rotunda 3 — 5 — partitis , partitionibus Y

lla -trifidis incumbentibus ; huc

borealis: : L in". lapp. t 3. f. L X signo hybriditati sum in sectione eecunda tertiaque,

pr dan, t. 2 et yoo W «if.

in Jacq. coll. I, t. 8. et 9. f£. 1,2. —

crithm ifolius : foliis a par-

T longius petiolnlatis triter=

Fis, 1. A. Part of the d scri tion of R lus glacialis from Reichenbach (1830—33 p. lo ae. in meca joe of ^R. gelidus Wffgg." as-

fR glacialisformaa B. A short pt from the"Praemonenda" of Reich p. iv), explaining the unusual

symbol in the discussion shown in part À.

unpublished name cannot be validated at the species level by its association with Reichenbach’s descrip-

tion of a forma, so the supposed name “Ranunculus gelidus Hoffmansegg ex Reichenbach” does not exist.

Ranunculus gelidus Karelin & Kirilov is therefore a valid name and is not a superfluous name as concluded by

Cherepanov. The editors of IPNI are correct in concluding that it is R. karelinii Czerep., not R. gelidus Karelin

& Kirilov, that is superfluous and illegitimate.

A second nomenclatural problem associated with these species is the correct name of the section to

which the species belongs. North American publications (Benson 1948; Whittemore 1997) refer to it as

Ranunculus sect. Epirotes (Prantl) Benson, while some recent European references (Tutin & Akeroyd 1993;

Hörandl et al. 2005) refer to it as Ranunculus sect. Auricomus Spach. However, Spach (1839, p. 210) published

this taxon at the rank of subgenus, not section, as already noted by Benson (1948). Ranunculus sect. Epirotes

is thus the correct name for this taxon at the rank of section.

TAXONOMY

North American and east Asian material of this buttercup was considered conspecific with the central Asian

Ranunculus gelidus by Ostenfeld (1909, p. 44). Benson, in his 1948 monograph of North American Ranunculus,

explicitly stated that he had not seen central Asian material of R. gelidus, and was following Ostenfeld in treat-

ing North American material under this name. In order to evaluate the relationship of the North American

buttercup to the type of R. gelidus, herbarium material from Central Asia (including isotype material of R.

gelidus) was compared with North American material for characters used by Asian authors (Ovchinnikov

1937; Borodina-Grabovskaya et al. 2001; Wang & Gilbert 2001) to distinguish R. gelidus s. str. from similar

species of Ranunculus. Several characters support the distinction of the Central Asian populations to which

the type of R. gelidus belongs from the North American and east Asian plant.

First, Asian references emphasize trifoliolate leaves with petiolulate leaflets in Ranunculus gelidus s. str.

North American and Siberian plants referred to R. gelidus s. lat. or R. grayi are described as having the leaf

blade 3-parted but not compound (Benson 1948; Voroshilov 1982; Tumokhina 1992; Whittemore 1997).

Leaf morphology is somewhat more variable than the literature suggests, and North American specimens

rarely may have the blade of the basal leaves divided to the base and thus technically compound, but the

leaflets are sessile, a condition not seen in specimens or in published descriptions of central Asian R. gelidus

Whittemore, Identity of Ranunculus gelidus 247

s. str. The shape of the leaf segments also differs, with the ultimate segments being oblanceolate to obovate

in Central Asian specimens and narrowly oblong or linear in North American specimens. Stem leaves often

han basal leaves, but they also never have well-defined petiolules in North American

material. The vanos of variation in the two regions is certainly different.

Second, North American and Siberian material that has been assigned to Ranunculus gelidus has the

petals only 3-6 mm long, scarcely longer than the sepals. Isotype material of R. gelidus and other Central

Asian specimens, however, have larger petals, 7-9 mm long (to 10 mm according to Wang & Gilbert 2001),

about twice as long as the sepals.

Third, the achenes of North American and Siberian material that has been assigned to Ranunculus

gelidus show a morphology similar to most members of Ranunculus sect. Epirotes (Prantl) Benson, with the

beak of the fruit parallel to the long axis of the achene. In Central Asian specimens, however, the beak is

displaced, originating on the adaxial side of the achene at an angle of ca 90? to the axis of the achene. This

character is unusual in the genus, and was emphasized by Ovchinnikov (1937) in erecting Ranunculus cycle

Gelidi Ovch. for two Central Asian species, R. gelidus and R. rufosepalus Franch. The unusual insertion of

the achene beak was not mentioned in the Flora of China (Wang & Gilbert 2001), but is well developed on

plants seen from the id narrow ee of the species in China (Yunatov et al. 948, MO).

Other cl t rlap, but still seem to show different ranges of variation in Cen-

tral Asia and North eee thus, Central pene lants always have solitary flowers, while North American

material has 1-5 flowers per stem.

It seems clear, then, that Porsild (1943), Benson (1948), and Whittemore (1997) were wrong in treating

the North American plant under the name Ranunculus gelidus, and recent authors who have used R. karelinii

are incorrect on both taxonomic and nomenclatural grounds. The correct name for this buttercup is R.

grayi, as used by Old World (and earlier North American) authors. It is unfortunate that it is necessary to

introduce a third species name for these plants to the North American literature, but the name change is

necessary to eliminate the incorrect application to North American plants of names belonging to a Central

Asian endemic and reflect the true relationships of the North American plants. The following taxonomy

ensues:

KEY TO RANUNCULUS GELIDUS SENSU FLORA OF NORTH AMERICA

. Achene beak sharply inflexed so it is + perpendicular to axis of achene. Petals 7-9 mm long, ca. twice as long

—

as sepals. Central Asia Ran aaa gelidus

1. Achene beak parallel to axis of achene. Petals 3-6 mm long, scarcely longer than sepals. North Americ

eastern Siberia ae grayi

Ranunculus grayi Britton, Bull. Torrey Bot. Club 18:265. 1891. (Fig. 2 E-H). Ranunculus pedatifidus Hook., Fl. Bor.-

Amer. 1:18, pl. 8 fig. B. 1829, not of Smith, The Cyclopaedia; or, Universial Dict. Arts,... 29: Ranunculus n. 72. 1818. Ranunculus

hookeri Regel, Reis. Sud. Ostsib. Bot. Abt. 1:47. 1861, not of en pud 210, 425. 1830. Ranunculus drummondii E.

Greene, Erythea 2:192. 1894, nom. superfl. R. gelidus subsp én, Ark. Bot. (n.s.) 7:59. 1968. Tyre: “Barren summits

of the Rocky Mountains, on tl f the ridge, lat. 52° to 55°, Drummond.”

Ranunculus verecundus B.L. Robinson ex Piper, Contr. U.S. Natl. Herb. 11:274-275. 1906. Tere: WASHINGTON. Yakima Co.: Mount

(=Mt. Adams), alt. 6000-7000 ft, WN. Suksdorf 93 (HoLotyPE: GH 38452)

Ranunculus ramulosus M.E. Jones, Contr. W. Bot. 14:47. 1912. Tyre: MONTANA: Border of Glacier and Flathead Cos.: Swift Current Pass,

Glacier National Park, 11 Aug 1910, M.E. Jones s.n. (HoLoTyPE: POM 100

Ranunculus gelidus var. shumaginensis Hultén nom. inval., Ark. Bot. (n.s.) 7:59. 1968, Madrono 19:223. 1968. Hulténs Arkiv for Botanik

paper has only the name and a Latin diagnosis, with no mention of a type specimen at all. The Madrono reference merely adds,

“Type in S," but gives no specimen information. Since there is no refer to a single specimen or gathering, the name is invalid

(McNeill et al. 2006, art. 37.1-37.2)

Plants erect or decumbent from short caudexes, 3-22 cm. Roots slender, 0.5—1.0 mm thick. Leaves and stems

glabrous or sparsely spreading-pubescent with long weak hairs, pedicels pubescent or glabrous. Basal leaves

cordate or reniform, 0.5—1.8 x 0.83.0 cm, 3-parted (rarely 3-foliolate with sessile leaflets), segments again

lobed, ultimat wly oblong or linear, 1.5-4 mm wide. Cauline leaves often larger, more deeply

248

Eal Das sa lO h ledit £T.

| | TL RAI

JUUI Tidi OF TEAS 211]

Fic? M hal fel

A-B basal leaves, Ca flower, D an achene. Ranunculus grayi, E-F basal leaves,

Gaflower, H an achene.

Whittemore, Identity of Ranunculus gelidus 249

lobed, and with narrower segments. Flowers 1-5 per stem. Sepals 3-5 x 1-4 mm, pubescent or glabrous;

petals 3-6 x 1-5 mm; receptacle glabrous or pubescent. Achenes 1.2-2.4 x 0.8-2.0 mm, glabrous, beak

0.4—0.8 mm, subulate, curved or hooked, parallel to axis of achene.

Seepy places in open rocky slopes and meadows, arctic and alpine. Widespread in western North

America (from Alaska south to Oregon and Colorado; Benson 1948; Hitchcock & Cronquist 1964; 5coggan

1978; Whittemore 1997) and eastern Asiatic Russia (from the Lena Valley and the mountains east of Lake

Baikal east to the Chukotka Peninsula; Voroshilov 1982; Tumokhina 1992; Cherepanov 1995).

1. CANADA Rritich Col ] ice, Tenquille Lake area, Crown Mtn., 6000-6500

ft, K Beamish & F. Vrugtman 60938 (US); alpine slopes of Chipuin Mtn., Marble Mts., 6000 ft, J.W. & E.M. Thompson 579 (US). Alpine

slopes of Bluster Mtn., Marble Mts., 7000 ft, J.W. & E.M. Thompson 384 (US). U.S.A. ALASKA: Yakutat Co.: head of Russell ul EV.

Coville & T.H. Kearney Jr. 940 (US). Southeast Fairbanks (ca) Co.: Weng on EM talus uo ea N-S ridg f

Mtn., Healy (D-4), 63? 45 1/2’ N, 148? 49’ W.S. Carwile 80-39 (US). IDAHO. B C lder Creek Gio

Sawtooth Mts., 10,000 ft, JW. Thompson 14100 (US). Custer Co.: Slide rock, 8000 ft, CHE IE Macbride & E.B. Payson 3394 (US).

MONTANA. Glacier Co.: moist rocky slopes, frequent; vicinity of Sexton Glacier, 1950-2200 m, Glacier National Park, P.C. Standley

17230 (US). Judith Basin Co.: Long Baldy, Little Belt Mts., 7000 ft, J.H. Flodman 469 (US). OREGON, BR Co.: apa mountain

adow, Dino Creek head I Steen's Mtn., 9000 ft, P. Train s.n. 31vii35 (NA).

inter glacier, Mt. Tahoma (=Mt. Ranier), 7500 ft, J.B. Flett 2177 (US). Yakima Co.: alpine slopes of Mt. Aix, 7500 ft, JW. Tami 15046

(NA, US); on volcanic sand and gravel, Mount Paddo (=Mt. Adams), ca. 2400 m, W.N. Suksdorf 4142 (US).

pen las gelidus. coun & Kirilov, Bull. Soc. Imp. Naturalistes Moscou 15:133. 1842. (Fig. 2 A-D).

herepanov, Sosud. Rast. SSSR. n 1981, nom. superfl. Tyre: In summis alpibus Alatau ad fontes fl. Lepsa,

ing — igi ME a f I 1841, Karelin et Kiriloff (soryre: NY - Torrey!)

Plant to 15 cm tall. Roots secos or fleshy, «1-2 mm thick. Leaves ( | d ti faces), stems,

petioles and pedicel ly spreading-pubescent with long weak Bars Basalleaves to 4—10 cm ine blade

to 7-23 mm ine. 12-25 mm wide, 3-foliolate, leaflets with well-defined petiolules; leaflets again 2-3-parted

and the parts lobed, ultimate segments oblanceolate to obovate, 1-3 mm wide. Cauline leaves similar but

shorter. Flowers solitary. Sepals 3-5 x 3-4 mm, spreading-pubescent with long weak hairs; petals 7-9 x

4.5-9 mm; receptacle pubescent. Achenes glabrous, 2.3 x 1.2 mm, very strongly asymmetrical (ventricose),

beak 0.4 mm long, straight, inflexed so that it is + perpendicular to axis of achene.

Alpine areas on stony soil and talus where watered by snowmelt. Endemic to the Jungarskiy Alatau and

Tien Shan, Central Asia: Kazakhstan (Cherepanov 1995) and Xinjiang, China (Wang & Gilbert 2001).

Additional specimens examined: KAZAKHSTAN: Kumdaban, Jungar Mts., 9000 ft, May 1879, A. Regel s.n. (US). CHINA. Xinjiang:

Tian Shan SW of Manas (W of Urumchi), on talus, 21 Jul 1957, A.A. Yunatov, Li Shi-In, & Yuan Y-Fen 948 (MO

ACKNOWLEDGMENTS

I would like to thank the curators of ALA, CAS, COLO, GH, MO, NY, UC, and US for providing specimen

loans, facilities for work at their herbaria, or information on specimens in their collections, and Tom Zanoni

for helpful discussions. Dave Murray and Bruce Ford provided useful comments on an earlier version of

the manuscript.

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Alma-Ata. Pp. 10-131.

Benson, L. 1948. A treatise on the North American Ranunculi. Amer. Midl. Naturalist 40:1—261.

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CHEREPANOV, S.K. 1981. Sosudistye Rasteniia SSSR [Plantae vasculares URSS]. Akademiia nauk SSSR, Leningrad.

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HORANDL, E., O. Paun, J.T. JOHANSSON, C. LEHNEBACH, T. ARMSTRONG, L. CHEN, AND P. LOCKkHART. 2005, Phylogenetic relation-

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Phylogen. Evol. 36:305-327.

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Carolum Cnobloch, Leipzig.

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Spach, E. 1839. Ranunculus. In: Histoire naturelle des vegetaux : Phanerogames, vol. 7. Librairie encyclopédique

de Roret, Paris. Pp. 203-220

TUMOKHINA, S.A. 1992. Ranunculus. In: Flora Sibiri vol. 6, ed. L.E. Malysheva and G.A. Peshkovoy. Science Press,

Novosibirsk. Pp. 99-198, 264-282.

Turin, T.G. AND J.R. AkeroYD. 1993. Ranunculus. In: Flora Europaea vol. 1, 2nd edition, ed. T.G. Tutin, VH. Heywood,

N.A. Burges, D.H. Valentine, and D.M. Moore. Cambridge University Press, Cambridge, UK. Pp. 269-286.

VonosHiLOv, V.N. 1982. Opredelitel' rasteniy Sovietskogo dalnego vostoka. Science Press, Moscow.

WANG, W.-C. AND M.G. GILBERT. 2001. Ranunculus. In: Flora of China vol. 6, ed. Z.-G. Wu and PH. Raven. Science Press,

Beijing. Pp. 391-431.

WHITTEMORE, A.T. 1997. Ranunculus. In: Flora of North America vol. 3, ed. Flora of North America Editorial Commit-

tee. Oxford University Press, New York. Pp. 88-135.

INFRAGENERIC TAXONOMY OF ASTROPHYTUM (CACTACEAE),

WITH REMARKS ON THE STATUS OF DIGITOSTIGMA

Richard R. Montanucci

Department of Biological Sciences

Clemson University

Clemson, South Carolina 29634-0314, U.S.A.

ABSTRACT

In a monograph of the genus Astrophytum, Megata (1944) proposed two infrageneric sections—Astrophytum sect. Austrastrophytum and

ag pik + 1 e. A cf 1 4. + A 2 i PAI 2 1 Pb. T x I S

ect ji j j yp of the

DAT Leal A 1 f dpa ll a 1 ; laid 1 f.L $ zn ESL P

CBN TE a rF de

unaltered. Backel (1950) pror ! b Astrophyt bg. E hyt d LA ae Neoastrophytum hich essentially

led to M f However. Backehers sÁ subs usina, ining ype st g violated Art.

22.2 a Art. 21.3 E the international ae

n 1961, Backeberg used N } in hi ftl but proposed Astrop) bg. Astrophy-

tum to replace A. subg. nasal: Terion, Backeberg's (1961) subgeneri lid, and! ived wide circulation, and

should be accepted; O two sections should be O he basis of ive fl 1 frui hol the species

ma oe sae Ve ee o & Nevárez appears to have evolved baee pue genis ri A ale and id a idest m

to A be } A. subg. Astrophytum. However, } habit, dimorphic

us pid fusiform root, and ly ci basal cleft and ), caput-medusae's pl in Ástro-

1 Ty LA LE LN y 1 1 11 1 4 221 y

PI J gng g )

a a pssebls Méleculas geneti di led to further clarify tl lutionary relationships of A. caput-medusae

RESUMEN

EL fin Jal nA A poni p Y i (10445 3 ` a a Av A bar Á +. Tan

g j; Megata (1944) prop g p y

A. sect. S iastrophyt A } Austrastrophyt i l tipo del género viola el Art. 22.2 del ICBN

L E Es r t r eo L A

A Anei&n d ^ Sas 1 ting del sé T ^ PE a

subdivisión g q y I» g g Bac-

keberg aie prop d bg s: Astrophyt bg. Euastrophytum y A a K I ial t d

l Megata. Sin M A subo Puas ropi Backers, ti la especie tipo del género, y viola los Art. 22. 2 y Art.

213 Ba iid international.

Lak 1 1 AT 1 7 Jal A l 1 phy-

tum para seme A. subg Fussropytum. Por tanto, los Epa ca an dn de Backeberg aen son válidos, s seda amplia

circulación g I flor y fruto, la

especie Desine caput-medusae V &N hal lucionad l género A strophytum j muestra una TAMEN más

grande co u n Á. subg Asiriphytum Sin embargo, debid

bg 1

ae ende sdmáriss raíz ancha mue y

ill 1 iid 1 1

testa verrucosa), el emplazamiento

o ES J 1 r Ya

hihrid Avi ^ do c It A; Ati 1 1 1 Lie 1 1 : Ta E}

O o

A. caput-medusae.

SUBGENUS VS. SECTION

The cactus genus Astrophytum comprises a small group of species occurring principally in the Chihuahuan

Desert region of central and northern Mexico, with one species entering southern Texas (Anderson 2001;

Hunt et al. 2006). On the basis of hybridization experiments involving the different species, Móller (1927)

recognized two ly natural groups icu the ies A decade later, the German horticulturalist, Curt

Backeberg (1937), p N a brief, d ipti ies of Astrophytum in Blåtter für Kakteenforschung,

a bulletin of cactus research. He also eee two PM within the genus—Group A having apical fruit

dehiscence and yellow flowers, and Group B with basal fruit dehiscence and red-throated flowers.

Moritane! Megata, a student of horticultural sciences at the Kyoto Imperial University in Kyoto, Japan,

'Moritane, i i i M i D

L L

ia.

J. Bot. Res. Inst. Texas 3(1): 251 — 256. 2009

252 t tanical Í i Texas 3(1)

published the first monographic treatment of the genus Astrophytum in 1944. Data from experimental

hybridization and comparative morphology, led him to propose two infrageneric sections—Astrophytum

sect. Septentriastrophytum (diagnosis: Flores floribus Sect. Austrastrophyti majores, aurea praeter intus basi

rubra; fructus carnosus, irregulariter dehiscens; semina copiosa, 80—300, nigra vel luteobrunnescentia.), and A.

sect. Austrastrophytum (diagnosis: Flores minores quam flores Sect. Septentriastrophyti, lutei et non rubric intus

ad basim; fructus siccus, stellatim dehiscens; semina 40—60, nigra). Megata recognized five species, placing A.

myriostigma and A. ornatum in A. sect. Austrastrophytum and A. asterias, A capricorne, and A. coahuilense in

A. sect. Septentriastrophytum.

In 1950, Curt Backeberg published two subgenera under Astrophytum which corresponded exactly to

Megata's (1944) sections. He popes? DU subg. Mn (diagnosis: Fauce lutea; fructu apice

stellariter dissiliente) with the type st iostigma Lem. (1839) and A. subg. Neoastrophytum

(diagnosis: Fauce rubra; fructu basi dissiliente), ai the type species Echinocactus asterias Zuccarini (1845).

Backeberg's six volume series Die Cactaceae was published in 1958 t01962, and the genus Astrophytum

was treated in his volume 5 (1961). Backeberg proposed Astrophytum subg. Astrophytum to replace A. subg.

Euastrophytum, and adopted his previously published A. subg. Neoastrophytum. He apparently realized that

Astrophytum subg. Euastrophytum violated Art. 22.2 of the ICBN which requires that a name of a subdivision

a a Te that includes the type of that genus repeat as its epithet the generic name unaltered, and therefore

should have been named Astrophyt bg. Astrophytum. Furthermore, Art. 21.3 forbids the

epithet of a subdivision of a genus to be ed from “Eu” plus the generic name. Backeberg separated the

two subgenera on the basis of fruit morphology, type of fruit dehiscence, and seedling color. He recognized

a total of six species, placing A. myriostigma and A. ornatum in Astrophytum subg. Astrophytum, and A. asterias,

A. capricorne, A. coahuilense, and A. senile in A. subg. Neoastrophytum. He followed the same taxonomy in

the first (1966), second (1970) and third (1976) editions of Kakteenlexikon (as well as the English version of

the third edition, Cactus Lexicon 1977). However, he abbreviated the subgeneric diagnoses, using only the

mode of fruit dehiscence as a diagnostic trait.

The fundamental dichotomy within the genus Astrophytum was also recognized by the Astrophytum

specialist Otakar Sadovský who carried out long-term hybridization experiments involving hundreds of

plants. However, he and hi thors did not give formal taxonomic recognition, either at the level of section

or subgenus, to these two clades (Haage & Sadovský 1957; Sadovsky & Schútz 1979). More recent evidence

from chloroplast DNA seems to further support this basic division within Astrophytum (Wallace 1995, Fig. 5).

As is evident from the foregoing discussion, essentially the same two groups of species in the genus

Astrophytum have been treated in the past both as sections and as subgenera by two different authors, creat-

ing a taxonomic problem requiring resolution. Furthermore, this problem has persisted in recent works.

In The New Cactus Lexicon (Hunt et al. 2006: 31), Backeberg's Astrophytum subg. Astrophytum and A. subg.

Neoastrophytum are recognized as well as A. subg. Stigmatodactylus D. Hunt. By contrast, in a new mono-

graphic treatment of Astrophytum, Hoock (2008) allocates five species to Megata's (1944) sections, A. sect.

Austrastrophytum and A. sect. Septentriastrophytum, and places a sixth species, Astrophytum caput-medusae

(Velazco & Nevárez) D. Hunt, in A. subg. eL A D. Hunt, following Hunt (2003a, b).

The decision whether to divide a g subgenera is subjective. In reviewing the question

in this case, it appears that the use el the subgenus is more appropriate and justifiable than the use of the

section for the following three reasons: 1) Megata's (1944) A. sect. Austrastrophytum is in violation of ICBN

Art. 22.2 because this section includes the type (Astrophytum myriostigma) of the generic name; it should

have been named A. sect. Astrophytum. 2) Backeberg's two subgeneric names, A. subg. Astrophytum and A.

subg. Neoastrophytum, are validly published and have received wide circulation, appearing in volume 5 of

his Die Cactaceae and in all three German editions of Kakteenlexikon as well as the English translation of his

third edition. 3) Hunt (2003a, b) proposed A. subg. Siiemalidaciyius to accommodate the morphologically

divergent A. caput-medusae, thereby setting further y for the use of the subgenus category as opposed

to the section, and as indicated above, the Bibscnus category was adopted in The New Cactus Lexicon.

Montanucci, | fi ict PP Peer »

STATUS OF DIGITOSTIGMA

Velazco and Nevárez (2002) described a new genus and species, Digitostigma caput-medusae from Nuevo León,

Mexico. Subsequently, Hunt (20032) wrote: "The authors of Digitostigma caput-medusae are to be congratu-

lated on a truly remarkable discovery, but the feeling of our group is that its peculiar habit is not sufficient

in itself to justify excluding it from Astrophytum, with which it shares unusual and significant features in

common, i.e., the floccose indumentum and eccentric (hat-shaped) seeds, as well as having very similar

flowers.” Hunt (2003a) therefore placed Digitostigma in the synonymy of Astrophytum and recombined the

species as Astrophytum caput-medusae D. Hunt.

Shortly thereafter, Hunt (2003b) wrote that Kanchi Gandhi of Harvard University had determined that

Velazco and Nevárez “had failed to indicate the type of the generic plant in the explicit manner required

for the ICBN,” thus invalidating both the name Digitostigma and the species name. Gandhi regarded Hunt

(20032) as the valid publication of the name Astrophytum caput-medusae as well as the subgeneric name, A.

subg. Stigmatodactylus. But according to Hunt (2003b), Nigel Taylor argued that Velazco and Nevárez had

failed to provide a validating Latin diagnosis for the species name n dde E original s and

Hunt's revision were invalid. To correct this error, Hunt (2003b) y ] formal

iol Á 1 l n Jie It il " LET,

“Astrophytum caput-medusae D. Hunt ab ali

Mexico, edo. Nuevo León, [locality withheld], ‘atonal espinoso Cuda 100-200 n m, 28 Aug 2001, Nevárez & Velaz

(UNL 023704 holo.; UNL 023705 iso.). Syn: Digitostigma caput-medusae Velazco & Nevárez, Cact. Suc. Mex. 47(4):81-82 cl

nom inval. (Art. 43.1)”

lus D. Hunt Replaced Digitostigma \ Ex Nevárez Cact. Suc. Mex. 47(4):79 (2002) nom inval. (Art.

37. 5), Buon. descr Lat. Type: Astrophytum caput-medusae D. Hunt."

Then in the editorial section of Cactaceae Systematics Initiatives 20:4. 2005, we are informed that Roberto

Kiesling and Detlev Metzing proposed a clause to ICBN Art. 37, that was approved by the International

Botanical Congress in Vienna (July 2005). The clause removes the "pitfall" introduced when said Article

was previously amended in 1990 which invalidated the generic name Digitostigma Velazco & Nevárez when

originally published, as well as the specific names D. caput-medusae Velazco & Nevárez and Astrophytum

caput-medusae (Velazco & Nevárez) D. Hunt. The draft Art. 37.7 states: "In the case of a new monotypic

genus (or monotypic infrageneric taxon above the rank of species), the correct mention of, or reference to,

the type of the species name is sufficient." As a result of the acceptance of the clause to Art. 37, the original

names in Cactaceas y Suculentas Mexicanas 47(4):76—86. 2002, and the recombination in Astrophytum (Cac-

taceae Systematics Initiatives 15:6. 2003a) retroactively became validly published, and the "validation" of A.

caput-medusae (Cactaceae Systematics Initiatives 16:4. 2003b) was judged superfluous. Consequently, if authors

prefer to classify the new taxon as a distinct genus, the correct name would be Digitostigma caput-medusae

Velazco & Nevárez. But if it is to be regarded as a member of Astrophytum, then the correct name would be

Astrophytum subg. Stigmatodactylus caput-medusae (Velazco & Nevárez) D. Hunt, and Digitostigma would be

placed in the synonymy of Astrophytum.

Detailed morphological comparisons between caput-medusae and Astrophytum have been lacking until

now. In an attempt to clarify caput-medusae's affinities and generic status, a summary of my comparative

study is presented below. Flowers, fruits, and seedlings of the following taxa were examined (number of

flowers, fruits, adult plants, and seedlings in parentheses): A. ornatum (3, 3, 2, 69), A. myriostigma (22, 41,

41, 249), A. asterias (29, 5, 5, 59), A. capricorne (40, 11, 14, 82), A. coahuilense (16, 20, 20, 120), and the spe-

cies caput-medusae (34, 8, 5, 14).

The flower of caput-medusae has a vivid orange color at the base of the inner perianth segments (Velazco

& Nevárez 2002) and in this respect it is similar to that of A. subg. Neoastrophytum species in which the

inner perianth segments vary from crimson to orange. By contrast, members of A. subg. Astrophytum have

entirely yellow flowers. The fruit of A. subg. Neoastrophytum is described as fleshy and turns pink, red, or

reddish purple when ripe, whereas that of A. subg. Astrophytum is less fleshy and remains green (Megata

1944; Hoock 2008). The fruit of caput-medusae is similar to that of A. subg. Neoastrophytum in that it usually

254 Journal of tani Insti Texas 3(

turns pink at maturity (Hoock 2008); but at maturity it appears less fleshy to nearly dry and in this respect

is similar to the fruit of A. subg. Astrophytum. The fruit of caput-medusae undergoes longitudinal dehiscence

(Velazco & Nevárez 2002) and hence differs from both Astrophytum subgenera. According to Velazco and

Nevárez (2002:82), the remains of the perianth are deciduous and leave a 3 mm diameter scar. However, in

all fruit that I observed the floral remnants persisted until dehiscence. In A. subg. Neoastrophytum the floral

remnants are strongly attached to the fruit, but in A. subg. Astrophytum they usually detach from the fruit

at the time of dehiscence, leaving a small scar.

The flower and fruit of caput-med di imilarities in particular with those of Astrophytum asterias.

In both taxa, the exterior perianth ta are Dosen dis and they usually lack, or have few, black

tips. In A. subg. Neoastrophytum capricorne and A. subg. Neoastrophytum coahuilense the exterior perianth

segments are usually yellow with black tips, and often there is a reddish line or spot near each black tip. The

bases of the inner perianth segments in caput-medusae are vivid orange; in A. asterias the color may vary from

dark red to pink or orange. In A. capricorne and A. coahuilense, the bases of the inner perianth segments are

usually dark crimson, but may vary from pale red to orange, or rarely, pure yellow in natural populations

(Hoock 2008). In A. asterias the mature fruit turns pink (Hoock 2008; this study) like that of caput-medusae.

Furthermore, in caput-medusae, the receptacle tube and pericarpel are covered with fewer lanceolate scales

and more white wool than in the species of both Astrophytum subgenera, except in A. asterias which displays

a further reduction in number and size of these scales (i.e., spinescent bracteoles) and more wool, especially

on the pericarpel.

Velazco and Nevárez (2002) stated that the seeds of D. caput-medusae are “hat-shaped” and compare

favorably with those of Astrophytum, and according to Hunt (20032) they resemble those of A. asterias.

However, my study suggests that the seeds of all Astrophytum species are very similar to one another and

differ collectively from those of D. caput-medusae in several ways. The seeds of all Astrophytum species have

a smooth testa whereas those of D. caput-medusae are verrucose (Velazco & Nevárez 2002). The seeds of

Astrophytum clearly fit the concept of “hat-shaped,” with a peaked, but rounded profile opposite the hilum

and a rather flat base (where the concavity of the hilum is situated). But the seeds of caput-medusae have a

deeply and acutely angled cleft at the concavity of the hilum and the lateral profile opposite the hilum is

especially enlarged and rounded.

The seedlings of caput-medusae are usually light green, but they can be dark violet in color. Seedlings

of all members of A. subg. Neoastrophytum are typically light green, whereas those of A. subg. Astrophytum

are generally russet or reddish brown, rarely green.

The distributional relationships of the taxa under consideration are also informative. Astrophytum subg.

Astrophytum has a southerly distribution with A. myriostigma centered on the plateau of San Luis Potosi and

adjacent areas including the Jaumave Valley, and A. ornatum occurring through the barranca region as far

south as the state of Hidalgo (Megata 1944; Hoock 2008). On the other hand, A. subg. Neoastrophytum has

a northerly distribution, ranging from extreme northern Zacatecas and adjacent Durango, northward and

eastward through Coahuila and adjacent Nuevo León into southern Texas (Hoock 2008). The species caput-

medusae also has a northern distribution, being restricted to northern Nuevo León (Velazco & Nevárez 2002)

adjacent to the range of A. capricorne and A. asterias.

In sum, comparative data from flower and fruit characteristics, and seedling color support the hypoth-

esis that caput-medusae is a member of the genus Astrophytum. Moreover, the evidence (including geographic

relationships) suggests that caput-medusae is more closely related to A. subg. Neoastrophytum than to A. subg.

Astrophytum and may have evolved from the former group after it had separated from the latter. Therefore,

accepting Digitostigma as a distinct genus to accommodate caput-medusae would render the genus Astro-

phytum a paraphyletic taxon, which would be undesirable in a classification system based on phylogenetic

principles. It is clear, however, that caput-medusae has undergone considerable morphological divergence

from the other Astrophytum species, possibly as a result of strong selection pressures for adaptation to the

Tamaulipan thornscrub habitat, perhaps as a shrub debris mimic.

Infrag > £ Ae EE 255

Montanucd,

As an alternative hypothesis, the kable di Iting in the unusual suite of morphological

characteristics (tuberculate habit, dimorphic areoles, leue jum. root, and large rounded seeds with a

verrucose testa and acute basal cleft) could be explained by intergeneric hybridization, a possibility previ-

ously raised by Hunt (2003a). If true, separate generic status for caput-medusae would then be justified. 1f

caput-medusae is an ancient hybrid, it is reasonable to assume that a member of A. subg. Neoastrophytum was

involved as one of its parents. The other parent could have had a pure yellow flower. One characteristic of the

flower of caput-medusae that suggests hybridization is the narrowness of zone of orange color surrounding

the stamens. A narrow zone of color is typical of Astrophytum hybrids produced artificially by crossing a pure

yellow-flowered species (e.g., A. myriostigma) with a species with red-throated flowers (e.g., A. capricorne, A.

asterias); see Hoock (2008) for examples. Without doubt, molecular genetic studies are needed to further

clarify the evolutionary relationships of caput-medusae.

CONCLUSIONS

It is recommended that A. sect. Austrastrophytum Megata nom. inval., (Art. 22.2), and A. sect Septentrias-

trophytum Megata should be suppressed. Backeberg's Astrophytum subg. Astrophytum and A. subg. Neoas-

trophytum should be accepted and used when necessary. The species Digitostigma caput-medusae Velazco &

Nevárez appears to have evolved within the genus Astrophytum, and shows a closer relationship to A. subg.

Neoastrophytum than to A. subg. Astrophytum. However, because of its aberrant morphology, caput-medusae's

placement in A. subg. Stigmatodactylus D. Hunt appears justified, though provisional; its status as an ancient

intergeneric hybrid remains a possibility. Molecular genetic studies are needed to resolve this question.

Protologues and a diagnostic key to the three subgenera follow:

Astrophytum subg. Astrophytum Backebg., Die Cactaceae 5:2655, fig. 2526. 1961. Subg. Euastrophytum Back-

ebg., Cac. Succ. J. (Los Angeles) 22:5. 154. 1950. Tyre species: Astrophytum myriostigma Lem., Cact. Gen. Sp. Nov. 4(-6). 1839. Tree

LOCALITY: not cite

F 1 ; f i llariter dissili . Distribution.—North-c ico (B berg 1950:154).

Diagnosis. —Frucl formig aufspri d, mit festerer Wand, Samlinge Mns nd 1961: 2653).

Note.—Distribution is central Mexico: not Bund central Mexico.

Astrophytum subg. Neoastrophytum Backebg., Cac. Succ. J. (Los Angeles) 22:5. 154. 1950. Tre species: Echi-

nocactus asterias Zucc., Abh. Math.-Phys. Cl. Kónigl. Bayer. Akad. Wiss. 4(2):13, t. 3. 1845. Tere LocaLrrY: Mexico.

I f asi dissiliente. Distribution — Central Mexico.

agnosis.

Note.—Distribution is north-central Mexico, not central Mexico.

Astrophytum subg. Stigmatodactylus D. Hunt, ae ae Init. pee 6. Us REPLACED SYNONYM: Digitostigma

Velazco & Nevárez, Cact. Succ. Mex. 47:79. 2002. Tyre tostig put-medusae , Cact. Suc. Mex. 47:81-82.

‘Digitostigma’ not only contravenes Linnaean canons for the formation of generic names but implies that

the plant is notable for its finger-shaped stigmas rather than its spotted tubercles, contrary to the authors'

explicit intention. For this reason the opp ity afforded by the change of rank has been taken to provide

an unambiguous name formed according to classical usage.

idée: caput-medusae (Velazco & Nevárez) D. Hunt, Cactaceae Syst. Init. 15:6. 2003. Digitostigma

medusae Velazco & Nevárez, Cact. Suc. Mex. 47:81-82. ir: MEXICO. Nuevo León: [locality withheld], matorral espinoso

a 100-200 m, 28 Aug 2001, Nevárez & Velazco s.n. (noLorvee: UNL 023704; isotype: UNL 02

Radix fusif di darians edens a ratione E pane aerea mpm dE brevis, So costas

iis

ES : +15] 1; in radiales vel

centrales non diferentes breves, non diia rectae, , aliquantum undulata, albidae, ad spiceum atrocastaneae. e dimorphae:

spinifera ad tuberculi apicem, albolanata ,in d ta. Flores in

li evolventium, non apices diurni, avi n g is 1 ioril cian aurantiaca, segmentis

“1 n sos as 1 EE Ec] latis napiracels CEA ERU PE 2115 Sel E

Eu L r E L E

lbi ientil i llo isd i ito. F j is viridis, carnosus isl latis ad axillis lanatis, ad matu-

Es i > , , L

ritate sicco, dehiscentia l linali i lari. Semi (A hyti similia), ad 3 mm longa, pileata 1 1 nigra vel

atrobrunnea; hilo basali profundissi icropyle ab hilo ext , sed ad id iguo (Vel Sr Nevárez 2002:79)

£l n.a + al D Lt "T £T.

256 Journal of Texas 3(

KEY TO SUBGENERA

: Fruits remain green, open apically; fl yellow subg. Astrophytum Backebg. (A. myriostigma, A ornatum)

. Fruits turn Bu. red or red- -purple when ripe, split basally | in penes fashion, disintegrate basally, or split

longitudinally in irregular usually with orange or red throat.

2. Body with ribs subg. Neoastrophytum Backebg. (A. asterias, A capricorne, A. coahuilense)

2. Body with elongate tubercles subg. Stigmatodactylus D. Hunt (A. caput-medusae)

ACKNOWLEDGMENTS

I wish to thank Nicholas J. Turland of the Missouri Botanical Garden for clarifications of the pertinent

articles of the ICBN. I also thank A. Michael Powell for reading and commenting on a previous draft pi this

N Urs Eggli (ZSS) and an anonymous reviewer provided helpful reviews. Detailed pl

d herbarium preparations of Astrophytum caput-medusae were obtained a ies Hund

and CA Hinton, us

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Backesenc, C. 1958-1962. Die Cactaceae. 6 volumes. VEB Gustav Fisher Verlag, Jena, Germany.

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Haace, W. AND O, SADOVSKY. 1957. Kakteen-Sterne, die Astrophyten. Neumann Verlag, Radebeul.

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IIl. Cactaceae Syst. Init. 15:1-2, 5-6.

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Digitostigma. Notulae systematicae lexicon cactearum spectantes Ill. Cactaceae Syst. Init. 16:4.

Hunr, D. 2005. Editorial: rule-changes validate Digitostigma. Cactaceae Syst. Init. 20:4.

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MYCOLOGICAL LITERATURE ON TEXAS FLESHY BASIDIOMYCOTA,

TWO NEW COMBINATIONS, AND NEW FUNGAL RECORDS FOR TEXAS

David P. Lewis Clark L. Ovrebo

262 CR 3062 Department of Biology

Newton, Texas 75966, U.S.A. University of Central Oklahoma

plewis@jas.net Edmond, Oklahoma 73034, U.S.A.

covrebo@uco.edu

ABSTRACT

A D 1: 1 J Pi : 13 hcl 3 7 c a sft cf]

basidiomycet-

ous fungi described from Texas. This search resulted in 202 articles on Texas fungi ih 89 new taxa described from Texas. We also

report on 52 new records of fleshy basidiomycetous fungi from Texas and propose two new combinations, Hygrocybe chamaeleon

and Hygrocybe mississippiensis.

RESUMEN

Texas. Esta bú da di Í i

nuevos taxa para Texas C primera vez. También I quí 52 pecies de hong basidiomi para Texas,

yala ei I 2 78 P rr

PART I——MYCOLOGICAL LITERATURE ON TEXAS FLESHY BASIDIOMYCOTA

David Lewis has been collecting fungi in southeast Texas for over 30 years. During this time he has at-

tempted to amass the literature that documents the fleshy Basidiomycota throughout Texas. Reported here

are the results of the literature search as conducted by both authors. The authors’ primary research interests

center around gilled mushrooms and boletes (Agaricales sensu lato — but see Part III of this paper for a newer

classification) so perhaps the list below is most complete for the fungi of these groups. We have, however,

attempted to locate references for other fleshy basidiomycetous fungi.

For a reference to be included in the list, a collection must be cited from Texas. For modern references

this is not difficult to discern because of the exacting requirements of locality information and herbarium

designations. pe that give the distribution as “as far as Texas” or something similar without docu-

mentation of coll not included. A little more liberty is yielded to the literature of the nineteenth or

early twentieth centuries where geographic information might not be as complete, nor is there a guarantee

that a herbarium specimen exists.

Following each reference is a number in parenthesis. The number indicates how many species, with at

least one Texas record, are reported in the article. For taxa that have the holotype from Texas, the binomi-

als and localities (sometimes only the state name was given in the protologue) where collected are provided

beneath the references, followed by the herbarium location of the holotype. We have relied on literature

references and online herbaria data to determine the location of the holotypes, but we have not confirmed

their continued existence in the designated herbaria. For a few of the older taxa we were not able to deter-

mine the location of the types. We are aware that a number of the newly described taxa have since been

nomenclaturally or taxonomically synonymized, but we feel that for the purposes of this paper it is best to

give only the basionym as the starting reference.

RESULTS

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J. Bot. Res. Inst. Texas 3(1): 257 — 271. 2009

258 ainia

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Murrill, W.A. 1907. Polyporaceae. N. Amer. Fl. 9(1):1-72. (3)

Murrill, W.A. 1908. Polyporaceae. N. Amer. Fi. 9(2):73-132. (8)

Inonotus juniperinus Murr. (HoLoTvee: TEXAS)

Pyropolyporus texanus Murr. (HoLoTYee: TEXAS; BPI)

Murrill, W.A. 1915. Southern polypores. Published by the author, New York. 66 p. (7)

Murrill, WA. 1920. Light-colored resupinate polypores Il. Mycologia 12:299-308. (1)

Nakasone, KK. Ms H.H. Burdsall, Jr. 1995. Phlebia species from the eastern and southeastern United States. Mycotaxon

54:335-359

Naohiko, S. 1984. a "Corpse Finder” Mcllvainea 6:7—9. (1)

Overholts, L.O. 1967. The Polyporaceae of the United States, Alaska, and Canada. The University of Michigan Press, Ann Arbor.

466 p. (55)

Ovrebo, C.L. 1988. Notes on the cultural characters, morphology and distribution of Ripartitella brasiliensis. Mycotaxon

: (1

229-237

Parmasto, E. 2001. Hymenochaetoid Fungi e of North America. ray 79:107-1 76. (7)

Pegler, D.N. 1983. The genus Lentinus—a world monograph. Kew Bull. Addit. Ser. X:1-281. (1)

Petersen, R.H. and KW. Hughes. 2004. A preliminary monograph of Lentinellus (R lales). Bibl, Mycol. 198:1—270. (1)

Punugu, A., M.T. Dunn, and A.L. Welden. 1980. The peniophoroid fungi of the West Indies. ors 10:428-454. (1)

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of the Atti del Convegno Internazionale; 1984, Sept. 10-15. “Borgo Val di Taro, Italy Centro Ed per la Flora Mediterranea.

p. 175-199. (1)

Redhead, S.A, J. Ginns, and R.A. Shoemarker. 1987. The Xerula (Collybia, Oudemansiella) radicata complex in Canada. Myco-

taxon 30:357-405. (1)

Restivo, J.H. 1971. A checklist of the Agaricales and BL Me id of the Stephen F. Austin Forest and the Nacogdoches,

Texas area [M.S. Thesis]. Stephen F. Austin State y, Nacogdoches, Texas. 49 p. (80

Ryvarden, L. 1984. Type studies in the Polyporaceae 16. Species Pace by J.M. Berkeley, either alone or with other mycolo-

gists from 1856 to 1886. Mycotaxon 20:329-364. (1)

Ryvarden, L. 1985. Type studies in the Polyporaceae 17. Species described by W.A. Murrill. Mycotaxon 23:169-198. (3)

Ryvarden, L. 1992. Type studies in the Polyporaceae-23. Species described by C.G. Lloyd in Lenzites, Polystictus, Poria and

Trametes. N n 44:127-136. (1)

Shaffer, R.L. 1964. The subsection Lactarioideae of Russula. Mycologia 56:202-231.

Shaffer, R.L 1970. Notes on the subsection als and other species of m Lloydia 33:49-96. (1)

Shaffer, R.L. 1970. Cuticular terminology in Russula les). Brittonia 22:230-239. (1)

Sims, K.P, R. Watling, and P. Jeffries. 1995. A revised kB to the genus Mae a Mycotaxon 56:403-420. (1 )

Singer, R, J. Garcia, and L.D. Gómez. 1990. The Boletineae of Mexico and Central America land Il. Beih. Nova Hedwigia 98:1-77. (2)

Singer, R., J. García, and L.D.Gómez. 1991.The Boletineae S and Central America Ill. Beih. Nova Hedwigia 1 102:1-128. (4)

Singer, R, C.L. Ovrebo, and DP. Lewis. 1990. A new species of Melanomphalia (Crepidotaceae). Mycotaxon 37:47-52. (1)

d R, CL. Dn 2s i 7 id eee d es of Phylloporus and Tricholomopsis from Colombia, with notes on

9. (1)

Smith A H. 1937. Studies on the genus Mycena M. Mycologia 29: s 354. (1)

Mycena texensis A.H. Smith (HoLorvre: TEXAS. Eastland Co.: Cisco; MICH)

Smith, A.H. 1939. Studies in the genus Agaricus. Pap. Mich. Acad. sa = 138. (2)

Agaricus ciscoensis A.H. Smith (HoLotyre: TEXAS. Eastland Co.: Cisc

Smith, A.H. 1944. New North American agarics. Mycologia 36: ee E a

eastlandensis A.H. Smith (HoLorvrE: TEXAS. Eastland Co.: Cisco; MICH)

Agaricus vinaceo-umbrinus A.H. Smith (HoLorvrE: TEXAS. Eastland Co.: Cisco; MICH)

Smith, A.H. 1947. North American species of Mycena. University of Michigan Press, Ann Py 521 p. (2)

Smith, A.H. 1960. Uh halle (Agaricales) in the Western Hemisphere. Brittonia 12:41—70. (2)

Smith, A.H. 1972. The North American species of Psathyrella. Mem. New York Bot. Gard. a -633. (7)

Psathyrella ids Smith (HoLotyre: TEXAS. Hidalgo Co.: Mission; MICH)

Psathyrella texensis A.H. Smith (uctorvre: TEXAS. Montgomery Co.: Richards; MICH)

feat aed thiersii A.H. Smith IA odd Sey a a MICH)

mith, A d L.R.Hesler. 1962. 5 ius IIl f section Plinthogali. Brittonia 14:369—440. (1)

Lactarius Meu var. texensis Smith & Hesler var. nov. (HoLoTyreE: TEXAS. Meal tgomery Co.: Richards; MICH)

Smith, AH. and L.R. Hesler. 1968. The North American species of Pholiota. Hafner Publ. Co., New York. 402 p. (2)

Smith, AH. and R. Singer. 1964. A monograph on the genus Galerina Earle. Hafner Publ. Co, New York. 246 p. (1)

Lewis and Ovrebo, Texas fleshy Basidiomycota 263

Smith, A.H. and H.D. Thiers. 1964. A contribution toward a monograph of North American species of Suillus. Privately published,

Ann Arbor. 166 p. (2)

Smith, H.V. and N.S. Weber. 1982. Selected species of L prinus fi | tl tern United States. Contr. Univ. Michigan

Herb. 15:297-309. (3)

Leucocoprinus breviramus H.V. Smith & Weber (HoLoryrE: TEXAS. Brazoria Co.: Liverpool; MICH)

Leucocoprinus magnicystidiosus H.V. Smith & Weber (HoLoTYPE: TEXAS. Brazoria Co.: Liverpool; MIC

Smith, H.V. and N.S. Weber. 1987. Observations on Lepiota americana and some related species. Contr. Univ. Michigan Herb.

16211-221.(

Lepiota besseyi H.V. Smith & Weber (uotorvre: TEXAS. Brazoria Co.: Lake Jackson

Stevenson, J.A. and E.K. Cash. 1936. The new fungus names proposed by C. e ba Bull. Lloyd Libr. Mus., Bull. No. 35, Mycol

Ser. No. 8. 209 p. (13

Thiers, H.D. 1956. The Agaricaceae of the pine belt and adjacent areas in eastern Texas [Ph.D. Thesis]. University of Michigan,

Ann Arbor. 568 p. (205)

Thiers, H.D. 1956. A preliminary report of the agaric flora of eastern Texas. Texas J. Sci. 8:257-263. (37)

Thiers, H.D. 1957. The agaric flora of Texas |. New species of agarics and boletes. Mycologia 49:707-722. (12)

Amanita alba Thiers. (Hototyee: TEXAS. Brazos Co.: College Station; MICH)

Amanita salmonea Thiers (notorvee: TEXAS. Brazos Co.: College Station; MICH)

Boletus roseibrunneus Thiers (HoLotyre: TEXAS. Grimes Co.: Navasota; MICH)

Boletus silvaticus Thiers (uoLorvee: TEXAS. Grimes Co.: Navasota; MICH)

Cystoderma texensis Thiers (HoLoTvre: TEXAS. Montgomery Co.: Richards; MICH)

Lactarius albolutescens Thiers. (HotLorvre: TEXAS. Montgomery Co.: id MICH)

Lactarius albus Thiers (noLoryrE: TEXAS. Walker Co.: Huntsville; MIC

Lactarius cystidiosus Thiers (HoLoTYPE: TEXAS. San Jacinto Co.: subi Lake; MICH)

eburneus Thiers (HoLoTYPE: TEXAS. Walker Co.: Huntsville;

Lactarius echinatus Thiers (HOLOTYPE: TEXAS. Walker Co.: Huntsville; MICH)

Lactarius mordax Thiers (HoLotyPE: TEXAS. Grimes Co.: Navasota; ipe

Lactarius obnubiloides Thiers (noLorvre: TEXAS. Brazos Co.: Millican; MICH)

Thiers, H.D. 1958. The agaric flora of Texas II. C taxa of white- and pink-spored agarics. Mycologia 50:5 14—523. (9)

Collybia delicata Thiers (HoLoTyPE: TEXAS. Brazos Co.: Wellbor

s iin Thiers ebd TEXAS. San bus Co.: cling MICH)

Marasmius puniceus Thiers (uotorvre: TEXAS. Brazos Co.: Millica

ee reai var. texana Thiers var. nov. e TEXAS. Brazos Co.: College Station; MICH)

Mycena radicata Thiers (HoLoryrE: TEXAS. Montgomery Co.: Richards; MICH

Mycena avellaneibrunnea Thiers (noLor*PE: TEXAS. San Jacinto Co.: Coldspring; MICH)

Mycena epipterygia var. Pise Thiers var. nov. (Hotorvee: TEXAS. Montgomery Co.: Richards; MICH)

Rhodophyllus verrucosus Thiers (HoLorvpE: TEXAS. Brazos Co.: Wellborn; pen

richolomopsis squamosa Thiers (notorvee: TEXAS. Walker Co.: Huntsville; MICH)

us H.D. 1959. The agaric fee ded lil. New taxa of brown- and e agarics. Mycologia 51:529-540. (11)

Agaricus placomyces var. flavescens Thiers var. nov. (noLoTYPE: TEXAS. Brazos Co.: Wellborn; MICH)

Conocybe antipoda var. humicola Thiers var. nov. (HoLorvee: TEXAS. Brazos Co.: Wellborn; MICH

Coprinus comatus var. texensis Thiers var. nov. (HoLoTYrE; TEXAS. Montgomery Co.: near San Jacinto River, MICH)

Coprinus subangularis Thiers (HoLorvrE: TEXAS. Montgomery Co.: Richards; MICH)

Coprinus subcurtus Thiers (nororyre: TEXAS. Brazos Co.: Millican; MICH)

Cortinarius acystidiosus Thiers (HoLoTYPE: TEXAS. San Jacinto Co.: M xus MICH)

Cortinarius paliformis Thiers. (oLoTyPE: TEXAS. Walker Co.: Huntsvi

Galerina autumnalis f. robusta Thiers f. nov. GHoLotyre: TEXAS. a Co.: Richards; MICH)

Gymnopilus luteoviridis Thiers. (HoLoryrE: TEXAS. Brazos Co.: near Navosata River, College Station; MICH)

Inocybe texensis Thiers (HoLoryPE: TEXAS. Walker Co.: Huntsville; MICH)

Psathyrella hymenocephala var. farinacea Thiers var. nov. (HoLoTYPE: TEXAS. Ni o Co.: Richards; MICH)

Thiers, H.D. 1959. Notes on the genus Boletus in Texas. Texas J. Sci. 11:314-319.

Thiers, H.D. 1963. The bolete flora cf the Gulf Coastal Plain. |. The ie J. Elisha Mitchell Sci. Soc. 79: 32-41. (5)

Torrwa-Torres, j G. and L. Guzmán-Dávalos. 2008. Taxonomic status and new localities for | . Mycotaxon

103:33-40.

T VE. and A. Smith. 1963. Protoalkaloids of Panaeolus species. o Physiol. Alkaloide Halle 21:47-54. (1)

aeolus texensis Tyler & A.H. Smith (Hororvre: TEXAS. Eastland Co.: Cisco; MICH)

Ui. RE. and D.P. Lewis. 1994. Amanita westii-taxonomy and AUR A rare species from ing the Gulf o

Mexico. Mycotaxon 50:131-138. (1)

264 i it Texas 3(

i (LI | FL Ax Essi 1 | i£ J;

n C. and M. Rajchenberg. 1999. Two North American Hy , Aphyllor

na. Mycotaxon 72:417-422. (1

m VS. 1901. The Tylostomaceae of North America. Bull. Torrey Bot. Club 28:421-444. (3)

White, V.S. 1902. The Nidulariaceae of North America. Bull. Torrey Bot. Club 29:251-280. (1)

Wright, J.E. 1987. Two new species of Tulostoma (Gasteromycetes) from Texas. Mycologia 79:155-157. (2)

Tulostoma clathrosporum Wright (koLorvee: TEXAS. Eastland Co.: Cisco; SFSU).

Tulostoma thiersii Wright (HoLoTYee: TEXAS. Kimble Co.: near Junction; BAFC).

Zeller, S.M. 1943. North American species of Galeropsis, Gyrophragmium, Longia, and Montagnea. Mycologia 35:409-421. (2)

DISCUSSION

We fear and predict that we have missed references for Texas because distribution reports are often found

in obscure journals. We welcome feedback on citations that we may have missed and it is anticipated that

a follow-up article with updated references will likely be forthcoming.

In addition to technical articles where collections are cited, there is additional literature in the format

of field guides that include fungi found in Texas. Important is the field guide by Metzler and Metzler (1992)

which includes 202 species primarily from southeastern Texas. Other useful field guides for the area include

those by Bessette et al. (2000), Bessette et al. (2007), Horn et al. (1993), Lincoff (1981), Phillips (2005) and

Weber and Smith (1985).

PART II—NEW COMBINATIONS

1 d on morphol-

The following binati di ] with modern g

ogy (Singer 1986) and supported by ala evidence (Matheny 2006). During the process of preparing

this manuscript and nim ont in the protologue (Cibula 1979) for Hygrophorus chamaeleon, it

was not in order. In the protologue,

came to our attention that tion regarding the type 1

the type collection information for chamaeleon was cited as follows:

"Specimen typicum Cibula n. 589 in Herb. Mass conservatum est; legit prope Saucier, MS, 8 August 1974."

The first thing noticed was that the date for 589 was inconsistent with other numbers in the 500' for col-

lections cited in the Materials Studied, and 589 was not again cited in the Materials Studied. Checking no.

589 in Cibula's field record book, which we have, 589 is listed as a Russula, and the date of that collection

is 9 Oct. 1975. No Hygrophorus species were collected on that date. Unfortunately, we have not been able to

confirm the whereabouts of 589 if it still exists as a collection. Checking the other collection numbers from

Materials Studied of chamaeleon in Cibula's record book, he has *type" written next to coll. no. 489 and 489

is also cited in the Materials Studied. Upon examination of collection no. 489, “type spec." is written on the

associated hand-written field label. The field label, record book and the original 5 x 8 in card with macro-

scopic description and collecting data all give the date for no. 489 as 20 Aug. 1974. Based on evidence from

the field book and field label, Cibula had intended to designate 489 as the holotype, but somehow mistyped

the collection information and date in the Latin description. We can find no violation of any articles in the

International Code of Botanical Nomenclature (McNeill et al. 2006) for this error. We correct this error and

conclude that Cibula coll. no. 489 is the holotype. Given below are the new combinations including the

corrected type information for H. chamaeleon.

Hygrocybe chamaeleon (Cibula) Lewis & Ovrebo, comb. nov. Basiony: Hygrophorus chamaeleon Cibula, Mycotaxon

10:109. 1979. HoLorvre: MISSISSIPPI. Harrison Co.: Saucier, Harrison Experimental Forest, De Soto National Forest, in humus

along broadleaf evergreen draw across from Block II Plot 1, loblolly pine (north of plot), Fertilization Study Plots, 20 Aug 1974,

WG. Cibula 489 (NY).

Hygrocybe mississippiensis (Cibula) Lewis & Ovrebo, comb. nov. Basionvm: Hygrophorus mississippiensis Cibula, My-

cotaxon 10:106. xdi HOLOTYPE: MISSISSIPPI. Harrison Co.: Saucier, Harrison Experimental Forest, De Soto National Forest, on

a raised mound of t and in leaf litter in mixed bottomland hardwoods, adjacent to Block IV, Plot 4, loblolly pine,

Fertilization Study Plots, 8 Aug 1974, WG. Cibula 485 (NYD.

The field label gives the habitat for H. mississippiensis as a mixed pine/hardwood ravine. Hygrocybe mississippiensis was previously

Lewis and Ovrebo, Texas fleshy Basidiomycota 265

reported from Texas (Lewis & Cibula 2000). The hol llecti fH. ch l 1 H. mississippiensis were originally deposited

at MASS, they are now at NY.

PART III—NEW RECORDS OF FLESHY FUNGI FOR TEXAS

The following records are a result of long-term efforts by D.P Lewis and C.L. Ovrebo of documenting the

mycota of southeast Texas, through organized amateur and professional forays, and individual outings. This

listing represents only a small beginning to the updating of fungus records for Texas and reflects only col-

lecting done in southeast Texas. As is the case with all biodiversity studies, knowledge of the biodiversity of

a region is a function of the efforts that have been mounted to conduct field work, and importantly, records

must be based on vouchered collections. With the exception of the researches of Harry Thiers and David

Lewis, no other mycologists have devoted equally the time and effort in cataloguing in a scientific way the

fleshy mycota for this region. For this reason, our knowledge of the diversity of fungi for southeast Texas,

let alone the entire state of Texas, is far from complete.

The orders and families listed below may not be the ones readers are most familiar with. Until recent

years, families DE m n and boletes were included in the Agaricales oe Singer ioi The order and

family ci hanged significantly as a result of recent pl lar analyses and

now Duc apice fungi and other fleshy fungi maybe placed in different orders and families to reflect

a more natural classification system. Most of the classification hierarchy below is taken from Matheny et al.

(2006) and the reader is directed to that entire Mycologia issue devoted to recent developments in elucidat-

ing our understanding of phylogenies of fungi.

BASIDIOMYCOTA: AGARICOMYCETES: AGARICALES

AGARICACEAE

Agaricus ee cn in a ee on pu

nta Fe, Santa F

29? 23.049", W 95? 06.383" 24 Jan 2008, DPL-8493 (F )

Agaricus pocillator Murr. Found in mixed pine-hardwood forest.

Voucl i Hardin Co.: Big Thicket National P Jack B 1 Unit, off FM 2937, 9 Nov 2002, DPL-6647 (F). Orange

y) 1

grounds, field of 6* Street and Warpath Street, N

Co.: Vidor, near 455 Virginia Lane, 15 Jan 1989, DPL-4232 (F).

CORTINARIACEAE

Cortinarius aureifolius Peck. Found in an arid sandyland with Pinus palustris and oaks. (Fig. 1).

Voucher specimens: Newton Co.: Lewis Chapel community, Sand Ridge Cemetery, CR 4045, N 30° 44.580', W 93° 46.127', 19 Dec

2007, Ovrebo 4718 & 4719 (CSU).

Cortinarius corrugatus Peck. Under Quercus laurifolia and in beech-oak fore

Voucher specimens: Newton Co.: Bleakwood, 262 CR 3062 & State Hwy 87, N 30° 42.509', W 93° 49.630', 19 Oct 2004, DPL-6959

(F). Mayflower, Canyon Rim Nature Trail, N 31° 06.927', W 93° 43.752', 27 Oct 2004, DPL- 7049 (E); 29 Jul 2005, DPL-7286 (F).

Cortinarius hesleri Ammirati & A.H. Smith. Found in hardwood bottomland and oak- -hickory forests.

Voucher specimens: Newton Co.: Toledo-Bend, Sam Forse Collins Recreational Area, N 31? 10.029, W 93° 34.620, 20 Jul 2007, DPL-

129 (F). Tyler Co.: Big Thicket National Preserve: Neches River Corridor Unit, Round Lake area, 13 Sep 2003, DPL-6807 (P).

Cortinarius Meu. (L.) S.E Gray. Found in older pine forests.

Voucher specimens: Hardin Co.: Big Thicket National Preserve, Turkey Creek Unit, Kirby Nature Trail, 3 Dec 1978, DPL-1670 (F);

Jack Gore Baygall Unit, Arco Oil Field Road, 28 Nov 1980, DPL-2451 (E); Jack Gore Baygall Unit, near FM 2937, 30 October 1993,

DPL-5092 (F); 9 Nov 2002, DPL-6645 (F); 27 Nov 2004, DPL-7104 (F); Saratoga, Lance Rosier Unit, Teel Road, N 30* 15.629", W

94? 31.532!, 7 Nov 2006, DPL-7608 (TAES); 11 Nov 2006, DPL-7643 (TAES). Newton Co.: Bleakwood, 262 CR 3062 & State Hwy.

87, N 30? 42.509', W 93? 49.630', 1 Nov 1997, DPL-5962 (F).

ENTOLOMATACEAE

Clitopilus prunulus (Scop.) P Kumm. Found in floodplains, oak and pine forest.

Voucher specimens: Hardin Co.: Big Thicket National Preserve, Saratoga, Lance Rosier Unit, Cotton Road, 10 Jul 1988, DPL-4135 (F).

Newton Co.: Bleakwood, 262 CR 3062 & State Hwy 87, N 30? 42.509', W 93° 49.630', 10 Jun 2001, DPL-6459 (E); 18 Jun 2003,

DPL-6687 (F); 12 Sep 2007, DPL-8274 (F).

Clitopilus hobsonii (Berk. & Br.) PD. Orton. Found on rotten wood. Previously known from Europe, Florida

and North Carolina.

L Dat . ID l.l bitit, f T,

266 Journal of t Texas 3(

Voucher specimen: Hardin Co.: Silsbee, residence on Pine Street, 31 Jul 1988, DPL-4163 (F).

Entoloma megacystidiosum Hesler. Found in mixed pine-hardwood and beech forests, previously known

e type in Tennessee.

Voucher specimens: Hardin Co.: Big Thicket National Preserve, Lance Rosier Unit, Siam Cotton Road, 13 Jun 2007, DPL-7906

(F); Turkey Creek Unit, Kirby Nature Trail, 22 Jul 2007, DPL-8141 (F). Newton Co.: Mayflower, Canyon Rim Nature Trail, N 31°

06.927", W 93° 43.752', 3 Jul 2007, DPL-8084 (FE); 20 Jul 2007, DPL-8122 i

HYGROPHORACEAE

Hygrocybe chamaeleon (Cibula) Lewis & Ovrebo. Found in moss and mixed pine and hardwood forests,

previously known from Harrison Co., Mississippi. (Fig. 3).

Voucher specimens: Hardin Co.: Big Thicket National Preserve, Saratoga, Lance Rosier Unit, Cotton Road, 13 Jun 2007, DPL-7911 (F);

42 (F).

Hygrophorus borealis f. salmoneus Coker. Known only from the type in North Carolina. Found in an

Hone pine | icis next i o grandifolia.

2 CR 3062 & State Hwy 87, N 30? 42.509', W 93? 49.630', 24 Nov 2008, DPL-8815 (F).

Hygrophorus caespitosus (Mur) Me Found in hardwood forests.

Voucher specimens: Newton Co.: Bleakwood, 262 CR 3062 & State Hwy 87, N 30° 42.509', W 93? 49.630", 28 Jun 2003, DPL-6716

(E); 5 in Pad DPL-6742 (F). Tyler Co.: Temple-Inland Forest Lake Experimental Forest, Quercus alba test plot 26 Jun 1994,

DPL-530

Hygrophorus A (Fr.) Fr. Found with Pinus taeda.

Voucher Specimens: Jasper Co.: Buna, E -Inland Black Branch Hunting Club, off CR 701 and FM 1004, 24 Dec 1991, DPL-4578

(F). Newton Co.: Lewis Chapel C Sand Ridge Cemetery, CR 4045, 30 Dec 2006, DPL-7798 (F). Tyler Co.: Big Thicket

National Preserve, Turkey Gek Unit, Müscadine Road, 15 Dec 1979, DPL-2147 (F).

Hygrophorus perplexus A.H. Smith & Hesler. Found in mixed pine and hardwoods.

Voucher specimen: Hardin Co.: Big Thicket National Preserve, Saratoga, Lance Rosier Unit, Teel Road, N 30? 15.629', W 94° 31.532',

11 Nov 2006, DPL-7623 (TAES).

HYMENOGASTRACEAE

Hebeloma sarcophyllum (Peck) Sacc. Found with Quercus alba and mixed pine-hardwood forests.

Voucher specimens: Newton Co.: Bleakwood, 262 CR 3062 & State Hwy e N 30? 42.509', W 93? 49.630', 13 May 2000, DPL-6265

(F). Tyler Co.: Temple-Inland Forest Lake E ] Forest, Q lot, 15 Jun 1994, DPL-5258 (F).

Hebeloma sinapizans (Fr) e [oum in mixed d md hardwood forests.

y Road, CR 4045, near cemetery, N 30° 44.572', W 93°

46. 094 21 Dec 2007, DPL-8438 (F). Tyler Co.: Big Thicket National CA Canyonlands Unit, near Spurger, N 30? 43.241, W

94? 08.404', 20 Dec 2007, DPL-8463, DPL-8464 (F).

LYOPHYLLACEAE

Tephrocybe palustris (Peck) Donk. Found in sphagnum moss

Voucher specimens: Orange Co.: Vidor, near Virginia Lane, 10 May 1979, DPL-1724, DPL-1725 (F); 22 Feb 1986, DPL-3954 (F); 28

Feb 1990, DPL-4355 (F).

MARASMIACEAE

Marasmius fulvoferrugineus Gilliam. Found in leaf litter.

Voucher specimens: Hardin Co.: Big Thicket National Preserve, Lance Rosier Unit, Saratoga, Teel Rd, N 30? 15.629", W 94? 31.532',

11 Nov 2006, DPL-7681 ee 19 Jul 2007, DPL-8107 (F); Turkey Creek Unit, Kirby Nature Trail, 16 Jun 2007, DPL-7989 (F).

Newton Co.: Bleakwood, 262 CR 3062 & State Hwy 87, N 30° 42.509', W 93° 49.630', 20 Nov 2004, DPL-7092 (F).

OMPHALOTACEAE

Gymnopus luxurians (Peck) Murr. Subcaespitose on pine mulch

Voucher specimen: Jefferson Co.: Beaumont, campus of Lamar University near John Gray Institute, 10 Jul 2006, DPL-7494 (F).

PLUTEACEAE

Volvariella pusilla (Pers.) Sing. Found on soil, with mixed pine and hardwoods.

Voucher specimen: Newton Co.: Bleakwood, 262 CR 3062 & State Hwy 87, N 30° 42.509', W 93° 49.630', 31 Aug 2003, DPL-6770

F).

Volvariella taylori (Berk. & Br.) Sing. Found in a dried up cypress swamp.

Voucher specimen: Hardin Co.: Big Thicket National Preserve, Saratoga, Lance Rosier Unit, Teel Road, N 30° 15.629', W 94° 31.532',

17 Jun 2006, DPL-7422 (F).

Lewis and Ovrebo, Texas fleshy Basidiomycota 267

Lo no

yrella rugocephala (G.F Atk.) A.H. Smith. Found in a hardwood forests.

Ed specimens: Fort Bend Co.: Brazos Bend State Park, Hale Lake Nature Trail, 18 Nov 2006, DPL-7759 (E TAES).

Psathyrella carbonicola A.H. Smith. Reported from the western U.S. and Michigan. On burned ground.

Voucher specimen: Newton Co.: Bleakwood, 262 CR 3062 & State Hwy 87, N 30? 42.509', W 93? 49.630', 27 Nov 2008, DPL-8818

STROPHARIACEAE

Hypholoma sublateritium (Fr.) Quél. Found on a well decayed hardwood stump.

Voucher specimen: Newton Co.: Bleakwood, 262 CR 3062 & State Hwy 87, N 30? 42.509', W 93? 49.630', 28 Nov 2008, DPL-8823

(F)

Phaeomarasmius curcuma (B. & C.) Sing. Found on wood (Fig. 2

Voucher specimens: Newton Co.: 3leakwood, 262 CR 3062 & State Hwy 87, N 30? 42.509', W 93° 49.630', 24 Dec 1999, DPL-6214

(F); 21 Nov 2007, Ovrebo y (CSU). Orange Co.: Vidor, near 455 Virginia Lane, 12 Apr 1977, DPL-915 (E); 14 May 1977, DPL-

949 (E); 17 May 1977, DPL-955 (F); 23 May 1977, DPL-973 (F); 21 Apr 1978, DPL-1361 (F); 4 May 1978, DPL-1372 (F); 29 Dec

1981, DPL-2856 (F)

Psilocybe caerulescens Murr. Found in a Pinus plantation. Known from Alabama, central Mexico, Venezuela

and Brazil

Voucher specimen: Tyler Co.: Big Thicket National Preserve, Spurger, in pine pl i Neches Ri orridor Unit, 13 Sep 2003,

DPL-6800 (E, XAL).

TRICHOLOMATACEAE

Clitocybe trulliformis (Fr.) P. Karst. Found in pine duff.

Voucher Specimen: Newton Co.: Bleakwood, 262 CR 3062 & State Hwy 87, N 30? 42.509', W 93? 49.630', 28 Jan 2001, DPL-6400

(F).

Tricholoma nii uis jd on a iue: nd soil with pines and oaks (Fig. 4

Voucher specim n Co.: Lewi munity, Sand Ridge Cemetery, CR 4045, N 30? 44.572', W 93? 46.094', 30 Dec

2006, M od MA 21 Dec 2007, pu s p (Fig. 4)

BOLETALES

Boletellus Ma is (Snell) Sing. Found in mixed pine-hardwood forests.

Voucher specimens: Hardin Co.: Big Thicket National Preserve, Turkey Creek Unit, Kirby Nature Trail, Big 19 Sep 1980, DPL-2314

(F); Jack Gore ul Unit, e FM 2937, 4 Jul 1981, DPL-2664 (F); Lance Rosier Unit, Cotton Road, near Saratoga, 28 May 1984,

DPL-3656 (F); 5 Aug 1984, DPL-3697 (F). Tyler Co.: Big Thicket National Preserve, Beech Creek Unit, Nature Trail, 4 Aug 1982,

DPL-3241 (F); Spurger, near Neches River, 10 Aug 1982, DPL-3276 (F

Boletus auripes Peck. Found with Quercus alba, Quercus fal cata var. leucophylla, Quercus nigra and other

oaks.

Voucher specimens: Hardin Co.: Big Thicket National Preserve, Jack Gore Baygall Unit, off FM 2937, 16 Aug 1982, DPL-3328 (E);

Saratoga, Lance Rosier Unit, Cotton Road, 5 Aug 1984, DPL-3700 (F). Tyler Co.: Temple-Inland Forest Lake Experimental Forest,

Sawtooth oak (Quercus sp.) test plot, 9 Jun 1992, DPL-4635 (F); Quercus alba test plot, 12 Jun 1992, DPL-4636 (F); Quercus falcata

geen test plot, 21 Jun 1992, DPL-4699 (F); Quercus nigra test plot, 7 Jun 1997, DPL-5783 (F).

Boletus led ii A.H. Smith & Thiers. Found in stream floodplain forest.

Voucher specimen: Newton Co.: Bleakwood, 262 CR 3062 & State Hwy 87, N 30? 42.509', W 93? 49.630', 1 Jul 1996, DPL-5672

(P).

Boletus hypocarycinus E Found in floodplains, oak and ue forest.

Voucher m Newton Co.: Bleakwood, 262 CR 3062 & State Hwy 87, N 30? 42.509', W 93? 49.630', 19 Oct 2004, DPL-6966

wis Chapel community, Sand Ridge Cemetery Road, CR 4045, 12 Jul 2006, DPL-7519 (F). Burkeville, along FM 692, 31 Jul

o DPL-8205 (F). Orange Co.: Vidor, near 455 Virginia Lane, 10 Oct 1993, DPL-5035 (F). Tyler Co.: Dam-B, residence on

CR 4136, off FM 92, N 30° 48.811', W 94° 12.600', 10 Jul 1995, DPL-5498 (F). Temple-Inland Forest Lake Experimental Forest,

Quercus alba test plot, 7 Aug 1995, DPL-5573 (F). Big Thicket National Preserve, Beech Creek Unit, south boundary road, 14 Jul

2006, DPL-7530 (F)

Boletus longicurvipes Snell & A.H. Smith. Found in mixed pine-hardwood forests, near baygall communi-

de andi in Ben ps

Big Thicket National Preserve, Jack Gore Baygall Unit, Arco Oil Field Road, 1 Aug 1982, DPL-3233 (F);

Sarton Lance Rosier Unit, ce Road, 20 Oct 1984, DPL-3750 (B; bd pim Unit, Kirby Nature Trail, 27 Oct 1984, DPL-3767

Co.: Conroe d, Big Rons C , N 30? 15.326', W 95? 25.481', 28 Jul 2007, DPL-8176 (F).

Newton Col Blessed. 262 CR 3062 & State Hwy 87, N 30? 42.509', W 93? 49.630', 26 Aug 2001, DPL-6518 (E); 24 Jul 2007,

DPL-8151 (F). Orange Co.: Vidor, near 455 Virginia Lane, 30 Aug 1979, DPL-1916 (F); 15 Aug 1982, DPL-3317 (F).

3/17)

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Lewis and Ovrebo, Texas fleshy Basidiomycota 269

Boletus morrisii Peck. Found in beech hammocks next to baygall communities.

Voucher specimens: Hardin Co.: Big Thicket National Preserve, Jack Gore Baygall Unit, Arco Oil Field Road, 5 Aug 1983, DPL-3584

(F); 25 Sep 1983, DPL-3617 (F).

Boles rufomaculatus Tan Poon in mixed e forests.

d, 262 CR 3062 & State H 30° 42.509', W 93° 49.630', 7 October 2004, DPL-6929 (F).

nitie

Tylopilus « eximius (Peck) Sing. Found in beech hammocks next to baygall commun:

Voucher specimens: Hardin Co.: Big Thicket National Preserve, Jack Gore Baygall Unit, Arco Oil Field E 25 Jun 1983, DPL-3532

(F); 5 Aug 1983, DPL-3585 (F), 25 Sep 1983, DPL-3618 (E); 18 May 1984, DPL-3651 (F); off FM 2937, 26 Jul 1985, DPL-3878 (F).

Nacogdoches Co.: Angelina National Forest, Stephen Austin Experimental Forest, 17 Sep 1994, DPL-5338 (F).

Xanthoconium Ln Lond Snell & Dick. Found in bottomland hardwood forests.

Voucher specimens: Newton Co.: Bleakwood, 262 CR 3062 & State Hwy 87, N 30? 42.509', W 93? 49.630', 1 Jul 1996, DPL-5677

Rhizopogon baxteri A.H. Smith. Known from Michigan and Mississippi.

Voucher specimen: Hardin Co.: Big Thicket National Preserve, Lance Rosier Unit, Saratoga, Teel Road, 15 Nov 2008, DPL-8800 (F).

CANTHARELLALES

Craterellus odoratus (Schwein.) Fr. Found in mixed pine and hardwood forests.

Voucher specimens: Hardin Co.: Big Thicket National Preserve, Saratoga, Lance Rosier Unit, Teel Road, N 30? 15.629', W 94° 31.532!,

un 2007, DPL-7904 (F). Newton Co.: Bleakwood, 262 CR 3062 & State Hwy 87, N 30? 42.509', W 93° 49.630',18 Jul 2003,

DPL-6758 (F). Polk Co.: Big Thicket National Preserve, Big Sandy Unit, off CR 1071, 24 Jul 1976, DPL-445 (F). Tyler Co.: Fred,

residence on FM 1943, 8 May 1976, DPL-23 (F).

HYMENOCHAETALES

Inonotus amplectans Murr. Found on living branch of Asimina parviflora.

Voucher specimens: Newton Co.: Lewis Chapel Community, Sand Ridge Cemetery, CR 4045, 30 Jul 2006, DPL-7553 (F); 21 Jul 2007,

DPL-8134 (F).

POLYPORALES

Laetiporus persicinus (B. & C.) Gilbertson. Found at the base of tree/dead stump.

Voucher specimens: Hardin Co.: Big Thicket National Preserve, Lance Rosier Unit, Saratoga, Teel Rd, N 30° 15.629', W 94? 31.532, 17

Jun 2006, DPL-7450 (TAES, PC); Turkey Creek Unit, Kirby Nature Trail, 22 Jul 2007, DPL-8145 (F).

Ecco rome Furio oy ) a A mo De found on small branches and twigs.

Unit, Saratoga, Teel Road, 15 Nov 2008, DPL-8797 (F); Big

Thicket National Preserve, Tuckey Creek Unit, Kirby Nature Trail, 16 Jun 2007, DPL-7999 (F); 22 Jul 2007, DPL-8144 (F); Lumberton,

Village Creek State Park, Nature trail, 23 Jun 2001, DPL-6479 (LSU). Newton Co.: Bleakwood, 262 CR 3062 & State Hwy 87, N

30° 42.509', W 93° 49.630', 11 Jul 2000, DPL-6342 (F)

RUSSULALES

PODOSCYPHACEAE

Cymatoderma caperatum (Berk. & Mont.) D.A. Reid. A tropical species found on wood.

Voucher specimens: Hardin Co.: Big Thicket National Preserve, Lance Rosier Unit, Saratoga, Teel Road, N 30? 15.629', W 94° 31.532,

13 Jun 2007, DPL-7905 (F). Harris Co.: Pasadena, Armand Bayou Nature Center, 17 Nov 2007, DPL-8325 (TAES).

RUSSULACEAE

Lactarius argillaceifolius Hesler & A.H. Smith. Found in mixed pine and hardwood forests.

Voucher specimens: Hardin Co.: Big Thicket National Preserve, Saratoga, Lance Rosier Unit, Teel Road, N 30? 15.629', W 94° 31.532!

11 Nov 2006, DPL-7670 (TAES). Newton Co.: Bleakwood, 262 CR 3062 & State Hwy 87, N 30° 42.509', W 93° 49.630', 7 Sep

1996, DPL-5709 (F); 29 Oct 1998, DPL-6116 (F).

pan Brevis bius Found in mixed pine and hardwood forests.

ig Thicket National Preserve, Lance Rosier Unit, Cotton Road, near Saratoga, 15 Oct 1983, DPL-3633

(P.N loches Co.: Angelina National Forest, Stephen E Austin Experimental Forest, 17 Sep 1994, DPL-5339 (E); 24 Sep 1994,

DPL-5340 (P. Newton Co.: Bleakwood, 262 CR 3062 & State Hwy 87, N 30? 42.509', W 93? 49.630', 17 Sep 1996, DPL-5339

(F); 4 November 1998, DPL-6121 (P); 11 Nov 2004, DPL-7083 (TAES). San Jacinto Co.: Sam Houston National Forest, Big Creek

Scenic Area, 10 Oct 1998, DPL-6096 (F). Tyler Co.: Town Bluff, residence off FM 92, 29 May 1994, DPL-5206 (F).

ME ctas Ma Found in mixed pine and hardwoods forests.

l d, 262 CR 3062, N 30? 42.509', W 93? 49.630', 30 August 1998, DPL-6071 (F); 1 Jun 2004,

DPL-6874 (P).

270 tani i Texas 3(

Lactarius gerardii Peck. Found in hardwood forests.

Voucher specimens: Hardin Co.: Kountze, by Beaumont Creek, 12 Aug 1982, DPL-3293 (F). Newton Co.: Bleakwood, 262 CR 3062 &

State Hwy 87, N 30° 42.509', W 93° 49.630", 28 Jun 2003, DPL-6727 (F); 17 Jun 1999, DPL-6187 (F); 23 Oct 2004, DPL-6983 (F).

Tyler Co.: Big Thicket National Preserve, Neches River Corridor Unit, Round Lake, 13 Sep 2003, DPL-6789 (F)

Lactarius imperceptus Beards. & Burl. Found in mixed pine and hardwoods.

Mosi RC: Angelina Co.: Angelina National Forest, FR 313A, 2 Jan 2007, DPL-7811 (F). Hardin Co.: Big Thicket National

rve, Jack Gore Baygall Unit, near FM 2927, 28 Oct 1979, DPL-2080 (F); Turkey Creek Unit, Kirby Nature Trail, 21 Dec 1982,

Dum (F); Big Thicket National Preserve, Saratoga, Lance Rosier Unit, Teel Road, N 30? 15.629', W 94? 31.532', 11 Nov 2006,

DPL-7672 (TAES). Jasper Co.: Buna, Temple-Inland Black Branch Hunting Club, road 2, off CR 701, off FM 1004, 5 Nov 1993,

DPL-5096 (F); 13 Nov 1993, DPL-5116 (F). Newton Co.: Bleakwood, 262 CR 3062 & State Hwy 87, N 30° 42.509', W 93° 49.630',

1 Nov 1997, DPL-5964 (F); 9 Nov 1997, DPL-5969 (F); 18 Dec 2006, DPL-7781 (F); Mayflower, Canyon Rim Nature Trail, 31.06.927

N, 93.43.752 W, 18 Dec 2007, DPL-8415 (F). Polk Co.: Big Thicket National Preserve, Big Sandy Unit, Sunflower Road, 29 Nov

1985, DPL-3932 (F). Tyler Co.: Big Thicket National Preserve, Turkey Creek Unit, off FM 1943, 15 Dec 1979, DPL-2144 (F);

tesa Inland Forest Lake Experimental Forest, Quercus nigra test plot, 24 Nov 1995, DPL-5594 (F); Big Thicket National Preserve,

yonlands Unit, N 30° 06.927', W 93? 43.752', 20 Dec 2007, Ovrebo 4720 (CSU) & DPL-8444 (F).

ns lignyotus var. canadensis A.H. Smith & Hesler. Found in sphagnum moss bog.

Voucher specimen: Newton Co.: Bleakwood, 262 CR 3062 & State Hwy. 87, N 30° 42. 509', W 93? 49.630', 27 Sep 2003, DPL-6824

(F).

Lactarius peckii Burl. Found in hardwood floodplain forests and with Quercus alba.

Voucher specimens: Hardin Co.: Big Thicket National Preserve, Jack Gore Baygall Unit, Timber Slough Road, 18 Jul 1981, DPL-2735

F wton Co.: Bleakwood, 262 CR 3062 & State Hwy. 87, N 30? 42.509', W 93° 49.630', 16 Jul 1995, DPL-5518 (F); 19 Jun

1996, DPL-5641 (E); 29 Oct 1998, DPL-6115 (F). Tyler Co.: Big Thicket National Preserve, Turkey Creek Unit, off FM 1943, 9 Aug

1981, DPL-2778 (F); Temple-Inland Forest Lake Experimental Forest, Quercus nigra test plot, 13 Jun 1993, DPL-5011 (F); 22 Jun

1996, DPL-5651 (E); 7 Jun 1997, DPL-5788 (F); Quercus alba test plot, 30 May 1994, DPL-5225 (E); 22 Jul 1995, DPL-5550 (F); 7

Jun 1997, DPL-5787 (F).

SEBACINALES

Sebacina concrescens (Schwein.) P. Roberts. Found in mixed pine and hardwood forests on a vine

Voucher specimen: Hardin Co.: Big Thicket National Preserve, Saratoga, Lance Rosier Unit Teel Road, N 30? 15.629', W 94? 31.532',

19 Jul 2007, DPL-8114 (P).

CORTICIALES

Punctularia strigosozonata (Schwein.) Talbot. Found on hardwood logs.

Voucher specimens: Hardin Co.: Big Thicket National Preserve, Turkey Creek Unit, Kirby Nature Trail, N 30? 27.738', W 94? 21.015',

un 2008, DPL-8670 (F). Newton Co.: Bleakwood, 262 CR 3062 & State Hwy. 87, N 30? 42.509', W 93? 49.630", 13 Jan 2008,

DPL-8491 (F); Mayflower, Canyon Rim Nature Trail, N 31? 06.927", W 93° 43.752', 18 Dec 2007, DPI -8427 (F).

ACKNOWLEDGMENTS

We thank the Big Thicket National Preserve for granting permission to collect in the BE Tee We are

grateful to Scott Redhead, Ronald H. Petersen, and John McNeill for providing

advice; Roy E. Halling and The New York Botanical Garden for loaning the Hygrophorus type collections;

Gastón Guzmán for confirming the identification of Psilocybe caerulescens; Sean Jones and Juan Luis Mata

who assisted in various ways during the preparation of this manuscript; and Juan Luis Mata and Andrew

Methven for reviewing the manuscript and providing helpful comments.

REFERENCES

Besserre, A.E., W.C. Roopy, AND A.R. Bessette. 2000. North American boletes-a colored guide to the fleshy pored

mushrooms. Syracuse University Press, New York.

Besserre, A.E, W.C. Roopy, A.R. Bessette, AND D.L. Dunaway. 2007. Mushrooms of the southeastern United States.

Syracuse University Press, New York.

Ciputa, W.G. 1979. Fungi of the Gulf Coast I. Two new species of Hygrophorus section Hygrocybe. Mycotaxon

10:105-115.

Horn, B. R. Kay, AND D. ABeL. 1993, A guide to Kansas mushrooms. University Press of Kansas, Lawrence.

Lewis and Ovrebo, Texas fleshy Basidiomycota 271

Lewis, D.P. ano W.G. Cura. 2000. Studies on Gulf Coast agarics (Basidiomycota: Agaricaceae); notes on some

interesting and rare species. Texas J. Sci. 52 (Suppl):65-78.

Lincorr, G. 1981. Audubon field guide to North American mushrooms. Alfred Knopf, New York.

Matheny, PB. (AND 24 OTHERS). 2006. Major clades of Agaricales: a multilocus phylogenetic overview. Mycologia

98:982-995.

McNett, J., ER. Barrie, V. DevouUuN, D.L. Hawksworth, K. MARHOLD, D.H. NicoLson, J. PRADO, PC. SiLva, J.E. Scoc, J.H. WIERSEMA,

AND N.J. TURLAND, eds. 2006. International code of botanical nomenclature (Vienna Code). Gantner Verlag,

Rugell, Liechtenstein.

METZLER S. AND V. METZLER 1992. Texas mushrooms. University of Texas Press, Austin.

PHILLIPS, R. 2005. Mushrooms and other fungi of North America. Firefly Books, Ltd., Buffalo, New York.

SiNGER, R. 1986. Agaricales in modern taxonomy. 4" ed. Koeltz Scientific Books, Koenigstein.

WEBER, N.S. AND A.H. SmiTH. 1985. A field guide to southern mushrooms. University of Michigan Press, Ann Arbor.

£ 4L D sont D hinetiéné £T

272 Journal of Texas 3(

BOOK REVIEW

Wi Gites. 2007. Encyclopedia of Exotic Pants for Temperate Climates. (ISBN 978-0881927856, hbk.).

Timber Press, 133 S.W. Second Ave., Suite 450, Portland, Oregon 97204-3527, U.S.A. (Orders: www.

timberpress.com, 1-800-827-5622, 1-503-227-3070 fax). $49.95, 440 pp., 8 3/4" x 11 1/4".

The Victorian era and Victorians ptivated by the mysterious world of s exotic.

With their bold foliage, ] l and luscious scents, p d 19th century curiosity and thrilled Victorian

senses. Fortunately for g in tl ld” I g it y in tl pi d operate a gree

1 2 11 s r lala 1

The E Lio Exotic Plants for Temperate Cli

inde HOME dude us o exotics can os persuaded to flourish in he mikel

who covet their

est nds He iun ae that are subire ical or t lly originate in cooler regions, as well as true

r rer 4 5

Pe |

I E g

15 i d culti described in th yclopedia, covering many plant groups, including aroids, bananas,

e. Det iler Ve z va EC LI A fA4n 1 1 1 f 1

ore p

gingers, bromeliads, cacti, yuccas, ferns, palms, and mor

Lei f. 1 ` 1 1 141 i qm n

ever o

hi igins, microclimates

f. f,

1 1 1 : 1 A dde

Kingdom and North A i i I ing I

the text and collected in a bibliography. A sources are supplied as well.

This book is richly illustrated ne over a color bd Most on; dis pos are Mee un a that Tu a oa A

ecommended

good number are half-page size or larger. Thei

for public libraries and research collections. It will l to all who enjoy l in their gardening.

Will -o a woes exotic E enthusiast, dus xin his own din Exotic Garden in an unlikely location: England's

east coa t Will has appeared on a wide range of British

ene propius i cues regularly to UK tadio shows and mapacines Ofivinally trained as an artist, he has worked as an

i he New Exotic Garden (2000).—Gary L. Jennings, Librarian, Botanical Research Institute

illustrator for I

of Texas, Fort Worth, Texas 76102-4025, U.S.A.

J. Bot. Res. Inst. Texas 3(1): 272. 2009

TWENTY-EIGHT NEW THREE-LETTER FAMILY ACRONYMS FOR

VASCULAR PLANTS (WITH COMPREHENSIVE LISTINGS ON-LINE)

Neil 5now

nidi Pacificum

vus Museum, 1525 Bernice Street

nolulu, Wee 817-2704 U.S.A.

cendi nda

ABSTRACT

Three-letter acronyms are proposed oe 28 newly M iex d i vascular plants. Complete listings alphabetically by family, and

at: |

by acronym, are posted online

RESUMEN

c Ami J 1 ^o L “|: J 1 1 Tios 3 A LE TL t dd 2:15

vasculares y

de eae tan di ibl li I jj " 1f/Plant%20Keys/ACROS edt

The use of three-letter acronyms (TLA) for plant families can expedite many routine aspects of curation

(Weber 1982; Snow € Holton 2000; Brasher & Snow 2004a, b). TLA are used by some institutions in as-

sociation with plant specimen labels, herbarium folders, filing systems, data bases, and plant checklists

(Weber & Wittmann 1992; Snow 2009). If combined with a unique namespace, each TLA has the potential

to be used as part of a globally unique identifier (GUID) for that family in bioinformatics applications.

Recent publications and online sources that synthesize the coverage of vascular plants at the family

level for angiosperms (Stevens 2001; Heywood et al. 2007; Haston et al. 2007; Mabberley 2008) and ferns

(Smith et al. 20062, b) have begun to recognize families for which TLA previously were unavailable. This

paper proposes 28 additional TLA for those families and integrates them into a comprehensive listing al-

phabetically by spelling of the family name and by three-letter acronym. The Dedi lists (by family

and by acronym) will also be available in pdf at: http://www.conps {/Plant%20Keys/ACROS. pdf.

The following four posed for families of ferns: C o (CIB), 1 t (LXM),

Saccolomataceae (SCC), ae (CD.

The following 24 y proposed for angiosy B lowi (BRW), Durionaceae (DUR),

Calophyllaceae (CLP), C ] (CTR),E lepid (ERE), Eriospermaceae (ERS), Guamatelaceae

(GUA), Haptanthaceae (HAP), Hostaceae (HOS), r (JHN), Leptaul (LEP), Limeaceae (LME),

Linderniaceae (LDR), Lophigcarpáceae VOD. as anand (NES), Pentapetaceae (PEN), Pteleocarpaceae

(PTE), S l (SMO), S R), Stixaceae (STX), Syphostegiaceae (SYP), Talinaceae (TAL),

Thomandersiaceae (THO), e (VIB). Gee Haston et al. (2007) for a recommended linear sequence

for filing families based on updated versions of the APG II (2003) classification.

APPENDIX

The full alphabetical listings by family.

Abolbodaceae ABB Adoxaceae ADX Alangiaceae ALG Amaranthaceae AMA

Acanthaceae ACA Aesculaceae AES Alismataceae ALI — Amaryllidaceae AML

ACE Aextoxicaceae AEX Alliaceae ALL Amborellaceae ABL

Achariaceae ACH Agapanthaceae AGP ALO mbrosi AMB

Achatocarpaceae AHT Agavaceae AGA Alseuosmiaceae ASM Amygdalaceae AMY

oraceae ACO Agdestidaceae AGD Alsinaceae ASN Anacardiaceae ANA

Actinidiaceae ACT Aitoniaceae AIT Alstroemeriaceae ALS Anarthriaceae ANR

Actiniopteridaceae ACP Aizoaceae AIZ Altingiaceae ALT Ancistrocladaceae ANC

Adiantaceae ADI Akaniaceae AKA Alzateaceae ALZ Androstachyaceae AND

J. Bot. Res. Inst. Texas 3(1): 273 — 277. 2009

274

Anemarrhenaceae

miaceae

Angiopteridaceae

Anisophylleaceae

Aphloia

ees

Apiaceae

Aquifoliaceae

Araceae

Araliaceae

Aralidiaceae

raucariaceae

elaceae

Atherospermataceae

Athyriaceae

ucubaceae

Austrobaileyaceae

Averrhoaceae

Avicenniaceae

zollaceae

Balanitaceae

Balanopaceae

Balanophoraceae

Balsaminaceae

Barbeuiaceae

Barbeyaceae

Barclayaceae

Barringtoniaceae

Berberidac

c sue

etulaceae

Biebersteiniaceae

Bignoniaceae

Bischofiaceae

Bixaceae

Blandfordiaceae

Boryaceae

Botrychiaceae

Brassicaceae

Bruniaceae

Brunoniaceae

Buddlejaceae

Burmanniaceae

Burseraceae

Butomaceae

Caesalpiniaceae

Calceolariaceae

Callitrichaceae

£v

Campynemataceae

Canellaceae

Cannabaceae

Cannaceae

Canotiaceae

Carica

Caremanrices

Caryophyllaceae

Cassythaceae

Casuarinaceae

Cecropiaceae

Celastraceae

Celtidaceae

Centrolepidaceae

Clethraceae

Clusiaceae

Cneoraceae

Cobaeaceae

Cochlospermaceae

Colchicaceae

Columelliaceae

Convolvulaceae

ceae

rylaceae

Corynocarpaceae

ostaceae

Crassulaceae

Cyclocheilaceae

Cymodoceaceae

Cynomoriaceae

Dasypogon

Datiscaceae

Davalliaceae

Davidiaceae

Davidsoniaceae

Dicrastylidaceae

Didiereaceae

Diegodendraceae

Diervillaceae

nía

Di Hen M

Dioscoreaceae

Dipentodontaceae

Dryopteridaceae

uckeodendraceae

Dulongiaceae

Durionaceae

Ebenaceae

Ecdeiocoleaceae

Ehretiaceae

etaceae

Eremolepidaceae

Eremosynaceae

Snow, Three-letter family acronyms

Ericaceae

opallacsae

Fouquieriaceae

Francoaceae

Frankeniaceae

Fumariaceae

Garryaceae

Geissolomataceae

Glaucidiaceae

Gleicheniaceae

Globulariaceae

Gnetaceae

ramineae

Grammitidaceae

Greyiaceae

Griseliniaceae

Grossulariaceae

Grubbiaceae

Guamatelaceae

Gunneraceae

Guttiferae

Gymno A

Gyrostemonaceae

Halesiaceae

dE NE

Haloragac

Hanguanaceae

Heliotropiaceae

Helleboraceae

Heiwingiac

He! merocalidaceze

Herreriaceae

Hesperocallidaceae

Heteropyxidaceae

imantandraceae

Hippocastanaceae

Hippocrateaceae

Hippuridaceae

Hoplestigmataceae

Hostaceae

Hypecoaceae

Hypseocharitaceae

Icacinaceae

llicaceae

Illecebraceae

Illiciaceae

Ixerbaceae

Ixioliriaceae

Ixonanthaceae

Japonoliriaceae

Johnsoniaceae

Joinvilleaceae

Juglandaceae

Julianiaceae

Juncaceae

Juncaginaceae

Kaliphoraceae

Kaulfussiaceae

Kingdoniaceae

Kirkiaceae

Koeberliniaceae

Krameriaceae

Labiatae

Lacistemataceae

Lactoridaceae

miaceae

Lanariaceae

Lardizabalaceae

Lauraceae

Laxmanniaceae

idaceae

Ledocarpaceae

Leeaceae

eguminosae

Leitneriaceae

Lemnaceae

aceae

Lentibulariaceae

mud

Lepuropetalaceae

Limnanthaceae

Limnocharitaceae

Limoniaceae

inaceae

Linderniaceae

Lindsaeaceae

Linnaeaceae

Lissocarpaceae

saceae

Lophiocarpaceae

Lophosoriaceae

Loranthaceae

Lowiaceae

Loxogrammaceae

Loxomataceae

Loxosomataceae

Luzuriagaceae

Mackinlayaceae

Maesaceae

Malac

Malesherbiaceae

Marcgraviaceae

Marsileaceae

Melanophyllaceae

Melanthiaceae

Melastomataceae

Meliaceae

Melianthaceae

Meliosmaceae

Memecylaceae

Mendonciaceae

Menispermaceae

Menthaceae

Menyanthaceae

Mesembryanthemaceae

Mespilaceae

Metaxyaceae

Metteniusaceae

Mimosaceae

Misodendraceae

Mitrastemonaceae

Molluginaceae

nachosoraceae

Muntingiaceae

Musaceae

Myodocarpaceae

Myoporaceae

Myricaceae

MAE

Myristicaceae

D em

Myrsinaceae

Myzodendraceae

jadaceae

Nartheciaceae

276

Naucleaceae

Negripteridaceae

Nelumbonaceae

Nepenthaceae

Nephrolepidaceae

Nesogenaceae

Neumanniaceae

Neuradaceae

Nitrariaceae

Octoknemaceae

Olacaceae

Oleaceae

Oleandraceae

Oliniaceae

Oncothecaceae

Onocleaceae

Ophioglossaceae

Peganaceae

Pellicieraceae

Penaeaceae

Pennantiaceae

Pentadiplandraceae

Pentapetaceae

Pentaphragmataceae

Pentaphylacaceae

Penthoraceae

Peperomiaceae

eraceae

Peridiscaceae

Periplocaceae

Petermanniaceae

Petiveriaceae

Petrosaviaceae

Phellina

Phila eee

Philesiaceae

Phyllanthaceae

Phyllocladaceae

Phylionomaceae

Physenaceae

Phytolaccaceae

Picramniaceae

Picrodendraceae

Pinaceae

Piperaceae

Pistaciaceae

Podophyllaceae

Poran I acca

Pteridac

RS. eese

Pterostemonaceae

Punicaceae

Putranjivaceae

Pyrolaceae

Quiinaceae

Quillajaceae

Quintiniaceae

Rafflesiaceae

Ranunculaceae

Rapateaceae

Restionaceae

Retzia

Rha Wo

Rhopalocarpaceae

Rhynchocalycaceae

Roridulaceae

Rosaceae

Rousseaceae

ubiaceae

Ruppiaceae

Ruscaceae

Sabiaceae

Saccolomataceae

Sarcolaenaceae

Sarcospermataceae

xaceae

Scheuchzeriaceae

Schisandraceae

Schizaeaceae

dioi d

Schoepfiac

Sciadopity:

Scyphostegiaceae

Scytopetalaceae

Selaginellaceae

Setchellanthaceae

imaroubaceae

Simmondsiaceae

Sinopteridaceae

Siparunaceae

Siphonodontaceae

Sladeniaceae

Smilacaceae

Solanaceae

Sonneratiaceae

Sphenostemonaceae

Spigeliaceae

Spiraeaceae

Stachyuraceae

plat Rhe uslaeede

Stegnospermataceae

Stemonaceae

Stemonuraceae

Stenomeridaceae

Sterculiaceae

Stilaginaceae

Mies

seduce

Strelitziaceae

Str es

Stromatopteridaceae

chnaceae

Sym BE

Symplocaceae

Taccaceae

Taenitidaceae

Talinaceae

Tamaricaceae

Tapisciaceae

Taxaceae

Taxodiaceae

Tecophilaeaceae

Tetracarpaeaceae

Tetracentraceae

achondraceae

Tetradiclidaceae

Tetragoniaceae

Tetramelaceae

Tetrameristaceae

Thalictraceae

Theaceae

Theligo

ied

Themidaceae

Snow, Three-letter family acronyms

Theophrastaceae TEO Tremandraceae TMD Umbelliferae UMB Wellstediaceae WLS

Thismiaceae THS Tribelaceae TRB Urticaceae URT Welwitschiaceae WLW

Thomandersiaceae THO Trichomanaceae TCM Uvulariaceae UVU Winteraceae WIN

Thunbergi THN Trichopodaceae TCH Vacciniaceae VAC | Woodsiaceae WDS

Thurniaceae THU Trigoniaceae TRG Vahliacea VHL Xanthophyllaceae XPH

elaeaceae THY Trilllaceae TRL Valerianaceae VAL anthorrhoeaceae XAN

Thyrsopteridaceae THR meniaceae TRM Velloziaceae VIL Xeronemataceae XER

Ticodendraceae TIC Triplostegiaceae TPS rbenaceae VRB Xyridaceae XYR

iliaceae TIL Tristichaceae TRS ce VIO Zamiaceae ZAM

Tmesipteridaceae TMS Triuridaceae TRI Virburnaceae VIB — Zannichelliaceae ZAN

Tofieldiaceae TOF Trochodendraceae TRC Viscaceae VIS A e ZIN

Torricelliaceae TOR Tropaeolaceae TRP Vitacea VIT Zoste ZOS

ovariaceae TOV — Turnerac TNR Vittariaceae VTT ada ZYG

Trapaceae TRA Typhaceae TYP Vivianiaceae VIV

Trapellaceae TPL Ulmaceae ULM Vochysiaceae VOC

ACKNOWLEDGMENTS

My thanks to K. Richards (Landcare Research, New Zealand), E. Haston (E) and W. Judd (FLAS) for their

comments, and to Al Schneider and the Colorado Native Plant Society for agreeing to post the complete

listings online.

REFERENCES

APG ll. 2003. An update of the Angiosperm Phylogeny Group classification for the | families of flower-

ing plants: APG ll. Bot. J. Linnaean Soc. 141:399-436

BRASHER, J.W. AND N. Snow. 2004a. Further updates to Weber's three-letter family acronym system. Taxon (online

version, http//www.botanik.univie.ac.at/iapt/taxon/index.htm). [online version no longer available]

BRASHER, J.W. AND N. a i Pi She enla ens Mud system of three-letter plant family acronyms.

http:// lu/nhs/! /'acronyms.pd

HASTON, E., J.E. RICHARDSON, PF. STEVENS, M W Cuase, AND D.J. Harris. 2007. A linear sequence of Angiosperm Phylogeny

Group Il families. Taxon 56:7-12.

Heywoon, V.H., R.K. BrummrT, A. CULHAM, AND O. St&ERG. 2007. Flowering plant families of the World. Royal Botanic

Gardens, Kew.

MasgerLeY, D.J. 2008. Mabberley's plant-book. Cambridge University Press, Cambridge.

Smith, A.R., K.M. Prayer, E. SCHUETTPETLZ, P. KORALL, H. SCHNEIDER, AND PG. WoLr. 2006a. A classification for extant ferns.

Taxon 55:705-731.

Smith, A.R., H.-P. Keeler, C.H. HAUFLER, T.A. RANKER, AND H. SCHNEIDER, 2006b. Serpocaulon (Polypodiaceae), a new genus

segregated from Polypodium. Taxon 55:919-930.

Snow, N. 2009, Checklist of vascular plants of the Southern Rocky ! Region. Version 3. http://www.conps.

org/ gui diim Ese 2009 Version3, Final9620(2).pdf

Snow, N. AND N. Hotton. 2000. Additions to Weber's three-letter family acronyms based on results of The Angio-

sperm DS Group. Taxon 49:77-78.

Stevens, PF. 2001 (and onwards). Angiosperm Phylogeny Website. Version 9, June 2008. http://

MOBOT/research/APweb/welcome.html (accessed 10 Oct 2008).

WEBER, W.A. 1982. Mnemonic three-letter acronyms for the famili

herbarium curation. Taxon 31:74-88.

Weeer, W.A. AND R.C. Wirrmann,. 1992. Catalog of the Colorado flora: a biodiversity baseline. University Press of

Colorado, Niwot.

ora/

f vascular pl device for more effective

278 [| Ingel

BOOK REVIEW

JANET CHUI AND Jason LUNDBERG. 2008. A Field Guide to Surreal Botany. (ISBN 978-9810810177, pbk.). Two

Cranes Press, Singapore Post Centre, PO. Box 034, Singapore 914002. (Orders: www.twocranespress.

com). $12.00, 70 pp., 5 1/2" x 8".

A Fi 14 sT 1 El A +} 1 fi

Another work in the same

genre is Parallel Botany, by Leo Lionni. This small peak isa bit of lunacy sure to apes to slightly twisted plant lovers and botanists

everywhere.

As they say, presentation is everything. This appears to be an old-fashioned t ical i ith its deli ill Latin

names, notes on und and life cycle, e uA d kat Dee Great care has been put into bes in entries read as real and

authentic, but tl i d literary they are in on some way-cool j with the . It is an example

of what can be döne when did Bur care di attention on a literary project. It literally oozes olle This is a small press title that

3:3 -] 1

exceeded my ext y level

d hically by regions of the world and fully illustrate d by Janet Chui. It is hard to cl favorite f

the any species Amongst E favorites are the Forget-me- pis which m causes itching, did and xd in Mt on he

1 1 1 1

g which they grow; the Time C y

-l JA! f. 1; 1 2 y 1 1 y 1 de f N J+} I7. 1 A

ULAUL y P

ing Aspen, which is the Sue

known tree with a mati RENE

ry.

anet kenn and Jason undue met at the 2002 Clarion Writers Workshop in Michigan. They formed Two Cranes Press, an in-

1 isl in 2003

Raleigh, North Carolina, to Singapore where it is now based. Two Cranes was set up to

publish CRIME fiction and has published three other works.— Gary L. Jennings, Librarian, Botanical Research Institute of Texas, Fort

Worth, Texas 76102-4025, U.S.A.

J. Bot. Res. Inst. Texas 3(1): 278. 2009

CHROMOSOME NUMBERS OF MISCELLANEOUS

ANGIOSPERM TAXA

Marc Baker, Jon Rebman’, Bruce Parfitt', Donald Pinkava,

Charlotte Christy!, Andrew Salywon!, Raul Puente-Martinez'

School of Life Sciences, Arizona State University

PO. Box 87501, Tempe, Arizona 85287-4501, U.S.A.

Allan Zimmerman J. Hugo Cota

N Como Drive SONG sb oo il of Biology

Tortilita, Arizona 85142, U.S.A. atoon, SK s7n 5e2, CANAD,

ABSTRACT

Cl p 1 for 68 individuals in 64 including one i pecific hybrid) in 36 genera of 15 families. Of these,

11 taxa are counted for the first time.

RESUMEN

Se recontaron ] i icos de 68 individ 64 taxones (incluyendo un híbrido interspecífico) pertenecientes a 36

géneros de 15 familias. De estos, 11 taxones se han do por primera vez aquí

METHODS

Meiotic counts (n) were obtained from microsporogenesis in flower buds according to methods of Pinkava

et al. (1998); mitotic counts (2n) were obtained from root tips according to Cota et al. (1994)

RESULTS

First counts are reported for: Agave felgeri, A. margaritae, A. mchelveyana (Agavaceae); Cylindropuntia ante-

ojoensis x C. leptocaulis, Escontria chiotilla, Stenocactus multicostatus, Stenocereus beneckei (Cactaceae); Mosiera

ehrenbergii (Myrtaceae), Eriogonum ripleyi (Polygonaceae), Nemeranthus validulus (Portulacaceae) and Ivesia

multifoliolata (Rosaceae). A new count was made for Prunus virginiana var. demissa (Rosaceae). Other counts

are listed below (Table 1).

Tape 1. Vouchered cl t f miscell ; f a A eae eu

es 2

i Mal

his paper. V | , PK etal 1990) appear in parentheses. Asterisks (* ) denote first counts; (**)

denote a new chromosome number for a taxon.

Taxon Chromosome Locality, notes

number

DICOTYLEDONAE

Apocynaceae

Amsonia tomentosa Torr. & Frémont n=11 USA, Arizona, Mohave Co.: NNW of Alamo Lake, jctn.

var. tomentosa Signal and Alamo Rds., Parfitt 4181B & Christy (ASU).

Asteraceae / Compositae

Coreocarpus parthenioides Benth. n=12 MEX, Baja California: 5.3 km S of town of Bahia de Los

var. parthenioides Angeles, Rebman 957 & Marsh (ASU)

1¢ id J.P. Ret S g P. 0. Box 1390, San Diego, CA 92112; B.D. Parfitt, Biology Dept., University of Michigan, Flint, MI 48502-

2186: C. Christy, Biol Dept., Augusta College 00 Walton G. gusta, GA 30904-2200, Andrew Saly R. Puente-M., D B ical Garden, 1201 N. Galvin

Parkway, Phoenix, AZ 85008.

J. Bot. Res. Inst, Texas 3(1): 279 — 283. 2009

280

Taste 1. (continued)

Taxon Chromosome Locality, notes

number

DICOTYLEDONAE

Lasthenia coronaria (Nutt) Ornduff =4 MEX, Baja California: 4.8 km S of Punta Baja

W of El Rosario, Rebman 927 & Marsh (ASU).

Perityle crassifolia Brandegee var. n=19 MEX, Baja California Sur: 3 km S of San

robusta (Rydb.) Everly Gregorio, Baker 8767 & Johnson (ASU).

Perityle lobata (Rydb.) I.M. Johnst. n=17 MEX, Baja California Sur: 0.8 km E of San José,

Comondú, Baker 8748 & Johnson (ASU).

Senecio californicus DC. n=20 MEX, Baja California: 4.8 km S of Punta Baja

W of El Rosario, Rebman 930 & Marsh (ASU).

Stephanomeria schottii A. Gray n=8 USA, Arizona, Yuma Co.: Mohawk Dunes, T9S R15W

S2, Salywon 557 € Shohet (ARIZ, ASU, DES, SD).

Viguiera laciniata A. Gray n=18 MEX, Baja California: N of El Junco, between

Ensenada and Tijuana, Rebman 2156 & Hirales (ASU).

Brassicaceae / Cruciferae

Cakile maritima Scopoli n=9 MEX, Baja California: Punta Morro, N of Ensenada,

Rebman 2153 (ASU).

Hesperidanthus linearifolius (A. Gray) Rydb. n=11 USA, Arizona, Coconino Co.: 32 km N of Flagstaff on

Hwy 180, Salywon 256 (ASU)

Cactaceae

Cleistocactus cf. baumannii Lem. n=11 Origin unknown; cultiv. at Univ. of Tennessee

and Arizona State Univ., Christy s.n. (ASU).

*Consolea corallicola Small 2n — 66 USA, Florida, Monroe Co.: Torchwood Hammock,

Little Torch Key, Bob Ehrig, TNC. Cultiv. Fairchild

Tropical Gardens (FTG 90-04), Miami, us Arizona

State Univ, Tempe; recollected 14 June

by Pinkava (ASU). [JHC] Previously de

(Austin et al. 1998) without details.

Consolea rubescens (Salm-Dyck ex DC) Lem. 2n- 132 West Indies, Guadeloupe: Pointe de la Grand Vigie,

*Cylindropuntia anteojoensis (Pinkava) E.F.

Anderson x C. leptocaulis (DC.)

EM. Kunt

Cylindropuntia imbricata (Haw.) F.M. Knuth

var. imbricata

Echinocereus pensilis (K. Brandegee) Purpus

[Morangaya pensilis (K. Brandegee)

C.D. Rowley]

Echinocereus scheeri (Salm-Dyck) Scheer

subsp. gentryi (Clover) N.P. Taylor

Echinocereus viridiflorus Engelm.

var. viridiflorus

Na intertextus (Engelm.) Britton &

se var. intertextus

E o (Engelm.) E.M. Baxter

Eriosyce subgibbosa (Haw. Katt. subsp.

clavata (Sóhrens ex Schum.) Katt. var.

nigrihorrida (Backeb.) Katt.

Grand Terre (R. ibn UN in GeminiGardens,

Manalapan, Florida, and at

EINER Tempe Prien (ASU). JHC]

n=22 MEX, C S side of Sierra de la Fragua, near

Cuatro Geneace Salywon e & Tiller (ASU).

n=11 MEX, S.L.P: Municipio Santa Catarina, Puente 1258 (ASU).

n=11 Origin unknown; cultiv. in Scottsdale, Arizona,

Zimmerman 2665 (ASU). [BP]

n=11 Origin unknown; cultiv. in Tucson, Arizona,

Zimmerman 2774 ).

n=11 USA, Colorado, Lincoln Co.: Punkin Center,

Highway 71, Salywon 110a (ASU).

n=11 USA, Arizona, Pima Co.: Cienega Creek,

Salywon 115 & Tiller.

n=11 USA, Arizona, Mohave Co.: vicinity of Lake Mead,

purple flowers, Chamberland 58 (ASU). [?BP]

n=11 Chile, Cogimbo, Ritter 218, [Dr. J. Barrow's collection

No. 36] Atlanta Bot. Gard., then cultiv. in Desert Bot.

Gard., Arizona (DBG 1992-0203-01-01 ), recollected

by EF. Anderson (DES). [BP]

Í., Cl L £

Baker eta

Taste 1. (continued)

281

Taxon

Chromosome

number

Locality, notes

*Escontria chiotilla (F.A.C. Weber) Rose

Hatiora salicornioides (Haw.) Britton & Rose

Opuntia aciculata Griffiths

Opuntia elatior P. Miller

Opuntia humifusa (Raf) Raf.

Opuntia humifusa (Raf.) Raf.

Opuntia megarhiza Rose [O. pachyrrhiza

3mez-Hin. & Bárcenas]

Opuntia salmiana Parm. ex Pfeiff.

Opuntia stricta (Haw.) Haw.

Stenocactus crispatus (DC) A. Berger

AWM. Hill [Echinofossulocactus

crispatus DC.) Lawrence

Stenocactus multicostatus (Hildeman ex K.

Schum.) A. Berger [Echinofossulocactus

multicostatus (K. Schum.) Britton & Rose]

*Stenocereus beneckei (Ehrenb.) A. Berger

& F. Buxbaum

Stenocereus pruinosus (Otto) F. Buxbaum

Stenocereus stellatus (Pfeiff) Riccobono

Crassulaceae

Dudleya arizonica Rose

Dudleya ingens Rose

Fabaceae / Leguminosae

Astragalus nutriosensis Sanderson

DICOTYLEDONAE

n=11

n=22

n=44

2n=22

Origin unknown; cultiv. at Desert Bot. Gard.,

Arizona (Bed D, DBG #1950-2731-01-01),

Chamberland s.n. (ASU). [DP, JR]

Origin unknown; aa in ASU greenhouse,

Tempe, Rebman s.n. (ASU, SD).

Origin freee in Desert Bot: cy Arizona

(Bed 57W, DBG 76-73), Hodgson 36

(ASU photo, DES). [DP]

Origin from International Succulent Institute,

possibly in Colombia; cultiv. at Desert Bot. Gard.,

Arizona (DBG 168-9268-01-01) recollected by

J. Ward s.n. (DES, 2 sheets). JR; meiosis irregular]

USA, Florida: pineland remnant (from Montgomery

Foundation). Cultiv. in Fairchild Tropical Garden,

Florida (FTG 90-107) and in ASU Greenhouse,

Arizona, Pinkava s.n. (originally labeled as

var. austrina (Small) Dress JHC].

USA, Georgia, Bullock Co.: 3.2 km N of Lotts Creek

Bridge, 9.7 km S of Statesboro, Christy 2741b (ASU, DES,

SD). Richmond Co.: Augusta, 3.2 km SW of jctn. l-20 & l-

in Christy 2748-A1 (ASU), -A12 (ASU, NY, US). South

Carolina, Lexington Co.: 48 km W of Columbia,

Christy 2745B (ASU, NY, US).

MEX, S.L.P: Mpio. Zaragoza, 1 km N of Valley de

S. Fantasmas, Puente 601 (ASU, two sheets)

Origin unknown; cultiv. in Scottsdale, Arizona,

Zimmerman s.n.

Origin unknown; cultiv. (as var. dillenii) in Scottsdale,

Arizona, Zimmerman 2667 (A

MEX, S.L.P,13 m : id Huizache Junction, Rod McGill

then cultiv., Desert Bot. Gard., Arizona (DBG

1962- 7221-01 P pubis by J. Ward (DES). [BP]

Origin unknown. Cultiv. in Mesa, Arizona, Puente

538 (ASU, two sheets, one box). [RP & JR]

[me

Origin unknown; cultiv. in Desert Bot. Gard.

Arizona (Webster Patio near "Ne Parfitt s.n. (ASU).

MEX, Colima: Manzanillo, C. Meig €: F. G. Hermann, in

1955. Cultiv. in Desert Bot. Gard, s (DBG 5143-

01-01), Chamberland s.n. (ASU, pe

MEX, PUEBLA: near Tehuacan, 4 ultiv

Desert Bot. Gard., Arizona (DBG 139- 0628- 0104):

recollected by J. Ward. (ASU, DES)

MEX, Baja California: 3.2 km SE of San Matias Pass,

San Pedro Mártir, Baker 3716 (ASU)

MEX, Baja California: 3.2 km Wercamalu on Rte. 1,

Gallagher 82-40 (ASU). [MB]

USA, Arizona, Apache Co: SE of Eager, Picnic Hill

just W of Rte. 666, Baker 11431.2 & Wright (ASU).

282

Taste 1. (continued)

Taxon Chromosome Locality, notes

number

DICOTYLEDONAE

Astragalus oophorus S. Watson var.

aulescens (M.E. Jones) M.E. Jones

Astragalus trichopodus (Nutt.) A. Gray var.

lonchus (M.E. Jones) Barneby

Astragalus troglodytus S. Watson

Lupinus latifolius Agardh subsp.

leucanthus (Rydb.) Kenny & Dunn

Psoralidium tenuiflora (Pursh) Rydb.

Lamiaceae / Labiatae

Salvia dorrii (Kellogg) Abrams subsp.

mearnsii (Britton) McClintock

Loasaceae

Eucnide rupestris (Baillon) Thompson & Ernst

Mentzelia multiflora (Nutt.) A. Gray

Myrtaceae

*Mosiera ehrenbergii (Berg) Landrum

Polygonaceae

*Eriogonum ripleyi J.T. Howell

Portulacace

ons validulus (Greene) Kiger

[Talinum validulum Greene

Rosaceae

*lvesia multifoliolata (Torr) Keck

**Prunus virginiana L. var. demissa

(Nutt) Torr.

Scrophulariacea

Penstemon je A. Nelson

Penstemon albomarginatus M.E. Jones

Agavaceae

Agave xarizonica Gentry & J. H. Weber,

o sp. (A. chrysantha Peebles x

A. toumeyana var. bella)

USA, Arizona, Coconino Co.: 5 km N of Jacob Lake,

Baker Mee & Trushell (ASU

n=15 MEX, Baja California; 2 krn E of Rosalallita, m

Domingo Wash, Baker 8701 & Johnson, Hwy 1, 86 km

S of San Quintin, Rebman 1646 & Delgadillo Tn

—

n=11 USA, Arizona, Yavapai Co.: NW of Prescott, 0.8 km

SW of Lone Pine Tank, Baker 10652 (ASU).

n=24 USA, Arizona, Yavapai Co.: NW of Prescott, 1.4 km

WNW of Stinson Mtn., Baker 11023.1 €: Wright (ASU).

n=11 USA, Arizona, Yavapai Co.: NW of Prescott, 3.7 km

SW of Connell Mtn. summit, Baker 10843 (YCH).

n=15 USA, Arizona, Yavapai Co.: 4 km WSW of leia of

Hell Canyon and Verde River, 1.5 km NW rde

Ranch Headquarters, Baker 15697 (ARIZ, jc ASU, DES).

MEX, Baja California, S of La Ventana between Mexicali

d San Felipe, Rebman 2091 & Hirales (ASU).

n=10 USA, Arizona. Yavapai Co: S of Ash Fork, 6 km SSE of

Rock Butte, Baker 11394, 1 (ASU, two sheets).

n=ca. 21

2n=22 MEX, S.L.P: Guadalcazar; cultiv. in ASU Greenhouse,

Tempe, Landrum 7375 (ASU). UHC]

n=16 USA, Arizona, Yavapai Co.: NW of Jerome, 4.2 km SSE of

Red Butte, Baker 11803 & Wright (ASU).

n=24 USA, Arizona, Yavapai Co.: ca. 9 km W of Jerome, W

base of Woodchute Mtn., Baker 9388 (ASU).

n=14 USA, Arizona, Coconino Co.: T19N R3E S15,

Tule Tank Wash, Baker 9847.

n=16 USA, Arizona, Yavapai Co.: Woodchute Wilderness

Area, 5 km NW of Jerome, Baker 9823 (ASU, two

sheets, second collection from same individual).

Previously counted as 2n = 16 (Wilken 1993).

n=8 USA, Arizona, Graham Co.: 2 km NNW of Solomon

Pass W of Rock Tank, Baker 11626 & Kertell (ASU).

n=8 USA, California, San Bernardino Co.: N34.8145E

W116.4065E, 55 km W of Barstow, Baker 16674.1 &

Silverman (ASU).

MONOCOTYLEDONAE

n=30 USA, Arizona: cultiv. in Desert Botanical Garden,

Arizona (Bed #40), Hodgson 4526 (ASU, DES). [MB]

Baker et al., Cl

Tase 1. (continued

Taxon Chromosome Locality, notes

MONOCOTYLEDONAE

USA, Arizona, Yavapai Co.: 5 km N of Camp Verde, just

W of Montezuma Castle Natl. Mon. boundary, Baker

11854 et al. (with morphological intermediacy to A.

delamateri Hodgson & Slauson (ASU, DES).

EX, Sonora: San Carlos Bay, Felger 637 [topotype]

(ASU, DES). [AS]

Agave glomeruliflora (Engelm.) A. Berger n=60 MEX, Coahuila, Los Cojos Minas, SW of limestone

slopes of Sierra del Carmen, Gentry & Engard s.n,

since cultiv. in Desert Bot. Gard., Arizona (DBG 1972-

0159-01-07), Hodgson 10196 (DES). [MB]

Agave aff. chrysantha Peebles

Agave felgeri Gentry n- 30

Agave margaritae Brandegee nz30 MEX, Baja Calif. Sur: Santa Margarita Island, Glass

s.n. et al. [topotype] (DES). [AS

*Agave mckelveyana Gentry n=30 USA, Arizona, Yavapai Co.: 15 km SW of Bagdad,

Baker 10813 (ASU, DES); Rugar Ranch, SW of Kirkland

Jct, W of Hwy. 89, N34E 21.765'W112E 44.816,

C.D. Avis s.n. et al. (DES). [AS €: Turcott]

Agave toumeyana Trel. var. bella Breitung n z 30. USA, Arizona, Yavapai Co: ca. 5 km N of Camp

Verde, Wright 1538 et al. ASU) [MB]

Agave utahensis Engelm. var. kaibabensis n=30 USA, Coconino Co.: Grand Canyon Natl. Pk., E of

(McKelvey) Gentry Hance Creek, Hodgson 9725 (DES). [MB

n=30 USA, New Mexico, Colfax Co.: N36.7819 W104.8619,

Yucca glauca Nutt.

Baker 12530 & Wright (ASU, DES).

Nolinaceae

Nolina bigelovii (Torr.) S. Watson n=19 USA, Arizona, Mohave Co.: Devil's Canyon, 15 km S of

Wickieup, Baker 10764 et al.. Yavapai Co.: 15 km SW of

Bagdad, Baker 10814 (ASU).

ACKNOWLEDGMENTS

Michael Chamberland, Tucson Botanical Garden and Wendy Hodgson, Desert Botanical Garden, Phoenix,

provided collections and bud material for some of the chromosome determinations. Gerald D. Carr and

L.M. Hill offered helpful comments on an earlier draft of the manuscript.

REFERENCES

Cora, J.H. AND C.T. Puitsrick. 1994. Chromosome number variation and polyploidy in the genus Echinocereus

(Cactaceae). Amer. J. Bot. 81:1054-1062.

Wiken, D.H. 1993. Prunus. In: J.C. Hickman, ed. The Jepson manual: higher plants of California. Univ. of California

Press, Berkeley. P. 970.

HoLmaren, PK., N.H. HOLMGREN, AND L.C. Barnett (eds.). 1990. Index Herbariorum. Part I. The herbaria of the World.

IAPT, New York Botanical Garden, Bronx.

Pinkava, DJ., J.P. Resman, AND M.A. Baker. 1998. Chromosome numbers in some cacti of western North America —

VII. Haseltonia 6:32-43.

284 Journal of tl n sD kiss f Texas 3(1)

BOOK NOTICE

GABRIELA G. HASSEL DE MENENDEZ AND MARTA E: Ruses. 2009. Catalogue of Marchantiophyta and Anthocero-

tophyta of southern South America (Nova Hedwigia, Beiheft 134). (ISBN 978-3-443-51056-5,

ISSN 1438-9134, pbk.). J. Cramer in der Gebrüder Borntraeger Verlagsbuchhandlung, Johannestr. 3A,

D-70176 Stuttgart, Germany. (Orders: www.schweizerbart.com, mail@schweizerbart.de, +49 (0)711

35 14 56 0, +49 (0)711 35 14 56 9 fax). $255.00, 672 pp., 3 b&w i 6 5/8" x 9 1/2".

C tri d Chile, Argentina and Uruguay, including E land (P Island), Malvi Island (Falkland Island), South

Georgia Island d th Shetland Island Scu Sandwich Island, and South Orkney Island.

Contents

Abstract & Resumen

Introductio

Ackno E

List of Marchantiophyta names and taxa

Annotations on Marchantiophyta names

List of Anthocerotophyta names and taxa

Annotations on Anthocerotophyta names

Gverematir lists of Marchantienl dA

Systematic and taxonomic changes

Geographic distribution of taxa

Patterns of distribution

References (43 pp.)

Index (42 pp. of scientific names)

From the Scd “The y t ] 1 he inf ion on the taxa known from Chile, Argentina, and Uruguay, and dis-

ld publications, mail he basis of modern literature and on studies by the senior author. The reasons

to present a joint catalogue for EM three eodem South American countries are that the literature for Chile and Argentina is to a

great extent the same, and Uruguay, although rather under collected [sic], has some phytogeographical relationship to Argentina, and

could therefore not be left out."

J. Bot. Res. Inst. Texas 3(1): 284. 2009

ANATOMÍA DE LA EPIDERMIS FOLIAR EN LAS ESPECIES MEXICANAS

DEL GÉNERO MUHLENBERGIA (POACEAE)

Octavio Roses Carrillo Yolanda Herrera Arrieta

Centro Bás Instituto Politécnico Nacional

id Autónoma P CS CHDIR Unidad Durango-COF,

ersidad s.n. Aguascalientes Sigma 119, Fracc. 20 de hell il

s. CP 20100, MÉXICO Durango, Dgo., MÉXI

orosales@correc.uaa.mx yherr eragipn. mx

RESUMEN

Se llevó a cabo tudio comparativo de la anatomía de la epidermis abaxial en láminas foli de 113 especies de dedi do =

las 120 estimadas bene en México. Se identificaron 21. racter iales y fueron sometidos a un análisis d

que resultaron tres grupos principales, imei se adi la Ancón de subgrupes Ses a pes propuestos puo otros autores con

uin en caracteres pees Con! I : ]

la a los epidérmicos, leli b 1 5mi d ió l

de lamina foliar, dado e no existe concordancia con P división del género en los subgéneros Muhlenbergia y Trichochloa, realizada

con base en anatomía de sección transversal de laminas en la mayoría de las especies del género Muhlenbergia. El perfil de caracteres

koi j -J A 1 i4] — E ; sad : ma ; 1 CN | J :

r li r ce r r

Se presentan microfotografías y se describen 1 t pidérmi n la superficie abaxial q teri l género y a grupos

con especies similares

PALABRAS Crave: Anatomía de epidermis, Muhlenbergia, Poaceae, México

ABSTRACT

1 E id 1 1 1 141 f. A ted t th iri 1A f, 7 1 1 1 hi 1

A I g I g I g 113 species of

Ahi 1 f. MA DI 1 31 1 L f. z

g y g PI g ps. These three

] ] ] ] based hol 1cl while some are not concordant

with che proposed subgenera: Muhlenbergia s Trichodla, ue on leaf BRE carecen of transversal sections. A profile of

s, but more characteristic for groups. Descriptions and photographs of the

main characters are given for the genus w groups of a species.

Key Wonps: Epidermal anatomy, Muhlenbergia, Poaceae, Mexico

INTRODUCTION

Las especies del género n poda ieee Du MM se localizan desde cerca del

nivel del mar hasta los ltmit en y idas del Nuevo Mundo desde

el sur de Estados Unidos de America hasta Apenina a especies se encuentran en Asia. La mayoría de

las especies (147 aprox.) se distribuyen en el sur de Norteamérica, por esta razón, el sur de Estados Unidos

y México, se consideran como su posible centro de origen (Peterson y Herrera 2001).

El género Muhlenbergia comprende aproximadamente 160 especies en el Mundo, en México cuenta

con cerca de 120 especies. Forma parte de los ecosistemas templados y secos con vegetación primaria de

pastizales, matorrales ads y bosques de encino y pino (Peterson y Herrera 2001). Es uno de los géneros

de gramíneas mayor ia económica para México por su alto valor forrajero, amplia distribución

y adaptabilidad. Las principales contribuciones al conocimiento sistemático parcial del género son los estu-

dios realizados por: Soderstom (1967), Morden y Hatch (1987), Peterson y Annable (1991), Herrera y Grant

(1994), Herrera (1998), Peterson (2000), Peterson y Herrera (001) y Herrera y Peterson (2006).

El género Muhlenbergia se caracteriza por presentar especies anuales o perennes; las especies perennes

son cespitosas, rizomatosas, o a veces decumbentes y enraizadas en los nudos inferiores (Herrera y Peterson

J. Bot. Res. Inst. Texas 3(1): 285 — 306. 2009

286 Journal of tl i i Texas 3(

2006). Hitchcock (1951), en sus estudios florísticos de las especies americanas, define que estos caracteres

morfológicos son de gran ayuda para dividir al género en grandes grupos y de esa manera hacerlo más mane-

jable, separando Muhlenbergia en cuatro grandes grupos morfológicos: 1) Las especies anuales, pequerias y

frágiles; 2) Las especies perennes rizomatosas; 3) Las especies perennes no rizomatosas, robustas, mayores

de 1.5 m de alto y 4) Las especies perennes no rizomatosas, no robustas, menores de 1 m de alto.

Peterson y Herrera (2001) en un estudio de anatomía foliar en sección transversal de las casi 160 espe-

cies Ponens en el Mundo e enero oda Enc que el género es divisible en cuatro grupos

gia y Trichochloa) y dos secciones (Epicampes

y PRI en el BUE do Trichochloa, y SHE el reconocimiento de estas entidades subgenéricas.

Estudios anatómicos de la epidermis abaxial de laminas han aportado información de calidad para la

clasificación de las gramíneas en diferentes niveles como son: Metcalfe 1960; Renvoize 1981, Stieber 1982;

Morden y Hatch 1987, Peterson y Eon 1989; a y Koch 1998; Vieria y col. 2002; Gómez y Téllez 2008.

Por esta razón diaron | del género Muhlenbergia para determinar si los caracteres

de la epidermis abaxial de an son consistentes en la delimitación de especies, así como para obtener

caracteres “diagnóstico” que permitieran inferir sus relaciones de similitud.

MATERIALES Y MÉTODOS

Las muestras biológicas de las especies de Muhlenbergia (ver nO mun principalmente de ejem-

plares depositados en los herbarios CIIDIR y UAA. O p de los herbarios: ENCB, GUADA,

IBUG, INEGI, MEXU, SLPM, 21 y US:

Para obtener f baxial, se utilizó la técnica de raspado directo, sobre la superficie

adaxial en la porción media de laminas foliares Bee desarrolladas, observando a través del microscopio

estereoscópico. Los fragmentos se colocaron en una solución acuosa de detergente comercial y se llevaron a

temperatura de ebullición durante 30 minutos para hidratar y ablandar los tejidos, en seguida se procedió a

raspar la superficie adaxial, con navaja bisturí, para eliminar tejidos y obtener la epidermis abaxial limpia.

Las muestras de epidermis se tiñeron con safranina y se eliminó el exceso de agua pasando los cortes por

una solución acuosa de alcohol a diferentes concentraciones (30%, 50%, 70% 95% y 100%). Las muestras

se montaron en resina.

Las ob j tómi hicieron con un mi pio de luz con objetivo de 10x y 40x y ocular

de 10x d to, | icrofotografí t con una cámara digital Canon Powershot S45 adaptada a

un microscopio Leica DMD. Las descripciones de la epidermis abaxial se hicieron siguiendo la terminología

propuesta por Ellis (1979).

El número de ejemplares analizados por especie fue variable (1 a 7, ver Apéndice), dependiendo de la

abundancia o rareza y disponibilidad de cada especie. Así, de las especies menos comunes solo fué posible

obtener una población para el presente análisis. En el caso de especies con distribución amplia se escogieron

ejemplares de localidades distantes. La información de epidermis generada para cada especie se sometió a

un análisis de similitudes. En el análisis cada especie (Apéndice), se consideró como una unidad taxonómica

operacional (UTO) y se registraron 21 caracteres anatómicos de epidermis (Cuadro 1). Se aplicó el Índice

de Euclidean (adecuado para el tipo de n con el fin de generar la matriz de similitudes a partir de

la cual se construyó el árbol de similitude mediante la utilización de UPGMA (unweighted

pair group mathematical average o du

RESULTADOS

Los resultados se interpretaron en dos formas: 1) la descripción general que se generó combinando los

caracteres de todas las especies de Muhlenbergia incluidas en este estudio y 2) resultados de un análisis de

similitudes.

Racalac 2 Herrera, A + tada | I *4 L I Iz fal An Monk L g 287

Cuapno 1. Lista d y estados d ter anat | análisis de congl d

Carácter Estados Codificación

1. Dif iación d Zonas costales e intercostales bien definidas 1

Zonas costales e intercostales no definidas 2

2. Pared anticlinal de las células largas Sinuosa 1

intercostales Lisa 2

Ondulada 3

3. Forma de las sinuosidades Forma de U 1

Forma de V 2

Forma de 3

4, Hileras de estomas Una hilera definida 1

Dos hileras definidas 2

Una hilera definida y algunos estomas dispersos 3

Mas de dos hileras 4

5. Células cortas intercostales Presentes 1

Ausentes 0

6. Micropelos cantidad por campo la 1

De6a10 2

Más de 10 3

7. Micropelos (longitud total) Hasta 9 1

De 10 a 20 2

Más de 20 3

8. Longitud célula basal micropelo Menos de 5 1

Más de 10 3

9. Longitud célula apical micropelo Hasta 12 1

De 13 a 40 2

10. Papilas Presentes 1

Ausentes 0

11. Distribución de las papilas Presentes en todas las células largas intercostales 1

Presentes en las células ineterestomáticas 2

12, Arreglo de las papilas Una papila por célula 1

Más de una papila por célula 2

13. Papilas cubriendo al estoma Una papila cubriendo en parte o en su m al estoma 1

Más de una papila rodeando al estom 2

14. Forma de las papilas Convexas e infladas 1

Pequeñas y cóncavas 2

Alargadas, distalmente engrosadas 3

15. Aguijones ausentes 0

presentes 1

16. Distribución de aguijones En zonas intercostal 1

En zonas ae 2

En ambas zo 3

17. Tamaño de la base del aguijón Base más corta aie la barba 1

con respecto a la barba Base más larga que la barba 2

18. Forma de la barba en relación Barba desarrollada desde la base hasta el ápice altamente elevado 1

a la base Barba no desarrollada desde la base hasta 2

el ápice, base cónica no elevada

19. Longitud de la barba en relación Longitud de la barba más corta que la longitud de la base 1

a di longitud de la base Longitud de la barba eno i. aa Arga Ba E rud de la base 2

20. los cuerpos Equidimensional, en for

» éflice constrenidos en la por ntral 1

Ae Lai en n forma a ies de montar, 2

redondeado al

21. Macropelos Presentes 1

288 t i i Texas 3(

DESCRIPCION GENERAL DE LA EPIDERMIS ABAXIAL

EN LAMINAS FOLIARES DE MUHLENBERGIA DE MEXICO

Zonas Costal e intercostal.—(Fig. 1). La epidermis en vista de superficie, esta dividida en zonas o bandas

conspicuas (Vieira y col. 2002). En las especies mexicanas del género Muhlenbergia, 103 especies presentan

las zonas bien definidas, mientras que en las 10 restantes no es posible diferenciar estas dos zonas.

hl las células]

Células largas intercostales.—(Fig. 2). En] p

presentan paredes anticlinales de tres tipos: Grupo I) 85 especies exhiben paredes sinuosas, 39 de ellas

en forma de U, otras 36 en forma de V y las 11 especies restantes en forma de. Grupo II) 12 especies con

paredes lisas y Grupo III) 16 especies con paredes onduladas (Ellis 1979).

Estomas.—En todas las especies examinadas se observaron aparatos estomáticos subdiscoidales. Las

células subsidiarias son redondeadas (en forma de domo), en una, dos o más hileras longitudinales a lo largo

de la zona intercostal.

Papilas.—(Fig. 3). La superficie externa de las células epidérmicas son aplanadas o convexas o presen-

tan abultamientos o salientes conocidos como papilas. Estas salientes de la pared externa pueden ser muy

diferentes, desde las simples cónicas a estructuras más largas, localizadas en el centro o en la parte distal de

las células intercostales largas, de acuerdo con Ellis (1979), las especies de Muhlenbergia presentan formas

convexas e infladas, pequeñas y hundidas, alargadas distalmente engrosadas.

Tricomas.—Las especies de Muhlenbergia, en su epidermis abaxial muestran dos tipos de tricomas:

Micropelos y Aguijones.

Micropelos.—(Fig. 4). Los micropelos consisten de dos células con paredes delgadas, una célula basal

y una distal. Las células del micropelo en conjunto pueden ser más largas o pueden ser más cortas que

el complejo estomático. En el presente estudio la mayoría de las especies presentan micropelos con una

longitud total de hasta 20. Se distinguieron 2 tipos característicos: i) célula distal inflada, de 1/3 ó 2/3 del

largo del micropelo, sin célula apical (Figs. 5a y 5b); ii) célula distal delgada (no inflada), 1/3 ó 2/3 del largo

del micropelo, con una célula apical corta (1/10 del largo del micropelo) o carente de ella. Pocas especies

poseen micropelos con una longitud total mayor a 20, su forma característica es ser micropelos delgados y

alargados con la célula apical más corta que la distal, (Figs. 5c y 5d).

Aguijones.—(Fig. 5). Los aguijones son fitolitos unicelulares, que se distinguen por su base dilatada y el

ápice afilado, se observan en la zona costal, en la zona intercostal, en ambas zonas o pueden estar ausentes.

De las especies estudiadas 56 tienen aguijones en la zona costal, 4 en la zona intercostal, 13 en ambas zonas

y 40 no presentan aguijones.

Células Cortas.—(Fig. 6). Las células cortas se dividen en células de sílice cuando el interior de la

célula esta ocupado por SiO, que se solidifica en formas variadas, y son llamados cuerpos de sílice. El otro

tipo de células cortas son denominadas células de corcho o süber, cuyas membranas están suberificadas y

a menudo tienen material orgánico sólido. La mayoría de las especies mexicanas del género Muhlenbergia

(96 especies) contienen cuerpos de sílice del tipo de silla de montar formando una doble equis, y las 17

restantes con cuerpos de sílice en forma de silla de montar constreñidos en la parte media con los extremos

redondeados o irregulares (Figs. 6a y 6b). Los cuerpos de sílice se localizan en las zonas costales. El otro

tipo de celdas son las células de corcho que se encuentran tanto en la zona costal como en la intercostal, sin

embargo en las especies estudiadas son raras en las zonas intercostales y comunes en la zona costal (Fig.

6d). Las células cortas se agrupan en hileras longitudinales ya sea en pares o solitarias, (Figs. 6c y 6d) entre

células largas consecutivas.

Macropelos.—(Fig. 7). Los macropelos generalmente se disponen en la zona intercostal de la epider-

mis. Solo en 18 de las especies estudiadas se observaron macropelos. Éstos son de diferentes tamarios desde

pequefios con un ápice punzante hasta largos y suaves al tacto como los que muestra M. pubescens.

La epidermis de las láminas foliares, en vista de la superficie abaxial, esta dividida en zonas o bandas

conspicuas. En general es posible distinguir dos zonas principales la zona costal y la zona intercostal. La

mayoría de las especies mexicanas de Muhlenbergia presentan esta organización, sin embargo, especies tales

1Z mas Lal: Jn Mull L H 289

como, M. alamosae, M. capillipes, M. Lois de i di M. firma, M. glabrata, M. gypsophila, M. laxiflora, M.

macroura y M. nigra no presentan di has zonas. En estos casos, la descripción de caracteres

epidérmicos se lleva a cabo localizando sus estomas y Pus s que son típicos de la zona intercostal.

ANÁLISIS FENÉTICO

En las especies analizadas (Cuadro 1) se determinaron 21 caracteres epidérmicos de calidad taxonómica

(diagnóstico). Dichos caracteres fueron utilizados para hacer una matriz (Cuadro 2) la cual se sometió a un

análisis de similitudes. El dendrograma resultante (Fig. 14) reveló la existencia de tres grupos principales

definidos por: la presencia/ ausencia de aguijones, la presencia/ ausencia de papilas, además de M. breviseta

y M. hintonii en una posición externa a ellos. El grupo I incluyó 47 especies que muestran aguijones y care-

cen de papilas, en el grupo II se agruparon 27 especies que muestran tanto aguijones como papilas, y en

el grupo III se concentran 37 especies sin papilas ni aguijones. Las 2 especies externas (M. breviseta y M.

hintonii) comparten con el grupo 3 la carencia de papilas y ee aguijones.

bie

Las especies RE po I (Fig. 14) incluy I n definidas] tales e int

tales, aguij lavana ly mes ilas; se redefinen en subgrupos: el subgrupo

IA se define io la cena de una sola iier de ase en la zona intercostal (Fig. 8), mientras que el

subgrupo IB lo confi ] t hil d por zona intercostal (Fig. 9).

El grupo II (Fig. 14) esta inicorado x especies que tienen papilas, aguijones, zonas costales e intercos-

tales bien definidas, estomas en una, dos o más hileras, mientras que son del grupo de plantas perennes, no

robustas. También se forman claramente dos subgrupos: el subgrupo IIA contiene 10 especies que disponen

de papilas y aguijones y la mayoría no presenta sinuosidades en las paredes anticlinales de las células largas

intercostales (Fig. 10). El subgrupo IIB contiene 16 especies que disponen de papilas y aguijones y además

muestran sinuosidades en forma de U o de V en las paredes anticlinales del las células largas intercostales

(Fig. 11)

El grupo III que se define por carecer de aguijones y de papilas, se divide en dos subgrupos: IIIA con la

mayoría de las especies anuales (Fig. 12), y IIIB con la mayoría de las especies perennes de vida corta (Fig.

13). Las especies que son externas a los grupos en el dendrograma, se asocian más con el tercer grupo por

compartir con este la carencia de aguijones y papilas.

DISCUSIÓN

En el grupo 1 (Fig. 14) se concentran especies perennes, la mayoría de ellas robustas, con algunas intrusiones

de especies no robustas, compartiendo los caracteres de epidermis abaxial anotados.

En el grupo II (Fig. 14) dominan especies perennes no robustas y rizomatosas por igual. Es interesante

observar que en el subgrupo IIB comprende ocho de las 13 especies del complejo Muhlenbergia montana

incluidas en este estudio (M. cualensis, M. eriophylla, M. jonesii, M. michisensis, M. virescens, M. quadridentata,

M. montana y M. watsoniana), reconocido por Herrera y Grant (1994), con base en caracteres morfológicos,

anatómicos foliares de sección transversal y del contenido de flavonoides. Como se observa en el dendro-

grama (Fig. 14), estas especies son también similares con base en anatomía de epidermis abaxial.

El grupo III esta constituido a su vez por dos subgrupos: el subgrupo IIIA formado por especies que

no presentan papilas ni aguijones (Fig. 14); especies la mayoría de hábitos anuales. Mientras que en el sub-

grupo IIIB con presencia de papilas y ausencia de aguijones, se encuentran en mezcladas especies perennes

rizomatosas, estoloníferas decumbentes y anuales en proporciones similares.

Las observaciones anteriores muestran la concordancia que presentan las especies de Muhlenbergia

en sus caracteres morfológicos vegetativos y epidérmicos en vista de superficie abaxial. Ásí, encontramos

que especies cercanas morfológicamente como M. texana y M. eludens (por caracteres de lema y cariopside),

anatómicamente también son similares (misma rama en ne iis m embargo, el caso contrario también

sucede, donde especies como M. rigens y i li muy disimiles, mientras

ue por su epidermis, presentan t il las coloca mas cerca entre sí que con otras especies.

290

Cuaoro 2, Matriz básica de datos.

E. Carácter

19 20 21

12 13 14 15 16 17 18

UTOs

M. aguascalientensis 1

M. annua

M. appressa

M. arenacea

M. arenicola

M. brandegei

M. brevifolia

M. breviligula

M. brevis

M. capillipes

M. ciliata

aie io e

M. crispiseta

M. cualensis

— — a Ko a.

M. densiflora

M. distichophylla

M. diversiglumis

M. dubia

mee — c 1.

M. durangensis

M. elongata

M. eludens

M. emersleyi

— Oe Oe -—

prar sey Wee See

M. eriophylla

M. firma

— — c— c— -—

== >= >. —

M. glauca

M. gooddingii

M. gypsophila

M. jaliscana

M. jonesii

291

Racalac 2 Harrara A

CUADRO 2. (continued)

E. Carácter

19 20 21

12 13 14 15 16 17 18

UTOs

. laxiflora

M. minutissima

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DES SO O O p ER RE MES Qe (eem OS OOS == OK OS Oo So

DTDOMDADAMM—MNMMDOMMADOM-—-_ BDAODAMNANNDAODGCGAGGO

OcOoOOoOOoo0o0-—o0-—o0-—nudoco-—-nuoooocoooocoo-—oooooooococosoc

VDO—Too-——-—--—--— NO0OOo0==00O0-——r0000-—=——-00000000

OQO—o00o0——-ANA—ANANA—=00-> 700-0000 N--00000000

VDO—TODOm-—m mm — COnr- oo 000000000000

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=sFae & 8 == US UY oad Bd T =.= m S ov x 64 6 O 4 gi

SuDEeEcecsSOgeunQnsaBeaegetrurnwmKBBeaesevrve GC ne LrHeu A

= Os evyvaoed Uo 3 au E 9 og SOGtvrtbt as Gsis § o

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SeStzszez2222S2S2S5S22 2222222222232 23223223232 222325322852

292 i itute of Texas 3(

Cuaoro 2. (continued)

E. Carácter

UTOs 1 2 3 4 5 6 12 13 14 15 16 17 18 19 20 21

M. utilis 1 1 1 2 1 2 2 2 1 1 1 1 0 2 0 0 0 0 0 2 O0

M. vaginata 1 1 1 2 1 2 2 3 2 1 1 1 O 1 0 0 0 0 0 2 OQ

M. versicolor 1 1 2 2 0 1 2 2 1 0 0 0 0 0 1 2 2 2 1 2 OQ

M. villiflora 1 1 1 2 3 3 3 2 1 1 1! 1 QO 2 1 1 1 1 1 1 0

M. virescens 1 1 1 2 0 1 2 3 T 1 1 1 2 3 1|! 2 2 2 1 2 O0

M. virletii 1 1 2 4 0 1 3 1 1 0 0 0 Q O 1 2 2 1 1 2 O0

M. watsoniana 1 1 3 2 0 1 1 2 1 1 1 1 2 3 1 3 1 1 2 2 0

M. wrightii 1 1 1 2 1 2 3 3 2 1 1 1 O 1 2 1 0 O 2 2 OQ

M. xanthodas 1 1 2 3 0 3 2 2 2 00000 1 2 1 1 2 2 Q

De los datos generados en las especies analizadas, se confirman las conclusiones obtenidas por Sod-

erstrom (1967), en lo relativo a los micropelos y cuerpos de sílice de los grupos Podosemum y Epicampes,

señalando que los micropelos del primer grupo son cortos y su célula distal es inflada (Fig. 4a), mientras

que los del segundo grupo son más largos y delgados (Fig. 4d). En cuanto a los cuerpos de sílice el autor

menciona que son del tipo de silla de montar formando una doble equis (Fig. 6b), este caracter de los cuerpos

de sílice lo comparten las 113 especies estudiadas del género, mientras que, los micropelos de las especies

que no se incluyen en estos dos grupos, presentan micropelos característicos de Podosemum o de Epicampes

en cuanto a forma y tamafio.

De acuerdo con Ellis (1979), las papilas en la familia Poaceae, ocurren en las células largas y cortas,

especialmente en las zonas intercostales, y el número puede variar de una a muchas por célula. En las espe-

cies estudiadas de Muhlenbergia la presencia o ausencia de papilas es un carácter importante para separar las

especies en grupos (IIIa y IIIb, Fig. 14), lo que por otra parte, confirma la propuesta de Vieira y col. (2002),

quienes señalan que la presencia o ausencia de papilas puede ser interpretado como un carácter de valor

taxonómico.

Los aguijones o ganchos son estructuras epidérmicas a las que se les ha dado poca importancia, en

algunos trabajos, sólo se menciona que son asperezas de la epidermis que se presentan tanto en la zona cos-

tal como intercostal (Vieira y col. 2002, Ellis 1979). Sin embargo, para las especies del género Muhlenbergia

fueron importantes, ya que de acuerdo a la presencia, ausencia, forma y tamaño (ver en Cuadrol), se forman

grupos definidos, por lo cual se consideran caracteres taxonómicos importantes en este género.

Las células cortas se presentan en pares o solitarias en hileras longitudinales, y se clasifican en células

silíceas o suberosas de acuerdo al contenido del lumen (Vieira y col. 2002). Los cuerpos de sílice dentro de la

célula silícea tienen formas características que han sido importantes para reconocer tribus o aun subfamilias

de la familia Poaceae (Ellis 1979). En las especies mexicanas del género Muhlenbergia se presentan cuerpos

de sílice del tipo característico en la subfamilia Chloridoideae, en forma de silla de montar formando una

doble equis, aunque en algunas especies se presentan cuerpos de sílice más largos O con la

] por el mismo

parte media más estrecha y los extremos red los o irregulares, son también

autor con forma de silla de montar.

La forma de estomas es otro carácter que se presenta en la epidermis. Los aparatos estomáticos son

subdiscoidales, compuestos de células subsidiarias redondeadas (en forma de domo). Se encuentran dis-

tribuídos en dos hileras longitudinales, una hilera a cada lado de la zona costal. Este carácter igual que los

cuerpos de sílice, son caracteres con valor taxonómico bajo por presentar formas con escasa variación en

todas las especies, por tal motivo no ayudan a la a de erupes:

q baxial en laminas de Muhlenbergia

Comparando | partit

con los de anatomía foliar en corte transversal de Muhlenbergia (Peterson y Herrera 2001), no se observan

RBacalac 2 Herrera, A + ía dal id : l ?4l limi Ens In Adsl L . 293

havial da Mortal hi : L P tenda el A

—

c) M. ciliata, M. emersleyi y M.

nnm con zonas definidas, (d—f) M. gypsophila, M. laxiflora y M. nigra con zonas no defi nidas.

Fc? P. J Anticlinalac ES Inl E O LE he og J zenada A J VAS A sdas £, je U. b) M.

pectinata, i idad fi le V. O M. ciliata, si idad fi le W. d) M. tenuifolia. Paredes lisas, ye)M. peruviana,

similitudes con las agrupaciones naturales (emanadas de un análisis cladístico) que dieron sustento a la

propuesta de reconocer los subgéneros (Muhlenbergia y Trichochloa) y las secciones (Epicampes y Podosemum)

en el subgénero Trichochloa. El dendrograma de epidermis abaxial (Fig. 14) muestra que las especies de los

subgéneros mencionados (Peterson y Herrera 2001) se intercalan en todos los [dein del dendrograma de

caracteres epidérmicos. En el primer y segundo grupos, por ejemplo, se asocia de Muhlenber-

gia, Epicampes y Podosemum. El tercer grupo está más definido, con la mayoría de las especies del subgénero

Muhlenbergia, sin embargo, también se agregan especies de las secciones Podosemum y Epicampes.

294 t i i Texas 3(

Fic 3 Tij In Danil Mahbhianh

Fl presente estudio es el primero que se hace con base en la epidermis abaxial de las especies del género

Muhlenbergia (Poaceae) de México, en el cual se estudiaron 113 de las cerca de 120 especies reportadas.

CONCLUSIONES

El conjunto d t idérmi hib especies del género Muhlenbergia de es demuestra

que hs epidermis por si Sola no sedes ser utilizada en la delimitación de especies, secciones y subgé de

Muhlenbergia. Podrían ser considerados caracteres adicionales a los morfológicos y los anatómicos de corte

transversal en láminas foliares, para la delimitación de especies.

De la formación de grupos epidérmicos diferentes a los anatómicos de sección transversal, en las espe-

cies del género Muhlenbergia de México, podría concluirse que la anatomía de epidermis foliar posiblemente

tuvo una evolución divergente a la de sus estructuras internas.

APÉNDICE

Esp de Muhlenbergia utilizad te estudio (colect grit ponden a las imágenes de las figuras

1-13).

M. aguascalientensis. Aguascalientes: Playa Mariquitas, M. alamosae lientes: B de los Lobos, Calvillo,

al NW de la Congoja, mpio. San José de Gracia, de la Cerda & Q. o 303 (UAA). Durango: Potrero la

García, Herrera y de la Cerda 5531 (UAA); de la Cerda Tinaja, 2 km de San Juan de Michis por el camino La

85 (UAA) Michilía, Súchil, YY. Pun 21 (CIIDIR). Jalisco: Arroyo del

Racalac R. Herrera, Anat, fn dal P| : L sal lim’ £41: 1, RAL hill “ 295

Fic. 4. Tipos de Micropelos, obtenidos por el método d do: a) M. strictior, pequeños e inflados < 30, b y «) 32 a 80, b) M. flaviseta micropelos

inflados célula distal 1/3 del total, c) M. quadrident icropelos ang y alargados, célula distal 1/10 del total, d) M. fragilis micropelos angostos

y alargados > 80.

ET; dni H

Ind n )M. . n la | I scj | [ ta, b) M. brevifolia con

aguijones de punta mas larga que la base.

296 Jou tani ti Texas 3(

Fic. 7. Macropelos, obtenid | método d I do: a) M. tenella, I | d istenci igid b) M. pubescens, con abundantes

macropelos largos y suaves.

Agua Caliente de los baños El Encanto, L.M. rd 14 M.annua. Durango: 14 km al NNE de San José de Avino, mpio.

(SLPM); ae ana à Tepetitián, Zapopan una 3746 Pánuco de Coronado, R. Serrano s.n. (INEGI).

(GUADA). N S p M. cacha Baja California: 1.5-2.5 mi upstream from

Nayar, mpio. El Nayar, Díaz Luna 9556 (GUADA). join Rincón, 4.5 mi NE of Sta. Catarina, 64 mi SE of Ensenada,

between La Fraguita and El Madroño, carr. Durango- RE. pue (MEXU); 10.5 km W of La Rumorosa along

Mazatlán, mpio. Concordia, Beetle M-3730 (ENCB). hwy 2, mpio. Tecate, Thorne 60422 (MEXU).

Rosales & Herrera, Anatomía de la epidermis abaxial en láminas foliares de Muhlenbergia

Fic. 8. Especies del género Muhlenbergia en el grupo lA: a) M. aguascalientensis, b) M. rigens, c) M. palmeri, d) M. pauciflora, e) M. capillaris, f) M. dubia,

9) M. glabrata, h) M. rigida, i) M. laxiflora. Nótese 1 sola hilera de estomas en la zona intercostal, aguijones en la zona costal y ausencia de papilas.

Fic. 9. Especies representativas de Muhlenbergia en el grupo IB: a) M. distichophylla, b) M. emersleyi, c) M. gooddingii, d) M. grandis, e) M. robusta, f) M.

xanthodas. Nótese 2 hileras de estomas en la zona intercostal, aguijones en la zona costal y ausencia de papilas.

M. atenacea. Coahuila: Cañón de la Vaca, sierra Santa Fé del M. arenicola. Aburto Chihuahua: 15 mi W of Jiménez, mpio.

Pino, mpio. Ocampo, J. Passini 5248 (ENCB). Nuevo León: López, Reeder & C. Reeder 4875 (ENCB). Durango: 9 mi N

2 km SSW Puerto Grande en el camino a Castillo, mpio. of Zacatecas state line along hwy 49, mpio. Sta. Clara,

Galeana, H. López s.n. (INEGI). Reeder & C. Reeder 6475 (ENCB). Nuevo León: 4 km al E

298

Fic. 10. Especies representativas del grupo IIA: a) M. appressa, b) M. microsperma, c) M. parviglumis, d) M. schreberi, e) M. polycaulis, f) M. brevifolia.

7 dea T 3 F

del ejido María de Ramos, mpio. Galeana, L. Ledesma s.n

(INEGI). San Luis Potosí: Estación Berrendo, mpio. Charcas,

Rzed 6575 (SLPM)

M. argentea. Chihuahua: 15.3 mi S of Mex 127 and 6.9 mi

E of la Bufa, mpio. Guachochi, PM. Peterson, Annable &

Sierra San Pedro Marti; MHG Mexicali, Moran aa

(ENCB); cerca Ob , Sierra

San Pedro Martir, mpio. Ensenada, F. Takaki s.n. (NEG).

M. articulata. Aguascalientes: Milpillas de Arriba, mpio.

Jesús María, Zúñiga 20 EXU); Arroyo El Terrero, mpio.

Y Herrera 971 (CIIDIR).

M. arizonica. Chihuahua: Chihuahua

Carretas, mpio. Janos, L.H. pe 1624 (TAES); Rocky hills

near Chihuahua, Pringle 402 (ENCB). Durango: about 41

mi N of Cd. Durango, mpio. Canatlán, Reeder & C. Reeder

6482 (ENCB).

M. arsenei. Baja California: E of Upper Vallecitos Meadow,

Anos border Rancho

Calvillo, Zúñiga 236 (MEXU). Hidalgo: 20 km al SSE de

Ixmiquilpan, mpio. Progreso de Obregón, L. González

31 ant MER S 4 km al W de Sta. Ma. Amajac,

mpio. At | , J. García 139 (MEXU). Jalisco:

km 219 dd i carr. México: Nuevo Laredo entre Zimapán

y Jacala, mpio. Zimapán, A. Cruz 1417 (ENCB). Querétaro:

1 km al S de Vizarrón, mpio. Cadereyta de Montes, Mora

Rosales 8: Herrera, Anatomía de la epidermis abaxial en láminas foliares de Muhlenbergia

Fic. 12. Especies del grupo IIIA: a) M. annua, b) M. implicata, c) M. minutissima, d) M. strictior, e) M. texana, f) M. fragilis. Nótese la ausencia de papilas

y de aguijones

Fic. 13. Especies incluidas en el grupo IIIB: a) M. depauperata, b) M. plumbea, «) M. repens, d) M. arsenei, e) M. asperifolia, f) M. arizonica, g) M. vaginata,

h) M. utilis, i) M. majalcensis. Nótese la presencia de papilas y la ausencia de aguijones.

300

qq AAA

PATA

. elongata (P)

, scoparia (E)

tie CA)

. sextonii m

. tenuissima

TILA

Fi. 14. Dendrograma agrupando 113

7 | breviseta (A)

E i i = hintonii (P)

H z BA LS £ . . J + tint

SA DNI g PE RENE | M uet cui

301

slac & Harrara A

Benítez 353-AMB (MEXU). San Luis Potosi: 10 km al W de

Guadalcázar, Rzed. 4930 (ENCB); 3 km al SW de Guadal-

cázar, Rzed. 6809 (TAES).

. asperifolia. Baja California: Sierra San Pedro Martir, mpio.

Mexicali, R. Morán 17973 (TAES). Chihuahua: about 9 mi N

of Las Delicias, Reeder & C. Reeder 2615 (ENCB). Coahuila:

Ferriho's laguna area, mpio. Cuatrociénegas, D. Pinkava &

E. Lehto & D. Keil 5409 (ENCB)

. brandegei. Baja California Sur: aprox. 25 km S of La Paz,

W side p Isla La Partida, mpio. La Paz, PM. Peterson &

Annable 4760 (US).

brevifolia. a Ejido El Zapote, mpio. Pénjamo, Mora

enítez 251-AMB (MEXU).

M. E Ha. C

to Pinola Las Rosas, Breedlove & Raven 13 135 (TAES); near

Venustiano Carranza along the road to pugiltic, Laughlin

2702 (TAES).

brevis. Chihuahua: 0.7 mi W of Nuevo Majalca, 8.5 mi W

of hwy 45, P.M. Peterson & Annable 4512 (ENCB); 15.6

mi NE of El Vergel on hwy 24, mpio. Balleza, Peterson &

Annable 4061 (US). Durango: 5.4 mi W of Navíos, 42 mi W

of Durango on hwy 40, Peterson & Annable 4586 (US).

breviseta. Jalisco: about 12 mi E of Zacapú on road from

Guadalajara to Morelia, Reeder & C. Reeder 1447 (ENCB).

Michoacán: Cerros Las Viudas y Las Flores, al SE del Rancho

Seco, A. Martínez 871 (ENCB).

brevivaginata. Aguascalientes: Laguna Seca, mpio. San

José de Gracia, Y. Herrera & O. Rosales 604 (UAA). Durango:

Potrero Las Escobas, San Juan de Michis, mpio. Súchil, J.

Alvarado 725 (CHAPA). Zacatecas: km 30 de la terracería

entre Tlaltengo y Jalapa, mpio. Tlaltenango de Sánchez

Román, Balleza 2720 (ENCB)

. capillaris. Jalisco: Las Popas de Arriba, 14 km al S de

Ojuelos, Santana-Michel 1507 (ENCB). Oaxaca: 4 km al

S de Ejido Benemérito de las Americas, camino a Flor

de Cacao, E. Martínez 10785 (MEXU). Querétaro: 3 km

al S de San Joaquín, mpio. San Joaquín, R. Fernández

3248 (ENC

capillipes. Chihuahúa Rancho “eseachic, mpio. Namig-

uipa, dia M 3266 o 22 mi SE of Chuichupa on

] o. Madera, Reeder & C. Reeder 2673

(US); 24 mi SW of A he on road to Creel, mpio. Guer-

rero, Y Herrera 953 (ENCB); 0.5 mi W of Flores Magón, mpio.

a, Reeder, & Soderstrom 3496 (US).

Buenaventu

. ciliata. fie km 29 por la carr. Zapopan-San Cristóbal

de la Barranca, Santana-Michel 1973 (IBUG); parte más

alta del Cerro Grande de Cujaruato, al SW de la Piedad,

Rzed. & McVaugh 515 (ENCB). Michoacán: Cerro Colorado,

mpio. Pátzcuaro, J.M. Escobedo 240-A (ENCB); 7 km al

E de Villa Jiménez, sobre el camino a Copándaro, Rzed.

40757 (ENCB)

. confusa. México: Terrenos de Juchitepec, mpio. Juchite-

ec, O. Domínguez (MEXU)

. crispiseta. Durango: 6 mi W of Río Chico crossing W of

Cd. Durango, Reeder & C. Reeder 4645 (CHAPA); Camino

los Herreras-Topia km 64, mpio. Santiago Papasquiaro,

M. González-Elizondo 3252-a (CHAPA).

cualensis. Jalisco: Minas Zimapán, por la brecha a Cuale,

M. dist

R. Guzmán 6100 (MEXU); Minas Zimapán, por la brecha a

Cuale, Santana-Michel 1217 (SLPM); El Caracol, 1 km al W

de la Mina de Zimapán, González.-Villareal 3283 (ENCB).

curvula. Uau 6 mi NE ona Rzed. 16133

(UAA) Dolores

Hidalgo, Rzed. 42071 (CHAPA). San Luis Potosí: Sierra de

San Miguelito, 30 km al WSW de San Luis Potosí, carr. a

Villa de Arriaga, Rzed. 29381 (CHAPA). Zacatecas: aprox. 38

km al W de Jalpa sobre la carr. a Tlaltenango, 30 km del

entronque con la carr. Jalpa-Juchipila, pe & McVaugh

974 (ENCB).

densiflora. Chiapas: about 15 mi SE of Teopisca, Reeder &

C. Reeder 2045 (ENCB).

depauperata. Chihuahua: just N of Villa Matamoros on

wy 45 towards Parral, PM. Peterson & Annable 4579 (US).

Hidalgo: about 8 mi S of Ixmiquilpan, mpio. Chilcuautla,

Reeder & C. Reeder 1611 (CHAPA). Jalisco: 1 ql al N de

Ventura, mpio. Villa Hidalgo, Rzed. 11455 (TA

ans. Aguascalientes: El Zapote, 1 km al S i Cosío, de

la Cerda4194 (UAA). D.F: base de la Sierra de Sta. Catarina,

al N de Tlatelolco, Delegación Tláhuac, Rzed. 37470 (SLPM).

Jalisco: Sierra de Cuale, entre Cuale y Talpa, mpio. Talpa

de Allende, J.V. de la Torre 8634 (ENCB). México: Sierra de

Guadalupe, base del cerro Tontepec al SW de Coacalco,

mpio. Coacalco de Berriozabal, Rzed. 38248 (TAES).

distichophylla. Jalisco: SSE of Puerto Vallarta, 14 km from

the opening ofthe Zimapán Mine, 1 km W of Providencia

OME El aus El Cuale, d -Michel 2348 (ENCB);

| Río Salado, 0 km al W de Guada-

lajara, Rzed. 15490 (ENCB).

diversiglumis. Durango: Canelas, 1 km por el camino a

UM i González- Elizondo 274- ENED). Med i

km al S lel M

mpio. Pere Rzed. 41323 (UAA).

dubia. Aguascalientes: Bajío La Canoa, Sierra San Blas de

Pabellón, mpio. San José de Gracia, de la Cerda 4085 (UAA).

Coahuila: 153.2 km NW of Müzquiz on hwy 53 towards

Boquilla del Carmen, PM. Peterson & Annable 10594 (US).

Guanajuato: Rancho de Enmedio, Sierra Sta. Rosa, Y.

Herrera & Cortés 1421 (UAA). Jalisco: 25 km al NW de

Bolafi or la brecha a Los Asmoles, mpio. Bolaños,

Satna Miche 1766 (IBUG). San Luis Potosí: Sierra La Viga,

We pat ce eee 1155 (INEGI).

, mpio. Comala, M. Navarrete

aprox

M.dum

596 EO Michoacán: about 25 km S of Arteaga, road

to Playa Azul, mpio. Lázaro Cárdenas, Díaz Luna 9300

(GUADA). Zacatecas: Florencia, 3 km al W del pueblo,

mpio. Benito Juárez, Díaz Luna 15291 (GUADA).

el camino a Platanitos, mpio. Mezquital,

Elizondo 1692 (US); bajada del Pam a San Fco. de Lajas,

mpio. Pueblo N 1 (ENCB). Jalisco: km

22.5 brecha Bolaños-Tenzompa, mpio. Mezquitic, Ramírez

& Guzmán 641 (IBUG).

elata. 2 km al E de El Limón, mpio. Santa Ma. del Oro, M.J.

Sepúlveda 987 (CIIDIR).

Nuevo, S.

M. elongata. Chihuahua: Cañón del 54, mpio. Chihuahua, J.M.

Peña s.n. (ENCB); 12 mi S of Villa Matamoros on hwy 45

302

towards Parral, Reeder & C. Reeder 4886 (ENCB); Batopilas,

mpio. Batopilas, R. Fierros 1663 (MEXU).

. eludens. Chihuahua: hills near Chihuahua, Pringle 3052,

3056 (TAES); El Tigre to El Rancho de Robles, NE Sonora,

mpio. Bavispe, J. Vera 1957 (CHAPA); Tomochi, 2 km al-

rededor del poblado, mpio. Guerrero, M. Vergara 153

(MEXU)

M. emersleyi. Jalisco: 7 mi SSW of Mazamitla, Reeder & C.

Reeder 6329 (ENCB). Durango: 4 km al W de La Parrilla,

mpio. Nombre de Dios, S. Acevedo & 5. González 2779

(ENCB). Querétaro: Puerto de Los Velázquez, mpio. Pinal de

Amoles, V. Jaramillo, G. Villegas & A. Miranda 808 (MEXU).

. eriophylla. Aguascalientes: lado SW de presa La Arana,

dun e. José de ae G García 1371 da Guana-

juato: 12 km al NE d , mp Enano ee

s (CD Burange: Cerro Bla

fera"La Michilía”, mpio. Súchil, M. E Elizondo 3740

(CIIDIR). Michoacán: Cerro de Ucareo, mpio. Zinapécuaro,

L. Oseguera 52 (MEX

M. firma. Chiapas: about 15 mi SE of Teopisca, mpio. Teopisca,

Reeder & C. Reeder 2045 (ENCB). Guanajuato: 53 km al E de

San Luis de la Paz, sobre carr. a Xichú, mpio. Victoria, Rzed.

41390 (CIIDIR). México: 6 km de Tlazala de Fabela por la

carr. a Nicolás Romero, mpio. Isidro Fabela, M. González-

Ledezma, J. García & |. Hernández 108 (SLPM).

M. flavida. Chihuahua: 15.3 mi S of Mex 127: and 69 mi NE ofla

Bufa, mpio Guachochi, M.E. S

7 (UAA); Sier Td

Milpillas 28.1 mi E of Los Tanques on road to Chinipas, 1.6

mi N of Los Chinipas, PM. Peterson & Annable 4174 (MEXU).

o 4 km W de El Talayote, 35 km E de Yécora, mpio.

ora, P Tenorio 4520 (ENCE)

M. E ta. Chihuahua: Parq ascada Basase-

achic, 1 km airline S of Cascada, mpio. Ocampo, M. Vergara

139 (MEXU). Durango: Predios da EIS La Victoria, mpio.

Pueblo Nuevo, D. A J); Potrero Chaides, 9

km al NE de La Mesa, brecha a ElYaqui, mpio.Tepehuanes,

P Tenorio & C. Romero 2080 me ^c

M. fragilis lod R li

mpio. Comonfort, JJ. Macías 591 (MEXU). Guerrero: 18 km

Chil de los Bravo,

O. Tellez 4552 (MEXU). Sonora: N sf Cananea e José

María Morelos, Beetle M-7867 (EN N de

Pueblo Viejo, sierra de Md cerro i Piñones, ladera

; mpio Juchip ila Bali Adamo 0102 (SLPM).

M. gigantea. Chiapas Kulaktik, mpio. Tenejapa, A.S. Ton 4570

(ENCB); 12 km S of Mexican hwy 190 near Rizo de Oro,

mpio. Cintalapa, Breedlove 20641 (TAES). Durango: orillas

del pueblo de Topia, rumbo a Cuevecillas, A. Benítez 1105

(CHAP). Jalisco: km 217 carr. Guadalajara-Barra de Navidad,

mpio. Cihuatlán, L.M. Villareal 8303 (IBUG). Michoacán:

5 km al N Tlalpujahua, mpio. Tlalpujahua, 16-Dic.-1986,

Rzed. 42265 (ENCB).

glabrata. Michoacán: 9 mi W of Zacapú, mpio. Zacapú,

Reeder & C. Reeder 4746 (MEXU). San Luis Potosí. 5 km E

Soledad de Zaragoza, mpio. Xilitla, Rzed. 7122 (ENCB);

Sierra de Monte Grande, arrcyo de Carbonerías, al NW

del cerro de Juan Chepe, mpio. Charcas, J.A. Reyes 375

(CHAPA)

Ina

al NW de Chilpancingo, mpio

T Af Dat H ID l.l EHE fT,

M. glauca. Aguascalientes: Bajío La Canoa, Sierra San Blas

de Pabellón, mpio. San José de Gracia, de la Cerda 4090

(UAA). Coahuila: Sierra de Zapalinamé, 5 km S Saltillo

in San Lorenzo Canyon, mpio. Saltillo, Hatch, Morden &

J. Valdés 4498 (MEXU). Durango: 15 mi W of El Pino, 29

mi W of Durango, Reeder & C. Reeder 3127 (ENCB). San

Luis Potosí: 12 km W Estación Berrendo, mpio. Charcas,

Rzed. 6576 (CHAPA); Sierra de Monte Grande, Tanque de

adentro, mpio. Charcas, J.A. Reyes 579 (CHAPA); 3 mi SW of

San Luis Potosi, Pringlei 1893 (TAES); km 30 carr. San Luis

Potosí-Ojuelos, 3 km entronque a la Amapola, mpio. Villa

de Arriaga, A. Romero s.n. (SLPM).

M. gooddingii. Sonora: 19 km alW de Yécora, Beetle M-6169

(MEXU); S of Ures, mpio Ures, Tañori 8310 (MEXU); El

Cañón de la Bellota, Sierra de la Cabellera, mpio. Agua

Prieta, J. Vera 2110 (US).

M. grandis. Guanajuato: 9 km WNW de la Valenciana, mpio.

Guanajuato, Rzed. 49767 (CIDIR). Jalisco: ca 13 mi NNE

of Guadalajara, mpio. Zapopan, Reeder & C. Reeder 4724

(US); cerca de El Corte Colorado, La Resolana, Rezedowski

15037 (ENCB).

I li |

, Q.Domín-

guez 1000, 1254 (MEXU); D. Fuentes s.n. (ENCB); La Joya,

Cuesta Blanca, 15 km de Aramberri rumbo a Zaragoza,

mpio. Zaragoza, J. Valdés VR-2554 (ENCB); 2 km al 5 del

Salero, mpio. Galeana, Valdés & Marroquín 1619 (ENCB).

San Luis Potosí: 12 mi NW of Matehuala on road to Cedral,

Reeder & C. Reeder 4812 (ENCB).

M. hintonii. México: on del Llano de la Horca, mpio.

Tlalmanalco, Rzed. 31662 (SLPM).

.implicata. vie toes Barranca El Temascal, mpio. Cal-

villo, L. Juárez 987 (UAA). Chihuahua: 0.7 mi W of Nuevo

Majalca, 8.5 mi W of hwy 45, PM. Peterson & Annable 4514

TAES); La Joya, Cuesta Blanca, 15 km de Aramberri rumbo

a 3 Zaragoza, e i inan J. Valdés VR-2554 (ENCB).

mascaltepec, Hinton 2079

(ENCB). 14.8 mi NE of El acd mpio. Balleza, PM. Peterson

& Annable 4079 (TAES). Michoacán: 15 mi W of Jiquilpan,

mpio. Jiquilpan, Reeder & C. Reeder 6326 (ENCB).

M. iridifolia. Jalisco: 15 km S Talpa, sobre camino a La Cuesta,

mpio. Talpa de Allende, Rzed. 15235 (ENCB).

M. jaliscana. Jalisco: camino Sta. Rosa, 1 km antes del Sal-

vador, mpio. Tequila, A. Navarro 213 (IBUG); San Cristóbal,

mpio. Zapopan, A. Navarro & E. Cervantes 158 (IBUG).

M. jonesii. California: 9.5 mi SE of hwy 97 on Military Pass Rd,

NE side of Mt Shasta, PM. Peterson & Annable 4857 (US).

M. laxa. Veracruz: Cerro Macuiltepec, just outside Jalapa,

mpio. Xalapa, Reeder & a pus 5991 (ENCB); Cerro Ma-

cuiltepec, mpio. Xalapa, V. Hernández 80 (MEXU

M. beds MU non a mi D iu El one iii Puce

CB) IW

of Oaxa ca, mpio. Oaxaca, Reeder & C. E (MEXU).

M. lindheimeri. Coahuila: 4 mi WSW of Nava, Reeder & C.

Reeder 4814 (ENCB).

M. longiglumis. Guanajuato: 28 km NNW Juventino Rosas

sobre carr. a Guanajuato, Rzed. 41197 (CHAPA). Jalisco:

RUN. mpio. Guadalajara, Pringle 2365 (US); Ayo El

Grande, mpio. Jesús María, A. Navarro 781 (MEXU).

Racalac 2, Harrara A + ia dal id i I ial

J Anhi k E 303

M. da Chihuahua: about 15 mi S of Madera,

adera, Reeder & C. Reeder 2639 (ENCB). Sonora:

N of E Ejido José María Morelos, Beetle M-7823

(MEXU).

M. lucida. Chihuahua: 10.7 mi S of Mexico 127 on road to

Batopilas, Y. Herrera, PM. Peterson & Annable 946 (CIIDIR);

e la Junta de Basaseachic, mpio. Guerrero, R.

Fierros 1691 (MEXU).

s. Ag Río Gil e ua

de la Ce rda. 4409 (UAA); B EIE

I s E d la Ed 3943 (UAA). ARa E im a E

s Joyas

mpio. Autlán, EJ. a TS. Cochrane 4818 (BUG).

. macroura. Aguascalientes: Bajío La Canoa, Sierra San

Blas de Pabellón, mpio. San José de Gracia, de la Cerda

4103 (UAA). Puebla: Campo experimental San Juan Tetla,

vertiente E a Ixtaccihuatl, mpio. San Martín Texmelucan,

A. May Nah 971 (ENCB).

M. majalcensis. Chihuahua:

Y mi E of Cumbres de Majalaca, mpio. Riva Palac

Peterson, Annable & Y. Herrera 949 (CIIDIR).

michisensis. Durango: Potrero Las Escobas, San Juan de

Michis, mpio. Súchil, J. Alvarado 723 (CHAPA)

M. ricus Michoacán: 51 km de Pátzcuaro rumbo a

Un ruapan, mp o. Ziracuaretiro, A. Díaz Osorno 358 (ENCB);

Cañada del río Grande, cerca de Cointzio, mpio. Morelia,

Rzed. 45281 (CIIDIR). Oaxaca: 6 km al E de Guadalupe

Cuautepec, mpio. Santiago Miltepec, A. Salinas & A.

Campos 3690 (MEXU). San Luis Potosí: Rancho El Palmar,

potrero Tortugas, mpio. San Luis Potosí, LV. Rivas € M.

González-Ledezma 226 (SLPM).

M. minutissima. Chihuahua: Cal del

goata, mpio. Bocoyna, B. Tah 44 (MEXU). Tamaulipas: km

26 carr. Victoria-San Luis Potosí, mpio. Victoria, G. Villegas

169 a Tlaxcala: camino de San Cosme Xalostoc

5.5 km de la carr. Apizaco-Huamantla, H.

as 20 (CHAPA).

. montana. Aguascalientes: La Ciénega, mpio. San José

de Gracia, de la Cerda 3989 (UAA). Chihuahua: carr. San

Juanito-La Junta, 17 km, mpio. Guerrero, B. Tah 8 (MEXU);

Chihuahua-Sonora border, Rancho Carretas, mpio. Janos,

Harvey 1639 (ENCB); 12 mi SW of Chuhichupa, mpio.

iE nis Spellenberg & Zimmerman 8951 (ENCB). Durango:

alW de anaran, “OE Mesa de Ae : di

s.n LES Oaxaca: 2 San Antonio Aba

Concepción Buenavista, R. Cruz 2322 E ).

monticola. Coahuila: 25 km S of Saltillo at the rancho

Recreacion El Chorro, mpio. ec Hatch, Morden & Valdés

4590 (TAES); 5 km S of Saltillo, in San Lorenzo Canyon,

mpio. Saltillo, Hatch, Morden & e 4495 (TAES).

. mucronata. Durango: 7.3 mi N of Charcos on road towards

San Juan de Michis, mpio. Mezquital, PM. Peterson, S.

González & Knowles 13646 (US).

nigra. México: Paso de Cortés, entre Popocatepetl e

Iztaccihuatl, mpio. Amecameca, A. Cortés s.n. (INEGI);

Parque Nacional Nevado de Toluca, mpio. Texcaltitlán,

Beaman 1708 (ENCB).

M. orophyila. México: Paso de Cortés, 13 km al E de Amec-

N of México 45 and about

io, P.M.

ced na R

ameca, mpio. San Nicolás de los Ranchos, Koch 76266

(CHAPA).

M. palmeri. Chihuahua: Sierra Madre Occidental, along río

Corareachi, P.M. Peterson & Catalán 17605 (CIIDIR);

Parque Natural Barranca del Cobre, 1.6 km E of La Bufa,

mpio. Guachochi, PM. Peterson & Knowles 13579 (US).

M. parviglumis. T Zempoala, mpio. Zempoala, A.

Ventura 1495 (ENCB).

M. pauciflora. Sonora: E of Cananea, Sierra de los Ajos, mpio.

Frontera, Beetle M-7869 Ee

i of Taxco on hwy 55, mpio.

Pilcaya, P.M. Peterson pui (ENCB).

M. peruviana. Aguascalientes: El Pandito de Vázquez, Sierra

de San Blas Pabellón, mpio. San José de Gracia, G. García

2335 (UAA). México: 2.3 miE of! y3on road to Nevado

de Toluca, mpio. Zinacantepec, PM. Peterson & Annable

4636 (TAES); 6 mi W of pass on road from Toluca to Te-

mascaltepec, mpio. ii ae 17197 (ENCB).

Michoacan: 9.3 mi ,mpio. Salvador

Escalante, PM. ad &Annable 4622 (ENCB).

M. plumbea. D.F.: Cuilotepec, Sierra de Ajusco, deleg. Tlalpan,

E. Manrique 1144 (MEXU). Hidalgo: Laguna de Apan, mpio.

Apan, R. Cruz 411 (ENCB). Tlaxcala: Camino de San

Cosme Xalostoc a Terrenate, 5.5 km de la carr. Apizaco-

Huamantla, H. Vibrans 2836 (ENCB).

M. polycaulis. lo Barranca El aa mpio.

an J 3943 Chihuahua:

6 mi E of Buenaventura o C ne 3200 (ENCB).

Sonora: El C erra de la Cabellera, mpio

Aqua Prieta, J. Vera 2097 ER o su Jeréz-

Guadalupe Victoria, mpio. Jerez, Balleza 2222 (CHAPA),

. porteri. Durango: 3.5 km al ESE de El a carr. 45 a

Durango, J. Blando s.n. (INEGI). San Luis Potosí: km 428

carr. Central Mpio 28m Fus Potosí, A. Gómez 925 (SLPM).

,La Mesa de Las Carreras,

W of Colonia Morelos J. Vera 2054 (US).

Hus San José de Gracia, G. García s.n. (UAA); 8 km al

W de la uc El Garabato, mpio. Jesús María, de la

Dos 3486 (UAA). Durango: 10 km al SW de Süchil, carr.

en construcción a Tepic, mpio. Süchil, S. González & Y.

Herrera 1512 (CIIDIR). San Luis Potosí: La Amapola, ejido

El Pachón, pu d Eus Fotos A. Avila 37 (SLPM).

M. pubiglum Demostrativo"Los Angeles"

48 km al S = Saltillo, J.A. Villarreal & J. Valdés 3370

)

(ENCB).

M. purpusii. San Luis Potosí: Entronque a San Agustín, mpio.

Matehuala, S. Villegas y E. García s.n. (TAES); San José del

Refugio, km 550 carr. central entre el entronque y Mate-

huala, F. Medellín 1130 (SLPM).

M. T México: 10 km al E de Amecameca, sobre

el camino a Tlamacas, Koch 76269 (TAES). Michoacán:

parte EN del cerro del Burro, mpio. Huiramba, Rzed.

44627 (UAA). Veracruz: Vidal Díaz Muñóz, mpio. Perote,

G. liso il NEC.

M.racem lel hoEl B

12 km E E headquarters, mpio. Acuña, J. Valdés & D.

Riskind VR-1277 (TAES).

304

M. ramulosa. Baja California: E of Upper Aeadow,

Sierra San Pedro Mártir, mpio. sia M. ind Real 300

(MEXU). Chihuahua: km 85 carr. Balleza-Gi i

Guachochi, M.E. Siqueiros 1606 (MEXU); 12 mi NW of M se

era, mpio. Madera, Reeder & C. Reeder 2649 (SLPM). Jalisco:

1 km al SW de la comunidad Maquixtla, mpio. Zacatlán,

S. Aguirre S-189 (MEXU). México: Volcán de Toluca, mpio.

Toluca, R. Sohns & E. Matuda 1006 (TAES); camino viejo al

be nia ae ju y Juchitepec, A. Manda E

Villega XU). Micl

ius e Zinapécuaro, Rzed. alee (MEXU).

reederorum Si

Durango, J.A Ochoa &R. A 315 (MEXU)

repens. Aguascalientes: La Ciénega, mpio. San José de

Gracia, dela Cerda 3981 (UAA). Jalisco: La Tapona, mpio.

Tula, M.H. Cervera 132 (MEXU). San Luis Potosí: Cerro San

mpio

Llanitos, mpio. Mexicali, Moran 28016 (U

M. rigens. Aguascalientes: Camino cerro del Tepózán (orilla de

arroyo), mpio. Calvillo, G. García 286 (UAA); Puentes Cuates,

mpio. Calvillo, de la Cerda 9398 (UAA). Durango: W de Sta

María de Ocotán, mpio. Mezquital, M. González-Elizondo

y S. Acevedo 12 "en "is

M. rigida

José de Gracia, G. García 1867 (UAA). Chiapas: about 2

mi SE of San Cristóbal de las Casas, Breedlove 8: G. Davidse

54703 (SLPM). Durango: pe ud al E de Durango, carr.

lulosa, Gould &

IW de La Estancia mpio San

a México

J. Morrow 7916 (TAES). Jalisco: 15 km al SW de Matanzas,

mpio. Ojuelos, Díaz (GUADA); 2 mi NW of Te-

quila at the edge of the Barranca Country, mpio. Tequila,

Cervantes y Castro 19 (MEXU). Michoacán: alrededores de

los filtros viejos, cerca de Morelia, A. Miranda, G. Villegas

&R. Mr M (MEXU). Tlaxcala: Hacienda San Francisco,

a de Hermenegildo Galeana, mpio. Huamantla, A.

rime 669 (MEXU).

M. robusta. Durango: km 60 carr. Durango-Mazatlán, mpio.

Durango, Beetle M-7745 (MEXU). Guanajuato: 2 km al NW

de La Gavia, sobre el camino a Cortazar, mpio. Cortazar,

Rzed. 40924 (SLPM). México: parte alta de la Sierra de

Alcaparrosa, mpio. Tepotzotlán, Rzed. 29939 (SLPM). Pue-

bla: carr. San Fco. Totimehuacán a Santo Tomás Chiautla,

mpio. Puebla, 5. Aguirre S-140 (MEXU). San Luis Potosí:

Cerro El Chiquihuitero, cerca de Tortugas de Arriba, mpio.

Santa María del Río, F. Tataki 1984 (UAA). Zacatecas: 3

km al E de Maravillas, mpio. Villanueva, Ma.C. Arteaga

722 (MEXU)

M. scoparia: A oe bn Caliente de los baños El

n, L.M. Villareal 7317

(IBUG); El Tuito, mpio. Cabo Corrientes, M del C. Mendoza

s.n. (IBUG). Michoacán: El Cerrito, 1.5 km E Jeráhuaro, mpio.

Zinapécuaro, M.J. Jasso 1709 (CilDIR)

M. schreberi. Tamaulipas: La Begonia, 2 km al S de San José,

mpio. San Carlos, Briones 1306 (SLPM).

M. seatonii. México: alrededor de Peñas Largas, mpio. Epa-

zoyucan, Rzed. 38485 (CIIDIR)

M. setifolia. Coahuila: 26 mi NW of Rancho Margareta, 90 mi

NW of Sabinas, mpio. Müzquiz Gould 10688 (TAES).

M. sinuosa. Sonora: E of Cananea, Sierra de los Ajos, mpio.

Frontera, Beetle M-7875, M-7876 (TAES).

M. prise Jalisco: 4-5 km SE de Platanarillos, 11-12 km

ESE de Minatitlán, R. Cuevas & L. Guzmán 1182 (IBUG).

o 24.4 mi NW of Surutato, 12 mi SW of Los Hornos,

mpio. Badiraguato, ie 16816 (TAES).

M. spiciformis. Chiapas: 3 mi S of La Trinitaria, mpio. La

Trinitaria, PH. Raven ~ (TAES). Querétaro: 2 km de

nal de Amoles carr. a Jalpan, mpio. Pinal de Amoles,

lo Rincón 206 (MEXU)

M. stricta. Durango: about 3 mi E of El Salto, mpio Pueblo

Nuevo, Reeder & C. Reeder 2551 (SLPM). Jalisco: 5 km al W

de Xalisco, J. Sepúlveda 846 (MEXU); La Primavera, en la

Arena, mpio. Zapopan, R. González 87 (MEXU). Zacatecas:

x oan SE el PHI ge State Ie on hwy 49, then 3 km to

los, mpio. Juan Aldama,

E Spellenberg & M. Mahrt 10725 (CIIDIR).

M. strictior. Chihuahua: Cusarare, JEN. Pas Guachochi,

R.A. Bye 5191 (MEXU); Parq ada Basase-

achic, 1 km airline S of Cascada, mpio. inc Torres 3777

(MEXU). Durango: about 9 mi E of El Salto, mpio. Pueblo

Nuevo, Reeder & C. Reeder 4676 (ENCB).

M. passis Durango: km 95 carr. Durango-Mazatlán, a 5

del Salto, mpio. Pueblo Nuevo, Miranda 1382 (MEXU);

E 09 carr. Durango-Mazatlán, mpio. Pueblo Nuevo,

Beetle M-7735 (MEXU).

M. tenella. Jalisco: Barranca El Huizichi, mpio. Tolimán, Díaz

Luna 20145 (GUADA). México: 1.75 km al NE de Nesón

Nuevo, mpio. Tejupilco, A. Ramírez 595 (MEXU). Nayarit:

alrededores de Tepic, F idcm i oe eee us

Linga, mpio Soledad de Doblado

El Aguacero en el rio La Venta, mpio. Ocozocuautia, B

Martínez 22006 (MEXU).

M. tenuifolia. Aguascalientes: 2 km al E de Tepezalá, mpio.

Tepezalá, de la Cerda 3894 (UAA); Puentes Cuates, mpio.

Calvillo, de fa Cerda jd (AA) W e. la ee de mi-

croondas, mpio. Ri 418 (UAA).

Guanajuato: Rancho Divino M km 21 carr. Dolores

Hidalgo-San Luis Potosí, mpio. Dolores Hidalgo, S. Rojas

174-SRV (MEXU). Querétaro: 1 km al S de Vizarrón, mpio.

Cadereyta de Montes, Zamudio 3356 (UAA

M. tenuísima. Jalisco: aprox. 45 km al E de El Aserradero,

sobre el camino a Manuel M. Diéguez, L.M. Villareal 5870

(IBUG). Nayarit: alrededores de Tepic, Beetle, G. Harding y

R. Guzmán M-6293 (MEXU).

M. texana. Chihuahua: 22 km al S de Humira, 21.8 km al SW

de la dev. a La Bufa, mpio. Batopilas, R. Torres 3690 (MEXU);

Sierra Milpillas, 24.1 mi E of Los Tanques on road to Mil-

pillas, mpio. Chinipas, Beetle, R. Alcaráz & R. Cuadra 9203

MEXU). Nayarit: Puente de Comatlán, mpio. La Yesca, A.

Castro 081 (MEXU

M. torreyi. Sonora: N of Cananea, Ejido José María Morelos,

Beetle M-7866 (TAES). San Luis Potosí: an Vicente Guer-

rero, mpio. Charcas, H. Bravo 0003 (MEXU).

M. utilis. Chihuahua: Mesa de El Poleo, ejido El Largo, mpio.

I F4 H Enl J RA. Ll Ļ = 305

Madera, Blanco 08/77 (IBUG). Guanajuato: Rancho El M. villiflora. Nuevo León: La Cardona, mpio. Bustamante, R.

Molino, mpio. Dolores Hidalgo, A. Santillán 673 (MEXU). López & A. Brito 042a (MEXU); Ejido El Jarro, 14 km del po-

Hidalgo: Pachuca, mpio. Pachuca de Soto, R. Guzmdn 3786 blado Dr. Arroyo carr. a Matehuala, J. Garza & M. Castillo 192

(MEXU). México: 1.25 km al SE Ge San Diego Linares, mpio. (MEXU). San p dd km 195 Carr. SUELO; San Luis

Toluca, A. Ramirez 497 (MEXU). San Luis Potosi: Ejido Cerrito Potosí, mpio. S , M.A. Reyes 64

Blanco, 10 km E Matehuala, A. Bolaños 203 (MEXU). (MEXU). Zacatecas: Palula, mpio. Villa de mos E. Manrique,

M. vaginata. Colima: Sierra de Halo, near a lumber road Beetle & G. Harding 1285 (MEXU).

leaving the Colima hwy 7 mi SSW of Tecalitlán and SEto M. virescens, Guanajuato: 6 mi NE of Guanajuato, Reeder &

San Isidro, Koch, J. Villa & J. Chávez 267 (SLPM). D.F.: Caldera C. Reeder 4750 (ENCB); 21 km al W de Dolores Hidalgo,

El Guarda, al S de Parres, deleg. Tlalpan, A. Miranda y P sobre la carr. a Guanajuato, Rzed. 41131 (IBUG). Michoacán:

Guerrero 101 (MEXU). Durango: km 115 carr. Durango- 19 km al E de Queréndaro, sobre la carr. a Maravatío, mpio.

Mazatlán, mpio. Pueblo Nuevo, A. Rangel 410 (MEXU). ie nee ie 40988 All

Jalisco: 1-2 mi E of Tapalpa, Santana-Michel1441 (IBUG). M. virletii. Mé Juchitepec, mpio. Juchitepec

México: San Miguel, mpio. Temascaltepec, E. Manrique, A. Ventura 818 (MEXU); Pedregal As Huisihuilango, mpio.

Jaramillo & Núñez 393 (MEXU). Sinaloa: between Mazat- Juchitepec, A. Ventura 1132 (MEXU); al N de Jilotepec,

lán and Durango, between La Fraguita and El Madroño, mpio. Jilotepec, V. Jaramillo, G. Villegas & A. Miranda 901

Durango-Mazatlán, mpio Concordia, Beetle M-3718 (MEXU).

(MEXU). M. watsoniana. Jalisco: Meza de León en el predio de La

M. versicolor. Chiapas: at Escopetazo, mpio. Ixtapa, Breedlove Primavera, mpio. Zapopan, L.M. Villareal 13196 (IBUG);

& G. Davidse 53960 (SLPM). Guerrero: Amojileca, mpio. Cerro El Huiluxtle, mpio. Zapopan, L.M. Villarreal & Pérez

Chilpancingo, U. González 1963 (MEXU). Puebla: carr. de la Rosa 2893B (IBUG). San Luis Potosí: cerro near San

San o desv. a Reynoso, 300 m o Luis, Schaffner 1067 (US).

adentro, mpio. Coatepec, R. Méndez 1268 (MEXU); 14 M. ie Baja California: Yerba Buena, mpio. Ensenada,

ele la an a a L. ARIAN 508 SOM oran & Thorne14236 (TAES). Chihuahua: 13 mi W o

„L. García ducens on hwy 16, mpio. Cuahutémoc, PM. Peterson

BE eie 5 km W of Ixtlan de Juárez, B. Benz 9586 (US).

& K. Benz 694 (IBUG). M. xanthodas. Chiapas: 18 km al NE de Tuxtla Gutiérrez rum-

bo al Cañón del Sumidero, A.J. Zenón 0043 (MEXU).

AGRADECIMIENTOS

Se reconoce y agradece el financiamiento recibido de la Comisión Nacional para el Uso y Manejo de la

Biodiversidad (CONABIO-V024), y del Instituto Politécnico Nacional (SIP-20010344). A las autoridades del

CIIDIR IPN Unidad Durango, les agradecemos el facilitar la tarea que nos permitió llevar a cabo y concluir el

presente estudio. A los curadores de los herbarios consultados, se les agradece sus atenciones. Se agradecen

las observaciones de 3 revisores anónimos y del Dr. Lipscomb, que ayudaron a mejorar el presente trabajo

sustancialmente. En el presente reporte se vierten resultados parciales, generados durante el desarrollo del

proyecto "Biosistemática del género Muhlenbergia de México" SIP-20010344.

REFERENCIAS

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in surface view. Bothalia 12:641—671.

Gómez S., M. v S.D. Koch. 1998. Estudio anatómico comparativo de la lámina foliar de Eragrostis (Poaceae: Chlo-

rideae) de México. Acta Bot. Mex, 43:33-56.

Gómez- SANCHEZ, M. v K.G. TéLuez-PIMIENTA. 2008. Anatomia foliar de algunas gramíneas alpinas y subalpinas del Eje

Volcánico Transversal, México. J. Bot. Res. Inst. Texas 2:495-515.

HERRERA A., Y. 1998. A revision of Muhlenbergia montana (Nutt.) Hitchc. complex. Brittonia Vol. 50:23-50.

HERRERA A., Y. AND W.F. GRANT. 1994. Anatomy of the Muhlenbergia montana (Poaceae) complex, Amer. J. Bot. Vol.

81:1038-1044.

HERRERA A., Y. Y P. M. Peterson. 2006. Muhlenbergia (Poaceae) de Chihuahua, México. Sida, Bot. Misc., No. 29:1-109.

Hrrcucock, A.S. 1951. Manual of the grasses of the United States. 2™ ed. Rev. A. Chase. U.S.D.A. Misc. Publ. no. 200.

Mkrcatre, C.R. 1960. Anatomy of the Monocotyledons. |. Gramineae. longon aa University Press.

MoRpbEN, C.W. AND S.L. HarcH. 1987. Anatomical study of the Muhlenbergia rer plex (Poaceae: Chloridoideae:

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Peterson, PM. 2000. Systematics of the Muhlenbergiinae (Chloridoideae: Eragrostideae). In Grasses: systematics

and evolution, eds. S.W.L. Jacobs and J. Everett. CSIRO, Melburne. Pp. 195-211.

Peterson, P.M. AND Y. Herrera A. 2001. A leaf blade anatomical survey of Muhlenbergia (Poac Muhlenbergiinae)

Sida 19:469-506.

PETERSON, P.M. AND C.R. ANNaaLE. 1991. Systematics of the annual species of Muhlenbergia (Poac Eragrostideae)

Syst. Bot. Mongr. 31:1-109.

PETERSON, PM., C.R. ANNABLE, AND C.R. FRANSESCHI. 1989. Comparative leaf y int! | Muhlenbergia (Poaceae).

Nordic J. Bot. 8:575-583.

Renvoize, S.A. 1981. The subfamily Arundinoideae and its position in relation to a general classification of the

Gramineae. Kew Bull. 36:85-102.

Soperstrom, T.R. 1967. Taxonomic study of subgenus Podosemum and section Epicampes of Muhlenbergia

(Gramineae). Contr. U.S. Natl. Herb. 34:75-189.

SriEgER, M.T. 1982. Revision of Ichnanthus sect. Ichnanthus (Gramineae, Panicoideae). Syst.c Bot. 7:85-115.

Vieira, R.C., M.S. Gomes, L.S. SARAHYBA, AND C.O. ARRUDA. 2002. Leaf anatomy of three herbaceous bamboo species.

Brazilian J. Biol. 62(4B):907-922.

PRIMER REPORTE DE CYSTOPTERIS REEVESIANA (ASPLENIACEAE)

PARA MEXICO

José A. Villarreal-Q. Celestino Flores-L.

eee de oe P aiii. ee ids

Ling raria Antonin Narra { ? Annin Narro

rre 1923, Saltillo 25. 315 Coahuila, MÉXICO Buenavista 1923, Saltillo 25 315 Coahuila, MÉXICO

RESUMEN

Se ci pri pieri iana Lellinger para México. La distribució i ida del ie es del

de los Estados Unidos de América, ánd trabajo q ha colectado d dos del de México

ABSTRACT

z TAI g ; I 15 1 fi : : gi Mexico T1 p : - 4] f. 1 1 r USA ,

but also has b llected in tl 1 Mexico

Durante la colecta de material botánico en el sur de Chihuahua, llamó la atención la presencia de un Cystopteris

diferente al C. fragilis (L.) Bernh., que por ser el mas frecuente, es el mejor conocido. Después de ubicarlo

taxonómicamente se encontró que es C. reevesiana Lellinger. Al parecer, es el primer reporte en el que se da

a conocer la presencia de la especie en Chihuahua.

1 ld her): MÉXICO. Caimuanua. M de Guadal Calvo: Cerro El Mohinora, 25? 5739.14" N, 107?

02’ 1. 59" W, 3142 m, 1 Oct 2007, C. Flores López 439, 460 (ANSM).

La especie crece en un bosque de oyamel con Pseudotsuga menziesii, Abies durangensis, Picea mexicana, Pinus

arizonica, Salix paradoxa, Ribes madrensis, Senecio toluccanus y Erigeron fraternus. Es poco común en el área y

forma colonias esparcidas en lugares con humedad. Cystopteris reevesiana fue descrito en 1981. Forma parte

del complejo de especies relacionadas con C. fragilis. Se caracteriza por presentar frondas 2-3 pinnadas,

con segmentos muy angostos, escamas del rizoma de color café claro y carencia de pelos glandulares en el

estipite.

Cystopteris cido de Arizona, Colorado, Nuevo México, Utah y de Trans-Pecos en Texas,

pero no había sido reportado de Chihuahua en México (Knobloch y Correll 1962; Mickel y Smith 2004).

Sin embargo al investigar si la especie no era conocida de México, encontramos que en algunos herbarios

existen colectas de Sonora (TEX, UC), de Baja California (UC) e incluso de Chihuahua (UC), por lo que la

especie extiende su distribución conocida del suroeste de los Estados Unidos al noroeste de México.

AGRADECIMIENTOS

Agradecemos la ayuda de los doctores J.T. Mickel y A. R. Smith en la identificación del material botánico y

proporcionar información sobre la especie. A Remedios Aguilar por la revisión del escrito.

REFERENCIAS

JoHNsTON M.C. 1990. The vascular plant of Texas: a list, up-dating the Manual of the vascular plants of Texas. 2nd

ed. Published by the author.

KNOBLOCH, IW. AND D.S. Corrett. 1962. Ferns and fern allies of Chihuahua, Mexico. Texas Research Foundation,

Renner.

LUNDELL, C.L. AND COLLABORATORS. 1955. Pteridophyta. In: Flora of Texas 1(1):1-121.

Micke, J.T. AND A.R. Smit. 2004. The Pteridophytes of Mexico. Mem. New York Bot. Gard. 88:1— 1054.

VILLARREAL-O, J.A. 2001. Flora de Coahuila. Listados florísticos de México XXIII. Instituto de Biologia, Universidad

Nacional Autónoma de México. México D.F.

J. Bot. Res. Inst. Texas 3(1): 307. 2009

308 i itute of Texas 3(1)

BOOK NOTICE

WILLIAM HAWTHORNE AND NTIM GYAKARI. 2006. Photoguide for the forest trees of Ghana: a tree-spotter's

field guide for identifying the largest trees. (ISBN 0-85074-164-5, pbk.). Oxford Forestry Institute,

Department of Plant Sciences, South Parks Road, Oxford OX13RB, United Kingdom. (Orders: www.

nhbs.com, +44 (0)1803 865913, +44 (0)1803 865280 fax). $30.00, 432 pp., color photos, b&w figures,

5 3/4" x 7 1/4".

From ub cio dd “A field guid helarg 1 in Ghanaian f It isl 14 p:

1f li il ical This nd ae 320 Species, virtually al ed S foreste canopy anden emer-

gent trees. The photos show La Ent, flowers, bark,

notes on a eee size, bark & slash, leaves, flowers, fruits, and simil Simil i i d and distinguished."

wings of botanical terminology are included on the iets front and batis covers of this book. A common name index,

ind name index, vand a are des e The front cover of the book boasts a grid of tiny color photographs, the very same

eaf, and riptions. But take a step back from this cover du dnd - see Paw ihe smaller images

hat tł ble f far a big eye—presumably the watchful, “tree-spotter.”

i=) 4

J. Bot. Res. inst. Texas 3(1): 308. 2009

FLORISTIC COMPOSITION, ABUNDANCE, AND DISTRIBUTION PATTERN OF

WOODY PLANTS IN A TROPICAL SAVANNA IN NORTHERN GHANA

Alex Asase, Patrick K. Ekpe, and John Y. Amponsah

Department of Botany

University of Ghana

PO. Box LG 55, Legon-Accra, GHANA

aasase@ug.edu.g

ABSTRACT

Tur Š > ae ‘om A jd: -1 : rÍ J 1 : Š 1 Fer I1 i Fo rest Reserve

in northern Giana was studied ae " circular vegetation plos ofi lo m a radius, In tonal, 2534 e of woody plants belonging

to 62 species in 19 families were identified. T} mbretaceae and Rubiaceae.

The abundant woody plants iue Tope cM cinerea (L.) Wright & Arn. (Fabaceae), Poo d Engl. & Diel. (Combreta-

ceae), eae egal Fresen. a xii paa G. Don. e. and diesen jus nus d d

ly I O

showed some degree of randomness. The mus has shown that Bs fone positi and abundance of plants is very vàriable in

different areas of the Guinea Savanna Zone.

Key Wonps: Woody plants; Gui Singsaglebini F R ; Ghana

RESUMEN

La P QE ea abundancia y patrón de distribución de pa leñosas en una sabana tropical en la Sinsabligbini Forest

Reserv se estudió usando 15 parcelas de t ula 10 m de Eo En total se OS 2534

divid de pl len I d 19 familias Las famili

Fabacaeae, Codi iore y Rubiaceae Las plantas lef 1 ] Dichrostachys ci ) Wright & Arn. (Fabaceae),

pa E mida s &D pn a miren ee Fresen. (Combretaceae) paraa ei G. Don. Menu y

s. (Combretaceae) ie i 8

de di il ió d i lon dod Fl 1: } 1 ] val

£ e e [3 Y

y y abusdan d de plantas es muy ae diferentes 4 de | de sabana de Gui

INTRODUCTION

The most extensive vegetation type in Ghana is the guinea savanna. It is estimated to cover about 60.77%

of the total land-mass of Ghana (Anonymous 2002). Previous publicati of studies on the guinea savanna

vegetation in Ghana include that of Vigne (1936), Taylor (1952), Baker (1962), Lawson et al., (1969), Houssain

and Hall (1969), Hopkins (1979), Oteng-Yeboah (1996) and recently Asase and Oteng-Yeboah (2007). Other

studies have examined the effect of fire on the savanna (Ramsay & Rose Innes 1963; Brookman-Amissah

et al. 1980). Despite these contributions, few studies have documented the diversity and abundance of

species of plants found in different areas in the savanna zone of Ghana. This current documentation is of

management i I as it il to a better understanding of the need to conser d sustain ably

utilize plant diversity.

The Sinsabligbini Forest Reserve in northern Ghana is one of the important protected areas in the

savanna zone of Ghana. The reserve was created in 1956 to protect the headwaters of Moya River and its

tributaries. To the best of our knowledge, however, there is no published information on the different species

of plants found in the reserve that could assist those involved in the management of the reserve to develop

appropriate management strategies for the conservation of the vegetation in the reserve.

The objective of the present study was therefore to (i) document the floristic composition, abundance

and distribution pattern of woody plants in the Sinsabligbini Forest Reserve in northern Ghana and (ii)

compare it with other areas in the Guinea Savanna Zone of West Africa particularly Ghana. It is hoped that

this information could assist in the development of strategic management plans for the reserve.

J. Bot. Res. Inst. Texas 3(1): 309 — 316. 2009

310 Journal of the Botanical h Insti Texas 3(

MATERIALS AND METHODS

Study area

The study area at the Sinsabligbini Forest Reserve is situated about 20 km from Tamale, the Northern Re-

gional Capital of Ghana. The reserve is located between latitude 09? 24 138 N and longitude 00? 38 377 W,

and covers an area m 72.72 km?. About 0.2 iol e the reserve area has been converted into a plantation of

Eucalytptus sp., grandis L. and A i Guill & Perr. The natural vegetation in the reserve

area is guinea savanna. The terrain is generally flat an the soil in most areas of the reserve is shallow with

exposed laterite. In seasonally flooded areas of the reserve the soil is mainly made up of clay and silts.

The reserve area experiences one rainy season between May and October each year with a peak period

in August and a dry season from November to March. The average annual rainfall and temperature are 1034.1

mm and 26.7? C, respectively. There are over 10 communities living around the reserve and the people are

mostly from the Dagomba ethnic group. Most of the people living around the reserve are farmers.

Methods

The woody plants in the natural vegetation area of the reserve were studied using vegetation plots. With

reference to Landsat EM 2000 satellite images and topographical maps, we located different habitat areas

in the reserve and randomly demarcated circular vegetation plots of 10 m radius (ca. 314 m?size) in each of

the habitat areas. In total, 15 vegetation plots were examined; at least three plots were demarcated for each

different habitat area. All species of woody plants found rooted within each plot were identified and their

individual plants counted.

The identification of the species of plants encountered was later confirmed using relevant literature

(Arbonnier 2000; Hutchinson & Dalziel 1957-1972), and by comparison with already identified specimens

at the Ghana Herbarium located at the Department of Botany, University of Ghana.

Data analysis

The individual-based rarefaction methodology described by Gotelli and Colwell (2001) was used to estimate

the expected number of species for the construction of species accumulation curve with 9596 confidence

interval. The free statistical software Estimates version 8.0 (Gotelli 2006) was used for the species accumu-

lation curve. The program was set to randomized samples with replacement and shuffle; the individual of

plants among plots within species 1000 times.

The variance: mean ratio of individuals per unit area can be used as a measure of the degree of random-

ness or aggregation in populations or whole communities (Pilou 1977; Sokal & Rolf 1981). Chi-squared test

was used to determine whether the pattern of distribution was significantly random or aggregated based

on the variance: mean ratio calculated for each species (Lambshead & Hodda 1994; Rice & Lambshead

1994

RESULTS

Floristic composition

In total, 2534 individuals of woody plants belonging to 62 species in 48 genera and 19 families were identified

in the reserve. The individual-based species accumulation curve shows that the majority of woody plants

in the reserve were sampled during the study as seen in the curve as it reaches the asymptote (Fig. 1). The

list of species, their families and growth-forms are presented in Table 1. The majority (40.396) of the species

were big trees, followed by small trees (29.096) and shrubs (27.496). Two species namely, Dalbergia afzeliana

G. Don. and Opilia celtidifolia Endl. ex Walp.were the only lianas identified. About 62.996 of the woody plant

genera in the reserve consisted of only one species. Genera with two or more species were Acacia (3 spe-

cies), Annona (2 species), Bridelia (2 species), Combretum (4 species), Gardenia (2 species), Lannea (2 species),

Strychnos (2 species), Terminalia (3 species) and Vitex (2 species).

The most species rich family (29%) was Fabacaeae, followed by Combretaceae (14.5%), and Rubiaceae

(11.3%). Other taxonomic families with many species were Verbenaceae (6.5%), Euphorbiaceae (6.5%), Anac-

ly plant composition in a tropical 311

Asase et al., W

704

Number of species

0 T T T T T

0 500 1000 1500 2000 2500 3000

Number of individuals

Fic. 1. Individual-t | speci las ly pl : ingsaglebini

ardiaceae (4.896) and Annonaceae (4.896) in decreasing order of number of species (Table 2). Ten (52.696)

of the families, namely, Bignoniaceae (Stereospermum kunthianum Cham.), Bombacaceae (Bombax costatum

Pellegr. & Vuillet), Celastraceae (Maytenus senegalensis (Lam.) Exell), Cornaraceae (Rourea coccinea (Schu-

mach. & Thonn.) Hook.f., Moraceae (Ficus sp.), Opiliaceae (Opilia celtidifolia Endl. ex. Walp.), Polygalaceae

(Securidaca longepedunculata Fresen.), Proteaceae (Protea madiensis Oliv), Sapotaceae (Vitellaria paradoxa

C.F. Gaertn.) and Tiliaceae (Grewia venusta Fresen.) were each represented by only one species indicated in

brackets. The families with highest number of individuals were Fabaceae and Combretaceae. These families

contributed about 68.796 of individuals to the woody flora in the reserve. Seven other families (Bombacaceae,

Celastraceae, Moraceae, Opiliaceae, Polygalaceae, Proteaceae and Verbanaceae) contributed less than one

percent of the total number of individuals per family. The families Moraceae and Polygalaceae were least

abundant with each one represented by only one individual plant recorded in the reserve.

Abundance and distribution pattern

The six most abundant woody plants in the reserve were Dichrostachys cinerea (L.) Wight & Arn. (Fabaceae),

Pteleopsis suberosa Engl. & Diels (Combretaceae), Combretum collinum Fresen. (Combretaceae), Dalbergia

afzeliana G. Don. (Fabaceae), Terminalia mollis S Vidal. (Combretaceae), and Vitellaria paradoxa C.F. Gaertn.

(Sapotaceae) in decreasing order of abundance (Table 1). These species contributed about 57.396 of the total

number of individual of plants identified. In contrast the least abundant species were Bridelia micrantha Baill.,

Daniellia oliveri (Rolfe) Hutch. & Dalziel, Ficus sp., Gardenia aqualla Stapf. & Hutch., Parkia biglobosa Jacq.)

R.Br.ex G. Don, Securidaca longepedunculata Fresen., and Vitex chrysocarpa Planch.Eight of the woody plants

in the reserve were frequently encountered, i.e., in 5096 or more of the plots studied. These species were

Combretum mollis S. Vidal, Crossopteryx febrifuga Benth., Vitellaria paradoxa C.F. Gaertn., Strychnos spinosa

Lam., Annona senegalensis Pers., Grewia venusta Fresen., Combretum sp., and Pterocarpus erinaceus Lam. lt

312

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Family Species richness Relative species Abundance ( number Relative

(number of species) richness of individuals) abundance

Anacardiaceae 3 48 32 13

Annonaceae 3 4.8 91 36

Bignoniaceae 1 1.6 29 1,1

Bombacaceae 1 1.6 4 0.16

Celastraceae 1 1.6 3 0.12

Combretaceae 9 14.5 854 33.7

Connaraceae 1 16 116 4.6

Euphorbiaceae 4 6.5 98 3.9

18 29.0 886 35.0

Loganiaceae 2 3.2 76 3.0

Meliaceae 2 32 49 1.9

rac 1 1.6 1 0.04

Opiliaceae 1 1.6 4 0.16

Polygalaceae 1 1.6 1 0.04

Poteac 1 1.6 19 0.75

Rubiaceae 7 11.3 127 5.0

Sapotaceae 1 1.6 84 3:3

Tiliaceae 1 1.6 41 1.6

Verbanaceae 4 6.5 19 0.75

thus follows that the most abundant woody plants in the reserve were not necessarily the most frequently

encountered plants.

The result of the analysis of the general pattern of distribution of the woody plant community in the

reserve shows that the species were highly aggregated (4? 212465.8, df = 930, P = 0). The distribution of the

individual woody plant species was highly discordant with respect to each other (y? = 11746.5, df = 916,

P= 0) whiles individual species were found to be highly aggregated in the plots (y? = 719.3, df =14, P = 0).

The majority (80.696) of the individual woody plant species were found to be aggregated in their pattern of

distribution in the reserve (Table 1). In contrast, twelve of the species, namely, ae eae epee &

Thonn., Bridellia micrantha (Hochst.) Baill. Daniellia oliveria, pos T , Gardenia aq

(Lam.) Exell., Hymenocardia acida Tul., Lannea barteri Engl., Parkia biglol a voa Taub. per

longependuculata, and Vitex ree showed some dene of randomness in the pattern of distribution in

the reserve.

DISCUSSION

Most of the species of plants identified in the Sinsabligbini Forest Reserve in northern Ghana are species

that have been reported in other guinea savanna areas elsewhere in Ghana (Lawson et al. 1969; Brookman-

Amissah et al. 1980; Oteng-Yeboah 1996; Asase & Oteng-Yeboah 2007). However, a few of the species such

as Rourea coccinea (Schumach &. Thonn.) Hook.f., Dalbergia africana, and Hoslundia opposita Vahl that were

identified in the reserve have not been reported in other areas in the Guinea Savanna Zone of Ghana based

on survey of the available literature. Similarly, some of the common woody plant species such as Balanities

aegyptiaca Sands. and Ximenia americana L. that have been reported in other areas of the Guinea ne

Zone of Ghana were not identified in the reserve. This finding has confirmed that the floristi of

the savanna is immensely variable even over relatively homogeneous areas (Hopkins 1979; Lanon 1985).

The high species richness and abundance of taxonomic families such as Fabaceae, Combretaceae,

Rubiaceae, and Anacardiaceae in the guinea savanna has been noted elsewhere (Hopkins 1979; Asase &

Oteng-Yeboah 2007). In a study of three traditional groves in northern Ghana, Oteng-Yeboah (1996) also

sn TT i 315

£T T T

Asase et al : Wi

lpi Papili , and Mimosaceae),

reported on the predominance of members of the Fabaceae (C

Combretaceae, Rubiaceae in the different groves which report is lr to ihe results of this study. It is

also interesting to note that even though some of the families such as Verbenaceae and Anacardiaceae were

found to have several species, their individual plant representation were very low. On the other hand, there

were more individual members of some of the families with lower species numbers such as Connaraceae

and Sapotaceae. It thus follows that not all the families with many species are abundant in the reserve. It

is important to note that some of the important guinea savanna trees such as Afzelia africana Sm., Daniellia

oliveria, and Parkia biglobosa (Hopkins 1979; Lawson 1985) were not abundant in the reserve.

In a study of the pattern of distribution of some woody plants in the Olokemeji Forest Reserve in Nigeria,

Greig-Smith (1991) noted that y could be explained as a response to factors such as soil differences,

pattern of previous farming, intensity of burning from wild fires and at the smallest scale interference be-

tween individual plants possibly due to competition for water. We found evidence of annual wild bush fires

and farming activities in the reserve and these factors might also explain the pattern of distribution of the

woody plants in the reserve. As yet there are no reports on the patterns of distribution of savanna plants in

Ghana and so no comparisons could be made.

To conclude, the present study is the first publication on the species of plants found in the Sinsablig-

bini Forest Reserve in northern Ghana. The study has shown that the characteristics of the vegetation in

the reserve were generally similar to that of other guinea savanna areas. It has also shown that the floristic

composition and abundance of species in different areas of the Guinea Savanna Zone is very variable. Thus

there is a case for the study of the diversity of plants found in the different areas of the Guinea Savanna Zone

of Ghana in order to support the conservation of savanna botanical resources especially for specific areas

of the Guinea Savanna Zone in Ghana.

ACKNOWLEDGMENTS

We are most grateful to th ities livi d the reserve for their hospitality during fieldwork. We

are also thankful to Messer’s Francis Chimsah añd john Baba of the University for Development Studies (UDS)

in Ghana for their assistance in the field. This study was supported by the Northern Savanna Biodiversity

Conservation Project (NSBCP) in Ghana funded by the World Bank. We also thank Alfred Oteng-Yeboah

for a critical review of this paper.

REFERENCES

ANONYMOUS. 2002. National biodiversity strategy for Ghana. Ministry of Environment and Science.

ARBONNIER, M. 2000. Arbres, arbustes et lianes des zones séches d'Afrique de l'Ouest. CIRAD, MNHN, UICN.

ASASE, A. AND A.A. OTENG-YesoaH. 2007. Assessment of plant biodiversity in the Wechiau Community Hippopotamus

Sanctuary in Ghana. J. Bot. Res. Inst. Texas 1(1): 549-556.

Baker, H.G. 1962.The ecological study of vegetation in Ghana. In: J.B. Willis, ed. Agriculture and land use in Ghana.

Oxford. Pp 151-159.

BROOKMAN-AMISSAH, J., J.B. HALL, M.D. Swaine, AND J.Y. ATTAKORAH. 1980. A re-assessment of a fire protection experiment

in north-eastern Ghana savanna. J. Applied Ecol. 17:85-99.

Gomlu, NJ. 2005. EstimateS statistical estimation of species richness and shared species from samples. http://

lu/EstimateS pages/EstimateS.flx.

Gortiu, NJ. AND R.K. CotwetL. 2001.Quantifying biodiversity: procedures and pitfalls in the measurements of spe-

cies richness. Ecol. Letters 4:379-391.

GreiG-Smrrh, P. 1991. Pattern in a derived savanna in Nigeria. J. Trop. Ecol. 7:491-502.

Hopkins, B. 1979. Forest and savanna. Heinemann Educational Books Ltd 2nd Edition.

Houssa, M. AND J.B. HALL. 1969. The tree of Mole National Park, Damongo, Ghana. Revised by Jongking, C.C.H.,

2" edition. University of Ghana.

HUTCHINSON, J. AND J.M. Datzie. 1957-1972. Flora of West Tropical Africa. Crown overseas agent, London.

viceroy eeb.uconn

316 Journal of tl tani Insti Texas 3(

LAMBSHEAD, P.J.D. AND M. Hoppa. 1994. The impact of disturbance on measurements of variability in marine nema-

tode populations. Vie & Milieu 44:21—27.

Lawson, G.W, J. Jenik, AND K.O. AaMsrRoNG-MkNsaH.1969. A study of a vegetation catena in guinea savanna at Mole

Game Reserve (Ghana). J. Ecol. 56:505—522.

OTENG-YEBOAH, A.A. 1996. Biodiversity in three traditional grooves in the Guinea Savanna, Ghana. In: LJ.G. van

der Maesen et al. eds. The biodiversity of African plants. Kluwer Academic Publishers, Dordrecht, London.

Pp. 188-197.

PieLou, E.C. 1977. Mathematicai ecology. 2™ edition. John Wiley and Sons, New York.

Ramsay, J. AND R. Rost Innes. 1963. Some quantitative observations on the effects of fire on the Guinea savanna

vegetation of northern Ghana over a period of eleven years. Sols Africains 8:41-85.

RICE, A. L. AND P.J.D, LAMasHEAD. 1994. Patch dynamics in the deep-sea benthos:the role of a heterogeneous o

oforganic matter. In: PS. Giller, A.G. Hildrew, and D.G. Raffaelli, eds. Aquatic ecology: scale, pattern

34th Symposium of the British Ecological Society. Blackwell Scientific Publications. Oxford. Pp. 469-499

SOKAL, RR. AND FJ. ROHLF. 1981. Biometry.2"* edition. W.H. Freeman and Company, San Francisco.

TavLoR, C.J. 1952. The vegetation zones of the Gold Coast. Bull. Forest Dept. Gold Coast 4:1-12.

VicNE, C. 1936. Forests of the Northern Territories of the Gold Coast. Empire Forest J. 15:210-213.

NEW STATE RECORDS OF VASCULAR PLANTS

FOR THE SOUTHERN ROCKY MOUNTAIN REGION

Pus F. Smith Donald L. Hazlett Neil Snow

Overhill Drive PO. Box 345 Herbarium Pacificum

Fort a. a 80526 U.S.A. Pierce, Colorado 80650 U.S.A. ]

Bishop Museum

1525 Bernice Street

Honolulu, Hawaii 96817 U.S.A.

ABSTRACT

Á +} Lf

recorded for Colorado and N New Mexi ico, whereas Euphorbia extipulata and Trifioltum Wenn: var. anemophilum are ‘documented jor

the first time in Colorado.

RESUMEN

Sei 1 los tada del t da] aea | NA ^ac D Á 7 AA La |

r £ iod r [7] J

nuevamente para Colorado y Nuevo México, mientras que Euphorbia extipulata y Trifiolium dasyphyllum var. anemophilum se documentan

por primera vez en Colorado.

INTRODUCTION

Routine identifications during the past few years of specimens housed in the herbarium at the University

Northern Colorado Herbarium (GREE), where the authors previously worked, revealed new state records

of ae oes for COSTADO and New Mexico, which we report here. All specimens are housed at GREE

ariu ing Holmgren & Holmgren 1998) except for the specimen of Trifolium (at COLO),

and were compared against holdings at GREE, CS, COLO, or RM. Nomenclature and family designations

follow Snow (2009).

AMARANTHACEAE

Amaranthus californicus (Moq.) S. Wats. This species is a glabrous, prostrate, monoecious annual that

occupies seasonally moist flats, shores and other disturbed habitats at elevations between 0—2800 m (Mos-

yakin & Robertson 2003; Welsh et al. 2003). We report it here for the first time from Colorado and New

Mexico; various sources also report the species from Canada, California, Idaho, Kansas, Montana, Nebraska,

Nevada, Oregon, South Dakota, Texas, Utah, Washington and Wyoming.

Voucher specimen: COLORADO. Weld Co.: TOON R65W S20 SE, «0.2 mi N of Weld County Road 102, ca. 6 air mi E of Hwy 85, elev.

ca. 5160 ft [1573 m], infrequent in alkali bed of Little Owl Creek, Pawnee National Grassland, 10 Sep 2004, D.L. Hazlett 12159. NEW

MEXICO. San Miguel Co.: elev. ca. 6000 ft in a dry lake be, 15 Aug 1944, A.L. Hershey s.n.

EUPHORBIACEAE

Euphorbia exstipulata Engelm. In the Southern Rockies this annual species was collected only once previ-

ously, in 1894, in Platte Co. Wyoming (Dorn 2001). It is noted as likely extirpated for the State of Wyoming

(Kartesz 2009). An additional collection near our area is known from Cimarron County, Oklahoma, but the

species otherwise occurs in the USA in UT, AZ, NM, TX, and CA (Kartesz 2009).

Voucher specimen: COLORADO. Baca Co.: Ravine 10 mi SW of Campo, May-Sep 1977-1978, W.E. Harmon 9507.

FABACEAE

Trifolium dasyphyllum Torr. & A. Gray var. anemophilum (Greene) J.S. Martin ex Isley. This native

perennial previously has been known only from Wyoming. The closest population occurs in the Laramie

Range in eastern Albany County, Wyoming (Isley 1980).

J. Bot. Res. Inst. Texas 3(1): 317 — 318, 2009

318 Journal of t tanical i Texas 3(

inf G 1 a Ni facina

Voucher specimen: COLORADO. Weld Co.: TION R60W S4, 7 mi E of Grover, elev. ca. 1646 m, ] facing

rocky ridge (private land), 04 Jun 1999, D.L. Hazlett 10924 (COLO).

ACKNOWLEDGMENTS

Support for curatorial work of Southern Rocky Mountain taxa at GREE was supported by NSF DEB-0237 149.

Thanks to an anonymous reviewer, Ronald Hartman, and Ben Legler for reviewing the manuscript, and to

John Kartesz for providing advance copies of Version 2 of the Floristic Synthesis.

REFERENCES

Dorn, R.D. 2001. Vascular plants of Wyoming. Third edition. Mountain West Publishing, Cheyenne.

HOLMGREN, PK. ano N.H. HOLMGREN. 1998 [continuously updated]. Index Herbariorum: A global directory of public

herbaria and associated staff. New York Botanical Garden's Virtual Herbarium. (http://sweetgum.nybg.org/ih/).

Istey, D. 1980. New combinations and one new variety in Trifolium ponies) Brittonia 32:55-57.

KARTESZ, J.T. 2009. A synonymized checklist and atlas with biol tes for the vascular flora of the United

States, Canada, and Greenland. Second edition. In: Kartesz, JT. Floristic synthesis of North America, Version

1.0 (in press).

Mosvaxin, S.L. AND K.R. ROBERTSON. 2003. Amaranthus L. In: Flora of North America north of Mexico. Vol. 4, Magno-

liophyta: Caryophyllidae, part 1. Oxford University Press, New York. Pp. 410-435.

Snow, N. 2009. Checklist of vascular plants of the southern Rocky Mountain Region. Version 3. (http://www.

botanicus.org/title/b1334416x) or (http:// ps.org/plant, lists keys.html).

Wers, S.L, N.D. ArwooD, S. GOODRICH, AND L.C. Hicains. 2003. A Utah flora. Third edition. Brigham Young University,

Provo.

PYRACANTHA KOIDZUMII (ROSACEAE) NEW TO THE ARKANSAS FLORA

Brett E. Serviss

Departmentof Biology

Henderson State University

ida: idus 71999-0001, U.S.A.

b@hs

rvisb@hsu.edu

ABSTRACT

Pyracantha koidzumii (Hayata) Rehder (Formosa eee) is here reported as new to the Arkansas flora. Additionally, it is also m

ably the e record a the genus Pyracantha E ST p in pu state. A key and illustrations of th y

RESUMEN

Pyracantha koidzumii (Hayata) Rehder eee ara) is here iUd as new to the Arkansas flora. Additionally, it is also prob-

ably the s cda of the genus Pyracantha g g SI y in put state. A key and illustrations of the three most commonly

teid

f Iti ti in rl TIG

INTRODUCTION

At present, non-native species comprise approximately 21% of the Arkansas flora (Arkansas Vascular Flora

Committee 2006), with new species of non-native plants continuously being documented and added (Peck

2003; Peck & Serviss 2006; Serviss et al. 2006, 2008a, 2008b; Serviss & Peck 2008). Many of the species

comprising these most recent records were well established when initially encountered; therefore, it is ex-

tremely important to record first encounters with escaped populations and monitor these new spontaneous

occurrences of non-native species to evaluate their potential for becoming the next wave of invasive species

(Yatskievych & Raveill 2001).

Pyracantha is a Eurasian genus consisting of about 10 species of woody, evergreen, thorny shrubs and

trees (Bailey 1971; Flora of China Editorial Committee 2003; Krússmann 1978). Species of Pyracantha are

important as ornamentals because of their evergreen habit, and bright red, Does or yellow-colored fruits

that persist well into winter. Several species of Pyracantha have become estal ious degrees across

much of the southern US from California to Florida and the Carolinas (NRCS 1999. but until now, have

not been recorded outside of cultivation in Arkansas.

Pyracantha coccinea (D. Don) MJ. Roem. (scarlet firethorn) was previously documented for Arkansas

(Arkansas Vascular Flora Committee 2006). However, this record now appears to be of questionable status

regarding its position in the flora, as further investigation has led to the possibility that the specimen cited

in the checklist may have simply B persisting from cultivation; and not actually spontaneous. As a result,

it will apparently not be included in the up las for the vascular flora of Arkansas (Brent Baker, pers.

comm.). Additionally, a second possible record oli Pyracantha in the state collected from Lafayette County in

1959 by Demaree (41944; UMO), was examined by the author and determined to be Crataegus (hawthorn).

Subsequently, the record of P. koidzumii presented here is currently the only definitive record of this genus

occurring outside of cultivation in Arkansas.

SPECIMEN RECORD OF P. KOIDZUMII FOR ARKANSAS

Pyracantha koidzumii (Hayata) Rehder (P. formosana Kanehira) (Rosaceae). Formosa firethorn is a large,

evergreen shrub or small tree to about 4(5.5) m in height that is native to Taiwan. In addition to Arkansas,

Formosa firethorn has been documented outside of cultivation in the US in Alabama, Arizona, Florida,

Georgia, and South Carolina, as determined from the national flora database kept by NRCS (1999).

J. Bot. Res. Inst. Texas 3(1): 319 — 321. 2009

3200 doma t tani i Texas 3(

Voucher specimen: Garland Co.: one EC b ides age plant Pon fruits present), steep bluff with rocky soil, semi-

, Quercus velutina, Nyssa sylvatica, Pinus echinata, Pinus

disturbed, upland woods habitat, ]

taeda, Prunus serotina, Ulmus alata, and tofP ll ca ore dat Black N of Central Avenue, 4500 block, Hot Springs,

1 Nov 2008, B. Serviss and A. Serviss 7396 (HEND).

Many species of Pyracantha are difficult to distinguish without careful scrutiny; thus the following key has

been provided as an aid to identification. In addition to the approximately 10 currently recognized species

in the genus, numerous interspecific hybrids and cultivars also occur. No attempt to treat these additional

taxa is made here (for a list and descriptions of many of the hybrids and cultivars of Pyracantha, see Krúss-

mann 1978).

KEY TO PYRACANTHA SPECIES LIKELY ENCOUNTERED IN ARKANSAS

At least three species of Pyracantha: P. coccinea, P. fortuneana (P. crenatoserrata), and P. koidzumii, (Fig. 1)

are cultivated in the state, and spontaneous seedlings are sometimes observed in the vicinity of cultivated

plants (Serviss 2006).

1. Leaves usually elipt sometimes narrowly so, apex acute, margins toothed P. coccinea

1. Leaves usually oblanceolate, or obovate-oblong, apex obtuse, rounded, or nearly t te, sometimes

emarginate, gi o or entire (leaves of P koidzumii can be elliptic, but "m entire margins).

2. Leaf margins on most or all leaves PEREN toothed, coe: ge widest lowe apex P. fortuneana

2. Leaf margins entire leaves may have a few widely si i g widest more toward

the middle P. koidzumii

Pyracantha can sometimes be confused with certain species of Crataegus, but differs from it by its evergreen

habit, leafy thorns, and two fertile ovules per carpel. Crataegus species are deciduous with leafless thorns

and only one fertile ovule per carpel. The generally narrow, unlobed, finely-toothed leaves of Pyracantha

can also be useful in distinguishing it from Crataegus.

ACKNOWLEDGMENTS

I would like to sincerely thank my daughter, Annabelle Serviss, who initially discovered and brought to

my attention the P. koidzumii plant, Brent Baker (University of Arkansas) for providing information regard-

ing the current status of Pyracantha in Arkansas, Robin Kennedy (University of Missouri at Columbia) for

providing a specimen loan, Guy Nesom (University of Texas at Austin), Victor Maddox (Mississippi State

University), and one anonymous reviewer for their helpful comments and suggestions regarding this paper,

and the Henderson State University Biology Department for supporting this research.

Serviss, Pyracantha koidzumii in Arkansas 321

REFERENCES

ARKANSAS VASCULAR FLORA COMMITTEE. 2006. Checklist of the vascular plants of Arkansas. University Herbarium, Depart-

ment of Biology, University cf Arkansas, Fayetteville.

Batey, L.H. 1949. Manual of cultivated plants. MacMillan. New York, NY.

Baker, B. (Brent). Personal communication. Botanist, University of Arkansas Herbarium, Fayetteville.

FLORA OF CHINA EDITORIAL COMMITTEE, 2003. Flora of China. Vol. 9 (Pittosporaceae through Connaraceae). Beijing (China):

Science Press, and St. Louis, MO: Missouri Botanical Garden Press. p. 108-110.

KRÜSSMANN, G. 1978. Manual of cultivated broad-leaved trees and shrubs. Vol. 3. Timber Press. Portland, OR.

Peck, J.H. 2003. Additions, re-instatements, exclusions, and re-exclusions to the Arkansas flora. Sida 21:

1737-1757.

Peck, J.H. AND B.E. Serviss, 2006. New and noteworthy collections for Arkansas. Sida 22:817-820.

Serviss, B.E. AND J.H. Peck. 2008. New and noteworthy records of several non-native vascular plant species in

Arkansas. J. Bot. Res. Inst. Texas 2:637-641.

Serviss, B.E., N. FREEMAN, J. HERNANDEZ, A. Legie, and C. TaLLey. 2008a. Tungoil tree (Aleurites fordii Hemsl.) (Euphorbi-

aceae) new to the Arkansas flora. J. Arkansas Acad. Sci. 61:128-130.

Serviss, B.E., N. FREEMAN, J. HERNANDEZ, A. LelBLE, C. TALLEY, AND B. BAKER. 2008b. Negundo chaste tree (Vitex negundo L.)

(Verbenaceae) new to the Arkansas flora. J. Arkansas Acad. Sci. 61:131-133.

Serviss, B.E. 2006. Non-native woody plants of Arkansas. Available at http://www.hsu.edu/servisb. Accessed on

14 November 2008,

Serviss, B.E., N. FREEMAN, AND S. MELANCEN. 2006. Chinese flame tree (Koelreuteria bipinnata Franch.) (Sapindaceae)

new to the Arkansas flora. J. Arkansas Acad. Sci. 60:197-199.

USDA, NRCS. 1999. The PLANTS database. National Plant Data Center, Baton Rouge, LA 70874-4490 USA. Avail-

able at http://plants.usda.gov. Accessed on 14 November 2008.

YATSKIEVYCH, G. AND J.A. RAVEILL. 2001. Notes on the increasing proportion of non-native angiosperms in the Missouri

flora, with reports of three new genera for the state. Sida 19:701—709.

322 J lofti

BOOK NOTICE

HIDEAKI OHBA, Yu IOKAWA, AND LOKENDRA Raj SHARMA (eds.). 2008. Flora of Mustang, Nepal. (ISBN 978-

4-906464-15-9, hbk.). Kodansha Scientific Ltd., 9-25 Shin-ogawa-cho, Shinjuku-ku, Tokyo, Japan

(Orders: ww co.uk). $106.00, 483 pp., index, color plates, b&w figures and line drawings,

7" x 10 1/4".

From the preface: “The Nepal Himalaya is of i i | ists due to its rich biodiversity. All global biocli

1 Mex th J f NT :

MA. A * rf

The g reg y

m 41 Ri La i

g vegetation are Tibetan in character.”

and high degree of

J Lr e r

floral endemism ... It is a nearly

Contents:

Foreword

Preface

List of Contribut

PRU of Color Eme ip a nae a aod M e type p 4 individual plant photos, 60 separate photos of

History of Botanical porto in Mustang

Vegetation of Must

Climatic Duk s Mustang

ipid Synopsis and Plant dd

Systematic List of the Collec

PTERIDOPH

LAO Rib

JR Parkeriaceae, Aspleniaceae, Dryopteridaceae, Woodsiaceae

SPERMATOPHYTA-GYMNCSPERMA

Ephedraceae, Pinaceae, ah era

DON

EAE-ARCHICHLAMYDEAE

Salicaceae, Betulaceae, Urticaceae, Santalaceae, Polygonaceae, Nyctaginaceae, Carypohyllaceae, Chenopodiaceae, Ranunculaceae,

Circaeasteraceae, Berberidaceae, Papaveraceae, Cruciferae, Crassulaceae, Saxifragaceae, Hydrangeaceae, Rosaceae, Leguminosae,

Geraniaceae, Zygophyllaceae, Linaceae, Euphorbiaceae, Polygalaceae, Balsaminaceae, Thamnaceae, Malvaceae, Thymelaeaceae,

Elaeagnaceae, Violaceae, Tamaricaceae, Onagraceae, Umbelliferae

SPERMATOPH

YTA-DICOTYLEDONEAE-SYMPETALA

Ericaceae, Primulaceae, Oleaceae, Gentianaceae, Asclepiadaceae, Rubiaceae, Convolvulaceae, Boraginaceae, Labiatae Solanaceae,

Scrophulariaceae, Bignoniaceae, Orobanchaceae, Lentibulariaceae, Plantaginaceae, Caprifoliaceae, Dipsacaceae, Campanulaceae

Compositae

SPERMATOPHYTA-MONOCOTYLEDONEAE

Juncaginaceae, P

, Liliaceae, Iridaceae, Juncaceae, Graminae, Araceae, Cyperaceae, Orchidaceae

New Names and Combinations Appearing i in Fl M g Nepal (2008)

Index to Botanical Names

J. Bot. Res. Inst. Texas 3(1): 322. 2009

MEDICAGO RIGIDULA AND M. TRUNCATULA (FABACEAE):

NEW TO THE CALIFORNIA FLORA

Richard E. Riefner, Jr. Ernest Small

Research Associate . Principal Research Scientist

Rancho Santa Ana Botanic Garden Agriculture and Agri-Food Canada

500 North College Avenue Central Experimental Farm

Claremont, California 91711-3157, U.S.A. Ottawa, Ontario, K1A 0C6, CANADA

rriefnergearthlink.net smalle@agr.gc.ca

ABSTRACT

Medicago rigidula and M. truncatula are reported for the first time for California. These species have likely become established from

rangeland, forage mnes cover e grown over the s Un They can be confused with the relatively common M. polymorpha, and

hahl

are probdabdly more $

Key Wonps: biological i i grasslands, Medicago rigidula, Medicago truncatula, medics, nonnative plants

RESUMEN

Medicago rigidula y M. —— se ee de eee vez en pena Probabl t i han establecido a través

de los pastizales de forraje y | siglo pasado. Medicago rip dila y M. truncatula pueden ser

confundidas con la especie relativamente común M. polymorpha, y probablemente están más generalizadas que lo indicado por los

registros actuales.

Annual species of Medicago, commonly known as medics, are indigenous to the Mediterranean region, and

some of these have become established in temperate and Mediterranean-type climate regions around the

world (Heyn 1963; Piano & Francis 1992). In this paper, we provide the first documented records of Medicago

rigidula (L.) All. and M. truncatula Gaertn. for California.

Medicago rigidula (Tifton medic, Tifton burclover, Tifton bur medic, rigid medic) and M. truncatula

(barrel medic, barrel clover) have not been reported previously in major publications addressing nonnative

species in California (Isely 1993; Bossard et al. 2000; Hrusa et al. 2002; Bossard & Randall 2007; DiTomaso

& Healy 2007; Jepson Flora Project 2008; USDA 2008a, b). They have oe not been included in treatments

of the Fabaceae in county floras or in recent local floristic studi tal southern California (Boyd

2001; Schneider-Ljubenkov & Ross 2001; Bowler & Bramlet 2002; Naher 2002; Bowler & Elvin 2003; Pyke

et al. 2003; Roberts et al. 2004; Rebman & Simpson 2006; Clarke et al. 2007; Roberts & Bramlet 2007;

Roberts et al. 2007; Roberts 2008).

Voucher specimens:

Medicago rigidula (Fabaceae, Trifolieae, Trigonellinae), U.S.A. CALIFORNIA. Orange Co.: City of San Juan Capistrano, San Juan

Creek Rd. at San Juan Creek Circle, UTM (NAD 83) 115 0439671E 3706917N, elev. ca. 24 m, uncommon in annual grassland, 18 Mar

2007, Riefner 07-150 (RSA); ee of ees iue ndn Niguel Regional Park, general vicinit La Paz Rd. at Aliso Creek, UTM (NAD

83) 115 0434480E 3712053N, ca. 70m mmon, disturbed roadside, 1 Apr 2007, Riefner 07-165 (DAO, RSA); City of San Juan

o San Juan Creek area, ca. 0.2 mi W a intersection of Paseo Tirador and Calle Arroyo St., UTM (NAD 83) 11S 0439010E

660N, elev. ca. 31 m, field, 1 May 2008, Riefner 08-73 (DAO, RSA); Ladera Ranch, Cecil Pasture at Jerome Rd.,

UTM (NAD 83) 115 0439962E 3713383N, dex. ca. 157 m, uncommon dioi. Bluff Top Trail in annual grassland, 12 May 2008, Riefner 08-75

(DAO); City of San Juan Capistrano, 0.2 Pkwy. on Hwy. 74 (Ortega iuis UTM M 29) 115 p de 3709137N, elev.

ca. 88m grassland and di bed i 19 May 2008, Riefner 0

the Cities of Lake F 1 Rancho S M ita, El Tor Rd ., UTM (NAD 83) 118 0442282E 3725738N, elev.

ca. 362 m, ] fire break with ruderal vegetation, 22 ins 2008, Ricfner 08 08- "120 0 (RSA).

Medicago truncatula (Fabaceae, Sdn Trigonellinae), U.S.A. CALIFORNIA Orange Co.: City of Laguna Niguel, Laguna Niguel

Regional Park, general vicinity of La Paz Rd. at Aliso Creek, UTM (NAD 83) 115 0434480E 3712053N, elev. ca. 70 m, uncommon in

Atriplex scrub, 1 Apr 2007, Riefner 07- 166 (DAO, RSA); City of Laguna aps Laguna pid dS HA iudi dia iind of La Paz

Rd. at Aliso Creek, UTM (NAD 83) 115 0434316E 3712082N, elev. ca. 63 m, uncommon in ] t

J. Bot Res. Inst. Texas 3(1): 323 - 329. 2009

| af al D H In LJ die £ T.

324 J t Texas 3(

reddish, diamond-shaped marl f each leaflet lamina), 1 Apr 2007, Riefner 07-168 (DAO); City of Orange, SW portion of Irvine

Regional Park, Irvine Park Rd., UTM (NAD 83) 11S 0431403E 373509N, elev. ca. 188 m, uncommon in disturbed and ruderal habitats,

12 Apr 2007, Riefner 07-178 (DAO, RSA); City of Rancho Santa Margarita, Tijeras Canyon, N ca. 0. - mi from oom Trails Rd. on

Antonio Pwky., UTM (NAD 83) 115 AL dd elev. ca. 221 m, d, 24 March 2008, Riefner 08-64

(RSA); City of San Juan Capistrano, 0.2 wy. 74 (Ortega Hwy.), UTM (NAD 83) 115 E 3709137N, elev.

ca. 88 m, uncommon, 1 grassland, 19 M 2008, oe 08-92 (DAO, RSA).

Medicago rigidula sensu lato is native to Eurasia and North Africa (Small & Jomphe 1989; Small et al. 1990).

The European and African populations have been segregated as M. rigiduloides E. Small, based particularly

on pollen morphology and fruit characters (Small 1990; Small et al. 1990). Additional studies confirming

separation of these taxa are needed (Heft & Groose 1996), so we have not attempted to identify the California

collections with respect to these groups. Medicago truncatula is also indigenous to Eurasia and North Africa

(Small & Jomphe 1989; Small et al. 1990; Small et al. 1991). With the exception of M. polymorpha L., it is

the most common weedy annual Medicago in the Old World (Small & Jomphe 1989).

Medicago rigidula and M. truncatula grow on a variety of soils, ranging in texture from sandy loams to

clay, and the former is adapted to rocky soils; both species are best adapted to neutral to somewhat alkaline

(pH 6 to 9) conditions (Small & Jomphe 1989; Nair et al. 2006; Frame 2008). The annual medics are not

cold hardy and will die after a killing frost (Quinlivan et al. 1986). However, M. rigidula is relatively cold

tolerant (Walsh et al. 2001; Small & Jomphe 1989; Krall et al. 1996). Although not widely recognized as

a halophyte, M. truncatula performed very well in an experimental study of cover crops suitable for saline

soils of California’s Great Central Valley (Mitchell 1996).

Medicago species are among the legumes that are highly selective of rhizobial bacteria, which have im-

portant nitrogen fixing properties (Allen & Allen 1981). The annual species of Medicago are highly adapted

to a wide range of environments and to new locations (Crawford et al. 1989). Accordingly, many annual

medics now play an important agronomic role in dryland farming around the world (Walsh et al. 2001).

The annual medics comprise the principal legume component of pasture lands on more than 20 million ha

in Southern Australia where they are utilized as self-seeding annuals to improve soil structure, increase soil

nitrogen, and provide forage for livestock (Cocks et al. 1980; Crawford et al. 1989; Squires & Tow 1991).

Medicago truncatula is a highly valued and widely cultivated species in Australia and other Mediterranean

climate regions around the world (Crawford et al. 1989; Walsh et al. 2001; Nair et al. 2006). There are more

cultivars of M. truncatula than of any other Medicago except M. sativa L. (alfalfa). Medicago truncatula is also

being developed as a model legume plant in both classical and molecular genetic studies to a the

functions of its genes (e.g., bacterial and fungal symbiosis, stress resistance, and plant archit ) and to

exploit its genome (e.g., improved seed quality and production of specific secondary metabolites); for recent

reviews see Thoquet et al. (2002) and Watson et al. (2003). By contrast, M. rigidula has attracted relatively

limited interest as an agricultural crop, and there are few cultivars.

In coastal southern California, ranchers have long prized burr clover (M. polymorpha) as valuable forage

(U.S. Coast & Geodetic Survey 1891). Several other annual medics, including M. arabica (L.) Huds., M. lupu-

lina L., M. rigidula, M. scutellata Mill., M. truncatula, and M. turbinata (L.) All. were tested in row nurseries or

broadcast plots in order to improve forage on California rangelands, including four sites in Orange County,

southern California (Jones & Love 1945). These species did not perform well during the early field tests,

and were not recommended for use on rangelands in the South bn pur (Jones E Love n However,

following t! ful breedi d development of tal genotypes, many

annual species of Medicago, including M. rigidula and M. truncatula, have been a re- a for

use and/or are currently utilized as cover crops or forage in the United States (Allen & Allen 1981; Zhu et

al. 1996; Shrestha et al. 1998; Fisk et al. 2001; Walsh et al. 2001; Krall et al. 2007). In California, the annual

medics are favored as cover crops in no-till orchards and vineyards, especially in the Great Central Valley

(Miller et al. 1989; UC SAREP 2008). Despite their widespread use, M. rigidula and M. truncatula have been

rarely reported growing outside of cultivation in the United States (Kartesz & Meacham 2005; Jepson Flora

Project 2008; USDA 2008a, b).

n:.£ ne li, T AM Ad: [ : £, Calif 1 325

There ar 1 reasons why the annual lics should be expected asi ] ] weeds in California.

All of the den or so cultivated medics are weeds in the Old World, and have the potential to be weedy

elsewhere. The cultivars that are often grown experimentally or for their proven value in California are

scarcely different from their wild progenitors. In addition, essentially wild plants are also often cultivated.

Accordingly, the annual medics have not been weakened by domestication.

Also, the majority of the annual species of Medicago have spiny pods adapted to adherence to fur and

feathers, which ily distributed by wild animals, and also by humans (for example, in wool). Their seeds

are well protected in indehiscent, spiny pods, and are long lived. A seed of M. polymorpha extracted from

adobe brick from Mexico, estimated to be of the order of 200 years of age, successfully germinated (Spira

& Wagner 1983). Therefore, it would not be surprising to find annual medics naturalized in California.

Southern Orange County supports ideal conditions that have facilitated the introduction and natu-

ralization of annual medics, which include: a Mediterranean-type climate; widespread neutral to alkaline

sandy loam to clay soils; extensive historic dd and supporting a grazing, dryland row crops, orchards,

and equestrian uses; and expanding resid ial lan I that has split up the historic

ranchos, increased disturbance, and facilitated invasions of new a species in the region (Wachtell 1978;

Hallan-Gibson et al. 2005; Orange County Historical Society 2005; Riefner & Boyd 2007). Notably, access

to historic grazing habitats in southern Orange County is now possible owing to lands dedicated to open

space and conservation, most of which have been poorly studied (Roberts & Bramlet 2007).

Some medic cultivars are characterized by single, reddish, adaxial leaflet marks (Nair et al. 2006), al-

though this phenomenon is relatively rare among most wild medic species. Some of the plants collected in

Orange County (Riefner 07-168, DAO) have such marks, which may reflect origin from agricultural cultivars.

Many of the medic cultivars have been selected for resistance to aphids and root-lesion nematodes, and adap-

tations to specific soil textures (Nair et al. ido Early introductions of M. rigidula and M. truncatula failed

to establish, perhaps because of lack of suitabl tations, but continued releases of cultivated strains may

have led to bd cañon and the production of ponia populations, aiding naturalization in southern

California. The introduction of cultivars has been hypothesized to contribute new genetic diversity that

could enhance or aid the establishment of nonnative plants (Cox 2004).

In Eurasia, it is extremely common for several species of annual Medicago to grow together as weeds.

In Orange County, M. rigidula and M. truncatula are closely associated with M. polymorpha, which is the

most commonly collected medic in southern California and the most likely to be confused with either M.

rigidula or M. truncatula. Medicago rigidula and M. truncatula are members of section Spirocarpos Ser. subsec-

tion Pachyspirae (Urb.) Heyn, which is characterized by pods that at maturity are extremely hard, and have

alveolar (appearing spongy) tissue on the coil faces (and at the base of spines) that often obscures the coil

venation (Small & Jomphe 1989). Medicago rigidula pods are usually covered with velvety-glandular hairs

(observable in young pods), are discoid, cylindrical, ovoid or spherical in shape, often have evident inter-coil

gaps on mature pods, and have radial veins EN are si curved with limited anastomosing (Heyn 1963;

Small & Jomphe 1989; Bena et al. 1998). Medica, go tula has cylindrical pods, usually with a few simple

trichomes (often on the spines), long curved spines that frequently point towards the ends of the pod, and

weakly to moderately curving radial veins on the coil faces, with limited anastomosing (Heyn 1963; Small

& Jomphe 1989; Bena et al. 1998). Occasionally, plants of M. truncatula and M. rigidula have pods without

spines, and both species have moderately pubescent herbage.

Medicago polymorpha, of section Spirocarpos (= subsection Leptospirae (Urb.) Heyn), is generally glabrous

to lightly hispid, and the pods, spiny or me merely with tubercles, generally have

venation on the coil faces. However, M. polymorpha frequently mimics Spes of Pachyspirae in developing

very hard pods with proliferating tissue on the fruit coil faces, making id li t (Small @ Jomphe

1989). This mimicry does not occur in young pods, which should be examined for positive identification.

Typical pods of M. rigidula, M. truncatula, and M. polymorpha, and venation on coil faces are depicted in

Figure 1.

retic ulate

326 tani i Texas 3(

Given the difficulty of discriminating M. rigidula and M. truncatula from the widespread M. polymor-

pha, it may be that the species reported here as new to California could be established in other agricultural

regions of the State, including the Great Central Valley and the Imperial Valley, where there are extensive,

suitable habitats.

Note.—After the manuscript was accepted for publication, we identified the two voucher specimens of

“Medicago muricata All” cited in Dean et al. (2008) (collections made by G. Butterworth in 2005 from San

Luis Obispo Co.) as M. truncatula Gaertn.

ACKNOWLEDGMENTS

We greatly appreciate the efforts of Harvey Brenneise and Irene Holiman (Library of Rancho Santa Ana

Botanic Garden) for assistance with document retrieval. George W. Cox (San Diego State University) and

an anonymous reviewer provided helpful comments that greatly improved the manuscript. We also thank

Fred Hrusa (California Department of Food and Agriculture, CDA) for a loan of specimens.

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330

BOOK NOTICE

DoucLas J. Furuyma, H. BRADLEY SHAFFER, AND DANIEL SIMBERLOFF (eds.). 2008. Annual Review of Ecology,

Evolution, and Systematics, Volume 39. (ISSN 1543-592X; ISBN 978-0-8243-1439-2, hbk). Annual

Reviews, Inc., 4139 El Camino Way, PO. Box 10139, Palo Alto, California 94303-0139, U.S.A. (Orders:

www.AnnualReviews.org, science&annualreviews.org, 800-523-8635, 650-493-4400). $84.00 indiv.,

674 pp., 7 5/8" x 9 3/8",

Contents of Volume 39:

1. Top Predators as Conservation Tools: Ecological E e a OS Sergio, Tim Caro, Danielle Brown,

Barbara d. oa Hunter, James Ketchum, Kath g

Evolution: From Populati ic Marl Dri f Adaptive Shif d Speciation?—

Ee Honmann: torent bid:

3. Radial Symmetry, t ior Axis, and Echinoderm Hox Genes— Rich Mooi, Bruno Dav

4. The Great American Schi fM O i After the Rise of the Central American Isthmus—H.A. Lessios

5. The Ecological Performance of Pieced TE J. Gaston, Sarah F. Jackson, Lisette Cantú-Salazar, Gabriela Cruz-Piñón

6. Morphological Integration and Developmental Modularity—Christian Peter Klingenberg

7. Herbivory from Individuals to Ecosystems—Oswald J. Schmitz

8. Stoichiometry and Nutrition of Plant Growth in Natural Communities—Géran I. PTT

9. Nous sd or MOSS Fish? - Review of ind Potopi 1 Impacts of Int I Graham H. Pyke

hil 1 Arabidopsis— Stephen I. Wright, Peter Andolfatto

Stability of Plant-Rhi here M li E. Toby Kiers, R. Ford Denison

11. EOS aoe and tl D

F N d Consequences—Fernando Valladares, Ulo Niinemets

12. Shade Tolerance, a Key Pl

13. The Impacts of Fisheries on Marine Ecosystems and the Transition to Ecosystem-Based Management—Larry B. Crowder, Elliott L.

ead eile ANSA dA Tone Catherine Latanich, Matthew B. Ogburn

14. g Act—Mark W. Schwartz

15. Phylogenetic Apr } he Study iof Extinction—Andy Purvis

16. ti M inal Habitats—Tadeusz J Kawecki

17. Gon pecile Brood! Parasitism in Birds: A Life-History Perspective—Bruce E. Lyon, John McA. Eadie

18. es d d. Temporal Data and Character Data in Phylogenetic Inference— Daniel C. Fisher

MD indies ian T Ender

mission— Michael E Antolin

20. nee cking B: Within-Hos of 1

21. Evolutionary cd of f and Their Associates: Recon Gage ind OUR Puzzles—Edward Allen Herre, K. Charlotte

Carl

andé

22. The Earliest Land om ipis G. Gensel

23. Spatial D fF Priyanga Amarasekare

24. Species Selections Theory and Data—David Jablo

25. New Answers for Old Questions: The a ice ane Genetics of Wild Animal Populations—Loeske E.B. Kruuk, Jon Slate,

Alastair J. Wilson

26. Wake Up and Smell the Roses: The Ecology and Evolution of Floral Scent—Robert A. Raguso

27. Ever Since Owen: Changing dedu on the Early Evolution of Tetrapods— Michael I. Coates, Marcello Ruta, Matt Friedman

28. Pandora's Box Contained Bait: The Global Problem of Introduced Earthworms— Paul F. Hendrix, Mac A. Callaham Jr., John M. Drake,

Ching-Yu Huang, Sam W. we lames, Bruce A. enya Weixin Zhang

29. Trait-Based C Ecology of Phy Elena ciel Christopher A. Klausmeier

30. What Limits Trees in C4 Grasslands and Savanmas? Wihan]. Bon

Cumulative Index of Contributing Authors, Volumes 35-39

Cumulative Index of Chapter Titles, Volumes 35-39

J. Bot. Res. Inst. Texas 3(1): 330. 2009

NEW AND NOTEWORTHY PLANTS FROM FLORIDA

John M. Kunzer! and Richard P. Wunderlin Loran C. Anderson

Department of Cell Biology Department of Biological Science

Microbiology,and Molecular Biology Florida State University

University of South Florida, Tallahassee, Florida 32306-4370, U.S.A.

4202 East Fowler Ave, SCA 110

Tampa, Florida 33620-5200, U.S.A.

James R. Burkhalter

788 Deedra Avenue

Pensacola, FL 32514, U.S.A.

ABSTRACT

T J e TT om a AE ed ] +1 ful ON | A H z * 1 asiatica Mol TE Afl Ochna

serrulata, and Passiflora vitifolia. The reports for the non-native Lami Sida urens, and Sp! teri i, and the native

Crataegus opaca, Cyperus bipartitus, Cyperus eragrostis, and Geranium paoli are the first for Florida Here fun occurrence of s

arvensis in Florida is verified (for a total of 12 new records for Florida), and 37 ji g sig I

RESUMEN

Se citan cuatro taxa como nuevos ps los Estados neni en la Led pum di el estado de Florida: Azolla pinnata subsp. asiatica

Melochia nodiflora, Ochna serrulata, ifl itif d Si s, y Sphaeropteris cooperi,

y las nativas Crataegus opaca, Cyperus part iim eragrostis, y Geranium aula son las primeras para Florida. Se verifica

aquí la presencia de Torilis arvensis in Florida (con un total de 12 nuevas citas para Florida), y se citan 37 taxa que representan colectas

importantes.

INTRODUCTION

Recent field work throughout the state of Florida has resulted in the discovery of 12 taxa previously unre-

ported for the state. An additional 37 taxa are reported here as significant collections that either confirm a

previously reported taxon's presence here (second and third collections), or represent a significant distri-

butional disjunction. Most of these are either protected by Florida law (Florida Chapter 5B-40) or are not

native to the state (Wunderlin & Hansen 2008). In order to prevent overcollecting, precise locality data for

protected taxa have not been included within the specimen citations.

NEW TO FLORIDA

Azolla pinnata R. Br. subsp. asiatica R.M.K. Saunders & K. Fowler (Azollaceae). This species, which has

a known range of much of the old world tropics (Saunders & Fowler 1992), was reported as new to Florida

in the Spring 2008 issue of Wildland Weeds (Brown 2008). Here, we provide voucher information and an

infraspecific determination.

Voucher specimens. Palm Beach Co.: Loxahatchee River watershed, along a N-5 canal, 26? 53' 47.6" N, 80° 8' 37.6" W (NAD 83), Jan

2008 [no day provided], Bodle s.n. (USF); unincorporated Jupiter area, canal 1/8 mi W of the intersection of 69% Dr and 149'^ Pl North,

26? 53! 47.6" N, 80? 8' 37.6" W (NAD 83), 25 Apr 2008, Bodle s.n. (FLAS)

Crataegus opaca Hook. & Arn. (Rosaceae). We consider C. opaca to be a distinct species and report it here

as new to Florida, although it is sometimes placed in synonymy under Crataegus aestivalis (Walter) Torr. &

A. Gray. It is currently known along the Gulf coast from Texas east to Alabama Md NRCS 2008).

Voucher specimens. Escambia C 2 mi SE of South Flomaton, al E I 4 bridge,

15 Mar 1981, Wilhelm 8553 (USF); W bank Escambia River N of Rte Old FL 4 bridge s a S of de concrete boat ramp (via

K Adress: John M. Kunzer, Park Biologist, Tomoka State Park, 2099 North Beach Street, Ormond Beach, Florida 32174, U.S.A.

J. Bot. Res. Inst. Texas 3(1): 331 — 337. 2009

332 tani it Texas 3(

Campbell Rd and Boat Ramp Rd), 24 Apr 1994, Burkhalter 14044 (UWFP); floodplain W side Escambia River N of boat ramp via Mystic

Springs Rd, 17 Apr 1988, Burkhalter 10914 (UWFP). Santa Rosa Co.: near E bank Escambia River N of FL 4 bridge, 21 Mar 1985, Bur-

khalter 9881 (UWFP); E bank of Escambia River N of FL 184 bridge, 300 ft N of bridge, 23 Oct 1986, Burkhalter 10472 (UWFP).

Cyperus bipartitus Torr. (Cyperaceae). This is the first report for Florida of this common species of the

continental United States and southeast Canada (USDA, NRCS 2008).

3 4269592 FfECTTY c : Ax ma = f 1

Voucher specimen. Wakulla Co.: Wakulla River, S of Hwy 98, 13 Sep 2007, A

by Richard Carter (VSC).

Cyperus eragrostis Lam. (Cyperaceae). This collection represents an apparently recent introduction; it

was previously known within the southeast United States from South Carolina and Alabama, west to Texas

(USDA, NRCS 2008).

Voucher specimen. Escambia Co.: corner of Nine Mile Rd and Untreiner Áve, near Pensacola, 28 Oct 2007, Burkhalter 21650 (UWFP).

Geranium maculatum L. (Geraniaceae). This is the first record for Florida of this common eastern North

American plant.

Voucher specimen. Gadsden Co.: bottom of steep ravine of Willacoochee Creek drainage, ca. 4.5 air mi NE of Quincy, 14 Apr 2008,

May s.n. (FSU).

Lamium purpureum L. (Lamiaceae). This represents the first record for Florida of an Eurasian weedy spe-

cies naturalized throughout much of North America (USDA, NRCS 2008).

Jack C iland } y Loop Rd, near NE end of Merritts Mill Pond, 7 Feb 2007, Anderson

22713 (FSU).

Melochia nodiflora Sw.(Malvaceae). This taxon, with a currently known range of Mexico, Central America,

the Caribbean basin and northern South America (Bornstein 1989), is new to the continental United

States.

I i Miami-Dade Co.: E lades National Park, “Hole-in-the-Donut” restoration project, 5 Dec 2007, Sadle 548 (FNPS,

USF).

Ochna serrulata (Hochst.) Walp. (Ochnaceae). This ly cultivated species is native to South Africa

and is new to the continental United States. This collection represents the first set of incontrovertibly wild

specimens we have seen from Florida, where it has escaped cultivation.

Voucher specimen. Palm Beach Co.: High Ridge Scrub Natural Area, N central portion, ca. 500 ft distant from the nearest cultivated

area (housing development), T45S, R43E, Sec. 9, 2 Mar 2007, Lietzky et al. s.n. (FLAS, USF)

Passiflora vitifolia Kunth (Passifloraceae). This taxon, which is native to Central and South America, is

new to the continental United States, where it has escaped cultivation.

Voucher specimen. Highlands Co.: spreading rampantly along the sh f Lake August, Placid Lakes Subdivision, Lake Placid, 8 Jul

2008, Pickert s.n. (USF)

Sida urens L. (Malvaceae). This is the first report for Florida. It was previously known from the Continental

United States only from Alabama (USDA, NRCS 2008). Its known range includes most of the Neotropics

and Africa (Fryxell 1988).

Voucher specimen. Broward Co.: Natural Area, 0.5 mi S of Orange Dri Flamingo Road, ca. 2.5 mi WNW of Cooper City,

T50S, R40E, Sec. 23, 26, 16 Jan 2008, ae 1048 (USF).

Sphaeropteris cooperi (Hook. ex F. Muell.) R.M. Tryon (Cyatheaceae). This is the first report from Florida

for this species and the second for the continental United States. In the United States, it is currently known

from Oregon (Wood 2008). It is native to tropical Australia.

Voucher specimen. Broward Co.: Tradewinds County Park, N and S of Sample Road, 0.5 mi E of Lyons Road; ca. 3 mi NW of Pompano

Beach, T485, R42E, Sec. 17, 20, 2 Apr 2004, Howell 779 (USF).

Kunzer et al., New vascular plant records for Florida 333

Torilis arvensis (Huds.) Link (Apiaceae). This species was reported for Florida by McGregor (1986), but

we have seen no specimens to document its occurrence. This validates its occurrence in the state.

Voucher specimen. Escambia Co.: Tarkiln Bayou State Park, SW of Pensacola, 10 May 2008, Burkhalter 21793 (FSU, UWFP).

SIGNIFICANT NEW COLLECTIONS

Asparagus aethiopicus L.(Asparagaceae). These are the first reports of this Category 1 species (FLEPPC

2007) for northern Florida and the panhandle. Wunderlin and Hansen (2003) list it for the central and

southern peninsula.

Voucher specimens. Nassau Co.: inside the St. iid River inlet, just w of Fort Clinch i dae stretches for more than 100 m, 17

Nov 2005, e s.n. disi Okaloosa Co.: South Santa Rosa Island, ca. 0.8 air mi W of Gulf Islands National Seashore, S side

of US 98, W side of , 30.39719908? N, 86.59745579? W, growing on a coastal dune with p vomitoria, d camara, and

Flaeagnus pungens, 4 Jan 2003, I ing 1643 (USF). Wakulla Co.: St. Marks National Wildlife Refuge, nat g by old

homesite near Wakulla Beach, 27 ul 2008, And 23990 (FSU).

Asplenium dentatum L.(Aspleniaceae). This northerly disjunction of this State of Florida endangered spe-

cies (Florida Chapter 5B-40) is at least 285 km.

oucher specimen. Volusia Co.: Bulcw Creek State Park, SSE of the jct. of Walter Boardman Ln and Old Dixie Highway, 20 Feb 2008,

Kunzer et al. 2268 (FLAS, USF

Asplenium erosum L.(Aspleniaceae). This county record represents a new northern limit for this State of

Florida endangered species (Florida Chapter 5B-40).

Voucher specimen. Flagler Co.: Bulow Creek State Park, ca. 2.6 km NNW of the jct. of Walter Boardman Ln and Old Dixie Highway,

26 Dec 2008, Kunzer et al. 2699 (USF).

Brickellia cordifolia Elliott (Asteraceae). These specimens represent a county record and a new locality

for a State of Florida endangered species (Florida Chapter 5B-40).

Voucher specimens. Wakulla Co.: edge of calcareous woods on N side Hwy 98, 4 Sep 2007, And 23463 (FSU); N side of US 98, ca.

40 plants at edge of mowed bahiagrass right-of-way against edge of mixed hardwoods, with Cercis, Carya, Fagus, Prunus serotina, and

Andropogon, 19 Sep 2007, Norris s.n. (USF).

Bromelia pinguin L. (Bromeliaceae). This is the second county record for Florida; the first, from Hillsbor-

ough Co., was in Wunderlin et al. (2002).

Voucher specimen. Volusia Co.: Tomoka State Park, between the youth camp and the park concession store, ca. 0.4 km due N of the

Old Dixie Highway bridge at the Tomoka River, 29? 20' 43" N, 81? 5' 13" W, 1 Oct 2004, Kunzer 684 (USF

Calibrachoa parviflora (Juss.) D'Arcy (Solanaceae). This is the third county record for this species from

Florida, all records of which are from panhandle counties.

Voucher specimen. Escambia Co.: W Pensacola Beach, Santa Rosa Island, 29 Apr 2007, Burkhalter 21362 (FSU, UWFP).

Campanula robinsiae Small dé MM MEA These collections represent a southern disjunction of over

50 km of this United States and Florida endangered ies (Florida Chapter 5B-40) from its only previously

known eu in the MN Hill area of Hernando LM FL.

Voucl Hills! h Co.: Hillsl h River State Park, | fa depressi h, 22 Mar 2006, vanHoek

et al. HRO442 (USF); Hillsborough River State Pak. 16 Mar 2007, Jensen et al. HR0452 (USF).

Casuarina glauca Sieb. ex Spreng. (Casuarinaceae). First reports of this Category I exotic (FLEPPC, 2007)

for northern Florida and the Panhandle. Wunderlin and Hansen (2003) list it for the central and southern

peninsula.

Voucl i Franklin C lized ( ] ites) on Alligator Point, 21 Jun 2008, Keys s.n. (FSU). St. Johns Co.:

13* Lane, St. A ine Beach, planted i d and dingt li near the beach, Meisenburg s.n. (FLAS).

e E / E o or he 4

334 t i i f Texas 3(

Cyperus involucratus Rottb. (Cyperaceae). This is the first report of this exotic species from the Florida

panhandle.

Voucher specimens. Wakulla Co.: St. Marks National Wildlife Refuge, in woods N edge of Panacea, 27 Sep 2007, A 1 23610 (FSU);

3 Jun 2008, Anderson 23869 (FSU).

Desmodium triflorum (L.) DC. (Fabaceae). This is the first report from west of the Apalachicola River in

the panhandle.

E bia Co.:1 f Uni ity of West Florid I near Pensacola, 5 Nov 2006, Burkhalter 21256 (FSU, UWFP).

ae Pi

Dyschoriste angusta (A. Gray) Small (Acanthaceae). In Florida, this species had previously been known

only from the southern half of the peninsula from southern Polk County southward. The Hernando County

specimen is a northwest disjunction of ca. 160 km, and the Wakulla County specimens are a northwest

disjunction of ca. 240 km from the Hernando County locality.

Voucher e mee Hem Co.: na Preserve, ca. 2.4 km W of the jet. of C-595 and US 19, ca. 3.3 km NNW of the June-

tion of the I with US 19, 28° 38’ 58” N, 82° 27 48" W, 10 Jun 2006, Kunzer et al. 1824 (USF). Wakulla Co.

t. Marks National Wildlife Refuge, g grassy adios (Rd 203) in coastal wetlands/flatwoods, 8 Sep 2006, Anderson 22360 (FSU); bas

ae Boa Rd ae just N of Rd oa e na ee Mia Pd (FSU); 0.2 mi N of Rd 202 on Rd 205, 6 Jun 2008, Anderson 23917

nd of Rd 04, 6 Jun 2008, Anderson 23924 (FSU).

Eragrostis unioloides (Retz.) Nees ex Steud. (Poaceae). This specimen represents a southern disjunction

of at least 130 km from the nearest known population in Polk County, Florida.

Voucher specimen. Lee Co.: ca. 6.5 mi E of the jct. of 1-75 and Corkscrew Road (C-850), S of Corkscrew Road, 26? 26' 31.2" N, 81?

4021.3" W, 17 Sep 2005, Kunzer 1137 (USF).

Ficus aurea Nutt. (Moraceae). This is a new northernmost station for this taxon in Florida.

Voucher specimen. Volusia Co.: Ponce Inlet, Lighthouse Point Park, ca. 250 m SE of the lighthouse, 29° 4' 49" N, 80° 55' 30" W, 12

Aug 2005, Kunzer et al. 1107 (USF)

Hypericum canadense L. (Clusiaceae). This species is new to the central panhandle; it was previously

known only from extreme northeast and extreme west Florida.

Voucher specimen. Calhoun Co.: locally common amongst grasses and sedges in wet sandy loam bordering ditch along Rte 167 in

(FSU).

extreme NW corner of county (Tenmile Creek drainage), 30.54583° N.; 85.369038"? W, 18 Aug 2008, Anderson 24119 (FSU

Jasminum mesnyi Hance (Oleaceae). This is the first record from the Florida panhandle.

Voucher specimen. Wakulla Co.: St. Marks National Wildlife Refuge, naturalized hedge spreading old shop area, Rd 132, 3 Apr

2008, Anderson 23743 (FSU)

Juncus bufonius L. (Juncaceae). This collection represents a new southernmost station for this taxon in

Florida.

Voucher specimen. Polk Co.: Colt Creek State Park, ca. 3.3 km SE of FL 471 bridge over the Withlacoochee River, ca. 4.6 km NE of jet.

FL 471 and US 98,28? 17' 13.4" N, 82° 2! 21.1" W, 15 Apr 2008, Kunzer & Hansen 2391 (USF

I tal thulatum (Muhl) Elliott (Parnassiaceae). These are the second and third county records

lor this State of Florida endangered species (Florida Chapter 5B-40), and all of these are considerably west

of its previously known records in Gadsten County.

oucher specimens. Escambia Co.: abundant in ditch near Pensacola Blvd, 29 Mar 2008, Burkhalter 21744 (FSU, UWFP). Walton Co.:

ditch beside Hwy, DeFuniak Springs, 15 Mar 2008, Burkhalter 21720 (FSU, UWFP

Listera australis Lindl. (Orchidaceae). This easily overlooked species is known sporadically throughout

north Florida and the northern Florida peninsula (Wunderlin & Hansen, 2008), and westward to Texas

and northward to Quebec, Canada (USDA, NRCS 2008). This station represents the new southern limit for

the species, which is threatened in Florida (Florida Chapter 5B-40).

Kunzer et al., New vascular plant records for Florida 335

Voucher specimen. Sarasota Co.: N cf Plantation Boulevard, 14 Jan 2008, Kunzer 2266 (FLAS, USF).

Mecardonia procumbens (Mill) Small (Veronicaceae). This specimen represents the first record of this

species from the Florida panhandle.

Voucher specimen. Escambia Co.: Hwy 29 just S of Church St, near Century, 19 Apr 2008, Burkhalter 21779 (FSU, UWFP).

Momordica charantia L. (Cucurbitaceae). This is the second report for the panhandle and represents a

new county record for the western panhandle. The first record for the panhandle was from Leon County

(Anderson 2007).

Voucher specimen. Escambia Co.: fence row at corner Gulf St and Jackson St, near Pensacola, 6 Oct 2007, Burkhalter 21638 (FSU,

Peltophorum pterocarpum (DC.) Backer ex K. Heyne (Fabaceae). Previously known from Miami-Dade

County and the Monroe County keys, this is the third county record from Florida.

Voucher specimen. Lee Co.: South Fort Myers (unincorporated), on the W side of Metro Parkway, ca. 0.4 km SSW of the jct. of Metro

Parkway and Colonial Boulevard (FL 884), just E of Tenmile Canal, 81? 51' 11" W 26? 35' 27" N, 29 Sep 2006, Kunzer 1900 (USP).

Phlebodium aureum (L.) J. Sm. (Polypodiaceae). This is the second county record from the Florida pan-

handle.

Voucher specimen. Wakulla Co.: St. Marks National Wildlife Refuge, fallen palm trunk beside Northline Rd (rd 200), 31 Aug 2006,

Anderson 22310 (FSU)

Polygala verticillata L. var. isocyta Fernald (Polygalaceae). This species was previously known in the

Florida panhandle from a site reported by Anderson (1986). That population has been extirpated, so these

new sites verify its continuance in the county (and the panhandle) in more natural settings.

Voucher specimens. Wakulla Co.: St. Marks National Wildlife Refuge, Rd 205 just N of Rd 206, 17 May 2007, Anderson 23062 (FSU);

edge of Rd 108 at junction Rd 107, 17 Jun 2008, Anderson 23951 (FSU

Ranunculus pusillus Poir. (Ranunculaceae). These collections, only ca. 25 km apart, represent a southern

disjunction of at least 140 km from the nearest known station in Alachua County. (Wunderlin & Hansen

Voucher specimens. Hillsborough Co.: Hillsborough River State Park, Riiie a oiu depression marsh on the S side of

the Fort King Trail, ca. 200 yds W of US 301, T27S, R21E, Sec. 17, SE% of NW, 22 Mar 2006, vanHoek et al. HRO436 (USF). Polk Co.:

Colt Creek State Park, ca. 4.8 km SE of FL 471 bridge over the Withlacoochee River, ca. 3.2 km NNE of jct. FL 471 and US 98, 28? 16'

4.1" N, 82? 2' 32.9" W, 3 Apr 2008, Kunzer et al. 2320 (USF).

Richardia grandiflora (Cham. & Schltdl.) Steud. (Rubiaceae). This species appears to be rapidly spread-

ing throughout peninsular Florida, and the Alachua County specimen represents the current northernmost

station in the state.

Voucher specimens. Alachua "ille, 2950 Archer Road, weed growing in lawn, 22 Jul 2008, Weaver 5176 (PIGH, USF). Volu-

sia Co.: South Daytona, "pi "s S side of Reed Canal Road, ca. d mE pi ie jet. of Reed Canal Road and Clyde Morris Boulevard,

T165, R33E, Sec. 6, NW% of NE, 24 Nov 2005, Kunzer 1221 (USF); al f Indian Lake Road, next to Volusia County Public

Library, ca. 0.5 mi N of US 92, 5 mi W of I-95, T15S, R31E, Sec. 36, 13 Jul 2007, Slaughter 15741 (USP).

Rumohra adiantiformis (G. Forst.) Ching (Dryopteridaceae). This is the first record from the Florida

panhandle.

Voucher specimen. Franklin Co.: naturalized under oaks along Bay Street, Apalachicola, 29 Dec 2007, Anderson 23624 (FSU).

Schefflera arboricola (Hayata) Merr. (Araliaceae). This collection represents a disjunction of over 200 km

and a new northern station for this taxon.

litt Darl 357 CE af the lah inl ls 1 IA tal

Voucher specimen. Volusia Co.: Ponce inlet, Li ca g j I

swale, 80° 55' 30" W 29° 4' 49" N, 12 Aug 2005, Kunzer et al. 1108 (USE).

336 tani i Texas 3(

Spigelia loganioides (Torr. & A.Gray ex Endl. & Fenzl) A. DC. (Strychnaceae). The vouchers for this

endangered (Florida Chapter 5B-40) endemic species represent an eastern disjunction of at least 85 km

from the eastern edge of this species’ known range of central Marion County south to north-central Sumter

County and west to southern Levy County.

Voucher specimens. Volusia Co.: Bulow Creek State Park, 22 Dec 2006, Kunzer & DuToit 2027 (USF), Bulow Creek State Park, 7 Aug

2008, Kunzer et al. 2027 (USF).

Spiranthes xitchetuckneensis P.M. Br. (Orchidaceae). This taxon is new to the Florida panhandle.

Voucher specimen. Wakulla Co.: St. Marks National Wildlife Refuge, cypress-maple-g p bordering Shepherd's Spring, 13 Nov

2006, Anderson 22637 (FSU).

Spiranthes lucayana (Britton) Cogn. (Orchidaceae). Endangered in the State of Florida (Florida Chapter

5B-40), this population and county record was first discovered by Charles DuToit, the Park Biologist for the

Tomoka Basin Parks State Park complex. Paul Martin Brown has visited this site and confirmed the presence

of this species within the park (C. DuToit, pers. comm.).

Voucher specimen. Volusia Co.: North Peninsula State Park, 13 Mar 2007, Kunzer et al. 2078 (USF).

Stachys crenata Raf. (Lamiaceae). This is the only known station outside of Gadsden County for this state

AM taxon NA eau 5B-40).

Hillst Hills) Park, roadside, 2 May 2008, vanHoek & Parsons 2405 (USF); Hillsborough

River State Park, 18 May 2008, YaniHaek & Parsons : 2406 (USF).

Tillandsia pruinosa Sw. (Bromeliaceae). This species is endangered in the State of Florida Florida Chapter

5B-40). In Florida this taxon has only been documented from conservation land in Collier County, with

vouchers only being known from the areas of Fakahatchee Strand Preserve State Park and Collier-Seminole

State Park. The closer of these, Fakahatchee Strand Preserve State Park, is approximately 60 km to the

southeast of this new locality. The collection represents a range extension that is geographically minor but

floristically significant.

Voucher specimen. Lee Co.: 18 Oct 2007, Kunzer et al. 2205 (USF).

Tridens carolinianus (Steud.) Henrad (Poaceae). This collection represents a southerly disjunction of over

235 km from the nearest known locality in Suwannee County, FL.

Voucher specimen. Hillsborough Co.: "Fishhawk" Preserve, ca. 0.5 mi S of the Alafia River, ca. 0.25 mi N of Fishhawk Boulevard and

N of powerline right-of-way, ca. 1 mi E of Bell Shoals, T30S, R21E, Sec. 19, 30 Nov 2006, Dickman s.n. (USF).

Urochloa piligera (Muell. ex Benth.) R.D. Webster (Poaceae). This is the first report for this species from

Florida west of the Apalachicola River.

Voucher specimen. Escambia Co.: along Palafox St near Pensacola, 22 Jul 2007, Burkhalter 21464 (FSU, UWFP).

Urochloa plantaginea (Link) R.D. Webster (Poaceae). This is the first report for this species from Florida

west of the Apalachicola River.

Voucher specimen. Escambia Co.: S side Nine Mile Rd near Bowman Ave, near Pensacola, 23 Sep 2006, Burkhalter 21096 (FSU,

Veronica polita Fr. (Veronicaceae). This is the second county of record for this species in Florida, the other

iii: Escambia County.

I Co.: | Robinhood Rd, Tallahassee, 14 May 2008, Anderson 23817 (FSU); edge of woods on Lakeshore

Dr, Tallahassee 14 May 2008, Anderson 23819 (FSU).

Xyris longisepala Kral (Xyridaceae). These represent a new county record for this Panhandle endemic

species, which is endangered in Florida (Florida Chapter 5B-40).

Kunzer et al., New vascular plant records for Florida 337

Voucher specimens. Wakulla Co.: St. Marks National Wildlife Refuge, ditch, 26 Jul 2007, Anderson 23379 (FSU); edge of Ilex pond,

ep 2007, Anderson 23470 (FSU); edge of Hypericum zone of pond, 4 Sep 2007, Anderson 23479 (FSU); cypress pond, 11 Sep 2007,

Anderson 23500 (FSU

ACKNOWLEDGMENTS

The authors would like to thank Patricia Howell, Stephen Dickman, Carmel vanHoek, Michael Bodle, Jimi

Sadle, Richard Weaver, Cecil Slaughter and Lee Norris for tl j , and Alice Bard (FDEP Florida Park

Service), Charles DuToit (FDEP Florida Park Service), and Mary Barnwell (Southwest Florida Water Man-

agement District) for the collecting opportunities. Our appreciation is also extended to the personnel at St.

Marks National Wildlife Refuge for providing logistical support. We also extend our thanks and appreciation

to Kent Perkins (FLAS) and an anonymous reviewer for their helpful comments and suggestions.

REFERENCES

ANDERSON, L.C. 1986. Noteworthy plants from north Florida Il. Sida 11:379-384.

ANDERSON, L.C. 2007. Noteworthy plants from north Florida VIII. J. Bot. Res. Inst. Texas 1:741-751.

BORNSTEIN, A.J. 1989. Sterculiaceae. In: R.A. Howard. Flora of the Lesser Antilles, Leeward and Windward Islands;

vol. 5. Arnold Arboretum, Harvard University, Jamaica Plain, MA.

Brown, K. ed. 2008. Internodes. Wildland Weeds 11:22.

FLORIDA Exotic Pest PLANT Council (FLEPPC). 2007. Florida Exotic Pest Plant Council's 2007 list of invasive plant spe-

cies. www.fleppc.org (accessed 3 Jul 2008).

FryxeLL, PA. 1988. Malvaceae of Mexico. Syst. Bot. Monogr. 25:1-522.

McGREGOR, R.L. 1986. Apiaceae. In: Great Plains Flora Association, eds. Flora of the Great Plains. University Press

of Kansas, Lawrence.

SAUNDERS, R.M.K. AND FowLER. 1992. A morphological taxonomic revision of Azolla Lam. sec. Rhizosperma (Mey)

Mett. (Azollaceae). Bot. J. Linn. Soc. 109:329-357.

USDA, NRCS. 2008. The PLANTS Database(http://plants.usda.gov, 4 November 2008). National Plant Data Center,

Baton Rouge, LA 70874-4490 USA.

Woop, W. 2008. Subtropical tree fern Sphaeropteris cooperi (Hook. ex F. Muell) R.M. Tryon, found modestly estab-

lished in Oregon. Amer. Fern J. 98:113-115.

WUNDERLIN, R.P. AND B.F. HANSEN. 2003. Guide to the vascular plants of Florida: 274 ed. University Press of Florida,

Gainesville, FL.

WUNDERLN, R.P. AND B.F. Hansen. 2008. Atlas of Florida vascular plants (http//www.plantatlas.usf.edu/).[S.M. LANDRY

and K.N. CameseLL (application development), Florida Center for Community Design and Research.] Institute

for Systematic Botany, University of South Florida, Tampa.

WUNDERLIN, R.P, B.F. HANSEN, AND L.C. ANDERSON. 2002. Plants new to the United States and Florida. Sida 20:

813-817.

BOOK NOTICES

T.G. LAMMERS. 2009. Augustus Green in the Lair of the Pye-a-Saw. (ISBN none, saddle-stapled). Published

by the author. (Orders: tlammersOnew.rr.com). $5.00 postpaid, 62 pp., illustrated, 8 1/2" x 5 1/2".

From the Publisher—An Amazing Fictional Advent the American Frontier!

Four years before the Louisiana Purchase, an intrepid g American botanist defies the Spanish authorities to study the flora

of what is today ] Iowa. A panied only by hi ful French-Canadian guide, lc boldly plunges into this IM

J: A 1 : 1 1 ; 1 14

land, gp g wilderness.

Then h t i f f antiqui ible 1 hat fed up he d d buffalo

LI 1 1; 1; E wma E 11 1 ane A Faura J = Tx E

and men! g P I g

1 pee f +1 J 1 1 LAA ERR di

path ... What will be the fate of the vods explorer when he y ying y

of his life?

EO. ZuLoAGa, O. Morrone, AND M.J. BELGRANO (EDS). 2008. Catálogo de las Plantas Vasculares del Cono

Sur: Argentina, Sur de Brasil, Chile, Paraguay y Uruguay - Volumen 1, 2, and 3. (ISBN 978-

1-930723-70-2, hbk.). Missouri Botanical mea Press (St. nd 4344 Shaw Boulevard, St. Louis,

Missouri 63110-2291, U.S.A. (Orders: v Org, @mobot.org, 314-577-9534, 314-

571-9594 B $375.00 set, Vol. 1: 984 p pp., Vol. 9» 1302 pp Vol. 3: 1602 pp., 8 1/2" x 11 1/4".

From the publisher: “The Catalogue of the Pl of the S hecklist of tł ] i ,000 ae found Reus southern

Brazil, Chile, Paraguay, and Uruguay. This is the first time tl h an international list h d. In three volumes, 17,697

accepted species are treated, organized alphabetically by family and genera, and documented with a citations, herbarium

specimens, and information on their distribution, habit, native or introduced status, elevation, and cross reference synonymy.” More

forthcoming!

J. Bot. Res. Inst. Texas 3(1): 338. 2009

VASCULAR FLORA OF CHURCHILL RANCH, SARASOTA COUNTY, FLORIDA

Alan R. Franck Richard P. Wunderlin

Department of Cell Biology, Microbiology, Department of Cell Biology, Microbiology,

Molecular Biology and Molecular Biology

University of South Florida University of me lorida

Tampa, Florida, 33620-5200, U.S.A. Tampa, Florida, 33620-5200, U.S.A.

ABSTRACT

Churchill Ranch, owned by Sarasota County, contains 170 hectares with eight plant communities. A survey of vascular plant species

conducted from May 2007 to August 2008 yielded 368 taxa, o 89 famili d 241 genera. Of these, 327 (88.9%) were native

taxa, 41 (11.1%) exotic, 15 endemic to Florida, four state-endanger d 29 new county o voucher records,

RESUMEN

El Rancho Churchill, propiedad del S ounty, contiene 1701 ] id les. Un estudio de las especies

de plantas vasculares dea lizsdo ind iun e a pd 2008 sumó 368 taxa, de d familias y 241 géneros. De estos, 327 (88.996) fueron

nativas, el 41 (11.196) exóticas, rida , y 29 nuevo citadas con testigos en el condado.

INTRODUCTION

Churchill Ranch (CR) is a 170 ha site acquired by Sarasota County through the Environmentally Sensitive

Lands Protection Program. The site, situated between Myakka River and Deer Prairie Slough, is managed

as part of the nearby Deer Prairie Creek county properties (totaling ca. 3000 ha). It is located about 2.3 km

ENE of the junction of Interstate 75 and County Road 777 (River Rd.), approximately 13 km ENE of the

gulf coast, 27°07'03"N, 82?19'40"W; T39S, R20E, Sec 5 (Fig. 1). Border Road marks the northern bound-

ary, and Interstate 75 marks the southern boundary of the property. Access to CR is from Border Rd. off

Jacaranda Blvd. The results of this research are intended to help Sarasota County make knowledge-based

management plans for its property.

SITE OVERVIEW

Geology, Topography, Hydrology, and Soils.—The mean elevation of Churchill Ranch is about 7 m above

sea level. The surface soils are mostly acidic, sandy spodosols low in fertility which were deposited by sea-

water in the Pleistocene during receding sea level. Herein lies the Surficial Aquifer System which contains

the water table 2-6 m below mean sea level (FDEP 2007). Below this is the Miocene Hawthorn Group (2-30

m below mean sea level) composed of the Peace River and Arcadia Formations which constitute the Inter-

mediate Aquifer System. Below this is the Florida Aquifer System (100—200 m below mean sea level) which

contains the Oligocene Suwannee Limestone, Eocene Ocala Limestone, and Avon Park Formation (Hyde et

al. 1991; Halford and Yobbi 2006). The surface soil is 42.796 Eugallie and Myakka fine sands characterized

by slash pine (Pinus elliottii) flatwoods; Pineda fine sand (21.5%) is largely mesic hammock; Wabasso fine

sand (12.796) is a mixture of mesic hammock and pine flatwoods; Holopan fine sand, (11.696) occurs in

herbaceous marshy depressions. Felda fine sand (7.196) marks hydric hammocks containing open marshy

sites as does Pople fine sand (3.096). Delray fine sand (1.396) is found partly in an herbaceous marsh in a

hydric hammock as well as a basin marsh which drains into an adjacent pond (USDA 2007).

Climate.—Sarasota County has a subtropical climate with a humid, rainy summer period (June—

September) and a cooler, drier period (October-May). The wet season averages 82.07 cm of rainfall and

the dry season averages 52.10 cm (SWFWMD 2008) with about 100 days per year receiving precipitation

(NCDC 2008). The year 2007 had the lowest amount of annual rainfall for Sarasota County in the last 92

years (SWFWMD 2008). Average wind speed annually is 13 km/hour with prevailing winds usually blowing

J. Bot. Res. Inst. Texas 3(1): 339 — 348. 2009

laf Dat 21D Is IRL £T,

—

LEGEND

Flatwoods Lake

Marsh Lake

Basin Marsh

Depression Marsh

Hydric Hammock

Mesic Hammock

Mesic Flatwoods

Xeric Hammock

Ruderal

Fig. 1 Tha | Pr £fL khi D Ltrs IL ha hls lr ct thin d f. ty, Florida TI

Ranch.

east. Daily maximum temperatures range from 32.69°C in August to 22.61°C in January. Daily minimum

temperatures range from 23.83°C in August to 11.92°C in January. During most years freezing

occur once or twice, only with occasional localized spots of frost. Relative humidity in the morning ranges

from 90% in August and 86% in April. Relative humidity in the afternoon ranges from 62% in August to 49%

in April. Climate data other than rainfall was averaged from weather stations in Tampa (north of Sarasota)

and Ft. Myers (south of Sarasota) as recorded by the NCDC (2008).

Franck and Wunderlin, Vascular flora of Churchill Ranch 341

Past Land Use.—Aerial photographs from 1948 and 1957 (UF 2004) suggest past logging and the

site likely has been cattle ranched for most of the remaining years. Concomitant with this study, intermit-

tently from December 2007 through July 2008, a cattle herd of 10-15 foraged on the land. Significant fire

has likely not occurred for 10 or more years.

METHODS

Vascular plant collecting occurred once every 1-4 weeks from May 2007 to August 2008. Permits for

endangered species were obtained from the Florida Department of Agriculture and Consumer Services.

Voucher specimens were deposited at the University of South Florida Du ad The nomenclature

used follows Wunderlin and Hansen (2003) with updat flected on the herba: ite Atlas of Florida

Vascular Plants (Wunderlin & Hansen 2008). Natural community descriptions follow Florida Natural Areas

Inventory (FNAI 1990).

Vascular plants of special interest include those taxa that are state-listed species and exotic species

listed by the Florida Exotic Pest Plant Council (FLEPPC 2007). Taxa unreported for Sarasota County as

determined by Wunderlin and Hansen (2008) are noted.

RESULTS

The inventory yielded 368 taxa representing 89 families and 241 genera at Churchill Ranch. Native species

comprised 327 taxa (88.9%) and 41 (11.1%) were exotic species. The greatest numbers of taxa recorded were

in the families Poaceae (59), Asteraceae (51), Cyperaceae (39), Fabaceae (18), Rubiaceae (11), Euphorbiaceae

(9), and Plantaginaceae (9). Rhynchospora, with 10 taxa, represented the largest genus, followed by Cyperus

(8), Hypericum (7), Quercus (7), Panicum (7), Andropogon (6), Dichanthelium 6), Fimbristylis (6), Tillandsia (6),

Eragrostis (5), and Polygala (5). The families containing the most number of exotics were the Poaceae (13),

Fabaceae (6), Asteraceae (4), and Amaranthaceae (3). Of the exotics, eight were listed as category I and three

as category II invasive plant species as designated by the Florida Exotic Pest Plant Council (FLEPPC 2007)

(Table 1). There were 29 new county records for Sarasota. Fifteen species (3.5896) are endemic to Florida, of

which three are in the family Campanulaceae. Four endangered species, Glandularia tampensis, Lythrum flagel-

lare, Tillandisa fasciculata, and T. utriculata, and two commercially exploited species, Encyclia tampensis and

Osmunda regalis var. spectabilis, were found. One hybrid, Quercus laurifolia x Q. pumila, was documented.

PLANT COMMUNITIES

The classification for plant communities found within the study (Fig. 1) following the Florida Natural Ar-

eas Inventory (FNAI 1990) is discussed below from the lowest elevation to the highest followed by ruderal

areas.

Flatwoods Lake.—One large pond covering 4.9 ha (2.996) of the site is dominated by Typha domin-

gensis with other herbs such as Eleocharis interstincta and Hydrocotyle umbellata growing along its perimeter.

A small island sits in the center, dominated by Polygonum glabrum and Salix caroliniana. It is accessible by

land when the water level is down.

Marsh Lake.—Marsh lakes characterize the other perennial bodies of water at the site, covering 0.2

ha («0.0196). In the extreme southeast corner a small pool of water next to Interstate 75 reaches into the

site. This contains the floating "ndn Leanna il and Wolfiella oblonga. Also along Border Rd. lies a

small elliptic pond where the native sp Azolla inensis and Nuphar advena, and the exotics, Ludwigia

peruviana, Panicum repens, and Salvinia minima, are found. During the peak of the drought, this pond was

nearly dry.

Basin Marsh.—A basin marsh occurs directly adjacent to the large pond, covering 3.3 ha (1.9%). The

ground is densely covered with herbs such as Iris hexagona, Phyla nodiflora, and Polygonum punctatum. Before

the summer rains of 2008, Baccharis halimifolia had successfully invaded and reached maturity. However

after the rains, the inundated conditions began to cause dieback of these shrubs.

342 tani i Texas 3(

EI EDD. Pte Ber NANTI

Tage 1. Exotic i

Category | Category Il

Dioscorea bulbifera Ricinus commu

Hymenachne Md Sphagneticola a

Ludwigia peruv. Urena lobata

Melinis repens

Panicum repens

Schinus terebinthifolia

Solanum viarum

Depression Marsh.—Depression hes occur on 15.1 ha (8.996) of the site. From the beginning of the

study and through the spring of 2008, the soils ively dry and never inundated. However, significant

summer rain in 2008 saturated and flooded these B S peta accum one marsh.

Other marshes are characterized | bs | I flag ij Others are

Lal th T dC T3 pa MES a A tin El 1 pl UPS

dominated by Cep

Hydric Haroitrock. —The hydric hammocks occur on 15.5 ha (9.196) of the site. The dou canopy

trees are Quercus laurifiolia and Q. virginiana with Ulmus americana as an occasional associate. An isolated

patch of hydric Hammock is oe pid ida caiman. Various, d d bp) herbaceous, marshy

sites occur here. A i I Y g JI Sisyrinchium

angustifolium, Cladium jamai Hibi difl andl Helianthus agrestis- Campanula a floridana. Epiphytes

such as Encyclia tampensis, Tillandsia S Vittaria lineata, and Pleopeltis polypodioides var. michauxiana are

abundant. The exotic Solanum viarum also occurs here.

Mesic Hammock.—The mesic hammocks occur on 57.9 ha (34.1%) of the site with Quercus virgini-

ana dominating the canopy and Sabal palmetto the subcanopy. Serenoa repens characterizes the understory,

interspersed with Callicarpa americana and Smilax bona-nox. Exotic species include Schinus terebinthifolia,

which is being treated for removal by the county, and Sphagneticola trilobata.

Mesic Flatwoods.—The mesic flatwoods is the dominant community, covering 69.7 ha (41.0%). The

canopy consists mainly of Pinus elliottii with a dense understory of Serenoa repens of 1-2m, reaching 3-4m

ina d pae TL reli nature of s flatwoods is obviously due to fire suppression. Other common ele-

ticulata, Ilex glabra, Lyonia fruticosa, Lyonia lucida, Q elliottii, Quercus

minima, and Vaccinium darrowii and the vines Galactia elliottii and mila auriculata. Common herbaceous

species include Aristida spiciformis, Fimbristylis puberula, Ludwigia maritima, Lygodesmia aphylla, Panicum

anceps, Polygala setacea, Pteridium aquilinum var. pseudocaudatum, and Solidago odora var. chapmanii.

Xeric Hammock.—A small ridge of land in the westernmost portion, wedged between a mesic ham-

mock and a marsh, represents a xeric hammock of 1.7 ha (1.0%). Here the canopy is composed of Quercus

nigra and Quercus virginiana. The understory has mostly Serenoa repens but its sparseness here allows open

sites to occur. In these open areas plants such as Gratiola hispida, Opuntia humifusa, Pityopsis graminifolia,

and Stipulicida setacea var. lacerata occur.

Ruderal.—The ruderal areas are found along the roads, trails, and firebreaks which make up about 1.7

ha (1.0%). Patches of vegetation along the roads and trails are seasonally mowed. Common in these areas

are the native species Bulbostylis stenophylla, Paspalum setaceum, and Setaria parviflora. Common exotics are

Commelina diffusa, Fimbristylis schoenoides, and Paspalum notatum var. saurae.

DISCUSSION

Disturbance, mainly fire, is a well-recognized component of Florida ecosystems. At Churchill Ranch, fire

suppression has allowed the flatwoods to become thickly overgrown with woody vegetation. Other distur-

bances which significantly impact the plant ities include foraging by domestic cattle, feral pigs, and

anthropogenic mechanical disturbance (mowing, roller chopping).

Franck and Wunderlin, Vascular flora of Churchill Ranch 343

Mowing has an essential role in maintaining paths and firebreaks. However, mowing maintains herba-

ceous communities as disturbed sites and may da wee short life cycle, high-fecundity species such as

the exotics Digitaria longi d Eragrostis at d the native Paspalum setaceum. It also may replace

the effects of grazing allowing the exotic Paspalum notatum var. saurae to continue. These disturbed sites

may also serve as corridors for the spread of invasive species.

Roller chopping has been implemented as a way to reduce shrub vegetation, mainly Serenoa repens as

well as Lyonia spp. and Quercus spp., to reduce fire intensity, and to increase light and resource availability

to understory herbs. Approximately 50 hectares of mesic flatwoods adjacent to Interstate 75 were roller

chopped at Churchill Ranch in March 2007, reducing fire hazard to drivers on the interstate. Serenoa repens

was predominantly affected with some saplings such as Pinus elliottii toppled as well. A thick layer of detritus

from affected plants, uncommon in flatwoods, was also left. As the roller chopping occurred near the end

of this study post-treatment effects were not assessed. Roller chopping without burning at a dry prairie in

nearby Myakka River State Park did not increase the abundance of native grasses (Watts et al. 2006).

Huffman (2006) estimates that most pine flatwoods have historically experienced natural fire, not

started by man, at least every five years. The lack of fire disturbance, or fire suppression, may have negatively

impacted the species richness of Churchill Ranch in plants as well as other organisms. Burning would be

ideal for this site but is problematic due to its location next to Interstate 75.

Exotic disturbances (cattle grazing and feral hogs) and the lack of natural disturbance (fire) have likely

played a large role in the ecology of CR. Plant herbivory by large animals has been intensive at CR and could

have quickly reduced or extirpated some taxa. Many taxa recorded at CR seemed to occur as one delicate,

reduced population. Some examples include species found exclusively along fencerows along the property

boundary such as Muhlenbergia capillaris, Nephrolepis exaltata, Osmunda regalis, and Thelypteris kunthii.

ANNOTATED LIST OF VASCULAR PLANTS

The list is artificially grouped into PTERIDOPHYTES, GYMNOSPERMS, MONOCOTS, and DICOTS. Within

these four groups, families are listed alphabetically, and within each family, the taxa are alphabetical by

genus and species. Following each species name is its authorship, habitat in the study area, and the senior

author's collection number. The habitat abbreviations are FL-flatwoods lake, ML-marsh lake, BM-basin

marsh, DM-depression marsh, HH—hydric hammock, MH-mesic hammock, MF-mesic flatwoods, XH-

xeric hammock, and RU-ruderal areas. Annotations preceding taxa are ^ for a new county record, * for

exotic species, and + for taxa endemic to the state of Florida.

PTERIDOPHYTES Salviniaceae

Azollaceae *Salvinia minima Baker —ML; 668

Azolla filiculoides Lam.—ML; 403 Thelypteridaceae

Blechnaceae Thelypteris kunthii (Desv.) CV. Morton—MH; 631

Blechnum serrulatum Rich.—MF; 279 Vittariace

Woodwardia virginica (L) Sm.—MF; 280 Vittaria mad (L) Sm.—HH; 227

Dennstaedtiaceae GYMNOSPERMS

Pteridium aquilinum (L.) Kuhn var. pseudocaudatum (Clute) i

Clute ex A. Heller—MF; 397 Pinaceae

Pinus elliottii Engelm.—MF; 217

Nephrolepidaceae

Nephrolepis exaltata (L.) Schott—MH; 444 MONOCOTS

Osmundaceae Alismataceae

Osmunda regalis L. var. spectabilis (Willd.) A. Gray-—MH; 445 Sagittaria graminea Michx.—DM; 824

Polypodiaceae Sagittaria lancifolia L.—DM; 264

Phlebodium aureum (L) J. Sm.—HH; 214 Amaryllidaceae

Pleopeltis polypodioides (c FE G. do E Windham Va. Crinum americanum L.—HH; 327

ict iana (Weath.) E MH; 199

Araceae

^Lemna dinis n —ML; 404

Pistia stratio 402

^Wolfiella Km cu ) Hegelm.—ML; 405

Are accan

(Walter) Lodd. ex Schult. & Schult. f—-MH; 662

n repens (W. Bartram) Small—MF; 100

de minc Schult. & Schuit. f —MH; 635

Tillandsia fasciculata Sw.—MH; 374

Tillandsia recurvata (L.) E MH; 361

Tillandsia setacea Sw.—MMH; 323

Tillandsia usneoides (L) L—MH; 111

Tillandsia utriculata L.—XH; 252

Commelinaceae

*Commelina diffusa e RU; 752

Commelina erecta L.—X

*Murdannia nudiflora (L 1 D RU; 394

Cyperaceae

Bulbostylis ciliatifolia (Elliott) Fernald—XH; 313

ola stenophylla (Elliott) C.B. Mm 257

Carex lupuliformis Sartwell ex Dewey—ML;

a exans FJ. vua .—FL; 209

Cladium jamaicense Crantz—HH; 144

Cyperus distinctus Steud.—BM; 270

Cyperus ligularis L.—RU; 350

Cyperus odoratus L.—BM; 191

Cyperus polystachyos Rottb. as a 349

o retrorsus Chapm

Cyperus surinamensis Rottb. —

Pens geniculata (L.) m Hum —FL; 452

Eleocharis interstincta (Vahl) Roem. & Schult.—FL; 197

Fimbristylis autumnalis (L) Roem. & Schult.—FL; 211

Fimbristylis cymosa R. Br.—RU; 150

^Fimbristylis dichotoma (L.) Vahl—RU; 345

Fimbristylis puberula (Michx) Vahl—MF; 712

*Fimbristylis schoenoides (Retz.) Vahl—RU; 443

Fimbristylis spadicea (L.) Vahl—HH; 218

Fuirena breviseta (Coville) Coville—RU; 454

Fuirena scirpoidea Michx.— pa , 654

Kylli

^Lipocarpha micrantha (Vahl) G. a Tucker—MH; 3

^*Oxycaryum cubense (Poepp. & Kunth) en m

Rhynchospora colorata (L.) H. Pfeiff —MH; 6

Rhynchospora divergens Chapm. ex M.A. NE 371

Rhynchospora fascicularis (Michx.) Vahl—MF; 205

^Rhynchospora fernaldii Gale—M

Rhynchospora globularis (Chapm.) Small—HH; 419

Rhynchospora inundata (Oakes) Fernald—DM; 363

Rhynchospora microcarpa Baldwin ex A. e ld 268, 419

Rhynchospora nitens (Vahl) A. Gray—ML; 4

ARhynchospora odorata C. Wright ex c 230

Rhynchospora plumosa Elliott—MF; 267

Schoenoplectus tabernaemontani (C.C. Gmel) Palla —BM; 193

Scleria reticularis Michx.—RU; 412

Scleria triglomerata Michx.—XH; 253, 428

Dioscoreaceae

*Dioscorea bulbifera L.—RU; 296

Eriocaulaceae

Eriocaulon decangulare L.—ML; 275

Lachnocaulon anceps (Walter) Morong—MF; 12

Syngonanthus flavidulus (Michx.) Ruhland—MF; ; 15

aemodoraceae

Lachnanthes caroliana (Lam.) Dandy—MF; 379

Hypoxidaceae

Hypoxis curtissii Rose—MM; 224

Hypoxis juncea Sm.—MF, 225

Iridaceae

Iris hexagona Walter—BM; 602

Sisyrinchium angustifolium Mill. —HH; 498

Juncaceae

Juncus effusus L. subsp. solutus (Fernald €: Wiegand) Hámet-

Ahti—BM; 174

Juncus marginatus Rostk.—DM; 206

Juncus megacephalus M.A. Curtis—DM; 781

Juncus dd Lam.—ME; 43

Marantacea

Thalia fru —BM; 401

Orchidaceae

Encyclia tampensis (Lindl.) Small—HH; 212

Habenaria floribunda Lindl.—MH; 489

Poaceae

Androgogon glomeratus (Walter) Britton et al. var. glaucopsis

(Elliott) C. Mohr—ME; 659

Andropogon glomeratus (Walter) Britton et al. var. pumilus

(Vasey) Vasey ex L.H. Dewey—MF; 331

AAndropogon longiberbis Hack.—RU; 286

Andropogon ternarius Michx.—MF; 447

^Andropogon virginicus L. var. glaucus Hack.—MF; 424

Andropogon virgincus L. var. virginicus—MF; 441

Aristida patula Chapm. ex Nash—HH; 34

Aristida purpurascens Poir. var. tenuispica (Hitchc) Allred—

431

Aristida spiciformis Elliott—MF; 355

Axonopus fissifolius (Raddi) Kuhlm.—HH; 289

Axonopus furcatus (Flüggé) Hitchc.—HH; 166

.—RU; 303

*Cynodon dactylon (L.) Pers.—RU; 202

id ic nd aegyptium (L.) Willd. ex Asch. & Schweinf.—

U; 340

A m aciculare (Desv. ex Poir) Gould & C.A. Clark—

H; 461

a) Eb fc Pal IA

J WOU

MH; 222

oe erectifolium (Nash) Gould & C.A. Clark—MF;

OLDER laxiflorum (Lam) Gould—-MH; 169

Franck and Wunderlin, Vascular flora of Churchill Ranch

Digitaria ciliaris (Retz.) Koeler—RU; 287

*Digitaria longiflora (Retz.) Pers.—RU; 288

Echinochloa muricata (P. Beauv.) Fernald—BM; 186

+Echinochloa paludigena Wiegand—

Echinochloa walteri (Pursh) A. Heller—DM; 192

*Eleusine indica (L.) Gaertn.—RU; 326

*Eragrostis atrovirens (Desf) Trin. ex Steud.—RU; 330

A*Eragrostis bahiensis (All) Vignolo ex Janch.—RU; 219

Eragrostis hypnoides (Lam.) Britton et al.—

Eragrostis spectabilis (Pursh) Steud.—MH; 223,449

Eragrostis virginica (Zuccagni) Steud.—RU; 655

Eremechloa ophiuroides (Munro) Hack.—RU; 271

Eustachys glauca Chapm.—HH; 229

Fastac peras (Sw) — po 302

) Nees—DM; 362

*Melinis repens (Willd. ) Zizka- AU 300

Muhlenbergia capillaris (Lam. Trin.—MH; 453

^Oplismenus hirtellus (L.) P. Beauv.—HH; 390

Panicum anceps Michx.—MF; 396

Panicum AN. ~ var. bartowense (Scribn. &

Merr.) Fernald—BM

Panicum hemitomon —ML; 185

Panicum hians Elliott—RU; 285

*Panicum repens L—-ML; 184

APanicum tenerum Be a Trin.—MF; 408

Panicum virgatum L— 18

APaspalum conjugatum : i Bergius—MF; 410, 459

Paspalum floridanum Michx.—HH; 231

*Paspalum notatum Flüggé var. saurae Parodi —RU; 237

adl ee MEI 0

Benth.) Hitchc.—DM;

272 458

Saccharum giganteum (Walter) a "d 407, 434

Sacciolepis striata (L) Nash—BM; 2

Schizachyrium scoparium (Michx.) us 370, 432

Setaria parviflora (Poir) Kerguélen—RU; 339, 382

Sorghastrum secundum (Elliott) Nash— u pm

Sporobolus Lu (Trin. one Kunth—

*Sporobolus indicu . Br. var. mau (P. Beauv.)

Tripsacum dactyloides (L.) L—HH; 663

A*Urochloa distachya (L) T.Q. Nguyen—RU; 376

Pontederiaceae

Pontederia cordata L.—ML; 660

Smilacaceae

Smilax auriculata ee 416

milax bona-nox L.—MH;

Typhaceae

Typha domingensis Pers.—FL; 101

Typha latifolia L.—BM; 194

Xyridaceae

Xyris caroliniana Walter—MF; 306

Xyris elliottii Chapm.—MF; 114, 269

345

DICOTS

Acanthaceae

ifolia (Mich Kuntze

MH L :265

+Justicia angusta (Chaprm ) Small—DM; 130

^Stenandrium dulce (Cav.) Nees—MMH; 196

Adoxaceae

Viburnum obovatum Walter—MH; 664

Amaranthaceae

*Alternanthera sessilis (L.) R. Br. ex DC.—HH; 622

*Chenopodium ambrosioides L.—RU; 360

*Gomphrena serrata L.—RU; 336

Anacardiaceae

Rhus cop —MF; 317

*Schinus terebinthifolia Raddi—MH; 175

Toxicodendron radicans (L.) Kuntze—MH; 142

Annonaceae

+Asimina reticulata Shuttlew. ex Chapm.—MEF; 611

Apiaceae

Eryngium baldwinii Spreng.—HH

Ptilimnium capillaceum (Michx.) Es eT 179

Apocynace

rds. DE Walter—MF; 261

Asclepias perennis Walter—HH; 129

fast

llex cassine L— 492

llex glabra (L.) A. E S 122

Araliaceae

Centella asiatica (L.) Urb.—FL; 442

Hydrocotyle umbellata L—FL; 200

Asteraceae

us Baba (Lam.) R.K. Jansen var. repens (Walter)

R.K. ; 620

Ambrosia an ia L—RU; 290

^Baccharis glomeruliflora Pers.—MH; 479

Baccharis halimifolia L—BM; 464

Bidens alba (L) DC.—RU; 242

Boltonia diffusa Elliott—ML; 281

+Carphephorus odoratissimus {J.F. Gmel.) H. Hebert var. sub-

tropicanus (DeLaney et al.) Wunderlin & B.F. Hansen—MF;

430

Chrysopsis mariana (L.) Elliott—HH; 470

^Conoclinium coelestinum (L) DC.—MMH; 143

Conyza canadensis (L.) Cronquist—RU; 338

+Coreopsis leavenworthii Torr. & A. Gray—DM; 134

Elephantopus elatus Bertol.—MF; 301

*Emilia fosbergii Nicolson—RU; 484

*Emilia sonchifolia (L) DC.——RU; 707

Erechtites hieraciifolius (L) Raf. ex DC.—MH; 235

Erigeron quercifolius Poir.—MH; 247

Erigeron vernus (L.) Torr. & A. Gray—MH; 346

346

Eupatorium capillifolium (Lam. Small ex Porter & Britton—

MH; 463

Eupatorium mohrii Greene—MF; 278

Eupatorium rotundifolium L—MF; 451

Euthamia caroliniana (L.) Greene ex Porter & Britton—MF; 429

Gamochaeta pensylvanica (Willd.) Cabrera—MH; 251

Helenium amarum (Raf) H. Rock—RU; 16

Helianthus agrestis Pollard —HH; 309

Heterotheca subaxillaris (Lam.) Britton & Rusby—RU; 809

Hieracium megacephalon Nash—xXH; 256

Alva microcephala Nutt.—DM; 353

Lygodesmia aphylla (Nutt) DC.—MEF; 618

Melanthera nivea (L) Smatl—HH; 329, 709

Mikania cordifolia (L. f) Willd.—MEF; 483

Mikania scandens (L.) Willd.—MH; 202

^Pectis glaucescens (Cass.) DJ. Keil —RU; 307

+Pectis linearifolia Urb.—RU; 366

Pectis prostrata Cav.—RU; 368

Pityopsis graminfolia (Michx.) Nutt.—XH; 327

Pluchea baccharis (Mill.) Pruski—HH; 246

Pluchea odorata (L.) Cass.—HH; 145

o d obtusifolium (L.) Hilliard € B.L. Burtt—

(RA)

n f A Elli dose

MH; 118

Dope. tortifolius (Michx) Nsescc ME 393

Solidago odora Aiton var. chapmanii (A. Gray) Cronquist —MF;

273

Solidago stricta Aiton—MMH; 481

"sphagneticola trilobata (L.) Pruski—MH; 348

Symphyotrichum bahamense (Britton) G.L. Nesom—HH; 480

Symphyotrichum carolinianum (Walter) Wunderlin & B.F.

Hansen—HH; 4

Symphyotrichum dumosum (L.) G.L. Nesom—HH; 478

*Tridax procumbens L.— been.

Verbesina virginica L—HH; 4

Vernonia blodgettii a 244

Bignoniaceae

Campsis radicans (L) Seernann—HH; 388

Boraginaceae

Heliotropium polyphyllum Lehm.—MF; 409

Brassicaceae

Rorippa teres (Michx.) Stuckey—HH; 603

Cactaceae

Opuntia humifusa (Raf) Raf.—XH; 328

Campanulac

+Campanula ee S. Watson ex A. Gray—HH; 621

+Lobelia feayana A. Gray— 6

+Lobelia homophylla E. Wimm. —MH; 139

Caryophyliaceae

Drymaria a (L.) Willd. ex Schult.—MH; 2

+Stipulicida setacea Michx. var. lacerata CW. Ja m 332

Chrysobalanaceae

Licania michauxii Prance—XH; 138

Cistacea

dM corymbosum Michx.—XH;

Lechea torreyi (Chapm.) Legg. ex Hon 471

Clusiaceae

Hypericum cistifolium Lam.—MH; 220

Hypericum fasciculatum Lam.—DM; 112

Hypericum gentianoides (L.) Britton et al.—MMH; 154

dn Loci (L) Crantz—MH; 245

Hypericu DM; 181

een E Pursh—MF; 105

Hypericum tetrapetalum Lam.—MH; 710

Convolvulaceae

Dichondra carolinensis Michx.—MMH; 493

Ipomoea cordatotriloba Dennst.—RU; 398

Ipomoea pandurata (L.) G. Mey—HMH; 633

Ipomoea sagittata Poir.—HH; 284

Cornaceae

Cornus foemina Mill.—HH; 248

Cucurbitaceae

Melothria pendula L.—HH; 310

Ebenaceae

Diospyros virginiana L.—MH; 708

Ericaceae

Bejaria racemosa Vent.—MF; 195

Lyonia fruticosa (Michx.) G.S. Torr.—MF; 104

Lyonia lucida (Lam.) K. Koch—ME; 119

Vaccinium arboreum Mee in 158

Vaccinium darrowii Camp—MF;

Vaccinium stamineum L—MF; 1 >

Euphorbiaceae

Acalypha gracilens A. Gray—

Chamaesyce hyssopifolia (L.) Ar E

Cha

amaesyce maculata (L.) Small—RU; 6

Cnidoscolus dcn piste. Lm 4 A. Gray—XH; 136

Croton glan

Croton din xii. E ic. m 203

*Phyllanthus Se Roxb.—MH; 488

*Ricinus communis L—RU; 297

Stillingia sylvatica L—MF; 108

abaceae

Amorpha herbacea Walter—XH; 1

o giant (L -) Sen a 254

ue rotundifolia J.F. Gmel Ru 110

*Crotalaria spectabilis Roth—RU; 6

Dalea carnea (Michx.) Poir.—MF; RE

mec incanum DC.—RU; 260

*Desmodium triflorum (L) DC.—RU; 377

Galactia elliottii Nutt—MF; 162

Galactia regularis (L.) Britton e E a 163

Indigofera caroliniana Mill—

ACIE tilium E : ub T 446

*Medicago lupulina L.—

AMimosa strigillosa Torr. ^ A an 147

*Senna obtusifolia (L) H.S. Irwin & Barneby—RU; 146

Franck and Wunderlin, Vascular flora of Churchill Ranch

Sesbania herbacea (Mill) McVaugh—BM; 341

ASesbania vesicaria Jacq.) Elliott —DM; 462

Vicia acutifolia Elliott—-HH; 601

d. I Michx.—HH; 325

Quercus minima (Sarg.) mE 103, 392

Quercus myrtifolia nee old 469

Quercus nigra L.—XH; 3

Quercus pumila eke ur 634

Quercus virginiana Mill —MH; 375

AQuercus laurifolia x Q. a F; 426, 711

Gentianaceae

Sabatia brevifolia Raf —MF; 365

Sabatia grandiflora (A. Gray) Small —FL; 236

Hydroleaceae

Hydrolea corymbosa J, Macbr. ex Elliott—ML; 298

amiaceae

d americana L.—MMH; 140

Hyptis alata (Raf) Shinners—HH; 315

Piloblephis rigida (W. Bartram ex Benth.) Raf. —MF; 421

Salvia lyrata L—MH; 12

Scutellaria e L.—MF; ag

ense L—RU; 2

lc) F pcm 151

Lauraceae

Cassytha filiformis L.—MMH; 266

Persea palustris (Raf) Sarg.—HH; 418

Loganiaceae

Mitreola petiolata (J.F. Gmel.) Torr. & A. Gray—HH; 226

Lythraceae

+Lythrum flagellare Shuttlew. ex Chapm.—DM; 117,615

Malvacea

Hibiscus pm Michx.—HH; 282

Kosteletzkya pentacarpos (L.) Ledeb—HH; 283

*Melochia corchorifolia L.—MWH; 316

Sida ulmifolia Mill.—RU; 438

*Urena lobata L.—MMH; 221

Melastomataceae

Rhexia mariana L—DM; 661

Myricaceae

Myrica cerifera L—MF; 216

MA apis

Nup hai V (Ait 1) Aiton—ML; 274

Oleaceae

Fraxinus caroliniana Mill—HH; 605

Onagraceae

Gaura angustifolia Michx.—RU; 240

Ludwigia maritima R.M. Harper—MF; 259, 455

*Ludwigia peruviana (L) H. Hara—ML; 657

^Ludwigia repens J.R. Forst.—DM; 171, 423

Ludwigia suffruticosa Walter—MF; 170

347

Orobanchaceae

Buchnera americana L—MF; 131

Oxalidaceae

Oxalis corniculata L.—RU; 153

Phytolaccaceae

Phytolacca americana L.—RU; 312

ela dea

copa monnieri (L) Pennell—DM; 1

e hispida (Benth. ex Lindl.) E RU; 155

Gratiola pilosa Michx.—XH; 6.

Linaria canadensis (L.) Chaz. —MF; 494

*Lindernia crustacea (L) F. Muell. —RU; 350

Lindernia grandiflora Nutt.—DM; 180

ecardonia acuminata (Walter Small var. peninsularis (Pen-

nell) Rossow—MF;

mos themum glomeratum (Chapm.) Shinners—HH; 385

Scoparia dulcis L.—MF; 250

Polygalac

Polygala neamata L. E 440

Polygala lutea L—

Polygala nana eru be —MF; 610

Polygala setacea Michx.—MF; 276

Polygala violacea Aubl.—RU; 109

Polygon

Poon p (Vent) Engelm. & A. Gray var. brachys-

a (Meisn.) Wunderlin—MF; 167

a glabrum Willd.—FL; 19

Polygonum punctatum Elliott—DM; BM; 176

Rosaceae

Rubus trivialis Michx.—RU; 495

Rubia

E al talis —DM; 183

Diodia teres Walter—RU; 2

Diodia virginiana a Ps 383

Galium tinctorium L.—FL; 60

Houstonia dia (J.F. ~ Standl—HH; 437

a repens L.—XH; 60

*Oldenla s ia m. T 357

Oldenlandia uniflora L.—MF; 417

Spermacoce remota Lam.—MH; 156, 395

Salica

i oe Michx.—DM; HH; 497, 653

Samolaceae

Samolus valerandi L. subsp. parviflorus (Raf) Hultén—DM;

Sapindaceae

Acer rubrum L.—MF; 389

Sapotaceae

Sideroxylon reclinatum Michx.—MF; 187

olanaceae

Physalis arenicola Kearney—XH; 320

348

Physalis pubescens L.—HH; 629

Solanum americanum Mill.—RU; 351

*Solanum viarum Dunal—HH; 623

£L Das *24l D

Journal Ol

Phyla nodiflora (L) Greene—DM; 120

Verbena scabra Vanl—HH; 141

Violaceae

Tetrachondraceae Viola sororia Willd.—MWH; 497

Polypremum procumbens L.—RU; 113 Viscacedë

Ulmaceae Phoradendron leucarpum (Raf) Reveal 8: M.C. Johnst.—MF;

Ulmus americana L.—HH; 387 381

Urtica Vitaceae

Ampelopsis arborea (L.) Koehne—MF; 172

Parthenocissus quinquefolia (L) Planch.—MF; 352

Vitis aestivalis Michx.—MH; 343

Vitis rotundifolia Michx.— MF; 135

Vitis shuttleworthii House—HH; 201

Hind m (L) Sw.—DM; 263

Parietaria floridana Nutt.—HH; 604

Verbenaceae

+Glandularia tampensis (Nash) Small—HH; 632

ACKNOWLEDGMENTS

We thank Bruce F. Hansen and John M. Kunzer for their i bl j with identification. We would

also like to thank Bruce F. Hansen and two anonymous reviewers for their comments on the manuscript,

and Kathleen Hotchkiss for her help with graphics. We thank the Land Manager, Debbie Blanco, of Sarasota

County, for access to the site. The senior author thanks his wife for her company on some of his collecting trips.

REFERENCES

FDEP (FLORIDA ON OF ON TA Protection). 2007. Florida Geological Survey - Hydrogeology Section.

http:// geology/programs/ hydrogeology/geographic_info_sys.htm Accessed September

9, 2008.

FLEPPC (Fiorina Exotic Pest PLant Council). 2007. List of Florida's invasive plant species. Florida Exotic Pest Plant

Council. http:// fleppc.org/07list.htrn Accessed June 15, 2008

FNAI (FLORIDA NATURAL AREAS INVENTORY). 1990. Guide to the natural communities of Florida. Tallahassee, FL.

HaLroro, KJ. anp D. Yossi. 2006. Estimating hydraulic properties using a moving-model approach and multiple

aquifer tests. Ground Water 44: Al

Hureman, J.M. 2006. Historical fi tl

Hype, A.G., G.W. Hurt, AND C.A. WETISTEN. 1991. Soil survey of Sarasota County, Florida. USDA, Soil Conservation

Service, Washington, D.C.

NCDC (Nationat Cumatic DATA CENTER). 2008. http://

Ph.D. dissertation, | State University.

dc.noaa.gov/oa/ mpp/freedata.html Accessed August

SWEWMD (Southwest FLonIDA Water MANAGEMENT District). 2008. Rainfall summaries: county totals. http://www.

swfwmd.state fl.us/data/wmdbweb/rainsum.htm Accessed August 6, 2008

UF (University or Floripa). 2004. Florida Aerial Photography. htto://www.uflib.ufl.edu/digital/ collections/flap/

Accessed September 4, 2008.

USDA (Unrrep States DEPARTMENT OF AGRICULTURE). 2007. Web soil survey. http://websoilsurvey.nrc

WebSoilSurvey.aspx Accessed September 5, 2008.

Watts, A., G. TANNER, AND R. Dye. 2006. Restoration of dry prairie using fire and roller chopping. Florida Land of

Fire and Water: The Florida Dry Prairie Ecosystem. Proceedings of the Florida Dry Prairie Conference, ed. R.F.

Noss. Pp. 225-230

WUNDERLIN, R.P. AND B.F. Hansen. 2003. Guide to the vascular plants of Florida, 2"4 ed. University Press of Florida,

Gainesville.

WUNDERLIN, R.P. AND B.F. HANSEN. 2008. Atlas of Florida vascular plants (http:// | | fedu/). [S.M. Landry

and K.N. Campbell (application development), Florida cn mind Design and Research.] Institute

for Systematic Botany, University of South Florida, Tampa.

gov/app/

VASCULAR FLORA OF A SALINE PRAIRIE IN WINN PARISH, LOUISIANA

Barbara R. MacRoberts Christopher S. Reid

and Michael H. MacRoberts and Patricia L. Faulkner

Bog Research, 740 Columbia Louisiana Natural Heritage Program

Shreveport, Louisiana 71104, USA. Louisiana Der f Wildlife and Fist

Herbarium, M PO. Box 98000 Baton Rouge, Louisiana 70898-9000, U.S.A.

Louisiana State University-Shreveport

Shreveport, Louisiana 71115, U.S.A.

ABSTRACT

We describe the mro ih of a saline pee in Winn Parish, da The prairi ins 59 species of which th federal

and/or state h in similariti her W lf 1 Plain

plant communities, notably OEN outcrops and barrens.

RESUMEN

Se d ibe la fl lar d prad li Winn Parish, Louisiana. La prad ti 59 especies de las q

en el estado. Mi que las prad h :dagd dif, + j itg

vegetales de la llanura costera del West Gulf, notabl fl i d i y tierras yermas.

INTRODUCTION

Recent research on saline prairies in Louisiana has greatly expanded our knowledge of this community (Reid

et al. in prep). This community occurs in central and northwestern Louisiana, eastern Texas, and southern

Arkansas (McInnis et al. 1993; Keith et al. 2004; Lester et al. 2005; Arkansas Natural Heritage Commission

2006; Diggs et al. 2006; Reid et al. in prep). In pre-settlement times, saline prairies were probably uncom-

mon. The estimate for Louisiana is fewer than 800 ha with only 10 to 25 percent remaining (Lester et al.

2005). Because of many factors, including urban sprawl, agriculture, grazing, oil exploration, off-road vehicle

activity, and agro-forestry, this community has been badly degraded and greatly reduced in extent and is

now ranked globally imperiled (GDL ester et al. 2005).

The earliest described sali in Louisiana are several in Winn Parish (McInnis et al. 1993; Lester

et al. 2005). Interest in the Winn Parish saline prairies initially focused on the federally-listed Geocarpon

minimum, which had not previously been found in Louisiana (McInnis et al. 1993).

A renewed interest in saline prairies since 2005 (MacRoberts et al. 2007; Reid et al. 2007; Reid et al. in

prep) and in Geocarpon minimum (MacRoberts & MacRoberts 2007, 2008) led to an interest in the floristic

and edaphic conditions of the Winn Parish saline prairies, which had not previously been thoroughly de-

scribed (McInnis et al. 1993).

n this paper we describe the floristics of Upper Weyerhaeuser Prairie (aka Castor Creek Saline Prairie

[McInnis et al. 1993]), one of the Winn Parish saline prairies.

General edaphic and community Mano A regarding E prairies including Upper Weyerhaeuser

Prairie as well as an extensive bibliography y and p is given in Reid et al. (in prep).

STUDY SITE AND METHODS

Figure 1 is an aerial view of Upper Weyerhaeuser Prairie (TIONRIES2). Figure 2 is a scenic view of the prairie.

The prairie measures approximately 3.9 ha. A road runs through its center with shallow ditches along its

sides. We excluded the road and the ditches from the survey. We also avoided prairie edges, staying at least

three meters from the edge. Upper Weyerhaeuser Prairie, unlike many other saline prairies, has no mima

J. Bot. Res, Inst. Texas 3(1): 349 — 354. 2009

350 | | | | | | Journalol t ical i Texas 3(

Fic. 1. Aerial view of Upper Weyerhaeuser Prairie.

(pimple) mounds or wet depressions. The soils are classified as Brimstone fine-silty, siliceous, thermic Glossic

Natraqualfs. They are high in sodium and are alkaline. They resist wetting, inducing droughty conditions.

Prairie substrait has physical properties similar to a dense clay hardpan that is resistant to root penetration

(McInnis et al. 1993; Boyd et al. 1998; Reid et al. in prep.). Two lichens, Cladonia caroliniana (Schwein.) Tuck.

and Cladonia sobolescens Nyl. ex Vain., are a common ground cover in some areas of the prairie (Fig. 3). The

surrounding forest is dominated by Pinus taeda, Quercus stellata, Quercus phellos, Ilex vomitoria, Crataegus

spathulata, and Vaccinium spp., with Chasmanthium laxum being the dominant understory species.

Upper Weyerl Prairie as a whole was collected monthly between 19 March and 5 November 2008.

Specimens are deposited at the Louisiana State University Herbarium (LSU) and Louisiana State University

in Shreveport Herbarium (LSUS). Nomenclature follows USDA NRCS (2008), Diggs et al. 2006), and Flora

of North America (1993-2006).

In order to investigate species richness, two 0.1 ha plots each with two nested 0.001 ha plots and two

nested 0.0001 ha plots were established within the prairie. All species in the plots were recorded.

RESULTS

Table 1 lists the vascular species found in Upper Weyerhaeuser Prairie. All collection numbers refer to B.R.

MacRoberts and M.H. MacRoberts. An asterisk * indicates an exotic species. Table 2 summarizes the plot data.

Upper Weyerhaeuser Prairie had 59 species. The two 0.1 ha plots had a mean of 50 species (range 47

to 53), the four 0.001 ha plots had a mean of 19.5 species (range 15 to 28) and the four 0.0001 ha plots had

a mean of 16.5 species (range 9 to 26). Only one exotic species was found in the prairie.

MacRoberts et al., Vascular flora of a saline prairie

3 ; ZO i

ES gor ant A. re

> de VAN d

i E y Le j E

Fic. 3. Lichens in Upper Weyerhaeuser Prairie.

DISCUSSION

Rare species at Upper Weyerl Prairie include G ini (G2S2), Schoenolirion wrightii (G3S2),

and Phemeranthus patios (G5S3) (Louisiana Natural Menige Program 2008). Geocarpon minimum was

first discovered at this prairie in 1991 when approximately 620 plants were found in four locations (un-

published field notes on file with the Louisiana Natural Heritage Program). Fewer plants were observed

in 1992, but the survey was less intensive. Surveys in 2007 and 2008 discovered the species at only one

location. Total number of plants was not determined but four 0.0001 ha plots had 141 and 357 plants in

those two years, respectively (MacRoberts & MacRoberts 2007, 2008). Geocarpon minimum is now known

from Caddo, DeSoto, and Winn parishes in Louisiana (MacRoberts & MacRoberts 2007, 2008). The pres-

ence of Schoenolirion wrightii in Upper Weyerhaeuser Prairie adds Winn Parish to the list of Caddo, Red

River, Natchitoches, De Soto parishes where it had previously been found (MacRoberts & MacRoberts 1993;

Louisiana Natural Heritage Program 2008). In Louisiana, Phemeranthus parviflorus is known from Caddo,

De Soto, Natchitoches, Red River, Sabine, and Winn parishes (Louisiana Natural Heritage Program 2008).

Both Schoenolirion wrightii and Phemeranthus parviflorus are known from both saline prairies and sandstone

outcrops in Louisiana (MacRoberts & MacRoberts 1993; Louisiana Natural Heritage Program 2008); whereas

Geocarpon minimum is known only from saline prairies.

Upper Weyerhaeuser Prairie is relatively uniform. The two 0.1 ha plots contained all of the species

found in the prairie although the plots constituted only about five percent of the prairie. Further, it was clear

from repeatedly walking the entire prairie that virtually the same species occurred throughout.

Compared with saline prairies in northwestern Louisiana (Reid et al. in prep), Upper Mi uini

prairie is relatively depauperate. It lacks mima mounds and wet depressi ] thus plant sq

with them. But also it lacks many other species found in the northwestern Louisiana saline prairies such as

Cooperia drummondii, Evax verna, Gratiola flava, Lechea san-sabeana, Marshallia caespitosa, Minuartia drum-

mondii, Minuartia muscorum, Opuntia humifusa, Phacelia glabra, and ae raza pile ie et a in i piep ^

A7 1

Nonetheless, floristically Upper \ Prairie most

prairies in De Soto and Caddo pushes (Reid et al. in prep.). We found that approximately 69 percent of

the Upper Weyerhaeuser flora occurred in them. Upper Weyerhaeuser next most resembles sandstone out-

crop communities on the Kisatchie Ranger District of the Kisatchie National Forest in Natchitoches Parish,

Louisiana, with approximately 50 percent of its flora occurring in them (MacRoberts & MacRoberts 1995).

Species in common between them include Bigelowia nuttallii, Evolvulus sericeus, Phemeranthus parviflorus,

Polygala verticillata, and Schoenolirion wrightii. Next in community resemblance is Black Branch Barrens on

the Angelina National Forest in Jasper County, Texas, which has just below 50 percent of the Upper Weyer-

haeuser flora (Marietta & Nixon 1983; Orzell 1990; MacRoberts et al. 1997). Interesting plants in common

between Black Branch Barrens and Upper Weyerhaeuser Prairie are Bigelowia nuttallii, Evolvulus sericeus,

Phemeranthus parviflorus, and Schoenolirion wrightii.

ACKNOWLEDGMENTS

This work was partly supported by contract CFM663087, Louisiana Department of Wildlife and Fisheries.

Alan Boyd, Land Use Forester, at Weyerhaeuser Company, aided with the research. Shirley Tucker identified

the lichens. Charles Allen made several valuable comments on the paper.

MacRoberts et al., Vascular flora of a saline prairie

TABLE 1. Species of Upper Weyerhaeuser Saline Prairie.

353

Acanthaceae

iue i Nutt., 8254

Alliac

w pM L., 8160, 8167

Asteraceae

Bigelowia nuttallii L.C. Anderson, 8313

lva angustifolia Nutt. ex DC., 8331

Krigia dandelion Nutt., 8154

Krigia occidentalis Nutt., 8129

Liatris pycnostachya Michx., 831 o 8326, 8329

Senecio tomentosus Michx.,

Solidago nitida Torr & A. Gray, 83 m 8330

Symphyotrichum dumosum (L.) Nesom, 8362

Brassicaceae

Lepidium densifl Schrad., 8168

Campanulaceae

Triodanis perfoliata L., 8158

Caryophyllaceae

Geocarpon minimum Mack, (no specimen

Cistaceae

ie HIE Michx, 8251

Clusia

seine aaa (Greve. €: Hook.) T.&G., 8243, 8311

Convolvulaceae

ties sericeus Sw. 8153, 8244

Cyper

Carex Ms PE. E & Hook., 8166, 8169

Fimbristylis puberula (Michx.) Vahl, 8163, 8246

Rhynchospora globularis (Chapm.) Small, 8240, 8150, 8252

Rhynchospora pusilla Chapm. ex M.A. Curtis, 8241

roseraceae

Drosera brevifolia Pursh, 8135

ricaceae

Vaccinium arboreum Marshall, 8308

Euphorbiaceae

Croton wildenowii G.L. Webster, 8310, 8332

Fabaceae

Neptunia lutea (Leavenw.) Benth., 8250

Gentianaceae

Sabatia const Nutt., 8242

Hyacinthaceae

steno ME

,8172,8131

Hypoxi

Hypoxis m T 8156

lridaceae

*Sisyrinchium rosulatum Bicknell, 8161

Sisyrinchium sagittiferum Bicknell, 8164

Lamiaceae

ipe hispida Pursh, 8165, 8247

Liliacea

atero bivalve (L.) Britt., 8132

aceae

us dd (Planch.) Britton, 8255

Onagra

i m Nutt., 8155

Orchidaceae

Spiranthes vernalis Engelm & A. Gray, 8249

alidaceae

Oxalis corniculata L., 8134

Pinaceae

Pinus taeda L., 8126

Plantagi

ginaceae

Plantago pusilla Nutt., 8133

Plantago virginica L., 8157

Agrostis elliottiana Schult., 8149, 8162

Aristida longespica Poir., 8365, 8382

Aristida oligantha Michx., 8369, 8381

Dichanthelium dichotomum (L.) Gould, 8170

Dichanthelium sphaerocarpon (Ell.) Gould, 8171, 8248

Eragrostis hirsuta (Michx.) Nees, 8322, 8360

E refracta (Muhl.) pl 8323, 8333, 8361, 8364

aspa setaceum Michx. 8

Schizac ee scoparium Re Nash, 8371

Tridens strictus ce Nash, 8366, 8368, 8383

Polygalac

aum. mariana Mill, 8239

Polygala verticillata L., 8245

Portulacaceae

diu parviflorus (Nutt.) Kiger, 8309

Rosa

Conte marshallii Egglest., 8173, 8174

Rubia

Ded teres eee 8312

oustonia micrantha ub. Terrell, 8127

furia 24 Schoepf, 81

otac

apo

Seon lanuginosum Michx, 8151

ceae

a: tenuifolia (Vahl) Raf, 8367, 8370

354 Journal of t ical Insti Texas 3(

Taste 2. Species richness in Upper Weyerhaeuser Prairie.

Plot and plot No. of Average species

size (ha) plots (range)

Plot A

0.0001 2 20.0(14-26)

0.001 2 22.5(17-28)

0.1 1 53.0

Plot B

0.0001 2 13.0(9-17)

0.001 2 16.5(15-18)

0.1 1 470

REFERENCES

ARKANSAS NATURAL HERITAGE COMMISSION. 2006. Warren Prairie Natural Area. www.naturalheritage.com

Boro, W.H., G.L. BowptN, AND M.J. BORDELON. 1998, Soil survey of Winn Parish. USDA Soil Service, Washington, D.C.

Dices, G.M., B. L. Liescome, M.D. Reep, AND R.J. O'KeNNON. 2006. Illustrated flora of east Texas. Sida, Bot. Misc. 26:1—1594.

FLORA OF NORTH America. 1993-2006. Edited by Flora of North America Editorial committee.Oxford Univ. Press,

New York

Kerm., E.L., J.R. SINGHURST, AND S. Cook. 2004. Geocarpon minimum (Caryophyllaceae), new to Texas. Sida 21:1165-

1169

Lester, G.D., S.G Sorensen, P.L. FAULKNER, C.S. REID, AND I.E. Maxit. 2005. Louisiana comprehensive wildlife conservation

strategy. Louisiana Department of Wildlife and Fisheries. Baton Rouge.

Louisiana NATURAL HERITAGE PROGRAM. 2008. Rare plant species of Louisiana. Louisiana Department of Wildlife and

Fisheries. Baton Rouge.

MacRoserts, M.H. AND B.R. MacRoserts. 1993. Vascular flora of sandstone outcrop communities in western Louisiana

with notes on rare and noteworthy species. Phytologia 75:463-480.

MacRoserts, M.H., B.R. MacRoserTs, AND R.E. Evans. 1997. Notes on Spiranthes parksii Correll (Orchidaceae) in deep

east Texas. Phytologia 83:133-137.

MacRoserts, M.H. AND B.R. MacRoserts 2007. Survey for pon minimum in northwest Louisiana. Unpublished

report. Louisiana Department of Wildlife and Fisheries, Baton Rouge, dn

MacRoserts, M.H. AND B.R. MacRoserTs 2008. Data collection for í Unpublished report. Louisiana

Department of Wildlife and Fisheries, Baton Rouge, Louisiana.

MacRoseats, M.H., B.R. MacRoseats, C.S. Rep, PL. FAULKNER, AND D. Estes. 2007. Minuartia drummondii (Caryophyllaceae)

and Gratiola flava (Plantaginaceae) rediscovered in Louisiana and Gratiola flava historically in Arkansas. J. Bot.

Res. Inst. Texas 1:763-767.

Marietta, K.L. AND ES. Nixon. 1984. Vegetation of an open, prairie-like community in eastern Texas. Texas J. Sci.

3625-32.

McInnis, N.C., L.M. SmitH, AND AB. Pirman. 1993. Geocarpon minimum (Caryophyllaceae), new to Louisiana. Phy-

tologia 75:159-162.

Orzett, S.L. 1990. Texas Natural Heritage Program inventory of National Forests and National Grasslands in Texas.

Texas Parks and Wildlife Department, Austin.

Remp, C.S., PL. FAuLKNer, B.R. MAcRoBERTs, AND M.H. MacRosenrs. 2007. Saxifrag (Saxifrag ) to Louisiana.

J. Bot. Res. Inst. Texas 1:1251-1252.

Rep, C.S., PL. FAULKNER, B.R. MacRosenrs, M.H. MACROBERTS, AND M. BORDELON. In prep. Characteristics and floristics of

saline prairies in Louisiana.

USDA, NRCS, 2008. The PLANTS Database (http://plants.usda). National Plant Data Center, Baton Rouge, LA

70874-4490 USA.

VASCULAR FLORA OF MORSE CLAY PRAIRIES IN NORTHWESTERN LOUISIANA

Barbara R. MacRoberts Christopher S. Reid

and Michael H. MacRoberts and Patricia L. Faulkner

Bog Research, 740 Columbia Louisiana Natural Heritage Program

a ua 71 104, idi Louisiana Der f Wildlife and Fish

Herbarit O. Box 98000 Baton Rouge, Louisiana 70898-9000, U.S.A.

A State University- "c

Shreveport, Louisiana 71115, U.

ABSTRACT

We describe the floristics, soils, smal!-scale species ricl 1 phytogeog ffinities of M Clay calcareous prairies in north-

western Louisiana. Four 0.1 ha plots had 159 sp ] 185 i eiat 0.0 l ged 27 species, and eight 0.0001

ha plots averaged 19.5 es. The most diverse families were ee (40 species), pr (23 species), and Poaceae (17 species).

The soils are y alkaline high-calcium silty clay loams. hos North American distribution of Morse Clay prairie species is

eastern and central. Whether Morse Clay prairies should b id distinct y type will ad on bcd research on

isolated prairies in nearby Texas, Arkansas, and Louisiana, but at the p i hey do not appear t least some

of the calcareous prairies in the region.

RESUMEN

Se describe la florística, zem riqueza de especies a pequeria escala, y afinidades fitogeográficas de las prad ] de Morse

Clay en el noroeste de L ana. oe fu B 1 ha tuvieron 159 especies y lia de 85 especies, och las de 0.001 ha

tuvieron una media de 27 especi 00 dia de 19.5 especies. Las familias con más diversidad

Tueton ub nd dpt Faces @ a di y Poaceae ve oe Los ae son p P ME manes oap

es oriental y central.

F 7 ly

Si las praderas Morse Clay deben, o no id ipo d idad diferente dependerá de i g p p

eras asiladas en las cercanas Texas, Arkansas, y encore p tualmente no parecen ser diferentes de al menos algunas praderas

calcáreas de la región.

INTRODUCTION

While prairies are one of the most endangered plant communities in North America, they are one of the best

studied (Sims & Risser 2000; Diggs et al. 2006). Thousands of books and papers have been written on the

floristics, management, and restoration of prairies. Louisiana prairies were documented in the eighteenth

and nineteenth centuries by land surveyors, cartographers, and explorers; however, botanists did not be-

gin to study them until the 1930s (Brown 1997; MacRoberts et al. 1997, 2003; MacRoberts & MacRoberts

2004 and references therein). Southeastern prairies, except for the prairie belt of Alabama and Mississippi,

have been woefully neglected in the literature (Leidolf & McDaniel 1998; Peacock & Schauwecker 2003;

Barone 2005; Barone & Hill 2007). However, such isolated prairies were extensive in pre-European times,

and there still exist many small, relatively natural prairies in the pineywoods region of Texas, Arkansas,

Louisiana, and Mississippi (Jordan 1973; Brown et al. 2002; Foti et al. 2003; Zollner et al. 2003; MacRoberts

& MacRoberts 2004).

Included in this group are the *Morse Clay" prairies of northwestern Louisiana and southwestern

Arkansas (McInnis 1997; Lester et al. 2005; Arkansas Natural Heritage Commission pers. comm.). These

are small, grassy openings that occur on reddish, fine textured, alkaline and calcareous clays and silt loams

(Chaffin et al. 1959). While the fire frequency was not different from surrounding woodland, these prairies,

like so many others, are maintained as openings in the oak-pine woodlands by soil-dependent variations

in grass biomass, resulting differences in fire intensity, and extreme soil properties (Diggs & Schulze 2003;

Lester et al. 2005; Diggs et al. 2006). Herbivory by large mammals may also play a role.

The purpose of this paper is to describe the floristics of Morse Clay prairies in northwestern Louisiana

and to compare them with prairies elsewhere in the southeast.

J. Bot. Res. Inst. Texas 3(1): 355 — 366. 2009

I Lattkhn D H ID Kl A pd. f T,

356 t Texas 3(1)

THE SETTING

Morse Clay prairies and soils occur on Pleist j liate t j tl d central Louisiana

and southwestern Arkansas. Prairies typically exist as small openings on side-slopes and flats in shortleaf pine/

oak-hickory forests (Figs. 1 and 2) (Lester et al. 2005). P 1 extent of Morse Clay prairies is estimated

to have been between 800 and 4,000 ha, with 5 to10 percent thought to remain (Louisiana Natural Heritage

Program 1993). A recent status survey of Louisiana Morse Clay prairies revealed 11 occurrences totaling a

mere 60 ha, and nearly half of that area was in poor condition (MacRoberts & MacRoberts 2007b).

The best known examples of Morse Clay prairies are found on the 9,000 ha Barksdale Air Force Base

and 14,000 ha Bodcau Wildlife Management Area in Bossier Parish, Louisiana. Barksdale AFB was purchased

in the late ae The western side is ci as an air Ie base; the eastern side is undeveloped and

pp g oak-pine woodlands with occasional [ ings. Prior to its establishment as an air

base, | le timber harvesting had eliminated all uu Dabe. When the Federal Government acquired

the DER it was generally open fields and cropland (McInnis 1997). Current management of the eastern part

is focused on wildlife, oil and gas exploitation, timber harvesting, and recreation. Consequently, few areas

remain in natural or semi-natural condition. All of the prairie area has been subject to disturbance, notably

fire suppression, food plot development with plowing, and part of it was at one time a bombing range with

large craters. Teague and Wendt (1994) and McInnis (1997) give an overview of the entire area.

Bodcau WMA, owned by the U.S. Army Corps of Engineers and leased by the Louisiana Department of

Wildlife and Fisheries, has similar habitat to that of Barksdale AFB with similar areas of Morse Clay prairie

(Lester et al. 2005). At Bodcau by the mid-1950s, prairies were cotton farmed and over-grazed by livestock

(John Haygood, pers. comm.). Some of the prairie area in Bodcau WMA is protected in registered natural

areas, but much of the area that was historically prairie is now plowed land managed for food plots. All areas

where Morse Clay prairie occurs have had ground disturbance in the past.

METHODS

In order to have comparable samples, we modified the plot design of Peet et al. (1998) used widely in the

southeast. We established four study plots in four prairies away from prairie edges (Fig. 3). Three plots were

established in closely adjacent prairies on Barksdale AFB, and one plot was established on a single prairie

on Bodcau WMA 32 km to the northeast. Each plot measured 20 m x 50 m or 31.6 m x 31.6 m (0.1 ha).

Within each 0.1 ha plot were two nested 3.16 m x 3.16 m (0.001 ha) plots. Each of these in turn had a 1 m

x 1 m (0.0001 ha) nested plot within it. Thus, there were four 0.1 ha plots, eight 0.001 ha plots, and eight

0.0001 ha plots. We surveyed each plot for total species. All species in the plots were recorded every month

over a year between 2006 and 2008 (except in mid-winter). A specimen of all species collected is deposited

at Louisiana State University Herbarium (LSU). Nomenclature follows Kartesz and Meacham (2005) and

Flora of North America (1993-2006)

Using our list of species found in the four Morse Clay prairies and data from Flora of North America

(1993-2006) and NatureServe (2008), we plotted the North American distribution of species by state, region,

or province to determine the geographic pattern of Morse Clay prairie species (Fig. 4).

We collected soil samples from the upper 20 cm within the prairies, in the woodlands above the prairies,

and in the woodlands below them at about 3 meters from the prairie edge. These we had analyzed for pH,

various elements, and soil texture at Louisiana State University Soil Testing and Plant Analysis Laboratory.

We made brief floristi veys of other Morse Clay prairies in Caddo and Bossier parishes (MacRoberts

& MacRoberts 2007b).

RESULTS/DISCUSSION

The four study plots had 159 species: North Pond had 105, Lower Green Tree had 85, Bomb Crater had 75,

and Nemastylis had 76 (Table 1). Asteraceae, Fabaceae, and Poaceae dominated with 40, 23, and 17 species,

respectively. The plot richness (Table 2) is about average for open plant communities (e.g., bogs and xeric

MacRoberts et al., Vascular flora of Morse Clay prairies 357

sandylands) throughout the West Gulf Coastal Plain with about 20 species per 0.0001 ha, 27 species per

0.001 ha, and 85 species per 0.1 ha) (MacRoberts et al. 2007). Table 3 gives Sorenson's Index of Similarity

among the four prairies. This level of similarity indicates that they are the same community. Soil texture

in the prairies was either silty clay loam or silty clay (Table 4). Soil textures of upslope woodlands ranged

from silty clay loam to very fine sandy loam. The range of pH value in the prairies was 6.8 to 8.3, n = 7. By

contrast, pH values ranged from 4.6 to 5.6, n = 4 in the woods above the prairie, and 5.2 to 74, n = 3in the

woods below prairies. Calcium concentration was substantially higher in prairie soils versus soils collected

from adjacent woodlands. Potassium concentration also seems to generally be higher in prairie soils.

We found five state rare species in the prairies. Carex meadii (S253) is found in several west Louisiana

parishes where it occurs in calcareous and coastal prairies and in sandstone glades. It occurred in or near

all of the study plots. Coreopsis palmata Nutt. (S1), found only in Bossier and Claiborne parishes, occurred

only in Bomb Crater Prairie, and only at its upper edge. At Bomb Crater, Coreopsis palmata appeared to be

associated mainly with upland woodlands. Houstonia purpurea var. calycosa (S2) occurs in Bossier, Caldwell,

Grant, LaSalle, and Winn parishes in calcareous prairies. This species was common in North Pond Prairie

but was not found at the other Morse Clay prairies. Nemastylis geminiflora (S2S3) is found in several West

Gulf Coastal Plain parishes. It was found in the Lower Green Tree and Nemastylis prairie plots and in the

prairies adjacent to both the North Pond and the Bomb Crater prairie plots. Koeleria macrantha Ledeb. (S1)

(Reid 5546 LSU) occurred in Bomb Crater Prairie but just outside of the plot. This is one of the only two

extant populations in Louisiana, the other being in a calcareous prairie in Winn Parish. Further informa-

tion on rare species and the explanation of "S" ranks can be found in Louisiana Natural Heritage Program

Q007)

Isolated prairies occur across the West Gulf Coastal Plain and eastward onini & c dd us n

MacRoberts & MacRoberts 2004). While few prairies have been the subject of

enough have been done to provide quantitative comparisons. Brown et al. (2002) studied Windham Prainie

in Polk County, Texas, 250 km southwest of Bossier Parish. An Index of Similarity (Sorenson's) between

Windham Prairie and our Morse Clay prairies is 37. Leidolf and McDaniel (1998) studied Sixteen Section

Prairie in Oktibbeha County, Mississippi, about 450 km east of Bossier Parish. An Index of Similarity be-

tween it and our Morse Clay prairies is also 37. Barone and Hill (2007) studied several blackland prairies in

Alabama and Mississippi about 500 km east of our study sites. An Index of Similarity between those sites and

the Morse Clay prairies is 31. Contrasted with these figures, the Keiffer prairies in Winn Parish, Louisiana

(Smith et al. 1989; MacRoberts & MacRoberts 1996), 125 km southeast of the Morse Clay prairies, have an

Index of Similarity of 51 with the Morse Clay prairies. This figure is especially interesting since North Pond

Prairie on the Bodcau WMA, only 32 km northeast of the three Barksdale AFB prairies, differed from them

as much as did the Keiffer prairies (Table 3). Unfortunately, there are no complete floristic i ies of the

blackland and Morse Clay prairies of southwestern Arkansas. However, Foti (1989) provides a preliminary

list of species in ten of the less disturbed blackland prairies approximately 120 km north of our Morse

Clay prairies. An Index of Similarity between those sites and our Morse Clay prairies is 44 a difference that

indicates the importance of additional study of southwestern Arkansas prairies, notably since such species

as Carex microdonta Torr. & Hook. and Spiranthes magnicamporum C. Sheviak have recently been found in

them (Theo Witsell pers. comm.).

Morse clay soils proved similar to soils of other calcareous prairies in Louisiana, Arkansas, and Texas

which are alkaline and are high in calcium. Of the properties tested for all prairies, phosphorus and potas-

sium were the same; only magnesium was higher (MacRoberts & MacRoberts 1996, 2004

Judging habitat condition is difficult, notably when there is no known "natural" habitat for comparison.

Unfortunately, this is the case for Morse Clay prairies, none of which are pristine; all have been subject to

various unnatural disturbances, notably clearing, grazing, plowing, fire suppression, and bombing. One can

only attempt to reconstruct what these prairies were like by studying those surviving remnants that most

closely resemble our best estimate of the structure and composition of presettlement plant communities.

Journal of the Botanical Research Institute of Texas 3(1)

A A qí _— M Az 42=<= OAK

Fic. 1. Morse Clay Prairie.

Fis. 2. Aerial photograph of Morse Clay Prairie.

MacRoberts et al., Vascular flora of Morse Clay prairies 359

ARKANSAS :

a | E

1. BOSSIER A

y )

Vicinity Map

à North Pond >o. 7]

Location of Prairies

Fic. 3 Louisiana. Unlabeled d I : Pee eee Y Clay y r^ EPUM

surveyed. Labeled d ha lacatinn nf tt : :

While it is clear that the best remnants of this prairie type occur on Barksdale AFB and Bodcau WMA (McIn-

nis 1997), these sites can scarcely be described as *relatively undisturbed." Thus, our best estimate becomes

largely subjective based on our experience with other prairies in the West Gulf Coastal Plain (including

other Morse Clay prairies surveyed as part of this study but not reported here [MacRoberts & MacRoberts

2007b]). Assessing such characteristics as native biotic diversity and presence/absence of exotics (McInnis

1997), we determined that the four Morse Clay prairies we studied had 10 (6%) exotics, which indicates

360 ical Insti Texas 3(

hi v sae Numb f It in state. province, or region. Bossier and

I | I Y g

Fic. 4

Caddo parishes,

thal 4 CAA rl

little disturbance (MacRoberts et al. 2008). The Barksdale AFB and Bodcau WMA prairies are therefore

important because they present a standard by which the quality of other Morse Clay prairies may be evalu-

ated, but care should be exercised in making general conclusions from these few scattered remnants, all of.

which have undergone a long history of anthropogenic disturbance.

The North American geographic distribution of Morse Clay Prairie species is decidedly eastern. However,

a comparison of their distribution with that of Caddo Parish flora as a whole (MacRoberts & MacRoberts

20072) sl their affiniti lightly more central and western than are those of Caddo Parish (Caddo

Parish is just west of Bossier Parish and also has Morse Clay prairies, see Fig. 3). This might be expected

since there are many habitats in Caddo Parish that are exclusively eastern (e.g., swamps), whereas prairies

generally have a central and western component.

ACKNOWLEDGMENTS

This work was partly supported by contract 64332 from the Louisiana Department of Wildlife and Fisher-

ies. Bruce Holland, Natural Resources Manager, Barksdale AFB, facilitated access to the study sites. We also

thank Suzanne Odom, Forester, U.S. Army Corps of Engineers, for permission to work on Bodcau WMA

and informing us about the management history of Bodcau. We thank Richard McMullen, formerly of

Louisiana Department of Wildlife and Fisheries, for taking us out several times on Bodcau WMA to show

MacRoberts et al., Vascular flora of Morse Clay prairies

361

Taste 1. Vascular flora of four Morse clay prairies (NP = North Pond Prairie, LGT = Lower Green Tree Prairie, BC = Bomb Crater

Prairie, and N = Nemastylis Prairie. * = exotic species

Acanthac

thaceae

Ruellia humilis Nutt., NP, LGT, BC, N, MM 7780, 7698, 7515,

7484

Agavaceae

Manfreda virginica (L.) Rose, NP, LGT, N, MM 7477

Yucca louisianensis Trel, LGT, MM 7857

nacardiaceae

Toxicodendron radicans (L.) Kuntze, NP, LGT, BC (no specimen

collected)

Apiaceae

all era ee BC, N, MM 7540

x DC, NB LGT, MM 7781

Aquifoliaceae

llex decidua Walt., NP, LGT, BC, MM 7512

Araceae

Sabal minor se Pers., NP, MM 7504

Ps cea

ASC. illata L., NP LGT, BC, N, MM 7786, 7560, 7961

hn viridis Walt, NP, MM 7689, 7562

Asteraceae

llected)

eo pole L, LGT, N (n

Ambrosia bidentata Michx., NP, MM 7523

Ambrosia n DC., NP MM 7522

Ambrosia trifida L, LGT, MM 7556

Arnoglossum plantagineum Raf, NP, LGT, BC, N, MM 7690,

7713

Baccharis halimifolia L., NP, LGT, BC, N, MM 7509, 7531

Boltonia diffusa Ell, N, MM 7965

Cirsium horridulum Michx., NP, LGT, BC, MM 7685, 7712

Coreopsis lanceolata L, NP, LGT, BC, N, MM 7783, 7688, 7718

Coreposis palmata Nutt, BC, MM 7793, 7845

Echinacea pallida (Nutt.) Nutt., NP, LGT, MM 7691

Echinacea sanguinea Nutt., BC, MM 7791, 7789

Erigeron strigosus Muhl. ex Willd., NP, LGT, BC, N, MM 7703,

7707, 7479, 7716

Erigeron tenuis Torr. & A. Gray, N, MM 7630

Eupatorium serotinum Michx., BC, N, MM 7536

Eurybia hemispherica Alexander) Nesom, BC, N, MM 7530

Gaillardia aestivalis (Walter) H. Rock, LGT, MM 7480

Helianthus hirsutus Raf., LGT, BC, N, MM 7794, 7846, 7858

Heterotheca subaxillaris (Lam.) Britt. & Rusby, LGT, MM 7558

Iva annua L., NP LGT, BC, N, 7557, 7840

Liatris aspera Michx., NP, LGT, BC, N, MM 7500, 7553, 7485,

8054

id aid (L) Michx., BC, MM 7454

ex DC, LGT, N, MM 7715

Parthenium integrifolium L, NP | 6 T, BC, N, MM 7460, 7853

Rudbeckia hirta L., LGT, N, MM 7859

Solidago nitida Torr. & A. Gray, NP, BC, N, MM 7498

Solidago rigida L., LGT, BC, N, MM 7557, 7849, 7841

Symphyotrichum Arammondii (Lindl) Neto LGT, MM 7472

Symphyotrichum laevis (L.) A. & D. Lóve, LGT, BC, N, MM 7478,

7572, /838, 8057, 8115

M lateriflorum (L.) Lóve & Lóve, NP, LGT, MM

Sm cm ontarione (Weigand) Nesom, LGT, MM

575, 7576

C. $ sab

,LGT, BC, N,

iti i 7966

(Ait) N , NP. LGT, BC, N, MM 7499,

7482, 7546, 8116

dc aad pilosum (Willd.) Nesom, NP. LGT, BC, N, MM

i mm praealtum (Poir) Nesom, NP, N, MM 7570

Symphyotrichum pratense (Raf) Nesom, BC, N, 7962, 8114

Vernonia texana (A. Gray) Small, NP, LGT, MM 7492

Bignoniaceae

Campsis deant .) Seem. ex Bureau, LGT, N, MM 7837

Boragin

Onasmadum bare DC. ex A. DC., NP MM 7519

mpanulac

pis "perdu A.DC., NP (no specimen collected)

Caprifolia

dd japonica Thumb, BC, MM 7535

, BG, MM 7527

Viburnum duum Raf, NP, BC, MM 7508

Convolvulace

Dichondra cain Michx., LGT, MM 7722

Corna

Cornus od CA. Mey. BC, MM 7532

Cupressaceae

Juniperus virginiana L., LGT, MM 7580

Cuscutaceae

Cuscuta ee Engelm., N, MM 7839, 8110

Cype

pa abscondit Mack, NP, n 7702

P LGT, BC, N, MM 7636, 7624

e rex meadii Dewey, NP, LGT, N MM 7632, 7622, 8136

Fimbristylis puberula (Michx.) Vahl, BC, MM 7708

Scleria ciliata Michx., NP, MM 7779b

Scleria oligantha Michx., NP, MM 7779a

Ebenaceae

Diospyros vi

Euphorbia

EE nutans (Lag.) Small, LGT, MM 7559b, 7582

Croton monanthogynus Michx, NP, LGT, N, MM 7520, 7469,

, NB. LGT, BC, N, MM 7503, 7534

7964

Euphorbia corollata L., LGT (no specimen collected)

Tragia urticifolia Michx., NP. LGT, BC, N, MM 7488, 7709

Fabaceae

Acacia angustissima (Mill) Kuntze, NP, LGT, BC, N, MM 7470

m ~ Torr. & A. Gray, NP, LGT, N, MM 7628, 7571,

soria oe Muhl. ex Elliott, NP, LGT, BC, N, MM 7629,

362

Centrosema virginianum (L.) Benth., LGT, MM 7854

Cercis canadensis L., BC, MM 7526

Chamaecrista fasciculata (Michx.) Greene, NP MM 7494

Dalea candida Willd., NP, BC, N, MM 7777, 7792

Dalea purpurea Vent., NP, LGT, BC, N, MM 7456

Desmanthes illinoensis (Michx.) MacM. ex B.L. Robins & Fern.,

T, BC, N,

Galactia volubilis (L.) Britt, NP, LGT, BC, N, MM 7851

Gleditsia triacanthos L, NP (no specimen collected)

espedeza cuneata (Dum. Cours.) G. Don, NP, BC, MM 7493

*Medicago lupulina L, NP, MM 7785

Mimosa nuttallii (DC.) B.L. Turner, N, MM 8183

Mimosa strigillosa Tort. & A. Gray, NP, LGT, BC, N, MM 7496

"o lutea (Leavenw.) Benth, NP LGT, BC, N, MM 7516,

molum, dubium Sibth., NP, MM 7693

Fagac

ae tlt Wangenh., NP, BC, MM 7843

Gentianacea

*Centaurium pu Re (Sw.) Druce, = MM 7852

a campestris Nutt, NP, N, MM 7

Iridac

Md geminiflora Nutt., LGT, N, MM 7619

Sisyrinchium albidum Raf., LGT, BC, MM 7620

iud lanlo Greene, NP MM 7695, 7633

acea

w m L., NB. LGT, BC, N, MM 7687, 7717

Pycnanthemum tenuifolium Schrad., BC, MM 7529

Salvia azurea Michx. ex Lam., LGT, BC, N, MM 7550, 7863,

8113

Salvia lyrata L., NP, LGT, BC, N, MM 7635, 7623

Liliaceae

Allium canadense L., NP, BC, N, MM 7699, 7706

Nothoscordum bivalve (L.) Britt, NP, LGT, MM 7637, 7626

Lythraceae

Lythrum alatum Pursh, BC, LGT, NP, N, MM 7856

Moraceae

Maclura pomifera (Raf) Schenid., NP MM 7501

Oleaceae

Fraxinus pennsylvanica Marsh, NP, LGT, BC, N, MM 7510, 7533

Onagraceae

Oenothera oe Nutt., NP. N, MM 7701

Orchidac

Fiir loci (Small) Ames, NP, MM 7788

Oxalidac

ie s L, NP MM 7638, 7700

Pinaceae

a E Miil, NB MM 7514

Poac

fare gerardii Vitman, NP, LGT, BC, N, MM 7545, 7537,

7466

Andropogon glomeratus (Walt.) B.S.P, NP, LGT, BC, N, MM

7538

Andropogon virginicus L, NP. LGT, MM 7567

Aristida oligantha Michx, NP, MM 7568

*Botriochloa ischaemum L., NP, MM 7583

Bothriochloa longipaniculata (Gould) Allred & Gould, LGT,

MM 7861, 7487

M n (Desv. ex Poir.) Gould & Clark, LGT, BC,

7719, 7710, 7694

5c Hs L., NP, LGT, BC, MM 7506, 7473

Eragrostis lugens Nees, N, MM 8056, 8112, 7963

Eragrostis spectabilis (Pursh) Steud., P 5 N, MM 7542

id ani zu anos Michx., NP, BC, MM

Aichx., NP LG a ji m 7544

‘Paspalum dilatatum Poir., LGT, sd alae 7586

Paspalum laeve Michx., NP, MM 7

a scoparium DAE m NP. LGT, BC, N, MM

7541

sen carr (Poir) Kerguelen, LGT, BC, N, MM 7474,

a -— (Poir) Roem. €: Schult, NP MM 7565

icd nutans (L) Nash, LGT, BC, N, MM 7539, 8117,

Sterol a (Michx) Scribn., NP LGT, BC, N, MM

moe asper (Beauv.) Kunth, NP (no specimen col-

lected

Sporobolus compositus (Poir.) Merr, NP, LGT, MM 7566, 7574,

Tridens strictus (Nutt) Nash, NP MM 7569

ica O ) L, NP, MM 7865

Polem

Phlox posa! hi RP MM 7696

Ranuncu

Amenone eee Pritz, NP, MM 7631

Delphinium carolinianum Walt., NP, MM 7784

Rhamnaceae -

Berchemia scandens (Hill) K. Koch, NP, LGT, BC, N, MM 7513

Rosaceae

Crataegus brachyacantha Sarg. & Engelm., NP, MM 7502,

7495

Crataegus marshallii Egglest., NP (no specimen collected)

Crataegus spathulata Michx. NP, MM 7505

Rosa carolina L, NP, MM 7490, 7778

Rosa setigera Michx., BC, N, MM 7790, 7850

Rubus O Michx., NP, LGT, N, MM 7634

Rubiac

nsus Ionen: (Lam. Fosberg, LGT, BC, N, MM 7547,

7476

Houstonia micrantha (Skinner) Terrell, NP (no specimen col-

lected)

Houstonia purpurea L. var. calycosa A. Gray, NP, MM 7497, 7697

Sapotaceae

Sideroxylon lanuginosum Michx., NP, LGT, BC, N, MM 7511,

7483

Scrophulariaceae

Agalinis heterophylla (Nutt.) Small, NP, LGT, MM 7525, 7555

Agalinis tenuifolia (Vahl) Shinners, NP, N, MM 7563, 8111

Aureolaria grandiflora (Benth.) Pennell, BC, MM 7467

Pedicularis canadensis L., BC (no specimens collected)

Penstemon laxiflorus Pennell, NP, BC, N, (no specimens col-

lected)

Penstemon tubaeflorus Nutt, N, MM 8182, 7686, 7705

MacRoberts et al., Vascular flora of Morse Clay prairies 363

Verbena

Smilaceae aceae

a canadensis (L.) Nutt, NP. LGT, BC, N, MM 7627,

Smilax bona-nox L., NP. LGT, BC, N, MM 7518, 7475

Ulm

Tuo UE (L) Michx., NP MM 7787, 7491

*Verbena brasiliensis Velloso, NP MM 7521

Verbena halei Small, NP, N, MM 7866

Ulmus alata Michx., NP. LGT, MM 7507

Ulmus crassifolia Nutt., LGT, BC, N, MM 7528

Valerianaceae

Valerianella radiata (L.) Dufr., NP, MM 7692 itaceae

Ampelopsis arborea (L.) Koehne, LGT, BC, N, MM 7860

Taste 2. Species richness in Morse Clay Prairies.

Plot and plot No. of Average species

size (ha) (range)

North Pond Prairie

0.000 2 2100-22)

0.001 2 30.5(28-33)

0.1 1 105

Lower Green Tree Prairie

1 2 2201-23)

0.001 2 28.5(28-29)

0.1 1 85

Bomb Crater Prairie

0.0001 2 11.5(5-18)

0.001 2 18(1 7-19)

0.1 1 75

Nemastylis Prairie

0,000 2 23.5(21-26)

0.001 2 30.5(25-36)

0.1 1 76

Total

0.0001 8 19.5(5-26)

0.001 8 26.9(17-36)

0.1 4 85.3(75-105)

(S A Morse Clay prairies (NP — North Pond Prairie, LGT — Lower Green Tree Prairie, BC — Bomb Crater Prairie, and N = Nemastylis

LE3.I

nt Formula is QUA t B) x 100 where C= number af species in common between two samples, A = total species in sample A, and B = total species in sample B.

NP LGT BC

NP > 7 -

LGT 56 - -

BC 49 61 -

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MacRoberts et al., Vascular flora of Morse Clay prairies 365

us prairies and share with us his knowledge of the plant life of the area. John Haygood, formerly a Louisiana

Department of Wildlife and Fisheries biologist, provided information on the condition of Bodcau prairies

in the 1950s. Bob O'Kennon, Botanical Research Institute of Texas, Guy Nesom, and Charles Allen aided

with plan: identification. Theo Witsell, Arkansas Natural Heritage Commission, provided information on

the Arkansas Morse Clay prairies. Latimore Smith, Dennis Bell, and an anonymous reviewer made useful

suggestions that improved the manuscript.

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BARONE, J.A. AND J.G. HiLL. 2007. Herbaceous flora of Blackland prairi tsin Mississippiand western Alabama.

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Brown, C.A. 1997. Preliminary report on the isolated prairies of Louisiana. Proc. Louisiana Acad. Sci. 60:10-19.

BROWN, L.E., K. HiLLHOUsE, B.R. MAcRoserts, AND M.H. MacRoserTs. 2002. The vascular flora of Windham Prairie, Polk

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CHAFFIN, B.F, J.L. MILLET, J.R., ScaLr, J.A. DEMENT, B.J. Gnirris, R.H. JORDAN, S.A. Lyre, B.F. GRAFTON, AND E.F. YOUNG. 1959. Soil

survey of Bossier Parish, Louisiana. USDA and Louisiana nent oe ea an Rouge.

Dices, G.M. AND PC. ScHuLtze. 2003. Soil-dependent fire frequency: a new hy for the distribution of prairies

and woodlands in north central and east Texas. Sida 20:1139-1153.

Dices, G.M., B.L. Liescome, M.D. REED, AND R.J. O’KENNON, 2006. Illustrated flora of east Texas. Sida, Bot. Misc. 26:1-1594.

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4, 5, 19, 20, 21, 22, 23, 24, 25, 26. Oxford Univ. Press, New York and Oxford.

Fon, T.L. 1989. Blackland prairies of southwestern Arkansas. Proc. Arkansas Acad. Sci. 43:23028.

Fon, T.L., S. SIMON, D. ZOLLNER, AND M. HATTENBACH. 2003. Blackland prairie landscapes of southwestern Arkansas. In:

Peacock, E. and T. Schauwecker, eds. 2003. Blackland prairies of the Gulf Coastal Plain. University of Alabama

Press, Tuscaloosa. Pp. 94-109

JORDAN, T.G. 1973. Pioneer evaluation of vegetation in frontier Texas. SouthW. Hist. Quart. 76:233-254.

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Garden. Chapel Hill.

Leipotr, A. AND S. McDanteL. 1998. A floristic study of black prairie plant communities at sixteen section prairie,

Oktibbeha County, Mississippi. Castanea 63:51-62.

Lester, G.D., S.G. SORENSEN, P.L. FAULKNER, C.S. REID, AND I.E. Maxit. 2005. Louisian | i ildlif vation

strategy. Unpublished report. Louisiana Department of Wildlife and Fisheries, Baton Rouge.

LOUISIANA NATURAL HERITAGE PROGRAM. 1993. Estimated presettlement and current acres of natural plant communities

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Louisiana Natura HERMTAGE PROGRAM. 2007. Rare plant species of Louisiana. Unpublished Report. Louisiana Natural

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tion. Cambridge Univ. Press, New York. Pp. 324—356.

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The Nature Conservancy, Baton Rouge.

ZoLuner, D., S. SIMON, AND T.L. For. 2003. A plant community classification for Arkansas's blackland prairie ecosys-

tem. In: Peacock, E. and T. Schauwecker, eds. 2003. Blackland prairies of the Gulf Coastal Plain. University of

Alabama Press, Tuscaloosa. Pp. 110-145

FLORISTICS OF UPLAND SHORTLEAF PINE/OAK-HICKORY FOREST IN

NORTHWESTERN LOUISIANA

Barbara R. MacRoberts and Michael H. MacRoberts

zs Research and Herbarium

40 Columbia useum of Life Sciences

nn. bed 71104, U.S.A. Louisiana State University in Shreveport

Shreveport, Louisiana 71115, U.S.A.

ABSTRACT

Th t t f 1 A cl 1 Poot f, A 1 A c 1 f. "n 1. 1:1 F 1 d

I I p y historically in the pepe West

Gulf Coastal Plain is poorly understocd. This is primarily | fi d d before 1 were able t

describe it. What little is | inly d is derived largely en historia information. The mo layer is even

Wed iL 1 1 1 1 rf 1 1 1

more poorly understood. y p y Louisiana ind study

maya oF PASS 1 PEE 1 1 "E cc A

Į y y represent what naturally f YI p

RESUMEN

La estructura del bosque de pino de hoj ta y Dusk nd de p i "e corta Hoe NUN que scd ette en la

zona alta de la llanura costera del V

los botánicos y ecólogos poe re lo poro que se sabe se centra principalmente e en los arboles y se reels ea nae

end dE El el p

roble g T te de Louisiana. Ell A PH fcri 1 há 1 1

a = og i

e £ r r r vt

1 : : 1 an

r r

INTRODUCTION

Virtually no upland shortleaf pine/oak-hickory forest survived late nineteenth and early twentieth century

logging and little is known about this ecosystem, which apparently was once common in the Upper West

Gulf Coastal Plain (southern Arkansas, southeastern Oklahoma, northern Louisiana, and northeastern Texas)

(Williams & Smith 1995; Carr 2000; Bragg 2002, 2003, 2008; Diggs et al. 2006; MacRoberts & MacRoberts

2008a, 2008b; Masters 2008 and references therein). Historical accounts and photographs indicate that

shortleaf pine often dominated and was sometimes found in pure or near pure stands on drier and fire-

prone upland sites, but that more often it occurred in association with oak and hickory (Bragg 2002, 2008;

Lester et al. 2005). Many stands appeared to be open—“park like”—with a rich “high-light” community of

herbaceous plants (Bragg 2002), but like most forest descriptions of the nineteenth and twentieth centuries,

the herbaceous layer was ignored and certainly never described in detail (Gilliam 2007). Consequently, not

only is little known about the over-story of these forests but even less is known about the herbaceous layer

(Carr 2000; Bragg 2002, 2008). The destruction of the shortleaf pine/oak-hickory forest community was

so complete, not only through logging but fire suppression and agriculture, that an understanding of this

community depends almost entirely on historical documents, notably land survey records and forester's de-

scriptions that emphasize trees (but not always species of trees)(Bragg 2002, 2003, 2004, 2008; MacRoberts

& MacRoberts 2005; Nowacki & Abrams 2008; see also Diggs et al. 2006; Van Kley 2006).

In the course of our work on another project on Barksdale Air Force Base, Bossier Parish, in northwestern

Louisiana, we found several areas where shortleaf pine was the dominant canopy species, where fire had

been used as a management tool, and where the herbaceous layer appeared to be intact.

We took the opportunity to study the flora of one of these sites, recognizing that it was not pristine

but that perhaps it was as close to “natural” as now exists. Our main interest was the herbaceous layer. In

the few places where shortleaf pine/oak-hickory forest exists, the understory has been radically altered by

J. Bot. Res. Inst. Texas 3(1): 367 — 374. 2009

368 tani Insti Texas 3(1)

fire suppression: fire intolerant and shade-adapted species have replaced the naturally occurring pyrogenic

heliophytes (Bragg 2002; Gilliam 2007; Nowacki & Abrams 2008).

STUDY SITE

The study area is located on Barksdale Air Force Base, Bossier Parish, Louisiana, (T17NR12WS2). Natural

Areas on Barksdale AFB have been described by McInnis (1997). Land plat records from the 1830s show that

the study area was, in general, a pine-oak-hickory forest (McInnis 1997; Lester et al. 2005). While the area

has been repeatedly logged, it is still oak-pine-hickory today but is predominantly shortleaf pine. Canopy

trees on the study plots today are only shortleaf pine (Fig. 1). In the mid 1990s, the area's pine was thinned

from approximately a basal area of 70 to about 45. Removal of pine increased hardwood growth, which was

subsequently removed by helicopter application of the herbicide Arsenal. The area is currently controlled

burned on a two-to-three year rotation with prescribed fire in the early months (generally January-February)

of the year. The area was last burned in February 2008. Soils are the Sacul series, typically moderately well-

drained, acid fine sandy loam, thermic Aquic Hapludults, with a clay underlayer along the ridgetops in the

uplands (Kilpatrick et al. 1990).

METHODS

To obtain a species list and to determine species richness, we established two 0.1 ha plots, each with two

nested 0.001 ha plots and two nested 0.0001 ha plots about 100 m apart on ridge tops (Fig. 1). All species

in the plots were recorded and most species in the plots were collected. Voucher specimens are deposited

at the Louisiana State University in Shreveport Herbarium LSUS). We visited the site monthly between 2

April 2008 and 21 October 2008.

To determine the geographical distribution of species in the sample, we established a grid of 114 km to

a side across the West Gulf Coastal Plain and, using Turner et al. (2003), Thomas and Allen (1993-1998),

and USDA NRCS (2008), plotted the distribution of species in the Barksdale AFB shortleaf pine/oak-hickory

forest study plots.

Soil samples were coll 1 from the upper 30 cm in the center of each 0.1 ha plot. These were analyzed

by the Soil Testing and Plant Analysis Laboratory, Louisiana State University, Baton Rouge.

RESULTS

The flora of the 0.2 ha study area (two 0.1 ha plots) is given in Table 1. Table 2 summarizes the species rich-

ness data from the plots. Table 3 gives the soil sample results. Figure 2 gives the results of the geographical

distribution analysis.

e two 0.1 ha plots contained 122 species. Taxa from the Asteraceae, Fabaceae, and Poaceae domi-

nated and accounted for 46 percent of the flora. The mean number of species in the two 0.1 ha plots was

99.5 species (92 and 107), the mean number in the four 0.001ha plots was 30.25 species (range 28 to 33)

and the mean number in the four 0.0001 ha plots was 17.5 species (range 15 to 19). The soils have low pH

(5.2 and 5.4 in two samples) and are generally low in nutrients (see Table 3). The data in Figure 2 indicate

that the species found in the study plots are not specific to any particular region of the West Gulf Coastal

Plain but occur throughout the area.

DISCUSSION

1-1

The Barksdale AFB shortleaf pine/oak-hickory forest flora i ly similar to upland longleaf pine forest/

savanna in central Louisiana and southeastern Texas (see Harcombe et al. 1993; Van Kley 1999a, 1999b and

especially Carr 2000 and references therein). O etal. (1999) listed ne species groups identified during

multivariate analysis to be important in disti ies in the longleaf pine ecoregion of

Texas and Louisiana. These species are abundan co iss occur consistently within a community, but are

specialized enough to be present in some communities but absent or markedly less abundant in others. We

MacRoberts and MacRoberts, Floristics of upland shortleaf pine/oak-hickory forest

Ar Y

Fic. 1. Barksdale Air Force Base shortleaf pine/oak-hickory forest study site.

370 I Ingel Dat sa D a. ee en f Texas 3(1)

^ y ES

Fi; 2 Pi + £Darlcedala AED el n] f pi Inak hu yf ly pl I : : hn UAL ener. IDin: The grids are 114

km on a side.

found that our shortleaf pine/oak-hickory forest plots had 71 percent of the species in their “Schizachyrium

group,” which characterizes the upland longleaf pine forest/savanna. This similarity between Barksdale

AFB shortleaf pine/oak-hickory forest and particularly upland longleaf pine forest/savanna is reinforced by

the plants listed by the Louisiana Natural Heritage Program (2008) as typifying western upland longleaf

pine forest: 63 percent occurred in the Barksdale AFB plots. Turning to more detailed studies, 53 percent

of the Barksdale AFB shortleaf pine/oak-hickory forest species occur in four 0.1 ha upland longleaf pine

forest/savanna plots in central Louisiana on the Kisatchie National Forest (MacRoberts et al. 2002), and

60 percent of the Barksdale AFB shortleaf pine/oak-hickory forest species occurred in eight 0.1 ha upland

longleaf pine savanna/forest plots in eastern Texas on the Sabine National Forest (Philipps et al. 2007). An

hortleaf pine/oak-hickory forest flora and the flora of

Index of Similarity TR between the Barksdale

upland longleaf pine f in both the Kisatchie National Forest in central Louisiana and the Sabine

National Forest in eastern Texas Mn et al. e Philipps et al. 2007) gives figures of 46.4 and

46.5, respectively. Considering the diff ksdale AFB sample, 122 species; Kisatchie

hickory forest 371

Tage 1. Species in the two 0.1 ha study plots. N

Acanthaceae

Ruellia pedunculata Torr. & A. Gray, 8186

diaceae

Rhus a L, 831

cala radicans (L) Kuntze

Aristolochiaceae

a LI Jacq.

Asclepiadac

Asclepias pes L., 8185

Asteraceae

Ambrosia artemisiifolia L., 8295, 8354

Antennaria parlinii Fern., 8146, 8221

Baccharis halimifolia L., 8265

Conyza canadensis (L.) Cronquist, 8340

Coreopsis lanceolata L., 8195

Erigeron strigosus Muhl. ex Willd., 8187

Eupatorium perfoliatum L., 8374

Eupatorium rotundifolium L., 8286

Eurybia hemispherica (Alexander) Nesom, 8338, 8359

Euthamia leptocephala (T. & G.) Greene 8336

Gamochaeta purpurea (L) Cabrera, 8208, 8222

Helianthus hirsutus Raf., 8262, 8277

Hieracium gronovii

actuca T

Liatris pycnostachya Michx.

Liatris squarrosa (L.) Michx., 8287

Pityopsis graminifolia (Michx.) Nutt., 8278,

Pseu rdi MN ) e a m 8349

Rudbeckia hirta

Solidago ue y ie

Solidago odora Aiton, 8291

Solidago petiolaris Aiton, 8356, 8376

Solidago radula L., 8317, 8357

Symphytorichum | dumosum (L.) Nesom

Symphyotrichum lateriflorum (L) A. Love & D. Lóve, 8380

Symphyotrichum patens sae Nesom,

Verbesina helianthoides Michx.,

Vernonia texana (A. Gray) Small ps

ampanulaceae

Lobelia appendiculata A. DC., 8193

Piel eae (L) Nieuwl.

om ic mE Michx., 8290

Clusiaceae

Hypericum drummondii (Grev. & Hook.) T. & G., 8355

Hypericum hypericoides (L.) Crantz

Commelinaceae

Commelina erecta L.

Tradescantia hirsutiflora Bush, 8142, 8190

Cyperaceae

Carex complanata Torr. & Hook., 8207, 8211

Cyperus echinatus (L.) Wood, 8202, 8213, 8272

Rhynchospora sides W. ta 8274

Scleria es Michx 3

Dennstedtiac

Pteridium ean (L.) Kuhn, 8141

Dioscoreaceae

Dioscorea pne L, 8137

Ebenaceae

SSA virginiana L., 8305, 8298

Ericace

a arboreum Marshall, 8276

Euphorbiaceae

Acalypha monococca (Engelm. ex A. Gray) L. Mill. Gandhi,

82,829

Chamaesyce nutans (Laq.) Small, 8341

Euphorbia corollata L., 8145, 8189

abaceae

Baptisia nuttalliana Small, 8197

Centrosema ge de Benth., 8260, 8321

Crotalaria sagittalis L

Desmodium ciliare ri ex Willd.) D.C, 8263, 8280

Lespedeza hirta (L) Hornem.

Mimosa nuttallii (DC) B.L. Turner, 8269

Stylosanthes biflora (L.) Britton, Sterns, & Poggenb., 8198

Tephrosia virginiana (L.) Pers., 8279

Vicia caroliniana Walt., 8149

agaceae

Quercus falcata Michx., 8373

Quercus marilandica Münchh.

Quercus phellos L., 8292

Quercus dde Mang, 8372

Hamamelid

uidanbar sac L., 8266

ridac

Sommehumc iud Raf, 8140

Juglandac

Carya sp., Er

Juncaceae

Juncus dudleyi Wiegand, 8212

Luzula bulbosa (AW. Wood) Smyth, 8147

Lamiaceae

Monarda fistulosa L., 8267

Salvia azurea Lam.,

Salvia lyrata L., 8139

Lauraceae

Sassafras albidum (Nutt.) Nees

Liliaceae

Allium canadense L., 8188

Loganiaceae

B sempervirens (L.) St. Hil., 8304

Oleacea

China een L, 8296

xalidac

son más Jacq. 8191

Oxalis a 8144, 8210

Pinace

Pinus uem P Mill, 8281

372

Taste 1. (continued)

Plantaginaceae

ca virginica L., 8200

Poac

ai el (Walt) Britton, Sterns, € Poggenb., 8201,

8218

Andropogon gerardii Vitman, 8315

Andropogon ternarius Michx.

Aristida purpurascens Poir., 8350, 8351, 8375

Chasmanthium sessiliflorum (Poir) Yates

828

Dichanthelium aciculare (Desv. ex Poir.) Gould & Clark, n 7

Dichanthelium dichotomum (L.) Gould, 8204, 8215, 82

Dichanthelium ravenelii (Scribn. & Merr.) Gould, rn

PE Sopa (Lam.) Gou ld, ins

CIHO UDI

Schizachyrium scoparium (Michx.) Nash

Sphenophoiis obtusata pera Scribn., 8205, 8220

Tridens flavus (L.) Hitch., 83

Tridens strictus (Nutt.) we lus 8346

Polemoniaceae

Rosa

os spathulata Michx.

Rubus argutus Li

Rubus og Michx., 8306

Rubia

peca teres e 8337

Galium obtusum Bigelow, 8196

Galium pilosurn Aiton, 8257

Houstonia E Schóepf

Scrophu

Agalinis pco (Vahl) Shinners, 8353

Aureolaria grandiflora ue : Pennell, 8335

Pedicularis canadensis L., 8148, 8192

Penstemon laxiflorus a 8285

Smilacaceae

Smilax bon

o 8261, ~ 8320

Smilax = Morong, 83

Ulmace

Ulmus Michx., 8300, 8378

Verbenaceae

PAD americana L., 8268

Phlox pilosa L, 8184 Violac

Polygalacea Viola po L, 8138

Polygala e L., 8199 Vitaceae

Portulacaceae Parthenocissus quinquefolia (L.) Planch.

aa virginica L., 8143 Vitis rotundifolia Michx., 8264

National Forest sample, 158 species; Sabine National Forest sample, 196 species) these figures indicate that

all three are probably the same or very similar communities. Van Kley and Welch (2003), in a study using

multivariate ordination and classification of 420 sites across the West Gulf Coastal Plain including shortleaf

pine/oak-hickory forest and upland longleaf pine forest/savanna, found that plant communities are very

similar throughout the region. Carr (2000) found that shortleaf pine/oak-hickory forest resembled pine

savanna elsewhere in the southeast United States and pointed out that southeastern pine savanna commu-

nities may not be most appropriately defined by dominant overstory species; rather, the hundreds of other

plant species present may provide a much better designation of community types. The conclusions of both

Carr and Van Kley are reinforced by our finding that none of the species in our shortleaf pine/oak-hickory

forest study site is restricted to the Upper West Gulf Coastal Plain.

Other similarities between upland longleaf pine forest/savanna and shortleaf pine/oak-hickory forest

are also interesting. The three families that dominate the Barksdale AFB shortleaf pine/oak-hickory forest

study area—Asteraceae, Fabaceae, and Poaceae—also dominate the upland longleaf pine forest/savanna in

both the Kisatchie National Forest and the Sabine National Forest and account for 43 percent and 42 percent

of the flora, respectively (Barksdale AFB shortleaf forest is 46 percent) (MacRoberts et al. 2002; Philipps et

al. 2007). Also, shortleaf pine/oak-hickory forest species richness is similar to upland longleaf pine forest/

savanna in both the Kisatchie National Forest, which averaged 100 species (range 82-113 species) in four

0.1 ha plots (MacRoberts et al. 2002), and the Sabine National Forest, which averaged 82 species (range

71-112 species) in eight 0.1 ha plots, 29 species (range 17-46) in sixteen 0.001 ha plots, and 12 species

(range 5—25) insixteen 0.0001 ha plots. Finally, the soils in upland longleaf pine f / d shortleaf

pine/oak-hickory forest are similar (MacRoberts et al. 2002).

RA Dal " ARA Dag P" Planes: £ J 3 -L al fpi " L hiel yf 373

Taste 2. Species richness in the two plots.

Plot and plot No. of plots Mean species

size (ha) (range)

Plot A

0.0001 2 16.5(15-18)

0.001 2 30.0(30-30)

0.1 1 107.0

Plot B

0.0001 2 18.5(18-19)

0.001 2 30.5(28-33)

0.1 1 920

Taste 3. Soil sample results for the two piots.

Sample (ppm) pH Calcium Copper Magnesium Phosphorous

Plot A 5.35 275.05 0.27 72.80 4,83

Plot B 5.24 222.69 0.20 45.63 25.01

Table 3 continued from right

Sample (ppm) Potassium Sodium Sulfur Zinc Soil Texture

Plot A 73.93 5.78 5.26 0.74 sandy loam

Plot B 38.48 474 4.86 0.65 fine sandy loam

ACKNOWLEDGMENTS

Bruce Holland, Natural Resources Manager, Barksdale AFB, and Matthew Stroupe, Forester, Barksdale AFB,

aided with the study. Charles Allen and Barney Lipscomb made several comments that aided the paper.

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ed. Freeman and Custis Red River expedition of 1806: two hundred years later. Bull. Mus. Life Sci. 14:55-79.

Masters, R.E. 2008. Shortleaf pine community description. Forest Encyclopedia Network. www.forestryencyclo-

pedia.net

Mclnnis, N.C. 1997. Barksdale Air Force Base—threatened and endangered species - natural areas survey. The

Nature Conservancy, Baton Rouge, Louisiana.

Nowacki, G.J. AND M.D. Asrams. 2008. The demise of fire and “mesophication’ of forests in the eastern United States.

BioScience 58:1 23-138.

Phiips, T.C., S.B. Waker, B.R. MacRoserTs, AND M.H. MacRoserts. 2007. Vascular flora of a longleaf pine upland in

Sabine County, Texas. Phytologia 89:31 7-338.

Thomas, R.D. AND C.M. ALLEN, 1993-1998. Atlas of the vascular flora of Louisiana. Louisiana Department of Wildlife

and Fisheries, Baton Rouge.

Turner, B.L., H. NicHoLs, G. Denny, AND O. Doron. 2003. Atlas of the vascular plants of Texas. Sida, Bot. Misc. 24:1-

888.

Turner, R.L., JE. Van Kuey, L.S. SMITH, AND RE. Evans. 1999. Ecological classification system for the National Forests and

adjacent areas of the West Gulf Coastal Plain. The Nature Conservancy, Nacogdoches, Texas.

USDA NRCS. 2008. The PLANTS Database. (http://plants.usda) National Plant Database Center, Baton Rouge.

Van Key, J.E. 1999a. The vegetation of the Kisatchie Sandstone Hills, Louisiana. Castanea 64:64-80.

Van Key, J.E. 1999b. The vegetation of the High Rolling Uplands, Louisiana. Castanea 64:318-336.

Van Kiev, J.E. AND M.L. WeLch. 2003. The vegetation of the north Louisiana clay hills. Big Thicket Science Confer-

ence, Abstracts. P. 28.

Van Kiev, J.E. 2006. The pineywoods. In: G.M. Diggs, B.L. Lipscomb, M.D. Reed, and RJ. O'Kennon, eds. Illustrated

flora of east Texas. Sida Bot. Misc. 26:76-106.

WILLIAMS, R.A. AND L.M. SmitH. 1995. A survey and description of the natural plant communities of the Kisatchie

National Forest, Caney District. Louisiana Department of Wildlife and Fisheries. Baton Rouge.

A REVISION TO THE VASCULAR FLORA OF CALCASIEU PARISH, LOUISIANA

Ray Neyland

Department of Biology and oa Health

McNeese State Uni

Lake Charles, ee -o U.S.A.

ABSTRACT

Th blicati f lar fl ys of Calcasieu Parish, Louisiana by Neyland et al. in 2000 and Neyland in 2002 d

£51] 11 1 ; $112 +} f I isl Additionally, 17 eee that

revised. Sut

previously had been reported by other sources to occur in the epee but: were not found in other Peinar or during the two previous

surveys have been discovered. T

RESUMEN

a) T 1 I a A us j A A; J f] 1 J 1 I 1 = A Calcasieu T I N 1 q 12000

y Neyland en 2002. 1 i tigació bsiguiente d po y herbario } Tad 14 hrimienta de 22 pecies p la

sik AS se han descubierto 1 17 especies que habían sido señaladas p f ] qui f

1 A A T h 1 A e pa | A iod 1

This paper updates and revises the inal vascular flora surveys of Calcasieu Parish, Louisiana by Neyland

et al. 2000) and Neyland (2002). Specifically, the purpose of this paper is to list new collections from the

parish, to list newly discovered sp t previously were reported from the parish but were unverified

and to correct identification errors.

METHODS

Since the publication of the original aA by None et al. (2000) and Neyland (2002), plant collections

from Calcasieu Parish have continued. \ were prepared usi lard herbarium practices

and are housed at the McNeese State University (MCN) and Louisiana State Dare (LSU) herbaria. Ad-

ditionally, specimens from the LSU herbarium have been examined. Nomenclature follows Flora of North

America Editorial Committee (1997+) and USDA, NRCS (2008).

RESULTS AND DISCUSSION

The following entries are new additions to the vascular flora of Calcasieu Parish. These taxa have not been

reported previously as occurring in the parish. Specimen references follow the same format as in the origi-

nal survey (Neyland et al. 2000) and in the first update (Neyland 2002). Following the collection data is an

abbreviation for the vegetative region where each species typically occurs: prairie (Pr); pineland (Pi); fresh

marsh (FM); saline marsh (SM); swampland (S); bottomland forest (B); disturbed sites including ditches,

pastures, lawns, roadsides etc. (D).

anthaceae Caryophyllaceae

Ruellia nudiflora (Engelm. & A. Gray) Urb.—R. Neyland 2238 Stellaria cuspidata Willd. ex Schltdl. ssp. prostrata (Baldwin) J.K.

(MCN) D Morton—R. Neyland 2264 (MCN) D

Asclepiadaceae Clusiaceae

Asclepias tuberosa L. ssp. interior Woodson—R. Neyland 2288. Hypericum galioides Lam.—R. Neyland 780 (LSU) B, S

(MCN) B, D, Pi C i

mmelinaceae

Brassicaceae Murdannia nudiflora (L.) Brenan—R. Neyland 2105 (MCN) D

Cardamine hirsuta L.—R. Neyland 2262 (MCN) D Cyperaceae

Burmanniaceae Kyllinga brevifolia Rottb. R Neyland 913 (LSU) D

Burmannia capitata (Walt. ex J.F. Gmel) Mart.—R. Neyland Euphorbiaceae

CN) Pi Euphorbia cyathophora Murr.—R. Neyland 2198 (MCN) D

J. Bot. Res. inst. Texas 3(1): 375 — 377. 2009

376

Fabiaceae

Medicago sativa L.—R. Neyland 2106 (MCN) D

Melilotus officinalis (L) Lam.—R. Neyland 2097 (MCN) D

Vicia sativa L—R. Neyland 2278 (MCN) D

lridaceae

Iris giganticaerulea Small—R. Neyland 2249 (MCN) FM, S

edi bel albidum Raf.—R. Neyland 1202 (LSU) D, Pr

Noms prm (Hegelm.) Hegelm.—R. Neyland 2377

(MCN) D, FM, S

Orchidaceae

Zeuxine strateumatica (L) Sthltr.—R. Neyland 2266 (MCN) D

Poaceae

Panicum hemitomon Schut.—R. Neyland 2163 (MCN) D, FM

Polygonaceae

Rumex altissimus Alph. Wood—R. Neyland 2289 (MCN) D, S

Rubiaceae

Oldenlandia corymbosa L.—R. Neyland 2252 (MCN) D

Richardia scabra L.—R. Neyland 2239 (MCN) D

Solanaceae

Physalis cordata Mill.—R. Neyland 2196 (MCN) D

Violaceae

Viola sororia Willd.—R. Neyland 2263 (MCN) B

The following entries previously have been reported in the parish but were neither collected in the previous

two studies (Neyland et al. 2000; Neyland 2002) nor discovered in physical searches of the herbaria listed

in the original study (Neyland et al. 2000). These entries provide evidence for the existence of previously

unverified species reported in Calcasieu Parish (Neyland 2000). The sources for this information are a recent

search of the LSU herbarium and from new field collections. Therefore, these taxa are added to the verified

total for Calcasieu Parish.

Lentibulariaceae

Utricularia macrorhiza Leconte—G.N. Montz 6191 (LSU) FM,

Asteraceae

bc pallida (Nutt) Nutt.—C.A. Brown 5871 (LSU) Pr

uthamia e (L) Nutt.—N.M. Gilmore 3457 (LSU) D

Loganiaceae

W.Stone—A. Spigelia e e ) LL.—R. Neyland 2268 (MCN) B

—

Cyper

Carex po Wahlenb. ssp. harperi (Fernald

eatherman s.n. (LSU) S

Euphorbiaceae

Chamaesyce prostrata (Aiton) Small—P Pias & R.D. Thomas

4477 (LSU) D

Nyctagin

Boerhavia difusa L.—P Pias 4489 (LSU) D

Poaceae

Aristida ramosissima Engelm. ex A. Gray—C. Brown 8641

Croton lindheimerianus Scheele—P Pias & R.D. Thomas 4483 SU) D,

Eragrostis elliottii S. Watson—N.M. Gilmore s.n. (LSU) D, Pr

Fabiaceae Rubiaceae

T ee Small—A. Featherman s.n. (LSU) D Houstonia purpurea L. var. purpurea—A. Featherman s.n.

Gentia LSU) B

ids angulars (L) Pursh—D.L. Hartnett 8 (LSU) D Solanaceae

Solanum americanum Mill.—R. Neyland 1431 (LSU) D, Pr

o intermedia (Nutt.) Engelm. & A. Gray—A. Feather- a enn Cav.—E.C. Wurzlow s.n. (LSU) D

man s.n. (LSU) D, FM, Pr Urtic

pod i" Muhl. ex Willd. G.N. Montz 3030 (LSU)

The following entries were misidentified in the two previous surveys.

1. The specimen reported as Nepeta cataria L. Walker s.n. (MCN) D is Stachys floridana Shuttlw. ex Benth.

Therefore, this taxon is removed from the survey.

2. The specimen reported as Tamarix ramosissima Ledeb.—R. Neyland 1303 (MCN) SM is Tamarix gallica

The most notable addition to the flora of Calcasieu Parish is the discovery of a healthy population of Zeuxine

strateumatica on the campus of McNeese State University. According to the range map published by the Flora

of North America Committee (2002), the range of this species extends from Florida and southern Georgia

westward to southeastern Louisiana. Therefore, the present collection of Z. strateumatica in Calcasieu Parish

represents a westward range extension of about 200 km Additional evidence of this westward movement

is supplied by Diggs et al. (2006) who reported that Z. strateumatica occurs in Harris and Brazos counties

in eastern Texas.

As a result of these revisions to the two previous vascular surveys of Calcasieu Parish, the number of

Neyland, Vascular flora of Calcasieu Parish, Louisiana 377

confirmed specific and subspecific entries for the parish is increased to 1,178. The number of reported but

unverified entries for the parish is reduced to 113.

ACKNOWLEDGMENTS

I thank Diane Ferguson from the Louisiana State University herbarium for her assistance and Christopher

S. Reid and Charles M. Allen who provided helpful reviews.

REFERENCES

DicGs Jr, G.M., B.L. Lipscome, M.D. REED, AND R.J. O'KeNNoN. 2006. Illustrated flora of East Texas, Vol. 1. Austin College

for Environmental Studies, Sherman, Texas and Botanical Research Institute of Texas, Ft. Worth.

FLORA of NORTH America EDITORIAL Committee. 1997+. Flora of North America north of Mexico. 10+ vols. Oxford Uni-

versity Press, New York and Oxford. Vol. 3, 1997; vol. 4, 2003; vol. 5, 2005; vol. 19, 2006; vol. 20, 2006; vol. 21,

2006; vol. 22, 2000; vol. 23, 2002; vol. 25, 2003; vol. 26, 2002.

NEYLAND, R. 2002. An update to the vascular flora of Calcasieu Parish, Louisiana. 2002. Sida 20:431-433.

NEYLAND, R, BJ. HOFFMAN, M. MAYFIELD, AND L.E. UrBATSCH. 2000. A vascular flora survey of Calcasieu Parish, Louisiana.

Sida 19:361-386.

USDA, NRCS. 2008. The PLANTS Database (http://plants.usda.gov), 31 December 2008). National Plant Data

Center, Baton Rouge, Louisiana 70874-4490 USA.

378 tani i Texas 3(

BOOK NOTICE

WiLLIAM K. CHAPMAN, VALERIE CONLEY CHAPMAN, ALAN E. BESSETTE, AND ARLEEN RAINIS BESSETTE, with botanical

drawings by Puiuippa Brown. 2008. Wildflowers of Massachusetts, Connecticut, and Rhode Island

in Color. (ISBN: 978-0-8156-3185-9, hbk.; 978-0-8156-0926-1, pbk). Syracuse University Press, 621

Skytop Road, Suite 110, Syracuse, New York 13244-5290, U.S.A. (Orders: www.SyracuseUniversity-

Press.syr.edu, email supress@syr.edu, 1-800-365-8929, 1-866-536-4771 fax). $59.95 hbk., $24.95

pbk., 182 pp., 6" x 9".

From blisher website.—This field guide will gi t husi i to the di d beautiful flora of these New

E 5

England states. Combining 400 1 pi grar I itl i d if i iti i i y-to foll nical 1 g

illustrations have been f 11; selected for thei i ifi 1 thei 1 i 1 y

C 1 : a q co nee £f, 1 : s F 1 A rare A J

r 4/ " F r

in other guides. TI I p ide key h species! d } ble c} isti fcolor, flower shape l leaf

1] : g 3 A I lil P. q PO ly identify flowers NT 1 1 1 I A £1 zT ETE D a

J. Bot. Res. Inst, Texas 3(1): 378. 2009

ADDITIONS AND EMENDATIONS TO THE VASCULAR FLORA

OF CADDO PARISH, LOUISIANA

Barbara R. MacRoberts and Michael H. MacRoberts

Bog Research

740 Columbia

Shreveport, Louisiana 71104, U.S.A.

and Herbarium, Museum of Life Sciences

Louisiana State University in Sd

Shreveport, Louisiana 71115,

Christopher S. Reid and Patricia L. Faulkner

Louisiana Natural Heritage Program

. TÉ LIAS HALL po ae ae eer

Baton Rouge, Louisiana 70898, U.S.A.

ABSTRACT

Ti 1 f1 Cl f:4l4 jt 1

illc

h has yielded 25 species new to the

of Caddo Parish, Louisiana, is updated

parish. Six species are deleted from the flora. The total parish count is now 1424 species, of which 1183 are native and 241 are exotic.

RESUMEN

Lafl lar de lat ia de Caddo, Louisiana, E i dia. Lai igació bsigui d po y I id

25 la parroquia. Seis esp limi de la fl El to actual es de 1424 especies, de las que 1183 son

nativas y 241 son exóticas.

INTRODUCTION

Since the publication of “An updated, annotated vascular flora of Caddo Parish, Louisiana, with notes on

regional phytogeography and ecology” (MacRoberts & MacRoberts 2006), plant collecting in Caddo Parish

has continued and species new to the parish have been found. Consultation of recent literature has resulted

in the addition of one species. Examination of herbarium collections of species reported for the parish has

also continued, resulting in a reevaluation of some taxa. These additions and emendations are given below.

We do not address nomenclatural changes.

ADDITIONS TO THE CADDO PARISH FLORA

ALISMATACEAE

us o (L) Buchenau, Raymond 2167 LSUS, Reid

5809 FSU, LSU. Recently reported as new to Louisiana,

Sagittaria subulata was found on the shoreline of Caddo

Lake at Oil City where it was locally abundant (Reid et

al. 2008).

APIACEAE

Lil j is J.M io Reid 5978 LSU Known

X +h

found on the banks of es Lake near TERRE

ARACEAE

cn A LSUS, in dd mis Kalinsky

L er lettuce is spreading in the state and has

semis E found in Cross Lake and Ps sites in

Caddo Parish

ASTERACEAE

Senecio ampullaceus Hook., Reid 6020 with Faulkner LSU.

Thought to be a Texas endemic (Carr 2002) but recently

found in southwestern Arl | southeastern Okla-

J. Bot. Res. Inst. Texas 3(1): 379 — 382. 2009

homa (Holmes et a!. 2006, Singhurst and Holmes 1998),

Senecio ampullaceus has recently been collected from

xeric sandylands in northern Caddo Parish (Reid et al.

2008).

CALLITRICHACEAE

Callitriche pedunculosa Nutt., MacRoberts & MacRoberts 7306

in southern Caddo Parish.

CISTACEAE

Lechea san-sabeana (Buckley) Hodgdon, MacRoberts & Mac-

Roberts 7830 LSU, 7834 LSUS. Previously considered a Texas

endemic (Carr 2002; Diggs et al. 2006), Lechea san-sabeana

nao! SI Eu I

ries in Caddo and De Soto parishes (Reid et al. 2008).

CYPERACEAE

Bolboschoenus robustus (Pursh) Sojak, Reid 6210 with Sibley LSU,

This species was found in marshy open ground in

an inlet of Caddo Lake. In Louisiana and Texas, it is mainly

380

another collection from tl Louisiana in Claiborne

Parish (Thomas & Allen 1

Carex alata Torr. Reid 6492 P SEG Brimer LSU, SWSL.

This species was collected in April 2008 from a shrubby

marsh, perhaps influenced by beaver, in northern Caddo

Parish, where it was growing with abundant Osmunda

regalis and Morella cerifera.

Carex O Steud., MacRoberts 1243 and 1281 LSUS,

aynes 5322 LSU. Collections of C. frankii at LSU and LSUS

dd to M recently described entity (Ford & Reznicek

2002).

Carex bushii Mack., MacRoberts & MacRoberts 7389 LSU. This

species was collected from a saline prairie in southern

Caddo Parish and was initially identified as Carex caro-

and ela

El INC It AA DAL Q AA DAL

72320

LSU, LSUS. Becchar tenuis was found in a saline prairie

in southern Caddo

BIGGER won A Gray, a 6083 a Eleocharis wo liis on

lar

tage di RA and Texas bi of w of-concern

leoct ed from wet

(Diggs et al. 2

depressions in a saline prairie in southern Caddo Parish.

Lipocarpha drummondii (Nees) G.C. Tucker, Thieret 32137 NCU.

This species is reported new to Louisiana by Sorrie and

LeBlond (2008) based on a Caddo Parish specimen

w globularis (Chapm.) Small, MacRoberts &

MacRo ber s 7428 LSU, LSUS. Rhynchospora TEN was

e prairie in southern Caddo Parish.

Scleria eee Steud., Reid 6741 LSU, cee species

line right-of-way near Mira in northern Caddo Parish.

GENTIANACEAE

Bartonia paniculata (Michx.) Muhl., Reid 4638 LSU, TEX-LL,

eid 64 2 Md Md E Eus eed here WEG

collec

ern Scan Parish. Collected i in ‘August 2003, Reid 4638 is

referable bsp. texana (Correll) K.

hee DU E. York, & Struwe, stat. nov. (Mathews et

id as determined i B. Teana Cor-

al.2

H fel J

the collector,

and a duplicate sent to TEX- LL was annotated ei ae

Singhurst as B. texana

is included in Caddo Parish by Mathews et al. acd on

the basis of the Reid 4638 specimen at TEX-LL. Collected

in October 2007, Reid 6405 is intermediate between

the descriptions of B. texana and B. paniculata (Correll &

Johnston 1970). Bartonia dE subsp. paniculata is

expected in Caddo Parish.

HYDROCHARITACEAE

Hydrilla verticillata (Lf) Royle, MacRoberts & MacRoberts

EXCLUDED FROM THE

ASTERACEAE

Solidago ulmifolia Muhl. ex Willd. is S. rugosa P. Mill.

BRASSICACEAE

Cardamine pensylvanica Muhl. ex Wilid. is C. parviflora L.

7464b LSUS, Kalinsky s.n. LSUS. Hydrilla verticillata recently

reached Caddo Parish, where it has now spread to many

lakes and m

LENTIBULARIACEA

anciano PUIpUIca 7 It, Raymond 165 mae Reld d E

Black Bayou Lake at Noah Tyson Park in northern eats

Parish. This species occurs in coastal areas of Louisiana

and Texas and must be a recent arrival to Caddo Parish

since Noah Tyson Park is a frequent stop for botanists

and the species a have been noticed if it were a

long-standin

Utricularia radiata Small, - 6488 with Faulkner and Brimer

LSU. This bladderwort is an aquatic and was found in a

beaver-influenced slough dominated by Nyssa aquatica

in northern Caddo Parish.

Itricularia subulata L., Reid 6486 with Faulk LSU. This species

was recently discovered in an open, sandy seep along a

stream bisecting a power line right-of-way near Mira in

northern Caddo Paris

LYCOPODIACEAE

diria Run (Chapm.) Cranfill, MacRoberts & Mac-

Rober 4 LSUS. Southern clubmoss was found to be

E. common in a small seepage area within a power

line right-of-way near Mira in northern Caddo Parish.

NAJADACEAE

Najas minor All., Kalinsky s.n. LSUS. This species was recently

found on the shoreline of Cross Lake. It is an Old World

introduction that is scattered in Louisiana, mostly in the

northern part of the state; it is not yet known for East

Texas but is expected (Diggs et al. 2006).

ALVINACEAE

Salvinia molesta D.S. Mitch., Reid 5798 with Jones LSU, LSUS,

Kalinsky s.n. LSUS. This is another invasive aquatic plant

that RPM a nen Caddo Parish very recently,

here it und growing abundantly in Caddo Lake.

Since its discoverys in the parishi in May 2006, ithas spread

within Caddo Lake and has appeared in Cross Lake (C.

Reid, pers. obs..

XYRIDACEAE

Xyris difformis (Chapm.) var. difformis, Reid 6745, 6746 LSU; 6747,

49 LSUS. This species occurs in an open seepage area

within a power line right-of-way near Mira in northern

Parish.

Xyris laxifolia Mart. var. iridifolia (Chapm.) Kral, Barbour 1099

LSUS. We had previously identified this as X. caroliniana

Walt.

CADDO PARISH FLORA

CYPERACEAE

Carex frankii Kunth. See Carex aureolensis in Additions list.

Carex caroliniana. See Carex bushii in Additions list.

Websteria confervoides (Poir) Hooper. Like others before us,

MacRoberts et l., Fl f Caddo P. ish, Louisiana 381

we misidentified a sterile aquatic species of Eleocharis XYRIDACEAE

as Websteria. Although widely reported from Louisiana Xyris caroliniana Walt. See Xyris laxifolia Mart. var. iridifolia

(Thomas & Allen 1993), this species probably does not (Chapm.) Kral under Additions.

occur in the state (Bruhl 2002).

NOMENCLATURAL ERROR

CHENOPODIACEAE

Bassia scoparia should be Kochia scoparia (L.) Schroder.

DISCUSSION

The Caddo Parish vascular plant count now stands at 1424, of which 1183 are native and 241 are exotic.

By Florida standards, Caddo Parish is well collected and exceeds predicted species richness (Williams &

Debilica 2008).

ACKNOWLEDGMENTS

We thank George Dickson and Ralph and Becky Dalton for allowing our surveys on Dickson and Dalton

saline prairies, respectively. Sidney Evans and James Taylor allowed us to survey Barron Road Prairie in

Caddo Parish. We thank D.W. and Patricia Little for allowing access to their property to collect Senecio am-

pullaceus. Ryan Hook allowed access to the Senecio ampullaceus location through his family's property and

we appreciate his kindness and interest. Jason Singhurst verified our first collection of Lechea san-sabeana.

We thank Robert Haynes for verifying a duplicate of Sagittaria subulata. Charles Bryson (SWSL) vetted a

number of Carex. We thank Craig Windham of Caddo Levee District for allowing access to Levee District

lands. We are indebted to Sam Brimer, Forester with Caddo Levee District, for taking us to Levee District

lands in north Caddo Parish. We thank Alan Boyd, Weyerhaeuser Company, for allowing access to the

seep with Lycopodiella appressa, Utricularia subulata, and others. Deltic Timber Company allowed access to

some of their lands in north Caddo. Larry Raymond facilitated access to Caddo Parish parks. The Nature

Conservancy allowed access to Caddo Black Bayou Preserve near Rodessa. Royal Tyler kindly hosted us on

his property near Rodessa. Charles Allen reviewed the paper.

REFERENCES

Brun, J.J. 2002. Websteria. In: Flora of North America Committee, eds. Flora of North America north of Mexico.

Vol. 23. Oxford Univeristy Press, New York. Pp. 120-121.

Carr, W. 2002. No place but Texas: an annotated list of plant taxa endemic to the lone star state. Unpublished

manuscript. The Nature Conservancy of Texas, Austin.

ConntLL, D.S. AND M.C. JOHNSTON. 1970. Manual of the vascular plants of Texas. Texas Research Foundation, Renner,

Texas.

Dices, G.M., B.L. Liescoms, M.D. Reen, AND R.J. O’KENNON. 2006. Illustrated flora of East Texas. Sida, Bot. Misc. 26:1-1594,

Forp, B.A. AND A.A. REZNICEK. 2002. Carex Linnaeus sect. Squarrosae. In: Flora of North America Committee, eds.

Flora of North America north of Mexico. Vol. 23. Oxford nión Press, New York. Pp. 518-519.

HoLmes, W.C., J. SINGHURST, AND S.G. Powers. 2006. S. io amp (Asteraceae): a West Gulf Coastal Plain endemic

new to Oklahoma. Phytologia 88:193- 196.

LOUISIANA NATURAL HERITAGE EPUM 2009. Pale Dan list. Department o ME and Fisheries, Baton Rouge. Avail-

able at: http//www.wlf.loui

MacRoserts, B.R., AND M.H. MacRoserrs, 2006. dm ed "modd cl flora of Caddo Parish, Louisiana,

with notes on regional phytogeography and ecology. Sida 22:1191-1219.

MarHEWS, K.G., N. Dunne, E. York, AND L. STRUWE, 2009. A phylogenetic analysis and taxonomic revision of Bartonia

(Gentianaceae: Gentianeae), based on molecular and morphological evidence. Syst. Bot. 34:162-172.

Remp, C.S., PL. Faulkner, B.R. MacRoserTs, AND M.H. MacRoserTs. 2008. Noteworthy vascular plant collections from

northwest Louisiana. J. Bot. Res. Inst. Texas 2:643-647.

Dag *.. 15 A £T,

382 Journal of tl

SINGHURST, J.R. AND W.C. Homes. 1998. Four new additions to the Flora of Arkansas. Castanea 64:276-277.

Sori, B.A. AND RJ. LEBLOND. 2008. Noteworthy collections from the southeastern United States. J. Bot. Res. Inst.

Texas 2:1353-1361.

Thomas, R.D. AND C.M. ALLEN. 1993. Atlas of the vascular flora of Louisiana. Louisiana Department of Wildlife and

Fisheries, Baton Rouge.

WiLuams, J.K. AND A. Deseuica. 2008. Analysis of the completeness of vascular plant records in Florida. J. Bot. Res.

Inst. Texas 2:1363-2371.

MENTHA SUAVEOLENS AND M. xROTUNDIFOLIA IN NORTH CAROLINA:

A CLARIFICATION OF DISTRIBUTION AND TAXONOMIC IDENTITY

Michael W. Denslow Derick B. Poindexter

LW. Carpenter Jr. Herbarium LW. Carpenter Jr. Herbarium

Department of Biology epartment of Biology

Appalachian State University Appalachian State University

Boone, North Carolina 28608, U.S.A Boone, North Carolina 28608, U.S.A.

md68135@appstate.edu poindexterdb@appstate.edu

ABSTRACT

A capace of the distribution and identity of Mentha suaveolens and Menta d is mue ee Tomates S

North Carolina for the first time. P p

n non- hybrid name e rotundifolia to M. suaveolens. Ment! lij lia i HOMES to be well aa in northwestern North

pop Fy AS ri EE 1970. This

I pp ly land in North Carolina ony to ee literature reports M. longifolia does not appear

to be present in North Carolina. A key and photograg of these taxa

RESUMEN

E È : A A rian do ladi ar bhann e enida dn 1 M lifolia. Se encuentran

poblaciones naturalizadas de M tundifoli Carolina del Norte por j Drum vez. Las citas anteriores ni M. xrotundflia estaban

la clasifi M. suaveolens. M

1 J

A" 33: 2-203 1 Jar 15 dol N - d: i p Porl Į i vez desde 1970,

24 =f: 11 : An AA suaveolens E 1; r^ X dal RT ARE Al : 1 I Hii 1

anteriores, M. lonatiolia no aparece en Carolina del Nore Se delayed una ea y fotos Į poyar la identificación de estos taxa

INTRODUCTION

The genus Mentha has traditionally been regarded as taxonomically difficult. High levels of morphologic

plasticity within this group have been attributed to rampant hybridization, polyploidy, colonial mutant

propagation, and the persistent/spreading nature of nothomorphs (Stace 1997; Tucker & Chambers 2002;

Bunsawat et al. 2004). In addition, the genus Mentha is extensively cultivated for its economic value (e.g.,

culinary use, essential oil production, ornamental value) (Mabberley 1997; Sutour et al. 2008). As a result

of such hybridization and the continued human introduction of multiple taxa, misidentification and un-

certainty in the literature is expected.

The primary purpose of this paper is to clarify the distribution and taxonomic identity of what has

traditionally been referred to as Mentha rotundifolia auct. non. (L.) Huds. in North Carolina. It is beyond the

scope of this paper to address all the taxa within the genus Mentha. Rather, we will focus on the state-wide

distribution and confusion of a single taxonomic concept that has been traditionally misapplied in this state

and elsewhere. The information presented will also assist in the identification of Mentha specimens in other

geographic areas where the concept of M. rotundifolia has been applied.

THE PROBLEM

While collecting in northwestern North Carolina we encountered a confounding member of the genus

Mentha at a several scattered localities. All of these specimens had terminal inflorescences and pubescent,

rugose leaves. These plants most closely matched the concepts of M. rotundifolia and M. longifolia (L.) L. in

the Manual of the Vascular Flora of the Carolinas (Radford et al. 1968) and the dnd of M. suaveolens Ehrh. in

the Flora of the Carolinas, Virginia, and Georgia, northern Florida, and ing (Weakley 2008). Mentha

J. Bot. Res. Inst. Texas 3(1): 383 — 389. 2009

384 tani Insti Texas 3(

rotundifolia is now considered to be a hybrid derived from a cross between M. longifolia and M. suaveolens

and is referred to as M. xrotundifolia (L.) Huds. (Tucker & Naczi 2007). The name M. rotundifolia has been

misapplied to M. suaveolens, M. xrotundifolia and M. xvillosa Huds. on herbarium sheets, in the literature, and

in the herb trade (A.O. Tucker, pers. comm.) In addition, M. rotundifolia has been erroneously considered

synonymous with M. suaveolens by some authors e B. E et al. 2008).

Mentha longifolia, M. suaveolens and M. xrot ll been considered infrequent escapes in North

Carolina and are only oe from a few counties (Wofford 1989; Radford et al. 1968; USDA, NRCS 2009).

The Mentha t ly become well established outside of cultivation in Alleghany, Ashe

and Watauga counties. For dis reason we set out to clarify the identity of these plants and to determine

where they had previously been documented in North Carolina.

METHODS

In addition to our own field collections, we requested all specimens labeled as M. xrotundifolia, M. suaveolens,

M. longifolia, including relevant synonyms, as well as Mentha specimens not identified below genus from

all major North Carolina herbaria. Additionally, we requested loans of the type specimens of M. suaveolens

and M. spicata L. var. rotundifolia L. from GOET and UPS respectively. Tucker and Naczi (2007) was used

for identifying basionyms and locating the institutional repositories for the type specimens. Specimens

were examined from the following herbaria: BOON, DUKE, NCSC, NCU, UNCC, WCU, and WNC. AII

herbarium acronyms follow Index Herbariorum (Holmgren & Holmgren 1998). High resolution digita!

photos of the lectotype specimens of M. suaveolens and M. spicata var. rotundifolia were also examined. In all,

we examined 10 vouchers and respective duplicates of M. xrotundifolia and 12 of M. suaveolens from North

Carolina. In addition, we ined many vouchers of cultivated Mentha hybrids from the herbarium of M J.

Murray supplied by A.O. Tucker.

RESULTS

The examination of type specimens and previously published literature (e.g., Stace 1997; Tucker & Naczi

2007), revealed 11 vouchers of M. suaveolens and one previous collection of M. xrotundifolia from North

Carolina. No ca. of M. longifolia were seen. Our own field work in northwestern North Carolina lo-

cated 9 new popul of M. xrotundifolia and one new population of M. suaveolens (Fig. 1). Our specimens

represent the first known accounts of naturalized populations of M. xrotundifolia in North Carolina and the

first collections since 1990. Prior to this study, M. suaveolens had not been collected since 1970. Mentha

longifolia has apparently not yet been legitimately documented for North Carolina.

A key, specimen citations, and photographs (Fig. 2) are provided below to facilitate the identification of

additional populations of M. xrotundifolia or M. suaveolens. We include M. longifolia, M. spicata and M. xvil-

losa in the key because they have been confused with M. xrotundifolia or M. suaveolens in the past. Mentha

longifolia and M. xvillosa have not been documented from North Carolina; however, these taxa have been

reported from the eastern United States (e.g., Rhoads & Block 2007; Weakley 2008, Tucker in press). We

therefore include these taxa in the key to increase its utility. Most of the specimens examined were readily

identifiable as either M. xrotundifolia or M. suaveolens. However, some specimens had primary affinities with

M. suaveolens, but exhibited some intermediacy. As pointed out by Stace (1997), these intermediate taxa are

usually hybrids involving M. suaveolens and can be difficult to distinguish. However, the following key can

be reliably used to separate M. xrotundifolia and M. suaveolens for a great majority of specimens. Back crosses

between M. xrotundifolia and it parents have been documented in common garden settings, though it is not

clear if this occurs in naturalized populations. The taxa included in the key can be distinguished from other

Mentha taxa in North Carolina by the presence of a terminal spike with densely crowded flowers. It should

be noted that 3 subsp. of M. suaveolens have been recognized by Tucker and Naczi (2007). In addition to

the type subsp. these authors recognized subsp. insularis (Req.) Greuter and subsp. timija (Coss. ex Briq.)

Harley ex Harley & Brighton. However, neither a key nor a description is presented by Tucker and Naczi

385

37*N 4

36°N 4

35°N 4

34°N 4 A^ M. suaveolens

M. x rotundifolia 0 50 100km

€ both taxa N

Y Y Y y LI

849W 82°W 80°W 78°W 76°W

Fic. 1 f. y I Lot iL 40 L£ A8 suaveolens and M Jc. E . a Lr. ge (I J | 2 £. DAMA DUKE, NCSC NCU UNCC, WCU

and WNC). Symbol pped ty d

(2007) to separate these 3 taxa. Harley (1972) expresses doubt as to the distinctiveness of M. insularis

Req. (= M. suaveolens subsp. insularis), while subsp. timija is not mentioned by Harley since it is outside the

geographic scope of this treatment. In short, it is difficult at this time to definitively identify which subsp.

of M. suaveolens is present in North Carolina, but after a review of existing literature, we have tentatively

defaulted with Weakley’s (2008) recognition of ssp. suaveolens. Future taxonomic studies across the native

and introduced ranges of M. suaveolens are needed to define these taxa.

KEY TO SELECTED TAXA OF MENTHA

Key adapted from Stace (1997), Tucker and Naczi (2007), Weakley (2008) and Tucker (in press). Note: Taxa

included within brackets are not currently known from North Carolina.

1. Leaves lanceolate to oblong-lanceolate

2. Abaxial leaf hairs unbranched, leaves widest near the middle, slightly rugose; fertile anthers 0.28-0.38 mm

long, generally musty-odored (2n = 24) [M. longifolia]

2. Leaves oblong-lanceolate, adaxially glabrous, rarely with tree-like hai d face, leaves widest near

se, conspicuously rugose; fertile anthers 0.38-0.52 mm, generally spearmint-odored (2n = 48) M. spicata

1, Leaves oblong to ovate

3. Leaves generally 1-2X as long as wide, ovate-suborbicular, broadly rounded to subcordate at the base,

apex obtuse; leaf serrations rounded and often turned downward (thus appearing somewhat crenate);

leaf surface strongly rugose, with scattered dendritic hai he abaxial surf. ^sickl t

179

=24) M. suaveolens

. Leaves generally 1-3x as long as wide, ovate to oblong, broadly cuneate to rounded at the base, apex

acute; leaf serrations sharp and patent, leaf surface moderately rugose; si int scented

4. Leaves generally oblong, with nearly parallel sides and a broad rounded base; each flower generally

with 4 fertile anthers (2n M. xrotundifolia

4. Leaves generally ovate, infrequently oblong; scattered fertile anthers sometimes present (2n = 36)

[M. xvillosa]

X7, 1 r AA 1. JA INA All h

peci if ghany Co.: Glade Valley, located along US 21 S, ca. 1 mi past

Glade Valley Rd., latitude 36.4651, longitude -81.0502, 28 Aug 2008, Poindexter 08-983 (BOON). Ashe Co.: ca. 1.25 mi down William

T. Calloway Road, Bluff M in Gameland, latitude 36.3684, longitude -81.5738, 22 Sep 2007, Zgieb s.n. (BOON); Orion, located

along NC 88/16, ca. 1 mi from 16 S divergence and the South Fork of the New River, traveling E just past Frank Dillard Rd., latitude

36.3956, longitude -81.4185, 28 Aug 2008, Poindexter 08-993 (BOON). West Jefferson, l datthel ÍM Jeff at the jct. of

Oakwood Rd. and Cottontail Trail, latitude 36.3878, longitude -81.4663, 7 Aug 2008, Poindexter 08-673 (BOON). West Jefferson, located

Mentha suaveolens Ehrh.

m anba xr tundzfofta (L) Huds,

AO, Tucker

Pal North Carolina 2 1 /

i . Fa is TIS .. foh Canela, U:

X Lanata a Co.

Eo Qdkuths suaveolens Ehrhan. [Apple Mint] "

m ^. Jeffers unt Jefferson ^ ane N aural Are; asd environs, Blue RÀ

p pu B idge Province; Southern Appalachian Mountain -: Along

X ix "ns Senec. 142709 ea, it. 56.262" and vnnamod tributary of Sowth Fark New River. Brookshire

d " i238: ner

` . S pun

tne la imme ddr recie,

inam ccm rue etu Ema rper

[ Derick B. Poindexter DIETA 21.0 uber 2007

APPALACHIAN BIATE VMIENSITY HERSARIGM [BODH]

DOORS, BORTH CAROLINA, U.S.A. 28504

Fic. 2. Specimen, leaves, and infl f A-C) Ment lens, and D-F) Mentha xrotundifolia.

along Mount Jefferson Rd., ca. 1 mi prior to Ashe County High School and Oal d Rd., latitude 36.3858, longitude -81.4804, 7

Aug 2008, Poindexter 08-674 (BOON). Wat Co.: bel Boone, poe a a large na E 421; a dirt Rd., 15 Sep 1990, Basinger 853

Boone, near US 421 bridge on Charlie Hollar Rd., rud the South k of the New River, latitude 36.2203,

NCC);

longitude -81.6399, 22 Oct 2005, Poindexter 05-2255 (BOON, DOV); Greenway Trail E of Boone, isis unnamed tributary of South

Fork New River, latitude 36.2114, longitude -81.6478, 18 Sep 2007, Denslow 2560 (BOON, DOV); along unnamed tributary of South

Fork New River, Brookshire Park ENE of Boone, latitude 36.2292, longitude -81.6411, 1 Oct 2007, Denslow & a 2561 (BOON,

DOV); Boone, located along Wilson Rd., just before the jct. with NC 105 (Linville Rd.), latitude 36.2025, longitude -81.6721, 14 Aug

2008, Poindexter 08-832 (BOON).

Habitat and Phenology.—Mentha xrotundifolia has so far been observed in wet depressions, and disturbed

areas including floodplains and streambanks. It has been collected between early August and late October,

n ri o PE led M + I JAA +1 + ditalia 387

with a majority of the vouchers made between mid August and late September. The average collection date

was the 9" of September.

Voucher specimens for Mentha suaveolens: NORTH CAROLINA. Ashe Co.: Jefferson, Mount Jefferson State Natural Area and en-

virons, S.R. 1152, growing along disturbed embankment adjoining woodlands, at latitude 36.3925 and longitude -81.4674, 11 Aug

2005, Poindexter 05-1695 (BOON); same as previous locality, 21 Oct 2007, Poindexter 07-874 (BOON, DOV). Beaufort Co.: Belhaven,

Salt marsh, 6 Jul 1958, Radford 36297 (NCU). Craven Co.: New Bern, waste place, 19 Jul 1958, Radford 37332 (NCU). Graham Co.:

meadow border, 2.5 mi N of Robbinsville, on [U.S.] 129, 20 Jun 1965, Crisp s.n. (WCU). Haywood Co.: Creek side, 5.2 mi ENE of

Crabtree, 5 Jun 1958, Ahles & LaDuke 42139 (NCU). New Hanover Co.: Wilmington, moist thicket, 7 Jul 1938, Godfrey 4880 (NCSC);

Wilmington, E bank of Cape Fear River off U.S. 421, marshy field, 6 Aug 1966, Bradley & Stevenson 3395 (BOON, DUKE, NCU, UNCC,

WCU, WNC not seen); along railroad tracks on E side of Northeast [Cape Fear] River at US 17, Wilmington, 23 Oct 1968, Leonard &

Radford 2. Wilmington, en area, d Da Fear] River, 22 Aug 1970, Leonard 3553 (WCU). Transylvania Co.:

Cascade L 15 Aug 1958, F 2 (NCU). V Co.: 0.4 mi E of Plymouth, fresh-water marsh, near Roanoke River, 5

Aug 1958, a 38817 (NCU).

Habitat and Phenology.—Mentha suaveolens is primarily restricted to moist or wet areas such as lake shores,

creek sides and salt marshes. It has also been observed in disturbed sites such as along railroads and road

embankments. This species has been collected between early June to late October, though a majority of the

specimens were vouchered between early July and mid August. The average collection date was the 5° of

August.

DISCUSSION

Naturalized populations of M. xrotundifolia sensu Tucker and Naczi (2007) are reported from North Carolina

for the first time. This hybrid has been reported from North Carolina previously (e.g., USDA, NRCS 2009);

however, as stated above, these reports were app ly based on misapplication of the name M. rotundifolia

to M. suaveolens.

Mentha xrotundifolia is widely cultivated and this is most likely the pathway by which this plant was

introduced into North Carolina. Horticulture and agriculture are common vectors of plant introduction to

new areas (Reichard & White 2001). Mentha xrotundifolia has been cultivated in the Boone area for at least

50 years. We examined a vouchered garden planting of M. xrotundifolia dated 1957 from Davidson (Mecklen-

burg County) that was ‘transplanted from Boone, N.C.’ [Brown 689, 10 July 1957, (UNCO)]. Today, cultivated

stems are sold at local farmer's markets in western North Carolina [Denslow 2591, 30 Aug 2008, (BOON)].

Regardless of the mode of introduction, this plant is now well established outside of cubano, Although

we have observed naturalized populations in close proximity to cultivated plants, most popul appear

to be established in more disparate localities. It is most successful in moist areas near water sources where

it can become the dominant plant.

In light of the prevalence of M. xrotundifolia in northwestern North Carolina, it is unclear why this

plant was only collected once prior to 2007. One possible explanation is that this plant has simply been

overlooked by collectors. Another possibility is that M. xrotundifolia has only recently become successful

outside of cultivation. Long periods from initial establishment to subsequent spread or ‘lag times’ are com-

monly observed for introduced species (Lockwood et al. 2007). Explanations for these lag times include

an increase in positive biotic interactions and post-establishment evolution that promotes range expansion

(Lockwood et al. 2007). It is also possible that the rate of introduction (i.e., propagule pressure, Lockwood

et al. 2007) has increased in recent years.

A previous report by Radford et al. (1968) of M. longifolia from Gates County, North Carolina was based

on a misidentification of M. spicata. Radford et al. (1968, p. 924) reports M. longifolia as a ‘very rare escape’

from a ‘roadside ditch’. This report is apparently based on a specimen collection by H.E. Ahles on 1 Aug

1958 and is now listed as M. spicata in the NCU database (http://www. herbarium.unc.edu/seflora/firstviewer.

htm). Thus, Mentha longifolia has not yet been collected in North Carolina.

The first report of M. rotundifolia for North Carolina appears to be from the Wilmington area (Curtis

1834). Curtis's (1834) description of this plant indicates that it may actually be M. suaveolens. Curtis (1834,

388 i Institute of Texas 3(

p. 26) states that the plants have leaves that are “roundish, rugose, crenate, sessile" and are “unpleasantly

scented." These characters are consistent with M. suaveolens, though voucher specimens for this report have

not yet been located.

Radford et al. (1968) apparently misapplied the name M. rotundifolia to specimens of M. suaveolens.

This misapplication may have caused other authors (e.g., NRCS-USDA 2009; Weakley 2008) to incorrectly

report M. x rotundifolia from North Carolina. Contrary to these reports we examined no previous specimens

labeled M. x rotundifolia from North Carolina. We did however locate a specimen of M. x rotundifolia collected

in 1990 [Basinger 853, (UNCC)] that was misidentified as M. spicata. This specimen seems to be the first

known collection of M. x rotundifolia from North deeds

Based on current herbarium records, M to infrequently establish in North Carolina.

It is possible however that this taxon is more frequent, but has been overlooked by collectors. Future plant

collecting and detailed habitat descriptions will help determine if this plant persists at the historic sites

where is has been documented. In particular, M. suaveolens should be sought in the Wilmington area where

it was collected several times in the past.

ACKNOWLEDGMENTS

We extend our appreciation to the curators and staff of DUKE, NCSC, NCU, UNCC, WCU, and WNC for

access to their specimens, as well as GOET and UPS for images of type specimens. We would like to thank

Arthur O. Tucker for verification of our original specimens, for generously providing specimens of Mentha

hybrids and valuable insights that improved the manuscript. We are especially grateful for the helpful sug-

gestions on this manuscript provided by Christine Edwards, Gabrielle Katz and the Spanish translation of

the abstract provided by L. Baker Perry. Funds to offset publication costs were graciously provided by the

Graduate Student Association Senate at Appalachian State University.

REFERENCES

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Curtis, M.A. 1834. Enumeration of the pl g g spont ly around Wilmington, North Carolina. Boston

J. Nat. Hist. 1:82-141.

Harley, R.M. 1972. Mentha. In: T.G. Tutin, VH. Heywood, N.A. Burgess, D.M. Moore, D.H. Valentine, S.M. Walters

and D.A. Webb, eds. Flora Europaea, Volume 3 Diapensiaceae to Myoporaceae. Cambridge University Press,

Cambridge. Pp. 183-186.

HOLMGREN, P.K. AND N.H. HOLMGREN. 1998 [continuously updated]. Index herbari global directory of public her-

baria and associated staff. New York Botanical Garden's Virtual Herbarium. http://sweetgum.nybg.org/ih/

LockwooD, J.L., M.F. Hoopes, AND M.P. MarcHetti. 2007. Invasion ecology. Blackwell Publshing, Malden.

MassenLEY, D.J. 1997. The plant book: a portable dictionary of vascular plants. Ed. 2. Cambridge University Press,

Cambridge.

RADFORD, A.E., H.E. AHLES, AND CR. Bett. 1968. Manual of the vascular flora of the Carolinas. The University of North

Carolina Press, Chapel Hill.

REICHARD, S.H. AND P. WHr. 2001. Horticulture as a pathway of invasive plant introductions in the United States.

BioScience 51:103-113.

RHOADS, A.F. AND T.A. BLock. 2007. The plants of Pennsylvania: an illustrated manual. University of Pennsylvania

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STACE, C. 1997. New flora of the British Isles. Ed. 2. University of Cambridge Press, Cambridge.

SUTOUR S., P. BRADESI, D. DE ROCCA-SERRA, J. CASANOVA, AND F. Tomi. 2008. Chemical composition and antibacterial activity

of the essential oil from Mentha suaveolens ssp. insularis (Req.) Greuter. Flav. Fragr. J. 23:107-114

Tucker, A.O. 1992. The truth about mints. Herb Companion 4:51—52.

n I l Daina 4 : AA ^L I JAA £L * Aifals 389

Tucker, A.S. in press. Mentha. In: The Jepson manual: higher plants of California Ed. 2. J.C. Hickman, ed. University

of California Press, Berkeley.

Tucker, A.O. AND H. CHampers. 2002. Ment densis L. (Lami ):a relict amphidiploid from the Lower Tertiary.

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Tucker, A.O., R.M. HarLey, AND D.E. FAIRBROTHERS. 1980. The Linnaean types of Mentha (Lamiaceae). Taxon 29:233-

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Tucker, A.O. AND R.F.C. Naczi. 2007. Mentha: an overview of its classification and relationships. In: Lawrence, B.M.

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Accessed 20 March 2009.

WeaxLer, A.S. 2008. Flora of the Carolinas, Virginia, Georgia, northern Florida, and surrounding areas. working

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—

390 Jo

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Environmental Issues in Russia—Laura A. Henry, Vladimir Douhovnikoff

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Cumulative Index of Contributing Authors, Volumes 24—33

Cumulative Ind Chapter Titles, Volumes 24—33

J. Bot. Res. Inst, Texas 3(1): 390. 2009

CASTILLEJA ANGUSTIFOLIA VAR. DUBIA (SCROPHULARIACEAB),

A NEW RECORD FOR SOUTH DAKOTA

Grace Kostel Lynn Hetlet

Black Hills State University USDA FS Nebraska National Forest

1200 University Street, Unit 9003 Buffalo Gap National Grassland

erbarium, J159 1801 Highway 18 Bypass

Spearfish, South Dakota 57799-9003, U.S.A. Hot Springs, South Dakota 57747, U.S.A.

GraceKostel@bhsu.edu thetlet@fs.fed.us

ABSTRACT

W he first d f Castill ifolia (Nutt.) G. Don var. dubia A. Nelson (C. chromosa A. Nelson) in South

Dakota: The species was found on the Buffalo Gap National e by pis Hetlet in e and nine Ms collected, identified,

and vouchered by Grace Kostel in 2006. Its distribution

RESUMEN

Citamos la primera ocurrencia decundentada a ee een ae - Don var. dubia A. Pads (C. d A. Puis en

Dakota del Sur. I I P ynn Hetlet en 200 ntificada

y comprobada por Grace Kostel in 2006. Se discuten la distribución l E fológi li yel el hábitat

INTRODUCTION

The Artemisia iaa Nutt. shrub habitat in southwest South M uie begins to DENN way to mixed-grass

prairie and dge of A. tridentata. Much of the i ion is under public

ownership aid administered by the USDA Forest Service Nebraska s Forest. Castilleja angustifolia var.

dubia (C. chromosa) is an associate of A. tridentata at lower elevations (Holmgren 1984). A floristic inventory of

the Buffalo Gap National Grassland and vicinity was conducted recently, and led to the discovery of several

other species new to the grassland (Kostel 2006).

Distribution.—Castilleja angustifolia var. dubia (C. chromosa A. Nels.) ranges throughout and peripheral to

the Intermountain region, central and eastern Wyoming, western Colorado, northwest New Mexico, central

and northern Arizona, and eastern California (Holmgren 1984; Chumley 1996).

Mind [i c U.S.A. i piis DAKOTA. nn River Co.: T8S R1E S34 N1/2 Mule Creek SE Quad.; 43.3169? N; -103.9876° W

lca. 8 air mi WNW of as and ca. 1 mi due N of SD Hwy. 18; Artemisia

Bid one plant with ca. 12 inflorescences, bright yellow; soil (sand with gravel), permeable, a overlie shale, 20

May 2006, Grace Kostel 10056 (BHSC, SD, USA)

Morphology.—Perennial herb from woody base, 1-4 dm tall, stems ascending to erect, usually several in a

ad herbage densely hispid with long, somewhat flat, multicellular hairs; leaves with 3—7 lobes distally;

often broad and compact, bright red to orange-red, occasionally yellow; bracts lanceolate with

1-3 pairs of rounded segments; calyx 20-27 mm long, the primary lobes 6-12 mm long adaxially, 4-10 mm

long abaxially; corolla 20-32 mm long, galea 10-18 mm long, about half the corolla length, the lower lip much

reduced with incurved teeth, the tube 10-13 mm long; capsule 9-15 mm (Holmgren 1984; Dorn 2001).

Soil.—The most influential feature affecting vegetation at the site of C. angustifolia var. dubia is soil. Soils

are of the Samsil series characterized by clay, smectitic, calcareous, mesic, shallow, and aridic ustorthent

found on gently sloping to very steep hills, ridges, and breaks of dissected shale plains. These soils formed

in alluvium or residuum weathered from shale. The loose texture makes it susceptible to localized wind

erosion of topsoil when it is grazed too heavily (Soil Survey Staff 1982).

Habitat.—The area supports dry upland shrub communities of Artemisia tridentata Nutt. ssp. wyomin-

gensis Beetle & Young, Ericameria nauseosa (Pall. ex Pursh) G.L. Nesom & Baird, Atriplex spp., and Sarcobatus

—

J. Bot. Res. Inst. Texas 3(1): 391 — 392. 2009

392 J t tanical Insti Texas 3(

vermiculatus (Hook.) Torr. These shrubs are interspersed with bare ground and graminoids, e.g., Bouteloua

gracilis (Willd. ex Kunth) Lab. ex Griffiths, Bouteloua curtipendula (Michx.) Torr., Hesperostipa comata (Trin.

& Rupr.) Barkworth, Pascopyrum smithii (Rydb.) A. Love, and Bromus spp. The forbs Thermopsis rhombifo-

lia (Nutt. ex Pursh) Nutt. ex Richardson, Draba nemorosa L., Lesquerella arenosa (Richardson) Rydb. var.

arenosa, Penstemon eriantherus Pursh, MUS ein Pursh, Lepidium densiflorum Schrad., Descurainia

pinnata (Walter) Britton, and Androsace occidentalis Pursh are associate species at this site. The Buffalo Gap

National Grassland is leased for cattle grazing. ibd the area is browsed by native fauna including

deer, antelope, and small mammals.

DISCUSSION

This is the first report for C. angustifolia var. dubia in South Dakota. It has conservation status in Montana

(S3) and Idaho (S4) and is SNR/SU throughout the remainder of its range (Nature Serve 2008). Castilleja

angustifolia var. dubia is not a rare species when its entire range is considered, and certainly it is not rare in

adjacent Wyoming (Chumley 1996); however, its occurrence in southwestern South Dakota is likely due

to the unusual habitat.

CONCLUSIONS

Upon review of specimens at BHSC and RM, it is clear that Castilleja angustifolia var. dubia occurs in south-

western South Dakota on the Buffalo Gap National Grassland in Fall River County. Additional floristic

surveys in the area might result in the discovery of additional plants.

ACKNOWLEDGMENTS

The authors wish to thank Ronald L. Hartman, B. Ernie Nelson, Gary E. Larson, Ronald R. Weedon, and

Steven B. Rolfsmeier for reviews of the manuscript, and the Rocky Mountain Herbarium (RM) and Black

Hills State University Herbarium (BHSC) for the use of specimens for identification and services. Sincere

appreciation is extended to Mark L. Gabel for the Spanish translation.

REFERENCES

CHuMtEv, T. 1996. Atlas of the vascular plants of Wyoming. Online species distribution maps of the Rocky Moun-

tain (RM) Wyoming Specimen Database. Available http://www.sbs.utexas.edu/tchumley/wyomap/list.ntm.

(Accessed: May 2, 2008).

Dorn, R.D. 2001. Vascular plants of Wyoming. 3* ed. Mountain West Publishing, Cheyenne, Wyoming, U.S.A.

Houmeren, N. 1984. Castilleja. In: A. Cronquist et al. Intermountain flora: vascular plants of the Intermountain

West, U.S.A. 4:488.

Kostet, G.M. 2006 A vascular plant inventory of the Buffalo Gap National Grassland (South Dakota) and Oglala

National Grassland (Nebraska). M.S. thesis, University of Wyoming.

Nature Serve, 2008. NatureServe Explorer: An online encyclopedia of life [web application]. Version 7.0. NatureServe,

Arlington, Virginia. Available http//www.natureserve.org/explorer. (Accessed: May 2, 2008).

Soll Survey STAFF. 1982. Soil survey of Fall River County, South Dakota. United States Department of Agriculture,

Natural Resources Conservation Service, Washington, D.C.

GLYCERIA DECLINATA (POACEAE) NEW TO THE FLORA OF TEXAS

Charles M. Allen Patricia & David P. Lewis

Colorado State Univ, Fort Polk Station 262 CR 3062

Newton, Texas 75966-7003, U.S.A.

1645 23rd St.

Fort Polk, Louisiana 71459, U.S.A.

ABSTRACT

NR 1 T, AA: T

Lid 5 F o

Resh: D

nv 7 Tark

species of Glyceria in that state.

RESUMEN

I^ J r

(73 à A ds Huila D S IN e;

otras tres especies de Glyceria en el estado.

Low glyceria (Glyceria declinata Brébiss) (Poaceae) (Fig. 1) is a European species that has become estab-

lished on the western seaboard of North America from southern British Columbia to southern California

(Barkworth & Anderson 2007; Whipple et al 2007)). It is also reported from northeastern Nevada, Arizona,

and Long Island, New York. Saichuk et al (1981) reported it new to Louisiana from Jefferson Davis Parish

and Allen et al (2004) lists it from nine additional parishes. Three species of Glyceria (G. arkansana Fern., G.

1 for Texas (Diggs et al. 2006; Barkworth

septentrionalis A. Hitchc., and G. striata (Lam.) A. Hitchc.)

* t

Vor

Ae o

tel oc

* .

B t

2 [b iE

Ped:

* 4j *

Nh.

gu.

ps .

TEE DA A A pne iam og,

Pe. nor

ta daneli Dohter £ I: - L (1950 86, fig 94)

Fic. 1. Gl)

J. Bot. Res. Inst. Texas 3(1): 393 — 394, 2009

1 af el D ? al D kh Boa ether ob, £T.

394 j Texas 3(

& Anderson 2007; USDA, NRCS 2009), This collection is apparently the first collection of G. declinata for

Texas. Glyceria declinata can be separated from G. striata by its erect rigid pedicels, spikelets longer than 5

mm with more than seven florets, and lemmas longer than 2.5 mm compared to weak flexuous pedicles,

spikelets shorter than 5 mm with seven florets or fewer, and lemmas shorter than 2.5 mm in G. striata.

Glyceria declinata is a smaller stature plant (shorter than 0.5 m) and has lemma apices that are irregularly

lobed compared to tall plants (mostly 1 m and taller) and lemmas with entire to slightly crenate apices in

both G. arkansana and G. septentrionalis.

Voucher specimens: TEXAS. Newton Co.: Caney Creek Park, S of Hwy. US 190 in Newton, 30° 50' 842"N 93° 45' 802"W, 17 Mar 2008,

Allen and Lewis 20732 (BRIT, FTPK).

REFERENCES

ALLEN, C.M, D.A. Newman, AND H.H. Winters. 2004. Grasses of Louisiana, 3" Ed. Allen's Native Ventures, Pitkin,

Louisiana.

BaakwonrH, M.E. AND L.K. ANDERTON. 2002. Glyceria. In: Flora of North America Committee, eds. Flora of North America

north of Mexico. Vol. 24. Oxford Univeristy Press, New York. Pp. 68-88.

Dicas, G.M, B.L. Lipscome, M.D. Reep, AND RJ. O’keNNON. 2006. Illustrated flora of East Texas. Sida, Bot. Misc. 26:

1-1594

Hrrcucock, A.S. (Rev. A. Chase). 1950. Manual of the grasses of the United States. USDA Misc. Publ. No. 200. Wash-

ington, DC.

SAICHUK, J.K., C.M. ALLEN, AND S. Hepert. 1981. Glyceria declinata Brebiss in Louisiana. Sida 9:19-20.

USDA, NRCS. 2009. The PLANTS database (http://plants.usda.gov/plants). National Plant Data Center, Baton

Rouge, Louisiana 70874-4490 USA.

Wuirete, 1.G., M.E. BarkworTH, AND B.S. Bushman. 2007. Molecular insights into the taxonomy of Glyceria (Poaceae:

Meliceae) in North America. Amer. J. Bot. 94:551-557.

VEGETATION AREAS OF TEXAS: CONCEPT AND COMMENTARY

Ray C. Telfair Il

117805. Hill Creek Road

Whitehouse, Texas 75791-8212, U.S.A.

ABSTRACT

1 ET 1 1 1 J f£. 1 Flifof 1 ; J A

and taxa (floristic composition). The aae ol plant Pai) me in T

refinements continue. Important results include identification of vegetation types as sell as tee ean of those Pel that are both

threatened and in need of protection. pour boundaries ca areas are auc to assess, their pou communities are

recognizable; thus, the concept of ti p y 8

RESUMEN

1 A larga hist | 1 Yad sd za Aa

Las áreas de vegetación de Tı I p

(fisonomía y estructura) y taxa (composición florística). El número de tip idades vegetales de Texas depende del sistema de

Misi us usado i los mo realizados. Los ee importantes melee Mb ADD bi = ps de vegetación así

i r 1 I E

pación DUILL

dificil d taHl idad 1 21.1 H 1 Po IN J P D

en la conservación y manejo de flora y fauna.

INTRODUCTION

MacRoberts and MacRoberts (2008) analyzed the vegetational species richness of Texas. They found that

plant species richness does not correlate well with primary productivity or size of currently recognized

vegetational areas, i.e., the majority of species are not confined within regional areas. They suggest that

the traditional vegetational areas of Texas should be reassessed to provide a simpler vegetational map more

reflective of current species distributional information. Therefore, I believe it is important to examine the

vegetational area concept; to provide some commentary about the ecological processes that produce these

areas; and, to discuss their relevance to conservation and management.

VEGETATIONAL AREAS VS. FLORISTIC AREAS

fl : J + N 1

Landscapes exhibit mosaics of vegetational un. net relate to life f

as trees vs. grasses and taxa (floristic composition, i.e., species pattern), e. 3 see Oosting (1956); Dansereau

(1957); Küchler (1964); Daubenmire (1968): eel (1971). These patterns are plant communities com-

posed of a limited number of life forms and taxa, some of which are more prominent than others, thus, are

recognizable. The presence and proportion of life forms and taxa give a given plant community its recog-

nizable characteristics. Only one life form is usually used to characterize a vegetational unit; but, in some

cases different life forms may intersperse such as in a savannah (trees and shrubs dominate in some areas,

grasses in others). Changes in floristic d ion can Boe a to subdivide a vegetational area. However,

ccepted; thus, vegetational maps based

no vegetational scheme of delineation or

on life forms and taxa are subject to continual interpretation sion: and application (e.g., Gould 1962;

Correll & Johnston 1970; McMahan et al. 1984; Diamond et al. 1987; Edwards et al. 1989; Hatch et al. 1990;

Bezanson 2000; Diggs & Schultze 2003; Griffith et al. 2004; Telfair 2006). Nevertheless, the concept of veg-

etational areas is important and necessary for landscape preservation and wildlife habitat conservation and

management in Texas (Telfair 1999). See Diggs et al. (2006) for examples of most of these maps in reduced

size and color as well as detailed discussion. The Figure 4 map (Telfair's Vegetational Regions of Texas) will

require use of a hand lens or magnifying glass to see some county boundaries and to identify small outlier

J. Bot. Res. Inst. Texas 3(1): 395 — 399, 2009

396 i i Texas 3(

areas of vegetation. However, the map is available online with a zoom feature—http-//artemis.austincollege.

edu/acad/bio/gdiggs/EastT X/introduction.pdf—page 6 (p. 23 of the pdf introduction).

Vegetational patterns reflect the influences of many geographic as well as climatic factors that have

interacted over long periods of geologic time to produce regions and subregions of characteristic vegetation.

In contrast, floristic regions largely reflect only the influence of climatic factors, mainly rainfall and tem-

perature (for comparison, see the 2 maps of vegetational and floristic regions, Turner et al. 2003). However,

vegetational patterns are also subject to many widespread human-caused influences, especially farming and

ranching, water resource development, forestry practices, mineral and energy production, urban/industrial

expansion, recreational/leisure developments, transportation facilities, introduced species, and land frag-

mentation (USFWS 1979; Gunter & Oelschlaeger 1997; Telfair 1999; James 2000).

COMMENTARY

Texas landscape regions and their included plant communities are more diverse than those of any other

state. These regions are defined by their prominent vegetation (vegetation areas) or physiographic features

(ecological regions). In most regions, there is close correlation between vegetational and ecological areas

except where names refer to physical features, e.g., Coastal Sand Plains, Llano Uplift, Rolling Plains, High

Plains, Edwards Plateau, and Trans-Pecos Desertic Basin, Plains, and Mountains. However, most subdivi-

sions within these regions are named for the prominent vegetation therein. These regions are a result of

millions of years of the interaction of geological, climatic, and other geographic influences. They form a

complex mosaic of odd shapes, sizes, and patterns some of which are interwoven. However, their natural

history can be interpreted ecologically.

Spearing (1991) provided maps and analysis of ecological processes that formed the physiography of

Texas. Seven basic geologic processes are involved: 1) mountain building and erosion, 2) basin formation

and filling, 3) uplifts, 4) volcanism, 5) wind erosion, 6) marine influences, and 7) stream entrenchment.

In combination with these geological processes, climatic processes (annual and seasonal rainfall and tem-

perature patterns) produced regional zones that influenced the development of soil associations which, in

turn, influenced the types and patterns of vegetation that developed co-regionally (Griffiths & Orton 1968;

Godfrey et al. 1973).

The combination of rainfall and temperature patterns, in relation to the influence of topography, forms

regional climatic zones of arid to humid, tropical to temperate. Thus, in general, there are 3 major climatic

zones in Texas with distinct vegetation: 1) the eastern 10% (humid, forested), central 80% (semi-humid

to semi-arid (primarily grasslands and savannahs), and 3) western 10% arid (desert vegetation). The major

environmental conditions of these zones are:

Forest.—Major influential environmental factors are precipitation and soil nutrients; fire in some areas

(e.g., longleaf pine and post oak savannah communities). Most nutrients are bound in woody tissues for

many years, thus, sandy forest soils are especially nutrient deficient.

Grassland.—Major influential environmental factors are drying winds, droughts, irregular rainfall,

and lightning-caused fires. Nutrients recycle annually between plants and soils, much being stored in the

soil. In relation to rainfall belts, grasses vary from short to tall along a gradient from west to east.

Desert. —Maj orinfluential l factors are water shor tage and d heat/cold stress. When deserts

formed, local plants adapted and some immigrated from more southern tropical climates.

Other important regional geographic infl rivers, elevations, soils, and standing water (Spear-

ing 1991). River drainages allow bottomland forests to extend far westward along riparian corridors into

grasslands; some smaller drainages also occur in desert areas. The Texas land surface rises gradually from

sea level northwestward to almost 1524 m. Abrupt changes in elevation occur at the Balcones and Caprock

Escarpments. In the Trans-Pecos Region, mountains rise to heights of 1524-2667 m from valley floors of

914-1219 m elevation. These elevation changes influence regional climates which, in turn, influence the

vegetation. Soils are of major influence in the development of growing conditions for vegetation. Worldwide

Telfair, Vegetational areas of Texas 397

there are 12 soil orders of which 9 occur in Texas. These 9 orders, in Texas, include 73 soil associations,

and about 1300 soil series (National Resource Conservation Service 2008). Thus, the great variety of Texas

soils reflects the diversity of geologic, climatic, and biologic conditions during which they developed. Also,

there is a strong correlation with vegetational patterns. Soil Order name derivatives (Godfrey et al. 1973;

Soil Survey Staff 1999) illustrate the basic soil differences and types of vegetation supported.

Alfisols Leached of aluminum and iron; subsurf: y lation; occur in semiarid to moist areas;

primarily in forest are

Aridisols Dry; lack of moisture dios accumulation of salts and minerals; deserts.

Entisols Recent soils of little development in areas of high erosion or deposition rates; occur in dunes,

steep slopes, and flood plains; good examples are Monahans and coastal sand dunes and is-

lands.

Histosols* ^ High fiber tissue content; mostly saturated bogs and marshes

Inceptisols Beginning to moderate development; semiarid to humid slo opes; XD PN flood plains.

Mollisols Soft; dark with dE organic matter; highly fertile; dinem

Spodosols* Ash-colored, acid, inf y subsoil s organic matter; conifer

ous forests of humid ar

Vertisols High in clays; swell Bn wet; crack when dry; d high in natural fertility; grasslands.

Ultisols Well-developed, maximum leached; pine forests

Surface water can provide many types of small, local, and diverse plant communities within each vegetation

region (e.g., swamps, iuc hes oxbow lakes, bogs, seeps, playa lakes).

Some the result of influencing factors that override or dominate other factors. The

Post Oak Savannah (or Clay Pan Savannah) occurs in a forest climate, but developed under the influence of

an edaphic factor (a shallow nearly impervious clay pan restricts water percolation during dry periods); so,

plant diversity is lessened. An unusual exception to the “Clay Pan" soils occurs in Bastrop County—home

of the "Lost Pines”, where there is a sandy inclusion of moist soils—the Carrizo Sands—which support a

unique community of loblolly pines, a western remnant once connected to the now more eastern pine for-

est. These relict pines have developed an ecotype more drought-tolerant than those to the east. Blackland

Prairies occur in a forest climate, but developed under the control of pyric and edaphic factors. The high

clay content of this soil causes it to swell during periods of high rainfall and to shrink during periods of

drought. The shrinkage opens wide, deep crack that damage tree roots. Also, most grassland plants are

fire-adapted; thus, resprout from below-ground root collars rather than the above-ground terminal buds for

most forest plants. The forested areas of the East and West Cross Timbers occur in a grassland climate; but,

deep, moist, sandy soil belts allow trees to dominate. Coastal region vegetation is highly variable (dunes,

maus prairies, and forests). It is influenced by many interacting marine influences (e.g., stream erosion,

meandering, and delta formation; wave and tidal action; and the influence of hurricane winds

and salt EN Wind erosion produced the Monahans Sands in west Texas, and the Coastal Sand Plains

between Corpus Christi and Brownsville in south Texas.

Differential regional blending or transition of environmental factors, the influence of locally control-

ling factors, and the adaptation of vegetation make difficult delineation of boundaries between regional and

subregional areas; and, in some areas, there are inclusions of one area within another (e.g., “islands” within

adjacent regions). Nevertheless, regional and subregional distinctions in vegetation are recognizable and

mappable.

Physical, floral, and faunal regions of Texas were analyzed in detail by the U.S. Fish and Wildlife

Service (1979). This reference is 30 years old, seldom cited, but important. It was prepared as a concept

plan to determine unique wildlife ecosystems of Texas and contains detailed maps and extensive regional

descriptions of flora and fauna with listings of many representative species. The wildlife section analyzes

general zoography, distributional trends, and regional faunas with lists of species of special concern and

threatened and endangered species.

398 Journal of the Botanical f Texas 3(

CONCLUSIONS

The number of plant community types in Texas is dependent upon the classification system used (Bezanson

2000). However, an important result has been not only the identification of vegetation types; but, those that

are both threatened and in need of protection. Thus, although boundaries of vegetational areas are difficult

to assess, their plant communities are recognizable; thus, the concept of vegetational areas is especially

beneficial in floral and faunal conservation and management. However, as proposed by MacRoberts and

MacRoberts (2008), a new vegetational map of Texas based on total flora may provide additional helpful

insights for establishing vegetational areas that better interpret the state's phytogeography.

ACKNOWLEDGMENTS

Iam sincerely grateful to Michael H. MacRoberts (Louisiana State University, Shreveport), Stephan L. Hatch

(Texas A&M University, College Station), and Barney L. Lipscomb (journal editor) who carefully reviewed

the manuscript, corrected errors, and provided helpful comments and suggestions that greatly improved

this paper.

REFERENCES

BEZANSON, D. 2000. Natural vegetation types of Texas and their representation in conservation areas. M.A. Thesis,

Univ. of Texas, Austin.

Corrett, D.S. AND M.C. JoHNsTON. 1970. Manual of the vascular plants of Texas. Texas Research Foundation,

Renner.

Dansereau, P. 1957. Biogeography: an ecological; perspective. The Ronald Press Company, New York.

Dausenmire, R. 1968. Plant communities: a textbook of plant synecolgy. Harper & Row, Publishers, New York.

DiamonD, D.D., D.H. Riskin, AND S.L. ORZELL. 1987. A fi for plant community classification and conservation

in Texas. Tex. J. Sci. 39:203-221.

Dices, G.M., JR., B.L. Liescoms, M.D. REED, AND did id Ban 2006. Illustrated flora of east Texas. Sida, Bot. Misc.

26:1-1594. Available: http:// ad/bio/gdiggs/EastTX/introduction.pdf.

Dicas, G.M., JR AND PC. SE 2003. Soil- -dependent fire frequency: a new hypothesis for the distribution of

prairies and woodlands in North Central and East Texas. Sida 20:1139-1153.

Epwanps, RJ, G. Lonatey, R. Moss, J. WARD, R. MatTHews, AND B. Stewart 1989. A classification of Texas aquatic com-

munities with special consideration toward the conservation of endangered and threatened taxa. Texas J.

Sci. 41:231-240.

GoprnEv, C.L, G.S. Mckee, AND H. Oakes. 1973. General soil map of Texas. Texas Agric. Exp. Sta., Misc. Publ. 1034,

College Station.

GouLn, FW. 1962. Texas plants—a checklist and ecological summary. Texas Agric. Exp. Sta, Misc. Publ. 585:1-

112.

GnirrITH, G.E., S.A. Bryce, J.M. Omernik, J.A. Comstock, A.C. Rocers, B. Harrison, S.L HATCH, AND D. Bezanson. 2004. Ecore-

gions of Texas (color poster with map, descriptive text, and photographs). Scale 1:2,500,000. U.S. Geol. Survey,

Reston, Virginia.

GRIFFITHS, J.F. AND R. Orton. 1968. Agroclimate atlas of Texas — Part |. Texas Agric. Exp. Sta., Misc. Publ. 888, College

Station.

Gunter, PA.Y. AND M. OELSCHLAEGER 1997. Texas land ethics. University of Texas Press, Austin.

HarcH, S.L., K.N. GANDHI, AND L.E. Brown, 1990. Checklist of the vascular plants of Texas. Texas Agric. Exp. Sta., Misc.

Publ. 1655:1-158, College station

James, L. 2000. Fragmented lands: changing land rship in Texas. The Agriculture Program, Agricultural Com-

munication. Texas A&M University System, MKT-3443 15M, 9/00, College Station.

KückLER, A.W. 1964. Manual to accompany the map—potential natural vegetation of the conterminous United

States. Amer. Geogr. Soc, Spec. Publ. No. 36, New York.

Telfair, Vegetational areas of Texas 399

MACMAHAN, C.A., R.G, FRYE, AND K.L. Brown (eds.). 1984. The vegetation types of Texas including cropland. PWD Bull.

7000-120. Tex. Parks and Wildlife Department, Austin.

MacRoserts, M.H. AND B.R. MacRoserrs. 2008. Species richness of vegetational areas of Texas: a first approximation.

J. Bot. Res. Inst. Texas 2:1373-1379.

NATURAL RESOURC 2008. U.S. Department of Agriculture, Temple, Texas. wwwtx.nrcs.usda.gov.

Oostine, H.J. 1956. The study of plant communities. 2nd ed. W. H. Freeman and Company, San Francisco, Cali-

fornia.

SPEARING, D. 1991. Roadside geology of Texas. Mountain Press. Publ. Co., Missoula, Montana.

SHimweLL, DW. 1971. The description and classification of vegetation. University of Washington Press, Seattle.

Soi Survey STAFF. 1999. Soil taxonomy: A basic system of soil classification for making and interpreting soil sur-

veys. Agricultural Handbook 436, 2*1 ed. U.S.D.A. Natural Resources Conservation Service, U.S. Government

Printing Office, Pittsburgh, Pennsylvania.

TELFAIR, R.C. II. (ed.). 1999. Texas wildlife resources and land uses. Univ. of Texas Press. Austin.

TELFAIR, R.C. II. 2006. Map of vegetational regions of Texas (based on dominant plants). Figure 4. In: G.M. Diggs, Jr.,

B.L. Lipscomb, M.D. Reed, and RJ. O'Kennon. Illustrated flora of east Texas. Vol. | (introduction, pteridophytes,

gymnosperms, and monocotyledons). Sida, Bot. Misc. 26: 1-1594.

Turner, B.L, H. NicHoLs, G. Denny, AND O, Doron. 2003. Atlas of the vascular plants of Texas. Vol. 1. Sida, Bot. Misc.

24:1 -888.

U.S. FisH AND WiLDLIFE Service. 1979. Concept plan: unique wildlife ecosystems of Texas. U.S. Department of the

Interior, Region 2, Albuqueraue, New Mexico.

Lel D . ID lL, | PF. £T,

400 Journal of Texas 3(

BOOK NOTICE

SABEEHA MERCHANT, WinsLow R. Briccs, AND DoNatp Ort (eds.). 2009. Annual Review of Plant Biology,

Volume 60. (ISSN 1543-5008; ISBN 978-0-8243-0660-1, hbk.). Annual Reviews, 4139 El Camino

Way, PO. Box 10139, Palo Alto, California 94303-0139, U.S.A. (Orders: www.annualreviews.org,

science@annualreviews.org, 800-523-8635, 650-493-4400). $89.00 indiv., 607 pp., cumulative indices,

7 1/2" x 9 3/8".

Contents of Volume 60:

1. My Journey from Horticulture to Plant Biology—Jan A.D. Zeevaart

2. Roles of Proteolysis in Plant Self-Incompatibility—Yijing Zhang, Zhonghua Zhao, Yongbiao Xue

3. Epigenetic Regulation of Mone Elements in Plants—Damon Lisch

4. 14-3-3 and FHA Domains M phoprot ieee ee Chevalier, Erin R. Morris ou C. Mid

5. Analyzingl pecific V: g Į s—Dan Kliebenstein

6. D from O 11 he Nucleus: The Idi fE biosi Tatjana Kleine, Uwe G. Maier, Dario Leister

7. The HSP90-SGTI Chaperone Complex for NLR Immune Sensors—Ken shite.

8. Cellulosic Biofuels—Andrew Carroll, Chris Somerville

9. Jasmonate Passes Muster: A Receptor and Targets for the Defense Hormone—John Browse

10. Phloem Transport: Cellular Pathways and Molecular Trafficking—Robert Turgeon, Shmuel Wolf

11. Selaginella and 400 Million Years of Separation—Jo Ann Banks

12. Sensing and Responding to Excess Light—Zhirong Li, Setsuko Wakao, Beat B. Fischer, Krishna K. Niyogi

13. Aquilegia: A New Model for Plant Development, Ecology, and Evolution—Elena M

14. Environmental Effects on Spatial and Temporal Patterns of Leaf and Root MP Walter, Wendy K. Silk, Ulrich Schurr

15. Short-Read Sequencing Technologies for Transcriptional Analyses—Stacey Á. Simon, Jixian Zhai, Raja Sekhar Nandety, Kevin P. Mc-

Cormick, Jia Zeng, Diego Mejia, Blake C. Meyers

16. Bi hesis of Plant Isoprenoids: P ives for Microbial Engineering—James Kirby, Jay D. Keasling

17. The Circadian System in Higher Plants—Stacey L. Harmer

18. A Renaissance of Elicitors: Perception of Microbe-Associated Molecular Patterns and Danger Signals by Pattern-Recognition

Receptors—Thomas es Georg Felix

19. Signal Transduction in Response UV-B Radiation—Gareth I. Jenkins

20. Bias in Plant Gene Content Following Different Sorts of Duplication: Tandem, Whole-Genome, Segmental, or by Transposition—

Michael Freeling

21. Pals nas Metabolism: Genes, Mutants, Energetics, and Redox Signaling— Christine H. Foyer, Arnold J. Bloom, Guill Queval

Graham

22. Roles of vis Small RNAs in Biotic S R Virginia Ruiz-Ferrer, Olivier Voinnet

23. i D dm dul = ide Á Scientists Analy sis of the Issues (Part I)—Peggy G. Lemaux

2 ela S. Soltis, Douglas E. Soltis

Cumulative Index Contributing Authors, Volumes 50—60

Cumulative Ind Chapter Titles, Volumes 50-60

J. Bot. Res, Inst. Texas 3(1): 400. 2009

VASCULAR PLANT SPECIES RICHNESS OF THE BIG THICKET, EAST TEXAS

Michael H. MacRoberts and Barbara R. la

Bog Research and Herbarium, A f Life Scie!

Columbia Louisiana State University in Shreveco

Shreveport, Louisiana 71104, U.S.A. Shreveport, Louisiana 71115, U.S.A

ABSTRACT

It has been claimed that the Big Thicket of oca Texas is icd ad species rich when compared to other areas of North

America. Extensive plant Felecdne in the Big Thic data than previously has been available. These data

indicate that the Big Thicl ] g I icl f parabl i United States.

RESUMEN

Se ha dicho q ] Big Thicket del sureste de T: pcional i peci parad de N América

que el Big Thicl imad 1 dia de riq de especies que á hisedel re de los Estados Unid

INTRODUCTION

For many years it has been claimed that the Big Thicket of southeastern Texas is exceptionally rich in plant

species (see Cozine 2004; MacRoberts & MacRoberts 2004; Diggs et al. 2006 for reviews and literature). This

claim has been difficult to test since until recently the Big Thicket region has been botanically understudied

and underdocumented (MacRoberts et al. 2002; MacRoberts & MacRoberts 2007). However, this problem

is now largely overcome by a floristic inventory of the major part of the Big Thicket National Preserve (Mac-

Roberts et al. 2002; Brown et al. 2005, 2006a, 2006b, 2008a, 2008b, in prep a, b), a reexamination of earlier

plant collections from the Big Thicket National Preserve (L.E. Brown pers. comm .), the recent publication of

Atlas of the Vascular Plants of Texas (Turner et al. 2003) and Illustrated Flora of East Texas (Diggs et al. 2006), a

floristic inventory of Walker, Polk, and San Jacinto counties (Nesom & Brown 1998), an inventory of a Polk

County prairie (Brown et al. 2002), and an inventory of the Roy E. Larsen Sandylands Sanctuary in Hardin

County (Matos & Rudolph 1985).

Using this information, we attempt to determine the species richness of the Big Thicket.

SETTING

It has long been realized that the Big Thicket is ill-defined and poorly delineated (McLeod 1971; Cozine

2004; Diggs et al. 2006). There are at least five delineations: 1) the "Broadest Conception" of the Big Thicket

(Diggs et al. 2006), 2) the "Biological Survey" of Parks and Cory (1938), 3) the "Ecological Area" of McLeod

(1971), 4) the "Traditional (Hunter's) Thicket" of the late nineteenth and early twentieth century (Diggs et

al. 2006), and 5) the "Artificial Delineation" (Diggs et al. 2006) (see Diggs et al. 2006 and their figures 88

and 94 for details). We will be concerned only with the Broadest Conception (“Broadest Thicket" hereafter)

and the Artificial Conception (“Artificial Thicket" hereafter) (Fig. 1), the reasons being that few pay attention

to the Traditional Thicket and that the Artificial Thicket basically encompasses both the Biological Survey

delineation and the Ecological Area delineation: the two delineations most frequently used.

The Broadest Thicket consists of part or all of 23 counties, a total of about 39,477 square km (Fig. 1)

that includes the area between the cities of Nacogdoches and Liberty on a north-south axis and the Sabine

River to the Brazos River on an east-west axis. The Artificial Thicket consists of seven and a half counties

(Hardin, Jasper, Liberty [north half only], Montgomery, Newton, Polk, San Jacinto, and Tyler) and totals 17,974

square km (Fig. 1). It is located entirely within the Broadest Thicket area. The Broadest Thicket includes not

only small isolated prairies, palmetto flats, oak floodplain, baygalls, wetland pine savanna, longleaf pine

J. Bot. Res. Inst. Texas 3(1): 401 — 405. 2009

Dat p ID hI bitit, £T,

402 Journal of tt

BROADEST

THICKET |

ARTIFICIAL

114

ICKET

Fic. 1. Locati £ Avtifi-tal Thiel ID lact Thikat in +} Big Thi

upland, beech-magnolia forest, and xeric sandylands of the Artificial Thicket, but also coastal prairie on its

southern edge, blackland prairie and post oak-hickory savanna on its western edge, and pine-oak-hickory

forest to the north (Ajilvsgi 1979; Diggs et al. 2006 and references therein).

The Big Thicket National Preserve is located within the Artificial Thicket and consists of 14 units ranging

from 2.2 square km to 100 square km and totaling 401 square km. Since the units were originally selected

to provide a representative sample of community structure and flora (Peacock 1994; Diggs et al. 2006), a

floristic inventory of the units should constitute a fair sample of the entire flora.

METHODS

Using the following sources (Matos & Rudolph 1985; Nesom & Brown 1998; MacRoberts et al. 2002; Brown

et al. 2002, 2005, 2006a, 2006b, 2008a, 2008b, in prep. a, b; Turner et al. 2003; Diggs et al. 2006), we

determined which species occurred in the Artificial Thicket and which occurred in the Broadest Thicket.

Obviously, if a species occurred in the Artificial Thicket, it also occurred in the Broadest Thicket since the

Artificial Thicket is encompassed entirely by the Broadest Thicket. If, however, a species only occurred in

AarDal I! Ms -DAL " ; Si " il £ + Big TI ich t, Fast Texas 403

Taste 1, Number of Big Thicket plant species.

Artificial Thicket Broadest Thicket

Ferns & Gymnosperms 48 58

Monocots 573 748

Dicots 1144 1549

Total 1765 2355

Native species 1518 1937

the Broadest Thicket outside the Artificial Thicket, it was counted as occurring only in the Broadest Thicket.

Only species were counted: if there were two or more varieties or subspecies, they were lumped.

RESULTS/DISCUSSION

Table 1 gives the number of ferns and gymnosperms, monocots, and dicots in the Artificial Thicket and

Broadest Thicket. The total is 1765 species in the Artificial Thicket of which 1518 are native and 2355 spe-

cies in the Broadest Thicket of which 1937 are native. Monocots account for 32.5 percent of the flora of the

Artificial Thicket and 31.8 percent of the Broadest Thicket.

Diggs et al. (2006, see also Diggs et al. 2003), using earlier sources (up to about 2003), determined that

the Artificial Thicket had 1826 species (native and naturalized) of which 31 percent were monocots. The

figure 1826 is very close to ours and is between ours and the figure for the Broadest Thicket. The figure of

31 percent monocots is again almost identical to ours. The high incidence of monocots is indicative of the

mesic to wet nature of the Big Thicket region of Texas (Diggs et al. 2006; MacRoberts & MacRoberts 2008).

Dryer areas, such as North Central Texas, have fewer monocots (Diggs et al. 2006).

These figures are interesting because they are comparable to figures from other places at the same or

approximately the same latitude in the southeastern United States. For example, the Florida Panhandle

consists of 38,628 square km and has 2359 species, of which 31 percent are monocots, and 1989 are native

(Clewell 1985). It is therefore al identical in size, species number, and percent monocots to the Broad-

est Thicket. The Artificial Thicket is also comparable to other areas of equal size. For example, Bienville,

Bossier, Caddo, Claiborne, Jackson, Lincoln, Ouachita, Union, and Webster parishes in northern Louisiana

taken together consist of 16,601 square km and have approximately 1780 species, 30.1 percent of which are

monocots (MacRoberts unpublished data developed from MacRoberts & MacRoberts 2006 and Thomas &

Allen 1993—1998). However, the Florida Parishes of Louisiana (East Baton Rouge, East Feliciana, Livingston,

West Feliciana, St. Helena, St. Tammany, Tangipahoa, and Washington), 200 km due east of the Big Thicket,

consist of only 12,129 square km but have 1705 native species, of which 33 percent are monocots. If exotics

are added, the total number probably would be between 2000 and 2100 species (MacRoberts unpublished

data developed from Thomas & Allen 1993-1998).

These findings support our previous analysis of species/area across the south, in which we used the

Diggs et al. (2006) figure of 1826 species, and found that the Big Thicket was not exceptional in species

numbers but is about average for the southeastern United States (MacRoberts et al. 2007). Thus, our current

analysis confirms our previous conclusion. Although the data are not available, it may transpire that the

Big Thicket is slightly below average for similar sized areas at the same latitude in the southeastern United

States. The data for the Florida Parishes of Louisiana suggest this, but until other areas such as southern

Mississippi, Alabama, Georgia, and northern Florida are sampled, no conclusion is possible.

While we are in process of redefining the Big Thicket, it should be remembered that the Big Thicket is

important to preserve because hundreds of southeastern species reach their range limit in the Big Thicket.

The Big Thicket is, therefore, a boundary area and is important for a variety of reasons: boundary plants

are often genetically unique and peripheral populations are often more sensitive to environmental change

and can act as ecological indicators (Diggs et al. 2006). But, the most important reason for considering the

3 L af ob D . D h Im etit ET

404 Journal of t l Texas 3(

Big Thicket immensely valuable was made by Donovan Correll years ago before the Big Thicket became a

preserve. Testifying before ine Senate committee in Beaumont in 1970, he pointed out that the vast pine

hardwood forest that cl 1th theastern United States had no areas that were preserved. He said:

“We ought to have a representation of the great Southeastern pine hardwood forests somewhere, and since

the development is so optimum here in Texas,” then the Big Thicket would be ideal for that representation.

ACKNOWLEDGMENTS

Larry Brown provided many useful comments on an earlier version of this paper including his personal ob-

servations on species richness on the southeastern United States. George Diggs and an anonymous reviewer

had a number of valuable comments about the manuscript.

REFERENCES

Auvsal, G. 1979. Wild flowers of the Big Thicket. Texas A&M University Press, College Station.

Brown, L.E., HILLHouse, K., B.R. MacRoserts, AND M.H. MacRoserts, 2002. The vascular flora of Windham Prairie, Polk

County, East Texas. Texas J. Sci. 54:22/—-240.

Brown, L.E., B.R. MacRoserTs, M.H. MacRoserts, PA. Harcomee, 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, Texas. Sida 21:1807-1827.

Brown, L.E., B.R. MacRoserts, M.H. MacRoserts, PA. HARCOMBE, W.W. Pruess, I.S. ELsik, AND S.D. Jones. 2006a. Annotated

checklist of the vascular flora of the Big Sandy Creek Unit of the Big Thicket National Preserve, Polk County,

Texas. Sida 22:705-723.

Brown, L.E., B.R. MacRoserTs, M.H. MacRosenrs, PA. HARCOMBE, W.W. Pruess, I.S. ELsik, AND S.D. Jones. 2006b. Annotated

checklist of the vascular flora of the Lance Rosier Unit of the Big Thicket National Preserve, Hardin County,

Texas. Sida 22:1175-1189.

Brown, L.E., B.R. MacRoserts, M.H. MacRoserts, PA. Harcomse, W.W. Pruess, l.S. Esik, AND S.B. WaLker. 2008a. Annotated

checklist of the vascular flora of the Beech Creek Unit of the Big Thicket National Preserve, Tyler County, Texas.

J. Bot. Res. Inst. Texas 2:651-660.

Brown, L.E., B.R. MacRoserts, M.H. MacRoserts, W.W. Pruess, I.S. ELsik, AND S.B. Waker, 2008b. Annotated checklist of

the vascular flora of the Loblolly Unit of the Big Thicket National Preserve, Liberty County, Texas. J. Bot. Res.

Inst. Texas 2:1481-1489.

Brown, L.E., B.R. MacRoserts, M.H. MacRogerTs, W.W. Pruess. In prep. a. Annotated checklist of the vascular flora of

the Menard Creek Unit of the Big Thicket National Preserve, Polk, Liberty, and Hardin counties, Texas.

Brown, L.E, B.R. MacRoserts, M.H. MacRoserts, W.W. Pruess, AND IS. Etsi. In prep. b. Annotated checklist of the

vascular flora of the Jack Gore Baygall and Neches Bottom units of the Big Thicket National Preserve, Jasper

and Hardin counties, Texas.

CueweLt, A.F. 1985. Guide to the vascular plants of the Florida Panhandle. University Presses of Florida, Gainesville.

Cozine, J. 2004. Saving the Big Thicket: From exploration to preservation, 1685-2003. Univ. North Texas Press,

Denton.

Dicas, G.M., R.J. GEORGE, AND L.E. Brown. 2003. Vascular plant species richness of the Big Thicket of East Texas. Big

Thicket Science Conference 2003, Abstract pp. 13.

Dias, G.M., B.L. Lirscome, M.D. Reen, AND R.J. O'KENNON. 2006. Illustrated flora of east Texas. Sida, Bot. Misc. 26:1—1594.

MacRoserrs, B.R, M.H. MacRosnrs, AND L.E. Brown. 2002. 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, 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.

MacRoserts, M.H. AND B.R. MacRoserTs. 2004. The Big Thicket: typical or atypical. East Texas Hist. Assoc. 42:42-51.

MacRoserTs, M.H. AND B.R. MacRoserts. 2007. Phytogeography of the Big Thicket, east Texas. J. Bot. Res. Inst. Texas

2:1149-1155.

AacRoberts and MacRoberts, Species rict f the Big Thicket, East Texas 405

MacRoserts, M.H. AND B.R. MacRogerTs. 2008. The Big Thicket as floristically unique habitat. J. Bot. Res. Inst. Texas

2:665-67 1.

MacRoserts, M.H., B.R. MACROBERTS, AND R.G. KaLinsky. 2007. Vascular plant species/area relationships (species rich-

ness) in the West Gulf Coastal Plain: A first approximation. J. Bot. Res. Inst. Texas 1:577-583.

Matos, J.A. AND D.C. Rupo.PH. 1985. The vegetation of the Roy E. Larsen Sandylands Sanctuary in the Big Thicket

of Texas. Castanea 50:228-249,

McLeon, C.A. 1971. The Big Thicket forest of East Texas. Texas J. Sci. 23:221-233.

NESOM, G.L. AND L.F. Brown. 1998. Annotated checklist of the vascular plants of Walker, Montgomery, and San

Jacinto counties, east Texas. Phytologia 84:107-153.

Parks, H.B. AND V. Conv. 1938. Biological survey of the East Texas Big Thicket area. Texas Agric. Exp. Sta., College

Station.

Peacock, H.H. 1994. Nature lover's guide to the Big Thicket. Texas A&M Press, College Station.

THomas, R.D. AND C.M. ALLEN. 1993-1998. Atlas of the vascular flora of Louisiana. Louisiana Department of Wildlife

and Fisheries, Baton Rouge.

TURNER, B.L., H. NichoLs, G. DENNY, AND O. Doron. 2003. Atlas of the vascular plants of Texas. Sida Bot. Misc. 24:1-888.

406 tani i Texas 3(

BOOK NOTICE

Hu REN-LIANG, WANG YOU-FANG, AND MansHALL R. Crosby (eds.-in-chief), with Si Hz (ed.). 2007. Moss Flora

of China, Vol. 7. English Version. Amblystegiaceae — Plagiotheciaceae. (ISBN Vol. 7: 978-1-

930723-77-1, hbk.). Science Press (Beijing) and Missouri Botanical Garden Press (St. Louis), 4344 Shaw

Boulevard, St. Louis, Missouri 63110-2291, U.S.A. (Orders: www.ml org, mbgpress@mobot.

org, 314-577-9534, 314-577-9594 fax). $85.00, 258 pp., 8 5/8" x 11 1/4".

From the D DNA s Moss Flora of ching; cia Version, is a seri ling with all known to be na-

identification keys, AMA M of Chi demi d East Asian taxa, habitat

adn listributional rang id f China, p d distributi ps of all pted The present volume

1 blished in tl ies. It treats 41 genera and 188 specifi linfraspecifi including binati

The last volume of the series, Volume 5, treating such | ] ick Leucodontaceae, Pterobryaceae, Meteoriaceae,

and Neckeraceae will be completed by the end of 2009.”

Contents:

Amblystegiaceae E Wu Yu-huan, Gao Chien, and Li Wei

Helodiaceae —by S

Brachytheciaceae i Wang You-fang and Hu Ren-liang

Entodontaceae —by Hu Ren-liang and Wang You-fang

see —by Li Deng-ke

bec d —by Li Deng-ke and Robert R. Ireland

Literature Cited

Index to Latin Names

Index to Chinese Names

J. Bot. Res. Inst, Texas 3(1): 406. 2009

A CLASSIFICATION OF GEOGRAPHIC ELEMENTS AND

ANALYSIS OF THE FLORA OF BIG BEND REGION OF TEXAS

Anna Saghatelyan

Department of Biology

McMurry edid sind Station

Abilen

ABSTRACT

oi ae o Pa Big Bend en in the Trans-Pecos Tepe oe ee uu uisi was studied ue a BU CD eque $

America

distribution of species were obtained from ie Weise the "Synthesis," aud other on-line databases. There are oe native e species

592 genera and 125 families in the flora. These species, based on their contemporary distribution outlines, 19

EN a elements. The genera were classified into 19 geographic groups based on their general distribution. Herein is

presented a aes scheme of probrapmic een b DER ei ME i o iol, Rd the geoelement descriptions,

and the analysis of the flora. The Bi g y ; Madrean and Boreal—has a

complex i of connections, sath predominance of th htl i oe from the Madro-Tertiary geoflora

stalk. Laurasin (Arcto-Tertiary) flora derivatives play a ad ii The strongest connections of the flora are AN Mesoamerica-South

y. play 8

America. The Madrean species (6696) and o the Ch g p

P and western North American, fol y sul ical hl ] th h and east

y tror F g I

g E ] Nortl Hemisphere' i he Atlantic, show three different patterns:

northern ea rly Ti i igrati fthel i hvll Aivereifieatinn nf alder Madre Teil 1 ot 1

J pical g 8

the Tethys nod a d migration westward along the spatter T Teths h f phyll ient t riginated or preserved in

RESUMEN

La flora de la región de Big Bend en el área de vegeta a ioe ee tudió desde la perspectiva biogeografi

Pri t + a tä] de la fl € "n American I Fl ra y j bI 1 la

distribución general d las especies, is "Synthesis," y otras bases de datos on-line. Hay 1587 pue nativas en 592 shes y 125 familias

n la flora. Estas especies, según sus perfiles de distribución contemporánea, se clasificaron en 19 elementos geográficos (floristicos).

Los géneros se clasificaron en 19 grupos geográficos según su distribución Bere n se ose un esquema de clasificación de

elementos geográficos (geoelementos), el catálogo acompañado por las d ntos, y el análisis de la flora. La

flora de la region Big Bend—en la frontera de = on el oe x d Bota iene un p SO de conexiones, con

predoininio m fíti Mo Las d de la flora Laurasina

(Arcto-Ti i pel I i As fi Je la fl Sur América 1

nas (66%) y BE demi Chihuahuanos (2696) son los más numerosos. En la composición genérica, m géneros del

J 1 i 1 f. T LT 1 4 1 t J 1 1 J l4l

+ ED o r E r J T 1 F

: E O E E eee ^c del Atlántico, lik ]

r oO

norte en el Terciari de gé 1 : afi As f eamAn do] 1 ul MR ee ete

a lo largo de de a ruta del Tetis, y y migración hacia el oeste a lo largo de la costa sur ddl Tetis de bata taxa viejos moles. nados o

preservados en g frica

INTRODUCTION

The Trans-Pecos region of Texas or West Texas Vegetational Area (Correll & Johnston 1970) is distinct

from that of surrounding areas in Texas because of its numerous mountain systems and low arid basins

with elevation ranges from 305 to 2388 m (1000-7835 ft) (Powell 1998). It lies on the northern edge of

the Chihuahuan Subprovince of the Sonoran Province of Takhtajan’s floristic system (Takhtajan 1986) and

thus on the boundary of two Subkingdoms of the Holarctic Kingdom: the Boreal and Madrean. An analysis

of such flora can provide a good opportunity to reveal proportions of different floristic elements, to look

for Arcto-Tertiary and Madro-Tertiary (Axelrod 1958; Raven & Axelrod 1987) relicts, and to specify the

J. Bot. Res, Inst. Texas 3(1): 407 — 441. 2009

408 I lafal Dat tani D h Inetiti t f Texas 3(1)

level and areas of endemism (Platnick 1991; Morrone & Crisci 1995). Northern and eastern limits of the

Madrean region are difficult to delineate and they were subject of debate in the North American literature

(Good 1974; Cronquist 1982; Morrone et al. 1999; McLaughlin 2007; see McLaughlin 2007:31 for com-

parison; also Katinas et al. 2004; and Fenstermacher et al. 2008). The Northern Madrean boundary is a

composite area were different distribution tracks overlap and partial floras of different types of vegetation

in the mountains represent historical elements of different origins, affinities, and age, both relictual and

progressive, autochthonous, and migrational.

The flora of three counties in the Big Bend region—Brewster, Presidio, and Jeff Davis—was chosen to

test the floristics in the Chihuahuan Subprovince. A major objective is to reveal the taxonomic proportions

and biogeographic affinities of the plant taxa in the region.

MATERIALS AND METHODS

Using primarily the Synthesis of the North American Flora (Kartesz & Meacham 2002; Kartesz 2008), a check-

list of the flora of Brewster, Presidio, and Jeff Davis counties in Trans-Pecos was compiled. The non-native

species were excluded. Several new species were added to the checklist draft after it was compared with the

one of the Dead Horse Mountains, Big Bend National Park (Fenstermacher et al. 2008). Taxonomic counts of

the families, genera, and species were performed. The distributional and other data were obtained from the

Synthesis of the North American Flora (Kartesz & Meacham 2002), Tropicos (Tropicos.org.), Flora of North

America, Digital Flora of Texas databases, the Onagraceae website (Wagner & Hoch 2005), the literature

(Correll & Johnston 1970; Powell 1998; Turner et al. 2003; Diggs et al. 1999; Villarreal 2001), and other

sources. The distribution outlines of all the species in the flora were studied. Congruent distributions of two

or more species were named after well known chorionomic units (Takhtajan 1986; Thorne 1993; Rzedowski

1978) or in geographic terms. Each species was thus referred to and treated as a particular geographic ele-

ment, or geoelement (Saghatelyan 1997a, b) of the flora. All the species were classified into 19 geoelements

according to their general distributional patterns revealed during the current study. The distribution data

on the genera were retrieved from Wielgorskaya (1995) and Mabberley (1997). The genera were classified

into 19 groups based on their distribution outlines.

The species list (Appendix 1) for the above-mentioned counties was prepared with the major objective

of defining the geoelements individually and collectively represented. Proportions of geoelements in a flora

are robust characteristics which are not sensitive to minor nomenclatural changes. "Good species" always

have "good ranges" and they usually serve as a basis of biogeographic analysis. The ranges of some species

could not be referred to a particular geoelement; they are noted with a question mark in the checklist and

are omitted from the analysis. Major outcomes expected from the analysis are:

1. Proportion of the species and genera of the Boreal and Madrean Subkingdoms and their provinces and

subprovinces in the flora.

2. Northern temperate versus southern subtropic & tropical connections of the flora.

3. Proportion of the species confined to one, two, or more subprovinces of the Sonoran Province.

4. The weight of the northern Madrean species in the Madrean element of the flora.

5. Connections of the Madrean and Tethyan subkingdoms.

6. Major migrational tracks.

RESULTS

A CLASSIFICATION SYSTEM OF GEOGRAPHIC ELEMENTS AND THEIR PROPORTIONS IN THE

BiG BEND REGION FLORA (TABLE 1)

Abbreviations: C—center; TX-Texas; CA- California; CO-Colorado; AZ-Arizona; OK-Oklahoma; MO-

Missouri; WO-Wyoming; SMO-Sierra Madre Oriental; Rocky M-Rocky Mountains Province; Mont-

montane; US—United States; Mesoam-Mesoamerican; Gulf Coast—Atlantic and Gulf Coastal Plain Province;

Tr—Pecos—Trans-Pecos; J Davis—Jeff Davis; c-county; cc-counties, Warm—warm temperate.

r g kien! nl " £L a £D:AD J gi Texas 409

Saahatelvan

» y r

Taste 1. Proportions of geographic elements in the Big Bend flora.

Geoelement Subelement Number of species

Madrean 192

Sonoran-Chihuahuan 80

Sonoran 82

Chihuahuan 253

Chihuahuan-Tamaulipan 50

SW North American 149

CE Mountain-Madrean 20

SWC US & SWC N American 64

pce US & SW N American 65

W US/W N American 133

WUS/W N American 76

WC US/WC N American 29

S Great Basin-Sonoran/Chihuahuan 9

Amphitropical 19

STX Endemic 101

Prairie 88

EN American 37

Comanchian/SC US 45

N American 98

Mesoamerican 84

Amer 101

American wide 96

Western American 5

Tropical/Subtropical 32

American-African 6

Tropical/Subtropical 26

Holarctical 21

Polichorous 14

Not established 25

Total 1587

1. Pon NU ERU on ida continents. There are 14 species (sp.) of herbaceous wetland

), and weedy (Plantago major) plants of this geoelement

in the fioi

2. Holarctical: wide ranges in north temperate latitudes of the New and Old World; 21 mesophytic species.

Among them are 6 sp. of grasses (Bromus ciliatus), 2 sp. of horsetails (Equisetum hyemale), and a fern

(Asplenium trichomanes). Other large north temperate genera have just 1—2 sp. each (Campanula rotun-

difolia, Ranunculus sceleratus, and Artemisia campestris) in the flora.

3. Trop/Subtr: wide distribution in tropical and d latitudes; oa cee vo aquae and

this

wetland species, especially of herbaceous habit (Thypha domingensi

group. There are also several shrubs (Sapindus pM Parhinsonia coa and caca farnesiana) of

pantropical or subtropical distribution. Two sub-elements of Trop/Subtr element are listed below.

3a. American-African: the previous type restricted to America and Africa. Seven species: three of

grasses (Echinochloa crus-pavonis), three of sedges (Cyperus squarrosus), and one fern (Cheilanthes

bonariensis).

3b. Warm Temperate/Subtropical: mostly in subtropical and warm temperate regions. Only 11 sp.

of grasses (Digitaria sanguinalis) are in this group.

4. American: wide distribution in the Americas; 101 sp. Here are mostly subtropical weedy grasses (29 sp.:

Bouteloa barbata) and sedges (8 sp.: Cyperus seslerioides), as well as temperate Asteraceae (6 sp.: Conyza

410 tani i Texas 3(

canadensis). Western American (Cordilleran) genera have six species (Muhlenbergia rigida, Epilobiium

ciliatum), while 5 species of Amaranthus and Heliotropium are mostly Caribbean.

4a. American Trop/Subtr: widely distributed in tropical and subtropical (warm temperate) parts of

the Americas. This sub-element has 30 of the 101 sp. of the American element with the grasses

(Cenchrus myosuroides) being especially numerous. Other examples are Tillandsia recurvata, So-

lanum elaeagnifolium, Phyla nodiflora, and Ipomoea cardiophylla.

5. North American: more or less wide ranges in temperate regions of North America; 98 mesophytic species.

They either belong to widely north temperate genera (Maianthemum racemosum, Carex hystericina, and

Vicia americana), north American genera (Solidago gigantea, Lobelia cardinalis, Monarda fistulosa), or cos-

mopolitan genera with large sections in temperate latitudes ee pia! Herbaceous habit,

especially in the grasses (18 sp.), sedges (8 sp.), Asteraceae, and Eug (6 sp. each) predominate.

A few older montane woody species have interesting ranges: either absent in ie south-eastern and

south-central region (Prunus virginiana), poorly represented in the western and absent in the central

(Cephalanthus occidentalis) region, or those that tend to the Rocky Mountain (Cheilanthes feei) Region.

6. East North American: wide ranges in the Atlantic North American Region of Cronquist (1982). Repre-

sentatives of ancient Laurasian genera are essential among the 37 sp. of this geoelement (Ostrya virgini-

ana, Carya ilinoensis, and Clematis pitcheri). A few species however have tropical connections (Cocculus

carolinus, Celastrus scandex, and Nothoscordum ENSE

6a. Gulf Coast-(Caribbean): with rang

6b. Appalachian: with ranges in the namesake province (Osa virginiana, Melica nitens).

7. Comanchian/ SC US: Edwards Plateau westward to the Trans-Pecos extending northward into central

Oklahoma or further to Ozark Plateau, southward into northeastern Mexico and western Lousiana,

mainly in the limits of the Comanchian Subprovince of McLaughlin (2007). There are 46 species in

this geoelement like Juniperus ashei, Lupinus texensis, and Scutellaria wrightii, with some having disjunct

relictual ranges (Leptopus phyllanthoides).

8. Prairie: wide ranges in the North American Prairies Province 2: Deals dl (1982); a PSY herbaceous

species. Biggest group of 24 sp. is in Ast (Liatris 1 ; DE aaa)

in the Heliantheae s.l. (Panero & Funk 2002). P (Bouteloua denia des Sched 1

and papilionaceous Fabaceae (Astragalus lotiflorus, Dalea aurea) have 7 sp. each. Beds EUR

are in large temperate genera (Oenothera triloba, Salvia azurea) with big centers of diversity in western

America and Mexico.

8a. S Prairie/SC North American: from Central and South Texas extending northward to adjacent

states and southward to northern Mexican Plateau (Eryngium leavenworthii, Argythamnia humilis,

Rhus lanceolata, and Gaillardia suavis).

9. W North American: wide ranges in the entire western North America or its parts mostly north of Mexico.

Among the 133 sp. of this geoelement 28 sp. of Asteraceae (Brichelia californica, Erigeron divergens)

prevail, followed by 18 P of Poaceae (Agrostis voe iani and 9 P of Brassicaceae (Stanleya pinnata).

Western American genera} , Cryptantha cinerea) including

the dominants in different types of communities. ponle e from those of montane forests (Pinus

ponderosa, Quercus gambelii, and Cercocarpus montanus), Rocky Mountain and Madrean woodlands and

shrublands (Juniperus scopulorum, Rhus trilobata, Holodiscus dumosus), to widely distributed xerophytic

(Opuntia polyacantha) as well as riparian (Salix exigua) species.

9a. WC US/ WC North American: includes western mountainous (not Pacific) region and western

half of the Prairies Province in the United States/or southward to central Mexico; this element

supports S. McLaughlins (2007) Western Region. Of the 133 sp. of W North American element,

29 sp. are in this sub-element. They iu to m npe genera (Astragalus, Cirsium, and

, Oenothera albicaulis), or a few tropical

[Th ahat qb. Aulas

Lithospermum), North American genera (

(Heliotropium convolvulaceum) genera.

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Fic. 1. Distribution of Yucca baccata. SW N American element. Source:

Tropicos.org. Missouri Botanical Garden. 27 Mar 2009 «http://www.

tropicos.org>.

, Texas 411

Fic 2. Distribution of f M | t. Source

Tropicos.org. Missouri Botanical Garden. 27 Mar 2009 <http://www.

tropicos.org>

Fic 3 Nietrihitian at alice, AA Mas Fun el +

16 4. Distributi Ef les Bn € h Darlin, AA + ye ES

Source: Tropicos.org. Missouri Botanical Garden. 27 Mar 2009 <http://

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element. Source: Tropicos.org. Missouri Botanical Garden. 27 Mar 2009

<http://www.tropicos.org>.

Fic & Dietributi £D : EMad 1 c T,

Missouri Botanical Garden. 27 Mar 2009 «http://www. tropicos, or.

Fic. 6. Distribution of Peganum mexicanum. Sonoran-Chihuahuan element.

Source: In E Missouri Botanical Garden. 27 Mar 2009 <http://

www.tropicos.o

10. SWC North American/ SWC US: more southern/eastern than the previous geoelement: from Colorado

Plateau, southern Rocky Mountains and south-western part of the Prairies Province southward to the US

border or Sierra Madre O

idental/ tern central Mexico. Among 64 species of this element those

of western and south-western North American genera predominate. The grasses (11 sp.), Asteraceae

(9 sp.), and Fabaceae (5 sp.) are followed by species of typical American desert genera in Agavaceae

(Nolina texana), Hydrophyllaceae, Onagraceae, Cactaceae, etc. Several species of this element are of

woody habit (Quercus mohriana, Lonicera albiflora).

412

11. SW North American/ SW US: southern part of the Rocky Mountain Province, Colorado Plateau, southern

and eastern Great Basin, southeastern California (not always), both Mohavean and northern Sonoran

Subprovinces of the Cronquists (1982) system eastward through New Mexico to southwestern Texas and

southward to adjacent northern Mexico. A group of 65 xerophytic species including 5 out 7 species of

Hydrangeaceae of the Big Bend flora, leido nios Yucca baccata (Fig. 1), Opuntia phaeacantha and

other desert dominants. All herl f desert genera (Mentzelia multiflora, Hachelia

pinetorum, 9 sp. each of Poaceae and seais etc.). There are two subtypes:

11a. Apachian: from central to southeastern Arizona, western New Mexico, to northeastern Sonora, and

northwestern Chihuahua as delineated by McLaughlin (2007). Among species of the southwestern

element 20 sp. are in this sub-element (Penstemon jamesii, Boerhavia torreyana, and Phlox nana).

11b. S Great Basin-Sonoran-Chihuahuan: ranges from the southern half of the Great Basin Prov-

ince southward into Sonoran and Chihuahuan Subprovinces of Cronquist (1982). Nine species

are restricted to this area, mostly those of south-western genera (Pennellia longiflora, Abronia

angustifolia).

12. Amphitropical: disjunctive ranges in warm temperate deserts of the western North and South America;

20 species. Among them are 6 sp. of grasses, 3 sp. each of Fabaceae and Asteraceae and one in each of

other desert genera (Aloysia gratissima, Kallstroemia parviflora, and Mentzelia albescens.)

13. Madrean wide: ranges embracing Takhtajan’s (1987) Madrean Region in some cases excluding the Cali-

fornia Province (Figs. 2, 3). Most spectacular group of 192 species with the highest number of woody

species-dominants in various types of mountain and desert communities. The prevailing families are:

Asteraceae (30 sp.), Poaceae (24 sp.), Fabaceae (15 sp.), Euphorbiaceae (15 sp.), and Pteridaceae (11

sp.). Very important are 4 sp. each of Cupressaceae, Rhamnaceae and Oleaceae, 3 sp. of Zygophyllaceae,

including Larrea tridentata, 2 species of Ephedra, Juglans major, Garrya ovata, Rhus virens and Morus

microphylla. About 30 species of the flora with wide Madrean distribution are absent in California and

most of the Great Basin Provinces. The madrean element has following subelements.

13a. S Rocky Mountain-Madrean: along the Rocky Mountains from Colorado Plateau southward

to Sierra Madre Occidental Province or further into the Madrean mountains (Fig. 4). There are

20 species in this sub-element mostly of Asteraceae (5 sp.), Brassicaceae (3 sp.), and Malvaceae

(3 sp.). The species of western American genera (Castilleja integra) predominate, however a few

species are of tropical/warm temperate genera (Cleome multicaulis).

13b. East Madrean: mostly in the Sierra Madre Oriental Province of Morrone et al. (1999) extending

northward to the Edwards Plateau and mountains in Trans-Pecos (Brewster, Jeff Davis, and Pre-

sidio counties). The following species highlight this sub-element: Rhus virens (Fig. 5), Fraxinus

cuspidata, E greggii, Croton incanus, Euphorbia bifurcata, Dalea gerggii, D. frutescens, Centaurium

calycosum, Oenothera calcicola, O. primiveris, and 6 sp. of Pteridaceae.

13c. North Madrean: ranges in northern, mostly US part of the Madrean Region of Takhtajan (1987),

southward to Mexican part of the Sonoran Subprovince and south-western Texas. This sub-

element has 57 sp. of the madrean element in the genera with following ranges: south-western

North American (Wislizenia refracta), central-to tropical American (Matelea parviflora), mostly N

American (Symphoricarpos palmerii), western N and S American (Hedeoma nana), and tropical

subtropical American (Proboscidea parviflora, Boerhavia wrightii). Several species are in cosmopoli-

tan (Chamaesyce, 5 sp.) and north temperate (Eriogonum, 3 sp., Galium) genera.

14. Mesoamerican: wide ranges in Mesoamerica and Caribbean Region extending to southern United States

and northern South America; 83 species. Most of them are among the Asteraceae (17 sp.), grasses (9

sp.), legumes (8 sp.), and ferns (5 sp.). The species of Mesoamerican-Madrean (Bouvardia ternifolia),

Mesoamerican-S American (Margaranthus solanaceus), tropical (Cyclanthera dissecta) and subtropical

American (Macroptilium gibbosifolium), as well as widely American (Oenothera hexandra) genera prevail.

Two species, Arbutus xalapensis and Helianthemum glomeratum, are in the Madro-Tethyan genera.

Saghatelyan, Geographical el ts of the fl f Big Bend region, Texas 413

Fic, 2 Dictributi £I hdi, £ hil

Fic. 7. Distribution of Condalia ericoides. Chihuahuan element. Source: Tamaulipan

Tropicos.org. Missouri Botanical Garden. 27 Mar 2009 «http://www. ^ element. Source: Tropicos. org. Missouri Botanical Garden. 27 Mar 2009

tropicos.org>. <http://www.tropicos.org>.

14a. Mesoamerican-Madrean: wide ranges in the Madrean Region and Mesoamerica southward to

Costa Rica. Of the Mesoamerican element, 19 sp. are in this subelement (Quercus rugosa, Chamae-

syce villifera, and Desmodium psilophyllum).

15. Sonoran wide (or Sonoran Province): embracing all subprovinces of the Sonoran Province of Cron-

quist (1982); 82 xerophytic species including many dominants of the sonoran desert communities. The

legumes (8 sp.: Calliandra humilis), Malvaceae, and Nyctaginaceae (6 sp. each) prevail among them.

Especially characteristic are those of western American (Aloysia wrightii, Garrya wrightii) and SW N

American (Chilopsis linearis) genera, and some of more widely American genera (Verbena perennis). Also

very important are four species each of Cactaceae (Opuntia) and Rhamnaceae (Condalia ericoides), as

well as 2 sp. of Rutaceae (Choysia dumosa).

16. Sonoran-Chihuahuan: ranges in the Sonoran (+Mohavean) and the Chihuahuan (mostly western part)

Subprovinces of the Sonoran Province (Figs. 6, 7). Of the 80 xerophytic species of this element, 18 sp.

are in Asteraceae (Acourtia nana), while Sonoran wide element has only three species of this family. The

6 sp. of Cactaceae (Mammillaria grahamii), 5 each of legumes (Dalea wrightii) and ferns (Cheilanthes vil-

losa), 4 sp. each of grasses (Muhlenbergia arenacea) and Polygala are followed by genera with 1-2 species

(Condalia viridis, Peganum mexicanum, Kallstroemia hirsutissima, and Salvia henrii).

17. Chihuahuan: ranges in the Chihuahuan Subprovince including Edwards Plateau (Fig. 8). This richest in

the flora geoelement has 253 species which reveals the high level of Chihuahuan endemism and diver-

sification in temperate and tropical-subtropical genera. There are 44 sp. of Asteraceae, 24 sp. of cacti,

18 sp. of legumes, 15 sp. of grasses, 12 sp. each of Boraginaceae (mostly in Heliotropium and Tiquilia)

and Brassicaceae, 8 sp. each of Nyctaginaceae and Lamiaceae, 7 sp. each of Agavaceae, Acanthaceae, and

Euphorbiaceae, 6 sp. of Convolvulaceae, 5 sp. each of Fagaceae, Scrophulariaceae and Amaranthaceae,

4 sp. each of Caryophyllaceae, Solanaceae, Polemoniaceae, etc. There are fewer shrubs and small trees

nudae speciosa, Diospyrüs oe ll minus, Holacantha stewartii, and Colubrina texensis)

but they dominate ities. Typical are the rosette forming lilioids (Dasylirion

leiophyllum, Nolina erumpens Agave and Yucca) and cacti (Echinocereus, Escobaria, etc.).

17a. North Chihuahuan: restricted to northern, mostly Unites States portion of the Chihuahuan

Subprovince southward to Coahuila, Mexico. There are 35 sp. in this subelement (Drymaria

pachyphylla, Phyllanthus ericoides, and Hedeoma pucaid

18. Chihuahuan-Tamaulipan: ranges in the Chihuahuan Subp tendi d to the Tamaulipan

Subprovince. Among 50 sp. of this geoelement 8 sp. are in Asteraceae, nowever they are of different

genera than those of the chihuahuan element. The tropic/subtropical genera in this element are more

pronounced (Forestiera angustifolia, Guajacum angustifolium, and Passiflora tenuiloba).

18b. Tamaulipan-Chihuahuan: ranges in the Tamaulipan Subprovince extending westward to the

414 J t ical i Texas 3(

Chihuahuan Subprovince. There are 5 sp. in this sub-element (Agremone sanguinea, Cynanchum

barbigerum, Ayenia pilosa, Meximalva filipes, and Thelesperma ambiguum).

19. South Texas-Coahuila: a part of the Chihuahuan element with narrow ranges in SW Texas-(Edwards

Plateau), and Coahuila- iid biu of ao Mexico. This is an interesting group of 103 species

(for example, Lecl f hoxylum parvum) which deserves a separate analysis.

A few subtypes are given below:

19a. South Texas Endemic: mainly in Edwards Plateau extending to Trans-Pecos and along the

Rio-Grande valley to N Tamaulipan Subprovince; 8 species (Lithospermum mirabile, Chamaesyce

angusta).

19b. Trans-Pecos-Coahuila: endemic or subendemic to Trans-Pecos and Coahuila; 29 species (Quercus

carmenensis, Fendlera rigida, Solanum leptosepalum, Eriogonum hemipterum, Phacelia robusta, and

Selaginella viridissima).

19c. Trans-Pecos (SW) Texas Endemic: endemic to the Trans-Pecos region or parts of it; 29 species

(Prunus havardii, Eriogonum suffruticosum, and Selinocarpus parviflorus).

19d. Brewster c Endemic: endemic to the Brewster county or only Chisos Mountains; 10 species

(Hectia texensis, Quercus tardifolia, and Bouteloua hayi).

TAXONOMETRIC ANALYSIS OF THE FLORA

1. FAMILIES

There are 1587 species in 592 genera, and 125 families of vascular plants in the Big Bend region flora. Most

specious are cosmopolitan families Asteraceae (230 species/107 genera), Poaceae (202/63), and Fabaceae

(107/37). They are listed first in the Tethyan (Saghatelyan 2006) floras as well, however, the first two families

have as much as 3096 of all species of the Big Bend flora. The leading position of Asteraceae reflects their

South American origin (Funk et al. 2005) and outstanding diversity in Mexico (Rzedowski 1995; Turner &

Nesom 1993; Valdés-Reyna and Cabral-Cordero 1993). The composition of legumes in the Big Bend flora

is very different from that in the eastern Tethyan floras as exemplified by the flora Armenia (Saghatelyan

1997a,b; 2006). vee absolute dominance of temperate papilionoid legumes in Armenia, the Trans-Pecos

flora has all th ilies well represented. Armenia has only Cercis griffitii of the basally branching Cae-

salpinioid legumes, the remaining genera being of more derived North Temperate papilionoid Hologalegina

(Wojciechowski et al. 2000; Schrire et al. 2005). The Trans-Pecos flora has eastern North American Cercis

canadensis and 6 genera/17 sp. of other basally branching clades of Caesalpinioideae (Schrire et al. 2005),

mostly in Cassieae. Their ranges are in more subtropical latitudes and more eastern parts of the Madrean

Region and eastern North America. Six genera and 27 species of the Mimosoideae in Trans-Pecos show only

southern connections with the ranges mostly in different, especially the Chihuahuan, parts of the Madrean

Region. Prevailing in North temperate floras Papilionoideae (62 sp. in Big Bend) are in the genera with mostly

western ranges in North America (Astragalus, Lupinus, Dalea, Sophora). Fewer genera have mostly eastern

North American ranges (Lespedeza, Desmodium), and a few have southeastern (Rhynchosia, Indigofera) ranges.

According to classification of Schrire et al. (2005), the legume genera of succulent or temperate biome affili-

ation and amphiatlantic or pantropical disjunction prevail in the Big Bend region flora.

High diversity of the next two families, the American Cactaceae (59/17) and Euphorbiaceae (59/10)

is due to the setting in the Sonoran Province, as well as (for Euphorbiaceae) on the southern border of the

United States. The middle sized families—Brassicaceae (43/21), Cyperaceae (40/10), Lamiaceae (38/11),

Boraginaceae (33/9), Solanaceae (32/10), Pteridaceae (33/8), Nyctaginaceae (32/13) Scrophulariaceae (29/13),

Onagraceae (28/4), Malvaceae (24/12), Asclepiadaceae (23/4), Amaranthaceae (22/7), Chenopodiaceae (21/6),

Verbenaceae (21/8), and Fagaceae (21/1)—reveal different connections: western and southern more than

northern and eastern. The high position of Fagaceae is due to the proximity to the center of diversity of

oaks in the highlands of central and eastern Mexico (Nixon 1993). If the Boraginaceae are counted with the

western American Hydrophyllaceae (14/2), the combined 47 species would be in fifth place after Cactaceae/

Saghatelyan, Geographical el ts of the fl f Big Bend region, Texas 415

Taste 2. Major groups of genera of the Big Bend flora.

North Temperte Old Worid 19

Relictual North Temperte 30

American-African 20

Madro-Tethyan 5

Fast Gondwanan 5

North American 37

EN American 7

W N American 57

Amphitropical 10

i 5

Chihuahuan 7

Mega Mexico Il 10

Mega Mexico III 26

SW N America 41

Tropic-Subtropical American 56

Tropic-Subtropical 47

Tropic-Warm Temperate 37

Tropical American 21

Tropical 6

Temperate-Subtr nical American

Temperate-Subtropical North and South 9

Total

TABLE 3. Genera combined.

North Temperate (wide and relictual) 54

North A icn G wide and east) 106

Madrean 98

Tropic/Subtropical/(Warm) Genera 212

West/SW/S North American Genera 122

American 288

Laurasian 160

ondwanan 306

Euphorbiaceae by their importance in the xeric vegetation. The Boraginaceae (Eritricheae) are especially

prominent in the California Province (Raven & Axelrod 1978), but also interesting in their geographical

connections and tribal representation in the Big Bend. The disjunctive North Temperate Boraginoideae

have 12 species of Eritricheae (Cryptantha, 9 sp.) and 7 sp. of Lithospermaeae (Lithospermum, 5 sp.) in the

Big Bend flora. Only slightly fewer species of the Boraginaceae in the flora represent two tropical tribes, the

Heliotropoideae (Heliotropium, 8 sp.) and Ehretioideae (Tiquilia, 5 sp.).

Paucity of species in typical North Temperate families like Caryophyllaceae (15/5), especially the older

ones, like Rosaceae (12/9) and Ranunculaceae (11/7), is due to the young age of this xerophytic flora and its

remoteness from eastern and western North American mountains. Among only 28 (!) species of petaloid,

monocot families (by contrast richly represented in Armenia) all, except for a few species of Allium, have

the Southern Hemisphere connections. In the Rhamnaceae (11/9) of the flora, the species of Madrean origin

and southern connections prevail.

The remaining 44 families have 3-10 species each and contribute only 264 (13%) species to the flora.

Sixteen families have only two species, and 27 families have only one species each. This type of spectrum

illustrates an arid subtropical character of the flora of a recent age. It supports the inclusion of the Big Bend

416 Journal of t tanical h Insti Texas 3(

—

Taste 4. Western North American genera (+ W S North American). Numbers in parentheses: (total sp./sp. in Big Bend

Fendlera (3/3 SW N Am) Giliastrum (3/3 SW N Am) Stephanomeria (24/2)

ma (45/7) Ipomopsis (24/5) Telesperma (12/5)

Phacelia (200/9) Eriogonum (250/15) Townsendia (25/1)

Cryptantha (100/9) Schoenocrambe (4/1) Xylorhiza (8/1)

Poliomintha (4/1 SW N Am) Stanleya (6/1) Trixis (60/1 SW N Am)

Salazaria (1/1 SW N Am) Karwinskia (17/1) Tridestomia (7/4 SW N Am)

Dasylirion (1/1 SW N Am) Holodiscus (8/1) Amsinckia (50/1)

Nolina (15/2 SW N Am) Purshia (7/1) Halimolobus (15/2)

Cevallia (1/1 SW N Am) Nerisyrenia (9/1) Cleomella (10/1)

Eucnide ( 8/1 SW N Am)

—

Aloysia (58/2, to S America

Ribes (150/3, temperate Eurasia, NW

Jamecia (12/1) Bahia (15/4) Africa

Rhynchosida (2/1) Encelia (15/1) Ibervillea (5/2 SW N Am)

Sphaeralcea (60/6) Flourensia (30/1) Dalea (160/17, chiefly Mexico, SW N

Acleisanthes (7/3 SW N Am) Gutierrezia (20-30/4) Am)

Selinocarpus (9/3) Helenium (40/3) Lupinus (200/3 4- Mediterranean, trop.

Camissonia (62/1) Heterosperma (5/1) i

Papaver sect. Agremone (28/4) Hymenoxis (28/1) Escobaria (16/8

Platystigma (9/4 Stevia (230/2 Blepharidachne (4/1)

Gilia (25/1) Macharanthera (35/5) Hesperostipa (1/1 SW N Am)

region in the natural chorion Mega-Mexico 2 (Rzedowski 1993), as well as its good placement in the Sonoran

Province (Thorne 1993) and the Western Region (McLaughlin 2007) of North America.

2. GENERA

Proportions of major geographic groups of genera, except cosmopolitan, in the Big Bend flora are illustrated

by Tables 2, 3, and 4. More than two thirds of the genera have their origins in different parts of the Americas.

Together with tropic-subtropical genera, the majority of which are also American, 78% of all the genera are

restricted to the Western Hemisphere. West-east connections across the Northern Hemisphere are much

weaker. Only 25% of all genera belong to northern (warm) temperate flora, while tropic-subtropical genera

comprise 37% and illustrate the essentially American and warm temperate-subtropical character of the Big

Bend region flora that has much more connections with the south, than with the north. Rzedowski (1993)

stated that in the entire Mexican flora the southern connections are four times stronger than the northern

ones.

1. North Temperate Old World Genera.—This group of 19 genera has 50 native species in the Big Bend

flora. These are Allium (700 world /6 Big Bend/ 102 USA), Linum ( 180/8/36), Limonium (350/1/10), Centau-

rea (500/1/4), Filago (35/1/3), Lactuca (100/2/8), Daucus (22/ 1/2), Lappula (60/1/2), Lithospermum (60/5/8),

Omphalodes (28/1/3), Erysimum (100/2/20), Noccaea (13/1/5), Silene (500/3/79), Bassia (26/1/6), Corispermum

(70/1/9), Krasheninnikovia (9/1/1), Helianthemum (110/1/14), Astragalus (2200/9/349), Polypogon (15/4/8), and

Arundo (5/1/1). Eurasian (EA) origin is well known from their distribution and is supported by phylogenetic

data for several genera, like Linum (McDill et al. 2008) and Limonium (Lledo et al. 2005). Eurasian origin

and migration via the Bering Land Bridge (BLB) with tive single colonization event to Andean South

America were stated for Astragalus (Scherson et al. 2008) and Silene (Popp & Oxelman 2007). The majority

of the genera in this group are of the young Neogene age (Popov 1963), with recent migration events to

North America (NA) and from there to temperate South America. Their centers of diversity lie in different

parts of the Tethyan Subkingdom: for Centaurea in Western Asia, for Silene in the Eastern Mediterranean-

Western Asia, for Limonium in the Mediterranean, while the largest papilionoid genus Astragalus is in the

first place in any Irano-Turanian flora. The genus Allium has a huge center in the Tethyan Subkingdom and

an essential one in western North America. Most of these genera probably reached the Big Bend region from

the north-west (via the BLB), fewer genera (Helianthemum, Linum, and Limonium) more likely crossed the

Saghatelyan, Geographical el ts ofthe fl f Big Bend region, Texas 417

Atlantic. Lledo et al. (2005) stated a close connection of the North American Limonium species with north-

ern European species of recent, Pliocene, age and an earlier migration of one species of a different clade to

South America. Thus, independent migrations of Limonium species from the Western Mediterranean to the

Americas in the Middle Miocene and in the Pliocene time could illustrate the recent pattern of Madrean and

Tethyan disjunctions, as opposed to that of older genera migrated earlier in the Tertiary in both directions

(Manchester 1999; Tiffney & Manchester 2001; Tiffney 2000).

2. Genera of Northern Latitudes and Mountains.—This group has genera with wide north temperate

distribution including those with arcto-alpine species. Many of them have their major center of diversity in

Eastern Asia (EA.), for example, Arenaria (China 102/world 300/Big Bend 5), Androsace (73/100/2), Anemone

(53/120/2), Clematis (147/300/2), Delphinium (173/350/2), Salix (275/520/5), and Gentiana (248/361/1). More

diverse in North America are Draba (world 350/ NA 102/ China 50/Big Bend 3) and Aquilegia (world 70—100/

NA 22/China 13/Big Bend 2). In North America they are mostly found in the western half, especially the

Rocky Mountains and Great Basin. A North American genus Epilobium (world 165/NA 42/Big Bend 1) has

seven sections most diverse in the Madrean Region or restricted to it, with the large section Epilobium di-

versified in all other continents except Antarctica (Katinas et al. 2004).

3. Relictual Widely North Temperate Genera.—This is a group of relictual genera, disjuncts of East-

ern Asian-Eastern North American, as well as several smaller southern Eurasian regions of preservation

of the Tertiary Laurasian flora. These are Juglans (world 21/Big Bend 2), Fraxinus (65/4), Crataegus (265/1),

Prunus (430/3), Ostrya (7/1), Lonicera (200/1), Vitis (65/1), Viburnum (225/2), Philadelphus (75/4), Cercis (6/1),

Pseudotsuga (6/2), and Lespedeza (40/1). It was suggested that Juglans, Fraxinus, and Acer have North American

origin in the Eocene (Manchester 1999), while Lonicera and Viburnum (Bell & Donoghue 2005), as well as

Mahonia (Wang et al. 2007) have E Asian origin and BLB crossing. Symphoricarpos (20/2) is mostly a NA

genus with only one species in EA. Pinus (100/3) and Mahonia (110/2/) have more species in the Western,

than the Eastern Region of North America. Together 30 north temperate genera of groups 2 and 3 have

only 60 species (4%) in the Trans-Pecos flora. They occur mostly at higher elevations or are scattered on the

canyon walls and along rivers (Powell 1998).

4. Widely Distributed Temperate Old World-New World Genera.—A group of 21 large genera of

different (mostly Laurasian) origin widely distributed in temperate regions of both North and South Hemi-

spheres. There are 80 species of the flora in these genera. Some of them have two centers of diversity: in

Asia and in Mexico. The latter center is reflected in the Trans-Pecos flora by the high numbers of species

in Quercus (500 world/150 Mexico/20 Big Bend), Salvia (900 world/325 Mexico/12 Big Bend/84 China/46

Ecuador, and a big center in Irano-Turania), and Lithospermum (60 World/5 Big Bend). A great antiquity

and Laurasian affinities for Mexican Salvia were stated by Ramamoorthy (1993). Other genera have a big

center in Asia and one (via the Beringian crossing) in western North America (Artemisia: 300 World/ 200

China/ 58 United States/5 Big Bend), or in Asia and in the Appalachian and Rocky Mountains (Thalictrum

(150 world/73 China/ 23 US). Fewer genera of this type are of North American origin: Solidago (100 world/5

Big Bend/ 72 US/ 5 China). The remaining genera are almost cosmopolitan: Marrubium, Lamium, Mentha,

Stachys, Polemonium, Hieracium, Bromus, Poa, Rhamnus, Paronychia, and others.

5. North American Genera.—This group has 37 genera/91 species (5.8% of the flora) of North American

and 7 genera/11 species of Eastern North American (Liatris, Ratibida, and Spermolepis) origin. Most of them

are of herbaceous habit and many are from Asteraceae. The larger genera have up to 8 sp. in the Big Bend

flora (Penstemon: 265 total /8 Big Bend; Erigeron: 200/8; and Castilleja: 200/4). The smaller genera have 3 to

6 sp. in the region (Matelea: 150/4, Lesquerella: 40/6, Helianthus: 49/4, Packera: 67/3, Gaillardia: 28/4, and

Gaura: 21/6).

6. Western North American Genera.—Table 4 has a list of 57 indigenous western NA wide genera

that are very important in the plant cover. These genera contribute 170 species (10.696) to the flora. This

group should include as well 12 species of five small, but peculiar Apachian genera (Fallugia, Engelmannia,

Pseudoclappia, Fendlera, and Berlandiera) and seven monotypous Chihuahuan genera (Amblyolepis, Psathy-

laf +l D * 4l D h bct. £ T.

418 J t Texas 3(

rotopsis, Emorya, Ariocarpus, Lophophora, Allolepis, and Holacantha). The Trans-Pecos flora has four of the

renouned Madrean genera: Foquieria, Chilopsis, Wislizenia, and Nectouxia. Altogether there are 73 western

North American genera with 193 sp. in the flora.

7. Amphitropical Genera.—There are 23 species of 10 amphitropial genera in the flora: Krameria: 18

total/3 Big Bend; Larrea: 5/1; Tiquilia: 20/5; Koeberlinia: 1/1; Hoffmanseggia: 28/3; Gottea: 1/1; Trichloris: 2/1;

Hedeoma: 38/7; and Scleropogon: 1/1. They grow in warm arid and semiarid regions of temperate North and

South America and have different evolutionary patterns involving long distance dispersals. For example,

Simpson et al. (2004) stated that Krameria has two major clad h containing a North and a South Ameri-

can subclade. They result from two independent dispersals, or two vicariant episodes, involving North and

South America. Hoffmanseggia arose in South America and had at least two dispersals to North America

at different times (Simpson et al. 2005). Tiquilia, as stated by Moore and Jansen (2006), exhibits a North

American origin and first divergence after greatest Cenozoic episode of rapid aridification near E Eocene/

Oligocene boundary, major diversification in the NA deserts in the Miocene and four long dist ispersal

to South America (Moore et al. 2006). The monotypic genus Koeberlinia of core brassicalean iy is of

North American deserts origin (Tobe & Raven 2008). For the American Larrea clade a separation from Old

World Zygophyllum by sea floor spreading in the Cretaceous was proposed (Porter 1972; Sheahan & Chase

1996). The South American-North American! arreoideae has two out of the five generain the Big Bend flora.

Sheahan and Chase (2001) stated that they are of South American origin and recent, Late Neogene, time of

arrival by long distance dispersal of the ancestors of L. tridentata to North American deserts.

Altogether, 82 western American genera with 204 sp. (14% of the flora) in groups 6 and 7 represent

the core of the autochthonous xeric floristic complex of the south-western deserts in the Big Bend region

flora. Many dominant and co-dominant species of the genera like Foquieria, Larrea, Koeberlinia, Nolina,

Dasylirion, Aloysia, Fallugia, and others, are in these two groups. Most of them arose from the ancestors

of Madro-Tertiary geoflora (Axelrod 1975; Raven & Axelrod 1978) affiliation. Fewer genera, for example,

Eriogonum, Nerisyrenia and section Argemone of Papaver, are Cenozoic western North American (Madrean)

derivatives of the ancestors from Arcto-Tertiary geoflora.

8. American-African Genera.—There are 20 genera with 57 sp. disjunctly distributed in America and

Africa: Thalinum, Thamnosma, Mimulus, Ambrosia, Coreopssis, Flaveria, Asclepias, Lupinus, Pomaria, Parkinsonia,

Heteranthera, Sageretia, Sideroxylon, Nicotiana, Bouchea, Lantana, Tagetes, Calliandra, Galactia, Mimosa, and

Sorgastrum. Some of them, like Mimosa (6 sp.), Asclepias (12 sp.), Lupinus (3 sp.) and Nicotiana (3 sp.) are

more specious in Big Bend and important in the plant cover. Transatlantic disjunctions of this group at the

genus level were attributed to long-distance dispersal in both directions mostly by water, while species level

disjunctions, to the anthropogenic factor (Renner 2004; Pennington et al. 2006; Simpson et al. 2006).

Madro-Tethyan Genera.—There are only 5 genera and 9 species that can be defined as Madro-Tethyan

in the Big Bend flora. Some of those analyzed by Axelrod (1975), for example Pistacia, are not in the region, but

grow not far from it, in the mountains of eastern-southern Mexico. Others, like Quercus, Juniperus, and Pinus,

were considered in the group 3 or 4 of this classification, even though they have entire vicariant sections in

the Tethyan and Madrean mountainous vegetation of low-middle altitudes. Some genera of Madro-Tethyan

origin have a wider range today (e.g., Cercis). Because the present scheme follows contemporary ranges of

taxa, these genera have been included in group 4 of north temperate relicts. Basalmost Cesalpinioideae was

important in the Tertiary floras of subtropical littoral type (Mai 1995) and Cercis is in the Mediterranean

element of many authors (Mai 1987; Kamelin 1983). In many cases, a convergent character of the Madrean

and Tethyan similarities, much discussed in the literature (Axelrod 1975; Raven 1971, 1973), stems from

the old age of such madro-tethyan genera. “The closer correspondence of woody taxa between the Madrean

and Tethyan regions in the Tertiary time stems chiefly from a more favorable climate and terrain for a wider

distribution” (Axelrod 1975:316).

The five Madro-Tethyan genera in the Big Bend flora are as follows:

Ephedra—35-—70 extant species in Central and SW Asia, SW North America, Mexico, Mediterranean,

Saghatelyan, G hical el ts of the fl f Big Bend region, Texas 419

at pl J

N Mediterranean Africa, and Andes from Ecuador to Patagonia. Four species grow in Trans-Pecos. Rydin et

al. (2006) stated a striking increase in gnetalean diversity and wide distribution of Ephedra and Ephedra-like

plants across Laurasia in the Early Cretaceous followed by a significant decline through the Late Cretaceous.

Major groups of Ephedra originated after the final rifting of Gondwana (Rydin et al. 2004) and have good

pollen record in Early Cretaceous paleoequatorial regions of Africa-South America (Crane 1996). Early

Cretaceous fossils from Brazil, USA (Virginia), China, and Portugal very similar to extant Ephedra, suggest

that crown group of Ephedra might be of Mesozoic origin (Rydin et al. 2004). North African-Mediterranean

species are in a basal grade whereas all non-African species belong to a clade of Asian origin and New World

species originated within the Asian clade (Rydin et al. 2004). Major center of diversity of extant Ephedra is

in Central Asia (18 sp.) with pollen recorded for SW Siberia since the Upper Cretaceous (Peshkova 2005).

Presence of an arid center in continental Asia since the Upper Carboniferous was stated by Kryshtofovich

(1955). “Apparently, the same flora of Angarida or Tungussian Flora is ancestral to both the Permian xeric

flora of Europe and Jurassic polichronous flora with its mesophytic characteristics. In the Upper Permian

and Lower Triassic extensive orogenesis and lava intrusions in Asia brought about even larger expansion

of the arid territories around the dry core of Asia” (Kryshtofovich 1957:46-47; cited and translated from

Peshkova 2005).

Cupressus—28 taxa, 16 in W USA, Mexico to Central America, the rest in NW Africa, Middle East

along Himalaya to SW and Central China (4 species) and N Vietnam (The Gymnosperm database). Recent

phylogenetic research (Little 2006) stated the polyphyly of Cupressus with the 12 Old World species being

sister to Juniperus. The 16 New World species of Cupressus plus Callitropsis were resolved as sister to the Old

World Cupressus plus Juniperus clade.

Arbutus—14-20 sp., W North America, Central America, West Europe, Mediterranean, and SW Asia.

A Laurasian origin for Ericaceae in the Late Cretaceous with the Arbutoideae in the basal clade was stated

by Kron and Luteyn (2004). “A widespread common ancestor may have occurred along the coastline of the

developing Tethyan seaway and subsequent vicariance may have been a major factorin the initial divergence

of the ancestors of Arbutoideae, Ericoideae, and Vaccinioideae due to the CORP ud uns apart of Africa,

Europe, and North America” (Kron & Luteyn 2004:498). The diversification of A | gthe Tethys

seaway stated also by Hileman et al. 2001) might explain th lisjuncti j of

the pattern in other older Madro-Tethyan genera, like Ephedra, Ce Eres pinions, and evergreen oaks.

Oligomeris—3 species, two of which are SW African, one O. linifolia from N Africa to W India, Canary

Islands and southern and south-eastern United States and Mexico, all growing in desert vegetation (Kubitzki

2003). Resedaceae, an Old World family of Capparalean affinities (Kubitzki 2003), has 6 genera and ca. 85

sp., with a major center of diversity in the arid regions of the Mediterranean basin and two centers of dif-

ferentiation, one in the western Mediterranean, and the other in the eastern Mediterranean and SW Asia

(Martin-Bravo et al. 2007). It might have crossed the Atlantic by long distance, as the access to post Eocene

NALB was restricted to cool-temperate and deciduous taxa (Tiffney 2000). Other younger Tethyan groups

since the Neogene time with concentration in the Mediterranean basin (like Cistaceae and Limoniaceae)

follow a similar pattern. Latest [ j lysis also stated for Oligomeris a long-distance dispersal

event from the Old World ius the sca. (Martin-Bravo et al. 2009).

Peganum—5-—6 species in the Mediterranean, southeast Europe, Transcaucasia, Middle Asia, Iran,

Afghanistan, Mongolia, China, India; one species in eastern and north-central Mexico. Recent placement

of Peganaceae in Nitrariaceae (Angiosperm Phylogeny Website) along with other Eastern Tethyan (Irano-

Turanian) genera, Nitraria and Malacocarpus, makes the Tethyan origin of Peganum even more obvious.

Mega-Mexico II Genera (Rzedowski 1993).—Ten genera with 21 species of the flora have their

ranges in Mexico and southern United States. All of them belong to Asteraceae, Cactaceae, and Poaceae.

11. Mega-Mexico III Genera (Rzedowski 1993).—Twenty six genera and 58 species have their ranges

in Mexico, extending to southern United States as well as tropical Central and northern South America.

Among them are Agave (100/, Agavaceae), Echeandia (12/1, Asphodellaceae), Pilostiles (25/1, Rafflesiaceae),

l ad ol D enli D h leti ET

420 J Texas 3(

(Bouwardia, 20/1, Rubiaceae), Leucophyllum (12/3, Scrophulariaceae), 8 small asteraceous genera, and one

of each family Cactaceae, Poaceae, and Amaranthaceae. Three small genera are present also in the West

Indies: Maurandella (Scrophulariaceae), Jefea (Asteraceae), and Garrya (13/2, Garryaceae). Among the three

Mesoamerican genera Hechtia (47/1, Bromeliaceae) extends northward to only 3 counties of southern Texas

in the United States.

Altogether, there are 23 genera with 68 species of groups 10 and 11 in the Big Bend flora. Species of

some of them (like Agave, Leucophyllum, Garrya, and Echinocereus) play a dominant role in the xeric com-

munities of the Madrean vegetation.

12. Western and south-western United States, Mexico (northern) and Texas (southern United

States) Genera.—Forty one peculiar genera with 70 sp. evolved in the Madro-Tertiary Geoflora have this

type of range. Among those, six genera of the Cactaceae are most diverse. The remaining genera are from

mostly tropical-warm temperate families like Apocynaceae (Haplophyton), Malvaceae (3), Nyctaginaceae (3),

Rhamnaceae (Adolphia), Solanaceae (Chamaesaracha), and Verbenaceae (Tetraclea); largely tropical Rutaceae

(Choisya, Ptelea) and Sapindoideae (Ungandia). The Asteraceae have 12 genera, papilionoid legumes have

three, and endemic western North American family Crossosomataceae is monotypic. Prevalence of warm

temperate-subtropical-tropical connections, some of those being very interesting, is evident from this list.

For example, in core Rutaceae (Groppo et al. 2008) Choisya (4—5) and Ptelea (1-3) form a tropical American

clade, Zanthoxylum is pantropical, and Casimiroa-Dictamnus clade has Dictamnus (1-2 sp. S Europe to N

China), Skimmia (4 sp. east of the Himalayas to S Vietnam and the Philippines) and Casimiroa (Mexico, Costa

Rica). Thus, core Rutaceae clade exemplifies both Madrean-tropical American connections and east to west

Old World-New World (madro-tethyan) connections along the southern Tethys shore.

Among primarily north temperate families, the Brassicaceae has two and the Rosaceae has three genera

(Cercocarpus, Malacomeles, and Vaquelinia) which exhibit this pattern of distribution. Six of the eight rosa-

ceous genera in Trans-Pecos arose in the Madro-Tertiary Geoflora, with the remaining two genera being in

common with the Old World.

13. Tropic-Subtropical American Genera.—There are 56 genera (119 sp.) in the flora with large distri-

bution in tropical and subtropical parts of the Americas. Some of those are very peculiar and important in

xeric desert and th hrub communities (Condalia, Holocantha, Guajacum, Calliandra, Proboscidea, Mentzelia,

Eryngium, etc). The largest family in this group is Asteraceae (12 gen.).

14. Tropic-Subtropical Genera.—There are 47 genera (102 sp.) of pantropic-subtropical distribution

in Trans-Pecos. The grasses (10 gen.) and the legumes (8 gen.) are most specious, mee dominant

] gen.), like Jatropa, | disj

species (Acacia, 9 sp., Prosopis, 2 sp). Several E

the same refers to the Rhamnaceae (Coba Ziziphus) and Buddleja. The genus Boerhavia, Bb spiele.

has 8 out of its 20 species in the Big Bend flora

15. Tropical to Warm Temperate Genera.—There are 42 tropic/subtropical genera (179 sp.) extending

to warm temperate countries in the flora. The majority of them are grasses with 15 gen. /83 sp. (Imperata,

Pappophorum, etc.), followed by Malvaceae (4/11), Cyperaceae (3/18), Amaranthaceae (2/12), Senna (10),

Heliotropium (8), Ipomoea (8), Kallstroemia (5), and Rhus (4).

16. Mostly Tropical American Genera.—Twenty one genera with 64 species. Here are the legumes (4

gen./7 sp.), Cucurbitaceae (2 gen.), and others (Tecoma, Tillandsia). Eleven more genera and 50 sp. have wider

distribution in the Americas, are important in the plant cover, and have their ranges centered primarily in

Central and South America (Baccharis, 6 sp; Opuntia, 16 sp; and Bouteloua, 15 sp.).

17. Tropical genera.—Six genera with 10 sp. of pantropical distribution: Cissus, Sesuvium, Rynchosia,

and three genera of the grasses.

18. Temperate-Subtropical American Genera.—Seven American genera with 41 sp. are widely dis-

tributed in north and south temperate and subtropical latitudes: Oenothera (1 1), Verbena (7), Physalis (6), etc.

19. Temperate-Subtropical Genera of North and South Hemispheres.—Nine genera with 21 species

absent from the tropics have this type of wide distribution. Five of them are grasses (Agrostis, Melica, and

Saghatelyan, G hical el ts of the fl f Big Bend gion, Texas 421

JO r »

Hordeum). Most specious in this group is Lycium (5 sp). It has 70 species disjunctly distributed in temper-

ate subtropical regions of South and North America, S Africa, Australia, and Eurasia. Fukuda et al. (2001)

stated that Lycium originated in the New World, all the South African, Australian, and Eurasian species

together are monophyletic and have a common ancestor from the New World. It was confirmed by recent

data (Levin & Miller 2005) that the Old world Lycium species clade is nested within the American species,

which comprise the rest of the Lycieae. This is another example of Madro-Tethyan connections, although

the genus has a wider distribution. Species of Lycium grow in littoral (turgay) communities of dry regions

in the Tethyan subkingdom.

20. Leptopus PUE tribe a Euphorbiaceae presents an interesting case.—An

endemic NA clade of And (Vorontsova et al. 2007) includes two species disjunctly

distributed in Trans-Pecos-Coahuila, Edwards Placa and Ozarks Plateau. Itis a relictual disjunctive type of

rune This clade is sister to a clade of a shrubs restricted to relictual humid forests of western

, Asia, and Indonesia plus a clade of evergreen rainforest understory trees of Asia and Australia

(Vorontsova et al. 2007). These authors state that tribe Poranthereae appears to be centered in Africa, with

movement to Asia, Australia, and the New World, and movement from subtropical arid habitats to wetter

temperate and tropical ones. The basalmost clade combines microphyllous procumbent species of semi-

deserts and has two subclades: the African-American (Somalia, Ethiopia/Mexico, and Peru) and the Asian/

African subclade (Horn of Africa and N Africa/ Irano-Turania). The New World subclade of the basalmost

clade indicates a single colonization and has two very similar amphitropical species of Andrachne in Baja

California and Peru. Thus, we see another example of colonization of North America from both the northern

route across Laurasia (section Phyllanthopsis), and along the southern Tethys shore across the Atlantic (the

Mexican-Peruvian species).

This latter migration track seems very feasible for termophyllous ancient taxa originated or preserved

in the Horn of Africa region. The importance of that region was stated by many researchers of Africa: "The

high endemism of Horn of Africa emerges clearly....The flora of the Somali Republic is a remarkable one

with very many outstandingly distinct species found nowhere else" (Brenan 1978:467). "The Somalian

phytochorion could be seen as a hub for the arid truck” (Thulin 1994). “In historical terms it suggests that

Horn of Africa would be a refuge area for the arid flora" (Friis & Balsev 2005).

The rest of the genera are cosmopolitan, or of wide temperate distribution, or have mostly non-native

species in the flora.

CONCLUSIONS

The setting of the Big Bend region flora on the boundary a two subkingdoms of the Holarctic Kingdom ex-

plains its mixed character with combination of li d either from Madro-Tertiary or Arcto-Tertiary

geoflora stock. However, the Madrean lineages RM and the flora is mostly south-western North

American by composition (Table 1). There are 1586 native species in 595 genera of 125 families in the Big

Bend region flora. Many genera and families in the flora are monotypic or oligotypic.

Three quarters of the genera are restricted to the Western Hemisphere, and tropic/subtropical, mostly

South American, genera have 3796 of the species diversity in Trans-Pecos. Taxa with southern connections,

especially the Madrean element which has 1056 species (or 6696) of the flora, prevail. The Chihuahuan

Subprovince has a high level of endemism (2696 of the flora) with many younger taxonomic groups, evolved

in situ since the Cenozoic rapid aridification episode. It is well placed in the Sonoran Province, however, its

boundary with the Tamaulipan Subprovince is very vague.

North Madrean and southern Rocky Mountains-Sonoran species (696 of the flora) are of either Madro-

Tertiary or Arcto-Tertiary geoflora affiliation. The Apachian element is well represented (996 of the flora)

which corroborates delineation of the namesake subprovince (McLaughlin 2007), while the Comanchian

(2.596 of the flora) probably is a chorion of a lower rank.

East to west connections across the northern Hemisphere have a lesser weight in the flora. The older

£ al Dat H ID i, | dei toe ke ET

422 Journal of Texas 3(

Laurasian genera that migrated in the early Tertiary via one of the two northern bridges are not many in

this younger flora. The NA-EA similarities are on the genus level and above. More recently migrated to the

region are mostly progressive eastern Tethyan (mostly Irano-Turanian) genera which came from Asia via

the BLB (Salsoleae, Astragalus).

Diversification of older Madro-Tethyan genera, like Ephedra, Cercis, Arbutus, Cupressus, pinions, and

evergreen oaks along the Tethys seaway and recent long distance dispersal of younger genera from the

Mediterranean (Oligomeris, Limonium) might explain contemporary pattern of the Madrean and Tethyan

disjunctions. Another pattern (core Rutaceae) shows both Madrean-tropical American connections and Old

World-New World connections E the Tethys shore. The southern east to west connections across the

Atlantic along the southern T ident from several striki in , tropi-

cal Boraginaceae, Rutaceae, Arda biome Fabaceae, Euphorbiaceae, A DIE AE and others.

APPENDIX 1

AREA DIAGNOSES FOR THE SPECIES OF BIG BEND REGION, TEXAS

The following list of species was extracted mostly from the Synthesis of the North American Flora (Kartesz & Meacham

2002) and it follows, with few exceptions, all the nomenclatural combinations as well as author citations of this source. The

nomenclature and distributional data of the Onagraceae follows the Onagraceae website (Wagner & Hoch 2005) and the

a clade follows Vorontsova et al. (2007). In the few cases where the nomenclature is different from that in the above

DHoneo sources, ds auinar nomes are niea The list below should be considered asa EOM er a puc of

| The families of vascular F

as are genera and species. Numbers by each family name indicat / ratios in the family.

ACANTHACEAE: 18/8 AIZOACEAE: 1/1

Anisacanthus linearis Chihuahuan Sesuvium verrucosum American Trop/Subtr

o dela poe n ALISMATACEAE: 1/1

nisacanthus quadrifidus Chihuahuan Sagittaria calycina N American

Carlowrightia arizonica Sonoran

ra dd O Apachian AMARANTHACEAE: 22/7

Alternanthera caracasana Tropical American

Carlowrightia a Tr-Pecos-Coahuila

Carlowrightia parv. ISI Tr-Pecos-Coahuila maranthis acanthochiton Chihuahuan

E dd Tr-Pecos- Lone Amaranthus albus Polichorous

Carlowrightia a Chihuahua Amaranthus arenicola Prairie with radiations

Dyschoriste ond SCU Amaranthus blitoides N American (WC)

Dyschoriste schiedeana Sonoran-Chihuahuan

Elytraria imbricata Mesoam- ean Amaranthus palmeri N American (SWC)

Justicia pilosella Chihuahuan-Tamaulipan Amaranthus polygonoides a

Justicia warnockii Tr-Pecos Amaranthus powellii Cordiller

Amaranthus retroflexus N Po (polychorous)

Amaranthus s Chihuahuan-Tamaulipan

Amaranthus torreyi W Sonoran

Froelichia arizonica soar Mc

Stenandrium barbatum Chihuahuan

Tetramerium Sonoran

Froelichia gracilis N Amer

ACERACEAE: 1/1 Gomphrena nitida a are

Acer gander N Madrean Mont. (S Rocky M-E Great Guüilleminea densa Madrean

Basin-Sonoran Province) Iresine heterophylla Sonoran wide

AGAVACEAE: 10/2 Iresine leptoclada Chihuahuan

Agave glomeruliflora Chihuahuan Tidestromia carnosa Chihuahuan

Agave gracilipes Chihuahuan Tidestromia gemmata Chihuahuan

Agave havardiana Tr-Pecos Endemic Tidestromia lanuginosa SW & SC US

Agave lechuguilla Chihuahuan Tidestromia suffruticosa Chihuahuan

a O ANACARDIACEAE: 4/1

TUTA DACA E 2 Rhus lanceolata SC US-Chihuahuan

er did Sonoran-Chihuahuan Rhus microphylla Sonoran-Chihuahuan wide

Yucca faxoniana Chihuahuan Rhus trilobata W N American (+ W Prairie)

Yucca thompsoniana Chinuahuan Rhüsvirens E Madresn

Yucca torreyi Chihuahuan

r kiral al $ £ sl £a £ D:4D

Saahatelvan

PJ , r

ANEMIACEAE: 1/1

Anemia mexicana Mesoam

APIACEAE: 7/5

Arnmoselinum popei S Prarie

Cyclospermum leptophyllum S N American

Daucus pusillus N American

Eryngium heterophyllum W & S Madrean

Eryngium leavenworthii S Prairie

Spermolepis echinata S N American

Spermolepis inermis C & S Prarie-Appalachian

YNACEAE: 8/4

Amsonia longiflora Chihuahuan

Amsonia palmeri N Sonoran Province

Apocynum cannabinum N Americ

Haplophyton crooksii " Sonoran- CEDERE

Telosiphonia hypoleuca J Davis c, Madrean Mont

Telosiphonia lanuginosa Madrean Mont (East)

Telosiphonia macrosiphon Tr-Pecos-Coahuila

ARALIACEAE: 1/1

Aralia racemosa ssp. bicrenata SW N American

ARISTOLOCHIACEAE: 2/1

Aristolochia coryi S TX Endemic (SW-SC TX)

Aristolochia wrightii Chihuahuan

ASCLEPIADACEAE: 23/4

Asclepias asperula SWC N American

Asclepias brachystephana Sonoran- oe

Asclepias engelmanniana Prairie-SW

oid glaucescens (Sonoran)- Em

Ie latifolia 5 "dex SW US

Subprovince

Asclepias iS 2 US- ow

o caposa Chi disjun

Asclepias sperryi Tr-Pecos-Coahuila

Asclepias serica SW US-Chihuahuan

Asclepias texana N Chihuahuan-Coahuila

Asclepias vido N American

Cynanchum barbigerum Tamaulipan

Cynanchum pringlei Chihuahuan

Cynanchum racemosum N Chihuahuan-Tamaulipan

Funastrum crispum SWC US-Chihuahuan

Funastrum cynanchoides N & W Madrean

Funastrum torreyi Chihuahuan-Tamaulipan

Matelea parvifolia N Madrean

Matelea producta N & W Madrean

Matelea reticulata Chihuahuan-Tamaulipan

Matelea texensis Tr-Pecos Endemic

Matelea sagittifolia S TX Endemic

ASPLENIACEAE: 3/1

Asplenium palmeri ) Davis c. Mesoam-Madrean Mont

Asplenium resiliens N American

Asplenium trichomanes J Davis c. Holarctical

ASTERACEAE: 230/107

Acourtia nana Sonoran-Chihuahuan

Acourtia runcinata Chihuahuan-Tamaulipan

Acourtia wrightii Sonoran-Chihuahuan

Texas

Ageratina herbacea Madrean (SWC US)

Ageratina rothrockii Sonoran-Chihuahuan

Ageratina wrightii Chihuahuan

Amblyolepis setigera Chinuahuan-Tamaulipan

Ambrosia confertiflora SW & SC US. N Madrean

Ambrosia psilostachya N American-Mesoam

Amphiachyris dracunculoides Prairie

Aphanostephus ramosissimus ?

Aphanostephus riddellii SC US (W TX-Chihuahuan)

Artemisia campestris Holarctical

detis Mint ME N American

Holarctical

me filifolia WC US

Artemisia ludoviciana N Amer

Astranthium robustum OAM de

Baccharis bigelovii Chihuahuan

Baccharis salicifolia American Trop/Subtr

Baccharis salicina Prairie. WC N American

Baccharis wrightii SW N American

Bahia absinthifolia Sonoran-Chihuahuan

Bahia bigelovii Tr-Pecos Endemic

Bahia dissecta WC US Mont (Rocky M)

Bahia pedata S Rocky M-Chihuahuan

Baileya multiradiata a (no Mexican Highlands)

Berlandiera lyrata M

Bidens bigelovii ae i N Chihuahuan ?

Bidens laevis American n

Bidens leptocephala Apachia

Borrichia frutescens Pecos c. Ht Coast

Brickellia californica W N Am

Brickellia conduplicata o Mont

rickellia V Sonoran Province

Brickellia eupatorioides N o (not Pacific)

Brickellia gra raWNA an

Brickellia inco Tr-Pecos eee

Brickellia laciniata Chihuahuan wide

Brickellia veronicifolia Chihuahuan-Mexican Highlands

alyptocarpus vialis Mesoam-Gulf Coast

Carphochaete a N Sonoran-Chihuahuan

W-SC US (S Prairie-Apachian)

Chaetopappa ellas Mel EM

Chae Ed appa ericoides WC N Am

Chaetopappa parryi oo Tarai

Chaptalia texana E Madrean Mon

Chloracantha spinosa ae MEE

Chrysactinia mexicana E ~ Mont

Cirsium ochrocentrum WC

Cirsium texanum dide

Cirsium turneri S TX-Coahuila Endem

Cirsium undulatum WC N dd ey US)

Conoclinium dissectum Chihuahua

Cosmos parviflorus Apachian-Mexican Highlands

Dyssodia papposa Mesoam-N American

423

424

ds prostrata American a radiations

celia a a N Chihuahua

nia peristenia S or Rocky M-Chihuahuan)

adrean

Erigeron divergens W N American

Erigeron eximius J ise C. dim American Mont (Rocky M-

Colorado-Apachian Mont)

Erigeron flagellaris W N mpm Mont

WC US ?

a SC US (S Prairie)

e od Mesoam M radiations

Flourensia cernua Sonoran

Flyriella parryi (ES) da

Gaillardia multiceps ?

Gaillardia pinnatifida SW US (not Cal)

a rdia pulchella N American (mostly S-C)

rindelia arizonica Apachian

Grindelia grandiflora J Davis c. N Chihuahuan

Grindelia ben SW TX & SW AZ Endemic

Grindelia US

Grindelia um S diede -(Mexican Highlands)

Grindelia scabra N entra ahua

irean (no Mexican Highlands)

Gutierrezia sarothrae W N American

Gutierrezia A Chihuahuan-Tamaulipan

Gutierrezia a Chihuahuan

Gymnosperma see osum Mesoam

a greggii Sl Prairie-Chihuahuan

Hedosyne ambrosiifolia sca Chihuahuan

Helenium amarum E N Am

Helenium elegans e A

Helenium microcephalum S Prairie-Chihuahuan-Tamaulipan

Helenium n m var. ooclinium Chihuahuan

Helianthus annuus N American- da

Helianthus o SC N Amer

Helianthus niveus Sonoran dide (W Madrean)

Helianthus paradoxus Pecos, Reevs cc. ?

Heliomeris longifolia Madrean disjunctive

Heliomeris multiflora Rocky M

Heliopsis parvifolia N Madrean Mont

Heterosperma pinnatum Mesoam-Madrean

Heterotheca canescens SC-SW US

Heterotheca fulcrata W US Mon

Heterotheca subaxillaris N American

Heterotheca villosa W N Am

Heterotheca viscida N a (+S AZ)

Hieracium carneum S Great Basin-N Chihuahuan (+ S AZ)

nt disjunctive

Hieracium schultzii Mesoam

Hymenoclea monogyra N Madrean

Hymenopappus flavescens SW US

Hymenopappus scabiosaeus Prairie

anophyllon solidaginifolium Chihuahuan

ae graminifolia dad (SW US)

Laennecia coulteri N

Laennecia filaginoides viu m-S American

Laennecia sophiifolia Amphitropical ?

Leuciva dealbata Chihuahuan

Liatris punctata Prairie

Lygodesmia ramosissima Sonoran-Chihuahuan

Lygodesmia texana S Prairie-Chihuahuan

Machaeranthera blephariphylla N Chihuahuan ?

Machaeranthera gracilis N Madrean

Machaeranthera gypsophila Chihuahuan

Machaeranthera pinnatifida Prairie-Madre

Machaeranthera tanacetifolia W Prairie-S c M-Madrean

Melampodium leucanthum SWC U

Melampodium strigosum J Davis c. Sonoran-Chihuahuan ?

(+ Colorado)

Nicolletia edwardsii Chihuahuan

Packera millelobata Chihuahuan

Packera neomexicana Apachian

Packera Ades Davis c. EC Prairie

Palafoxia callos hian extending to MO

Palafoxia e Chihuahuan

Palafoxia sphacelata E Prairie

Parthenium argentatum Chihuahuan-Mexican Highlands

Parthenium confertum Chihuahuan-Tamaulipan

Parthenium incanum Madrean wide

Pectis angustifolia WC N America

Pectis filipes Sonoran-Chihuahua

Pectis papposa Madrean (no Mex. Highlands)

Pectis prostrata Mesoam-Central American

Pericome caudata N Madrean

Perityle aglossa Tr-Pecos-Coahuila

Perityle angustifolia SW TX Endemic

Perityle bisetosa Chihuahuan

Perityle cinerea Tr-Pecos Endemic

Perityle rupesttis Chihuahuan

Perityle vaseyi Chihuahuan

bns vitreomontana Brewster c Endemic

ropappus roseus Madrean-Mesoam

drean

orophyllum scoparium ~

Psathyrotopsis scaposa Chihua

Pseudoclappia arenaria Pecos c. fe Rocky M-N Chihuahuan

(Coahuila)

Pseudognaphalium arizonicum Sonoran- role

Pseudognaphalium canescen de N Amer

Pseudognaphalium jaliscen

Pseudognaphalium zu ca Chihuahuan

f. khi-al al + Eel A £D:4 D J

Saahatelvan

> , F

, Texas 425

> T E E

Pseudognaphalium stramineum W N American

seudognaphalium viscosum Mesoam

Psilactis asteroides Sonoran-Chihuahua

Psilactis brevilingulata Chihuahuan (onm

Psilostrophe gnaphalioides Chihuahuan-Tamaulipan

Psilostrophe tagetina SWC US

Ratibida Seen Prairie wide

Ratibida tagetes

Sanvitalia procum UN am

Schkuhria multiflora Re MN

Senecio flaccidus Madrean wide

Senecio parryi N Sonoran -Chihuahuan

Senecio riddellii W Prairie

enecio wootonii J Davis, Pecos cc. SWC N American

Simsia calva Chihuahuan-Tamaulipan

Solidago velutina W N American (not Pacific)

Solidago wrightii S Rocky M-W Madrean

Stephanomeria minor W N American

Stephanomeria pauciflora W N American

Stevia ovata Mesoam-S American

Symphyotrichum expansum ?

Symphyotrichum falcatum WC N American (W Prairie wide)

Symphyo otrichum lanceolatum J Davis c. N Amer

Symphyotrichum subulatum American (coastal) ae

Tagetes micrantha Mesoam-Madrean

Tetragonotheca texana Chihuahuan-Tamaulipan

Tetraneuris linearifolia S Prairie-Chihuahuan-Tamaulipan

Tetraneuris scaposa W Prairie-Chihuahuan-Tamaulipan

Thelesperma ambiquum Tamaulipan

Thelesperma filifolium Prairie

Thelesperma longipes Chihuahuan (4- S AZ)

Thelesperma megapotamicum Prairie, secondarily Amphi-

tropical

o. simplicifolium ed (Tamaulipan ?)

Thymophylla acerosa N Madrean wide

Thymophylla aurea ?

Thymophylla a rial

Thymophylla pentachaeta Madre

no ind a

Townsendia N ce

Trixis al N Ma dre

Verbesina encelioides pec

Verbesina nana Chihuahuan

Verbesina oreophila Chihuahuan Mont

Verbesina virginica J Davis c. E N American

Vernonia larsenii Chihuahuan

Vernonia marginata W Prairie-Chihuahuan

Viguiera cordifolia N Sonoran-Chihuahuan

Viguiera dentata Mesoam-Madrean

Viguiera stenoloba Madrean (Chihuahuan wide)

Wedelia texana Chihuahuan-Tamaulipan

Xanthocephalum gymnospermoides J Davis c. Sonoran-

Chihuahuan

Xylorhiza wrightii Chihuahuan

Maii triantha Chihuahuan

Zinnia grandiflora S Rocky M-Sonoran wide

AZOLLACEAE: 1/1

Azolla mexicana C & N American

BERBERIDACEAE: 2/1

Mahonia haematocarpa Sonoran

Mahonia trifoliolata Sonoran

BETULACEAE: 1/1

Ostrya virginiana Appalachian. E N American

BIGNONIACEAE: 2/2

Chilopsis linearis Sonoran w

Tecoma stans American la

BORAGINACEAE: 33/9

Amsinckia menziesii W US

Antiphytum floribundum E € S Madrean

Antiphytum heliotropioides S edad (Coahuila)

Cryptantha albida Sonoran-Chihuahua

Cryptantha angustifolia SW N American

Cryptantha cinerea W N Am

Cryptantha ipes Endemic for n NM, & Brewster, TX,

Chihuahuan

Coptantha crassisepala Sonoran wide

Cryptantha mexicana Chihuahuan

ae oblata Endemic S NM — SW TX

Cryptantha palmeri Chihuahuan

Cryptantha pusilla Sonoran-Chihuahuan

Hackelia pinetorum SW US (AZ, NM, SWTX)

Heliotropium t Chifuahuan

Heliotropium convolvulaceum in N American

Heliotropium curassavicum Am

Heliotropium pues rim

Heliotropium procumbens Caribbean with irradiations to N &

erica. American

Heliotropium torreyi Chihuahuan-Tamaulipan

Lappula occidentalis W N American

Lithospermum multiflorum S WC US

Lithospermum viride Chihuahuan rare

Omphalodes aliena Chihuahuan narrow

Tiquilia canescens Madrean

Tiquilia gossypina Chihuahuan

Tiquilia greggii Chihuahuan

Tiquilia hispidissima Chihuahuan

Tiquilia mexicana Chihuahuan

RASSICACEAE: 43/21

Arabis fendleri W N American (Great Basin-S Rocky M)

ardamine macrocarpa Chihuahuan, rare

426

Descurainia incana J Davis c. W N American

Descurainia pinnata N American (mostly W)

Dimorphocarpa wislizeni N Sonoran-S Rocky M

Draba cuneifolia W & SC N American

Draba platycarpa W & SC US poa

Draba standleyi Apachian ? Rare

Erysimum gane Prairie iud ie

Erysimum capitatum W N Amer

SEDE diffusa. TA Me

Lep ocky M-E Great Basin

pora austrinum SC N American

Lepidium densiflorum N American

Lepidium lasiocarpum Great Basin-Sonoran

Lepidium oblongum S & C N American

pid sordidum Chihuahua

pidium virginicum N i

ae argyraea Chihuahua

Lesquerella fendleri S Rocky M-(SE Great Basin)-Sonoran

Lesquerella gordonii Sonoran-SE Prairie

Lesquerella mcvaughiana Brewster & Pecos cc Endemic

Lesquerella ovalifolia C Prairie-N Sonoran

Lesquerella purpurea N Sonoran proper

Mancoa pubens N Sonoran-Chihuahuan disjunct

O a Chihuahuan spam

US Mont not Cal Provi

sed en Davis c. S Great Basin- sae

Pennellia micrantha S Great Basin-Sonor

Rorippa ramosa laa rare

dius teres Gulf Coa

enocrambe inca W N American

pns dissecta N Sonoran-Chihuahuan disjunct

Sisymbrium auriculatum EE

Stanleya pinnata W N American

Streptanthus carinatus N am O

Streptanthus cutleri Brewster c. Endem

Streptanthus platycarpus SW TX P

Synthlipsis greggii Chihuahuan

Thelypodiopsis purpusii SW US ( ?)

Thelypodium tenue Tr-Pecos Endemic

Thelypodium texanum Tr-Pecos Endemic

Thelypodium wrightii SW US (Apachian ?)

BROMELIACEAE: 2/

Hechtia texensis Tr-Pecos Endemic (Big Bend)

Tillandsia recurvata American Trop/Subtr

BUDDLEJACEAE: 3/2

Buddleja agone m

Buddleja scordioides E Madre

morya suaveolens M huila

CACTACEAE: 56/17

Ariocarpus fissuratus Chihuahuan

Coryphantha echinus Chihuahuan

Coryphantha macromeris Chihua

Coryphantha ramillosa Chihuahuan (Coahuila)

Coryphantha robustispina Sonoran-Chihuahuan

Cylindropuntia davisii Chihuahuan+ SWC US

ylindropuntia imbricata S N American

Cylindropuntia kleiniae Sonoran-Chihuahuan

Echinocactus texensis Chihuahuan-Tamaulipan+ SWC US

Echinocereus chisoensis ?

Echinocereus coccineus SW US-Sonoran wide (no Cal)

Echinocereus dasyacanthus Chihuahuan

Echinocereus enneacanthus Chihuahuan-Tamaulipan

Echinocereus pectinatus Chihuahuan-Tamaulipan

Echinocereus viridiflorus W Prairie-Chihuahuan

Echinomastus intertextus Sonoran-Chihuahuan

Echinomastus mariposensis Chihuahuan

Echinomastus warnockii Chihuahua

Epithelantha micromeris Sonoran- db

Escobaria duncanii Chihuahuan

Escobaria hesteri Tr-Pecos Endemic

Escobaria minima Brewster c. Endemic

Escobaria tuberculosa Chihuahuan

Escobaria vivipara W N American

Ferocactus hamatacanthus Chihuahuan-Tamaulipan

Glandulicactus uncinatus var. wrightii Chihuahuan

Grusonia aggeria Tr-Pecos-Coahuila

Grusonia grahamii Tr-Pecos

Grusonia schottii Chihuahuan-Tamaulipan

Lophophora williamsii Chihuahuan-Tamaulipan

Cane abita Sonoran-Chihuahuan

n wide (+Tamauilipas)

Mammillaria heydeity var. PA Apachian

Mammillaria lasiacantha er Chihuahuan

Mammillaria pottsii Chihuahua

Neolloydia conoidea chat -Tamaulipan

Opuntia atrispina STX Endem

Opuntia aureispina Brewser c. I.

Opuntia engelmannii Sonoran wide

Opuntia engelmannii var. lindheimeri SC N American

Opuntia phaeacantha SW N American

Opuntia polyacantha W N American

Opuntia rufida Chihua

Peniocereus greggii ne ran rude

Thelocactus bicolor Chihuahuan-Tamaulipan

CALLITRICHACEAE:1/1

Callitriche heterophylla ) Davis c. C&N American

CAMPANULACEAE: 6/3

ampanula rotundifolia Holarctical (Palearctical)

Lobelia berlandieri Chihuahuan

Lobelia cardinalis N American

Lobelia fenestralis Tr-Pecos, AZ, Mexico, SW NM Disjunct

Mont

r kiral al + £ el £1 {Din D ry

, Texas 427

Saahatelvan

- q d

Triodanis holzingeri CS Prairie

Triodanis ii N American

CAPPARA 6/5

Cleome nulia Disjunct: AZ. CO. TX, WO. S Rocky M-W

Madre

Cleomella m Sonoran-Chihuahuan (W)

Koeberlinia spinosa Amphitropical (NW Sonoran-

Chihuahuan

Polanisia dodecandra N American

Polanisia uniglandulosa Madrean (Mexican Highlands-N&E

Madrean

Wislizenia refracta Sonoran

CAPRIFOLIACEAE: 5/3

Lonicera albiflora SWC US disjunct mont

Viburnum australe J Davis c, rare, also in Mexico ?

Viburnum rufidulum J Davis c. E N American

CARYOPHYLLACEAE: 15/5

Arenaria benthamii SC US. C-S Texas

Arenaria fendleri ) Davis c. S Rocky M-S Great Basin-Apachian

Arenaria lanuginosa Davis c. N American ?

Arenaria livermorensis ) Davis c. Tr-Pecos Endemic

Arenaria ludens Tr-Pecos- nana

Cerastium axillare Chihua

Drymaria laxiflora Chihua tee

Drymaria leptophylla SWC N American (Sonoran ?)

Drymaria molluginea J Davis c. SWC N American

Drymaria Aid N Chihuahuan

a onychia jamesii SC US

Paronychia mon E Tr-Pecos -Coahuila Mont

Paronychia wilkinsonii Chihuahuan Mont

Silene antirrhina N American

Silene laciniata Madrean wide

CELASTRACEAE: 4/3

Celastrus scandens N American (NE-NC)

Mortonia sempervirens Chihuahuan

Mortonia scabrella N Sonoran- Chinuahuan

Schaefferia cuneifolia Chihuahuan

CERATOPHYLLACEAE: 1/1

Ceratophyllum demersum Polychorous

HENOPODIACEAE: 21/6

Atriplex acanthocarpa Chihuahuan

Atriplex canescens W US (+ Chihuahua)

Atriplex elegans N Madrean

Atriplex obovata SE E RM S in M-Chihuahuan

Atriplex wrightii SW-C US + N Mex

Chenopodium LT A eee

Chenopodium cycloides E CS Prairie

Chenopodium fremontii W N American

Chenopodium incanum W N American

Chenopodium graveolens Amphitropical

Chenopodium leptophyllum W N American

Chenopodium neomexicanum SW US

eae murale Polichorous (Palearctic)

Chenopodium pallescens Presidio c. ?

Chen T prateri am N American

Corispermum americanum WC N American

eee Sean N American

Krascheninnikovia lanata W N American

Monolepis nuttalliana W erican

Suaeda suffrutescens var. suffrutescens Apachian-Chihuahuan

(+ OK)

CISTACEAE: 2/2

Helianthemum glomeratum C American

Lechea mensalis Chisos Mont. Brewster c Endemic

COMMELINACEAE: 6/2

e — S o M- Madrean

mmelina E N Am

o a Tecos Mont

Tradescantia leiandra Chihuahua

Tradescantia occidentalis Prairie S

Tradescantia wrightii N Chihuahuan

CONVOLVULACEAE: 16/6

Bonamia ovaliifolia N Chihuahuan

Bonamia repens N Chihuahuan

Convolvulus equitans SWC N American

Cressa truxillensis Madrean

Dichondra argentea Chihuahuan

Dichondra brachypoda Chihuahuan

Evolvulus alsinoides Madrean

Evolvulus nuttallianus WC N American

Evolvulus sericeus J Davis c. American Trop/Subtr

Ipomoea capillacea C America

Ipomoea cardio T Re Trop/Subtr

omoea costellata Sonoran

Ipomoea cristulata a

Ipomoea lindheimeri RORUN

Ipomoea rupicola N Chihuahuan

Ipomoea tenuiloba Sonoran

CRASSULACEAE: 6/3

Echeveria strictiflora Chihuahuan

Sedum cockerellii ) Davis c. AZ+NM

edum havardii Tr-Pecos-Coahuila

Sedum nanifolium Pecos- o Endemic

Sedum wrightii SWC N Amer

Villadia squamulosa a

ROSSOSOMATACEAE: 1/1

Glossopetalon spinescens W N American

d 8/6

anthera undulata Td Chihuahuan

Get dou NE rea

Y Y

n wide (radiations to WC US)

a dissecta Miser M a

Ibervillea lindheimeri Comanchian- Cem ?

Ibervillea tenuisecta Chihuahuan

Melothria pendula E N American (Gulf Coast)

Sicyos microphyllus Mesoam

CUPRESSACEAE: 8/2

Cupressus arizonica Madrean wide disjunct

428

Juniperus ashei Comanchian

Juniperus coahuilensis idi Chihuahuan (wide)

uniperus deppeana Madrea

Juniperus flaccida Brewster c. a in US. Madrean

Juniperus monosperma Madrean

Juniperus pinchotii SC US

Juniperus scopulorum W N American

CUSCUTACEAE: 6/1

Cuscuta applanata ?

Cuscuta indecora N American

Cuscuta leptantha ?

Cuscuta pentagona American

Cuscuta squamata Chihuahuan

Cuscuta umbellata :

rro 40/10

illaris American pl

Bulbostylis Juncoides S Meso

Carex agrostoides Sonoran- v: (N)

Carex athrostachya J Davis c. W N American

Carex frankii EN Am

Carex obl gt € da een

rex mesochorea E N Amer

Can E ne CS Prairie

Carex muriculata Pecos c. Chihuahuan

Carex occidentalis J Davis c. W N Am

Ps planostachys SC US-E Me + «Florida

ium mariscus Trop/Subtr

Gp erus acuminatus J Davis, ~ cc. N American

Cyperus echinatus E N Americ

s ala la S HM

erican African (Polich Ic

Aft ican (FOIICI \orous)

aie flavicomus J Davis c ?

Cyperus laevigatus Trop/Subtr

Cyperus niger American c en

Cyperus odoratus Am

Cyperus pci sc - (TX, N Mexico)

Cyperus retroflexus S N Am

Cyperus seslerioides En a btr)

Cyperus spectabilis Madrean wide

Cyperus sphaerolepis Madrean

rrosus American-African

Eleocharis macrostachya Am

Eleocharis montevidensis nine can

Eleocharis parishii Presidio c ee wide

Eleocharis parvula Holarctical

Lipocarpha aristulata i American

Lipocarpha micrantha American

Schoenoplectus acutus J Davis c. N American

Schoenoplectus maritimus Holarctical

Schoenoplectus pungens Polichorous

DENNSTAEDTIACEAE: 1/1

Pteridium aquilinum Holarctical

DRYOPTERIDACEAE: 4/2

Phanerophlebia umbonata Tamaulipan- -Chihuahuan

Woodsia neomexicana A

s phillipsii Sonoran Subprovin

sia plummerae Sonoran-N ENTIER

EBENACEAE: 1/1

Diospyros texana Chihuahuan proper

EPHEDRACEAE: 4/1

Ephedra antisyphilitica leal (+NE TX to OK)

Ephedra aspera Madrean

Ephedra torreyana SW N pend

Ephedra trifurca N Madrean

EQUISETACEAE: 3/1

quisetum arvense Holarctical

Equisetum hyemale Holarctical

Equisetum laevigatum N American

ERICACEAE: 1/1

Arbutus xalapensis Mesoam

EUPHORBIACEAE: 59/10

Acalypha monostachya Chuhuahuan

Acalypha neomexicana Sonoran

Acalypha ostryifolia American a American)

Acalypha phleoides Madrean w

Andrachne arida Tr-Pecos- quum

Argythamnia humilis S Prarie

Argythamnia neomexicana N Madrean

Bernardia obovata Chuhuahuan

hamaesyce acuta N Chuhuahuan

Chamaesyce albomarginata N Madrean

Chamaesyce angusta S TX Endemic

d. arizonica me Madrean (Sonoran)

ubprovince

peal chaetocalyx Apachian

Chamaesyce cinerascens Chinuahuan- oli

Chamaesyce dioica American Trop/Sub

Chamaesyce fendleri WC US

Chamaesyce geyeri Prairie

Chamaesyce glyptosperma N American

hamaesyce golondrina SW Texas Endemic

Chamaesyce hyssopifolia American Trop/Subtr

Chamaesyce jejuna N Chihuahuan

Chamaesyce lata SWC US

Chamaesyce micromera N Madrean

Chamaesyce missurica Prairie

Chamaesyce nutans C&N American

Chamaesyce perennans SW Texas Endemic

Chamaesyce serpyllifolia N American with E

amaesyce serrula Sonoran- a huahua

Chamaesyce setiloba N Madre

Croton dioicus Mexican-Chihuahuan

, Texas 429

Saahatelvan

- q r

T ai hi Lal + £L fi £D: D J

Croton fruticulosus Chihuahuan-Tamaulipan

Croton incanus E Madrean (Tamaulipan)

Croton lindheimerianus S Prairie

Croton monanthogynus E N American (E Prairie-Gulf Coast)

Croton pottsii Sonoran-Chihuahuan

Croton sancti-lazari Chihuahuan

Euphorbia brachycera W N American

Euphorbia cyathophora N American

Euphorbia davidii N American

Euphorbia dentata N American

Euphorbia eriantha N Madrean, disjunct

Euphorbia exstipulata N Madrean

Euphorbia spathulata N American

Jatropha dioica (only TX in US) Madrean

Phyllanthus ericoides N Chihuahuan

d nii polygonoides SC N American

Tragia amblyodonta Sonoran wide

Tragia ramosa: SWC U

FABACEAE: 107/37

Acacia angustissima American Trop/Subtr

Acacia berlandieri Chihuahuan

Acacia constricta Madrean wide

Acacia farnesiana Pantropical

Acacia greggii Madrean

Acacia neovernicosa Sonoran

Acacia rigidula Chihuahuan Pd

Acacia roemeriana Chihuahua

Acacia schottii (Brewster & sae cc) N Chihuahuan

Amorpha fruticosa Presidio c. N American

Astragalus crassicarpus Prairie wide

~ giganteus N Chihuahuan

agalus humistratus S Great Basin-N Sonora-Chihuahuan

haraga lotiflorus Prairie

Astragalus missouriensi S Rocky M-Prairie

Astragalus mollissimus WC N American

Astragalus e ies WC N American

us nuttallianus var. austrinus SWC N American

Endem

Brongniartia minutifolia ire Encemic

Calliandra conferta Madrean (S, C&E)

Calliandra humilis Sonora

Calliandra iselyi Tr-Pecos Pee

Cercis canadensis E N American

Crotalaria du American Trop/Subtr

Dalea aurea

Dalea pd oa Endemic

Dalea bicolor Sonoran

Dalea candida Prairie

Dalea enneandra Prairie

Dalea formosa SWC N American

Dalea frutescens E Madrean

Dalea lachnostachys N Sonoran-Chihuahuan

Dalea lanata SWC

Dalea laniceps N Chihuahuan

Dalea leporina American, mostly Mesoam

Dalea nana Sonoran wide

Dalea neomexicana BR Sonoran-Chihuahuan

Dalea neomexicana var. cM 5 Chihuauan-Tamaulipan

Dalea pogonathera Son

Dalea wrightii Sonoran- chat

Desmanthus cooleyi Apachia

Desmanthus glandulosus N ee Chihuahuan

Desmanthus illinoensis Prairie

Desmanthus obtusus N Chihuahuan

Desmanthus velutinus N Chihuahuan

Desmanthus virgatus American (except N)

Desmodium grahamii Madrean

J FA hi : n

Desmodium psilophyllum Mesoam-E Madrean

Eysenhardtia spinosa Presidio c. Chihuahuan ?

Eysenhardtia texana Chihuahuan

Galactia canescens J Davis c. Chihuahuan

o wrightii Sonoran

enistidium dumosum Chihuahuan local

chon lepidota WC N American

Hoffma

ggia oxycarpa Chihuahuan

Indigofera lindheimeriana Chihuahuan

Indigofera miniata Mesoam (Caribbean)

Lespedeza texana Comanchian

Leucaena retusa pasate

Lotus plebeius SW N American Mont

Lupinus concinnus N Madrean

Lupinus havardii SW TX, N Tamaulipan, MO ?

Macroptilium gibbosifolium Mesoam

Mimosa aculeaticarpa Madrean wide

Mimosa borealis SWC US

Mimosa dysocarpa Sonoran w

Mimosa emoryana Chihuahuan: N Highlands disjunct

Mimosa texana Chihuahuan ?

MIT iChihiiahi

ran dicitinct ?

Nissolia platycalyx Chihuahuan disjunct, local

Parkinsonia aculeata em

Peteria scoparia Apachia

Phaseolus oe Subprovince

Pomaria jam C US wide

Pomaria t Tr-Pecos-Coahuila

Prosopis glandulosa W N Sisi (mostly S)

Prosopis pubescens N Madre

Psoralidium tenuiflorum EM SWC US

Rhynchosia senna Amphitropical

Senna alata Trop/Subtr

430

Senna bauhinioides Madrean

Senna durangensis Chihuahuan-Tamaulipan disjunct local ?

Senna lindheimeriana Chihuahuan-Tamaulipan

Senna orcuttii Sonoran-Chihuahuan local

Senna pilosior Chihuahuan

Senna pumilio (S Prairie-Chihuahuan) SC N American

Senna roemeriana (NM & W TX) SWC US

Senna ripleyana Brewster c. ?

Senna wislizeni N & E Madrean (disjunct)

Sesbania herbacea Mesoam

Tephrosia vicioides Mesoam

Vicia americana N American (mostly W & N)

Vicia ludoviciana N American (S)

FAGACEAE: 20/1

Quercus arizonica Sonoran-Chihuahuan

Quercus carmenensis Chihuahuan local

Quercus depressipes J Davis c. Chihuahuan local

Quercus emoryi Sonoran-Chihuahuan

Quercus gambelii W N American

Quercus graciliformis Brewster c. Endemic

Quercus hinckleyi Presidio Endemic

Quercus hypoleucoides Sonoran-Chihuahuan

Quercus intricata Chihuahuan

Quercus laceyi E Madrean

Quercus mohriana SWC US

Quercus muehlenbergii EN American

Quercus oblongifolia W Madrean

Quercus pungens Madrean (E)

Quercus robusta Brewster c. Endemic

Quercus rugosa Mesoam-Madrean

Quercus tardifolia Brewster c. Endemic

Quercus turbinella SW N American

Quercus vaseyana Chihuahuan

FOUQUIERIACEAE: 1/1

Fouquieria splendens Sonoran-Chihuahuan (N&W Madrean)

FUMARIACEAE: 2/

Corydalis aurea Prairie N American (WC)

Corydalis anus Prairie N American (C-SW)

GARRYACEAE: 2/1

Garrya ovata uc Madrean

Garrya wrightii Sonoran

GENTIANACEAE: 6/3

Centaurium arizonicum N Madrean

Centaurium beyrichii Comanchian (SC US ?)

Centaurium calycosum E Madrean

Centaurium glanduliferum Tr-Pecos Endemic

Eustoma exaltatum Mesoam-N American

Gentiana affinis W N American

GERANIACEAE: 3/2

Erodium texanum SWC US ?

Geranium caespitosum W N American

Geranium carolinianum J Davis c. American

GROSSULARIACEAE: 3/1

Ribes aureum N American (not E & SE

Ribes leptanthum J Davis c. SW N American (US)

Ribes mescalerium SWC US (SW TX-SC NM)

HYDRANGEACEAE: 8/2

Fendlera rigida INE Coahuila

Fendlera denne Apachian

Fendlera wi

Fendlerella pU SW US

Philadelphus crinitus ) Davis c. Endemic ?

Philadelp (ond mearnsii Amphitropical ? (SW US)

Philadelphus microphyllus SW US wide

Philadelphus po Sw US disjunct

HYDROPHYLLACEAE: 15/2

Nama dichotomum W American

Nama havardii N Chihuahuan

Nama hispidum SWC N American

Nama parvifolium Chihuahuan-Tamaulipan

Nama torynophyllum Chihuahuan

Nama undulatum Chihuahuan-Tamaulipan

Phacelia caerulea Sonoran Subprovince

Phacelia congesta SC N American

Phacelia crenulata SW US

Phacelia infundibuliformis Tr-Pecos

Phacelia integrifolia SWC U

Phacelia pallida Tr-Pecos, rare

Phacelia popei S

Phacelia robusta Tr-Pecos-Coahuila

Phacelia i. N Sonoran-Chihuahuan

IRIDACEAE

esee o. m)?

Sisyrinchium cernuum J Davis c. W Madrean

JUGLANDACEAE: 3/2

Carya illinoinensis J Davis c. EN American

Juglans major Madrean wide

Juglans microcarpa SC N American

JUNCACEAE: 8/1

Juncus acuminatus N American-C American

Juncus arcticus J Davis c. Polichorous (Arcto-Mont)

Juncus bufonius Polichorous

Juncus ensifolius W N American

Juncus interior Prairie wide

Juncus marginatus J Davis c. American

Juncus nodosus N American

Juncus torreyi N American

KRAMERIACEAE: 3/1

Krameria erecta Sonoran (mostly W)

Krameria grayi Sonoran

Krameria lanceolata SC N American (4- Florida)

LAMIACEAE: 38/11

Agastache micrantha Chihuahuan (mostly W)

(ed de Sonoran

a E Madrean-Caribbean

He "n ma pid WC N American

Hedeoma mollis Tr-Pecos Endemic

r Le Lal + £ al £l fD: D

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, Texas 431

Hedeoma reverchonii Comanchian (TX, S OK)

Hedeoma reverchonii var. t UEM ? STX

Mentha arvensis Holarctical

Monarda citriodora A US

Monarda fistulosa J Davis c. N American

Monarda pectinata WC US

Physostegia virginiana J Davis c. E N American

ACE glabrescens Chihuahua

NM A

n (mostly Mohavean)

Salvia arizonica Sonoran -Chihuahuan disjunct

Salvia az

Salvia age SC US (S Prairie)

Salvia greggii Chinuahuan-Tamaulipan

Salvia henryi Sonoran- Chihuahuan (W)

Salvia lycioides Chihuahua

Salvia reflexa N ed onm Prairie)

Salvia regla M

Salvia roemeriana Us proper

Salvia texana Pecos c. SC N American

Salvia tiliifolia Mesoam-S American (NW)

Salvia whitehousei N Chihuahuan

Scutellaria drummondii SC-SE N American

Scutellaria potosina Sonoran-Chihuahuan

Scutellaria texana Chihuahuan

Scutellaria wrightii SC N American (TX-OK)

Stachys bigelovii Chihuahuan

Teucrium canadense J Davis c. N American-Mesoam

Teucrium coahuilanum Tr-Pecos- ee

Teucrium cubense Mesoam-Madre

Teucrium laciniatum SWC US

LEMNACEAE: 3/2

Lemna minuta J Davis c. American

Lemna valdiviana American

Spirodela polyrrhiza ) Davis c. American

LILIACEAE: 16/8

Allium cernuum N American

Allium glandulosum Sonoran-Mexican Highlands

Allium drummondii SC US (Prairie)

bit did Prairie

ooperia drummondii Pecos c. SC US (+NE Mexico)

uda eosin Chihuahua

Dasylirion texanum Chihuahuan ae

ea flavescens J Davis c. Sonoran (W)

ianthemum racemosum J Davis c. N American Mont

No s erumpens Chihuahuan

Nolina texana SWC U

Nothoscordum bivalve EN American-Mesoam

Schoenocaulon texanum Chihuahuan

Zephyranthes longifolia N Sonoran-Chihuahuan

LINACEAE: 8/1

Linum aristatum SW N American

Linum berlandieri Prairie (SC)

Linum hudsonioides S Finis

Linum lewisii W N American

Linum puberulum W 3 A em

Linum rupestre Madrean wide

Linum schiedeanum E Madrean-Mesoam

Linum vernale Chihuahuan

LOASACEAE: 11/3

Cevallia sinuata Sonoran wide

cnide bartonioides Chihuahuan-Tamaulipan

Mentzelia albescens Amphitropical

Mentzelia asperula N Sonoran disjunct

E lindheimeri Chihuahuan-Tamaulipan

ntzelia mexicana Chihuahuan

e multiflora Madrean (N)-S Rocky M

Mentzelia oligosperma Prairie

Mentzelia pachyrhiza Chihuahuan

Mentzelia reverchonii SC

Mentzelia saxicola Presidio c. Chihuahuan

LYTHRACEAE: 2/2

Lythrum californicum W N American

Nesaea longipes N Chihuahun

MALPIGHIACEAE: 1/1

Janusia gracilis Sonoran (+W Chihuahuan)

VACEAE: 24/1

Abutilon fruticosum Tropical

Abutilon malacum Sonoran

Abutilon parvulum Sonoran

Abutilon wrightii Chihuahuan

Allowissadula holosericea Chihuahuan-Tamaulipan

Batesimalva violacea Chihuahuan-Tamaulipan

Callirhoe involucrata Prairie

Herissantia crispa Trop/Subtr

Hibiscus coulteri Sonoran wide

Hibiscus denudatus S California-Sonoran

Malvella lepidota Sonoran

Malvella leprosa Amphitropical ?

Malvella Rn dies Sonoran

Meximalva filipes Tamaulipan

Mo P caroliniana American deed

Sida longipes Chihuahuan- dm

Sida neomexicana Sonoran-Chihuahuan ?

Sphaeralcea angustifolia Madrean

Sphaeralcea coccinea W N American

Sphaeralcea digitata S Great Basin-Chihuahuan

Sphaeralcea hastulata S Great Basin-S Rocky M-Sonoran wide

DUREN incana S Great Basin- 5 Rocky M-Chihuahuan

hyll S Rocky M-N Chihuahuan

MARSILEACEAE: 2/1

Marsilea mollis Mesoam-S American

Marsilea vestita N American (+ Peru)

MENISPERMACEAE: 1/1

Cocculus carolinus E N American

MOLLUGINACEAE: 1/1

Mollugo verticillata American (Trop/Subtr)

432

MORACEAE: 2/1

Morus microphylla S Great Basin-Sonoran-N Chihuahuan

NAJADACEAE: 1/1

Najas guadalupensis American

NYCTAGINACEAE: 32/11

Abronia angustifolia S Great Basin-Sonoran

Acleisanthes acutifolia Chihuahuan

Acleisanthes longiflora Sonoran wide

Acleisanthes wrightii SW TX Endemic

mocodon C

join eriosolenus Chihua

Anulocaulis leiosolenus di N aco

Anulocaulis reflexus N Chi an loc

Boerhavia anisophylla Madrean

Boerhavia coccinea Trop/Subtr

Boerhavia erecta American bd

Boerhavia gracillima Sonoran

Boerhavia intermedia Sonoran Med (N&W Madrean)

Boerhavia linearifolia Chihuahuan (+N NM)

Boerhavia torreyana Apachian

Boerhavia spicata Sonoran (W)

Boerhavia wrightii N Madrean

mm ndens American uem

e crassifolia E Chihua

Cyphomeris gypsophiloides Ca ee

Mirabilis albida N American (Prairie-SW N een

Mirabilis linearis W N American (+Prairie)

Mirabilis longiflora Sonoran wide

Mirabilis multiflora SW N American

Mirabilis a SW N American

Mirabilis texensis SW TX

Nyctaginia p Chihuahua

Selinocarpus angustifolius an

Selinocarpus diffusus Pecos c. SWC US (W TX-NM)

Selinocarpus parvifolius Tr-Pecos Endemic

OLEACEAE: 8/3

Forestiera angustifolia Chihuahuan-Tamaulipan

Fraxinus cuspidata N & E Madrean Mont

Fraxinus greggii E Madrean Mont

Fraxinus papillosa N Sonoran-Chihuahuan

raxinus velutina N&W Madrean Mont

Menodora heterophylla N Chihuahuan (S TX)

Menodora longiflora N Chihuahuan (S TX)

Menodora scabra Madrean (no Mexican Highlands)

ONAGRACEAE: 28/4

Camissonia chamaererodes N Madrean

Ludwigia peploides American (Polichorous)

Ludwigia repens E N American (with radiations)

Oenothera arida pred

Oenothera albicaulis WC US

alode boquilensis eminus nuan

n (no Mexican Highlands)

Oenothera calcicola E Madea

Oenothera coccinea N American-Mesoam

Oenothera elata subsp. hirsutissima W N American

Oenothera engelmannii SWC US ? (E NM-W TX)

Oenothera grandis Prairie

Oenothera hartwegii SWC US

Oenothera havardii Chihuahuan

Oenothera hexandra Mesoam-Madrean

Oenothera kunthiana Mesoam-Madrean

Oenothera lavandulifolius W N American

Oenothera macrocarpa Tr-Pecos

Oenothera pallida W US (not Pacific)

Oenothera primiveris N Madrean

Oenothera rosea Trop/Subtr (mostly n

Oenothera speciosa N American-Meso

Oenothera suffulta subsp. nealleyi N ies

Oenothera texensis Chihuahuan

Oencthera triloba Prairie

Oenothera tubicula N Chihuahuan

ORCHIDACEAE: 9/4

Dehomantus une E ue

Epipactis gi a W N America

Hexalectris go! Davis c. — disjunct

Hexalectris nitida SE NM-S TX rare

Hexalectris Eus ? (S AZ-SE NM-S TX)

Hexalectris spicata EN American (also S N America)

Hexalectris warnockii ? (S AZ-SW-SC TX)

? Malaxis erhenbergii Chisos, Brewster c. Endemic

Malaxis soulei J Davis c. American Trop/Subtr

OROBANCHACEAE: 4/2

Conopholis alpina Madrean-Mesoam (C American)

Orobanche ludoviciana N American (US)

Orobanche multicaulis Mexican Highlands-Chihuahuan ?

Orobanche uniflora J Davis c. N American

OXALIDACEAE: 3/1

Oxalis albicans Midi wide

Oxalis dillenii N Am

xalis drummondii E R Chihuahuan

PAPAVERACEAE: 4/1

Argemone aenea J Davis N in -Tamaulipan

Argemone chisosensis Chihuahua

Argemone polyanthemos J Davis c. or irie

Argemone sanguinea E Chihuahuan-Tamaulipan

Argemone squarrosa SWC US

SSIFLORACEAE: 1/1

Passiflora tenuiloba Tamaulipan-N Chihuahuan

PEDALIACEAE: 4/1

o althaeifolia Sonoran

roboscidea louisianica N American (mostly S US)

sn cidea parviflora N n

Proboscidea spicata N a (local)

PHYTOLACCACEAE: 2/2

Phytolacca americana N American (mostly E)

Rivina humilis American Trop/Subtr

PINACEAE: 5/2

Pinus cembroides Madrean wide

r biral anl $ £l a D:n D J

P Texas 433

Saahatelvan

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awe 2

Pinus ponderosa W N American

Pinus remota Trans-Pecos-Coahuila

Pinus strobiformis J Davis c. Apachian (AZ, NM)

Pseudotsuga menziesii W N American

PLANTAGINACEAE: ~

Plantago helleri SC U

Plantago hookeriana A US (TX + Coahuila)

da ovara S Palearctica Ace

SIAM

entive in SW)

P Tali (adv

Plantado major Polichorous

Plantago patagonica American

Plantago wrightiana S N American

PLUMBAGINACEAE: 2/2

Limonium limbatum Pecos c. SWC US (Apachian-N Chihuahuan)

Plumbago scandens Trop American

POACEAE: 202/63

Achnatherum aridum N Madrean

Achnatherum curvifolium N Chihuahuan

Achnatherum eminens CS Madrean (proper)

Achnatherum lobatum N oe

Agrostis exarata NAm

Agrostis hyemalis

Agrostis Pala al (Boreal)

Allolepis texana Tr-Pecos-Coahuila

Aristida adscensionis n (Trop/Subtr)

Aristida arizonica N

Aristida desmantha = ES d TX €: Tr-Pecos)

Aristida divaricata Mesoam-Madrean

Aristida havardii Madrean proper

Aristida pansa Madrean proper

Aristida purpurea N American (W-C)

Aristida purpurea var. nealleyi SW N American

Aristida purpurea var. wrightii SW N American

Aristida schiedeana Mesoam-N S American

Aristida ternipes Mesoam-Madrean-N S American

Blepharidachne bigelovii Tr-Pecos-Coahuila

Blepharoneuron tricholepis S Rocky M-Madrean

Bothriochloa alta Mesoam-Amphitropical Mont

Bothriochloa barbinodis American Trop/Subtr

Bothriochloa laguroides American (subtr)

Bothriochloa springfieldii American (subtr)

Bouteloua aristidoides American (subtr)

Bouteloua barbata American (subtr)

Bouteloua breviseta Chihuahuan

Bouteloua curtipendula American

Bouteloua dactyloides Prairie wide (+ Mexico)

Bouteloua eriopoda SWC US (N Madrean)

Bouteloua gracilis WC N American

Bouteloua hirsuta Prairie-N Madrean

Bouteloua simplex J Davis c. American (Warm-Trop/Subtr)

Bouteloua trifida N Madrean

Bouteloua uniflora Chihuahuan

Bouteloua warnockii N Chihuahuan

Bromus anomalus Mesoam-Madrean

Bromus arizonicus J Davis c. California-Sonoran

Bromus ciliatus Holarctical

Bromus lanatipes SW ic oe -Apachian)

romus marginatus W N Americ

Bromus polyanthus W N nine Mont

Bromus rubens Pecos c. W N American

?Cathestecum erectum S AZ-Tr-Pecos-Guatemala ?

Cenchrus longispinus N American

Cenchrus haa AmericanTrop/Subtr

Cenchrus spinifex

Chloris cucullata 2 N American

Chloris submutica CS Madrean

Chloris texensis SW & SETX nndis

Chioris a Prairie (+WN American)

gata American-African (o uH

Cot el e American (Amphitropical)

SE pulchella Madrean wi

Dichanthelium acuminatum n American-Mesoam

Dichanthelium oligosanthes N American

PII d 2m pee al

-Trop/ AMICI

Digitaria n E N American

Digitaria hitchcockii S TX- Coahuila

Digitaria patens SW-SC TX Endemic

Digitaria pubiflora SWC US-Chihuahuan (+Tamaulipan)

Digitaria sanguinalis Warm/Subtr

Distichlis spicata American

Echinochloa colona Trop/Subtr/Warm

Echinochloa crus-galli Subtr/Warm

Echinochloa crus-pavonis Presidio c. Trop/Subtr/Warm

(American-African)

Echinochloa muricata N Amer

Elionurus barbiculmis DE TEn Highlands

Elionurus tripsacoides American Trop/Subtr

Elymus arizonicus Sonoran-Chihuahuan

Elymus canadensis N American

Elymus elymoides W N American

o N American ~

eapogo auxii Warm/Su

o nk Warm Temp/ ae (+ Trop Africa)

Eragrostis cilianensis Polichorous

dees TR SC US

Eragrost a J Davis, Presidio cc. N Chihuahuan

SERA nde SW-SC US-Madrean

Eragrostis lehmanniana Sonoran wide

Eragrostis lugens American Trop/Subtr

Eragrostis mexicana W American

Eragrostis palmeri Chihuahuan wide

Eragrostis pectinacea American

Eriochloa acuminata ?

Erioneuron avenaceum Am a

Erioneuron nealleyi Chihua

Erioneuron pilosum WC N NM

Festuca arizonica SW

Festuca ligulata Chihuahuan

Festuca rubra Holarctical

—

434

Hesperostipa neomexicana SW US

Heteropogon contortus Trop/Subtr

Hilaria belangeri Chihuahuan wide

Hilaria swallenii N Chihuahuan

Hordeum jubatum Holarctical

Hordeum pusillum N American

Imperata brevifolia N&W Madrean

Koeleria macrantha Holarctical

Leersia oryzoides Holarctical

Leptochloa dubia American Warm/Subtr

Leptochloa fusca Polichorous

eptochloa panicea Trop/Subtr (America-Asia)

Lycurus phleoides American Trop/Subtr

Melica nitens Appalachian (E N American)

Melica porteri SW US (Colorado-Apachian)

Microchloa kunthii Trop/Subtr (American-African)

Monroa squarrosa WC US

Muhlenbergia N Sonoran-Chihuahuan

Muhlenbergia arenicola SWC US-Chihuahuan

Muhlenbergia asperifolia W American (N Temp)

Muhlenbergia brevis SWC US-Chihuahuan

Muhlenbergia depauperata W did Mont

Muhlenbergia dubia Madrea

Muhlenbergia eludens es ed huahuam Mont

veda emersleyi Madaan wide

Mont (mostly W)

Muhlenbergia glauca Madrean

uhlenbergia minutissima W N American

Muhlenbergia montana W N American

Muhlenbergia pauciflora SW US-W Madrean Mont

Muhlenbergia polycaulis W Madrean Mont

Muhlenbergia porteri N8WC Madrean

Muhlenbergia repens Madrean

Muhlenbergia rigens NC Madrean

Muhlenbergia rigida W American Mont

Muhlenbergia schreberi EN American (+S America)

Muhlenbergia setifolia Chihuahuan

Muhlenbergia spiciformis ? Caribbean

Muhlenbergia tenuifolia Madrea

Muhlenbergia texana W Madrean

Muhlenbergia torreyi SW US

Muhlenbergia villiflora Chihuahuan Mont

Nassella leucotricha da E&S adrear

Panicum bulbosum ATE ric

Panicum capillare N Modo (+Temp S America)

DCUM pap E nA

J. JC ba

lily ‘Trop/Subtr)

Panic obtusum SWC N ae

Panicum virgatum N American wide

D le bicolor O Mexican Highlands

phit ropical

B smithil WUS

Paspalum distichum Warm Temperate/Trop/Subtr

Paspalum pubiflorum E N American-Mesoam

Phalaris angusta American (coastal)

Phalaris caroliniana N American (coastal)

Phragmites australis Polichorous (warm)

Piptochaetium fimbriatum Madrean wide

Piptochaetium pringlei ) Davis c. W Sonoran

Pleuraphis jamesii Presidio c. SW US

Pleuraphis mutica Sonoran wide

Poa arachnifera S Prairie

Poa bigelovii N Madrean (SWC US)

Poa fendleriana W N American

Poa strictiramea Brewster c Endemic Mont

Polypogon elongatus American Trop/Subtr

Polypogon interruptus American (W American)

Pseudoroegneria spicata W N American

Schedonnardus paniculatus Prairie

Schizachyrium cirratum Madrean (+N S America)

Schizachyrium sanguineum m oem

Schizachyrium scoparium N A

Schizachyrium spadiceum E Coahuila

Scleropogon brevifolius Madrean-Mesoam

Setaria grisebachii Madrean-Mesoam

Setaria leucopila SWC US-E Madrean

Setaria parviflora Trop/Subtr-Warm Temperate

Setaria reverchonii SC

Setaria scheelei Chihuahuan-Tamaulipan

Setaria villosissima Chihuahuan very local

Sporobolus contractus J Davis & Presicdio cc. SW US

Sporobolus cryptandrus N American

Sporobolus flexuosus SW US

Sporobolus pyramidatus American

Sporobolus texanus S

Sporobolus wrightii Madrean wide

Tridens eragrostoides Gulf Coast- oe Chihuahuan

een autau pane Hee d

marican m" NA S American

Trisetum interruptum SWC US (S Prairie-Gulf Coast ?)

Urochloa arizonica Sonoran wide-Mexican Highlands

Urochloa fusca American Trop/Subtr

Vulpia octoflora N American

POLEMONIACEAE: 10/4

Gilia stewartii Chihuahuan wide

Giliastrum acerosum SWC US

Giliastrum incisum Chihuahuan wide

Ipomopsis havardii A oe

lpomopsis laxiflora SWC U

Ipomopsis longiflora e US-E Prairie

Ipomopsis pumila WC US

Phlox nana Apachian

Fal [9 al 4 £l n Din D J

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POLYGALACEAE: 9/1

ala alba Prairie-Madrean wide

Polygala barbeyana Madrean wide

Polygala hemipterocarpa J Davis c. Sonoran-Chihuahuan

Polygala Wane uut Chihuahua

de Meyi

an Hinhlands<

Polygala maravillasensis Chinua local

Polygala nudata Chihuahuan

Polygala obscura J Davis c. N Sonoran-Chihuahuan

Polygala scoparioides Sonoran-Chihuahuan wide

POLYGONACEAE: 20/3

Eriogonum abertianum (S Great Basin)-Sonoran-Chihua-

huan

Eriogonum alatum WC US

Eriogonum annuum Prairie

Eriogonum havardii N Chihuahuan

Eriogonum hemipterum Tr-Pecos-Coahuila

Eriogonum hieraciifolium S Great Basin-Chihuahuan low

unt

Eriogonum jamesii SWC US

Eriogonum longifolium Pecos c SC US

Eriogonum nealleyi Pecos c. S TX Endemic

Eriogonum polycladon J Davis c. S Great Basin-Sonoran

Eriogonum rotundifolium Chihuahuan

Eriogonum suffruticosum Tans Pecos Endemic

Eriogonum tenellum Colorado, E New Mexico, W OK, N, W,

C& SWTX ?

Eriogonum wrightii N Madrean

Polygonu ac var. omissum ?

Polygonum punctatum Am

Polygonum un EM 7 ai

Polygonum glabrum Atlantic-Gulf Coast-Caribbean

(+Madagascar&China)

Polygonum hydropiperoides American

Rumex maritimus N &W erican

PODIACEAE:

Pleopeltis riograndensis N Sonoran-Chihuahuan

AE: 3/1

Heteranthera dubia N American

Heteranthera reniformis Mesoam-S American

Heteranthera rotundifolia Mesoam-5 American

PORTULACACEAE: 9/4

Phemeranthus aurantiacus Apacian-E Madrean

Phemeranthus brevicaulis N Chinuanuan+ NE NM

Phemeranthus longipes ? (Apachian)-Chihuahuan-Tamaulipan

Phemeranthus paie ~

Portulaca pilosa T

Portulaca drei Sonoran

Portulaca umbraticola American Trop/Subtr

Talinopsis bcd: Chihuahuan

alinum culatum American

POTAMOGETONACEAE: 2/1

Potamogeton foliosus Presidio c. N American

Potamogeton nodosus Polichorous

PRIMULACEAE: 3/2

Androsace occidentalis N American

, Texas 435

Androsace septentrionalis Holarctical

Samolus ebracteatus Mesoam-Madrean

PTERIDACEAE: 33/8

Adiantum capillus- veneris Trop/Subtr/Warm

Madrean wide

Argyroch

Astrolepis cochisensis Madrean wide

pd oe ea wide

nuata American Trop/Subtr/Warm

Bommeria hispida Sonoran

Cheilanthes alabamensis Appalachian-N Madrean

Cheilanthes bonariensis American-African

Cheilanthes eatonii Madrean wide

Cheilanthes feei N Americ

Cheilanthes horridula E m

Cheilanthes kaulfussii Mesoam-Central American

Cheilanthes lendigera Mesoam-Central American

Cheilanthes lindheimeri Madrean

Cheilanthes tomentosa S& SEN American

Cheilanthes villosa Sonoran-Chihuahuan

Cheilanthes wrightii Sonoran

Notholaena aliena Chihuahuan ?

Notholaena aschenborniana E Madrean

Notholaena copelandii E Madrean

Notholaena grayi Sonoran-Chihuahuan

Notholaena greggii Chihuahuan

otholaena nealleyi E Madrean ?

Notholaena neglecta Chihuahuan

Notholaena standleyi E Madrean-Apachian

Pellaea atropurpurea N American-Mesoam

Pellaea intermedia Sonoran-Chihuahuan

Pellaea ovata Mesoam-S American

Pellaea ternifolia Mesoam-S American

Pellaea truncata J Davis c. N Madrean

Pellaea wrightiana ? (N American, sporadic)

RAFFLESIACAE: 1/1

Pilostyles thurberi SW N American disjunct

RANUNCULACEAE: 11/7

je chrysantha Apachian-N Sonoran SW US

Aquilegia longissima N Sonoran ? N Chihuahuan ?

SE drummondii SOlgiad wide

X. EN American (wide)

Delphinium wootonii Apac

Delphinium madrense Hin NU T

Thalictrum fendleri Presidio, J Davis W N American

RESEDACEAE: 1/1

pres linifolia Madro (N Madro)-Tethyan ?

RHAMNACEAE: 11/9

Adolphia infesta C-S Madrean Vai Highlands-Sonoran)

Ceanothus greggii Madrean M

Colubrina texensis | one,

436

Condalia ericoides Sonoran Mont

Condalia viridis Chihuahuan (+Sonora)

Condalia warnockii Sonoran

Frangula betulifolia Madrean wide Mont

Karwinskia humboldtiana Caribbean-Sonoran

Ziziphus obtusifolia Madrean

ROSACEAE: 12/9

Cercocarpus montanus W N American Mont

Crataegus tracyi S TX-Coahuila

Fallugia paradoxa NC Madrean

Holodiscus dumosus W N American (not coastal)

Malacomeles denticulata ? Mesoam Mont disjunct

Petrophyton caespitosum W N Am

Prunus havardii Tr-Pecos Endemic eee

Prunus murrayana Tr-Pecos Endemic Mont

Prunus serotina ? American-Europea

Prunus virginiana J Davis c. N American (not SE)

Purshia ericifolia Tr-Pecos-Coahuila

Vauquelinia corymbosa Chihuahuan-Tamaulipan disjunct

Mont

RUBIACEAE: 17/6

ridi ternifolia Mesoam Mont

ephalanthus occidentalis N American (not Central)

re md Chihuahuan local

alium mexicanum Mesoam

Galium ie eal (not Cal)

Galium proliferum N Ma E.

Galium uncinulatum Mes

Galium virgatum SC US cora

Galium wrightii N Madrean Mont (mostly Sonoran)

Hedyotis intricata Chihuahuan

Houstonia acerosa S Rocky M-Chihuahuan Mont

Houstonia humifusa SC US

Houstonia wrightii ) Davis c. NW Sonoran

Stenaria butterwickiae Tr-Pecos Endemic

Stenaria mullerae N Chihuahuan local

Stenaria nigricans E N American

Stenaria rupicola N Chihuahuan (Tr-Pecos ?)

RUTACEAE: 4/4

Choisya dumosa Son

diis trifoliata N n E -Europea

nosma texana N Sonoran rarer

ee parvum Tr-Pecos Endemic Mont

SALICACEAE: 6/1

Salix a da C. N American

Salix exiqua W N Am

Salix Bore N ea

Salix iix N American

Salix nigra EN A

Salix taxifolia alus

SAPINDACEAE: 2/2

Sapindus saponaria Trop/Subtr

Ungnadia speciosa Chihuahuan wide? (+SCW Africa + SW China)

J Lanf tha Dat : ID hI "rn £T,

SAPOTACEAE: 1/1

Sideroxylon lanuginosum SC& SE US

SAURURACEAE: 1/1

Anemopsis californica W Madrean

SAXIFRAGACEAE: 1/1

euchera rubescens W Madrean

SCROPHULARIACEAE: 29/13

Agalinis calycina Chihuahuan local

Bacopa monnieri Trop/Subtr

Bacopa rotundifolia ? American temperate

uchnera americana E N American

Castilleja sessiliflora Prairie

Castilleja mexicana Chihuahuan

Castilleja integra S Rocky M-W Madrean

Castilleja lanata Madrean

Castilleja rigida Chihuahuan

Epixiphium wislizeni SWC US

Leucophyllum candidum Chihuahuan

Leucophyllum frutescens Chihuahuan-Tamaulipan

d d Eua

Mesoam-Madrean

Mecardonia procumbens American Trop/Subtr

Mimulus dentilobus W M iie rean

Mimulus glabratus Americ

Nuttallanthus texanus N oum

Penstemon ambiguus SWC US

Penstemon n Chihuahuan

Penstemon barbatus SW US-Madrean Mont

Penstemon desp Sonoran-Chihuahuan (W)

Penstemon fendleri SWC US

Penstemon havardii TrPecos Coahuila Mont

Penstemon jamesii Apac

Penstemon ramosus ded

Penstemon wrightii Tr-Pecos Endemic

Seymeria scabra Chihuahuan

Veronica peregrina Polichorous

SELAGINELLACEAE: 9/1

aginella arizonica Sonoran

a. a E&S Madrean Mont

ela ica Rocky M

Sela ee aie ona Arerian

Selaginella ru Tr-Peco taro disjunct

Selaginella rupincola PE Highlands- rn Mont

Selaginell oodii W N American (Rocky M)

a underw

Selaginella uu Tr-Pecos-Coahuila

Selaginella wrightii Chihuahuan-Tamaulipan

SIMAROUBACEAE: 1/1

Holacantha stewartii Chihuahuan

SOLANACEAE: 32/10

Calibrachoa parviflora American (SN, Mesoam)

Chamaesaracha coniodes SC N American

Chamaesaracha villosa Chihuahuan

Datura quercifolia ?

r Lt Lal > £ al ii £D:

Texas

Saahatelvan

» P4 ,

Datura wrightii N American (mostly W)

Lycium nd vir wide

Lycium pallidum

ycium la a

Lycium texanum Tr-Pecos Endemic

Lycium torreyi N Madrean

Margaranthus solanaceus Madrean

Nicotiana obtusifolia Madrean (N+W)

Nicotiana repanda Chihuahuan ?

Physalis angulata American Trop/Subtr

Physalis cinerascens SC N American-Mesoam

Phys alis hederifolia Madrean- Prairie

Solanum citrullifolium ?

Solanum davisense ir-Pecos-Coahuila

Solanum douglasii American Trop/Subtr

Solanum elaeagnifolium American

Solanum Karo adu Tr-Pecos-Coahuila

Solanum ptychanthum N American

Solanum tenuipes pen

Solanum triquetrum A

STERCU vibe 4/1

onoran

Ayenia il Sonoran-Chihuahuan

Ayenia pilosa Tamaulipan

Melochia a American Trop/Subtr

THELYPTERIDACEAE: 1/1

Thelypteris ovata Gu!f Coast-Caribbean

TYPHACEAE: 1/1

Typha domingensis Trop/Subtr

—

ULMACEAE: 3/1

Celtis pallida Torr. Mesoam-S American

Celtis laevigata EN American

Celtis reticulata W N American wide (not Pacific)

URTICACEAE: 1/1

Parietaria pensylvanica N American

VERBENACEAE: 21/8

Aloysia gratissima Amphitropical

Aloysia wrightii Sonoran wide

Bouchea linifolia Chihuahuan (S TX- Coahuila)

Bouchea spathulata Chihuahuan (Tr-Pecos-Coahuila)

Glandularia bipinnatifida N American (Prairie)

landularia pumila SC U

Glandularia d o -Tamaulipan

graveolens Mesoam-C Am

antana aa qune s Pm (subtr

a antana urticoides ? Madrean with radiations

Phyla cuneifolia WC US. Prairie

Phyla fruticosa American Trop/Subtr

—

Verbena bracteata J Davis & Presidio cc. N American

Verbena canescens E Madrean

Verbena halei E Madrean-Gulf Coast

Verbena neomexicana Sonoran

Verbena perennis Sonoran

juin plicata SWC US

Verbena scabra Mesoam

VIOLACEAE: 1/1

Hybanthus verticillatus SWC US-Prairie

VISCACEAE: 6/1

Phoradendron coryae Son

Phoradendron ed * Chihuahuan

Phoradendron juniperinum W N American

Phoradendron macrophyllum MM ‘Sonoran

Phoradendron tomentosum E-S N Am

Phoradendron villosum ?

VITACEAE: 2/2

Cissus trifoliata Mesoam-S N American

Vitis arizonica Madrean wide

ZANNICHELLIACEAE: 1/1

Zannichellia palustris Polichorous

ZYGOPHYLLACEAE: 8/4

Guajacum angustifolium Chihuahuan-Tamaulipan

Kallstroemia californica N & W Madrean

Kallstroemia grandiflora W Madrean

Kallstroemia hirsutissima Sonoran-Chihua

437

Kallstroemia parviflora Madrean pns Amphitroi (Peru)

Kallstroemia perennans SW TX E

Larrea tridentata Madrean wide (W a

Peganum mexicanum Chihuahuan-S Sonora

ACKNOWLEDGMENTS

I deeply thank Michael Powell and Steven McLaughlin for review and helpful comments, S. McLaughlin, S.

Manchester, and M. Lavin for the offprints of their works, and my husband A. Karabegov for the help with

the wording. This work was supported by a grant from Sam Taylor Foundation. I am very thankful to all

the contributors to the Tropicos and Flora Texas Consortium as well as other botanical databases for the

data available online. My deepest gratitude goes to John Kartesz and Christopher Meacham for the software,

Synthesis of the North American Flora.

ln el D «onl D h Imetitiit tT.

438 J Texas 3(

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442 I Lat +k Dat H ID hi PA f Texas 3(1)

BOOK NOTICE

FLORA OF CHINA EDITORIAL COMMITTEE. 2008 (April). Flora of China, Vol. 11. Oxalidaceae through Acer-

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wp] f,

From iik aim wus five p ili l Among these, the A include 99

nut and a the places CAN 204 Dens : ays the nid include the hos

woods; the Cel include E

y ntain some

400 species in 75 genera, including rubber, d is Wide cultivated in a areas ie its latex; the es comprise mainly

i ined eins paige ornamental ae the I flax, ins is ba ida i^ its stem Re and e oik S

T Perr

Hon hich i i i heni ts of the world.”

ors Wu Zhengyi, Peter H. Raven, and Hong Deyuan state, “[This] is the i of a 25-volume work. It includes 35 families,

212 genera, and 1444 species, among which six Een and 615 Pe (42.696) China, and one family, 17 genera and

49 td O 4%) are introduced to China.” Th are oa alphabetical order): Aceraceae, Anacardiaceae, Aquifoliaceae,

, Burseraceae, Buxaceae, Callitrichaceae, Cardi

Celastraceae, Cneoraceae, Coriariaceae, 2 iphyllaceae,

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Surianaceae, Tapisciaceae, Tropaeolaceae, and Zygophyllaceae. Al Preface, Introduction

of Nomenclatural las cea in this Volume of the Flora of d n to E Names, Index to Pinyin Nazas Index to

Scientific Names, Ind he Flora of China and the Fl I Si and aay Mun of ju dio

of hi d th f China Ill i A large map of China and its BS Admini i Jivisi

1 A i

POr ree

J. Bot. Res. Inst. Texas 3(1): 442. 2009

ANNOTATED CHECKLIST OF THE VASCULAR FLORA OF THE

MENARD CREEK CORRIDOR UNIT OF THE BIG THICKET NATIONAL PRESERVE,

POLK, LIBERTY, AND HARDIN COUNTIES, TEXAS

Larry E. Brown Barbara R. MacRoberts and Michael H. MacRoberts

Spring Branch Science Center Herbarium Bog Research, 740 Columbia

8856 Westview Drive Shreveport, Louisiana 71104, U.S.A.

Houston, Texas 77055, U.S.A and Herbarium, Museum of Life Sciences

larry-theplantman@att.net Louisiana State University in Shreveport

Shreveport, Louisiana, 71115, U.S.A.

"ees W. Mee

ry £E, E DA

Rice > Uni iversity

Houston, Texas 77005, U.S.A.

ABSTRACT

An annotated, vouchered checklist is provided of the vascular pl f the 1,537 hectare Menard dh pied Unit, Big Thicket

70(

National Preserve, in Polk, Liberty, and Hardin counties, Texas. D unit are 689 taxa 1096) are exotics.

RESUMEN

Se realiza un catálogo co estigos de los taxa de plantas vasculares de las 1,537 hectáreas de la Menard Creek Corridor

Unit, Big Thicket Tum Preserve, en e condados de Polk, Liberty, y Hardin, Texas. Se documentan para la unidad 689 taxa de los

que 70 (10%) son exóticos.

INTRODUCTION

This is the 7th part of an inventory, based upon available voucher specimens, of the vascular flora of the

Big Thicket National Preserve, southeastern Texas. In six previous papers, we provided annotated check-

lists of the Hickory Creek, Turkey Creek, Big Sandy Creek, Lance Rosier, Beech Creek, and Loblolly units

(MacRoberts et al. 2002; Brown et al. 2005, 2006a, 2008, 2008b). In this paper, we provide an annotated

checklist of the Menard Creek Corridor Unit.

SITE AND METHODS

The Big Thicket is located in the West Gulf Coastal Plain in southeastern Texas (see Diggs et al. 2006 for

literature and description). Peacock (1994) reported 12 units for the Big Thicket Biological Preserve. Since

then the Big Sandy Creek Corridor (1,798 hectares), the Canyonlands (350 hectares), and the Village Creek

Corridor (2,059) have been added. The addition of these 3 units has added 4,207 hectares to the 34,000

reported by Peacock in 1994. However, Deanna Boensch (per. comm.) indicated the total hectares are now

40,088. The units range from 222 to 10,000 hectares and are scattered over 7 Texas counties. The Menard

Creek Corridor Unit (1,537 hectares) follows Menard Creek through parts of Polk, Liberty, and Hardin

counties (Fig. 1). It is located on the Deweyville/Fluviatile formation within a small channel and narrow

bottomlands consisting of Bibb sandy loam and Hatliff fine sandy loam soils that are frequently flooded

(Deshotels 1978; Aronow 1981; Shelby et al. 1992; Griffith 1996). Elevation ranges from about 46 meters at

the Highway 943 bridge over Menard Creek in Polk County to 13 meters at the confluence with the Trinity

River in Liberty County. The unit consists of bottomland forest next to the stream and lower slope-hardwood

pine forest and upper slope pine-oak forest (Harcombe & Marks 1979; Marks & Harcombe 1981). Some

other small distinct habitats that are mostly in Liberty County are Beech-Magnolia slopes where Prenanthes

J. Bot. Res. Inst. Texas 3(1): 443 — 455. 2009

444

laf el Dat

ALABAMA - COUSHATTA \

INDIAN RESERVATION

WOODVILLE fib

-LE SP

enm

& Beech

dg * Creek Unit

HILLISTER@

NECHES BOTTOM

& JACK GORE

BAYGALL UNIT

quel

SILSBEE Y

(527) T one

=I Loblolly \ dome zl 287 Y LOWER NECHES RIVER

J Unit BATSON Ug pee n A j . GORRIDOR UNIT

"d d

ORANGE CO.

LITTLE PINE ISLAND

BAYOU CORRIDOR UNIT I

BEAUMONT UNIT

So, Mi]

HARDIN CO. T

LIBERTY CO. ^| s [SOUR LAKE

BEAUMONT a

- pra

! H-7

a P ¡TE

- JEFF

Fic. 1. Locati Etha M TS

Texas

barbata, Viburnum acerifolium, Cynoglossum virginianum, and Carex abscondita are found; areas of deep sands

where Sida lindheimeri, Setaria corrugata, Mitracarpus hirta, and Silene subciliata grow; alkaline soils with

heliophyte plants such as Verbesina helianthoides, Stenanthium gramineum, Liatris asper, Phyllanthus pudens,

and the 2 Silphium taxa. Another small habitat with a distinct flora is the alluvial soil exposed by receding

water during late summer and fall at the confluence of Menard Creek and the Trinity River. Some of the

plants only or mostly on these drawn-down sites are Fimbristylis vahlii, Eragrostis reptans, Leptochloa panicea

subsp. mucronata, Cyperus squarrosus, Cyperus odoratus, Eragrostis japonica, Marsilea vestita, Panicum virgatum,

and Spermacoce glabra.

Like all units of the Big Thicket National Preserve, the Menard Creek Corridor Unit had not previously

been the subject of a detailed floristic inventory.

Brown et al., Vascular flora of Menard Creek, Big Thicket National Preserve 445

Because our purpose is to produce a list of taxa known to occur on the Menard Creek Corridor Unit, a

vouchered specimen was considered to be the only evidence acceptable for inclusion in the list. In all, 1,137

herbarium specimens form the data for this report. All specimens have been annotated by Larry Brown.

Larry Brown collected 1,020 specimens beginning in the late 1980s and early 1990s and during 16

field days in 2007 and 2008. Geraldine Watson collected 106 specimens over several years in the late 1970s

and early 1980s. The collection total of a few others is 13 and is designated by the appropriate abbreviation

below.

In most cases, nomenclature follows Jones et al. (1997), Kartesz (1999), and Diggs et al. (2006). In cases

of multiple collections, no more than 4 are listed for each taxon.

RESULTS

A J

The following is an annotated list of the Menard Creek Corridor Unit taxa iati ar n below.

ER<r

CP = Charles Peterson. His one Azal 11

DB = Deanna Boensch. Big Thicket fire ecologist. She collected

and those are at T

GW = Geraldine Watson. Her specimens are at TAES.

JL = Joe Liggio. His orchid specimens are at SBSC.

JS = Jeff Schultz. His one orchid collection is at SBSC.

LB = Larry Brown. His specimens with. collection numbers

above 23,000 are at TAES and those with numbers below

23,000 are at SBSC.

MM = ein. "s Michael MacRoberts. Their specimens are

at TAES.

PR - Paul M His collections are at TAES.

SH - Shawn Harper couse : id

SM = larper (

WH = Walter Hesson. His one pus collection is at SBSC.

WRC - Bill Carr. His specimens are at TEX

* — exotic

Sam H

+ = see note at end 2 a

' 1st 311 Jin (ho M ]

Creek Unit when compared sn i» Tother surveyed units

list of the Jack Gore-

which includes a preliminar y Sp

Neches Bottom unit.

A collection number followed by a (P) is from Polk County. A

collection number followed by (H) was collected in Hardin

County. All others are from Liberty County.

Acanthaceae

IDicliptera brachiata (Pursh) Spreng., GW 3293, LB 32845

Hygrophila lacustris (Schlecht. & Cham.) Nees, LB 18779,

26104 (P)

Justicia ovata (Walter) Landau var. lanceolata (Chapm.) Long,

LB 32694

Ruellia caroliniensis (Walter) Steud., LB 32913 (P)

IRuellia pedunculata Torr. ex A. Gray, PR s.n.

IRuellia strepens L., LB 18705, 33097

Aceraceae

lAcer negundo L., GW 3236; LB 32523

Acer rubrum L., LB 33147, 26111 (P)

Agavaceae

Manfreda virginica (L.) Rose, LB 32563, 32879

Alismataceae

Sagittaria latifolia Willd. LBJL 32970, 32982 (P), LB 26113 (P),

5829 (P)

ISagittaria montevidensis Cham. & Schltdl. subsp. calycina

(Engelm.) Bogin, LB 18831

Alliac

Ern e L. var. canadense, LBSH 332

Allium canadense L. var. mobilense (Regel) bs LBSH,

33434 (P)

Nothoscordum bivalve (L.) Britton, LB 32847

Amaranthaceae

[Amaranthus tuberculatus (Moq.) Sauer (including A. rudis

Sauer), LB 18827; LBJL 32985

Froelichia floridana (Nutt.) Moq, LB 28810 (SBSC)

Iresine rhizomatosa Standl., LB 32884

nacardia

Pin VM L., LB 32700

Toxicodendron radicans (L.) Kuntze, LB 32528

Annonaceae

Asimina parviflora (Michx.) Dunal, LB 32574; LBSH (H)

Apia

Charpy tainturieri ha var. tainturieri, LBSH 33212

Cicuta m L., LBSM 3

“elspa | atom eS Sprague ex Britton & P.

Wilson,

Cynosciadium psu DC, 3247, 3255; LBSH 33419

IDaucus pusillus Michx., GW

[Eryngium hookeri Walp., LB de 9

Eryngium prostratum Nutt. ex DC, LB 18616

Hydrocotyle verticillata Thunb., LB 28243 (P

Limnosciadium pumilum (DC.) Mathias & Constance, LBSH

33420

eo. capillaceum (Michx.) i LBSH 33415

Sanicula canadensis L., LB 32505; LBSH 3 c a

a divaricata (Walter) Raf. LBPR 3

éd inermis (Nutt) Mathias & ons LBSH

33451

iod laum (L) A. EA var. aureum Britton, GW 247;

477, 32691; LBSH 3

p "d arvensis (Huds.) Link, En 3248; LBDB 33655

Trepocarpus aethusae Nutt. ex DC., GW 3262; LBSH 33245

Apocynacea

nia tabernaemontana Walter, GW 3

Amso

Trachelospermum difforme (Walter) A. d LB 32714 (P),

32823, 25185

446

Aquifoliaceae

llex ambigua (Michx.) Torr., LB 18846; LBJL nee

llex decidua Walter, LB 25179, 32908 (P); GW 3

llex longipes et ex Trel., GW 3157; EE 255 (P), LB

33128; L 3461 (P)

llex opaca Aiton, 32534, 22564

llex vomitoria Aiton, LB 32636

Araceae

Arisaema dracontium (L) Schott., LB 23880; LBPR 3

Arisaema triphyllum (L.) Schott., LB 18525, 23880a, pum (P);

Araliaceae

Aralia spinosa L., LBSH 33332

*Hedera helix L, LBSH 33399

Arecaceae

Sabal minor (Jacq) Pers., LB 32856

Aristolochiaceae

Aristolochia serpentaria L., LB 33168 (P)

Asclepiadac

Asclepias perennis a LB 18574

Asclepias tuberosa L., LB 32565

ASCIcpids variegata E idi 33351

) Shinners, LB 32519

Asplenium platyneuron (L.) Britton, Sterns € Poggenb., GW

3217; LB 32717

Asteraceae

Acmella repens (Walter) Richard, LB 18860

Ageratina altissima (L) R. King & H. Rob. var. altissima, LB18832,

33149, 33174 (P); LBJL 32957

Ambrosia artemisiifolia L, LB 32854

jn trifida L., LB 32844, 32820, 32874

Bidens aristosa (Michx.) Britton, LB 18553, 32920, 33105; LBJI

32954

Bidens bipinnata L., GW 3286; LB 32930, 32916

Bidens discoidea (Torr. & A. Gray) Britton, Ps 18820b (P)

Bidens frondosa L. LB 18511, 18859,1

Bradburia pilosa (Nutt) Semple, - 2255 18743, 26108 (P)

ICalyptocarpus vialis Less., LBSH 3

Chrysopsis mariana (L.) Elliott, en a LBSM 33540

Cirsium boleta Michx. var. megacanthum (Nutt.) DJ. Keil,

LBSH

mos UE (L) DC., LB 32878

bu onyza bonariensis (L) Cronquist, GW 3251

onyza canadensis (L.) Cronquist, LB 32654

o basalis (A. Dietr.) S.F. Blake, GW 3277; LB 18606,

32463

Coreopsis lanceolata L., LBSH 334

Coreopsis tinctoria Nutt. var. om LB 32938; LBSM 33525

(P)

Croptilon divaricatum (Nutt.) Raf, LB 32575, 32576

Echinacea sanguinea Nutt,, LBSH 33516

Pus dod (L) L, LB 18856, 32825

El topus carolinianus Raeusch, LB 33155 (P)

Ele, Bern udatus A. Gray, GW 3284

Elephantopus tomentosus L., LB 32575, Es

Erechtites hieraciifolia (L.) Raf. ex DC, LB 3274

Erigeron annuus (L) Pers., GW 3276; LBSH T 33443 (P);

LBSM 33544

Erigeron philadelphicus L, LBSH 334

Erigeron strigosus Muhl, ex Willd., C 3267; LBSH 33512,

iss tenuis Torr. & A. Gray, LBPR 33353; LBSH 33392,

33416

Eupatorium capillifolium (Lam.) Small, LB 33129

Eupatorium semiserratum DC., LB 32675, 33153

Eupatorium serotinum Michx., LB 32877

Eurybia hemispherica (Alexander) G.L. Nesom, GW 3473; LB

32812

xe leptocephala (Torr. & A. Gray.) Greene, LB 32753,

cll pulchella Foug. var. pulchella, LB 32688

chaeta coarctata (Willd.) Kerguelen, LBSH 33378

ds pensylvanica (Willd.) Cabrera, "e 33244 (P)

Gamochaeta purpurea (L.) Cabrera, LBSH 33431,33449a (P)

Heleni Rock var. amarum, | A 33260, 32706,

32928; LBSH 33438 (P)

Helianthus hirsutus Raf, GW 3512; LB 32705

Heterotheca subaxillaris (Lam.) Britton & Rusby, LB 32924

ieracium gronovii L, GW 3311; 2504

Iva annua L., LB 32906 (P)

Krigia cespitosa (Raf) K.L. Chambers var. cespitosa, GW 3369;

SH 33240

Krigia cespitosa (Raf) K.L. Chambers var. Gracilis (DC) K.L.

mbers, LBSH 33247 (P

IKrigia dandelion (L.) Nutt. LBSH 33325

Krigia virginica (L.) Willd. LBSH 33316

E A (A. Gray) K.L. Chambers ex K.J. Kim LBSH

Lac > e AEN LB 32656

Lactuca floridana (L.) Gaertn., LB 32846

lLactuca hirsuta Muhl., LBSM 33535

ILiatris asper Michx., GW 3503, 3504, 3505; LB 32562

Mikania scandens (L) Willd, LB 32750

d Misi (Poir) C. Jeffrey, LBSH 33225a, 33231

ovata (Muhl. ex Willd.) W.A. Weber & Á. Lóve, GW

2 LBSH 33321

Packera tampicana (DC.) C. Jeffrey, LB 18532

Pluchea camphorata (L.) DC., LBJL 32977

Pluchea foetida (L) DC., LB 32939

H-Prenanthes barbata (Torr. & A. Gray) W.L. Milstead ex Cron-

quist,

Pyrrhopappus carolinianus (Walter) DC., LB 33103

Pyrrhopappu ips (D. Don) DC., LBSH 33249 (P)

Rudbeckia hirta L, LB 32697

ISilphium ea L. var. asteriscus L, LB 32744

ISilphium asteriscus L. var. simpsonii (Greene) J.S. Clevinger,

LBSM 33538

Solidago altissima L., LB 328

Solidago ludoviciana (A. "e V" LB 5839a

Brown et al., Vascular fl , Big

Solidago rugosa Mill. var. asper (Aiton) Fernald, LB 18854,

18865

Solidago tortifolia Elliott, LBJL 32979

*Soliva sessilis Ruiz & Pavon, LBPR 33338

*Sonchus asper (L.) Hill, LBSH a

*Sonchus oleraceus L., LBSH 3

Smallanthus uvedalia K. ME ex m LB 18707

ISymphyotrichum drummondii (Lindl.) G.L. Nesom, LB 32508;

GW 3238

Symphyotrichum dumosum (L.) G.L. Nesom, LB 32873, 32872

ISymphyotrichum ericoides (L.) G.L. Nesom, LB 33154 (P)

ISymphyotrichum lanceolatum (Willd.) G.L. Nesom., LB 33083,

33130, 33133, 33145

Symphyotrichum lateriflorum (L.) Á. Lóve & D. Lóve, LB 18863,

18862

Symphyotrichum patens (Aiton) G.L. Nesom, LBSM 33541

Symphyotrichum racemosum (Elliott) G.L. Nesom, LB 18826,

19767, 33104; LBJL 32955

+Verbesina helianthoides Michx., LB 32568

Verbesina virginica L, LB 32564

Vernonia missurica Raf., LB 26102 (P), 32640; GW 3280

Vernonia texana (A. Gray) Small, sl 32566

Xanthium strumarium L., LB 3282

*Youngia japonica (L) DC., GW 25

Berberidac

I*Nandina e Thunb., LB 32830

Podophyllum peltatum L., PR s.n.

Betulaceae

Betula nigra L., LB 18603, 33122 (P), 33159 (P), 28249 (P)

Carpinus caroliniana Walter, LB 18600a

Ostrya virginiana K. Koch, LB 32628

Bignoniaceae

Bignonia capreolata L., LB 32517, 18502, 27743 (P)

Campsis radicans (L.) Seem. ex Bureau, LB 32689

Catalpa bignonioides Walter, LBSH 33460 (P)

Blechnaceae

Woodwardia areolata (L.) T. Moore, LB 26118 (P), 32555,

18520, 18744

Woodwardia virginica (L) Sm., LB 19025, 33158 (P)

Boraginace

Cyn des virginianum L., GW 3195, 3239; LBPR 33358

*Heliotropium indicum L., LB 18825, 32842, 17515 (P)

Heliotropium procumbens Mill., LB 18842, 32887

lyosotis macrosperma Engelm., LB 18497

Brassicaceae

*Cardamine hirsuta L., LBSH 3320

Cardamine parviflora L. var. LEM (Britton) O.E. Schulz,

, 3205

Lepidium virginicum L., LB 32647; LBSH 33317

Rorippa sessiliflora (Nutt.) Hitchc., GW 3189, 3204b; LB 18829,

Buddlejaceae

iiti procumbens L., LB 32935

Burman

reo ped L., LB 18778

447

Callitrichaceae

Callitriche heterophylla Pursh, LBSH 33253 (P)

Campanulaceae

Es oe A. DC. LBPR 33352, 33440 (P); LBSM

mos NUR L. var. rea LB 32935

Lobelia puberula M 7; LBJ! ka

Triodanis biflora um E E, di LBSH

ITriodanis lamprosperma McVaugh, LBSH 2 Dem (P)

adi perfoliata (L) Nieuwl., GW 3234, 3235; LBSH 33398

Wahlenbergia marginata (Thunb.) A. DC., LBSH 33448 (P)

Caprifoliaceae

*Lonicera japonica Thunb., LB 32718

Lonicera sempervirens L., LBSH 33236

Sambucus nigra L. subsp. canadensis (L.) Bollii, LB 32516;

LBJL 32962

ISymphoricarpos orbiculatus Moench., LB 18529, 18708

Viburnum acerifolium L., 18516, 24130, 33125

Viburnum dentatum L., LB 32634, 32624

Viburnum nudum L., LB 18499

Viburnum nidum Raf, LB 32812

Caryophyllaceae

*Cerastium glomeratum Thuill., LBSH 33208

Loeflingia squarrosa Nutt., LBPR 33355

Sagina decumbens (Elliott) s a 2 LBPR 33353

*Stellaria media (L.) Vill, LBSH

[Silene antirrhina L., LBSH ie e

I*Silene gallica L., LBSH 33457 (P)

+Silene subciliata Robins., LB 18727, 18162, 32633; LBJL

32953

Celastraceae

Fuonymus americana L., LB 32551

Chenopodiaceae

Dysphanis ambrosioides (L.) Mosyakin & Clemants, LB 33138

Cistaceae

Helianthemum georgianum Chapm., LBSH 33511

Commelinaceae

Commelina difusa Burm. f, LB 3289

Mao ce E var. En (Michx) Fernald, LB

M 33548

man vies L, LB 18597, 32893

Convolvulaceae

Dichondra is Michx., LB 32837

Evolvulus sericeus Sw., G

Ipomoea cordatotriloba Dennst. var. cordatotriloba, LB 32849

Ipomoea lacunosa L., LB 32827

Cornaceae

ICornus drummondii C.A. Mey., GW 3254; LB 32524, 18530

Cornus florida L., LB voee 18595

Nyssa aquatica L., L

Nyssa sylvatica al i 32540, 18518, 18735

Crassulaceae

Penthorum sedoides L., LB 18851, 18780

448

Cucurbitaceae

Cayaponia quinquefolia i n LB 32836

Melothria pendula L., LB 3

Cupress

a iu dno L., LBJL 329

Taxodium distichum (L.) Rich., UR

Cuscutaceae

Cuscuta gronovii Willd. ex Schult., GW 3294, LB 32929

Cyper

E nal (Rottb.) C.B. Clarke, LB 28808

Carex abscondita Mack., LB 27762, 29426; LBSM 33537

Carex albicans Willd. ex Spreng. var australis (L.H. Bailey) Ret-

tig, LBSH 33374

Carex albolutescens Schwein., LB 18507, 18575

Carex atlantica L.H. Bailey subsp. capillacea (L.H. Bailey

Reznicek, LB 27724 (P), em LBSH 33334, 33404 (H)

Carex basiantha Steud., LB 27765, 29429

Carex blanda Dewey., LB 27759, 27752, 25183

Carex brevior (Dewey) Mack. ex Lunell, LB 18505b

Carex caroliniana Schwein., LB 27740(P); LBSH 33376

Carex cephalophora Muhl. ex Willd., LBPR 33359

Carex sia Schwein., L8 32506

Carex complanata Torr. & Hook., LB 18523; LBPR 33344

Can da aan LB 27767, 18534

ae crebriflora Wiegand, LB 27764, 2774; LBSH 33259 (P)

33369

Carex crus-corvi Shuttlew. ex Kunze, LBSM 3351

Carex debilis Michx. vat. debilis, LB 27758, 23878, o

Carex digitalis Schkuhr ex Willd. var. diode a H. Bailey) Naczi

& Bryson, LB 27761, 27760, 277

Carex ER Schkuhr ex Willd., i M LBSH 33425

Carex flaccosperma Dewey, LB 27749, 27738; LBPR 33362

Carex frankii Kunth, LB 15433(P), 28249 (P)

Carex glaucescens Elliott, LB 33161 (P)

ICarex hyalina Boott, LBSM 33523

Carex intumescens Rudge, LB 27722, 18576

Carex joorii L.H. Bailey, LB 18734, n 15441 (P)

Carex leavenworthii Dewey, LB 2

Carex leptalea Wahlenb. subsp. id (Fernald

27728(P), 33163 (P); LBJL 19024

Carex longii Mack., LBSH 33389

Carex lupulina Muhl. ex Willd., LBSH 33463 (P)

Carex oxylepis Torr. & Hook., LB 27754, 18537; LBSH 33426

Carex planispicata Naczi, LB 27757; LBPR 33357

a tineis x Bailey) Small, LBPR 33343; LBSH 33407,

"n5. sees an ex Willd., LB 1

Carex striatula Michx., LB 29427, m. on 18501

P triangularis Boeck, LBPR 33341; LBSH 33381

N; lokey, LB28245

W. Stone, LB

—

(P), 18507, 24134

I+ Carex t a Michx., LBSM 33518

Cyperus acuminatus Torr. x ao ex Torr, LB 32875, 33114 (P)

Cyperus croceus Vahl,

LB 2

Cyperus erythrorhizos t. E uo 32989

*Cyperus esculentus L., LB 331

Cyperus haspan L., LB 18613, VM (P), 32677, 18100a (P)

Cyperus odoratus L., LB 18839b, 33085, 33095

Cyperus retroflexus Buckley var. SUE LB 32573

Cyperus squarrosus L., LB 3309

pn strigosus L, LB 1 e

Cyperus surinamensis Rottb., LB 3

cin thyrsiflorus Jungh., LB n i BSM 33532

679

Eleocharis montana (Kunth) Roem. & Schult., LB 32285

Eleocharis montevidensis Kunth, LBSH 33414

Eleocharis tuberculosa (Michx.) Romer & Schult, LB 18611,

32676, 15434a (P), 26615 (P)

Fimbristylis autumnalis (L) Romer & Schult., LB 32902

*Fimbristylis miliacea (L.) Vahl, LB 32681, 33094

Fimbristylis vahlii (Lam.) Link, LB 18833, 33087,17510 (P),

Fuirena breviseta Coville, LB 26125 (P)

lFuirena pumila (Torr.) Soreng., LB 22566 (P)

Isolepis carinata Torr., LB 18510

Rhynchospora caduca Elliott, LBSM 33531

Rhynchospora corniculata (Lam.) A. Gray, LBSM 335

Rhynchospora globularis (Chapm.) Small var. Fon LB

32707

ri ue ati (L.) Vahl, LB 15429 (P), 26122 (P),

Poncio ee (Michx.) Vahl, LB 32936, 15432 (P);

iaa mixta Britton, LB 28248 (P), 22862 (P); LBJL

32966, 19176

Scirpus cyperinus (L.) Ku p" um ~

Scleria oligantha Michx,

Scleria triglomerata ve Mad : 1349: LB 20592

Cyrillaceae

Cyrilla racemiflora L., LB 26114 (P), 18873, 33142

Dennstaedtiaceae

Pteridium aquilinum (L.) Kuhn var. Pseudocaudatum (Clute)

A. Heller, LB 32556

Dioscoreacea

Dioscorea ee L, GW 2288, 3194, 3282; LB 32831; LBSH

Droseraceae

Drosera brevifolia Pursh, LBSH 32262, 33257 (P)

Dryopteridaceae

Athyrium filix-femina (L) Roth subsp. oo (Michx.)

Hulten, LB 28250 (P), 32554, 18517

Onoclea sensibilis Ly u o (P), 27735 M m 33108 (P)

Polystichum (Michx) Schott, LB 29422, 32553,

18515, 18604

Ebenaceae

Diospyros virginiana L., LB 32860

Ericaceae

Rhododendron canescens (Michx.) Sweet, LB 27747 (P),

18748,18487, 32625

Rhododendron viscosum (L) Torr, LB18512, 207 70 (P), 33151(P);

Brown et al., Vascular fl M i ig

Vaccinium arboreum Marshall, LB 18580, 33173 (P); LBSH

33455 (P)

Vaccinium corymbosum L., LB 32548, 18489, 18586

Euphorbia

Acalypha S A. Gray, GW 3230, 3288

Acalypha rhomboidea Raf., LB 32904 (P), 32946, 33110 (P)

Chamaesyce humistrata (Engelm.) Srnall, bi Due (P)

a maculata a mi LBDB 3

mall, LB ed m

us capitatus MESS ae 11350; LB 32861

Croton glandulosus L. var. o 3291, LB 32941a

roton monanthogynus Michx., L

Euphorbia corollata L, LB 32561

Euphorbia dentata Michx., GW 3432; LB 32693

Euphorbia spathulata Lam., LB 18532b

Phyllanthus caroliniensis Walter, LB 18857a, 33115 (P)

IPhyllanthus pudens Wheeler, LB 18818b (P)

*Phyllanthus urinaria L, WRC 11355; LB 18866

Sebastiania fruticosa (Bartr.) Fernaid, LB 27732 (P), 32547,

18485, 18598

*Triadica sebiferum (L.) Smail, LB 32544

Fabaceae

Aeschynomene indica L., LB 17513 (P)

*Albizia julibrissin Durazz., LB 32699; LBSM 33543

Baptisia nuttalliana Small, LB 18539, 24237

Centrosema virginianum (L.) Benth., LB bd

Cercis canadensis L. var. canadensis, LB

Chamaecrista fasciculata (Michx.) Sere ji 18729, 33136

Chamaecrista nictitans ME

Clitoria mariana L., LB 3

Crotalaria sagittalis L., i s 5; LBSH 33452 (P)

Dalea phleoides (Torr. & A. Gray) Shinners var. phleoides, GW

3509

Enn nu canescens (L) DC., LBDB d.

modium ciliare (Willd) DC., GW 3313b; L

Demodun glabellum (Michx) DC., GW 3290, od ee

328

on paniculatum (L.) DC., LB 18161, 32922; LBDB

33665

IDesmodium pauciflorum de utt) DC, LB 29428

Erythrina herbacea L., LB 32

Galactia volubilis (L.) ron on 3285, 3304, 3312; LB 18244

Gleditsia triacanthos L.

lottidiurn vesicarium ree jum LB 32840

*Indigofera suffruticosa Mill., LB 32570, 28805 (SBSC)

*Kummerowia striata (Thunb.) a LB a 32811a

Lespedeza procumbens Michx., LB 3

Lespedeza repens (L.) Barton, LB id

a virginica (L.) Britton, GW a 1; LBSM 33524; LBDB

a arabica (L) Huds., LBSH 33223

*Medicago lupulina L., GW 3190a, 3219a, LBSH 33210

*Medicago polymorpha L., GW 3219b; LBSH 33233a

Mimosa hystricina (Small) B.L. Turner, LB 32858a

Mimosa latidens (Small) B.L. Turner, LBSH 33514

Mimosa strigillosa Torr. & A. Gray, LB 32712 (P), 33090

IRhynchosia minima (L.) DC, LB 32822

449

Senna marilandica (L.) Link, GW 3227; LB 18828, 32745

Senna obtusifolia (L.) H.S. Irwin & Barneby, LB 18828, 17514 (P)

Sesbania drummondii (Rydb.) Cory, LB 33106

Sesbania herbacea (Mill.) McVaugh, LB 32918

*Sesbania punicea Benth., LB 22567 (P)

Sophora affinis Torr. & A. Gray, LB 32503, 33102

Strophostyles helvula (L.) Elliott, LB 32637

Strophostyles umbellata (Willd.) Britton, LB 32719

Stylosanthes biflora (L.) Britton, Sterns, & Poggenb., GW 3305;

LB 32811

*Trifolium campestre Schreb., LBSH 33227

*Trifolium dubium Sibth., LBSH 33251 (P)

*Trifolium lappaceum L., LB 29420

*Trifolium repens L., GW 324

I*Trifolium resupinatum L., GW 3219c

Vicia ludoviciana Nutt. subsp. feavenworthii e & A. Gray)

setter & C.R. Gunn., LBSH 33224, 33235a; LB 18443

Vicia ludoviciana Nutt. subsp. ludoviciana, LBSH mid

Vicia minutiflora Dietr, GW 3203, 3225, 3300; LBSH 33217

*Vici ed tiva L. subsp. nigra (L) Ehrhart, GW 3226; LBSH

332

*Vicia mad Roth, LBSH 33246 (P)

Wisteria frutescens (L.) Poir, LB 18478, 18733, 32907 (P)

Fagaceae

Castanea pumila (L.) Mill, LBSH 33512

Fagus grandifolia Ehrh., LBJL 19170; LB 18163, 32557, 27730

(P)

Quercus alba L., LB 32538a, 18483

Quercus falcata Michx., LB 32814

Quercus hemisphaerica Bartram ex Willd., LB 32538, 32987,

9, 24128

Quercus lyrata Walter, LB 32710 (P), 33124 (P)

Quercus michauxii Nutt, LB 18541; LBJL 32968

Quercus nigra L., LB 32695

Quercus pagoda Raf., LBDB 33657

Quercus phellos L., LB 32815, 27718 (P); d ~ (P)

B 3275 56

IQuercus texana Buckley, LBPR 33347

Geraniaceae

Geranium carolinianum L., LBSH 33396

Grossulariaceae

Itea virginica L, LB 18488, 33140, 26123 (P); LBJL 32974

Haloragaceae

Proserpinaca palustris L, LB 32275

Hamamelidaceae

Hamamelis virginiana L., LB 32650, 18242, 18608, LBSH

(

Liquidambar styraciflua L., LB 29291, 32650

Hippocastanaceae

Aesculus pavia L. var. pavia, LBSH 33248 (P)

Hydrophyllac

Hydrolea ee i 821

INemophila aphylla (L BLU GW 1346, 3190b; LBSH

33215

450

IPhacelia hirsuta Nutt., LB 18442; LBPR 33339

Hyperic

E p (Grev. & Hook) Torr. & A. Gray, LB 32664

Hypericum galioides Lam., LB 18740

Hypericum hypericoides (L.) Crantz, LB 32629

Hypericum mutilum L, LB 18522, 33151

Triadenum walteri (J.G. Gmel) Gleason, LB 18684, 27725 (P)

Triadenum tubulosum (Walter) Gleason, LB 32621

Hypoxidaceae

Hypoxis curtisii Rose, LB 18578; LBSH 33391, 33458 (P)

Iridace

rains ome (Molina) a LBSH 33408

Iris pseudacorus L., LB 27739 (P)

Sisyrinchium m Mill., LB 18505; LBSH 33465

Sisyrinchium rosulatum E.P. Bicknell, GW 3249, LB 18524;

SH 33387

Juglandac

Carya ie (Michx. f) Nutt., LB 18855, 32708 (P)

Carya cordiformis (Wang.) K. Koch, LB 32892

Carya illinoensis (Wang.) K. Koch, LB 32509; LBDB 33654

ICarya ovata (Mill) K. met GW 3191, 3196; LB 32858

Carya texana Buckley, 2883

Carya tomentosa iin NE LB 18481, 33127

Juncaceae

Juncus acuminatus Michx., LBSH 33428

Juncus anthelatus (Wiegand) R.E. Brooks, LBPR

Juncus coriaceus Mack., LB 27748 (P)

Juncus dichotomus Elliott, LBSM 33521

uncus diffi ipud LB 18609; LBSH 33436 (P)

Juncus effusus L., LB 328

Juncus marginatus inm ism 33413, 33427

Juncus scirpoides Lam., LB 1861

Juncus tenuis Willd., LB 27746 (P), 33172 (P)

Juncus validus Coville, LB 18610, 32937

Lamiaceae

Hedeoma hispida Pursh, LBSH 33430; PR s.n.

(P)

Lycopus rubellus Moench, GW 3309

Lycopus virginicus L., LB 18850, 32933

Monarda fistulosa L., GW 3510

Monarda punctata L. var. lasiodonta A. Gray, LB 18685, 28804,

32932

*Perilla frutescens (L.) ME » 32619

i al vulgaris L, LBPR 333

anthemum ud a & A. Gray, LB 32671

bs lyrata L, LBSH 33213

Scutellaria elliptica a GW 3258a; LB 18593, 18240

Scutellaria integrifolia L, LB 18592; LBSH 33412

IScutellaria ovata Hill, GW 3258b

ics parvula Michx. var. parvula, LBPR 33360; LBSM

suchst ue md e 32620, 26103 (P)

8251 (P), 33165 (P)

Wiehostenma Pulp e L., LB 32933

Lauraceae

Persea borbonia (L.) Spreng., LB 29423, 32558

Sassafras albidum (Nutt.) Nees, LB 32549, 18864

Lemnaceae

*Landolta punctata (G. Mey.) Les & DJ. Crawford, LB 32863

Lentibularia

Utricularia a 9 LB 26117 (P), 25516 (P)

Linaceae

(Plar icd |) Br ittor 1 Val. A EAU (Plar cl |) Fer I ald,

LBSH 33450 (P); PR s.n.

Linum striatum Walter, LB 18612, 24132

Loganiaceae

elsemium sempervirens St.-Hil, LB 32638

ISpigelia texana (Torr. & A. Gray) A. DC, GW 3259; LB 32838, 18704

Lygodiaceae

*Iygodium japonicum (Thunb.) Sw., LB 32537; LBSH 33241

Lythrac

Ammannia coccinea Rottb., LB 32896

*Cuphea carthagenensis Jack.) J.F. Macbr., LB 18618

Rotala ramosior (L.) Koehne, LB 33112

Magnoliaceae

Magnolia grandiflora L., LB 32545

Magnolia virginiana L., LB 26112 (P), 18777; LBJL 32963

Malvaceae

Hibi / L. subsp. lasiocarpos (Cav.) OJ. Blanchard,

33107

Malvastrum coromandelanum (L.) Garcke, LB 32947

Malvaviscus arboreus Cav. var. drummondii (Torr. & A. Gray)

Schery, LB 33145; LBJL 32971

Modiola caroliniana (L) G. Don, LBSH 33324

ISida lindheimeri Engelm. & A. Gray, LB 32569

Sida rhombifolia L., LB 32641

Sida spinosa L., LB 32910 (P)

Marsileaceae

IMarsilea vestita Hook. & Grev. subsp. vestita, GW 3155

Melastomataceae

Rhexia mariana L. var. mariana, LB 32669

Melanthiaceae

"s MUI (Ker-Gawl.) Morong, GW 3500;

LBSH 3

Menispermaceae

Cocculus carolinus (L.) DC., LB 32835

Menyanthaceae

INymphoides aquatica (J.G. Gmel.) Kuntze, LB 25517 (P)

Molluginaceae

*Mollugo verticillata L., LB 32901

Moraceae

Morus rubra L., LB 32525

Myricaceae

Morella caroliniensis (Mill) Small, LB 26124 (P), 19774, 33161

(P); GW 3310

451

Brown et al., Vascular fl

Morella cerifera (L.) Small, LB 282471P)

Nymphaeaceae

Nuphar advena (Aiton) WT. Aiton, GW 3289; LBSH 33251 (P)

Oleaceae

Chionanthus virginicus L, LB 18503, 20593, 33176 (P; LBJL 32976

Forestiera acuminata (Michx.) Poir., LB 32843; LBJL 32981 (P);

Fraxinus americana L., LB 29431, 32502, 18590

Fraxinus pennsylvanica Marshall, LB 32711(P)

*Ligustrum sinense Lour., LB 32652

Onagrac

Ludwigia nen L. LB 26120a (P), 18589, 18728, 32658

Ludwi wigi ia decurrens Walter, LB 18730, 32900

Ludwigia glandulosa Walter var. A LB 32716

E hirtella Raf., LB 1873

Ludwigia leptocarpa (Nutt.) A LB 33121 (P)

Ludwigia microcarpa Michx. LB 26120 (P), 25518 (P)

Ludwigia octovalvis (Jacq.) PH. Raven, LB 18101 (P), 32848

Ludwigia palustris (L.) Elliott, LB 25182, 33116 (P)

Oenothera biennis L., LB 32649; LBSH 33250 (P)

Oenothera laciniata Hill, LB 27717 (P), 20169 (P)

Oenothera speciosa Nutt., LBSH 33406

Ophioglossaceae

Botrychium biternatum A S 18744, 33123 (P)

lOphioglossum vulgatum L., GW

Orchidaceae

IHabenaria repens Nutt., LB 5825 (P); JS 280 (P)

Platanthera clavellata (Michx.) Luer, LB 20591; LBJL s.n.

Platanthera cristata (Michx) Lindl., LB 15431 (P); JL s.n. (P)

Spiranthes cernua (L.) Rich., LB 18861, 33137

ISpiranthes ovalis Lindl., LBJL 32959

Spiranthes praecox (Walter) S. Watson, GW 3257, 3380; LBSH

33367

Tipularia discolor (Pursh) Nutt., LB 18492, 18514, 18731; JL s.n.

Orobanchaceae

Epifagus virginiana (L) W.PC. Barton, LB 18163; LBSH 33329

Osmundaceae

Osmunda cinnamomea L., LB 1

Osmunda regalis L. var. Se T ) A. Gray, LB 18579

Oxalidaceae

Oxalis corniculata L. var. wrightii (A. Gray) B.L. Turner, LB32654a,

33117 (P), 27715 (P); LBSH 33319

ios Re Kunth var. corymbosa (DC.) Lourteig, LBSH

m o Pursh, LBSH Lm

Oxalis violacea L., LB 3274

Passiflorac

Passiflora incarnata L., LB 32657, 32662

Passiflora lutea L., LB 32542, 18738

Phytolaccaceae

Phytolacca americana L., LBSH 33366

Pinaceae

Pinus echinata Mill, LBSM 33546

Pinus taeda L., LB 32

Plantaginaceae

Plantago aristata Michx., LBSM 33536

Plantago major L, LBSH 33367

Plantago rhodosperma -A 33384

Plantago virginica L, LBSH 3

Plata

fias c xw L., LB 32894

Poaceae

Agrostis hyemalis (Walter) Britton, Sterns & Poggenb., LBSH

333

Agrostis perennans (Walter) Tuck. var. perennans, LB 18167,

33143, 33156 (P); LBJL 32956

Andropogon gerardii Vitman, GW 3210

Andropogon glomeratus (Walter) Britton, Sterns & Poggenb,,

LB 33160 (P)

Andropogon gyrans Ashe var. 2A GW 3209

Aristida oligantha Michx., GW

Arundinaria gigantea (Walter) ut LB 32539

— fissifolius (Raddi) Kuhim, LB 32666

xonopus furcatus ace Hitchc., LB 32943

ce minor L, L

*Bromus catharticus on LBSH 33393

*Bromus japonicus Thunb. ex Murray, LBSH 33380; LBSM

33550

IBromus pubescens Muhl. ex Willd., LBSH 33453 (P)

I*Bromus secalinus L., ps 33545

Cenchrus spinifex Cav., LB 32949

Chasmanthium latifolium Em H.O. Yates, LB 26107 (P),

] , 13

Chasmanthium laxum (L) H.O. Yates, GW 2314; LBDB 33663

Chasmanthium sessiliflorum (Poir.) H.O. Yates, LB 32514,

*Cynodon dactylon (L) Person, LB 32715 (P)

Dichanthelium acuminatum (Sw.) Gould & C.A. Clark var.

acuminatum, LBSH 33459 (P)

Dichanthelium acuminatum (Sw.) Gould & C.A. Clark var.

lindheimeri (Nash) Gould & C.A. Clark, LB 33093; LBSM

33528 (P), 33519

Dichanthelium boscii (Poir) Gould & C.A. Clark, LB 29430,

29424, 29425; LBSH 33401 (H)

tatum, LB 27742a (P), 18498

Dichanthelium dichotomum (L.) Gould subsp. dichotomum,

LB 18602, 22562

Dichanthelium dichotomum (L) Gould subsp. microcarpon

iie ex Elliott) Freckmann & Lelong, LB 32818; LBJL

19177; LBSH 33371, 33435a (P)

Di&hentielum laxiflorum (Lam.) Gould, LB 27744 (P), 18504,

LB 32661,18581

subsp. commu-

/'C-L

It) Gould var. oligosanthes,

LB 32824a; LBSM 33549

Dichanthelium oligosanthes (Schult.) Gould var. scribneri-

anum (Nash) Gould, LB 18521; LBJL 32965; LBSH 33314,

33454

Dichanthelium scoparium (Lam.) Gould, LBJL 32965

Dichanthelium tenue (Muhl.) Freckman & Lelong, LBSH

33435 (P)

452

Digitaria ciliaris (Retz) Koeler, LB 32529a, 32645, 33092

*Digitaria ischaemum (Schreb.) Muhl., LB 32747a

IEchinochloa crus-pavonis (Humb., Bonpl. & Kunth) Schult. var.

m egand) Gould, LB 32944

Fchinochloa walteri (Pursh) Heller, LB 32911 (P), 33095

*Eleusine indica (L mc " 32631, 32530

lymus virginicus L, LB 3

*Eragrostis japonica EUM LB 18836, 33089

IFragrostis reptans (Michx.) Nees, LB 18834, 32903, 33119 (P)

Eragrostis spectabilis (Pursh) Steud., LB 32942

Eustachys petraea Re Desv, ut ~ (P)

Hordeum pusillum N H3

Leersia lenticularis Tus LB iun

eun virginica Willd., LB 32690

ochloa panicea (Retz) Ohwi subsp. mucronata (Michx.)

TN ,

ILimnodea arkansana (Nutt.) LH. Dewey, LBSH 33394

*Lolium perenne L., LB 33231a

Melica mutica Walter, LB 18482; LBSH 33218

IMuhlenbergia schreberi J.F. Gmel., LB 33170 (P)

Oplismenus hirtellus (L) P. Beauv. subsp. setarius (Lam.) Mez,

2626

Panicum anceps Michx., LB 18706, 32862

Panicum brachyanthum Steud., LB 32925

Panicum dichotomiflorum Michx., LB 32895

Panicum gymnocarpon Elliott, LB 32834

*Panicum repens L., LB 32642

Panicum rigidulum Bosc. ex Nees, LB 26106 (P), 33139

Panicum verrucosum Muhl., LB 18849

Panicum virgatum L., LB 33086

Paspalum laeve Michx., LB 32674

Paspalum langei (Fourn.) Nash, LB 32529

aspalum notatum Flüggé, LB 32670

Paspalum plicatulum Michx., LBSH 33445 (P), 33513

Paspalum be PJ. Bergius, LB 18824, 17512 (P), 32909 (P);

LBJL 329

Paspalum setaceum Michx. LB 32670a

*Paspalum urvillei Steud., LB o LBSM 33527 (P)

Phalaris angusta Trin., LBSH 3

Phalaris caroliniana Walter, pe T 8

Piptochaetium avenaceum (L.) Parodi, LB 27720 (P), 18486;

LBSH 33318

*Poa annua L., LBSH 33207

Poa autumnalis Muhl. ex Elliott, LB 27742 (P), 18533; LBSH

33331

Saccharum baldwinii Spreng., LB 32921

Saccharum giganteum (Walter) Pers., LB 18858

+*Schedonorus arundinaceus (Schreb.) Dumort., LBSH 33383

Schizachyrium scoparium (Michx.) Nash var. divergens (Hack.)

Gould, GW 3208

a scoparium (Michx.) Nash var. scoparium, LB

32917

n corrugata (Elliott) Schult., 1B 28807

l+Sphenopholis dell ad ex LH. Dewey) Hitchc., LB

18542, 32263; LBSM

+Sphenopholis ae ee Scribn., LB 18526; LBSH

33470

Sporobolus indicus (L.) R. Br., LB 32672

Steinchisma hians (Elliott) Nash, LBSM 33520

Tridens flavus (L.) Hitchc. var. flavus, LB 32665

Tridens strictus (Nutt.) Nash, 2915

Vulpia octoflora (Walter) Rydb., LB 18519; LBSH 33446 (P)

Polemoniacea

Phlox pilosa L., v 33326

Polygalaceae

Polygala polygama L., LBSH 33439 (P)

Polygonacea

Brunnichia ovata a a LB 32507

Polygonum densiflorum Meisn 284

Polygonum hydropiperoides s LB 32828a; LBSH 33421

Polygonum punctatum Elliott, GW 3288a; LB 32692, 32828

Polygonum ee " s 32522

*Rumex crispus L., LBS

Rumex hastatulus "ed p 33315

Polypodiaceae

Pleopeltis polypodioides (L) E.B. Andrews & Windham var.

michauxianum (Weath.) E.B. Andrews & Windham, LB

32510, 32961

Portulacaceae

¡Claytonia virginica L., LBSH 33216

Potamoget

be od gae Raf., LBSH 23353 (P)

Primulac

"Anagalls an arvensis L., LBSH 33232a

Anagallis minima (L.) Krause, LB 32680

Lysimachia iue. Hook., WRC 11354; LB 28170, 18682,

18170

Ranunculaceae

a crispa L., GW 3303; LB 32512; 18862 (P); MM 8224

Clematis reticulata Walter, LB 32749

rose M eue s var. 4 (Chapm.) T. Duncan,

a (P); LBSH 33225, 332

iin parviflorus L., T oe

Ranunculus pusillus Poir, LB 18496

IThalictrum dasycarpum Fisch. 8: Ave-Lall., GW 3231; LBSH

33232

Rhamnaceae

Berchemia scandens (Hill) K. Koch, LB 32914 (P)

Frangula caroliniana (Walter) A. Gray, LB 18513

Rosaceae

Crataegus brachyacantha Sarg. & Engelm., LB 32816; LBSH

Crataegus marshallii Egglston, LB 32833

Crataegus spathulata Michx., GW 3216; LB 32511

Crataegus viridis L., GW 3298; LB 32712 (P), 32857

*Duchesnea indica (Andrews) Focke, LB 33169 (P); LBSH

33255

IGeum canadense Jacq. var. camporum (Rydb.) Fernald &

th., LB 32527, 18543; LBSH 33424

IPrunus angustifolia Marshall, LB 18685

Prunus caroliniana (Mill) Aiton, LB 32926

IPrunus mexicana S. Watson, LBPR 33349

£A IF L Dia Thiel nt Alag’ I

Brown et al., Vascular fl

Prunus serotina Ehrhart, LB 32651

Prunus umbellata Elliott, GW 3328, 3334

Rubus argutus Link, LB 32622; LBSH 33234

Rubia

LE eae p LB 18490

Diodia teres Walte

*Galium aparine E GW m 9

Galium obtusum Bigelow var. obtusum, LBSH 33410

Galium pilosum Aiton, LB 3257; LBSH 33469; LBSM 33533

Galium tinctorium (L.) Scop., GW 3202,3220, 3286; LBSM

33526 (P)

Houstonia pusilla Schoepf., LBSH 33206

Mitchella repens L., LB 32546, 18493, 29421

I+*Mitracarpus hirtus DC., LB 18847, 32644

Oldenlandia boscii (DC.) Chapm., LB 18168, 22570

Oldenlandia uniflora L., LB 18171, 22569

*Sherardia arvensis L., LBSH 33228

ISpermacoce glabra Michx., LB 17511 (P), 33096

Rutaceae

Ptelea trifoliata L var. iod Torr. & A. Gray, LB 22571, 18479,

32931; LBSH 3337

Zanthoxylum calva- mu L., LB 28254 (P)

Salicaceae

Populus delt p. deltoides, LB 32518;

af E

GW 3232

Salix nigra Marshall, LBDB 33656

apindaceae

ICardiospermum halicacabum L., LB 32839, 32905 (P)

Sapotaceae

Sideroxylon lanuginosum Michx., GW 3268; LBPR 33346

Saururaceae

Saururus cernuus L., LB 18577; LBSH 33437 (P)

Scrophulariaceae

Agalinis d (Elliott) Raf, LB 32871

Agalinis purpurea (L.) Penn., LB 33135

Aureolaria a (Benth.) Penn, LB 26109 (P)

Gratiola brevifolia Raf., LB 18619

Gratiola neglecta Torr, LB 18538

Gratiola pilosa Michx., LB 18617

Gratiola virginiana L., LB 27726 (P), 18500; i 33402 (H)

Leucospora multifida (Michx.) Nutt., LB 18

Lindernia aunta (L) id bá 18830, LB 331 i: (P)

I*M is, LBSH 33227a

Mecardonia acuminata (Walter) Small, ee 11351; LB 33148

Mecardonia procumbens (Mill) Smali, GW 3233

Micranthemum umbrosum (J.F. Gmel) de Blake, LB 18852a

33166 (P)

Mimulus alatus Aiton, LB 18737, 5838 (P); LBJL 32973

Nuttallanthus texana (Scheele) mu LBSH 33320

IPedicularis canadensis L., LBSH 33322

H-Penstemon tenuis Small, LB 18535; LBPR 33348; LBSH

33422

Scoparia dulcis L, LB 18732, 32941

*Veronica arvensis L., LBSH 33203, 33239

453

Veronica peregrina L. subsp. xalapensis (Kunth) Pennell, GW

3314, 3366

Veronica persica Poir., LBSH 33229

Selaginellaceae

Selaginella apoda (L.) Spreng., LB 27738 (P), 20168 (P)

Smilacaceae

Smilax bona-nox L., LB 32627

Smilax glauca Walter, LB 18584, 32753

Smilax laurifolia L, LB 27736, ced 18746

Smilax pumila Walter, LB 18599; LBJL 32967

Smilax rotundifolia L, LB 33175 (P)

Smilax smallii Morong, LB 32639

Solanaceae

IPhysalis angulata L., LB 18841, 33120 (P)

Physalis heterophylla Nees, LB 25168, 32886; LBSH 33238;

LBPR 33345

Solanum ,LB 33144

Sparganiaceae

Sparganium americanum Nutt., LB 15436 (P), 33171 (P),

25516a

Sphenocleaceae

*Sphenoclea zeylandica Gaertn., LB 18822

Styracaceae

Styrax americana Lam., LBJL 32980 (P), 33164 (P), 33141; LB

18165

Symploc

DOM. tinctoria (L) L. Her, LB 32535, 18508

Thelypteridaceae

Thelypteris kunthii (Desv.) Morton, LB 32557a, 28244 (P)

Tiliaceae

Tilia americana (L) var. caroliniana (Mill) Castig, LB 32531

Typhaceae

Typha latifolia L., LBSH 33441 (P)

Ulmaceae

Celtis laevigata Willd., LB 32891

Planera aquatica L., LB 18819 (P), LB 32748, 27745 (P)

Ulmus alata Michx., LB 32655

Ulmus americana L., LB 32912 (P)

tUlmus crassifolia Nutt., LB 32709 (P)

Urtic

iras ia (L.) Sw., LB 26105, 32889

lUrtica chamaedryoides Pursh, LB 25180; GW 3156; LBSH

33243

Valerianaceae

Valerianella radiata (L.) Dufr., LB 18475; LBSH 33237

Verbenaceae

Callicarpa americana L., LB 32623

*Glandularia pulchella (Sweet) Tronc., LBSH 33234a

erbena brasiliensis Vell, LB 32701

Verbena halei Small, LBSH 33385

*Verbena rigida (L.) Spreng., LBSH 33442 (P)

454 J t tani i Texas 3(

Violaceae Parthenocissus quinquefolia (L) Planch., LB 32521

Viola lanceolata L., LB 27721(P) Vitis aestivalis Michx., LB 32660

Viola palmata L., GW 3192, 3306, 3307; LBSH 33214 Vitis cinerea (Engelm.) Engelm. ex Millardet var. cinerea, GW

Viola primulifolia L, LB 27731 (P), 27716, 18495, 33167 (P) 3269

a roria Willd. oe (Greene) McKinney, GW !Vitis mustangensis Buckley, LBDB 33654a

8, 3302; LBSH 33226 Vitis palmata Vahl, LB 18820a (P); GW 3244

TER sororia Willd. var. sororia, LBDB 33659 Vitis rotundifolia Michx., LB 32635

Vitaceae Xyridaceae

Ampelopsis arborea (L.) Koehne, LB 32658 Xyris jupicai L. Rich., LB 26121, 26116 (P), 32668

lAmpelopsis cordata Michx., LB 32515, 32658; LBPR 33342;

NOTES

Eupatorium semiserratum. See Brown et al. (2007) regarding the status of this species in Texas.

Prenanthes barbata. Poole et al. (2007) mapped it in 12 Texas counties. Thisis the first record for Liberty County.

Verbesina helianthoides. See Brown et al. (2007) regarding this species in Liberty County.

Silene subciliata. Poole et al. (2007) mapped it in 8 Texas counties. All known Liberty County records

are from the Menard Creek Unit.

Carex typhina. Both Stanley Jones in Diggs et al. (2006) and Turner et al. 2003) mapped it in 7 north-

east Texas counties. However, in Diggs et al. (loc. cit.) there is one record farther south in Jasper County.

Our collection is a new county record farther south then those previously recorded. There were numerous

individuals on sandy alluvial soil along Menard Creek a short distance upstream from the confluence with

the Trinity River.

Schedonorus arundinaceus. We are following Darbyshire (2007). Other synony

Schreb. and Lolium arundinaceum (Schreb.) Darbysh.

Sphenopholis longiflora. Both this grass and S. obtusata are present in this unit. In many publications,

S. longiflora is treated as a variety of S. obtusata. At least in this unit, species rank is justified for various

reasons including no plants were found with int diate morphology; habitat difference, with S. longiflora

in bottomland habitats and S. obtusata in upland habitats; and anthesis period, with S. longiflora in flower in

March and shedding mature spikelets when S. obtusata is coming into flower in April.

Penstemon tenuis. Some authors merge this species with P. laxiflorus. All of our plants had glabrous stems

or sparse hairs in thin lines, the flowers were pink, and were collected in bottomland hardwoods; thus are

referable to this species. We found no penstemons that we could call P. laxiflorus. The differences between

the 2 species are readily discernable when both are observed in the field.

Mitracarpus hirtus. See Brown et al. (2007) regarding the presence of this species in east Texas.

DISCUSSION

There are 127 families and 396 genera for the 689 taxa (619 native) on the Menard Creek Corridor Unit list.

Eight families: Asteraceae (88 taxa), Poaceae (82 taxa), Cyperaceae (67 taxa), Fabaceae (51 taxa), Scrophu-

lariaceae (21 taxa), Lamiaceae (18 taxa), Euphorbiaceae (16 taxa), and Fagaceae (13 taxa) account for 51% of

all taxa collected. The genera with 10 or more taxa are Carex (35 taxa), Cyperus (12 taxa), Quercus (11 taxa),

Dichanthelium (11 taxa), and Juncus (10 taxa).

While it is the case that our list is incomplete, as are all floras, we believe that we have collected 90+ %

of the taxa. We estimate that the Menard Creek Corridor Unit has about 625 to 650 native species/taxa.

Of the 7 Units inventoried so far, this unit has 86 taxa that have not been found in the other surveyed

units including a preliminary checklist of the Jack Gore Baygall/Neches Bottom Unit. Thus 12% of the

Menard Creek Unit taxa have not been found in the other inventoried units.

Brown et al., Vascular fl fM | Creek, Big Thicket National P 455

ACKNOWLEDGMENTS

This study was supported in part by an All Taxa Biological Inventory grant (ATBI) from the Big Thicket As-

sociation of Saratoga, Texas. We thank Ray C. Telfair II and Barney Lipscomb who carefully read the paper

and pointed out a number of errors that hopefully we were able to correct.

REFERENCES

Aronow, S. 1981. Notes on the geologic units: Big Thicket National Preserve. Big Thicket National Preserve,

Beaumont, Texas.

Brown, L.E., B.R. MacRoserTs, M.H. MacRoserts, PA. 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, Texas. Sida 21:1807-1827.

Brown, L.E., B.R. MacRoserTs, M.H. MacRosenrs, PA. HARCoMBE, W.W. Pruess, l.S. ELsik, AND S.D. Jones. 2006a. Annotated

checklist of the vascular flora of the Big Sandy Creek Unit of the Big Thicket National Preserve, Polk County,

Texas. Sida 22:705-723.

Brown, L.E., B.R. MacRogerTs, M.H. MacRoserts, PA. Harcomee, W.W. Pruess, I.S. ELsik, AND S.D. Jones. 2006b. Annotated

checklist of the vascular flora of the Lance Rosier Unit of the Big Thicket National Preserve, Hardin County,

Texas. Sida 22:1175-1189.

Brown, L.E., B.R. MacRoserTs, M.H. MacRoserts, PA. HARCOMBE, W.W. Pruess, l.S. ELsIK, AND S.B. WaLker 2008. Annotated

checklist of the vascular flora of the Beech Creek Unit of the Big Thicket National Preserve, Tyler County, Texas.

J, Bot. Res. Inst. Texas 2:65 1-660.

Brown, L.E., B.R. MacRoserts, M.H. MacRoserts, W.W. Pruess, I.S. ELsik, AND S.B. WALKER. 2008b. Annotated Checklist of

the Vascular Flora of the Loblolly Unit of the Big Thicket National Preserve, Liberty County, Texas. In press, J.

Bot Res. Inst. Texas 2:1481—1489

BROWN, L.E., E.L. Kern, DJ. Rosen, AND J. Liccio. 2007. Notes on the flora of Texas with additional and other significant

records. Ill. J. Bot. Res. Inst. Texas 1:1255- 1264.

Danavsuine, S.J. 2007. Schedonorus. In: The flora of North America. Vol. 24. Oxford University Press, New York.

DeshoteLs, J.D. 1978. Soil survey for the Big Thicket National Preserve, Texas. USDA Soil Conservation Service and

Texas Agricultural Experiment Station, College Station.

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.

GRIFFITH, K.L. 1996. Soil survey of Liberty County, Texas. USDA and Texas Agricultural Experiment Station, College

Station.

Harcome, PA. AND P.L. Marks. 1979. Forest vegetation of the Big Thicket National Preserve. Report to National Park

Service, Santa Fe, New Mexico.

Jones, S.D., J.K. WipFF, AND P.M. MONTGOMERY. 1997. Vascular a of Texas. Univ. of Texas Press, Austin.

KARTESZ, J.T. 1999. A synonymized checklist and atlas with biological attributes for the vascular flora of the United

States, Canada, and Greenland. 1st ed. In: J.T. Kartesz and CA Meacham. Synthesis of North American flora..

Version 1.0. North Carolina Botanical Garden, Chapel Hill.

MacRoserts, B.R., M.H. MACROBERTS, AND L.E. Brown. 2002. Annotated checklist of the vascular flora of the Hickory

Creek Unit of the Big Thicket National Preserve, Tyler County, Texas. Sida 20:781-795.

Marks, P.L. AND PA. HARCOM&E. 1981. Forest vegetation of the Big Thicket, southeast Texas. Ecol. Monogr. 51:287-305.

Peacock, H.H. 1994. Nature lover's guide to the Big Thicket. Texas A&M University Press, College Station.

Poo e, J.M., W.R. Carr, D.M. Price, AND J.R. SINGHURST. 2007, Rare plants of Texas. Texas A&M University Press, College

Station.

SHELBY, C.A., M.K. McGowen, S. ARONow, W.L. FisHER, L.F. Brown, J.H. MicGowen, C.G. Groat, AND V.E. Barnes. 1992. Geologic

atlas of Texas: Beaumont Sheet. Bureau of Economic Geology, University of Texas, Austin.

456 I Ll a€ ol Dat «oat D h lInrti tt, f Texas 3(1)

BOOK NOTICE

FLORA OF CHINA EDITORIAL COMMITTEE. 2008 (December). Flora of China, Vol. 7. Menispermaceae through

Capparaceae. (ISBN Vol. 7: 978-1-930723-81-8, hbk.). Science Press (Beijing) and Missouri Botanical

Garden Press (St. Louis), PO. Box 299, St. Louis, Missouri 63166-0299, U.S.A. (Orders: www.mbgpress.

org, mbgpress@mobot.org, 1-877-271-1930). $125.00, 499 pp., 8 3/4" x 11 1/4".

From the publishers.—“Eleven plant families are described in this volume. Among these, the Capparaceae include the caper (Capparis

` . y 1 cy LAE y > A f

spinosa); theI include tł f pl 1 } js C i i verum),

and the cultivated bay laurel (L bilis); the Magnoli include the Asi lip tree (Liriodend hi ), found only in China

and Vietnam, and some 1 j gnolias tl leit} 12 ser g llin Magnolia; the Myristi includ

th f g and (Myristica fragrans) 1 tk include the horticul lly imp poppies, Meconopsis and

Papaver, as well as the very di Corydalis, which has 357 species in Chin

Editors Wu Zhengyi, Peter " Raven, and Hong Deyuan state, "[This] is the 1 of the 24 text volumes in the series. It includes

11 families, 95 genera, and 1226 species, sees four gue and 778 P o are anions to China, and eight genera and

16 species (1.3%) are introduced to China.” The f: Iphab canthaceae dd Cleomaceae,

Hernandiaceae, Illiciaceae, Lauraceae Magnoliaceae, M MEDEE Papaveraceae ] Also included

in the text: Preface, Introduction, lla List ol Nomenclatural Novelties Published in this Volume of the Flora of China,

Index to Chinese Names, Index to Pinyin Names, Index to DA oan He to Habes in the Flora of China and the Flora Rei-

publicae Populari: Sinicae, and Published Vol A large map of China and its 35

th tl 1 aq f. t IE- J j «t T 2 1 1 fj!

r * C r £

Administrative D

J. Bot. Res. Inst. Texas 3(1): 456. 2009

CYPERUS PILOSUS (CYPERACEAE) NEW TO THE FLORA OF TEXAS

Richard Carter Charles M. Allen

Herbarium (VSC) Colorado State University

Department of Biology Fort Pol ji

Valdosta State University 1645 23rd Street

Valdosta, Georgia 31698-0015, U.S.A. Fort Polk, Louisiana 71459, U.S.A.

rcarter@valdosta.edu

Patricia and David P. Lewis

262 CR 3062

Newton, Texas 75966-7003, U.S.A.

ABSTRACT

Cyperus pilosus is reported new to Texas and a dic! key is p 1 to distinguish it f g in th

RESUMEN

Cyperus pilosus se cita nuevo para Texas y se [ lave dicotómica para dif: iarlo d gé l estad

Cyperus pilosus Vahl is widely distributed in tropical or subtropical areas of the Old World (Kúkenthal

1935-1936). It is a common weed of rice in Asia (McGivney 1938; Bryson & Carter 2008) and is known

from Hawaii based upon a single historical collection made in 1916 (Wagner et al. 1999). Cyperus pilosus was

first reported in the continental United States from Louisiana, where it was apparently introduced from Asia

as a contaminant of rice seed (O'Neill 1938). Subsequently, it has been reported from Florida (Burkhalter

1985), Mississippi (Bryson & Carter 1992), South Carolina (Tucker et al. 2002), and Georgia (Carter et al.

2009). Cyperus pilosus inhabits hydric soils of wet ditches, rice fields, edges of ponds, and wetlands. It has

not previously been reported from Texas (Jones et al. 1997; Tucker et al. 2002; Diggs et al. 2006), and the

nearest mapped locality is in nearby Calcasieu Parish, Louisiana (Thomas & Allen 1993).

The specific epithet pilosus is descriptive of the hispidulous rachis. The species is illustrated in Figure

1. If inserted in the key to Cyperus immediately after the first lead of couplet 8 on page 1133 in Diggs et al.

(2006), the E Ee will enable the identification of C. M RM in eastern Texas.

8. Plant w | ted sl harply triquetrous, C. pilosus

8. Plant cespitose, “without elongated rhizomes; culm bs iS to A. P not easily compressed

ntinue with couplet 9

Voucher specimens: TEXAS. Newton Co.: Caney Creek Park, S of Hwy. US 190 in Newton, 30° 50' 842"N 93° 45' 802"W, 30 Oct 2008,

Allen and Lewis 21096 (VSC), Allen and Lewis 21098 (BRIT, FTPK, VSC).

REFERENCES

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.

Bryson, C.T. AND R. CARTER 2008. The significance of Cy Is. In: R.F.C. Naczi and B.A. Ford, eds. Sedges:

uses, diversity, and systematics of the Cyperaceae. Monogr. Syst. Bot. Missouri Bot. Gard. 108:15-101.

BURKHALTER, J.R. 1985. Aletris farinosa, Cyperus difformis, and Cyperus pilosus new for Florida. Sida 11:247-248.

CARTER, R., W.W. Baker, AND M.W. Morris. 2009 Contributions to the flora of Georgia, U.S.A. Vulpia. In press.

Dices, G.M., B.L. Lirscoms, and RJ. O'KeNNoN. 2006. Illustrated flora of east Texas. Sida, Bot. Misc. 26:1-1594.

Jones, S.D., J.K. Wiprr, AND PM. Montcomery. 1997. Vascular plants of Texas. Univ. of Texas Press, Austin.

KUKENTHAL, G. 1935-1936. Cyperaceae-Scirpoideae-Cypereae. In A. Engler, ed., Pflanzenreich IV. 20 (Heft)

101:1-671.

J. Bot. Res. Inst. Texas 3(1): 457 — 459. 2009

Journal of the Botanical Research Institute of Texas 3(1)

Fig. 1. Cyperus pilosus. 1a. inflorescence habit. 1b. Detail of triquetrous culm showing antrorse prickle hairs on wing-edges. 1c. Inflorescence ray with

spikelets. 1d. Hispidulous rachis. 1e. Rhizomes (arrows) and connected bases of culms. Photographs from A. Carter 16081, Bacon County, Georgia.

Carter et al., Cyperus pilosus in Texas 459

McGivney, M.V. 1938. A revision of the subgenus Eucyperus found in the United States. Cathol. Univ. Amer. Biol.

Ser. 26:1—74.

O'NeiL, H.T. 1938. Cyperus pilosus Vahl in the United States. Rhodora 40:74.

THoMas, R.D. AND C.M. ALLEN. 1993. Atlas of the vascular flora of Louisiana, vol.1: Ferns & fern allies, conifers &

monocotyledons. Louisiana Department of Wildlife & Fisheries, Baton Rouge.

Tucker, G.C., B.G. Marcks, AND J.R. CARTER. 2002. Cyperus. In: PW. Ball, K. Gandhi, RW. Kiger, D. Murray, J.L. Zarucchi,

AA. Reznicek, and J.L. Strother, Flora of North America, vol. 23. Oxford Univ. Press. New York. Pp. 141-191.

WaGner, W.L., D.R. HEnssT, AND S.H. SoHMER. 1999. Manual of the flowering plants of Hawai'i (rev. ed.). 2 vols. Bishop

Mus. Spec. Pub. 97. Univ. Hawaii and Bishop Museum Press. Honolulu.

I | af ab D sal D hi PO £T.

460 Texas 3(

BOOK NOTICE

FLORA OF CHINA EDITORIAL COMMITTEE. 2008. Flora of China Illustrations, Vol. 12. Hippocastanaceae through

Theaceae. (ISBN Vol. 12 illustrations: 978-1-930723-79-5, hbk.). Science Press (Beijing) and Missouri

Botanical Garden Press (St. Louis), PO. Box 299, St. Louis, Missouri 63166-0299, U.S.A. (Orders: www.

mbgpress.org, mbgpress@mobot.org, 1-877-271-1930). $125.00, 475 pp., 8 3/4" x 11 1/4".

3 cp BE | A | 1

From the publishers.— “Volume 12 of the illustrations series is devoted to 18 plant familie

dd Sio PR source of jo kiwi E pisc with is colo Impatiens ee Malvaceae, s bin ndn co

Theaceae

cae aaa Vitaceae, aotable for Vitis acies the Ga

Editors Wu Zhengyi, Peter H. Raven, and his publication is the 13'* in a series of 24 volumes of Flora of China

Illustrations. It iN MR ide an China TEN Volume 12, ne was IS pusliste ed in 2007 and treated 125 genera with 1275 species in

18 families. This D I g l of 937 species, 14 subspecies, and 202 varieties in 123

genera in 18 families." Also included in tl Preface, Acl ledg Ud to oe Names, Index to Pinyin Names, Index to

Scientific Names, Index to dual in Fus Flora of China and the Flora Rei Sinicae, and Published er Ln e He

eee and ai of Cl A large map of China and its 35 5 Administrative Divisi p

ree

J. Bot. Res. Inst. Texas 3(1): 460. 2009

CRYPTOSTEGIA GRANDIFLORA (APOCYNACEAE: ASCLEPIADOIDEAB),

A NEW NON-NATIVE WEED FOR TEXAS

Thomas F. Patterson Guy L. Nesom

Dept. of Biology, South Texas College 2925 Hartwood Drive

1 3167 Fort Worth, Texas 76109, U.S.A.

Rio Grande City, Texas 78582, U.S.A. www.guynesom.com

tfpatt@southtexascollege.edu

ABSTRACT

Cryptostegia grandiflora is naturalized in Starr County, Texas, where it has been observed at two localities along the Rio Grande. It is

1 A JA po ate 4 1 :

RESUMEN

difl A lizada en el condado de Starr, Texas, donde ha sido observada en dos localidades a lo largo del Rio

J h E, A yh 1

L oOo o

Grande. Es abundant bos] y

Cryptostegia grandiflora (Roxb. ex R. Br.) R. Br. (Apocynaceae, Asclepiadoideae; Palay rubbervine) has

been discovered growi tside of cultivation in several places in Starr Co., Texas. It has not been previously

reported or unequivocally confirmed as a naturalized member of the Texas flora gd 2009).

TEXAS. Starr Co.: R rig g g le below Ot Deck of R Bluffs World Birding Center, completely

lensel (El bano) up to 10 m high, fruiting, 17 Jan 2009, T.F. Patterson 2009-1 with E.C. Patterson (BRIT,

TEX) Rio Grande Ci p: forest along the Rio pd pound Ft. Bis oe mr 17 Jen nid T.F. Patterson nd 2 with E.C.

Patterson (BRIT, TE Tex

outside of cultivation: Cameron Co.: il sandy soil, alt. 30 ft, scandent shrub, 20 Oct 1923, Runyon 566 (TEX).

At the Roma Bluffs, Cryptostegia grandiflora densely covers 13 trees of popinac (Leucaena leucocephala), Ber-

landier ash (Fraxinus berlandieriana), and ebano (Ebenopsis ebano). Over an area of ca. 1 acre, growing from

the banks of the Rio Grande up the slope to the backyards of houses lining the bluff. This Roma neighbor-

hood may have been the source of the escaped Palay rubbervine.

The Ft. Ringgold site occurs ca. 22 miles downriver from the Roma Bluffs in a riparian forest far away

from any houses in Rio Grande City. Cryptostegia grandiflora densely covers over 60 trees of salt cedar (Tama-

rix aphylla) and sugar hackberry (Celtis laevigata), spread over ca. 3 acres. Most of the rubbervine drapes

salt cedars that occur on the first terrace above the river. The species also covers hackberries mostly in the

riparian forest at the level of the river inland from the Phragmites-covered bank. Many of the most densely

covered hackberries have died. Both areas are periodically flooded by the Rio Grande and it may have been

during these floods that logs with attached C. grandiflora and the copious wind- and water-dispersed seeds

transported the rubbervine downriver to Ft. Ringgold.

Cryptostegia grandiflora is native to Madagascar, where it occurs in riverine and seasonally flooded forests

in the dry, southwest part of the country, and it is widely adventive in other parts of the world. It is reported

to be invasive in the United States in the Florida Keys, Monroe Co. (Wunderlin & Hansen 2003, 2008)

and it also is naturalized in Mexico, is: West Indies, Australia, and Pacific Islands (PIER 2007). Standley

(1924) reported that C. grandiflora ghly naturalized in Sinaloa, and a *distributed query" to major

herbaria with Mexican specimens (REMIB 2008) shows subsequent collections from Baja California Sur,

Guerrero, Jalisco, Michoacan, Nayarit, Puebla, Sinaloa, Sonora, Tamaulipas, and Yucatán.

The genus Cryptostegia includes only two species, C. grandiflora and C. madagascariensis Bojer ex Decne.

ane & is dd Sidi both endemic to Madagascar and both widely cultivated as ornamentals. Both

land invasive in Florida and in Australia. In Hawaii, “Both species are usually identified

J. Bot. Res. Inst. Texas 3(1): 461 — 463. 2009

462 J lof the Botanical h Instit Texas 3(1)

as C. grandiflora R. Br., but nearly all Hawaiian cultivated plants proved on careful inspection to be C. mada-

gascariensis” (Staples et al. 2006). Apparently all or most reports for the Pacific area have been re-identified

as C. madagascariensis in the Pacific Island Ecosystems at Risk account (PIER 2007) and previous reports of

naturalized C. grandiflora should be evaluated in light of the possibility of confusion with its closely similar

congener. Descriptions of both species are given on the pages in the PIER online database, and color photos

of both also are available (PIER 2007; HEAR 2008). The following key, based on the work of Marohasy and

Forster (1991), can be used to distinguish the species:

1. Stems with prominent, few lenticels; leaf blades with 14-16 pairs of secondary veins; corollas 3-4 cm long;

corona TIGER entire; folie 5. m 9.9 cm long EOD mue

]. f blades with 11-13 pairs of secondary veins;

corona filaments bilobed; follicles 10- 15.4 cm long Ciyptostegia grandiflora

Diagnostic features of Cryptostegia grandiflora: Climbing, lactiferous subshrubs; leaves opposite, simple,

thick, evergreen, glabrous, blades broadly oblong to elliptic, 4-10 cm x 3—5 cm, apices abruptly acuminate;

flowers 6-12 in terminal cymes, corollas reddish purple to pale pink, tubes campanulate, 2-4.5 cm long,

lobes spreading, 2-4 cm long; follicles fusiform-ovoid, 10—15.4 cm long. “Large C. grandiflora can produce

more than 8000 wind dispersed seeds in a single reproductive episode and can set seed at least twice per

year. More than 9096 of seeds will germinate within 10 days of moisture becoming available. Few, if any,

seeds survive for more than 12 months in the soil" (Grice 1996).

In Australia, EUR Eon is *found in Queensland in the dry tropical areas often fringing

streams and river systems i j t hills and pastures. It smothers vegetation replacing native spe-

cies, particularly in areas degraded! by stole [It] Hinders pastoralism and reported to be toxic to livestock"

' (Smith 2002, p. 47). A remarkable photo shows a vast area in north Queensland, including ground, shrubs,

and trees, completely covered by the viny growth (Invasive Species Council 2008), and accompanying notes

describe the species as *arguably Australia's worst weed." A small area at the Ft. Ringgold riparian site appears

similar to the Queensland photo with C. grandiflora completely covering trees, shrubs, and ground. The viny

undergrowth is impenetrable. The spread of Palay rubbervine should be of concern to both ranchers and

the Lower Rio Grande Valley National Wildlife Refuge with refuge tracts from Falcon Dam to the mouth of

the Rio Grande. Pastures and refuge tracts along the Rio Grande should be monitored for the first arrival of

eee so it can be eradicated pee it becomes established.

ly are no rest the sale of either species of Cryptostegia in the United States, and

both are included on recent versions of the “Nursery eligible plant list and plant price schedule" for Texas,

Oklahoma, and New Mexico (FCIC 2008). Cryptostegia grandiflora is offered for sale by nurseries in Florida

and Texas and was offered for *swap" on a 2006 Texas gardening forum on the internet.

ACKNOWLEDGMENTS

We are grateful to David Lemke and Barney Lipscomb for their helpful review comments and to Elena

Castelo Patterson for her help in the field.

REFERENCES

FCIC. 2008. Nursery Crop Insurance Program, Federal Crop Insurance Corporation. <http://www.ag-risk.org/

FCICDOCU/NURSERY/NUR2008/nursery.htm>

Grice, A.C. 1996. Seed production, disp land lifl | Zizipl iti

two invasive shrubs in tropical woodlands B northern dla Australian J. Ecol. 21:324- 331.

HEAR. 2008. Hawaiian ecosystems at risk project. Invasive species inf tion for Hawaii and the Pacific. <http://

www.hear.org/>

INVASIVE SPECIES COUNCIL. 2008. A rogues gallery (images of serious invasive pests and weeds of Australia). <http://

. “ / I / H AA htra!

/ yp14.htmi

Datt, JA r. tart H Hal * T, 463

MaroHasy, J. AND Pl. Forster. 1991. A taxonomic revision of Cryptostegia R. Br. (Asclepiadaceae: Periplocoideae).

Austral. Syst. Bot. 4:571—577.

NESOM, G.L. 2009. Literature reviews of the Texas flora: non-native species recently discovered to be naturalized

in the state; Non-native species in the state, complete list. <http://guynesom.com/Texasplantsweb.htm>

PIER. 200 Epoca grandiflora; Cryptostegia madagascariensis. Pacific Island ecosystems at risk. <http://

/pier/sp eee Du

REMIB. 2008 Med Biodi rl iodiversity Inf ion Network. <http//www.

conabiaaobimremib. Ina es doctos remilb: ing Premi

SmitH, N.M. 2002. Weeds of the wet/dry tropics of Australia - a field guide. Environment Centre NT, Inc., Darwin,

Northern Territory, Australia.

STANDLEY, PS. 1924. Asclepiadaceae. Trees and shrubs of Mexico. Contr. U.S. Natl. Herb. 23(4):1166-1194.

SrapLEs, G.W., D.R. HeresT, AND C.T. ImaDa. 2006. New Hawaiian plant records for 2004. In: N.L. Evenhuis and L.G.

Eldredge, eds. Records of the Hawaii Biological Survey for 2004—2005. Part 2: Notes. Bishop Mus. Occas. Pap.

WuNDERLN, R.P AND B.F, HANSEN, 2003. Guide to the vascular plants of Florida (ed. 2). Univ. of Florida Press, Gainesville

WUNDERLIN, R.P. AND B.F. HANSEN. 2008. Atlas of Florida vascular plants. [S.M. Landry and K.N. Campbell (application

development), Florida Center for Community Design and Research.] Institute for Systematic Botany, Univ. of

South Florida, Tampa. «http;/www.plantatlas.usf.edu/»

464 Journal of tani it Texas 3(

BOOK NOTICE

FLORA or CHINA EDITORIAL Committee. 2008. Flora of China Illustrations, Vol. 13. Clusiaceae through

Araliaceae. (ISBN Vol. 13 illustrations: 978-1-930723-80-1, hbk.). Science Press (Beijing) and Missouri

Botanical Garden Press (St. Louis), PO. Box 299, St. Louis, Missouri 63166-0299, U.S.A. (Orders: www.

mbgpress.org, mbgpress@mobot.org, 1-877-271-1930). $125.00, 491 pp., 8 3/4" x 11 1/4".

From the publishers.—“Volume 13 of the illustrations series is devoted to 33 plant families, a number of which are of horticultural or

agricultural importance or are otherwise distinctive: Violaceae, with its white, purple, and yellow violets; Passifloraceae, the pass

flower es kena of which Bn genus ee furnishes us with the papaya dins "Besonidceae ln E m ii its orna-

£1

; Thymelaeaceae, i

used for the e of paper e medianes ee the water chestnuts; esca ini Camptotl source of

the anticancer drug ot ak Davidia, the dove tree, ird Hane occurs only in China; Onagraceae, dos evening

primrose and Epilobium ili d Araliaceae ell as Panax or ginseng.”

ditors Wu Zhengyi, Peter H. Raven, and H e, "This Sus is the 14^ in a series of 24 volumes of Flora of China

Illustrations. It ium Flora of China (FOC) Volume 13, which was published in 2007 and treated 151 genera with 1288 species in

volume of ursi ME 470 plates fepe a total of 774 species, 12 subspecies, and 18 varieties in 157

" Also i Preface, Aci g Index to Se Names, Index to Pinyin Names, Index to

to Families in the Flora of China and the Flora Reipubli 1 Sini and Published bia Di i oe

of China and ther Flora of China Illustrations. A large map of Chi d its 35 Admini ive Divisi I

/f4 1 1 «1 D 2212531 1 A 1 A

aes as 33 families.

J. Bot. Res. Inst. Texas 3(1): 464. 2009

TOURNEFORTIA HIRSUTISSIMA (BORAGINACEAE)

NEW TO THE FLORA OF TEXAS

Alfred Richardson W. Ken King

Departmentof Biological Sciences 1008 W 5% Street

The University of e at eee Weslaco, Texas 78596, US.A

80 Fort Bro

Brownsville, Texas 28 USA

ABSTRACT

Tournefortia hirsutissima (Boragi ) is rep 1f th Texas, a new record for the state of Texas.

RESUMEN

Te fortia hirsutissima (Boragi ) se cita del ext de Tejas, es el primer registro para el estado de Tejas.

Tournefortia hirsutissima L. (Chiggery Grapes, Mata de Nigua) is native to various parts of Mexico, Central and

South America, and the West Indies (Standley1924). It also grows in Florida (Wunderlin & Hansen 2003) but

has not been reported for Texas (Turner et al. 2003). It is a sprawling and climbing shrub reaching at least 5

meters in height (Fig. 1). The stems and leaves are covered with stiff hairs which can cause irritation to the

skin. The leaves are alternate, the blades ovate to elliptic and up to 15 cm long. The flowers, in branching

cymes, have a fragrance somewhat like that of orange blossoms. The corollas are white, actinomorphic, ca.

1 cm broad, the tubes 4-5 mm long. The fruit are white drupes ca. 8 mm broad, each with one small black

dot where the style has fallen, and containing 2-4 mericarps (Fig. 2).

Tournefortia hirsutissima has been given many common names in different countries. Both of the two

that we have listed refer to the belief that rubbing an infected area with a leaf will kill or remove chiggers.

The Spanish name means “chigger killer.”

The plant was called to our attention by Mary Ann and David Sato through a mutual friend, Mike Heep.

It was growing on the Sato property along Hudson Street in San Benito, Cameron County, Texas. We visited

their home in March and April of 2007. The liana was growing next to the road. It was very large, covering

an estimated 7 meter square area and growing 5 meters into the trees. Three more individuals were growing

at the back of the property. The Satos said that they had not planted them and that they grew without any

care. We suspect that the plants arrived by natural means, probably as seeds deposited by birds. Llera and

El Encino, two collection sites in Tamaulipas, Mexico, are less than 400 km from San Benito.

Since the plant we examined was so robust, flowering and fruiting freely, it seems that it will gradu-

ally spread to other areas, at least in years when rains are abundant. It appeared to be old enough to have

survived some of our drier years, although its native habitat generally receives more rainfall than Cameron

County. It probably would not be so lush during the dry times.

Voucher specimens: TEXAS. Cameron Co.: S of San Benito on Sam Houston, E on Hudson. One large sprawling liana, at roadside,

occupying an area about 20! x 20', climbing 15’ high into trees, 31 Mar 2007, Richardson and King 3315 (BRIT, TEX); S of San Benito on

Sam Houston, E on Hudson, one large sprawling liana, at roadside, occupying an area about 20’ x 20’, climbing 15” high into trees, 9

Apr 2007, Richardson and King 3320 (BRIT, TEX).

ACKNOWLEDGMENTS

We thank Mary Ann and David Sato and their friend Mike Heep for noticing and bringing Tournefortia

hirsutissima to our attention. Monique D. Reed and an anonymous reviewer provided helpful reviews.

J. Bot. Res. Inst. Texas 3(1): 465 — 467. 2009

Journal of the Botanical Research Institute of Texas 3(1)

Fic. 1. Tournefortia hirsutissima. Flowers.

Fic. 2. Tournefortia hirsutissima. Fruit.

Richardson and King, Tournefortia hirsutissima new for Texas 467

REFERENCES

STANDLEY, P.C. 1924. Tournefortia. In: Trees and shrubs of Mexico. Contr. U.S. Natl. Herb. 23:1229-1234,

Turner, B.L., H. NicHoLs, G. Denny, AND O. Doron. 2003. Atlas of the vascular plants of Texas. Vol. 1. Botanical Research

Institute of Texas, Fort Worth.

WUNDERUN, R.P. AND B.F. HANSEN. 2003. Guide to the vascular plants of Florida. Second Edition. University Press of

Florida, Gainesville.

468 Journal of the Botanical Research Institute of Texas 3(1)

ANNOUNCEMENT

THE 2009 APPLICATIONS FOR THE DELZIE DEMAREE TRAVEL AWARD

Applications for the 2009 Delzie Demaree Travel Award should include a letter from the applicant telling

how symposium attendance will benefit his/her graduate work and a letter of recommendation sent by the

major professor. Please send letters of application to: Dr. Donna M.E. Ware, P.O. Box 8795, Herbarium, Biol-

ogy 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 applications are 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 2009 are 9-11 October 2009.

The Delzie Demaree Travel Award was established in 1988 honoring Delzie Demaree who attended

35 out of a possible 36 symposia 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.

Anyone interested in making a contribution to Delzie Demaree Endowment Fund, which supports the

travel award, may make contributions by VISA or MasterCard or by a check, payable to Botanical Research

Institute of Texas, to Barney Lipscomb, 500 E 4th Street, Fort Worth, TX 76102-4025, U.S.A. 1-817-332-

7432; Email: barneyObrit.org. Thank you.

J. Bot. Res. Inst. Texas 3(1): 468. 2009

AVAILABLE TITLES FROM THE BRIT PRESS

SBM 01 | A Quantitative Analysis of the Vegetation on the Dallas County White Rock Escarpment

SBM 02 | The Vascular Flora of St. Francis County, Arkansas

SBM 04 | Asteraceae of Louisiana

SBM 05 | The Genus Mikania (Compositae: Eupatorieae) in México

SBM 06 | Frontier Botanist William Starling Sullivant's Flowering-Plant Botany of Ohio

SBM 07 | A Taxonomic Revision of the Acaulescent Blue Violets (Viola) of North America

SBM 08 | Aster & Brachyactis (Asteraceae) in Oklahoma

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SBM 20 | Generic Conspectus of the Tribe Asterae (Asteraceae) in North & Central America

SBM 21 | A Comparative Checklist of the Plant Diversity of the Iwokrama Forest, Guyana

SBM 22 | Lloyd Herbet Shinners: By Himself

SBM 23 | Taxonomy, Distribution, & Ecology of the Genus Phaseolus (Leguminosae)

SBM 24 | Atlas of the Vascular Plants of Texas, Volume 1 & 2

SBM 25 | Los Géneros de Léguminosas del Norte de México

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SBM 28 | Wild Flowers of Mombacho (Nicaragua)

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SBM 30 | Tundra to Tropics: The Floristic Plant Geography of North America

SBM 31 | A Fifth Checklist of Tennessee Vascular Plants

OTHER PUBLICATIONS

Keys to the Vascular Plants of the Black Gap Wildlife Management Area, Brewster County

Mosses of Texas: A Manual of the Moss Flora

Eriocaulaceae of Continental North America, North of México

Trees in the Life of the Maya World

Violets (Viola) of Central & Eastern United States: An Introductory Survey

The Ecology of our Landscape: The Botany of Where we Live

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