JOURNAL OF THE BOTANICAL RESEARCH INSTITUTE OF TEXAS

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

VOLUME 3 NUMBER2 25 NOVEMBER 2009

ac Botanical Research Institute of Texas (BRIT)

E 500 East 4th Street

am 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

Electronic mail: barney@brit.org; jbrit@brit.org

HISTORY AND DEDICATION

1962—Lloyd H. Shinners

(left), a member of the

Southern Methodist University

(SMU) faculty and a prolific

researcher and writer, published the first issues of Sida,

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

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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BIBLIOGRAPHICAL

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

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Electronically tinted botanical illustration

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BRIT

PRESS

TABLE OF CONTENTS

SYSTEMATICS

Xyris spathifolia (Xyridaceae), a new xyrid from the Ketona Dolomite-Limestone glades

of Alabama

ROBERT KRAL AND J. Mincy MOFFETT, JR.

Eriogonum cedrorum (Polygonaceae: Eriogonoideae), a new species from northwestern

California

JAMES L. REVEAL AND ROGER RAICHE

A new narrowly endemic species of Dirca (Thymelaeaceae) from Kansas and Arkansas, with a

phylogenetic overview and taxonomic synopsis of the genus

AARON J. FLODEN, MARK H. MAYFIELD, AND CAROLYN J. FERGUSON

Two new combinations in Tryphane (Alsinaceae)

W.A. WEBER AND R.C. WITTMANN

Two new species of Solanum from Ecuador and new combinations in Solanum section

Pachyphylla (Solanaceae)

STEPHEN R. STERN AND LYNN BoHs

Three new species of Solanum section Herpystichum (Solanaceae) from Ecuador

Eric J. TEPE AND LYNN Bonus

Stenaria sanchezii (Rubiaceae), a new and endangered species from Sonora, Mexico

DaviD H. LORENCE

Muhlenbergia tarahumara (Poaceae: Chloridoideae: Cynodonteae: Muhlenbergiinae),

a new species from Chihuahua, Mexico

PAUL M. PETERSON AND J. TRAVIS COLUMBUS

Una nueva especie y novedades nomenclaturales en el género Meliosma (Sabiaceae)

J. FRANCISCO MORALES

Estudios en las Apocynaceae neotropicales XXXVII: monografía del género Rhabdadenia

(Apocynoideae: Echiteae)

J. FRANCISCO MORALES

Estudios en las Apocynaceae neotropicales XXXVIII: tres nuevas especies de Mandevilla

(Apocynoideae: Mesechiteae) para Colombia y Venezuela

J. FRANCISCO MORALES

Studies of neotropical Compositae—II. A new combination in Chromolaena (Eupatorieae)

JOHN F. PRUSKI

Folia taxonomica 13. Passiflora curva (Passifloraceae), a new species from French Guiana

in subgenus Passiflora supersection Coccinea

CHRISTIAN FEUILLET

Folia taxonomica 14. Notes on Passiflora supersection Coccinea (Passifloraceae) from the

Guiana Shield

CHRISTIAN FEUILLET AND JOHN VANDERPLANK

Folia taxonomica 15. Five new species of Paradrymonia subgenus Paradrymonia (Gesneriaceae:

Episcieae) from the Venezuelan Guayana

CHRISTIAN FEUILLET

Folia taxonomica 16. Dilkea (Passifloraceae) 1. Epkia, a new subgenus and five new species

from western Amazonia and the Guianas

CHRISTIAN FEUILLET

A new species of Schefflera (Araliaceae) from the Venezuelan Andes

L.J. Dorr AND BASIL STERGIOS

New species of Eugenia (Myrtaceae) from Ecuador

MARIA LUCIA KAWASAKI AND BRUCE K. HOLST

469

479

485

501

503

511

521

527

535

541

565

573

577

581

583

593

605

Revision of Bauhinia subgenus Phanera section Schnella (Cercideae: Caesalpinioideae:

Fabaceae)

RICHARD P. WUNDERLIN AND RICHARD M. EILERS

Revision of Bauhinia subgenus Bauhinia section Pauletia series Ariaria (Cercideae:

Caesalpinioideae: Fabaceae)

RICHARD P. WUNDERLIN

Eriogonum villosissimum (Polygonaceae: Eriogonoideae), a new species endemic

to Acker Rock, Oregon

JAMES L. REVEAL, DANA YORK, AND RICHARD HELLIWELL

Nomenclatural innovations in Phlox (Polemoniaceae), with updated circumscription

of P. caespitosa, P. douglasii, P. missoulensis, and P. richardsonii

JAMES H. LOCKLEAR

Validation of the name Symplocos pentandra (Symplocaceae)

Joáo Luiz M. ARANHA FILHO, PETER W. FritscH, FRANK ALMEDA, AND ANGELA B. MARTINS

Clarifications concerning the nomenclature and taxonomy of Oxypolis ternata (Apiaceae)

Mary ANN E. FEIST

Neotypification of Artemisia carruthii (Asteraceae: Anthemideae)

CALEB A. MORSE AND JAMES L. REVEAL

A phylogeny of Arctostaphylos (Ericaceae) inferred from nuclear ribosomal ITS sequences

GREGORY A. WAHLERT, V. THOMAS PARKER, AND MICHAEL C. VASEY

The reinstatement of Beautempsia (Capparaceae) and a key to the genera of neotropical

Capparaceae with variously stellate or peltate indumenta

XAVIER CORNEJO AND HUGH H. ILTIS

Revisión de las especies sudamericanas del género Schwartzia (Marcgraviaceae: complejo

Norantea)

DIEGO GIRALDO-CAÑAS

Again: taxonomy of yellow-flowered caulescent Oxalis (Oxalidaceae) in eastern North America

Guy L. Nesom

Clarification of the typification of Michaux names in eastern North American Vitis (Vitaceae)

CONNIE FISK AND ALEXANDER KRINGS

Additional morphological measurements of Arceuthobium siskiyouense and A. monticola

(Viscaceae)

ROBERT L. MATHIASEN AND CAROLYN M. DAUGHERTY

Additional observations about Phaseolus rotundatus (Fabaceae), an endemic bean species

from western Mexico

Jesús SALCEDO C., ROGELIO LÉPIZ I., NORA CASTAÑEDA A., CESAR OCAMPO N., AND DANIEL G. DEBOUCK

BOTANICAL HISTORY

Discovery of a Lewis and Clark grass collection, Poa secunda (Poaceae) sensu lato, at the

herbarium of the Royal Botanic Gardens, Kew

JACQUES CAYOUETTE, JAMES L. REVEAL, AND GARY E. MOULTON

FLORISTICS, ECOLOGY, AND CONSERVATION

Pinus pinceana (Pinaceae), nuevo registro para el estado de Nuevo León, México

SUSANA FAVELA LARA, CARLOS G. VELAZCO MACÍAS Y GLAFIRO J. ALANÍS FLORES

Diversidad y distribucion de las gramineas (Poaceae) en el estado de Zacatecas

YOLANDA HERRERA ARRIETA Y ARMANDO CORTÉS ORTIZ

Lipocarpha micrantha (Cyperaceae), nueva cita para la flora de Argentina: micromorfología

del fruto

PRISCILA R. VAN DE VELDE Y M. GABRIELA LÓPEZ

619

629

639

645

659

661

667

673

683

691

727

739

741

751

763

TEI

775

793

Index to Volume 3 (2009)

Titles of Articles with Authors—998

Authors—1001

Botanical Names and Subjects—1002

New Names and Combinations—1007

INDEX to new

J, Bot. Res. Inst. Texas 3(2), 2009

Appunia longipedunculata (Steyerm.) Delprete,

comb. n 9

V—

Appunia peduncularis (Kunth) Delprete, comb.

v.—809 .

Bauhinia esmeraldasensis Wunderlin, sp. nov—

631

Bauhinia lilacina Wunderlin & Eilers, sp. nov—

621

Bauhinia trichosepela P 29 Wunderlin &

Eilers, comb. nov. —

Chromolaena Eis (DC.) Pruski, comb.

nov.—573

Dilkea Heeh Feuillet, sp. nov.—595

Dilkea subg. Epkia Feuillet, subg. nov.—594

Dilkea lecta Feuillet, sp. nov.—597

Dilkea ovalis Feuillet, sp. nov.—599

Dilkea tillettii Feuillet, sp. nov.—599

Dilkea vanessae Feuillet, sp. nov.—602

Dirca decipiens A. Floden, sp. nov. —494

Eriogonum cedrorum Reveal & Raiche, sp.

nov.—479

Eriogonum villosissimum Reveal & D.A. York, sp.

nov.—639

is M.L. Kawasaki & B.K. Holst,

sp. nov.—609

Eugenia castaneiflora M.L. Kawasaki & B.K. Holst,

sp. nov.—6

Eugenia crassimarginata M.L. Kawasaki & B.K.

Holst, sp. nov. —611

Eugenia oe M.L. Kawasaki & B.K. Holst,

sp. n

pa a M.L. Kawasaki & B.K. Holst,

nov.—6

Mandevilla albifolia J.E Morales, sp. nov.—565

Mandevilla caquetana J.F Morales, sp. nov.—567

Mandevilla sancta-martae J.F Morales, sp.

nov.—5069

Meliosma isthmensis J. E nid sp. nov.—537

M. Peterson & Colum-

bus, sp. nov.—

Paradrymonia glandulosa Feuillet, sp. nov.—583

Paradrymonia hamata Feuillet, sp. nov.—585

Paradrymonia lutea Feuillet, sp. nov.—585

Paradrymonia tepui Feuillet, sp. nov.—588

Paradrymonia yatua Feuillet, sp. nov.—588

Passiflora curva Feuillet, sp. nov.—577

Phlox austromontana subsp. Carr (Wherry)

Locklear, comb. et stat. no 5

| ee (S.L. Welsh)

Locklear, comb. et stat. nov —646

Phlox a ta subsp. € (E.E. Nelson)

Locklear, comb. et stat.

Phlox pilosa subsp. longipilosa (Waterf. ) Locklear,

omb. et stat. nov.—

Phlox richardsonii subsp. borealis (Wherry) Lock-

comb. nov.—647

Phlox subulata = setacea (L.) Locklear; comb.

et stat. 49

Phlox “dio sia mid latisepala (Wherry)

Locklear, comb. nov.—649

Schefflera vanderererftil Dorr & Stergios, sp.

nov.—605

faha Dal

ae Lay

A aca han

us nov.—509

Solanum crassinervium Tepe, sp. nov. —514

Solanum diversifolium subsp. chloranthum

a Bohs, comb. nov.—509

malha nal

hi 5 ?

comb. n

ban erdum Bohs, nom. nov.—509

Solanum limoncochaense Tepe, sp. nov.—516

Solanum manabiense S. Stern, sp. nov.—504

Solanum pacificum Tepe, sp. nov. —512

Solanum zumbense Bohs, sp. nov.—506

Stenaria sanchezii Lorence, sp. nov.—522

Symplocos pentandra (Mattos) Occhioni ex Aranha,

comb. et stat. nov.—660

Tryphane rubella (Wahlenb. 4 A. & D. Love ex WA.

Weber, comb. nov.—

Tryphane rubella subsp. ie (Richardson) A.

D. Love ex WA. Weber, comb. nov.—502

Xyris spathifolia Kral & Moffett, sp. nov.—472

XYRIS SPATHIFOLIA (XYRIDACEAE), A NEW XYRID FROM THE

KETONA DOLOMITE-LIMESTONE GLADES OF ALABAMA

Robert Kral J. Mincy Moffett, Jr.

Botanical Research Institute of Texas Georgia Department of Natural Resources

500 E. 4" Street Wildlife Resources Division

Fort Worth, Texas 76102-4025, U.S.A. Nongame Conservation Section

Georgia Natural Heritage Program

5 U.S, Highway 278 S.E.

Social Circle, Georgia 30025-4743, U.S.A.

Mincy.Moffett@dnrstate.ga.us

ABSTRACT

O qiu: : Vion 1 ry 1; TE T Kon nha ral l4] ti (of T3 Y fen in Bibb C ty, Alabama

El [e] M ^ e ME

TI lty g longside tl 1 endangered X. tennesseensis, but i istently distinct from it. We distinguish this putative new

[. "S 1-2 NE. TN : 1- 1 ] 1 + ff: 74 11 J T T A —

» EYES LL SẸ a g , and

herbarium surveys. The new species, X. spathifolia is herein described, figured, and i lationship with X. tennesseensis discussed.

RESUMEN

C1 3 X. E = 1 141 Al e; on ry] 1 A 15 da 5 at ^ 1 A | da 1:11

$! peq (. )( : , Alabama. Esta

novedad crece junto con la rara y amenazada X. tennesseensis, Į distinta de ella. Distingui t putativo, dándole

] E d 1 A 1 1 ‘ ] 21 . 1 1 1: 3 . 3 is "P

g E g po, | , y revisiones de

herbario. Se d ibe aquí 1 pecie, X. spathifolia, se ilustra, y se discute su relación con X. tennesseensis.

INTRODUCTION

Despite the pervasive notion that cataloging of the vascular plant flora of North America is complete, new

species continue to be discovered. Frequently, these discoveries occur in spatial or temporal clusters, re-

vealing a botanical “hot spot,” as in the case of the Ketona Glades of Bibb County, Alabama (Ertter 2000),

located at the extreme southern edge of the Ridge and Valley Ecoregion (Fig. 1). During the last 10-12

years, nine vascular plant taxa new to science (including the one described herein) have been identified

from this area, as well as seven state records including some regional disjuncts, and more than 60 taxa of

conservation concern (Allison 2001). With a total flora of over 420 species occurring within an area of ca.

125 ha (Alabama Water Watch 2002), it is one of the most botanically diverse areas in the eastern United

States (Allison 2001).

During a 1999 field survey of populations of the rare and endangered Xyris tennesseensis Kral in the

Ketona Glades, a markedly different xyrid, shorter in stature and smaller in vegetative and floral features,

was found admixed with the former in a small, intermittent seep at the headwaters of a tributary to Alliga-

tor Creek (M. Moffett s.n., 7 Aug 1999). On the date of the discovery, approximately 900 flowering spikes of

the novelty were tallied, emanating from approximately 200 clumps. Individuals of the putative new xyrid

showed consistent differences from X. tennesseensis the mixed population. This smaller and different

xyrid, thus far known only from the type locality, was once relatively abundant. However, severe drought

during the summers of 1999 and 2000 decimated the population. The drastically reduced population size

and fluctuating local climate have limited the collection of additional voucher material.

The site of this mixed Xyris population, containing the novel xyrid, is a fen/seep complex at the head-

waters of Alligator Creek, within an area mapped as “Alligator Glades West” by the Alabama Natural Heritage

Program (Alabama Natural Heritage Program 1999), and referred to as “Enchanted Glade” by Allison (2001).

Alligator Glades West is one of ca. 45 such sites comprising the Bibb County Ketona Glades and contains

the sole known population of this new taxon. The Ketona Glades are shallow-soiled, herbaceous communi-

J. Bot, Res, Inst, Texas 3(2): 469 — 478. 2009

53/5)

470 tani titute of Texas 3(2)

Lo) le tg ae

v T 0 50 100 — 150 200 Km

em Coastal P! bi titi ti |

Griffith et al. Ecoregions Level ill (1987)

Fic. 1. Map of the study area.

ties atop outcrops of Ketona dolomite, an unusually pure dolomite (i.e., few siliceous contaminants) with

relatively high levels of Mg (> 1,100 ppm). The seep itself is very small, ca. 24 m? (3 m x 8 m) in size. It is

located 1.7 km N of Bulldog Bend (Little Cahaba River) in Bibb County, Alabama (Lat. 33? 04"; Long. 87°

01). Ownership of this site has changed several times during our investigation. It is currently owned by

Forest Investment Associates, and previously by Great Eastern Timber Company, and prior to that by U.S.

Alliance/Coosa Pines Corporation, all of whom we are indebted to for access. Extensive searches made along

Alligator Creek, and of the six glades (and their ecotones) in Bibb County known to contain X. tennesseensis,

have yielded no other populations.

METHODS

Common Garden Experiment

The common garden experiment initially developed from an attempt to prevent the extinction of this in-

teresting new taxon. During the severe summer drought of 2000, and following the severe drought in late

summer and fall of 1999, there was concern regarding the future viability of this sole known population of

X. spathifolia. In August 2000, as a hedge against possible future extirpation, a mature fruiting spike was

collected from each of the 13 surviving clumps sporting at least two such spikes. A mature fruiting spike

also was collected from 13 of the surviving 21 flowering clumps of X. tennesseensis. Spikes were stored in

separate brown paper sacks in an unheated outdoor shed in Opelika, Lee County, Alabama.

In February 2001, seeds from each spike were sown onto a 60:40 mixture of Pro-Mix (Premier Horti-

culture, Dorval, Canada) and sand in separate 15.2 cm diameter (6-inch) clay pots and grown in a heated/

evaporatively-cooled greenhouse at the Auburn University Plant Science Research Center (PSRC). Ap-

proximately 50-100 seeds germinated in each pot by June. Plants were thinned to about 10 per pot during

Ww [| IRS ff. VM 5 Py? ee 1? "JE Alak ea

r AYTIS 5| , 471

the second year, and their parental identity was carefully maintained. Seedlings removed from pots (and

hence the experiment) were grown commingled in flat trays in a separate part of the greenhouse. Annual

greenhouse temperatures typically range from ca. 18.3-29.4°C, per PSRC records. During the growing

season (March to October), the average daily photosynthetic photon irradiance at bench level (adjusted for

60% shade cloth) ranged from 3 mol m-2 d-1 to 15 mol m-2 d-1, on overcast and sunny days, respectively

(Elkins and Wallace 2000). Plants were fertilized every two weeks with liquid 20-20-20 (N-P-K) during the

growing season. During the subsequent three years, the seedlings matured and reproduced vegetatively,

generating clumps of ca. 30-40 ramets and 30-60 flowering spikes per pot by the summer of 2003

During the summer of 2004, we realized that what began as a “rescue propagation” could provide data

for a limited morphometric analysis of the principal characters distinguishing these syntopic xyrids. The

3-year-old plants of known parentage grown from seed in a common environment constituted a “common

garden” experiment. Sample size was N = 25; 12 pots of X. spathifolia and 13 pots of X. tennesseensis.

The nine morphological characters selected for measurement are aspects of the new taxon that, in the

field, appeared most obviously different from X. tennesseensis, and were also easily quantifiable. The charac-

ters are: leaf length, leaf width, plant height, spike length, lateral sepal length, petal length, anther length,

stamen length, and seed length. In all cases, measurements were taken at the widest (or longest) part of a

structure along the appropriate axis. With regard to plant height, the longest or “tallest” plant structure for

each ramet was measured. This was usually a measure of maximum scape length, although in some cases

the structure of greatest length was a leaf. Flower petal length was measured from base of claw to blade

apex. Stamens were measured from the point of basal fixation with the anther to the lowest point of basal

adnation to the corolla. Seeds were measured along the major axis (i.e., tip to tip). Data were not collected

on several other important characters used in North American lla Species classification because of the

high similarity of the two taxa for these characters. Excluded luded: keel features of the lateral

sepal, position of the lateral sepal relative to the bracts (i.e., inserted/exserted), color and texture of the leaf

sheath, and the presence/absence of a mealy oa o ia ind

Samples were generated by first ] in a pot, and then randomly selecting

five ramets from each pot. Sample values for pad pot were created by averaging the values recorded from

the five randomly selected ramets. Since most of the plants resulting from the original propagation did not

flower and produce seeds until summer/fall 2003, it is certain that most ramets selected for measurement in

this experiment were the result of asexual, vegetative reproduction. Thus, each sample represented a mean

of the progeny of a single distinct, field-identified clump. Samples for different characters were developed

using different approaches.

Measurements of leaf length and lia were gathered from the four outermost leaves of each selected

ramet. Spike lengths were obtained spikes from a selected ramet. Measurements of floral structures

were recorded from the single open [lower on Pac Spe (itis usually the case iiis idis inal only one flower

]

h CTY1 TE

opens each day per spike). For seed length

and numbered. One capsule was randomly selected fror each spike and then five seeds were subjectively

chosen from each capsule for measurement. Plant height ts involved but a single value for each

ramet. Length measurements for petals, sepals, stamens, anthers, and seed length were made using a light

microscope with a stage micrometer.

Differences in the nine character means for the two taxa were statistically analyzed using the Student’s

t-test, except when data “eae ay assumptions and were not subsequently improved using either

square root or logarit! This was the case involving seed size, where the non-parametric

Mann-Whitney U-test was employed. Normality was assessed using the Kolmogrov-Smirnov and Wilk-

Shapiro tests. Homogeneity of variance pl were evaluated using Levene's test, with Levene's cor-

rection applied to violations. Effect sizes for t-tests are expressed as eta” values, and for the Mann Whitney-U

test by an effect size correlation (r yA) using Cohen’s D. Statistical software utilized was SPSS 16.0. Analysis

adhered to procedures found in Pallant (2001) and Shannon and Davenport (2000).

472 t tani titute of Texas 3(2)

Been en Hera Records

, regional, and national herbaria

ihis project Taxa selected for the review process were either part of the X. difformis “complex” (Kral 0 or

known to occur syntopically with X. tennesseensis. The X. difformis complex comprises X. difformis (varieties

curtissii, difformis, and floridana), X. tennesseensis, and X. torta. Xyris jupacai is not considered a member of

this complex, but does co-occur with X. tennesseensis at a few locations. Only those specimens collected from

Alabama, Georgia and Tennessee (the tri-state range of X. tennesseensis) were examined. The purpose of this

investigation was to determine if specimens of X. spathifolia had been collected previously under a different

name, thereby providing not only ana specimens for study, but also expanding the known range of

the new taxon. Approximately 900 specimens of X. difformis, X. jupicai, X. tennesseensis, and X. torta were

examined from vascular plant collections at AUA, DUKE, GH, JSU (Jacksonville), JSU (Anniston Museum

of Natural History), MO, NY, US, TROY, UNAF, HALA, UNA, FLAS, GA, MICH, NCU, TENN, and VDB at

BRIT. Herbarium acronyms follow Index Herbariorum (Holmgren 2008).

RESULTS

Common Garden Experiment

Analysis of morphological characters from greenhouse grown plants in a common garden setting revealed

eae differences in all nine size-related traits. In all cases, for X. tennesseensis specimens

y larger than tl for X. spathifolia (Table 1). Significance values ranged from « 0.001 to 0.03,

with effect sizes indicate that 39-93% of the variance in each character was explained by taxon identity.

Table 2 provides an assessment of character state differences between the two taxa based on vouchered

herbarium specimens and field-collected material.

Review of Hear oe

Review of pr j OLX difformis, X jupicai, Xt j 1X. torta from 19 national,

regional and state herbari "located only a single record fitting the concept of X. spathifolia, collected by James

R. Allison in 1993 from Alligator Glades West, the sole known extant site of this new taxon.

IAAUNOMIC DESCRIPTION

¥wri thifolia Kral & Moffett, Sp. NOV. (Fig. 2). Tyre: U.S.A. AragAMA. Bibb Co.: Alli Glad t, small seep over Ketona

Dolouiite at headwaters of W branch of Alligator Creek (Lat 33° 04'; Long 87°, 01°), 7 me 1999, M. Moffett s.n. (HOLOTYPE: VDB).

Xyris spathifolia, Kral et Moffett, sp. nov., X. tennesseensi Kral (Fig. 3) similli 1 st Í i 5-40 (non aie e) a aa

Ng ecu cine valde o lev id. papillosis, scapis apicem versus valde 5—7-costatis (non bicostatis),

, differ

Perennial, cespitose, 15—40(—50) cm tall, roots fibrous. Stems short, or present as slender, ascending, rhi-

zome-like bases on innovations (result of burying of clump by seciment). Leaves ascending to erect, 15—30

cm long, soft, sheaths often fully as long as blade or longer, gradually widening from keeled apex to convex

base, proximally multi-costate, smooth, pale tan to brown, upsheath with tints of red or pink, progressively

increasingly papillose, densely so and green distally, margins pale, scarious, gradually narrowed to blade

(this often most of its length infolding ait leaf of innovation); blade linear, mostly 1-2 mm wide, slightly

to very twisted, flattened, lly to medially strongly papillose, toward apex smooth or nearly

so; apex narrowed, excetitedi cU. its incurved tip incrassate, blunt, smooth; surfaces densely pale-

granular-papillose and interruptedly rugulose proximally, progressively smoother up-blade. Scape sheaths

shorter than principal leaves, mostly open, short-bladed. Scapes erect or ascending, DUNS aded ca.l

mm thick, twisted, pale green, proximally subterete with (mostly) 5—8 papillate costae vals smooth

or nearly so, distally sharply unequally angulate-ribbed, with up to 57) poete malis the angles, 2-3 most

raised, all densely papillate-tuberculate, the intervals sulcate to broadly concave, slightly to very papillose

or papillose-rugulose in short lines. Spikes ovoid to ellipsoidal or lance-ovoid, 5-8 mm long, blunt, several

flowered, bracts in flat spiral, loosely imbricate, the proximal 23 sterile, narrowly ovate-triangular, 2.53 mm

yrid from Alat 473

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X. spathifolia X. tennesseensis

(n= 12) (n= 13) Test Effect

Character (mean + S.E.) (mean + S.E.) d.f. Statistic Sig. Size

Leaf length (cm) 24.1 + 1.57 48.9 t 2.3 20.8 -8.87 * «0.001 0.774

Leaf width (mm) 1.51+0.16 6.98 + 0.69 13.3 -7.77 * <0.001 0.733

Piant height (cm) 27.5 + 3.54 48.6 + 3.83 23 -4.05 0.001 0.413

Spike length (mm) 6.75 +0.42 130 +067 19.8 -7.93 * <0.001 0.732

Petal length (mm) 3.46 + 0.03 4.44 + 0.04 22.2 -17.5 * <0.001 0.930

Sepal length (mm) 4.27 + 0.06 4.77 X 0.07 23 -5,59 <0.001 0.575

Stamen length (mm) 1.01 + 0.02 2.14 + 0.06 157 -17.1* «0.001 0.926

Anther length (mm) 0.55 + 0.02 1.74 + 0.07 13.8 -17.1 * <0,001 0.927

Seed length (mm) 0.52 + 0.01 0.55 + 00] 22.7 -2.18 0.03 0.39

Character State Xyris spathifolia Xyris tennesseensis

15~40(—50) cm

May be short-ascending-rhizomatous

(ascending lateral offshoots)

15-30 cm x 1-2 mm

Plant height 30-70(-85) cm

Stem base Lacks rhizomes

Principal leaves

x width)

Leaf surfaces

40—60 cm x 4-10 mm

Sheath papillate medially and distally,

blade edges papillate, proximal

surfaces papillate

Sheath smooth or finely papillate, sheath keel

sometimes papillate, blade edges

mostly smooth, surfaces smooth

Scapes Subterete and multicostate proximally, ca. Subterete and multicostate proximally, 2-3 mm

1 mm wide distally, sharply 5- wide distally, often compressed, 3-5 ridged,

isodiametric, ridges densely papillate- with 2 ridges flattened and wing-like, all

tuberculate, some sharply raised, intervals papillate at edges, intervals smooth, level to

sulcate or deeply , papillate shallowly concave

Spikes Ovoid to ellipsoidal or lanceovoid, 5-8 mm Ovoid, 10-15 mm

Lateral sepals

Petal blades

Stamens

Seeds

4.0-4.5 mm long, lanciform, keel

ascending-lacerate apically

Obovate, ca. 3.5 mm

1-1.1 mm long, anthers 0.5-0.6 mm

Ellipsoidal, ca. 0.5 mm, slightly farinose

slightly to very farinose

4.5—5.0 mm long, lanciform, keel ascending-

lacerate apically

Obovate, ca. 4.5 mm

2-24 mm long, anthers 1.5-2 m

Ovoid to broadly lo ca. DE 5-0.6 mm,

long, keeled, the fertile ones broadly obovate, 4.5-5 mm long, convex, apex broadly rounded, entire, ag-

ing erose, surface a lustrous rich brown with a green, ovate, subapical, granular dorsal area. Lateral sepals

free, equilateral, oblanceolate (viewed from side), ca. 4-4.5 mm long, acute, the thin keel entire proximally,

progressively widening and shallowly ascending-lacerate distally. Petals distinct, blades obovate (measured

from claw apex), ca. 3.5 mm long, apically erose; staminodia bibrachiate, staminodial hairs pencillate,

slightly clavate. Capsule ellipsoid-cylindric, 2-2.5 mm long, placentation 3-parietal; seeds broadly ellipsoid

to narrowly ovoid, ca. 0.5 mm long, apiculate, with 8—10 ribs/side, slightly farinose.

Etymology.—The epithet “spathifolia” is based on a peculiar morphology of lower leaves of innovations.

These are spathe-like, their margins conduplicate and enfolding inner leaves.

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PARATYPE: ALABAMA. Bibb Co.: Alligator Glades Wes

33° 04'; Long 87°, 01°, 7 Aug 1999, R. Kral & M. q. 90103 Md

Other specimens examined: ALABAMA. Bibb Co.: Alligator Glades Wi

Alligator Creek (Lat 33? 04'; Long 87°, 01), 5 Sep 1993, James R. Allison 7963 (AUA).

OE A A TA] ite at headwat fW branch of

KEY TO XYRIS TENNESSEENSIS SYMPATRIC XYRIDS, INCL. XYRIS DIFFORMIS COMPLEX

1. Keel of laeral sepals usually firm, papilate, ciliate, ciliolate, or fimbriate, or in various combinations of these

Xyris torta

. Keel of lateral Sepa scatious lacerate to (rarely) nearly entire.

eaf sheaths or sheath base light green straw-colored, or dull brown Xyris jupicai

2. eaS arida or mE base with red, pink, or purple tints, or glossy brown or red-brown.

opaque o

£ pus of mature e plant en

5. Plant 30— f sheat! | blad f tly smooth or just slightly papillat

along keels and edges; scapes witl hat flatt | and wing-like distally, all papil-

late at edges; stem base lacks rhizomes _ Xyris tennesseensis

5. Plant 15-40 cm; leaf sheaths and blade surfaces papillate; scapes sharply 5-7 angled distally,

irn oxi papillate-tuberculate; stem base possesses what may be short ascending

rhizo Xyris spathifolia

4. Base of Mim plant not bulbous Xyris difformis var. floridana

3. xc all

6. AS +ļ | ++ mu | iol | i tally;

2 ribs comparably wider, making v wings smooth or papillate Xyris difformis var. difformis

nutely | much y;

al equally oie somewhat DION Xyris difformis var. curtissii

DISCUSSION

As mentioned in the prefatory paragraph, this plant, thus far known only from the type locality, was once

abundant (over 200 clumps on the lower side of a small open seep). However, on our return to the site in

August 2000, severe drought had drastically reduced numbers (thus eliminating the chance for additional

voucher material). We hope to provide more vouchers once the population has recovered.

The question of what rank to assign this novelty is intriguing. Certainly, careful attention should be

given to the plant at its sole known location, as well as searching for it in other calcareous fens of the area. Of

particular interest is the fact that, prior to this discovery, only Xyris tennesseensis of North American xyrids

was known to occupy fen-like habitats. One would be tempted to consider the new morphology a simple

anomaly, perhaps a reaction to extreme habitat by X. tennesseensis, were it not for the side-by-side mixing of

id two, cs Sina are in Sonne on the basis of several characters.

thifolia and X. tennesseensis were manifest primarily

in terms of leaf and scape surface features (i.e., ridges, wings. papillae), and the size of individual characters,

as well as the overall plant. There was also a tendency for some X. spathifolia specimens to possess small

lateral ascending offshoots reminiscent of rudimentary rhizomes, a feature not known from other North

American xyrids. The differences exhibited by the new taxon are of a nature and degree that are consistent

with species status distinctions recognized for this genus (Kral 1966; Kral 2000).

CONSERVATION CONSIDERATIONS

Considering the documented rarity of X. spathifolia, its association with and similarity to X. tennesseensis,

and the specific threats associated with its single known extant site, a strong argument can be made for a

“high” conservation ranking. This sole isolated site is located within a fire-suppressed loblolly pine planta-

tion with severe woody tree/shrub encroachment. The overstocked plantings along (and within) the inter-

mittent stream course supplying the fen are capable of drastically reducing groundwater levels Jackson et

al. 2005). Moreover, this site lacks formal legal protection (i.e., it is privately owned with no conservation

easements/covenants in place). Thus it would seem to satisfy the requirements of both extreme rarity and

Fa [| Fr Els V. Li PI A "JE AlaLlos mm

¡ 477

threat necessary to justify a critically imperiled ranking both globally and in Alabama (G1/S1), using the

NatureServe (2008) system

The rarity and threat to X. al ifolia Id tits stat the xyrid of greatest conservation concern

in North America (north of Mexico). The ento E Xyris species of greatest tion concern

is X. tennesseensis, known from approximately 25 sites in diees states (AL, GA, and TN) with a NatureServe

global ranking of imperiled (G2). Xyris tennessensis also has status and receives protection under the federal

Endangered Species Act of 1973, under state law in Georgia through the Georgia Wildflower Preservation

Act of 1973 (Georgia Dept. of Natural Resources 2008), and under state law in Tennessee through the Rare

Plant Protection and Conservation Act of 1985 (Crabtree 2008). Conversely, as X. spathifolia is only now

being recognized as a new species, it enjoys no legal protections at either the federal or state level. Further

contributing to this precarious situation is the lack of any state laws protecting rare plants in Alabama.

The conservation situation involving X. spathifolia is obviously dire. Currently, there is some living

material remaining from the common garden study that will be safeguarded at Georgia Plant Conserva-

tion Alliance (GPCA) member institutions and the Auburn University Donald E. Davis Arboretum. Visits

to obtain additional X. spathifolia seed from the Alligator Glades West site, in order to bolster the existing

ex situ collections, will be made in August/September of this year (2009). Efforts will also be made to work

cooperatively with the new property owners of the Alligator Glades West site, Forest Investment Associates,

to better manage the site for the benefit of both xyrids.

ACKNOWLEDGMENTS

The authors wish to express their gratitude to the U.S. Alliance/Coosa Pines Corporation, and the Great

Eastern Timber Company for their access to the site, and to Jim Allison, Georgia Department of Natural

Resources, Natural Heritage Program, for his 1998 tour of all known Bibb County Glade sites containing

X. tennesseensis, including the site for this research. Two anonymous reviewers provided constructive and

helpful reviews. We also wish to acknowledge Curtis Hansen, Curator, John D. Freeman Herbarium, for

his assistance with loan material.

REFERENCES

ALABAMA NATURAL HERITAGE PROGRAM. 1999. Element oci Xyris t s. [he Nature Conservancy,

Montgomery, AL.

ALABAMA Water WarcH. 2002. Citizen guide to Alabama rivers (volume 1): Black Warrior and Cahaba. Alabama

Water Watch website, an on-line report series. http//www.alabamawaterwatch.org/awwnp/publications/

report, series/cahwar/war-cah.pdf (Accessed 11/3/04).

ALLISON, J.R. AND T.E. Stevens. 2001. Vascular flora of the Ketona dolomite outcrops in Bibb County, Alabama.

Castanea 66:154—205.

CRABTREE, T. 2008. Tennessee Natural Heritage Program: rare plant list. Tennessee Department of Environment

and Conservation, Division of Natural Areas. Updated January 2008. http://www:state.tn.us/environment/

na/pdf/plant list.pdf. (Accessed: May 10, 2008).

Enxins, C. AND B. WaLLace. 2000. PPFD and PAR data for PSRC greenhouse complex. Unpublished data, Plant

Sciences Research Center, Auburn University, AL.

ErTTER, B. 2000. Floristic surprises in North America North of Mexico. Ann. Missouri Bot. Gard. 87:81-109.

GEORGIA DEPARTMENT OF NATURAL CU 2008. Tue AE Division: Georgia rare species and natural com-

munity information. <http://georgiawildlife.org geredspecies_conservation.aspx (Accessed:

July 17, 2008).

HOLMGREN, PK. AND 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/. (Accessed May 10, 2008).

JACKSON, R.B., EJ. JosgAGy, R. Avissar, S.B. Roy, DJ. Barrett, C.W. Cook, K.A. Farley, D.C. LeEMAITRE, B.A. MCCARL, AND B.C.

Murray, 2005. Trading water for carbon with biological carbon sequestration. Science 310:944-1947,

478 [| 3 Eal D ak . 1m L I d den ndis fTexas 3(2)

KraL R. 1966. The genus Xyris (Xyridaceae) in the southeastern United States and Canada. Sida 2:177—260,

KraL, R. 2000. Xyris. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America north of

Mexico. 12+ vols. New York and Oxford. 22:154-167.

NatureServe. 2008. NatureServe Explorer: An online encyclopedia of life [web application]. Version 7.0. NatureServe,

Arlington, VA. http//www.natureserve.org/explorer. (Accessed: May 8, 2008).

PALLANT, J. 2001. SPSS survival manual. Open University Press, Buckingham, GreatBritain.

SHANNON, D. AND M. Davenport. 2001. Using SPSS to solve statistical problems. Prentice Hall Inc., Upper Saddle

River, NJ.

ERIOGONUM CEDRORUM (POLYGONACEAE: ERIOGONOIDEAE),

A NEW SPECIES FROM NORTHWESTERN CALIFORNIA

James L. Reveal Roger Raiche

L.H. Bailey Hortorium Herbarium Planet Horticulture Garden Design

Department of Plant Biology 3485 Old Lawley Toll Road

Cornell University Calistoga, California 94515, U.S.A.

412 Mann Library roger@planethorticulture.com

Ithaca, New York 14853, U.S.A.

jir326@cornell.edu

ABSTRACT

Eriog d fd bg. Oligog is pro] land ill 1 for a localized endemi finedt Due CMS area of Sonoma

Co., California. The new species, allied to E. nervulosum, differs f that species in having bright yell on shorter

stipes but associated with longer involucres.

RESUMEN

Se propone y se ilustra Eriogonum cedrorum del subg. Oligog demi finado al área de The Cedars del condado

de Sonoma, California. I E : p ntada con E A difiere d alri p fT “11, 1 illa te que

į R | = 1

i i

The Cedars wild buckwheat, Eriogonum cedrorum, has been known to botanists since first collected by

Freedom W. Hoffman (1880-1959) in 1947. In the mid 1980s Raiche made additional collections, which

Reveal mistakenly assigned to E. nervulosum (S. Stokes) Reveal, leading to confusion in the description of

that species (e.g., Reveal 2005). An opportunity to study the two species in the field in late July of 2009

showed that the two were not the same and The Cedars plant is now described formally.

Eriogonum cedrorum Reveal & Raiche, sp. nov. (Figs. 1-2). Tree: UNITED STATES. CALIFORNIA: Sonoma Co.: The Cedars,

Central Canyon area NE of Layton Mine and S of the “upper mine" on steep, gravelly, serpentine, N-facing slopes at 410 m elev.,

38°37'16"N, 123°07'37"W, TON, R12W, sec. 13SENE, 28 Jul 2009, J.L. Reveal & R. Raiche 8989 (HOLOTYPE: NY; isotypes: BH, BM,

BRY, CAS, GH, LL, MO, OSC, RM, RSA, TEX, UC, US, UTC).

A Eriogono nervuloso floribus flavis ( Ibis), invol longioribus (4—6.5 mm nec 3-4 mm) et stipitibus brevioribus (0.1-0.3 mm

nec 0.5—0.8 mm

Plants low spreading synoecious herbaceous perennials, 1-4(-10) dm long, 1-3(-5) dm across, composed

of loosely arranged rosettes of tufted leaves at tips of slender, woody caudex branches arising from a stout

woody taproot. Leaves tightly arranged in small, more or less well defined basal rosettes; petiole 0.3—0.8(-1.1)

cm long, densely tannish-white tomentose, arising from a narrowly elongate triangular petiole base, 2-3.5

mm long, 1.5-2 mm wide, tomentose and minutely villous abaxially, glabrous and tannish adaxially; blade

broadly elliptic to ovate, 0.7—1.5 cm long, 0.4-1 cm wide, tannish-white lanate abaxially, thinly tomentose

to glabrate or glabrous and greenish to light olive-green adaxially especially with age, with a broadly obtuse

to rounded base not tapering to the petiole, an entire, plane margin, and a mostly obtuse to broadly obtuse

apex; midvein slightly raised, obscured by tomentum. Aerial flowering stems erect to slightly spread-

ing, 2-8 cm long, villous, bractless. Inflorescences compound umbellate, slightly open, mostly 1-2 cm

long, 1-3 cm across; branches villous, grayish to greenish white, not drying blackish; bracts 4-6 at base

of inflorescence, spreading, sessile or nearly so with elliptic blades, 4-10 mm long, (1.5-)2-4 mm wide,

usually white tomentose abaxially, thinly tomentose and slightly greenish adaxially; centrally positioned

peduncle 5-10 mm long, villous; lateral branches 3-8 mm long, these terminated by a whorl of 4-6 linear

to narrowly oblanceolate bracts, 2-8 mm long, 0.7-1.2 mm wide; peduncles atop lateral branches 2-5C-7)

J. Bot. Res. Inst. Texas 3(2): 479 — 483. 2009

[| ful Dos b In LE "EC di nio

480 J t f Texas 3(2)

mm long, villous. Involucres solitary, broadly turbinate, 4—6.5 mm long, 3.5—5.5(-6) mm wide, densely

villous abaxially, glabrous adaxially; teeth 6-8, erect, acute (when long) to obtuse or even rounded apically,

0.3-0.6(-1) mm long Pedicels erect to slightly curved with age, 4—6.5 mm long, glabrous; bractlets linear,

4-6(-6.5) mm long, densely villous. Flowers bright yellow at early anthesis, rapidly becoming fused with

red and ultimately red to reddish maroon with an undertone of yellow, 2-3 mm long when yellow becom-

ing 4.5—6 mm at maturity on a 0.1-0.3 mm long stipe, glabrous; tepals monomorphic, obovate; stamens

slightly exserted, with 2-4 mm long, slightly pilose basally filaments, and oval, yellow, 0.4-0.6 mm long

anthers; pistil with styles 1-1.3(-1.5) mm long. Achenes light yellowish brown, narrowly trigonous, 4.5—6

mm long, glabrous; embryo straight.

Distribution.—Known only from serpentine talus slopes and rock crevices at The Cedars, 1200-1800

ft elev, Sonoma Co., California (Fig. 3). Jun-Sep.

Additional collections EE U.S.A. CN Sonoma Co.: Layton Mine, Austin Creek, 29 Sep 1947, Hoffman 507 (CAS,

RSA, UC, UTC); on King R , The Cedars, 5 elev., 23 Jun 1984, Raiche 43234 (CAS); near

Layton Mine N of Cazadero, The Cedars, 370 m Ja 10 Aug 1985, Raiche 50716 ee

Eriogonum cedrorum (from the Latin cedrorus, of cedars) is a member of subg. Oligogonum Nutt., a taxon typi-

fied by E. umbellatum Torr., that presently is composed of some 35 species of subshrubs and herbaceous

perennials that occur from Alaska to central Mexico, and from Virginia and West Virginia to the Pacific

Coast. The new species belongs to a subgroup characterized by an involucre with erect teeth and smallish

leaves arranged into rosettes that result is low, spreading mats. Therefore, within the subfamily The Cedars

wild buckwheat may be quickly distinguished by a whorl of bracts immediately below the inflorescence,

glabrous flowers on short stipes that are initially yellow but rapidly change to a red or reddish-maroon, a

synoecious habit, villous involucres, and leaf-blades that are lanate abaxially. Based on the yellow flower

color, E. cedrorum is allied to E. ternatum, but that species differs in its compact mat of densely arranged

rosettes of leaves terminated by long (10-30 cm long) flowering stems that bear an umbellate inflorescence

with sulphur-yellow flowers that remain distinctly yellow even at maturity. Eriogonum cedrorum differs from

the more closely allied E. nervulosum of nearby Colusa, Glenn, and Lake counties, California, by its yellow

(not white) flowers, and while both quickly become reddish, the yellow and white undertone is retained in

each species. Both of these species occur on serpentine but E. nervulosum tends to be more compact than

E. cedrorum. Both species have relatively short aerial flowering stems and similar leaves, but they differ in

lengths of their involucres (4-6.5 mm in E. cedrorum versus 3-4 mm in E. nervulosum) and in the length

of their stipes (0.1-0.3 mm versus 0.5-0.8 mm, respectively). Also individual plants of E. cedrorum tend

be larger than those of E. nervulosum, but this may be the result of the loose talus slopes where E. cedrorum

is found rather than any significant genetically-fixed morphological difference. Both species differ from E.

ternatum in having compound umbellate inflorescences (Fig. 2), with that of E. cedrorum more open (and

thus more obviously branched) than that of E. nervulosum, which often appears to be subumbellate.

The Cedars wild buckwheat joins a small number of other relatively restricted species found within

the more or less 7000 acres that defines The wei ae area is an Eos block of ultramafic mantle rock

(Harzburgite), and its derived soils, which is only f he moist, fog-dominated coastal climate

influenced by the Pacific Ocean; nonetheless, the area is dominated by a hot dry summer more typical of the

interior California climates well to the east. Six other plant species are endemic here: Arctostaphylos bakeri

Eastw. ssp. sublaevis PV. Wells, Calochortus raichei Farwig & Girard, Epipactis gigantea Douglas ex Hook. f.

rubrifolia P.M. Br., Erigeron serpentinus G.L. Nesom, Streptanthus glandulosus Hook. ssp. hoffmanii (Kruckeb.)

M. Mayer & DM. Taylor, and S. morrisonii F.W. Hoffm. ssp. hirtiflorus F.W. Hoffm. In addition we are aware

of an undescribed Holodiscus taxon.

While The Cedars terrain is dominated by Sargent cypress woodland (Hesperocyparis sargentii Jeps.)

Bartel) and chaparral, the extremely steep yen Spe: are mostly nn ies and talus forming extensive

serpentine barrens. This open rock and tal ly place that Eriogonu rum grows, and only a small

portion of this habitat ble for this wild buckwheat. The ere of plants grow in loose gravelly

tay

A a p

. ii

| th

Sm. * *

Fic. 2. Eriogonum cedrorum, details of inflorescences.

Fic. 1. Eriogonum cedrorum, habit.

482 lof the B IR h Institute of Texas 3(2)

Tombs Creek Quadrangle Warm Springs Dam Quadrangle

tain Ridge $1 a > 14 A F, 5

* j Er

fi 25 am. 7 J - >

FT “ A ; lx j 2. A i Zá . S

Fort Ross Quadrangle 127 07 30 Cazadero Quadrangle

Purple shading shows general outcrop of peridotite and serpentine

0 ann |

Base map courtesy of i ee ———-

Dr. Robert Coleman, Scale

Stanford University. . 683 Elevation in meters. Major Faults =

Fi 2K lictriburti f Eri g l t The Cedars, Sonoma Co., California

talus but a smaller number find bare rock crevices suitable; however, those on the talus are typically larger

mats. There are roughly three zones where it occurs (Fig. 3); (1) the northwest part of The Cedars in upper

Danfield Creek, (2) the central upper canyons of Big Austin Creek which contains the largest population,

and (3) the east side near Red Slide above East Austin Creek which also harbors a large population. There

seems to be no clear reason why this taxon is so restricted to these sites while not occurring nearby. While

slopes with N-facing aspect account for the majority of the plants, some are on E-, 5- and W-facing slopes.

There are +1500 to 2000 plants in existence. A recent survey of the Central Canyon sites shows the

population to be extremely stable. While there were few small young plants, not a single dead mat was

noted. A few plants had their crowns elevated >1.5dm above the current rock surface, showing they had

undergone that much erosion and survived. Erosion of the substrate is constant but not rapid. The lack of

Reveal and Raiche, Eric — pecies from Californi 483

any significant human disturbance at any site, the lack of senescence or death, and the persistence of plants

in MPs. Hun sites je dil Mus taxon is ERE of great age.

ypica Woodson, Aspidotis densa (Brack.) Lellinger, Cardamine

ca lifomica (Nutt.) Greene var. sinuata (Greene) O.E. Schulz, Epilobium minutum Lindl. ex Lehm., Eriogonum

luteolum Greene, E. nudum Douglas ex Benth. var. auriculatum (Benth.) J.P. Tracy ex Jeps., Hesperolinon

spergulinum (A. Gray) Small, Minuartia douglasii (Fenzl ex Torr. & A. Gray) Mattf., Phacelia corymbosa Jeps.,

Sairocarpus vexillocalyculatus (Kellogg) D.A. Sutton, Streptanthus morrisonii, and S. barbiger Greene. Allium

falcifolium Hook. & Arn., Eriophyllum lanatum (Pursh) Forbes, and Eschscholzia caespitosa Benth. are more

restricted to certain sites. The Holodiscus taxon mentioned above is the only hard-woody plant to occur

within populations of E. cedrorum, but Arctostaphylos bakeri ssp. sublaevis and A. viscida Parry ssp. pulchella

(Howell) PV. Wells may occur peripherally.

ACKNOWLEDGMENTS

We wish to thank Dan Segal of the Plantsmen Nursery of Ithaca who brought us together with the idea that

The Cedars wild buckwheat was “interesting.” We also wish to thank Mrs. Bette Campbell of Cazadero for

kind assistance not only to us but to a wealth of scientists over the years who have visited The Cedars. We

thank Robert Coleman, Professor Emeritus of Geology at Stanford University, Palo Alto, California, for his

geological map of The Cedars area that served as a base map for the plant’s distribution, and Orion Johnson,

a doctoral candidate at the University of Southern California, for help with the graphics. Guy Nesom kindly

provided a prompt and useful review of the manuscript for which we are grateful.

REFERENCE

REVEAL, J.L. 2005. 44a. Polygonaceae Jussi bfam. E id Arnott, Encycl. Brittannica (ed. 7), 5:126. 1832

e Wild buckwheat subfamily. Fl. N. Amer. 5: 218-478,

484 tani itute of Texas 3(2)

BOOK REVIEW

Jim Enperssy. 2008. Imperial Nature: Joseph Hooker and the Practices of Victorian Science. (ISBN 978-

0-226-20791-9, hbk.). The University of Chicago Press, 1427 East 60th Street, Chicago, Illinois 60637,

U.S.A. (Orders: www.press.uchicago.edu, 1-773-702-7000, 1-773-702-9756 fax). $35.00, 429 pp., 49

halftones, 1 line drawing, 6" x 9".

n Jim E 7 1 1 m at Joseph LT 1 E] 1 > 1 : Ton lx 19 Oth- -Cer L = ] 41 D E E 14 1 Pa

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J. Bot. Res. Inst. Texas 3(2): 484. 2009

A NEW NARROWLY ENDEMIC SPECIES OF DIRCA (THYMELAEACEAE)

FROM KANSAS AND ARKANSAS, WITH A PHYLOGENETIC OVERVIEW

AND TAXONOMIC SYNOPSIS OF THE GENUS

Aaron J. Floden', Mark H. Mayfield, and Carolyn J. Ferguson

is al and Division of Biology

as State University

TES pnm 66506-4901 U.S.A.

ABSTRACT

Dirca decipiens A. Floden, sp. nov., is d ibed f tern K ] ] Arkansas, and ised t

and key to al ae is pou eee a new pee is eae distinguished from D. puo its nearest a cine by its

iall D. mexicana and D

abaxially F r L os

15 a H Il ¿l +l ME A Pear A E J C4} 1 ] YEclmnz RL el

r i J F D/ E T

highligl its distincti f D palustris Dd lated ] habi 1] £1 1

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r e

spacer ena intron regions (trnD-trnT ae trnL- trnP). The vlslogenstie data support 1) the monophyly of the widespread eastern North

American D. palustris, 2) a cl D. mexicana and D. decipiens, and 3) a well ] hvl Fr]

of the western D. occidentalis from the

three eastern species that is sister to the Californian ne D. occidentalis. Early d

o Ii i a ao E i

RESUMEN

Di d A. Floden, nueva especie, es d ta de K yel de Arlan € — sro

rina clave nara el T a TM 22 1 dJioti 17] FA^] t Pa e T) I f h

) p género palustris, su vecina geográl ica más cercana, por sus xs

pubescentes en el lado abaxial, y flores : frutos "sg y de D. mexicana y D. occid Į g I y su posesión de

tricomas en el ápice del ovario y el fruto. Un ip de la Mp de el hábitat de | pecie clarifi ás la disti

ELK y D. palustris: D. deci] p liza, y empieza floreciendo aproximad

i D. palustris. I ] fil sti del vé fueron Inadas t ] A

i [e]

secuencia "de ADN (ITS; y de] cloroplasto , trnD- -trnT y trnL- trnF) Ect dar tent 131 filia de 1

rango en el este de Moftesimeden D. vakisiris 2) una e cercana entre D. mexicana y D. Pan y 3) un grupo monefiletieo Big

apoyado de p orientales q de hermana con la esp de California, D. ide La

li iat de | i idental, D. occidentalis, de las d i tád d j

o In ’ , E E

anteriores.

Key Wonps: Arkansas, Dirca, decipiens, Kansas, phylogeny, Thymelaeaceae

The genus Dirca L. is the DER extant native North American temperat t within the largely tropical fam-

ily Thymelaeaceae. This g t recently considered to include tl imilar but geographically

isolated species of North. American shrubs Gean & Mayfield 1995). Dirca palustris L. occurs infrequently

over most of eastern North America (from southern Ontario and northern Florida west to eastern North

Dakota, eastern Oklahoma, and central Louisiana). Dirca occidentalis A. Gray and D. mexicana G.L. Nesom

& M. Mayfield are both narrowly endemic species, the former occurring locally in the San Francisco Bay

and Central Coast floristic provinces of California (Berman 1993; Schrader & Graves 2008) and the latter

at a single locality in montane northeastern Tamaulipas, Mexico (Nesom & Mayfield 1995; Graves 2008).

In the present study, we describe a fourth species of Dirca from Kansas and Arkansas populations that have

heretofore been identified as the broadly distributed eastern species D. palustris.

Morphological taxonomic work on the genus Dirca has been extensive despite the small size of the

tadd Herbari iD t t of Ecol 1 Evoluti Biology, Uni itv of T , Knoxville, Tennessee, 37996-1610,

U.S.A. (afloden@utk.edu)

J. Bot. Res. Inst. Texas 3(2): 485 — 499. 2009

486 t tani titute of Texas 3(2)

genus. Following a period of lcit towards taxonomic recognition of D. occidentalis, Vogelmann

(1953) presented a detailed morphological comparison between D. palustris and D. occidentalis in which he

demonstrated the taxonomic utility of foliar and stem vestiture, perianth morphology, position of filament

insertion, and pedicel elongation for distinguishing these species. Vogelmann also suggested there was an

overall leaf size distinction between these species but stated that additional study would be necessary to

conclusively document those differences. Subsequent treatments have once M arr in br dp

both of these species. M tly, Nesom and Mayfield (1995) employ f characters

when they compared a newly discovered species—D. mexicana—to the other species in the genus. In the

present study we additionally assessed ovary vestiture as a taxonomic character within the genus, though

there has been little discussion regarding presence/absence of trichomes associated with the gynoecium

(Small [1903], however, noted the ovary is glabrous in D. palustris).

In 1997, AJF first noticed an unusual population of Dirca in Johnson County, Kansas, around the time

of its iee pe Meat discovery and documentation by Freeman et al. (1998). In the latter, a presentation of

lation as D. palustris without further

new K , Freeman et al. identified the single K

consideration of its taxonomic position within the genus. ce of living plants of both D. palustris

and the Kansas pean in a field and under cultivation over several years led AJF to pursue a more

thorough dy of tl hology. Through the course of herbarium and field studies, at least

two anal ARE from Midas an morphological and life history traits similar to the Kansas

plants were discovered. Here we present and develop these findings, which support the conclusion that

these populations represent a unique species, herein described as D. decipiens A. Floden. We also present a

taxonomic synopsis of the genus and phylogenetic data on relationships among the four Dirca species.

MATERIALS AND METHODS

Study of herbarium specimens.—Morphological comparisons were made between D. decipiens and the

other three species of Dirca. A total of 561 specimens were examined (D. decipiens, 67 specimens; D. mexi-

cana, 9; D. occidentalis, 28; D. palustris, 457). This included material from the three known populations of

D. decipiens, material from the single known population of D. mexicana, specimens of D. occidentalis from

all of the six counties from which it is reported, and specimens of D. palustris from across its broad range.

A subset of specimens representing mature plants (collected six or more weeks after anthesis) was selected

(D. decipiens, n = 12; D. mexicana, n = 5; D. occidentalis, n = 4; D. palustris, n = 342), and morphological mea-

surements were made with emphasis on characters discussed in the literature (Small 1903; Holm 1921;

McMinn & Forderhase 1935; Vogelmann 1953; Nesom & Mayfield 1995). Morphology assessed included

the following: density and color of pubescence on twigs, leaves, involucres, ovaries, fruiting pedicel, and

fruit; leaf length and width, shape of the blade and number and length of cilia on the margins (all based

on the two uppermost mature leaves of each branch, as per Vogelmann [1953]); involucral bract shape and

size; calyx tube and limb lengths, lobing and margin of the limb, and position of filament insertion; pres-

ence of peduncle (vs. sessile fruits), and fruit length and color. Exsiccatae were used in the study of floral

characters, as possible: flowers of all species except D. mexicana were re-hydrated, dissected, and opened

to enable morphological assessment. In addition, fresh flowers from the Kansas population of D. decipiens

were studied, and flowers of D. decipiens and D. palustris from Arkansas were field collected in ethanol for

later observation. Limited material of D. mexicana was available (see Nesom & Mayfield 1995); however,

photos of flower dissections previously made by MHM were examined and revealed a lack of trichomes on

the ovary apex (a character directly observable on specimens at the fruiting stage).

Fieldwork.—Fieldwork was poneucied | in the spring of 20 06 in Kansas, and in spring 2007 in Arkansas,

Kansas, and Missouri in order to allow si n of flowering material of typical D. palustris

in close geographical proximity to D. decipiens. The two extant populations of D. decipiens in northwestern

Arkansas were compared in terms of morphology and phenology to D. palustris in close geographical prox-

imity (< 25 km by air).

Floden et al., A ies of Di ] | is of tl 487

Phylogenetic data.—Eleven samples of Dirca were selected for DNA seq

relationships among members the genus: a sample from each of seven populations of the side: "o D.

palustris, two Pa of D. decipiens (one each from Kansas and Arkansas), and one each of the narrow

endemics D. mexica d D. occidentalis. A species of Daphnopsis Mart. was utilized as the outgroup for the

present study. While the relationship of Dirca to other members of the Thymelaeoideae is not well resolved

(Van der Bank et al. 2002), Daphnopsis is a New World group in which the sequence regions considered in

this study are readily alignable with Dirca. All voucher information is presented in Table 1.

Total DNA was extracted from silica-dried leaves or from material from herbarium specimens using

a DNeasy Plant Mini Kit (Qiagen, Valencia, CA) and stored at -20°C. The internal transcribed spacer (ITS)

region of the nuclear ribosomal DNA (nrDNA) was employed in this study, largely because of ease of use

but also to enable comparison with a previous study in Dirca (Schrader & Graves 2004). Following prelimi-

nary study of several non-coding chloroplast DNA (cpDNA) sequence regions for their utility in phylogeny

reconstruction for Dirca, two regions were selected for use: the trnD°S-trnT°°" intergenic spacer region,

hereafter trnD-trnT (see Demesure et al. 1995), and trnLV^^-trnLV^^-trnF9^^. hereafter trnL-trnF (see Taberlet

et al. 1991). PCR and sequencing conditions followed those described by Ferguson et al. (2008; for ITS,

the forward modified primer ITS5 of Downie and Katz-Downie [1996] and the reverse ITS4 of White et al.

[1990] were used, and the annealing temperature was 50°C)

Both forward and reverse strands were obtained for each product, and sequences were edited using

Sequencher 4.5 (Gene Codes Corp., Ann Arbor, MI) and aligned manually with the aid of Se-Al (Rambaut

2002). Gaps were coded as a “fifth rae for insertion-deletion (indel) events longer a one bp. al DEED one

of the affected cl l f Parsimony analy

searches in PAUP* 4.0b10 (Swofford 2002), with all characters weighted equally. Support for branches was

evaluated by bootstrapping (Felsenstein 1985) using full heuristic Sena with 10,000 replicates. A com-

bined data set (ITS, trnD-trnT, trnL-trnF) was constructed and an i length difference (ILD) test

(Farris et al. 1995) was conducted to test for homogeneity among the ITS versus cpDNA data (implemented

in PAUP*, the partition homogeneity test, branch and bound search). An analysis was then conducted on

the combined data following the approach described above.

RESULTS AND DISCUSSION

Dirca decipiens—Compared to other species.—Dirca decipiens was found to differ notably from D. palus-

tris, its closest geographical neighbor (see Taxonomic Treatment, below). One of the first and strongest clues

to its separation from the latter was its larger, sessile fruits, a feature it shares with the other two species,

D. occidentalis and D. mexicana. As a group, these three sessile-fruited species in a a ales a

abaxial leaf surf: tricl youngest twigs. However, D 1

species of the genus in having ovaries with apical trichomes (visible in both flower añ fruit) and pubescent

pedicels (Fig. 1; Table 2). A suite of quantitative characters further differentiates D. decipiens from the other

sessile-fruited endemics (e.g., compared to D. occidentalis and D. mexicana, D. decipiens has white abaxial leaf

pubescence of higher density, generally larger leaves, and distinctive perianth traits; Table 2).

Phylogeny. cun data set and tree statistics are po in Table 3, and trees resulting from

analyses of the indi data set ted in Figure 2. All phylogenetic data support a sister relation-

ship between the Californian — D. occidentalis and the —— of the genus, a finding consistent

with the phenograms of Schrader and Graves (2004). Sampled populations of D. palustris form a clade in

both cpDNA phylogenies, with D. decipiens and D. mexicana exhibiting an unresolved relationship relative

to D. palustris (trnD-trnT) or forming a clade sister to D. palustris (trnL-trnF; Fig. 2). However, the ITS tree

differs in that one sample of D. decipiens is placed in the D. palustris clade (with weak support, 6396 BS; Fig.

2). The ILD test was non-significant (P = 0.22); therefore an analysis of combined data was carried out. The

combined data set consisted of 3,004 bases of aligned sequence data with 43 parsimony informative char-

acters (Table 3). Missing data accounted for 3.796 of the combined matrix (Table 3), and most of this (3.696)

was due to a single missing sequence (trnD-trnT for D. palustris LA).

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489

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TABLE 2. Comp |

are given in parentheses.

D. occidentalis D. mexicana D. palustris D. decipiens

Carriage sessile-1 mm sessile-1 mm pedunculate, 7-20 mm sessile-1 mm

Calyx

length 8-10 8-10 5-8 7-11

tube 2-4 5-6 2-4 5-7

limb lobed lobed unlobed lobed

sinus depth 1-3 0.5-2 N/A 1-3

limb width 3-4 2-4 ].5-3 2.5-4

limb margin entire undulate crenulate erose

Filament insertion proximal distal distal distal

Ovary/fruit glabrous glabrous glabrous pubescent apically

Fruit length 7-8 [7.4] 7-10 [8.6] 7-9 [8.1] 8-9 [8.25]

Involucre

pubescence white white brown white

shape obovate-oblong elliptic-obovate elliptic obovate-oblong

apex roun acute acute

immature size 4-46 x 6-12 7-2 x 3-1 1-17 x 6-8 6-18 x 5-8

mature size 4—6 x 6-12 4-6 x 6-12 2-35 x 8-15 18-5 x 8-24

Stem

indumentum pubescent pubescent glabrous pubescent

thickness 2-4 2-4 — 3-6

internode 24 26 17 28

eaves

cilia per mm 6-9 8-12 0-3 (3-8) 11-18

cilia length 0.2-0.5 (1) 0.5-1.2 0.2-0.8 0.5-2

adaxial pubescent glabrous glabrous pubescent

abaxial pubescent pubescent glabrous (pubescent) jee

length 40-70 40-80 45-100 75-90

width 25-45 25-60 45-65 45-60

Figure 3 presents the strict consensus of two most parsimonious trees resulting from analysis of the

combined (ITS, trnD-trnT, trnL-trnF) data. The sister relationship between D. occidentalis and the remainder

of the genus is strongly supported (100% BS). All samples of D. palustris form a moderately well supported

clade (87% BS; within D. palustris, the samples from FL and NJ group with 99% BS), and D. decipiens and D.

mexicana form a models in AU ponen ap Ee BS, with D. mexicana nested within D. decipiens, sug-

gesting insufficient ti o result in reciprocal monophyly). The sequence data for sampled

populations of D. palustris exhibited variation in all three sequence regions studied, and phylogeographic

and/or population genetic studies of this wide- angie peces Hay be informative. With regard to the new

species, it is noteworthy that D. palustris AR is roximate to D. decipiens AR (« 25 km by air).

The present study advances our snderstandine of rvioseny of this poorly known genus. Schrader

and Graves (2004) presented phenograms based on ITS sequences and inter simple sequence repeat (ISSR)

data for Dirca. As in the present study, they inferred a sister relationship between D. occidentalis and the

remainder of the genus. The addition of data from the chloroplast and from additional samples (including

lations of D. palustris from different parts of its range) along with cladistic analysis

the new species and [

further refine our understanding of the phylogeny. We suggest Daphnopsis as an appropriate outgroup for

future studies of Dirca (although Daphne has been utilized with some success by earlier workers, we were

unable to align Daphne sequences with confidence to Dirca).

It is intriguing to consider morphology (Table 2) in light of the strict consensus tree (Fig. 3). Many of

491

Floden et al

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Table 3. Data set and tree statistics for the phylogeneti ly

Data set ITS trnD-trnT trnL-trnF Combined

# samples 12 11 12 12*

Sequence lengths 700-709 1260-1273 916-995 =

Aligned length 717 1287 1000 3004

% missing data 0.4% 0 0 3,7%*

# indels; no. characters excluded 17713 7,32 12; 80 36; 125

# variable (included) Enatactels: 125 53 41 219

# parsimony INTOrmaltive {I !) characters 25 9 10 46

most parsimonious trees 2 2 1 2

MP tree length 135 54 43 237

Cl 0.99 0.98 0.98 0.96

Cl excluding uninformative characters 0.93 0.90 0.91 0.84

RI 0.93 0.93 0.96 0.87

*One trnD-trnT sequence was unobtainable (D. palustris LA), and this missing sequence accounts for the vast majority of

missing data in the combined matrix.

the characters shared by the three rare taxa but not exhibited by D. palustris can be viewed as pleisiomor-

phic: more rounded leaves; abundant white pubescence on the twigs, abaxial leaf surfaces and involucres;

a perianth with a more definite lobed limb; and sessile to sub-sessile fruits. The hoary, persistent, abaxial

leaf pubescence, especially along the veins, has been previously noted as a characteristic of D. mexicana and

D. occidentalis (Vogelmann 1953; McMinn & Forderhase 1935; Nesom & Mayfield 1995). Dirca decipiens

is similar in this regard but the trichomes are more evenly distributed over the entire abaxial leat surface.

Dirca palustris is usually completely glabrous abaxially; however, we noted rare individuals that exhibit

sparse, brown pubescence on the abaxial leaf surfaces and stem (see Appendix 1). Of note, the sample of D.

palustris from Michigan (Table 1) exhibited this slight, brown pubescence, yet it grouped, as expected, with

other samples of D. palustris. Several characteristic features of the widespread D. palustris are autapomor-

phic: elliptic-obovate leaves with cuneate bases and an acute to obtuse apex; dark brown/bronze pubescent

elliptic-obovate involucres with an acute apex; an unlobed, crenate perianth margin; and pedunculate

inflorescences (in fruit, both the peduncle and pedicels elongate).

Biogeographical patterns.—It is likely that the present day distribution of the genus Dirca is the

result of range contraction in the family following a period of expansion in the Eocene and early Miocene

periods when global cooling allowed elements of a widespread temperate forest to expand across the North

American continent (Graham 1993; see Nesom & Mayfield 1995). Biogeographical affinities within much

of the Thymelioidaeae remain to be discerned (Van der Bank et al. 2002), and our data are equivocal with

regard to resolving the origin of the genus. Whatever accounts for the initial invasion of Dirca across North

America, the drying of western North America and the uplift of the Rocky Mountains surely played a role in

isolating D. occidentalis from eastern elements ol Dirca, which likely remained in long-term contact during

f contact during wet, cooler periods of the Pleistocene.

the late Miocene followed b

This is shown by marked es benseen D. occidentalis and the other three eastern species (Fig. 3; see

also discussion by Graves & Schrader 2008). Mid-Miocene conditions may have contributed to the migra-

tion of an ancestral stock along the western edge of the Ozark Plateau southward into eastern Mexico (or

vice versa) and potep Hay eastward through the northeastern Ozark Plateau (see Graham 1993). During

this time, cooling t d increased aridity provided the conditions responsible for expansion of

grasslands, which would have resulted in the isolation of Dirca into central riparian and eastern deciduous

woodlands ond) ee Takhtajan ee ien EE

1 1 1 241] 1

Current distri patterns may | | expansion of the Great Plains

and the glacial maxima. Increasing aridity in Texas and Tuna during interglacial periods would have

z £n: l y | e £ «l g 493

Floden et al : n

Dirca decipiens KS

D. mexicana

D. decipiens AR

D. palustris MO

100

— [) palustris TN

——4— D. palustris AR

— |). palustris MI

D. palustris LA

D. palustris NJ

[ 99 |

D. palustris FL

D. occidentalis

Daphnopsis

philippiana

bined (ITS, trnD-trnT, trnL-trnF) data. Num!

Fi 2 Strict Ë in

are bootstrap values > 50.

494 t tani itute of Texas 3(2)

severed the connection between the Ozarks and the forests of northeastern Mexico (see Takhtajan 1986),

isolating D. mexicana. Dirca mexicana occurs in an area known for its concentration of relict species, which

are uns with wider distril inthe t aed ae E g., il ala americana, ue ide ovata,

urbiniana, H. li i Illi

J r El

and T, dd 1] present in ] I : yto en ; lau NI & Mayfield 1995:

see on 2008 for ecological habitat a for D. DS o the eastern United States,

the present day distribution of D. palustris and its floristic associations suggest that it is a Pleistocene relict.

At the southern periphery of its distribution, D. palustris is rare and present mostly along Pleistocene river

systems-the Rn Chattal hee, Flint, and Apalachicola (Delcourt & Delcourt Pu Dirca palustris,

bit tum, Magnolia ashei, M li Paci bens, Phlox

c+ z - L

J 4 Dl: ml... il os Sud ss

A quilegi rl rr

A x ,and Sangu 10 701Y1 7H

aana Podophyllum peltatum, T

relicts that have greater distributions further north and a need distribution in the Marianna lowlands, Red

Hills of Georgia, and the Apalachicola bluffs (Delcourt & Delcourt 1975; James 1961; Kurz 1928; Mitchell

1963; Thorne 1954).

A new species in the central United States.—This study reports the unusual biological discovery

of a distinctive undescribed shrubby vascular plant species from a relatively well-studied floristic region.

Although two of its three known populations had peen previously collected by others, D. decipiens had

escaped taxonomic recognition despite recent t attention. Several factors likely contributed to the

failure of earlier workers to recognize D. decipiens, including 1) its extreme rarity (apparently even prior to

extensive habitat fragmentation in the region), 2) limited collecting of early spring flora, resulting in very

few reproductive specimens for botanical study, and 3) a lack of close observation of living material and

exsiccatae by previous workers, along with a difficulty in discerning floral characters on dried specimens.

This discovery underscores the importance of continued biological collecting and study of museum collec-

tions, even in relatively well-studied regions, so that we may be able to better appreciate biodiversity (see

Prather et al. 2004).

TAXONOMIC TREATMENT AND KEY TO SPECIES OF DIRCA

Dirca L., Sp. Pl. 1:358. 1753. Leatherwood. [From the Greek, Dirce, wife of Lycus. Dirce was a follower of

Dionysus who caused a ps to flow at the site of her death.]

Infl , projecting out of the i bracts; adaxial surf f involucral

“bracts with dark brown, rarely light I lobed, th gi te, undulate, or erose;

leaves usually gl , rarely OMENAN ntc pubescent D. palustris

. " FE el $ I I | 4 a |

ducral Bie with white to light tan tomentum; Ea mostly 4-lobed, the margins entire to crenate; ies

ubescer it LU UVP o O E SG

2. Apex of ovary sud fruit lacking trichomes; leaf margins with 12 or less cilia/mm

3. Calyx tube (2-4 mm) shorter than the gradually flaring limb ee E mm), Marnen noenten prona

stems reddish brown; adaxial surface g the veins; lea

margins with 6-9 cilia/mm; loca! ir in California (San Francisco Bay area) D. occidentalis

3. Calyx tube (5-7 mm) longer flaring limb (2-4 mm), filaments inserted distally; stems

gray-brown; adaxial surface of the proximal half of the leaves glabrous, or rarely the lower thir

puberuient along the veins; leaf margins with 8-12 cilia/mm; rare and local in Tamaulipas, Mexico

(mountains) D. mexicana

2. Apex of ovary and fruit with a few trichomes; leaf margins with 11-18 cilia/mm; rare in Kansas and Arkansas

(and possibly also Missouri) D. decipiens

Dirca decipiens Á. Floden, Sp. nov. (Fig. 1). Tree: U.S.A. Kansas. Jol Co.: Overland Park Arboretum, S side of Wolf Creek

on bluff, under Quercus sp., Cercis canadensis, Carya ovata, with Staphylea, Asimina, Corylus, Erythronium albidum, Maianthemum,

Polygonatum, Arisaema, Aquilegia, 2 Apr 2007, A.J.Floden, L. Nevling & M. Mayfield 162 (notorvee: KSC; isorYpEs: BRIT, KANU, MO,

NEB, NY, OKL, TENN, TEX, UARK, US).

Thi | ee | ; ca ee UE 4 1; wallie F 1 1:5 Joerg

Floden et al., A ies of Di ] is of tl 495

Soft woody shrub 2-3 x 2 m, bark tan to grayish, current growth 3—5 mm thick and flexuous, bearing

persistent appressed white pubescence, sparse lenticels. Leaves: alternate, obovate to elliptic-oblong, 8 x 5

cm, base round, apex round, margins entire, ciliate (11-18 / mm,1-2 mm long), persistent heavily sericeous

pubescent on abaxial lamina and veins, mostly glabrous above, pubescent on primary veins below mid-

lamina, petiole to 4 mm long, heavily sericeous, deciduous. Involucres: obovate-oblong, cuneate base and

rounded apex, first two deciduous, second two persistent and becoming leaf-like, abaxially hoary to light

brown, adaxial primary veins bearing whitish sericeous hair. Flowers: concurrently with the leaves, sessile

to sub-sessile (if so, borne on individual pedicels < 1 mm, pubescent), deflexed fascicles of 3(2—6), calyx

greenish to pale yellow; narrowly cylindrical-campanulate 7-11mm long (9 mm avg.), tube 4-8 mm, limb

3-4 mm, flared, 2.5~4 mm wide, calyx limb most often divided into 4 distinct but uneven lobes 1-3 mm

sinus (avg. 2 mm), occasionally unlobed, margin erose. Stamens: 8, distally inserted at the throat at one level,

extending unequally 3-5mm beyond the calyx limb margin, pollen yellow. Style: 12-16 mm long, extending

past stamens, stigma capitate, extending 1—5(-8) mm beyond the stamens. Ovary: ovoid, 2-3 mm x Imm,

10—17 trichomes to 0.5 mm in length at the apex of ovary. Fruit: a one seeded drupe, sessile to subsessile,

pyriform to ovoid, green, yellowing at apices at maturity, (drying reddish-brown), 8-11 mm long, 4-5 mm

wide (avg. 9 x 5 mm), trichomes present on apex.

Phenology.—Flowering before the canopy has begun to leaf out in late March to early April. Dirca

palustris in adjacent populations in Arkansas begins flowering approximately one week before D. decipiens.

Fruits ripen sequentially from terminus of branch down over a few days, 8-10 weeks after anthesis.

Distribution.—Dirca decipiens is presently known from three non-contiguous locations all on northeast

facing bluffs and slopes above rivers within the Appalachian Region, as classified by Takhtajan (1986). The

Johnson County, Kansas, population occurs on the eastern boundary of the bluestem prairie where Ozarkian

woodland elements are present in mesic valleys along riparian systems. Large prairie expanses separate D.

decipiens in Kansas from the nearest known population of Dirca—a site a D. palustris in Benton County,

of D. decipiens. Many eastern

Missouri. Glaci íi maxima reached the area just north of the K |

] 1 with the K lati 1 reach their western limits in this

]

deciduous ¥

I L i

region (e.g. Panax quinql efol ius , Aralia racemosa, Act ; Sanguin aria canaden sis, Trillium sessile, and

tł t Arkansas in the

of D dec ipiens alt in

Geranium M HAN The two other known p

Ozark Highlands in Carroll County, both les: dian 25 km distant from the nearest D. palustris populations.

One might seek additional populations of D. ea in Sasa Missouri, extreme eastern Kansas,

ly assigned to D. decipiens

and northwestern Arkansas. In fact, t

based on vegetative features (see below); further fieldwork will be necessary to obtain reproductive material

and definitively confirm D. decipiens from Missouri. Furthermore, suitable habitat in northwest Arkansas

should be explored to determine whether the two known Arkansas populations may be contiguous.

Conservation.—The only known Kansas population, which is also the largest population, is located

on public property designated as parkland. This population is healthy and composed of several thousand

plants, both mature and abundant immature recruits. Both Arkansas populations are smaller in size, and

both are on property that was for sale as of March, 2007. The locality near Elk Ranch seems most threatened

by development due to nearby tourist destinations. The population is scattered along a rocky roadside bluff

along Leatherwood Creek. One hundred to 150 plants estimated present at the site. Very few seedling

recruits were present. The population south of Metalton consists of approximately 40 individuals in a lim-

ited and rural area along a narrow gravel road. This area seems unlikely to be developed. Plants from the

Kansas population have been brought into cultivation by AJF and have been distributed to correspondents

in Europe and the United States and to one nursery in the United States.

Reproductive success.—Vegetative reproduction has not been documented in D. decipiens, although D.

mexicana, D. occidentalis, and D. palustris do reproduce by rhizomes (McMinn & Forderhase 1935; Graves

2004). No rigorous collection of data on seed germination in D. decipiens has been kept by AJF, but seed

propagation estimates for three years are near 9596 germination. In contrast, germination rates for the other

[| [| £ al Das - ID LJ PHI £T. il

496

species are surprisingly low: 4.6% in D. mexicana, 60% in D. occidentalis, and 39.5% in D. palustris (Schrader

& Graves 2005); and 61% in D. palustris (Del Tredici 1981). Methods of Del Tredici (1981) are effective for

D. decipiens (AJF, unpubl.). These include allowing the ripe fruit husk to rot for one to two weeks, wash-

ing the seeds clean, potting the seeds, and leaving them to stratify over the winter (with some immediate

germination, < 5%).

Ftymology.—The specific epithet decipiens, deceiving, was chosen because of Eie SUME misin-

terpretation of the morphology that defines this aban as ae ius its geogra[ y close ii MA It

4 “4 ric

could equally be viewed as deceiving in

D. mexicana and D. occidentalis.

1/A Thad * YC; ape eres + 1 1 1 J

[* are vegetative, ii "——— here is tentative): U.S.A. ARKANSAS. Carroll Co.: N of Bla Springs, Beav er/Elk RRA Área,

12 Apr 1899, W. a ees = May HA B.F. Bush 1525 (MO); arado e 22 -ap 1913, EJ. Palmer 4444 (MO); 13 May

1914, E.J. Palmer 5578 , Beaver, 27 Apr 1926, E 29 & 29 Apr 1926, EJ. Palmer 29837 (MO),

Leatherwood Creek, 19 Mar 1927, D. D 2700 (MO); 16 Apr 1928, D. Demaree 4785 (BRIT: Elks Ranch, 5 Apr 1942, D.M. Moore

420010 (TEX/LL); banks of White River, near Beaver, 23 Oct 1925, E J. Pal En oa of bonn i: near ni icu. a

1931, EJ. Palmer 39454 (MO); on Hwy. 187 dh

bluff top [N 36?27'57" W 93°45'32" det. by ACME Moa 2 0- WGS84], 22 Mar 2007, A. Floden 150 (MO, TENN, TEX, Medos East Fork

ry Creek, Metalton, S on Hwy 21, W on CR 524 about 3/4 mi [near N 36? 10' 15" W 93? 32! 51", det. by ACME Mapper 2.0 — '

22 Mar 2007, A. Floden 151 (KSC, UARK, MO). Independence Co.: beside Cave Creek N.E. of Chinn Springs, T14N, vna SEC2],

17 jun 1968, R.D. Thomas bio. class 8733* (BRIT). KANSAS. Johnson Co.: Overland Park Arboretum, 38 47'45"N, 94 41'30"W det by

topo map, T145 R24E Sec 25 SYSEM, Sec 36 NYNE, Jct US 69 & 169, 6 m S, 1 m W, 900—1000 ft, 26 Mar 2000, C.C. Freeman & J.A.

Freeman 14078 (KANU) and 8 May 1998, C.C. Freeman 10700 (KANU); 9 Sep 2005, A. Floden 1; 1 Apr 2006, A. Floden 24; 8 Apr 2006,

A. meee 30,23 icd e A. e» add 42; n M 2006, A. Floden 92; 20 May 2007, A. Floden 94. MISSOURI. Gasconade Co.: W-facing

43N R6W NE sec 5 and [T44N R6W] SE sec 32, 2 2d mi i E Perstuny, Upper pani of

rocky slopes, rare, 4 Sep 1951, , J.A. Steyermark, 72691* (MO). Oregon Co.: base of wooded li

“the Narrows,” W of Calm, 11 May 1935, J.A. Steyermark 18902* (MO).

pica” mexicana G. L dn > ah das Sida. n Mid s on Tree: MEXICO. TAMAULIPAS. a Hidalgo:

o obscuro, 2.0 mi NE of Los Caballos toward

TNR Ei Mimbre, 15. 0 road mi W of | | El Mimbre, ca 1800 m, 3 Mar 1995, G. Nesom, M. Mayfield, &

G. Anderson 7863 (HOLOTYPE: MEXU; IsoTYPES: BRIT! - KANUI, MO!, NY [internet Hue, TENN!, TEX!).

1 1 T 1 11 11 22 4 1: gl th

Shrub to 2 m tall, branches grev-brown, lightly sericeous.

ovate- BODEN 4—8 cm long, 2 = : cm wide, base and apex round, sana glabrous except near gee

abaxially led, petiole 2-4 mm. Flowers (1)-3—(6) in axillary and terminal

fascicles, deflexed, sessile to sub-sessile (if so, individually pedicellate, and glabrous), calyx clear yellow, tube

longer than limb, filaments distally inserted, distinctly 4-lobed, lobes flaring at a 45 degree angle. Fruits, a

drupe, ovoid, 8-10 mm long, 4-5 mm wide, yellowish on the apices.

Phenology.—Flowering early spring, late February—early March. Fruiting eight to ten weeks after an-

thesis, May—June.

Distribution.—Moist rocky slope in western Tamaulipas, Mexico, ca. 1800 m.

ntati ined: MEXICO. Tamautiras. Municipio Hidalgo: 3 Mar 1995, G. Nesom, M. Mayfield, & G. Anderson

7863 (BRIT, KANU, MO, NY, TENN, TEX).

Dirca occidentalis A. aes Proc. Amer. Acad. 8:631. 1878. Te: U.S.A. C Oakland Hills, 13 Mar 1869, A. Kellogg

z, selected by Nevling 1964: GH; DUPLICATE OF LECTOTYPE: NY [internet image!]).

Shrubs to 2 m tall; branches red-brown, lightly sericeous. Involucres hoary. Leaves obovate to broadly ovate,

4-7 cm long, 2.5-4.5 cm wide, base rounded, apex rounded, adaxially pubescent on primary vein to mid-

lamina, abaxially sparsely sericeous mostly on primary veins, petioles 2-6 mm long. Flowers 1-3(-6) in

axillary and terminal fascicles, deflexed, sessile to sub-sessile (if so, individually pedicellate, and glabrous),

calyx clear yellow, tube shorter than limb, distinctly 4 (3) lobed, filaments proximally inserted. Fruits, a

drupe, ovoid, 8-10 mm long, 4-5 mm wide, yellowish on the apices.

Floden et al., A ies of Di | is of tl 497

Phenology.—Flowering early spring, sometimes winter, November—March, mostly late February—March.

Fruiting eight to ten weeks after anthesis, (April)May—June.

Distribution —Rich moist slopes in Alameda, Contra Costa, Marin, San Mateo, Santa Clara, and Sonoma

counties around the San Francisco Bay in the range of the coastal fog belt.

Representative specimens examined: CALIFORNIA, Alameda Co.: flowers and fruit, 3 Feb 1920 and 8 Apr 1920, C.C. Hall 10998

(MO). Contra Costa Co.: 3 Feb 1920, C.C. Hall 10998 (MO). San Mateo Co.: 29 Mar 2007, R. Patterson & M. Meshriy 1984 (KSC); Apr

1951, AJ. Sharp (TENN); 15 Feb 1958, J.H. Thomas 6980 (TX/LL); 9 Mar 1964, L.S. Rose, 64004 (TX/LL); 20 Feb 1977, J.H. Thomas 18470

(MO); 7 Feb 1964, D.E. Breedlove 5916 (MO).

Dirca palustris L., Sp. Pl. 1:358. 1753. Tree: U.S.A. Vircinu: (Lectorvr, selected by Nevling 1964: LINN 501.1).

T1 f. T ] 1

Shrub to 3m tall; branches brown-gray, g ely pel sistently 1 pubescent ) pu bescent

Leaves obovate to ovate, 4.5—10 cm [ons 4.5-6. 5 cm wide, pubescent, glabrescent above, the base cuneate,

the apex acute or obtuse, the petiole 2-5 mm long. Flowers (2-)3(—6) in axillary and terminal fascicles,

pendent, peduncles 5-13 mm, pedicel 2-10 mm long, calyx yellow-green, tube slightly longer than limb,

crenulate, undulate, or erose, filaments distally inserted. Fruits, a drupe, ovoid, 8-12 mm long, 4-5 mm

wide, yellowish on the apices.

Phenology.—Flowering early spring, March—May, fruiting eight to ten weeks after anthesis, May-July.

Distribution.—Rich mesic slopes above streams, lowland along rivers, calcareous woodland; N.B., Ont.,

Que.; Ala., Conn., Del., Fla., Ga., Ill, Ind., Iowa, Ky., La., Mass., Md., Maine, Mich., Minn., Mo., N.C., N.D.,

N.H., NJ., N.Y., Ohio, Okla., Pa., R.I., S.C., Tenn., Va., Vt., Wis. Dirca palustris occurs infrequently within a

broad area mostly within the bounds of the Appalachian province of eastern North America but is widely

scattered and of local occurrence.

Common Names.—American mezereon, eastern leatherwood, harbinger of spring, leatherwood, moose-

wood, rope bark, e

tati ined: U.S.A. ALABAMA nid Co.: 27 Mar 1969, R. Kral, 34034 EUM Monroe Co.: 17 Apr 1983,

B. E Wofford & Z. Murrell 83-42 (TENN). ARKANSAS. Newt 22 Mar 2007, A. Floden 156 Floden 154 (KSC).

22 Mar 2007, A. Floden 153 (KSC). Stone Co.: 1 Jul 1969, E.M. Browne and E.T. B 69G4.8 (TENN). KENTUCKY. F

May 1879, R.H. niis 2451 RN cae pui 22 May 1965, E.M. Browne & E.T. Browne 10151 (TENN). LOUISIANA. Catahoula

Parish: M.H. Mayfield & C ARYLAND. Mont y Co.: 4 Apr 1920, S.F. Blake 8062 (TX/LL). MICHIGAN.

Ingham Co.: 20 Apr 1963, L.C. Anderson 2228 eee MINNESOTA. Becker Co.: 6 May 1986, V. McNeilus (TENN). MISSISSIPPI.

Marion Co.: 1 Apr 1984, S. A 9 (TENN). Wayne Co.: 16 Mar 1975, K.E. Rogers 9428-A (TENN); 19 Mar 1977, S. McDaniel

21171 (MO). NORTH CAROLINA. Chatham Co.: 9 Apr 1960, H.E. Ahles and J. Haesloop 53191 (TENN) OKLAHOMA. McCurtain

Co.: 28 Aug 1976, J. Taylor 23358 (TENN). TENNESSEE. Lewis Co.: 23 Mar 2006, M.H. Mayfield 3649, (KSC). VERMONT. Orange

Co.: 4 Jun 1966, FC. ela 23613 (TENN). Caledonia Co.: 11 Jun 1982, D.E. Boufford and D.S. Conant 22913 (TENN). VIRGINIA.

ae Co.: 17 Apr 1975, R. Kral 55124 ieu) WIESE VIRGINIA. bii Co.: = id 1955, E.A. Bartholomew W-4024 (TENN). Wayne

0.: 8 Ápr 1938, F. Gilbert € Willi TENN ADA May 1935, F. Marie-Victorin & F. Rolland-

Germain 43268 (TX/LL). Ontario: 11 May 1941. H.A. em & M.N. Zinck 162 (TENN).

APPENDIX 1

[ee iss l sl | I TERN Si "E FI! I | L L

f pubescence; Y ANA OI AI

List of

U.S.A. DELAWARE. HP 1876, G.W. Letterman (MO). MASSACHUSETTS (PI. H. Kohankie, Painesville, O., 1928): Jamaica

Plain, Arnold Arboretum, Harvard University, 16 Apr and 25 Jul 1941, R.B. Clark 20656 MO): MICHIGAN. »Cheboygan Co.: in

the beech-maple forest on Colonial a 9 Aug [n ne ae TOR Yoc Iron Co.: Lake,

15 Aug 1970, EA. Bourdo Jr. 22197 (MSC). MINNESO lake fi Ingersan d, Sand | Bus 24 "n 1926,

AM ahasan 2019 (MO). NEW a an Co.: bluffs of Delaware River, Phillipsburg, 5 Jun 1910, KK. Mackenzie 4584

(MO). Salem Co.: wet woods along Major Run, 1.5 mi S of Sharptown, 1935, J.M. Fogg Jr. 8844 o Salem Co.: ich

wooded slope along Culliers Run, 1.5 mi NE of leal 29 Apr 1936, J.W. Adams 2846 (MO). N K. Washin

Co.: woods, Pilot Knob, E side of Lake, George, 30 Aug 1944, H.D a 28695 (LL/TX). RHODE END 2 Fls e

Lvs Fr. Charlotte, May 1877 and 20 Jun 1876, C.S. Pringle (MO). VERMONT: rich woods, Brandon, alt. 500 ft, 30 Apr and 8 Sep

924, D.L. Dutton (KSC). Chittenden Co.: woods, Essex, 3 Aug 1911, S.F. Blake 2511 (LL). WEST VIRGINIA. Monroe Co.: bank

of Second Creek, 2 Jul 1941, AJ. Sharp

498 Journal of tl tanical Institute of Texas 3(2)

CANADA. ONTARIO. Halton Co.: Loc. 443 2 mi E. of R

6192 (MO).

Blue Springs, 24 May 1956, J.H. Soper & G. Fleischmann

ACKNOWLEDGMENTS

We thank Eduardo Estrada, John Lonsdale, Matt Meshriy, and Bob Patterson for collecting and sending im-

portant plant material; personnel of the Overland Park Arboretum (Johnson Co., KS) and San Mateo County

Parks (CA) for permitting assistance; the staff of MSC for allowing us to Sape tissue from an herbarium

specimen; the staff of MO for sending silica-dried material of Dapl ; Shannon Fehlberg and

Mary Andreopolous for assistance with molecular work; and Deyka Garcia for assistance with the spanish

abstract. We are grateful to Lorin Nevling for discussion of Dirca biogeography and morphology, to Bill

Graves, Guy Nesom, and Gene Wofford for valuable comments on an earlier version of the manuscript, and

to Guy Nesom and Bob Patterson for valuable comments during manuscript review. We acknowledge the

following herbaria for loans of specimens: BRIT, KANU, LL/TEX, MSC, MO, and TENN. This work was

partially supported by NSF DBI-0544980 (CJF and MHM) and the Kansas Agricultural Experiment Station

(Contribution No. 10-011-J).

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BOOK REVIEW

C.T. Bryson AND M.S. DeFetice (eps.), with A.W. Evans (PHOTOGRAPHS). 2009. Weeds of the South. (ISBN

978-0-8203-3046-4, flexibind). The University of Georgia Press, 330 Research Drive, Athens, Georgia

30602-4901, U.S.A. (Orders: www.ugapress.org, 1-800-266-5842). $39.95, 498 pp., 1500 color pho-

tographs, 125 illustrations, 405 maps, 7 1/2" x 10°.

vo =e the South is a pene book, “ue M 3 pages, pd produced and iut itd pel the P ae are oe to aj con-

Su h

isingly, thoug

families). By my count, the book includes 400 ies in 65 f: The geographi g “extends fom Virginia along the Ohio River

i to Missouri, across ee and en to T 1 Florida", and almost all i The "pasos E id

cy 4 E a

for ed species. Al ided is ini ti ler headi f "Synonyms," “Habit, Habitat, nE Origin,” “Seedling Chartes"

“Mature Plant Cae aes ” “Special Identifying Features,” and “Toxic Properties.”

From the brief Introduction—Weeds are “plants that alter the structure of natural communities; interfere with the function of

a or have can effects on DACH, agriculture, or other societal interests. Any plant can ue a weed in one situation and

j i di 's wild ” “Weeds are usually plants

other words, one person’s weed is ano s wildflower.

that grow spontaneously and. ogee in a habitats that id En modified D human a activity.” ? “Clearing for cultivation and urban

Although not explicit anywhere,

development

a pua criterion i adasi in ue book perhaps is a need for management "Weed management nes ii and accurate

l! In most Ca ses, 1 IL LS LlIELCeSSsal M LU identify weeds early 5 5 when

they are easier to manage.”

To the extent — Weeds el the South is intended to provid ns of identification for plants growing in disturbed sites, it will be a

good help. B t's intent t t which should | ged and controlled Pihon caveat, the book is misleading. 193 of

ti ies is a good sample, but tl is no perspective to guide

ae 400 total Po are native to the USA 207 herbaceous non

native invacives

Of de 69 agn o Dnes mentioned, 44 are natives, ee ici altissima, Coreopsis tinctoria, Dracopis amplexicaule,

5 nia ppm and Vernonia gigantea. OUCH!

Erigeron philadelphicus, leli , Rudbeckia hirta,

ue ese are native BM conspicuously b beautiful. d ] to be bool hri je rth America. Here's the

1 irii. A 4 dq -q 113 13 ] nmativese

5 y f

: id A tina altissima, Ambrosi temisiifoli LA Vudentzld. A. grayi, A trifida, Amphiachyris d loides, Bidens

bipinnata, B. frondosa e discolor, C Menden Conyz 1 d ramosissima, C tinctoria., D i pl icaul , Eclipta

prostrata, Erig lelphicus, E. strigosus, Eupat illifoli Fl i Grindelia ciliata, G. squarrosa, Gut t}

H autumnale, Heli ,H ciliaris, H aa H. grosseserratus, H petiolaris, H. tuberosus, Heliopsis helianthoid

Lact floridana, Packera gl ella, Pyrri lini , Rudbeckia hirta, Solidag lensis, S. altissima, Symphyotrich livaricat

] a A i Tha IR nan rn Abi 211, £ 1 fiel de

-j 3

>: pilosum, Verbesina encelioides, dien baldwinii, V. gigantea,

Anthemis cotula, Arctium minus, Carduus nutans,

C I t} l , Matricaria

Cent biel teinii, C. cyanus,

[e]

C. solstitialis, Ci ium intyl Meny arvense, C ele

discoidea, Porfhenium OM Mie Senecio vulgaris, burton marianum, Soliva sulle Sonchus arvensis, S. asper, S. dari Tanacetum

edi Taraxacum officinale, Tragopogon dubius, and Xanthium spinosu

e the a ian hold so obviously in other ee Let’s see. ae ME 5 SAREA species, all are natives: Ludwigia

decurrens, I peploides th , O. laciniata, and O speciosa Oft I 11 are native. Beans, grasses, and

1 1 1 41. m atixvrec M ] f A al] din 1 Andropogon

Lo! r r a d

virginicus, Brunnichia ovata, C l li D thus illi is. C i li Hoff ia el Lupinus perennis. Mvosurus

1 , t E JJ DO o ? t L iain:

miminus, Nuttall lensi n t) i quinquefolia, Passiflora i ta, P. lutea, Phytol i S , Tripsacum

PE Verbena bracteata. The first t] ies in the bool Equisetum arvense, E. hyemale, and Pteridium aquilinum

any of these natives s often abe in disturbed R n does ¥ eeds of the So uth ply I d treated tl

came ac the nan hh} i ; Ji ] t t 1

t I pes] , with no

1 jee » ] 1 1 And then tl T he S I ] ld I lerful Vernonia

I 14 = H _] fe 1 Pe PNE Finally 1 , colo call 1 gJ 1 1 1

i Zt in d O” Pi o e El O

the South: to fill he space of this boo Salvinia mimi d S. molesta, Myriophyll ti Lygodium japonicum, Bothriochloa

ischaemum, Centaurea melitensis, Torilis arvensis, and others?—Guy Nesom, botanist, Fo: com, Fort Worth, Teds. U.S.A.

J. Bot. Res. Inst. Texas 3(2): 500. 2009

TWO NEW COMBINATIONS IN TRYPHANE (ALSINACEAE)

W.A. Weber and R.C. Wittmann

University of Colorado Museum

B 265

Boulder, Colorado 80309, U.S.A.

billweber@colorado.edu

-—

ABSTRACT

Two scientific names in the genus Tryphane are validly published.

RESUMEN

Dos nombres científicos del género Tryphane se publican como válidos.

Áskell and Doris Love (1976) treated Minuartia as several smaller y hyletic genera, including Lidia, Alsi-

nanthe, and Tryphane. cme their arguments for this have been ignored. Their discussion follows:

The redefinition of the strictly Medi Minuartia L. by Hiern eee mes made it pone to accommodate within

its limits even arctic species, m was caus ised b ] g gnitude, although later authors and

even two see apa Sabe TAN 1922; -— 1962) to 1 pted this without hesi The dici SO dur]

defined is highly f view it is looked I t clearly 7

ties d by McNeill in dividing it into sub , sections, and even cio and series, some ai qu are not only

morphologically het but al -— acterized by distinct basic chromosome numbers and by s different

from that " cs supposedly opar lanys.

Minuartia s. st } | 15. wl ti 11

been reported to have basic numbers as ipu as X = li 9, 10, 11, 12, 13. dh 23. We have had the a to ea oe

AA sncieS: still ne unpublished,

Eurasia and North A d al f lati f the Medi d th th Aciatic area of

E Iu

: s 1 = Mar J ge En | 4 1 .

the calicis genus. These studies, which ]ud b poll ] phology, have convinced

al a Lot 4 7 1.1 J 1 A: Des | e a + el 1 £^. ] 3 Lo... ` 1 Jf P E em EU d

[e] [e] Y [m] [e]

Therefore, in the Cytot ic Atlas of the Arctic Flora (Love & Lóve 1975) we | i ] long ig 1 but well-

. : ] ZI Je. Gl 4 ] Fae ra RA A lands ] 1 r p

[e] I a

y | ds 3 + +l +1 1 Ln pra a |

SiPLupo 1UL

The authors went on to discuss Porsildia, Wierzbickia, Lidia, Alsinanthe, Tryphane and Neumayera, noted their

chromosome numbers, and provided a useful key to the genera, but while they made new combinations

in Alsinanthe, Porsildia, and Lidia, they evidently neglected making any in Tryphane, a genus proposed by

Reichenbach (1841). We adopted three of these genera in the Colorado Flora (Weber & Wittmann 2001) and

followed the nomenclature as given in Lóve (1983

In his Icelandic flora Á. Lóve (1983: 194) seed the combination Tryphane rubella, attributing the au-

thorship to Reichenbach. He also listed a subspecies propinqua (Richardson) Love & Love. Neither name

appears to have been published hitherto. Search as we might in his bibliography (Weber 1995), we have

not been able to find a place wherein Askell and Doris made the combinations. As Askell often said, “Even

Jove nodded.”

Rabeler et al. (2005) cite, without discussion, Tryphane rubella (Wahlenberg) Reichenbach as a synonym

of Minuartia rubella (Wahlenberg) Hiern. We agree that the various varieties and subspecies mentioned as

synonyms are trivial.

To resolve this problem and use the names, we propose to validate them as follows:

Tryphane rubella (Wahlenb.) A. & D. Lóve ex WA. Weber., comb. nov. Basionym: Alsine rubella Wahlenb., Flora Lapp.

128, pl. 6. 1812

| Rot Rac Inst Tewac 3(2): EnM — 502. 2009

502 J | of the Botanical R h Institute of Texas 3(2)

Tryphane rubella subsp. propinqua (Richardson) A. & D. Love ex WA. Weber, comb. nov. Basionwx: Arenaria

propinqua Richardson, in J. Franklin, Narr. Journey Polar Sea 738. 1823.

ACKNOWLEDGMENTS

We are indebted to Kanchi Gandhi for bringing this problem to our attention and for commenting on an

earlier draft of the paper.

REFERENCES

Love, A. 1983. Iceland's flora. Reykjavik.

Love, A. AND D. Love. 1975. Cytotaxonomic atlas of the arctic flora. J. Cramer, Vaduz.

Love, A. AND D. Love. [1975] 1976. Nomenclatural notes on Arctic plants. Bot. Not. 128:497—523.

Raseter R.K, eT AL. 2005. Minuartia. In: Flora of North America Editorial Committee, eds. Flora of North America

North of Mexico. 5(2):134.

REICHENBACH, (H.G) Lupwic. 1841. Flora Germanica. 28:28-29, 32.

Weeer, W.A. 1995. A bibliography of the published works of Áskell Love. Acta Bot. Islandica 12:6-34.

Weser, W.A. AND R.C. WITTMANN. 2001. Colorado flora. Eastern and Western Slopes. Third edition, two volumes. Univ.

Press of Colorado.

TWO NEW SPECIES OF SOLANUM FROM ECUADOR AND NEW COMBINATIONS

IN SOLANUM SECTION PACHYPHYLLA (SOLANACEAE)

Stephen R. Stern and Lynn Bohs

Department of Biology

University of Utah, 257 S. 1400 E.

Salt Lake City, Utah 84112, U.S.A.

af È Tet a

ABSTRACT

I o4 f Tara ee ana HA e rl here Cal Ls Cc. "ce à 1 fe des r

Two new species

totrichum. It is most similar to S. deflexum, S. turneroides, and S. hoffmanseggii but differs in a unique combination of straight hairs, a

| BS 1 pers J ] 1 1.: 11 x A arn pical Serene) T 4 1 A = P +1 T ss + fu J

D e rr

] 1 T J f 1 AA Le il 1 11 + 1 Canl 1 nzal ] £ 1 LF 1

1 fcil Te : + 2 21 + obliqi e ARE = 1 1 4 1

in southern Ecuador, i Pachy sok ylla

hairs on the foliage and axes, acute fruits, purplish, nearly glabrous corollas, and very us filitienis: One new name add three new

hylla

combinations are M validated in Solanum section Pachyphy

RESUMEN

i uo sl :T * Hd : dax M ula 7l ER pa | Eal hi C Cr i h As Cal 3 £27

i E

natotrichum. Est i imil S. deflexum, S. turneroides Y S. hoffmanseggii, pero se dif ia por tener bi

de pelos rectos, un hábito r rizomatoso, y ill g pical. Parece q émi la costa pacífica

SA yt ae Y i j ES IO: Bs] 1 HA 1 tig Caf 1 Dal 1 - | r J

Zumba del le E lor; esta especi iembro de Solanum ión Pachyphylla. Es parecida a S. obliquum, pero

lif ia [ f puntiagudos, f! d l la y casi glabra, y filamentos largos, También se publican un nombre y

t þi i Solanum sección Pachyphylla

Two New Species of Solanum from Ecuador

A comprehensive project to complete a species-level taxonomic treatment of the entire genus Solanum, sup-

ported by the National Science Foundation Planetary Biodiversity Inventory program, has facilitated detailed

taxonomic study of many little-known groups within this giant genus (Knapp et al. 2004; http://www.nhm.

ac.uk/solanaceaesource). Herbarium study and field work associated with this project have uncovered two

undescribed Solanum species from Ecuador, which are treated below.

HE of the species, Solanum manabiense S. Stern, pene to S. section Gonatotrichum, a group of about

zx aoe eC ok RIAYA

disiunct distribution between C

dum incide 5, diam Greenm. and S. lignescens Fernald from Central America, S. turneroides Chodat and

S. hoffmanseggii Sendtn. from northern Argentina, Bolivia, Paraguay, and into the Mato Grosso do Sul region

of Brazil, ano S. adscendens Sendtn., endemic to Southern brazil; Molecular phylogenetic analyses indicate

that of S. section G ichum form a letic group that is part of the Brevantherum clade

of Bohs (2005) and Weese and Bohs (2007). Species al this group are distinctive in the genus due to their

explosive fruit dehiscence. The fruits are wale, with a thin exocarp, and build up sian posue as d

mature until they burst, explosively | ds. Plants of this group are generally diminutive annua

or herbaceous perennials with short, fewdlowered inflorescences and simple, Sometimes geniculate hairs.

This group was studied by Nee (1989), who recognized two species. However, Nee annotated a specimen

of the new species described below as S. manabiense, as S. deflexum vel. aff., which he later synonymized

with Solanum adscendens. Detailed taxonomic study has reevaluated the ci iption and species limits

of section Gonatotrichum and has revealed S. manabiense to be a distinct species.

J. Bot. Res. Inst. Texas 3(2): 503 — 510. 2009

504

: M

Fis. 1. Sol bi S. Stern. A, B. Habit. C. Bud (left) and i fi (right). D. Fl tion. E. Stamens. All from 7. Plowman & P

Alcorn 14334 (NY)

Solanum manabiense S. Stem, Sp. nov. (Figs. 1-2). Tree: ECUADOR. Manasi: Pacoche Reserve, road from Manta to San

Lorenzo, ~2 km W of El Aromo, 01°04'09.5"S, 80°52'32"W, 350 m, 8 Feb 2009 (fl, fr), S. Stern & EJ. Tepe 377 (uoLorvre: QCNE;

ISOTYPES: BM, E MO, NY, QCA, UT).

Solano deflexo et S. hoffmanseggii affinis sed a S. deflexo habitu rhi t t inil i gine tumido et incisum apicali, a S

hoffmanseggii pilis non geniculatus differt.

Rhi t herl ti lightly woody at the base, 1.5—4 dm tall. Stems sparsely to densely pubescent

: ce El

Stern and Bohs, N i | binations in Sol (sect. Pachyphylla) 505

Fic 7 nL de Es il A frst LE c Starn A Hahit R PI rt L H rt Al rrr ia

r * n n (A | 1 Lana? AB L i, E [| J P | AN 4 I a, Canalan LEE A 10 cm. B,C=5mm.

with two celled unbranched, straight hairs, unarmed. Sympodial units difoliate, geminate. Leaves simple,

1-6 x 0.5-3 cm, elliptic to elliptic-ovoid, chartaceous to membranaceous, nearly glabrous to sparsely pu-

bescent adaxially and abaxially with one- to two-celled, unbranched, straight hairs, these lying flat along

the blade, denser along veins; base rounded to obtuse, often decurrent into petiole; margin entire and ciliate

with unbranched hairs; apex acute to obtuse; petioles 0.5—1 cm, moderately pubescent with unbranched

straight hairs. ee EE extra- exa Ty or subopposite the leaves, unbranched, with 1-5 flowers,

all flo I tely pubescent with unbranched hairs; peduncle absent; rachis

absent to ca. 1 mm; Wedel: 5- 15 mm in flower 10-20 mm in fruit, nearly contie nons; articulated at the

base. Flowers homostylous, 5-merous. Calyx 3-10 mm long, the tube 1-3 mm, the lobes 2-7 x 0.5-1.5 mm,

linear-lanceolate, moderately to densely pubescent; fruiting calyx not accrescent, not completely covering

the fruit. Corolla 0.4—1 cm in diameter, 2-3 mm long, rotate with abundant interpetalar tissue, chartaceous

to membranaceous, white, the tube 2-3 mm long, the lobes very short, 1-2 x 0.5-1 mm, triangular, acute

at apices, sparsely to moderately pubescent abaxially and on margin with 2-3-celled unbranched, straight

hairs, glabrous adaxially. Stamens 2-4 mm long; filaments up to 1 mm long, glabrous; anthers 1.5-3 x

0.5-1.5 mm, oblong, connivent, yellow, the base cordate, the apex emarginate, the pores directed introsely

and subapically, not opening into longitudinal slits. Ovary glabrous; style 4-6 x 0.5-1 mm, equal to or

exserted beyond stamens, cylindrical, glabrous; stigma up to 1 mm wide, capitate. Fruit a globose berry,

B || Eal ra " Im L I Side Sf)

506 f Texas 3(2)

5-12 mm in diameter, white to yellow when immature, maturi it t, drying brown, glabrous,

the mesocarp watery and held under pressure until decime eds at maturity. Seeds 10-35 per

fruit, ca. 2.5 x 1.5 mm, somewhat flattened, with a small notch where connected to placenta, the margin

not swollen, the surface with fine raised ridges radiating from the center to the edges and shallow ridges

running parallel to aA

Known only from the central coast of Ecuador in Provs. Manabi and Guayas

MALAS YY LL bama A

from sea level to 400 m. Flowering and ee Spence Muse a collected in did and July.

difoliate, occasionall inate sympodia; | lal the abaxial surface; unbranched, straight an

geniculate) hairs; small flowers viti equal cds and seeds with unexpanded margins and no obvious

notch at the attachment point to the placenta. It is apparently restricted to coastal Ecuador in the Guayas

and Manabi provinces. It is named for the latter province where the type was found and the species is ap-

parently a common roadside weed

Solanum manabiense resembles S. hoffmanseggii, but the hairs of S. hoffmanseggii are geniculate whereas

those of S. manabiense are straight. Solanum manabiense also resembles S. turneroides, but S. turneroides is a

more robust herb with heterantherous flowers that are much larger than those of S. manabiense. Solanum

manabiense is also similar to S. deflexum, but the latter has densely pubescent leaves, is not rhizomatous,

and has seeds with a swollen margin and pronounced notch where they connect to the placenta.

Parsimony analyses of sequence data from three molecular markers (nuclear waxy or GBSSI and ITS

and chloroplast trnT-F) place Solanum manabiense in the Gonatotrichum clade (S. Stern, unpub. data). The

waxy and combined analyses place S. manabiense sister to a clade containing the South American species S.

hoffmanseggii and S. turneroides. The Central American S. deflexum and S. lignescens clade is more distantly

related. Solanum manabiense is the only species of S. section Gonatotrichum known from northern South

America.

Pararvres: ECUADOR. Guayas: 2 to 4 km E from Recinto Olon, ca. 10 km N of Manglaralto, 19 Feb 1974, Gentry 10068 (MO). Manabi:

Montecristi, Cerro Montecristi, eastern slopes above town, 18 Jul 1986, Pl an € Alcorn 14334 (F, NY, QCA

The second new species, Solanum zumbense, belongs to the Cyphomandra clade of Solanum along with other

species traditionally oe in S. sections ii n and Kies e Bitter poe E 2007).

cium Be La rc

Many members of this clade he gem homandra due to their

connectives, which function as csmephores i he attraction of male euglossine bees (Bohs 1994). Subse-

quent phylogenetic research revealed that Cyphomandra is deeply nested within Solanum, necessitating the

transfer of all of its species to Solanum (Bohs 1995). Monophyly of the group of species with enlarged anther

connectives has not been conclusively demonstrated, but monophyly of the more inclusive group of species

comprising the Cyphomandra clade is well-supported (Bohs 2005, 2007; Weese & Bohs 2007).

ip Lio have been published for spen of the cae clade (Bohs 1994, 2001), but novel-

ties t discovered. Specimens of the following new species were annotate d as S. sp. aff. obliquum

Or $a sp. nov. by Bohs during ihe preparation of her 1994 treatment. Some collections recognized here as

a zumbense were noted in the monograph as being anomalous within S. obliquum Ruiz & Pav., but at that

time theres doubt about their status as a distinct species. Subsequent assessment of herbarium and

living e torem that these collections represent a new species.

Solanum zumbense Bohs, Sp. nov. (Fi 1g. 3). Tree: ECUADOR. ZamorA-CHINcHIPE: road between Zumba and Amaluza, 8-10

km W of Zumba, 04?50'07"S, 79°09'50"W, 1500-1700 m, 31 Mar 2005 (fl, fr), L. Bohs et al. 3366 (HOLOTYPE: Q : BM,

LOJA, NY, QCA, UT).

ci Li: . 1 J 11: :1 f=1 "LI : PEN M +31 qe oi! pene

Shrub or small tree 1-3 m tall, from a single trunk with a spreading crown. Stems densely puberulent with

unbranched glandular and eglandular hairs and often also sparsely to moderately pilose with eglandular

hairs 1-3 mm long. Sympodial units 3—4-foliate, geminate or not. Leaves simple, the blades 4-25 x 2.3—24

cm, 1-2.5 times as long as wide, subcoriaceous, ovate, sparsely to densely puberulent adaxially and abaxi-

Stern and Bohs, N i ] binations in Sol (sect. Pachyphylla) 507

2) ie.

28 a 2

Fic. 3. Sol bense Bohs. A. | | infl B. infi letail. C. Bud. D. Flower in face-on (left) and side view (right). E. Flower

cross-section. F. Stamens. G, H. Fruits, All from Bohs et al. 3366.

508 t tani titut Texas 3(2)

ally with unbranched eglanduiar hairs and often also sparsely to densely pilose with unbranched hairs 1-3

mm long, the hairs more dense on veins; major veins ca. 5 per side; base truncate to deeply cordate, often

oblique, with basal lobes (0-)0.5-3.5 cm; margin entire; apex acuminate; petioles 3-10 cm, densely pu-

berulent and often also sparsely to moderately pilose. Inflorescences 3-12(-25) cm long, leaf-opposed or in

a branch fork, unbranched, forked, or rarely further branched, with 15-40 flowers, all flowers perfect, the

axes densely puberulent (pilose); peduncle 1.5-5 cm; rachis 1-6.5 (20) cm; pedicels 15-25 mm, 25-40

mm in fruit, spaced 1-6 mm apart, articulated near the base and leaving pedicellar remnants ca. 1 mm

long. Buds ellipsoidal or ovoid, acute at apex. Flowers homostylous, 5-merous. ae 3-5 mm long, the

tube 2-4 mm, the lobes 1-2 x 2-3 mm, deltate to very shallow, apiculate at tips, sul , moderately

to densely puberulent, with hairs denser toward tips of lobes; fruiting calyx not accrescent. cla 1.5-2

cm in diameter, 12-16 mm long, stellate, coriaceous, dull greenish, violet, or brownish-purple, the tube

1-2 mm, the lobes 9-14 x 3-4 mm, narrowly triangular, acute and slightly cucullate at apices, glabrous

abaxially, glabrous to sparsely puberulent adaxially on midrib and toward apices, the margin tomentose to

ciliate. Stamens with the free part of filaments 2-4 mm long, the filament tube 1-2 mm; anther thecae 4-5

x 1.5-2 mm, lanceolate, loosely connivent, cream to purplish, the pores directed abaxially and distally,

not opening into longitudinal slits; connective 4-5 x 1.5-2 mm, lanceolate, purple or brownish, abaxially

slightly shorter than thecae at apex, slightly exceeding them at base, adaxially produced as a swelling ca.

3 x 1 mm. Ovary glabrous or finely puberulent; style 4-7 x 0.5-1 mm in diameter at base, 1.5-2.5 mm in

diameter at apex, equal to or shorter than the stamens, umbrella-shaped, strongly dilated distally, glabrous;

stigma truncate, 1.5—2.5 mm in diameter, with two prominent apical glands. Fruits 6-7 x 1.5-4.5 cm, el-

lipsoidal to fusiform, acute to apiculate at apex, pale yellow when mature, glabrous to finely puberulent;

stone cell aggregates large. Seeds 5-6 x 3-4 mm, flattened, reniform, rugulose to pubescent along margin

with white pseudohairs.

Distribution and phenology.—Clearings and open places in tropical rain forest, (400—)1500-2255 m in

elevation, eastern Andean slopes in southern Ecuador and Peru, with an outlying collection from Brazil.

Flowering specimens have been collected in March, July, September, November, and December. Fruiting

specimens have been collected in March through June and December.

Local names.—Peru: chupo sacha (Schunke 5866); tomato del campo (Mexia 8235). Brazil: tsetsepere

(Deni; Prance et al. 16402).

Uses.—Mexia 8235 describes the fruits as having a tomato-like taste, and Bohs et al. 3366 describe them

as sweet-sour. In Amazonian Brazil, the leaves are Bead in water and used to bathe babies, both to keep

them healthy and to cure fever (Prance et al. 16402). Schunke 5866 reports that an infusion is given in an

enema to combat the grippe.

Within the Cyphomandra clade, S. zumbense is most similar to S. obliquum. Both species have very

broad stigmas with two apical glands, short anthers with the connective not prolonged below the bases

of the anther thecae, and stellate, coriaceous corollas with relatively broad and spreading lobes. Solanum

zumbense differs from S. obliquum in having abundant long hairs on the foliage and axes, pointed and often

pubescent fruits, and purplish, nearly glabrous corollas. The filaments are quite long and are a distinctive

eae Solanum ii is aon at ee elevations ait 1500 as ee pue peus ofS. obliquum.

la the town of Z in southern border,

where the Brae collection was made. Although this locality is in the northern extremity of the range of this

species, the euphonious epithet was too good to pass up. “Zumbar” is also the Spanish word for “buzz,” a

common pollination mechanism in many species of Solanum.

PARATYPES. ECUADOR. Zamora-Chinchipe: trail between Mirador and Pallas, 2010-2255 m, 9 Sep 1943 (fl), Steyermark 54279 (NY).

Peru. Amazonas: Prov. Amazonas, Dist. Vista EISE Sector Poe ca. 1 km de Vista ding 06?10'S, 77*18'W, 1540-1720 m, 1

Jul 1998 (fD, Sánchez Vega & Zapata 9595 (F). Huánuco: ba, trail Puent to, 1625 m, 26 Sep 1936 (fl, fr),

Mexia 8235 (B, BH, BM, F Pe K, MO, NY, S, U, MP Prov. Leoncio Prado, Dist. Hermilio Valdizán, La Divisoria, 21.8 km E of Puente

Pumahuasi (Río Tul Pucall 5°S, 75?52"W, 1550 m, 27 Dec 1981 (fr), Plowman & Schunke

11724 (F, GH, MO); m Planos (carretera Monge): 22 Apr 1962 i ea 5866 (F, US); Prov. Huánuco, km 474 on Lima-Tingo

Stern and Bohs, N peci ] binations in Sol (sect Pachyphylla) 509

Maria road, Huánuco, 1650 m, 2 Jun 1981 (fr), Young & Sullivan 634 (NY). Pasco: Oxapampa, ca. 2 dei E O village, 10?35'S, 75?35"W,

1850 m, 10 Mar 1984 (fl, fr, Knapp et al. 6315 (BH, F, G, GH, MO, NY, US); Prov. Oxapampa, 51 , 75°23 W, 10?36'S,

1850 m, 13-16 Dec 1982 (fl, fr), D. Smith 2959 (F, NY): O pa Prov., Palcazu valley, Río San José in the Ri io Chuchurras drainage,

75°20'W, 10°09'S, 400—500 m, 14 May 1983 (fr), Smith 4023 (NY). San Martin: prope Tarapoto, 1855-6, Spruce 4941 (NY). Ucayali:

Prov. ~ Portillo, nea Azul, km 43 on Tingo María-Pucallpa road, ca. 1500 m, 5 Jun 1981 (fr), Young & Sullivan 741 (MO).

BRAZ Rio Cunhuá at Canacá, 06°34'S, 66°27'W, 27 Nov 1971 (fD), Prance et al. 16402 (NY).

NEW COMBINATIONS IN SOLANUM SECTION PACHYPHYLLA

Bohs (1995) transferred tl ithets of all Cypl l ies to Solanum. However, transfers of infraspecific

taxa were not "A at that Lime, The combination Selena corymbiflorum subsp. mortoniana (L.B. Sm. €r

Downs) Bohs was published in Zuloaga et al. (2007) and the citation isincluded here for completeness. The fol-

lowing new combinations will validate the remaining infraspecific taxa belonging to S. sect. Pachyphylla.

J 7 NA

Solanum circinatum Bohs subsp. ramosum (Bohs) Bohs, comb. no iers) Walpers

subspecies ramosa Bohs, Revista Acad. Colomb. Ci. Exact. 16:73. 1988. Tyre: COLOMBIA. Hunt Fundación O ca. 1300

m, 18 Aug 1981 (fl, fr), L. Bohs 1644 (HoLotyre: COL; isotypes: CAUP, GH

Solanum corymbiflorum (Sendtn.) Bohs subsp. oro (L.B. Sm. € Downs) Bohs, Darwiniana

45:24 1. 2007 (as dari mortoniana”). B toniana L.B. Sm. & Downs, Phytologia 12:250. 1965

Cypl a corymbifl itn. subsp. mortoniana (L.B Sm pM Bohs, Fl. Neotrop. Monogr. 63:68. 1994. Tree: BRAZIL.

SANTA C São J h juei 7 km E of São Joaquim), 1100-1200 m, 16 Jan 1957, L.B. Smith & R. Reitz 10219

(HOLOTYPE: US; ISOTYPES: GH, HBR, R).

Solanum ee Dunal Sue: laran a Pons. EOS Nov. BASIONYM: i dapi chlorantha

r. S. Amer. Pl. 116. 1920. Cy Fl. Neotrop Monogr.

63:82. 1994. Type: COLOMBIA. SANTA MARTA: Hr in d learings, flowers Jan to Apr 4500 ft, 26 Jan-25 Feb 1899 (fl,

fr), H.H. Smith 1180 (Lecrotyee, designated by Bohs 1994: NY; ISOLECTOTYPES, A, E G, GH, K, L, LL, MO, NY, P S, U, US, W, WIS).

Solanum endopogon nn ne subsp. guianense (Bohs) Bohs, comb. nov. B

otrop. Monogr. 63:90. 1994. Tyre: FRENCH GUIANA. Saul, Monts A Fumée, 3037N, 53912

200-400 m eden 16 Aug 1982 a S. Mori & B. Boom 14709 (HoLoTyrE: F).

In addition to validating these infraspecific names, molecular evidence in Bohs (2007) indicates that S.

circinatum subsp. ramosum might be better recognized as a distinct species. The combination S. ramosum

Lam. has already been published; therefore a new epithet must be coined for this taxon. The epithet huilense

commemorates the predominant distribution of the plants, in Dept. Huila, Colombia.

Sol huil Bohs, nom. nov. for Sol circinatum Bohs subsp (Bohs) Bohs. Basionym: Cyphoman-

dra hartwegii (Miers) Walpers subsp. ramosa Bohs, Rev. Acad. Colomb. Ci. Exact. 16:73. 1988. Tyee: COLOMBIA. Huma: Fundación

Merenberg, ca. 1300 m, 18 Aug 1981 (fl, fr), L. Bohs 1644 (notorvee: COL; isotypes: CAUP, GH).

ACKNOWLEDGMENTS

We thank J. Bennett, J. Clark, and E. Tepe for field assistance; the herbaria listed for loans of specimens;

and Bobbi Angell for the drawings. This work was supported by NSF grant DEB-0316614 (PBI Solanum: A

Worldwide Treatment) to LB. Gregory J. Anderson and an anonymous reviewer provided critical reviews.

ROPFERENCES

Bons, L. 1994. Cypt ira (Sol ). Fl. Neotrop. Monogr. 63, New York Botanical Garden, Bronx, New York.

Bous, L. 1995. Taree of Dvohomandia (Sclanacee) and its species to Solanum. Taxon 44:583-587,

Bous, L. 2001. A revision of Solanum section Cyphomandropsis (Solanaceae). Syst. Bot. Monogr. 61:1-83.

BoHs, L. 2005. Major clades in Solanum based on ndhF Aene gala In: Keating, R.C., V.C. Hollowell, and T.B.

Croat, eds. A festschrift for William G, D'Arcy: the | t. Monogr. Syst. Bot. Missouri Bot. Gard,,

Vol. 104. Missouri Botanical Garden Press, St stig Missouri Pp. 27-49,

Bons, L. 2007. Phylogeny of the Cyphomandra clade of the genus Solanum (Solanaceae) based on ITS sequence

data. Taxon 56:1012-1026.

510 tl tanical h Instit Texas 3(2)

Knapp, S., L. Bos, M. Nee, AND D.M. Spooner. 2004. Solanaceae — a model for linking genomics with biodiversity.

Comp. Funct. Genomics 5:285-291

Nee, M. 1989. Notes on Solanum section Gonatotrichum. Solanac. Newsl. 3:80-82.

Weese, T.L. AND L. Bons. 2007. A three gene phylogeny of the genus Solanum (Solanaceae). Syst. Bot. 32:445-

] J T H | I ] £l

ZULOAGA, F.O., O. MORRONE, AND M.J. BELGRANO. 2007. N y nomenclat | vascular

del Cono Sur de Sudamérica. Darwiniana 45:236-241.

THREE NEW SPECIES OF SOLANUM SECTION HERPYSTICHUM

(SOLANACEAE) FROM ECUADOR

Eric J. Tepe and Lynn Bohs

Department of Biology, 257 S. 1400 E.

Salt Lake City, Utah 841 12, U. 7 d

A 4L £,

¿ULT TUN poy Ls ere Ll

ABSTRACT

Three new species of Solanum sect. Herpystichum from Ecuador are avenue and illustrated: $. AS endemic to the

eles of eastern ee S. pacificum, endemic to the Pacifi lands, and S. crassinervium, f Ecuador and

bles the Colombi les S dalibardiforme, | liff ily in having stel

[s E i e

1 1 1 t

late corollas. Sol i is similar to S. dolichorhachis, |

very tl

leaves with maed» abiicure bases. Lastly, S. crassinervium is most anlar to 5. deca qs but can b differentiated by its larger,

ee ad cin with rs veins. Sol 1S. pacij pp

RESUMEN

um = Ma | (que | Seid LI I 13-17 4 A Daj :] " J cC is 1 i= D ]

bajas del Ecuad paras S. pecu endémica de las tierras ae del EU a y > pcia, del noroeste del

Ecuador y el suroeste de g on I S. dalibardiforme, de la cual se

dice: ] ae | Cal f; : . L p: MEE E

5 In $ tj -) 5 dias pero tiene noras MUY imas Y simeltricas en vez qe

hojas d tá iá las | ] te obli Finalmente, $ i i I S. evolvulifolium, pero se puede

xen . lots j E 1 1 ; | dietrihuci le S S pacificum

T m 1 4

INTRODUCTION

The genus Solanum has been the focus of intense research because it contains several important crop spe-

cies, ee the potato (S. tuberosum), tomato (S. lycopersicum), and eggplant (S. melongena). Despite the

of the genus, however, many groups among its ca. 1500 species remain poorly known.

The “PBI Ta a world-wide treatment” project (www.solanaceaesource.org), an NSF-funded initiative

to provide online descriptions and keys to all Solanum taxa, has allowed for increased exploration and dis-

covery of new Solanum species. Ecuador is home to 183 species of Solanum, including 31-34 endemics, and

is one of the highest centers of diversity for the genus (Jørgensen & León-Yánes 1999; Montúfar 2000). This

paper, along with three others funded by this project (Knapp 2007, 2008; Stern & Bohs 2009), adds seven

new species, five of which are endemic, to the list of Ecuadorian Solanum.

Solanum sect. dial i Bitter is a group of ca. ten species found from Central America to northern

Peru, and is a member of the i lly-named “Potato clade" (Weese & Bohs 2007). Molecular phylogenetic

analyses indicate that sect. Herpystichum constitutes a monophyletic group sister to sect. Pteroidea Dunal

(Weese & Bohs 2007; Tepe & Bohs, unpublished manuscript). Species of sect. Herpystichum are trailing,

terrestrial or climbing, herbaceous to woody vines rooting at the nodes. They are primarily plants of the

rainforest understory, but several ae are BER found in cada or other Spes areas. Like sect. Pteroidea,

they have perfect flowers, but can | 1 by the usually extra-axillary i (tl in sect

Pteroidea axe axillary in position). Some species have flattened fruits that are unique in Soldmum: This group

has never been revised and has received little attention since Bitter described most of its species in 1912

and 1913 (Bitter 1912, 1913a, b). Detailed study of sect. Herpystichum has revealed three additional species,

described herein, that have been collected since Bitter's focus on the genus. Two of the species are endemic

J. Bot. Res. Inst. Texas 3(2): 511 — 519. 2009

512 t tani i f Texas 3(2)

to Ecuador and occur in small reserves in areas under intense pressure from large-scale agricultural or oil

exploration. They should both be added to the Red List for Ecuadorian Endemics (Knapp et al. 2007).

Maximum parsimony analysis of sequence data from nuclear ITS and GBSSI (or waxy) sequences place

all three species in sect. Herpystichum (Tepe & Bohs, unpublished manuscript). Solanum crassinervium and

S. pacificum are in a clade with S. evolvulifolium Greenm., a widespread species that occurs from Central

America and Andean South America and S. loxophyllum Bitter from the Pacific lowlands of Ecuador. Solanum

limoncochaense is in a clade with the morphologically similar Central American species S. phaseoloides Polak.

and S. trifolium Dunal, a high-elevation species known only from Bolivar Province, Ecuador. We were unable

to obtain material for molecular analysis of S. dalibardiforme Bitter and S. dolichorhachis Bitter, the species

most similar morphologically to S. limoncochaense and S. pacificum, respectively.

Solanum pacificum Tepe, sp. nov. (Fig. 1). Ter. ECUADOR Los Rios: Ce ífico Rio Palenque, in secondary forest,

15 m, 5 Feb 2009 (fl, fr), Ej. Tepe et al. 2696 (HOLOTYPE: Q ‘PES: BM!, MOI, NY!, QCA!, UT).

C ud J4Il3«L TL 1 J-F -1 : lal = 1:1 E 1 1 a £1 1 -1

e oF

Vine, climbing other vegetation via adventitious roots at the nodes. Stems slender, weakly herbaceous,

glabrous, densely gland-dotted; fertile branch tips pendent. Sympodial units plurifoliate, not geminate.

Leaves simple, 14-19 x 4.5-8 cm, 23 times as long as wide, lanceolate to ovate, membranaceous to thinly

chartaceous, glabrous adaxiaily and abaxially, densely gland-dotted, the base rounded to obtuse, more or

less symmetrical, the margin entire, the apex acuminate; petioles 1-1.5 cm, glabrous, densely gland-dotted.

Internodes 1.5—7 cm. Inflorescences 4-10 x 23 cm, slender, extra-axillary, unbranched with 17-58 flowers

(scars), all flowers apparently perfect, the axes glabrous, very slender; peduncle 2-4.5 cm long; rachis ca.

6 cm; pedicels 8-10 mm in flower, slender, 15-20 mm in fruit, enlarged apically, glabrous, spaced nearly

contiguously to 12 mm apart, articulated at base. Flowers with the calyx 1-1.2 mm long, glabrous to minutely

and sparsely ciliate along margins, the tube 0.5-0.7 mm long, the lobes 0.5-0.6 x 0.8-1 mm, rounded,

rounded to weakly acuminate at tips; fruiting calyx somewhat accrescent, the lobes 0.6-0.8 x ca. 1 mm.

Corolla 0.8—1 cm in diameter, ca. 5 mm long, stellate, membranous, green to wale near ieee margins of the

petals, the lobes 45 x 1.2-2.5 mm, reflexed at maturity, acute at apices, glal 1 abaxially, the

margins ciliate. Stamens subequal, with filaments ca. 0.8 mm long, glabrous, free; — 1.522 x 0.7-1.2

mm, oblong, not connivent, yellow, the pores directed distally, opening into short longitudinal slits with

age. Ovary glal ; style 4—4.5 x 0.1-0.2 mm, glabrous, slightly clavate; stigma truncate. Fruit (immature)

ca. 0.9 x 0.6 cm, ovoid, pointed at apex, green, glabrous. Seeds unknown.

Additional specimens examined: ECUADOR. Junction of the provinces Bolivar, Caüar, AS ans eer ee of

the western a near the eee of Pu 1000-1250 ft, 8— opua 1945 auk W.H. Camp E-3782 (A

Q t km 47, 1.7 km S of REN Pilar,

0°35! 5s 79?2]"W, 220 m, 9 Apr 15 1992 2 (9, T.B. Croat ME Río o Palenque uL Statim km 56 Rd. Quevedo-Sto. Domingo,

150—220 m, 26 Oct 1974 (sD, C ison 5663 (SEI , km 56 Rd. Quevedo-Sto. poen 150-220 m,

7 Aug 1975 (fl), C.H. Dodson 5933 Sab MO, QCA, Pun Río Palenque Field Station, "s way between Que and Santo Domingo

de los Colorados, 200 m, 22 Feb 1974 (fl), A. Gentry 10109 (MO).

Cal ifi is a striki ies with ayes that are ee PUE green above with whitish veins, and

: ly

ewtarnciuislir

weakly [c MEN purple bawi is Sp d has not been colle

but it is clearly distinct from other members of ERE Hv fant It is recognizable by its e habil.

completely glabrous vegetative parts, its large, very thin leaves and slender, weakly herbaceous stems.

Solanum pacificum specimens have been identified as S. dolichorhachis, but S. dolichorhachis has wiry

woody stems, chartaceous to coriaceous leaves with el ra leaf bs and ena ce oe

2-3 times longer than wide vs. the weak-!

at the base, and globose to somewhat conical (but less than twice as long as wide) fruits of S. pac cum It

is also related to S. crassinervium, S. evolvulifolium, and S. loxophyllum, but is easily distinguished from these

species by the texture, shape, and size of the leaves. Solanum pacificum also bears a resemblance to the sym-

patric S. leptorhachis Bitter (sect. Geminata) due to its simple leaves, long, slender inflorescences, and small,

Tepe and Bohs, TI

eas

LI RE

BAV

ext

pt ppa

C. Inflorescence. D. Rud. E. Fl

513

E Stamens. 6. Inflorescence

A. Habit. B. |

514 | l of the Botanical R h Institute of Texas 3(2)

stellate, greenish-white flowers. However, S. leptorhachis is a free-standing shrub 1-2 m tall with unifoliate

sympodial units on flowering stems vs. a vine with plurifoliate sympodial units in S. pacificum.

The lowlands of Ecuador between the Pacific Ocean and the Andes are under large-scale cultivation

of cacao, papayas, bananas, African oil palm, etc., and very few areas of undisturbed habitat remain intact.

Most collections of S. pacificum are from the small, but well-preserved Centro Científico Río Palenque reserve.

There are no large reserves in western Ecuador (like Yasuní in the east) and it is likely that S. pacificum and

other plant species endemic to western Ecuador are surviving in small, isolated reserves. As a result, S.

pacificum is in critical danger of extinction due to habitat loss and every effort should be made to maintain

the reserves in this region.

Etymology.—Sol ificum i 1 after the Pacific lowlands of Ecuador where it is endemic, and

as a tribute to ET’s wife and frequent field companion María Paz Moreno.

Habitat and Distribution.—Solanum pacificum occurs in rainforest habitats in the Pacific lowlands of

Ecuador, 50—300 m in elevation.

Phenology.—Flowering specimens have been collected from Feb—Aug; the type collection, collected in

Feb, is the only fruiting specimen seen. It is likely that fruiting is more frequent than the collection record

indicates.

Solanum crassinervium Tepe, sp. nov. (Fig. 2). Tyre. ECUADOR. eee near Lita, 600 m, wet evergreen forest,

19 May 1987 (fl, fr), H.H. van der Werff 9496 (HOLOTYPE: QCNE!; i NY!)

eee | 1 fade fr: a A Gots Fe + | +1 “1 p. 5 n ax 141 Sd : 4 Pee D beet 1 1, E 1

J J > i ? I I L D

notabilis.

Vine or scandent shrub, climbing pee trees as high as 4 m via adventitious roots at the nodes; leafy

branches spreading to pendulous. St erbaceous to hase! NOD somewhat fleshy, glabrous to

very sparsely | pubescent and soon glabrescent Sympod ial unit ly unifoliate, not geminate.

Leaves simple, 3.5-14 x 1.5-8 cm, ca. 1-2 times as long as vids: gradually reduced in size toward the inflo-

rescence, ovate to ADOS somewhat fleshy, glabrous adaxially and abaxially, the secondary veins 5-7 per

side, conspicuous and prominent abaxially, the base rounded to truncate to cordate, sometimes oblique, the

margin entire, the apex shortly acuminate; petioles (0.2—)1—1.5 cm, glabrous. Internodes 1.5—5.5 cm. Inflo-

rescences 13 x 1-4 cm to ca. 6 x ca. 5.5 cm in fruit, stem-terminal to axillary to extra-axillary, unbranched

to branched, with 2-16 flowers (scars), all flowers apparently perfect, the axes glabrous; peduncle 0.1—0.5

cm long; rachis 0.1—2 cm; pedicels 4-12 mm in flower, 9— IS mm in fruit, only slightly enlarged apically,

glabrous to rarely very sparsely pubescent, ee ni ly, articulated at base. Flowers with the

calyx 2.5-3.5 mm long, glabrous, tl ] 1, the tube 2.5—3 mm long, the lobes 1.5-2.5 x 1.5-2

mm, deltoid, acute to acuminate at tips, white to pale pink; fruiting calyx somewhat accrescent, the lobes

1.5-2 x 1.5-2.5 mm. Corolla 1.5-2 cm in diameter, 5-8 mm long, stellate, somewhat fleshy, white, the lobes

5—8 x 2.5-3 mm, planar at anthesis, acute to acuminate at apices, glabrous adaxially, sparsely pubescent

near the apex abaxially, the margins densely ciliate. Stamens equal, with filaments 1-1.5 mm long, glabrous,

free; anthers 3-4 x 1.2-1.5 mm, oblong, not connivent, yellow, the pores directed distally, opening into

short longitudinal slits with age. Ovary glabrous; style 4—6 x ca. 0.3 mm, glabrous, cylindrical, sometimes

deflected to one side of flower (Fig. 1D); stigma capitate. Fruit (immature) 0.7-1 x 0.7-0.9 cm, ovoid to

nearly globose, slightly flattened, somewhat pointed at apex, green to pale orangish to brownish at maturity,

nde Seeds 2-2.2 x 1.8-2 mm, flattened-reniform, tan, the surface minutely reticulate-rugulose.

idi ined: COLOMBIA. Nariño: Mpio. Barbacoas, Corregimiento Altaquer , Vereda El Barro, Reserva Natural Río

Nambi, -— derecha del Río Nambi, 1°18'N, 78°08'W, 1325 m, 1 Dec 1993 (f, P. Franco et di 4707 (NY); Mpio. Tumaco, Resguardo

de Albi, lado izquierdo del Río Albi, 1°22'N, 78°28'W, 220—280 m, 12 Nov 1995 (fl, fr), B.R. Ramirez et al. 8826 (NY); Mpio. Barbacoas,

Resguardo Indígena de Saundé, 1?30'N, 78?20'W, 350 m, 21 Jan 1996 (tr), B.R. Ramírez et al. 9699 (NY). ECUADOR. Carchi: Cantón

Tulcán, Parroquia Tobar Donoso, Reserva Indígena Awa, Centro EI Baboso, 0°53'N, 78°25'W, 1800 m, 17-27 Aug 1992 (fr), G. Tipaz

et al. 1813 (BM, NY, QCNE); border area between Prov. Carchi and Esmeraldas, 20 km ues Lita on Has i Alto f 550 m, 25

Jun 1991 (£D, H. van der Werff et al. 11972 (MO, NY, QCNE). Esmeraldas: Bilsa Biolog Mach km NW of

Quinindé, 3 km W of Santa Isabel, 0°22'N 79°45'W, 600 m, 26 Sep 1994 (fr), J.R. Abbott 15256 ( 10); San Lorenzo, Reserva Étnica Awá,

Tepe and Bohs, TI j f Sol ( t Herpystichum)

[c mH

ura Iu PH

AD OP

Fic. 2. Solanum crassinervium Tepe. A. Habit. B. Inflorescence. C. one E HOWER E. PONER ; longitudinal Pec F. Stamens. G. Infructescence biu

immature). (A, G lili WO van aer Meri ff 11972 [M0] [photo]; pe 2729 [photo]; D d tern

[p fi [M0] Fj fepe 2729 [photo].

Parroquia Alto Tambo, Centro de la Union, Cañon del Río Mira, 0°52'N 78°26'W, 250 m, 22 Mar 1993 (fl), C. Aulestia & M. Aulestia

1431 (QENT); Cantón Q d ab Bilsa Biological , de Macl 5 km W of Quinindé, 5 km W of Santa Isabel, reserve

d,1 the Río Cube tributar and the E-bearing boundary crossing the Río Cube, 0°21'N, 79°44'W,

400—600 m, 26 Sep 1994 (fr), M. S. Bass 6 N. Pitman 68 (BM, NY): Bilsa Biological Station, Montañas de Mache, 35 km W of Quinindé,

5 km W of Santa Isabel, 0°21'N 79°44'W, 400—600 m, 6 Dec 1994 (f£D, M.S. 5n & N. T 289 P Se km 321 Sene iiis

from Ibarra to San Lorenzo, 1°N, 78°W, 350 m, 5 May 1982 (fD, B.M. Boom 1374 (N CA); C é, Bilsa B

Montañas de Mache, 35 km W of Quinindé, 5 km W of Santa Isabel, Monkey Bone Trail Q?21'N, 79944 W, 400— 600 m, 15 Sep 1994

heavy LANA AL yl

f Texas 3(2)

516

(fr), J.L. Clark & B. A 55 (QCNE); Canton Quinindé, Bilsa Biol 1 St Mache Mountains, 35 pe aad ee

Santa Isabel, 0°21'N, 79°44 Ww. 400- 600 m, 24dan 1995 (£D, J.L. Clark 412 (BM, NY, Q OCNE): Cant

ache Mountains, 35 km W of Sa 5 im W 2s Santa pee ona idi W, 500m m, “18 Feb 1996 > 69. ] L. Clark 2121 (BM, NY,

ee Cantón Quinindé, Mache-Chindul W of Quinindé, 0°21'N,

79°44'W, 500 m, 1-10 Jan 1997 (fr), J.L. Clark et a 3762 MO, NY, |, QCNE) 10 km W al Lita on road to San Lorenzo, 0°55'N, 78°30'W,

800 m, 12 May 1991 (fD, A. Gentry et al. 69984 (MO, NY) tal 1, finca of Dr. La Lama, 13.5 km S of

Lita, 0°49'N, 78°26'W, 1220-1350 m, 2 Nov 1992 (fl, D, J.L. Luteyn et al 14744 (MO, NY, QCA); Cantá El

Páramo d PR Estación Biológica Bilsa, dd 7"W, 79°42'40.4"W, 580 m, 18 Feb- 5 Mar 1995 (fo. W. Palacios et al. 13548 (MO,

Y, , Bilsa Biol de Mache, 35 km W of Quinindé, 5 km W of Santa Isabela, Monkey

ay OPZIN, 79*44W, 400—600 m, 1 Dec 1994 (£D, N. Pitman & M. Bass 1091 (MO, NY, QCNE); San Lorenzo, Territorio Awa,

centro Mataje, 1°11'44"N, 78°34'29"W, 200 m, 17 Nov 2000 (£D, W. Ramirez et al. 12 (NY), 15 (NY); Bilsa Biological Station, 5 km W

Sta. Isabel, 0°20'49"N 79°42'41"W, 540 m, 14 Feb 2009 (fl, fr), S. Stern 400 (QCNE, UT); Bilsa Biological Station, 5 km W of Sta. es

0°20'49"N 79°42'41"W, 540 m, 13 Feb 2009 (fl, fr), EJ. Tepe & S. Stern 2729 (BM, NY, QCA, QCNE, UT)

Solanum crassinervium is the most robust species in sect. Herpystichum and is recognizable by the somewhat

fleshy texture of the leaves, stems, and flowers, its ovate to elliptical leaves with conspicuous secondary

veins in fresh and dried material, and occasionally branched inflorescences. Its distribution is restricted to

the Chocó region of SW Colombia and NW Ecuador, a biodiversity hotspot with one of the highest species

diversities TA area en et a. 2000), but it is apparently quite common in some parts of its range.

ly related to S. evolvulifolium and S. loxophyllum, but differs from both species

in its robust habit, ovate leaves (vs. mostly oblong), fleshy calyx and corolla, and inflorescences that may be

simple and branched within an individual. The leaves, petioles, and internodes are many times larger than

those of S. evolvulifolium. Solanum loxophyllum is larger in stature than S. evolvulifolium, but S. crassinervium

is easily distinguished from the former based on leaf and inflorescence characters. The leaves of S. crass-

inervium, like those of S. loxophyllum, are somewhat fleshy, but S. crassinervium has more secondary veins

(5—7 vs. 3-4 pairs) and these are prominent abaxially, whereas those of S. loxophyllum are obscure within

the fleshy leaf blade. These leaf characters are apparent in both fresh and dried material. The ER stout

Als um «1 1 1 m

inflorescences of S. crassinervium differ from those of S. loxophyllum, whi PI

very slend ] delicate. Furthermore, inflorescences of S. loxophyllum are typically beak on Wien les

parts of the stem, as compared to those of S. crassinervium, which appear to always be produced in the leafy

parts of the stem. Solanum crassinervium has sometimes been identified as S. siphonobasis Bitter, a member of

Solanum sect. Anarrhichomenum Bitter, but differs from this species in having > 2 flowers per inflorescence

(S. siphonobasis has 1-2 flowers per inflorescence), and in the fleshy texture of the leaves, stems, and flow-

ers. Species of sect. Anarrhichomenum are also node-rooting vines, but they differ from sect. Herpystichum

primarily in possessing pseudostipules at the base of the petioles (Correll 1962).

Etymology.—The epithet crassinervium describes the prominent secondary veins that help distinguish

this Pina cvs its Pos pues

anum crassinervium occurs in low land and { ] { habitats in

+l +

the Chocó region of Colombia and Ecuador, including the Mache-Chindul mountain range in

Ecuador, 150-1800 m in elevation.

Phenology.—Flowering apparently occurs year-round; fruiting specimens have been collected from

Jan—Feb, and Sep—Dec.

Limoncocha,

Solanum limoncochaense Tepe, sp. nov. (Fig. 3). Tee. ECUADOR. S

in wet primary forest near NW corner of lake, 250 m, 22 Jan 2009 (fl, fr), EJ. Tepe & S. Stern 2627 (HOLOTYPE: QCA!; , BM!,

MO!, NY!, QCNE!, US!, UT!)

ct? I 4:4 LE , 11 :1 = "n 1.1 x ry] :1 ra b eec: 42 ff,

J o o >

Terrestrial herb, trailing, rooting at the nodes. St lend

ate, not geminate. Leaves simple, the blades 3.5-7 x 3.5-8 cm, slightly wider than long, mound, coi

fleshy, glabrous; venation palmate with 5 (—7) veins, the base cordate, the margins entire, slightly revolute

on some leaves, the apex rounded to obtuse to shortly acuminate, petioles 3-16 cm, glabrous. Internodes

, glabrous Sympodial it y plur ifoli-

Tepe and Bohs, TI ies of Sol (sect Herpystichum) 517

Fic. 3. Solanum limoncochaense Tepe. A. Habit. B. Inflorescence. C. Flowers. D. Stamens. E. Infructescence. F. Fruit (immature). (All drawn from M.T.

Madison et al. 5327 [MO]).

2.5-20 cm. Inflorescences 4-8 x ca. 3 cm, extra-axillary, unbranched, with 23 flowers, all flowers appar-

ently perfect, the axes glabrous; peduncle 1.3-4.5 cm, slender; rachis 0.9-1.5 cm; pedicels 15-25 mm in

flower, 25-30 mm in fruit, slender, glabrous, spaced ca. 15 mm apart, articulated at the base. Flowers with the

calyx 2.5-4.2 mm long, the tube 1-1.5 mm long, the lobes 1.5—2 x ca. 1.2 mm, rounded, acuminate at tips,

glabrous to sparsely pubescent abaxially, densely pubescent adaxially, purplish; fruiting calyx very slightly

accrescent, the lobes 1-1.2 x 1.5—2 mm, truncate—acuminate. Corolla 1-1.6 cm in diameter, 5-8 mm long,

stellate, membranous, white, the tube 1.5-2 mm, the lobes 6-10 x 1.5-3 mm, lanceolate, narrowly acute

J J £ al Daa a Ip LI B8. don ndo Siri

518 f Texas 3(2)

at tips, the apex papillose adaxially and abaxially, the margins ciliate apically. Stamens equal, the filaments

ca. 1 mm, glabrous; anthers 2-2.5 x ca. 1 mm, slightly tapered, sagittate basally, not connivent, yellow, the

pores large and directed distally, opening into longitudinal slits with age. Ovary sparsely papillose; style

2-2.5 x 0.3 mm, straight, cylindrical, stout, sparsely papillose in lower half; stigma capitate, somewhat

2-lobed. Fruits 1-3 x 0.6-3.2 cm, ovoid-rhomboid, flattened, greenish-brown to purplish near apex when

immature, bronze-brown when mature, sparsely pubescent with hairs < 0.1 mm to glabrous when mature,

the apex truncate to emarginate, strongly fragrant with a sweet, heavy scent, very juicy, the flavor sweet.

Seeds 2—2.5 mm in diameter, lenticular, light reddish-brown, the surface minutely rugose.

Additional specimens examined: ECUADOR. Napo: Limoncocha on Rio Napo, 300 m, 18 Oct 1974 (f1), B.A. Drummond 7350 (MO);

environs of Limoncocha, 240 m, 16 Jun 1978 (fl, fr), M.T. Madison et al. 5327 (AAU, F, K, MO, NY, QCA, SEL); near northwest corner of

lake, Limoncocha, Sep 1969 (f1), R.N. Mowbray 699104 (MO).

LS | LT 1-1

Solanum li ] ] ] t a3 dictinctive orem of ies f 1 Solanum cect b x AA

D o r E L

ized by herbaceous, ground-trailing stems that root at the nodes and bizarre, flattened, arrowhead-shaped

fruits; this group includes S. pentaphyllum, S. phaseoloides, S. dalibardiforme, and S. trifolium. Solanum limon-

cochaense can be distinguished from other species in this group by its simple leaves and stellate flowers.

Vegetatively, it is most similar to S. dalibardiforme from central Colombia, and most specimens have been

annotated as this species. Both species have simple, cordate leaves, but S. limoncochaense is distinguish-

able because all vegetative parts are glabrous, and has stellate rather than rotate corollas. Furthermore, S.

limoncochaense is found at low elevations, whereas S. dalibardif is found at elevations » 2000 m. Solanum

limoncochaense grows in dense, presumably clonal patches on uibs rainforest floor and over fallen trees. It is

apparently a weak climber and living plants in the field were seen climbing trees to no more than 1 m from

the ground (EJ. Tepe & S. Stern, pers. obs).

Mature fruits are bronze-brown in color, very juicy, and have a strongly sweet aroma and flavor. The

fruits lie on the ground when stems are trailing, or are pendulous on climbing stems. The dispersal agents

of these fruits are unknown, but based on the strong scent, color, and presentation, it is likely that they are

eaten by rainforest rodents such as the guatusa or agouti (genus Dasyprocta; W. Haber, pers. comm.).

Solanum limoncochaense is endemic to Ecuador and all known collections are from the ‘terra firme’ for-

ests at the northern side of the Limoncocha lake within the Reserva Biológica Limoncocha, a small reserve

of 4613 hectares (including the lake which is ca. 250 hectares). Much of the rest of the reserve is made up

of swamps and seasonally flooded forests, and S. limoncochaense was not encountered in any of the flooded

forest habitats. The reserve is surrounded by cultivated land and oil platforms, whose effects on the habitats

within it are unknown. If this area is truly the only locality for this species, then it should be considered

critically endangered; however, the Reserva de la Biósfera Yasuní is nearby, and much of this enormous re-

serve has not yet been explored botanically. It is hoped that this species is more widespread than it appears

to be based on the ESOS currently available.

1 "ni LO

the I] aguna de Li Ecuador,

the geographic locality of all known collections of the species.

Habitat and Distribution This species is apparently endemic to Napo and Sucumbios Provinces,

Ecuador, near the Laguna de Limoncocha, where it grows in primary forests and clearings, 240—300 m in

elevation.

Phenology.—Flowering specimens have been collected in Jan, Jun, Sep, and Oct; fruiting specimens

have been collected in Jan and Jun.

ACKNOWLEDGMENTS

We thank Stephen Stern and Freddy Villao for assistance in the field; the herbaria AAU, BM, F, K, MO, NY,

and SEL for loans of specimens; Ann Kelsey from UT for assistance with loans; David Neill and the staff at

QCNE for assistance with logistics in Ecuador; Katy Coral and Hendry Moya of the UISEK Estación Amazónica

at Limoncocha; Freddy Villao and the staff at the Centro Cientifico Rio Palenque; and Carlos Aulestia and

Tepe and Bohs, TI i f Sol ( t Herpystichum) 519

the Fundación Jatun Satcha for access to the Bilsa Biological Station; Bobbi Angell for the illustrations; and

the two anonymous reviewers whose questions and comments improved this manuscript. This work was

supported by the NSF through the PBI: Solanum grant, DEB-0316614, to LB.

REPERENCES

Biter, G. 1912. Solana nova ve: minus cognita ll. Repert. Spec. Nov. Regni Veg. 11:1-18.

Brrrer, G. 1913a. Solana nova vel minus cognita VII. Repert. Spec. Nov. Regni Veg. 11:481—491.

Brrrer, G. 1913b. Solana nova vel minus cognita X. Repert. Spec. Nov. Regni Veg. 12:49-90.

CorreLL, D.S. 1962. The potato and its wild relatives. Contr. Texas Res. Found. Bot. Stud. vol. 4. Texas Research

Foundation, Renner.

JØRGENSEN, PM. AND S. LEON-YANEZ (EDs.) 1999. Catalogue of the vascular plants of Ecuador. Monogr. Syst. Bot., Mis-

souri Bot. Gard. 75:1-1182.

Knapp, S. 2007. Solanum coalitum (Solanaceae), a new endemic species from southern Ecuador. Novon 17:

212-216.

Knapp, S. 2008. A revision of the Sol pecies grout | taxonomic additions to the Geminata

clade (Solanum: Solanaceae). Ann. Missouri Bot. Gard. 95:405—458.

MONTUFAR, R. 2000. Solanaceae. In: R. Valencia, N. Pitman, S. León-Yánes, PM. Jørgensen, eds. Libro Rojo de las

plantas endémicas del Ecuador 2000. Herbario QCA, Pont. Univ. Católica del Ecuador, Quito.

Myers, N., R.A. MiTTERMEIER, C.G. MITTERMEIER, G.A.B. DA FONSECA, AND J. Kenr. 2000. Biodiversity hotspots for conservation

priorities, Nature 403:853-858.

STERN, S.R. AND L. Bous. 2009. Two new species of Solanum from Ecuador and new combinations in Solanum sec-

tion Pachyphylla (Solanaceae). J. Bot. Res. Inst. Texas 3:503-510.

Weese, T.L. AND L. Bous. 2007. A three-gene phylogeny of the genus Solanum (Solanaceae). Syst. Bot. 32:

445—463.

520 I I nfahkn Das ‘in le Imctituitn AET

BOOK REVIEW

MARGARET B. GARGIULLO. 2007. A Guide to Native Plants of the New York City Region. (ISBN 0-8135-

4042-9, hbk.). Rutgers boni Min 100 Joyce Kilmer Ave # 1, Piscataway, New Jersey 08854, U.S.A.

(Orders: ww g , 1-800-848-6224, 1-800-272-6817 fax). $49.95, 306 pp., 131

line drawings, 6 3/8 "x 9 1/2"

Presented in this unique guide is a "detailed look at the full scope of flora that i j this regi 1 available for propagati ipi

+l NES NC e F xi : : ta NI th America © T

that ld pl tificial limit on the pl I ld 1 ble in this day andit time. But, as New York Citv has lost neatly 43%

of its native planes, d this actually Id to i tł ] mE f

ed into two basic parts. Part 1 is an dn M at the planis of New York Ey and vicinity, focusing on trees,

shrubs, vines, herbs, graminoids, and ferns g I i ypes. These are followed by

A J Al:

a p til L VV RAY EM LAA VM

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From the e publisher: g I , ranging

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size, flower color, blooming time, 1 its y ib] i tland mitigation, erosion oand ] natural torati Some plants

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Pennsylvania, Delaware; i^ Maryland.”

Recommended for p h collections, especially those dealing with landscape design and preservation of

habitat. This book is not ae geographically to the New England area but is applicable to use elsewhere, as inspiration if nothing

else.

ra ined B. Gargiullo has a Ph.D. ini lant l gy dI tł d l articl i ive Į lant 1 the historical fl f

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2006.— Gary Jennings, Librarian, Botanical R h Institute of Texas, Fort Worth, Texas 76102-4025, U.S.A.

J. Bot. Res. Inst. Texas 3(2): 520. 2009

STENARIA SANCHEZII (RUBIACEAE), A NEW AND ENDANGERED SPECIES

FROM SONORA, MEXICO

David H. Lorence

National Tropical Botanical Garden

apalina Road

3530 Pap

Kalaheo, Hawaii 96741, U.S.A.

lorence@ntbg.org

ABSTRACT

4 f .1 A 1 A c DAFYT 11 ] 1 J f. i , Mexico Ca 1

i e L

T + f FAT 1 1 11 +l kaa 0 mrn leno A Akl 1; ] T^ t Ix TEL DNA d. oer seers | ta

4 * 2

small population size, and vulnerability to threats from grazing animals and Habitat ies it i idered to be critically endangered

(CR) based in IUCN Red List criteria.

Key Wonps: Stenaria, Rubiaceae, Hedyotideae, Spermacoceae, Sonora, Mexico, critically endangered, IUCN Red List

RESUMEN

Se describe una nueva dn p ume norteamericano Stenaria (Raf). Terrell de Sonora, México. Stenaria sanchezii Lorence se

l ; I 9 mm de largo, y semillas de oblongas a elipsoides. Debido a

ingida con solamente dos individ i debido a la v oo al ganado y pérdida de ae se

caracter iza dd

T h ítico (CR) ] los criterios de la tista Roja de IU

Veet JAJAL

During the course of field work in Sonora, Mexico by Jesús Sanchez-Escalante, Richard Felger, and collabo-

ns were made new species of Rubiaceae belonging to the Spermacoceae s.

lat. ER AMA tribe. Based on gor bod characters it is referable to ix Pens Stenaria and is here

ii Lorence. The combination of woody herb bit, thin non-leathery

leaves, foliaceous calyx dies tapering to a narrow, attenuate base, and corollas with PN long

tubes to 9 mm Une ID odia 5. Sdn eee EUM its congeners

Based on to the group of Mexican and North American

species in the tribe Hedyotidese. currently incude d by some VPE ME in Spermacoceae s. lat. (Groeninckx et

al. 2009). Members of this alliance form a relatively large and taxonomically complex group that is not yet

well resolved. Historically, Standley (1918), in his treatment for the North American flora, recognized 48

species which he assigned to either Houstonia L. or Oldenlandia. Subsequently Lewis (1961, 1962) merged

all North American (including Mexican) members of the alliance into Hedyotis L. However, Hedyotis in the

strict sense is an Old World genus ranging from SE Asia to the Caroline Islands of Micronesia with two

widespread species in western Polynesia (Church 2003; Terrell et al. 2005). Terrell and Robinson (2003)

concluded that the genus is restricted to Asia and the Pacific region with the type species, Hedyotis fruticosa

La ae from Sri ee

TNR t referable to Hedyotis, Houstonia, or Oldenlandia,

Terrell and collaborators resurrected or seda new genera including Carterélla Terrell, Oldenlandiopsis

Terrell & W. Lewis, Stenaria (Raf) Terrell, and Stenotis Terrell (Terrell 1987, 1990, 1996, 2001a, 2001b;

Terrell & Lewis 1990; Terrell & Robinson 2006).

Although generic delimitations in this complex have not been fully resolved, recent morphological and

molecular studies are contributing to a greater understanding of the group: Initial studies of iral a

chloroplast DNA by Church (2003), suggest that the North American I

but belong to the same lineage as Stenaria. The picture is complex, however, with radiations in dn Pease

accompanied by descending aneuploidy and apparently other characters such as annual versus perennial

J Rot Rec Inct Tavac 3{2): E71 — 525. 2009

522

habit, heterostylous versus monostylous breeding systems, and evolution of self fertilization, which seem

to have originated multiple times (Church 2003). More recent phylogenetic studies of the Hedyotideae

s. lat. and Spermacoceae based on chloroplast, plastid, and nuclear DNA regions (Karehed et al. 2008;

Groeninckx,et al. 2009) support Hedyotis as being restricted to the Paleotropics and Houstonia and Stenaria

as being American.

Floral, mii and un TIGE, of this new species indicate it is referable to the group of species

which includes d related species. These are currently placed in Stenaria (Raf.) Terrell,

a genus of six species (indludine this new one) restricted to Mexico and the southwestern United States

(Terrell 20012). The phylogenies of Karehed et al. (2008: 850) and Groeninckx et al. (2009: 118) based on

plastid DNA data place Stenaria nigricans (Lam.) Terrell in the Arcytophyllum-Housonia clade.

Until the generic delimitations of this complex group are more clearly resolved, it was decided to place

the new species in Stenaria. Based on its woody perennial habit, thin textured leaf blades, and subglobose

capsules, Stenaria sanchezii seems closest to S. nigricans, which differs in having shorter corollas 2-8.5 mm

long with tubes (1.5-)2— 4(-5. 5) mm long (Terrell 20012). Corollas are much longer in the new species, al-

though only two mature corollas were available for study. The foliaceous calyx lobes tapering to a narrowly

attenuate base in S. sanchezii are also very distinctive and in combination with the other characters readily

separate it from its congeners.

Stenaria sanchezii Lorence, sp. nov. (Fig. 1). Tyre: MEXICO. Sonora: Municipio de uaa Aguaje de Robinson, 7 km (by

air) north of San Carlos; 28°2'8"N, 111%6'34"W 80 m, 1 Feb 2001, R.S. Felger, J.J. Sdncl , L. Moreno-Moreno, O. Gutiérrez-

Rochin, & M. Espericueta-Betancourt 01-111 (Hototyre: PTBG 44233; ISOTYPES: ARIZ, USON).

Species Stenariae i dapi ud br t ae obis calycinis foliaceis anguste obl latis att tis, 33.5 mm longis,

0.5—0.8 mm ate

Woody perennial herbs or subshrubs 20—60 cm tall, spreading to 120 cm across, basal stems woody, 2.53

mm in diam., bark pale brown, exfoliating, leafy twigs 0.4-0.8 mm diam., densely hirtellous-puberulent

with short white trichomes to 0.1 mm long. Leaves opposite, sometimes pseudoverticellate with 1-2 smaller

leaf pairs in axils, petiolate; petioles 1-3 mm long, 0.15—0.3 mm diam., winged distally, adaxially flattened,

minutely puberulent; blades oblanceolate to oblong-elliptic, base cuneate, attenuate and decurrent along

pos apex abruptly us acuminate to apiculate, thinly chartaceous, both surfaces minutely puberulent

Costa d base, glabrescent, venation inconspicuous, secondary veins 3—5 per

side, —M— higher order venation loosely reticulate; stipules interpetiolar, narrowly triangular,

sheath 0.2-0.3 mm long, body c. 1 mm long, apex slender, glandular-tipped, margins with 2-4 gland-tipped

setae. Inflorescences terminal, monochasial, corymbiform, 3—5-flowered, axes puberulent, pedicels 1-4.5 x

0.2 mm, bracteoles subulate or fimbriate, 0.3-0.7 mm long; flowers apparently hermaphroditic; short-styled

flowers with hypanthium broadly ellipsoid-compressed, 0.8-1 mm long and wide, sparsely puberulent,

8-ribbed; calyx tube lacking, calyx lobes 4, equal or subequal, narrowly oblanceolate, 3-3.5 x 0.5-0.8 mm,

tapering to narrow base 0.5—0.6 mm wide, apex acuminate, Bands venose; corolla in bud (possibly not

fully expanded) 10 mm long including the lobes, obtuse, at antl tube 9 mm long, 0.8 mm wide medially

and 1.3 mm wide distally, lobes 4, narrowly ovate- te-oblong, 2-2.5 x 1-1.1 mm, acute; stamens 4, attached just

below mouth of tube, subsessile, antl llipsoid, 1-1.2 mm A Hpse exserted; style included, 6 mm long,

puberulent in basal 1/3, sti 0.8 mm long, slightly bilobed; long tseen Cas 2-3 x 2-2.8

mm, subglobose, dehiscence loculicidal, cont g ds; calyx lol tent, 3.5-4 x 1.4-1.5

mm; seeds (possibly not fully mature) non-crateriform, moderately copiis ced. with puncniorm hilum,

oblong to ellipsoid or broadly ellipsoid, 0.6-0.8 x 0.3—0.5 mm, testa foveolate-reticulate, dark brown.

Etymology.—This new species is named for its first collector, J. Jestis Sánchez-Escalante, Curator of

Herbario de la Universidad de Sonora (USON) and founder of the Asociación para las Plantas Nativas de

Sonora, A.C. in Hermosillo, in recognition of his efforts to develop USON as a major botanical resource in

northwestern México to study and conserve the flora of Sonora, and to inspire young biologists to study

plants.

Lorence

PF GE aur! i

PTT

rap 5 6 683 tt

je ez,

job, P, Bb, Su iE I Iu Mu Iw.

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UPDATED

ENTERED IN

DATABASE

523

Steunnacia. Sew Ce 2 otme

Wots

DET: D. Lorence (PTBG) 2009

National Tropical Botanical Garden Herbarium (PTBG)

PLORA OF SONORA, MEXICO

Hedyotis sp. nov.

Aguaje de Robinson, 7 km (hy air) north of

= Carlos: north- 2 palm oasis with

randegeei oran desertscrub;

or 02 ‘a aios’ uw 80 = elevation.

se 2 plants) subshr e 1,2 m across,

0.6 m tall, the eter alter, scandent

iat dead palm

R.S, Felger ĝl- il, ie Sanchez E., b.

Moreno H., 0. Gutierres R., M.

Espericueta B,

: Bunicipio de Guaymas

1 February 2001

i. 01-111, PTBG).

Fic. 1. Hol

[1 fal Dos a it LI PAN ET >

524 Jou lexas 3(2)

Distribution and habitat.—Known only Hom the type d in Sonora. This new species is extremely

rare and localized in microhal n slopes on rth side of the Sierra El Aguaje. The general vegetation

is desertscrub in the Central Gulf Coast subdivision of the Sonoran Desert (Shreve 1964). Felger's (1999)

flora of Nacapule in the southeaster n Sierra El Aguage provides a good general description of the vegetation

and associated species. Stenaria sanchezii occurs on rocky substrate in a shady canyon bottom and also in

a north-facing palm oasis with Brahea brandegeei (Purpus) H. E. Moore and Ficus petiolaris (Kunth) subsp.

palmeri (S. Watson) Felger. Four to five plants were seen in 2000, but only two plants were observed at the

same locality in 2001. The area is heavily grazed by cattle, a primary threat to this new species.

Conservation status.—When evaluated using the IUCN criteria Lior Gona (IUCN 2001, see also

www.iucnredlist.org/info/categories_criteria2001), St into the Critically Endangered (CR)

category, which designates species facing the ches: risk of extinction in the wild. IUCN Red List Category:

Critically Endangered (CR; Bla; B2a, B2b i-iii): Bl, extent of occurrence estimated to be less than 100 km’,

and Bla known to exist at only a single location. B2, area of occupancy estimated to be less than 10 km’,

and B2a, a single population known. B2b G-iii), habitat continuing decline inferred. The suitable habitat

for S. sanchezii at a single locality is indicated as an endangered environment, threatened by human activity

(firewood harvesting and fire), grazing, and invasive plants, reducing the extent of the Sonoran desert scrub

habitat.

PARATYPE: MEXI CO. S 1 io de Guaymas de Robinson, 7 km (by air) N of San Carlos, 28?2'8"N, 111°6'34"W, 80 m,

2 Mar 2000, J. Sanchez, L. Moreno, M. Esvericuptasm (PTBG 44234, USON).

ACKNOWLEDGMENTS

I wish to thank Jesús Sánchez-Escalante for fieldwork leading to the discovery of this new taxon, Tom

Van Devender for bringing this new species to my attention, Richard Felger for information on habitat and

threats, and the reviewers Edward Terrell and Charlotte Taylor for valuable discussions and comments on

the manuscript. Curators of the herbaria ARIZ and USON are thanked for making the specimens available

for study.

REFERENCES

Church, S.A. 2003. Molecular phylogenetics of Houstonia (Rubiaceae): descending aneuploidy and breeding

system evolution in the radiation of the lineage across North America. Molec. Phylogen. Evol. 27:223-238.

Feicer, R.S. 1999. Flora of Cañón Nacapule: a desert-bounded tropical canyon near Guaymas, Sonora. Proc. San

Diego Soc. Nat. History 35:1-42.

GROENINCKX, l., S. Dessein, H. OCHOTERENA, C. Persson, TJ. Money, J. KAREHED, B. BREMER, S. HUYSMANS, AND E. Smets. 2009.

Phylogeny of the herbaceous Tribe Spermacoceae (Rubiaceae) based on plastid DNA data. 2009. Ann.

Missouri Bot. Gard. 96:109-132.

KAREHED, J., |. GROENINCKx, S. DESSEN, TJ. MOTLEY, AND B. Bremer. 2008. The phylogenetic unity of chloroplast and nuclear

DNA markers and the phylogeny of the Rubiaceae tribe Spermacoceae. Molec. Phylogen. Evol. 49:843-866.

Lewis, WH. 1961. Merger of the North American Houstonia and Oldenlandia under Hedyotis. Rhodora 63:

216-223.

Lewis, WH. 1962. Phylogenetic study of Hedyotis (Rubiaceae) in North America. Amer. J. Bot. 49:855-865.

Sureve, F. 1964. Vegetation of the Sonoran Desert. In: F. Shreve and I.L. Wiggins. Vegetation and flora of the Sonoran

Desert. Stanford Univ. Press, Stanford, CA. Pp. 9-186.

STANDLEY, PC. 1918. Rubiaceae (pars). North American Flora 32(1):1-86.

TerRELL, E.E. 1987. Carterella (Rubiaceae), a new genus from Baja California, Mexico. Brittonia 39:248-252.

TerreLL, E.E. 1990. Synopsis of Oldenlandia (Rubiaceae) in the United States. Phytologia 68:125-133.

TerreLt, E.E. 1996. Revision of Houstonia (Rubiaceae-Hedyotideae). Syst. Bot. Monogr. 48:1-118.

Terret, E.E. 2001a. Taxonomy of Stenaria (Rubiaceae: Hedyotideae), a new genus including Hedyotis nigricans.

Sida 19:591-614.

Lorence, St i hezii, | les f Mexi 525

TERRELL, E.E. 2001 b. Stenotis (Rubiaceae), a new segregate genus from Baja California, Mexico. Sida 19:899—91 1.

TERRELL, E.E. AND W.H. Lewis. 1990. Oldenlandiopsis (Rubiaceae), a new genus from the Caribbean Basis, based on

Oldenlandia callitrichoides Grisebach. Brittonia 42:185-190,

TeRRELL, E.E. AND H. ROBINSON. 2003. Survey of Asian and Pacific species of Hedyotis and Exallage (Rubiaceae) with

nomenclatural notes on Hedyotis types. Taxon 52:775—782.

TERRELL, E.E., H.E. ROBINSON, W.L. WAGNER, AND D.H. Lorence. 2005. Resurrection of genus Kadua for Hawaiian Hedyo-

tidinae (Rubiaceae), with emphasis on seed and capsule characters and notes on South Pacific species, Syst

Bot. 30:818-833.

TERRELL, E.E. AND H. Rosinson. 2006. Taxonomy of North American species of Oldenlandia (Rubiaceae). Sida

22:305-329.

526 ] l af tha Rat izal D kh Imetitut 3/5)

f Texas 3(2)

BOOK REVIEW

G. LEDYARD STEBBINS, WITH V.C. HoLLowELL, V.B. SMocovitis, AND E.P. DUGGAN (eps). 2007. The Ladyslipper and

I. (ISBN 978-1-930723-65-8, hbk.). Missouri Botanical Garden Press, PO Box 299, St. Louis, Missouri

63166-0299, U.S.A. (Orders: www.mbgpress.org, 1-314-577-9547, orders@mbgpress.org). $35.00,

173 pp., 24 b&w photos, 7 1/4" x 10 1/4".

G. Ledyard Stebbins—one doesn’t read far in biology without coming across this name. The Ladyslipper and I is his autobiography,

nuce LRL m like his e on Pih m ie own ee uiid id ns was a manuscript that he had

Fomrritta

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y biology.—Robert J George, Botanical Research Institute of Texas, Fort a Texas 76102-4025,

U.S.A.

J. Bot. Res. Inst. Texas 3(2): 526. 2009

MUHLENBERGIA TARAHUMARA (POACEAE: CHLORIDOIDEAE: CYNODONTEAE:

MUHLENBERGIINAE), A NEW SPECIES FROM CHIHUAHUA, MEXICO

Paui M. Peterson J. Travis Columbus

Department of Botany Rancho Santa Ana Botanic Garden and

National Museum of Natural History Claremont Graduate yin

Smithsonian Institution 1500 North College Ave

Washington, DC 20013-7012, U.S.A. Claremont, California 91711 e e USA,

peterson@si.edu j.travis.columbus@cqu.edu

ABSTRACT

Muhlenbergia taral P.M. Peterson & Columbus, sp. nov., is described and ill ] i new Le occurs on n i

ridgetops, and white-tuff rock outcrops in the Si Madre Occidental and is | ly f

huahua, Mexico. Based phological and phylogeneti lysis of plastid and ncs DNA Sequens aes we conclude that the

new I 3 1 1 I A egopog 1:31 +l es 4 ] eL 153 r I LA. T g g Muhle nbergia Muhlenbergia

" 1 Bere by f A gopog y aj był i gI S 1.1 1 E 13 per culm, 1 Leg 1 _one-f]

glumes 3-5.8 mm long, shorter than the | d ly 1 late; | 3.6-7.3 mm long, lanceolars paleas 3-4.6 mm e

and anthers 2.2-2.8 mm long, yellowish orange.

RESUMEN

Se Scere e ilustra O NENAS P.M. ida & Columbus, sp. nov. La nueva es] hal cuestas r en

eae AAA mi | 1 1 4 "m APA m

OS y la Sierra dental y p

A Y 1 q. nl -:1 1 2 L T E 1 ej: E. E:1 eye i| 1 = J A TAAT 1 A | 1

, México g y g p y nuclear ‘

lias 1 : 1 dubbed dc tees d ] Jad es s S ] 1,1

Or dc

bonu subgen Muhlen bergia Muhlenberei } lifi le A braid m qe Imo, cada 3

espiguillas unifloras terminales; anta de 3-5.8 mm de largo, más as que el lema y estrechamente lanceoladas lemas de 3.6—7.3

mm de largo, lanceoladas; páleas de 3-4.6 mm de largo; y anteras de 2.2-2.8 mm de largo, amarillento-anaranjadas

On a 2003 collecting trip to the Sierra Madre Occidental in Chihuahua, Mexico, two unusual specimens

were found and initially given fieldnames of Aegopogon (Peterson & P. Catalán 17542) and Bouteloua (Peterson

& Catalán 17621). Later, when PMP was preparing treatments of Bouteloua and the tribe Muhlenbergiinae

for northeastern Mexico (Peterson et al. 2007b: Herrera Arrieta et al. 2008), these specimens were reevalu-

ated after seeing them in the United States National Herbarium (US) in separate unidentified folders of

Aegopogon and Bouteloua, respectively. Upon closer inspection, these collections represented the same new

species that could not be easily placed in Aegopogon or Bouteloua, nor any of the remaining genera currently

placed in the Muhlenbergiinae (Peterson et al. 2001b, 2007a). Individuals of this new species are unique in

having terminal, raceme-like panicles with only two spikelets at the end of each primary branch. A second

opinion seemed warranted; PMP then sent a duplicate off to JTC for examination and incorporation into

DNA sequence data sets that could possibly elucidate affinities of this enigmatic species. Subsequently, a

third specimen was found (Peterson et al. 8032). As discussed below, analyses of DNA sequences from the

new species placed it in subtribe Muhlenbergiinae.

Members of subtribe Muhlenbergiinae are grasses (Poaceae) in the subfamily Chloridoideae, tribe Cy-

nodonteae (Peterson et al. 20015, 2007a, in review). Muhlenbergiinae are highly variable morphologically,

although the group can be generally characterized as follows: ligule a ae ey a line en ARA

inflorescence a panicle, Md. Or Mu oe only of primary branches; s

¿l

pairs or triads, cleist casionall y present in the leaf sh doel a mere perfect,

staminate, or sterile; slimes dened or unawned; lemmas 3-nerved, awned or unawned; base chromosome

number x = 8-10 (Peterson 2000; Peterson & Herrera Arrieta 1995; Peterson et al. 1997, 2007a, b; Colum-

J, Bot. Res. Inst. Texas 3(2): 527 — 534. 2009

528 Jou

E ( . PEL | b. E Ee RC E PA xac

bus et al. 2007). Two subtypes of C, photosynthesis, NAD-M

malic enzyme) and PCK (phosp! lp; te carboxyki

cal assays to occur in Muhlenbergiinae (Gutiérrez et al. 1974, Brown 1977; Hattersley & Watson 1992).

By far, the largest genus in the subtribe is Muhlenbergia, which has 154 species including the important

North American range grass M. montana (Nutt.) Hitchc., hitropical disjuncts M arenicola Buckley and M.

torreyi (Kunth) Hitchc. ex Bush, and seven species located in southeast Ásia (Herrera Arrieta 1998; Peterson

& Ortíz Diaz 1998; Peterson 2003; Wu & Peterson 2006; Herrera Arrieta & Peterson 2007; Peterson et al.

2007b). Species indigenous to North America number 127 (8696), and 125 of these occur in Mexico, where

56 species are endemic (Espejo Serna et al. 2000; Dávila-Aranda et al. 2004, 2006; Peterson & Herrera Ar-

rieta 2005; Peterson et al. 20072).

The remaining nine genera have four or fewer species, and four are monotypic. All are limited to the

New World except for an occurrence of Aegopogon cenchroides Humb. & Bonpl. ex Willd. in Papua New

Guinea (Veldkamp 1985). Five are endemic to North America. Apart from its ce in Papua New Guinea,

se), have been found and verified by biochemi-

Ts i

Aegopogon (four species) is distributed in North and South America. B ( I

ern Mexico (Peterson 1989; Peterson et al. 1993). Blepharoneuron (two species) is found in North America

and includes B. tricholepis (Torr.) Nash, an important range grass in the southwestern U.S.A. and northern

Mexico (Peterson & Annable 1990, 2003). Chaboissaea (four species) has three species in central Mexico

and C. atacamensis (Parodi) P.M. Peterson & Annable in Argentina and Bolivia (Peterson & Annable 1992;

Peterson & Herrera Arrieta 1995; Sykes et al. 1997). Lycurus (three species) has one species limited to North

America and two amphitropical disjuncts, including L. setosus (Nutt.) C. Reeder (Reeder 1985; Sánchez &

Rügolo de Agrasar 1986; Peterson & Morrone 1998). Pereilema (four species) is distributed in North, Cen-

tral, and South America. Redfieldia (one species) is endemic to the U.5.A. (Reeder 1976). Schaffnerella (one

species) is known only from San Luis Potosi, Mexico (Columbus et al. 2002). Schedonnardus (one species) is

yet another genus with an amphitropical distribution.

Analyses of cpDNA ndhA-intron, ndhF, rpl32-trnLV^9, rps3, rps16-intron, rpsl6-trnK, trnL-F, and nrDNA

ITS sequences of species in the Muhlenbergiinae (Columbus et al. in press; Peterson et al. in prep.) indicate

FE A ye ee -— EE EI hvletic | tl ine, smaller genera

l.i +l

to norin-

are nested within it; therefore, expanding the circumscription of Muhlenbergia to include these nine genera

seems warranted (Duvall et al. 1994; Peterson et al. 2001a, 2004; Peterson & Herrera Arrieta 2005; Peterson

et al. 2007a). In this paper, in addition to describing a species new to science, we present a phylogenetic

hypothesis and support for the generic placement of this unusual taxon.

Muhlenbergia tarahumara PM. Peterson & Columbus, sp. nov. (Figs. 1A-J; 2). Tee: MEXICO. CumuaHua. Muni-

cipio Guachochi, Sierra Madre Occidental, 2 km W of Rio Coraréachi and E of Osichi (27°28'15.0"N, 107°31'5.0"W), 1960-2040

m, 30 Aug 2003, PM. Peterson & P Catalán 17621 (HOLOTYPE: US-3470469!; isorvpes: CIIDIRI!, KI, MO!, RSA!, USD.

(1 J 1 4 2 1:1 : £1 -1

Ab Aegopogon cenchroides Humb. & Bonpl. ex Willd. paniculis ramis 5-13 per culmo, ]

spiculis; glumis 3-5.8 mm longis, brevioribus quam lemmatibus, ang lanceolatis; lemmata 3.6-7.3 mm longis, lanceolatis; paleae

3-4.6 mm longis; antherae 2.2-2.8 mm longis, tangerinis, recedit.

1.41

Caespitose perennials. Culms 18-55 cm tall, erect, terete near base, g below the nodes, usually 3 nodes

per culm; internodes glabrous and shiny. Leaf sheaths 0.8-8 cm long, shorter than the internodes above,

pubescent above and mostly glabrous below; ligules 1-1.8 mm long, membranous, abaxially pubescent or

glabrous, apex acute, often erose, minutely ciliolate; blades 2-)3.5-15 cm long, 0.2-1.3 mm wide, flat to

tightly involute, apically acuminate, som ewhat sinuous, antrorsely hirsute on both surfaces, the hairs 0.1-0.4

mm long. Panicles 3-6 cm long, 0.7-1.6 cm wide, narrow, terminal with 5-13 racemosely arranged primary

branches, 1 per node; branches 0.5-1.4 cm long, with two terminal spikelets, deciduous, disarticulation

near base, branches first ascending then spreading (bending sharply or curling near base) from the culm

axis, usually secund, antrorsely hirsute, the hairs 0.2-0.4 mm long; inflorescence axis flattened, ending in

terminal branch, margins hirsute; pedicels fused or 0.2-0.6 mm long, one slightly longer than the other,

tightly appressed. Spikelets 4.5-7.8 mm long, appressed to one another, 1-flowered; glumes 3-5.8 mm long,

RI pL. =

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Fic. 1. Muhlenbergia tarahumara [PM. Peterson &

nm". rz

D. Branch with

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17621 (US)]. A. Habit. B. Sheath, ligule, and blade. C. T ired spil

1) G. Lemma. H. Paleas. |

Stamens and pistil. J. Lodicules.

Aa r L]

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530 f Texas a2)

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sheath of vascular bundle.

1 t chartaceous

narrowly lanceolate, usually equal in length, shorter than the lemma, chartaceous, 1-veined,

pubescent, apex acuminate, awned, the awns 1.4-3.2 mm long; lemmas 3.6-7.3 mm long, lanceolate, awned,

distinctly 3-veined, membranous to chartaceous, appressed pubescent, apex acute to acuminate, bifid, the

central awn 2-3.7 mm long, straight or slightly recurved, the lateral veins extending into awns 0.6-1.5 mm

below and appresse d pubescent

long; paleas 3-4.6 mm long, shorter than the lemma,

near apex, 2-veined, apex acuminate, each vein extending as a mucro or awn 0.2-0.9 mm long stamens 3,

anthers 2.2-2.8 mm long, yellowish orange; lodicules 2, membranous; ovary glabrous with two styles and

two stigmas. Caryopsis not seen.

Comments.—The new species can be distinguished from all other species of Muhlenbergia by having a

terminal, raceme-like panicle with only two spikelets at the end of the branch. It closely resembles species

of Aegopogon, which have the same kind of inflorescence, including the deciduous branches, but with three

spikelets per branch, and some species of Bouteloua. The species is named after the Tarahumara people who

are indigenous to the northern Sierra Madre Occidental.

Distribution and Habitat —HMuhlenbergia tarahumara is known only from three locations in the Sierra

Madre Occidental in Chihuahua, Mexico, where it occurs on rocky slopes, ridgetops, and white-tuff rock

outcrops with species of Pinus, Quercus, Cupressus, Arctostaphylos, Vaccinium, Comarostaphylis, Aristida, Ae-

gopogon tenellus (DC.) Trin., A. cenchroides, Muhlenbergia lucida Swallen, M. montana, M. pauciflora Buckley,

M. polycaulis Scribn., and M. rigida (Kunth) Kunth; 1880-2075 m.

> fal [| ,M LI L H 4 L ] H £ MA 5 531

Additional specimens examined. MEXICO. Chihuahua: Municipio Guachochi, Sierra Madre Occidental, 41.3 km S of Creel on road

to aciei 2075 m, 10 Sep 1989, P.M. Peterson, C.R. Annable & Y. Herrera Arrieta 8032 (RSA, US-3513942); Sierra Madre Occidental,

Ya mi E of hwy N of Río Urique crossing towards Basihuare and Creel, 27°23'59.4"N, 107°29'20.4"W, 1880-1900 m, 26 Aug

2003, P.M. Peterson & P. Catalán 17542 EUIS RSA, US-3468887); 5 Sep 2008, P.M. Peterson & J.M. Saarela 22053 (RSA, US); 6 Sep

2008, P.M. Peterson & J 9 (US).

Leaf anatomy.—Cross-sections of the leaf blade were done with a rotary microtome on fresh-field fixed mate-

rial (Peterson & Saarela 22053), stained, and mounted pense to DEREN (Sharman 1943; Columbus 1999;

Peterson & Herrera Arrieta 2001). P t slides are dep d at RSA and US. The following description

employs much of the standardized terminology proposed by El Ellis (1976).

The leaf blade in transverse section displays K y, which is associated with C, photosynthe-

sis (Fig. 2). It is nearly flat apad. melts on drying) and both surfaces have rounded or flat-topped,

longitudinal costal (al /below the vascular bundles) ribs alt ting with V-shaped intercostal furrows; a

conspicuous rounded midrib projects abaxially. The furrows are up to 1/2 the thickness of the blade; those

on the abaxial surface are more aaa The innen punas (VBa have two sheaths, an inner (mestome,

XyMS+; Hattersley & Watson 1976) and an outer (parenchymatous, PCR

tissue; Hattersley et al. 1977) sheath of eases amt an outer dan is uneven in outline and its cells

contain round chloroplasts that are distributed ‘peripherally within the cell (Prendergast et al.

1987). The VBs differ in size, develog t of metaxylem, whether or not the outer sheath is continuous,

and the amount of scl present on the ada d abaxial sides. There are four primary VBs (with

large metaxylem vessels; includes the midvein), four secondary VBs, and four tertiary VBs arranged as fol-

lows: iii-i-iii-i-ii-ii-i-ii-ii-i-iii-iii. The phloem is not sclerosed. In the innermost five VBs, the outer sheath is

extended adaxially into the rib. The outer sheath is interrupted by sclerenchyma on the abaxial side of all

VBs except three of the four tertiary VBs, and on the adaxial side of bd two "un pus VBs nearest E

J: oi. 1 el

midvein. A narrow layer of smaller, isodiametric, indistinctly

outer sheath (PCA tissue; Hattersley et al. 1977). Sclerenchyma girders (in — with the outer us are

present on the adaxial and abaxial sides of all VBs. The largest girder (many cells in five series/rows) forms

the midrib, whereas those of the tertiary VBs are smallest (« 10 cells in one or two series). Abaxial girders

associated with primary and secondary VBs, as well as those on the adaxial side of the two lateral primary

VBs nearest the pd are broadest near the epidermis. Tue PIRE ee has two or cee subepidermal

fibers. Bulliform t adaxially adjacent to tl an y smaller towards

the margin; those on the basal side are smaller than their adaxial SOUTE DATIS. eee cells one or

two columns wide extend between bulliform cells on both sides of the blade, dividing the chlorenchyma

associated with adjacent VBs.

Molecular analyses.—DNA was obtained from field-collected leaf material (Peterson & Saarela 22053)

and was sequenced for ndhA-intron, ndhF, rpl32-trnL™©, rps3, rps16-intron, rps16-trnK, trnL—F, and ITS. To

determine the phylogenetic position of the new To a Chloridoideae, the sequences were aligned

with the Columbus et al. (2007) data set, Te] ridoid genera an d the Peterson et a m Evien)

data set representing 96 chloridoid genera. Upon au the new species resolved in the }

clade. To determine its position within Muhlenbergiinae, the sequences were aligned with the two pecu aie

data sets that included all ten genera in the subtribe and 80-90% of the species in Muhlenbergia (Columbus

et al. in prep.; Peterson et al. in prep). Analyses of the Muhlenbergiinae data sets revealed that the new

species is a member of a subclade that includes Aegopogon, M. subgen. Muhlenbergia, and Pereilema. Its posi-

tion within the subclade was not resolved in the trnL—F phylogeny (Columbus et al. in prep.), although in

the combined ndhA-intron, ndhF, rpl32-trnL™®, rps3, rps16-intron, rps16-trnK phylogeny (Peterson et al. in

prep.) the new species was supported as sister to Aegopogon cenchroides and A. tenellus. However, in the ITS

phylogeny the new species is supported as the sister of Aegopogon (Columbus et al. in prep; Peterson et al.

in prep.).

532 J | of the Botanical R h Institute of Texas 3(2)

DISCUSSION

The new species g members of the Muhlenbergiinae in having only two spikelets per branch,

these racemosely amagi along the panicle axis. Reduction of the number of spikelets per branch is also

seen in Muhlenbergia diversiglumis Trin. (a member of M. subgen. Muhlenbergia) where there are 2-5 spike-

lets per branch EOSDEM & Annable [on PONE er, M. focum poemas most closely Aegopogon in

morphology, which sup the phylogenetic hypothesis that these taxa are sister (Columbus et al. in prep.;

Peterson et al. in prep.). Aeon differs hom M. tarahumara in having three spikelets per branch. The

three spikelets are dimorphic—one is larger and hermaphrodite and two are smaller (sometimes rudimen-

tary; one often not developed in A. bryophilus Doll) and staminate or neuter. In M. tarahumara, no spikelet

dimorphism is evident.

Leaf anatomy suggests M. tarahumara undergoes the PCK subtype of C, photosynthesis. The presence

of two bundle ae uneven pou of the outer shea, oe ak of the chloroplasts in the outer

sheath, and the heral position of these cl ts within each cell are together predictive

of PCK (Fig. 2; prendersasta et al 1987). Aegopogon likewise has PCK anatomy (Columbus 1996). Peterson

and Herrera Arrieta (2001) reported that the species in M. subgen. Muhlenbergia possess PCK-like anatomy,

whereas the remaining species have NAD-ME anatomy. However, M. tarahumara differs from species of Ae-

gopogon and M. subgen. Muhlenbergia by having mesophyll chlorenchyma that forms a narrow layer around

the outer sheath and is not continuous between adjacent vascular bundles, being separated by columns of

colorless cells. This po is P seen in M. pauciflora and M. polycaulis (Peterson & Herrera Arrieta

2001), where the vascular | the middle of the amd db columns of colorless cells that separate

adjacent vascular bundles. However, near the leaf blad tiary vascular bundles of M. pauciflora

and M. polycaulis have chlorenchyma cells that are continous between each adjacent vascular bundle. In

addition, Muhlenbergia pauciflora, like M. tarahumara, has well developed abaxial scl hyma girders that

are broadest near the epidermis (Peterson & Herrera Arrieta 2001).

ACKNOWLEDGMENTS

We thank the Nati 1G hic Society C ittee for R h and Exploration (g t number 8087-06)

for field and laboratory po the taa Institutions, Restricted Endowment Fund, the Scholarly

Studies Program, Research Opportunities, Atherton Seidell Foundation, and Biodiversity Surveys and Inven-

tories Program, all for financial support; Alice R. Tangerini for illustrating the new species; Pedro Acevedo

and Ida C. López for correcting the Spanish resumen; Adolfo Espejo Serna and Yolanda Herrera Arrieta for

providing helpful comments on the manuscript; and a number of colleagues who assisted with fieldwork.

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2315-525

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(Poaceae: A Oricola) aopa on internal transcribed spacer region sequences (nrDNA). Botany 2001

abstract: http:// y2001.org/section12/abstracts/33.shtml

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Science Press and Missouri Botanical Garden Press, Beijing and St. Louis. 22:486-487

UNA NUEVA ESPECIE Y NOVEDADES NOMENCLATURALES

EN EL GENERO MELIOSMA (SABIACEAE)

J. AED Morales

instituto Nacional de Bi lad (INBio), Apto 22--3100

Santo Domingo de Heredia, COSTA RICA

imorales@inbio.ac.cr

RESUMEN

Se describe Meliosma AA una nueva caver de Costa Rica ieee Panamá y se discuten sus afinidades taxonómicas. Meliosma

hartshornii A.H. Gentry

ABSTRACT

Meli ist] i ies f C Rica, and Panama, is d ibed and i ic relationshi di d. Meliosma

J J +1 SA » C. 11

hartshornii A.H. Gentry y ym)

PALABRAS CLAVE: Sabiaceae, Meliosma, Costa Rica, Panama

Meliosma (Sabiaceae) es un género de distribución amfipacifica con una interesante disjunción entre el SE

de Asia hasta Malasia y la America tropical, presentando una mayor diversidad en Centro América y los

Andes tropicales (Kubitzki 2004, 2006). En el tratamiento de la Flora de Panama, Gentry (1980) registró

un aproximado de 40 especies en América (vs. 15 en el Asia). Desde entonces, se han realizado numerosas

publicaciones de novedades taxonomicas (e.g., Arbeláez 2004; Aymard & Cuello 1994; Cornejo & Bonifaz

2006; Cuatrecasas 1988; Cuatrecasas & Idrobo,1988; Gentry 1986, 1992; Menjivar et al. 2008; Morales

2003; Steyermark & Gentry 1992) y compilaciones en floras y catálogos (e.g., Flora de Venezuela (Steyer-

mark & Gentry 1992); Flora de Nicaragua (Gentry 2001); Flora de la Guyana Venezolana (Aymard & Cuello

2005), por lo que actualmente el número total de especies del género se ubica entre 80 y 90, con cerca de 70

restringidas al Nuevo Mundo. Sin embargo, ante la carencia de trabajos monográficos recientes, que traten

en forma total o parcial las especies de Meliosma en el neotrópico, es probable que el número real pueda ser

menor, sobretodo ante la presencia de posibles sinónimos nomenclaturales en taxones de Sur América.

La separación infragénerica más utilizada ha sido la propuesta por Van Beusekon (1971), en la cual

subdivide el género Meliosma en dos subgéneros: Meliosma y Kingsboroughia (Liebermann) Beusekom. El

primero incluye la secciones Meliosma y Lorenzanea (Liebermann) Beusekom, mientras que Kingsboroughia

esta oe en nas secciones Sk wana dd y Hendersonia Beusekom. El subgénero Meliosma se diferencia

de Ki jas y I simples ( puma) cinco sépalos (vs. cuatro), pétalos

ee de nal MN E bred los hasta anchamente elípticos (vs. más

anchos que largos, anchamente imbricados, reniformes) y presencia de un haz vascular (el cual conecta el

pedicelo de la semilla) situado en un canal ind ii denitro Be endocarpo, (vs. situado fuera del endocarpo

o libre en la pulpa del mismo). Todas las tran incluidas en la sección Loren-

zanea, excepto M. alba (Schltdl.) Walp., (ume del SE ide México y SE de Asia) la cual pertene a la sección

Kingsboroughia.

Durante el proceso de elaboración de una revisión del género en México y Centroamérica, se han

encontrado una serie de novedades taxonómicas, entre las que se TUR varias especies sin describir, así

como la necesidad de reducir a la sinonimia A ion se describe una especie nueva

para Costa Rica y Panamá, discutiéndose las ones con las especies más relacionadas y se propone un

nuevo sinónimo nomenclatural.

J. Bot. Res. Inst, Texas 3(2): 535 — 540. 2009

536

SEE s

rre :

E TOT. .

II rem ft

1 & Keefe 4177, INB; H de Chacon 135, CR). A. R

wag. Jl

E da dail

iyo H

1 Li

. fA

V y L

flaral € Sánala victa ahavial D. Sénalo vista adayial. E Pátaln externo. vista Ahavial E Ectaminndia G Ectambrac

f Li f , 7 r

por una flecha).

Morales, N i lad laturales in el Meli 537

Meliosma isthmensis J.F Morales, sp. nov. (Fig. 1). Tro. COSTA RICA Limón: cantón de Pococí, llanura de Santa Clara,

finca La Suerte, 50 m, 8 jul 1995 (fD), R. Aguilar, C. Woodward & K. Keefe 4177 (HoLoriro: : CR, MO)

1: + [Anl : POEM E. S

A Meliosma schilimii (Turcz.) Triana, cui affinis,

Árboles de 7-14 m de altura, abes PRA a simian con lenticelas pares y más o menos

tulo, los tallos

circulares, las yemas y tallos j pubescentes, el i

viejos con el indumento más esparcido y algo ao Hojas alteras; a veces más o menos aglomeradas al

final de las ramitas, cortamente pecioladas, el pecíolo 1,1-1,9 cm de largo, esparcidamente hirsutulo hasta

glabrescente, pulvinulo presente, pero apenas evidente; lámina 35-55 x 8-17,5 cm, obovada a angostamente

obovada, el ápice agudo, la base angostamente cuneada, a veces prolongándose tenuamente en el peciolo,

no revoluta, el margen entero y no revoluto, no bulladas, glabrescentes en el haz, la pubescencia restringida

sobre la costa de las nervaduras, densa a moderamente puberulentas o tomentulosas en Rue envés, la pubes-

cencia màs o menos distribuida por toda la lámina, con 37 a 46 (o más) pares de ias, las venas

terciarias conspicuamente reticuladas en la superificie abaxial, las venas apenas visibles en la ere

adaxial. Inflorescencias panículas, laterales o axilares (ay t les en las ramas jóvenes),

tripinadas, 13-40 cm de largo, esparcidamente puberulentas, el indumento incoloro, brácteas 1-3 mm

de largo, filiformes, bracteólas 0,4—0,6 mm de largo, angostamente ovadas, escariosas; flores densamente

aglomeradas, sésiles a aia con pedicelos (cuando presentes hasta o 7 mm BS longitud), eae 5, de

1,3-1,9 mm de largo, anch e ovados el ápice agud u obtuso,

el margen ciliado; pétalos crema a verde—crema, 5, desiguales en an los 3 externos 1,9—2,2 mm de ae,

anchamente ovados a suborbiculares, glabros, exceptos por unos cuantos cilios esparcidos en el margen,

los 2 internos 0,9-1,2 mm de largo, ubicados dorsalmente sobre los filamentos y sobrepasándolos, lineares,

glabros; estambres 5, pero solo 2 fértiles, las tecas 06-0,9 mm de largo, los estaminodios 1,1-1,4 mm de

largo, angostamente subovados, glabros; ovario 0,3—0,6 mm de largo, glabro. Frutos 2,2-2,6 x 1,9-2,2 cm,

obovoide-piriformes, blanco verdosos al madurar.

Distribución, habitat y ecología. —Restringida a la vertiente caribe de Costa Rica y Panamá, donde crece

en bosques muy húmedos en elevaciones de 100-1000 m. Especímenes con flores han sido recolectados en

febrero, junio y julio. Frutos se han observado en marzo, mayo, junio y de agosto a octubre.

Meliosma isthmensis habia sido identificado como M. schlimii (Turcz.) Triana en la sinopsis ah pue

Meliosma para Costa Rica y P á (Morales 2003). Sin embargo, a través de

(adicional a la colección tipo) debssitadiss en diferentes herbarios colombi y t

que M. schilimii es un taxón restringido a las zonas altas de los Andes en Colombia y Feader y la especie

presente en Costa Rica y Panamá representa un taxón diferente, restringido a bosques muy húmedos, de

elevaciones bajas y medias Bajo los ee m de abi Meliosma P difiere de M. schlimii por sus

tallos jóvenes y yemas con el ind tul fi 1 zs. puberulento e incoloro), láminas

foliares con la base no revoluta (vs. revoluta), no bulladas, así como fees densamente aglomeradas (vs. no

aglomeradas), y habitats diferentes, ya que M. isthmensis se encuentra confinada a bosques muy húmedos

en zonas bajas (100-1000 m) de la vertiente atlántica en Costa Rica y Panamá, mientras que M. schlimii está

diia a ees montanos, sobre los 2200 m en Colombia y Ecuador.

, Se conis ve:

los. COSTA RICA. Cart Chitaria, Turrialba, 22 jun 1972 (fl, fr), Poveda 183 (MO). Limón: Reserva

indiana Talamanca, quebrada Heliotropo, entre Sukut y nt 9 jul 1989 (fr), ee 135 (CR, m al SO de Bribri, 4 may 1983

(fr), Gomez et al. 20469 (MO); Almirante, entre ríos Xichiary y Boyei, 11 ago 1995 (fr), Herrera 8390 (CR ); El Cedral, al N de Cariari, 16

sep 1994 (fr), Thomsen 1213 (CR); Talamanca, fila Carbon, Buena Vista, 28 mar 1997 (fr), Valverde 27 (CR). PANAMA. Coclé: río San

Juan, 12 jun 1978 (fr), Hammel 3450 (MO). Colón: fil Rita, al SE de Colon, 21 may 1986 (fr), McPherson 9207 (MO); Santa Rita,

29 may 1975 (fr), Mori & Crosby 6411 (MO). PANAMÁ: cerro Jefe, cerca del río Indio, 17 feb 1968 (fl), Duke 15213 (MO). San Blas:

entre Yannuadi y la costa, Nargana, 25 oct 1992 (fr), Herrera 1260 (MO)

Meliosma irazuensis Standl., Arch. Bot. Sist. 9(3—4):194. 1933. (Fig. 2). Tiro: COSTA RICA. Canraco: Volcán Irazü,

cerca de Guayabillos, 2500 m, 22 may 1930 (fl), G. Cufodontis 451 (moLoriro: F [fotocopia, MO]; isoripos: EAP [fragmento], W).

Meliosma hartshornii A.H. Gentry, Arn. Missouri Bot. Gard. 73:820. 1986 [1987], syn. nov. Tipo: COSTA RICA. Herpa: camino al Volcán

Barba, 2 km al N de Sacramento, 2750 m, 29 dic 1974 (fD, G. Hartshorn 1608 (uoi CR)

538

l cm

L PA f 5|

LC Sépalo, victa

Fic. 2. Meliosma irazuensis (A—F de Herrera 3631, INB; G de Morales 8664, INB). A. R

adaxial D. Pétalo externo, vista adaxial. E. Estaminodio. F. Estambres, At

flect ). G Fruto

e W

Morales, N i lad laturales in el Meli 539

Árboles de 8-18 m de altura, tallos EH à SUCHE Tios, con lenticelas esparcidas, y más o menos

circulares, elevadas, las yemas y tallos j es adpreso-pubescentes, el indumento

amarillo, los tallos viejos gldbrascentes Hoji alternas, a veces más o menos aglomeradas al final de las

ramitas ey pare cenas electa o subverticiladas, cortamente pecioladas, el pecíolo 0,5-1,2 cm de largo,

glabro o glabrescente, el pulvinulo presente, evidente; lámina 3,5-11,5

16 ,5) x (1142, 4- 4,6(-5,5) cm, eliptics bounds: «iis a obovada, el ápice agudo, la base angostamente

cuneada, prolongándose en el peciolo, usualmente revoluta, el margen entero a espaciadamente dentado,

no bulladas, glabras en el haz, esparcida e inconspicuamente pubescentes abaxialmente, con 8 a 14 pares

de venas secundarias, las venas terciarias finamente reticuladas en la superificie abaxial, las venas apenas

visibles en la superficie adaxial. Inflorescencias panículas, terminales o subterminales en ramas jóvenes,

volviéndose axilares con la edad, tripinadas, (4—7—15 cm de largo, densa a moderadamente puberulentas,

el indumento algo ferrugíneo, brácteas 1,2-2,1 mm de largo, angostamente ovadas, bracteólas 0,8-1,4

mm de largo, ovadas, escariosas, dispuestas inmediatamente debajo del cáliz y semejando sépalos; flores

densamente aglomeradas, sésiles a subsésiles, con pedicelos (cuando presentes) hasta 1,3 mm de longitud,

sépalos 5, de 1,5-1,9 mm de largo, anchamente ovados, el ápice obtuso, glabros externamente, el margen

conspicuamente ciliolado; pétalos verde-crema a crema, 5, desiguales en tamano, los 3 externos 1,8-2,2

mm mm de largo, anchamente ovados, glabros, los 2 internos 0,5-0,8 mm de largo, ubicados dorsalmente

sobre los filamentos, subcuadrados, glabros; estambres 5, pero solo 2 fértiles, las tecas 07-0,9 mm de largo,

los estaminodios 1,1-1,3 mm de largo, subcuadrados, glabros, ovario 0,3—0,4 mm de largo, glabro. Frutos

1,2-1,5 x 1-1,2 cm, subglobosos, rojizos o blanco-rojizos al madurar.

Distribución, habitat y ecología.—Honduras, Costa Rica y Panamá, en bosques muy húmedos, bosques

nubosos y formaciones de roble (Quercus sp., Fagaceae), en elevaciones de 2250-3300 m. Especímenes con

flores han sido recolectados entre octubre y marzo. Frutos se han observado intermitentemente durante

todo el año.

En la descripción de Meliosma hartshornii, Gentry (1986) comentó que ese taxón estaba cercanamente

relacionado con M. irazuensis Standl., de la cual difería por las flores pediceladas (vs. sésiles), así como in-

florescencias más anchas, con las ramas lateral g las y ame diferente. El estudio de

dichos caracteres en las colección tipo, colecciones adicionales y ol | en el campo, han puesto de

manifiesto que las flores en una misma rama, pueden ser eese subsésiles o cortamente pediceladas, con

pedicelos que no exceden (cuando presentes) 1,3 mm de longitud. Las inflorescencias inmaduras tienen

un indumento denso, el cual llega a ser insignificante con el desarrollo de la inflorescencia. Finalmente,

el especimen tipo de M. irazuensis tiene un indumento similar al encontrado en algunos especimenes de

M. hartshornii. Ante la inconsistencia de los caracteres utilizados para separar ambos taxa, M. hartshornii

se propone como sinónimo, en virtud que sus hojas, inflorescencias y flores coinciden dentro del rango

a Md penado para M. irazuensis.

p conocida solo de Costa Rica y Panamá (Morales 2003), pero es reportada

por primera vez para Honduras, basado en colecciones realizadas en el departmento de Lempira, lo que

incrementa a cinco el número de especies de Meliosma presentes en Honduras (Morales & Nelson, datos sin

pons

los. HONDURAS. Lempi I N jo, al SE de Gracias, parq ional de Celaque, 31

ene e 1992 (fr), Mejía 21 (EAP, INB, MO, TEFH).

AGRADECIMIENTOS

Quiero agradecer a los siguientes herbarios por permitir el estudio de sus colecciones: COL, CR, DUKE,

EAP, F, HUA, JAUM, LAGU, MEDEL, MHES, MO, NY, TEFH, W. De igual forma, agradezco las facilidades

brindadas por Cirilo Nelson (TEFH) para el estudio de especimenes en Honduras

540 J t tanical Insti Texas 3(2)

RETERCNCIAS

ArseLágz, A.L. 2004. Two new species of Meliosma (Sabiaceae) from Bolivia. Novon 14:12-16.

AYMARD C, G. & N. CueLto.1994. Meliosma gentryi Aymard & Cuello (Sabiaceae) una nueva especie para la flora de

la Guayana venezolana. BioLiania 10:1-3.

AYMARD C, G. 8: N. CueLLo. 2005. Sabiaceae. En: J.A. Steyermark, P. Berry & B. Holst, eds. Flora of the Venezuelan

Guayana. Timber Press; Missouri Botanical Garden, Portland; St. Louis, Estados Unidos. 9:39-43,

Cornejo, X. & C. Bonifaz. 2006. Meliosma stellata, una nueva especie de Sabiaceae de Ecuador. Novon 16:

328-330.

CUATRECASAS, J. 1988. Miscellaneous notes on neotropical flora XVII. New species of Meliosma. Phytologia

Cuatrecasas, J. & J. M. loroBo.1988. Tres nuevas especies de Meliosma Blume (Sabiaceae) de Colombia. Ernstia

Gentry, A.H. 1980, Sabiaceae. En: R. Woodson, R. Schery, et al, eds. Flora of Panama. Ann. Missouri Bot. Gard.

67:949-963

Gentry, A.H. 1986. New neotropical species of Meliosma (Sabiaceae). Ann. Missouri Bot. Gard. 73:820-824.

Gentry, A.H. 1992. Four new species of Meliosma (Sabiaceae) from Peru. Novon 2:155-158.

Gentry, A.H. 2001. Sabiaceae. En: W. Stevens, C. Ulloa, A. Pool & O. Montiel, eds. Flora de Nicaragua. Monogr. Syst.

Bot. Missouri Bot. Gard. 85:2303-2306.

Kuerrzx, K. 2004. Sabiaceae. En: N.P. Smith et al., eds. Flowering plants of the neotropics. Princenton University

Press, Princeton, New Jersey, Estados Unidos. Pp. 335-336.

Kuarrzki, K. 2006. Sabiaceae dd K. Kubitzki, ed. The families and genera of vascular plants. Volume IX. Flowering

plants. Eudicots. Berl idales, Buxales, Crossosomatales. Springer, Berlín, Alemania. Pp. 413-417.

MEnuivar, J., G. CERÉN & J.F. Mones 2008. Sinopsis del género Meliosma (Sabiaceae) en El Salvador. Anales Jard.

Bot. Madrid 65:389-392.

Morales, J.F. 2003. Sinopsis del género Meliosma (Sabiaceae) en Costa Rica y Panamá, con 3 nuevas especies.

Sida 20:931—943.

STEYERMARK, J. & A.H. Gentry. 1992. Sabiaceae. En: G. Morillo, ed. Flora de Venezuela 5(1):221-267.

Van Beusekom, C.F. 1971. Revision of Meliosma (Sabiaceae), section Lorenzanea excepted, living and fossil, geog-

raphy and phylogeny. Blumea 19:355-529.

ESTUDIOS EN LAS APOCYNACEAE NEOTROPICALES XXXVII: MONOGRAFÍA

DEL GENERO RHABDADENIA (APOCYNOIDEAE: ECHITEAE)

J. ANG Bolsos

Instituto Naci Bj | (INBio), Apto 22--3100

Santo Domingo de Heredia, COSTA RICA

fmorales@inbio.ac.cr

RESUMEN

del género Rhabdadenia (A chiteae). En total, tres especies son aceptadas (R. biflora Jacq.)

Mall. ATE. R. madida (Vel) in de R. ragonesei Woodson), yee en una clave, RESP iir d Meme T Estado de

rostoma (Benth.) Múll. Arg yR pohlii Mull.Arg. bajo | de R. madida, asi le | E. biflorus Jacq.,

E. madidus . lindeniana Müll.Arg., R. polli Müll.Arg. y R. pohlii Müll.Arg. var. iberia Müll.Arg., y la selección de un neotipo

Vell.,

para Echites ee Schltd

PALABRAS CLAVE: Apocynaceae, Apocynoideae, Echiteae, Rhabdadenia, Echites

ABSTRACT

A 1 Fl n1 La J y FA ees Ts

ted TI f pt ](R biflora (Jacq.) Müll Arg.,

Té E

R. mo. (Vell) Mieres and R. ragonesei IE NEM and a key, üesen some. illustrations,

Rhabdadenia macrostoma e Müli.Arg. and R. pohlii Müll.Arg. are placed in synonymy of R. madida Lectotypes are selected for E.

biflorus Jacq., E. madidus Vell., R. E Mull.Arg., R. pohlii Müll.Arg., and R. pohlii Müll.Arg. var. suberecta Müll.Arg. and a neotype

is chosen for Echites uc Schlt

Key WoRps: Apocynaceae, Apocynoideae, Echiteae, Rhabdadenia, Echites

Rhabdadenia Mull.Arg. es un pequeño género de tres especies distribuido prácticamente en todas las áreas

tropicales y subtropicales del continente americano, desde el S de Florida (Estados Unidos) y las Antillas

hasta el N de Argentina (excepto Chile), con una de las distribuciones geográficas más amplias de las Apo-

cynoideae Neotropicales, compartida únicamente por los géneros Forsteronia, Mandevilla, Mesechites, Odon-

tadenia, Prestonia y Tabernaemontana (Hansen 1985; Leeuwenberg 1994 a; Morales 1998, 1999; Woodson

1936). Este género fue descrito por Muller (1860 a), quien describió una especie, R. pohlii, incluyendo tres

nuevas variedades, y transfirió otra previamente tratada en el englobador género Echites (E. biflorus Jacq.).

En un trabajo subsecuente y publicado casi en forma simultánea el mismo Muller (1860 b), transfirió E.

macrostomus Benth. a Rhabdadenia y publicó tres taxones adicionales (R. cubensis Müll.Arg., R. lindeniana

Müll.Arg. y R. wrigthiana Mull. Arg.), aumentando el número de especies en el género a seis. Posteriormente

Miers (1878) describió o transfirió diferentes taxones descritos en otros end a Ree y e un

total de 11 especies. Finalmente Woodson (1936), clarificó y definió

un total de tres especies, número que se incrementó a cuatro con la deserción de R. — ligeramente

más tarde (Woodson 1940). Continuando con la actualización de monografías de los géneros de las tribus

Echiteae y Mesechiteae (Apocynaceae, Apocynoideae), se presenta a continuación una revisión del género

Rhabdadenia.

MORFOLOGÍA

Habito.—Las especies de Rhabdadeni hi tas, escandentes o lianas, usualmente restringidas

a manglares o áreas anegadas, tales como sabanas o Bagtinos Aunque tanto R. biflora como R. ragonesei

son consistentemente lianas, una alta variación en relación al hábito está presente en R. madida, de manera

que tanto for tas o escandentes son encontradas en la misma población. Aunque estos extremos

J. Bot. Res. Inst. Texas 3(2): 541 — 564. 2009

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542

de variación pueden ser totalmente diferentes, la morfología floral es prácticamente la misma. La misma

variación está presente en algunas especies de otros géneros de las Apocynoideae, como Mandevilla, donde

algunas especies como M. pentlandiana (A. DC.) Woodson (Ezcurra 1981) y M. sancta (Stadelm.) Woodson

pueden comportarse como lianas o arbustos erectos, incluso en la misma población y localidad (como en

Morro del Chapeu, Bahia, Brasil). Por lo tanto, el reconocimiento de variedades basado en este carácter no

es aceptable.

Tallos.—Los tallos son algo aplanados en ramas jóvenes, volviéndose cilindricos o subcilindricos con

la edad. Los coléteres son glándulas secretoras (Fahn 1979) que son comunes a muchos miembros de e

Echiteae. En Rhabadenia los coléteres interpeciolares son diminutos y rápidamente deciduos, por lo que

siempre están ausentes en tallos maduros; aunque en R. biflora son inconspicuos, en R. madida son relati-

vamente conspicuos en tallos jóvenes (Fig. 1A). Por otro lado, los coléteres intrapeciolares son diminutos e

inconspicuos y están presentes en la base del pecíolo. La presencia de coléteres intra e interpeciolares han

sido utilizados en algunos casos para separar géneros (e.g., Woodson 1933; Simões & Kinoshita 2002; Mo-

rales 2005), ya que mientras en algunas especies de géneros de las Mesechiteae éstos llegan a desarrollarse

notablemente (e.g., Mandevilla, Mesechites), en muchos otros géneros de las Echiteae éstos son inconspicuos.

Al cortar los tallos, ramitas y en general cualquier parte de la planta, se da la presencia de secreción lechosa.

Es importante tomar en cuenta que aunque el tipo de secreción aco vs. acuosa) ha sido considerado

un carácter de relativa importancia taxonómica en las A iteae y Echiteae), y que si bien

iw ser Ronde en varios grupos, existen otros géneros dondea a a og la prerane de secreción

i I a (e.g., A (Hemsl.) Woodson) y

incluso generas (como Echites), donde al ies ti secreción acuosa (E onarena] F. Morales)

y otras lechosa (como E. umbelatus mes

Hojas.—Las hojas son pecioladas, con los pecíolos largos o muy cortos y entonces las hojas parecen

subsésiles, y carecen de coléteres en el nervio central. La variación en la forma de la lámina foliar en Rhab-

dadenia es extrema y puede ser comparada con el que se presenta en algunos grupos de especies de otros

géneros de las Apocynoideae, como Echites (e.g., E. umbelatus Jacq.), Mandevilla (e.g., M. lancifolia Woodson)

o Mesechites (e.g., M. repens (Jacq.) Miers). Esta variación abarca tanto la morfología de la lamina foliar como

la forma de la base (Fig. 2). El análisis aislado de estos extremos de variación morfológica puede llevar a la

errónea conclusión de reconocer más taxones, pero un análisis integral demostrará que no son sostenibles.

En términos generales, R. i table en relación a la morfología foliar que las otras dos especies

del género.

La pubescencia está usualmente ausente, si bien un indumento leve e inconspicuo puede encontrarse

en R. madida, principalmente en la cara abaxial. La presencia o ausencia de indumento en Rhabdadenia no

es un carácter significativo para la distinción de taxones a nivel específico. La venación secundaria está

usualmente impresa, pero en forma leve, mientras que las venas terciarias usualmente no son evidentes.

Asimismo, los márgenes son siempre planos, no revolutos.

Inflorescencias.—Las inflorescencias son cimas dicasiales muy reducidas (Woodson 1935 a, 1936),

algunas veces con apariencia umbeliforme, usualmente con dos flores (Fig. 6A, C), pero con un rango que

oscila entre una a cinco. Como en muchos otros géneros de las Echiteae, las brácteas son siempre escariosas

y los pedicelos carecen de bracteolas. Los sepalos son esencialmente iguales, membranáceos, foliáceos y

carecen de coléteres en la base de su cara adaxial. Tanto Múller (1860 a) como Woodson (1936) propusieron

separar especies de Rhabdadenia (e.g., R. madida y R. pohlii) basado principalmente en el largo de los sépalos,

sin ERI se presenta una alta variación en relación al tamano y forma (Fig 2, lo cual A usar

Aas re +ralariao

el específico. Aún más, no existe ni

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sobre cada lóbulo (Fig. 3A—B); bajo este punto se puede trar un indumento denso de pelo

Morales, Monografia del género Rhabdadenia

Fic. 1. Caracteres morfológicos en Rhabdadenia madida (señalados por una flecha). A. Coléteres interpeciolares. B. Detalle de las hojas cordadas basal-

mente. C. Nectarios y ovario. D. Detalle de la pubescencia infraestaminal.

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rígidos, el cual se prolonga más o menos por un tercio dentro del tubo hacia la base (Fig. 1D). La superficie

externa es usualmente glabra, aunque si bien un leve indumento de pelos simples y diminutos puede estar

presente en R. madida. Los estambres están totalmente incluidos dentro del tubo y las anteras tienen un

indumento denso en su región dorsal y apical, el cual sobrepasa el ápice (Fig. 4A). Este tipo de indumento

es característico de Rhabdadenia, ya que mientras en otros géneros de las Echiteae (e.g., Peltastes, Prestonia)

pueden presentar indumento en las anteras de algunas especies, éste Imente se encuentra restringido a

la mitad basal y no a la región apical. Las anteras están unidas a la cabeza estigmática en la región del anillo

membranoso basal, por una agrupación de pelos en forma de herradura (Fig. 4B), en contraposición con los

miembros de las Mesechiteae, en los cuales las anteras se encuentran unidas por fusión celular a la cabeza

estigmática. De igual forma, las anteras se encuentran unidas medialmente a la cabeza estigmática por un

anillo de pelos corto. La cabeza estigmática es fusiforme, con un anillo membranoso conspicuo en la base

U te en el ápice (Fig. 4B). El ovario es invariablemente apocárpico y los carpelos se hallan fusiona-

dos en forma postgenital al estilo y se encuentran rodeados por cinco nectarios, los cuales usualmente no

sobrepasan en longitud al ovario (Fig. 1C). Muchos otros géneros de las Echiteae tienen nectarios anulares,

los cuales pueden ser casi enteros o irregular y variadamente pentalobulados. Aunque si bien el número y

Morales, Monografia del género Rhabdadenia

545

R biflora. B. Detalle

546 tani itute of Texas 3(2)

la lobulación del nectario ha sido usado para distinción a nivel específico, es un carácter que cuya utilidad

es cuestionable, dada la alta variación intraespecífica presente en algunos géneros (Morales 2004). Tanto el

ovario como los nectarios carecen de un indumento visible.

Foliculos.—Los folículos de Rhabdadenia son siempre fusiformes, continuos y algo divergentes hacia

su región apical, glabros o glabrescentes, más raramente diminutamente puberulentos ys son esencialmente

aja] en ] pos neq Taa Apo

lisos externamente, careciendo de costillas longitudinales. En forma gen

cynoideae los folículos están unidos en su ápice, aunque si bien en algunos pocos (e.g., Prestonia) falignlss

divaricados y divergentes estan presentes en unas cuantas especies (Morales 1997; Woodson 1936). Las

semillas son numerosas, rostradas y con coma (penacho de pelos) apical, con el rostro delgado y prolon-

gado, carácter compartido con muchos otros géneros de las Echiteae neotropicales. Algo peculiar para este

género es que el rostro de la semilla se prolonga más alla del punto de donde nace la coma en el extremo

micropilar, de manera que este se extiende de manera muy fina casi hasta el extremo terminal y distal de

la coma. En la mayoría ee los poe de las Echiteae y Mesechiteae, la coma nace aproximadamente de un

mismo punto, indey de que la semilla sea rostrada o truncada. A través de las descripciones,

las medidas de las illas abarcan desde el ápice chalazal hasta el punto donde nace la coma en el extremo

micropilar, por lo tanto, no incluyen la dins MN pe extremo micropilar más alla de este punto.

Relaciones intergenéricas.—En la clasificación de las d Bruyns

(2000), Apocynaceae y Asclepiadaceae fueron tratados de nuevo como un dol € taxonómico. En este

sistema de clasificación, Rhabdadenia fue ubicada en la subfamilia Apocynoideae, tribu Echiteae, grupo

conformado por cerca de 22 géneros de los cuales 19 están restringidos al Neotrópico (Endress & Bruyns,

2000). Otros sistemas propuestos anteriormente solo trataron los géneros de las Apocynaceae s. str. (e.g.,

Leeuwenberg 1994b; Pichon 19482, 1948b, 1948c, 1949, 1950 a, 1950b) y agruparon a Rhabdadenia con

algunos de los géneros de la tribu Echiteae sensu Endress and Bruyns (2000). En el sistema propuesto por

Endress and Bruyns (2000), algunas inconsistencias en la conformación de los miembros de las diferentes

tribus han sido corregidas en forma paulatina en los últimos años (e.g., Simóes et al. 2004, 2006, 2007).

Finalmente, en forma más reciente, Endress et al. (2007) propusieron nuevas modificaciones que impli-

caron la transferencia de cinco géneros a otras tribus (Amalocalyx Pierre [Apocyneae], Cycladenia Benth.

[Odontadenieae], Neobracea Britton [Malouetieae], Pottsia Hook. & Arn. [Apocyneael, Stipecoma Müll.Arg.

doce y la cases de 3 poneros recientemente descritos (Bahiella J.F. Morales, Mon K.

Williams) o i idos a la sinonimia (Rhodocalyx Mull.Arg.), de manera que la tribu Echiteae

ELLA ALA AS

quedó color por un total de 20 géneros, 17 de ellos Neciropicales:

A pesar de que se han ido conformando grupos más naturales, todavía falta trabajo por hacer, sobre

todo en definir las posible relaciones intergenericas prevalecientes en algunas tribus (como en Echiteae) y

establecer la conformación definitiva de otras. De hecho, algunos géneros de la tribu Echiteae se mad al

rado en forma adicional por solo un carácter morfológico (como Echites y Fernaldia) y

basados en indi lares y caracteres fológicos (e.g., Livshultz et al. 2007) han mostrado que

algunos deben ser reducidos a la sinonimia, apoyando lo sugerido en forma previa por otros autores (e.g.,

Williams 2004).

Existen pocos estudios filogenéticos que traten parcial o totalmente los miembros de las Echiteae, por

lo que las afinidades intergénericas de los miembros de la tribu no son claras. Algunos de estos estudios

en no ras incluido Mee (e. Bs Wiens dd mientras que los resultados de un análisis

filogenético basado 06)

Sues que pievaiece una m homoplasia y que por el momento, son ios SUI ica para

poder t dete mas certeza | ] : ] tes. En forma

más reciente, Livshultz et al (2007) realizaron un estudio filogenético er en WE moleculares y

caracteres morfológicos y sugieren que Rhabdadenia conforma una politomía con algunos miembros de las

Apocynoideae (clado del Nuevo Mundo [New World Clade] fide dco et al. 2007) y un clado compuesto

A 3 : Tasg C

por miembros de las subfamilias I y

Morales, Monografia del género Rhabdadenia 547

A nivel morfológico, Rhabdadenia se puede reconocer del resto de géneros de las Apocynoideae por la

siguiente combinación de caracteres: nee usualmente el a zonas anegadas estacionalmente o

manglares, tallos con coléteres inter[ ápid ] pareciendo ausentes, hojas eglandu-

lares, intl s. de ienci belif ae ee spal ] ] sin coléteres

4 r

en la base de la cara al amenas con indumento en la porción apical, esu compe por cinco

glándulas individuales, y folículos usualmente divergentes en su región apical, con semillas rostradas.

Notas sobre el tratamiento taxonómico.—Para cada especie y para cada país solo se citan tres

especímenes representativos por cada unidad geográfica (i.e., Provincia, Departamento, Estado), pero un

listado completo del material estudiado se cita al final. Los usos mencionados en algunas especies han sido

tomados directamente de las etiquetas de herbario, tal y como han sido citados por cada colector, por lo

tanto, no ha sido posible verificar su efectividad. Las distintas partes de las corolas (tubo basal y superior)

siguen lo descripto por Ezcurra (1981), Morales & Fuentes (2004) y Simóes & Kinoshita (2002).

TRATAMIENTO TAXONÓMICO

Rhabdadenia Müll.Arg., Fl. Bras. 6(1):173. 1860. Two: Rhabdadenia pohlii Múll.Arg. (Lecroriro, designado por Britton &

Millspaugh 1920).

Lianas o hierbas eae tallos con secreción lechosa, glabros o glal tes a puberulentos, aplanados

euando joven. lvié cerca de eso con la edad. lét int iol evidentes,

iol i i Hojas opuestas, raramente algún nudo con Hojas verticiladas, eglan-

dulares, sin calétenes en el nervio central adaxialmente, usualmente glabras, raramente puberulentas en la

superficie abaxial, membranáceas, sin domacios, márgenes no revolutos, pecioladas a subsésiles. Inflores-

cencias cimas reducidas, con menos de 5 flores, usualmente solo 2 ó 3 presentes, axilares o subterminales,

glabras o inconspicuamente puberulentas, brácteas escariosas e inconspicuas, bracteolas ausentes; cáliz

con 5 sépalos esencialmente iguales, diminutos a foliáceos o subfoliáceos, glabros o puberulentos en la

superficie externa, el ápice no reflejo, sin coléteres en la base de la cara adaxial; corola infundibuliforme,

internamente pentalobulada, con una cresta y un lóbulo diminuto sobre cada lóbulo alrededor del punto

de inserción de las anteras, purpura, rosada, blanco-rosada a blanca, raramente roja, usualmente glabra

en la superficie externa, con numerosas y conspicuas venas longitudinales, el tubo recto, limbo con 5

lóbulos, con estivación dextrorsa; estambres incluidos, insertos en la base de la parte superior del tubo,

anteras conniventes y aglutinadas a la cabeza estigmática, la base no auriculada, filamentos inconspicuos

o no evidentes, cabeza estigmática con un anillo membranoso conspicuo en la base y la parte distal más o

menos fusiforme, pubescente en el ápice; gineceo bicarpelar, apocárpico, con numerosos óvulos, nectarios

5, separados o irregularmente connados basalmente y formando un nectario pentalobulado. Frutos dos

folículos LA ar CHOR dc y más o menos paralelos entre sí, continuos, membranáceos, no

divergentes; , rostradas, comosas

Fhicionad

en el extremo a el pene "em aea más m del punto de inserción de la coma,

en forma de una cerda inconspicua.

Género con tres especies distribuidas desde el SE de Estados Unidos (Estado de Florida) hasta el N de

Argentina (excepto Chile) y las Antillas.

CLAVE PARA LAS ESPECIES DE RHABDADENIA

1. Parte inferior del tubo 31-34 mm ae tango; brácteas florales 8-11 mm de largo R. ragonesei

1. ids Mis del tubo (6-)8-20 m 1—6,5 mm de largo

2. Corola con la parte superior del tubo obcónica; lóbulos de la corola blancos, blanco-rosados o rosado

pi, Sh oa 1-1.8 i de argo colle nee tubo 12-20 mm de largo; lámina foliar con

e la bas S 6 de largo R. biflora

2. Corolar con la pare superior ae tubo dinare | lada; lóbulos de la co a pürpura, genes ro-

A lae DERE la | ta inf, 12mm

de largo; lámina foliar con la base la base cordada, subcordada 2 a redondeada O buda pecíolo

2-5(-6) mm de largo R. madida

[| I fal Dos * In LI PA fT e F óg b

548 J

1. Rhabdadenia biflora Jaca.) Müll.Arg., Fl. Bras. 6(1):175. 1860. (Figs. 3B, 4A, 5 y 6A—B). Echites biflorus

Jacq., Enum. Syst. Pl. 13. 1760. Tiro: T. 96 en Plumier, Descr. Pl. Amer. 1693 (Lecroriro, designado aqui).

Echites paludosus Vahl, Eclog. Amer. 2:19. 1798. Exothostemon paludosum (Vahl) G. Don, Gen. Syst. 4:83. 1838. Rhabdadenia paludosa

Eu Miers, uc S. Amer.: 119. 1878. Tiro: TRINI ini Pains Perle: (£D, von Rohr 21 Ene C a cartulinas] )

E dl., Linnaea 26:666. 1853. R Müll.Arg. ex Griseb. Fl. B I. 415, Linnaea 30:454.

1860. Tro: REPUBLICA DOMINIC ingo, fecha ueni G bag (no localizado). bos Micre, 1910 (fI),

uertes 220 (NeoTIPO: designado aquí, US; ISONEOTIPOS: MO. PU

Dipladenia billbergii Beurl., Kongl. Vetensk. Acad. Handl. 40:137. 1854 [1856]. Tiro: PANAMA. Coton: Portobello, abr 1826 (fl), Billberg

254 (HOLOTIPO: S).

(Mill) Miers, Apocyn. S. Amer.: 122. 1878. Apoc) Mill., Gard. Dict. (ed. 8) n. 9. 1768. Tiro: COLOM-

BIA. ATLÁNTICO: € , sin fecha (fl), Miller s.n. (HoLoTIPO: BM).

pI £a EMT Mie Apocyn. S. Amer.: 122. 1878. Apoc latum Mill., Gard. Dict. (ed. 8) n. 10. 1768. Tipo: MÉXICO.

VERACRUZ: datos senlidos (£D, Miller s.n. (4oLoTIPO: BM).

Rhabdadenia macrantha Donn. Sm., Bot. Gaz. 40:7. 1905. Teo: HONDURAS. Cortez: Puerto Sierra, ene 1903 (fl), Wilson 244 (HoLorro:

US; Isoripo: NY).

Lianas o arbustos escandentes, tallos glabros a glabrescentes. Hojas con el pecíolo 6-18 mm de largo, lámi-

nas (2.7-)3-11.2 x 0.5-4.2 cm, angostamente elípticas, elípticas, linear-elípticas a obovado-elipticas, más

raramente lineares, el ápice obtuso-mucronado a agudo-mucronado, la base obtusa o cuneada, glabras, las

venas secundarias usualmente impresas, aunque a veces algo inconspicuas, venas terciarias evidentes o

no evidentes. Inflorescencia con 1 a 3(5) flores, usualmente glabra, pedúnculo 1.7-6.8(-8.4) cm de largo,

pedicelos 12-21 mm de largo, brácteas 1-1.8 x 1-1.3 mm, angostamente ovadas; sépalos (1.5-)2-11 x

0.8-3.5 mm, ovados, dad T el aes Agudo a aul -mucronado u obtuso-mucronado, foliáceos o

subfoliáceos, más raramente diminutos ola con el tubo blanco en su parte externa,

los lóbulos blancos, blanco-rosados o rosada pálidos, parte mone del tubo 12-20 x 23.5 mm, la parte

superior obcónica, 19-28 mm de largo, con un diámetro de 1.3-1.6 cm en la boca, el ápice del botón floral

apiculado o cortamente acuminado, lóbulos 1. ño: 4x : mU 1 cm, obovados; anteras 4.1—4.7 mm de largo,

glabras dorsalmente, excepto por la pul pical da; ovario 1.2-1.6 mm de largo, ia

cabeza estigmática 1.4—1.7 mm de fara — 1/2-2/3 de la longitud total del ovario, lil algunos

ellos irregularmente connados basalmente. Folículos 9.1-18.2 cm x 1.5-4.5 mm, ¿os semillas 2.4-4.1

cm de largo, glabras, la coma 3-4.1 cm de largo, color crema

Distribución, habitat y ecología.—S de Estados Unidos (estado de Florida) a través de Mesoamérica hasta

el N de Ecuador y el N de Brasil y las Antillas, donde crece en manglares o formaciones vegetales asociadas,

en elevaciones de 0-10 m. Especimenes con flores y frutos han sido recolectados durante todo el año, pero

las flores son más abundantes durante el período de lluvias.

Nombres comunes y usos. —“bejuco lechero” (Venezuela, Monagas); “cai seca” (Brasil, Pará, Marajó);

“cipó de carangrejo” (Maranhao, río Maracassumé); “dub inanusu” (Panamá, San Blas, Isla Urbili, nombre

Kuna): “lecho de mar” (Colombia, Nariño, Tumaco); “liane mangle” (Guyana Francesa, Cayenne, Mahury);

“mangrove vine” (Belize, Belize city); “palo de leche” (República Dominicana). Esta especie ha sido usada

como destumescente, mediante un baño con decoción y aplicándola sobre el área afectada (Arvigo et al. 365,

MO, NY). También ha sido usada en baños contra dolores de cabeza y mareos.

DL5alhdss4 : lu ] A 4 IE ta Asare O A as

[^ |

J h E e 7 r

por la siguiente combinación de caracteres: hojas conspicuamente pecioladas, con los pecíolos usualmente

de 17-84 mm de largo, y corolas con la parte inferior del tubo de 12-20 mm de longitud, la parte superior

obcónica. Aunque en términos gente y sobre todo, comparándola con la alta variación intraespecifica

en cuanto a la variación morfológica de los hojas, la

presente en R. madida, R. bifl

misma variabilidad en en el tamaño, forma y e de los sépalos presente en R. madida se encuentra en

] sobrepasen los 4 mm de largo, se

este taxón. De esta manera, aunque es comün que los sépalos

pueden encontrar numerosos especímenes, sobre todo provenientes de las Antillas, donde los sépalos rara-

mente sobrepasan los 3 mm de longitud, encontrandose en un rango de 1.5-2.5 mm. El pretender reconocer

estos especímenes como un taxón aparte no parece correcto, por que no existen diferencias adicionales en

el resto de caracteres morfológicos que puedan justificarlo, aün a nivel de variedad.

Morales, Monografia del género Rhabdadenia 549

Fic. 5. Rhabdadenia biflora (A de Aguilar 6205, INB; B—G de Morales 3070, INB; H- de Cediflos 3371, INB). A. Ramit infl ias. B. Ramita con

inflorescencias. C. Vista de 1 laxial de d ápal i trando | ia da colát la | n Tuha de | la 2g mE

trando i ició | t E. Antera, vista dorsal. F. Cab tig ética. G. Nectari y ia H Falículas | Semilla

3 I fl n.a H In LI PE a E fT 379)

550

Se designa un neotipo para Echites ehrenbergii Schltdl., dado que la única colección citada en la descri-

pción original (C. Ehrenberg s.n.) no fue localizada. Es probable que este espécimen haya sido destruido en

Berlin durante la Segunda Guerra Mundial. En la monografía de Woodson (1936), se citó Echites billbergii

Beurl. como un sinónimo de R. biflora; sin embargo, en la publicación original (Beurling 1856), dicho taxón

fue descrito como Dipladenia (Echites?) billbergii, por lo cual es citado de manera correcta aca.

Especimenes representativos examinados: ESTADOS UNIDOS. Florida: Eo ied o n Rein 27 jul 1966 (fI),

Almeda & Lakela 285 (US, USF); Miami, jun 1880 (fl, fr), Curtiss 2266 (BR, LV, P, US) Dade, 4 mar 1930 a

Moldenke 725 (MO, NY, US) MEXICO. Campeche: Sanación, Ius 1932 (f. St 1910 (MO); Calki EAE Isla Arena, 28 no

1993 (fl, m Tun et al 126 y Mog Quintana R i lombia, isla Cozumel, 10 jul 1984 a

rera & Cabrera 6762 (MEXI los, S de Xel-ha, 6 may 1980 a pa 2050 (MEXU, MO); Vigía Chico, 11 may 1980

m Téllez 2238 (MEXU, MO). Tubascd: carretera Mecoacan-Chiltepec, 23 ni 1981 (fl), Magaña & Zamudio 309 (MO); Centla, al 5 de

Puerto San Pedro, Laguna El Cometa, z Rp 1998 r Novelo & Ramos 3105 (MO); Centla, laguna Pez Espada, 14 dic 1998 (fD, Novelo

& Ramos 3618 (MO). Vi ha, 20 jun 1972 (fl), Dorantes et al. 1201 (MEXU, U); Laguna de Sontecoma,

Catemaco, 25 feb 1973 (ID, Menéndez 97 (MEX, MO), Suite istmo de pea 12 ene 1895 (fl), den 1080 (EAP). Yuca-

tán: Puerto Silam, 1895 (fl), Gaumer 653 (US Silam, may 1916 (fl), Gaumer 23335 (EAP, F, MO, US); S de

E] Progreso, 23 mar 1970 (fD, Utley 39 (MO, USE, WAG). BELICE. Belice: ciudad Belice, ue highway, 30 mar 1990 (fD, Arvigo et

2 365 us NY}; St. cam A 15 mar ibd d aD Mi id et al. 11 (MO); entre ciudad de Belice E v MA 9 bi 1992 oe Wor-

, 17 mar 1987 (fl), Davidse & B Chunox,

18m mar 1987 (fl), Davidse & Brant 32622 (MO). St k: | ia, 30 dic 1985 (i. fr), Walter 1107 (MO). Toledo: río Monkey

1 dic 1941 (£D, Gentle 3826 (MO, US). GUATEMALA. Izabal: Puerto Berrios, 2 abr 1922 (fl, fr), Greenman & Greenman 5986 (EAP, MO);

Machacas, a Chocón, 6 mar 1993 (fI), Päll et al. s.n. INB, UVAL); is dis Laguna aue 28 ne 1988 n fr), Tenorio et al.

14941 (IN XU, MO). Petén: San Pedro, 8 ene 1970 (fl), Contreras 9471 (MO). HONDURA Micos, NO de

Tela, 10 nov y 1988 (fl), MacDougal et al. 3435 (MO, WAG), Puerto Escondido, O x Tela, 11 feb 982 (fD, Nelson 7854 (MO, TEFH), entre

barras de Cuero y de Salado, May—Ago 1989 (f1), Soto 485 (TEFH). Colón: Puerto Castilla, 17-20 jul 1973 (ED, Nelson & Martinez 1154

(EAP, TEFH); NE de Trujillo, camino viejo a Castilla, 23 sep 1981 (fl), Saunders 619a (MO); entre Trujillo y Castilla, 10 ene 1981 (D,

Saunders et al. 854 (MO, TEFH). Cortés: playas de Tulián, 16 jun 1952 (fl), Molina 5208 (EAP, US); entre Tulián y Cortes, 12 may 1950

(fD, Molina 6951 (EAP); laguna de Vedián, al O de Barra de Ulúa, 26 mar 1993 (£D), Nelson & Andino 15500 (EAP). Gracias a Dios: río

n villa de Ras, 23 may 1973 (fl), Gentry et al. 7553 (MO, TEFH); Tuas, O de Brus, abr 1971 (f1), Nelson & Hernández 968 (EAP, MO),

osquitia, Puerto Lemira, laguna Caratasca, 17-21 jul 1977 (fl), Nelson & Romero 4245 (EAP, TEFH). Islas de la Bahia: Isla de

oatan, 19 abr 1967 (fD), Molina 20648 (EAP, US); Flowers Bay, 17 sep 1982 (fl), Nelson 8503 (TEFH, US); on de Barbareta, 9 may 1984

(fl), Nelson & Cruz 8895 (TEFH). EL SALVADOR. La Unión: Isla Martin Pérez, Golfo de Fonseca, 4 dic 2006 (st), igi 14599 (INB,

MHES). NICARAGUA. Zelaya: Corn Island, 9 mar 1971 (fl), Atwood 4393 (MO); cercanías de ede 13a 961 (ID, Bunting &

Licht 1278 (F, US); o in 23 jun 978 nx e 4563 T m COSTA RICA. Guanacaste: Palo Verde, río o 23 jun

1993 (fl), Chavarría 823 (CR, F, INB), P Nicoya, 13 UP. id (fD, Gómez 20567 (CR). Limón: parque nacio-

nal Cahuita, Puerto Viejo, Es 2000 (fl), Acosta et dl 2202 (INB, MO); ent y quebrada Ernesto, 2 ago 1984 (I), Grayum

& Sleeper 3658 (CR, MO); Manglar Gandoca Manzanillo, 24 jun 1 983 (fD, Soto s.n. (CR). Puntarenas: parque marino Ballena, 29 jun

1994 (fI), aes 399 ow INB, MO); Quepos, 7 jul 1994 (fl), Morales 3010 (CR, INB, MO); Golfito, Golfo Dulce, mar 1896 (fI), Pittier 9903

(BR, CR, US). P MÁ. Bocas del Toro: S de Chiriquí Grande, 25 mar 1985 (fl), Hampshire & bi 948 (BM, MO); S de Tigre

Key, 25 feb 1989 ipd Peterson & Annable 7129 (MO, US); Water valley, 8 nov 1940 (fl), Wedel 1546 (MO, US). Chiriquí: río de David,

dic 1881 (fD, Lehmann 1000 (US); Estero Rincón Largo, SSE de Horconcitos, 22 sep 1983 (fD, Schmalzel & Stedtman 1789 (MO); San

Bartolomé, Península de Burica, 28 Jul-1 ago 1940 (fl), Woodson & Pa ons b Colón: pup pai ae 25 ago 1978 (fl), Hammel

4572 (MO, WAG); playa Langosta, al E de Colón, 12 ago 1979 (fl), Meije , 30 may 1971 (fl), Thoms

2 (MO, PMA). Darién: La Miel, 18 jul 1985 (ID, Cuadros 2239 (COL); Eb Carey, 22 ago 1964 (1D, iel 5145 (MO); E de Santa

Fé, 16 jul 1966 91D, Tyson et al. 4670 (MO). Panamá: cerca de la boca del rio Chagres, 15 mar 1935 (£D. Allen 885 (MO, P, US); entre

Balboa y Chamé, 2 dic 1934 (fP), sd et al. 16727 nA un Barro oe 16 ene 1982 e e. 316 (MO); Coco Solo, 27 sep

1972 (fl), Tyson 6806 (TEFH). S y Cangandi, 10 dic 1985 (fl), de Nevers et al. 6441 (MO); Mu-

latuppu, río Ibedi, 7 ago 1966 a Duke 8477 (MO); San Blas, Bc oct 1978 (m. Hammel & ac 5038 (MO). Veraguas: Isla de Coiba,

Santa Cruz, 17 oct 1997 (fl), Ibañez et al. 543 (MA); bahía pm Lerén, 7 ago 2002 (fD, Ibañez et al. 2141 (MA), Isla de Coiba, rio Santa

Cruz, 21 nov 1994 (fD, Martin-Ballesteros et al. 624 (MA). C MBIA. Antioquia: Turbó, Bocas del Atrato, Vereda Turbito, Bahia Co-

lombia, Golfo de Urabá, 4 ago 1987 (ID, na et al. 5030 MA MO); río ~, a zu 1985 (fl), Renteria et al. 4285 (JAUM, MO);

Arboletes, 30 sep 1986 (fl, fr), Roldán et al. 516 (COL, HUA [2 cartulinas], M ico: Cartagena, 1805 (fl), Bonpland 1354 (P);

Barranquilla, Las Flores-La Playa, 13 sep 1959 (fI), Cuatrecasas 24295 (INB, US): e E ene 1928 (fl), Elias 371 (US). Chocó: Bahía

Solano, corregimiento El Valle, entre del Valle y Almejal, 15 abr 1989 (£D), Espina et al. 2453 (CHOCO, HUA, INB); boca del río San Juan,

9 ene 1947 (fl), Haught 5416 (COL, MO, US); Playa de Togoramá, 12 jun 1944 (£D), Killip & Cuatrecasas 39078 (US, VALLE). Magdalena:

Ciénaga, 30 mar 1942 (fl), 13 feb 1950 (£D, Romero-Castañeda 1925 (COL, MEDEL); Santa Marta, 1898-1901 (fD, Smith 1664 (G, NY, P,

US). Nariño: bocas del río Sanquianga, La Tula, 12 abr 1976 (fI), Cabrera & Escallón 4013 (CUVC); Tumaco, río Chagui, entre Vereda

Morales, Monografia del género Rhabdadenia 551

Chajal y Guadual, 16 jun 1989 (fl), Madrigal & Mesa 661 (COL, HUA). San Andrés: Isla de San Andrés, 22-27 abr as D, Proctor 3312

(US). Sucre: Tolú, Arroyo Villero, carretera Tolú-Coveñas, 18 d xis d Bee ets Berrio 1983 (COI

19 ago 1979 (tl), Bernal 133 (COL, P); San Onofre, g yó Socó y man glares, 7 de 1996 (fl, 2 ed 170 0 HUA

Valle: boca del río Yurumangui, 24 feb 1983 (fl), G & 40726 (COL, MO); Punta Arenas, N 2 jun 1944 (D,

Killip €» Cuatrecasas 38622 (MEDEL, VALLE, US); bahía Málaga: vereda La Plata, 18 abr 1992 (£1), Otero & Sandino 157 (CUVO. VENEZU-

ELA. Amazonas: Delta del Orinoco, piene icd due d 0), Midi 593 (US, VEN). Delta Amacuro: río Amacuro, cerca de

San José de Amacuro, 21 nov 1960 (fl), S , boca del Caño Guiniquina, 18 oct 1977 (1D), Steyermark &

Liesner 114892 (MO, VEN); río Cuyubini, bs 1955 CD, Wurdack 324 (US, VEN). Falcón: Hacienda Guacabana, base O del cerro de

Chichiriviche, parque nacional Morrocoy, 23 feb 1993 (f1, E mete & du i id Mn 2 cerro de Chichiriviche, S de

Golfete de Guare, 13 ago 2002 (fD, Dorr & Niño 9100 (INB, | t sta Playa Sur, 24

sep 1993 (fl), Martin-Ballesteros 151 (MA). Miranda: Paparo, boca del Rio Grinde: Barlovento, 11-16 jun 1913 (El, E Pittier 6352 (US):

Silva, caño Ramadita, al N de Boca de Aroa, 1 sep 1974 (£D), Steyermark & Manara 110529 (VEN); Silva, La Peñita, Golfete de Guare, 30

ago 1974 (fl), Steyermark & Manara 110893 (VEN). Monagas: reserva florestal de Guarapiche, 18-20 jun 1965 (fl), Berti 699 (VEN).

Sucre: Península de Paria, San Juan de las Galdonas, 1995 (fl), Allorge 1023 (P); cercanías de Cristobal Colón, 5 Ene-22 feb 1923 (fD,

Broadway 428 (GH, US); Quebrada El Purgatorio, jul 1955 (fl), Lasser & Vareschi 3863 (VEN). wes eer La Ceiba, 10 op 1978 =

Stergios 1375 (VEN). Zuliá: entre Río Grande y Mej S Península de Paria, 22 ago 1961 (fD, A (VEN); entr

Santa Barbara y San Carlos del Zuliá, ensena 30 nov 1966 (fl), Bruijn 1301 (U, US, WAG, VEN): e entre Bella Vista y —

Rosa, cerca Maracaibo, 7 oct 1922 (£D, Pittier 10480 (US, VEN). TRINIDAD: Manzanilla, sep 1940 (fl, fr), Baker & Broadway 412 (MO):

carretera Cocos, suampo Nariva, 21 jul 1977 (fI), Philcox et al. 8045 (K, P); Manzanilla, 12 mar 1950 (fl), Simmuels 448 (K, MO). PUER-

TO RICO: Playa de la Carmelita, abr 1883 (fl), Eggers 836 (BR, P [3 cartulinas], W); Bayamon, 22 mar 1885 (fl), Sintenis 935 (S, US);

Martín Peña, 25 ene 1914 (fl), oe 433 (US). BAHAMAS: Cerca de Providence, Lake Cunningham, 23 mar 1974 (fl), Correll &

Correll 41941 (FTG, MO) Providence 14 mar 1946 (fl), Degener 18738 (MO, US); Nueva Providencia, lago Cunning-

ham, 11 jul 1960 (£D), Webster et al. 10410 (US). GRAND CAYMAN: Sureste de Georgetown, 6 Ja 1938 CD, Kings 199 (BM, MO); Snug

Harbour, 16 jun 1974 (fl), Stoddart 7014 (MO, US); West Bay, 17 jun 1974 (£1), Stoddart 7039 (US). L : Suroeste de Mary's

Bay, 5 ago 1975 (fL), Proctor 35116 (US). CUBA. Granma: 5 de a 12 jul 1985 (fl, S: abis 51015 (MO). Habana: Habana,

s.d. (£D, Boldo s.n. (MA); Vedado, 1904 (€D, Hermann 427 (P); al N Isla de Pinos, 6—10 feb 1956 (£D, Morton apie (US).

Isla de Pinos: río Las Casas, cerca de Nueva Gerona, 3 feb 1953 (fl), Killip 42583 (US); río pid 27 feb 1953 (fl), Killip 42954 (US);

camino a Bibijagua, 21 mar 1953 (ŒI), Killip 43152 (US). Matanzas: Buena Ventura, 10 abr 1994 (fl), Acevedo-Rodriguez et al. 6347 (US);

cero de Cananova, ago 1948 (fl), Clemente 6201 (US); entre Varadero d , 20 jun 1988 (fL), ibd qa ee

10638 (MA). Oriente: Ensenada de Mora, 26-29 mar 1912 (fl, fr), Britai " al. 23070 (US). P del rí Afuera

Bahía Honda, 14 dic 1910 (fl), Y s 9264 (U). Santa Clara: Cien Fuegos, río Dauji, 23 may 1895 (fD, Combs 78 (MO, P, US); Gavilan.

jun 1941 (D, Howard 4941 (MO, US); Cien Fuegos, Belmonte, Soledad, 6 ago 1927 (fl), Jack 5242 (P). REPÚBLICA DOMINICANA.

Distrito Nacional: Santo a Maimón, jun 1887 ab, Eggers 2676 ©), ey ipium o ene 1823 (ID, Ritter s.n. (W). Espaillat:

cerca de vee Hernández, 5 abr 1959 (fD, Jiménez 1959 (US). María Trini gua, 20 mar nied a ee 14529

(NY, US). S ta, al S de Sánchez, 1 may 1921 (fl), Abbott 1444 (US) de Sánchez, penínsu , 24 may

1922 (fr), Abbott 2417 (US); entre a, y Punta Rey, 29 nov 1923 (fl), Abbott 2847 (US). HAITÍ. Grand Anse: Grande Cayemite,

20 ago 1927 (fl), Eyerdam 321 (MO, P, US); Miragoane, 15 sep 1927 (fl, fr), Eyerdam 559 (P, US). Nord Ouest: cercanías de Basse Terre,

Isla Tortue, 22 mar 1929 (fl), Leonard & Leonard 13916 (US). Ouest: Massif de la Pelle, 5 ene 1927 (fl), Ekman 8053 (S, US); Port au

Prince, 17 may 1942 (ID, Holdridge 1222 (MO, US); Papet, entre Leogane y Grande Goave, 6 jul 1941 (fl), Seibert 1719 (MO, US). JA-

MAICA. Clarendon: rocky point Bay, 21 nov 1963 (fT), Proctor 24219 (U). Manchester: base NO de Round Fu, Miu 1980 (f), Gentry

& Kapos 28373 (MO). Portland: Puerto iria 28 jun 1897 (fr), Fredholm 3071 (US). St. And ts Bay, 22 feb

(fl), Maxon & Kllip 321 (P, US). St. A B St. Annes, 9 mar 1916 (fL), Killip 533 (US). St. TI il

14 ene 1958 (fl), Yuncker 17961 nen MARTINICA: Lamantin, río Saleé, 1883 b xiii: uid AUN rio eds. 14 j on 1940 om, Stehlé 6 &

Stehlé 4562 (US). GUADALUPE: Grande Terre, petit canal, nov 1990 (fI), Billiet € o Salec, 1893

(£D, Duss 7 (P); Marécages, 2 feb 1939 (f), s 4188 (US). ANTIGUA: Millars Old Works, M 1937 (fl, fr), Box 576 um GRENADA:

St George, dic 1887 (fl), Eggers 6510 (P, US) GUYANA: Demerara-Mahaica, entre Cane Grove y Lama, 10 feb 1988 (f1), Hahn et al. 3808

(P, U, US, WAG); río Berbice, al 5 de New ned 5 oct 1981 (£D, Maas et al. 5548 (MO, P, U, WAG, VEN); Pointe Macouria, Cayenne,

21 ene 1978 (ID, Raynal-Roques eni e U). GUYANA FRANCESA: ae pda dic 1989 (11), Billiet & Jadin 4734 (BR, MO);

playa de Susini, entre Bourd Montjolv, 21 dic 1986 (£1), C / NY, P, U, US); I , Mahury, La Levée,

18 ene 2001 (fl), Prévost 4103 ce MO, K, P, D. T Peperrot, 5 mar 1951 (£D, Florschüt — Gn 16 77 (BR, oo Paramaribo,

24 mar 1961 (fD, Kramer & Hekking 3130 (U, , camino a Carl Francois, 2 jun 1944 (fI), iin i & s 23596 bd

U, US). ECUADOR. raldas: La Tola, 22 feb 1994 (fl), Cornejo 1890 (GUAY, QCNE); N de Atacames, carretera Esmeraldas-Muisne

31 jul 1980 (fl), Hansen et aa 7931 (MO, USF); San José de Chamanga-Salina, 28 feb 1993 (£D, Jaramillo 15203 (QCA). BRASIL. Amabi

NO de Amapá, 7 nov 1979 (fl), Austin et al. 7339 (MG, US, WAG); Amapá, 3 jun 1944 (fl), Baldwin 4096 (US). Pará: Ilha do Marajó, 12

nov 1948 (fl), Black 3428 (IAN, MO); Marapánim, Camara, al NO de Marudá, 3-4 abr 1980 (£1), Davidse et al. 17838 (US, WAG); Belém,

27 may 1947 (tl, fr), Pires & Black 1619 (P, US). Maranhão: rio Maracassumé, 9 sep 1932 (fl), Froes 1886 (P, U); localidad perdida, 1856

(fl) Gardner 6060 (BM, CGE, P, US, W [2 cartulinas).

552 Į | £a nas s Im LI di dan ndi ETa as 3(2)

2. Rhabdadenia madida (Vell.) Miers, Apocyn. S. Amer. 121. 1878. (Figs. 1, 2, 3A, 4B, 6C-D y 7). Echites ma-

didus Vell., Fl. Flum. 112. 1829. Tiro: BRASIL. Rio De Jankmo: Vell., Fl. Flumin., Icon 3: tab. 42. 1827 (ectoriro, designado aquí).

Rhabdadeni (Benth.) Müll.Arg., Linnaea 30:435. 1860. Rhabdadeni (Benth.) Müll.Arg. var. glabrata Müll.Arg.,

innaea 30:435. 1860. Echites macrostomus Benth., J. Bot (Hooker) 3:248. 1841. Tiro: GUYANA: localidad perdida, 1837 (fl, fr),

nne 329 me K; isompos: BM [fotografias, INB], CGE, G-DC [foto F neg. 7564], P [2 cartulinas, foto F neg 38759], P,

US, W [2 cartulinas}).

Rhabdadenia macrostoma Sena ) Müll.Arg. var. pubescens Múll.Arg., Linnaea 30:435. 1860. Tiro: SURINAM: Paramaribo, jun 1830 (1D),

Splitgerger s.n. (HOLOTIPO: W).

Rhabdadenia pohlii Müll. ds Fl. Bras. 6(1):174, pl. 52. 1860. Rhabdadenia pohlii Müll.Arg. var. volubilis Müll.Arg., Fl. Bras. 6(1): 174.

1860. Tro: BRASIL. Esprerru Santo: localidad exacta perdida, 1816-1821 (fl, fr), St. Hilaire 328 (Lecroriro, designado aquí, P [2

tulinas]).

ia Mull Arg. var. suberecta Müll.Arg., Fl. Bras 6(1):175. 1860. Rhabdadenia latifolia (Múll.Arg.) Mal ! ta (Mall

Arg.) Malme, Ark. Bot. 21A(6):18. 1927. Tipo: BRASIL. Minas Gerais: localidad perdida. 1816-1821 (fl), St. Hilaire s.n. omo

designado aquí, P [2 cartulinas]).

D Arg. var latifolia Mull Arg , Fl. Bras. 6(1):175. 1860 Rhabdadenia latifolia (Müll Arg.) Malme, Ark. Bot. 21A(6):17

rasil. Rio GRANDE DO Sur: sin localidad precisa, 1835 (fI), Isabelle 6 eens P.

E mamorensis Rusby, Mem. New York Bot. Gard. 7:326. 1927. Tiro: BOLIVIA. Santa Cruz: cerca de Trinidad, 4 mar 1922 (fl),

Cárdenas 24 (HoLoripo: NY; 1sormo: US).

Lianas, hierbas suberectas o erectas, tallos glabros, glabrescentes a esparcidamente puberulentos. Hojas

con el pecíolo 2-5(-6) mm de largo, laminas (2.5-23.2-8.2 x 0.6-2.7(-2.9) cm, angostamente ovadas,

angostamente ovado-elípticas a elípticas, más raramente obovado-elípticas, el ápice obtuso-mucronado,

redondeado-mucronado, agudo-mucronado o agudo, algunas veces muy cortamente apiculado, la base

cordada, subcordada a redondeada o subtruncada, glabras, glabrescentes a densa a moderadamente pu-

berulentas en ambas caras, las venas secundarias usualmente impresas, aunque a veces algo inconspicuas

adaxialmente, venas terciarias no evidentes o levemente impresas abaxialmente. Inflorescencia con (1223)

flores, glabras o densa a moderadamente puberulentas, pedúnculo 2.5—8.7(-15.5) cm de largo, pedicelos

5-15 mm de largo, brácteas 3.5-6.5 x 0.7-1 mm, angostamente ovadas a linear-ovadas; sépalos (1.5-)2-10

x 0.6-2.2 mm, angostamente ovados, angostamente elipticos, linear-ovados a lineares, el ápice acuminado,

foliáceos, glabros a puberulentos externamente; corola con los lóbulos pürpura, rosados a rosado intenso,

raramente rojas (Guareco 566, INB), la parte inferior del tubo angostamente cilíndrica, (6-)8-12 x 1.1-2.1

mm, la parte superior cilindrico-campanulada, 22-35 mm de largo, con un diámetro de 1-1.4 cm en la

boca, el ápice del botón floral acuminado, lóbulos 1.5-2.5 x 1.2—2.1 cm, obovados; anteras (3.5-)5-7.5 mm

de largo, glabras dorsalmente, excepto por la pubescencia apical, la base redondeada, cabeza estigmática

1.3-2 mm de largo; ovario 1.3-1.6 mm de largo, glabro, nectarios 1/2—2/3 de la longitud total del ovario,

usualmente libres. Folículos 7.5—8.8 cm x 13.5 mm, glabros; semillas 2.1-2.4 cm de largo, glabras, la coma

1.7-3.9 cm de largo, blanca a blanco-crema.

Distribución, habitat y ecología.—Ampliamente difundida en casi toda Sur América (excepto Chile),

donde crece en áreas inundadas, orillas de ríos, formaciones arbustivas en áreas arenosas, así como pan-

tanos asociados a sabanas y bosques de Chaco asociado a zonas fangosas arenosas y bordes de esteros, en

elevaciones de 0—500 m. La floración y fructificación ocurre durante todo el año.

Nombres comunes.—“huevo de gato” (Perú, Loreto, Bella Vista).

Esta especie se puede separar ee ids aa ind ragonesei por sus flores con la parte inferior del tubo de

1 2

la corola más pequeño, así como brác pequeñas y hojas usualmente más anchas. Rhabdade-

nia madida es una especie polimórfica, tanto en lo referente al hábito y las hojas, como en el tamaño de los

sépalos, lo cual ha sido comentado por varios autores (Ezcurra 1981; Ezcurra et al. 1992; Woodson 1936).

De esta manera, esta especie puede ser una liana o comportarse como una hierba escandente o suberecta

(Ezcurra 1981; Ezcurra et al. 1992; wee a eae oe en bw general es más común el hábito de

liana en todo el rango de distribución, los hál en las poblaciones presentes en el

S de Brasil, E de Bolivia, Paraguay, Uruguay y Argentina. En todo caso, en algunas poblaciones presentes en

determinadas regiones de Paraguay se pueden encontrar ambos hábitos. Al estudiar en forma aislada cada

Morales, Monografia del género Rhabdadenia

Fic. 6. Inflorescencias en Rhabdadenia. A-B. R. biflora. C-D. R. madida.

554 [| || £ al Dang H ID LI F e Y f Texas 3(2)

LEE a | a ros J I J s 1

estigmática. G. Nectarios y ovario. H. Folículos. I. Semilla.

al J [| i J | : J lia || L

D. Corola. E Antera, vista dorsal. F. Cabeza

Morales, Monografia del género Rhabdadenia 555

una de estas variaciones sin un análisis morfológico exhaustivo en todo el rango de distribución geográfica,

puede llevar a la conclusión errónea de reconocer diferentes taxones (a nivel de especie o varietal) (Múller,

1860), pero al examinar la gran cantidad de estados intermedios es claro que esto no es posible.

Otro carácter que tiene una alta variación dentro del género y en particular en esta especie se refiere

a la longitud de los sépalos. Woodson (1936) separó R. madida de R. pohlii basado en el indumento de los

tallos (esparcidamente pilosos vs. glabros) y en la longitud de los sépalos, de manera que en R. madida dio

una longitud con un rango de 2-4 mm, mientras que en R. pohlii la variación que citó era de 8—12.5 mm.

Es probable que estos rangos se E E en e apenas pocos peces Finds por 2 (6

especímenes de R. madida y 9 de R. pohlii), | material

en día, no son sostenibles ni excluyentes, ni siquiera a nivel de distribución geográfica. Esta seen había

sido comentada previamente por Monachino (1958), quién cuestionó la ee de los carac teres utilizados

rnlersrión

por Woodson. Todavía más concluyente es la variación presente en los sépalos de g

Smith & Klein 8305), donde la longitud de los sépalos varía en un rango de 2-7(—9) x 0.5-1 mm. En todo

caso, la misma variación en E ¡SABINA de ne sépalos esta ete en R. ke donde algunas poblaciones

de las Antill relación a | n el Continente Americano). Por

lo tanto, el mantener Eu R. madida de R. polii basado en la longitud de los sépalos no es justificable.

Situaciones similares se han presentado en géneros como Odontadenia (Apocynoideae, Apocyneae), donde

diferentes taxones fueron separados basados en variaciones mínimas de ese carácter (Woodson 1935b), los

que pO ne han ED ae ser ados en monogra: recientes (Morales 1999). Por otro

lado, aunqu e , este carácter

varía de forma ind liente en relación a B longitud de los sépalos y aneoce es consistente o excluyente

para justificar la aceptación de R. pos

En orma g neral es comün Í de Mandevilla tifolia (Mal ) Woodson identi-

ficados como Rhabdadenia madida, ya que MC especies comparten hojas con ilias foliares de similares

formas y dimensiones y convergen en su distribución en algunas regiones geogfráficas, lo cual había sido

anteriormente mencionado por Ezcurra et al. (1981, 1992). Sin PUDE a dn vegetativo y sin entrar en

un análisis detallado de las flores, los especímenes de M. angustifoli separar con facilidad por sus

inflorescencias racemosas, usualmente con numerosas flores, así como yor sus tallos viejos con los coléteres

interpeciolares comúnmente desarrollados en tallos viejos y simulando pequeñas garras, lo cual no se pre-

senta en Rhabdadenia (Ezcurra 1981, 1992). Otro carácter adicional también útil, se refiere a la presencia de

coléteres en la base del nervio en la superficie adaxial de las hojas, lo cual es algo común en Mandevilla. Sin

embargo, este tipo de coléteres son rapidamente caducos y a veces parecen estar ausentes en M. angustifolia

(e.g., Morales 9325, INB), por lo que este carácter debe utilizarse en combinación con el resto de caracteres

anteriormente mencionados.

En su monografía de Rhabdadenia, Woodson (1936) relegó el taxón Echites madidus Vell. a la sección

de especies dudosas o excluidas y sugirió la posibilidad de que se tratara de una especie de Mandevilla. Sin

embargo, rand en E MIR de la MUCH que Reap E pd RAE original, donde se muestran

que ocurre ), hojas con la base cordada (Fig.

a aL.

1B), sépalos sin coléteres y el ovario con cinco nect te dimi y separados, se deduce que

este epíteto específicorepresenta el nombre correcto a ser usado para Rhabdadenia pohlii, por tener fecha de

publicación más antigua.

La colección Saint Hilaire 328 es seleccionada como el lectotipo de Rhabdadenia pohlii, ya que es la

colección mejor preservada y con mayor nümero de duplicados (2), mientras que el resto de los sintipos

citado por Müller (Gaudichaud 85, Pohl 5170) son conocidos por una ünica colección, con la excepción de la

colección realizada por Sello (s.n.) que no pudo ser localizada. En todo caso, es importante aclarar que existe

una colección adicional (St. Hilaire 527) en el herbario de París (P) y anotada por Müller como R. pohlii, que

también consta de dos duplicados, pero representa una planta suberecta (como se infiere fácilmente de la

colección), por lo que dado su discrepancia con el protólogo original (donde se especifica que el taxón es

“voluble”), no es escogida como el lectotipo.

|| [| £ al n.a H ID L. ] rr lo F da kl

556 f Texas 3(2)

De las colecciones citadas por Müller (1860 a) en la descripción de Rhabdadenia pohlii var. suberecta Müll.

Arg., se selecciona a Saint Hilaire s.n. como el lectotipo. Esta colección está representada por dos duplicados,

ambos en buen estado de conservación y que representan en forma clara los caracteres diagnósticos del

protólogo. En todo caso, las restantes colecciones citadas por Múller (Sello 799, Riedel 614, Langsdorff s.n.),

no fueron localizados.

los: COLOMBIA. A P Narif ional natural Amacayacú, jul 2000 (fl),

p maya & Melendez 173 en cercanias de Leticia, 18 nov an a, K Gentry 12741 (MEXU, MO, US); Leticia, vía a Tarapacá, 28 oct

1991 (£D, Pipoly et al. 15434 (MO, VEN), A la Quintero, 6 sep 2000 (fD, Fonnegra et al. 7233 (HUA); Puerto

Berrio, 14 jul 1917 (ID, Pennell & Pu 35 (NY): Atlántico: región de Manati, 29 mar 1935 (fl), Dugand 743 (F); Barranquilla, ago 1927

(£D, Elias 295 (US). Bolivar: río Mara, ene 1918 (£D, Pennell 3956 (MO, NY, US); Mompós, a dic 1982 (fl, fr), Ramírez 4 (COL);

Palotal, Ayapel, 12 may 1949 (fl, fr), Romero-Castañeda 1635 (COL, MO). Chocó: Riosucio, rio Truandó, entre Riosucio y Calle Larga, 21

ago 1986 (fl), Espina & Mosquera 2097 (COL, CHOCO); río Atrato, cerca de río Sucio, 16 ene 1974 e Cute 9243 (MO [2 cartulinas],

- río Cacarica, 1 jul 1957 (f), Romero-Castañeda 6336 (COL, MO, NY, P). Córdoba: ciénaga de Ayapel, 10 jul 1973 (f), Idrobo & bri

572 (COL, U, WAG); Ayapel, ciénaga de Ayapel, 6 dic 1989 (fl, fr), Marulanda 1626a (HUA). Magdalena: Barro Blanco, 30 n

PA Haught 4752 (US). Meta: cercanías Villavicencio, hacienda Mozambique, 7 mar 1971 (fD, Sastre & Pinto 1020 (COL, MO, ld parque

nacional natural Tinigua, Serranía Chamusa, ene 1995 (fI), Stevenson 1357 (COAH, NY). VENEZUELA. Apure: entre río Orinoco y

Piedra La Villa, 27 ene 1956 ŒD, Wurdack & Monachino 41391 (MO, NY, P, U, US, W, VEN). A : Orinoco, Sacupana, abr 1886

(£D, Rusby & Squires 20 (MO, NY, US, W, WU). basic carona, Matanzas, antes de Puerto Ordaz, 15 may 2001 (£1), Díaz & Aguilera 5172

(VEN); entre San Félix y Puerto Ordaz, ie 1964 ( $ 94620 (NY, US, VEN). Cojedes: San Carlos de Cojedes, 9 may 1946

(fl), Velez 2735 (US, VEN). Delta A Cocoina, 23 jun 1993 uM ee 1358 MIA a entre ee Mulas,

13 oct 1977 (ID, Steyermarh et al. 114584 (MO, NY, VEN); Tucupita, Caño buca , 24 oct

1977 (fD, Steyermark et al. 115321 (MO, VEN). Guarico: Miranda, parque je máciotal me" Guaita, SE de la montaña de Guarda-

humo, dic 1981 (fl), Delascio et al. 11374 (VEN); SO de Calabozo, Hato Masaguaral, 1 sep 1983 (fI), Rondeau 456 (ARIZ, MO, VEN).

Mérida: entre Santa Bárbara y Encontrados, 12 feb 1953 (fl), Bernardi 362 (NY). Monagas: La Pica, cerca de Maturín, jul 1959 (fl),

Aristeguieta 3929 (NY, VEN); río Guarapiche, al SSO de Jusepín, 22 feb 1967 (fl), Pursell 8131 (VEN). Táchira: Caño La Ceiba, NE de

Orope, 15 nov 1979 (fD, eco et al 120423 (MO, VEN). Trujillo: El Cenizo, 28 abr 1951 (ID, Lasser 2833 (VEN); la Ceiba, 3 dic

1922 (fl), Pittier 10887 (NY, US, VEN). Zulia: S de río Catatumbo, 27 mar 1982 ©), Liesner & González 13264 (MO, WAG, VEN); Perijá,

s.d. (fl), Tejera 209 Ait GUYANA: Entre Berbica y Rut , 20 jun 1919 (fL 09 (NY); Mulson creek, Berbice, 10 feb 1977

(1D, Grewal 267 (U). SURINAM: río Coppename, 23 dic 1948 (£5, e 1555 (NY, U); Nickerie, entre Hampton Court

Henar-polder, 8 may 1949 (fl), Lanjouw & Lindeman 3178 (NY, U); Paramaribo, 1851 (fr), Wullschlagel 326 (BR, NY, W [2 ere

GUYANA FRANCESA. Alrededores de Kourou, jun 1969 (fl), Petitbon 121 (P); rio de Kaw, 5 sep 1979 (f1), Prévost 747 (P. ECUADOR

Sucumbios: NO de Añangu, cerca del río Napo, 8 jun 1995 (£D, Clark et al. 1070 (MO, US). PERU. A : Caballococha, 14 ago

1989 (fl, fr), Vásques & Jaramillo 12716 (MO). Loreto: Iquitos, ago 1929 (fl), Killip & Smith 26911 (MO, NY, US); Maynas, rio Ampiyacu,

Pebas, 27 abr 1977 (fI), Plowman et al. 7066 (MO, US, USM, Z), Quebrada de Maquia, Contamina, 16 dic 1964 (fI), Schunke 6676 (MO,

UC, US, USM). Madre de Dios: Laguna Coco Cocha, 3 jun 1986 oe ied et mae 8410 i Tope cerca a la unión de il ríos

Tambopata y La Torre, 26 jul 1985 (fD, "s et al. 51387 (MO [2 cart

Tambopata y río La Torre, 16 oct 1985 (fl), Smith et al. 776 (MO, NY, US). Ucayali: Coronel Portillo, liac Yarinacocha, Cashivo cocha,

25 nov 1997 (fl, fr), Graham & Vigo 381 (MO, NY). BRASIL. Amapá: Rio Jari, Santo Antonio da Cachoeira, 30 jul 1961 (fr), Egler & Irwin

46056 (NY, U, US, VEN); Ariramba, 23 jun 1982 (fl), Rosa & Alfeu 4395 (MG, NY). Amazonas: Lago de Janauacá, S de Río Solimões,

Varzea, 28 ago 1973 (fl, fr), Berg et al. 19785 (INPA, MO, NY, U, US); Igarapé Belém, 24 dic 1948 (fl, fr), Fróes 23781a (IAN, MO); Río

Solimões, cerca de Codajaz, 19 may 1933 (fl), Krukoff 4500 (MO, NY, U, US). Brasilia: Recreio dos bandeirantes, 29 oct 1938 (£D, Pulle

& Lutz s.n. (R, U). Mato Grosso: Cáceres, cerca de rio Paraguai, 8 nov 1996 (fD, Hatschbach et al. 65385 (MBM, WAG); Rio Alegre, Vila

Bela da Santissima Trinidade, barra do rio Guaporé, M nov 1996 (fD), Hastchbach et al. 65647 (INB, MBM, U, WAG); Poconé, Pantanal,

camino a Porto Jofre, 25 oct 1985 (fl, fr), Thomas et a 63 (NY). Minas Gerais: Canandaí, 22 dic 1949 (ID, Duarte 2284 (MO, RB);

Joao Agres, 31 dic 1866 (fl), Glaziou 16248 (P); inn 5 e. 1997 (fl), Neto 2602 (BHCB, INB). Pará: Oriximiná, Rio Trombetas, lago

Iripixi, 12 sep 1980 (fl), Cid et al. 2406 (INPA, MO, NY, US, WAG); Ihla do Marajó, río Mocóes, Anajás, 28 oct 1984 (fl, fr), Sobel et al.

4839 (MO, NY, US); Rio Maicurú, 31 jul 1981 (fr), Strudwick et al. 3936 (MO, NY, US, USF). Paraná: São José dos Pinhaes, Rincão, 22

ene 1950 (fD, Hatschbach 1792 (MBM, US); Contenda, camino do Xisto, 31 dic 1967 UD, ee nd ae NY, P, US); Guarapu-

ava, 14 dic 1965 (fD, Reitz & Klein 17622 (P, US, WAG). Rio Grande do Sul: Corumbá, 7 nov 1980 (tl), Mizoguchi 2420 MO); pene

5 ene 1947 (fl), Rambo 34990 (MO, SD. Rio de Janeiro: Mage, Piedade, 14 ai 1869 (fl), aay 4082 (P); entre e y Morro do

Coco, 8 dic 1964 (fD, Trinta & Fromm 1029-2105 (HB, NY); R Bandeirantes, ene 1935 (fl, fr), Lutz s.n. (R, US). Santa Catarina:

mpo do Massiambú, Palhoça, 19 dic 1953 (f), Reitz 4949 (NY); iiri y Campos Novos, 5 dic 1956 (fl), Smith € pis 8305 (MO,

es US); Chapecó, Fazenda Campo Sao Vicente, 26-28 dic 1956 (ID), Smith & Klein 11588 (US). Sao Paulo: Parajussará, 29 dic 1920 (ED,

Gehrt 4741 (NY); Moji-Guacu, NNO de Padua Sales, sep 1960 (fl), Mattos & Mattosu 8243 (US); Bairro Caete, São Roque, al O de São

Paulo, 2 dic 1987 (fl), Tsugaru & Otsuka 2263 (MO, NY). BOLIVIA. Beni: Marban, San Rafael, Curiche, 6 oct 1979 (11), Beck 2665 (LPB,

MO, P, SI, WAG): Ballivián, río Yacuma, 21 oct 1980 (fl), Beck 5257 (LPB, MO, SD, Moxos, O de Trinidad, 21 oct 1984 (1D, Greg & Schmitt

151 (LPB, MO, NY, U, USF). Cochabamba: Carrasco, Valle de Sajta, 20 mar 1995 (fl), Ritter 1688 (MO). La Paz: Iturralde, Luisita, al O

Morales, Monografia del género Rhabdadenia 557

del rio Beni, 11 sep 1984 (fl, fr), Haase 533 (LPB). Santa Cruz: Nuflo de Chavez, Estancia San eae 24-26 jul 1995 (fl), Fuentes

996 (LPB, USZ); Velasco, El Refugio, 3 may 1995 (fl), Guillén & Choré 3484 (INB, MO); Guarayos, río Negro, 12 jun 1990 (fr), Vargas 595

(LPB, NY, USZ). Paracuay. Alto Paraguay: frente a Valle Mi, Riacho Mosquito, 17 oct 2000 (fI), siad et al. 9696 (SD; Olimpo, nov

1991 (£D, Schmeda 1449 (US); Cucarani, dic 1951 (fD, Schmeda 1590 (US). Alto Paraná: rio Alto Paraná, 1909 (fD, big. 6451 (G, SD;

Estancia río Bonito, 28 feb 1996 cb, icd et al. 44707 (AS, MO). A pa y río Aquidabán, Estrella, dic sin año (£D),

Hassler 7816 (BM, G, NY). C , 16 nov 1874 (fn, Balansa 1355 (G, P). Caazapá: Tavai, ed Bogado,

20 dic 1988 (fL), Meda 2146 (FCQ, G, INB, SD); Tavai, oe Shini, 3 dic 1966 (fl), Ortiz 855 (FCQ, MO), Estancia Tapytá, 13 dic

1999 (fl), Zardini & Brítez 52758 NS "rà winds cerca de Curuguaty, Arroyo Schini, 3 feb 1982 (fl, fr), Fernández-Casas &

Molero 5874 (G, MA, MO, NY); Si i, río Jejui guazú, 1904 (ID, Hassler 5731 pen , NY, P, W); reserva natural Mbaracayú,

14 ene 1998 (fl), Zardini & Vera 47799 (INB, MO). Coat Luque, Tarumandy, ago 1991 (fl), Meas 4062 (FC . Concepcion:

Sargento José E. López, 16 ene 2000 (fl), Zardini & Guerrero 53938 (INB, MO). ais Tucanguá, feb 1898— pue (£D, Hassler 3841

(G, NY, P). Itapua: reserva Yacyreta, 5 dic 2002 (fI), Zardini & Gamarra 59320 (INB, MO). Gran Chaco: Loma bien 1903 (fD, Hassler

2470 (BM, G, NY, P, M Misiones: e olas, 4 dic 2000 ie Zardini & Guerrero 55892 (INB I y, sin fecha (fl)

Page s.n. (US). Presid , 11 nov 1985 (£D, ae 1358 ys MO, PY, SD; Puerto Militar, río Paraguay,

Casuariaga, 9 dic 1989 (fI), Vanni et al. 1226 (CTES, MO); P Colorado-Monte Lindo, 30 jul 1997 (fl), Zardini & Guerrero 47263 (INB,

©). San Pedro: Lima, estancia Carumbé, 6 oct 1967 m Pedersen 8516 (C, P); entre Santa Rosa y Santa Bárbara, 29 oct 1996 (fl),

Zardini & Guerrero 45517 (INB, MO); Antequera, 23 jun 2001 (fD, Zardini & Guerrero 56633 (INB, MO). URUGUAY. Colonia: río de la

Plata, 1853-1856 (£D, Page s.n. (US). ARGENTINA. cornes Santo Tomé, Kira 12 nov 1974 (fI), Schinini & Carnevali 10299

Paso Crucecita, 6 ene 1960 (ID, P US) a Puerto San Ramón, 18 nov 1991(

Fortunato et ài 2429 (BAB, MO, SI). Misi E pció 28 ene 1926 (fl, fr), Clos 1979 ys San Ignacio, ene 198 De Hauman 3591

(MO)

3. Rhabdadenia ragonesei Woodson, Lilloa 5:19. 1940. (Fig. 8). Tiro: ARGENTINA. Santa Fe: General Obligado, entre

Villa Guillermina y Rabon, 14 feb 1938 (fl), Ragonese 3627 (HoLotipPO: MO; isoripos: LIL, MO).

Lianas, tallos glabros. Hojas con el pecíolo 1.5—5.5(-8) mm de largo, láminas 7.1-10.9 cm x 1.8-7.5(-12.5)

mm, lineares a linear-elípticas, el ápice acuminado, la base angostamente cuneada, angostamente obtusa

a angostamente subcordada, glabras, venas secundarias inconspicuas o no evidentes, venas terciarias no

evidentes. Inflorescencias glabras con 1--2(3) flores, pedúnculo 4.1-9.2 cm de largo, pedicelos 4—10(-12) mm

de largo, brácteas 8-11 x 0.4—0.7 mm, lineares; sépalos 11-15 x 1-1.3 mm, lineares a linear-ovados, el ápice

acuminado, subfoliáceos, glabros externamente; corola rosada, la parte inferior del tubo 31-34 x 1.1-1.9

mm, la parte superior obcónica, 26-31 mm, con un diámetro de 1.3-1.6 cm en la boca, el ápice del botón

floral acuminado, lóbulos 2.8-3.5 x 2.4—2.8 cm obovados; anteras 4.5—5 mm de largo, glabras dorsalmente,

excepto por la pubescencia apical, la base inconspicuamente auriculada, con las aurículas agudas; cabeza

estigmática 1.8-2 mm de largo; ovario 1-1.3 mm de largo, glabro, nectarios ca. la mitad de la longitud del

ovario, usualmente libres. Folículos 7.1-8.8 cm x 1-1.5 mm, glabros; semillas 2.2-2.7 cm de largo, glabras,

la coma 1.5-3.4 cm de largo, blanca a blanco-crema.

Distribución, habitat y ecología.—Restringida al S de Brasil (Estado de Mato Grosso do Sul), Paraguay,

y el N de Argentina, donde crece en pantanos, bancos de ríos y sabanas inundadas, en elevaciones bajo los

500 m. Especímenes con flores han sido recolectados entre noviembre y marzo. Frutos se han recolectado

en enero y oe

] fundirse fácil te con la polimórfica R. madida, pero se puede separar

con facilidad por sus llores con la parte inferior del tubo de la corola más largo, de 31-34 mm de longitud

(vs. (6-)8-12 mm), así como por sus hojas usualmente más angostas. Otro carácter útil para reconocer esta

especie se refiere a la longitud de las brácteas florales, ya que aunque todos los taxones del género tienen

brácteas angostas y escariosas, R ragonesei tiene las Si más ips on con un rango de 8-11 mm de largo

(vs. 1-6.5 mm). Aunq I 5n de R. madida, hasta

el momento no se encuentran NOT que reflejen un — nnmis de los caracteres diagnósticos

utilizados para separalas.

Existe una colección de Argentina de la provincia de San Juan (Kermes s.n., BAB), una región relativa-

mente austral para el ámbito de Ancón geográfica conocida de este taxón, donde hasta el momento

no había sido reportada esta especie ie (Ezcurra 1981; Woodson 1936). Ese espécimen puede representar una

colección con una localidad o etiqueta errónea.

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Morales, Monografia del género Rhabdadenia 559

E dos: BRASIL. Mato Grosso do Sul: Bataguacu, río Parana, 21 nov 1992 (fl), Cordeiro et al. 1028

(MO, S SP). PAR AG UAY. Caaguazu: Arroyo Yakare’i, 8 feb 1989 a Zardini & Velásquez 10815 (FCQ, MO). Chaco: Puerto Casado, dic

1916 (fD, Rojas 2095 (SD. Central: Estero del Ypoá, al SO de Nueva Italia, Isla Guazú, 10 feb 1990 (1D, Zardini 6 Velásquez 18744 (SD.

Cordillera: río ee al W de oe y Esteros, 23 dic 1989 (fl), Zardini & Velásquez 17209 (SD. Guairá: Itapé, ene 1901 (ID), Jór-

gensen 4038 (NY, SI, US). Itap Y ta, 21 ene 1997 (fl, fr), Zardini & Villate 46237 (AS, MO), 23 oct 1999 (fI), Zardini & Gamarra

51849 (INB, MO). Misiones: río MM entre Ayolas y San Cosme y Damián, 22 ene 1997 (tI), ina & Guerrero 46399 ipis n

Paraguari: Lago Ypoá, al E de Cerro Taruma, 4 ene 1994 (fI, ipud Zardini & Tilleria 37786 (INB

2 nov 2001 (fD, Zardini & Vera 57163 (INB, MO). San Pedro: Sierra de Maracayú, río Tassen 1904 (fD, Hassler 4174 (NY,P, W):

oe woe Primavera, snes 22 mar 1955 (ID, nd i inm MO, SI, U). ARGENTINA. Chaco: Colonia Benitez, 1931 (fL,

P Schulz 8388 (BAB, CTES, LPB, À Corrientes: San Roque, estancia San

onia Benitez, dic 1952 (£D,

Antonio, camino a Coenai 4 dic 1996 (fl), Arbo et al. 7010 (CTES, CUVC, LPB, MEXU); Mercedes, macrosistema Ibera, estancia

Rincón del Diablo, laguna Yacaré, 30 nov 1998 (fD, Arbo et al. 8050 (CTES, CUVC, HUEFS); Mburucuyá, Estancia Santa Teresa, 9 feb

1951 (st), Pedersen 1004 (BR, C, MO, NY, P, U, US). Formosa: parque nacional Pilcomayo, Puerto de Gendarmeria La Angela, 18 mar

1992 (fl), Fortunato 2811 (MO); Pilcomayo, N de Riacho He He, 6 ene 1980 (fl), Guaglianone et al. 408 (INB, MO, SI). San Juan: Ubivurá,

Bermejo, 29 ene 1907 (fl), Kermes s.n. (BAB).

ESPECIES EXCLUIDAS O DUDOSAS

Echites lucidus Willd. ex Roem. & Schult., Syst. Veg. 4:795. 1819. Rhabdadenia lucida (Willd. ex Roem. & Schult.) Miers,

123. 1878. Tiro: VENEZUELA. Amazonas: río Orinoco, sin fecha (fl), von Humboldt & Bonpland s.n. (HOLOTIPO:

P-H ndn nitida t Müll.Arg.

Echites oras Desv. ex Ham., Prodr. Pl. Ind. Occid. 30—31. 1825. Rhabdadenia barbata (Desv. ex Ham.) Miers, Apocan

S. a a Tiro: B sin datos (fD, Hamilton s.n. en herb. Desvaux (HOLOTIPO: P) = Penta NN luteum (L.) B.E Hansen

se compe Vell., Fl. Flumin. 113. 1829. Amblyanti tris (Vell.) Müll.Arg., Fl. Bras. 6(1):149. 1860. Rhabdadenia

pestris (Vell.) Miers, Apocyn. S. Amer.: 121. 1878. Tiro: >: BRASII Rio DE JANEIRO: Vell., Fl. Flumin. , Icon 3: tab. 43. 1827 (LECTOTIPO,

[2007]) ) K. Schum

Echites ere DC., "dim 8:447. 1844. Rhabdadenia berterii (A. DC.) Mull.Arg., Linnaea 30:435. 1860. Angadenia berterii (A

Miers, Apocyn. S. Amer.: 180. 1878. Tiro: REPUBLICA DOMINICA? ing. 21 (fl), Bertero s.n. (HOLOTIPO: G- DC

F neg 33879]) = Angadenia berterii (A. DC.) M

Echites sagra A. DC., Prodr. 8:450. 1844. ee sagraei (A. DC.) Múll.Arg., Fl. Brit. W 1. 415. Angadenia sagraei (A. DC.)

Mie ii S. mer: 181. 1878. Tiro: CUBA: datos perdidos, 1899 (fl), Ramón. de la Sagra 81 (notorio: G-DC [foto F neg

: P) = Angadenia berterii (A. DC.) Miers.

blades ae Müll.Arg., Linnaea 30:435. 1860. Angadenia + is Miers, Apocyn. S. Amer.: 181. 1878, nom. illeg.

Tiro: "um Ho cu: Sabanas de Saltadero, may 1844 (Il), Linden 1716 (HoLotiro: G-DC; isoripos: BR, P) = Angadenia berterii (A.

Rhabdadenia ia Müll.Arg., Linnaea 30:437. 1860. Angadenia lindeniana (Mull Arg.) Miers, Apocyn. S. Amer.: 180.

UBA. SANTIAGO DE CUBA: cerro Líbano, may 1844 (fl) Linden 1823 (Lecroripo, designado aqui, BR; isolectotipos, G-DC,

P) = Angadenia lindeniana (Müll.Arg.) Miers.

De los dos sinti los por Muller (1860 b) se designa la colección Linden 1823 depositada en el herbario

de Bruselas (BR), Bélgica, como el lectotipo, dado que esta colección posee más duplicados y está en mejor

estado de conservación que el otro sintipo (Linden 1700, P [3 cartulinas]).

Rhabdadenia lindeniana Müll.Arg. var. angustifolia Müll.Arg., Linnaea 30:437. 1860. Tipo: CUBA: datos de localidad es 1843-1844

(fl), Linden 1699 (notorio: W; isoriros: BM, G, NY iu iS MO) = Angadenia lindeniana (Mull. A

Rhabdadenia wrightiana Müll.Arg., Linnaea 30:438. fandevilla wrightiana (Mull.Arg.) Benth., Gen. i 2:727. 1876. Tiro: Cuba.

ORIENTE: entre Paso Ene dune n) 1860 o. Li 399 (notorio: G; isoripos: B [destruido], GH, MA, MO [2 cartulinas],

NY, P US, W [2 cartulinas]) ich.) Urban.

d ie ip Miers, Daa S. Amer. 120. 1878. Tro: CUBA: sin localidad ni fecha (fl), Wright 400 (HoLotiro: BM; Isotipos: G,

cartulinas L.) B.F Hansen € Wun

ies ala Small, Bull. New York Bot. Gard. 3:434. 1905. T F C G y Cutler,

3] oct-4 nov 1903 (fl, fr), Small & Carter 714 (HoLotipo: NY; isorieo: NY) = A l berterii (A. DC.) Miers

Rhabdadenia polyneura Urb., Symb. Antill. 7:337. 1912. Odontadenta polyneura (Urb. ) Woodson, "ma Missouri Bot. Garden 18:546.

1931. Tiro: REPÚBLICA DOMINICANA: Santo DOMINGO: entre Constanza y Río Jimenoa, Loma del Hato quemado, 1400 m, jun

1910 (f), von Tuerkheim 3341 (notorio: NY NEN INB]; isoripos: BR, G, GH, M, U, US, W, WU, Z) = Odontadenia polyneura

(Urb.) Woodson.

|| || £ all n.a H In LI di dian adis il

560 | f Texas 3(2)

APÉNDICE 1

ÍNDICE A NOMBRES EN EL TRATAMIENTO SISTEMÁTICO

Apocynum ehrenbergii (Schltdl.) Müll. ida ex Griseb. (= R. biflora)

cordatum Mill. (= R. biflora) latifolia (Müll.Arg) Malme (= ida

nervosum Mill. (= R. biflora) macrantha Donn. Sm. (ER. bi hd

Dipladenia macrostoma (Benth.) Müll.Arg. (= R. madida)

billbergii Beurl. (= R. biflora) t (Benth.) Müll.Arg. var. glabrata Müll.Arg. (=

Echites R. madida)

biflorus Jaca. (= R. biflora) madida (Vell) Miers

ehrenbergii Schltdl. (= R. biflora) mamorensis Rusby (= R. madida)

macrostomus Benth. (= R. madida) nervosa (Mill) Miers (= R. biflora)

ria Vahl (= R. biflora) paludosa (Vahl) Miers (= R. biflora}

Exothost pohlii Müll.Arg. (= R. madida)

UR (Vahl) G. Don (= R. biflora) pohlii MúlLAro. var. latifolia Müll.Arg. (= R. madida)

Rhabdadenia pohlii Müll.Arg. var. suberecta Müll.Arg. (= R. madida)

biflora ene Müll.Arg. pohlii Müll.Arg. var. volubilis Müll.Arg. (2 R. madida)

cordata (Mill) Miers (= R. biflora) ragonesei Woodson

APÉNDICE 2

ÍNDICE A LOS EJEMPLARES ESTUDIADOS

Rhabdadenia biflora (Jacq.) Müll.Arg.

Rhabdadenia madida (Vell) Miers

Rhabdadenia ragonesei Woodson

Abbott pe a a ); Aoranami 209 (2); Acevedo-Rodríguez 414 (1), 415 (1), 6535 (1); Acosta 2202 (1); Afanador

s.n. (1), s.n. (1); Agostini 2720 (1); Aguilar 5679 (1), 6205 (1); Alain 2535 (1); Allen 885 (1), eee tl , dde 1023 (1); Almeda 285

(1); Ama a 173 5 Arbo 602? (2) 652? ? (3), 7010 (3), 8050 (3, Aranacen (2); Aricta-lacank Arictanmiota 2044 (1), 3929

(2), 4773 (1); Arvigo 365 (1); Asplund 4964 (1); Atha 948 (1); Atwood 4393 (1): Austin 7339 (1 y Ayala 5571 (2).

Baker 412 (1); Balansa 1355 (1); Balcazar 125 (2); Baldwin 4096 (1); Bamps 5107 (1); Benoist 451 (1); Berg 19785 (2); Bernardi

362 me ed Ls (1), ER 8061 (1); Beck 2665 (2), 5161 (2), 5257 (2), 5361 (2), 5914 (2), 6575 (2), 16671 (2), 27985 (2);

Beetle 26219 (1 s.n. (1), 2427 (1); Bernal 133 (1); Berti 699 (1); Billiet 4296 0A i di (1), 5136 (1); Pace n (2), pels »

ae ps (3) Baldo s.n. (1); Bonpland 1354 (1); Boom 8028 (1); Bordenave 102 (1)

Britton 582 (1), 23070 (1); Broadway 365 (1), 428 (1); Bruijn De 1301 (1); Brumbach 9072 (1); na 1358 (2); Bunting 1278

(1); Burchell 10057 (1); Burger 12298 (1); Burkart 30041 (3), 31025 (3); Byron, 158 Q2).

Cabrera 117 (1), 4013 (1), 6762 (1); Callejas 5030 (1); Cano 793 (3); Cárdenas 24 (2), 6191 (2); Cardona 593 (1); p 745 (2);

Castillo 2720 (1); Castro 99 (2); Cavaler iE 1810 (2); Cedillo 3371 (1); Chapman s.n. (1), 165 (1); Chavarría 223 (1); Chessman

389 (1); Cid 1021 (2), 2406 (2); Clark 1070 (2); Clemente 6201 (1); Clos 1979 (2); Colonello s.n. (1), 1358 (2), 1375 (2), bed (2),

2005 (2); Combs 78 (1); Contreras 9471 (1); Cordeiro 1028 (3); Core 1570 (1); Cornejo 1890 (1); Correll 41941 (1); Craighead s.n.

(1); Cremmers 9523 (1); Croat 248 (1), 267 (1), 6574 (1), 8296 (1), 9994 (1), 11067 (1), 36871 (1), 36895 (1), 76261 (1); Cruz 945

(1), 1417 (1), 3274 (1); Cuadros 2239 (1), Cuatrecasas 20002 (1), 24295 (1); Curran 789 (1), 1341 (1), 2266 (1).

Darcy 2020") I), om ), n ); Davidse 17838 (1), 32588 (1), Aaaa ); Lavia d (1); Degener 16128 UJ Derascio naa

D pic 28252); 2251 d Den Outer 930 (1); D 926 (1); Di

(1); Dionizio 3360 (2); Dobbeler 5218 (1); Dod 727 (1); Dodson 2984 (2), 14569 (1); Don 25 (1); D'Orbygny s.n. 3); Dorr 9100

(1); Duarte 569 (2), 2284 (2); Duchafraing s.n. a ate 1565 (2); Dugand 743 (2); Duke 5759 (1), 6636 (1), 8477 (1), 10995 (1);

Dusén 3005 (2), 7781 (2), 14025 (2); Duss 7 (1), 1867 (1), 2840 (1); Dwyer 11 (1), 1013 (1), 14999 (1).

Eggers 836 (1), 1155 (1), 2676 (1), 6510 (1); Egler 46056 (2); Eiten 1649 (2); Ekman 8053 (1); Elias 295 (2),

(1), 1291 (1), 1697 (1); Encarnación Mejía, (2); Espina 2097 (2), 2453 (1); Eyerdam 321 (1), 559 (1).

Feddema 2026 (1); Fendler 257 (1), 619 (1); Fernández 357 (1); Fernández-Casas 5874 (2), 10638 (1); Feuillet 892 (1), 1586 (1);

Fiebrig 6451 (2); Florschút 914 (1), 1677 (1); Fonnegra 7233 (2); Fortunato 2429 (2), 2811 (3); Fredholm 3071 (1), 5573 (1); Frey

760 (2); Fróes 1886 (1), 23781a (2); Fryxell 2821 (1); Fuentes 220 (1), 996 (2), 1386 (2), 1589 (2), 2028 (2), 2464 (2); Fuertes 220

(1); Funk 8410 (2).

Garner s.n. (1); Gardner 6060 (1); Gaudichaud 85 (2); Gaumer 653 (1), 23335 (1); Gehrt 4741 (2); Gentry 1524 (1), 28373 (1),

9243 (2), 12741 (2), 18296 (2), 28373 (1), 32134 (2), 40726 (1), 51015 (1); 51387 (2); Gentle 826 (1), 3921 (1); Geyskes s.n. (1);

Gillespie 1196 (1); Ginzberger 350 (2); Glaziou s.n. (2), 4082 (2), 4086 (2), 5941 (2), 16348 (2); Godoy s.n. (1); Gómez 20567 (1);

Gémez-L. 397 (1)3862 (1), 5539 (1), 10186 (1); Goodal 263 (3); Graham 381 (2); Grainville 5153 (1), 5760 (1), 6850 (1); Grayum

371 (1), 517 (1), 585

Morales, Monografia del género Rhabdadenia 561

AMO fD\. 67 Ezx FM Fina

^ fe ^ ra 2333

3658 (1), 4042 (1); Greg 151 (2); Greenman 5986 (1); Grewal 267 (2), 296 (1); G

(2), 2853 (2), 2914 (2), 2937 (2), 3214 (2), 3484 (2); Gutiérrez 1643 (2), 1848 Qy

Haase 533 (2); Hahn 123 (1), 3525 (1), 3808 (1); Hammel 4572 (1), 5038 (1), 21886 (1); Hammit 189 (1), 193 (1); Hampshire 948

(1); Hansen 7931 (1); Harvey 3436 (2); Hassler 2470 (2), 3841 (2), 4174 (3), 5731 (2); Hatschbach 1792 (2), 6641 (2), 15449 (2),

end (2), 202030 ) d (2), 28217 (2), 65385 (2), 65647 (2); Haught 4752 (2), 4841 (1), 5416 (1); Hauman 3591 (2); Hayes 91

427 (1); Herrera 277 (1), 559 (1); Heyde 468 (1); Hicken s.n. (3), 1619 (2); Hill 27161 (1); Hitchcock

; n. (1), s.n. (1), s n. (1), 202 0, 16713 (1); Hoffman 750 (1); Holdrigde 1222 (1); Hooker s.n. (1); Hostmann 494a (2), 831 (2);

sete 4941 "d Hunter 70a (1); Huston 507 (1).

Ibañez 2141 (1); Idrobo ZI») (2); | x 9.0L (1); Isabelle, ^ (2).

Jack 5242 (1); Jaeger s.n. (1); Jahn 1217 (1), 1230 (1); Janzen 1177 (1); Jaramillo 15203 (1); Jardim 3725 (2); Jenman 6808 (2);

Jiménez 1235 (1), 1429 (2), 1959 (1); Johnson 1153 (1), 1290 (1); Jonker-Verhoef 89 (1); Jordán 3 (2); Jórgensen 3450 (2), 4038

(3); Juncosa 2522 (1); Junk 59 (2)

Kermes, s.n. (3); Kiesling 9696 (2); Killen 6892 (2); Killip 533 (1), 26911 (2), 30464 (1), 33010 (1), 38622 (1), 39078 (1), 42500 (1),

42583 (1), 42954 (1), 43152 (1), 44215 (1), 44280 (1), 44509 (1); Kings 199 (1), 348 (1); Klug 762 (2), 1290 (2); Kramer 3130 (1)

Krapovickas 13757 (3), 20246 (3); Krukoff 4500 (2), 5915 (2), 5931 (2).

Labat 89 (1); Lane s.n. (1); Lanjouw 1053 (1), 1070 (1), 1082 (2 ), er Y indi (2), ea ip ), 3178 (2), 3430 i o

2833 (2), 3863 (1); Ledru 16 (1); Leewenberg 11600 (1), 11851 ( (1); Lehmann 1000 (1); León 268

(2), 644 (2); Leonard 4176 (1), 4207 (1), 13916 (1); Lépiz 399 (1 ; oe 72 (1); Lewis 87 (1), 1862 (1); 37625 on lens

1923 (1), 13264 (2); Lindeman 5728 (2), 15597 (2); Linden 57 (1); Liogier 14529 (1); López 5798 (2); Lundell 62 (1), 1813 (1),

4055 (1); Lut (2), 685 (2

Maas 5548 (1); MacDougal 3435 (1); Madrigal 70 (1), 661 (1); Magafia 309 (1); Maguire 23596 (1); Martin-Ballesteros, 151 (1),

624 (1); Marulanda 1626 (2), 1627 (2); Mathews 171 (1); Mattos 8243 (2); Maxon 321 (1), ips Melanie mid (1); Meijer 332

(1); Mejía 406 97

—

amm.

1); Meleady 219 (1); Melinon 293 (2); Mell s.n. (1); Mello 3494 (2); Mendoz (2) (1); Mennega

166 (1); Mereles 2146 (2), 2265 (2), 3452 (2), 4062 (2); Miller s.n. (1); Millspaugh 974 (1); T 2420 (2), 2426 (2); Molas

1098 (2); Moldenke 725 (1); Molina 2048 (1), 5208 (1), 6951 (1), 20648 (1); Moore 1007 (2); Moraes 1138 (2); Morales 3010 (1),

6558 (1), 7004 (1), 14599 (1); Moreno 13199 (1); Moreno 11989 (1), 12312 (1); Moretti 186 (1); Mori 20317 (2), 21218 (2); Morton

10002 (1); Mouret 229 (1); Nee 40175 (2); Nee 41657 (2).

Neill 4563 (1); Nelson 968 (1), 1154 (1), 4245 (1), 7854 (1), 8503 (1), 8741 (1), 8895 (1), 11118 (1), 15500 (1); Neto 2602 (2); Nevers

de 4880 (1), 6441 (1), 6590 (1); Novelo 2976 (1), 3105 (1), 3618 (1).

Oldeman 32 (1), 796 (1), 1064 (1), 1396 (1); Orell 858 (2); Ortiz 855 (2); Otero 157 (1).

age s.n. (1), s.n. (1); Pamer 2o (1), FI a), hans Paul 1511 Pedersen 1004 (3), 5328 (2), 5375 (3), 8516 (2), 10839 (3); Pennell

35 (2), 3738 (2), 3956 (2); Persaud 212 (1); Peterson 6355 (1); Peterson 66 6667 (1),6770 (1), 7129 (1); Petitbon 93 (1), 121 (2); Philcox

8045 (1); Pipoly 9038 (1 », 15434 (2); Pires 757 01), 1619 (1), 4801 © >, 6533 (2 ); Pittier 379 (1), 2629 (1), 4106 (1), 4359 (1), 6352 (1),

9903 (1), 10480 (1), 10887 (2), 14034 (1); Plée sn, (1), s.n. (1), 17 (1); Plowman 7066 (2); Poeppig 2946 (1); Pohl 5170 (2), 11574

(1); Poiret s.n. (1); Poiteau s.n. (1); Pohl s.n. (2); Pall s.n. (1); n 1024 (1); Prance 2542 (2), 8032 (213714 (2), 14780 (2), 16747

(2); Prévost 702 (1), 747 (2), 3288 (1), 4103 (1); Proctor 3312 (1), 24219 (1), 35116 (1); Pulle s.n. (2); Pursell 8131 (2).

Quesada 685 (1); Questel s.n. (1), 4188 (1).

Rambo 34990 (2); Ramirez 4 (2); Ramos 113 (2); Ramsammy 2 (1); Raven 21514 (1); Raynal-Roques 19814 (1); Realpe, 170 (1);

Reijenga 623 (2); Reitz 4949 (2), 14545 (2), 17622 (2); Renteria 4285 (1); Revilla 2043 (2); Reyes s.n. (1); Rimachi 6526 (2); Ritter

s.n. (1), 1688 (2), 3763 (2), 4495 (2), 4631 (2); Robertson 254 (1); Rodríguez s.n. (2); 3118 (1), 3198 (1), 5584 (1); Rohr von 21 (1);

Rojas 2095 (3); Roldán 516 (1); Romero 1574 (3); Romero-Castañeda 65 (1), 1635 (2), 1946 (1), 1925 (1), 6336 (2); Rondeau 456

(2); Rosa 4395 (2); Rudas 4263 (2); Rugel 114 (1), 694 (1); Rusby 20 (2); Rutten 204 (1).

Sachet 449 (1); Sáenz 5a (1); Sag (1), s.n. (1), s.n. (1); Saint Hilaire 328 (2), 400 (2), 491 (2); Salas 1333 (1), 2761 (1); Saldías 572

(2), 2916 (2); Sandino 2199 (1), 2225 (1); Santa 343 (1); Santos 491 (2); Sargent 23 (1); Sastre 1020 (2), 4162 (1); Saunders 619a (1),

854 (1 » il 10299 (2); Schipp 148 (1); Schmalzel 316 (1), 1789 (1); Schmeda 1499 (2), 1590 (2); Schomburgk 329 (2); Schultes

7158 (2); Schulz 238 (3), God 9, 10525 (1); Lbs ), 960 9, mo 6676 (2); uh 213 (1); Seibert 1719 (1); Silva 168

(1), 2141 (2); Simmuels 448 Si 252 (1 is 935 (1 ll (1), 4591 (1); Smith 776 (2), aoe ), 1080 (1),

1664 (1), 8305 (2), 9465 (2), 1 1077 (2), 11588 (2), 12881 (1); Sobel 4839 2); ee 100 (1); Soria 1820 (2); S (isn

(1), 485 (1), 748 (1), 822 (1), 1074 (1), 3393 (1); Sousa 10867 (1); Spegazzini s.n. (2); Spellman 1443 (1), 2432 (1); Splitgerger s.n.

(2); Spruce s.n, (1), 516 (2); Standley 373 (1), 20059 (1), 24827 (1), 25141 (1), 28748 (1), 31133 (1), 72863 (1); Steere 1910 (1), 1937

(1); Stehlé 4562 (1); Stergios ie (1); Sterringa 12418 (1); Stevens 1044 (1), 7870 (1), 10465 (1), 17765 (1), 19619 (1), 20043 (1),

20665 (1); Stevenson 433 (1), 1357 (2); Stevermark 87705 (1); 94620 (2), 107867 (1), 108393 (1), 110529 (1), 110893 (1), 114584 (2),

114842 (2), 114982 (1), 115321 d 120423 (2); Stier 4 (1),61 (1); Stoddart 7014 (1), 7039 (1); Strudwick 3349 (2), 3936 (2), 4131 (2).

562 J ti tanical h Insti Texas 3(2)

Tamayo 4730 (1); Taylor 1055 (1); Tejera 209 (2); Téllez 2050 (1), 2238 (1); Tenorio 14941 (1); Tessmann 3547 (2), 3668 (2); Teunis-

sen 13201 (2), 15105 (2); Thomas 4563 (2); Thoms 2 (1); Thorne 14935 (1); Toledo 284 (2), 578 (2); Tonduz 7081 (1); Townsend

39 (2); Triana 3409-11 (1); Trinta 1029-2105 (2); Tsugaru 2263 (2); Tun 126 (1); Tyson 4670 (1), 6806 (1).

Utley 39 (1).

Vanni 1226 (2); Vargas 595 (2), 6675 (2); Vreden 14075 (1).

Wacket s.n. (2): Walter 1107 (1); Webster 10410 (1); Wedel 496 (1), 591 (1), 1546 (1), 1978 (1); Weissenhofer 35 (1); Went, 334

(1), 573 (1); Wessels 507 (1); White 39 (1), 1522 (2); Wilbur 23973 (1), 32827 (1); Wiley 143 (1); Williams 15539 (1); Willemse 1 (1);

Wilson 244 (1), 264 (1); Wood 13216 (2), 17363 (2); Woodroffe 602 (1); Woodson 764 (1), 933 (1), 1682 (1); Woolston 483 (3);

Worthing, 21223 (1); Wright 2594 (1); Wullschlágel 326 (2), 342 (1); Wurdack 323 (1), 41391 (2); Wydler s.n. (1)

Vanni 1226 (2), 4133 (3); Vargas 595 (2); Vasquez 923 (2), 4373 (2), 12716 (2); Vélez 2735 (2); Vincelli 652 (1), 668 (1), 670 (1),

670a (1); Vreden 14075 (1).

Yepes 27 (1); Yuncker 17961 (1).

Zainum 1680080 (2); Zardini 10815 (3), 17209 (3), 18744 (3), 37523 (2), 37786 (3), 41979 (2), 42854 (2); 44707 (2), 45517 (2),

46237 (3), 46399 (3), 47263 (2), 47281 (2), 47799 (2), 48138 (2), 51849 (3), 52758 (2), 53938 (2), 54670 (2), 55634 (3), 55892 (2),

56502 (2), 56633 (2), 56637 (2), 56910 (2), 57613 (3); 59320 (2), 59381 (2), 59888 (2). Zuloaga 1485 (2), 3128 (3), 3145 (2).

Pam

APÉNDICE 3

INDICE DE NOMBRES COMUNES

bejuco lechero (A. biflora, Venezuela, Monagas) ne de mar (f. iae Colombia Nariño, Tumaco)

cai seca (A. biflora, Brasil, Para, Marajo) , Cayenne, Mahury)

cipó de caraugrejo (R. biflora, Brasil, Maranhao) mangrove ' vine 2 pias Belice, ciudad de Belice)

dub inanusu (R. biflora, Panamá, San Blas, nombre kuna) palo de leche (R. biflora, Republica Dominicana)

huevo de gato (R. madida, Perü, Loreto, Bella Vista)

AGRADECIMIENTOS

Quiero agradecer a los herbarios A, AAU, ALCB, AS, ASE, B, BAB, BEREA, BHCB, BIGU, BM, BOLV, BR, C,

CAS, CAY, CEN, CEPEC, CGE, CHAPA, CICY, CIIDIR, CIMI, CM, COAH, COL, CR, CVRD, CUVC, CUZ,

DPU, DS, DUKE, E, EAP, ECON, ENCB, ESA, ESAL, F, FCQ, FDG, FI, FI-W, FUEL, G, G-BOIS, G-DC, GB,

GFJP, GH, HAL, HB, HBG, HERZU, HOXA, HRB, HRCB, HST, HUA, HUEFS, HUQ, IAN, IBGE, INPA, IPA,

JAUM, JEPS, JPB, K, LAGU, LD, LE, LIL, LP, LPB, LZ, M, MA, MBM, MBML, MEDEL, MEXU, MG, MHES,

MICH, MO, MOL, NY, O, P, P-BA, P-HB, P-JU, P-LA, PEL, PEUFR, PMA, PORT, PR, Q, QAME, QAP, QCA,

QCNE, QPLS, R, RB, S, SI, SP, SPF, SPFR, TEFH, TRIN, TULY, U, UB, UC, UCOB, UDCB, UFMT, UFP, UPS,

US, USF, USJ, USM, USZ, UVAL, W, WU, WAG, WIS, VALLE, VALLE, VEN, VIC, XAL, Z, y ZT por proveer

material en préstamo o por facilitar el acceso a sus colecciones, así como a los herbarios BHCB, ESA, LAGU,

MO, UVAL por el numeroso envío de material como regalo por identificación. Se agradece la colaboración

de Michael Grayum (MO), quien suministró referencias bibliográficas claves y colaboró en la tipificación

de algunos taxones, así como a Barry Hammel (MO), quien revisó By oe el resumen en inglés. Se desea

reconocer de manera muy especial las atenci y | las por Paul Maas e Hiljte Maas (U),

que Bn posible Bev visita vde varios AEDA en Holanda. Otras personas a las que quiero expresar mi

gratitud por brindar f ilit la visita de varios herbarios en Centro y Sur América

son Alessandro Rapini (HUEFS), Aledo Fuentes (USZ), Alvaro Cogollo (JAUM), Ana Luc McVean (UVAL),

Andre Amorin (CEPEC), Andre Fontana (MBML), Asunción Cano (USM), Cirilo Nelson (TEFH), Eunice

Echeverria (MHES), Frank Sullyvan Cardoza (EAP), Gabriela Jolochin (MVFA), Homero Vargas (QCNE),

Joaquina Alban (USM), Jorge Monterrosa (LAGU), Julio Betancur (COL), Marccus Alves (UFP), Mario Véliz

(BIGU), Mauricio Bonifacino (MVFA), Norma Degninani (SD, Phillip Silverstone-Soukup (CUVC), Reneé

Fortunato (BAA), Ricardo Callejas (HUA), Rocio Rojas (HOXA), Rodolfo Vasquez (MO) y Stephan Beck

(LPB). Finalmente, quiero agradecer de manera muy especial al personal de los herbarios de la Universidad

de Antioquia (HUA) y del Jardín Botánico Javier Antonio Uribe (JAUM), en Medellín, Colombia, por las

facilidades brindadas y el excelente trato humano brindado en mis continuas visitas a esa ciudad. Mary E.

Endress y un revisor anónimo contribuyeron con la revisión del manuscrito.

Morales, Monografia del género Rhabdadenia 563

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ENDRESS, M., S. LiEpE-SCHUMANN AND U. Meve. 2007. Advances in Apocynaceae: The enlightenment, an introduction.

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50:214-233

MoRALES, J.F. 1999. A synopsis of the genus Odontadenia (Apocynaceae), Series of revisions of Apocynaceae XLV.

Bull. Jard. Bot. Nat. Belg. Bus ame

MoRALES, J.F. 2004. Estudios Neotropi en Prestonia (Apocyn

aceae, Apocynoideae) para Colombia y Ecuador, con tari bre el grado de lobulación del nectario.

Candollea 59:159-165,

Monats, J.F. 2005. Estudios en las Apocynaceae neotropicales XIX: La familia Apocynaceae (Rauvolfioideae,

Apocynoideae) en Costa Rica. Darwiniana is 20 91.

Moarates, J.F. 2006. Estudios en las Apocynace icales XXIV: Bahiella (Apocynoideae: Echiteae) un desa-

percibido género endémico de Bahia, Brasil Sida 22: 333-353

Mora es, J.F. 2007. Estudios en las Apocynaceae Neotropicales XXXI: el complejo de Mandevilla hirsuta y cuatro

nuevas especies. J. Bot. Res. Inst. Texas 1:859-869.

MORALES, J.F. AND A. FUENTES. 2004. Estudios en las Apocynaceae Neot ll: nuevas especies de Mandevilla

(Apocynoideae, Mesechiteae) para Perú y Bolivia, con notas sobre: la moralo floral en corolas infundibu-

liformas. Candollea 59:167-174.

MULLER Arcoviensis, J. 1860a. Apocynaceae. En: C.EP. von Martius (ed.), Flora Brasiliensis 6(1):1-180

MOLLER Arcoviensis, J. 1860b. Species novae nonnullae americanae ex ordine Apocynearum et observationes

quaedam in species generis Echites auctorum earumque distributio in genera emendata et nova. Linnaea

30:387-454

PicHon, M. 1948 a. Classification des Apocynacées. |. Carissées et Ambelaniées. Mém. Mus. Natl. Hist. Nat., Sér.

B., Bot. 24:111-181.

PicHon, M. 1948 b. Classification des Apocynacées. V. Cerbéroidées. Not. Syst. Paris 13:212-229.

564

Pichon, M. 1948 c. Classification des Apocynacées. XIX. Le rétinacle des Echitoidées. Bull. Soc. Bot. France

95:211-216.

Pichon, M. 1949. Classification des Apocynacées. IX. Rauvolfiées, Alstoniées, Allamandées et Ta bernaémontanoi-

dées. Mém. Mus. Natl. Hist. Nat. 27:153-251.

Pichon, M. 1950a. Classification des Apocynacées. XXV. Echitoidées. Mém. Mus. Natl. Hist. Nat., Sér. B., Bot.

1:1-143.

Pichon, M. 1950b. Classification des Apocynacées. XXVIII. Supplément aux Plumérioidées. Mém. Mus. Natl. Hist.

Nat., Sér. B., Bot. 1:145-173.

Simoes, A.Q. AND L.S. KinosHita. 2002. The Apocynaceae s. str. of the Carrancas region, Minas Gerais, Brazil. Darwini-

ana 40:127-169,

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Simoes, A.O., M. ENDRESS, T. VAN DER Niet, L.S. KINOSHITA, AND E. Conti. 2006. Is Mandevilla (Apocynaceae, Mesechiteae)

monophyletic? Evidence from five plastid DNA loci and morphology. Ann. Missouri Bot. Gard. 93:565-591.

Simoes, A. O, T. LivsHuttz, E. Conti, AND M.E. Enpress. 2007. Phylogeny and systematics of the Rauvolfioideae (Apoc-

ynaceae) based on molecular and morphological evidence. Ann. Missouri Bot. Gard. 94:268-297.

WiLLIAMs, J. 2004. Polyphyly of the genus Echites (Apocynaceae: Apocynoideae: Echiteae): evidence based on a

morphological cladistic analysis. Sida 21:117—131

Woobson, R. 1935 a. Observations on the inflorescence of Apocynaceae (with special reference to the American

genera of Echitoideae). Ann. Missouri Bot. Gard. 22:1-48.

Woobson, R. 1935 b. Studies in the Apocynaceae. IV. The American genera of Echitoideae XXVI. Ann. Missouri

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Woopson, R. 1936. Studies in the Apocynaceae. IV. The American genera of Echitoideae. Ann. Missouri Bot. Gard.

23:169-438.

Woopson, R. 1940. Una nueva especie santafecina de Rhabdadenia. Lilloa 5:199-200.

ESTUDIOS EN LAS APOCYNACEAE NEOTROPICALES XXXVIII:

TRES NUEVAS ESPECIES DE MANDEVILLA (APOCYNOIDEAE: MESECHITEAE)

PARA COLOMBIA Y VENEZUELA

J. Francisco —

i E IE Be q AT » 1H Df: je i FIN IP? pto o 22--3100

Santo Domingo de Heredia, o RICA

fmorales@inbio.ac.cr

RESUMEN

Se describen e ilustran Mandevilla albifolia, M. caquetana y M. sanctae-martae, tres nuevas especies de Venezuela y Colombia y

se discuten sus relaciones con especies afines.

PALABRAS CLAVE: Apocynaceae, Apocynoideae, Mesechiteae, Mandevilla, Colombia, Venezuela

ABSTRACT

Mandevilla albifolia, M caquetana, and M t tae, tł i ] ; V, 1 | hi 4 e RN

ill " tar A laws Jar Ll: "41 1 ] : J: 3

ds I

Trabajando en la db dd J las Lu ir pata el Catálogo de las Plantas de Colombia y Venezuela,

se describen tr as especies, dos de ell Colombia y la ultima conocida solo del estado

del Táchira en Venezuela. En las descripciones, las panies de la corola siguen a Morales (2005).

Mandevilla albifolia J.F Morales, sp. nov. (Fig. 1). Tiro: VENEZUELA. Tacuira: Uribante, Siberia, camino a Pregonero, 20

nov 1985 (fl, fr), van der Werff & Ortega 8086 ( : INB, isoripros: MO, WAG

AAA J :11 1 Af 311 Hu E 1 C “Ts Pote 172 - 1.7%: r 1: ` 1 1 LEN x

t J.F.M M cui similis

L > 2 i e

Liana; ramitas aplanadas cuando jóvenes, cilíndricas a subcilíndricas con la edad, usualmente sólidas,

diminuta y densa a moderadamente papilado-puberulentas, el indumento más esparcido con la edad; colé-

teres interpeciolares inconspicuos, hasta 0.8 mm de largo. Hojas opuestas; pecíolos 2. c 5 mm de largo;

láminas foliares 4.5—9.5 x 0.4—0. im Be cm, amid a near elipaeas, el ápi acuminado, la

base cuneada a obtusa, los colé id lo largo del nervio central, membranáceas,

glabras en la de adaxial, densamente glauco-tomentosas abaxialmente, el indumento no visible a

simple vista, el margen no revoluto, las venas secundarias apenas visibles abaxialmente, inconspicuas en

la superficie adaxial, venas terciarias no impresas. Inflorescencia más corta o igualando las hojas subya-

centes, axilar, esparcidamente puberulenta a glabrescente, el indumento restringido mayormente al eje,

con muchas flores, pedúnculo 4-17 mm de largo, pedicelos 3-4.2 mm de largo, brácteas 13 x 0.7-1.3

mm, linear-ovadas, escariosas; sépalos 1.1-1.4 x 1-1.2 mm, similares en longitud, angostamente ovados,

acuminados apicalmente, el ápice no reflexo, escariosos, glabros o glabrescentes externamente, el coléter

solitario, irregularmente laciniado apicalmente; corola infundibuliforme, amarilla o anaranjado-amarilla,

glabra externamente, el tubo basalmente giboso, la parte inferior 6.5-7.6 x 1.5-2.1 mm, la parte superior

13-14 mm de largo, campanulada, 9-10 mm de diámetro en el orificio, el apie a poles floral obtuso a

redondeado; lóbulos 13-15 x 9-10 mm, obovados, aj extendidos; la base

de la parie M del tubo, anteras 3.5—4 mm de largo, glabras doresirmetire la base auniculaaa, con las

eza estigmática 1.4—1.7 mm de largo; ovario 1-1.3 mm de largo, glabro; neci anular,

modom TEN ca. la mitad de la longitud del ovario. Foliculos 15.3-20.8 cm x 1-3 mm, no

fusionados longitudinalmente, algunas veces solo unidos en el ápice, glabros, moniliformes; semillas 8—9

mm de largo, glabras a glabrescentes, la coma 1.3—1.9 cm de largo, canela-amarilla.

J. Bot. Res. Inst. Texas 3(2): 565 — 571. 2009

566 | t tanical i Texas 3(2)

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Fic. 1 LY TEMS Vul UCI y , INB; l J WAG) A Ramita con flores B f

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J F ; T 4

da | | ial te abi t I tubo ail lab F Antera. vista dorsal. G. Cab tiamática. H

pod r pr]

Nectario y ovario. I. Foliculos. J. Semilla.

Morales, N peci le Mandevill | Colombi y Venezuela 567

Distribución, hábitat y ecología.—Endémica al estado de Táchira, Venezuela, donde crece en vegetación

secundaria y vegetación asociada a sabanas, en elevaciones de 600—1300 m. Flores se producen de mayo a

agosto y noviembre. Frutos se han ccna en ne julio y noviembre.

evilla albifolia se n M. caquetana, M. obtusifolia y M. pachyphylla Woodson,

un grupo de taxones que se reconocen por sus inflorescencias con las brácteas florales escariosas y laminas

foliares con la venación secundaria levemente impresa y las venas terciarias no impresas. Sin embargo, M.

albifolia se puede diferenciar de estas especies por sus láminas foliares lineares a linear-elípticas, con la

superficie abaxial glauco-tomentosa.

Especímenes examinados: VENEZUELA. Táchira: carretera Fundación-Siberia, sector La Idea, 17 Ago 1988 on Badillo et al. 7912 (F,

MY); Uribante, camino a Presa La Honda, 24 Jun 1990 (fl, fr), Dorr & Barnett 7177 (VEN); E de La Fundación, represa Dorada, 1 May

1981 (ID, Liesner et al. 11649 (INB, MO, VEN); La Fundación, camino a Pregonón, 13 nov 1980 (fl, fr), Maas & Pais 5273 (WAG); Cerro

Las Minas, SE de Santa Ana, 10 nov 1979 (fl), Steyermark et al. 119835 (MO, VEN); Uribante, cerca de La Fundacion, 6 Jul 1983 (fl, fr),

van der Werff 4902 (MO, VEN).

Mandevilla caquetana J.F Morales, sp. nov. (Fig. ud Tiro: COLOMBIA. Caquetá: Solano, sabanas del Yari, Chorro de la

Gamitana, 7—14 abr 1997 (ID, Arbeláez et al. 832 (HOLOTIPO: COAH, HUA).

A Mandevill ifolia M h. cui similis, petiolis 1-2.5 mm longis (vs. 4—5 mm), pedicelis 1-1.5 mm longis (vs. 3 mm), corollae tubo

20—23 mm longis d 45 mm) et faucibus 5—6 mm diametro (vs. 15 mm) differt.

Liana; ramitas PDOs: a Super ignicas, solidas, no bod: pense a mogeragamente puberulentas a

papilado-pul de largo. H tas; pecío-

los 1-2.5 mm de lares: láminas foliares 1.54 x 1-2. 9. Gi, elites: el ápice sedondestio- nado, más

raramente emarginado-mucronado, la base cordada, los coléteres distribuidos irregularmente a lo largo del

nervio central, membranáceas, glabras, levemente revolutas marginalmente, venación secundaria impresa

en ambas superficies, las venas terciarias no evidentes. Inflorescencia igualando o ligeramente más larga que

las hojas subyacentes, axilar, inconspicua y esparcidamente puberulenta a glabrescente, con muchas flores,

pedúnculo 18-21 mm de largo, pedicelos 1-1.5 mm de largo, brácteas 1.5—2 x 0.5-1 mm, angostamente

ovadas a linear-ovadas, escariosas; sépalos 1-1.5 x 1-1.2 mm, angostamente ovados, similares en longitud,

cortamente acuminados apicalmente, el ápice no reflexo, escariosos, glabros a glabrescentes externamente,

el coléter solitario, subentero a diminuta e irregularmente lacerado apicalmente; corola infundibuliforme,

amarillo-verdosa, glabra externamente, el tubo giboso basalmente, la parte inferior 12-13 x 1.4—2.1 mm, la

parte superior 8—10 mm de largo, cónica, con un diámetro de 5-6 mm en el orificio, el ápice del botón floral

obtuso-apiculado o redondeado-mucronulado; lóbulos 5-6.5 x 4—5.4 mm, obovados, extendidos; estambres

insertos en la base de la parte superior del tubo, anteras 2.9—3.2 mm de largo, glabras dorsalmente, la base

auriculada, con las aurículas obtusas a redondeadas; cabeza estigmática 1-1.3 mm de largo; ovario 1-1.3

mm de largo, glabro; nectario anular, irregularmente lobulado, ca. la mitad de la longitud total del ovario.

Folículos 8.3-10 cm x 1-4 mm, no fusionados longitudinalmente, algunas veces solo unidos en su región

apical glabros, moniliformes; semillas desconocidas.

Distribución, hábitat y ecología. —Endémica al Departamento de Caquetá, en la Amazonia Colombiana,

donde crece en vegetación asociada a afloramientos rocosos de cuarzita del escudo Guyanes (sierra de

Chiribiquete), en elevaciones de 350-750 m. Especímenes con flores han sido recolectados en abril y de

noviembre a diciembre. Material con on se conoce en a y de noviembre a diciembre.

andevilla l subgénero 1 Woodson, 1933) y ha sido confundida

con M. obtusifolia, taxón endémico al Estado de Amazonas, Venezuela y conocido hasta el momento solo por

la colección tipo, la cual se perdió en 1978 durante un envío de material del New York Botanical Garden a

Venezuela. Sin embargo, basado en la extensa descripción original y en las ilustraciones que la acompañan

(Monachino, ina in d on ade se puede ea con facilidad de esa especie por la siguiente combi-

res 1 n los pecíolos 1-2.5 mm de largo (vs. 4-5 mm), láminas

foliares con la base e ea (vs. led y subcordada), flores cortamente pediceladas,

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Nectario y ovario. I. Foliculos.

Morales, N ies de Mandevill Colombia y Venezuela 369

con los pedicelos de 1-1.5 mm de largo (vs. 3 mm), sépalos 1-1.5 mm de longitud (vs. 2.5 mm), corolas con

la parte inferior del tubo de 12-13 mm de longitud (vs. 30 mm) y la parte superior de 8-10 mm de largo

(vs. 25 mm), con un diámetro de 5-6 mm (vs. 15 mm) en el orificio y anteras ligeramente más cortas, de

2.9-3.1 mm de add (vs. 4.3 mm).

COLOMBIA. € í te, 26 nov 1992 (fl), Bar? c ones 8084 (COL); sierra aa pera

campamento Norte, 6 Dic 1990 a a in ji al. 1021 (COL, MA [2 d 7 Di

(fl, fr), Castroviejo et al. 11989 (CO de Chiribiquete, 19 nov 1992 (tb, Franco et al. 4127 (C OL, MA); si d Chine

ae sur, 11 Dic a mr Pieres et al. 1144 (COL); sierra de Chiribiquete, 10 Dic 1990 (fL), Ella et al. 2286 (COL, M

sierra de Chiribiquete, cueva de las Pinturas 1 Ago 1992 (fr), Palacios et al. 2404 (COAH, COL, P); sin localidad precisa, 26 nov 1992

(fr), Velayos et al. 6386 (COD.

Mandevilla sancta-martae J.F Morales, sp. nov. (Fig. 3). Tiro: COLOMBIA. }

E de quebrada Indiana, 1 sep 1972 (fl), Kirkbride 2049 GaoLotipo: INB; isoripos: COL, NY, US).

A Mandevilla schlimii (Mall. Arg.) Wood ilis, foliiis glabris (vs. tomentosis), tubo proprio 17-18 mm (vs. 10-14 mm) et faucibus

16-17 mm diametro (vs. 8—10 mm) differt.

Liana; ramitas cilindricas a subcilindricas, a veces algo aplanadas antes de los nudos en ramitas tiernas,

sólidas, glabras; coléteres interpeciolares inconspicuos, hasta 0,6 mm de largo. Hojas opuestas; pecíolos

11-17 mm de largo; láminas foliares 5.5—10.2 x 1.8—3.7(—4.1) cm, elípticas a ovado-elípticas, el ápice larga-

mente acuminado, la base cordada, los coléteres irregularmente distribuidos a lo largo del nervio central,

membranaceas, glabras, no bulladas, el margen no revoluto, las venas secundarias y terciarias impresas

en ambas superficies, pero usualmente las venas terciarias algo inconspicuas en la superficie adaxial, con

domacios diminutos a lo largo del nervio central abaxialmente. Inflorescencia igualando a más larga que las

hojas subyacentes, axilar, glabra, con relativamente pocas flores ete 12), pedunculo 10-17 mm de largo,

pedicelos 3-5 mm de ALES. brácteas 3-6 x 0.6-0.9 mm, ang ovadas, ; sépalos 3.2—4.5 x

1.1-1.5 mm, ang ovados, similares en longitud, el ápice acuminado, no reflexo, escariosos, glabros

externamente, el coléter solitario, abener o inrecularmenie lacerado; ol infundibuliforme, amarilla,

los botones con manchas rojizas, glabra externamente, el tubo algo giboso en la base, la parte inferior 9-10

x 3.7-5 mm, la parte superior 17-18 mm de largo, anchamente campanulada, con un diámetro en la boca

de 16-17 mm, el ápice del poen floral e oana a obtuso; lóbulos 16-21 x 12-16 mm, obovados,

a base de la part ior del tubo, anteras 4.8-5.2

mm de largo, sabres dorselmente la base auriculada, con las aurículas beds: cabeza estigmática

2-2.2 mm de largo; ovario 1.2-1.4 mm de largo, glabro; nectario anular, pentalobulado, ca. la mitad de la

longitud del ovario. Folículos desconocidos.

Distribución, hábitat y ecología.—Endémica al Departamento de Magdalena en Colombia, donde crece

en bosques muy húmedos en la Sierra Nevada de Santa Marta en elevaciones de 1600—1700 m. Especimenes

con flores han sido recolectados entre agosto y septiembre.

Mandevilla oie ae pues ser d con M. schlimii (Müll. Arg.) Woodson, pero se puede

reconocer por s con la y del tubo 17-18 mm de largo y con un diámetro de 16-17 mm

en la boca (vs. 10-14 mm de jango y 8- 10 mm de diámetro), láminas foliares con la superficie abaxial glabra

y con domacios a lo largo del nervio central en la superficie abaxial (vs. glauco-tomentosas y sin domacios).

Otro grupo de especies relacionadas incluyen a M. mollissima (Kunth) Woodson, M. scabra (Hoftmanns. ex

Roem. & Schult.) K. Schum., M. speciosa (Kunth) J.F. Morales y M. symphitocarpa (G. Mey.) Woodson, pero

M. sanctae-martae se puede separar de estas especies por la presencia de domacios en sus hojas y corolas

con la parte inferior del tubo inferior a 1 cm de largo. El epiteto de esta especie hace referencia a la Sierra

Nevada de Santa Marta en Colombia, una zona con un alto indice de endemismo.

T ] Lal CURADIA AR 1.1 [e "AT J -l c " Ja Dant Dad Azul [at

29. sep 1972 (ID, Kirkbride 2277 (COL, US); c camino de San Javier a San Andrés, por a filo del Cerro Ratón, 23 Ene 1959 (fl), ROET:

Castañeda 7055 (COL)

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Morales, N ies de Mandevill Colombia y Venezue la 571

AGRADECIMIENTOS

Quiero agradecer a Julio Betancur y Rodrigo Bernal (COL) por las atenciones brindadas en las visitas al

Herbario Nacional de Colombia (COL), así como a Ricardo Callejas, Francisco Roldán y Ramiro Fonnegra

del herbario de la Universidad de Antioquia, Colombia (HUA) y a Eliana Noguera, Shingo Nozawa y Sirli

Leython en el Herbario Nacional de Venezuela (VEN). Se agradece a los siguientes herbarios por el envio de

material en préstamo o por el préstamo de sus colecciones: BM, C, CGE, COAH, COL, F, G, G-DC, HUA,

K, M, MA, MO, NY, P, P-JU, P-LA, US. Thanks to Raul Gutierrez and Mary E. Endress for reviewing an

earlier draft of the manuscript.

REFERENCIAS

MONACHINO, J. 1961. Apocynaceee. En, B. Maguire, J. Wurdack y colaboradores, eds. The botany of the Guayana

Highland IV(2). Mem. New York Bot. Gard. 10(4):59-65

Morales, J.F. 2005. Estudios en las Apocynaceae Neotropicales XIX: la familia Apocynaceae (Apocynoideae,

Rauvolficideae) en Costa Rica. Darwiniana 43:90-191.

Woobson, R.E. 1933. Studies in the Apocynaceae IV. The American genera of Echitoideae XXVI. Ann. Missouri

Bot. Gard. 20:605-790.

572 [| [| E al n.a H In Li FE £f'T 3(2)

BOOK REVIEW

Jerry A. POWELL AND Paur A.OpLER. 2009. Moths of Western North America. (ISBN 978-0-520-25197-7,

hbk.). University of California Press, 2120 Berkeley Way, Berkeley, California 94704-1012, U.S.A.

(Orders: www.ucpress.edu or California-Princeton Fulfillment Services, 1445 Lower Ferry Road, Ewing,

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of at least 2596 of the species i h family. Every y superfamily subfamily, a and family ibed as to basic cl teristics, followed

by ] ipti h ref to tl j I , some plates witl tl 50 images. Plates

59 ugh 64 life cy f selected lepidopt 1 the | f “macros' O on their food plants. Western

species predominate in coverage, but the West extend the Rocki d int as, Texas, etc., and there are many

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Karges, Texas Christian University Library (retired), Fort Worth, Texas, U.S.A.

J. Bot. Res. Inst. Texas 3(2): 572. 2009

STUDIES OF NEOTROPICAL COMPOSITAE-II.

A NEW COMBINATION IN CHROMOLAENA (EUPATORIEAE)

John F. Pruski

Missouri Botanical Garden

PO. Box 299

St. Louis, Missouri 63166-0299, U.S.A.

ABSTRACT

RESUMEN

Ir I

Most species of Chromolaena DC. (Compositae: Eupatorieae: Praxelinae) have been treated historically as

members of Eupatorium sect. Cylindrocephala (e.g., Robinson 1918, 1919). id al as d by

King and Robinson (1987), however, differs from Eupatorium L. by glabrous (vs. setul , distinct

(vs. indistinct) popon and S M dads of deciduous (vs. persistent les Chiomalastia

differs from l corolla lobes, that are usually flat on top,

long pappus bristles, and BEA branches "hat are cj n ndrical and never clavate (King & Robinson 1987).

In a revision of Peruvian Eupatoriums, B.L. diobinson (1919) recognized the distinctive Eupatorium

clematitis DC. (see Fig. 1), whereas King and Robinson oe and Robinson and Holmes (2008) list it in

synonymy of C. odorata (L.) R.M. King & H. Rob. Eupatorium titis was not listed as occurring in Ecuador

by Robinson (1918). Chromolaena odorata is common and widespread throughout much of De neotropics,

and is known also as an introduced weed in the paleotropics (King & Robinson 1987; Pruski 1997). My

treatment of Compositae for the Río Cenepa region of northern Peru recognizes C. clematitis as endemic to

Ecuador and Peru, where it occurs in low foothills of the eastern Andes. The purposes of this paper are to

provide a new combination for this species and a description of it for use as an aid in identifications.

Chromolaena clematitis (DC.) Pruski, comb. nov. (Fig. 2). Bastonym: Eupatorium clematitis DC., Prodr. 5:144. 1836.

Osmia clematitis (DC.) Sch. Bip., Jahresber. Pollichia 22—24:252. 1866. Tier: “Peruvia?,” 1834 [sic], Poeppig 3108 (HoLorvee: G-DC

[photograph sub F neg. 8150 in MO, US; IDC microfiche 800. 803.111.6]; isotypes: E W

Perennial herbs to vining shrubs, 0.7-2 m tall; stems few-branched distally, subterete, puberulent to glabrate.

Leaves simple, opposite, long-petiolate; petiole 0.8-3 cm long, slender, puberülent: blade deltate to hastate,

2-7 cm long, 0.6-5.5 cm wide, chartaceous to eu so, 3-veined from base, base hast t t ae

c ] lent alc : i glabr

entire to few-serrulate, apex long inate to attenuate, adaxial

abaxial surface densely red-glandular, otherwise irc to Eibrous: the non- Sn LES very

short. Capitulescence terminal, open, thyrsoid-paniculate with ultimate branches umbelliform to corymbi-

form, 2—21-headed, to 15-20 cm tall, ca. 15 cm wide. Capitula discoid, 23-26-flowered, 9.5—12.5 mm high,

2.83.5 mm wide; involucre cylindrical, 9—11.5 mm long, corollas slightly exserted; phyllaries imbricate,

graduated, 5—7-seriate, tightly appressed, deciduous in fruit, scarious with apex subherbaceous, generally

3-nerved, most or at least the mid-series phyllaries truncate-mucronate apically, glabrous to outer series

sparsely puberulent distally, outer ones elliptic-lanceolate to ovate, 1-2 mm long, 1-2 mm wide, grading

into inner ones, these lanceolate, 9—11.5 mm long, 1-1.5 mm wide; receptacle epaleate, shortly clavate but

flat on top, ca. 1.6 mm tall and wide, glabrous; peduncle 1—2.5 cm long, striate, often 1- or 2-bracteolate,

puberulent, main capitulescence stalk to 6 cm long. Florets bisexual; corolla white, funnelform, 4.8—5.7

mm long, glabrous, lobes subequal, ca. 0.5 mm long, erect to slightly spreading; anthers included; style

J. Bot. Res. Inst. Texas 3{2): 573 — 576. 2009

1 q £ sl n.a H in L Boden aiio 3/3)

574 f Texas Ji]

En 1 PL i i aL’ (DC.) Pru cli A a P.L L Fi A L (P. i LEM LLYEF.P : MO) n P. aa a L L] iat de 1 L L Hf

ith | | lat itul (Rueda et al. 1043, MO).

base glabrous, branches cylindrical, greatly exserted from corolla, 3-4 mm long, the proximal half of each

branch with paired ventro-marginal stigmatic lines, the distal half with minutely papillose sterile append-

age with an acute to obtuse apex. Cypselae narrowly obconical, brown with tan angles, 3-3.6 mm long,

3-4-angled, glabrous; carpopodium distinct, tan, circular; pappus bristles many, elongate, stramineous,

ca. 5.5 mm long.

en a examined: papi ten Napo: Estación Biológica Jatun Sacha, Río Napo, 8 km al E de Misahuallí, bosque muy

1°04'S, 77°36'W, 450 m, 23-27 Jun 1987, Cerón 1678 (MO); Tena Canton, Aliahui,

I húmed dad o pa asia 450 m, 1°04'S, 77°36'W, Jul 1994, Palacios 12395 (MO); Tena Cantón, Esta-

ción Badean. sucha, sendero 1 y 3, 1*04' s. 77°36'W, 380 m, 9 Aug 1992, Rueda et al. 1043 (MO). PERU. Amazonas: Bagua Prov.,

Yamayakat, bosque secundario, ne "e 78°19'W, aoe m, ario 1995, Vasquez et al. 20117 (MO). San Martin: Mariscal Caceres, Tocache

Nuevo, Puerto Pizana, má H ga, 350—400 m, 22 Jun 1974, Schunke V. 6995 (M

]4 34 rl Hticicl tr fl f June to AYeust at ] ti ]

Distribution and ecology.—C} g

350-450 meters in the eastern Andean foothills of Ecuador and Peru om about 1—8? south latitude.

Chromolaena clematitis is known only from a few collections in northern Peru and eastern Ecuador, from

where the type was presumably collected. By gestalt and reddish sessile glands of the abaxial leaf surfaces,

C. clematitis is similar to C. odorata, under Had it was SURE by King and Robinson (1987) and

Robinson and Holmes (2008). Chromola l easily, however, by its leaves with hastate

to truncate bases and capitulescences with ultimate brariches of long- -pedumcüluie typically umbelliform

capitula. Although across its broad range in both hemispheres C. odorata is somewhat variable in shape

and pubescence of leaves and phyllaries, its variation is minor and continuous, while C. clematitis remains

—

Pruski, Anew combination in Chromolaena 575

| Esparta Cea y De f "

Fic > Phot L valk. Lobos iP, 39770 fh} ALC Ew j PETHE Ya

LI F-a E f

576 j i tanical h Institute of Texas 3(2)

consistently distinctive. Chromolaena clematitis differs from C. odorata by basally hastate to truncate (vs. at-

tenuate to subtruncate) leaves with blades glandular, otherwise puberulent to glabrous (vs. glandular and

pubescent) abaxially, by ultimate branches of the capitulescences with long (vs. short)-pedunculate typically

umbelliform (vs. corymbiform) capitula, and by most or at least the mid-series phyllaries with truncate-

mucronate (vs. acute to rounded) apices.

ACKNOWLEDGMENTS

idus. i re = ] j e

I would like to thank Rosa Ortiz (MO) for translating the abstract i p for helping prey

1, Walter Holmes (BAYLU), Rosa Ortiz (MO), and Harold Robinson (US) for comments on Le mamiseript

Lucia Kawasaki and Anna Balla (F) for sending me an image of the holotype, and Christine Niezgoda (F)

for permission to publish this photograph.

REFERENCES

King, R.M. AND H. Rosinson. (1987). The genera of Eupatorieae (Asteraceae). Monogr. Syst. Bot. Missouri Bot. Gard.

22:1—58]

Pruski, J.F. 1997. Asteraceae. In: J.A. Steyermark, PE. Berry, and B.K. Holst, eds. Flora of the Venezuelan Guayana,

vol. 3. Missouri Botanical Garden, St. Louis. Pp. 177-393.

Rosinson, B.L. 1918. Keyed recensions of the Eupatoriums of Venezuela and Ecuador. Proc. Amer. Acad. Arts

54:331-307.

Rosinson, B.L. 1919. A recension of the Eupatoriums of Peru. Proc. Amer. Acad. Arts 55:42-88.

Rosinson, H. AND W.C. HoLmes, 2008. 190(3). Compositae-Eupatorieae. Flora of Ecuador 83:1-347.

FOLIA TAXONOMICA 13. PASSIFLORA CURVA (PASSIFLORACEAE),

A NEW SPECIES FROM FRENCH GUIANA

IN SUBGENUS PASSIFLORA SUPERSECTION COCCINEA

Christian Feuillet

Department of Botany, MRC-166

Smithsonian Institution

20. Box 3701

Washington, DC 20013-7012, U.S.A.

feuillec@si.edu

ABSTRACT

A new species, Passiflora curva, is described in subgenus Passifl I tion Coccinea. It was eallecied? in French Guiana, in the

lower Sinnamary River basin. Passifl i ily set apart from the other species persection by its leaves with margins

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outer corona filaments white and erect.

RESUME

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vers la base et serrée vers na ses bricos longues de moins de 3 cm, ses fleurs avec un take beue et le verticille mm de la

couronne blanc et érigé.

During a field trip in French Guiana in September 2008, I had the Pp nani to visit the DE ind of

Passiflora aimae Annonay & Feuillet (1998) and collected two good sr that clearly

ina also in Passiflora DEDE cue Feuillet SJ. M. MacDougal. e new pea Pc curva, and P. aimae

ost i iew sp color of the bud and floral

tube, and the morphology of the two-row corona in wilüch " t of fil t, leaning neither

inward nor outward. Pictures of both species have been published; Passiflora aimae in Ulmer & MacDougal

(2004: pl. 161, p. 212; photo H. Annonay); and Passiflora curva (as P. aimae) in Ulmer & MacDougal (2004:

pl. 8, p. 165; photo H. Gelewsky) and in Ulmer & Ulmer (2005: p. 9; photo H. Gelewsky). Some have been

uploaded and are available on the web: Passiflora aimae on C. Houel's website (photo C. Houel); and Passiflora

curva on C. Houel's web site (as “sp. Distephana”; photo C. Houel); and on Flickr in cpfl’s photostream (as

P. curva; photo C. Feuillet). Christian Houel ("Collection Nationale de Passiflores", France) became aware

there were two different species when he found the two growing side by side. As far as I know, encounters

with this species by various persons previous to the type collection were not documented by herbarium

specimens, but it may be in cultivation in Europe or in Martinique where H. Annonay has his collection.

Passiflora curva Feuillet, sp. nov. (Fig. 1) n GUIANA l Si River basin, ca. 20 km SW of Sinna

on the roadside of Route de St Elie (D 21), 13 km S oft] 1 from Cay to St.-Laurent du-Maroni (N 1), 5°18'20"N en

35 m, 28 Sep 2008, fl., Feuillet 17049 (noLorvrE: US; : BRIT, CAY).

Passiflora cui 1 E D ifl I à K^ ie I j I H p. gi 1 1 S 1

(ab basi ad ci lium) vel serratis (ab ci dio ad apicem), bracteis minus 3 cm longis, hypanthio extus stramineo differt.

Climber with tendrils; plant with short tric] on vegetative parts, the outide of the sepals, the androgy-

nophore, and the gynoecium, the flower otherwise glabrous. Stem reddish, somewhat 5-angular, reaching

at least 6 m long. Tendrils axillary, reddish. Stipules setaceous, reddish, 5-6 mm long. Leaves alternate

petiole 1-1.8 cm long, reddish, with a pair of glands at the base, glands round and plump, green when

young becoming reddish; blade ovate, 7-12 x 3.5—7 cm, base truncate to slightly cordate and shortly cune-

J. Bot. Res. Inst. Texas 3(2): 577 — 580. 2009

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ate, apex rounded to slightly acute, short acuminate, margin reddish-brown, slightly and loosely crenate

in basal third and serrate in apical two thirds, adaxial surface yellowish-green when young becoming dark

green, abaxially pale grey becoming light green, veins reddish, 5-6 veins each side of the midrib. Peduncle

axillary, solitary, erect, 4-5 cm long, reddish; bracts (1 bract & 2 bracteoles) verticillate, inserted 1 mm

below the Joni, similar in shape and size, 1. n 2 x 0.6—0.8 cm, red, green at base, 4—5 pairs of glands at

margin, gl 1, green, larger at | the joint, floral stipe 0.4—0.5 cm long. Flowers erect floral

tube campanulate, base invaginate Around the apex of the stipe, forming inside a 10-lobed nectar chamber

at the bottom, 10-12 cm long, greenish straw-colored suffused with pink and with 10 green veins outside,

inside pink to red; sepals red but the parts exposed in the bud straw-colored abaxially, narrow oblong,

carinate, with a 2-4 mm long subterminal red awn curved inward, round at apex, 3 x 0.6 cm; petals red

all over, without carina or awn, otherwise similar to sepals; corona in 2 rows, the outer row of white erect

filaments about 1 cm long, laterally flattened, subulate, apex reaching the level of the base of the stamens,

but not leaning on the androgynophore, inner row about as long, membranous in the basal half, oblique

toward the androgynophore, white suffused with pink, and filamentous in the apical half, white, point-

ing toward the base of the tube and leaning on the androgynophore, then becoming straight and erect;

operculum born 2/3 up the tube, membranous, parallel to the inner corona, bent about 2/3 from the base,

short fimbriate at margin, white suffused with pink; nectary ring on the bottom of the tube reddish; limen

pinkish, membranous, toothed at margin, surrounding the base of the androgynophore, curved outward

at margin; androgynophore 2.5 cm long, white at base, cream dotted with red above; stamens 5, shortly

fused at base, hiding the short gynophore and the base of the ovary, filaments dorsiventrally flattened, 15

x 1.5 mm, cream heavily spotted with red, emarginate at apex, anthers 9 x 2.5 mm, nearly rectangular,

dorsifixed, held transversly, white, slightly greenish-cream near margin, pollen white; ovary ovoid, 9 x 2

mm, cream, styles slightly S-shaped, 6—7 mm long, diameter slightly increasing toward the apex, stigmas

globular, 1.5-2 mm in diam., stigmatic surface cream, otherwise bright red. Fruits not seen; (the following

from a photograph by C. Houel) unripe fruit green, dotted with white, ovoid.

By its red bracts with marginal glands, red-faced flowers, white erect straight outer corona filaments,

and leaves crenate to serrate and glandular at margin, P. curva belongs in subg. Passiflora supersect. Coc-

cinea. It differs from the other species by the combination of the following characters: the leaf blade mar-

gins are loosely and slightly serrate in the apical half and loosely and slightly crenate in the basal half, the

bracts are much shorter than the flowers, the floral tube and the parts of the calyx exposed in the bud are

straw-colored, and the corona has 2 rows, the outer row erect and comprised of white filaments not lean-

ing on the androgynophore. Lastly, the inner corona row partly membranous, initially held horizontally

(unpublished photograph) and later upward oblique. In both stages it is bent in the middle with the apical

filamentous half held downward oblique, (Fig. 1, both photographs taken around 8:00 am, 28 Sep 2008).

Ultimately it unfolds to become erect and parallel to the outer row (unpublished drawing of the paratype

by J. Vanderplank, around 10:30 am, 17 Mar 2009). The basal membrane of inner row of the corona is an

effective, although temporary, barrier keeping pollinators or nectar robbers away from the second obstacle,

the operculum, on the way to the nectar. The flowers of the type collection had the inner row of the corona

bent when split open and placed in the loosely tied field press, but the filaments straightened before being

placed in the drier.

In the last published key of the supersection (Feuillet 2007), P. curva would be in a new third arm of

bracket 4, reading: “Leaf margin slightly crenate in basal third and slightly serrate in distal two third,” as

opposed to “Leaf margin crenate” or “Leaf margin dentate or serrate.” The second group includes 9 species

with largely or densely serrate or biserrate leaf margins. The first group includes 2 species with crenate

leaf margins from the same small area of forested hills west of the Lower Sinnamary River, P. aimae with a

much shorter corona, 0.5—0.6 mm versus] cm, whose inner row is red, and P. longicuspis Vanderpl. with a

much longer corona, 1.8—2.4 cm, in 3 rows whose 2 outside rows are red or deep purple, and both with the

corona filaments leaning on the androgynophore.

Feuillet, Pacciflora curva, 4 f. E Lf. A

Fic. 1. Passiflora curva. A. Flower buds

beginning to open; B. Flower (lon-

gitudinal cut). Both photographs of

the type plant, Feuillet 17049. (Photo-

graphs C. Feuillet)

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Distribution and pl Passiflora curva is | only from the type locality west of the Sinnamary

River, 13 km south ol Road N 1, on the first hills south of the littoral savannas. It was found blooming in

March and September.

Etymology.—The epithet, from the Latin adjective curvus, means bent, referring to the morphology of

the inner row of the corona at M anthesis.

ParaTYPE: FRENCH GUIANA: ca. 20 km f Si ide of Piste de St Elie, 13 km from Rd 1, 5° 18'20"N 53° 02'21"W,

35 m, 17 Mar 2009, fl., R.J.R. SIE C. Feuillet & M. Medea 1620/09 (CAY, Nat Coll. nd UK).

ACKNOWLEDGMENTS

I thank John MacDougal and Steve Tillett for their review of the manuscript. I am indebted to Christian

Houel and John Vanderplank for sharing their data. I thank Sophie Gonzalez, director of the herbarium in

Cayenne (CAY) for providing the ideal conditions to work with the collections in her care. This is number

154 in the Smithsonian's Biological Diversity of the Guiana Shield Program publication series.

REFERENCES

dd H. AND He FEUILLET. 1998. Passiflora aimae (Passifloraceae), une espèce nouvelle de Guyane francaise.

EU C. 2007. Folia taxonomica 2. New species of Passiflora subgenus Passiflora (Passifloraceae) from the

Guianas. J. Bot. Res. Inst. Texas 1:819—825.

FeuiLLET, C. in Flickr website, flick /ohotos/68587888@NO0 g /, accessed 20 June 2009.

Hout, C. website http://www. puede fr/, accessed 5 Apri 2009.

ULMER, B. AND T. UL MER. 2005. Farbatlas passionsblumen. Formosa Verlag, Witten.

ULMER, T. AND J.M. MacDoucat. 2004. Passiflora, passionflowers of the world. Timber Press, Portland, Oregon.

FOLIA TAXONOMICA 14. NOTES ON PASSIFLORA SUPERSECTION COCCINEA

(PASSIFLORACEAE) FROM THE GUIANA SHIELD

Christian Feuillet John Vanderplank

Department of Botany, MRC-166 National Collection of Passiflora

Smithsonian Institution Lampley Road, Kingston Seymour

PO. Box 37012 Clevedon, North Somerset BS21 6X5, U.K.

Washington, DC 20013-7012, U.S.A. johnvanderplank@yahoo.co.uk

feuillec@si.edu

ABSTRACT

ERE dd 1 I 1.3 E TES ees gi r 1 : fp ifl I t i F he Gui Shi ld, P. compar,

P. longicuspis and P. tecta.

RÉSUMÉ

D llecti lles étentent la répartition de trois espèces de Passiflora supersect. Coccinea du Bouclier Guyanais, P. compar, P.

longicuspis, et P. tecta.

During a field trip in French Guiana in March 2009, we had the opportunity to study the collections at CAY.

Some interesting specimens of Passiflora species belonging in subg. Passiflora supersect. Coccinea Feuillet

=. M. MUD are worth OEA as sha peu Inu ne and poony collected Apo

plank 2006; Feuillet 2007, 2008)

cies of supersect. Coccinea with unlobed leaves at US and in the herbarium associated with the OS

holding the National Collection of Passiflora (United Kingdom). The new data presented here expand the

known distribution area for those species.

1. Passiflora compar Feuillet, J. Bot. Res. Inst. Texas 1:821. 2007.

Passiflora compar is easily separated from other species of su] tion Coccinea by its leaves that are much

thinner and from most other species by the corona with 2 rows of red or pink filaments 5-7 mm long.

Add 1T All ERENCH CUTAN +} Arataye Piran NE NI around camp f hel 1,200

m, 6 Jan 2003, fl., S.A. Mori et al. 25527 (CAY, NY); Saul, Hen of Paine Claires, 3°37 N, 53*12'W, ca. 300 n m, 16 Feb 1993, il, S.A.

Mori et al. 23004 (NY, US).

Distribution and phenol Passiflora compar is known in Guyana from the Upper Takutu-Upper Essequibo

Region, la Mts., from the Petar eae Region, near Kato, and in French Guiana from the upper

Maroni basin, Saut Lavaud, from the Satil area, Eaux Claires, and from Montagne des Nouragues. It inhabits

moist forest below 400 m of elevation, near rivers or on slopes. It is expected to be present in Surinam.

Passiflora compar was blooming in January, February, March, September, and November; it was fruiting in

February and March.

2. Passiflora longicuspis Vanderplank, Curtiss Bot. Mag. 23:232; pl. 563. 2006.

Passiflora longicuspis is characterized by the corona filaments, 22-26 mm long, reaching the ovary on top of

the androgynophore. It is also one of the few with crenate leaf margins, one of the few with small bracts,

and one of the few with colored outer corona filaments.

4A Aditi 1 eal] ICH GUIANA Cayenne, Mont Baduel, 10 m, 2 Jun 1987, fl ,M.H if 5222 (CAYY Pi le St Élie, Mar 2003,

fl., RR. Vanderplank & S.E. Vanderplank 1762/09 (Nat l} )

Distribution and phenology —The type of P. longicuspis from the British National Collection of Passiflora origi-

nated in French Guiana from Route de St Elie, near the Si y River. The new collections are from Mont

J, Bot. Res. Inst. Texas 3(2): 581 — 582. 2009

[| [| £ all n.a H ID LI PF ai”

582 f Texas 3(2)

Baduel situated inside the administrative limits of the city of Cayenne and from the hills SW of Sinnamary

neat the type locality. It was blooming in March and June.

3. Passiflora tecta Feuillet, J. Bot. Res. Inst. Texas 2:281. 2008.

In the supersection, P. tecta is characterized by its leaf blades that are bi-serrate with the serrations tipped

with glands, its corona in 3 rows of filaments, and the 2 outer rows that are deep red with the outmost row

12 mm long and the middle row 9 mm vod

Additional collections: VENEZUELA. Bolí Roscio Distr., | SanF i le Y í and Chirimatá, 5°02'N 61°00'W, 1100

fl., O. Huber 9154 TUS GUYANA. Cuyuni — Mazaruni Ayanganna Plateau, 740 m, 19 Jul 1960, f1., S.S. Tillett et al. 44873

(NY, US); Groet ibo River, 26 Jun 1944, fl., D.B. Fanshawe 4728 (F1992) (NY, US); Ridge of Eagle Mountain, 5?12'N

59°07'W. 304-685 m, 10 Oct 1990, a T. McDowell 3472 (US). Essequibo Isl. — W Demerara: Lower Tiger Creek, 6°30'N 58°39'W,

0-15 m, 12 Dec 1992, fl., TW. Henkel 485 (US). Potaro — Siparuni: Essequibo River, Karupukari Crossing, 4°40'N, 58°41'W, 60-75

m, 19 Apr 1992, fl. & fr., B. Hoffman 1345 (US); Iwokrama Rainforest Reserve; N of Surama, 4?10'N, 59°3'W, 200 m, 20 May 1995, fl.,

C. Ehringhaus 89 (CAY, US). Upper Demerara — Berbice: 16-22 km from Ituni along Ituni - Kwakwani Road, 5°254N, 58°10'W, 70 m,

19 Jan 1990, fl., LJ. Gillespie et al. 3086 (CAY, US). SURINAM. Sipaliwini: vicinity of Blanche Marie Waterfall, 4°45'20"N 56°54'19"W,

60 m, 3 Feb 1998, fL, J.S. Miller et al. 9304 (MO, US

Two more specimens from plants having a pale band on the outer corona row probably represent the same

species.

RINAM. Brokopondo: Brownsberg, 1997, fl., NCP 1086/06 (National Collection of Passiflora herbarium). UNITED KINGDOM:

cultivated from seed collected in Surinam, National Collection of Passiflora greenhouses, 2000, NCP 1302/08 (National Collection of

Passiflora herbarium).

Distribution and phenology.—Passiflora tecta is now known from the State of Bolivar in Venezuela, from

Guyana, and from northern Sipaliwini and perhaps Brokopondo in Surinam. It was blooming in January,

February (photograph only), April to July, and September; it was in fruit in February (photograph only),

April, and October.

ACKNOWLEDGMENTS

We are grateful to John MacDougal and Steve Tillett for reviewing the manuscript. We thank Sophie

Gonzalez, director of the herbarium in Cayenne (CAY) for providing the ideal conditions to work with the

collections in her care. This is number 155 in the Smithsonian’s Biological Diversity of the Guiana Shield

Program publication series.

REFERENCES

FeuiLLET, C. 2007. Folia taxonomica 2. New species of Passiflora subgenus Passiflora (Passifloraceae) from the

Guianas. J. Bot. Res. Inst. Texas 1:819-825.

Feuer, C. 2008. Folia taxonomica 8. Passiflora tecta, a new species in subgenus Passiflora from the Guianas. J.

Bot. Res. Inst. Texas 2:281-283.

VANDERPLANK, J. 2006. Plate 563. Passiflora longicuspis (Passifloraceae). Curtis's Bot. Mag. 23:23 1-236; plate.

FOLIA TAXONOMICA 15. FIVE NEW SPECIES OF PARADRYMONIA

SUBGENUS PARADRYMONIA (GESNERIACEAE: EPISCIEAE)

FROM THE VENEZUELAN GUAYANA

Christian Feuillet

Department of Botany, MRC-166

Smithsonian Institution, PO. Box 37012

Washington, DC 20013-7012, U.S.A.

feuillec@si.edu

ABSTRACT

GA i ID A : 1 D 3 E A cae i T : ^ A 47, 21-3. +1 pata al AM ee exu Venezuela: Par-

adrymonia glandulosa, P. hamata, P. lutea, P. tepui, and P. yatua.

RÉSUMÉ

de P ia subg. P ia (G j : Episcieae ) sont décrites de l'état d'A , Venezuela: Paradry-

inn oct» noc

monia slanduloaa: P. Tamata P. a, P. tepui et P. yatua.

RESUMEN

C: A EUER ia (G Episcieae) descritas del estado Amazonas, Venezuela: Paradry-

monia glandulosa, P. hamata, P. lutea, P. tepui y P. yatua.

Continuing the publication of new species of Paradrymonia Hanst. from the Venezuelan Guayana (for an

introduction cf. Feuillet 2009), I propose here five new species in subgenus Paradrymonia.

1. or eens Pewien, Spunoy (Fig. 1. i VENEZUELA. Amazonas. Dep

“Sima Camp”, central portion of foreste g ran año Negro, 3°43'N, 65°31'W, 1140 m, 21-24 Feb

1985, fl., J.A. Steyermark & B. Holst 130443 (ism : MO, VEN).

Paradrymoniae ciliosae affinis; lamina ovata vel late elliptica, sepalis apice linearibus, corolla 3.5 cm longa, corollae lobis brevis, non

fimbriatis differt.

Terrestrial, saxicolous herb or basal trunk epiphyte. Stem sappy becoming subwoody at base, procumbent,

hirsute, glabrescent t ds | about 0.5 cm thick. Leaves strongly unequal in a pair, the smaller one early

deciduous; larger leaf: petiole 8— 15 cm long, densely to slightly asa pee blade chartaceous when

dry, 20-28 x 10-13 cm, broadly ovate to elliptic, base cuneate, caudate, margin glandular

serrate, above appressed-pilose to glabrous, beneath minutely pubescent to ane ne axillary,

fasciculate; ele 0.1-0.6 cm long, densely pilose, tricl ereen bordered with

maroon, free, subulate, 8-20 x 1-1.5 mm, apex long linear-acuminate about bali the total length, margin

loosely glandular serrate, pilose; corolla y white with purple lines, oblique in the calyx, basal gibbosity

prolonging the tube, curved, 3 x 2 mm, tube 2.5 cm long, narrow for 1.8 cm, then widely funnel-shaped,

with red trichomes, lobes shallow. Fruits not seen.

Distribution —Paradrymonia glandulosa was collected on forested slopes of Cerro Marhuaca and Cerro

Yapacana, department of Atabapo, state of Amazonas (Venezuela), between 1140-1200 m. It was in bloom

in January—February.

Paradrymonia glandulosa resembles P. ciliosa by its general appearance: thick stem with usually short

internodes, creeping on rocks or climbing lower tree trunks, large leaves with long petioles, and very short

inflorescences in the leaf axils. It differs from P. ciliosa and other species of subg. Paradrymonia from Ven-

ezuelan Guayana by the leaf blade ovate to widely elliptic with teeth wide-glandular at the tip, the sepals

subulate and linear-acuminate, the corolla with a curved basal gibbosity, and short, non fimbriate lobes.

J. Bot. Res. Inst. Texas 3(2): 583 — 592. 2009

F H F EE) fa Lalo Lasa 437/445 0g: d Bari L ri J:

leaf in inset. idem (US)

f the inset = 12 cm; corolla inset, Maguire, Cowan & Wurdack 30657 (NY), total length of the object = 3.5 cm.

Feuillet, Fi ies of Parad ia f the V | G 585

Although the flowers pressed in situ seem erect in a calyx, a separate corolla in the pocket of Maguire

et al. 30657 clearly shows a lateral insertion at the | t the curved gibbosity and the tube suggesting

that the corolla is strongly oblique in the calyx Ma e the na is ad a a on the pedicel. A situation

possibly imposed by the multi-flowered fasci g between the stem and the long

petioles. This species was “sp. C” in Feuillet & Steyermark (1999).

Etymology.—The epithet glandulosa refers to the glandular teeth on the margin of the leaf blades, the

bracts, and the calyx lobes.

PARATYPE: VENEZUELA. Amazonas. Depto. Atabapo: Cerro Yapacana, on slopes and cumbre, 1200 m, 3 Jan 1951, fl., B. Maguire, R.S.

Cowan & J.J. Wurdack 30657 (NY

2. Paradrymonia hamata Feuillet, sp. nov. (Fig. 2). Tree: VENEZUELA. Amazonas. Depto. Río Negro: Cerro de la Neblina,

Río Yatúa, 140-1700 m, 31 Dec 1957, fl., B. Maguire, J.J. Wurdack & C.K. Maguire 42563 (HoLotyre: NY).

Paradrymoniae ciliosae affinis; lamina ovata vel orbiculata, pedicello usque ad 4.5 cm longo, sepalis in dimidio superiore linearibus,

a 5-6 cm longa differt.

Terrestrial (?) herb; stem about 0.5 cm thick; trichomes red. Leaves so unequal in a pair that they look

alternate, small one obsolete or early deciduous, lamina not developing; larger one: petiole 5-12 cm long,

pubescent; blade broadly ovate to orbicular, base rounded or obtuse, then abruptly cuneate and narrowly

somewhat decurrent, apex broadly and shortly acuminate, margin obscurely to clearly glandular-serrate,

above minutely puberulent to glabrous, beneath appressed-pubescent on mains veins, trichomes scattered

between, margin densely appressed pubescent. Inflorescence axillary, fasciculate, pedicel 1-4.5 cm long,

long-hirsute. Flowers: calyx with long, erect, trichomes, sepals narrowly triangular-elliptic and for half its

length long linear-acuminate, 15-20 x 2-3 mm, with 13 teeth on the linear tip and near its base; corolla

white or hyaline with scattered trichomes outside, basal gibbosity 0.4—0.5 x 0.4 mm, tube narrowly tubular

at base, abruptly widening, then broadly tubular, 4-4.5 cm long, with white trichomes, lobes suborbicular,

0.9-1.2 cm, the ventral lobe shortly fimbriate, the others crenate to entire at margin; stamens borne near base

of the tube, seen only recoiled to the base of the wide portion of the tube, anthers thecae widely divergent,

about 1.5 mm long. Fruit not seen.

Distribution. —Paradrymonia hamata was collected partly buried in duff at 1200 m on Cerro de la Neblina

(Department Río Negro) and Cerro Yapacana (Department Atabapo) in the state of Amazonas (Venezuela).

It was in bloom in January and December.

Paradrymonia hamata resembles P. ciliosa by its general appearance: thick stem with usually short in-

ternodes, creeping on rocks or climbing lower tree trunks, large leaves with long petioles, and very short

inflorescences in the leaf axils. It differs from P. ciliosa by the leaf blade ovate to orbicular, pedicels up to

4.5 cm long, versus up to 2.5 cm, the sepals linear-acuminate in the apical half, the corolla tube 4-4.5 cm

long, versus 2-3 cm, and a much smaller basal gibbosity. Paradrymonia hamata can be separated from P.

lutea by its white coralla versus yellow, from P. tepui and P. yatua by its calyx lobes partly linear acuminate

versus elliptic, from P. glandulosa by its white corolla trichomes versus red, and from the other species of

subg. Paradrymonia from Venezuelan Guayana by its leaves broadly ovate to orbicular versus oblanceolate

to elliptic. This species was “sp. B" in Feuillet & Steyermark (1999).

— epithet hamata refers to the barb-like trichomes on the linear tip of the sepals.

PARATY UELA. Amazonas: Depto. Atabapo: Cerro Yapacana, Rio Orinoco, 1200 m, 2 Jan 1951, fl., B. Maguire, R.S. Cowan &

Ja ui den m ).

3. Paradrymonia lutea Feuillet, sp. nov. (Fig. 3). Tree: VENEZUELA. Amazonas. Depto. Rio Negro: Neblina Massif, Canyon

Grande, along Río Mawarinuma, ca. 7 km ENE of Puerto Chimo, 0°50—51'N, 66°02-06'W, 300 m, 9-14 Jul 1984, fl., G. Davidse &

J.S. Miller 27212 (notorvpe: US; isorYPES: MO, NY, VEN n.v.).

11 A E 1 1.5 1 Ps a E ie

P 1 i ili ffini iolo | i , corolla lutea,

ELL O

Herbaceous lithophyte, or low epiphyte. Stem densely pilose, trichomes reddish. Leaves opposite, very

unequal in a pair, the small one scale-like, mostly early deciduous; the larger one: petiole purple, appressed

[| [| Pal n.a 4 In Rog Et au ul af)

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pubescent to glabrous, 15-38 cm long; blade broadly ovate to ovate elliptic, 13—32 x 6-21 cm, cuneate and

shortly decurrent at base, abruptly acuminate at apex, margins obscurely and loosely glandular serrate,

glabrous, sometimes the veins sparsely appressed-pubescent abaxially, margin sparsely ciliate, with 9—13

main veins on each side of the midrib, impressed adaxially, conspicuous abaxially, tertiary venation not

evident. Inflorescences axillary, fasciculate or short pedunculate, 12-25-flowered, peduncle 3—5 mm long,

appressed-pubescent, bracts oblanceolate or lance-elliptic, acute at apex, 10-12 x 1.53 mm, minutely

appressed-pubescent, pedicels densely appressed-pubescent, 2-3 cm long. Flower: calyx lobes free, 15-25

x 3-6 mm, narrowly to broadly lanceolate or lance-elliptic, narrowed to an acuminate attenuate apex, en-

dured at the tip, sparsely ciliolate in the upper half, entire in the basal half, with a few endured blunt teeth

toward the apex; corolla bright yellow, tubular, 4.3—4.5 cm long, glabrous inside except for the verrucose

throat, pilosulous outside except toward the base, basal gibbosity 3.5 x 5 mm, ovate-oblong, glabrous, base

narrowed, broader above to 7-8 mm diam., lobes suborbicular, 6 x 7 mm, glabrous inside, ventral lobe

appressed-pilose outside, the others appressed-pilose toward base, otherwise glabrous outside; stamens

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4, inserted on the corolla, anthers 2 x 2 mm, cells pilose at base; disc reduced to a dorsal gland, glabrous;

ovary densely sericeous, style 25 mm long, glabrous at base, sparsely pilose toward apex, stigma verrucose.

Fruits not seen.

Distribution —Paradrymonia lutea grows on boulders in forest or as a low epiphyte at 150-1250 m on

the Neblina Massif, Amazonas (Venezuela).

I have not seen a single corolla and rely on careful notes taken by Julian Steyermark in the description

of the corolla and androecium. dci Cid ne resembles P. ciliosa from which it differs mainly by the

bright yellow (vs. white corollas), the | sepals oblanceolate or lance-elliptic (vs. narrow triangular),

and the leaf blades that are broadly ovate to sue elliptic (vs. oblanceolate) and cuneate (vs. long attenu-

ate) at base. The bright yellow corollas separate P. lutea from the other species of subg. Paradrymonia from

Venezuelan Guayana. This species was “sp. A” in Feuillet & Steyermark (1999).

Md c ane lutea refers to the bright yellow corolla.

PARATYPES: VENEZUEI

camp, 0°50'N, 66°07'W, 150—500 m, 14 Feb 1984, fL, R.L. Liesner 15896 (MO, US, VEN): C

0°49'N, 66°0'W, 1250 m, 21-24 Mar 1984, fl., R.L. Liesner & B. Stannard 16895 (MO, US, VEN).

Je as Kal]: f, +1 571 [c +1 1 WT +l 1 D 4 .

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4. iie m tepui Feuillet, e nov. (Fig. 4). Tire: VENEZUELA. AMAZONAS Depto. Rio Negro: Cerro Aracamuni, sum-

; Camp, in ravines and near edge of tepui, 01?32'N, 65°49'W, 1400 m, 31 Oct 1987, fl. & fr., RL. Liesner & G. Carnevali

22679 DES US; ISOTYPE: NO.

D 4 ; “J. a | q $ Ilat Tots 1 1] e J 1 1;

I r r

Terrestrial herb, lithophyte, or low epiphyte. Stem densely pilose, trichomes reddish. Leaves opposite, very

unequal in a pair, the small one scale-like, mostly persistent; the | one: petiole 14—23 cm long, glabrescent;

blade obovate to broadly elliptic, 14-30 x 6.5—13, cuneate and brielly decurrent at base, obtuse to acuminate

at apex, margin minutely and loosely serrate, glabrous except abaxially on veins, margin ciliate, 7-10 main

veins on each side of the midrib, midrib raised on both surfaces, veins raised or impressed on both surfaces.

Inflorescences axillary, densely fasciculate on short peduncle; peduncles 1-1.5 cm long; bracts similar to

the sepals; pedicels sparsely appressed pilose. Flowers: sepals greenish tan tinged with red, oblanceolate to

lance-elliptic, narrow-triangular acuminate at apex, 1.7 x 0.4 cm, sparsely pilose, few glandular teeth limited

to the acumen; corolla (seen only in bud) white, hyaline-pilose, basal gibbosity 5 x 3 mm, tube 2 cm long,

edge of barely opening lobe undulate. Fruit subglobose, greenish tan, tinged with red.

Distribution.—Paradrymonia tepui has been collected on slopes and summit of a tepui, Cerro Araca-

muni, in the Dept. Río Negro (Amazonas, Venezuela) between 600 and 1400 m. It was in flower and fruit

in October.

Paradrymonia tepui resembles P. ciliosa by its general appearance: thick stem with usually short inter-

mds ee on the substrate, strongly unequal leaves in a pair, the larger ones with long petioles, short

les in the leaf axils, and creamy white corollas. Paradrymonia tepui differs from P. ciliosa

by the leaf Bisa ovate to broad-elliptic, briefly decurrent at base, the inflorescence densely multiflowered,

and the sepals linear-acuminate. The shape of the leaves and the sepals oblanceolate to lance-elliptic and

narrow-triangular acuminate separate P. tepui from other species of subg. Paradrymonia from Venezuelan

Guayana. This species was “sp. D” in Feuillet & Steyermark (1999).

Etymology.—This species is named tepui, a noun in apposition, after the location of the three known

collections at the summit or on the slopes of a tepui.

Iconography.—See Fig. 493 in Feuillet and Steyermark (1999).

PARATYPES: VENEZUELA. Amazonas: Depto. Rio Negro: Cerro Aracamuni summit, Proa Camp, near edge of tepui, 1°32'N, 65°49'W,

1400 m, 27 Oct 1987, calices, R.L. Liesner &G. C 22530 (MO, US); Dept. Rio Negro, slopes of Cerro Aracamuni, Q

Laja area, 1?24'N, 65°38'W, 600 m, 22 Oct 1987, fr., R.L. Liesner & G. Carnevali 22324 (MO, US).

5. Paradrymonia yatua Feuillet, sp. nov. (Fig. 5). Te: VENEZUELA. Amazonas. Depto. Río Negro: Rio Yatúa, at base of

Piedra Arauicaua, 100—140 m, 3 Feb 1954, fl., B. Maguire, J.J. Wurdack & G.S. Bunting 37466 (HoLoTYPE: US; isoryee: NY).

Feuillet, Fi

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Herb saxicolous. Stem short pilose, trichomes hyaline. Leaves very unequal in a pair, the small one scale-

like, often early deciduous; the larger one: petiole 13-30 cm long, 1 cm wide (life-measurement), glabrous;

blade broadly obovate to elliptic or lanceolate, cuneate and then long decurrent into the petiole, acuminate

at apex, margin entire or serrulate toward apex, glabrous both sides, 40-55 x 13-18 cm, midrib strikingly

raised below, 12-13 main veins on each side of the midrib. Inflorescence A dense ia fasciculate; bracts

similar to the sepals; pedicels 1.5—3 cm, sparsely pilose. Flowers: sepals green lance-elliptic

aa owe y

long linear at apex, 2.5-2.8 x 0.9-1.1 cm, margin entire loosely ciliate, Pd pilose; corolla creamy white,

6-6.5 mm long, basal gibbosity 8 x 3 mm, curved, tube 2-2.8 cm long, pilose outside with hyaline trichomes,

lobes suborbicular, entire, 10 x 10 mm, undulate at margin, pilose at base outside with hyaline trichomes,

otherwise glabrous, dorsal gland, entire, rounded, 2 mm long, 1 mm wide; anthers reniform-suborbicular;

ovary slender ovoid, densely short strigose, trichomes purple, style 2.5 cm long, conspicuously short hirsute,

trichomes dark. Fruit not seen.

Distribution —Paradrymonia yatua has been collected at the base of Cerro Arauicaua, near Rio Yatúa

(Amazonas, Venezuela), between 100 and 150 m elevation.

Paradrymonia yatua resembles P. ciliosa by its general appearance: thick stem with usually short inter-

podes FIERA on s substrate, strongly unequal leaves in a pair, the si ones dn long petioles, short

the leaf axils, and creamy whit corollas. P differs from P. ciliosa

by the leaf blade broadly obovate to elliptic or lanceolate (rather than cblanbedlate) the sepals oblanceolate

to lance-elliptic, long linear at apex (rather than linear-acuminate), the corolla lobes undulate at margin

(rather than the ventral lobe and two dorsal lobes fimbriate). ac yatua ane from P. lutea by its

white corolla versus yellow, from P. tepui by its le te tol olate and its green

calyx versus tinged with red, and from the other species of subg. Bxadrymonia from Venezuelan Guayana

by its calyx lobes lacking a linear apex.

Etymology.—This species is named yatúa, noun in apposition, after the Rio Yatúa near which the known

specimens have been collected.

Pararvres: VENEZUELA. Amazonas: Depto. Rio Negro: Cerro Arauicaua, al pie y en las faldas inferiores, Rio Yatúa, 1°35'N, 66?10"W,

125-150 m, 11 Apr 1970, fl., J.A. Steyermark & G.S. Bunting 102546 (MO, NY, NY, US, VEN)

KEY TO THE SPECIES OF PARADRYMONIA SUBG. PARADRYMONIA IN THE GUIANA SHIELD AREA

EXCEPT P. CILIOSA, ALL RESTRICTED TO VENEZUELA, AMAZONAS)

1. Corolla bright yellow (Neblina Massif) Paradrymonia lutea

1. Corolla white.

2. a lobes elliptic.

eaf blades ovate or broad-elliptic; calyx tinged with red (Cerro Aracamuni) Paradrymonia tepui

i pe blades oblanceolate; calyx green (Río Yatua Paradrymonia yatua

2. Calyx lobes partly linear-acuminate.

4. Corolla white with red trichomes (Cerro Marhuaca & Cerro Yapacana) Paradrymonia glandulosa

4. Corolla white with nyaline t Teno:

. Leaf blades ovate to orbicular, sud tl le; calyx lo! =

filiform; estalla wenittal lobe fimbriate, there crenate to entire at margin (Río Yatúa & Cer

Yapacana) Par dcos hamata

. Leaf blades oblanceolate to elliptic, progressively long decurrent to the petiole; corolla 3 lobes

fimbriate.

6. Calyx lobes margin loosely glandular toothed, acumen short or long linear; corolla with the

ventral and dorsal lobes fimbriate (large distribution in Central & northern South America

Paradrymonia ciliosa

6. Calyx lobes margin (including the long linear acumen) loosely fimbriate; corolla with only the

ventral lobe fimbriate (Cerro Cuao) Paradrymonia sp. 1

592 J i h Institute of Texas 3(2)

ACKNOWLEDGMENTS

This work could not have been completed without the help of the curators of the herbaria MO, NY, and

VEN who made available to me the material in their care. Alain Chautems (G), Raul Gutierrez, John Clark

(UNA) and Marcela Mora (UNA) provided helpful reviews. This paper is published as number 146 in the

Smithsonian’s Biological Diversity of the Guiana Shield Program publication series.

REFERENCES

I P» . Iu tet ME ri REY aa hrala O

FeuiLLET, C. 2009. Folia taxonomica 12. Paradrym

a new species from Amazonas, and disuibution and floral mbipholoay of P naculata J. Bot. Res. Inst. oe

3:133-138

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

FOLIA TAXONOMICA 16. DILKEA (PASSIFLORACEAE) 1. EPKIA,

A NEW SUBGENUS AND FIVE NEW SPECIES

FROM WESTERN AMAZONIA AND THE GUIANAS

Christian Feuillet

Department of Botany, MRC-166

Smithsonian Institution

PO. Box 37012

Washington, DC 20013-7012, U.S.A.

lec

feuillec@si.edu

ABSTRACT

An introduction to Dilkea (Passifloraceae) is given and the new ae Ep à a new P from western Amazonia are

desemiped: Dilkea a D. iir us D. tillettii; as well as two new st Dilkea lecta and D. vanessae.

1 1 f: : ] f. +l 1 Jl 1 £ d +l 1 " A leew

D E +

to the subgenera af Dilkea: is also given.

RESUME

IT S ] "E a Tull D ET ^ t5 t | EnL: tt 1 1] JA am ta]

décrits: pa we D. ne et D. ‘tillettii: ainsi que deuxe DRE des e anes: Dilkea piis et D. vanessae. Les espéces du

Une clé des

sous-p y F eo E 5

sous-genres de Dilkea est aussi présentée.

RESUMEN

] Ced ^ T^: Tl (D $] ^ J 21 1 1 Cal : ] 1 Ama

J de las Gui Dilkea lecta y D. vanessae.

zonia occidental: Dilkea cuneata, D. ovalis y D. tillettii;

l subgénero Ep! i | lo tras las escamas de la vema más largo que los siguientes. Se

r E E

: 1 1

1 1 A da Py

INTRODUCTION

Dilkea was named by Masters (1871) to honor his late friend Charles Wentworth Dilke, 11 (1810-1869), made

baronet by Queen Victoria in 1862. Dilke was a patron of arts and sciences. He died in St Petersburg while

representing England at the horticultural exhibition held there. His father, C.W. Dilke, I (1789-1864), was

a founder of the Gardener's Chronicle.

Dilhea, with 7 Species, isa genus du tribe Passifloreae. The species can easily be separated from the other

American genera of the tri] tl j t ite: flowers with 4 sepals, 4 petals, 8 stamens,

: carpels um e stamens inserted at the bottom of the tube or halfway up the short, 0-2 mm long an-

ither flatt ted (Masters 1872; Killip 1938; Holm-Nielsen

et al. 1988; Tillett 2003; Feuillet & MacDougal 2007). Dilkea has a large lowland distribution in Panama

and South America, from Colombia to Mato Grosso and from Peru to Amapá, but has been little collected.

Dilkea Mast., Trans. Linn. Soc. 27:627. 1871. Te species: D. retusa Mast., Trans. Linn. Soc. 27:628. 1871. (for typification,

see Killip 1938)

Shrubs, small trees, or lianas. Leaves always simple and unlobed, margin entire; perio with a basal pulvi-

nus. Flowers white; sepals 4; petals 4; operculum and limen lacking; and shorter than wide or

lacking; stamens 8, inserted on the bottom of the floral cup or on a 1-2 mm ileng androgynophore; ovary

sessile or usually on a short 1-2 mm long gynophore, styles 4, united at base. Fruits yellow to orange, bac-

cate, ovoid or globose, apically rounded or with a conical acumen, pericarp coriaceous, placentas 4, seeds

J. Bot. Res. Inst. Texas 3(2): 593 — 604 . 2009

Lo £al n L in LI rr ET 22

594 Jour t t lexas 3(2)

LP |

sclerified

few, slightly bean-shaped or straight, not flattened, both ends similar, and sarcotesta thin,

nor ornamented.

There is a great similarity in floral morphology within Dilkea; thus, one cannot separate species in the

genus based on floral characteristics alone. Killip (1938) described Dilkea parviflora and commented: "I am

inclined to think that further SR and collecting will show that the four last species of the present treat-

ment should be merged in one.” Part of the problem, as suggested by Killip, resided in the small number of

field collections, but instead of getting a sy g SI a result of better sampling, it appears that

increased sampling is instead MS new eds

Historically, nine species have been described within Dilkea. Masters (1871) published D. acuminata

and D. retusa from Spruce’s collections as new species within this new genus. Later, Masters (1872) de-

scribed D. wallisii Mast., only after an illustration by Wallis, in his treatment of the Passifloraceae for Flora

Brasiliensis. Few taxa have been published since then: D. johannesii by Barbosa Rodrigues (1888), D. ulei by

Harms (1906), D. glaziovii (nom. nud.) by Glaziou (1909), D. johannesii var. parvifolia by Hoehne (1915), D.

parviflora by Killip (1938), D. magnifica by Steyermark (1968), and D. margaritae by Cervi (1991). For more

than 70 years, D. ulei has been treated as a synonym of D. johannesii. Dilkea glaziovii Mast. ex Glaz. (nom.

nud.) was ignored, as Mitostemma glaziovii Mast. was considered to be in the right genus. Thus, at present

seven species and one variety are circumscribed within Dilkea.

Most recently, Holm-Nielsen et al. (1988) placed D. acuminata, D. magnifica, and D. wallisii in synonymy

of D. retusa, keeping D. johannesii and D. parviflora as separate species on the basis of petiole, pedicel, and

gynophore sizes. They had not seen the type of D. parviflora and therefore based their identification on Kil-

lip’s description. In fact, the material cited from Ecuador and the illustration mostly belong to a new species

described below as D. cunvena,

Recent gifts of Dilkea tions to the Smithsonian Institution herbarium (US) revealed an unexpected

diversity of vegetat hology from the Loreto region in Peru and from the Guianas. This was confirmed

in February 20 09 bya visit to the deed collections at the Missouri Botanical Garden, especially rich in

Dilkea from western Amazonia. S woody vines that exhibit ti dial growth

and climb by the mean of strong els apically 3-fid but early deciduous if they do not Cad a support.

They represent what is considered here to be Dilkea subg. Dilkea. Another group of species, including the

five new ies described below, have mM flower and fruit morphology, but are small trees and exhibit a

unique “modular” growtl tl that the P Tl t | ( gh modular grow th;

the modules have at the rin very short internodes with bud scales, then a long internode, and apically a

swelling with short internodes, leaves, and inflorescences and ramifications. In most cases, the terminal

meristem aborts and the growth continues through 1(-3) subterminal vegetative buds. This results in the

overall appearance of a treelet with one stem and tiers of leaves, evidence of rhythmic or modular growth

(field studies are needed to ascertain this) and sometimes a few modular branches 1-2 meters above ground.

The plants are self standing and do not have tendrils.

In the text below, the world "lacking" means *not present in the taxon" and the expression "not seen"

means “not present in the collections studied."

Subgenus Epkia

Dilkea subg. Epkia Feuillet, subg. nov. Tee: Dilkea cuneata Feuillet (see below).

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itecnodus brevissimis distimetus:

The species of subgenus Epkia are mostly small trees (or shrubs or scandent shrubs). In each branch, the first

node above the numerous acicular bud scales is much longer than the following internodes and there is no

transition in leat shape año id Toe de eus become shorter in the branches of higher rami-

fication order. The | a ter on the club-shaped apical part of every growth

unit. The flowers are mula? to those of the other species of Dilkea by their color and general morphology.

f Dill 595

Feuillet, ü

Collection labels usually say “tree” or “small tree" with noted sizes from «1—8 m tall, but are occasion-

ally labeled as “shrub.” I think this is indicative of a small size rather than a clumping habit of the plants.

From pictures taken in the wild, the young plants 0.5-1.5 m tall have a main stem about 1 cm in diameter,

showing many leaves in a few tiers. A 1.5-m tall plant of Dilkea cuneata photographed in Peru has 4 tiers

of 7-20 large leaves. The lower is at ground level and the upper one is at the apex of the plant. The picture

shows the whole plant and it is not possible to decide if the stem has a monopodial and rhythmic growth or

a sympodial, modular growth. The branches, as demonstrated by herbarium specimens, are modular and

little branched constructions. Each module exhibits a unique morphology. At the base are usually more

than 10 acicular bud scales followed by a long internode and the apical part with short internodes bearing

the leaves and axillary flowers or short inflorescences.

The name of subgenus Epkia honors through his initials Ellsworth Paine Killip (1890-1968) author of

The American Species of Passifloraceae (1938), a two-volume masterpiece covering 366 species.

KEY TO THE SUBGENERA OF DILKEA

dde Or Scanaent shrubs; tendrils us at agen first internode shorter than or as long as the e

| at t] BUE hypanthium. Dilkea subg. Dilkea

. Small trees or shrubs o t shrubs; without tendrils; first internode much longer than the following

internodes; stamens E on a 1-2 mm m long androgynophore. Dilkea subg. Epkia

New species

Three new species from western Amazonia and two from the Guianas are here described in subgenus Epkia.

More species of Dilkea are still undescribed.

1. Dilkea cuneata Feuillet, sp. nov. (Fig. 1). Tre: PERU. Junin: Cahuapanas, on Rio Pichis, 340 m, 30 Jul 1929, fr., E.P Killip

& A.C. Smith 26730 (HOLOTYPE: US).

= Dilkea parviflora sensu Holm-Nielsen, Jorgensen & Lawesson 1988, in part; not Killip 1938.

LI ] 1 g E] 1-2 I . Det) 1 p h 1-11 :11 as Sii: T . Els 25-50 x 10-15 cm, anguste oblanceo-

lata, e 1. ‘oye -_ r x ` "ES 1 . 31 : 1] a dx 1 ] An x

Treelets 1—5 m tall, monocaulous or branched. Plant glal tl t. Stems with a long internode 6-20

cm long (2.5 cm in Gentry et al. 54258), 0.4—0.6 mm thick, follwedl by short ones 0.2-0.5 cm long. Tendrils

lacking. Stipules not seen. Leaves in terminal clusters; petiole reduced to the pulvinus 0.8-1.5 cm; blade

narrowly cuneate to cuneate, 28-53 x 10—13.5 cm, short to long attenuate and then abruptly ending at base,

apex acute to rounded and acuminate, acumen 5-15 mm long, margin entire, drying olive-green adaxially,

yellowish brown abaxially, midrib and veins raised on both surfaces and yellowish when dry, 12-25 main

veins on each side of the midrib. Inflorescences ous a about 5 mm long, elongating 7-10

mm under the fruit; bracts not seen. Fl ; hypanthium 0.5-1.8 cm long, funnel-shaped, 0.4—0.8

cm diam. at throat; sepals 4, white, oblong, 2.5-3. 5 x 0. 7-1 cm, obtuse to acute at apex, ciliate at margin;

petals 4, white, oblong, 2.5-3.5 x 0.5-0.7 cm, obtuse to acute at apex, united at base, thinner than sepals;

corona white, in 2 rows, outer one of free straight filaments, 13 cm long, seldom curled at apex, inner one

membranous, laciniate at margin, laciniae floccose; androgynophore 1-2 mm long; stamens 8, filaments

white, ca. 3 cm long, united at base for 1 mm, anthers sulphur yellow, 8 mm long, dorsifixed near base;

gynophore 1-2 mm long; ovary ovoid, 5-6 mm long, styles 4, ca. 3.5 cm long, united for 1/3-1/2 of their

length, stigmas capitate, sulphur yellow. Fruits spherical or slightly ovoid, apiculate, 2.5—3 cm long, yellow,

pericarp coriaceous, about 0.5 mm thick, ees iip few, in A 7 5-1.7 x 0.7-0.8 cm, brown.

sti ma from Ecuador (Morona-

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Santiago) and Peru into, Loreto, and Pasco), in primary forest on ae firme, at 100—700 m of elevation.

It is documented blooming in February, June, July, and November, and fruiting in January to April, June,

July, and ds

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fulgens Hook. F. (including by Killip in 1931), Clavija lehmannii Mez, Clavij igii Mez, to Passiflora L., and

even as a member of the Ranales. Some specimens have been identified as s Dilkea parviflora (Holm-Nielsen

et al. 1988) and Dilkea wallisii (Killip 1938) although those two Dilkea species are lianas. Nevertheless, D.

cuneata clearly represents a new species from western Amazonia. Interestingly, D. cuneata has some of the

largest leaves and of the smallest flowers in the genus The growth units have a 15-25 cm long internode

between the bud scales and the] lall tl led toward the apex with internodes shorter

than or equal to twice the diameter of the petioles. The congested inflorescence at the end of a branch is not

terminal, but in fact consists of 2 or 3 axillary, subsessile, pauciflowered inflorescences. Aestivation of the

two outer sepals is valvate, this pair surrounding the two inner sepals

Etymology.—The epithet “cuneata” refers to the leaf blades basal half that are long triangular.

PARATYPES: ECUADOR. Morona-Santiago: Taisha, 500 m, 14 Feb 1962, fl. & fr., P.C.D. Cazalet & T.D. Pennington 7771 (B, K, NY, US);

8-10 km NNW of military camp, 2?21'5, 77731 W, 650-700 m, 16 Jun 1980, fr., J. Brandbyge & E. Asanza 31937 (AAU). PERU. Loreto:

Biological Station Río Blanco, 4?20'S, 72%45'W, 150 m, 12 Sep 1985, fr., R. Vasquez, J. Ruiz & N. Jaramillo 6714 (MO, MYF); Maynas, Las

Amazonas, Explornapo Camp, 3?20'S, 72?55"W, 100—140 m, 3 Mar 1991, fr., JJ. Pipoly, R. Vásquez, N. Jaramillo, C. Grández, J. Ruiz &

R. Ortiz 14165 (MO); Maynas, Las Amazonas, Explornapo Camp, 3°15’S, 72%54'W, 140 m, 20 Nov 1991, bud, R. Vásquez 17568 (MO);

Maynas, Las Amazonas, Quebrada Sucusari, Explornapo Camp, 3?20'S, 72°55’W, 140 m, 17 Apr 1991, fr., R. Vásquez & N. Jaramillo

16095 (MO); Maynas, Iquitos, Alpahuayo, 73°30’W, 4°10’S, 150 m, 6 Jun 1985, fl., R. Vásquez, J. Ruiz & N. Jaramillo 6608 (HUA, MO,

MYF, US); Maynas, Iquitos, Alpahuayo, 73°30’W, 4?10'S, pu m, 24 Mar 1992, st., R. Vasquez, S.A. es eae & N. duum 18135 (MO);

Maynas, Iquitos, Santa Maria de Ojeal, 10 km below ío Nanay, 16 Jun 1976, fr., S. McDaniel & M 5 (MO); May-

nas, Yanamono, Río Amazonas, 3°25’S 72°50’W, 130 m, 12 Jun 1986, fr., A.H. Gentry, R. Vásquez & N. Jaramillo 54258 (MO); vicinity of

Requena, Fundo Canama, E of Río Ucayali, 17 Jul 1961, bud, M.E. Mathias & D. Taylor 5552 (F, LA, MO); Maynas, Las Amazonas, Río

Sucusari, 3?20'S 72?55'W, 116 m, 7 Nov 1989, fl., R. Vásquez & N. Jaramillo 13077 (MO, MYF, P). Pasco: Oxapampa, Palcazu Valley,

Iscozacin, 75°15’W, 10?12'S, 400 m, 26 Jan 1984, st., R.B. Foster 9545 (M

2. Dilkea lecta Feuillet, sp. nov. (Fig. 2). Tree: FRENCH GUIANA: near Organabo, primary forest on white sand, 20 Jun 1995,

fr., D. Loubry 2422 (uororyre: US; rsorvees: CAY, MPU)

=? Dilkea johannesii var. parvifolia Hoehne, Comm. Linh. Tel. Matto Grosso ao Amazonas, Annexo 5, Bot. pt. 5:73; pl. 111. 1915. Tyre:

IL. Maro Grosso: Rio Juruena, Comm. Linh. Tel. Matto Grosso Expedition 5433 (SP).

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Treelets with | I ti ive climbers, up to 3 m tall. Whole plant glabrous. Stems with a long

internode 1.3-5 cm long, isliowed by short ones about as long as once their diameter or less. Vegetative

bud ca. 2 mm above the leaf axils, is scales ace to narrow-triangular 3-5 mm long. Tendrils lacking.

Stipules not seen. Leaves in termina ole usually reduced to tł lvinus, occasionally up to 0.6

cm long, glands not seen; blade narrow okence cuneate, 8—17 x 2s. cm, long attenuate, apex acute,

long acuminate, margin entire, midrib and veins raised on both surfaces and yellowish when dry, 8-12

main veins on each side of the midrib. Inflorescences subterminal; pedicels not seen. Flowers not seen (see

below); gynophore 1-2 mm long under the fruit; styles 4, united only at base. Fruits spherical, apiculate,

33.5 x 2.53 cm, color not documented, pericarp coriaceous, about 0.5 mm thick; developed seeds not

seen, ovules flat, 2 per placenta.

Distribution and nid diss lecta is known from the Lely Mountains in eastern Surinam and from

central and nort] Guiana, and Brazil (Matto Grosso). The ecology is different between the four

collections. Loubry 2422 was collected in primary forest on white sand, on the coastal plain, at low elevation

(according to the map the whole area is below 50 m); Larpin 1058 is from low forest island on the northern

slope of a granite outcrop (inselberg) at 340 m; Mori & Bolten 8541 was collected between 2 plateaus of a

mountain range between 500 and 700 m, and Granville & Crozier 13648 is from middle to high ridge forests

on schist, quartzite, and conglomerate at 400 m. The two localities in French Guiana share a low amount of

available water due to the texture of the soil, white sand without visible amount of humus causing excellent

and fast drainage and maximum light reflection toward leaf undersurface in one case and the dark granite

slope aca water enue and favoring evaporation.

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larity so characteristic of subg. Epkia. While this branching pattern is less obvious than in the other species

of subgenus Epkia, it is still visible. Plants of Dilkea lecta are shrubs likely with one main trunk (Granville &

Crozier 13648 and Loubry 2422), yet able to passively climb on small trees (Larpin 1058 and Mori & Bolten

8541).

Hoehne (1915) gave a brief description and a diagnostic illustration of Dilkea johannesii var. parvifolia.

This is most likely a synonym of D. lecta. Based on Hoechne’s description one can infer the following: leaf

blades between veins, with small red dots underneath, pedicels slightly more than 1 cm long, flower

t base), sepals and petals of similar length, corona in 2 rows, at least

with hypanthium slightly asy

the inner one inserted low in den hypanthium, outer row of filaments MR Henny saan (more so than

the stigmas), inner t base, flamentos t apex, filame g t basal 1/3 and

upper 2/3 papillose; stamens shorter than the petals, inserted on a short and short gynophore;

ovary wide elliptic, styles united for 2/3 of their length (unlike Mori & Bolten 8541), ee spherical. Thus,

based on these features, this species fits within my current circumscription of D. lecta.

Etymology.—The specific epithet “lecta,” Latin for gathered, refers to the leaves grouped at the apex of

the branches.

PARATYPES: SURINAM: Wl Mountains, 175 km SSE of Paramaribo, 500—700 m, 19 Oct 1976, fr., S.A. Mori & A. Bolten 8541 (NY, US).

er 42"W, 340 m, 20 Apr 1992, st., D. Larpin 1058 (CAY, US); Montagne

de la Trinité, zone sud, Mana River basin, 4234 N, 53°27 W, 400 m, 16 Jan 1998, fr., J.-J. de Granville & F. Crozier 13648 (CAY).

3. Dilkea ovalis Feuillet, sp. nov. (Fig. 3). Tre: PERU. Loreto: Maynas, Las Amazonas, 72%33'W, 3925'S, 120-130 m, 25 Mar

1991, fr., C. Grandes, G. Criollo & W. Criollo 2291 (HoLotyPE: MO).

T J] Ent: Tanta tel lliptica, lata, 10-18.5 x 5-9 cm differt.

Shrub or small tree, 4-6 m tall; plant glabrous throughout. Stems with a long internode 3.5-12 cm long,

followed by short ones 0.1-0.3 cm long. Tendrils lacking. Stipules not seen. Leaves in terminal clusters;

petiole 0.5—2.3 cm long, pulvinate for less than half its length, glands not seen; lamina elliptic to slightly

obovate, 7.5-25 x 3.5-10 cm, widely acute and then short cuneate at base, apex rounded with acumen

0.6-1.2 cm long, margin entire, minutely frilled when dry, midrib and veins raised on both surfaces, 6-12

main veins each side of the midrib. Inflorescences subterminal; pedicels under fruit 7-9 mm long; bracts

not seen. Flowers not seen; gynophore under fruit about 1 mm long. Fruits spherical or slightly ovoid, api-

culate, about 2.5 cm diam., yellow, pericarp coriaceous, about 0.5 mm thick, glabrous; seeds few, elongate,

1.8—2 x 0.8—0.9 cm, brown.

Distribution and ecology.—The three known collections have been made in northeastern Peru, Region

of Loreto, Province of Maynas, at 140—180 m. It was in flower buds in July and fruiting in March.

Dilkea ovalis is easily set apart from other species in subgenus Epkia by leaf shape. In D. ovalis, the

laminas are wide elliptic rather than cuneate or narrow elliptic. The pulvinus is a third to half the petiole

total length. In the other long-petioled species, D. vanessae, the pulvinus represents less than the fifth of the

total length.

Etymology.—This species is named ovalis because of the oval shape of its leaf blades.

PARATYPE: PERU. Loreto: Maynas, Iquitos, Allpahuayo—IIAP, 73?25'47"W, 3%57'19”S, 140 m, 26 Apr 1997, st., R. Vásquez, O. Phillips,

R. Rojas & A. Peña 23599 (MO); Maynas, Iquitos, Allpahuayo-IIAP, 73?30"W, 4°10°S, 150-180 m, 8 Jul 1991, fl. Buds, R. Vásquez & N.

Jaramillo 17188 (MO).

4. Dilkea tillettii Feuillet, sp. nov. (Fig. 4) Tree: PERU. Loreto: Florida, Río Putumayo at mouth of Río Zubineta, ca. 200 m,

Mar-Apr 1931, fl., G. Klug 2100 (HOLOTYPE: 1 O, S)

[Dill lifolia] Killip ex Tillett, invalid: in sched., handwriting un! 1986 on Gentry et al. 37992 (F) “Tillet” and Tilletts hand-

p 1993 on C. Klug 2100 (MO) "ined. AN

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node, the only partially visible >12.5 cm long (Gentry et al. 37992 (F)), followed by short ones 0.2-0.5 cm

long. Tendrils lacking. Stipules not seen. Leaves in terminal clusters; petiole 1.5-3 cm long, usually reduced

to the pulvinus, glands not seen; lamina long oblanceolate, 35-43 x 12-15 cm, basal half long triangular

regularly narrowing to the short pulvinate petiole, rounded at apex, blunt or with an acumen up to 1.5 cm

long, margin entire, midrib and veins raised on both surfaces a drying more or leos = same color as

the rest of the leaves, 8-12 main veins each side of the midrib. I 1; pedicels 0.5 cm

long, 1 cm under the fruit, joined 0.2-0.3 cm from the base; pracls not seen. Flowers white, with a scent of

lilac (Barrier 84); hypanthium 17-19 mm long, perianth sey g and hypanthium getting shorter during

late anthesis; sepals and petals 20-35 mm long, about 5 mm wide, two outer sepals green outside, white

inside, aestivation valvate, two inner sepals white on both surfaces, aestivation valvate; corona white in two

rows 3.5-4 cm long, inserted near base of hypanthium, tubular at base, outer row filaments slightly wavy,

inner series heavily branched in the uppermost 1 cm, lateral segments very thin and tightly curled; stamens

inserted in the bottom on the hypanthium, filaments about 3 cm long, anther 11x 1 mm, apiculate, base 2

mm sagitate; gynophore about 2 mm long; ovary 5 mm long, tapering at base, wider at apex, styles 3.5-4

cm long, stigma capitate. Fruits globose, 2.5-3 cm diam., apiculate, yellow, pericarp coriaceous, about 0.5

mm thick, glabrous; seeds few, elongate, 1.8 x 0.8 cm, brown.

Distribution and ecology —Dilkea tillettii is known from Colombia (Amazonas & Caqueta), Ecuador

(Morona-Santiago, Napo & Pastaza), and Peru (Loreto & Pasco) between 120—700 m. It was blooming in

January and March—April, and fruiting in June to September and November.

Dilkea tillettii has obovate leaves that are cuneate in the basal half unlike D. lecta and D. ovalis. It is

distinct from the other large-leaved species, D. cuneata, by the shape of the petioles and the shape of the

leaf base. Although the pulvinus constitutes more than half the length of the petiole, it does not always

encompass the entire petiole. The base of the lamina is the continuation of the narrow triangular leaf shape,

neither suddenly ending as in D. cuneata nor decurrent. Dilkea lecta also has petioles more or less reduced

to the pulvinus, but its laminas are at most half the size of those in D. tillettii.

Etymology.—Dilkea tillettii is named to honor Steve Tillett, botanist in Caracas (at MYF), and a specialist

on the Passifloraceae who MAE this was a new species.

P COLOMBIA. A A National Park, Quebrada de A dre, 3?47'S, 70?15"W, 200-220 m, 15 Nov

1991, fr., J.J. Pipoly 16039 (MO). Caqueta: LIEU S troche a Yari, 0?25'S 72220" W, 200 m, 23 Jan 1989, st., A.H. Gentry & M. Sanchez

64981 (MO). ECUADOR. Morona-Santiago: Taisha, 8-10 km NNW of military camp, 2°21’S, 77*31'W, 650—700 m, 16 Jun 1980, fr.,

J. Brandbyge & E. Asanza 31937 (AAU); Napo: along Rd. between Coca & Río Tiguino, 76°52’W, 1°10’S, 300 m, 1 Mar 1992, fr., T.B.

Croat 72576 (MO); 77°45’W, 0°59’S, 200—300 m, 12 Apr 1997, fr., K. Wide M. Bass, G. Villa & C. Vriesendorp 2841 (MO, QCA),

Yasuni National Park, 0°55’S, 76?11W, 200 m, 26 May-8 Jun 1988, fr., C. Cerón & F. Hurtado 4258 (MO). Pastaza: Lorocachi, 3 km S

of the military camp, 1°38’S, 75*58"W, 200 m, 23 May pini: i XJ. oye Asanza 30659 ple PERU. Loreto: Mariscal Ramón

Castilla, Río Yuvineto, affluent of the Putumayo, territory í ya Indians, in ta, 17 Jan 1978, fl., S. Barrier

84 (P); Maynas, Las Amazonas, 72°33 W, 3°25’S, 120-130 m, 17 p 1991, fr., C. Grind & N. Jaramillo 2847 (MO, MYF); Maynas,

Mishana, Río Nanay, 130 m, 20 Sep 1978, fr., C. Díaz & N. Jaramillo 577 (MO); Maynas, Río a deci uda tip e Ru Dn

15 May 1978, fr., A.H. Gentry, C. Diaz & N, Jaramillo 21931 (MO); Maynas, Yanomono, Río Ama Río

Napo, 3928'S, 72°48'W, 130 m, 27 Jul 1982, fr., A.H. Gentry, D. Alfaro & N. Jaramillo 37992 (E, MO). P Si , ca. 20 km 5

of Iscozacin, Río Palcazu Valley, 10°20°S, 75°10’W, 300 m, 9 Jul 1988, st., A.H. Gentry, C. Díaz & O. Phillips 63506 (MO).

5. Dilkea vanessae Feuillet, sp. nov. (Fig. = Tyre: FRENCH GUIANA. Pic Matecho, camp d it, 500 m, 14-15

Dec 2000, fr., V Hequet 1000 (uotorvPE: US; rsoTYP Y).

Haec ad subgenero Epkia pertinens. Petiolo basi cum pulvino, lamina foliorum 20-21.5 x 7-8.5 cm, oblanceolata, cuneata differt.

Treelet 0.8-2 m tall, monocaulous or branched. Stems with a long a 2-20 cm long, followed by short

ones 0.2-0.3 cm long. Tendrils lacking. Stipules not seen. Leaves in terminal clusters; petiole 2-2.5 cm long,

including the pulvinus 0.3-0.4 cm jng; glands not seen; blade oblanceolate, cuneate, 19-21 x 7-8 cm, basal

half triangular and then with a very narrow wing along the petiole until the swollen base, apex rounded

and long-acuminate, margin entire, midrib and veins raised on both surfaces and light brown when dry,

7-9 main veins on each side of the midrib. Inflorescences subterminal; pedicels ca. 6 mm long in bud, joint

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604 J f Texas 3(2)

2 mm from the base; bracts not seen. Bud 1.8 x 0.9 cm. Flowers not seen; under the fruit, androgynophore

and gynophore each 1 mm long. Fruits spherical, apiculate, about 3.5 x 3 cm, yellow, pericarp coriaceous,

0.2—0.3 mm thick, glabrous; seeds up to 8, elongate, 1.3~1.6 x 0.5-0.7 cm, reddish brown.

Distribution and ecology.—The only known collections were made in the rainforest, on Pic Matecho and

Montagne des Nouragues, granitic outcrops in central French Guiana. They were fruiting in November and

December (see discussion).

Dilkea vanessae has obovate leaves with a cuneate basal half.

The holotype label at US does not provide information about the date of collection. Vanessa Hequet,

the collector of the holotype, noted that it was collected on 14 Dec 2000 (pers. comm), but AUBLET, the

database of the herbarium in Cayenne (CAY), lists the collection date as 15 May 2001. According to AUBLET

and the few specimens I saw, Hequet 972 to 1003 were all collected on 15 Dec 2000 at the same locality.

Thus, the date 15 May 2001 represents an error in data entry.

Etymology.—The specific epithet is named after the collector of the type specimens, Vanessa Hequet,

currently at NOU.

E ENCH GUIANA: D Top Zone 5, Road Régi ac 4?7'N, 52°1’W, 100 m, 25 Nov 1995, fr., G. Cremers & J.-J.

de Granville 14236 bis (CAY); Mont CI il] I tl k, 3?49'N, 52°44’W, 100m, 21 Apr 1997, fr., G. Cremers

& F. Crozier 15124 (CAY): Mousse des MEA 29242 W 43 N, 7 Nov 1994, fr., B. Riéra & A. Joly 2031 (CAY, P, U); Montagne des

Nouragues, 52°42 W, 4°3’N, 150 m, 24 Oct 1995, fr., O. Poncy 973 (P).

ACKNOWLEDGMENTS

I am grateful 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 AAU, CAY, MO, NY, and P who hosted

me in their institution or made available to me the material in their care. I would like to thank Véronique

Guérin (CAY), Stephanie Kiel (MO), and Ingrid Pol-Yin Lin (US) who provided the scans of the type speci-

mens used for the illustrations. Reviews by Paul Hiepko (B), Shawn Krosnick (RSA), and Silvia Pérez (VEN)

are greatly appreciated. This is number 151 in the Smithsonian’s Biological Diversity of the Guiana Shield

Program publication series.

REFERENCES

BARBOSA RODRIGUES, J. 1888. Ordo Passifloreae Endl. Vellosia 1, ed. 1:24—31; ed. 2 (1891):21-31.

Cervi, A.C. 1991. A study on Brasilian Passifloraceae, Dilkea margaritae A.C. Cervi, spec. nov. Candollea 46:

FEUILLET, C. AND J.M. MacDoucaL. 2007. Passifloraceae. In: K. Kubitzki, The families and genera of vascular plants.

Springer, Berlin. 9:270-281.

GLaziou, A. 1909. Plantae Brasiliae centralis a Glaziou lectae. Liste des plantes du Brésil central recueillies en

1861-1895. Bull. Soc. Bot. France 56, Mém. 3d:297-392.

Harms, H. 1906. Passifloraceae. In: E. Ule, Il. Beiträge zur flora der Hy! hdebS von Ule's Amazonas

Expedition. Verh. Bot. Vereins Prov. Brandenburg 48:184-186. ["1906/ publ. 8 Mar 1907]

HOEHNE, F.C. 1915. Passifloraceae. Comm. Linh. Tel. Matto Grosso, Annexo 5:72-111.

HoLm-NIELSEN, L.B., PM. JØRGENSEN, AND J.E. Lawesson. 1988. Passifloraceae. In: G. Harling and L. Andersson (eds.), Flora

of Ecuador 31:1-129,

Kur, E.P. 1938. The American species of Passifloraceae. Publ. Field Mus. Nat. Hist., Bot. Ser. 19:1-613.

Masters, M.T. 1871. Contributions to the natural history of the Passifloraceae. Trans. Linn. Soc. 27:593-645.

Masters, M.T. 1872. Passifloraceae. | In: PM von Martius, ed. n. nee oo 13, pt. 1:530-654.

STEYERMARK, J.A. 1968. N erra Imataca y Altip e Nuria del Estado Bolívar y del Territorio

Delta Amacuro. E Acta Bot. Ven. 3:186- 190.

Titett, S.S. 2003. Passifloraceae. in: PE. Berry, K. Yatskievych, and B.K. Holst, eds. Fl. Ven. Guayana, vol. 7:625-667.

A NEW SPECIES OF SCHEFFLERA (ARALIACEAE)

FROM THE VENEZUELAN ANDES

L.J. Dorr Basil Stergios

National Museum of Natural History Universidad Nacional Sani tal

Dept. Botany, MRC-1 de los Llanos Occi es

Smithsonian Institution "Ezequiel Zamora" ( DENS

^O. Box 37012 Mesa de Cavacas, Guanare

Washington, DC 20013-7012, U.S.A. Estado Portuguesa 3323, VENEZUELA

dorrl@si.edu basilvengcantv.net

ABSTRACT

3 EE: T3 Cost s L ] =1 J 1:11 H aoo CPLS E CHAR 1 f. +1 D ]

Anew species of Schefflera, S

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c laxinsc and A 1 S. pittieri and 5

r L E L e

de Guaramacal in the Venezuelan Andes

Key Wonps: Araliaceae, Schefflera, Venezuela, Andes

RESUMEN

J EE: T5 C C : ] E I 4 J Te H | E 2 Pane A |

Una nueva especie de Schefflera, S. glos, y

Ramal de (65 1 los And nos. Se discuten | finidad lif

S. pittieri y S. quinquestylorum.

PALABRAS CLAVE: Araliaceae, Schefflera, Venezuela, Andes

Schefflera J.R. Forst. & G. Forst. (Araliaceae) as it is currently circumscribed is a very large genus compris-

ing ca. 600 described species, but there is ample evidence that the genus is polyphyletic and divisible into

at least five clades none of which correspond neatly with previously recognized segregate genera (Plunkett

et al. 2005). Molecular data suggest that the Neotropical species of Schefflera form a single clade (Fiaschi et

al. 2007), which in turn is Ep of a broader "Asian Palmate clade" (Plunkett et al. 2004, 2005; Wen et al.

2001 as “Eleutl Schefflera group") that is at present defined more by molecular and

geographic than morphological o

Schefflera is well represented in Venezuela. Species diversity is greatest in the Venezuelan Guayana

where ca. 60 species occur (Frodin 1997, 2008) and relatively low in the Andes, Cordillera de la Costa, and

adjacent Llanos. Presently, only five species of Schefflera are found in the Venezuelan Andes (Frodin 2008;

see also Bono 1996; Dorr et al. 2000; Huber 1977); S. cuatrecasasiana Steyerm., S. jahnii (Harms) Steyerm.,

S. rufa Frodin, S. karsteniana (Marchal) Harms, and S. tamana Steyerm. Two of these five (S. cuatrecasasiana

and S. tamana) are localized in their distribution and only one (S. karsteniana) could be considered to be

widespread. Three species of Schefflera are found in the Cordillera de la Costa (Frodin 2008; see also Meier

1998; Steyermark & Huber 1978); S. glabrata (Kunth) Frodin, S. karsteniana (Marchal) Harms, and S. pittieri

Harms. Two of these three have fairly wide distributions, while the third (S. pittieri) is known only from a

few, adjacent localities in Aragua. (The Mérida record for S. pittieri cited by Frodin (2008) is almost certainly

an error). The sole species of Schefflera, S. morototoni (Aubl.) Maguire et al., found in the Venezuelan Llanos

also occurs in the Venezuelan Guayana (Frodin 2008; see also Duno de Stefano 2007).

The following new species can be added to the small number of species found in the Venezuelan An-

des. We were aware that it was undescribed when we published an enumeration of the vascular plants of

Guaramacal National Park (Dorr et al. 2000), but only subsequently did we collect material adequate for the

description that follows.

oe — por de eee sp. nov. ov. (Fig. 1). Ter: A TRUJ Lo: Mpio. Boconó, Parque Nacional

Gua sua Negra Oda. Saly aje (UTM: 1027793 N; 371576

E), 2000-2350 m, 15 Apr 2003 (fr), B. Stergios, LI Dorr & R. PM 20271 ae PORT: IsOTYPES: K, MO, US-2 sheets, VEN).

J. Bot. Res. Inst. Texas 3(2): 605 — 608. 2009

606 j tl tanical Institute of Texas 3(2)

Schefflera sp. A, Dorr et al., Contr. U.S. Natl. Herb. 40: 66. 2000 [2001].

Species nova a Scheffl I | f] ta et styli 5 differt.

Scandent shrubs or small trees, 3-4 m tall; di 5-7 mm in an a: andromonoecious. Leaves

digitately compound, (6—)7-foliolate, the leaf] ingle plane, | ate to ovate, chartaceous,

margin hyaline and revolute, apex long acuminate, base attenuate to e slichis obliasie sparingly

pubescent especially along midrib and secondary veins below, glabrous above; venation brochidodromous,

primary vein raised above and below, secondary veins slightly raised below, not noticeably raised above,

8-14 per side, arcuate-ascending. Petioles (10—)16.5—27 cm long, slender, terete, sparingly pubescent when

young, glabrescent in age. Stipular ligules 1.8-2.4 cm long, margin entire or sparingly ciliate, apex entire

or rarely slightly bifid. Median leaflets: petiolule (3.3-)4.5—7 cm long, blade 12.5-22.5 cm x 4.2-8.2 cm,

apical acumen 1.5-2.8 cm long. Basal leaflets: petiolule 1-3.5 cm long, blade (7.5212-13.5 x Q.5-)3.7 x 5.7

cm, apical acumen 1.5-3 cm long. Inflorescence appearing subterminal, paniculate, erect, G— 4-branched;

peduncle up to 7.5 cm long; rachis up to 5 cm long, bracts subtending rachis up to to 1 cm long; primary

branches 7 to 12, a Ta cm long ün um bract ca. 3 mm long; secondary branches 4 to 6, 1-1.7 cm long

bellules; each umbel-

(in fruit); ultimate inflorescence units umbe llate, often with 1—2 subtending lateral

lule with 4 to 6 flowers subtended by a ca. 1 mm long. Pedicels 1-2 mm long (in bud), 3-4 mm

long (in fruit), pubescent to hirsute when young, glabrescent in age, not articulate. Flowers bisexual and

staminate. Calyx obconic, ca. 1 mm long, with 5 minute teeth; corolla calyptrate, 3-5 x 3-4 mm, pubescent

to hirsute, glabrescent in age; stamens 5, filaments ca. 0.25 mm long (in bud); anthers ca. 1.5 mm (in bud),

arcuate. Styles 5, free (in bud) or very slightly connate basally (in fruit), strongly diverging in fruit and ca. 2

mm long, stigmatic surface capitate. Fruit strongly 5-ribbed when dried, urceolate, sparingly pubescent to

glabrescent; pyrenes (evidently immature) 5, each 1-seeded, ca. 6 x 2.5 mm, flattened.

Distribution and ecology.—Schefflera vanderwerffii is known, at present, only from north- and south-

facing slopes of the Ramal de Guaramacal in the northeastern corner of Guaramacal National Park where it

is found in lower montane or cloud forest; 1700-2350 m

Etymology.—The specific epithet honors Henk van der Werff of the Missouri Botanical Garden who

first collected this species. Early in his career he lived in Falcon and in addition to working on the flora of

that state he also made many valuable collections in the Venezuelan Andes.

PARATYPES. VENEZUELA. Port Sucre, La Divi le la C ión, 9°18'N, 70°06'W, 1700 m, 23 Oct 1985 (fl bud), H.

van der Werff et al. 7535 (PORT), 24 Oct 1985 (fl WES H. van der Werff et al. 75 76 (MO, PORT).

Schefflera vanderwerffii b perfici bl to S. pittieri, MÀ i nid to the size, shape,

and texture of its leaves. The new species differs from S. pittieri, however, in] ore congested

inflorescence (ca. 10-12.5 versus 20-40 cm long), lamer fruit (6 mm versus 3s mm long), 2nd TN styles

(5 versus 6—8). Huber (1977: 65), in his enumeration of the woody plants of Mérida and adjacent states,

considered S. pittieri to be a synonym of S. quinduensis (Kunth) Harms, but as noted by Harms (1927: 300)

the two differ in the size ei their flowers and the number of styles; 6-8 in the former species, 8-10 in the

latter. In addition, S. pitti to be a local endemic confined to the Cordillera de la Costa of Venezuela,

while S. ind is — known only from the Andes of Colombia and Ecuador.

Schefflera vanderwerffii might be more closely related to S. quinquestylorum Steyerm., which is found in

the Venezuelan Guayana (Bolívar state) and adjacent Guyana. Here, too, the size, shape, and texture of the

leaves are similar as is the number of styles, and shape of the fruit (see Frodin 1997, hg. e Pene d od

questylorum, however, has a much longer (19-20 cm long), less divided infl nd the style |

in fruit are completely free from the base.

We believe that Schefflera vanderwerfii is andromonoecious as are many species of Araliaceae. There

are clearly two floral morphs. Our description of the flowers, however, is based solely on staminate ones

as among the limited flowering material available to us for analysis (two collections in bud) we found only

flowers with well-developed anthers and vestigial styles and ovaries. The well-developed styles in the one

fruiting collection available for our study strongly suggest that there are also bisexual flowers.

| Stergios, A new species of Schefflera from the Venezuelan Andes 607

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ACKNOWLEDGMENTS

Field work was supported by funds from the Biological Survey and Inventories Program of the National

Museum of Natural History. David G. Frodin made Mei suggestions i pun the affinities of this new

species. Gregory M. Plunkett made helpful tions for improving the technical description of the

same. We are indebted to the staff of Guaramacal National Park for assistance in the field, the Ministerio del

Ambiente and INPARQUES for permits, and Cathy Pasquale for the illustration.

REFERENCES

Bono, G. 1996. Flora y vegetación del Estado Táchira, Venezuela. Museo Regionale di Scienze Naturali (Torino),

Monografie 20:1-951.

Dorr, L.J., B. STtERGIOS, A.R. SMITH, AND N.L. CueLLO A. 2000 [2001]. Catalogue of the vascular plants of Guaramacal

Nationai Park, Portuguesa and Trujillo states, Venezuela. Contr. U.S. Natl. Herb, 40:1-155.

Duno DE STEFANO, R. 2007. Araliaceae. In: R. DUNO DE STEFANO, G. AYMARD, AND O. Huser, eds. Catálogo anotado e ilus-

trado de la flora vascular de los Llanos de Venezuela. FUDENA-Fundación Empresas Polar-FIBV, Caracas. Pp.

358-359.

FIASCHI, P, G.M. PLUNKETT, AND PP. Lowry II. 2007. What do we know about Neotropical Schefflera (Araliaceae)?: a first

molecular phylogenetic analysis. Plant Biology & Botany 2007 Joint Congress. [Botanical Society of America,

St. Louis]. P. 294

FroDIN, D.G. 1997. Araliaceae. In: PE. Berry, B.K. Horst, AND K. YarskievrcH, eds. Flora of the Venezuelan Guayana

3:1-31

FRODIN, D.G. 2008. Araliaceae. In: O. HokcHE, PE. Berry, AND O. Huser, eds. Nuevo catálogo de la flora vascular de

Venezuela. Fundación instituto Botánico de Venezuela, Caracas. Pp. 215-218.

Harms, H. 1927, Araliaceae americanae novae. |. Repert. Spec. Nov. Regni Veg. 23:299-301.

Huser, H. 1977. Gehólzflora der Anden von Mérida. Teil |. Mitt. Bot. Staatssamm!. München 13:1-127.

Mezer, W. 1998. Flora und vegetation des Avila-Nationalparks (Venezuela/Küstenkordillere) unter besonderer

Berücksichtigung der Nebelwaldstufe. Diss. Bot. ar -X, as

PLUNKETT, G.M., PP. Lowry Il, D.G. Fronin, AND J. WEN. 2005. Phylogeny hy of Schefflera: | ive polyphyly

in the largest genus of Araliaceae. Ann. Missouri Bot. Gard. 92: 202- 224.

PLUNKETT, G.M., J. WEN, AND PP. Lowey II. 2004. Infrafamilial classifications and characters in Araliaceae: insights from

the phylogenetic analysis of nuclear (ITS) and plastid (trnL-trnF) sequence data. Pl. Syst. Evol. 245:1-39,

STEYERMARK, J.A. AND O. Huser. 1978. Flora del Avila. Publicación Especial de la Sociedad Venezolana de Ciencias

Naturales, Caracas. Pp. 1-971.

Wen, J, G.M. Plunkett, A.D. Micheli, AND S.J. Wacsrarr. 2001. The evolution of Araliaceae: a phylogenetic analysis

based on ITS sequences of nuclear ribosomal DNA. Syst. Bot. 46:144-1

NEW SPECIES OF EUGENIA (MYRTACEAE) FROM ECUADOR

Maria Lucia Kawasaki Bruce K. Holst

Department of Botany Marie Selby Botanical Gardens

Museum of Natural History 811 South Palm Avenue

1400 South Lake Shore Drive Sarasota, Florida 34236, U.S.A.

go, Illinois 60605-2496, U.S.A. bholst@selby.org

er eldmuseum.org

ABSTRACT

Five new species of Eugenia (Myrtaceae) from Ecuador are described and illustrated: Eugenia aequatoriensis, E. castaneiflora, E.

crassimarginata, E. longisepala, and E. pusilliflora.

RESUMEN

Se describ ilustran cinco especi de Eugenia (M ) del Ecuador: Eugenia aequatoriensis, E. castaneiflora, E.

whi AL Lad Pi E ML

BM MMC E. barea y E. pusilliflora.

INIRODUCIMON

In the checklist of Myrtaceae of Ecuador (Holst 1999), ca. 60 species of Eugenia

30 species represented possible new taxa and some of these have been recently bel dla &

Holst 2009). In preparation of the Myrtaceae treat t for the Flora of Ecuador, five additional new species

of Eugenia are here described: Eugenia aequatoriensis, E. castaneiflora, E. crassimarginata, F. longisepala, and

E. pusilliflora.

Eugenia aequatoriensis M.L. Kawasaki & B.K. Holst, sp. nov. (Fig. 1). Tres: ECUADOR. Loja: Bosque ‘El Tundo’, property

f ARCOIRIS foundati ff Macará, ca. Km 1, 4°19'29.4"S, 79%49'13.9"W 1750 m, 12 Jun 1996 (fI), B.B.

Klitgaard, B. Stáhl, P Losami T. Delgado, F. Flizalde, H pase & E Tinitana 173 (noLorvre: AAU; isotypes: NY, QCA, SEL).

Frutex vel arbor; foliis ellipticis vel anguste ellipticis; racemis pubescentibus, indumento brunneo-rufescenti vel rubro-flavescenti,

BE 11; 1 a r1 a Un | 1 s -7 f. £ 11; Lh MI 1 :

GALAS SAL o

Shrub or tree 3-25 m tall; trichomes, where present, brownish- or yellowish-red. Leaf blades elliptic to

narrowly elliptic, chartaceous to coriaceous, 9-13.5 x 3-6.5 cm, mostly glabrous, drying olive-green or

land tiform, indistinct above, dark and numerous below, midvein

greenish-brown above, paler below; g

sulcate on the upper surface; lateral veins 10—15 pairs, indistinct or slightly convex above, convex below;

] +]

marginal veins 2, the innermost slightly arched, 1-3 mm from the margin; shortly acuminate;

base cuneate; petioles 1-1.5 cm long, sulcate, glabrous. Inflorescences T short racemes, with up to ca.

6 flowers, axillary or subterminal, the main axis 3-13 mm long, the pedicels 4-13 mm long, appressed-

pubescent; bracteoles triangular, acute, to 1 mm long, appressed- PU PE Flowers 4-merous;

buds 3-4 mm long; hypanthium ca. 2 mm long, appressed-pubescent; lar, 2-3 mm long,

obtuse, appressed-pubescent; petals suborbicular, ca. 4 mm diam., lue disk ca. 3 mm diam, , glabrous;

stamens ca. 4-6 mm long; style ca. 6 mm long; ovary 2-locular, with several ovules per locule. Fruits ber-

ries, ellipsoid or obovoid, 1.3-1.5 x 1-1.1 cm, puberulous to glabrous, reddish; seed 1, ca. 10 x 8 mm, the

seed coat membranous; embryo eugenioid, the cotyledons fleshy, connate, the radicle indistinct.

Distribution. —Known only from the region of Reserva Natural El Tundo, Sozoranga, Loja province, in

wet premontane forests at 1750-1800 m elevation.

Among the species of Eugenia with racemose inflorescences in Ecuador, E. aequatoriensis is readily

distinguished by the short, brownish- or yellowish-red pubescent racemes, with the main axis 3-13 mm

long and a few (up to ca. 6), long-pedicellate flowers, the pedicels 4-13 mm long, often longer than the main

J. Bot. Res. Inst. Texas 3(2): 609 — 618. 2009

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axis of the inflorescences. While E. aequatoriensis is distinct among Neotropical Myrtaceae, its affinities are

not clear.

Additional collections examined: ECUADOR. Loja: Sozoranga, Reserva Natural El Tundo, dic ee ARCOIRIS foundation, Km 6

along track from Sozoranga-Macará road, 4°19'S, 79°49'W, 1800 m, 19 Aug 1997 (fr), G.P. Lewis ga, El Tundo,

Mar 1983 (fl), F. Vivar C. et al. 1727 (AAU, LOJA).

Eugenia castaneiflora M.L. Kawasaki & B.K. Holst, sp. nov. (Fig. 2). Tre: ECUADOR. Manasi: Pedernales, 34 km N

of Pedernales along new coastal highway, near village of Chindul, 00°14'S, 79°52'W, 60 m, 28 Aug 1998 (buds), D. Neill & QCNE

botany interns 11447 (HoLotypE: MO; isotypes: E QCNE).

Frutex vel art folii lliptici l oblanceolatis, 27-39 cm longis, coriaceis, glabrescentibus, acuminatis,

convexo vel Eno: TOM lana castaneo-tomentosis; fructis ellipsoideis.

Shrub or tree 3—7 m tall, the trichomes on branchlets, petioles, and inflorescences chestnut-brown, yel-

lowish-brown on leaf blades. Leaf blades narrowly elliptic or oblanceolate, coriaceous, 27-39 x 7-10 cm,

drying greenish-brown or brownish, much paler below, glabrescent, the few trichomes by the midvein on

the lower surface; glands punctiform, almost indistinct on both surfaces or slightly convex below; midvein

convex or biconvex on the upper surface; lateral veins 20-25 pairs, convex on both surfaces; marginal veins

2, the innermost slightly arched, 2-5 mm from the margin; apex acuminate; base obtuse; petioles 1-1.5 cm

long, flattened, puberulous to tomentulose. Inflorescences of abbreviate racemes, with up to 10 flowers,

the main axis inconspicuous; flowers appearing fasciculate at leafless nodes, the pedicels 1-1.5 cm long,

tomentose; bracteoles lanceolate, 2-4 mm long, tomentose, persistent. Flowers 4-merous; buds 9-10 mm

long, tomentose; hypanthium ca. 4 mm long; calyx lobes suborbicular, 5-7 mm long, obtuse, apparently

deciduous in fruit; fully developed petals, stamens, and style not seen; disk ca. 3 mm diam., glabrous; ovary

2-locular, with several ovules per locule. Fruits berries, ellipsoid, 3-4 x 1.5-1.8 cm, puberulous, glabres-

cent, gland-dotted, crowned by a tomentose collar; seeds 1(2), (1-22.5 x (0.521 cm; embryo eugenioid, the

cotyledons fleshy, connate, the radicle indistinct.

Distribution —Eugenia castaneiflora is known only from Manabi province, occurring in wet forests at

60-250 m elevation.

In leaf morphology and type of infl Fugenia castaneiflora is similar to E. tumulescens McVaugh,

a species from northern Amazonian Brazil and sauter Jenez In E. castaneiflora, however, the glands

on leaves are punctiform and mostly indistinct (vs. prominently convex) on both surfaces, the petioles are

smooth (vs. corky), the flowers are characteristically chestnut-brown (vs. yellowish-brown) and borne at

leafless nodes (vs. axillary), the pedicels are 10-15 (vs. 8-10) mm long, and the calyx lobes are suborbicular

(vs. oblong), 5-7 (vs. 6-8) mm long.

Additional collection examined: ECUADOR. Manabi: Pedernales Cantón, Reserva Ecológica Mache-Chindul, comunidad Ambacha

(Chindul coastal highway), 250 m, 00°15'N, 79°48'W, 25 Mar 1997 (fr), J.L. Clark et al. 4152 (F, MO, QCNE, US).

Eugenia crassimarginata M.L. Kawasaki & B.K. Holst, sp. nov. (Fig. 3). Tee: ECUADOR. Coropaxi: Tenefuerste,

Rio Pilalo, Km 52-53, Quevedo, Latacunga, 750-1300 m, 7 Feb 1982 (old fl), C.H. Dodson & A.H. Gentry 12168 (notorvre: MO;

ISOTYPES: SEL, US

Ll L e I r

1 Il

Arbor; foliis elliptici llipticis vel obovatis, chartaceis, pl labris vel ] pubescentibus, abrupte acuminatis,

p L 1 E [. Pa 11: >] .

T pu

"— venis márcialibus 4—8 mm remotis, d pra Į

Tree 5-15 m tall; trichomes, where present, yellowish-white to coppery. Leaf blades elliptic to narrowly

elliptic or obovate, chartaceous, 10-16 x 4.5-7 cm, drying olive-green, brown or dark-brown above, light

greenish-brown below; upper and lower surfaces mostly glabrous to sparsely appressed-pubescent; glands

punctiform, dark and convex on both surfaces; midvein convex on i upper niai lateral veins 8-12

xxx F]

pairs, convex on both surfaces, but impressed o y sulcate; marginal

veins 2, the innermost similar to the lateral veins in prieta slightly ied. 4— 8m mm from the margin,

the leaf blades appearing thick-marginate; apex abruptly acuminate, the acumen 1-2 cm long; base cune-

ate to obtuse; petioles 4-8 mm long, flattened, glabrous. Inflorescences of abbreviate racemes, with up to

375)

f Texas 3(2)

“tof eld

6 7 8 9 10

ECUADOR

MYRTACEAE

. Mangbi: Pedernales

34 ka north of Pedernales along nee

" coacta) niahwday, near village of

indul Mesi wert forast.

00" 145 079" CO n

peel trec 7 m tall. Unasratory in

primary fores. Flora) buds with

Pecani ah pubaescente.

28 August 1998

N? 2255781

J: JJ b i UE jd ji

FIELD z EUM Field Museum of Natural History (P)

Isotype of:

NATURAL HISTORY

Engen cadtancylora M E. Kawasaki & B.K. Holst David Neiti & QCNE betany interna

HERBARIO NACIONAL GEL ECUADOR (0 Chey

MISSOURI, BOTARICAL GARDEN HERBARIUM (40)

iflora M.L. Kawasaki €: B.K. Holst (Neill et al. 11447: Isotype, F; fruit f Clark et al. 4152, F).

Fic. 2 Fug

"3. o. LI F r1

LI JI

613

n OUR:

CAL GARUER

HERBARIUM

Frold Miuscom of Natural ilrzory tb)

ali sow oi

AS tpi) M E

PLANTS GF ECUADOR

Eugenie

Eur wate. file Pilato, Xm betj

Vi T5U 3300 mi, Peay Cut piri

(eli. A Doreen & A. H Gentry

Vusussk: & LC

3, Quevedo, Levens

highs

Mey ovem PUn?

121455

LEIA CP THOM MARIE SELEY SOTASICAL G SEDENS

¡putos br a HAST Geint es TERTII 999

E togada i etr Xoaeheimrfe, cap

MO; inset from isotype at SEL)

Fic. 3 Eug

Kawasaki & B.K. Holst (D

614 J | of the Botanical R h Institute of Texas 3(2)

8 flowers, the main axis inconspicuous; flowers appearing fasciculate at leafless nodes, the pedicels 1-2.5

cm long, slender, appressed-puberulous to glabrous; bracteoles ovate, ca. 2 mm long, deciduous. Flowers

4-merous; buds not seen; hypanthium ca. 2 mm long, appressed-pubescent; calyx lobes ovate, ca. 2 mm

long, obtuse, glabrous; petals not seen; disk ca. 2 mm diam., glabrous; stamens and style not seen; ovary

2-locular, with several ovules per locule. Fruits berries, ellipsoid, 1.5-2.5 x 1-1.5 cm, glabrous, greenish-

brown, gland-dotted; seed 1, ca. 15 x 9 mm, the seed coat membranous; embryo eugenioid, the cotyledons

fleshy, connate, the radicle indistinct.

Distribution.—Eugenia crassimarginata has been collected in the provinces of Cotopaxi, Los Ríos, and

Pichincha, in forests at 500-1300 m elevation.

Eugenia crassimarginata is Vision qu. by the upper surface of leaves with convex venation but im-

mersed in the leaf blade, ap] g gly sulcate, the HUM veins ae in a continuous inner marginal

vein at 4-8 mm from margin, and by the relatively small fl elongate, slender pedicels. Eugenia

churutensis X. Cornejo, a species know only from lowland forests of Guayas provincë: has similar flowers and

type of inflorescences, but each fascicle has 8-30 (vs. to 8) flowers, the bracteoles are narrowly triangular,

0.5-1 mm long, probably persistent (vs. ovate, ca. 2 mm long, deciduous), and the lateral veins join in ir-

regular as not MER. the characteristic marginal vein of E. crassimarginata.

A Mie 1 "ECUADOR TI R O do Cantón, Cerro Centinela, Montañas de Ila, 10 km E f Patricia Pil

na imal 500 m, EU. p ee W. Pal acios E. Freire 7452 (SEL). Los Rios/Pick icia Pilar, at 45 kon road

Centinela, FIA le Tl 1f P Pilar to 24

de Mayo at Km 12, 600 m, a, 10 in 1979 dis C.H. Do etal. ea (Es a SEL) Patricia ua at 45 km on road from Santo Domingo to

Quevedo, pati ela icia Pilar to 24 de Mayo at Km

m, 2 Oct 1979 (fo, C.H Dodson et al. 8686 (MO, SEL) Pichincl ntinela, rod Patricia Pilar-24 de Mayo, Km 12, ca. Km 47 of road

Santo Domingo-Quevedo, at le Ila, 650 m, 26 " 1984 (old fl/young fr), C.H. Dodson et al. 14523 (MO, SEL).

Eugenia longisepala M.L. Kawasaki & B.K. Holst, sp. nov. (Fig. 4). Tre: ECUADOR. Sucumstos: Sacha Lodge, 3 km

N of the village Añangu, near the Napo river, 00°39'S, 76°26'W, 200 m, 12 Jun 1995 (fr), J.L. Clark, L. Demattia & 1 Miller 1176

(HOLOTYPE: MO; isotypes: E, US).

Arbor; foliis anguste ellipticis vel lanceolatis, 23-36 cm longis, coriaceis, pl labri di minente;

racemis pubescentibus, indumento brunneolo vel brunneo-flavescenti, calycis lobis eds 10-13 mm loügis: fructis ellipsoideis.

Tree 7-12 m tall; trichomes, where present, brownish to yellowish-brown. Leaf blades narrowly elliptic to

lanceolate, coriaceous, 23-36 x 7-13 cm, drying olive-green to greenish-brown above, paler below; upper

surface mostly glabrous; lower surface glabrous to thinly sericeous; glands punctiform, indistinct on both

surfaces or slightly convex below; midvein plane to convex on the upper surface; lateral veins 17-20 pairs,

convex on both surfaces; marginal veins 2, the innermost slightly arched, 3-7 mm from the margin, apex

abruptly short-acuminate; base cuneate to obtuse; petioles stout, 2—2.5 cm long, sulcate, wrinkled, sparsely

tomentulose, glabrescent. Inflorescences of short racemes, axillary, the main axis in fruit 1-2.5 cm long,

the pedicels in fruit 6-10 mm long, pubescent; bracteoles not seen, deciduous. Flowers (loosen) 4-merous;

buds not seen; hypanthium ca. 6 mm long, pubescent; calyx lobes lanceolate, 10-13 x 5 mm long, foliaceous,

acute, pubescent; petals obovate, ca. 1 cm long, glabrous; disk 3-4 mm diam., glabrous; stamens ca. / mm

long; style ca. 7 mm long; ovary 2-locular, with several ovules per locule. Fruits berries, ellipsoid, 2.53 x

1-1.5 cm, pubescent, greenish-brown, crowned by the elongate, involute calyx lobes; seed 1, ca. 2.3 x 1 cm,

the seed coat membranous; embryo eugenioid, the cotyledons fleshy, connate, the radicle indistinct.

Distribution.—Eugenia longisepala is known from the provinces of Orellana and Sucumbios, occurring

in wet forests at 200-360 m elevation.

Eugenia longisepala is similar to Eugenia macrocalyx (Rusby) McVaugh, a species from northern South

America to Bolivia, in the uncommonly long and foliaceous calyx lobes. In E. longisepala, however, the inflo-

rescences, pedicels, and flowers are stout (vs. slender), the trichomes are brownish to yellowish-brown (vs.

whitish) and the leaves are much larger (23-36 x 7-13 cm vs. 6-13 x 3-5 cm), with a convex (vs. sulcate)

midvein on the upper surface.

j ki and Holst, New species of Eugenia from Ecuad 615

ao Y

m, Lm

"A — ECUADOR

Field Museum of Natural History (F)

HYRTACEAE

Eugenia

Eugenia longisepala ML, Kawssski & B.K. Holst

d in Sucumbios

Sacha Lodge, 3 ae NY a il village

Añangu, near the Napo

' Tropical sp Forest

39' GW

N? 2257296 00°39'S 076'2 200 m

ERLO MUOSEOM Tree, 12 m tall. Fruits green a

or covered with brown pubescence. .

i2 June i995

NATURAL HISTORY

John L. Clark,

Liz Demattia, & Tim Miller 1176

HISSOURT BOTANICAL GARDEN HERBARIUM (MO)

Fic. 4. Fugeni longi fa M.L. Kawasaki €: B.K. Holst (Clark et al. 1176: Isotype, F; fl f Kornina € TI 58651, AAU)

|| || £ al nm Li nm Li 8.2.4. F1 a fmt

616 t t f Texas 3(2)

Additional collections examined: ECUADOR. Orellana: Añangu, NW corner of the Parque Nacional Yasuní, surroundings of the SEF

terra firme line and KTH hectar plot, 00°33'S, 76°22'W, 260-360 m, 1-30 Apr 1985 (fl), J. Korning & K. Thomsen 58651 (AAU, QCA,

QCNE); Estación Científica Yasuní, Río i bane NW of confluence with Río Tivacuno, E of road Repsol-YPF, Km 7 on spur road to oil

well Tivacuno, sendero Napo at 140 ft side), 00°38'S, 76°30'W, 200-300 m, 23 Jul 2002 (fr), G. Villa & P. Alvia 1824 (AAU, QCA, US).

Eugenia pusilliflora M.L. Kawasaki & B.K. Holst, sp. nov. (Fig. 5). Ts ECUADOR. Naro: Tena Cantón, Biological

Station Jatun Sacha, 8 km E of Misahuallí, near oe stream, permanent parcel 03, 01°04'S, 77°36'W, 400 m, 16 Feb 1990

), C.E. Cerón, M. Crizón & C. Iguago 8667 (HOLOTYPE: Q EE

YAGQOIMINna

Arbor, folii Hipti „elli lol tis, chartaceis, puberulis, } pt dat

is, subtus TE e e neryo

{i leat 1 H] TT: , adpresso-pu theerentibie 2151 Ile: 11: 1 `

1 LES Y oOo I e oO

Tree 6-15 m tall; trichomes, where present, yellowish-white; branchlets smooth, puberulous, dark-dotted.

Leaf blades narrowly elliptic to elliptic or obovate, chartaceous, 5.5-7.5 x 2-3.3 cm, drying olive-green or

greenish-brown above, light yellowish-green below, puberulous on both surfaces, especially by the midvein,

glabrescent; glands dark, convex on both surfaces or indistinct above; midvein sulcate on the upper surface;

lateral veins ca. 10 pairs, indistinct above, slightly convex below; marginal veins G the innermost similar to

the lateral veins in prominence, slightly arched, ca. 1 mm from tl tly caudate-acuminate,

the acumen 1-1.5 cm long; base cuneate to obtuse; petioles 4-6 mm ime okas a to glabrous.

Inflorescences of short racemes, with 4-8 flowers, axillary or subterminal, the main axis 1-4 mm long,

quadrangular in cross-section, the pedicels 4-7 mm long, slender, appressed-pubescent; bracteoles ovate,

to 1 mm long, appressed-puberulous, persistent. Flowers 4-merous; buds ca. 1.5 mm long; hypanthium

ca. 1 mm long, appressed-pubescent; calyx lobes ovate, 0.4—0.8 mm long, in slightly unequal pairs, acute to

obtuse, appressed-puberulous without, ciliate; petals suborbicular, ca. 2 mm diam., ciliate; disk ca. 1 mm

diam., puberulous; stamens ca. 3 mm long; style ca. 3 mm long; ovary 2-locular, with several ovules per

locule. Fruits berries, globose, 7-9 mm diam., glabrous, dark-red to purple, gland-dotted; seed 1, ca. 6-7 x 5

mm, the seed coat membranous; embryo eugenioid, the cotyledons fleshy, connate, the radicle indistinct.

Distribution.—Eugenia pusilliflora has been collected in the provinces of Napo, Orellana, and Pastaza,

in primary rain forests at 200-430 m elevation.

Eugenia pusilliflora is characterized by the small, elliptic to obovate, abruptly caudate-acuminate leaves,

dark-dotted below and with sulcate midvein on the upper surface; the inflorescences are short racemes

with very small flowers and slender pedicels that are longer than the main inflorescence axis. In the shape

of leaves, type of inflorescence, and size of flowers, Eugenia pusilliflora is similar to E. protenta McVaugh, a

species from Venezuela and northern D ix ed The two PORE may be odd) c dde by Prol

glabrous), « inctly dark-dotte

characters: the leaves of E.

(vs. glands indistinct), piston (vs. Kors ou. with sulcate a (vs. sulcate to plane); the axis of

the inflorescences and bracteoles are yellowish-pubescent (vs. glabrous to sparsely rufous-pubescent), and

the vee is Eee -pubescent (vs. peus

A dAAitinnal 11 J. z^ TTA TA£ ID. AT a 71 D A 1 (21 cC [r1 1 : 1°05'S,

77°36'W, 430 m, 10 Mar 1985 (fl), D. Neill et al. 6063 (MO, SEL); Tena Cantón, Jatun Sacha Biological a: 8 km E of Misahualli,

01°04'S, 77°36'W, 400 m, 21 Sep 1990 (fr), D. Neill 9465 (F, MO, SEL); Tena Cantón, Jatun Sacha Biological Station, 5 of Rio Napo, 8

km E of Misahuallí, 01°04'S, 77°36'W, 400 m, 17 Sep 1992 (fr), D. Neill Zuleta 10131 (F, MO, SEL); Tena Cantón, Ahuano, 01°03'S,

77°32'W, 380 m, 10 Sep 1992 (fr), W. Palacios et al. 10438 (F, MO, QCNE, SEL). Orellana: Yasuni Forest Reserve, along road between

Km 70 and 100, E of PUCE e Station, 00°50.014'S, 76°20.518' Wto 00°50.730'S, 76°13. o 200 m, Bede 1995 (fr), P. Acevedo-

Rodriguez & J.A. Cedeño 7633 (SEI ) Aguarico, R Km Pad 00°56'S,

76°13'W, 248 m, 24-31 Jul 1994 (fr), A. Dik 1491 (F, MO, SEL); Aguarico, R Etnica H i, | I

project, Km 99—100, 00°56'S, 76°13'W, 250 m, 9 Jul 1994 (fr), N. Pitman 510 (SEL); Aguarico, R H

pipeline construction project, Km 99-100, 00°56'S, 76°13'W, 250 m, 10 Jul 1994 (fr), N. Pitman 515 (F, MO, SEL); Estació ntifica

Yasuní, Río Tiputini, NW of the confluence with Río Tivacuno, E of inn Repsol-YPF, Km 7, detour to Tivacuno well, 00?38'5, pen W,

200—300 m, 12 Feb 2002 (fl), G. Villa et al. 1302 (SEL) Yasuni, Río Tiputini, NW of confluence with Río Tivacuno, E of

road Repsol-YPF, Km 32 to NPF, Guiyerc, 00°38 S, 76°30' W, 200-300 m, 17 Jun 2002 (fn), G. Villa et al. 1540 (F, QCA, SEL). Pastaza:

Oil well “Golondrina”, PETRO-CANADA, 30 km NW of Curaray, 01°10'S, 77°06'W, 400 m, 10 Jul 1989 (fr), E. Gudiño 8 (F, MO, SEL).

i, Maxus road and

re TP Ist, N | . £T AP r J 617

iid Mescum ol ius Histeiy id]

otype: at:

natiiiaa MI. Kawa & AL, Halt

ECUADOR

Cant

Estación Biológica Jatun Sacha. 8 km

al e de Misahualli. Boat o zu

———————

= w

— inei Propi Mi une Bosque prinariv

cerca al riachueio Chirjui Pim. Parcel

- Permanente TAN

M? 2258152 77*36'W 01*04*8 400 f

Arbol de 15 m se epee

FIELD MUSAUM Florez verde-ugua

NATURAL HISTORY Carlos E. Cerin; i6 febrero 1990

E. Crizon & C. Iquago 8667

HERBARIO NACIONAL DEL ECUADOR (QCNE)

HISSOURI BOTANICAL GARDEN HEREARIOM (MO)

Cri

Ec 5 F j Thi MI Kawasaki & R.K. Holst (G

[| [| £ al n. H 1m La PEA ass)

618 f Texas 3(2)

ACKNOWLEDGMENTS

We thank the curators and staff of the herbaria cited in t! ipt for providing the collections used in

this study, especially Henrik Balslev and Benjamin Øll 1 from AAU, and Ron Liesner and Jim Solomon from

MO. We also thank Fred Barrie and Les jas m for helpful suggestions to improve the manuscript.

REFERENCES

Horst, B.K. 1999, Myrtaceae. In: PM. J | S. León-Yánez, eds. Catalogue of the vascular plants of Ecuador.

Monogr. Syst. Bot. Missouri Bot. Gard. 75:618-622.

Kawasaki, M.L. ano B.K, Host. 2009. Three new species of Eugenia (Myrtaceae) from Ecuador. Selbyana 30:

101-106.

REVISION OF BAUHINIA SUBGENUS PHANERA

SECTION SCHNELLA (CERCIDEAE: CAESALPINIOIDEAE: FABACEAE)

Richard P Wunderlin and Richard M. Eilers

institute for alles ele

Department of Cell Biology Molecular Biology

panca of South Flor rida

4202 zu aa doi BSF 218

0, U.S.A.

dn e E

ABSTRACT

A = 4 4 cp Lis bg Di ti C-l 71 En ESI 4 +L, E lilacina Wun-

derlin & Eilers from Bahia, Brazil and tl bination B tüchssendla (L.P (citas Wonderin & Eilers as new. Keys, descriptions,

and distribution data are provided.

RESUMEN

Se presenta un tratamiento taxonomico de Bauhinia subgénero Phanera sección Schnella. Se reconocen ocho especies con B. lilacina

Wunderlin & Eilers de Bahia, Brasil y se hace la nueva nan B. trichosepala (L.P. Queiroz) Wunderlin éx Eilers. Se aportan

claves, descripciones, y datos de distribución.

The pantropical genus Bauhinia L. (Cercideae: Caesalpinioideae: Fabaceae) with approximately 330 species

consists of four subgenera: Bauhinia L., Piliostigma (Hochst.) Kurz [-Elayuna (Raf.) Wunderlin], Barklya (F.

Muell.), and Phanera (Lour.) Wunderlin (Wunderlin et al. 1987). Bauhinia subgenus Phanera, as defined by

Wunderlin et al. (1987), is a pantropical group consisting primarily of tendriled lianas in 11 sections, of

which two, Schnella and Caulotretus, are confined to the New World tropics. The other nine sections are

Old World. Section Schnella is distinguished from Caulotretus by its five-nerved or inconspicuously nerved

calyx, an indehiscent or tardily dehiscent, thin-valved fruit, the stigma usually peltate or capitate, and the

pollen sexine with conspicuous supratectal processes. In contrast, Caulotretus has a conspicuously 10- or

15-nerved calyx, a dehiscent, usually woody fruit, the stigma usually oblique, and the pollen sexine merely

rugulose or perforate. Bauhinia herrerae (Britton & Rose) Standl. & Steyerm., a species placed in section

Caulotretus by Wunderlin et al. (1987) differs in several characters which otherwise distinguish the two

sections by having a peltate stigma and an indehiscent, thin-valved fruit as found in section Schnella, but

has the distinctive calyx and pollen characters that clearly place it in section Caulotretus.

Recent molecular data based on ITS sequences of nuclear ribosomial DNA (Hao et al. 2003) suggests

that subgenus Phanera is polyphyletic and that the New World Phanera forms a clade distinct from the rest

of subgenus Phanera in the Old World. Recently Lewis and Forest (2005) recognized Barklya F. Muell.,

Gigasiphon Drake, Lasiobema (Korth.) Miguel, Phanera Lour., Piliostigma Hochst., and Tylosema (Schweinf.)

Torre & Hillc. as genera distinct from Bauhinia. However, that classification is only partly supported by

molecular data (Lewis & Forest 2005). Since further study is needed to resolve this issue, it seems best at

this time to recognize Bauhinia s.l. rather than several segregate genera.

Two groups within section Schnella can be distinguished, although they are not formally recognized

here. Bauhinia lilacina, B. radiata, and B. trichosepala differ from the others in characters of the calyx and

pollen morphology. The n of these species is prominently 5-nerved and the pollen with the supratectal

ned on rid the colpi margins. The second group consists of B. flexuosa, B. maximiliani,

B. microstachya, B. poiteauana, and B. smilacina. d have an inconspicuously 5-nerved calyx and the pollen

with tl jually disposed on the tectum between the colpi and with no prominent

ridges ione the adini margins.

J. Bot. Res. Inst. Texas 3(2): 619 — 628. 2009

620 [| | £a D.s . in Le EF "IP £T. ia

Bauhinia L. subg. Phanera (Lour.) Kurz sect. en (Raddi) Benth., i ,in Benth. & Hook. f., Gen. Pl. 1:576.

1865. Schnella Raddi, Quar. Piant. Nuov. Bras. 32. 18 th. Tyre: Schnella macrostachya Raddi

[=Bauhinia radiata Vell.]. Lectotype designated by Britton and Rose (1930).

Caulotretus Rich. ex Schott, in Sprengel, Syst. Veg. 4(2):406. 1827. Caulotretus sect. Eucaulotretus Endlicher, Gen. Pl. 1317. 1840, nom.

inadmiss a El Fad 3. 4 s x (un "n" [ D dto. a] : (S 1 t) Ce J ]

23 on | ed

Woody vines with woody, nodal dd s. 1 pu minute intrastipular trichomes, the blade en-

tire or bilobate, t the midrib with ] tely d y

caducous. Inflorescences terminal or al and me spiciform racemes, these often paniculiform

through reduction of the leaves. Flowers with the hypanthium turbinate, nectariferous disc absent; calyx

veins, the stipules linear-lanceolate,

cupulate or campanulate, either conspicuously 5-nerved with the nerves extending beyond the lobe apex as

a short mucro or as a setaceous projection, or inconspicuously 5-nerved and with the nerves not extending

beyond the lobe apex, the calyx at anthesis either splitting irregularly into 23 lobes as the expanding pet-

als force the calyx to rupture or expanding into 5 lobes, nectariferous; petals 5, clawed, subequal; stamens

10, free, all fertile, in two ranks with the alternate ones (inner whorl) slightly shorter, the filaments slightly

upward curved, the anthers oblong, opening by longitudinal slits; pollen spheroid, 3-colporate, with the

tectum dam or psa uA often microperforate, with spheroid supratectal processes; ovary with a

short gyno thium, the style terminated by a small, peltate, and asymmetric stigma.

Fruit med samaroid, ihdeltiscent or tardily dehiscent, with the calyx persistent; seeds 102), with 2

short, funicular, aril-lobes.

1. Leaves otnowenng or as SIE entire, E —)5(-7)-nerved.

2. Fruit narrowly elliptic (at least 4 t wide), 6.5-7 cm long, chartaceous B. smilacina

2. ips elliptic- citan (2 times as long as wide), 3-6 cm long, subcoriaceous

Flower buds ca. 3 mm long; fruit 2.5-3.5 cm long B. poiteauana

: Flower buds 4-5 mm long; fruit 4.5-6 cm long B. maximiliani

. Leaves of flowering or Pa stems obse, 7- to oF nerved.

4. Calyx inconspicuously 5-nerved, t uds 2-5 mm long.

5. Branches ir cide Pede leaves (1.5-)2.5-3. Bi cm long; petals glabrous: p 2.5-3 cm long

B. flexuosa

5. Branches not flexuous; leaves (4-)7-8(-10) cm long; petals strigose along the median nerve on the

outer surface; fruit 4-7 cm long B. microstachya

4. Calyx conspicuously 5-nerved, the nerves extending beyond the apex as a mucro or a setaceous pro-

jection; flower buds 6-15 mm lon

6. Petals white; leaves 9-11 cm long B. radiata

6. Petals pink or lilac; leaves 4-8 cm lon

7. Calyx nerves undulate, winged toward the apex, extending 3-5 mm beyond the apex B. trichosepala

7. Calyx nerves not undulate nor winged, extending no more than 0.5 mm beyond the apex B. lilacina

Bauhinia flexuosa Moric., Pl. Nouv. Amer. 80, t. 53. 1840. Schnella pou inii) ware aaa Bot. Ml 5:572.

1846. d Pen (Moric. ds LR pai a : 6. 2006. Tyre: BRAZII ra Acurua,

E t G studied by Moricand: G EML Gl,

KI, NY!, P!; photo ak US). ! :

Woody, tendriled vine climbing to 5 m; young branches subterete, flexuous, strigose, becoming glabrate in

age, with conspicuous lenticels, the older stems flattened. Leaves with the blade bilobed 1/8—1/3 the length,

suborbicular, (1.5—-)2.5-3.5 cm long, (1.5-2)2-4.5 cm wide, 7-nerved, chartaceous, the lobes parallel or

slightly divergent, the i apex adis the blade base rounded to cordate, the upper surface glabrous or

glabrate, the or hirsute, especially along the veins, glaucous, the petiole 1-1.5

cm long, D ii do the sones Bose: DNA ca. 0.5 mm Ru SEDE Inflorescence terminal or

short, dense, spiciform racemes, if 2-10 cm long, densely

red-brown strigose; bids see 23 mm long and wide, densely dose m the pedicel 1.5—2

mm long, the bract and bracteoles lanceolate, ca. 0.5 mm long; hypanthium depressed-cupulate, ca. 1 mm

long, the inner surface glabrous; calyx cupulate, 2-3 mm long, inconspicuously 5-nerved, the nerves not

eee ae d Eilers, Bauhinia sut pi ct. Schnella 621

extending beyond the apex, 2- to 5-lobed, the lobes triangular-ovate, the inner surface sparsely red-brown

strigose; petals white, broadly spatulate, 6-7 mm long, ca. 4-5 mm wide, the claw 1-1.5 mm long, glabrous,

the blade concave, the apex rounded, glabrous; stamens 3-5 mm long, the filament 3—4 mm long, glabrous,

the anther oblong, ca. 1 mm long, glabrous; pollen spheroid, 3-colporate, the tectum microrugulate, often

microperforate, with depressed, spheroid supratectal processes; gynoecium ca. 5 mm long, the gynophore

ca. 1 mm long, red-brown short-hirsute, the ovary 2.5-3 mm long, red-brown short-hirsute, the style ca.

1.5 mm long, glabrous. Fruit elliptic-reniform, 2.5-3 cm long, 1.2-1.4 cm wide, the apex apiculate with the

persistent, somewhat oblique style or style base, the valves chartaceous, reticulate, light brown or reddish

brown strigose, especially on the margin, 1-seeded, the gynophore 3-4 mm long, densely red-brown strigose;

seeds suborbicular to broadly reniform, ca. 9 mm long, ca. 11 mm wide, ca. 1 mm thick.

Distribution and ecology.—Brazil (Bahia and Minas Gerais). In Caatinga and seasonally dry forests at ca.

450 m. Flowering and fruiting in April.

This rare species is most closely related to B. microstachya but is distinguished by its smaller leaves

(1.5—)2.5-3.5 cm long vs. (4—)7-8(-10) cm long), its smaller fruit (2.53 cm long vs. 4-7 cm long), and by

its maaan! flexuous stems.

ac] nT rm T 1T I A dean Tl pi

J. PDD AFTT Rah: 1 : [4 Dor T .

rama, ca. 450 m, 19 id 1980, Labo et al. 21551 (K, MEXU).A G | Infici ] 1C 12 Apr 1882, Glaziou 12622

(G, K, P); between d Caraça, s.d., Glaziou 13739 (P); between Inficionado and Caraga, 1883, Clio 13740 (G, K, P).

Bauhinia lilacina Wunderlin & Eilers, sp. nov. (Fig. 1). Te: BRAZIL. Bara: Km 5 on Núcleo Colonial de Una-Sao José

Highway (BR 101), 4 Aug 1977, Silva Mattos & Hage 81 (HoLotyre: CEPEC!; 1sorYPES: K!, US!).

Differt a Bauhinia radiata Vell. petalis lilacinis, florum bracteis et bracteolis triangularibus | ibus i picuis, stylis dense ru-

brotomentosis.

Woody, tendriled vine Einb gto T m; Li a SMS IS ce oe tomentose, becoming

sparsely so in age, with te. Leaves with the

blade bilobed 1/3 the ieie. siboroicular, ca. " 3cm as ca. 7 cm vide, 9 merged subcoriaceous, the lobes

slightly divergent, the lo! the blad cordate, the upper surface sparsely white-pilose, the

lower surface red-brown iomietitése; especially on the veins, the petiole ca. 2.6 cm long, red-brown tomen-

tose, the stipules lanceolate, 2 mm long, red-brown tomentose. Inflorescence terminal or subterminal and

axillary, in spiciform racemes, 10—14 cm long, red-brown tomentose; buds broadly elliptic, 6-7.5 mm long,

4—6 mm wide, red-brown tomentose, with 5 mucronate nerve tips extending ca. 0.5 mm beyond the apex,

the pedicel ca. 1 mm long, the bract and bracteoles linear-lanceolate, 2-2.5 mm long; hypanthium cupulate,

ca. 2 mm long, the inner surface glabrous; calyx campanulate, ca. 4 mm long, conspicuously 5-nerved, the

nerves usually extending ca. 0.5 mm beyond the apex, irregularly splitting into 5 lobes, the lobes triangular,

the inner surface red-brown strigose; petals lilac-colored, spatulate, 13-15 mm long, 4-6 mm wide, the

claw 1 mm long, glabrous, the blade concave, the apex emarginate, with a tuft of trichomes at the apex and

scattered trichomes near the base; stamens 10-12 mm long, the filament 7-9 mm long, glabrous, the anther

oblong, ca. 3 mm long, glabrous; pollen spheroid, 3-colporate, the tectum microrugulate, with supratectal

processes along the marginal ridges of the colpi; gynoecium ca. 11 mm long, the gynophore 1-2 mm long,

densely red-brown tomentose, the ovary ca. 4.5 mm long, densely red-brown tomentose, the style 4—4.5

mm long, densely red-brown tomentose except near the stigma. Fruit not seen.

PARATYPE: BRAZIL. Bahia: without ific locality, s.d., Belém 3675 (CEPEC)).

Etymology.—The specific epithet is in reference to the lilac-colored petals.

Distribution and ecology.—Brazil (Bahia). In secondary forest. Flowering in August.

This distinctive species is one of two species of section Schnella in Bahia with colored petals. It is

distinguished from B. trichosepala, the other species with pink- or lilac-colored petals, by its smaller petals

(13-15 mm long vs. 25-30 mm long), its sepals not elongated into setaceous tips, and its tomentose style.

Bauhinia maximiliani Benth., in Martius, Fl. Bras. 15(2):207. 1870. Binaria maximiliani (Benth.) A. Schmitz, Bull.

622

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Fic. 1. Bauhinia lilicina (isotype, US).

Wimdorlj d Eilers, Bauhinia sul PI t. Schnella 623

Jard. Bot. Natl. Belg. 43(3—4):404. 1973. Tyre: BRAZIL. Espirito Santo: Villa Nova de Almeida, s.d., Wied-Neuwied s.n. (HOLOTYPE:

BR; 1soTYPE: M; photo ex M: RB!).

Woody, tendriled vine; young branches subterete, red-brown strigose, soon becoming glabrate, the older

stems flattened, undulate, sometimes becoming perforate. Leaves with the blade entire, elliptic to broadly

lanceolate, 6-12 cm long, 3—6.5 cm wide, (3-)5-nerved, chartaceous to subcoriaceous, the apex obtuse to

acuminate, the base cuneate to subcordate, the upper surface glabrous, the lower surface sparsely short-

hirsute to XE ONE lighter in mer iun the upper surface, the n 0.5-1 cm Ac pa Spaey Sabri ALUA

f t glabra a x] "n

"E te, lanceolate, ca. 1 mm lon

or subterminal and axillary, in Le racemes, often iia, 10-25 cm long, densely red-brown

strigose; buds subglobose, 4-5 mm long and wide, densely red-brown strigose, the pedicel ca. 1 mm long,

the bract and bracteoles ovate-lanceolate, 0.5-1 mm long; hypanthium depressed-cupulate, ca. 1 mm

long, the inner surface glabrous; calyx cupulate, 3-4 mm long, inconspicuously 5-nerved, the nerves not

extending beyond the apex, (2-)3-lobed to about the middle, the lobes triangular-ovate, the inner surface

sparsely brown-strigose; petals white, spatulate, 4.5—6 mm long, 3.5-4 mm wide, the claw 1-1.5 mm long,

glabrous, the blade slightly concave, the apex rounded, sparsely red-brown strigose on the outer surface or

glabrous; stamens 4-6 mm long, the filament 4-5 mm long, glabrous, the anther oblong, ca. 1.5 mm long,

glabrous; pollen not seen; gynoecium ca. 5 mm long, the gynophore 2-3 mm long, red-brown short-hirsute

to tomentose, the ovary 2.5-3.5 mm long, red-brown, short-hirsute, the Der ca. 5 mm long, aan

Fruit (immature ?) elliptic-reniform, 4.5—6 cm long, 3.5-4.5 cm wide, tl with t]

oblique, style base, the valves subcoriaceous, inconspicuously reticulate, red: brown strigose, 1- à the

gynophore 4—7 mm long, red-brown strigose; seeds not seen.

Distribution and ecology.—BRAZIL (Rio de Janeiro, Bahia, and Espírito Santo). In wet forest. Fruiting in

June and July.

Originally described from flowering material and presently represented by only a few mainly fructifer-

ous specimens, B. maximiliani is still poorly known. It resembles B. poiteauana and B. smilacina by its entire

leaves and small, white flowers. It differs from B. poiteauana by its larger fruit (4.5—6 vs. 3.5—4.5 cm long),

though similarly reniform, and from B. smilacina by its distinctive reniform fruit (vs. narrowly elliptic in B.

smilacina). Fruiting material from Rio de e reported as B. smilacina by Vaz (1989) is placed here.

d: BRAZIL. Bahia: al C to Itabuna, 29 Jul 1965, Belém 1409 (CEPEC, IAN, NY, US); Rio

Gongog on 100-500 m, 1 O Nov 1915, "m 174 (US); between Camaca and Itabuna, 29 Jul 1965, Lanna Sobriho 972 (US);

Km 17 on BR 101, ca. 45 km S of Itabuna, 2 Jun 1979, Mori 11897 (CEPEC, K, US);

EEs Una and Rio en 16 Jun 1971, Pinheiro 1363 (CEPEC, K, US); Km 27 on Sáo José highway (junction of BR 101), Una, near

Fazenda Piedade, 20 Jul 1981, Silva et al. 1296 (CEPEC, K). Rio de Janeiro: Serra da Estrella, 2 May 1880, Glaziou 11912 (G, K, P).

Bauhinia microstachya (Raddi) J.E Macbr., Contr. Gray Herb. 59:22. 1919. Schnella microstachya Raddi, Mem.

Mat. Fis. Soc. Ital. Sci. Modena, Pt. Mem. Fis. 18:418. 1820. Phanera microstachya (Raddi) L.P. Queiroz, Neodivers. 1:6. 2006. Tyre:

BRAZIL. Rio DE JANEIRO: Raddi s.n. (HOLOTYPE: FI).

T) T r

Bauhina tomentosa Vell., Fl. Flumin. 171. 1829 («1825»); non L., 1753. Tree: BRAZII J [ , here designated: Vellozo,

Lu E. id 4:t. 84. 1831 («1827»)].

Acad. Imp. Sci. St -Pétersbourg, Sér. 6, Sci Math., Seconde Pt. Sci. Nat. 4:109. 1838. Tire: BRAZIL.

Rio wen near ane 1823, Riedel s.n. (HOLOTYPE: LE; isoTYPES: G!, P!)

Bauhinia bahiensis Bong., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 4:114. 1838. Bauhinia langsdorffiana

r. bahiensis (Bong.) Benth., in Martius, a Bras. bun 204. ok 0. eae microstachya (Raddi) J.E Macbr. var. bahiensis (Bong.)

He Macbr., Contr. Gray Herb. 59:22. 1919 bahiensis (Bong.) A. Schmitz, Bull. Jard. Bot. Natl. Belg.

Bauhinia e Vogel, Linnaea 13:310. 1839; non Rottler ex Sprengel, 1807; nec Kónig ex Wallich, 1831. Bauhinia vogeliana Steud.,

nel. Bot., ed. 2. 1:192. Ho Schnella spicata A. Schmitz, Bull. Jard. Bot. Natl. Belg. 43:402. 1973, nom. illegit. Tyre: BRAZIL

SS (HOLOTYPE: B, d here designated: K!; isotyres: G!, P!; photo ex B: US!; photo ex K: US!).

Bauhinia polyantha Vogel, imie 13: 310. 1839. Tree: BRAZIL. Rio DE Janeiro: near Rio de Janeiro, s.d., Richard s.n. (HoLotyre: B, de-

stroyed).

Bauhinia na Harms, Verh. Bot. Vereins Prov. Brandenburg 48:165. 1907. Tyre: BRAZIL. Amazonas: near Juruá, May 1901, Ule 5481

LoTYPE: B, destroyed).

I [| £ sl n.a Li In LI PEA iT. im”

624

Bauhinia n Lundell, Wrightia 3:120. 1965. Tyre: BELIZE. ToLeno District: between Condemn Branch Pine Ridge and Moffredye

La , in cohune ridge, 15 Sep 1946, Gentle 6047 (HoLotyeE: LL!; isorvee: MO!)

Bauhinia rra (Raddi) J.E Macbr. var. massambabensis Vaz, Pesquisas, Bot. 44:100. 1993. Tyre: BRAZIL. Rio DE J

987, M. Gomes 201 (HOLOTYPE: RB

Woody, tendriled vine climbing to 20 m or more; young branches subterete, densely red-brown strigose,

soon becoming glabrous, with somewhat conspicuous lenticels, the older branches flattened, with the center

undulate and sometimes perforate. Leaves with the blade bilobed 1/8—7/8 the length, suborbicular to ovate,

(4-)7-8(-10) cm long, (4.6—)5—6(8) cm wide, 9-nerved, chartaceous to coriaceous, the lobes parallel to widely

divergent, the lobe apex rounded, acute, or acuminate, the blade base cordate to rounded, the upper surface

with a few scattered white to tan trichomes or glabrous, the lower surface red-brown tomentose, especially

when young, the petiole (0.8—)1-3(-5. Ss cm uL. red- prow) tomentose, the ces lanceolate, ca. 3 mm

long, red-brown tomentose. Inil I land axillary, in dense, spiciform racemes,

often paniculiform, 10-23 cm long, densely red-brown tomentose; buds subglobose, slightly asymmetric, ca.

5 mm long and wide, red-brown tomentose, the pedicel ca. 0.5 mm long, the pant añ bracteoles triangular,

ca. 1 mm long, ca. 0.5 mm wide; hypanthium shallowly cupulate, ca. 1 mm long, the inner surface glabrous

or with a few trichomes at the base of the gynophore; calyx campanulate, ca. 5 mm long, inconspicuously

5-nerved, the nerves not extending beyond the apex, (2)3-lobed, the lobes triangular, the inner surface

with short, appressed trichomes; petals white, spatulate, 5-7 mm long, 2.5-4 mm wide, the claw ca. 2 mm

long, red-brown strigose, the blade slightly concave, the apex rounded to obtuse, usually red-brown strigose

along the median nerve on the outer surface,; stamens 7-10 mm long, the filament 6-8 mm long, glabrous,

the anther oblong, ca. 2 mm long, glabrous; pollen spheroid, 3-colporate, the tectum microrugulate, often

microperforate, with spheroid, supratectal processes; gynoecium 4.5-5 mm long, the gynophore to ca. 0.5

mm long, sparsely red-brown tomentose, the ovary ca. 2 mm long, densely red-brown tomentose, the style

ca. 2 mm long, sparsely red-brown tomentose. Fruit elongate-reniform, (4-)5—7 cm long, 1.1-1.8 cm wide,

the apex and base acute to rounded, the valves chartaceous, conspicuously reticulate, sparsely to densely

red-brown or sometimes red-tomentose, particularly on the margins, 1(2)-seeded, the gynophore 2-5 mm

long, red-brown strigose; seeds elliptic-reniform, 18-20 mm long, 8-10 mm wide, ca. 1 mm thick.

Distribution and ecology.—Mexico, Guatemala, Belize, Panama, Venezuela, Bolivia, Peru, Paraguay,

and Brazil. In wet to moist high forest at 0-1,200 m. Flowering March to September; fruiting January to

November.

Although several segregates at the species or varietal level have been proposed by previous workers,

B. microstachya is treated here as a single, wide-ranging, polymorphic species. It is most closely related to

B. flexuosa of the caatinga of Bahia and Minas Gerais, Brazil, but is distinguished from that species by its

larger leaves (4-9 cm long vs. 1.5-3.5 cm long), its larger fruit ((4—)7-8(10) cm long vs. 2.5-3 cm long), and

its lack of nd flexuous stems.

+ = ] ME CO cL: p 7 i Chansayab, JE Dal 1 1 ET] T ,340

m, 7 Nov 1985, Martes 15072 (WIS). Oaxaxa: Cerro Buenos Aires, W side of the Temazcal Dam (Miquel Aleman), 60 m, 6 Mar 1986,

Cortes et al. 182 (MEXU); 500 m SE of the spillway of the Temazcal Dam, 70 m, 24 Nov 1986, Cortes et al. 573 (MEXU). Veracruz: along

road 5.5 km E of Tebanca (5.5 km E of east side of Lago Catemaco), 800 m, 5 Jul 1980, Nee & Hansen 18853 (F, USF). GUATEMALA

El Petén: Km 72 of Poptun-Santa Elena Road, in high forest, 24 Oct 1966, Contreras 6495 (US). PANAMA. Darién: Rio Cerro Yaviza,

near Yaviza, near sea level, s.d., Duke & Bristan 422 (MO, NY, US). VENEZUELA. ee Rio inar area, edel the Yanomani,

5 Oct 1978, Fuentes 1103 (US). Miranda: del Bachiller, near east end, between Q dQ Santa Cruz, S

of Santa Cruz, 18-20 m, 18-19 Mar 1978, Steyermark & Davidse 116484 (MO, USF). PERU. Loreto: Florída, Río Putumayo, at mouth

of Río Zubineta, ca. 200 m, Mar-Apr 1931, Klug 2067 (BM, K, US). Madre de Dios: edge of Cocha Cashu, ox-bow lake of Río Manu,

delia Nacional del Manu, 12 eas 1973, Foster 2571 (K, as = m Chazuta, Río Huallaga, ca. 260 m, Apr 1935, Klug 4060 (BM,

US). BRAZIL. A mazon O y 1851, 2d 1522 nad K, B ne ca. 28 km N of

Seabra, road to Agua de Rega, ca. id ,000 m, 27 Feb 1971, Irwin et al. 31164 (K, Us Espirit tama, 9-1

Aug 1965, Lanna Sobriho 4598 (USF). Minas Gerais: State Agricultural School, Vicosa, 6 Nov 1958, invi 2029 (NY). Paraná: oe =

Cedro, 7 Mar 1968, Hatschbach 18664 (US, USF). Rio de Janeiro: Jurujuba, 19 Nov 1873, Glaziou 6841 (K

S of São Miguel d'Oeste, 600—700 m, 23 Feb 1957, emu dices 11715 (G, K, P, d BOLIVIA. La Paz: ien ca. 600 m, May 1886,

Rusby 952 (BM, K, US). PARAGUAY. San Ped g the Rio Alto Parana , 1909-1910, Fiebrig 6201 (G, K, US).

Wunrarl; ! Fil , Bauhini l PI t. Schnella 625

Fasi

Bauhinia poaa ice mae 13:309. 1839. Tee: F 1844, Poiteau s.n. (HOLOTYPE: B, destroy

here designated: K!; i USI; photo ex K: US!).

Woody, tendriled vine "e to 25 m; young branches subterete, red-brown strigose or puberulent, soon

becoming glabrate, with * conspicuous lenticels, the older stems flattened, undulate, sometimes becoming

perforate with age. Leaves with the blade entire, elliptic to broadly lanceolate, 8-12 cm long, 3.5-6 cm

wide, 3- to 5-nerved, subcoriaceous, the apex acuminate, the base rounded to subcordate, the upper surface

glabrous, the lower surface with scattered erect or appressed trichomes, the petiole 7-10 mm long, red-

brown tomentellous, the stipules ovate, ca. 1 mm long, red-brown tomentellous. Inflorescence terminal or

subterminal and axillary, in spiciform racemes, sometimes paniculiform, 11-12(—25) cm long, red-brown

tomentellous; buds subglobose, ca. 3 mm long and wide, red-brown tomentellous, the pedicel ca. 1 mm

long, the bract and bracteoles triangular-lanceolate, ca. 0.5 mm long; hypanthium depressed-cupulate, ca.

1 mm long, the inner surface sparsely appressed red-brown strigose apically; calyx campanulate, 3-4 mm

long, inconspicuously 5-nerved, tl t extending beyond the apex, 3-lobed, the lobes triangular, the

inner surface sparsely red-brown strigose; petals white, broadly spatulate, 4-6 mm long, 3-4 mm wide, the

claw ca. 1 mm long, sparsely red-brown strigose on the inner dase thg dn concave, ini as rounded,

sparsely red-brown pilose on the outer surface near the apex and

6-8 mm long, the filament 5-6 mm long, glabrous, the anther - oblong, ca. 2 mm long, aloe pollen

spheroid, 3-colporate, the tectum mi late, often mi forate, with spheroid, supratectal processes;

gynoecium 4-5 mm long, the a mophóres to 0,5 mm long, densely red-brown hirsute, the ovary ca. 1 mm

long, densely red-brown hirsute, the style 2-3 mm long, glabrous. Fruit elliptic-reniform, 2.5—3.5 cm long,

1.5-1.7 cm wide, the apex obliquely apiculate with the persistent style base, the valves subcoriaceous,

inconspicuously reticulate, reddish, sparsely red-brown strigose, 1-seeded, the gynophore ca. 2 mm long,

red-brown tomentose distally; seeds suborbicular, ca. 10 mm long, ca. 9 mm wide, ca. 1 mm thick.

Distribution and ecology.—French Guiana and Brazil (Amapá). In moist to wet forests at 0-200 m.

Flowering October to December; fruiting in January and February.

This species is distinguished from the other entire-leaved species, B. maximiliani and B. smilacina, by

its smaller fruits.

1l. FRENCH GUIANA Cayenne, 5 d., Martin s.n. (BM, K, P); Approuague, l ee bani on the Crique

Al 25 Jan 1967, Oldeman 2320 (US); C ¡ River, between he Habillé ] t Chi c 1967, Oldeman 3547

ed US); bank of the Camopi River (tributary of FO iver), near Saut Tampac, 6 Dec 1967, Oldeman 2567 din bank of the Crique

Inipi 3 km from the Camopi River, 7 Feb 1968, Old & Sastre 71 (US); Cayenne, s.d., Perrottet s.n. (P). Without precise locality: 1838,

Lepri: 348 (P); s.d., i inn s.n. (P); di ipii 188 (G); 1821, Perrottet 196 (G). BRAZIL. Amapá: along the Rio Oiapoque, 4 Feb

50, Fróües 25839 (P) i At. Tipac, 200-250 m, 15 Oct 1960, Irwin 48740 (K, US).

Bauhinia radiata Vell., Fl. Flumin. 170; 4: pl. 81. 1829 (“1825”). Ter: BRAZIL: (lost).

Bauhinia acuminata Vell., Fl. Flumin. 171. 1829 (“1825”); non L., 1753. Tyee: BRAZIL. Rio DE Janeiro: Rio Taguaí (lost).

Bauhinia raddiana Bong., Mém. Acad. Imp. Sci. St-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 4:111. 1838. Tyre: BRAZIL

(LECB?)

Raddi, Mem. Reale Accad. Sci. Modena 18: Fis. 411. 1820. Tyre: BRAZIL: without precise location: s.d., Raddi s.n.

OLOTYPE: FL: 1 ISOTYPE: GI).

Bauhinia verrucosa Vogel, Linnaea 13:311. 1839. Tree: BRAZIL. R J I Rio de Janei d Cabo Frio, Sello s.n. (HOLOTYPE:

B, destroyed).

Bauhinia raddiana Bong. var. brachystachya Benth., in Martius, Fl. Bras. 15(2):203. 1870. Schnella macrostachya Raddi var. brachystachya

(Benth.) A. Schmitz, Bull. Jard. Bot. Natl. Belg. 43:402. 1973. Tyee: BRAZIL. Rio DE JANEIRO: base of Monte Corcovado, May 1839,

oe m (HOLOTYPE: K!; isotyes: G!, P!)

, in Martius, Fl. Bras. 15(2):209. 1870. Binaria leiopetala (Benth.) A. Schmitz, Bull. Jard. Bot. Natl Belg 43:406.

1973. Tre: BRAZIL. M G d

Velhas, s.d., Warming [301] (HoLorvre: K; isoryPE: P!).

Woody, tendriled vine climbing to 8 m; young branches subterete, densely red-brown tomentose, becom-

ing glabrate in age, with conspicuous lenticels, the older stems flattened and sometimes perforate. Leaves

with the blade bilobed 1/4 to 1/2 the length, suborbicular, (8.1-)9-11 cm long, (6.6—)8-11 cm wide, 7- to

9-nerved, subcoriaceous, the lobes divergent, the lobe apex rounded, the blade base shallowly cordate,

626 Journ t tani i Texas 3(2)

the upper surface sparsely white-pilose, the lower surface short red-brown tomentose, especially on veins,

usually glaucous, the petiole 2-6 cm long, red-brown tomentose, becoming glabrate with age, the stipules

Pm to ovate, ca. 2 mm, d brown tomentose. Inflorescence terminal or subterminal and axillary, in

liform, 8-15 cm long, red-brown tomentose; buds ovate, ca. 6 mm

long, ca. 14 mm wide, ced brown re with 5 mucronate nerve tips extending ca. 0.5 mm beyond the

apex, the pedicel ca. 1.5 mm long, the bract lanceolate, ca. 2 mm long, the bracteoles triangular-lanceolate,

ca. 1 mm long; hypanthium cupulate, ca. 2 mm long, the inner surface sparsely red-brown tomentose near

the base of the gynophore; calyx campanulate, ca. 4 mm long, strongly 5-nerved, the nerves extending

0.5-1 mm beyond the apex, conspicuously 5-lobed, the lobes triangular, the inner surface red-brown to-

mentose; petals white, broadly spatulate, 10-11 mm long, 4-5 mm wide, the claw ca. 5 mm long, glabrous,

the blade concave, the apex emarginate, with short, red-brown trichomes along the top margin, stamens

12-13 mm long, the filament 10-11 mm long, glabrous, the anther oblong, ca. 2 mm long, glabrous; pollen

spheroid, 3-colporate, the tectum microrugulate, with supratectal processes along the marginal ridges of

the colpi; gynoecium 10-12 mm long, the gynophore ca. 2 mm long, red-brown appressed short-hirsute

on the distal 1/2, the ovary ca. 4 mm long, appressed, short-hirsute, the style ca. 6 mm long, red-brown

short-hirsute predominantly along the distal, ventral margin and the base. Fruit subelliptic, 6-10 cm long,

3.5-3.8 cm wide, slightly asymmetric, the apex acute with the oblique persistent style or style base, the

valves coriaceous, conspicuously reticulate, glabrous or with a few appressed, red-brown trichomes near

the gynophore, 1(-2)-seeded, the gynophore 1.5-2 cm long, appressed red-brown hirsute toward the base;

seeds suborbicular, 8-12 mm long and wide, 0.5-1 mm thick.

Distribution and ecology.—Brazil (Minas Gerais and Rio de Janeiro). In moist forests at 0-700 m. Flower-

ing in April and May.

This species is most closely related to B. lilacina and B. trichosepala from which it is easily distinguished

by its white petals.

Specimens examined. BRAZIL. Minas Gerais: Vicosa, Agricultural College Lands, Campo Alegre, Alto de Samambaia, 685 m, 30

May 1930, Mexia 4772 (BM, G, P, US, WIS). Rio de Janeiro: Rio de Janeiro, 1849, Armstrong s.n. (K); Jun 1767, Commerson 201 (P); Rio

de Janeiro, 1867, Cunningham s.n. (KY, Estado do Guanabara, Morro de D. Marta, 4 Apr 1961, Duarte 5532 (K); s.d., Endlicher s.n. (P);

Larangeiros, near Cosme Velho, 20 Apr 1867, Glaziou 1144 (P); s.d., Glaziou 3720 (K, P); Corcovado, 6 May 1873, Glaziou 6513 (G, K,

P); Rio de Janeiro, s.d., Graham s.n. (K); s.d., Lelande s.n. (P); s.d., Lépine s.n. (P); s.d., Loeferen 1263 (P); Rio de Janeiro, s.d., Miers 3113

(K); Rio de Janeiro, s.d., Miers 3536 (K); Corcovado, s.d., Miers 4645 (K); s.d., Nadeaud s.n. (P); Rio de Janeiro, May 1832, Riedel s.n. (G);

1816-1821, Saint-Hilaire 804 (P); 1836, Vauthier 81 (K, P); 1836, Vauthier 149 (G, P); s.d., Vauthier s.n. (P); 1844, Widgren s.n. (US).

Bauhinia smilacina (Schott) Steud., Nomencl. Bot. ed. 2, 1:192. 1840. Caulotretus smilacinus Schott, in Sprengel, Syst.

16. 4(2):406. 1827. Schnella smilacina (Schott) G. Don, Gen. Hist. 2:459. 1832. Tyre: BRAZIL. Rio DE Janerro: s.d. (fD, Schott

5632 (LecrorwrE, here designated: W!; isorvpe: US).

Woody, tendriled vine; young | bteret , strigose, soon | i labrate, with “ conspicuous len-

ticels, the older stems flattened, undulate, and sometimes perforate. Leaves ; with the blade entire, narrowly

ovate to lanceolate, 5-8 cm long, 2.5-3.5 cm long, (—)5(-7)-nerved, chartaceous to subcoriaceous, the

apex acuminate to mucronate, rarely obtuse, the oa Alo to cala the upper surface glabrous,

slightly rugose, the lower surface with a few appressed tri the base, otherwise glabrous, slightly

lighter in color than the upper surface, the petiole 7-1500) mm ie with a few appressed trichomes or

glabrous, the stipules obovate, ca. 1 mm long, glabrate. Inflorescence terminal or subterminal and axillary,

in spiciform racemes, 3-10 cm long, red-brown strigose; buds elliptic-globose, ca. 3 cm long, ca. 2.5 cm

wide, red-brown tomentose, the pedicel ca. 1 mm long the bract and bracteoles triangular-ovate, ca. 0.5

mm long; hypanthium depressed-cupulate, ca. 1 mm long, the inner surface sparsely red-brown pilose;

calyx campanulate, 3-3.5 mm long, inconspicuously 5-nerved, the nerves not extending beyond the apex,

3-lobed, the lobes triangular, the inner surface sparsely red-brown strigose; petals white, spatulate, 6-7

mm long, 2-3 mm wide, glabrous, the claw 1-2 mm long, the blade concave, the apex rounded; stamens

5-7 mm long, the filament 4.5—5.5 mm long, glabrous, the anther oblong, 1.5-2 mm long, glabrous; pollen

Windarli d Eilers, Bauhinia sul PI t. Schnella 627

spheroid, 3-colporate, tectum microrugulate, often microperforate, with depressed, spheroid supratectal

processes; gynoecium 4-6 mm long, the gynophore ca. 1 mm long, red-brown short-hirsute on the upper

part, the ovary ca. 1.5 mm long, red-brown short-hirsute, the style ca. 4 mm long, glabrous or with a few

scattered, appressed, red-brown trichomes. Fruit narrowly elliptic, 6.5-7 cm long, ca. 1.5 cm wide, slightly

curved, the apex with an oblique, persistent style base, the valves chartaceous, inconspicuously reticulate,

sparsely appressed, red-brown short-hirsute, 1-seeded, the gynophore ca. 7 mm long, red-brown hirsute;

seeds 15-16 mm long, 9-10 mm wide, 0.5-1 mm thick

Distribution and ecology.— Brazil (Espirito Santo and Rio de Janeiro). In wet forest. Fruiting in August.

This species differs from the other two entire-leaved species, B. poiteauana and B. maximiliani, by its

narrowly elliptic, chartaceous fruit which more closely resembles that of B. microstachya, to which it is ap-

parently closely related. The fruits of B. poiteauana and B. maximiliani are reniform or elliptic-reniform and

subcoriaceous.

|. BRAZIL. Espirito Santo: R Florestal de S , 10 Aug 1965, Belém c pue: NYL Linhares, Refügio

Biológico Sooretama, 9—10 Aug 1965, I Sobo 1056 (US, USF). Rio d i io de J , s.d., Burchell 1343 (K);

Janeiro, s.d., Burchell 1565 (K). Without precise isiy s.d., Elsa n. (P); s.d., Glaziou 2554 (K, P); s. d. pecie d n. (P).

Bauhinia €—— (L.P. Queiroz) Wunderlin & Eilers, comb. nov. Basionym: Phanera trichosepala L.P. Queiroz,

Neodivers. 1:7. 2006. Tere: BRAZIL. Banta. id Barbosa, Serra do Orobó, trilha para o Patio das Orquídeas, 12E 18"S, 40E 29"W, 8

Feb 2005, D. Cardoso 281 (HOLOTYPE:

Woody, tendriled vine; young | I bterete, red-brown tomentose, glabrate in age, with “ conspicuous

lenticels, the older stems flattened, undulate, often perforate. Leaves with the blade bilobed 1/3 to 1/2 the

length, suborbicular, often broader than long, 4-8 cm long, 5-10 cm wide, 7- to 9-nerved, coriaceous, the

lobes slightly divergent, the lobe apex rounded, the blade base deeply cordate, the upper surface usually

tomentellous on the midvein and the pulvinus, otherwise glabrous, rugose, the lower surface red-brown

tomentellous, especially on the nerves, subglaucous, the petiole 13.5 cm long, red-brown tomentose, the

stipules linear, ca. 8 mm long, red-brown tomentellous. Inflorescence terminal or subterminal and axillary,

in spiciform racemes, 15-20 cm long, red-brown tomentose; buds elliptic-ovate, 1-1.5 cm long, ca. 1 cm

wide, red-brown tomentose, 5-angled toward the base, undulate-winged toward the apex, each wing end-

ing in a twisted, setaceous tip 3-5 mm long, the pedicel 2-3 mm long, the bract and bracteoles linear, 8-10

mm long, hypanthium cupulate, 2-3 mm long, the inner surface glabrous; calyx campanulate, 8-9 mm

long, conspicuously 5-nerved, irregularly splitting to half the length into 2 lobes, the lobes triangular, the

inner surface red-brown strigose; petals bright pink, spatulate, 2.5-3 cm long, 0.7-1.4 cm wide, the claw

ca. 5 mm long, glabrous, the blade elliptic-ovate, the apex rounded, the margin slightly crisped, red-brown

tomentellous toward the apex on the outer surface; stamens 2-2.5 cm long, the filament 1.5-2 cm long,

glabrous, the anther oblong, 3-4 mm long, glabrous; pollen spheroid, 3-colporate, tectum microrugulate,

often microperforate, with spheroid supratectal processes on the marginal ridges of the colpi; gynoecium

22-25 mm long, the gynophore ca. 4 mm long, sparsely red-brown tomentellous distally, the ovary ca. 5

mm long, sparsely red-brown tomentellous toward the base and along the ventral margin, the style ca. 14

mm long, glabrous. Fruit (immature) elliptic, 8.5—9.5 cm long, 3-3.5 cm wide, glabrous, 1-seeded, the

gynophore ca. 2.5 cm long, glabrous; mature seeds not seen.

Distribution and ecology.—Brazil (Bahia and Minais Gerais). In caatinga seasonally dry forests at

800-1,000 m. Flowering Feburary to May; fruiting April to July.

A distinctive species and one of two taxa of section Schnella with non-white flowers. It is distinguished

from B. lilacina, its most closely related species, by its larger petals clea mm long vs. 13-15 mm long in

B. bM its undulate winged sepals elongated into conspicuous setaceous tips, and its glabrous style.

1. BRAZIL. Bahia: Encruzilhada, margin of the Rio Pardo, 23 May 1968, Belém 3598 (CEPEC); Ibotirama, 14 Apr

1971, dos iui "dd pi a KM 20 on n dde from Conquista to Barra do Choca, 19 Feb 1973, dos Sant 76 (CEPEC) 27 km

BAO32 t 00 m, 4 Mar 1977, Harley et al. 19397 (K, US). Minas Gerais: Serra da

Alegria, near aca s.d. P 14623 (K, P).

r n. H IP LI FTE

628 J lofti te of Texas 3(2)

ACKNOWLEDGMENTS

This paper is based in part on an unpublished masters thesis completed at the University of South Florida in

1991 by the junior author. We thank the curators of BM, BR, CEPEC, F, G, IAN, K, M, MO, P, US, USF, and

WIS for loan of the specimens used in this study, and Bruce F. Hansen (USF) for his insightful comments

and suggestions which greatly added to this work. We also thank Rafael Torres Colín (MEXU) for his com-

ments on the manuscript and information on collections at MEXU and an anonymous reviewer.

REFERENCES

Britton, N.L. and J.N. Rose. 1930. Caesalpiniaceae. N. Amer. Fl. 23:201-349.

Hao, G., D.X. ZHANG, M.Y. ZHANG, L.X. Guo, and S.J, Li. 2003. Phylogenetics of Bauhini Phanera (Legumino

sae: Caesalpinioideae) based on ITS sequences of nuclear ribosomal DNA. Bot. Bull Acad. Sci. 44:223-228.

Lewis, G.P and F. Forest. 2005. Tribe Cercideae. In: G. Lewis, B. Schrire, B. Mackinder, and M. Lock. Legumes of the

world. The Royal Botanical Garden, Kew. Pp. 57-67.

Vaz, AMS. pa F. and H.C. be Lima. 1986. Bauhinia smilacina (Schott) Steudel (Leg. Caes.)-uma espécie POUCO

conhecida do Etado do Rio de Janeiro. Atas Soc. Bot. Brasil, Secc. Rio de Janeiro 3:69-75.

WUNDERLIN, R.P, K. Larsen, and S.S. Larsen. 1987. Reorganization of the Cercideae (Fabaceae: Caesalpinioideae).

Biol. Skr. 28:1-40,

REVISION OF BAUHINIA SUBGENUS BAUHINIA SECTION PAULETIA

SERIES ARIARIA (CERCIDEAE: CAESALPINIOIDEAE: FABACEAE)

Richard P. Wunderlin

TA a Pa ee

Department of Cell B cular Biology

a of South Florida

4202 East Fowler Avenue, BSF 218

Tampa, Florida 33 150, U.S.A.

rwunder@cas.usfedu

ABSTRACT

Korece arn

A taxonomic treatment of Bana Ea aoa section dM series ai is RES Seven paper are recognized with

" :

RESUMEN

de R hini } R hini Ye Panletia «serie Arinrin S iet 1 la

E e

descripción de Bauhini ld is Wunderlin del Ecuador. Se incl l de distril

The pantropical genus Bauhinia Linnaeus, with approximately 350 species, consists of four subgenera:

Bauhinia, Piliostigma (Hochst.) Kurz [=Elayuna (Raf.) Wunderlin et al], Barklya (F. Muell.) Wunderlin et al.,

and Phanera (Lour.) Kurz (Wunderlin et al. 1987). Bauhinia subgenus Bauhinia, as defined by Wunderlin et

al., consists of nine sections of which three (Bauhinia, Pauletia (Cav.) DC., and Amaria (S. Mutis) Endl. are

New World. A revision of section Amaria was recently pubrenee by Wunderlin (2006).

Section Pauletia, the largest section of the genus in 1 j s defined here, consists of five series

(Aculeatae Vaz & A.M.G Azevedo, Cansenia (Raf.) Wunderlin et al. Perl on (Mart.) Wunderlin et al., Pentandra

Wunderlin et al., and Ariaria (C. Cuervo Márquez) Wunderlin et al.), all New World. Series Acuminatae,

consisting of two Old World species and placed in Pauletia by Wunderlin et al. (1987), is transferred to

the Old World section Telestria. Series Aculeatae was recently described by Vaz and Azevedo Tozzi (2003).

This revision of series Ariaria is one of several proposed taxonomic treatments of the New World species of

Bauhina.

KEY TO THE SERIES OF BAUHINIA SECTION PAULETIA

. Plants unarme

2. Leaves Em the flower oin much reduced or absent, inflorescence a terminal raceme or

panicle, the plants never cauliflor er. Cansenia

2. Leaves subtending the flower ien normally developed, inflorescence leaf-opposed or the um

cauliflorous ser. Ariaria

1. Plants armed wit! | i | infrastipular excrescences.

3. Fruit indehiscent ser. Perlebia

3. Fruit dehiscent.

4. Fertile stamens 5, or if 10, then the alternate ones much reduced ser. Pentandrae

4. Fertile stamens 10, of slightly unequal length ser. Aculeatae

Bauhinia section Pauletia series Ariaria (C. Cuervo Márquez) woe et al., Biol. Skr. 28:13. 1987. Ariaria

C. C Márquez, Prehist. & Viajes 219. 1893. Tyre: Ariaria superba C. C juez [=Bauhinia tarapotensis Benth.].

Unarmed trees or shrubs. Leaves bilobed; adpetiolar intrastipular e enlarged, forming a laterally

flattened, obtuse-tipped, lanceolate structure. Calyx at first spathaceous, soon splitting to the hypanthium

into several lobes; fertile stamens 10; pollen 5—7-porate or -colpate, the sexine reticulate with infratectal

processes. Fruit dehiscent.

J. Bot. Res. Inst. Texas 3(2): 629 — 637. 2009

Pal Da a im LI Ma dion ndis £T ls F áo

Sa i

630 j l

. Plants wiin cauloneds i racemes.

C pane

2. n long; fruit 20-22 cm long B. flagelliflora

2. Cauliflorus racemes to d cm ‘ong; fruit 10- E em long B. esmeraldasensis

1. P Plants with ter minal MV Y fy brancl IT.

3. Leaf lobes with the apex acute or obtuse B. tarapotensis

3. Pa lobes with the apex rounded

eaf blades ovate 5-91 1)

E Lower surface of the leaf blade with pl | trichomes B. multinervia

5. Lower surface of the leaf blade with erect trichomes B. eilertsii

4. Leaf blades supomicilal 11-13- nerve

6. Leaf blades 8-13 cm long; y densel y tomentose; Colombia B. conceptionis

6. blades 14 24 cm long; ovary B. pichinchensis

1. Bauhinia conceptionis Britton & Killip, Ann. New York Acad. Sci. 35:160. 1936. Tyre: COLOMBIA. Choco: La

Conceptión, 15 km E of Quibdó, ca. 75 m, 20 Apr-23 May 1931, WA. Archer 2086 (HOLOTYPE: NY!).

Tree to 9 m; branches minutely tomentulose soon becoming glabrate. Leaves with the blade chartaceous,

suborbicular, 8-13 cm long, 10-13 cm wide, bilobed ca. 1/4 their length, the lobes slightly divaricate, the

base shallowly cordate, the lobe apex rounded, the upper surface glabrous, the lower surface minutely

tomentulose, at least on the nerves, de dd in CODE than the upper surface, 11-nerved, the petiole

3.5-4 cm long, minutely t tul g te; stir lanceolate pec HB) dni M ri

intrastipular excrescence to 2 mm long, the ot! | Infl | xillary,

2—4-flowered, the fl ired on a common peduncle, the rachis tomentulose; buds linear-lanceolate, ca.

5 cm long, obscurely 5 "e the apex apiculate, the peduncle 1-1.5 cm long, the pedicel 1.5-2 cm long;

bracts and bracteoles lanceolate, 2-3 mm long, caducous. Hypanthium tubular, 10-13 mm long, 10-nerved;

calyx at first spathaceous, soon irregularly splitting to the hypanthium into 2-5 lobes; petals white, sub-

equal, linear to linear-spathulate, ca. 5 cm long, 6-7 mm wide, glabrous or glabrate (sometimes with a few

trichomes on the claw), the margin slightly crisped, the claw ca. 1 cm long; fertile stamens 10, the filaments

of the outer whorl ca. 5 cm long, subequaling the petals, the inner whorl slightly shorter, irregularly con-

nate at the base to 5 mm into a staminal sheath, the staminal sheath tomentulose on the inner surface, the

anthers not seen; gynoecium ca. 5 cm long, subequaling the androecium, the ovary ca. 1.5 cm long, densely

tomentose, with sparse glandular trichomes, de stipe ca. 1.5 cm long, glabrate, the style ca. 1.5 cm long,

tomentose proximally, sparsely tomentulose to glabrate distally, the stigma obliquely capitate. Fruit not seen.

Distribution and ecology. —Colombia in Choco. Known Dus from the type collection. Habitat unknown.

Flowering in April.

Bauhinia conceptionis is closely related to B. tarapotensis, but differs by its suborbicular leaves with

rounded lobe apices and smaller flowers (6—7 cm long vs. 14—21 cm long in B. tarapotensis).

2. Bauhinia eilertsii Pulle, Recueil Trav. Bot. Néerl. 6:269. 1909. Tyre: SURINAME: Upper Suriname River near Dotti

Bergi, 9 Jul 1908, J. Tresling 86 (HOLOTYPE: Un.

Tree to 22 m: branches hirsute-tomentulose to glabrate when young, soon becoming glabrous. Leaves with

the blade chartaceous, suborbicular, 10-21 cm long, 10-21 cm wide, emarginate or bilobed to 1/3 their

length, the base cordate, the lobe apex rounded, the upper surface glabrous, the lower surface light brown

hirsute-tomentulose, slightly lighter in color than the upper surface, 9—11-nerved, the petiole (2. 54-56)

cm long, brown hirsute-1 tulose; stipules lanceolate, ca. 2 mm SUE Ben adpetiolar infrastipular

excrescence ca. 2 mm long, the otl te. Infl ] xillary, the flowers

paired on a common peduncle, sometimes short-racemose, rarely paniculate, or less commonly solitary by

abortion, the rachis brown-tomentose; buds linear, 7-9 cm long, 10-nerved, the apex acute, the peduncle

ca. 1 cm long, the pedicel 0.5-2.0 cm long; bracts and bracteoles ovate-lanceolate, ca. 1 mm long, persistent.

Hypanthium short-tubular, 1.5-2.5 cm long; calyx at first spathaceous, soon irregularly splitting to the

hypanthium into 2-5 lobes; petals white with wine-red veins, subequal, linear to linear-spatulate, 8-11 cm

long, 6-10 mm wide, glabrous, the margins slightly crisped, the claw ca. 1 cm long; fertile stamens 10, the

Wunderlin, Bauhinia sul Bauhini t. Pauletia ser. Ariaria 631

filaments wine-red, the outer whorl 8-11 cm long, subequaling the petals, the inner whorl slightly shorter,

irregularly short-connate at the base for 5-20 mm into a staminal sheath, the staminal sheath tomentose

on its inner surface, otherwise glabrous, the anthers linear-oblong, 1-1.5 cm long, glabrous; gynoecium

7-9 cm long, subequaling the androecium, the ovary 1.5-2 cm long, reddish brown-tomentose, the gy-

nophore 3-4 cm long, reddish brown-tomentose near the ovary, glabrate below, the style 4-6 cm long,

reddish brown-tomentose proximally, sparsely short-hirsute distally, the stigma obliquely capitate. Fruit

linear, apiculate with the persistent style base, 20-30 cm long, 2.5-3 cm wide, reddish brown-tomentulose

or -velutinous, the stipe ca. 4 cm long, glabrate; seeds oblong to suborbicular, ca. 15 mm long, ca. 12 mm

wide, the surface brown, dull, the funicular aril-lobe scars subequal, ca. 1 cm long, extending ca. 1/3 the

way around the seed.

Distribution and ecology.—Venezuela in Bolivar and Monagas and in Suriname. In forest at 100—800 m.

Flowering January to October; fruiting January to July.

This species is most closely related to B. multinervia, but is easily poles LA its erect tricl on

] 1

the lower leaf surface and the blade suborbicular (vs. appressed trichom y suborbicular

in B. multinervia).

If TT d J 11 1 1 oo =]

Specimens examined. VENEZUELA. Boli I

the La Lara and Corazon de Jesús mines, Aug 1966, Blanco 537 (MO, NY); ca. 12 km S of El Dorado (06%43'N, 61°37 W), 19 Feb 1983,

Hokche 25 (USF); near sansa, ca. 100 m, ca. 50 km NE of upata, oer M 66°7'48"W), 19 Jun 1983, Hokche 39 (USF); Reserva

Forestal Imataca, Carret San Martín de Ti Río Cuyunf, 21 May 1982, Stergios et al. 3863 (MO); along

road between km 11 and 18.5, S of El Dorado, 215 m, 23 Jul 1960, Steyeinidrk 86620 (US). Monagas: ca. 2-3 km NE of Santa Ines,

790-800 m, 6 Jun 1967, Pursell et al. 9188 (NY, US). SURINAME. Nature Park Brownsberg, near Irene Falls, 21 Jan 1974, Roberts s.n.

(L.B.N. No. 16305) (K, US); Nature Park Brownsberg, 10 Oct 1969, Tawjaeran s.n. (L.B.N. No. 12584) (K, US); Nature Park Brownsberg,

25 Jan 1974, Tawjoeran s.n. (L.B.N. No. 13721) (K, US); Nature Park Brownsberg, 2 Jul 1924, Wullschlaegel s n. (L.B.N. No. 6126) (K);

Nature Park Brownsberg, 24 Mar 1924, Wullschlaegel s.n. (L.B.N. No. 6421) (NY); Nature Park Brownsberg, 30 Jan 1925, Wullschlaegel

s.n. (L.B.N. NO. 6760) (NY)

3. Bauhinia esmeraldasensis Wunderlin, sp. nov. (Fig. 1). Tw: ECUADOR. EsmeraLoas: Bilsa Biological Station, Mon-

tañas de Mache, 35 km W of Quinindé, 5 km W of Santa Isabela (00°21'N, 79?44"W), 400—600 m, 3 Jan 1955, N. Pitman 1173

: USF!).

MO!;

(HOLOTYPE:

2T AS

A D lcu £1 lifl X17 ] Ts fx pes , fructibus, q f1

Tree to 6 m; branches minutely stigulose to glabrate, soon becoming glabrous. Leaves with the blade sub-

membranaceous to chartaceous, suborbicular, 8-13 cm ac 8-13 cm wide, bilobed 1/4—1/3 their length,

the lobes divaricate, the base cordate, the lobe ap to apiculate or slightly caudate, the upper surface

glabrous, the lower surface minutely tomentulose, at least on iip nerves, ee au in color than the

upper surface, 9—11-nerved, the petiole 4-6 cm long, minutely st rate; sti lanceolate, ca.

2 mm long, caducous; adpetiolar intrastipular excrescence ca. 2 mm lona: the others minute. Inflorescence

cauliflorous racemes to 4 cm long, 10—12-flowered, the flowers fasciculate, the rachis tomentulose; buds

linear-lanceolate, 3-4 cm ron: d menea, c apex acuminate, the peduncle ca. 5 mm long, the pedicel ca.

5-10 mm long; bracts and ovate, ca. ] mm ong, persistent Hypanthium short-tubular,

6-10 mm long, 10-nerved; calyx at first ee soon splitti thium into 2—5 lobes; petals

white, subequal, liearspatinulate, 5-7 cm long, 0.5-1 cm mide drous. the margin slightly crisped, the

claw ca. 5 mm long; fertile stamens 10, the filaments of the outer whorl 4-6 cm, slightly shorter than the

petals, the inner whorl ca. 1 cm shorter, irregularly connate at the base 10-15 mm into a staminal sheath,

the staminal sheath sparsely pilose on the inner surface, the anthers linear-oblong, glabrous, those of the

inner whorl 5-6 mm long, those of the outer whorl 3-4 mm long; gynoecium 6-7 cm long, subequaling the

androecium, the ovary ca. 1 cm long, tomentose, the gynophore ca. 2 cm long, glabrous, the style 3-4 cm

long, tomentulose proximally, the stigma obliquely capitate. Fruit 10-15 cm long, 2-2.5 cm wide, brown,

apiculate with the persistent style base, sparsely tomentulose to glabrate, 3—4-seeded, the stipe ca. 2 cm

long, glabrous; seeds suborbicular, 12-14 mm long, 9-11 mm wide, the surface dull brown, the funicular

aril-lobe scars 2-3 mm long.

t tani titute of Texas 3(2)

632 Jo

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Fic. 1. Bauhinia esmeraldasensis (isotype, USF).

Wunderlin, Bauhinia sui Bauhini t. Pauletia ser. Ariaria 633

P ECUADOR. E ldas: Bitsa Biol IK M le Mache, 35 km W Vola edet MO 79*44"W), 400—600

m, 28 Sep 1994, M.S. Bass, L. Kueppers, & N. Pitman 88 (MO, USF); Bilsa Biol

5 km W of Santa Isabel, (00°21'N, 79°44'W), 400—600 m, 14 Nov 1994, J.L. Clark, H. Lintz, & S. Mora 234 (MO, USF), Bilsa Biological

Station, Mache Mountains, 35 km W > oe : km W of Santa Isabel, (00°21'N, 79°44'W), 400—600 m, 7 Oct 1994, J.L. Clark &

H. Lintz 162 (MO, USF) Bilsa Biologi t , 35 km W of Quinindé, 5 km W of Santa Isabel, (00°21'N, 79°44'W),

400—600 m, 10 Apr 1995, J.L. Clark & Y. Troya 696 (MO, USF); Bilsa Biol l Station, Montañas de Mache, 35 km W of Quinindé, 5

km W of Santa Isabela, (00?21'N, 79°44'W), 400—600 m, 7 Dec 1994, N. Pisa & M. Bass 1039 (MO, USF).

5 km W of Quinindé,

Etymology.—The name is derived from the Ecuadorian Province Esmeraldas.

Distribution and ecology.—Endemic to Ecuador in Esmeraldas. In mature premontane wet forest at

400-600 m. Flowering January to October; fruiting September to December.

Bauhinia emeraldasensis is closely related to B. flagelliflora in being cauliflorous, but differs in its smaller

inflorescences (4 cm long in B. esmeraldasensis vs. 80 cm in B. flagelliflora), its smaller fruit (10-15 cm long

in B. esmeraldasensis vs. 20-22 cm in B. flagelliflora} and its thinner and smaller leaves (8-13 cm long in B.

esmeraldasensis vs. 20-26 cm in B. flagelliflora).

In the Cercideae, cauliflory occurs in the temperate northern hemispheric genus Cercis Linnaeus and

the African genus Adenolobus (Harvey) Torre & Hillcoat. It is unknown in the west African genus Griffonia

Baillon and the Madagascaran genus Brenieria Humberet and is rare in Bauhinia. In the new world species

of Bauhinia it is known to occur only in B. cookii Rose (subgenus Bauhinia section Amaria) and in two species

of subgenus Bauhinia section Pauletia series Ariaria (B. esmeraldasensis and B. flagelliflora).

4, Bauhinia flagelliflora Wunderlin, Brittonia 35:335. 1983. Tree: ECUADOR. P C

of Montafias de Ila on road from Patricia Pilar to 24 de Mayo at km 12, 600 m, 6 Feb 1979, C.H. Bodin 7345 (HOLOTYPE: SEL!;

isotypes: MO!, USF!)

Tree to 9 m; branches glabrous. Leaves with the blade reo amas to chartaceous, broadly ovate,

20-26 cm long, 17-20 cm wide, bilobed to ca. 1/4 their length, the lol ghtly divaricate, the base cordate,

the apex lobe attenuate, the upper surface glabrous, the lower surface amenas at least on the nerves,

slightly lighter in color than the upper surface, 9—11-nerved, the petiole 7-9 cm long, sparsely tomentulose

to glabrate; stipules triangular-lanceolate, ca. 2 mm long, caducous; adpetiolar intrastripular excrescence

ca. 1 mm long, the others minute. Sinaia cauliflorous racemes to 80 cm long, 20—40-flowered, the

flowers paired or solitary by abortion, t luncle adnate to the rachis (stem), emerging just above the next

higher node, the rachis tomentulose; buds linear, ca. 6 cm long, ecostate, the apex apiculate, the peduncle

10-20 cm long, the pedicel ca. 1 cm long; bracts and bracteoles triangular-ovate, ca. 1 mm long, persistent.

Hypanthium tubular, ca. 1.5 cm long, obscurely 10-nerved; calyx at first spathaceous, soon irregularly splitting

to the hypanthium into 2—5 lobes; petals white, iera mnohe ca. 5 cm long, ca. 6 mm wide, glabrous,

the claw ca. 5 mm long; fertile stamens 10, the filaments of the outer whorl 3.5—4 cm long, the inner whorl

slightly shorter, irregularly short-connate at the base for 1-5 mm into a staminal sheath, the staminal sheath

pilose-tomentose on the inner surface, the anthers linear-oblong, 6-8 mm long, glabrous; gynoecium ca. 6

cm long, subequaling the androecium, the ovary ca. 12 mm long, tomentose, the gynophore 2.5—3 cm long,

glabrous, the style ca. 2 cm long, tomentulose proximally, eres glabrate, the stigma obliquely capitate.

Fruit Gi ture) linear-oblanceolate, apiculate with tl t style base, 20-22 cm long, 2—2.2 cm wide,

bie, sparsely tomentulose to glabrate, the stipe ca. 3 cm long, glabrous; seeds not seen.

Distribution and ecology.—Ecuador in Pichincha. Known only from the type collection. In subpluvial

forest at 600 m. Flowering in ey

With its large leaves, B t first resembles B. pichichensis and B. tarapotensis, but is prob-

ably more related to B. ld i þei lifl , a feature it shares with that species (see above).

It differs from B. esmeraldasensis by its La rescates (80 cm long in B. flagellliflora vs. 4 cm long in B.

esmeraldasensis) and its larger leaves (20-26 cm long in B. flagelliflora vs. 8-13 cm long in B. esmeraldasensis)

which are also chartaceous (vs. submembranaceous in B. esmeraldasensis).

5. Bauhinia multinervia (Kunth) DC., Prodr. 2:515. 1825. Pauletia multinervia Kunth, Nov. Gen. Sp. Pl. 6:316. 1825.

634 tani i f Texas 3(

Type: VENEZUELA. Mrranpa: Curiepe and Montaña de eee 16-25 Nov 1799, H.EWH.A. von Humboldt & A.J.A. Bonpland 576

( E: a Es 1 TTA A 56/C7!; RIVAIS ( À F! , Gl, MO!, NY! , USD).

Pauletia glaucescens Kunth, Nov. Gen. Sp. Pl. 6:317. Bauhini (Kunth) DC., Prodr. 2:515.1825. Tree: iiec aper MONAGAS:

near Caripe, 4-24 Sep 1799, H.LEWH.A. von Humboldt 6 A] A. Boo 221 (LECTOTYPE, | g IDC 157/A1!;

ISOTYPE: P microfiche IDC 157/A21).

Bauhinia megalandra Griseb., Fl. Brit. WI. 213. 1860. Type: ST. VINCENT: L. Guilding s.n. (Lectotype, here designated: K!; photo ex K:

USD.

Large shrub or tree to 10 m; branches ferruginous-tomentose when young, soon becoming glabrate. Leaves

with the blade chartaceous, ovate, rarely suborbicular or elliptic-oblong, 7-18 cm long, 6-14 cm wide,

bilobed 1/4 to 1/2 their length, the lobes slightly divaricate, the base cordate to rounded, the lobe apex

rounded or obtuse, the upper surface glabrous, the lower surface minutely strigose, occasionally glaucous,

ferruginous-tomentulose on the nerves and sometimes on the blade, lighter in color than the upper sur-

face, (5—)7-9(-11)-nerved, the petiole 2-4 cm long, ferruginous-tomentose to glabrate; stipules ovate, ca.

1 mm long, persistent; adpetiolar intrastipular excrescence to 2 mm long, the others minute. Inflorescence

terminal or subterminal and axillary, racemose or paniculate, 3-20-flowered, sometimes paired, the rachis

ferruginous-tomentose, becoming glabrate in age, buds linear, 8-11 cm long, the apex rounded, the pedicel

13 cm long; bracts and bracteoles ovate, ca. 1 mm long. Hypanthium short-tubular, 2-3 cm long; calyx

at first spathaceous, soon irregularly splitting to the hypanthium into 2-5 lobes; petals white, subequal,

linear, 6-8 cm long, 3-6 mm wide, glabrous, the claw 6-8 mm long; fertile stamens 10, the filaments of

the outer whorl 6-8 cm long, equaling or exceeding the petals, the inner whorl slightly shorter, irregularly

short-connate at the base for 5-15 mm into a staminal sheath, the staminal sheath with a laciniate collar on

the adaxial side, reddish tomentose at the base, the filaments sometimes tomentulose proximally to nearly

their length, the anthers 1.5-2 cm long, glabrous; gynoecium 8-10 cm long, subequaling the androecium,

the ovary 2-3 cm long, reddish brown-tomentose, the gynophore 4-5 cm long, reddish brown-tomentose,

the style ca. 3 cm long, tomentose or tomentulose below, glabrate on the upper 1/2, greenish white, the

stigma oblique-capitate. Fruit linear, apiculate with the persistent style, (15~—)20-30 cm long, 2-3 cm wide,

dark brown, sparsely tomentulose to glabrate, the stipe 4-68) cm long, reddish brown-tomentose; seeds

suborbicular or oblong, 15-18 mm long, 10-15 mm wide, the surface dull, dark brown, the funicular aril-

lobe scars 12-15 mm long, extending 1/2 way around the edge of the seed.

Distribution and Ecology.—Lesser Antilles in Antiqua, Guadeloupe, Martinque, and St. Vincent, and in

Venezuela, Trinidad, and Suriname. Along stream der in eena forests and moist evergreen forests

at 0—1,200 m. Flowering and fruiting all year. Someti ated as a novelty, but apparently not natural-

izing.

Bauhinia multinervia is most closely related to B. eilertsii, but is easily distinquished by the appressed

trichomes of the lower leaf surface (vs. erect trichomes in B. eilertsii).

Representative specimens examined. ANTIQUA. Northwest Sie district), Fig Tree Hill, m Nov 1937, Box 1259 (BM, UC, US);

without precise locality, 14—16 Feb 1913, Rose et al. 3401 (US). GUADELOUPE. li

839, Beaupertuis s.n. (P);

without geu locality, 1787, Isert s.n. (C); without precise locality, 4 Mar 1938, Questel 690 (US); witl ty, s.d., Richard

s.n. (F). MARTINIQUE. without presse Deeds 1857, kon 496 (P); without precise locality, 1882, jus: 1124 (NY); St. Luce, 5 m,

2 Jan 1939, Stehle & Stehle 3550 (US); Mois s.d., Terrasson 44 (P). ST. VINCENT. without precise locality, Mar 1890,

Smith & Smith 273 (K, NY). VENEZUELA. Bolí ] Km 11 and 18.5, S of El Dorado, d 1960, cid 86620

(US). Cojedes: near Tinaquillo, 24 Dec 1925, Pittier 11989 (US). Distrito Fed

11 Feb 1973, Croat 21557 (MO); road to electricity plant aaa eee near sea level, 1 12 Mar 1974, — & Morillo 10331 (MO);

along the Río Los Caracas, 0.8 m, 4 Nov 1982, Hokche 24 (USF); along road from Country Club to Río Chacaito y 1942, Lazzer 646

(US); 1-2 km S of Los Caracas, 40 km E of La Guaira, 18 Aug 1979, Nee 17552 (WIS). Miranda: along the Rio RA 400—60

30 Nov 1956, B li 5755 (G, MO); anes Nacional MS near ie om at 5 entrance, 13 Feb 1973, Croat 21729 m.

3 km SW of Araguita along road between Caucagua an racia de Orituco, 130 m, 17 Nov 1973, Davidse 4127 (MO); ca. 1-2 km

upstream from the mouth of the Río Chiquit its i ion with the Río C , S of » SOS ca. we m, 3 Jun 1977, Davidse &

González 13600 (MO, USF, VEN); along bea Chaguarama, 0-4 km SW of Palo Q Cúpira, ca. 50-100 m, 5 Mar

een Santa "lerpca ann Alta

1980, dd & Coens a en USF, TEN daa Pind Soins Siquire Valley, 500- 1,000 m, 19-24 Mar 1913, Pittier 5982

LS L I [-]

Wunderlin, Bauhinia sul Bauhini t. Pauletia ser. Ariaria 635

from Los ES 12kmf Rancl iQ ia, 520 m, 23 Nov 1961, Steyermark 89949 (NY, US); Cerros del Bachiller, near E end,

da C 1, S of Santa Criz 10 10 km W of Capira 20-65 m, 16-17 Mar 1978, Steyermark & Davidse 116243 (MO, USF,

VEN) Monagas: Montaña de Aquacate, al dade Pajarral, tributary to Río Caripe, NE of Alto de Aquacate, between Caripe

and Caripitc, 600-900 m, 19 Apr 1945, Serna 62177 (F). Sucre: vicinity of casei eon (Macuro), 5 Jan—22 Feb 1923, Broadway

633 (NY, US). Yaracuy: Montana de Maria Lionza, vicinity of Quebrada Qui fo Yaracuy, S of Chivacoa, 250-260 m, 12

Mar 1981 pan et al. 124894 (USF, v TRINIDAD. without precise bu pa 1857, Crueger s.n. (K); without precise locality,

1877-1780, Fendler 328 (BM, K, P). SURINAME. National Reserve Brownsberg, 10 Oct 1969, Tawjoeran s.n. (US).

6. Bauhinia pichinchensis Wanderlin, PESOS m 338. 1983. Tee: ECUADOR. Picuincua: Cooperativa Santa Marta

Rio Verde 2 | dos, 530 m, 5 Feb 1979, C.H. Dodson, A.H. Gentry, and J.A. Duke 7594

(HoLotyrE: SEL!; isotypes: MO!, USF!).

Tree to 10 m; branches reddish brown-tomentulose when young, glabrate in age. Leaves with the blade

chartaceous, oblong-suborbicular, 14-24 cm long, 13-21 cm wide, emarginate or bilobed to 1/4 their

length, the lobes slightly divaricate, the base cordate, the lobe apex rounded, the upper surface glabrous,

the lower surface reddish brown short-tomentose or -tomentulose, at least on the nerves, conspicuously

ad HENO enten in color than the upper surface, H the Lii 4—5 cm long, reddish brown-

a. 2 mm long, decid t 2 mm long,

the others Wn Infl I land axillary hare -Tacemose, 2- 4-flowered, the flowers paired,

the rachis reddish brown short-tomentose; buds lincar nico, ca. 5 cm long, the apex acute, obscurely

5-nerved, the peduncle ca. 1 cm long, the pedicel 2-2.5 cm long; bracts and bracteoles triangular-ovate,

1-1.5 mm long, persistent. id ode) nd dM: ca. 1 cm long, 5- or obscurely 10-nerved; calyx at

first spathaceous, soon irregularly splittin thium into 2—5 lobes; petals white, subequal, linear,

4-5 cm long, 3-6 mm wide, the las 1-1. 5 cm ona fertile stamens 10, the filaments of the outer whorl 3-4

cm long, the inner whorl slightly shorter, irregularly connate at base for 1-2 mm into a staminal sheath, the

staminal sheath glabrous, the anthers ca. 1 cm long, glabrous; gynoecium ca. 5 cm long, subequaling the

androecium, the ovary ca. 1.5 cm long, tomentose, the gynophore ca. 1.5 cm long, glabrate, the style ca. 1

cm long, tomentulose, the stigma obliquely capitate. Fruit linear oblanceolate, apiculate with the persistent

style base, 20-25 cm long, 33.5 cm wide, light brown, glabrate, the stipe ca. 3 cm long, glabrate; seeds

16-18 mm long, 14-16 mm wide, the surface dull brown, the funicular aril lobe scars extending about 1/2

way around the edge of the seed.

Distribution and Ecology.—Known only from Ecuador in Carchi, Esmeraldas, and Pichincha. In forest

at 250—1,200 m. Flowering and fruiting all year.

First collected by Raymond Benoist in 1930, the plant was rediscovered by Calaway H. Dodson in 1979

and recognized as a new species by Wunderlin (1983). It most closely resembles B. conceptionis of Colombia

in having large suborbicular, emarginate to slightly bilobed leaves, but differs from that species by having

larger leaves (14-24 cm long in B. pichinchensis vs. 8-13 cm long in B. conceptionis) and the ovary tomentulose

(vs. densely tomentose in B. conceptionis). Bauhinia conceptionis is related more closely to B. tarapotensis than

to B. pichinchinensis.

Specimens examined. ECUADOR. Carchi: Reserva Indígena Awá, San Marcos, 25 km NW of El Chical (01°02'N, 78°14'W), 1,800 m,

15-28 Jun 1991, Rubio et al. 943 (MO, USF); Reserva Indígena Awá, Gualpi Alta (01°02'N, 78°14'W), 1,800 m, 15-28 Jun 1991, Rubio et

al. 1730 (MO, USF). Esmeraldas: dd cad Awá, Centro de La Unión, Cañón del Rio Mira (00°52'N, 78?26'W), 250 m, 22 Mar

1993, Aulestia & Aulestia 1467 (MO, I , 10 km N of Lita, 11 May 1987, Acevedo & Daly 1684 (NY, USF);

Reserva Ethnica Awa, Centro Pambilar 1°08 N, 78°36'W), 500 m, 21 Jan 1993, Aulestia & Aulestia 1052 (MO, USF); Reserva Indigena

Awa, Cañón del Río Mira, 10 km E of Alto Tambo (01°02'N, 78°26'W), 250 m, 16-26 Mar 1991, Rubio et al. 1141 (MO, USF). Pichincha:

Santo Pas, Charco Vicente, Rio San ba Sag N, 78753 PME 200 m, eg Sep 1993, Tirado et al. t Ma WEM viis ene de

Las Colorados, 8 Sep 1930, B t 30 t km 3 W of!

530 m, 22 Jul 1979, Dodson et al. 8541 (MO, SEL. USE): Centinela, 12 km E of Patricia Pilar at Km 45 between Santo Domingo de les

Colorados, 600 m, 2 Feb 1985, Dodson & Neil 15525 (MO, USF).

7. Bauhinia t tensis Benth. in Mart., Fl. Bras. 15(2):198. 1870. Tree: PERU. San Martin: near Tarapoto, 1855—1856,

R. Spruce 4417 (HOLOTE: K!; ISOTYPES: B- destroyed. BMI, K!, Fl, G!, Pl; photo ex K (holotype): F!; photo ex B: F!, MO!; pl

Fl, MO

, J fall n.a a in LI L'AL ET * F da h

636

Ariaria a E ee cn i Prehist. & Viajes 219. 1893. Tyre: PERU. I th of Rio Santiago, 200 m, 25 Nov 1931,

Y M!, G!, K!, MO! ae unos here e No sepenmens were cited in the protolog.

har

name was overloglead until Fo

Bauhinia baina J.E Macbr., Field Mus. = Hist, Bot Ml TE 211. 1943. T PERU. Lots Rio Mazán, 10 May 1929, LI. Williams

186 (noLorveE: Fl; isotyee: USD. M id li l The handwritten label on

the aie says ARio Masana@ ee Rio A and a date 10 May 1929. The printed label with the specimens states

Aalong Río Itaya@ AMay, 19290,

Bauhinia amplifolia Ducke, Bol. Tecn. Inst. Agron. N. 2:16. 1944. Tyre: BRAZIL. Amazonas: near Tabatinga, margin of the Paraná de Ara-

massa, 4 Mar 1944, A. Ducke 1594 (HoLotype: IAN; isotypes: F!, MO!, US!).

Large shrub or tree to 10(19) m; branches strigulose to tomentulose, soon becoming glabrate. Leaves with

the blade chartaceous, ovate, 10—20(—25) cm long, 7-15 cm wide, bilobed 1/4—1/2 their length, the lobes

slightly divaricate, the base rounded to cordate, the lobe apex acute to obtuse, the upper surface glabrous,

the lower surface minutely strigose, often reddish brown-tomentulose toward the base and on the nerves,

slightly lighter in color than the upper surface, 9—11-nerved, the petiole 2.5-4(-6) cm long, tomentulose

to glabrate; stipules ovate to lanceolate, ca. 2 mm long, caducous; adpetiolar intrastipular excrescence to 3

mm long, the as minute. Inflorescence terminal or subterminal and axillary, 2-20-flowered, the flowers

paired, the rachis strigose to tomen tose; buds linear, 11—13(-18) cm long, the apex acute, obscurely 5-nerved,

the peduncle ca. 5 cm in lone the pedicel 5-10 mm long; bracts and bracteoles lanceolate, 2-3 mm long,

persistent. Hypanthium long-tubular, 4—6 cm long, 10-nerved; calyx at first spathaceous, soon irregularly

splitting to the hypanthium lip into 2-5 lobes; petals white, subequal, glabrous or glabrate, linear-filiform

to -spathulate, 9-15 cm long, 3-10(225) mm wide, the margin crisped, the claw 1.5-2 cm long; fertile sta-

mens 10, the filaments of the outer whorl 7-13 cm long, the inner whorl slightly shorter, occasionally with 2

reduced, irregularly connate at the base for 7-18 mm into a staminal sheath, the staminal sheath glabrous or

tomentulose on the inner surface, the anthers linear, 10-17 mm long; gynoecium 8-13 cm long, subequal-

ing the androecium, the ovary 23 cm long, with glandular trichomes or rarely tomentulose to tomentose,

the gynophore 6-10 cm long, glabrous, the style 4-6 cm long, glabrous, the stigma obliquely capitate. Fruit

linear, apiculate with the persistent style base, brown, 15-25 cm long, 2.5-3.5 cm wide, the stipe 4—5 cm

long, glabrous; seeds elliptic-obovate to suborbicular, 15-20 mm long, 11-18 mm wide, the surface dull,

brown, the funicular aril-lobe scars ca.1.5 cm long, extending 1/3-1/2 the way around seed.

Distribution and ecology.—Colombia in Caqueta, Meta, and Putumayo; Ecuador in Napo, Morona-Santiago,

and Pastaza; Peru in Amazonas, Huánuco, Loreto, San Martín, and Ucayali; Brazil in Acre and Amazonas. In

high primary or secondary rainforest, usually along rivers at 90—1,000 m. Flowering and fruiting all year.

Bauhinia tarapotensis is closely related to B. conceptionis, a species restricted to Colombia, but differs in

having ovate leaves (vs. suborbicular in B. conceptionis) with acute to obtuse lobes apices (vs. rounded in B.

conceptionis) and larger flowers (14—21 cm long in B. tarapotensis vs. 6—7 cm long in B. conceptionis).

Representative specimens examined. COLOMBIA. Caquetá: 10 km below Puerto Rico, 350—400 m, 27 Sep 1975, Cabrera et al. 3522

(USF); Florencia, Río Orteguaza, 400 m, 21 Mar 1965, Garcia-Barriga 18198 (US); 10 km S of San José de Fragua, 320 m, 11 Jan 1974,

Gentry et al. 9145 (MO). Meta: Puerto Losada, Mina Blanca-Puerto Losada road (16 km), 360—450 m, 23 Feb 1988, Callejas & Maru-

landa 5889 (USF); margin of the Río Guayabero, Macarena rapids, 380 m, 23 Jan 1959, Pinto et al. 304 (P); N side of river near Lis

of Guejar and Río Zanza, N end of Cordillera Macarena, ca. 500 m, 20 Aug 1950, Smith & Idrobo 1482 (COL, MO, US). Putumayo: E

slope of Cordillera Oriental, tributary of n Río Mocoa, near Mocoa, 700 m, ojan. 1945, Ewan 16723 (BM, D a 325 m, Oct-Nov

1930, Klug 1830 (BM, F, K, MO, NY, US). ECUADOR. Napo: from Coca to 1 | N 0°26-27'S,

76?41—57'W), 200 m, 18 Mar 1980, diis et al. 30232 (AAU, USF); Taisha, Río Guambime, (02°23'S, 77°30'W). 400 m, n 1980,

Brandbyge et al. 322601 (AAU, USF); Río Wai si ayá, a northern tributary to Río Aguarico, ca. 6 km upriver from San Pablo, (00?51'S,

76°21'W), 300 m, 10 Aug 1980, Brandbyge & Asanza 32764 (AAU, USF); Río Wai si ayá, 1 km upstream from the outlet in Río Aguarico,

(00°15'S, 76°21'W), 300 m, 6 Aug 1981, Brandbyge et al. 33260 (AAU, USF); Reserva Biológica Jutan Sacha, Río Napo, 8 km E of Misa-

hualli, (01°04'S, 77°36'W), 450 m, 24 Apr-5 May 1987, Cerón 1329 (MO, pein Reserva ones Jutan a 8 km E of Misahualli,

bank of Río Napo, (01°04'S, 77°36'W), 450 m, 4 Sep 1987, Cerón et al. 2162 (MO, -al d Misahuallí

(junction of Río Misahuallí and Río Napo) at Vereda Venesia, 3.8 km W of Misahuallí, (ca. 01°02! S, 77°42'W’), 370 m, 2 May 1984, Croat

58889 (MO, USF); between Cotapino (Concepción) and Río Bueno, ca. 400 m, 21 Feb 1968, Harling et al. 7154 (GB, USF); Latas, Rio

Napo, 12 Aug l pos im de 260 s idi n ca. 6 km from Puerto Napo, 14 Aug 1968, Lugo 298 (GB, USF); southern side of Rio

Napo, 3-41 ), 8 Jan 1973, Lugo 2546 (GB, USF); Tierra Colorada, ca. 3 km N of Coca (Puerto

Wundertin, Bauhinia sul Bauhini t. Pauletia ser. Ariaria 637

Francisco de Orellana), 22 Jan 1973, Lugo 2916 (GB, USF); San Pablo at Río Napo, 6-7 km SW of Coca (Puerto Francisco de Orellana),

28 Jan 1973, Lugo 3012 (GB, USF); road Pu imd cd aud Bl Cae 23 sen 1973, » Lugo 3487 (GB, USF); near Tena, 400 m, 2-11

Apr 1935, Mexia 7142 (US); ; Afiangu, l Yasuní, ca. 250 m, 30 Jun—9 Jul 1982,

SEF 10261 (AAU). Marona-Santiago: Centro Shur Yukutais, ee S of GEHEN (03°30! S. 78°10'W). 22 Apr 1989, Bennett & Andrade

3795 (NY, USE) Mendez, | tl P Upano, 530-760 m, 12 Nov 1944, Camp E-949 (K, US); Taisha, banks of

Rio Guay, 460 m, 6 Feb 1962, Cazalet es 7681 (K, US); vicinity i Mendez, ca. 620 m, 14 Oct 1989, Luther et al. 27124

(SEL, USF). Pastaza: Lorocachi, 1-5 km up Río Curaray from military camp, (01°38'S, 75°58'W), 200 m, 30 May 1980, Branbyge &

Asanza 31407 (AAU, USF); Curaray, Valle de la Muerte, (01?25'S, 75°52'W), 240 m, 22 Mar 1980, Holm-Nielsen et al. 22531 (AAU, USF);

Montalvo, along Río Bobonaza 0-1 km N of the military camp, (02°05'S, 76°58'W), 250 m, 16 May 1979, Løjtnant & Molau 13300 (AAU,

USF); Montalvo, within the military camp, (02°05'S, 76°58'W), 250 m, 17-22 May Md Parar & Molau 17470 setae, Rio ae

in the vicinity of Canelos, 15 Mar 1971, Lugo 1689 (GB, USF); Río Bobon , between D Cabo

Pozo and La Boca, (02°30'S, 76°38'W). ca. 275 m, 21 Jul 1980, a et 2d 34917 (AAU, USF); Río Pastaza, river banks between the

outlets of Río Bobonaza and Río Ishpingo, (ca. 02°34'S, 76°43'W), ca. 275 m, 22 July 1980, @llgaard et al. 34983 (AAU, USF). PERU.

Amazonas: near Kusu, Río Numpatkin, 340—400 m, 10 Mar 1973, Ancuash 79 (MO); Río Cenepa, vicinity of H pmi, ca. 3 km from

the mouth of the Río Huampami, 200-250 m, 25 Jul 1978, Ancuash 1115 (MO, USF); Quebrada "s Po Río pce vicinity of

Huampami, ca. 5 km E of Chávez Valdivia, (04°30'S, 78°30'W), 200—250 m, 11 Aug 1978, Ancuash 1 enepa, vicin-

ity of Huampami, ca. 5 km E of Chávez Valdivia, (04°30'S, 78°30'W), 200—250 m, 14 Aug 1978, "— 473 (MO, USF); Río Cenepa,

vicinity of Huampami, Quebrada Kachaig, ca. 5 km E of Chávez Valdivia, (04°30'S, 78°30'W), 200-250 m, 15 Aug 1978, Ancuash 1506

(MO, USF); island 1 km below La Poza, Río pee 180 m, 8 Aug 1979, iue 13 en nu Huánuco: Tet N o Puerto Inca, (09°18'S,

74°58'W), 250-300 m, 15 Sep 1982, Foster 8811 (MO, USF). Loreto: along Witoyacu, (04°15'S,

76°35'W). 31 Jul 1979, Diaz et al. 1313 (MO h of the Rio G a. 200r m, 18 May 1978, Gentry et al. 22080 (MO, USF);

Florida, Río Putumayo, at mouth of Río Zubirieta: 180 m, May—Jul 1931, Klug 2169 (BM, F, K, MO); Balsapuerto, ca. 220 m, Mar 1933,

Klug 2974 (BM, F, MO); Maucallacta, Río diia gie ca. 200 m, a 1935, Klug 3941 uaa F, K, s as near mouth a Río Santiago,

200 m, 25 Nov 1931, Mexia 6158 (BM, F, G, K, MO, WIS). San M E of Tarapoto 8 km C Creek, 1,000

m, 15 Aug 1937, Belshaw 3234 (K); Juan Jui, upper Rio ne ca. 400 m, Oct 1934, Klug 3851 (BM, F, MO, WIS); Juan Jui, Alto Rio

Huallaga, 400-800 m, Jan 1936, Klug indi (BM, K, MG) Te km E of at Snape. on road to Chazuta, (06°39'S, 76°30'W, 250 m, 25 May

1986, ned et al 7271 (MO, USF); Quel Mayo, 9.6 km W of Flores-Mamonaquihua (turnoff is

m 24 from Tarapoto), (06/3655, 76?10—11"W), ca. 400 m, 15 May 1986, Knapp et al. 7406 (MO, USF); Chara del Sr. Manuel Arévalo

es along the Río Huallaga, 400 m, 13 Jul 1970, Schunke Vigo 4106 (F, G); Río de la Plata, 600—700 m, 27 Mar 1975, Schunke Vigo 8169

(MO, US, USF); mouth of the Río Tocahe, 400 m, 29 Apr 1975, Schunhe Vigo 8329 (MO, USF). Ucayali: Quebrada Shesha, tributary of

Río Abujao, 1-2 days upriver by “peki-peki”, ca. 60-70 km NE of Pucallpa, (08°02'S, 73°55'W), ca. 250 m, 18 Jun 1987, Gentry & Diaz

58412 us pud Bose Nen Eumbolee. along Quebrada Talos ca. 200 m, 20 Jun 1981, Young 937 (MO, USF). BRAZIL. Acre:

Aug 1933, K 293 (BM, F, G, K, MO). A Rio Solimóes, 2 km below

Tabatinga, 24 Jul 1973, — et al. 16771 OK, MO, USF).

ACKNOWLEDGMENTS

I gratefully acunoWIenee the curators of AAU, BM, i COL. FG, GB, IL, E, LE. MO, P, SEE, NY, UG. US.

VEN, and WIS for mal ilable for study. Ithank Bruce Hansen (USF) for his helpful sugges-

tions and Alan Franck (USF) for assistance with breue I also acknowledge Rafael Torres Colín (MEXU)

and Howard L. Clark (DUKE) for review of the manuscript and helpful suggestions.

REFERENCES

Forero G. E. 1966. La identidad de Ariaria superba Cuervo Márquez (Caesalpiniaceae-Bauhinieae). Mutisia

3011-7].

Vaz, A.M.S. DAF. AND A.M.G. DE AzevEpo Tozzi. 2003. Aculeatae,

Caesalpinoideae, Cercideae). Novon 13:141-144.

WUNDERLIN, R.P. 1983. Three new species of Bauhinia (Fabaceae) from Ecuador. Brittonia 35:335-340.

WUNDERLIN, R.P. 2006. Revision of Bauhinia subgenus Bauhinia section Amaria (Cercideae: Caesalpinioideae:

Fabaceae). Sida 22: 97-122.

WUNDERLIN, R.P, K. LARSEN, AND S.S. LARSEN, 1987. Reorganization of the Cercideae (Fabaceae: Caesalpinioideae). Biol.

Skr. 28:1—40.

tion Pauletia (Leguminosae,

laf the Batanical D h Institute of Texas 3(2)

638

BOOK REVIEWS

LAWRENCE E. ESTAVILLE AND RICHARD A. Ear. 2008. Texas Water Atlas. (ISBN 978-1-60344-020-2, flexbound

with flaps). Texas A&M University Press, John H. Lindsey Building, 4354 TAMU, College Station, Texas

77843-4354, U.S.A. (Orders: www.tamu.edu/upress, 1-800-826-8911). $24.95, 152 pp., 94 color maps,

17 color photos, 3 tables, 9 graphs, 1 appendix, bibl., index, 8 5/8" x 11 1/4".

Lawrence E. Estaville and Richard A. Earl h iled and compared eee environmental, water de weather data, population

and human acu to Ep usd eod rocky y relationship itl ge-p k, Texas Water Sm

Et. d. dur A 1 114.4 E " 1 1 J 1

i E oO i r [e]

or unexplored. Each cl ] with an easy to understand histori iew based ig holarship by the authors,

1 D- c " T FILES US ba! C qp cce 4 T^ . | A

both SO of geography. The book, sp y y y, 1 g

1 T I ES: | h i : - g 3. a1 ese Bh | "mi gi t Texas water

Atlas i luabl ional ty of fields, educators and

supply, "s and quality. Texas Wat

the lay person.— Becky Brandenburg, Volunteer, Botanical Research ieat of Texas, Fort Worth, Texas 76102- 4025, U.S.A.

Jonn Tyrer Bonner. 2009. The Social Amoebae: The Biology of Cellular Slime Molds. (ISBN 978-0-691-

13939-5, hbk: alk. paper). Princeton University Press: 41 William Street, Princeton, New Jersey 08540-

5237, U.S.A. (Orders: http://press.princeton.edu, 1-800-777-4726, 1-800-999-1958 fax, 1-609-258-6305

overseas fax OR orders@cpfsinc.com, California-Princeton Fulfillment Services, 1445 Lower Ferry Road,

Ewing, New Jersey 08618, U.S.A.). $19.95, 156 pp., 34 b&w line illustrations, 5 1/2" x 8 1/2".

Tun es Pu has spent more c 60 years of study devoted to the biology of the cellular slime molds. His 18 previous books

a, beginning in 1959 we = First Edition of Tr aia ue Molds, which was not do

bet ic with d iptive list ] l. Bonner

Pu I ^ E oy

states his goal in this current book, “This book is not an encyclopedia, a eriba; or a monograph, it covers Sanly a fraction of all we

know about slime molds. Rather, it i he big l l 1, with a n unconventional ane new O ” d in

addition, the narrative is " enum at e curious layperson .. lin this B I g I g

4 ] = + J " A 1 J Ls :11 LEN ] A e A

L i P [e] Pu ri

4l + 4 L 1 1 1. 1 fn 2 + zz]

oo "o PLUIE

igh ical headi 11 Introducti "hel ife Cycle, Evolution, Ecology, Behavior of A | d Cell M

Nisrohocenesis Differentiation, an The Future. These sections are rather short, varying from 5 to 16 LK in length. The Evoliition

section makes for fascinating reading, RÁ the recent a dou an ee ae rDN ates i du into

pe +

four groups as shown on a phylogram. TI gh sp

chimaeras, heterokaryons, and | ial ti Pues is one ears vignette entitled “Slime Mold Aggression"

yons, y E jn

that ibes the di y of a cellular sli ld in a bat that al killer compound that destroys and engulfs

+ 1 s A + 11 [- PL J al] 4 Japi E .

E ) PF y Ter E In the Ecology topic there is a section on

“Latitudinal Diversity” where species diversity in the warmer tropics is hig! d is 1 t higl Pd ae in the tropics or

1 x A «1 1 Joel: FEN ot £ = I: CST Fr_11 +1 J]J: Il x

going I I y gI role of cyclic AMP in

m +l : » ros F +1 1 iw] 1 k 1 r1 x JR tha *

g g ggreg p I eed Ao

3 = el L T5. xl: Ls] 1 + tlh 1 4 4 £ : E A The

ee = is Pond selective e = 10 0 pages ane ae references are sprinkled ine. ane and newer references 2 I ln to

university, this ld 1 piq y this f. ing group of organi Harold W Keller, ey ere

—€— Research Institute of Texas, Fort Worth, Texas 76102-4025, U.S.A.

J. Bot. Res, Inst. Texas 3(2): 638. 2009

ERIOGONUM VILLOSISSIMUM (POLYGONACEAE: ERIOGONOIDEAB),

A NEW SPECIES ENDEMIC TO ACKER ROCK, OREGON

James L. Reveal Dana York

Cornell University California eto - LUE

L.H. Bailey Hortorium

idee of Plant Biology Eureka, pda = 0 : USA.

Mann Library dana_York@dot.ca.gov

hae aca, New York 14853, U.S.A.

jir326@cornelledu

Richard Helliwell

Umpqua National Forest

2900 NW Stewart Parkway

Roseburg, Oregon 97471, U.S.A.

rhelliwell@fs.fed.us

ABSTRACT

Te: CS F | z s Dan] ^ s i J A ^T D 1 1 1 A e E : E 1 Xl7

[ = ] x A Q +l E 13 Y

£^ ua f T^ ] r7 +l 4 my Li A -1 4 A :11 + *:x Tai : 1 £

—

Oligogonum and

is most closely allied to E. alpinum, a narrow endemic confined to north-central California.

RESUMEN

Se describe e ilustra Eriogonum villosissimum (Polygonaceae), una nueva especie restringida a Acker Rock, una roca volcánica casi

4 1 1 r^ -A1;11 -l 1 rn” A Ls | Paes A (ens | P we y | Ti 1 1 z E na 2 7 = L 1

A 1 f; A ] + P OE bt,

del sul Oli tá tad E. alpinum

SAM EE TEL O VIHEUSUTEATTE y i E ’

INTRODUCTION

In 2005, while rock climbing in the Western Cascades of southwestern Oregon, York discovered a previ-

ously unknown Friogonum Michx. (Polygonaceae Juss.: Eriogonoideae Arn.) growing in crevices and small

shelves of a volcanic rock formation known as Acker Rock. The plants form clumps that are scattered over

the south-facing vertical faces of the formation. Nearby outcrops were searched in 2006 with no success of

finding another population.

TAXONOMY

Eriogonum villosissimum Reveal & D.A. m sp. nov. (Figs. 1-2). T Orcos. Douglas Co

Rock, Umpqua National Forest, mainly S of td kout, 1275 m elev., 528800 E, 4766674 N, Zone 10, 43?03'08"N, 122°38'47"W,

T295, RIE, sec. 13NWNE, 25 Jul 2009, York et al. 2989 NS NY; isotypes: BH, CAS, HSC, OSC, US).

AL Eo JSfíf..r AZ A d 1:1 A xc :11 - f ea, pian tpl : -1 fn

g p ( yet C ), sti[ 1-0.3 mm, nec 0.5-0.8 mm),

RN +3 : Lar 1: Jeg ey

Plants (Fig. 1) low, spreading, compact herbaceous perennials (0.5—)0.8—2.5(—3) dm across, composed of

numerous loosely arranged rosettes of tufted leaves at tips of slender, woody caudex branches arising from

a stout taproot. Leaves in well defined rosettes; petiole 0.5-1.5 cm long, densely white tomentose and ap-

pressed to slightly alla! M. DE from a broadly elongate triangular petiole base 4-8 mm long,

23.5 mm wide, tomentose a ially, sparsely villous to nearly glabrous or glal 1 tannish

brown adaxially; blade mostly ovate, 0.5-1.5 cm long, 0.5-1 cm wide, densely white tomentose and ap-

pressed to slightly spreading villous abaxially, greenish white and slightly less densely t tose adaxially,

broadly acute to obtuse but tapering abruptly to the petiole, entire, flat or somewhat or slightly enrolled

J. Bot. Res. inst. Texas 3(2): 639 — 643, 2009

640 Journal of t tani itute of Texas 3(2)

marginally, broadly acute to obtuse or occasionally nearly rounded apically; midvein slightly raised and

obscured by tomentum. Flowering stems (Fig. 2, left) scapose, spreading or somewhat erect, (5—)7-9(-1])

cm long, tomentose and spreading villous, becoming thinly so with age, with a whorl of 3 foliaceous bracts

about 1/3 up flowering stem; ian petiolate (0.5-2 mm long) un ovate blades 5-10 mm long, 4-7 mm

wide, densely whitet ially, less so and more vill r: lower stem portion 1-3 cm long;

peduncle 3.5-7 cm long. Involucres colar, campanulate, 4.5—7 mm lus 5—8 mm wide, tomentose and

densely villous abaxially, glabrous adaxially; teeth 5-8, erect, broadly acute, 0.5—0.9 mm long. Pedicels

erect to slightly curved with age, 4-6 mm long, glabrous; bractlets linear, 4-6 mm long, villous. Flowers

(Fig. 2, right) bright yellow, 2.5-5 mm long on a stipe (0.1-)0.2—0.3 mm long, glabrous; tepals monomor-

phic, oblong-obovate; stamens slightly exserted with 2.5—5 mm long, slightly pilose basally filaments and

oval, yellow anthers 0.2-0.3 mm long; pistil with styles 1.5-2 mm long. Achenes light brown, narrowly

trigonous, 4—5 mm long, glabrous; embryo straight.

Additional material seen. U.S.A. Oregon. Douglas Co.: Umpqua National Forest, on the sl f Acker Rock, 528938 E, 4766230 N,

Zone 10, 43*03'01"N oe (NAD 27), 1045 m, 28 Sep 2005, York & Rusch 2899 (HSC); SE side of Acker Rock, 528988 E, 4766416

N, Zone 10, 43°03'06"N, 122°38'39"W, (NAD 27), 1185 m, 9 Aug 2006, York & Helliwell 2912 (BH, HSC).

Distribution, habitat, and phenology.—Eriogonum villosissimum (Acker Rock wild buckwheat) is a rare taxon

found in Western Cascade Range physiographic province (Fig. 3), near where it interfaces with the Klamath

physiographic province. The area is extremely complex geologically. The only known population occurs

on the southwest to southeast exposures of the vertical faces of Acker Rock in the South Umpqua River

watershed. The site is 30.5 kilometers due west of the northwest corner of Crater Lake National Park. This

formation towers over 600 meters above the surrounding landscape. Acker Rock is located in the older

Western Cascade Range and is not capped by younger volcanic material like the High Cascade Range

(Kays 1970). It is pyroclastic rock formed between the late Eocene to late Miocene according to Kays who

mapped Acker Rock as andesite rhyodacite. Orton (2007) refers to Acker Rock as quartz latite, a specific

type of andesite rhyodacite.

The plants thrive in full sun where there is little competition from mosses or other vascular plants.

Acker Rock wild buckwheat becomes sparse or EON pus we - vegetation is dense due to

shade or gentler slopes. There appear to be severa 5

to 1253 m in elevation. Acker Rock is bordered by forest dominated by Douglas: -fir (Pseudotsuga menziesii

(Mirb.) Franco) and madrone (Arbutus menziesii Pursh), with shrubby openings consisting of greenleaf

manzanita (Arctostaphylos patula Greene), and Fremont silk tassel (Garrya fremontii Torr.). Other associates

include Brodiaea elegans Hoover ssp. elegans, Bromus tectorum L., Cheilanthes gracillima D.C. Eaton, Elymus

elymoides (Raf.) Swezey, Eriogonum compositum Douglas ex Benth., Eriogonum nudum Douglas ex Benth.,

Holodiscus cae (Pareh) Maxim., Koelerig daba (Ledeb.) Schnift. Lomatium hallii (S. Watson) J.M.

Coult. & Rose, I Benth P (Piper) Howell, aa imbricans (D.C. Eaton)

D.H. Wagner, Selasinela wallacei Hieron., and Vulpia myuros (L.) C.C. Gm

Flowering occurs from June through August. Although no seedlings have been observed, the authors

have seen juvenile plants as young as four or five years old.

Relationship —Eriogonum villosissimum belongs to subg. Oligogonum Nutt., a taxon of some 35 species

(Reveal 2005) that ranges from Alaska to Mexico and from Virginia and West Virginia to the Pacific coast.

While many of the species are widely distributed (e.g., E. umbellatum Torr., E. heracleoides Nutt., E. flavum

Nutt.), several have a limited distribution; however, few can match the restricted distributions of E. vil-

losissimum and its near relative, E. alpinum Engelm. of Scott Mountain, Cory Peak and Mt. Eddy along the

Siskiyou and Trinity county line of northern California. The new species may be distinguished by its single

involucre with erect teeth atop a stem-like peduncle that is separated from a true aerial stem by a whorl of

foliaceous bracts, its glabrous, short-stipitate flowers, villous leaves, stems, peduncles and involucres, and

by its branched, matted habit of numerous, loosely arranged rosettes bearing tufts of ovate leaves.

In the subg. Oligogonum treatment in Flora of North America (Reveal 2005: 331) the new species will key

immediately to lead 27:

641

Reveal et al

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Fic. 1. Habit of Eriogonum villosissimum.

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m (Acker Rock) |

EN Ashland | | i

== | T | T T |

0 75 150 300 Kilometers

Fic, 3. Map of Oregon with location.

27. Perianths sparsely pubescent abaxially; North Coast Ranges, Trinity and Tehama counties, California

112. Eriogonum libertini

27. Perianths glabrous abaxially; Cascade Range of o County, Oregon, Mt. Eddy and Scott Mountain,

Siskiyou and Trinity counties, a or mete ione

28. Stipe 1-1.5 mm; leaf blade ti rra Nevada 116. Eriogonum prattenianum

28. Stipe (0.1-)0.2-0.8 mm; leaf blades ovate or r oval to round-oval; Cascade Range of Douglas County,

Oregon, or Mm Paay and Scott Mountain, Siskiyou and rà counties, California.

1-)0.

| E LI

eal mm; Douglas County, Oregon

beiow 2000 m elev. 139a. Eriogonum villosissimum

29. Leaf-blades oval to round-oval, densely | il tipe 0.5-0.8 mm; Mt. Eddy an

Scott Mountain, Siskiyou and Trinity counties, California, above 2000 m elevation 140. Eriogonum

alpinum

Conservation.—Acker Rock wild buckwheat, previously unknown and uncollected, is rare due to its lith-

ophytic nature on old volcanic rock. The area is managed by the United States Forest Service as a timber

emphasis area. There is an unmanned lookout on Acker Rock available for seasonal use by the public as a

rental; it is accessible by way of a short hiking trail through the forest on the northeast side of Acker Rock.

Because the Eriogonum is confined mainly to nearly vertical rock faces, technical rock climbing is the only

way of reaching most of the population. As a preface to his description of the 20 established climbs on Acker

Rock, Orton (2007) points out to climbers that the rare buckwheat is "easily avoided while climbing." An-

thropogenic impacts are more likely from climbers establishing new routes, a factor that will be critical in

Reveal et al., Eriog illosissi : peci f 0 643

future management decisions. Although seedlings are the most vulnerable to impacts, the seasonal closure

(1 January to approximately 15 July) for nesting peregrine falcons on Acker Rock affords some protection

during the early growing season.

ACKNOWLEDGMENTS

We wish to acknowledge, with thanks especially by the senior author, the assistance given to us by Gregg

Orton, Bobbi Orton and Harold Hall, all adventurous climbers, who aided us in obtaining a small collection

of this rare SPECIES: Without d efforts far ba observations would have been made. For views of Acker

Rock, see http://w : in/rock/153184/acker-rock.html and in particular http://

www. oT enaa 1640) 153184/acker-rock. html, a most interesting place to collect a plant.

We are grateful to Kenton L. Chambers for his prompt and useful review and, as always, Barney Lipscomb

for his comments and help.

REFERENCES

Kays, M.A. 1970. Western Cascades volcanic series, South Umpqua Fall region, Oregon. The Ore Bin 32:1-94.

Orton, G. 2007. Rock climbing western Oregon. Volume 2. The Umpqua. Mountain N' Air Books, La Crescenta,

ERE J.L. 2005. 44a. Polygonaceae Jussieu subfam. Eri id Arnott, Encycl. Brittannica (ed. 7), 5:126. 1832

- Wild buckwheat subfamily. Fl. N. Amer. 5: 218-478.

IR h Institute of Texas 3(2)

BOOK REVIEW

Kevin WiLLiams. 2007. Seed to Elegance: Kentia Palms of Norfolk Island, South Pacific. (ISBN 978-0-

9775121-1-9, pbk.). First published by Studio Monarch, Norfolk Island, 2899, South Pacific. (Orders:

in com, 1-248-885-9255). $18.95, 72 pp., numerous color and b&w images, 7" x 9".

th rigins , industry,

Ç Tta E] ] f£ Klavfalb Iceland , South Darifir 4 tn th that dat

J d b icc I

and discs of the kentia palm. The fi | ides tl d th detail he hi f the kentia palm f yon

ne Howe n to its RH, at various locations MM the world. This species was initially thought to I gn of wealth and wa

tant that d landscaping. TI thor tl 1 look at the specific ste]

that are involved: in da industry of dd , seed slanting: exporting, tribution. He provides tl ler with a history of how

the nsu grew and has Capana to Į tructi he prog juired by kentia palms.

4 J; 3 a ES f. T us E de CH 11 TL 1 ] +l : 1

ALLA BML LACE Lnd &e F J PARAR CLIC LUUAL Lite VILA

"E : 204 1,11 , -03 ... "n - 1 p A Aienorceal nf the

history of this s particular planis This book was a very q Į g y p t

If Ts +1 is 1 qs x 1 : I 1 E avi

ntia p g y I g I I Keri

McNew, MS S Prog Manager, Botanical R h Institute of Texas, Fort Worth, Texas 76102-4025, U.S.A.

J. Bot. Res. Inst. Texas 3(2): 644. 2009

NOMENCLATURAL INNOVATIONS IN PHLOX (POLEMONIACEAE),

WITH UPDATED CIRCUMSCRIPTION OF

P. CAESPITOSA, P. DOUGLASII, P. MISSOULENSIS, AND P. RICHARDSONII

James H. Locklear

7431 Briarhurst Circle

Lincoln, Nebraska 68506, U.S.A.

jimlocklearegmail.com

ABSTRACT

[ f +l LES mL? 1 1 $ J (TL mr “11

jonesii M Esas comb et stat. nov.; Phlox austromontana subsp. lutescens (S.L. ee Pone e et stat. nov.;

i (E.E b

et stat. nov.; Phlox pi osa L. su sp. 1 gip ockle ear,

comb. et stat. nov.; Phl ichard i Hook cibsn I lis (wl klear, comb. nov.; Phl bulata I ee setacea (L.)

Locklear, comb. et stat. nov.; Phlox villosissi bsp. latisepal (Wh ) NEN: comb. nov. Taxonomic circumscriptions of P.

caespitosa, P. douglasii, and P. richardsonii are provided in support of tH l changes, along with an updated circumscrip-

tion of P. missoulensis.

RESUMEN

Codd ` Td 1 1 | A RAR AD | ai " + z + m DLI DL! 4 4 r~ -11

pora vec Locklear, comb. et stat. nov.; ape austromontana subsp esten (S.L. Welsh) Locklear, comb. et stat. nov.;

covillei (E.E. Nel kl comb. et stat. nov.; Phlox pilosa I subsp

comb. et stat Phl ick i Hook sies l lis (WI ) klear, comb Pl bulata I RUBER setacea (I

Locklear, comb. et stat. nov.; Phlox villosissima subsp. latisepala (Wherry) Badar oak: nov. Se aportan las circunscripciones

taxonómicas de P. caespitosa, P. douglasii, y P. richardsonii en apoyo de estos cambios nomenclaturales, junto con una circunscripción

puesta al día de P. missoulensis.

Phloxes are ip aon ofa ane diverse array of plant communities i idc systems across

North America, and are ofte tcommon |

which they

occur. le include Phlox diffusa in subalpine grasslands of the Olympic Mountains, P. muscoides in

Wyoming Basin rock outcrop communities, P. pulvinata in Rocky Mountain alpine fell-felds, and P. subulata

in barrens and glades in the eastern United tales:

he most recent hon the genus P} (P ] | 1S that coe ERU A forthcoming

manuscript on Phlox a in prep.) latural cl ber of taxa, proposed here

to make these names available for the won Four of these are E panes in rank from variety to subspecies

for consistency with concepts of rank in the forthcoming work and those of Wherry, who used “subspecies”

in preference to “variety.” The concept of subspecies is used here to delimit population systems that exhibit

geographic variation and distinctive ecological associations

Justification for the proposed new combination of P. condensata subsp. covillei requires updated cir-

cumscriptions of P. He re i E mae are Doce’ here; Justification for the proposed new

combination of P. richard tion of P. richardsonii, also provided.

Additionally, an updated P Un of P ES is —

NOMENCLATURAL INNOVATIONS IN PHLOX

Phlox austromontana Coville subsp jonesii (Wherry) Locklear, oe et stat. NOV. Basionym: Phlox jonesii Wherry,

Notul. Nat. Acad. Nat. Sci. Philadelphia 146:8, fig. 5. 1944; P} Wherry) S.L. Welsh, Great Basin

Naturalist 45:792. 1985. Type: UNITED STATES. Utan. Washington Co.: Zion Canon, 07 Mi 1923, M.E. TE s.n. (HOLOTYPE: USI;

isotypes: CAS, PH, digital image! RSA-POM, digital image!).

J, Bot. Res. inst, Texas 3(2): 645 — 658, 2009

646 t tani titute of Texas 3(2)

Phlox austromontana Coville subsp. lutescens (S.L. Welsh) Locklear, comb. et stat. nov. Basionym: Phlox aus-

tromontana Coville var. lutescens S.L. Welsh, Great Basin Naturalist 45:792. 1985; Phlox lutescens (S.L. Welsh) S.L. Welsh, Utah Fl.

ed. 3. a 2003. Tree: UNITED STATES. Urau. Garfield Co.: T33S, R14E, SW1/4 Sec. 1, along Orange Cliffs Rd, E of hwy 95, 4500

ft elev., Rimrock — blackbrush, ash, squawbush, comm., 11 May 1983, S.L. Welsh, B. Welsh & M. Chatterley 21972 (HoLotyPE: BRY;

isotypes: NY, internet amd

In his paper on the P. austromontana complex, Welsh (1985) recognized four varieties—the first-described

and most widespread form (var. austromontana) plus three intergrading expressions (vars. jonesii, lutescens,

and prostrata) of much more pee RISEEIDUCIOR occurring in the dcin country of southern Utah and ad-

jacent Arizona. These taxa are hed at the rank of subs of their discrete distributions

and unique ecological associations. The combination P. austromontana subsp. prostrata has been published

previously (Wherry 1939).

Phlox condensata (A. Miri E. E. Nelson ond covillei (E.E. Nelson) Locklear, comb. et stat. nov. Basionvm:

Phlox covillei E.E. Nelson, Revis. WN. Amer. Phlox 15. 1899. Tree: UNITED ~ CALIFORNIA: Inyo Co.: Black Canon, White

Mountains, 08 Jul 1891. EV Be with F Funston 1801 ( y , designated by Wherry 1942: US!;

isotypes: B, GH! K, NY, internet image! PH, digital image).

The taxonomic history of P. condensata is complex. Gray (1870) described P. caespitosa var. condensata from

type material collected by Charles Christopher Parry in alpine habitat in the Southern Rocky Mountains

of Colorado (Hololtype: GH!; Isotypes: ISC NY, internet image! PH, digital image!). Elias Nelson (18992)

elevated it to species rank as P. condensata. Wherry (1941, 1955) reduced it to P. caespitosa subsp. condensata,

but Cronquist (1959) elevated it back to species rank. As detailed in the circumscription of P. caespitosa that

follows, P. condensata is treated here at the rank of "Renee

Elias Nelson (1899) described P. covillei from ty] ial collected in the White Mountains of eastern

California, and subsequent exploration of the Intermognrain West revealed this entity to occur in alpine/

subalpine situations in a number of other high elevation mountain ranges of the Great Basin as well as in the

Sierra Nevada. Wherry (1942, 1955) affirmed recognition of this taxon at the species level, but Cronquist

(1959) reduced it to synonymy under P. condensata. Wherry (1969) disagreed, defending recognition of P.

covillei on the basis of morphological differences and complete geographic separation from P. condensata

of the Rocky Mountains, but Cronquist (1984) was not convinced, noting, "I am unable to distinguish the

Colorado plants from more western materials." While morphological differences cited by Wherry are not

profound (see key below), the taxon covillei warrants recognition as a subspecies because of its entirely

separate range, a relationship paralleled in the similar bi-centric distribution pattern of the bristlecone pine

complex, with the Southern Rocky Mountain expression recognized as Pinus aristata and the Intermoun-

tain expression as Pinus longaeva (Bailey 1970). Phlox condensata is strongly associated with bristlecone pine

subalpine woodland, subsp. condensata with Pinus aristata in the Rockies of Colorado and New Mexico and

subsp. covillei with Pinus longaeva in Great Basin mountain ranges in California and Nevada.

KEY TO THE SUBSPECIES OF PHLOX CONDENSATA (A. GRAY) E.E. NELSON

P| i | ti rse, leaves linear-subulate, ciliate; inflores-

cence herbage UE pubescent; Southern Rocky a in Colorado and New Mexico

subsp. condensata

Plant composed of interlacing shoots; pubescence rather copious; leaves narrow-elliptic, copiously coarse-

ciliate, the margin markedly thickened; inflorescence herbage densely pubescent with coarse hairs, only

some gland-tipped; Sierra Nevada of California and mountains of the Great Basin in California and Nevada

subsp. covillei (E.E. Nelson) Locklear

Interpreting floristic and ecological literature in relation to P. condensata is made difficult by the confused

T sd: 1: PN NE a ] :. 1 s LT 4 CE : P" y AS ] , f 11

as P. condensata) includes these: Baker 1983; Boyce et al. 2005; Cockerell 1903; Cox 1933 (as P. caespitosa);

Hartman & Mitchell 1979; Hartman & Rottman 1985, 1988 (as P. pulvinata); Kiener 1967; Langenheim 1962

Locklear, Nomenclatural innovations in Phlox 647

(as P. caespitosa); Shepherd 1975 (as P. caespitosa), Weber & Wittmann 2001a, 2001b; Wooton & Standley

1915 (as P. caespitosa). Literature pertaining to subsp. covillei (generally as P. covillei) includes these: Billings

& Thompson 1957; Botti & Sydoriak 2001 (as P. condensata); Chabot & Billings 1972; Clokey 1951; Goodrich

1981 (as P. condensata); Kartesz 1988; Kimball et al. 2004 (as P. condensata); Krantz 1994 (as P. condensata);

Linsdale et al. 1952; Lloyd & Mitchell 1973; Major & Taylor 1977; Marchand 1973; Mooney et al. 1962;

Munz 1974; Munz & Keck 1959; Nelson & Kennedy 1906 (as P. dejecta); Patterson & Wilken 1993 (as P.

condensata); Peinado et al. 2005; Pemble 1970; Sharsmith 1940 (as P. caespitosa var. condensata); Spira 1991

(as P. condensata), Wenk & Dawson 2007 (as P. condensata), Wherry 1942; Wright & Mooney 1965.

Phlox pilosa L. subsp. longipilosa (Waterf.) Locklear, comb. et stat. nov. Basion: Phlox longipilosa Waterf., Rhodora

73:577. 1971; Phlox pilosa L. var. longipilosa (Waterf.) J. & C. Taylor, Sida 9:183. 1981. Type: UNITED STATES. OxLaHomMa. Kiowa

Co.: N slopes of granite mountains S of Lake Altus, 29 May 1957, U.T. Waterfall 13134 (HoLorvrE: OKLA!; isotypes: GH! KANU,

OKL, PH, digital image!).

Waterfall (1971) recognized this entity as related to but distinct from P. pilosa. Taylor and Taylor (1981)

reduced it to a variety of P. pilosa. Phlox pilosa subsp. longipilosa is endemic to granitic soils in the Wichita

Mountains region of southwestern Oklahoma, remote from the main distribution of P. pilosa to the east. It

is distinguished from the broader P. pilosa Na RN as shown in the adn: key.

Stems a piani numerous, with 10-18 leaf. ; Up] ; y

t | calyx densely pilose ith long jointed hai tly 2-4 mm long;

SE -lobes apically bearing relatively long, somewhat twisted awn Phlox pilosa subsp.

ongipilosa (Waterf) Locklear

Stems per plant few, ca. 6-14 leaf-nodes; uppermost leaves and bracts mostly opposite; hairs of upper stems

and calyx less than 1 mm long; calyx-lobes aristate Phlox pilosa sensu lato

Phlox ios. subsp. borealis (Wherry) Locklear, comb. nov. Basionym: Phlox borealis Wherry, Gen. Phlox: 126.

5; Phlox sibirica L. subsp. borealis (Wherry) shaver Luan 65:220. 1963; Phlox d L. var. edu (Wherry) B. Boivin,

ici Canad. 93:1062. ies ALASKA: Nome, Alaska o ridge, altitude 500 ft, slope 20%, moisture

fair, rocky lo I pilobium, Salix reticulata, Cassiope tetragona, A lticeps, 2 1929, WB. Miller 121 — c

(HOLOTYPE: BA

Hooker (1838) described P. richardsonii from type material collected by John Richardson while serving on

Captain John Franklin’s Second (1825-1827) Overland Expedition to the Arctic (HOLOTYPE: K; ISOTYPES: BM,

digital image! GH! NY, internet image!). Richardson collected the type in 1826 near Cape Bathurst on the

Arctic seacoast of northwest Canada.

The identity of P. richardsonii has long been tied up with that of P. sibirica, described by Linnaeus (1753)

from original material collected in “Asia boreali,” most likely in southern Siberia. Although Hooker (1838)

considered the two species distinct, he identified specimens of P. richardsonii collected in 1826 by Lay and

Collie at Kotzebue Sound on the Alaskan coast as P. sibirica, noting the plants differed “in no dala ' from

material collected in Asia. The treatment of phloxes from the North American Arctic as P. sibirica continue

well into the twentieth century, as exemplified by the works of such notable students of the Arctic flora as

Porsild (1939a, 1939b) and Hultén (1948, 1968).

Wherry (1955) separated the phloxes of the North American Arctic from P. sibirica, recognizing P. bo-

realis, P. richardsonii subsp. alaskensis, and P. richardsonii subsp. richardsonii. Major morphological differences

cited by Wherry are reflected in the key below. Not noted by Wherry are significant ecological differences,

P. sibirica being associated with mid-elevation, graminoid-dominated steppe communities, primarily in the

cold, dry interior plateaus of Middle Asia (Ermakov et al. 2002, 2006; Grubov 2001; Knyaszev et al. 2007,

Krasheninnikov 1937; Mirkin et al. 1985; Peshkova 1972, 2001; Sochava 1945; Yurtsev 1963, 1967, 1972), and

P. richardsonii occurring almost entirely above the Arctic Circle in association with alpine dwarf-shrubland

and alpine herb communities.

Plant a tandi ffrut t i perennial, sparingly rane bird branched, forming tufts 8-15 cm tall from

li ith ca. 5 leaf-nodes below the inflorescence;

d -

a3 taproot, tha arart

648 tani Institut Texas 3(2)

leaves linear, long-acuminate, falcate, thinnish, i length 30-60 cm; inflorescence (one-)

three- to six-flowered, maximum pedicel length 20-40 mm; ovary/capsule typically with 2 ovules/seeds per

pene Phlox sibirica

Pla . “EL lu E H j | te 2219 tal E l^ -l ln m

flowering shoots 2-6 cm long; leaves linear to subulate, maximum length 5-15 mmy inflorescence one- to

three-flowered, pedicel length 1-8 (up to 25) mm; ovary/capsule typically 1 ovule/seed per locule Phlox

richardsonii complex

Phlox richardsonii is recognized here as a complex of intergrading forms largely limited to HN western North

American Arctic, differing morphologically in stature a nd size of floral parts and inct floristic

regions of the Arctic. As shown in the key below, t I ized here under the first-published

name of P. richardsonii, necessitating the new combination of P. cians subsp. borealis.

KEY TO THE SUBSPECIES OF PHLOX RICHARDSONII HOOK.

—

. Longest pedicels (4-)8-25 mm long; corolla-lobe 8-14 mm long; styles 7-12 mm long; associated with

coastal areas of the Penna Strait — subsp. borealis (Wherry) Locklear

. Longest pedicels 1—6 ro 6-9 mm long; styles 4-8 mm |

2. Leaves flattened, thinnish, 8- 15 | ith ratl hai lyx 7-10 mm long; corol-

la-limb diameter up to 2.2 cm; macsecinicd with higher interior foothills and mountains of Alaska, Yukon,

—

and the Northwest Territories subsp. alaskensis (Jordal) Wherry

2. Leaves awl-shaped, thickish, 5-10 mm long; pubescent with fine, arachnoid hairs; on 5-7 mm long;

corolla-limb diameter ca. 1 cm; associated with the Arctic seacoast subsp. richardsonii

The distribution of the P. richardsonii complex ranges from the Bering Sea region of Alaska east to northwest

Canada, mostly north of the Arctic Circle, with a few highly limited occurrences reported from Asia. The

first-described expression (subsp. richardsonii) is known from relatively few stations in northwest Canada

and northern Alaska. The core of its range is along the Arctic seacoast of Canada, from Cape Bathurst in

the District of Mackenzie of the Northwest Territories, east to Clifton Point, a mainland promontory on

the coast of Nunavut Territory. It also occurs on Banks Island of the Canadian Arctic Archipelago. Floristic

and ecological literature pertaining to subsp. richardsonii (generally as P. richardsonii) includes these: Aiken

et al. 1999; Freedman et al. 1990 (as P. sibirica), Holm 1922; Johansen 1924; Kuc 1974, 1996; MaCoun @

Holm 1921.

Phlox richardsonii subsp. alaskensis occurs in the higher interior foothills and mountains of Alaska,

Yukon, and the Northwest Territories. It occurs most abundantly in the Brooks Range of northern Alaska,

but also in the Alaska Range of central Alaska and in the Mentasta, Nutzotin, and Wrangell mountains in

south-central Alaska. It also occurs in the Ogilvie Mountains and Richardson Mountains of the Yukon Ter-

ritory. Floristic and ecological literature pertaining to subsp. alaskensis includes these: Carlson et al. 2006a

(as P. alaskensis); Cody et al. 2002 (as P. alaskensis); Cook & Roland 2002 (as P. richardsonii); Cooper 1989

(as P. sibirica); Drew & Shanks 1965 (as P. sibirica), Jordal 1952 (as P. alaskensis), Juday 1989 (as P. sibirica);

Porsild 1972 (as P. alaskensis); Spetzman 1959 (as P. sibirica); Walker et al. 1989, 1991, 1994 (as P. sibirica).

Phlox richardsonii subsp. borealis is distributed along the coast of the Seward Peninsula of northwest

Alaska, in the vicinity of the Bering Strait. It also has been reported (as P. alaskensis) on the Asiatic side of the

Bering Strait in the far southeastern portion of the Chukotka Peninsula of the Russian Far East (Balandin &

Razzhivin 1980; Razzhivi 1986, 1994). On American side of the Strait, occurrences of P. richardsonii subsp.

borealis are known from coastal areas of Norton Sound and Kotzebue Sound, and from Cape Krusenstern,

cape PisbUmIE; Cape Prince of wee Cape Sabine, and Cape Thompson. Floristic and ecological literature

tai to subsp. borealis incl tl Carlson et al. 2006b (as P. alaskana), Hansen 1951 (as P. sibirica);

Hutchison 1934 (as P. sibirica); Johnson et al. 1966 (P. sibirica), Kelso 1989 (as P. sibirica); Muir 1917 (as P.

sibirica); Porsild 1939a, 1939b (as P. sibirica), Razzhivin 1986, 1994 (as P. alaskensis); Shetler 1963 (as P.

sibirica subsp. borealis).

The geographic distribution of these subspecies show parallels with patterns of speciation in the genus

Douglasia (Primulaceae) (Kelso 1992; Kelso et al. 1994). It also roughly corresponds to the extent of three

Locklear, Nomenclatural innovations in Phlox 649

floristic subdivisions of the Arctic described by Yurtsev (1994), each reflecting regional differences in envi-

ronment and climate. Subspecies richardsonii is associated with the Central Canada subprovince, which is

markedly continental. Subspecies alaskensis is associated with the North Alaska subprovince, which encom-

passes the more continental parts of the Brooks Range and its northern foothills, the Richardson Mountains,

and the Arctic slope of Alaska with its cold, true Arctic climate, a region which experiences colder winter

temperatures than coastal areas but also experiences more summer heat and dryness. Subspecies borealis

is associated with the Beringian Alaska subprovince, which encompasses the Alaskan coast of the Bering

Strait, notably the Seward Peninsula and coastal areas of Kotzebue Sound, Norton Sound, and the Chukchi

Sea. This subprovince has an oceanic climate due to the influence of a warm sea current. While the sum-

mer climate along the Bering Strait is generally colder, windier, and more overcast than continental areas

of Alaska, the overall climate is less harsh than in the interior mountains due to the proximity of the sea.

Species borealis also occurs on the Asiatic side of the Bering Strait on the coast of the Chukotka Peninsula

in the parallel Beringian Chukotka subprovince. A number of vascular plants are considered “Beringian”—

characteristic of or endemic to the Bering Sea region on both the American and Asiatic sides of the Strait

(Kelso 1987, Yurtsev 1994), with the floras of these coastal areas being more diverse than that toward the

interior (Kelso 1989).

Phlox subulata L. subsp. setacea (L.) Locklear, comb. et stat. nov. Basion: Phlox setacea L., Sp. Pl. 1:153. 1753; Phlox

mee L. ps eu-subulata arn) var. setacea dad Beane dc o al 78. 1907. LECTOTYPE, MH cis by Reveal

8, fig s

: Al magestum

GO DAT CT

ee 233. 1696; Tyroryre, designated iy Reveal et a 1982: Herbarium Sl 90: 59 (BM-SL

Linnaeus established the name P. setacea in Species Plantarum (1753) based on a figure (table 98, figure 3)

published in Plukenet's Phytographia (1691). The epithet setacea was subsequently misapplied by Curtis

(1798) in the Botanical Magazine for what was an illustration and description of P. nivalis (Loddiges 1823;

Sweet 1827) and the two names became entangled until Wherry (1929, 1937) sorted out the confusion and

reinstated P. nivalis. Wherry 1623) == P. setacea as a glabrate expression of P. subulata and later

(1955) 1 placed the Linnaean y nder P. subulat ] I subulata. Reveal et al. (1982) located

a dear specus in the Sloane Herbarium of the Natural History Museum, London (BM-SL), which they

onize of the Plukenet figure. Noting that tl learly matched the attributes of

wat Wherry (1929) eed as P. subulata var. australis, Reveal et al (1982) demonstrated that the Linnaean

epithet setacea has priority over australis, and resurrected P. subulata var. setacea, a name proposed by Brand

(1907). This taxon is recognized here at the rank of subspecies.

Phlox villosissima (A. Gray) Small subsp. latisepala (Wherry) Locklear, comb. nov. Basiovvw: Phlox pilosa L. subsp.

latisepala Wherry, Gen. Phlox 47. 1955. Tyre: UNITED STATES. Texas. Kerr Co.: 1600-2000 ft, 19-25 Apr 1894 A.A. Heller 1641

(HOLOTYPE: MO; isotypes: G, GH! NY, internet image! PH, digital image! US!).

Gray (1870) described P. drummondii var. villosissima f terial collected by Charles Wright in 1851

on the “Pebbly bars of the Nueces [River]” in southern Tass (HOLOTYPE: GH!). Small (1903) elevated it to

species rank, but still considered P. villosissima an annual like P. drummondii. Whitehouse (1935) relocated

the likely site of Wright’s collection by searching crossings of the Nueces River near Uvalde, Texas where

she found “numerous plants that closely matched the type” growing in the chalky gravel beds of the river

and determined that the entity is a perennial rather than an annual, providing further merit for recognition

as a species. Wherry (1955) recognized the distinctiveness of this taxon, but treated it as P. pilosa subsp.

riparia.

Levin (

ogy, flowering season, and incompatibility era Levin initially supported his view with evidence from

karyology (Levin 1966; Smith & Levin 1967) and experimental crosses (Levin 1966). Phlox villosissima is a

tetraploid (2n = 28) and appears to be an allotetraploid. These data, coupled with those from studies of seed

protein chemistry (Levin & Schaal 1970) and plant pigment (flavonoid) chemistry (Levy & Levin 1971,

1974) indicate P. villosissima could be the stabilized derivative of past hybridization between P. pilosa subsp.

Tfxz O^

) reinstated P. villosissi ies distinct from P. pilosa, reproductively isolated by ecol-

|| || £ al nm H in L | + "mir

650 t t f Texas 3(2)

pilosa and P. drummondii subsp. mcallisteri. Evidence from more recent molecular studies generally support

this hypothesis but also indicate more complicated relationships (Ferguson et al. 1999; Ferguson & Jansen

Elias Nelson (1899) described P. aspera from type material collected by Amos Arthur Heller in central

Texas. Brand (1907) discerned a relationship with P. villosissima, treating the taxon as P. drummondii subsp.

drummondii var. villosissima subvar. aspera. Wherry (1955, 1966) also recognized a close relationship be-

tween the two, but, as with P. villosissima, treated P. aspera as a variant of P. pilosa, proposing P. pilosa subsp.

latisepala (Wherry noted Nelson’s aspera was a homonym and unavailable as a subspecific epithet). Levin

(1968) reinstated P. aspera, citing the same justifications as for P. villosissima and noting the entity occupied

an intermediate position between P. villosissima and P. pilosa in terms of morphology and ecology. Both Levin

and Wherry considered the combined range of this pair of taxa to be centered on the Edwards Plateau of

Texas, with P. villosissima/P. pilosa subsp. riparia most common in the western portion of the plateau and P.

aspera/P. pilosa subsp. latisepala occurring more to the east and north.

Ferguson (1998) cited possible local integration of these two entities wate each other and with P. pilosa

(also noted by Wherry [1966]) as reason for maintaining them at the subspecies level pending further study.

In the most recent treatment of Phlox in Texas, Turner (1998) estbined both under P. villosissima. While

these taxa may intergrade where their ranges overlap, they show distinctive morphological differences and

ecological associations toward the outer portions of the overall range of the complex (see key below). The

taxon formerly treated as P. aspera and P. pilosa subsp. latisepala is recognized here as a subspecies of P. vil-

losissima, necessitating the new combination.

KEY TO THE SUBSPECIES OF PHLOX VILLOSISSIMA (A. GRAY) SMALL

| I I] * IH

ant with \ ell

to be branched with nodes Sr crowded 12- 25 cm tall inflorescence herbage densely ra

pubescent; xeric habitat in western portion of species range, notably riverwash habitat and talus slopes

subsp. villosissima

Plant ane mlizommetous, i stema tending to be simple, OO ou 22- 45 cm tall; pubescent eed

t n east

portion of species range subsp. ade on Locklear

CIRCUMSCRIPTION OF PHLOX CAESPITOSA NUTT.

Nuttall (1834) described P. caespitosa from type material collected by Nathaniel Jarvis Wyeth in 1833 on

the return (eastbound) leg of his first Oregon Expedition (1832-1833). Nuttall described the type locality

as “Flat-Head river, on the sides of dry hills,” adding, “Flowering about the 20" of April.” The date “April

227^" is written on the holotype specimen label (BM) and on isotypes (GH, PH) of P. caespitosa. According

to Wyeth’s journal, 22 April 1833 was the day he departed “Flathead Post,” also called “Flathead House,”

a trading post historians place not on the Flathead River but on the Clark Fork River near the present-day

town of Thompson Falls in northwestern Montana (Sanders County) (McKelvey 1955). A number of more

recent collections of P. caespitosa have been made in the vicinity of Thompson Falls.

Wherry (1941, 1955) interpreted P. caespitosa as a complex consisting of a taller, open-growing form

(subsp. caespitosa) occurring at middle elevations in the Rocky Mountains, plus three shorter, more compact

expressions (subspp. condensata, platyphylla, and pulvinata) occurring at higher elevations under alpine/

subalpine conditions. Cronquist (1959) revised this complex, raising the high elevation forms to the rank

of species as P. condensata and P. pulvinata (incorporating subsp. platyphylla) and interpreting P. caespitosa

as a taller, open-growing species of lower elevations, primarily in Pinus ponderosa woodland and savanna.

Wherry (1962, 1965a, 1965b, 1969) took ption with these revisions, considering P. condensata and

P. pulvinata mere high altitude ed pressionei or "EROS of P. caespitosa. However, the ecological divergence of

these alpine/subalpine entities from P. caespit 1, and, following Cronquist, P. condensata and

P. pulvinata are recognized here as separate sree

Locklear, Nomenclatural innovations in Phlox 651

. Plant a subshrub, eee m diffusely EREN aua open tufts 5- 15 cm tall, the aloe

Nos branche nd long, yl pl th montane wo

savanna and grassland Phlox caespitosa

1. Plant a caespitose lit herb, growth habit pulvinate, forming more or less compa mee or cushions

2-7.5 cm tall, the flowering shoots 1-3(-5) cm long; associated with alpine or upper subal pine ecological

systems.

2. Plants forming dense, tight cushions, es closely packed ther; | pI |, erect, 5-10 mm

long; corolla-tube 6-10 mm long, lobes 3-5 mm long; styles 1 a 3 mm long Phlox condensata

2. Plants forming loose, open cushions, shoots more spreading; leaves not appressed to stem, 6-12 mm

long; corolla o. larger, the tube 7-14 mm long, lobes 6-7 mm long; styles 2.5-5.5 mm long

Phlox pulvinata

While bringing this helpful clarification to the P. caespitosa complex, Cronquist (1959) unfortunately added

confusion by reducing the name P. douglasii to synonymy under P. caespitosa. These are recognized here as

se Spon See discussion of P. douglasii that follows).

| here, P. caespitosa is a species of the Rocky Mountain System, chiefly in the Northern

Rocky Mena in southeastern British Columbia, northern Idaho, aan western Montana. k occurs most

abundantly in Montana, particularly on the eastern slope of the Bitterroot } t

tain valleys of the Rocky Mountain Trench. It occurs at lower to midd ti osa 800-1500 m, but

up to 2040 m) in the piedmont and mountains, and in association with morainal features of river valleys.

Phlox caespitosa is strongly associated with Pinus ponderosa woodland and savanna, but ad is common in

montane, foothill, and valley grasslands in the region where P spicata, Fest is, and

Festuca campestris are the chief graminoids.

SUCTUS floristic and ecological literature in relation to P. caespitosa is made difficult by the con-

fused ling this plant, particularly in tion with the name P. douglasii (discussed

below), but with added obleas coming from the literature of alpine ecology where the name P. caespitosa

has been misapplied to P. condensata and P. pulvinata. Literature pertaining to P. caespitosa includes these:

Lackschewitz 1991; Lesica 2002; Root & Habeck 1972 (as P. pulvinata); Shaw & On 1979.

CIRCUMSCRIPTION OF PHLOX DOUGLASIT HOOK.

Hooker (1838) described P. douglasii in his Flora Boreali-Americana. As one of the first of the small-leaved

cushion-forming phloxes described from western North America, the name P. douglasii subsequently served

as catch-all for many later-discovered, superficially similar species, a number of which were treated as va-

rieties or subspecies of P. douglasii at some point in their nomenclatural histories, including P. andicola, P.

austromontana, and P. diffusa. Further confusion was caused by the emergence of the term “Douglasii Hybrids

in horticultural literature in the early 1900s, which was applied to a wide variety of cushion phlox cultivars

of probable hybrid origin. Wherry brought much needed clarity to this situation through his papers on the

western phloxes (1938, 1941) and his 1955 monograph in which he recognized P. douglasii as a distinct

species with a ee aoe on the Columbia Plateau in the Pacific Northwest.

Cronquist (19 luced P. d y under P. caespitosa, justifying this treatment under his

discussion of P Dalmata which he senate’ Dom P. caespitosa. Cronquist described the type of P. caespitosa

as “a compact plant of the taxon usually known as P. douglasii.” Since the name P. caespitosa was published

by Nuttall (1834) four years before Hooker (1838) published P. douglasii, Cronquist reasoned that “the latter

name [P. douglasii].. must subside.”

Wherry (1962, 1965a) idered this “a serious misunderstanding,” calling P. douglasii and P. caespitosa

“about as distinct species as can exist among the Microphloxes.” Indeed, Wherry (1955) placed these two

species in n subsections of the genus—Douglasianae and Mn tas pee Wherry’s protests,

most subsequent floristic and ecological workers in the Pacific Northwest | l Cronquist. Based on

examination of the type material of both P. douglasii (HOLOTYPE: K, digital image!; isotypes: BM, digital image!

GH! NY, internet image! PH, digital image!) and P. caespitosa (HoLotyre: BM, digital image!; isotypes: GH!

NY, internet image! PH, digital image!), plus study of these entities in the field (P. douglasii in Washington,

652 || I £ al m FI b In LI FEF ET "fm

1%;

Oregon, and California; P. caespitosa in Montana), they are recognized here pecies. As noted by

Wherry (1962, 1965a, 1969), the leaves of P. douglasii are dark green, slender and Heels: shaped (acicular

or acerose), and covered by long gland-tipped hairs, while the leaves of P. caespitosa are pale green, relatively

broad and flat (linear-oblong), thickish, and bear coarse glandless cilia along the margins. The distribution

of P. douglasii is centered on the Columbia Plateau, while that of P. caespitosa is centered in the Northern

Rocky Mountains.

Interpreting floristic and ecological literature in relation to P. douglasii is made difficult by the con-

fused nomenclature surrounding this species. Prior to Cronquist (1959), this taxon was treated as P.

douglasii or P. rigida, the latter recognized by Wherry (1938, 1955) as a reduced expression of the former

under the name P. douglasii subsp. rigida. Following Cronquist's revisions, most workers abandoned these

names in favor of P. caespitosa. Literature pertaining to P. douglasii includes these: Applegate 1938; Culver

1964 (as P. diffusa); Daubenmire & Daubenmire 1968 (as P. caespitosa); Driscoll 1964a, 1964b; Eckert

1957 (as P. diffusa); Hall 1967; Howell 1903; Munz & Keck 1959 (as P. douglasii subsp. rigida); Peck 1941;

Piper 1906 (as P. rigida); Piper & PESE 1914 (as P. rigida); St. John 1937 (as P. douglasii var. rigida). The

following key separates P. doug P pit d, additionally, from P. diffusa, a similar-appearing

species that occurs in range.

l. Leaves pale green, lati | br oad (linear bl g) i fl t, thickish, with gl {| ili NI +| J

Middle Rocky Mountains Phlox caespitosa

1. Leaves dark or bright green, relatively narrow and slender, cilia lacking or fine; mostly west of the Rocky

untains.

2. Pubescent with glandular hairs throughout; leaves somewhat acicular (needle-shaped) and stiff, their

surfaces pilose to glabrate with fine gland-tipped hairs; primarily dry to xeric habitat on the Columbia

Patean Phlox douglasii

| j | [| Er "l EM | Il £ | LA + |:

( TALI NP, NY PIT bulate

“and only moderately rigid; primarily moderate to high elevations one the Cascade/Sierra axis

Phlox diffusa

CIRCUMSCRIPTION OF PHLOX MISSOULENSIS WHERRY

Wherry (1944) described P. missoulensis from the type collected by Merton J. Reed in 1939 on Waterworks

Hill near Missoula, Montana (HoLorYPE: PH, digital image!). Cronquist (1959) reduced this entity to P. kelseyi

var. missoulensis, a relationship and reduction with which Wherry (1962, 1965b) strongly disagreed, calling

P. missoulensis “a striking endemic. ..wholly unrelated" to P. kelseyi. In her thesis on the P. helseyi complex,

Campbell (1992) followed Cronquist in treating P. missoulensis as a variant of P. kelseyi (proposing but not

publishing P. kelseyi subsp. missoulensis), but cited “extremely different habitats" plus “complete geographical

isolation coupled with Hc in 10 popu Pince in e ol lia of missoulensis

as a distinct entity within the P. kelseyi complex ] here as sepa-

rate species (see key below). Phi

endemic to the cals Rocky Mountans) in west-central

Montana, chiefly along or just west of the Continental Divide where it occurs in the Anaconda, Garnet,

Lewis and Clark, and Sapphire ranges and on small mountains in the vicinity of Missoula, with outlying

occurrences east of the Continental Divide in the Little Belt Mountains.

Leaves dark green, linear to linear-lanceoiate, thinnish | ith sub Hat | surficial! with

fine gland-tipped hairs; exposed mountain slopes and ridges; habitat dry to xeric; soils stony and Shallow

Phlox missoulensis

Leaves pale green, linear-lanceolate inat | idate, thickis! | lent, fleshy (I

on drying), the margin slightly thickened, basally coarse- ciliat te, surficially gla! t | ly pubescent

mountains valleys and intermountain basins and flats; habitat seasonally moist to wet; soils alluvial, ine

textured, alkaline to highly-alkaline Phlox kelseyi

ACKNOWLEDGEMENTS

The author wishes to acl ledge the curators a nd staff of the following herbaria for the loan of specimens

Locklear, Nomenclatural innovations in Phlox 653

and/or providing images and label data f i ALA, BM, CAN, K, MONTU, OKLA, PH, RSA-POM,

WS. Robert Kaul, curator of NEB, was aly helpful in securing these loans. Thanks also to the curators of

BRIT, GH, RM, TEX/LL, US for access to their collections. Tis work has benefited greatly from conversations

with Carolyn Ferguson. Finally, thanks t well as Andrew Hipp, Barney Lipscomb,

Tony Reznicek, and Guy Nesom for valuable review and helpful comments on the manuscript.

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VALIDATION OF THE NAME SYMPLOCOS PENTANDRA (SYMPLOCACEAE)

Joao Luiz M. Aranha Filho Peter W. Fritsch and Frank Almeda

ot Department of Botany

rsity 4 Campinas California Academy on

CP.6109, ed "pide 0, Campinas, SP BRAZIL 55 Music Concours

a cal com oM OnE California on < 03, U. 5. A.

Angela B. Martins

Department of Plant Biology

University of Campinas

C.P 6109, CEP 13083-970, Campinas, SP BRAZIL

amartins@unicamp.br

ABSTRACT

lid ] S lra (M ) Occl ¡(Ss ion Barberina (Vell) A. DC , Symplocaceae), a name not

ay eee in 1974 because when it was s raised to the ut di species the basionym was not fully "I The name S. pentandra

1 las S. itatiaiae Wawra var. pentandra Mattos.

F de te iginally

Key Wonps: Brazil, nomenclature, Symplocaceae, Symplocos

RESUMO

lid $ tandra (Mattos) Occhioni (Sympl cão Barberina (Vell) A. DC., Symplocaceae), um nome

náo re públiadoa em 1974, j que quando foi elevad ivel de espéci itado. O nome

S. pentandra representa uma nova combinado e um novo status para a entidade originalmente descrita como $. itatiaiae Wawra var.

pentandra Mattos.

PALAVRAS CHAVE: Brasil, nomenclatura, Symplocaceae, Symplocos

Symplocos section Barberina (Vell) A. DC. (sensu Fritsch et al. 2008) comprises ca. 25 species distributed

in eastern Asia and the Americas. Species within this section are recognized by their hermaphroditic or

dioecious breeding system, basally connate corolla, stamens that are adnate to the corolla only at the base,

terete filaments that are either distinct or united only at the base, 3-carpellate ovary, 1-(2-3) locular fruit

with 1(2—3) seeds, and an embryo with the radicle usually longer than the [oo pias shorter; pian

1901; Fritsch et al. 2008). One of the species within this section is the B

(Mattos) Occhioni. This species was originally described as a variety of S. itatiaiae Wawra by Mattos (1971)

and subsequently raised to species rank by Occhioni (1974). Specimens of this species are housed in many

herbaria and the name S. pentandra is commonly used in publications on the flora of Brazil (e.g., Sobral et

al. 2006).

In the course of revising the South American species of sect. Barberina we have agreed with Occhioni

(1974) that S. pentandra should be treated as a species and not a variety of S. itatiaiae. Therefore, we accept

S. pentandra in our treatment. When raising S. itatiaiae var. pentandra to species rank, however, Occhioni

(1974) did not indicate the page where it was validly published and therefore did not fully cite the reference

of the basionym. Thus, under article 33.4 of the International Code of Botanical Nomenclature (ICBN; McNeill

et al. 2006), S. pentandra cannot be considered a validly published name. In his unpublished revision of the

Brazilian species of Symplocaceae, Bida (1995) suggested that Occhioni’s name S. pentandra can be treated

as a validly published new species because Occhioni provided a holotype that was different than that of S.

itatiaiae var. pentandra and described the plant in Latin. It is clear, however, that Occhioni intended to raise

S. itatiaiae var. pentandra to the species rank, because of the author citation “Symplocos pentandra (Mattos)

J, Bot. Res. Inst, Texas 3(2): 659 — 660. 2009

660 J the Botanical h Institute of Texas

Occhioni, stat. nov.” Nonetheless, due to his omission in providing the page where the | was validly

published and his direct reference to the basionym work, S. pentandra is consequently still hot validly pub-

lished. This is because, according to the article 33.8 of the ICBN, on and after 1 January 1953 if an author

claims to be publishing a new combination but fails to provide all the requirements under article 33.4 the

name is not validly published even if the author provided at the same time other information that would

have resulted in valid publication as the name of a new taxon (McNeill et al. 2006).

We also studied specimens identified as e ca by pos XM has been considered a

validly published name (e.g., Mattos 1969) and cited as x Gürke. Our observations

of these specimens suggest that 2 copaiaoensis makes a to the same entity as S. pentandra. However,

Gurke (1891) provided neither d illustration with analysis for S. caparaoensis, which is therefore

a nomen nudum. Thus, S. sanardoensis is not validly published under articles 32.1(d) and 42.3 of the ICBN,

respectively (McNeill et al. 2006).

We here validate the name S. pentandra. We choose to validate this name instead of S. caparaoensis

because S. pentandra is based on a validly published name.

Symplocos pentandra (Mattos) Occhioni ex Aranha, comb. et stat. nov. B

Mattos ["Symplocos itatiaiae Wawra var. pentrandra Mattos"], Loefgrenia 54:1. 1971. Tyre: BRAZIL. PARANÁ: Piraquara, Florestal, 930

m, 7 Nov 1946, G. Hatschbach 358 (HOLOTYPE: MBM; isorvPES: PACA, SPI).

ACKNOWLEDGMENTS

The first author is grateful to Conselho Nacional do Desenvolvimento Científico e Tecnológico (CNPq) for

a Ph.D. grant and the Lakeside Foundation for support during a six-month research visit to the California

Academy of Sciences.

REFERENCES

Bind, A. 1995. Revisão taxonómica das espécies de Symplocos Jacq. (Sympl ) do Brasil. Unpublished Ph.D.

thesis. Universidade d: Sáo Paulo, SP, Brazil

BRAND, A. 1901. Symplocaceae. In: A. Engler, ed. Das Pflanzenreich, IV.242 (Heft 6):1-100. Engelmann, Lipsia,

Germany.

FritscH, PW, L.M. KeLLv, Y. WANG, F. ALMEDA, AND R. KriegeL. 2008. Revised infrafamilial classification of Symplocaceae

based on phylogenetic data from DNA sequences and morphology. Taxon 57:823-852.

Gurke, M. 1981. Symplocaceae. In: A. Engler and K. Prantl, eds. Nat. Pflanzenfam., Vol. 4(1):165-172. F. Fleisher,

Lipsia, Germany.

Mattos, J.R. 1969. Observações sobre Symplocaceae. Arq. Bot. Estado São Paulo 4:269-272.

Martos, J.R. 1971. Novidades taxonómicas. Loefgrenia 54:1-2.

McneL, J., ER. Barrie, H.M. BurDer, V. Demoutin, D.L. HawksworTH, K. MARHOLD, D.H. NICOLSON, J. PRADO, PC. SitvA, J.E. SKOG,

J.H. WIERSEMA, AND J. TURLAND. 2006. International code of botanical nomenclature (Vienna Code). A.R.G. Ganter

Verlag KG, Ruggell, Liechtenstein.

OccuioN,, P. 1974. As espécies de Symplocaceae da flora do Paraná. Leandra 4-5:31-52.

SosRAL, M., J.A. JARENKOW, P. Brack, B. IRGANG, J. LAROCCA, AND R.S. RobRiGues. 2006. Flora arbórea e arborescente do Rio

Grande do Sul, Brasil. Rima Editora e Novo Ambiente Editora, São Carlos, Brazil.

CLARIFICATIONS CONCERNING THE NOMENCLATURE

AND TAXONOMY OF OXYPOLIS TERNATA (APIACEAE)

Mary Ann E. Feist

Department of Plant Biology

University of Illinois at Urbana- La

Urbana, Illinois 61801, U.S.A

mfeistaillinois.edu

ABSTRACT

John R. Edmondson made ME na id S pda. inda n R. VEM aoa on the pace a Sium denticulatum

cialis was sy um mous wi Hel l history of

, however, that S. denticulatum Baldwi ] O. ternata (Nut À Hell y y In addition, the name

O. aa (Baldwin) J.R Edm. is an ison; f O. denticulata (Bald

Key Wonps: Apiaceae, isonym, Oxypolis denticulata, Oxypolis rigidior, Oxypolis ternata, Peuced Sium denticulat

RESUME

John R. Edmondson hizo | binación O is d lata (Baldwin) J.R. Edm. de acuerdo con la Nicol ae Sium ii

] de

ni era inónimo Jue O. ternata | Nun) A n i igación detallada de la ]

lacionados sin embargo o que 5 dentie ulatum Baldwin y P. ternata £5 (Sut) A. Heller no son sinónimos.

es el nombre O. denticulata Baldwin) ] LR. Edm. es un isónim

In 2005, J.R. da ne what X one to be a new ae Oxypolis denticulata (Baldwin)

J.R. Edm., t dentict m Baldwin, P. tum Nutt., and O. ternata (Nutt.)

A. Heller. He argued that the name O. denticulata dpereded O. ternata, the ommon cited modern name

for this species. Further investigation into the nomenclatural and taxonomic history of these and related

taxa has shown, however, that Edmondson made two important errors in proposing this combination: his

name is an isonym, and S. e a is not T ae P. a aig O. cla

11.11

In 1898, A

modern name for this species. Edmondson (2005) found an visotybe of Sium denticulatum in the bears

of the Linnean Society of London (LINN). He made the assumption that P. ternatum was synonymous with

S. denticulatum. Since Heller's nae was based on Ne (1818) epithet published after Baldwin’s

a Teh), Edmonson reasoned that t li ] riority and a new FORGDIDAUOR was warranted.

TI proposed the combination O. denticulat id

(LINN-Smith no: 508.5) labeled “Sium denticulatum nov. sp.” and celles by William Baldwin in 1817.

Edmondson went on to explain that when Rafinesque established the new genus Oxypolis (Rafinesque

1825), he included Sium denticulatum under Oxypolis but failed to make a new combination for the species.

Edmondson was correct that Rafinesque did not make the new combination in 1825, but he over] 1 that

Rafinesque did make EE CORA non five years later in the first volume of Bulletin oes (Bull. Bot.

l: 218. 1830) Rafinesque ] Was clearly | based on $ denticulatum, tl i y upon which

etbtatI1o

Edmondson based his combination. Consequently, the Edmondson isonym is without

and should be disregarded according to Article 6 Note 2 of the International Code of Botanical Nomenclature

aay et al. oe

cerrar errari i EA 1 L i f : : dhar TP C PUE ET

with Peucedanum ternatum, when in actuality, S. denticulatum is a synonym of Oxypolis rigidior (L.) Raf. The

recognition of this synonymy is not unprecedented for in 1840 Torrey and Gray recognized S. denticulatum

as a synonym ot S. rigidius L., the basionym for O. rigidior.

J. Bot. Res. Inst. Texas 3(2): 661 — 666. 2009

| fal Moa è Im LI PE L£ETrras 3/79)

662 Journal of t Texas 312)

TI tol for Sium denticulat nd P ] | | ide the first clue that tl species

are not os Baldwin (Elliott 1817) described S. alain as having “leaves unequally pinnate;

leaflets oval, toothed, acute; ... generally with three pairs of leaflets and an odd one.” In contrast, Nuttall

(1818) characterized P. ternatum as having: “leaves all ternate, upon very long common petioles; partial

leaves entire, long, linear, acute, and attenuated below... perfectly entire and scarcely three lines wide.

Comparing the lectotype of Sium denticulatum (Fig. 1) with the lectotype of S. rigidius (Fig. 2), and

comparing these with herbarium specimens and published descriptions of Oxypolis ternata and O. rigidior,

provide further clues and convincing evidence that S. denticulatum is a synonym of O. rigidior not O. ternata.

Differences are especially apparent in the leaf morphology (Figs. 13). These and other morphological differ-

ences between the species are summarized in Table 1. Information in Table 1 was compiled from a number

of sources (Weakley 2008; Radford & et al. 1968; Rodgers 1950; Mathias & Constance 1945; Torrey & Gray

1840), as well as from personal examination of numerous specimens from the following herbaria: DUKE,

F, ILL, ILLS, NCU, UGA, USF, and USCH.

Oxypolis ternata and O. rigidior can be distinguished by the following: The leaves of O. ternata are either

unifoliate or ternate and palmately disposed. The leaflets are petiolulate, parallel-veined, always entire, and

filiform to linear, typically being no more than 6 mm wide. The number of rays on the compound umbel is

5-10. In contrast, the leaves of O. rigidior are pinnately disposed with 5-13 sessile leaflets, reticulate-veined,

remotely dentate or entire (it is rare to find a plant without some leaflets that are dentate in the upper part

of the leaf), linear-lanceolate to elliptic-lanceolate, and 5-45 mm wide. The number of rays is 12-45.

No type has been located for Peucedanum ternatum. Inquiries or searches were made of all herbaria

listed by the Harvard University Herbarium Index of Botanists to have known Nuttall collections. Nuttall

gave North and South Carolina as the locality for P. ternatum. A specimen collected in South Carolina by

AE. Radford is herein designated as the neotype (Fig. 3).

Oxypolis ternata (Nutt.) A. Heller, Cat. N. 2mm ón 5. doen exire ternatum Nutt., Gen. N. Amer. Pl. 1:182. 1818.

Sataria linearis Raf., New Fl. 4:21. 1838 (nom illeg.) tt.) Nutt. in Torr. & A. Gray, Fl. N. Amer. 1:631. 1840.

O ternata (Nutt.) J.M. Coult. & Rose, Bot. Gaz. 12:74. 1887. Tret: U.S.A. Souta CAROLINA. Georgetown Co.: ditch, near

US 701, S-SW of Yauhannah, 20 Oct 1957, A.L. Radford 31381 (wEorYre, designated here: NCU!; DUPLICATES OF THE NEOTYPE:

UC!, UF!, Pp

1: x oe: ad len + waive a lararet

Sataria linearis Raf., var. longipes Raf., New Fl. 4:21. 1838. Rafi

Neurophyllum longifolium Torr. & A. Gray, Fl. N. Amer. 1:613. 1840. Types: U.S.A. Nort CAROLINA. [Cravern Co.]: “Swamps near New-

bern, North Carolina, Mr Croom! Dr. Loomis! Middle Florida, Mr. Croom! Sept.” (LECTOTYPE, designated here: NY!). Syntype: NORTH

CAROLINA: New Bern, s.d., H.B. Croom s.n. (PH!).

Oxypolis rigidior (L.) Raf., Bull. Bot. 1:218. 1830. Sium rigidius L. P Pl. 1:251. 1753. Oenanthe rigidius (L.) Crantz, Cl.

Umbell. Emend. 85. 1767, "rigide. Pastinaca rigidior (L.) Spreng. in Roemer & Schult Veg. 6:586. 1820, Ha clay

i oni (L.) DC., Prodr. 4:188. 1830, ‘rigida’. Peucedanum rigidius A.W. Wood, Amer. Bot FI 136 1870, nom. illeg.,

‘rigidum’. Tiedemannia rigidior (L.) J.M. Coult. & Rose, Bot. Gaz. 12:74. 1887, ‘rigida’. Tyre: U.S.A. VIRGINIA: J. Clayton 279 (econ

designated by J.L. Reveal in C.E. Jarvis et al., Taxon 55:215. 2006: BM, digital image!).

Archemora serrata Raf., Herb. Raf. 78. 1833. Tyre: U.S.A. Kentucky & TENNESSEE: specimen not located.

plaice ede Raf., Pp Raf. 78. 1833. Tyre: U.S.A. MISSOURI: Speebnen not located.

n. N. Amer. Pl. 1:189. 1818. P. } (Nutt.) Torr., Fl. N. Middle United States 315. 1824. Archemora

ambigua (Nutt. DC., Prodr. 4:188.1830. Archemora rigidior (L.) DC. var. ambigua A. Gray, Manual 158. 1848, ‘rigida’. Peucedanum

rigidius A.W. Wood var. ambiguum A.W. Wood, Amer. Bot. Fl. 136. 1870, ‘rigidum’. Tiedemannia rigidior (L.) J.M. Coult. & Rose var.

ambigua (Nutt.) J.M. Coult. & Rose, Rev. N. Amer. Umbell. 47. 1888, ‘rigida’. Oxypolis rigidior (L.) Raf., var. ambigua (Nutt.) B.L.

Rob., Rhodora 10:35. 1908. Tyre: U.S.A. PENNSYLVANIA. Philadelphia Co.: banks of the Delaware River near Philadelphia, s.d., T.

Nuttall s.n. (PH, digital image!).

Oxypolis turgida Small, Man. S.E. Fl. 986. 1933. Tyre: U.S.A. Vircinia. Staunton Co.: Staunton, 2 Oct 1895, WA. Murrill s.n. (LECTOTYPE,

designated here

Sium eek Baldwin i in S. oe Sketch Bot. S. Carolina 1:354. 1817. eiie angie (Baldwin) DC., Prodr. 4:188. 1830.

Bull. Bot. 1:218. 1830. Pasti Dietr., Syn. Pl. 2:971. 1840. Tyre: U.S.A.

com 1817, W Baldwin S.n. v— designated by Edmondson 2005: LINN- Smith, digital image!).

Sium longifolium Pursh, Fl. Amer. Sept. 194. 1813. Oxypolis rigidior (L.) Raf., var. longifolia (Pursh) Britton, Mem. Torrey Bot. Club 5:239.

Feist, T.

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TABLE 1. M

lis rigidior (L.) Raf.

Characters

Oxypolis ternata

Oxypolis rigidior

Petiole length

Leaf disposition

Leaflet number

Leaflet veination

10-30(-35) cm

palmate

(1)3

parallel

ntire

5-10 cm

pinnate

(5-)7-11(-13)

iculate

(entire) remotely dentate

Leaflet shape linear to filiform linear-lanceolate to elliptic-lanceolate

Leaflet length 5cm

Leaflet width 1-6 mm 5-45 mm

Leaflets petiolulate sessile

Ray number -10 12-45

1894, ‘rigidus var. longifolius’. Oxypolis longifolia (Pursh) Small, Fl. S.E. U.S. 875, 1336. 1903. Oxypolis rigidior (L.) Raf. subsp. longifolia

(Pursh) W. Stone, Pl. S. New Jersey 2:600. 1911. Tyre: U.S.A. New JERSEY: (Lectotvre, designated with reservations by Ewan 1979:

PH, digital image!).

Sium irn Elliott, Sketch Bot. S. Carolina 1:354. 1817. Arc! ta (Elliott) DC., Prodr. 4:188. 1830. Oxypolis tricuspi-

a (Elliott) Raf., Bull. Bot. 1:218. 1830 lata (Elliott) D Dietr.. Syn pl 2:971. 1840. Tyre: U.S.A. SOUTH CAROLINA:

oe CHARL)).

ACKNOWLEDGMENTS

I would like to thank Emilie Bess, J. Leland Crane, Stephen R. Downie, Geoffrey A. Levin, Anthony R.

Magee, Brenda Molano-Flores, and James R. Reveal for helpful discussions or comments relating to this

manuscript.

REFERENCES

EDMONDSON, J.R. 2005. A new combination in Oxypolis Raf. (Apiaceae). Novon 15:109.

E.uotr, S. 1817. A sketch of the botany of South Carolina and Georgia. 2 vols. J.R. Schenck, Charleston, South

Carolina.

Ewan, J. 1979. Introduction to the facsimile reprint of Frederick Pursh, Flora Americae Septentrionalis (1814). In:

FT. Pursh, Flora Americae Septentrionalis. Facsimile reprint. J. Cramer, Braunschweig, Germany.

Mathias M. AND L. Constance. 1945. Umbelliferae. N. Amer. Fl. 28B:43-295

McNett J., ER. Barrie, H.M. Bunper, V. Demoutin, D.L. HawksworTH, K. MARHOLD, D.H. NicoLson, J. PRADO, P.C. Sit va, J.E. SKOG,

J.H. Wiersema, AND NJ. TURLAND. 2006. International code of botanical nomenclature (Vienna Code). Regnum

Veg. 146.

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

Heartt, Philadelphia, Pennsylvania.

RADFORD, A.E., H.E. Aes, AND C.R. Bett. 1968. Manual of the vascular flora of the Carolinas. University of North

Carolina Press, Chapel Hill, North Carolina.

Rarinesque, C.S. 1825. Neogenyton, or indication of sixty-six new genera of plants of North America. Published

by the author, Lexington, Kentucky.

Rarinesque, C.S. 1830. Extract d'une lettre de M.S.C. Rafinesque, professeur à Philadelphie, addressée a M. le pro-

fesseur De Candolle. (Mai 1830). Bull. Bot. 1:214—221.

Ropeers C.L. 1950. The Umbelliferae of North Carolina and their distribution in the southeast. J. Elisha Mitchell

Sci. Soc. 66:195-266.

Torrey, J. AND A. Gray. 1840. A flora of North America. Wiley & Putnam, iid vale

WEakLEY, A.S. 2008. Flora of the Carolinas, Virginia, and Georgia, and

Herbarium, North Carolina Botanical Garden, University of North Cola, at Chapel Hill.

University of North Carolina

NEOTYPIFICATION OF ARTEMISIA CARRUTHII (ASTERACEAE: ANTHEMIDEAE)

— A. Morse James L. Reveal

McGregor Herbarium L.H. Bailey Hortorium Herbarium

Division E "en Natural History Museum Denar rtment of Plant Biology

and Biodiversity Research Center Cornell University

University of Kansas 412 Mann Library Bidg

2045 Constant Ave Ithaca, New York 14853-4301, U.S.A.

Lawrence, Kansas 66047-3729, U.S.A. jir326@cornelledu

cmorse@ku.edu

ABSTRACT

1 r 1 1 f. 11: el fal A 1; 1 : 1 k]

Authentic material of Artemisia carruthii Alph. W

Pa of names now cone LARA ie ue that p A. wright A. mar A. kansana Britton, A olovadensis Osterh.,

] with t A

”

roloradensis and A haberi and a “step 3”

ri Greene, and A pringlei Gree

ne eue for A. a

RESUMEN

El material cient de ln carruthii ee Wood ex Carruth se perdió después de la publicación del bre. Por ell ]

eotipo y id d peci A Me Gray, A. kansana Britton, A. colo-

radensis Osterh., A. bakeri Greene, y A. unde Greenm. Se revisan con “paso 2” lectotip g A. coloradensis y A. baheri y

un "paso 3" se designa lectotipo para A. wrightii.

James Harrison Carruth (1807-1896) published three new species attributed to Alphonso Wood in his

"Centennial catalog of the plants of Kansas" (1877). The disposition of one of these, Cyperus spiculatus Alph.

Wood ex Carruth (Cyperaceae), likely a later synonym of C. erythrorhizos Muhl., has been discussed by

McGregor and Brooks (1982). It is not clear that Carruth actually intended to publish the second species,

Muhlenbergia innominata Alph. Wood in Carruth (Poaceae) as new to science, rather than merely indicating

that Wood had identified it ] ibed. Although it | ts for publication, the descrip-

tion was reproduced almost verbatim from Carruth (1873), where de taxon was first described without an

epithet. The status of this name was not addressed by McGregor and Brooks (1982), and the name has been

overlooked by other authors. Original material has not been located. The third species, Artemisia carruthii

Alph. Wood ex Carruth, was published with this statement as its apis

481. tA. Carruthii sic vult Mr. Wood, nov. sp. Erect, simple, clothed all ] I tely parted, with linear

segments, the lower Nui i with al ie a all close- P 1' [inch] and less long. Heads E small, 1" Ll 7-lined

flowered, homogamous, 11 le. Plant 1-2 feet

Carruth did not cite type material or "i of collection, and the species is not figured in the publication.

Here, we discuss the source and fate of the original material, which appears to have been lost around the

time of publication. To fix application of the name, we designate a neotype.

In an 1892 letter to Bernard B. Smyth (1843-1913), Carruth suggests that original material was in

existence at the time he published these names, reporting that he had sent "several specimens to [Wood]”,

and that names published in his 1877 list were based on th i Carruth 1892). That Carruth took

pains to indicate that the description of Artemisia carruthii was stis own ‘and not merely transcribed from

correspondences from Wood, as he did in the case of Muhlenbergia innominata, suggests that a least one set

of specimens was in his possession at or near the time of publication. However, original material was almost

certainly not collected by Carruth himself. Carruth (1877) noted that the he had “not traveled widely, but

[was] greatly aided by the observations of others.” Carruth was stationed in Lawrence, Kansas, at the time

J. Bot. Res. Inst. Texas 3(2): 667 — 672. 2009

668 | | tanical h Institute of Texas 3(2)

of the publication of his Centennial Catalogue, and it does not appear that he made any excursions to the

western half of the state prior to 1881 (Carruth 1879, 1881), when he reported having travelled as far west

as Barton County (Carruth 1881-1882). As a consequence, many of the taxa included in his floristic lists

of 1872, 1873, 1874, and 1877 were based on nb M contributed by a “corps of observers” (Carruth

1873). Carruth i I 1 into his li i ided by these observers, but he also received

hundreds of specimens for determinati pr it is likely that the first specimen of A. carruthii was taken by

one of several collectors who lived in or traveled through western Kansas in the 1870s.

Carruth frequently failed to cite collectors of voucher specimens by name, but did include a station

for most specimens, from which it is often possible to deduce the donor. Unfortunately, the usual cita-

tion of a place of collection is omitted for Artemisia carruthii. Of the individuals who had collected in the

western part of the state prior to its publication, Carruth reported receiving specimens from areas within

the range of A. carruthii from three individuals: the paleontologist Benjamin Franklin Mudge (1817-1879)

is credited with contributing five specimens from “S. W. Kansas" and “W. Kansas" (Carruth 1872, 1877);

Louis Watson (1817-1894), a medical doctor and cousin of Sereno Watson (1826-1892), is presumed to

have taken most if not all of the specimens Carruth recorded from Ellis, Ellis County (Carruth 1873, 1874,

1877) and contributed *400 to 500 species, often many of each" (Carruth 1877); and Francis Huntington

Snow (1840-1908), Professor of Natural History (and eventually Chancellor) at the University of Kansas,

and his students were credited with specimens and eee from Russell County among other stations

(Carruth 1874, 1877). A single specimen from Ellis att ed to a collector cited as “UHer” (Carruth 1877)

is presumed here to be a typographical error. ME. Edwin Alonzo Popenoe (1853-1913), professor

at Kansas State Agricultural College, who traveled throughout the western part of the state and on whose

authority Carruth (1877) credits stations in fourteen counties, seems to have contributed only reports of

species, but no specimens (Carruth 1873, 1874, 1877).

Carruth's own herbarium was reputed to be poorly maintained and disposed of when he departed

Lawrence in 1892 (McGregor & Brooks 1982), and any material in existence is likely to have been discarded

at that time. However, it is possible that he had deposited some specimens in one of several botanical in-

stitutions in the state. In addition to serving as state botanist, Carruth held appointments as Professor of

Natural History at Baker University in Baldwin City and as a lecturer at Washburn University in Topeka

(McGregor & Brooks 1982), and he may have deposited some vouchers in the botanical collections of either

institution. In addition, a small number of specimens collected by Carruth, Snow, and Watson remains at

KANU, and a few collected by Snow and Watson remain at ksc; several specimens gathered by Watson are

also at cH. Alphonso Wood’s herbarium and types were deposited at ny sometime after his death in 1879

(Stafleu 1967), although material from his herbarium has been identified in other institutions.

Whatever the fate of Carruth's holdings of Artemisia carruthii, it appears that the material was lost to

students of the Kansas flora soon after publication. The name seems to have fallen into obscurity almost im-

mediately after Carruth's departure from Kansas. It was overlooked by authors of Sues quen compilations

of, and novelties to, the flora of the state. Artemisia was not included in the t of Kellerman

and Kellerman (1888), Smyth (1889, 1892b), or Hitchcock (1899), or listed among the 145 species excluded

by Smyth (1890). Smyth (1892a, 1892b) cited A. wrightii A. Gray for several K s counties and later (1898)

suggested that A. kansana a d be a form of A. frigida," which he had previously cited (Smyth 1890,

1892a, 1892b) for stations out nge of that species in Kansas. Indeed, no early specimens deposited

in KANU, KSC, OT WASH were iaa B the collectors to be A. carruthii, these being variously named A.

filifolia Torr., A. frigida Willd., A. kansana, or A. wrightii as late as 1931. That Smyth, who corresponded rather

extensively with Carruth, did not even discuss A. carruthii suggests that he had not seen original material

in Kansas herbaria, particularly xsc and wasn, which he was likely to have examined during the preparation

of his floristic lists.

Artemisia carruthii was first resurrected for the Missouri flora by Mackenzie (1902), who listed A. kan-

sana as a synonym without comment. In the ensuing debate about the correct name for this taxon, no type

thii 669

material was cited—most likely because it was not seen—by Rydberg (1910), Britton and Brown (1913), Hall

and Clements (1923), Bush (1928), Gates (1939), or Keck (1946). Notable among these authors were Britton

and Brown, whose listing of A. carruthii as only a questionable synonym of A. kansana suggests that Britton

had not seen material at ny referable to A. carruthii at the time of publication. Hall and Clements discuss A.

carruthii at length, but—unusual for these authors—also do not cite a type specimen. Acknowledging that

A. carruthii was the earliest available name for the element they recognized as a subspecies of A. vulgaris L.,

Hall and Clements (1923) nevertheless chose to employ Asa Gray’s epithet in publishing the combination A.

vulgaris var. wrightii, citing the earlier name as “not well established” (p 81), commenting (p. 82) that Gray

“did not intend to establish a species distinct from A. carruthii, the earlier publication of which he appar-

ently overlooked.” Results of biosystematic studies by Ray (1971) and Bol dd supported Keck's (1946)

circumscription of A. carruthii, and most authors of recent floras and checklists have accepte name (e.g.

Martin & Hutchins 1981, Barkley 1991, Cronquist 1994, Jones et al. 1997, Welsh 2003, Kaul et al. 2006,

Voss 1996, Yatskievych 2006, Shultz 2006). However, none of these authors have resolved typification of

the species. It was not accepted by Ling (1995), who treated A. wrightii as having priority, apparently based

on an error aes Mie pees date of the latter name.

y distributed from the soutl t Great Plains though the eastern part of the

Great Basin, New Mexico, Arizona, and northern Mexico (Shultz 2006), and it is reported to be adventive

in Michigan (Voss 1996) and Missouri (Yatskievych 2006). Welsh (2003) and Shultz (2006) reported that

A. carruthii intergrades with A. ludoviciana Nutt., so it is important to fix application of the name by select-

ing a type. The senior author attempted in October 2007 and January 2008 to locate original material of

A. carruthii for designation as a lectotype. Requests to identify specimens collected in Kansas in or before

1880 were sent to staff at Baker University, FHKSC, ISC, KSTC, GH, NY, PH, and Yu. Holdings at KANU, ksc, and

WASH were examined by the senior author; holdings at Mo were examined on the authors’ behalf by Craig

Freeman (Curator, KANU). These searches proved unsuccessful, and it seems doubtful that original material

remains. eee a none is designated below in accordance with Art. 9.15 of ICBN (McNeil et al.

2006). F g ion first proposed by Keck (1946) and accepted by subsequent

authors, we also di the typification of idered ymous of A. carruthii, designating "step 2"

lectotypes for A. coloradensis Osterh. and A. bakeri Greene, aud a "step 3" lectotype for A. wrightii (discussed

by McNeil et al. [2006] in Art. 9, Ex. 8, and by Anderson [2007]).

Artemisia carruthii Alph. Wood ex Carruth, Trans. Kansas Acad. Sci. 5:51. 1877... Artemisia vulgaris L. var. carruthii

(AI ha EC. ES Trans. Kansas inn m a 138. 1939. T S. Kansas. Hamilton Co.: 3.25 mi

& Wildlife Area, T23S R41W sec 28 SE of NE, 38.02517°N,

YY LLC Filio,

acuse

101. ens elev. 3340 ft, disturbed honras a on eroded nid slopes aan Plum Creek below dam, 9 Oct 2007, C.A.

TYPE

Y, COLO, CS, FHKSC, ISC, K, KANU, KSC,

Morse 16097 (NEOTYPE: KANU sheet 361746, designat

KSTC, GH, MEXU, MICH, MO, ND, NEB, NY, OKL, OKLA, PH, RM, TEX, WASH, WIS, US, UTC, a

Artemisia wrightii A. Gray, Proc. Amer, Acad. Arts 19:48. 1883. Artemisia Boh L; Pe a (A. nek H.M. Hall & Clem., Publ.

Ew oE Inst. jin) is 326:80, pl 9. 1923. Artemisia vulgaris var. wrightii (A. Gray) E.J. P Ann. Missouri Bot

935. Artemisia carruthii var. wrightii (A. Gray) S.E ate i Vigas ne a 30: 472. 1940. n a

Gard. 22:6

STATES. New Mexico. Grant Co.: Santa Rita - Cobre, 1851, C. cian

a [1910: a) as “type, ” here corrected; NY, I tion designated by Hall d Clemens (1923: ae as

e” h 8580, a third step lectotyy ificati deime ] [th heet

RE by Asa Gray v ds NY, US).

Art Brit: NI Britton & A. Brown, Ill. Fl. N. U.S. 3:466. 1898. Tyee: U STATES. kansas. Lane Co.: plains, 15 Aug

1895, A.S. Hitchcock 302 € GH, KSC, RM, US)

Artemisia coloradensis Osterh, Bull. es Bot. Club 27:506. 1900. Artemisia e var. EM (Osterh.) A. Nelson, New Man

Bot. Centr. PUE Mts. 568. 1909. Tyee: ae STATES. COLORADO. Larimer Co.: Da eek, 7 Sep 1899, G.E. Osterhout 2010

(LECTOTYPE I yl g y Keck [1946: 440] as “type,” here iie RM sheet 170628, a second step

ification designated | , RM [2 sheets])

Artemisia bie Greene, PL Baker 3: 31. 1901. Artemisia mexicana Willd. ex Spreng. var. id rc A. Nelson in New Man. Bot.

Centr. Rocky Mts. 569. 1909. Tyre: oa e CoLorapo. Montrose Co

1896, E.L. Greene s.n. (LECTOTYPE, a fi typi g y Hall and Cl ts [1923: 81] as “type,” here corrected;

|| ELL Da4 H ip LJ ata "1 LT le Vds A!

Vo,

ND-G sheet 062066, ] l ification designated here; DUPLICATE OF THE LECTOTYPE: -G).

Artemisia pringlei Greenm., Proc. Aster. Acad. 40: 50. 1904. Tire: MEXICO. CHIHUAHUA: plains near a 11 Sep 1885, C.L. Pringle

625 (LECTOTYPE: GH designated by Hall and Clements [1923: 81] as “type,” here corrected; DUPLICATES OF THE LECTOTYPE: Ds, NY [2

sheets], PH, US).

e (1910) merely restricted application of Artemisia wrightii to the Wright collection, whereas Hall and

Clements (1923) designated one of two specimens at NY as the ‘type.” A more logical choice would have been

one of the three sheets annotated by Gray at GH. We hereby restrict the choice to a single sheet at ny.

Keck annotated a duplicate of Osterhout 2010 (171095) as the “type” perhaps because the original Oster-

hout specimen (170628), marked with a small, red, preprinted label with the word “Type” was not seen by

Keck in 1939. Because there were three sheets of Osterhout 2010 at RM prior to 1946, however, we consider

Keck's actions a first step lectotypification as he did not distinguish between the three sheets. It should be

noted that Hall and Clements (1923) did not cite collection information of Artemisia coloradensis, other than

to remark (p. 82) “Type locality, near Dale Creek, Larimer County, Colorado." This by itself does not qualify

as an effective lectotypification.

Hall and Clements (1923: 81) stated that the type of Artemisia bakeri “was collected by Greene in the

canyon of the Gunnison, near Cimmaron, Colorado. This has not been seen [by us], but the above notes are

based upon other specimens cited by Greene, namely, Black Canon, Gunnison Watershed, Colorado, Baker

698 (Gr, UC, US) As they did not indicate where the Greene specimen was to be sought, our second step

lectotypification completes their designation of the lectotype.

ACKNOWLEDGMENTS

We thank James Estes, Craig Freeman, and Leila M. Shultz for reviewing the manuscript and providing

helpful comments. Carolyn Beans (cH), Roger Boyd (Baker University), Tom Eddy (kstc), Alina Freire-Fierro

(pH), Barbara Hellenthal (wp), Jill Larson (rm), Deborah Lewis Qsc), Mark Mayfield (xsc), Melissa Rossow (ny),

Patrick Sweeney (Yu), Joe Thomasson (FHksc), and Mark Allen Wetter (wis) supplied data on the holdings at

their respective institutions. Craig Freeman (KANU) examined holdings at mo. In addition, John Boggan (us),

cae pou MS Mayela, ane Melissa Rossow kindly provided images of type specimens not already

website. Lisa DeCesare (Botany Librairies, Harvard Eve) and Marie

Long (Mertz Library New York Botanical Garden) hed their holdings fo Car-

ruth and Wood. Ronald Hartman (rm) provided insights into the annotation isses of Osterhout's types.

Vic Landrum provided both information about and access to the collection at wasH. Curators OÍ KSC, ND,

RM, and wasH kindly provided specimen loans. The senior author's heldwork was made possible through an

anonymous contribution to the McGregor Herbarium endowment.

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A PHYLOGENY OF ARCTOSTAPHYLOS (ERICACEAE) INFERRED

FROM NUCLEAR RIBOSOMAL ITS SEQUENCES

ey A. Sene V. Thomas Parker

lant Biology Department of Biology

Ohio University San Francisco State University

irs Ohio 45701, U.S.A. San Francisco, California 94132, U.S.A.

7105gohio.edu parker@sfsu.edu

Michael C. Vasey

Department of Environmental Studies

University of California, Santa Cruz

Santa Cruz, California 95064, U.S.A.

ana ucedi

ABSTRACT

SA " " Aaa FT us ^ M ] A 1 +1 lTTc T F (0 : Lr j HE

y phylos ( ; g I I 3 were unique

Leese] 1 f j e 11 iJ] | 1 1 1 11 LM SN | L FA

taxa ii I y I ie g y I uva-ursi, A patula, A. pungens, A

manzanita, A. Po: m mentos Forty-four ingrour hared an identical seq The remaining species showed pairwise

1 ] jl ] 1.2 Lien AA E e 1 d ] 1

SUBEHURBORE rates of 2.4% or less, and 17 ] phic f j? y y g

Tad Á ] e] d dat t thr 17 ] } 1 | T clade tanninow

poly bcc i VY BLL Lili. Lf F Pi F NA la "VTEV-MPB

ld; 1 | PEERS “Le: L nalada y : 1 j T f 1 |] J Jipii . -r

X 3 4 oo

la origin. A proposed hypothesis js ET rates of nucleotide substitution in species that are Pit t ders vs. those that are

I 1 shared sequences. Low interspecifi variabili I ] LITS

region in Arctostaphyl v be due to severa l factors, i g diversification of the genus in Califonia bus Eran time

cx 1 Jan Ez eee tard ITI f ] il m arrays in both

Iu D L i I

Key Wonps: Arctostaphylos, generation time, i , molecular phylogeny, ] il LITS, polyploidy, super-

imposed elsa a patterns.

RESUMEN

Una filogenia de Arctostaphylos (Ericaceae) se Senado utilizando secuencias ES ADN Dun as ITS de 98 especies de las

L m4

-l ioc D zf- [y i Í Fa FA uvri- -ursi,

a. Le Á. PUNES, A. manzanita, A. glandulosa, udin i. tomentosa). C Cuarenta y cuatro t part ia idéntica. Las

mE 177 1 ] A wl. TI ] J i

I ,yen 17 I

A e AA al L 1; d P| 1 t Ve cau et ] de 1 z A] 22] ] L

L o I 3 e 17 r 2 1 qa

+ LS | Cera | = E 1 1 1 T 1 1 P ak 1 » J yeas LE J ] ez: ^al J

un origen hibrido Heee en La similitud de las secuencias de iudi tbe se es estrictamente por semillas vs las

que se aon por oe y: a la I p yor tasa de sustitución nucleotídica en

bs ned baj pecífica que muestra el ITS en Arctosta] hyl puede del varios factores, como la reciente

5 Td fep d 11 a ] + 2 " 1 4 J J sm 1 1 1 aed url

[us] a i" I] i be

AA ] 1 z 1 i E | t ] PE E "n 1 aa 1 a E PA | 4 a esc |

rápida del ITS en las por I I p y tetraploides.

INTRODUCTION

Arctostaphylos Adans. (Ericaceae) is composed of ca. 108 taxa and is the most species-rich genus of subfamily

Arbutoideae Nied. (Wells 2000). Except for Arctostaphylos uva-ursi (L.) Spreng.—which has a circumboreal

distribution—the remaining species are confined to western North America and Mexico. In California,

species of Arctostaphylos occur in a variety of plant communities, including chaparral, montane scrub,

mixed-coniferous forests, serpentine barrens, and oak savannas. At low latitudes, Arctostaphylos uva-ursi

occurs in montane regions (e.g., Rockies, Appalachians, Alps, Himalayas) and in coastal pine barrens and

l Rat Bac Inet Tavas 3(2): T? - 682. 2009

674 Jo tl tanical Insti Texas 3(2)

dunes (e.g., New Jersey and Michigan, USA). A. uva-ursi subsp. cratericola (Donn.Sm.) PV.Wells represents

a disjunct population in the volcanic highlands of Guatemala.

Arctostaphylos is placed in subfamily Arbutoideae with five other genera: Arctous Nied., Arbutus L.,

Comarostaphylis Zucc., Ornithostaphylos Small, and Xylococcus Nutt. (Diggs & Breckon 1981). Molecular

and morphological phylogenetic studies in the Ericaceae place the Arbutoideae between the basalmost

Monotropoideae and the rest of the family (Kron et al. 2002). Markos et al. (1998) presented phylogenetic

trees for 16 species of Arctostaphylos inferred from nuclear ribosomal internally transcribed spacer (orl TS)

and 28S nuclear ribosomal sequences. The two-clade topology recovered in their trees was not congruent

with the two subgenera of Wells (2000) nor did it support the monophyly of the A. hookeri G.Don species

complex in section Pungens (Markos et al. 1998). Boykin et al. (2005) presented a nrITS phylogeny of 38 taxa

that also recovered a two-clade topology. Further, these two-clade phylogenies are congruent (with respect

to species composition) with a phenogram constructed from flavonoid data for 20 species of Arctostaphylos

(Denford 1981).

The center of diversity for Arctostaphylos is centered about San Francisco, CA in the California Floris-

tic Province. The coastal areas and Coast Ranges of California, from Mendocino County to Santa Barbara

County, are particularly rich in taxa (Markos et al. 1998). Arctostaphylos has a high percentage of endemic

taxa, many of which are edaphic specialists (Wells 2000). Nearly half of all California taxa are considered

rare, threatened, or endangered (CNPS 2009). The diversity of modern taxa may be the result of a radiation

of Arctostaphylos since the end of the Miocene (5.3 Ma) (Edwards 2004). Factors contributing to the diversi-

fication of the genus include local adaptation to California's diverse substrates, an increase in fire frequency

at the end of the Pliocene, the onset of a Mediterranean climate, and the contemporaneous expansion of

chaparral into forested areas (Raven & Axelrod 1978; Axelrod 1989; Edwards 2004).

neon ane peyote ses are two processes driving speciation in Arctostaphylos. The base

] | 13, with most species being either diploid n = 26) or tetraploid

(2n = 52). Schierenbeck et al. (1992) provide morphological and cytological evidence for the allopolyploid

origin of A. mewukka Merriam. Many other taxa have been proposed ising via homoploid hybrid specia-

tion (e.g., Dobzhansky 1953; Gottlieb 1968; Schmid et al. 1968; Kruckeberg 1977, Knight 1985; Keeley &

Massihi 1994; Parker & Vasey 2004). For a group of 46 edaphic specialists or endemic taxa, Wells (2000)

proposed a hybrid origin for 37. Differing ploidy levels observed in Arctostaphylos uva-ursi subsp. coactilis

Fernald & J.F.Macbr. (2n = 26, 39, 52 and 78) suggests recent and recurring polyploidization, which is

common in arctic paa a 1981; epu et al à 4).

f hybrid origin (e.g., Rieseberg 1995)— par-

ticularly i in Arctostaphylos. 1t i is is likely d aon introgression and d polyploidization in AreOStAp IOS has lead to

re based on

In perentual species of Paeonia L. and Sidalcea A.Gray, the escenas inherited, niTS -— has been

used successfully to corroborate parentage of allopolyploid taxa (Sang et al. 1995; Whittall et al. 2000). In

angiosperms, direct sequencing of the maternally inherited chloroplast may provide additional evidence

of hybrid origin by identifying the maternal parent (Koontz et al. 2004). In Arctostaphylos, Wahlert (2005)

sequenced five chloroplast regions (trnL intron and the intergenic spacers atpB-rbcL, trnL-trnF, trnS-trnG

and trnH-psbA) for 12 morphologically divergent and geographically distant species. The resulting combined

alignment was al ne pa pairs (bp) ong P had just three parsimony dal positions: Thus, direct

ld not be used to inf hips or to confirm maternal

MM in hybrids species.

Different life history st ies in Arct I to inf the rate of speciation. In California’s

aid prar chaparral. ales Arctostaphylos a añ Ceanothus Es (Rhamnaceao) are the only genera of

olved an oblig ling mode of reproduction (Wells

1969). in these genera, in about M donee of the species, dult killed in to fi ] populations

recover by recruitment from seed banks. Most chaparral shrubs re-sprout after à stand: replacing fire, but

Wahlert et al., ITS phyl g y JA a "1 pl [| 675

in species of Arctostaphylos and Ceanothus, seeds are stored in a persistent soil seed bank Hom which ey

germinate only after fire (Parker and Kelly ie. Therefore, each Best fire gen eratio

product of sexual reproduction, wl ily from vegetative re-sprouts

(but also to a lesser degree from seed) (Keeley & Keeley 1981). A Heber rate af morphological diversifica-

tion and local endemism for seeders is supported by the fact that ca. 70% of Arctostaphylos taxa are obligate

seeders (Wells 1969). Generation-time theory predicts that taxa with shorter generation times will have

higher rates of molecular evolution in neutral DNA regions compared to taxa with longer generation times

le 8 Andreasen & Baldwin 2001). For that reason, it is de eee that in ita cae ne will be a

gher rate of interspecific ITS sequence variability d to

The primary goal of this study was to estimate the phylogeny of f Arctostaphylos with "— taxon

ecies composition of each clade,

RATAA [eae

sampling using nrITS sequences. Increased taxon s

might reveal additional subclades, and could allow for more rigorous testing of the monophyly of certain

infrageneric taxa. In addition, we sought to corroborate hybrid origin in putative hybrid species and to test a

hypothesis of higher rates of nucleotide substitution in obligate seeders compared to facultative sprouters.

MATERIALS AND METHODS

Taxon Sampling

The ingroup included a total B iia accessions representing 83 unique taxa. The remaining 15 accessions

were multiple samples of mor l/or widely distributed species: Arctostaphylos uva-ursi,

A. patula Greene, A. pungens HBK, A. manzanita Parry, A. glandulosa Eastw., and A. tomentosa (Pursh) Lindl.

The outgroup contained six species representing the other five genera in the Arbutoideae. All sequences

generated for this study were new, except for two accessions of Arctostaphylos and five outgroup taxa, which

were downloaded from GenBank (Appendix 1). Taxonomy follows Wells (2000) with names, geographic

origin, voucher specimen data, and GenBank accession numbers given in Appendix 1. All accessions se-

quenced for this study are represented by voucher specimens deposited at the Harry D. Thiers Herbarium

at San Francisco State University (SFSU), except for Arctous erythrocarpa, which is deposited at MO.

DNA Extraction, PCR Amplification, and Sequencing

Genomic DNA was extracted from herbarium material using DNeasy™ Plant Mini Kits (QIAGEN Corp.,

Alameda, CA, USA). The nrITS region was amplified using Arctostaphylos-specific primers designed for this

study (5’ to 3): ITS 4-M = GGAAGTAAAAGTCGTAACAAGG and ITS 5-M = CTGGGGTCGCAATCAAGG

PCR amplifications for the ITS region were carried out under the following conditions: an initial denatur-

ation step (92°C, 2 min) followed by 35 cycles of denaturation (92°C, 30 sec), annealing (58°C, 30 sec), and

elongation (72°C, 2 min), with a final extension step (72°C, 5 min). PCR amplifications were performed in

25 pl reactions containing 10-100 ng genomic DNA and 0.25 pl Expand Taq High Fidelity PCR System™

with buffer and Mg? (Roche Diagnostics, Indianapolis, IN, USA). PCR products were visualized on an

agarose gel and purified using MO BIO UltraClean™ PCR Clean-up Kits (MO BIO Laboratories, Carlsbad,

CA, USA). Cycle-sequencing reactions used the same primers, annealing temperature, and additive as for

PCR and employed BigDye? fluorescent dye-labeled chemistry (BigDye Terminator Cycle Sequencing Kit,

Applied Biosystems, Foster City, CA, USA). Cycle-sequenced products were separated and visualized on an

ABI 377 DNA sequencer (Applied Biosystems, Foster City, CA, USA). When a polymorphic nucleotide posi-

tion was detected, the taxon was sequenced a second time to confirm the polymorphism. DNA sequencing

was performed at the San Francisco State University Conservation Genetics Laboratory.

Phylogenetic Analyses

Due to low rates of nucleotide substitution and few insertions/deletions, ITS sequences were easily aligned

automatically using Sequencher version 4.1 (Gene Codes, Ann Arbor, MI, USA). Maximum parsimony

(MP) analysis was conducted on two data sets: 1) the total data set for all 98 Arctostaphylos sequences, and

2) a reduced data set with 17 taxa removed that had polymorphic nucleotide positions. Forty-four ingroup

676 Journal of ti tanical h Insti Texas 3(2)

taxa, which shared identical ITS sequences, were treated as a single “taxon” (the “Gray Leaf” haplotype),

therefore the actual number of taxa in both MP analyses was 55 and 38, respectively. MP analyses used

the heuristic search strategy of Olmstead and Palmer (1994) to search the tree space for multiple islands of

shortest trees. This search strategy, as implemented in PAUP* ver. 4.0b10 (Swofford 2002), was carried out

in four steps, with the saved trees from each step used as starting trees in subsequent searches: 1) 1,000

random addition sequence replicates, nearest neighbor interchange (NND, MulTrees Off, 2) tree-bisection

reconnection (TBR), MulTrees Off, 3) NNI, MulTrees On, and 4) TBR, MulTrees On. A strict consensus tree

was calculated from the shortest trees saved from each search. Internal branch support of phylogenetic trees

was estimated with 100 bootstrap (BS) replicates (Felsenstein 1985) using a full heuristic search with TBR

branch swapping, MulTrees on, and 10 random stepwise addition replicates. Results from the BS analysis

were summarized in a strict consensus tree. Nucleotide characters were equally weighted and unordered,

and indels were treated as missing data.

RESULTS

The aligned sequence matrix for the 98 ingroup accessions and 6 outgroup taxa was 600 bp in length (0%

missing data), with 127 variable positions and 62 parsimony informative characters. A one bp deletion at

position 403 was detected for Arctostaphylos malloryi (W.Knight & Gankin) PVWells, A. nissenana Merriam,

and three subspecies of A. viscida Parry. Forty-four taxa shared an identical nrITS sequence—the “Gray

Leaf” haplotype—and are listed in Appendix 2 (the phrase “Gray Leaf haplotype” is used throughout this

study to refer to this shared sequence and does not imply a uniform morphology among taxa sharing this

identical sequence). The uncorrected pairwise distances for the ingroup nrlTS sequences ranged from 0 to

2.4%. Parsimony analysis of the complete ITS data set reached a maximum of 100,000 equally parsimoni-

ous trees of length 178 before ending initial tree generation, with a consistency index (CD of 0.837 (0.753

excluding uninformative characters) and retention index (RD of 0.881. A strict consensus tree (not shown)

revealed a large polytomy and three small clades: 1) a clade of all six accessions of North American A, uva-

ursi (58% BS), 2) a clade containing A. pumila Nutt. and A. tomentosa (1) (59% BS), and 3) a clade containing

A. ells a A. pane M. E itd & "d iw M uS

+1 1 1 f. A

patterns iat nucisende addi ity (SNAPs) and were enfe in 17 taxa (18% E ingroup): A. ravens A.

pringlei ssp. drupacea Parry (B.VWells), A. mewukka, A. pilosula (Jeps. & Wiesl.), A. viscida ssp. viscida, A.

manzanita ssp. manzanita from Humboldt Co., CA, A. manzanita ssp. roofti (Gankin) PV.Wells, A. manzanita

ssp. laevigata (Eastw.) Munz, A. manzanita ssp. bowermanii, A. pungens from San Diego Co., CA, A. X pacifica

Roof, A. nissenana, A. edmundsii J.T. Howell, A. glandulosa ssp. glandulosa from San Diego Co., CA, A. myrti-

folia, A. nummularia (1) and A. hispidula Howell. The number of polymorphic nucleotide positions within

an individual ranged from 1 to 6 (0.7% to 4.3% of variable positions). Of these 17 taxa, 12 are tetraploids

and/or hybrids. In four putative hybrid-derived or introgressed taxa (A. pringlei ssp. drupacea, A. manzanita

ssp. roofii, A. manzanita ssp. laevigata, and A. hispidula) sequence electropherograms showed near-perfect

patterns of additivity between synapomorphic nucleotides of suspected pens ane » aisi (Table 1).

A SE ORAN M and yes was conducted with the 17 taxa with p

TI trix had 126 variable positions Bod 56 parsimony informative characters.

MP analysis if the Y ITS data set recovered a two-clade topology in the strict consensus of 350 equally

parsimonious trees (Fig. 1) (tree length = 169, CI = 0.817, CI excluding uninformative characters = 0.677,

RI = 0.858). Clade 1 was weakly supported (55% BS) and Clade 2 was moderately well supported (77%

BS). Within Clade 1, two subclades were recovered: 1) a clade containing the six accessions of A. uva-ursi

(59% BS) and 2) a clade containing three subspecies of A. stanfordiana Parry, A. nummularia A.Gray, and A.

mendocinoensis PV.Wells (76% BS). In Clade 2, the Gray Leaf “taxon” representing the 44 taxa with identical

sequences fell out in a polytomy while three clades were resolved: 1) A. pungens from Arizona, A. australis,

Wahlert et al., ITS hyl geny 1 Arctostanhvl 677

Taste 1. Four putative hybrid taxa showing superimposed nucleotide additivity patterns (SNAPs) bet pected parents and hybrid offspring. 21 = ploidy level (26 = diploid,

52 = tetraploid). N | i iti

Taxon Relationship 2n Nucleotide position

54 363 556

A. glauca parent 26 G A G

A. pringlei subsp. drupacea hybrid 26 G/A A/G G/T

A. pringlei subsp. pringlei parent 26 A G T

121 154

A. patula parent 52 T

A. manzanita subsp. roofii hybrid 52 C/T C/T

A. manzanita subsp. manzanita parent 52 C E

121 154

A. stanfordiana subsp. stanfordiana parent 26 T

A. manzanita subsp. laevigata hybrid 52 C/T C/T

A. manzanita subsp. manzanita parent 52 C E

500

A. viscida subsp. viscida parent 26 C

A. hispidula hybrid 26 C/A

A. stanfordiana subsp. stanfordiana parent 26 A

and A. pringlei subsp. pringlei (71% BS), 2) A. pumila and A. tomentosa subsp. tomentosa (65% BS), and 3) A.

malloryi and A. viscida subsp. mariposa (59% BS).

Of the 44 species sharing an identical Gray Leaf haplotype, five are facultative sprouters (A. glandulosa

ssp. cushingiana, A. glandulosa ssp. glaucomollis, A. tomentosa ssp. daciticola, A. tomentosa ssp. insulicola, and

A. tomentosa E roseii) e ai rest are eee nn "is low variability of the nrITS region—and the

fact that both obligate s t an identical sequence—precluded a test of higher

nucleotide substitution rates in seeders "t to sprouters.

DISCUSSION

The phylogeny inferred from the nrITS data sets reveals less resolution than had been expected for this mor-

phologically diverse and widely distributed genus. The species composition of Clades 1 and 2 recovered in

the MP analysis of the reduced nrITS data set is largely congruent with the two-clade molecular phylogenies

of Markos et al. (1998) and Boykin et al. (2005), and the flavonoid phenogram of Denford (1981).

None of the subclades recovered in MP analyses have a discrete pattern of geographical distribution

that can be mapped onto the phylogeny. Although the circumarctic distribution of Arctostaphylos uva-ursi is

unique in the genus, this species is also distributed along the coastal areas and mountains of western North

America where it co-occurs wath oier taxa (e. 8» e CA FEIER s imbricata Eastw., A. patula). Neither

hies that can be mapped

are there discrete and readily i

onto this phylogeny to dead the present ono m of Wells (2000) c or r the PUEDES n were

recovered in tl Boykin et al. (2005) found that the cl

bark, distribution af suia, and ploidy level were not consistent with monophyly of certain A s usegeni:

groups and that some states have arisen more than once in the genus. Because Wells’ (2000) classification

of Arctostaphylos is based on potentially homoplasious morphological characters, discordance between his

classification and the nrITS phylogeny is not unexpected.

The occurrence of polymorphic nucleotide positions showing patterns of additivity between synapo-

morphic nucleotides of putative a and E poems is ue of hybrid origin in four taxa

(Table 1). However, many ity between parents and hybrid

offspring would be needed to eonim hybrid origin lor: the taxa in Table 1 (e.g., Sang et al. 1995). Such

678

100

e»

lexas 512)

A fall n FI > Im L || ro. WT

uva-ursi Alaska

uva-ursi Michigan

uva-ursi New Jersey

uva-ursi Washington

uva-ursi Colorado

uva-ursi Wyoming

mendocinoensis

nummularia

stanfordiana subsp. decumbens

stanfordiana subsp. raichei

stanfordiana subsp. stanfordiana

ohloneana

patula Nevada

rudis

hookeri subsp. hookeri

patula Calaveras Co., CA

pungens California

hookeri subsp. hearstiorum

sensitiva

densiflora

patula Plumas Co., CA

uva-ursi Guatemala

pungens Arizona

71 [— | |

. australis

A. pringlei subsp. pringlei

3. RN A. pumila

A. tomentosa subsp. tomentosa

EN a A. malloryi

A. viscida subsp. mariposa

DPRPAAAPAAAAAAAAAAA AAA ASAS

tomentosa subsp. crustacea (1)

tomentosa subsp. crustacea (2)

manzanita X A. stanfordiana

tomentosa subsp. subcordata

klamathensis

>b>b>bbrho»>»

100

viscida subsp. pulchella

ED Arctous rubra Alaska

Arctous erythrocarpa Siberia

Comarostaphylis diversifolia

Ornithostaphylos oppositifolia

Arbutus menziesii

Xylococcus bicolor

Fic. 1. Strict f 350

€

The taxon “G

" A

ol uii "-— ee BE aF

Wahlert et al., ITS phyl g JA a. 4 L I 679

polymorphisms suggest: 1) recent hybrid origin where both parental alleles are maintained in the hybrid

progeny, 2) different alleles existing between parental genomes in tetraploids, 3) recently substituted nucle-

otides that have not been fully homogenized via concerted evolution, or 4) intra-individual nrITS paralogs

(Wendel et al. 1995; Koontz et al. 2004; Bailey et al. 2003).

The 44 taxa ne d Gray Leaf haplotype are morphologically diverse and are composed of both

obligate seeders and facultative sprouters (Appendix 2). Sprouting species sharing the Gray Leaf haplotype

are represen by five tetraploid taxa of the Arctostaphylos glandulosa-tomentosa complex, whereas most of

the other taxa are diploid obligate seeders. As a result, a hypothesis of greater nrITS nucleotide substitu-

tion rates in short generation-time seeders vs. long generation-time sprouters could not be tested. These

results suggest common € of all isi "ad " taxa, and Py extension, Due remaining taxa in Clade 2.

of common ancestry,

I fI

then here has been much vam in Alle morphology, cytology, and ecological life-histories of taxa

in Clade 2 without corresponding nrITS sequence divergence.

Resolving species-level phylogenies is difficult in perennial, reticulating, and putatively recently di-

versified genera such as pide Bue other genera containing iene that are mney distributed and

morphologically diverse (e.g., Ceanothus ( ), Hardig et al. 200 ), Bailey et al.

2004, Inga (Fabaceae), Richardson et al. 2001). Low interspecific sequence dI. in the niTS region in

Arctostaphylos may be due several factors, including recent diversification in California since the end of the

Miocene, long generation time between sexual reproduction events, genome homogenization via introgres-

sion, ama Enid sales evolution of nuclear ribosomal ITS aras in both diploids and Md d

that di een region

L [ato] e I

J 4 1 1 1 re a ee s A " " 1...1

provide insufficiently va riable DNA

Other PCR-based techniques have been used in Arctostaphylos without success. For M the M

of endophytic fungi in the sclerophyllous leaves of Arctostaphylos prevented the use of RAPDs (randomly

amplified polymorphic DNAs) and inter-simple seq ts (ISSRs) (C. nes pers. comm., Wahlert

unpubl. data). Another approach using RFLPs (restriction fragment length pol ) of the chloroplast

genome may prove useful in ML) the has origin of A. meda (K. Schierenbeck, ges comm...

acr 1 LR y

Bi Pealeliidaily inherited nuclear BOLCS Val ability alciiccucil y in

Arctostaphylos. Sequences from single- or Ipw-copy €—— genes or from anonymous nuclear a using

sequence characterized amplified regions (SCARs) may provide a means to resolve the phylogeny and to

tease apart patterns of reticulation in Arctostaphylos. Until then, many fascinating questions of polyploidiza-

tion, hybrid and ecological speciation, biogeography, and ecology will go unanswered in this remarkable

genus.

APPENDIX 1

List of taxa used in this study with voucher f ti | GenBanl ion numbers for nrITS sequences. All

vouchers are deposited in the Harry D. Thiers Herbarium at San Francisco State University GESU) except for Arctous —

rocarpa, which is deposited at MO. Numbers in parentheses indicate different accessions of the same taxon. * Denote

sequence obtained from GenBank

n Arbutus menziesii Pursh*, AF086828; Arctous rubra (Rehder & E.H.Wilson) Nakai*, AF091944; Arctous oe

mall., Russian Siberia, MO accession number 5196245, GQ281006; eeu diversifolia (Parry) Greene*, AF091947;

Otho oppositifolia (Parry) Eu AF297794 (ITS 1 and 5.85), AF091962 (5.8S and ITS 2); Xylococcus Bieler Nutt*,

AF091

ingroup. Arct hyl d A.Gray, San Mateo Co., CA, USA, VT.Parker & M.C Vasey 764, GQ280910; A. auriculata Eastw.,

Contra Costa Co, CA, USA, M.C. Vasey 840, GQ280911; A. australis Eastw., Baja Norte, Mexico, M.C. Vasey Ma e A.

bakeri Eastw. subsp. bakeri Eastw., Sonoma Co, nh USA, M.C. Vasey 71, GQ280913; A. bakeri su N oma

O, CA, USA, V.T.Parker & M.C.Vasey 548, 60280914 tw., Marin Co., CA USA, VT.Parker & M C Vas

443, GQ2809 15; A. canescens subsp. sonomensis oa PVWells, Lake Co, CA, USA, M.C Vasey 282, GQ280916; A. caine

PV.Wells, Santa Barbara E CA, US, M E y Pa GQ280917; A. columbiana Piper, Humboldt Co., CA,

M.C. Vasey 1030, GQ280918; A. confertifiora Eastw., Santa Barbara O., CA, USA, VT.Parker & M.C Vasey 900, GQ280919; A. cruzen-

680 t tani i Texas 3(2)

sis Roof, San Luis Obispo Co., CA, USA, VT.Parker et al. 12, GQ280920; A. densiflora M.S.Baker, Sonoma Co., CA, USA, M.C.Vasey

69, GQ280921;A. edmundsii J.T.Howell, Monterey Co, CA, USA, M.C. Vasey 819, ii A. a, Parker & M.CVasey,

San Benito Co., CA, USA, VT.Parker & M.C.Vasey 1086, GQ280923; A. gl cushingiana (Eastw.) Adams ex McMinn,

Santa Barbara Co, CA, USA, M.C.Vasey 1002, GQ280924; A. glandulosa subsp. glandulosa Fastw. (1), San Diego Co., CA, USA,

Me Vasey 724, GOZED A. PENAMAA a glandulosa (2), San Benito Co, CA, USA, VT.Parker & M. E Vasey 456, GQ280926;

N s Angeles ce; CA, USA, M.C. Vasey 1000, GQ280927; A. glauca Lindl., Los Angeles

iu CA, USA, M.C. Vasey 809, GQ280928; A. g ib., Santa Cruz Co., CA, USA, M.C. Vasey 401, GQ280929; A. hispidula

Howell, Del Norte Co., CA, USA, M.C. Vasey 360, GQ280930; A. hookeri subsp. franciscana (Eastw.) Munz, cultivated material;

Strybing Arboretum, Golden Gate Park, San Francisco, CA, USA, GQ280931; A. hookeri subsp. hearstiorum (Hoover €: Roof)

PV.Wells*, San Luis Obispo Co., CA, USA, AF1 ud. A TOON subsp cp Santa Cruz Co, CA, USA, V.T.Parker &

M.CVasey 55, GQ280932; A. hookeri subsp Wells, n Co, CA, USA, S. Markos 613, GQ280933; A. hooke-

ri subsp. ravenii PV. Wells*, San Francisco Co, CA, USA, AF106819; A. | Dg. Monterey Co., CA, USA, M.C.Vasey 1050,

GQ280934; A. imbricata Eastw., San Mateo Coh CA, USA. M.C. d 18, GQ280935; A Greene, Santa Barbara Co. CA,

USA, Anonymous s.n., GQ280936; A. | W. Keeler-Wolf & W.Knight, Siskiyou Co, CA, USA, M.C. Vasey 756,

GO280937: A. luciana PV Wells, San Luis Obispo Co., CA, USA, o 1046, "0070008; A. malloryi (W.Knight € Gankin)

PV. Wells, Shasta Co., CA, USA, pa Casey 83 834, ( GQ280939; bowermanii Roof, Contra Costa Co.,

CA, USA, M.C.Vasey 169, GQ2809 ) PV. Wells, Glenn Co, CA, USA, M.C.Vasey 216, GQ280941;

A. manzanita subsp. o eS Sonoma Co, CA, USA, VTParker & M.C. Vasey 508, GQ280942; A. manzanita subsp.

laevigata (Eastw.) Munz, Contra Costa Co., CA, USA, - E lcd mi A. rre subsp. manzanita Parry (1), Men-

docino Co, CA, USA, VT.Parker & M.CVasey 369, GQ280 a (2), Humboldt Co., CA, USA, M.C. Vasey

851, GQ280945; A. manzanita subsp. manzanita (3 g pet Co., CA, USA, Ve a 767, GQ280946; A. manzanita subsp.

manzanita (4), Sonoma Co, CA, USA, M.CVasey 424, GQ280947; A. manzanita subsp. roofii (Gankin) PV.Wells, Glenn Co., CA,

USA, M.CVasey 883, GQ280948; A. manzanita x A. stanfordiana, sonoma Co; E JS ME M 425, GQ280949; A. x media

Greene, Mendocino Co., CA, USA, M.CVasey 253, GQ280950; A Nells, ocino vOv CA, USA, G.Wahlert

s.n., GQ280951; A. mewukka Merriam, Tuolumne CO, A USA, M.C.Vasey 575, GQ280952; A f, San Mateo Co,,

CA, USA, M.C.Vasey 743, GQ280953; A , Monterey Co., CA, USA, M.CVasey 656, GQ280954; A. morroensis

Wiest. & B. Schreib., San Luis Obispo Co., CA, USA, K. Bode 823, GQ280955; A. myrtifolia Parry, Amador Co., CA, USA, M.C.Vasey

733, GQ280956; A. nevadensis A.Gray, Siskiyou Co, CA, USA, M.CVasey 757, GQ280957; A. nissenana Merriam, Placer Co., CA,

USA, M.CVasey 892, GQ280958; A. nummularia A.Gray, Mendocino Co., CA, USA, VT.Parker & M.CVasey 516, GQ280959; A. obis-

poensis Eastw., Monterey Co., CA, USA, M.CVasey 1052, GQ280960; A. ohloneana M.CNasey & V.T.Parker, Santa Cruz Co., CA, USA,

VT.Parker & M.CVasey 111, GQ280961; A. osoensis PV.Wells, San Luis Obispo Co. CA, USA, K. Bode 780, (30280962; A. otayensis

Wiesl. & Schreib., San Diego Co., CA, USA, M.C Vasey 748, GQ280963; A. x pacifica R f San Mateo Co., CA, USA, M.C Vasey 20,

Exp A. pajaroensis J.E.Adams, Monterey Co., CA, USA, V.T.Parker & M.C.Vasey 459, (30280965; A. pallida Eastw., Alameda

; ,M.CVasey 436, GQ280966; A. patula Greene (1), White Pine, Co., Nevada, USA, M.C. Vasey 626, GQ280967; A. patu-

(2), Plumas Co. CA, USA, M. Wood 316, GQ280968; A. patula (3), Calaveras Co., CA, USA, VT.Parker & M.C. panne GQ280969;

A. pilosula Jeps. & Wiesl. San Luis Obispo Co, CA, USA, M.CVasey 233, GQ280970; A. pringlei subsp. drupacea Parry, Riverside

Co, CA, USA, M.C.Vasey 993, GQ280971; A. pringlei subsp. pringlei Parry, Pima Co., Arizona, USA, MC Vasey 232, oe 2; A.

pumila Nutt, Monterey Co., CA, USA, V.T.Parker & M.C. Vasey 677, GQ280973; A. pungens HBK (1), San Bernardino Co., CA, USA,

M.CVasey 194, 60280974; A. pungens (2), San Diego Co., CA, USA, M.C. Vasey 696, GQ280975; A. pungens (3), Pima Co., Arizona,

USA, M.C. Vasey 228, GQ280976; A. purissima PV.Wells, Santa Barbara ra CA, ust UAE 2,GQ280977; A. rainbowensis

JE Keeley & Massihi, San Diego Co, CA, USA, K. Bode 775, GQ280978; , San Mateo Co., CA, USA, M.C. Vasey

557, (30280979; A. x repens J.T. Howell, Marin Co., CA, USA, VOR C Vasey 276, GQ280980; A. rudis Jeps. & Wiesl., San Luis

Obispo Co, CA, USA, VT.Parker & M.C Vasey 151, GQ280981; A. sensitiva Jeps., Marin Co, CA, USA, VT.Parker 306, GQ280982; A.

silvicola Jeps. & Wiesl., Santa Cruz Co., CA, USA, V.T.Parker & M.C.Vasey 865, GQ280983; A Parry subsp. decumbens

(PV. Wells) PV.Wells, Sonoma Co., CA, USA, M.C. MISC HAUS A.stanfordiana Parry subsp. raichei W.Knight, Mendocino

Co., CA, USA, M.CVasey 277, GQ280985; A. st stanfordiana, Napa Co., CA, USA, M.C.Vasey 392, (0280986;

A. tomentosa subsp. crustacea (Eastw.) PV.Wells (1), Monterey co, CA, UA ee 584, (11230207; és tomentosa subsp.

crustacea (2), Santa Cruz Co., CA, oe id E vas» 786, GQ28098 PA n Luis Obispo Co.,

A, USA, M.CVasey 1008, GQ280989; la PV.Wells, Sant Barbara Co., CA, USA, mi

903, GQ280990; A. tomentosa ene rosei (Eastw.) PV. Wells, Monterey Co., CA, USA, M.C.Vasey 822, GQ280991; A. tomentosa

subsp. subcordata (Eastw.) PV.Wells, Santa Barbara Co, CA, USA, VT.Parker & M.CVasey 901, GQ280992; A. tomentosa subsp.

tomentosa Pursh, Monterey Co., CA, USA, me Vds yud3, eb. A. uva-ursi (L) Spreng. Ocean Co., New Jersey, USA, V.T.Parker

437, GQ280994; A. uva-ursi, Sleeping Bear ! | Lal e, Michigan, USA, McEaghan 933, GQ280995; A. uva-ursi, Yellows-

tone National Park, Wyoming, USA, VTParker & M CVasey 942, pierces A. uva-ursi, Gunnison Co., Colorado, USA, M.C.Vasey

3, GQ280997; A. uva-ursi, Alaska, USA, J. Bishop 684, GQ280998; A. uva-ursi, Clallam Co., Washington, USA, M.CVasey 340,

GQ280999; A. uva-ursi subsp. cratericola (J.D.Smith) PV Wells, cultivated material from Guatemala, UC. Botanical Garden, 92-

0545, GQ281000; A. virgata Eastw. Marin Co., CA, USA, M.C. Vasey 783, GQ281001; A viscida subsp ) PV.Wells,

Tuolumne Co., CA, USA, M.C.Vasey 499, GQ281002; A. viscida subsp. ; Ichella (H PV. Well Sonoma Co., CA, USA, M.CVasey

Wahlert et al., ITS phyl g 1 Arctostaphvl 681

+ LI +

470, GQ281003; A. viscida subsp. viscida Parry, Eldorado Co., CA, USA, VT.Parker & M.CVasey 849, GQ281004; A. wellsii W.Knight,

San Luis Obispo Co, CA, USA, M.C.Vasey 1047, GQ281005.

APPENDIX 2

de taxa with identical nrITS seq (i.e, the “Gray Leaf" haplotype).

A hyl d ij, A. auriculata, A. bakeri ssp. bakeri, A. bakeri ssp sublaevis, A. canescens ssp. canescens, A. canescens ssp

sonomensis, A. catalinae, A. iui A. confertiflora, A. cruzensis, A. gabilanensis, A. glandulosa ssp. cushingiana, A. glandulosa

ssp. glaucomollis, A. glauca, A. glutino A. hookeri ssp. franciscana, A. hookeri ssp. montana, A hookeri ssp. ravenii, A. hooveri, A

imbricata, A. insularis, A. Fera A. manzanita ssp. elegans, A. manzanita ssp. nina A. manzanita ssp. manzanita (1), A.

manzanita ssp. manzanita (3), A. manzanita ssp. manzanita (4), A. x media, ontaraensis, A. montereyensis, A. morroensis, A

obispoensis, A. osoensis, A. pajaroensis, A. pallida, A. purissima, A. regismontana, re x repens, A. silvicola, A. tomentosa ssp. daciticola,

A. tomentosa ssp. insulicola, A. tomentosa ssp. roseii, A. virgata, A. wellsii.

ACKNOWLEDGMENTS

We thank Robert Patterson (SFSU) and Harvey Ballard, Jr. (Ohio University) for reviewing this manuscript

and providing comments that improved it. David Keil (Cal Poly State University) and an anonymous re-

viewer also provided helpful commentary that enhanced the paper. Frank Cipriano (Conservation Genetics

Lab, SFSU) and Juniper Scribner provided assistance with sequencing. Oriane Hidalgo (Ohio University)

translated the abstract into Spanish.

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THE REINSTATEMENT OF BEAUTEMPSIA (CAPPARACEAE) AND A

KEY TO THE GENERA OF NEOTROPICAL CAPPARACEAE WITH

VARIOUSLY STELLATE OR PELTATE INDUMENTA

Xavier Cornejo

The New York Botanical Garden

200th St. and Kazimiroff Avenue

Hugh H. Iltis

Department of Botany,

University of Wisconsin

Bronx, New York 10458-5126, U.S.A. 30 Lincoln Driv

xcornejo@nybg.org, cid Wisconsin ae U.S.A.

xcornejoguay@gmail.com wis@charter.net

ABSTRACT

Beautempsia (Benth. & Hook. f.) Gaudich. i ted pecific g f pical Capt It is represented only by

n 4 : : E FI thy) Alla : : ] 1 ] 31 4 A +1 Esp Im Jg t

Í , Jo

in Peru Its] (^ : iif Tia IL zl J i; ) y Fan Di : Ils a Dam Ce D ex DC ¡PRES ty ] ad The studied

specimens are fully cea. ‘nd a key t Í g pp vith variously stellate or p

Key Wonps: Beautempsia, Capparaceae, coastal dry forests, Ecuador, Peru

RESUMEN

R j R [ ia (Benth. & Hook. f} Gaudich., gé pi lde C ini | B

ener (Kunth 4 Alleiz. p g y secos del 1 occidente d de Ecuador i Ds desiertos de Perú. Se

fo E 17 el E :

E I"

eau dos de esta especie ysep presenta una clave para | é | ical de Capparaceae ti ind t trellado de

tado

r [^] I rr t

varios tipos o pelta

PALABRAS CLAVE: Beautempsia, Capparaceae, | , Ecuador, Perú

Beautempsia was proposed by Gaudichaud (1842) as a new generic name with a single species based on

Capparis avicenniifolia Kunth (1821), a distinctive species of neotropical Capparaceae, restricted to the very

dry woodlands and thorn scrub of western Ecuador and deserts in Peru. Ancón by ill health Johnston

1944), Gaudichaud was able to only publish plate 56 with tl Beaut iaefolia, without text.

Gaudichaud’s plate 56 consists of a leafy branch bearing inflorescences and a fruit, and a detailed group

of separate figures, all without captions or descriptions (Fig. 1). Therefore, Beautempsia Gaudich. (1842) is

not valid because Art. 41.2 of the ICBN (McNeill et al. 2006) states, a description or diagnosis is required

for valid publication of a genus

Subsequently, Bentham and Hooker f. (1862: 109) formally established Beautempsia as a section of

Capparis, with a short latin description, citing “Gaud. Voy. Bonite, t. 56,” and without citation of any type

species. That is the earliest legitimate name that has priority and must be followed in accordance with Arts.

41.2, 45.1, 45.3 of ICBN (McNeill et al. 2006). Three years later Eichler, in Martius’ Flora Brasiliensis (1865:

267, 269), raised it to Capparis subg. Beautempsia, citing Bentham & Hook f., “Gaud. Voy. Bonite t. 56,” with

a Latin diagnosis, and only represented by a single species, Capparis avicenniifolia. The respective captions

for Gaudichaud's plate 56 were published in French 14 years after the publication of Gaudichaud's Atlas,

12 years after Gaudichaud's death, and after the Bean 2 ME t ee of Capparis sect.

Beautempsia. That work (Gaudichaud 1866) of Charles D’Alleizette,

who provided explications and descriptions for the plates, validating the generic name and the respective

new combination (Gaudichaud 1842). Although Gaudichaud (1842, 1866) did not cite the basionyms, Art.

33.3 of the ICBN support them as valid taxa those published before Jan 1953.

Hutchinson (1967: 307) attempted to realign the taxonomy of the genus Capparis, and correctly re-

J. Bot. Res. Inst. Texas 3(2): 683 — 689. 2009

684

Voy. de ia Bonite.

e F ds hl

de £ T.

Texas 3(2)

CAPPARIDÉES.

Lave a ta

BEACTEMPSIA AVIGESANTSEOLLA va.

Fic 1 n " L] p ss ye FW al.) AN ni rad J Ig "m l HA | In Aal (1842) 1 Rranch J Elawar ? A | I L J

Flower bud, adaxial view. 6. Polien grains. 7—9. Anthers, dorsal, land lateral views. 10 ction of anther. 11. Ovules. 12

sa M.sl ¡ a 14 Avilac 15 || git ye I PEI £ y 1h F a EL ?4. 17 || é fy

LE ” J Ln ye I £ L

4. Young fruit. 5.

fruit. 18. Seed. 19 Embryo 20.1

Cornej | litis, Reinstat nf Raat ; 685

garded Beautempsia as a valid genus. In this paper, Beautempsia is again recognized as a neotropical genus

of Capparaceae.

Beautempsia (Benth. & d - Gaudich., Voy. Bonite, Bot. 4:38. 1866. Teus: Camas avicenniifolia Kunth. BasioNvw:

apparis sect Hook. f., Gen. Pl. 1:109. 1862. Capparis subg. B th. & Hook. f.) Eichl. in Mart.,

Fl. Bras. 13(1):276. 1865. t Gaudich., Voy. Bonite, Bot. 3 (Atlas), pl. 56. 1842, nom. el

Unarmed shrubs or small trees, with stellate trichomes throughout. Leaves simple, spirally arranged, with

petioles of similar length, lacking a pulvinus and stipules. Inflorescences terminal short racemes, flowers

pedicellate, zygomorphic, linear-bracteate, the bracts soon deciduous, the flowerbuds globose-obovate just

preceding anthesis. Calyx with open aestivation, gamosepalous, hemisphaerical-cyathiform, tetralobulate

to slightly tetracrenate or subtruncate. Hypanthium present. Nectary scales 4, deltoid, mostly free, fused

at the base, rounded at the apex. Corolla spirally twisted (torsivus), petals 4, unguiculate, sessile. Stamens

16 to 18, the outer provided with short staminal nectaries at base of filaments, the inner with glandular

tissue at base of filaments, all perfectly articulated with each other at the base, and forming a nectariferous

dish-shaped digitate pseudotorus supported by a common androphore. At anthesis the stamens project

outward and arch upward, with the gynophore abaxially strongly off-center. Ovary 2-locular. Fruits small

pepos (122-4 x 1-2 cm, containing a sticky orange pulp (in vivo); seeds cochlate-reniform, rounded to

bean-shaped, slightly flattened laterally, the testa light brown (in vivo), with a sarcotesta infiltrated by hairs,

the cotyledons convolute, the embryo white.

Etymology.—The generic name is formed by the French adjective beau, which beautiful, and the

French noun temps, which means weather. That refers to the beautiful weather that impressed Gaudichaud

during his visit to the coast of Peru where he found this genus.

Beautempsia is characterized by stellate trichomes, gamosepalous hemisphaerical-cyathiform calyces

with very short to highly reduced lobes, presence of staminal nectaries articulated to the base of the fila-

ments, forming a nectariferous dish-shaped digitate pseudotorus, and pepo fruits.

Due to the presence of relatively shal zygomorphic flowers with off center gynophores, unguiculate

petals, hypanthium, and seeds with white embryo, covered by a sarcotesta, B Atamisquea

Miers, another bee-pollinated and xerophytic specialized monotypic genus (Camejo & Iltis 2008). But the

latter mainly differs from Beautempsia by the presence of 2-seriate calyces with well developed dimorphic

decussate sepals (vs. calyces l-seriate with small or reduced lobes), ovary at the apex and style sharply

acute to subrostrate (vs. ovary at the apex thick, without style and with the stigma capitate, sessile), and the

absence (vs. presence) of a dish-shaped pseudotorus.

This is a monospecific genus restricted to the dry coastal woodlands and thorn scrub of Ecuador and

deserts in Peru. It is represented by the following species.

Beautempsia avicenniifolia (Kunth) Gaudich. ai Voy. Bonite, Bot. 4:38. 1866. (Fig. 1).

Basionym: Capparis avicenniifolia Kunth l'avicennifolia'], Nov. Gen. Sp. 5:94. 1821. Colicodendron avicenniifolium (Kunth) Seemann

[‘avicenniaefolium’], Bot. H.M.S. Herald 1:78. 1853. Tyre: PE bus in ete arenoso Maris Pacifici, inter Cascas et Trujillo

Peruvianorum, Floret Augusto,” Humboldt & ; g THE LECTOTYPE: B-10 0242517

[WIS photo

Capparis ovalifolia Ruiz & Pav. ex DC., Prodr. 1:253. 1824. C lifolia Ruiz & Pav. ex E.A. López, Ann. Inst. Bot. AJ. Cavanilles

16:384, fig. 432a. 1958, nom. illegit. Tre: ECUADOR. Pat MU [Huayaquil], Morro, 1803, J Tafalla (=Ruiz) s.n. (LECTOTYPE,

here designated: MA).

Speci ined: ECUADOR. Manabi: Bahía de Cará , 12 May 1968, Harling et al. 9454 (GB, US, WIS), Punta de Jaramijó, ca.

7 m, 5 May 1985, Harling & Andersson 24852 (GB, NY, QCA, WIS): Manta, 6 Apr 1955, Asplund 15972 (B, LD, NY, S, TEX, Z); vicinity

of Manta, 80°41'W, 0°57'S, ca. 5 m, 1 Oct 1985, Madsen 61025 (AAU, QCA, CENE 14 n SW > Manta on new road to San Lorenzo,

80°47'W 1%01'S, 300 m, 2 Jun 1997, Neill & Asanza 10708 (DAV, MO, QCNE, WIS); an de La Armada, 80?44"W,

0?7'S, 10 m, 24 Nov 1992, Alvarez (collected by ibt 747 (AAU, MO, QCNE); road Monissisti iia 13 May 1968, Harling et al.

9488 (GB, GH, QCA, WIS); 20 km N of Jipijapa, Pennington & Tenorio 10696 (K, QCA, QCNE); 6 km SW of Jipijapa on road to Cayo

río seco de Jipijapa drainage, 225 m, 2 Nov 1965, Anderson 2501 (RSA, WASH); Machalilla National Park, Playa Los Frailes, 80°44'W,

1°31'S, 125 m, 4 Aug 1990, Cerón et al. 11760 (MO, QCNE, WIS); 5 m, 23 Jan 1991, Gentry & Josse 72701 (MO, QCNE); km 19, bosque

686 t tani titute of Texas 3(2)

muy seco tropical, ca. 30 m, Aug 1994, Bonifaz & Cornejo 2447 (GUAY); Isla de La Plata, 81°13'W, 1916'S, 26-27 Jun 1973, Holm-Nielsen

et al. 7286 (AAU, NY, QCNE); 81°04'W, 1°17'S, ca. 25 m, 5 Feb 1992, Josse 760 (AAU); 81°06'W, 1°18'S, ca. 35 m, 18 Aug 1993, Núñez

127 (MO, QCNB); 81°04'W 1?17'S, ca. 60 m, 6 Apr 1991, Cerón et al. 13986 (QCNE); 81°06'W 0°26'S, 70 m, 21 Nov 1993, Woodruff &

Núñez 580 (NY, QCNE). Guayas: Guayaquil, pas Hinds s.n. a [in Sinclair Bot. Voy. Sulphur]; Colonche, 2 May 1956, Asplund 20404

(NY, S, TEX, US); Manglaralto, ca. 5 m, 10 Feb 1966, Sánchez 10 (GUAY, WIS); between Salinas and Manglaralto, 19 Nov 1952, Fagerlind

& Wibom 208 (LD, MO); Punta Ayangue, 80°46'W 157s, 19 Mar 1941, Svenson 11343 (BKL); Ayangue, 10 m, 26 Oct 2007, Cornejo &

Grochowski 7998 (GUAY, NY [samy yangue, entre Palmar y Valdivia, Aug 1980, Valverde 4826 (GUAY,

L, MO); small hill at Pacoa, 10 km N of San Pablo, ca. 2 km f the sl 80°45'W, 2°06'S, ca. 50 m, 26 Mar 1973, Holm-Nielsen et

al. 2532 (AAU, MO, NY); Salinas, 80°58'W, 2°13'S, 0 m, 2 Nov 1983, Barford et al. 48448 (AAU, QCNE, WIS); ond Guayaquil-Salinas,

2-3 W of Progreso, 80°25'W, 2°23'S, 100 m, 4 Feb 1987, Bohlin et al. 1167 (GB, QCA); km 89-90 from Guayaquil, just E of village

of Buenos Aires, 35 m, 17 Jul 1986, Plowman €» Alcorn 14312 (F, QCA, WIS); km 80 W of Guayaquil, 25 Aug 1965, Játiva & Epling 914

(LA, NY); along highway 17 km W of Progreso (Gómez-Rendón), 12 km E of Zapotal, 80°30'W, 2°22'S, ca. 100 m, 25 Jun 1977, Iltis

& Iltis E-1 (QCA, WIS). ca. 2.5 km SE of Salinas, just S of highway to Santa Elena, 80°57'W, 2°12'S, 2 m, pee Jun 1977, Iltis & Iltis E-20

(QCA, WIS); Punta de Santa Elena, 81°W, 2°12'S, 21 Mar p Holm- ae et al. 2353 (AAU) ta Flena and Punta Blanca,

80°50'W, 2°11'S, 22 Mar 1973, Holm-Nielsen et al. 2402 (AAT ta Elena and Ancón `, 80049! W, 2°13'S, 26 Feb 1941, os

11171 (BKL, GH, NY, US); 4-6 km S of Santa Elena on ue to oe sea level, 18 Feb 1974, Man 10010 ipe WIS); near Ancón, 2

pos 1965, Játiva & Epling 916 (LA); Lad le P ta Elena, entre Punta A t Ipa, 30 m, 19 Feb 1982, Baslev 2242

A, QCNE); Chanduy, in litore Maris Pacifi, Jun 1862, S} 6402 (BM, CGE, K. LE, OXF, an 1 km NW of Chanduy towards

pence 80°41'W, 2°23'S, ca. 5 m, 18 Mar 1973, Holm-Nielsen et al. 2147 (AAU, NY); Punta Carnero, 0 m, 20 May 1981, Dodson &

a 10994 (MO, QCNE, SEL, WIS); Pacific coast inmediately W of Punta Carnero, 80°55'W, 2°15'S, 17 Mar 1973, Holm-Nielsen

000 (AAU, MO, NY, QCNE, WIS); 400 m E of Punta Carnero, 80°55'W, 2°15'S, 17 Mar 1973, Holm-Nielsen et al. 2052 (AAU, MO,

NY, a Vía Playas—Posorja, May 1978, eee 2541 (GUAY, MO, SEL); Ayalán, 80°20'W, 2°40'S, 1 Mar 1990, Bonifaz 682 (GUAY, WIS);

Morro, cercanías, 5 m, 20 Aug 2002, C 85 (AAU, COL, GUAY, MO, QCA, QCNE, US, WIS [sample for DNA studies]); Isla Puna,

Puna Nueva to Santa Teresa, 79°57'W, 2°43) S, 0 m, 3 Dec 1987, Madsen 64140 (AAU, QCA, QCNE, MO); midway between Puna Nue

and Zapote, 79°56'W, 2°44'S, 0 m, 3 Sep 1987, Madsen 63879 (AAU, K, MO, NY, QCA, QCNE); Zapote to Estero de Lagarto, ES

2°43'S, 0 m, 4 Aug 1987, Madsen 63826 (AAU, GUAY, K, MO, NY, QCA, QCNE); entre Puerto Grande y Puna Nueva, 80°7'W, 2°57'S, 0

m, 8 Jul a ae 18175 (QCA); Insula Puna, Oct 1892, Eggers 14756 (GH, LE, M, US, K); ibidem, Mille 785 (US). El Oro: ca. 1 km

N of Huaquillas, 1 May 1980, Harling & Andersson 18812 (GB, dud ve anc 14 Mar 1955, Asplund 15743 (S, LD, NY); entre

Huaquillas y i d al lado de la carretera, 15 Apr-15 May 19 1262 (QCA, WIS); Isla Santa Clara o del Muerto,

80°28'W, 3?12'S, ca. 30 m, 4 May 1997, Vargas & Canaday 1444 e NY, QCNE, WIS); Isla Bellavista, 80?13'15"W, 3°24'40"S, 5 m, 19

Mar 1997, Van den Eynden & Cueva 946 (LOJA, QCA, QCNE). Loja: poa ca. p W, bn 17"S, Vivar & Merino 3430 (LOJA,

QCA, QCNE); road Catacocha-Macará, km 42, 79°50'W, nae 800 m, 2 169 (NY). PERU. Tumbes: Prov

Tumbes, Distr. Tumbes, Bosque Nacional de Tumbes, E tre El Caucho y Pampas del Hospital, 8 Jan 1968, Simpson

& Schunke 600 (NY). Piura: Talara, 9 Apr 1929, Hunt 12 (NY); Dept. Paita, Sech desert between Paita and Pi

to Sechura, ca. 100 m, 1 Jan 1964, Hutchinson & Wright 3356 (UC, NY). Laml Dept. Lambayeque,

2 km E of bridge, 450 m, 9 Feb 1988, Gentry et al. 61379 9 (MO, NY); entre Hda. ee Cl 200-300 n m, 13 Aug 1952, Ferreira

8384 (NY); Carretera entre Cayaltí y Oyotún, 150 m, 10 Oct 1986, Diaz 2086 (MO, NY); 28 kti of Olmos, 125 m, 9 Jun 1978, Gentry

et nus did es, e Prov. Contumazá, 700 m, 29 Dec 1983, lian drin a i La ee Prov. E

m, 20 Jun 1986, Mostacero et al. 1117 (F, MO

Lloc, « 500 m, 2 Jan 1983, St 21985 (MO, NY). Ica: without b Ruiz. s.n., 1788 (B. MA iot seen], MO [ex EM Pavón]).

Etymology.—The specific epithet refers to the bl of the leaf blades of this species with those of the

genus Avicennia L. (Acanthaceae).

Chromosome number.—n = 8, 2n = 16 [! Pazy, ! Przywara]. Ecuador: Guayas, ca. 2.5 km SE of Salinas,

H.H. Iltis & M. Iltis E-20 (WIS). Ecuador: Guayas, along highway 17 km W of Progreso H.H. Iltis & M. Iltis

E-1 (WIS).

Local names.—ECUADOR. Bichaya (Indigenous?, Madsen 64140, 63826); Vichayo (Indigenous?, Van den

Eynden & Cueva 946). PERU. Guayabito de gentil, Guayabo de gentil (Spanish, Mostacero et al. 1117); Vichayo

(Indigenous?, Simpson & Schunhe 600).

ses.—The fruits at maturity have a sweet pulp, reported as edible (Svenson 11343, Núñez 127, Van den

Eynden & Cueva 946; Van den Eynden et al. 1999). However, there are also reports that the *leaves and

fruits are toxic, produce dizziness and hallucinations" (Woythowshi 7068, MO, WIS, in Dpto. Lambayeque,

Peru).

Phenology and Ecological interactions.—This species flowers during the day (Cornejo pers. obs.). This is

an uncommon feature in neotropical Capparaceae, which in the coasts of Ecuador and Peru is also shared

r j | Iltis, R A "un taft D 4 H 687

] m taat scias siat , ' Jae dll

by pr th) Iltis & Cornejo. The small flowers with short stamens and

gynoplotes of Beautempsia avicenniifolia are exollinaied by bees (Madsen et al. 2001; Madsen 63879; Cornejo

pers. obs.). The fruits are eaten and the seeds dispersed by the Sechura fox, Lycalopex sechurae Thomas (Huey

1969), a canid with similar distributional pattern to that of Beautempsia avicenniifolia.

Distribution.—Locally dominant along and close to the Pacific coast, from Cabo Pasado in Ecuador

southward to Ica south of Lima, Peru, often found in the Ecuadorian coastal dry scrub forests, from sea level

to 230 m (to 800 m in the very dry western Andean slopes of Loja), and in the Peruvian deserts, up to 700

m, often forming thickets and thus one of several dune stabilizers, apparently not salt tolerant.

This species has been misreported in Flora of Panama (Woodson et al. 1948) under the name of Cap-

paris ovalifolia. However, Beautempsia avicenniifolia does not occur in that country. It is most likely that the

single specimen cited for Panama (Barclay s.n., *Isthmus of Darien") was collected on the coast of Ecuador

(perhaps by Hinds?, see cited specimen), on the Voyage of H.M.S. Sulphur.

KEY TO THE AMERICAN GENERA OF CAPPARACEAE

WITH VARIOUSLY STELLATE OR PELTATE INDUMENTA

1. mryallavnith cl | +i +i f£

OFrDolld WILI m buds , Stamens not quicl t

2. Calyx ! Seriar palatal or spathaceous or 4-valvate with the sepals rom or fused and the lobes regular, if

irregular g at or nearly t is into 2 to 4 segments.

3. Calyx at anthesis splitting to the base E 1 or : Tua finaly palate or dis Belencita

3. Calyx at anthesis tubular, subcylinaric dish- or/to cup-shaped or

d (the remaining iniu. 24 sepals, lobes o or segment

4, Calyx tubular, subcylindric or fünnetorm, aval Gee arene to orange -red (alive), the apical Es

2-4, indistinct at the early bu ruits i t, decomposing at ma

rity, linear, + torulose, oblong or cylindric to thickly com Steriphoma

4, bd Du ds ud shaped or arde green, ioi Or r brown (alive), the 4 sepals or cays

2 to 4 segments (Morisonia L); stamens T staminódes if any i to > 60, fruits globose, oblong to

linear, if linear often torulose, capsular, pepos or amphisarca.

3. Sepals entirely fused, irregularly cpm i ante into a 3(-4) segments, inflorescences

| Jy | VET y thick altra Ev

eral times shorter than fruit bod Morisonia

5. Sepals or calyx lobes 4, distinct from bud or opening by linear sutures at or nearly to anthesis;

inflorescences terminal; fruits es oblong to lineal, i meai oren torulose paa

pseudoamphisarca, pepos or capsular, p res elongat

6. Sepals widely spaced, e icy naa on and hiding the petal sutures. exposing the

petals and not exceeded by the corolla in buds; corolla with valvate aestivation; cotyledons

somewhat folded, 4-7 mm thick Calanthea

dee not widely spaced, entirely hidding the petals in young buds or if exposing the petals

n buds, then never arranged on the petal sutures and greatly exceeded by the corolla (Qua-

drella [DC] J. Presl, Beautempsia); corolla with torsivus or imbricate aestivation; cotyledons

ip pale convolute, 0.7-1.5 mm thic

. Leaves ovate to ovate-cordate, as wide or wider than long, with (sub) palmate venation at

oe base of the blade; MOE mun yen petals, = stamens, the eneclaly aln

fal. | 1

4, linear to lineal

the soft and very flexible fruit-wall splitting and falling apart in 2 to 4 segments at dd

Capparicordis

7. Leaves elliptic to obovate or oblong, two to several times longer than wide, with pinnate

venation from leaf base; flowers with white to cream or greenish petals, 8 to 60 stamens,

the nectary scales 4, d omo oeltoid, or od o al or eee mesa dish or

ary-cup present, | codendron,

^i receptacle (Quadrel la); IHE indehiscent or if dehiscent (Quaarelia) shel the rula

g gyno; t maturity.

8. Calyx dialisepalous, hypanthium absent; fruits capsular. le leelo a

Quadrella

8. Calyx gamosepalous, the sepals fused at least at the base, hypanthium present; fruits

amphisarca, ESEUHOHIR PASTE: Or pepos

9. Calyx + cup-shaped, v tion, tl pals devel |, their margins and

a renomi dd ene from bud u ul thesis; nectary of a 4-dentate dish or

ora3 ts of t! itl | taries, therefore the

dish- -shaped digitate pseudotorüs:l is absent ‘inflorescences bearing showy flowers

pollinated by vertebrates (bats); fruits large amphisarca or pseudoamphisarca

Colicodendron

9. Calyx hemisphaerical-cyathiform, mie e aestivation, the lobes very short to

highly reduced, their margins not 1 ach other from bud; nectary of 4 scales,

mostly f tatthe ex of the filaments forming a nectarif-

erous dish- -shaped digitate pseudotorus; inflorescences bearing small flowers polli-

ated by insects (bees); fruits smal! | pe pos Beautempsia

2. Calyx 2-seriate, the oak imbricate, partially exp i ds or the outer pair of

c pude du UE ur inner Lad of sepals in m ani Or nearly to anthesis.

. Leaf blades hisarca or pune niece nt) pepos, 4-20

x 4-8 cm, containing 10to = iuge is; ind ta stellat orto. candelabra throughout.

1 ie Flower

metric, petals imbricate in bud, at anthesis adaxially and laterally Mm and the stamens

all arranged; staminodia present; nectary of 3- to 4-dentate nectary cup present

ELO

—

— |

. Flower buds spherical to ovate-globose, not fl |

torsivus (spirally twisted) in bud, petals and stamens radially arranged at anthesis; staminodia

s nectary of 4 scales nt.

eee te tall trees; inflorescences Usa! y A. to baile floral bracts present, de-

ate-connate or valvate- pl

wider and entirely ‘enclosing the narrower and more Wr inner pair of sepals until o

nearly to anthesis; fruits amphisa Neocalyptrocaly

12. Lianas or scandent shrubs; rowers alway solitary i in leaf axis, ebracteate; sepals + equal im

bricate in bud, the outer pai |

overlapping ti pair, the inner pair exposed from

very young bud; fruits pepos.. Mesocapparis

| | +: Ll -l Aza i

- (tardily dehiscent), 4- 6 x 4- 6 mm, containing 1 to 2 pel seeds; indumenta peltate throughout

Atamisquea

1. Corolla with open aestivation from buas, st lily quickl t just bef tl

FTIR INE UI IA]

ACKNOWLEDGMENTS

Thanks are due to the following herbaria for sending their collections as loan or gift for this study: AAU, B,

BKL, CGE, DAN: E, eB GH, GUAY, K, LA, LD, LE, MS NY OXF, P, QCA, OCNE, S, SEL, TEX, UC, US,

WIS, Z. The aut iding a critical review of the manuscript, to John

McNeill, Werner Greuter, John Wiersema and Gandhi Masc for their very helpful nomenclatural discus-

sions, to Marie Long, Chris Pierce and Emma Antobam (NYBG Mertz library) for helping in different ways

to obtain a scan of Gaudichaud's Plate 56 reproduced in figure 1 of this article, and to Barney Lipscomb for

his efficient editorial work.

REFERENCES

GAUDICHAUD, CH. 1866. Voyage autour du monde éxécuté pendant les années 1836 et 1837 sur la corvette La

Bonite, commandée par M. Vaillant Capitaine de Vaisseau, Botanique. Paris, Arthus Bertrand.

BENTHAM, G. AND J.D. Hooker. 1862. Capparideae. In: Genera Plantarum 1. London. Pp. 103-110.

Cornejo, X. AND H.H. ILns. 2008. Two new genera of Capparaceae: Sarcotoxicum and Mesocapparis stat. nov. and

the reinstatement of Neocalyptrocalyx. Harvard Pap. Bot. 13:103-116.

EicHLER, A.W. 1865. Capparideae. in CEP. von Martius, ed., Flora brasiliensis 13(1). München. Pp. 237-292.

GAUDICHAUD, CH. 1842. Voyage autour du monde sur la corvette La Bonite, Atlas. Paris.

Huey, R.B. 1969. Winter diet of the Peruvian desert fox. Ecology 50:1089-1091.

Comejo and Iltis, Reinstat t of Beautempsi 689

HUTCHINSON, J. 1967. In: The genera of flowering plants (Angiospermae) Dicotyledones. Vol. 2. Clarendon Press,

Oxford. Pp. 303-317

JOHNSTON, I.M. 1944. Publication dates of Gaudichaud's botany of the Voyage of The Bonite. J. Arnold Arbor. 25:

481-487.

KuNrH, C.S. 1821. Capparideae. In: von Humboldt, FW.H.A., AJ.A. Bonpland, and C.S. Kunth. Nova genera et spe-

cies plantarum [quarto ed.], 5. Paris. Pp. 82-98.

Mapsen, J, H. Bastev, AND R. Mix. 2001. Flora of Puna Island: plant resources on a neotropical Island. Aarhus Uni-

versity Press.

McNett, J., F.R. Barri, H.M. BurDEr, V. DEMOULIN, D.L. HAwksWORTH, K. MARHOLD, D.H. NicoLson, J. Prano, P.C. Silva, J.E. Skoc,

J.H. WIERSEMA, AND N.J. TURLAND. 2006. international code of botanical nomenclature (Vienna Code). Adopted by

the 17th International Botanical Congress Vienna, Austria, July 2005. Gantner Verlag, Ruggell, Liechtenstein

VAN DEN EYNDEN, V., E. Cueva & O, Casrera. 1999. Plantas silvestres comestibles del Sur del Ecuador - Wild edible

plants of southern Ecuador. Ediciones Abya-Yala, Quito, Ecuador.

Woobson, R.E. 1948. Capparidaceae. In: Woodson, R.E., RW. Schery, and collaborators, eds. Flora of Panama, Part

5. Ann. Missouri Bot, Gard. 35: 75-99.

690 Jour t tani i exas 3(2)

BOOK REVIEW

JANE MANASTER. 2008. Pecans: The Story in a Nutshell. (ISBN 978-0-89672-640-6, pbk.). Texas Tech University

Press, Box 41037, Lubbock, Texas 79409-1037, U.S.A. (Orders: www.ttup.ttu.edu, 1-800-832-4042).

$19.95, 104 pp., 9 color nih 9 b&w pus 1 map, per 22x 9".

a TA A Y JE maus. | Jae fl +

Pecans: The Stor} ry in a Nuts P F e r

the U.S. as well as information on the cultivation, industry, and nutrition of "North Ambien: favorite native nut.” Manaster candidly

writes bou ni ene of the a "a explains zw Eu native peus ne P isi its MD iik ee is

cC clc t e pl ¿E I

e MEHR helped t te t ieties of hat thrive in tl ] United States today. Cultivation led to

de formation i id large ae now a "nn of our silio landscape. Many of the owners or workers developed new grafting

L o I r

NA 4 E 1 J ] a ^el Tes É J - +} ] 1 1 PE E 4 : 1 +

4 L n

pecan logs, and died iced , pies, cal d I The author used hi ical i t ifically show the reader

the RM of me particular species. n book would be a good nds for dnd marae or co xis is egi to inn more about

the life history i McNew, MS Biology, Prog ger, h Institute of Texas, Fort Worth, Texas

76102-4025, U.S. Pu

J. Bot. Res. Inst. Texas 3(2): 690. 2009

REVISION DE LAS ESPECIES SUDAMERICANAS DEL GENERO SCHWARTZIA

(MARCGRAVIACEAE: COMPLEJO NORANTEA)'

Diego Giraldo-Cañas

Instituto de Ciencias Naturales

cultad de Ciencias

Universidad Nacional de Colombia

Apartado 7495, Bogotá, D.C, COLOMBIA

dagiraldoc@unal.edu.co

Se presenta el ane taxonómico para | peci d del género Schwartzia. Se analizan di pect lati la

clasificación, | 1 ri S tan] ] 5 15] loin N

e 1 I L J 4

las 14 esp le Scl tzia y t Sudamérica, así las d ipci le l pecies, sus sinónimos, distril

y se comentan Igun I i fol Sal y C oldgic ao S F d establecer Y UE Col bia es el país con el may for número de

* J Cl PER f. El - t - [62 : y | A e] na 35 " j 4 ] J I f.

E C N E 3 "EI 5 n t > 3 F

-] 1] ds E ), P 3 (d a dA : ) y, Bolivia y V 1 3 J : As

ABSTRACT

Ag : esa] falo C ET » : Cal C. P JOA ] ] } ] i fi nomenclature

4 e i I T t

and generic affinities are discussed. Keys to the fi of the Norantea-Compl 1 the 14 South American species of Schwartzia

= 11 A ES fh : , synonymy, 1,4 T Asters Aas ] 1 T - ] A ] 1 T icueriratireme

i e e E m 1 [^j

1 1 3-5 a4 : : apum | q D “41 : s f£. J L X Enl]

Brazil

(four species, all Enden) Ecuador (three en one Endemio) Peru (two species, one DS. and Bolivia aaa Venezuela with

one species each one.

La familia Marcgraviaceae es de distribución neotropical y consta de 110-130 especies (Cronquist 1981;

Bedell 1989; Dressler 2001, 2004a, b), distribuidas en ocho géneros (Giraldo-Cañas 2007). Su distribución

más A la MM E los estados de Chiapas, Oaxaca y Veracruz en México, mientras UE la más

austral de Paraná y Santa Catarina en Brasil. S

restringidos a bis bosques húmedos a pluviales de las regiones amazónica, andina, saayana. así como en

el Chocó biogeográfico y Mesoamérica (principalmente Costa Rica y Panamá), aunque presenta un mayor

endemismo hacia el noroeste de Sudamérica (Amazonia occidental y los Andes). Cabe destacar que de las

110-130 especies que se conocen para la familia, 56-60 están en Colombia, constituyendo el país con el

mayor número de especies de Marcgraviaceae (Giraldo-Cañas 1999).

La familia Marcgraviaceae se caracteriza porque sus miembros corresponden a lianas epífitas o ter-

restres, o a arbustos epífitos y hemiepífitos, los cuales presentan dos tipos de nectarios, los que, según la

terminología de Schmid (1988), reciben los nombres de nectarios extra-reproductivos laminales (glándulas

presentes en la superficie abaxial de las hojas = “hypophyllous glands”) y nectarios reproductivos (brácteas

nectaríferas presentes en los pedicelos, en la base de éstos, en el raquis o en la base de la flor). Estos últimos

corresponden a los nectarios nupciales extraflorales (sensu Schmid 1988 y Weberling 1992) y son los que

más llaman la atención por su gran tamaño, posición, color y variedad de formas. Además, estos últimos

son de Eran ponia taxonómica en las aii andes

Las} han sido di Su conjunto, aunaue se cuen tacon lg trata entos

sepionalésv coma revisiones del complejo 1 tea (Bedell 1985; no is [ blicada)

y de los géneros Ruyschia Jacq. y ome Aubl. (de Roon 1975). Es por esto que es findamental empren-

Ww teeth 4 NC J Lal m j: -a fat I "TI ý A LE FAL 2 à nm... IIA J Eme .. 4

sidad Nacional de Colombia, Bogotá, D.C.

J. Bot. Res. Inst. Texas 3(2): 691 — 725. 2009

| EN al m . In LI PR £T

602 J t Texas 3(2)

der estudios integrales de la familia a nivel continental mas que regional. Por otra parte, se hace necesario

esclarecer la taxonomia del complejo Norantea, al ae pound los géneros Marcgraviastrum (Wittm. ex

Szyszyl.) de Roon & S. Dressler, Norantea Aubl., P ldo-Carias, Sarcopera Bedell y Schwartzia

Vell., ya que dicho complejo no es wotistilérico (Ward & Price 2002; Giraldo-Cañas 2007).

El género Schwartzia fue considerado por mucho tiempo parte de Norantea (Triana & Planchon 1862;

Wittmack 1878; Szyszylowicz 1893; Gilg @ Werdermann 1925; Macbride 1956; Reitz 1968; de Roon 1970,

1975; Punt 1971; Kummrow 1977; Ferreira 1982, 1995; Peixoto 1985; Pinheiro et al. 1995). Ambos géneros

presentan inflorescencias racemosas, pero difieren basicamente en la longitud del pedicelo, el cual es largo

en Schwartzia (1,4—9 cm) y corto en Norantea [0,3—0,7(-1,0) cm]. Por otra parte, en Schwartzia los nectarios

reproductivos pedicelares son sésiles o cortamente pediculados (pediculo hasta de 7 mm de longitud) y se

ubican generalmente en la porción media a proximal del pedicelo, mientras que en Norantea presentan un

pedículo de 5-20 mm de longitud y nacen en el tercio proximal o en la porción media a distal del pedicelo.

Schwartzia se distribuye ih ds Rica y Bra a del Caribe y Mount hasta iin y el sur de

rada

Brasil, en donde crece en y maduras o g

de alteración (e.g., bosques de montaña, TON aA manglares, campos rupestres, afloramientos

rocosos, restingas, matas, matorrales, cerrados, yungas, riberas de ríos), entre el nivel del mar y los 2500 m

de altitud.

En este trabajo se presenta la revisión de las especies sudamericanas del género Schwartzia, como con-

tinuación del estudio de las especies del complejo Norantea que viene adelantando el autor (véanse Giraldo-

Cañas 2003, 2005, 2006, 2008, Giraldo-Cañas & Fiaschi 2005). El presente aporte pretende contribuir al

estudio de la rica flora neotropical y en especial al conocimiento de la familia Marcgraviaceae.

MATERIALES Y MÉTODOS

Las técni tilizad den a las e das clásicamente en taxonomía y sistemática vegetales. Se

consultaron | lecci ] itad les EN CEPEC, COAH, COL, CR, CUVC, CHOCO, F, HUA,

IBGE, JAUM, MEDEL, MEXU, “MO, MOL, MPU, NY, PSO, RSA, SI, SP, UDBC, UIS, US y VEN (abreviados de

acuerdo con Holmgren et al. 1990). La terminología para las estructuras morfológicas esta basada en Font

Quer (1993), sia en Me referente con d ae ee de los nectarios, siguiéndose en este caso a Schmid

(1988) y para lat ía sexológica iguió a Cocucci (1 980). Por otra parte, las palabras

holotipo, isotipo, lectotipo. isolectotipo ye Gb similares, están escritas de acuerdo con Kiesling (2002: xi),

autor que adopta y aconseja la f cento ortográfico. Los dato ientes al material tipo han sido

tomados de varias fuentes bibliográficas, pene de Bedell (1985), así como de otras fuentes más

recientes y de comunicación personal con curadores de diferentes herbarios (e.g., MEL, MOL, MPU).

RESULTADOS Y DISCUSIÓN

ee actual de los cinco ences s compe Norantea (Tabla 1).

está basado en ^ MuR ex cs ii Maipo soppen.

a Delpino, = Norantea subgen. Marceraviastrum Wittm ). Este g

hasta Bolivia y Brasil, y comprende 15 especies, las que se caracterizan por sus inflotescéntias usibus us

a AS p aue pon pedicels pose una flor fértil 7 un nectario reproductivo. En virtud de sus

o con Marcgravia, pero éste último

posee flores tetrámeras (tairo sépalos: cuatro pétalos) vs. flores pentámeras en Marcgraviastrum, pétalos

connatos a una EA caduca en Marcgravia vs. pétalos libres o sólo basalmente connatos en

flor fértil (en la mayoría de los casos la flor aparece en forma

— en da porción distal ‘del nectario) en Marcgravia vs. cada nectario reproductivo posee una flor

fértil en Marcgraviastrum; además, Marcgravia presenta hojas dísticas, en tanto que en Marcgraviastrum son

espiraladas.

2. Norantea es reconocido actualmente como un género monotípico y corresponde a Norantea subgen.

Giraldo-Cañas, Estudio taxonómi | i lameri Schwartzi 693

Tasta 1. Características de los cinco gé lel complejo A

Género Tipo de Forma y posición del Ovario y nümero Distribución

Inflorescencia nectario reproductivo de estambres aia cay numero

de espec

Marcgraviastrum Umbelada Tubular a pipi. en los Ovario 5-9-locular; | Desde Nicaragua hasta

n pedicelos. Cada nectario estambres 10-75 Perú, Venezuela

reproductivo o posee una flor fértil y Brasil. 15 especies

Norantea Racemosa Tubular, sacciforme, cilíndrico Ovario 5-locular; Sudamérica tropical,

o clavado; en los pedicelos. estambres 15-38 Una especie

Cada nectario oe

posee una flor fértil.

Pseudosarcopera Racemosa Sacciforme a Edo en el Ovario 3 ó 4 locular; Colombia, Ecuador,

raquis, justo por debajo de la estambres 8-14 Perú y Bolivia.

inserción del pedicelo al s Dos especies

Cada nectario reproductiv

posee una flor fértil.

Sarcopera Espiciforme Ciatiforme, cocleariforme, Ovario 2-3(-5)- Desde Honduras hasta

(flores sésiles) cupuliforme o sacciforme; en el locular; estambres el norte de Bolivia, así

raquis (en la base de la unión de 6-30 como en Venezuela y

la flor al raquis). Cada nectario Brasil. Ocho especies

reproductivo posee una flor fértil.

Schwartzia Racemosa Sacciforme, tubular, cimbiforme, Ovario 3-5- ia Desde Costa Rica hasta

ciatiforme, cocleariforme estambres 10- olivia y Brasil, así

galeado, cuculado, esla O (sólo una inis como en V uela y

ventricoso; en los pedicelos. presenta 5) las Antillas Menores

Cada nectario reproductivo 17 especies

Posee una flor fértil.

Sacciophyllum Delpino [= Norantea sect. Sacciophyllum (Delpino) Gilg & Werderm.]. Norantea se caracteriza

poro su Hiorescencia ues y largamente racemosa, por sus flores cortamente pediceladas y por sus

de color rojo encendido, los cuales se ubican en el pedicelo. En Sudamérica

se e distribuye en Colombia, Venezuela, Guayana FISHER urnam, Guyana, Trinidad y Tobago, Ecuador,

Perú, Bolivia y Brasil; al parecer, las pocas colecciones y jamaiquinas provienen de mate-

rial cultivado. Su gradiente allitudinal está Comor aae principalmente entre el nivel del mar y los 1000

m, aunque puede haber algunas poblaciones creciendo cerca de los 2000 m.

3. Pseudosarcopera. Este género es fácilmente reconocible en el complejo Norantea, ya que es el único

que presenta inflorescencias racemosas y a la vez, nectarios reproductivos que siempre nacen en el raquis,

mientras que en los demás géneros los nectarios reproductivos siempre nacen en el pedicelo cuando las

inflorescencias son racemosas, en tanto que en Sarcopera — el género más cercano a Pseudosarcopera - si bien

los nectarios reproductivos nacen en el raquis, sus inflorescencias siempre son espiciformes (flores sésiles).

Hasta el momento, sólo se conocen dos especies para Pseudosarcopera: Pseudosarcopera diaz-piedrahitae

(Giraldo-Cañas) Giraldo-Cañas, una especie endémica de la región del Chocó Biogeográfico de Colombia y

Ecuador, y Pseudosarcopera oxystylis (Baill.) Giraldo-Cañas, endémica de la región andina oriental de Bolivia

y Perú.

4. Sarcopera está basado en las especies anteriormente ubicadas en Norantea subgén. Pseudostachyum

Delpino [= Norantea subsect. eas ae, bae - Norantea sect. Pseudostachyum (Delpino)

Gilg & Werderm.]. Este género s ribi ta Bolivia y Brasil y posee ocho especies, las

cuales se caracterizan por sus inf ias espiciformes (fl 5siles) y por tari productivos,

los que se originan en el raquis justo por and de la inserción de las flores a éste.

5. Schwartzia consta de las especies previamente reconocidas en Norantea subgén. Cochliophyllum y de

694 J t tani ti Texas 3(2)

otras ubicadas en Norantea subgén. Byrsophyllum. Este género fue considerado por mucho tiempo parte de

Norantea, a pesar de que habia sido propuesto por Vellozo en 1825 (publicado en 1829). Ambos géneros

presentan inflorescencias racemosas, pero difieren básicamente en la longitud del pedicelo, el cual es largo

en Schwartzia (1,4-9 cm) y corto en Norantea [0,3—0,7(-1,0) cm]. Por otra parte, en Schwartzia los nectarios

reproductivos son sésiles o cortamente pediculados (pedículo hasta de 7 mm de longitud) y se ubican gen-

eralmente en la porción media a proximal del pedicelo, mientras que en Norantea presentan un pedículo de

5-20 mm de longitud y nacen en el tercio proximal o en la porción media a distal del pedicelo. Schwartzia

incluye 17 especies y se distribuye desde Costa Rica y algunas islas del Caribe hasta Colombia, Venezuela,

Ecuador, Perú, Bolivia y el sur de Brasil, en donde crece en diferentes formaciones vegetales de áreas húme-

das y maduras o con diferente grado de alteración, entre el nivel del mar y los 2500 m de altitid. De las 17

especies del género, 14 están en Sudamérica, siendo 13 especies exclusivas de es | te y una [S.

spiciflora (A.L. Juss.) Bedell] es compartida con las Antillas Menores (Dominica, Guadalupe, St. Vincent y

Trinidad).

CLAVE PARA SEPARAR LOS CINCO GÉNEROS DEL COMPLEJO NORANTEA

1. Nectario reproductivo naciendo en el raquis justo por debajo de la unión del pedicelo o de la flor al raquis.

2. Fl iles (inf j ifi ) tario reproducti iatiforme, cocleariforme, cupuliforme

O sacciforme; ovario 2-3(-5) 5)-locular; estambres 6- 30. Sarcopera

Fl Ta: | J A a | | ips Pie nae. Hi TE EN | ] 3 Ô

* I X y ps di ,

locular; estambres 8-14 Pseudosarcopera

1. Nectario reproductivo naciendo directamente en el pedicelo, nunca en el raquis.

3 El : l I j l l I ) io J "ES «rv ] Ha i E 9-lo cu llar

Marcgraviastrum

TE if, ‚cilíndrico, clavado,

I El H . 4 "n A J

cimbiforme, ciatiforme, cocleariforme, cuculado, galeado, urceolado o ventricoso; ovario 5n 5- [oed

4. is largamente pedice'adas, pedicelo de 1,4-9 cm

n la porción media a proximal del pedicel: E en la porción distal pero nunca cercanos al

cli con un pealculo: de oe : mm ae ong tud Schwartzia

4. Je AF 1 A | T j . : A

en el tercio oroximal o en la porción — a distal del pedicelo, con un pedículo de 5-20 mm de

longitud Norantea

Schwartzia Vell., Fl. Flumin. 5:221. 1825 [1829]. Esrecie to: Schwartzia glabra Vell. [= S. brasiliensis (Choisy) Bedell ex

Gi nas

Arbustos escandentes, lianescentes, epífitos a hemiepífitos, raramente terrestres. Hojas enteras, espiraladas,

cortamente pecioladas, raras veces te pecioladas, coriáceas, glabras, con glándulas laminales

Y Xen Sur er id

en el envés, generalmente xibmáreitidles venación secundaria débil, broquidódroma, en ocasiones supri-

mida en una o en ambas caras. Inflorescencia terminal, [excepto en la especie de Costa Rica $. jimenezii

(Standl) Bedell, en la que siempre es lateral], racemosa, pauciflora o más comünmente multiflora, laxa o

más frecuentemente densa; pea eo oR dea alargados, de 1,4—9 cm de longitud, dispuestos en

forma espiralada en el eje de la i | reproductivos sacciformes, tubulares, cimbiformes,

conformes, cocleariformes, galeados, cuculados, urceolados o ventricosos, uno por pedicelo, pedículos hasta

de 7 mm de longitud, ubicados en la porción media a proximal del pedicelo o raramente cercanos al cáliz

aunque no pegados al mismo; en ocasiones el pedicelo más proximal con una bráctea foliosa y el terminal

sólo con el nectario sin llegar a desarrollar flor alguna. Flores largamente pediceladas, dispuestas en forma

horizontal u oblicua en el pedicelo, perfectas, actinomorfas; bractéolas 2, alternas, subopuestas a opuestas,

sepaloides, orbiculares, obtusas a emarginadas o deltoides, persistentes, sosteniendo el cáliz o alejadas

de éste varios milímetros; sépalos 5, imbricados en dos series, suborbiculares a orbiculares, persistentes;

pétalos 5, libres o connatos basalmente, reflexos en la antesis; estambres generalmente numerosos (más de

10, aunque muy raramente puede haber sólo 5), en uno o varios verticilos, filamentos lineares a aplanados,

libres o basalmente connatos y/o adnatos a la base de los pétalos, anteras introrsas, basifijas a subbasifijas,

Giraldo-Cañas, Estudio taxonómico para las especies sudamericanas Schwartzi 695

subcordatas o subsagitadas; ovario súpero, globoso a cónico o piriforme; estigma mamiforme, lobulado o

radiado, subsésil. Frutos capsulares, coriáceos, globosos, apiculados; semillas reniformes, falcadas, semilu-

nares, elípticas u oblongas, reticuladas, negruzcas, brillantes, numerosas, raramente escasas.

Etimología.—Género dedicado al botánico sueco D. Schwartz (Vasconcellio Souza 1881).

Observaciones. —Con relación a la venación secundaria, Gentry (1993: 595) anotó que en todas las

especies de Schwartzia ésta es suprimida, lo cual no coincide con mis observaciones, ya que en este género

puede haber especies con venación secundaria débil o en ocasiones, suprimida en una o en ambas caras.

Cabe destacar que en vista del gran tamaño de las estruct

florales de las especies del género Schwartzia,

se elimina toda posibilidad de polinización entomófila, pues los insectos, incluso los más grandes, no po-

drían simultáneamente alimentarse BEN néctar Ja a vez tocar oe LE OS jaan una visita

(Pinheiro et al. 1995). Por lo tanto, los ú

de pájaros, murciélagos u otros pequeños janie ése Gentry 1991; Dene 1992: Savina et al.

1993, Pinheiro et al. 1995, Arruda et al. 2007). No obstante, las infl 1

Y 1sitadas

especies de abejas, avispas, hormigas y mariposas.

CLAVE PARA SEPARAR LAS ESPECIES SUDAMERICANAS DE SCHWARTZIA

1. Nect product bif , cuculados, coclearif ciatiformes o ventricosos.

2. Nect product cuculados o cocleariforme ar de Brasil 4. S. brasiliensis

2.N | bif: ciatiformes o ventricosos.

3: Doce estambres por nol Plantas de Ecuador 11. S. pterosara

3. flor. Plantas de Col de las Antillas M y el norte de Venezu

ela (Estado Sucre).

. 48 estambres por flor. Plantas de colomnle 10. S. parrae

= 22 estambres por nor Plantas de las Antillas M y el norte de Venezue la (Estado Sucre) __ 13. S. spiciflora

N , tubulares, globosos o ad

5, Bractéolas deltoides.

alo os de 2 8 p.d J I a l | J Z A | ] HN L

ubicadas a 14n mm del cáliz; a 12 pares de alándulas laminales por hoja. Plantas de Bras] |^ 1. S. adamantium

. Sépalos de 1,8-2 mm de longitud; pétalos de 4,7-5,2 mm de longitud; bractéolas opuestas, soste-

niendo el cáliz; 2-7 pares de glándulas laminales por hoja. Plantas de Brasil

7. S. jucuensis

7 Hojas con venacié ER H Ted ws AAA GEN EE Smeg £ | O Beara eee te

8. Hojas con venac ión daria suprimida en ambas caras Plantas de Brasil 6. S. geniculatiflora

8. | lojas con ¡ión secun daria si ¡primi ida sólo en el envés.

Dix ma ] I de + ] LE ]

10. Estambres 20, adnat la | d lo de 4-6,5 de | itud: 4-9

de glandulas laminales por hoja. Plantas de Colombia

10. Estambres 13, libres; pedúnculo de 10-14 cm de longitud; 15-24 pares de glándulas lamina-

les por hoja. Plantas de Perú 9. S. magnifica

9. Boca de los nectarios reproductivos con las márgenes conspicuamente revolutas. Plantas de

2. S. andina

Bolivia y Pe Per 14. S. weddelliana

vs el Va ua MU uy

E 1. Hojas ede ias emarginadas Plantas de Colombia 3. S. antioquensis

11. g obtusas

12. Nectari | lucti | ical T; tubulares o gl | I] .

I E I: I AM Pe J T% L 95

13. Sépalos de (4-)5-7 mm de longitud; bractéolas de 2,5-5 mm de longitud; estambres 50;

ovario de 3-8 mm de longitud; fruto de 14-17 mm de diámetro; urnas de los nectarios

fepioductives peaicelares ge 15- mm | de longitud con el pedículo de 1-2 mm de lon-

it | ladas, pecíolo de 3-10 mm de longitud. Plantas

de Colombia y Ecuador 558.

. Sépalos de 2,2-2,8 mm de longitud; bractéolas ca. a mm mias t a ca. 25;

ovario ca. 2,5 mm de longitud; fruto ca. 3 e

ductivos pedicelares de 9-17 mm de m con el pedículoc Ca. jd mm de longitud; hojas

conspicuamente pecioladas, pecíolo de 9-32 mm de longitud. Plantas de Colombia y

chocoensis

I |] .£aL s A rn LI Medion nidis iT.

696 Jou t Texas 3(2)

Ecuador 8. S. lozaniana

con la boca: no rostrada; estambres 70. Plantas de Colombia 12. S. renvoizei

1. Schwartzia adamantium (Cambess.) Bedell ex Gir.-Cañas, Hickenia 3(32):121. 2002. (Fig. 1). Norantea

adamantium Cambess., Fl. Bras. Mer. O aes 1:312, t. a 1828. ae iaa Steud., Nomencl, bot. ed. 2, 1:145. 1840,

nom. superfl. Tiro: BRASII I amantes, St. Hilaire s.n. [noLortro: MPU; IsoriPOS:

Fl, P véase Bedell (1985: 143).

Arbustos escandentes, muy ramificados, postrad ] bentes, hasta de 4(—7) m de altura, terrestres

o rupícolas; tallos cilíndricos a subteretes, corteza castaño, estriada longitudinalmente. Hojas sésiles a sub-

sésiles o cortamente pecioladas, cuando pecioladas entonces el pecíolo engrosado, recurvado, canaliculado,

levemente estriado, castaño oscuro, din i mm mee inh e láminas ovadas a tas, de 3,5-10,5 cm de

¡Sagra x 2-7,5 cm de ancho, d la y ápice obtuso y/o dimi te emarginado

7 1 1 . EL CREER. AA 1 iE

con | tas; 710 medio conspicuo en ambas

ed maaan ata A Ks PR ne

te elevado en el envés, liso;

su poción distal plano en la haz yl

caras; 3—8(—12) pares de glándulas laminales en la superficie abasial dispuestas en una fila más o menos

regular a ambos lados de la cara abaxial, distanciadas 5-11 mm de la margen, elipticas, conspicuamente

distanciadas entre sí, ca. 1 mm de largo x 0,5 mm de ancho, con su orificio eliptico, negruzco. Inflorescen-

cia terminal, racemosa, multiflora, levemente laxa a más comünmente densa; pedunculo de 4—14 cm de

longitud; pedicelos de 3,5-8 cm de longitud, levemente curvos a más frecuentemente rectos, oblicuos con

relación al eje de la inflorescencia, lisos a ligeramente estriados, castafio oscuros. Nectarios reproductivos

pedicelares sacciformes a tubulares, péndulos, pedículos de los nectarios de 1-6 mm de longitud, urnas de

5-18 mm de longitud x 3-7 mm de ancho, con la boca circular dispuesta hacia el eje de la inflorescencia y

con las márgenes revolutas y generalmente rostrada; los nectarios naciendo en el cuarto o quinto proximal

del pedicelo. Flores 18—70; bractéolas sepaloides, subopuestas a alternas, deltoides, carnosas, de 2 mm de

longitud, ubicadas a 1-4 mm del cáliz; sépalos orbiculares, carnosos, de 2 = ,6 mm de Mdb pétalos

libres, oblongos a ovados, carnosos, de 6-9 mm de longitud x 4-5 mm d ; estambres ca. 20, filamentos

ca. 3 mm de longitud, aplanados, anchos en la base y angostos en su ápice, lisos, libres, anteras oblongas,

subsagitadas, amarillentas, ca. 2,5 mm de longitud x ca. 0,7 mm de ancho; ovario cónico, de 4—5 mm de

longitud; estilo cónico, ca. 1 mm de longitud; estigma mamiforme, negruzco. Frutos globosos, apiculados,

ligeramente rugosos, de 12-14 mm de diámetro; semillas oblongas a reniformes, de 2,53 mm de longitud,

Hines puedas negruzcas, brillantes, numerosas.

t n áfica y ecologia.—Sch

oeste E Brasil; se tienen registros de los estados de Bahia, Goiás, Minas Gerais y el Distrito Federal. Esta

rece en cerrados, serranías, matorrales de suelos arenosos

1.1 : s eml

Es A A PARO

Mir LAA

e en áreas boscosas, entre los 500 y las 1500 m.

Observaciones.—Schwartzia adamantium puede ser confundida con la Eee n p S. brasiliensis.

Sin embargo, éstas pueden diferenciarse fácilmente por los nectarios rme

tubulares en S. adamantium vs. cuculados a cocleariformes en S. brasiliensis, por la longitud de los pecíolos

(0-3 mm en S. adamantium vs. 5-15 mm en S. brasiliensis), por la longitud de los pedicelos (3,5-8 cm en

S. adamantium vs. 1,5—4 cm en S. brasiliensis), por la forma de las bractéolas (deltoides en 5. adamantium

vs. ovadas a oblanceolas en S. brasiliensis), por la longitud de los sépalos (2,8-3,6 mm en S. adamantium

vs. 1,5-2,1 mm en S. brasiliensis), por la longitud de los pétalos (6-9 mm en S. adamantium vs. 5-6 mm en

S. brasiliensis) y por el tamafio de las semillas (2,5-3 mm de longitud en 5. adamantium vs. 5-6 mm en S.

brasiliensis).

E i inados: BRASIL. Bahia: Mun. Rio de Contas, Pico das Almas, vertente este, 13-14 km ao NO da cidade, 1180-1200

m, 28 oct 1988, R. Harley et al. 25726 (NY). Mun. Agua Quente, pico das Almas, vertente norte, vale ao noroeste do Pico, 1400 m, 1 dic

1988, R. Harley et al. 26538 (NY). Mun. Agua Quente, pico das Almas, vertente oeste, t ilho a Rosa, 23 km ao O da

cidade, 1200 m, 1 dic 1988, R. Harley & N. Taylor 27042 (NY), valley of the Rio das idas Cerrado on slopes of the Espigáo Mestre,

ca. 8 km NW of Barreiras, road to Santa Rita da Cassia, 600 m, 3 mar 1971, H. Irwin et al. 31442 (NY). Distrito Federal: Brasília D.F.,

Giraldo-Cañas, Estudio t jmico para | i lameri Schwartzi 697

aui

(

I MO "Tu L ADA r:

Fic. j

(H. Irwin et al, 9778).

Cañac A Rama florifara R Botón floral tari lIucti licelar. C. Flor en antesis

| n

Córrego Pipiripau, afluente do Rio Sao Bartolomeu, 8 ago 1980, E. Heringer et al. 5402 (US). Goiás: Serra dos Cristais, 5 km by road

E of Cristalina, 1200 m, 5 abr 1973, W. Anderson 8209 (NY); Chapada dos Veadeiros, 23 km de Alto Paraiso em direção a Teresina de

Goiás, Parque Nacional da Chapada dos Veadeiros, Cruzeiro, 1410 m, 12 nov 1996, M. Aparecida da Silva & A. dos Santos 3201 (IBGE);

J F a 1: 2A] J + ra | ya , Alt I5] s J i m 1990,

Parque Nacional da Chapada dos Veadeiros,

698 || | | £ al Dag - Im LJ dee ndis f Texas 3(2)

T. Barbosa Cavalcanti et al. 665 (SP); Alto Paraíso de Goiás, Parque Nacional da Chapada dos Veadeiros, no caminho para a Cachoeira

do Rio Preto, 20 nov 1987, I. Cordeiro et al. 434 (COL). Mun. Alto Paraiso, Fazenda Santo António, 6 jul 1996, J. Felfili et al. 354 (IBGE);

Cristalina, 12 jul 1988, T. Filgueiras & D. oe 1427 (IBGE); Parque Nacional Chapada dos Veadeiros, Alto Paraiso-Cruzeiro, 1126

m, 28 sep 1995, M. Fonseca et al. 657 (IBGE), 6 dic 1988, M. Pereira Neto 106 (IBGE); Serra de Biribiry, 7-10 km W of Diamantina, 1200

m, 15 jul 1965, S. Glassman & J. Gomes Jr. 8154 (F, SP); Estrada Alto Paraiso-Teresina, 10 oct 1979, E. Heringer et al. 2357 (IBGE, uie

Alto da Serra de Jaraguá, 30 sep 1974, E. Heringer 14026 (IBGE); Chapada dos Veadeiros, ca. 12 km NW of Veadeiros, 1200 m, 19 o

d H. Irwin et al. dauid bi Mun. Cristalina, Serra dos Cristais, 5 km S of CE 1175 m, 1 nov 1965, H. Irwin et al. 9718 o

istali mar 1966, H. Irwin et al. 13353 (NY d ineus, ca. 20 km

NW of C ba de G 1 to Niquelandia, ca. 1400 m, 28 ene 1968, H. iun et al. 19355-A (NY); Serra dos Pireneus, ca. 20

km E of Pirenópolis, 1000 m, 14 ene 1972, H. Irwin et al. ed ds NY). Mun. oe m dos Pirineus, 18 oct 1956, L.B. S. &

A.M. 4803 *Herbario A. Macedo" (US-2248310 P como A. Macedo & L.B. Smith 4803); 21

jul 1955, A. Macedo 3853 (MO). Mun. Posse, im 16 da estrada Posse- CRM de Goiás, via serra, 10 ago 1994, B. Pereira & D. Alvarenga

2632 (COL, IBGE); C lina, ca. 8 km da cidade, estrada para Paracatu, 1050 m, 4 feb 1987, J. Pirani et al. 1586 (SP); Pirineus, sin fe-

cha, Rizzo 5890 (NY), 4 sep 1971, Rizzo 6713 (NY); Cristalina, 1250 m, 15 oct 1960, Colector (?, letra ilegible) 7767 (NY). Minas Gerais:

Serra do a ca. i oa 7 i 2 ae Diamantina on road to ee 1400 m, 10 abr 1973, W. Anaercon 8517 (NY); Serra do

M nhac y roa Jeq tina, 1150 m, 14 abr 1973, W. And 8890 (NY); 20

j I! da Mata, 17 may 1977, P. Gibbs et al. 5268 (NY). Mun. Diamantina, Estrada para

Mendanha, a 15 km a Diamantina, 1200- 1225 m, 31 oct 1988, R. ee et al. 25479 (NY); Córrego do Tigre, Mun. Gouveia, campo

rupestre, 14 sep 1985, G. oe & F. Zelma 49657 (MO); Entre Brasília e João Pinheiro, 30 ago 1979, E. Heringer & C. Rizzini 17432

(IBGE, MO, NY, US); 25 km W of Diamantina on road to Conselheiro Mata, 18 ene 1959, H. Irwin 2482 (NY, US); Serra do Espinhaco

at tapaba, ca. 18 km N ee Sérro on road (MG 2) to Diamantina, 1200 m, 24 feb 1968, H. Irwin et al. 20754 (NY); Serra do Espinhaco,

ca. 15 km NE of Diamantina, on road to Mendanha, 1300 m, 26 ene 1969, H. Irwin et al. 22582 (NY); Serra do Espinhaco, ca. 16 km

W of Gráo Mogol, 950 m, 17 feb 1969, H. Irwin et al. 23452-A M Mun. Gouveia, campo rupestre, 1100 m, 3 jun 1985, H. Leitdo et al.

17194 (IBGE) ; 49 miles from Diamantina, ca. 1200 m, 22 dic 1959, B. Maguire et al. 44739 (NY); on sandstone just west of Serra, road

from Concencáo to Diamantina, 9 AM Eun B. bis and et al. 49129 25 49138 (NY). Mun. Diamantina, 19 dic 1979, G. Marninelli 6272

(MO); Diamantina, E Mata, km 174, 4 feb 1987, S. Mayo et al. s.n. (NY); Grao-Mogol, Trilha da Tropa, no atto

da serra, ca. 1000-1200 m, 11 dic 1989, P. Sano et al. 12433 (MO).

2. Schwartzia andina Gir.-Cañas, Caldasia 25:5. 2003. (Fig. 2). Tiro: COLOMBIA. Antioquia: Mun. Valdivia, vereda

San Fermín, Alto de Ventanas, 145 km al NE de Medellin, finca La Soledad, 2000 m, 7 ene 1994, R. Callejas & A. Bornstein 11073

(HoLoTIPO: HUA!; isortros: COL!, MOD.

Arbustos escandentes, epífitos a hemiepífitos; tallos cilíndricos, angulosos en las porciones distales de las

ramas, levemente estriados a lisos, corteza castaño claro a negruzca. Hojas cortamente pecioladas; pecíolo

recurvado, negruzco, de 3—7 mm de longitud x ca. 3 mm de ancho; láminas oblanceoladas a oblongas, de

(8311,5-19 cm de longitud x (2,8-)4-6,5 cm de ancho, de base cuneada y ápice ligeramente apiculado,

márgenes ligeramente revolutas; nervio medio conspicuo en ambas caras, plano a levemente hundido en

la haz y ligeramente abultado en el envés, liso, ca. 3 mm de ancho en su porción basal, desvaneciéndose

is ue la región distal de la lámina; venación secundaria apenas perceptible en la haz (raras

eces suprimida), siempre suprimida en el envés; 4—9 pares de glándulas laminales en el envés, localizadas

a 5— 10n mm de la — cados entre sí 0,8-3,5 cm, elípticas, 1-1,8 mm de largo x ca. 0,4 mm de

ancho, con su orificio elíptico, negruzco, de 0,8—1,4 mm de largo x ca. 0,25 mm de ancho. Inflorescencia

terminal, cortamente racemosa, multiflora, densa; pedúnculo de 4—6,5 cm de longitud; pedicelos de 5-6,5

cm de longitud, cilíndricos, rectos, ensanchándose en su porción distal, lisos a levemente estriados o con

pocas y sobreelevadas verrugosidades sólo en su extremo distal, cobrizos a castaño oscuros. Nectarios re-

productivos pedicelares sacciformes, tubulares, péndulos, verde claros a purpúreos, reticulados, pedículos

de los nectarios 1-2,3 mm de longitud, urnas de (7110-12-14) mm de longitud x 3,3-)4-5(-6) mm de

ancho, boca rostrada, con el orificio elíptico a circular, de 1,7-2,8 mm de ancho, dispuesta hacia el eje de la

inflorescencia y ocupando una posición transversal respecto a su eje, los nectarios insertos por debajo del

1/3 proximal del pedicelo. Flores 16-26, inserción pedicelo-flor horizontal a levemente oblicua; bractéolas

sepaloides, opuestas, orbiculares, — rugosas, carnosas, de oat 8-5 AS, 5) mm de d sépalos

orbiculares, rugosos, carnosos, de 5,3—6,6 mm de longitud, con | las y translúcidas; pétalos

connatos basalmente ca. 2/5 de su a ovados, os en la "n de 9- 11 mm de longitud x ca. 4

mm de ancho, carnosos, verde claros; estambres 20, filamentos ca. 4 mm de longitud x ca. 0,6 mm de ancho,

Giraldo-Cañas, Estudio taxonómico para las especies sudamericanas Schwartzi 699

Fic. 2. Sch val lina Gir -Canas. A. Rama florifera. R. Botón floral tari jucti liralar €. Ovario (R Callejas 11073)

aplanados, lisos, adnatos a la base de los pétalos, anteras oblongas, subbasifijas, ca. 3 mm de longitud x ca.

1 mm de ancho; ovario globoso a cónico, rugoso, de 5-6,5 mm de longitud x 3,5-5 mm de ancho; estigma

mamiforme, subsésil, negruzco. Frutos desconocidos.

Distribución geográfica y ecología.—Schwartzia andina crece en bosques premontanos a montanos bajos,

húmedos a pluviales y en bosques de niebla, entre los 1700 y los 2400 m. Esta especie sólo ha sido regis-

trada en la región andina colombiana, en los departamentos de Antioquia, Caldas y Cauca. De esta especie

700 tani itute of Texas 3(2)

sólo se conocen unas cuantas colecciones, las cuales proceden de áreas que están siendo deforestadas y/o

transformadas drásticamente por el hombre, lo que podría significar que la especie se encuentra en peligro

bservaciones.—Schwartzia andina si ta el envés foliar con la venación secundaria suprimida.

Por otra parte, ] presenta unas verrugas dbultadas en la porción distal de los pedicelos.

F dos: COLOMBIA. Antioquia: Mun. Briceño, vereda San Fermin, via Ventanas-Briceño, km 2.5, 2100-2300 m,

n nov 1987, R. Callejas et al. 5569 (HUA, NY). Mun. Urrao, corregimiento La Encarnación, camino hacia el Parque Nacional Natural

Las Orquídeas, primera loma, 2200—2400 m, 8 feb 1995, J. Pipoly et al. 18621 (MO). Caldas: Manzanares-Pensilvania, 1750 m, 5 oct

Pis ii Mie & M. i Fraume 264 aaa Cauca: Mun. El Tambo, corregimiento 20 de Julio, carretera abandonada, cerca de 2 km

el INDERENA, iento Tuntas, limite con el Parq ional Natural Munchique, 2200

m, 6 ene 1991, R. Ruiz et al. 1201 (MO)

3. Schwartzia antioquensis Gir.-Cañas, Caldasia 25:7. 2003. (Fig. 3). Tiro: COLOMBIA. Antioquia: Mun. Urrao,

vereda Calles, río Calles, Parque Nacional Natural Las Orquideas, 1400-1500 m, 4 may 1995, R. Fonnegra & Grupo de Palinología

5569 (noLorIPo: COLI; isoriros: COL!, HUA!, MOD.

1 ? sall ax |Z

ificad , hemiepifitos, ndricos, levemente

Arbustos lianescentes, [

angulosos en las porciones distales de las ramas, huecos, corteza fuertemente "MÀ y S did i estrías

tranversales. Hojas pecioladas, pecíolo recurvado, canaliculado (el canal muy angosto y p ), leve-

mente estriado, castaño oscuro, de (8—10—13(-16) mm de longitud x 3-6 mm de ancho; Vues bodas

de 9-14,5 cm de longitud x 4,5—7(-8) cm de ancho, de base redondeada y ápice levemente mucronado, con

márgenes ligeramente revolutas; nervio medio conspicuo, liso o levemente estriado y abultado en el envés,

apenas perceptible y plano por la haz, desvaneciéndose hacia la porción distal de la lámina, ca. 2 mm de

ancho en su porción proximal; venación secundaria levemente impresa en la haz, un poco más fuerte en el

envés; 6-12 pares de glándulas laminales en la superficie abaxial, submarginales, localizadas a 1-3 mm de

la margen, distanciadas entre si 0,7—3 cm, circulares, ca. 1 mm de diámetro, con su orificio circular a elip-

soide, negruzco, ca. 0,5 mm de diámetro. Inflorescencia terminal, racemosa, multiflora, densa; pedúnculo

de (5,528—12(—14) cm de longitud; pedicelos de 6-8,5 cm de longitud, cilíndricos a levemente angulosos,

lisos a estriados, ocasionalmente verrugosos, ensanchándose en su porción distal. Nectarios reproductivos

pedicelares sacciformes, tubulares, péndulos, verdes a rojizos, rugosos, carnosos, pedículos de los nectarios

de 1-3 mm de longitud, urnas de 24-32 mm de longitud x 8-11 mm de ancho, boca circular de 4-5 mm

de diámetro, dispuesta hacia el eje de la inflorescencia y ocupando una posición oblicua respecto a su eje,

los nectarios insertos por debajo del 1/3-1/4 proximal del pedicelo. Flores 18-24, inserción pedicelo-flor

horizontal; bractéolas sepaloides, opuestas, orbiculares, emarginadas, carnosas, rugosas, de (3-)4-5,5

mm de longitud, con las márgenes delgadas y translúcidas; sépalos orbiculares, rugosos, carnosos, con las

márgenes delgadas y translúcidas, de (6,5—)8-10C-11) mm de longitud; pétalos connatos basalmente ca.

2/5 de su longitud, ovados, carnosos, reflexos en la antesis, de 7-9 mm de longitud x ca. 4 mm de ancho;

estambres ca. 22, filamentos ca. 2 mm de longitud x ca. 0,7 mm de ancho, aplanados, lisos, adnatos a la base

de los pétalos, anteras subcordatas, oblongas, blanquecinas, ca. 2,5 mm de longitud x ca. 1 mm de ancho,

ovario globoso a piriforme, rugoso, verde claro a amarillento, de 7-12 mm de longitud x 11-13 mm de an-

cho; estigma mamiforme, subsésil. Frutos globosos, carnosos, apiculados, rugosos, rojos cuando maduros,

de 18-25 mm de diámetro; semillas elípticas, ca. 4,5 mm de longitud x ca. 2 mm de ancho, reticuladas,

negruzcas, brillantes, numerosas.

Distribución geográfica y ecología.—Schwartzia antioquensis crece en bosques húmedos a pluviales y en

bosques de niebla de áreas premontanas, entre los 1200 y los 1700 m. Esta especie sólo se conoce de unas

cuantas colecciones de la región de Murrí en el municipio de Frontino y de la vereda Calles del Parque

Nacional Natural Las Orquídeas unoque SIME

Observaciones.—En vista de su hábit „S$ is es fácilmente confundida con S. chocoensis

No obstante, estas especies pueden separarse Belmene a la forma de las bractéolas (emarginadas en S.

antioquensis vs. obtusas en S. chocoensis), el número de estambres (ca. 22 en S. antioquensis vs. ca. 50 en 5.

chocoensis), el tamaño del ovario (7-12 mm en S. antioquensis vs. 3-8 mm en S. chocoensis), la boca de los

701

IP.

Giraldo-Cañas,

A Rama florifera. B. B

lanae

. Il

Fic. 3. Seh

C Ovarlo (R Fonneara et al 5569)

[| Il £a n.a LI 1m 1 | d. nnd. ET Sia

* ,

702

nectarios reproductivos pedicelares (no rostrada en S. antioquensis vs. rostrada en S. chocoensis) y las dimen-

siones del fruto (18-25 mm en S. antioquensis vs. 14—17 mm en S. chocoensis). Véanse las observaciones dadas

para S. renvoizei.

EC Pi J Sii AARAA DITA A 4 . AA Frontino, road to Murrí, 22 ^] km W [WT ;1 ] 3 LA z , 1330 m,

19 oct 1987, A. Brant & G. Martinez 1413 (HUA MO) Mun. Frontino, il lta del río C , 1560 m, 10 jul

1986, D. Sánchez et al. 282 (COL, MEDEL, MO, NY). Mun. Frontino, 22 of road Notibara: Murri, 1290 m, 21 sep 1987, J. Zarucchi et

al. 5512 (COL, CHOCO, MO, NY). Mun. Frontino, km 18 La Blanquita, región de Murrí, 1620 m, 4 nov 1988, J. Zaruc-

chi et al. 7117 (COL, HUA, MO). Mun. Urrao, Parque Nacional Natural Las Orquídeas, sector Cabaña de Calles, quebrada La Agudelo,

1300-1400 m, 31 mar 1992, D. Cárdenas et al. 3171 (MO). Mun. Urrao, vereda Calles, Parque Nacional Natural Las Orquídeas, margen

derecha del río Calles, NW de la Cabaña de Calles, 1450 m, 6 dic 1993, A. Cogollo et al. 7891 (MO). Mun. Urrao, Las Orquídeas, vereda

Calles, río Calles, Parque Nacional Natural Las Orquídeas, b den al NW de la cabaña Calles, 1340 m, 7 dic 1992, J. Pipoly et

al. 16715 (MO), 1 dic 1993, J. Pipoly et al. 17484 (MO), 9 dic 1993, J. Pipoly et al. 18034 (MO).

4. Schwartzia brasiliensis (Choisy) Bedell ex Gir.-Cañas, Caldasia 23:341. 2001. (Fig. 4). Norantea brasiliensis

Choisy in DC., Prodr. 1:566. 1824. Tiro: BRASIL: Rio de Janeiro, año 1814, Langsdorff s.n. (HoLotIpO: P isoripo: Fl, fotografía del

holotipo en F! y MOD.

Schwartzia glabra Vell., Fl. Flumin. 5:221. 1825 [1829]. Tiro: BRASIL: Vellozo s.n. [HoLotipo: no localizado, probablemente en MEL,

véase Bedell (1985: 142)].

Ascium selloi Spreng., Syst. Veg. 2:599. 1825. Norantea selloi (Spreng.) G. Don, Gen. Hist. 2:599. 1825. Tiro: BRASIL: Sello 178 [HoLorIPO:

W, véase Bedell (1985: 134)].

Arbustos escandentes, epifitos a más comúnmente terrestres, profusamente ramificados, ramas decumben-

tes hasta de 8 m de altura, tallos subteretes, lisos a levemente estriados longitudinalmente, corteza castaño

a ferrugínea. Hojas pecioladas; pecíolo recurvado, canaliculado, ligeramente estriado, castaño oscuro, de

5-14-15) mm de longitud x ca. 2 mm de ancho; láminas ovales, obovadas a oblanceoladas, de 5-16 cm de

longitud x 3-6 cm de ancho, de base redondeada a cuneada y ápice obtuso, emarginado, retuso o mucro-

nado, con márgenes planas o ligeramente revolutas y engrosadas; nervio medio conspicuo en ambas caras,

desvaneciéndose hacia su porción distal, plano o levemente hundido en la haz y abultado en el envés, liso

a estriado longitudinalmente; venación secundaria débilmente impresa en ambas caras; 3-20 pares de

gándulas laminales en la superficie abaxial, elipsoides, de 0,5-1 mm de longitud x 0,25—0,5 mm de ancho,

unas submarginales, a 1-2 mm de la margen, otras distanciadas a 5-16 mm de la margen y dispuestas en

filas más o menos uniformes a ambos lados del envés, aunque también puede haber glándulas laminales

distribuidas en forma irregular y próximas a la parte media del envés. Inflorescencia terminal, racemosa,

multiflora, laxa a densa; pedúnculo de (9-)13-35 cm de longitud; pedicelos de (1,5-)2-3,4(-4) cm de

ongia. rectos a Eee CUrvos, pi a ee estriados ope eee castaño oscuros.

, rojos, pedículos de los

nectarios de (Q-)3- 6(-7) mm de longitud, urnas de 5-11 mm de longitud x 4—/ mm de ancho, dispuestos

hacia el eje de la inflorescencia; los nectarios naciendo en el tercio, cuarto o quinto inferior del pedicelo.

Flores 38-160; bractéolas sepaloides, subopuestas o alternas, ovadas a oblanceoladas, 1-2 mm de largo x

ca. 1 mm de ancho, sosteniendo el cáliz o distanciadas 3—4(—6) mm de éste; sépalos orbiculares a suborbi-

culares, carnosos, de 1,5-2,1 mm de longitud; pétalos libres, ovados a oblongos, reflexos en la antesis, 5-6

mm de longitud x 2,5-3 mm de ancho, carnosos; estambres (1418-22, filamentos ca. 1,5 mm de longitud,

aplanados, haciéndose más angostos hacia su porción distal, lisos, adnatos a la base de los pétalos, anteras

introrsas, oblongas a ovales, subcordatas, ca. 2 mm de longitud x ca. 1 mm de ancho; ovario piriforme, liso

a débilmente surcado, 2-3 mm de longitud; estigma mamiforme, subsésil, negruzco. Frutos globosos, api-

culados, levemente rugosos, de 7-12 mm de diámetro; semillas semilunares, 5-6 mm de longitud x 1-1,5

mm de ancho, rugosas, reticuladas, negruzcas, brillantes, numerosas.

Distribución geográfica y ecología. —Schwartzia brasiliensis es endémica de la región oriental de Brasil

(Bahia, Distrito Federal, Minas Gerais, Paraná, Pernambuco, Rio de Janeiro, Rio Grande do Norte, Santa

Catarina y São Paulo) y crece en manglares, matas, matas de restinga, bordes de mata de restinga, aflora-

mientos rocosos y en campos rupestres, entre el nivel del mar y los 1000 m.

PNE ómico para las especi ; : "T — 703

Giraldo-Cañas,

RY,

ce A

to Ye

U AN /

lem

Fic. 4. Sct tzia brasiliensis (Choisy) Bedell ex Gir. -Cañas. A. Rama florifera. B. Fl tari lucti licelar. (Y. Pirani & 0. Yano 561)

I 1 Fal n.a H Im LI rr 3/5)

704 Jou 0 f Texas 3(2)

Observaciones.—Schwartzia brasiliensis es la especie que presenta las inflorescencias mas largas y las

semillas mas grandes entre todas las especies del género. Por otra parte, las inflorescencias de esta especie

son visitadas por anon a y peque Andas em cee y su polinización se da por algunas especies de

aves (Sazima et al. 1993). Véanse las observaciones dadas para nial Cabe destacar que el binomio

Norantea bahiensis Mart. ex Wittm. (Fl Brasil. 12:245. 1878), es que ponde ala misma

S. brasiliensis.

Especímenes examinados: BRASIL. Bahia: Região Cacaueira, Mun. Valença, estrada Valenca-Guaibim, km 10 ap., 8 ene 1982, A. de

Carvalho & G. Lewis 1119 (US); on the coast road between Alcobaca and Prado, 10 km NW of Alcobaca, and 4 km N along road from

the Rio Itanhentinga, sea level, 15 ene 1977, R. Harley et al. 17940 (NY, dos ejemplares). Mun. Maracas, 8 a 18 km ao S de Maracas, pela

"RÀ rodovia para Jequié, iia de Mata de Cipó, 900—1000 m, 15 feb 1979, T. dos Santos et al. 3481 (SP); Sin localidad y sin fecha,

fio 1841, Martius 1048 (MO, NY). Distrito Federal: Praia de — Recreio dos Bandeirantes, sea level, ^ abr Eu E E

et al. 6407 UN. ue) bow erue Rio Preto, 16 km da praca centra locali

Gabriel 5 N , divisa com Lima Duarte, 820 m, 26 feb 1999, R. Mello-Silva et al.

1644 (SP). Paraná: pw ad marginem PM 12 may 1909, P Dusén 8116 (MO, NY, US); Jacarahy, 24 mar 1911, P. Dusén 18196

(SD; swampy brejo, N of Río Nunes, N of Morretes, 40 m, 24 ene 1985, A. Gentry & E. Zardini 49801 (MO); Paranagua, Matinhos, 7 mar

951, G. Hatschbach 2149 (SI). Mun. Paranaguá, Rio Cachoeirinha, 10 m, 19 jun 1970, G. Hatschbach 24409 (NY). Mun. Guaratuba, rio

Quiririm, mata pluvial, 50 m, 1 abr 1973, G. Hatschbach 31796 (COL). Mun. Guaraquecaba, Morro do Quitumbe, 9 feb 1994, R. Lima

228 (NY); Guaratuba, restinga, 3 m, 21 feb 1952, P. Reitz 4369 (RSA). Mun. Guaratuba, low coastal forest, Guaratuba, 2 m, 21 feb 1952,

L. Smith & P. Reitz 5754 (US). Pernambuco: Sin localidad, años 1836-1841, G. Gardner 942 (NY, dos ejemplares); in a thicket, Tapera,

21 dic 1934, D. Pickel 3737 (F, NY). Rio de Janeiro: Restinga al S de Copacabana, 7 feb 1974, A. Burkart 30358 (SI); Rio de Janeiro,

restinga de Jacarepaguá, 0-10 m, 3 nov 1962, J. Cuatrecasas & A. Duarte 26633 (US); Rio de Janeiro, Recreio dos Bandeirantes, 7 feb

1964, W. Hoehne 5585 (COL); Recreio dos Bandeirantes, 30 km W - Rio - Janene, 5 mar 1964, K. Lems s.n. (NY); Furnas, nos blocos

de pedras, 19 dic 1928, A. Ducke 12801 (US). Mun. Maricá, resti j S. José do Imcagai, 23 nov 1983, C. Farney & E.

Handal 375 79 4E Mun. bed Frio, Massambaba, a 39 km paria de iguala ea 114 km do Trevo de Cabo Frio, 5 feb 1986, Fontellah et al.

2294( Om, 11 ene 1985, À ee et aes nn (MO); Pedra da Itauna, 9 oct 1977, P. Maas & P. Carauta 3135

(NY). Mun Para Na beira d di d Mine n n 5 feb 1989, R. e 206 (F, IBGE);

Macaé, entre Macaé e Rio das sitas: 15 may + 1993, R. Mello- Silva & J. Pirani 868 (?, letra ilegible), prox. Pedra do

panas, 2E ene 1977, P. Occhioni s.n (OGE 1020): imi ay of Angra ae Reis, 1 feb 1968, L Smith & E. McWilliams 15365 (US).

Along sand ta N , S of Natal, 8 sep 1946, J. Wurdack 165 (NY). Santa Catarina:

Mun. Ilhota, Morro do Baú, 30 ene 1964, E. Pereira & G. Pabst 8783 (NY): Brusque, 50 m, 23 feb 1950, P. Reitz 3359 (SI, i ejemplares);

Braco Joaquim, Luis Alves, Itajaí, 450 m, 16 feb 1956, Reitz & Klein 2712 (NY, US); Piloes, Palhoca, 350 m, 23 feb 1956, Reitz é Klein 2850

(NY, US); Salto de Piláo, Lontras, Rio do Sul, 300 m, Reitz & Klein 8626 (US). Mun. Pórto Belo, st ky woods, eastern shore

a nx João » Cunha, 1-5 m, 31 mar int E, a et E 12293 wn Isla do Francés, feb 1947, B. de dodi 42 eL dos ejemplares).

ntre ca. 600 m, 25 ene 1992, F idi P); "a ie

242 da rodovia Sd Paulo pwe 26 feb 1964, N. Cruz 86 (SP). Mun. Cananga. island of Cananéia, about 1 |

Cananéia, al g land, sea level, 17 feb 1965, G. Eiten & W. Clayton 6186 (MO, NY, SP: t l ). Mun baba

à 20 m da ponte do Rio Escuro, 9 dic 1961, J. Fontella 119 (SP, US); Cubatáo, 15 mar 1923, A. Gehrt 8223 (SP) São Sebastian, Bertioga,

10 nov 1976, P. Gibbs et al. 3513 (NY); Santos-Sáo Sebastian, km 40, rio Guaratuba, 1 abr 1976, S. Hallard 4 (NY); Sao Paulo, nativa no

Jardim Botánico, 10 feb 1935, O. Handro s.n. (SP: 32835, dos ejemplares). Mun. Santo André, Ribeirão Pires, 16 ene 1949, G. Hashimoto

586 (SP, US); Bertioga, bairro de Boracéia, Condomínio d be dn ca. km wae 5 E ips Rio-Santos, 24 mar 2000, M. Kirizawa

] Rin Dreta na pete ada p nara Rin an Bun

et al. 3406 (COL); Vilha Atlantica, 7 jui 1951, J. Morello 63 (S éia to do Rio Coruja, 29 nov

1966, J. Mattos & N. Mattos 14253 (SP); cultivada em pizseiedba: na Escola ala Luiz d d 26 mar 1934, P. de Oliveira Santos

s.n. (SP: 31673); Cananéia, mata de restinga, 25 feb 1983, J. Pirani & O. Yano 561 (COL); Ilha Anchie restinga, 10 e

1991, M. Robim 666 (COL); Sin fecha, H. Mosén 3160 bind Sin Estado y sin localidad, afios 1822— 1828, L. Riedel 831 (NY): Sin ae

sin localidad y sin fecha, Santos 75 (NY); Sin Estado, sin localidad y sin fecha, senate 137-159 (MO).

5. d chocoensis Gir.-Cañas, Revista Acad. Colomb. Ci. Exact. 25(97):478. 2001. (Fig. 5). Two:

Croco: Carretera Ansermanuevo-San José del Palmar, 8,4 km B Alto del Galápago, 1600 m, 19 feb 1977, E. Forero,

A. een Sugden & D. Daly 3000 (HoLotiro: COL!; isoriros: CHOCO!

C-I z ER TY & Bedell, in F & G + Ta ay tul l4] 1 t Jal A + ta nal 71 md A E

Į , Colombia. Biblioteca

José oe Triana 10:103. 1989, nomen mum

Arbustos escandentes, epífitos a hemiepífitos, profusamente ramificados, ramas robustas; tallos cilindricos,

huecos o fibrosos internamente, entonces las fibras lanuginosas, corteza castaño a ferrugínea, fuertemente

estriada y rugosa, las estrías tranversales. Hojas cortamente poa ped cea canaliculado (el

canal muy angosto y poco profundo, ca. 1 mm de ancho), levemente te liso, castaño

705

Giraldo-Cañas,

X i

o i >

A ML "iab

a Ds

E a dE

5cm

D. Ovario. (F

Cañas. A. Rama florifera. B. Bota

Fic. 5. Sct

Forero et al. 3000).

3 I ELl n.a a Im LI dis dien adis £'T ai”

706

oscuro, de (324—6(—10) mm de longitud x 5-6(-7) mm de ancho; lami lipti oblongas, de (12-)15-22

cm de ee x (3, a 5 EB 2) cm de ancho, d l t lata y api tamente apiculad

con y engrosadas, ae pruinosas en la ha nervio medio pica

en aii caras que poco notorio hacia la porción distal de la haz), plano a levemente hundido en la

haz y abultado en el envés, liso, de 1-2 mm de ancho; venación secundaria débil en ambas caras; 2—7 pares

de glándulas laminales en la superficie abaxial, submarginales, localizadas a 0,5—1,5(3) mm de la margen,

circulares, levemente elevadas, de ca. 0,8 mm de diámetro, con su orificio circular, negruzco, de ca. 0,15

mm de diámetro, distanciadas entre si 1,5—6(—7) cm. Inflorescencia terminal, racemosa, multiflora, densa;

pedúnculo de 5-30 cm de longitud; pedicelos de 4—5,8(—7) cm de longitud, rectos a ligeramente sinuosos,

lisos a levemente estriados, castaño oscuros. Nectarios reproductivos pedicelares sacciformes a tubulares,

péndulos, rojizos, estriados longitudinalmente, ligeramente rugosos, pedículos de los nectarios 1-2 mm de

longitud, urnas de (15220—33 mm de longitud x 5,5-10 mm de ancho, con los orificios circulares, de 1,5—3

mm de diámetro, dispuestos hacia el eje de la inflorescencia y ocupando una posición oblicua respecto a

su eje, el orificio usualmente con un pequeño apéndice en forma de espolón (rostrado) de 0,5-2,5 mm de

longitud; los nectarios naciendo en el tercio, cuarto o quinto inferior del pedicelo. Flores 5—54, inserción

pedicelo-flor horizontal a levemente n bractéolas sepaloides, opuestas, orbiculares, obtusas, rugosas,

carnosas, breves, de 2,5—5 mm de longitud; sépalos orbiculares, rugosos, carnosos, de 4—7 mm de longitud;

pétalos connatos basalmente ca. 1/6 de su a ovados, reflexos en la antesis, de 6-12(-13) mm de

longitud x 3,8-6 mm de ancho, carnosos, verde claros a Felizos; estambres 50, filamentos de 1-3 mm de

£a]

,anteras

longitud x ca. 0,5 mm de ancho, aplanados, levemente , adnatos a la b

oblongas, basifijas, subcordatas, blanquecinas, 3-5 mm de Iongitud x 0,5-1 mm de — ovario cónico,

ligeramente estriado, 3-8 mm de longitud x 2,5-6 mm de ancho; estigma mamiforme, subsésil, negruzco.

Frutos globosos, carnosos, apiculados, lisos, de 14-17 mm de diámetro, verde claros; semillas reniformes,

3-4 mm de longitud x ca. 1,5 mm de ancho, rugosas, negruzcas, brillantes, numerosas (ca. 30 por fruto).

Distribución geográfica y ecología.—Schwartzia chocoensis crece en bosques maduros, en manglares, en

bosques secundarios, en riberas de grandes ríos y en bordes de bosques de áreas muy húmedas a pluviales,

entre el nivel del mar y los 1600 m de altitud. Hasta ahora sólo ha sido recolectada en Colombia y Ecuador.

Observaciones. —Los nectarios reproductivos pedicelares de S. chocoensis producen una gran cantidad

de néctar, pues éste alcanza a llenar casi completamente el volumen de éstos (vistos en corte longitudinal).

Véanse las observaciones dadas para S. antioquensis y S. renvoizei.

3 ral CRRADTA Choci EP ES 3. Month 1 Pal 1 114. dJa] p—— ] LI d ,Cà 50 m, 9

"i 1982, L.. mE p Escobar et il cni HUA; Carretera Quibdó- Gaava, Diaa, margen derecha déls río Duatá, 40 m, 1,27 abr 1975,

E. Forero et al. 1293 MO, N da Togoromá, 13 jun 1944, E. Killip & : Cuatrecasas 39159 (US). Valle del Cauca:

vicinity of Bussssentiti: is Calima Region, pe Buenaventura and Río Calima, km 22 on main road to Canalete, ca. 6 km N of

main road, 50—60 m, 21 jul 1988, T. dus ied MOL PS Calima, ca. 100 m, 25 mar ue I. dis da s Bajo Calima, ca. 10

km due N of Buenaventura, Cartó I ?56'N-77*08'O, ca. 50

m, 7 dic 1981, A. Gentry 35421 (COL, MO); Bahía Málaga, mangroves, th of Quebrada Si APOIN- 779150, sea level, 17 feb 1983,

A. Gentry et al. 40423 (COL, MO); Bajo Calima, Dindo z area, 3°59'N-76°58'0, ca. 100 m, 11 jul 1984, A. Gentry et al. 47886 (COL, MO).

Mun. Buenaventura, quebrada San Joaquin, Concesión Pulpapel, 100 m, 7 may 1968, J. Idrobo et al. 6195 a dos ais ania Costa

Pacifica, Bahia Malaga, qom npe 2 e 1987, L. Millán 61 (CUVC); Río El Chanco,

Aguabonita, subi C] 400 m, 3 abr 1985, R. Ruiz et al. 94 (COL) ECUADOR. Pichincha: km 110

Quito-Nono-Tandayapa-Mindo-Puerto Quito, 1100 m, 1-24 jul 1982, C. Dodson & A. Embree 13091 (MO).

6. Schwartzia geniculatiflora Gir.-Cañas & Fiaschi, ien an 185. dud (Fig. e) Tipo: BRASIL. Banua:

Urucuca, 7,3 km north of Serra Grande on road to Itacaré, Fazenda Lagoa d t southern Bahian wet

forest, 14?25'24"S, 39°03'38"W, 15 nov 1995, WW. Thomas, A.M. de Carvalho, J. domar & G. Pitanga 11029 aoia: COLI; ISOTIPOS:

CEPEC!, COL!, NY).

Arbustos uel dud una ramas Gecumbentcs tallos cilíndricos a subteretes, corteza castaña

clara, estriada long ladas, pecíolo engrosado, recurvado, canaliculado,

liso a levemente estriado, castaño oscuro, hasta da 8 mm de longitud; láminas obovadas, de 5-8 cm de

longitud x 2,5-3,5 cm de ancho, de base redondeada a levemente asimétrica y ápice obtuso y/o diminuta-

Giraldo-Cañas,

707

Cañac & Fiacchi

ru L E Fm

A. Rama florifera. R. A

J H | fFuanSEF laf

Thomasetal 1 1029)

708 t tani titute of Texas 3(2)

mente mucronado, con márgenes planas o levemente revolutas; nervio medio conspicuo en ambas caras,

desvaneciéndose hacia su porción distal, plano en la haz y ligeramente elevado en el envés, liso; venación

secundaria suprimida en ambas caras; 1-4 pares de glándulas laminales en la superficie abaxial, ubicadas

en la parte media a distal de la lámina sin presentarse en la parte basal de la misma, dispuestas en una

fila más o menos regular a ambos lados de la cara abaxial, distanciadas 1-6 mm de la margen, elípticas,

conspicuamente ió entre sí, ca. 1,5 mm de largo x ca. 1 mm de ancho, con su mee eliptico,

negruzco. Infl inal, racemosa, pauciflora, laxa; pedúnculo de 6-6,5 cm de longitud; pedicelos

de 1,7-2 cm de longitud; rectos, casi PAS con relación al eje de la inflorescencia, lisos, castaño

oscuros. Nect leados, péndulos, sésiles, urnas de ca. 5 mm de longitud x ca.

4 mm de ancho, con la boca elpsoide ua hacia el eje de la infl ia; los nectarios naciendo en el

cuarto proximal del pedicelo. Flores 8-10, inserción pedicelo-flor oblicua; bractéolas sepaloides, opuestas,

orbiculares, carnosas, ca. 2,5 mm de diámetro, sosteniendo el cáliz; sépalos orbiculares, carnosos, ca. 3,5

mm de diámetro; pétalos libres, ovados, carnosos, ca. 5 mm de longitud; estambres 18, filamentos ca. 2,7

mm de longitud, aplanados, libres, anteras oblongas, subsagitadas, ca. 2,6 mm de longitud x ca. 1,5 mm de

G + re E E

ancho; ovario cónico, ca. 3,2 mm mm de longitud; estigma subsésil, mamiforme, negruzco

conspicuamente apiculados, rugosos, ca. 10 mm de diámetro, más la porción apiculada de ca. 5 mm de

longitud; semillas oblongas a reniformes, rugosas, reticuladas, negruzcas, brillantes, numerosas

Distribución geográfica y ecología.—Schwartzia geniculatiflora es endémica de los bosques húmedos del

sur de Bahia, E 010 ue la eae EDO

Obsen vIrTiOnec

ESCF VOACIOTICS,

"1,1 la L 1,1

reproductivo

pedicelar y, A n por ih -—" conspicuamente oblicua de la flor en el pedicelo, característica

poco rd en el oo encontrandose, aunque no tan notoriamente como en S. geniculatiflora, en las

ocoanas S. andina Gir.-Cañas, S. cl is Gir.-Cañas, S. parrae Gir.-Cañas y S. renvoizei

rca

7. d jucuensis Gir.-Cañas, Darwiniana 42:173. 2004. (Fig. 7). Tiro: BRASIL. Espirito Santo: Mun. Domingos

ns, Chapéu, Rio Jucu, Braço Norte, margens de rio, sobre rochas, 8 nov 1993, G. Hatschbach, M. Hatschbach & J. Silva 59728

a US!; otros: CEPEC, MBM).

A y] Getty peace d SEPT a +411 q oe ] ] ; EE Er P: Ar PR leve-

mente estriados; corteza cusan FM en las porciones proximale s de "m ramas, negruzca en las porciones

distales. Hojas pecioladas; pecíolo recurvado, canaliculado, ligeramente estriado, negruzco, de 10-23 mm

de longitud; láminas oboelípticas a obovadas, de 7-8,5 cm de longitud x 3,8—4,2 cm de ancho, de base

cuneada y ápice obtuso y retuso, con márgenes planas o ligeramente revolutas y engrosadas; nervio medio

conspicuo en ambas caras, desvaneciéndose hacia su porción distal, plano o levemente hundido en la haz y

abultado en el envés, liso a estriado longitudinalmente; venación secundaria débilmente impresa en ambas

caras; 1-7 pares de gándulas laminales en la superficie abaxial, circulares, ca. 1 mm de diámetro, submar-

ginales, a 0,5-3(-7) mm de la margen. Inflorescencia terminal, racemosa, multiflora, densa; pedünculo

de 4 cm de longitud; pedicelos de 3,9—4,8 cm de longitud, capilares, rectos, lisos a débilmente estriados

longitudinalmente, castafio oscuros. Nectarios reproductivos pedicelares tubuloso-sacciformes, pediculos

de los nectarios de 3-6 mm de longitud, urnas de s mm de wees x 2-2,5 mm de ancho, con la boca

dispuesta hacia el eje de la intl los nect iendo en el cuarto o quinto inferior del pedicelo.

Flores 22; opu. M Mad pies deltoides, de 1,7-2 mm de PRU x 1,4-2 mm de ancho, soste-

niendo el lares, carnosos, de 1,8—2 mm de longitud; pétalos ovados, libres, carnosos,

de 4,7-5,2 mm de longitud; estambres ca. 15, filamentos ca. 1,3 mm de longitud, a lisos, adnatos

a la base de los pétalos, anteras introrsas, oblongas, subcordatas, ca. 2,5 mm de longitud x ca. 1 mm de

ancho; ovario cónico, débilmente estriado longitudinalmente, ca. 3 mm x ca. 2 mm; estigma mamiforme,

subsésil, negruzco. Frutos desconocido

Distribución geográfica y ecologia.—S tzia j is sólo se conoce de la localidad tipo, en la región

sudeste de Brasil, estado de Espírito Santo, — en las márgenes rocosas del río Jucu.

709

Giraldo-Cañas, Estudio t

licelar. (G. Hatschbach et al. 59728)

A. Rama florifera. B. Botó

Cañar

En 7 Sri

I [| £ al > H 1m L. | di ea adis »Jhi

710 f Texas 3(2)

Observaciones.—Esta especie es única en el género por su inflorescencia densa y cortamente racemosa

(pedúnculo de 4 cm de longitud y TAM ca. 2,5 cm de longitud), sus delicados pedicelos de 3,9—4,8 cm de

longitud, sus pequeños nectarios rep pedicelares sacciforme-tubulares y con pedículos hasta de

6 mm de longitud, sus pequeñas flores (4,7-5,2 mm de alto x 3,8—4 mm de ancho), sus bractéolas deltoides

y por sus largos pecíolos (hasta de 23 mm).

8. Schwartzia lozaniana Gir.-Cañas, Caldasia 23:384. 2001. (Fig. 8). Tiro: COLOMBIA. Nano: Junin-Tumaco road,

6-11 km W of Junin, roadside thickets and forest edge, 850-1030 m, 27 feb 1979, J. Luteyn & M. Lebrón-Luteyn 6880 (HOLoTIPO:

COL); isoriros: MOI, NY!)

iss ida con ramas pee hasta de 4 m de nee t i lese reci à nee o İr-

a nn earrerta=na AACA

peddle ente edo recurvado, liculad tl canal muy — ca. 1mm de anche) castaño ENS

a negruzco, de 9-32 mm de longitud; nou n a elípticas, de (7)13-19 cm de longitud x 4-10 cm de

ancho, de base redondeada y ápice | te asimétrico, con márgenes ligeramente revolutas

y engrosadas; nervio medio conspicuo en ambas caras, elevado en la haz y plano a levemente abultado en el

envés, estriado, de 1-1,3 mm de ancho; venación secundaria débil; 0-6 pares de glándulas laminales en la

superficie abaxial (algunas láminas no presentan glándulas), submarginales, —— a 2-4,5 mm de la

margen, elípticas, de 1,5 mm de largo x 0, des mm de ancho, con su orifi de 0,5 mm de largo x 0,25

mm de ancho, distanciadas entre si 1—5 cm. Infl i inal, racemosa, multiflora, densa; pedúnculo

de 6,5-18 cm de longitud; pedicelos de G—)5—6(-7) cm de lonsimd: rectos a flexuosos, estriados, un poco

más ene en su pon distal, castaño oscuros, cada uno con una bráctea nectarífera (nectario). Nec-

, tubulares, péndulos, purpüreos, estriados longitudinalmente,

pedicles de los Mecano ca. ] mm de ind. urnas de (9213-15(-17) mm de longitud x 3-6 mm de

ancho, boca rostrada, con el orificio elíptico a circular, de 1,3—2,8 mm de ancho, dispuesta hacia el eje de la

inflorescencia y ocupando una posición transversal respecto a su eje; los nectarios Haud por debajo del

1/3 proximal del pedicelo. Flores 18—68, inserción pedicelo-flor horizontal; bractéol , opuestas,

orbiculares, obtusas, rugosas, carnosas, breves, ca. 2 mm de longitud, márgenes inui didus pde or-

biculares, rugosos, carnosos, 2,2-2,8 mm de longitud, márg translucidas; pétalos connatos basalmente

por ca. 2/3 de su longitud, obovado-elipticos, reflexos en la antesis, de 6,5-10 mm de longitud x ca. 3 mm

de ancho, carnosos; estambres ca. 25, filamentos de 3—5 mm de longitud, aplanados, ]

a la base de los pétalos, anteras oblongas, subcordatas, de 2-3 mm de longitud; ovario cónico, ligeramente

estriado, ca. 2,5 mm de longitud x ca. 1,5 mm de ancho; estigma mamiforme, subsésil, negruzco. Frutos

inmaduros globosos, apiculados, sulcados ais ib M c de 2,23 mm n dramaga; i Ed

, adnatos

Distribución geográfica y ecología.—Scl 5lo se conoc ce de la vertient e occidental andina

en el departamento de Nariño (sur de UMEN y de las provincias sie Carchi y Esmeraldas (Ecuador), areas

pertenecientes al Chocó Biogeográfico. Esta especie, como muc has otras de la familia Marcgraviaceae, crece

en bordes de bosques de zonas húmedas a muy húmedas, entre los 500 y los 1600 m.

Observaciones. —Schwartzia lozaniana es la especie que presenta los pecíolos más largos (hasta de 32

mm), mientras que en las demás especies éste sólo alcanza hasta los 1423) mm de longitud. Los nectarios

reproductivos pedicelares de S. RII. proces una Lun Canaan Ge néctar, alas se infiere de las

manchas dejadas por éste en el interior de los nectarios (vistos n ndo a llenar un

poco más de la mitad de su volumen. Los binomios buon venusta Bedell (in gud ) y Schwartzia venusta

de Roon & Bedell (in sched.) son nombres no ii que pid A a S. lozaniana.

Especímenes examinados: COLOMBIA. Nariño: Mun. Ricaurte, corregimiento de Altaquer, 1600 m, 19 feb 1977, O. de Benavides 893

(PSO). Mun. Barbacoas, to de Junín, 1200 m, 7 oct 1988, O. de Benavides 10252 (COL, PSO), 2-8 km E of Junín on Tumaco-

Tügueres road, 1100 m, 26 jul 1986, A. Gentry et al. 55249 (MO, PSO). Mun. Barbacoas, dM e El Diviso, camino al Carafio,

580—600 m, 23 ago 1995, M. González et al. MA Mun. Barbacoas, km 2 tray Barbacoas, 1150 m, 14 abr 1989, B. Ramírez

1334 (PSO). ECUADOR. Carchi: Trail to Río p ; g ridge li A I t, 1330 m, 17 ene gen W. Hoover et

al. 2538 (MO). Bord Prov. Carchi and E Idas, about 201 t Lit d Lita-Alto Tambo, 550 m, 25 jun 1991, H

van der Werff et al. 11992 (MO). E Idas: Quinindé Cantón, Bilsa Biological Station, R Ecológica Mache-Chindul, 40 km NW

Giraldo-Cañas, Estudio t bmi | i | i Schwartzi 711

Fig. 8. Sch ia | 7 Gir-Cañas. A. Rama florifera. B. Botón fl ly tari I lurcti I licelar. €. Detalle de | tami D. Ovario Y

Luteyn 6880).

of Quinindé, I le los G ill f , 650 m, 12 mar 1997, J. Clark et al. 3974 (COL); Lita-San Lorenzo road, 18

km W of Río Lita Bridge, on old road below Lita, 6.6 km W of bridge over Río Chuchubí, site near Alto Tambo, 425 m, 10 jul 1998, T.

Croat et al. 82631 (MO). Lita-San Lorenzo road, 10-20 km NW of Lita, 800 m, 12 may 1991, A. Gentry et al. 70088 (MO).

712 t tani titute of Texas 3(2)

9. Schwartzia magnifica (Gilg) Bedell, Monogr. Syst. Bot. Missouri Bot. Gard. 45:1256. 1993. (Fig. 9). No-

rantea magny Gilg, Bot. Jahrb. pe = ee p Tiro: PERÚ. Huánuco: near Monzón, 2000 m, A. Weberbauer 3528 [LECTOTIPO:

MOL, AL ignado Era Giraldo-C A FL O!]

Arbustos escandentes, hee ramas robustas, tallos levemente angulosos. Hojas cortamente pecio-

ladas; pecíolo recurvado, canaliculado, levemente estriado, castafio oscuro, de 3-4 mm de longitud x ca.

3 mm de ancho; láminas obovadas, de 9-12,5 cm de longitud x 5,5—6,8 cm de ancho, de base cuneada

y ápice redondeando y cortamente apiculado, con márgenes ligeramente revolutas y engrosadas; nervio

medio conspicuo en ambas caras, aunque se va desvaneciendo hacia la porción distal, plano a levemente

hundido en la haz y ligeramente abultado en el envés; venación secundaria apenas perceptible en la haz y

suprimida en el envés; 15—24 pares de glándulas laminales en la superficie abaxial, submarginales hacia la

parte distal de la lamina, localizadas a 2-13 mm de la margen, elípticas, ca. 05 mm, distanciadas entre si

0,1-0,4(-9) mm. Infl ia terminal, racemosa, multiflora, densa; pedunculo de 10-14 cm de longitud;

par de4 i 5 cm de hd NN on de 9 cm a Moe en fruto), sinuosos, lisos, castaño oscuros.

trados, péndulos, pedículos de

los medianos 1-5 mm de longitud, urnas de 12-17 mm de ae x 4,5-7 mm de ancho, con los orificios

dispuestos hacia el eje de la inflorescencia; los nectarios naciendo en el tercio inferior del pedicelo. Flores

18-28; bractéolas sepaloides, orbiculares, carnosas, de 5-6 mm de longitud; sépalos orbiculares, carnosos,

de 6,5 mm de longitud; pétalos ovados, connatos basalmente, carnosos, ca. 9 mm de longitud x ca. 3 mm

de ancho; estambres 13, filamentos ca. 3 mm de longitud x ca. 0,6 mm de ancho, aplanados, lisos, libres,

anteras oblongas, ca. 2,5 mm de longitud x ca. 1,5 mm de ancho; ovario cónico, de 3 mm de longitud; estilo

ca. 1 mm de longitud; estigma mamiforme, ca. 1 mm de diámetro. Frutos inmaduros globosos, ca. 1 cm de

nue sce aan E: apículo ca. 0 cm de ongir.

Jz zaan dad 1 J Ela de la región

andina de neo, y el sur de la Cordillera Vileabaniba (Perú) y crece entre los 2000 y los 2500 m. Esta

especie, al igual que otras especies de los bosques montanos peruanos, se encuentra bajo amenaza por la

deforestación y la expansión agrícola (León & Monsalve 2006).

Observaciones. —El holotipo depositado en el herbario de Berlin (B) fue destruido y no se conocía

ningún otro material de la especie, a excepción del isotipo de MOL y del ejemplar A. Weberbauer 7071 de

Perú (B destruido; fotografías F!, MO!, NY!) correspondiente a Norantea thyrsiflora Gilg (in sched., igual a S.

magnifica), hasta que estudié los ejemplares P. Núñez et al. 23484 (NY, US), los cuales estaban determinados

como Norantea macrocephala G. Don en NY y como Norantea macrocarpa G. Don en US. Después del estudio

detallado de estos especímenes, pude advertir que se trataba de Schwartzia magnifica, lo que se convierte en

el redescubrimiento de la especie.

Especímenes examinados: PERÚ. Cuzco: Prov. La Convención, Distr. Echarati, east Río Apurimac NE Pueblo Libre, up mountain of

Anchihuay and Bella Vista, south Cordillera Vilcabamba, 2445 m, 3 ago 1998, P. Núñez, W. Nauray, R. de La Colina & S. Udvardy 23484

10. Schwartzia parrae Gir.-Cañas, Novon 12:456. 2002. (Fig. 10). Tro: COLOMBIA. Antioquia: Mun. Yolombó, vereda

El Bote, sitio Alto del Bote, 8,1 km SO de Yolombó, bosque perturbado, 75?01'O, 6°32’N, 1550 m, 1 jul 1989, R. Callejas, J. Betancur

& O. Escobar 7957 (HoLoriro: HUA!, isoriros: MO!, NY!)

1 a aplanados, T PS PON Lae AOR DR NR ESL NUT AAPA Dad A A Uno QURE

Arbustos escandentes, tallos cilindricos ai

ferruginea a cobriza. Hojas subsésiles a ee pollas er ligeramente estriado, recurvado,

canaliculado (el canal muy angosto, ca. 1 mm de ancho), castaño claro a negruzco, de 2-7 mm de longitud;

láminas oblongas a obovadas, de (5-)9-22 cm de longitud x (1,3—)2-5,2 cm de ancho, de base redondeada

a ligeramente cordata y ápice acuminado prolongado en un apículo negruzco de 1-1,5 mm de longitud,

con márgenes ligeramente revolutas o involutas y engrosadas; nervio medio conspicuo en ambas caras,

levemente hundido en la haz y Mendes abultado en el envés, liso, de p mm dui Se en la pe

proximal de la lámina foliar, d ia la porción distal de la lámina foliar; venación secundaria

suprimida o apenas perceptible en la haz, débil en el envés; 14-28 pares de plandas laminales en la su-

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perficie abaxial, localizadas a 1,5-11 mm de la margen, elípticas, de 1 mm de largo x 0,5 mm de ancho, con

su orificio negruzco de 0,2 mm de largo x 0,15 mm de ancho, distanciadas entre si 1-5 cm. Inflorescencia

terminal, racemosa, ERE ME densa; pedunculo de 12-19 cm de longitud; pedicelos de 4—8 cm de longi-

tud, cilíndricos a ] ados, rectos a levemente flexuosos, lisos a ligeramente estriados, un

poco más gruesos en su porción rd es cada uno con una bráctea un (nectario). Nectarios

reproductivos pedicelares cimbiformes, levemente rostrados, péndulos, verdes a purpüreos, ligeramente

estriados longitudinalmente, pedículos de los nectarios 1,5-3 mm de longitud, urnas de (7-)9-18 mm de

713

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Fic. 10. Sch

Giraldo-Cañas, Estudio taxonómi | i lameri Schwartzi 715

longitud x (3-)4-8 mm de ancho, los orificios dispuestos hacia el eje de la inflorescencia y ocupando una

posición transversal respecto a su eje, los nectarios naciendo por debajo del 1/3-1/6 proximal del pedicelo.

Flores 20-32, inserción pedicelo-flor horizontal a moderadamente oblicua; bractéolas sepaloides, opuestas,

orbiculares, obtusas, rugosas, carnosas, 5-7 mm de longitud, márgenes translúcidas; sépalos orbiculares,

rugosos, carnosos, 7-8 mm de longitud, márgenes translúcidas; pétalos connatos basalmente (antes de la

antesis) por ca. 2/3 de su longitud, obovado-elipticos, reflexos en la antesis, de 8,5~11 mm de longitud x ca.

3 mm de ancho, carnosos, rugosos; Ei 48, etiam: ca. 1,5 mm de longitud (antes de la antesis),

aplanados, , adnatosalab ] s elíptico-oblongas, subcordatas basalmente,

de 3-4 mm de longitud; ovario cónico, j'oesemerite Eso do. ca. 3, 5 mm de longitud x ca. 3 mm de ancho;

EE mamiforme, subsésil, negruzco. Frutos desconocidos.

ribución geográfica y ecología. —Esta especie sólo se conoce de la cordillera Central andina (departa-

fica y ecología.

mento de Antioquia, Colombia) y crece entre los 1100 y los 2000 m. De esta especie sólo se poseen cuatro

ejemplares colombianos, por lo que se infiere que ésta se encuentra en vías de extinción, dada la acelerada

fragmentación y/o destrucción de los bosques en donde ella vive.

Especimenes examinados: COLOMBIA. Antioquia: Mun. Amalfi, 8-27 km NE de aa en la via V an preven sitios saat y

Marengo, 1150-1450 m, 7 dic 1989, R. Callejas et al. 9102 (HUA). Mun. Guatapé, sect 520510).

6°10’N, 1800-2000 m, 22 jun 1997, F. Alzate et al. 247 (F, HUA). Mun. San Luis, la Manizal fi le Ramón J ill .1460— 1760

m, 11 oct 1981, C. Orozco et al. 549 (COL)

11. Schwartzia pterosara de Roon & Bedell ex Gir.-Cañas, Novon 15:123. 2005. (Fig. 11). Tiro: ECUADOR.

Naro: Along road between Quito and Baeza, 3 mi W of Baeza, 0°25'S, 77°51'O, 2000 m, 2 oct 1980, T.B. Croat 50284 (HoLoTIPO:

MO!, isotiros: GH, MARY

Arbust dentes, epífitos, bteret lenticelas | itudinales. Hoj t lad

pedio eani canaliculado, castaño oscuro, de 3-6 mm de longitud x 2 mm de udo fenis

obovadas, de (528-12,5 cm de longitud x (2,2)3,8-6,2 cm de ancho, de base aguda a cuneada y ápice

obtuso a redondeado, mucronado (el mucrón ca. 1,5 mm de longitud) u ocasionalmente cuando el mucrón

se ha caído entonces el ápice se ve retuso; nervio medio liso, conspicuo en ambas caras, aunque se va

desvaneciendo hacia la porción distal, plano a levemente hundido o sobreelevado en la haz y ligeramente

elevado en el envés; venación secundaria débil en ambas caras; 25—34 pares de glándulas laminales en la

superficie abaxial, elípticas, 0,7-1,2 mm de largo x 0,5-0,8 mm de ancho, dispuestas en dos filas longitu-

dinales en ambos extremos de la lámina, una fila submarginal ubicada a 1—7 mm de la margen, la segunda

fila a 5-11 mm de la margen. Inflorescencia terminal, racemosa, multiflora, densa; pedúnculo de 10-17,2

cm de longitud; pedicelos de 3,5-5,5 cm longitud, rectos, formando con el raquis un ángulo de 45-83",

lisos a levemente estriados, cobrizos. Nectarios reproductivos pedicelares ciatiformes a ventricosos, pén-

dulos, verdosos a rojizos, levemente estriados, pedículos de los nectarios de 3—4 mm de longitud, urnas de

6-10 mm de longitud x 6-8 mm de ancho, con el orificio dispuesto hacia el raquis; los nectarios naciendo

en el cuarto o quinto inferior del pedicelo. Flores 18—42; bractéolas sepaloides, adpresas al cáliz, opuestas,

elípticas a obovadas, carnosas, de (428-11 mm de longitud x (4-)7-9 mm de ancho; sépalos orbiculares,

carnosos, con márgenes membranáceas, de (45-7 mm; pétalos libres, carnosos, elípticos a oblanceolados,

de 8-11 mm de longitud x 3-5 mm de ancho; estambres 12, de 6-8 mm de longitud, filamentos aplanados,

adnatos a la base de los pétalos, anteras subsagitadas, amarillentas; ovario piriforme o cónico, de 4-5 mm

de longitud; estilo cilíndrico, ca. 1 mm de longitud; estigma liso, mamiforme, negruzco. Frutos globosos,

apiculados, de 0,8—1,1 cm de diámetro, rojizos; semillas reniformes a elípticas, de 1,2-1,5 mm de longitud

x ca. 0,7 mm de ancho, reticuladas, aane numerosas.

Distribución geográfica y ecología. 5lo se conoce de los bosques andinos húmedos

(ca. 2000 m) en los alrededores de Baeza en los ere de Napo (Ecuador). No obstante, puede haber mas

especímenes en otros herbarios de Ecuador (e.g., QCA, QCNE, GUAY), pero desafortunadamente no tuve

acceso a esas colecciones. Por otra parte, Bedell (1985) citó otro espécimen de esta especie para Colombia

[Cuatrecasas 23618 (F, US)], pero tampoco tuve acceso a este ejemplar, a pesar de que dichos herbarios

716 || [| E al n.a H In LI FEE F1 iT la F ka A!

Lr 4

la R & Bedell ex Gir-Canas. A. Rama florífera. B. Botón floral tari ducti dicelar. €. Detalle de los

Fic. tzia pt

estambres. (7. Croat 50284).

Giraldo-Cañas, Estudio taxonómico para las especies sudamericanas Schwartzi 717

me enviaron sus colecciones, entre las que no se encontraba la mencionada por Bedell (1985). Asimismo,

Stefan Dressler (FR, com. pers.) me mencionó la existencia de otros especímenes de Perú depositados en

los herbarios FR y MO (Campos & Núñez 4612), pero desafortunadamente nunca ae vere Asi, si estas

colecciones pertenecen a S. pterosara, su gradiente de distribución se ampliaría sig

Observaciones. —Esta especie está más relacionada con S. magnifica, pero también exhibe similitudes

con S. adamantium y S. spiciflora. No obstante, S. pterosara difiere de éstas por su gran número de glándulas

laminales, la densa inflorescencia y el tamaño y la forma de los nectarios reproductivos pedicelares.

Especimenes examinados: ECUADOR. Napo. Vicinity of Baeza, wet montane forest, 2000 m, 28-29 jun 1982, L. Besse, H. Kennedy &

R. Baker 1504 (M

12. Schwartzia renvoizei Gir.-Cañas, Caldasia 26:424. 2004. (Fig. 12). Tiro: COLOMBIA. Vare pet Cauca: Cordillera

Occidental, vertiente occidental, dies del río Sanquinini, lado izquierdo, La Laguna, bosques, 1250-1400 m, 10-20 dic 1943, J.

OL!, FP.

Cuatrecasas 15456 (HOLOTIPO:

Arbusto epífito, escandente, muy ee ramas Pda robustas; tallo terete a subterete, rugoso,

corteza escamosa , gris oscura, estriada | 5siles, peciolos engrosados, recurvados,

rugosos, castaño oscuros, hasta de 56) n mm de LN yc ca. o mm "m ancho; laminas oblongo-elípticas,

de 13-14 cm de di ds x 5-6 cm de ERN | data y ápice obtuso a redondeado,

Fa - Li - ]

ATI IER ATITA ata mirada

con

su porción diel plano en la haz y hene to en nile envés, liso a levemente estriado; venación

secundaria muy débil a suprimida en la haz, notoria aunque débil en el envés; 45—57 pares de glándulas

laminales en la superficie abaxial, dispuestas en una fila más o menos regular a cada lado del nervio medio

(aunque puede haber algunas glándulas dispuestas en forma aleatoria) y separadas de éste entre 5 y 10

mm, y de la margen entre 10 y 15 mm, las glándulas basales distanciadas entre sí 5-9 mm, las mediales

y las distales muy próximas las unas a las otras (entre 0,5 y 1,5 mm), elípticas, levemente elevadas, ca. 0,6

mm de longitud x ca. 0,4 mm de ancho, con su orificio eliptico, ca. 0,1 mm de longitud. Inflorescencia

terminal, racemosa, multiflora, densa; pedúnculo de 10—11,6 cm de longitud; pedicelos de 4—6,9 cm de

longitud, levemente curvos, oblicuos con relación al eje de la inforescencia, lisos a ligeramente estriados,

castaño oscuros a cobrizos. Nectarios reproductivos pedicelares galeados, robustos, péndulos, pedículos de

los nectarios de 2-3,5 mm de longitud, urnas de 10-14 mm de longitud x 8-11 mm de ancho, con la boca

de contorno ovado y márgenes planas a débilmente revolutas, dispuesta hacia el eje de la inflorescencia;

los nectarios naciendo en el tercio inferior del pedicelo. Flores 20-26, presentando maduración basípeta,

inserción pedicelo-flor horizontal a levemente oblicua; bractéolas sepaloides, opuestas, sosteniendo el cáliz,

orbiculares, carnosas, de 5-6 mm; sépalos orbiculares, carnosos, de 5-7(-7,5) mm; pétalos aparentemente

libres y sólo una mínima porción basal de la corola es connata, reflexos en la antesis, oblongos, carnosos,

de 15-16 mm de longitud x 4—5 mm de ancho; estambres 70, filamentos ca. 9 mm de longitud, aplanados,

anchos en la base y angostos en su ápice, lisos a levemente estriados, adnatos a la base de los pétalos y

dispuestos en tres verticilos, anteras oblongas, sagitadas, amarillentas a castañas, ca. 4 mm de longitud x

ca. 0,7 mm de ancho; ovario piriforme, tetra-locular, ca. 8 mm de alto, rugoso a estriado, castafio oscuro a

negruzco; estigma mamiforme, negruzco. Frutos desconocidos.

Distribución geográfica y ecologia.—Schwartzia renvoizei sólo se conoce de la localidad tipo, la que cor-

responde a bosques andinos premontanos de una región pluvial de la vertiente pacífica de la Cordillera

Occidental en el departamento del Valle del Cauca (Colombia). En vista de que no se conocen más recolec-

ciones de la especie y dada la acelerada deforestación de la región en donde ésta vive, se presume que la

especie se encuentra en un "nes eee de extinción.

1 E |

, venación secundaria débil a suprimida,

Observaciones.—El

pedúnculo corto y número de flores por inflorescendia, S. renvoizei podría confundirse con S. antioquensis

y S. chocoensis. No obstante, S. renvoizei es única en el ee y! EE reconocible por sus nectarios ga-

leados, sus numerosos estambres y p glán El binomio Schwartzia

micradenia de Roon & Bedell (in sched.) corresponde a s renvoizei.

718

£T. 3/4)

ICAd» 12)

"ur è

IG. ri, Gir.-Cañas. A. Rama florifera. B. Detalle de | les de la | C. Flor y lucti licelar. (J.

Cuatrecasas 15456).

Giraldo-Cañas, Estudio t omi | i i i Schwartzi 719

13. Schwartzia spiciflora (A.L. Juss.) Bedell, Fl. Lesser Antilles 5:309. 1989. (Fig. 13). Marcgravia spiciflora A.L.

Ol Ann. Mus. Natl. Hist. Nat. 14:402, t. 25. 1809. Norantea spiciflora (A.L. Juss.) Krug & Urban, Bot. Jahrb. Syst. 21:520. 1896.

DELOUPE: Richard s.n. (HoLoriro: P-JU; isoripos: C, P; fotografías del holotipo en F!, MOD.

Ascium berteroi ae a Veg. eh ns lle berteroi T G. Don., Gen. Hist. 1:624. 1831. Tro: INDIAS OCCIDEN-

TALES: Bertero s. o TO, véase Bedell (1985: 152)].

Arbustos escandentes, Mm lianescentes, hasta de 6 m de altura, con ramificación profusa, ramas ro-

bustas; tallos subteretes, corteza lisa a ligeramente estriada longitudinalmente, castaño a ferrugínea. Hojas

cortamente pecioladas; pecíolo recurvado, canaliculado, engrosado, liso a débilmente estriado, castaño

oscuro, de 2-7 mm de longitud x 2,5-4 mm de ancho; láminas obovadas u oboelípticas, de (6-)8,4-11,2

cm de longitud x (324,6—5,9 cm de ancho, de base simétrica o levemente asimétrica, cuneada, ápice obtuso

o agudo y en ocasiones ligeramente asimétrico, mucronado, retuso cuando se ha caído el mucrón, con las

márgenes lisas o débilmente revolutas; nervio medio centrado a más comünmente desviado, lo que hace

que una de las mitades de la DA sea más ancha e la otra, Became MPH i4 FORSE en ambas

Caras, auaqu en la porci ela] lámina;

caras O debitiente perceptibles 0-14) pares de glándulas laminales en la superficie eer dispuestas

irregularmente, circulares, ca. 0,5 mm de diámetro. Inflorescencia terminal, racemosa, multiflora, densa;

pedúnculo de (4,525,5-10 cm de longitud; pedicelos de 1,6-3,5 cm de longitud (hasta 4 cm de longitud

en fruto), rectos a ligeramente curvos, lisos o débilmente estriados longitudinalmente, con lenticelas o sin

las mismas, castaño oscuros. Nectarios reproductivos pedicelares cimbiformes, rojizos, rugosos, con la

boca dirigida hacia el eje de la inflorescencia, pedículos de los nectarios de 0-1 mm de longitud, urnas de

(2,533,5-5,57) mm de longitud x ca. 3 mm de ancho, Tost AdAS; los nectarios naciendo a 0,4-1,3 cm de

la base del pedicelo. Hew MM da Peas bractéol i borbicul , opuestas, sosteniendo el

cáliz, de 2-3 mm del rbiculares, carnosos, ae 3-3, one de biai pétalos connatos hasta

su porción media, obovado- Duces a oblongos, 6-8 mm de longitud; estambres 22, filamentos ca. 2 mm de

longitud x ca. 0,5 mm de ancho (antes de la antesis), basalmente adnatos a los pétalos, aplanados, rugosos,

anteras subsagitadas, ca. 2,5 mm de longitud x ca. 0,8 mm de ancho; ovario globoso a cónico, rugoso, de

2,5—4 mm de diámetro; estilo estriado a rugoso, de 1-1,5 mm de longitud; cdd aM e negruzco.

Frutos globosos, conspicuamente apiculados, rugosos, de 9—10,5 mm de diámetro; il , de

33,5 mm de longitud x ca. 1,5 mm de ancho, reticuladas, rugosas, negruzcas, brillantes numerosas.

Distribución geográfica y ecología.—Schwartzia spiciflora es propia de bosques de montaña, tanto húme-

dos como nublados, caracterizados principalmente por Clusiaceae, Lecythidaceae y Podocarpaceae y crece

entre los 400 y los 1500 m. Ésta está restringida a las Antillas Menores (Dominica, Guadalupe, St. Vincent

y Trinidad) y al norte de la costa venezolana en el estado de Sucre.

Observaciones.—de Jussieu (1809) y Triana & Planchon (1862) comentaron que la corola de S. spiciflora

eS los pétalos connatos aaa) i qe la acercaba al ei Mu. ok Be un dd

4-09 A440 d oxrS6S APS P CS P DES

no en flores en antesis), en los que los uds —Á estar constituyendo una caliptra. El binomio bone:

jussiaei Triana & Planch. (Ann. Sci. Nat., Bot., sér. 4, 17:373. 1862) es un nomen nudum que corresponde a

la misma S. spiciflora.

Especimenes examinados: VENEZUELA. Sucre: Península de Paria, cerro de Humo, laderas de bosque húmedo nublado que miran

hacia La Roma e Irapa, entre la cumbre y La Laguna, noroeste de ms 1060- d m, i mar idi - Steyermark 94954 (NY, VEN);

Península de Paria, virgin cloud forest, above Las Melenas, N of Río Grande rro de Humo, 730-1050 m, 1 dic

1979, J. Steyermark & R. Liesner 120918 (MO).

14. Schwartzia weddelliana (Baill.) Bedell, Monogr. Syst. Bot. Missouri Bot. Gard. 45:1256. 1993. (Fig.

14). Norantea weddelliana Baill., Adansonia 10:242. 1872. Tiro: BOLIVIA: Yungas, Weddell 4318 (noLoripo: P; 1soTiro: Fl, fotografía

del holotipo en F!

Marcgravia ma ae ex ere Mem. ey Bot. di 3:8. Mn Tiro: BOLIVIA: Tune año 1890, M. Bang 390 [HoLoTiPo:

NY; isotipos: F! (d , NY! (d ), USI], pp. ex Wittm. [2 Marcgraviastrum

mixtum ise & Planch. ) un

720

ai”

lexas 3(2)

5mm

5 mm

EMI

p DAP ODe

Fic 12 el de? > FT

E (A.L. Juss.) Bedell. A. Rama florífera. B

120918).

C. Ovario v nedicelo. (1 Stevermark

Giraldo-Cañas, Estudi ómico para | i i i Schwartzi 721

Ni à | Gilg, Bot. Jahrb. Syst. 25, Beibl. 60:30. 1898. Tiro: BOLIVIA: Yungas, año 1890, M. Bang 390 | B?

F! (dos ejemplares) “MOI, NY! (dos ejempl

Norantea sandirst Gilg, Bot. Jahrb. Syst. 42: 126. 1908, Tiro: mo Puno: Prope Tambo Azalaya ad viam i Sandia et Chunci j

in fruticetis subapertis, 1500-1600 m, A. Weberbauer 1343 : g lel hol ip MO!, NY?).

Arbustos hemiepífitos, escandentes, prof t ificados, hasta de 4 m de altura, ramas medianamente

robustas; tallos teretes a subterestes, corteza débilmente estriada longitudinalmente, castaña o ferruginea.

Hojas cortamente pecioladas; pecíolo recurvado, canaliculado, levemente estriado, castaño oscuro, de 2-8

mm de longitud x 2-3 mm de ancho; láminas obovadas o oboelípticas, ocasionalmente elípticas, de 6-12,5

cm de longitud x (3, adl 53,25, 8) cm Be id nk base Fund y pice oe a iaa! nervio medio

conspicuo en aml | el envés tal; venación

secu ndari imida en amb is, MUY l M perceptible en la haz y/o envés; (5-)8-14(-19)

pares de glándulas laminales en la superficie abaxial, dispuestas en una fila más o menos regular a ambos

lados de la lámina, raramente en dos filas a ambos lados de la lámina, localizadas a 4-13 mm de la margen

y distanciadas entre si 3-17 mm, circulares a elípticas, ligeramente elevadas, crateriformes, de 0,9-1,4

mm de longitud x ca. 0,8 mm de ancho. Inflorescencia terminal, racemosa, pauciflora, laxa; pedúnculo

de (3—)3,5-7 cm de longitud; pedicelos de 3,5—7 cm de longitud, rectos a levemente curvos, débilmente

estriados longitudinalmente, castaño oscuros a cobrizos. Nectarios reproductivos pedicelares largamente

tubulares, péndulos, estriados longitudinalmente, pedículos de los nectarios de 1-7 mm de longitud, urnas

de (20230-40(-43) mm de longitud x 3—/ mm de ancho, con la boca dispuesta hacia el eje de la inflores-

cencia y con las márgenes te revolutas; los nectarios naciendo a 1,3—2,6(—3) cm de la base del

pedicelo. Flores 6—10(—12); braciédlas sepaloides, orbiculares, opuestas, carnosas, sosteniendo el cáliz, de

4—5,8 mm de longitud; sépalos orbiculares, carnosos, de 6—6,8 mm de longitud; pétalos libres, obovados,

carnosos, rojos, de 14-17 mm de longitud x ca. 8 mm de ancho; estambres ca. 18, filamentos ca. 7,5 mm

de longitud x ca. 0,7 mm de ancho, aplanados, lisos a ligeramente estriados longitudinalmente, adnatos a

la base de los pétalos, anteras ca. 4,5 mm de longitud x ca. 1 mm de ancho, subsagitadas; ovario globoso,

ligeramente rugoso, ca. 4,5 mm de diámetro; estilo ca. 1 mm de longitud; estigma mamiforme. Frutos glo-

bosos, apiculados, rojos, débilmente rugosos, de 10-16 mm de diámetro; semillas falcadas, de 2 ,83,5 mm

de longitud x ca. 1 mm de ancho, ae castafio oscuras, brillantes, reticuladas, numerosas.

ribución geográfica y ecología. 4 es propia del sur de Perü y el norte de Bolivia,

y crece en bosques húmed luros o secu cundarios, laderas pedregosas y en bosques ralos, entre los 500 y

los 2000 m. Merec destacarse la cita ade s weddelliana (tratada como Norantea sandiensis) para San José del

Guavire (norte de la Amazonia colombiana) realizada por Schultes (1951: 404) sobre la base del espécimen

Cuatrecasas 7685. Desafortunadamente dicho espécimen no fue localizado en ninguno de los herbarios y

por tal razón, se pone en duda la cita oc la ia pa paa one

Observaciones.—Esta especi d tari productivos, los cuales

presentan la boca con las márgenes conspicuamente ero las. Por otra parte, se observó en el ejemplar

Davidson 4779 (US) una variación morfológica poco usual, en la que un pedicelo distal se ramificaba en dos

ejes Mb a su vez dos flores fértiles) justo en el punto de origen del nectario reproductivo.

E inados: BOLIVIA. La Paz: Prov. Nor li oe i bei Coroico, 1100 m, oct-nov 1912, O. Buchtien 6020 (US).

Biev Nor Yungas, carretera fund 13, between Coroi road parallel to Rio Coroico, 700-1300 m, 6 nov

1976, C. Davidson 4779 (MO, NY, RSA-dos ejemplares-, US). Brow Nor Yungas, below San Pedro, Yolosa-Caranavi road, valley of Rio

Coroico, 760—950 m, 15 ene 1984, A. Gentry et al. 44215 (MO). Prov. Nor Yungas, 4.5 km below Yolosa, then 10 km W on road up the Río

Huarinilla, 1450 m, 19-20 oct Fin J Solomon n TO NY Prov. a Der 4.5 km below Yolosa, then 14 km W on road up Rio

Huarinilla, along old, abandoned road to Hacienda Sandillani, on sout 1200—1300 m, 24 ene 1983, J. ERA 9370 (MO).

Prov. Nor Yungas, 10.2 km SW of Yolosa on road to Gli ubi 20 km NE of Chintipata: 1700 m, 5 abr 1984, J. Solomon & M. Uehling

12250 (MO, NY, = ce ca. 2000 m, año 1885, H. Rusby 1362 (NY, dos ejemplares); Sin localidad: ca. 500 m, jun 1886, H. Rusby

2707 (NY). Santa E Ichilo, cerro Amboro, 23 sep 1990, M. Lewis & R. Clark 37736 (SD. PERÚ. Cuzco: Prov. La Convención,

139 km de Cuzco en Quell yo, subiendo hacia la “ceja”, entre Santa Teresa y Chaullay, 13 ene 1988, P. Núñez & F. Motocanchi 8811

(MO, NY). Prov. La Convención, Hisadquiña, km 127.5, 1700 m, 13-14 sep 1962, C. Vargas 13917 (US). Prov. La Convención, Colca,

1300 m, sin fecha, C. Vargas 20790 (NY). Junin: 38.1 km NE of Tarma on Carretera 20-B to Oxapampa, above Río Placa, ca. 2250 m, 13

722

L nm I

A Rama florifera. R. B

Fic. 14. Sch

Solomon 8526).

Giraldo-Cañas, | ómico | |

T 5.536 1] NIG EA Ü VOB ta Oxana mna

DIVA AdpPalilDa,

nov 1979, J. Jones 9090 (RSA, d

ca. 2000 m, 14 nov 1979, J. Jones 9149 (RSA).

EJEMPLARES EXCLUIDOS Y REGISTROS DUDOSOS

Bedell (1985) citó un espécimen del departamento del Cauca (Colombia) y lo asignó a la especie Schwartzia

pterosara |Cuatrecasas 23618 (F, US)]. No obstante y a pesar de disponer de los préstamos de Marcgraviaceae

de estos dos herbarios, dicha colección no se pudo localizar.

Como dato curioso, merece destacarse la cita de Sc]

sis) para San José del Guavire (norte de la Amazonia solemn) realizadas por Schultes (1951: 404) sobre

la base del espécimen Cuatrecasas 7685. Desafortunadamente dicho ejemplar no fue localizado en ninguno

de los herbarios y por tal razón, se pone en duda la cita de la especie para Colombia. Del mismo modo

S. weddelliana fue citada para Brasil por Szyszylowicz (1893: 163) (como Norantea weddelliana Baill.), pero

realmente ésta sólo se distribuye en el sur de Perú y el norte de Bolivia.

331_7l: Fa 4 ] papa a. NL, E EE

ESPECIES EXCLUIDAS

LISTA DE LOS SINÓNIMOS DE LAS ESPECIES SUDAMERICANAS DE SCHWARTZIA:

los binomios de la derecha son los nombres actualmente aceptados.

A (C ] Y Bedell ex Cir -Cafias

EL rim

Ascium adamantium Steud.

Ascium berteroi S. ere oo ze E: eS ined

Lanas

Ascium selloi Spre

Marcgravia M dn Poepp. ex Rusby - la liliana (Baill.) Bedell

Marcgravia spiciflora A.L. Juss. = SOANA pee n L. an inis

pare 3.11 r7: Cañas

Norantea adamantium Cambess. = Schwartzia adamantium (Cambess.) Bedell ex Gir.-Cañas

Norantea bahiensis Mart. ex Wittm

is ió re: G. ad ciar pco is L Juss Bedell

ffs Ale ) Bedell ex Gir -Cañas

Norantea j yu Triana & Planch ie spiciflora al L n ) Bedell

1 *

Iz] afi Ce Lee

n”-1 CL L3 ff 7-1 law} DBD 3-11

[o J [=]

qs E PLE

3.7.0012 ‘TD BM

Norantea selloi (Spreng) G. Don - Schwartzia brasiliensis (Choisy) Bedell ex Gir.-Cañas

Norantea Td (s L ns iip & Urban - oe tzia ne (A.L. Juss.) Bedell

cia weddelliana Baill = pipe ls a Baill js Bedell

aTiac

valid»

pai

)

Jı

Schwartzia "m Vell. = S tzi iliensis ( y

Cl : ; i A Ta T Kv Barlall Chl f Es Cañas

C-l hot p.4.11 cil it 1 p Caña

C-] z doa R &r Rerlell Gehwartria ln2aniama (ir -Cañas

AGRADECIMIENTOS

Qui ifest i pr pide l Instituto de Ciencias Naturales de la Universidad Nacional

de Colombia por todas las facilidades que me brindó para la preparación de este trabajo. A los curadores de

los herbarios CEPEC, CHOCO, COAH, COL, CR, CUVC, HUA, F, IBGE, JAUM, MEDEL, MEXU, MO, MOL,

MPU, NY, PSO, RSA, SI, SP, UDBC, UIS, US y VEN por los préstamos enviados o por su grata colaboración

talaci A Stefan Dressler (FR) y Adrian de Roon (U) por su valiosa ayuda y por

ies. Asimismo, a Peter A. Schafer del Institut de Botanique - Université

durante = visita asusi

Montpellier (Be:bario MPU) p por su valiosa información acerca de material tipo depositado en MPU. A Rosa

Ortiz-Gentry (MO), Olga Martha Montiel (MO), Victoria Hollowell (MO), Carlos Parra (COL), Enrique Forero

(COL), Paul M. Peterson (US), Orlando Rivera (COL), Orlando Rangel (COL) y Travis Columbus (RSA) por

724 || I al a. a || LI = Texas 3(2)

su valiosa colaboración. A Stefan Dressler (FR) y Alvaro Idarraga (HUA) por el obsequio de valiosa biblio-

grafia. A los herbarios de los jardines botanicos Rancho Santa Ana “RSA” (Claremont, California, EE.UU)

y Missouri “MO” (St. Louis, Missouri, EE.UU), así como al “Smithsonian Institution-National Herbarium”

(Washington D.C., EE.UU) por las facilidades económicas brindadas para las visitas a sus instalaciones. Á

Dubán Canal y Henry Arellano por la elaboración de las ilustraciones, A Barney Lipscomb por su valiosa

colaboración y amable ayuda para llevar a feliz término esta publicación.

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726 Jou

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ls A : "ES 1-335 1 E AAA | zie 1 1 L

suc g g p

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1 utilize the | herl den.—Tracie Harris, eee — Rescate jnstitüte 2d Texas, Fort Worth, Texas 76102- 4025, U.S.A.

LPL EL

A——————————————M——————————

J. Bot. Res. Inst. Texas 3(2): 726. 2009

AGAIN: TAXONOMY OF YELLOW-FLOWERED CAULESCENT OXALIS

(OXALIDACEAE) IN EASTERN NORTH AMERICA

Guy L. Nesom

2925 Hartwood Drive

Fort Worth, Texas 76109, U.S.A.

guynesom@sbcglobal.net

ABSTRACT

The taxonomy of Oxalis cect f^ z Tu: : E ] f£. NI^rth^ A » " + J “41 I : 1 Ui ed ] s

i Mu parecian those of Eiten and Lourteig. Eight taxa, previously gnized bspecies or varieties, are recognized

Oxalis stricta L. and O. dillenii Jacq ict ppropriately i ified in tl hat Fi 1 tł

Grate florida Salisb. = O. dillenii subsp filip ipes) inct ily of the Atlantic stat 1 Gulf 1

inland regions. Oxali distinct f tl theastern USA wit] ifi lisj li

in northeastern Medco. Oxalis texana (Small) Fedde (= Oxalis priceae na e) is — asa distinct species ofe east Texas and

adjacent Louisiana and Arkansas. Oxalis texana is close in MK MM ds dilleni ymp with it but

distinct in geography n de ad from 2 Peak sensu stricto, illi is Schwegm. is geographically and hologicall

distinct from O. grandis yI gnated for O. lyonii and O. illinoensis.

RESUMEN

c s lat fa su f p P a : lat 1 + da NT £n EA + " 1 E a

4 Lu Á4

nes, particularmente las de Eiten y Lourteig. Ocho taxa, alg idos previ bespeci binis se reconocen

aquí en el rango específico. Oxalis nd y O. dillenii Jacq trict identit propiad l 1 »

los nombres. Oxalis Lied Salisb. (= O. dillenii ED Jur ) pecie distinta primari de | tad lánticos y la costa del

L 2 C 11 es Fees | to qo TTCA : ' CF p:

Golfo, m g i I I

1] s J: 7 cada Mt f^ Las rc TX Gade. A L : 1 "n + I

E r L

dioni del este T Texas ee con las a Louisiana y Arkansas. O alis t tá próxi l fologia a O dillenii se sensu

stricto y I p y fología de O. pri trict Oxalis i illinoensis Schwegm.

ef: pes vh ERES NE 2 A x C 3 a E y re 211

es geográfica y g grandis Small.: g pos p lyonii y O. illinoensis.

Yellow-flowered caulescent Oxalis (sect. Corniculatae) of North America has been the object of interest and

study for more than a century. Small (1898, 1903, 1907) described a number of new taxa and brought the

names into wide usage; he also treated sect. Corniculatae as a separate genus, Xanthoxalis Small. Wiegand

(1925) provided a detailed and formal overview of the group and recognized many minor variants with

formal names, with an emphasis of study on northern North America. Eiten’s (1955, 1963) studies distilled

the taxonomy to a set of more realistic taxa based on biological concepts rather than typological ones.

Lourteig (1979) restudied the group and applied a very different set of names to the taxa, differing in part

in the selection of types and in part in concepts of relationships and taxonomic ranks. Useful perspectives

on problems of typification and nomenclature have been provided by Watson (1989) and Ward (2004), as

noted below. Still, significant problems in identification have remained and confusion persists about which

names are appropriate. In restudying, again, the eastern North American pate toward development of a

treatment of Oxalidaceae for the Flora of North America series, I have come to a still different view, although

it is far closer to Eiten’s than Lourteig’s. In fact, Lourteig’s beon € nomenclature appears to have been

retrograde in many ways compared to Eiten's.

In the taxonomic summaries below, synonymy is representative, showing the more commonly used

names and some that allude to aspects of variability within the species. Additional synonymy can be found

in Eiten (1955), Lourteig (1979), and Watson (1989). Closely related taxa of the western USA are Oxalis cali-

fornica (Abrams) R. Knuth, O. albicans Kunth, O. pilosa Nutt. ex Torr. & Gray, and O. suksdorfii Trel.—these

J. Bot. Res. Inst. Texas 3(2): 727 - 738. 2009

728 [| | -.fekn Das : iD a ee ORA

f Texas 3(2)

also have been treated in vari i (e.g., Wiegand 1925; Lourteig 1979; Turner 1994).

Useful illustrations are Proen by Mns (1958) a and Eiten oe

In the present study, s] ically int ling little or not at all with

other taxa—sympatry of such taxa is taken as evidence of their pre elation: With further study,

beyond the scope of this overview, complexities in reproductive biology probably will show the taxonomic

framework advanced here to be aig: P in eae aa a and O. dillenii, pee pr odd-

polyploid, even-polyploid, an reported; polyploid

reported in O. stricta. Eiten's x ai i MM ina ey homostylous species (O. corniculata, O.

dillenii, O. florida, O. stricta) are p

ts when self-pollinated—pollination and pollen tube

growth resulting only in ERC pera fertilization. Among these, at e - florida appears to be a facultative

apomict—emasculated fl t ndal

producing sexual seeds when

pollinated with acceptable M Gave Doust et al. 1981). Sexual seed production from outcrossing may

occur in the other species as well. Species with strong morphological heterostyly (e.g., O. grandis, O. priceae)

are consistently outcrossers. Artificial hybrids are easily produced and natural hybrids in various combina-

tions have been identified by Eiten (1963).

pup 0313713 a

Ir y tively estimate ene specimens os did

ts for herbaria consu ted). Both Eiten and I tated many s

here, especially at GH, SMU, and TEX-LL, so the current “author is (aciiliattr in detail with their concepts of

the taxa involved. In 2009, the author has studied natural populations of sect. Corniculatae in Arkansas,

Louisiana, Massachusetts, Missouri, Oklahoma, and Texas.

KEY TO THE YELLOW-FLOWERED CAULESCENT OXALIS SPECIES OF EASTERN NORTH AMERICA

1 [qo J ine.

«LIII VVVII ly

2. Flowers 1 or 2- 3(-5, ius to 8) ir in “‘umbelliform cymes, homostylous; petals 5-11 mm long, completely

ellow, without red lines 2. Oxalis dillenii

2. Flowers (2-)3-5(-8) in umbelliform cymes, distylous; petals (10-)12-16(-17) mm long, yellow, s

prominent e stripes at the base (corolla Taraa

4. Oxalis texana

J +1. F| J ]

y on pedut r pedicels.

4- +1

E^ Petals 10- 201 mm nlons yellow with p inent red SHE at the |

4. Stems densely and pilose wi eadit hai

an manvidLa! plant and lignescent. Bi ligneous flowers 1 or (2 a 2 li umbelliform cymes above the

level of t

ith i —

kl ALII

| y red-lined within; petals 0 mm long " Pens priceae

. a 4 EU AON

J | a H A tor L

LET

enm» OF uelisely p

"and nonsaptate hairs; stoloniform rhizomes usualy n ed y few. herbaceous Or ignescent; flowers Eei

2-4(-8) in regular or irregular cymes, above o

a to strongly red-lined within; petals 10- 18 mm long.

Plants arising from sender lignescent, A OROT Talzomies without tubers; leaflets pal upper

| shoulders usually roun

ievel of the leaves; ien 10- 14 mm long, throat yellow to very faintly or weakly red-lined within

7. Oxalis grandis

5. Plants arisi gfi lender, herbaceous, stolonif hi tint l luci hite, horizon-

€

tal, f L PESSE a val fl--—- ^l

Pi f el

nee produced esty within the level lof the Ne petals 12-18 mm long, throat strongly red-

lined within

3. Petals 4-9(-11) mm long, yellow, without red lines.

6. alll un and raGildting | from taproot, Toong 2H most nodes; iih brown, transverse bus not

8. Oxalis illinoensis

Oxalis corniculata

6. Stems eee usually arising singly from the base, rarely leaning a and decumboent. not trong at the

nodes o!

without free margins or distal sudan.

7. Stems a acc cm eno sparsely to VEDI Spare pilose wi with ne hairs or a mixture of

densely rising singly from the base from

a short herbaceous to lignescent rhi NES usually G -)5- ae in regular (rarely irregular)

cymes; capsules villous to puberulent lous to glabra 6. Oxalis stricta

reduced al id

Nesom, T f vellow-fl | caul t Oxali 729

J "££

7. SI es ca oe cm, sparsely pilose itl ptate hai | glabrous, aris-

g t stol fl 1 or 2(E3, aya in umbelliform

y i | gl t | y puberulent, not villous 5. Oxalis florida

1. Oxalis corniculata L., Sp. Pl. 1:435. 1753. Xanthoxalis corniculata (L.) Small, FL. S.E. U.S. 667. 1903. ` ‘Tre: ITALY: From a

wild source in Florence (“Florentia spunti,” as recorded by Burser),” Herb. Burser XVIII(2), fol. 60 (UPS-BUR , Watson

1989; see Jarvis 2007: Photo in Watson 1989, p. 357, Fig. 7). The protologue notes “Habitat in Italia, Sicilia.”

Oxalis corniculata var. atropurpurea Planch. in Van Houtte, Fl. Serres Jard. Eur. 12:47, pl. 1205. 1857. Tyre: Lourteig 1979, pl. 205 in

Van Houtte ).

Oxalis aaa var. ee (Small) Wieg., Rhodora 27:121. 1925. Oxalis € d Fedde d Bot. -o 32:410. 1905.

667, 1332. 1903. Tyee: U.S.A. LOUISIANA artinsville, in low woods,

15 Mar pa A B. Langlois s.n. (4OLOTYPE: NY, NY digital image!).

repens (Thunb.) Zucc., Denkschr. Koeingl. Akad. Wiss. München, ser. 2, 1:230. 1831. Oxalis repens Thunb., Oxalis

16. 1781. Tyre: Africa. Cultivated at Uppal, Thunberg 11118 (HoLoTYPE: UPS-THUNB?).

Oxalis corniculata var. viscidula Wieg., Rhodora 27:121. 1925. Tyre: U.S.A. Massacuusetts. Hampshire Co.: Northampton, 1902, Mrs.

£: GHI).

inue LOTY

Bieb.) Hohen., Bull. Soc. Imp. N li M 11:395. 1838. Oxalis villosa M. Bieb., Fl. Taur.-Caucas.

1:355. 1808. Tyre: ALB. ANIA Protologue: “Habitat in Albania ibericae lapidosis i is. D. Steuen

P] ial, caulescent, arising from thin to lignescent taproots. Stems 4—10(—30) cm, several and radiat-

ing all fom the taproot, creeping, rooting at the nodes and stolonlike, prostrate to distally ascending-

erect, initially herbaceous but lignescent, sparsely and loosely strigose to strigose-villous. Stipules present,

membranous, margins with free flanges, distal auricles free. Leaves basal and cauline; leaflets 3, obcordate,

lobed 1/5-1/3 length, (4-)6-12 mm, green on both surfaces or bronze-purple to maroon, margins often

prominently villous-ciliate, petioles 1-5 cm. Flowers 1 or 2-36) in irregular or umbell cymes, mostly

homostylous; peduncles (1-)2—-4(-8) cm; pedicels in fruit horizontal to deflexed; petals 4-8 mm, yellow.

Capsules angular-columnar, gradually or abruptly tapering to apex, 8—17(-20) mm, sparsely puberulent to

glabrate or glabrous. Seeds uniformly brown, transverse ridges brown. 2n = 24, 36, 42, 44, 48.

Flowering Mar—Aug, sporadically all year in Florida. Gardens, greenhouses, lawns, fields, roadsides,

hammocks, beach margins, open pine woods, grasslands; 10—500(-2500) m; Nfld.!, Ont.!, P.E.I.!; Ala.!,

Ark.!, Calif.!, Colo.!, Conn.!, D.C.!, Fla.!, Ill.!, Ind. (fide Kay Yatskievych), La.!, Me.!, Mass.!, Mo.!, N J.!, N.C.!,

Oreg.!, Pa.!, S.C.!, Tex.!, Vt.!, Va.!, W.Va.!; introduced; native to Mexico, West Indies, Central America, South

America; introduced Europe, Asia (India, China, Japan), Africa, Pacific Islands, Australia. The PLANTS

Database shows records for Ariz., Ga., Nebr., Ohio, Okla, S.Dak., and Wash.

Many infraspecific taxa of Oxali iculata in the broad have | described over its cosmopoli-

tan range, but their taxonomic status is uncertain and only the single entity is treated here. The reported

variation in ploidy level is a concomitant of the complex morphological variation.

Oxalis corniculata in the USA is recognized by its relatively small flowers, sparsely hairy stems creeping

and rooting at nodes, all procumbent and radiating from the taproot, and its well-developed stipules with

broad, free marginal nanges im auricled apices. Peduncles and E or 2-3 leaves are eae at the nodes,

short erect stems rarely ls but often | t-lived y tl gh the colonial

habit. Stems of O. dillenii sensu stricto may be decumbent or prostrate and rooting at the nodes, but they

almost always are ligneous to lignescent, not evidently radiating from the taproot, and erect stems charac-

teristically arise from the nodes. According to Eiten (1963), O. dillenii is able to form “vigorous, floriferous,

but sterile hybrids with O. corniculata.”

Plants of Oxalis corniculata with bronze-purple to maroon leaves and pubescent capsules have been

aaa as O. corniculata var. ince age ae (e.g., in Florida, Ward 2004; in California, Abrams 1951). Such

tly occur sy lly with the typical expression and it is not clear whether they are popu-

ladional v variants or whether hey are at least partially reproductively isolated. In Malaysia, var. atropurpurea

differs from typical O. corniculata in karyotype as well as in floral and vegetative morphology and is isolated

by post-pollination reproductive barriers (Nair & Kuriachan 2004)—at least in that region its biological

behavior indicates that var. atropurpurea should be treated at specific rank. Distinctive Australasian vari-

730 t tani itute of Texas 3(2)

ants sometimes identified as O. corniculata have recently been a as distinct species (e.g., de Lange et

al. 2005). A form of O. corniculata is common in central Mexico (coll from Hidalgo, Jalisco, Edo.

Mexico, Michoacan, Morelos, and Querétaro)—these plants produce large, prominent stipules and nearly

glabrous stems, but the habit varies from puteum to ascending and the stems rarely root at the nodes.

Most of the closest IEEE of Oxalis (in its American expression) occur in the Americas and

West Indies and the sp bably is native tl though probably seuils of the United States. In the USA,

O. corniculata occurs Boss in ban and highly disturbed habitats, but along the Gulf coast it occasion-

ally grows in less obviously disturbed sites. North of the Gulf Coast, its occurrence is mostly restricted to

greenhouses and horticultural sites (lawns and gardens), suggesting that it is repeatedly introduced rather

than persisting (and expanding) through seed production.

2. Oxalis dillenii Jacq., Oxalis Mon. 28. 1794. Oxalis corniculata L. var. dillenii Jacq.) Trel. in A. Gray, Syn. Fl. N. Amer.

1(1):365. 1897. Xanthoxalis dillenii Jacq.) Holub, Bot. Kózlem. 59:38. 1972. Tree: Jacquin nee in the Lines Moss non

vidi. Omnia ex Dillenio desumpsi" (Plant not seen. It is enti s ]

Pd il — d AE in den PEE a » 298, t. 221. 1732. ui pau uic pesa Was ines from. seed

] lf Dillenius),

4,1

Neil Anas (Reveal 1983) Tt tl i in Dilleniuss herbari ler his pł "Dye lutea ameticab

humilior et annua,” Dillenius s.n. (oror: OKE).

1 Ts;l1 x 1 b "m TH

Eiten dnd » "di PESTE Bap" I

y the plant Dilleni 1 (D & Vines 1907, bs 176), and therefore if daria is mE areal

Watson (1989, p. 361) al 1 tl Dill | Į

work [i.e., cited Eine: ” Lourteig (1979) cited “Tiro: America, Hort. Elth. OXE”

Oxalis dillenii Shi , Field & Lab. 24:39. 1956. Tyre: U.S.A. Texas. Freestone Co.: 4.8 mi SSE of Streetman, 28 Apr 1956,

T H C): 440m f TET Uy F

Oxalis lvonii Pursh, Fl. Xie Sept. 1: 322. 1814 118131. Oxalis doia L. var. lyonii (Pursh) Zucc., Denkschr. Königl. Akad. Wiss.

Munchen ser. 2, 1: 230. 1831. Tyee: U.S.A. Georaia. [Camden Co.]: Cumberland Island, Jun [ca. 1800-1813], John Lyon s.n. (herb.

Lyon, lost or destroyed, fide Ewan & Ewan 1963). Neorvre, designated here: Florida. [Duval Co.:] Near Jacksonville, “188-”, A.H.

Curtiss s.n. (NCU 69362, digital ae Curtiss made ati na Runs ione cmd at least in 1893, 1894, ae 1897, ma

1898; the NCU sheet has only “188” printed, lac! ] te. Jacksonville, in Duval Co., Fla

Co., Ga., separated only by about 25 mi, pe in aS Co. Fla.

The EU ai dela Hee ird E one ocius E MODE did a collection selected from southern Mississippi, in

conflict y in southeastern Georgia (see comments by Ward

E 6 5 7 E IE

2004) 1; 1.11 T : A a fry texana, :4 roe ae | A + Texas, ennthern Arkansas,

and west and central Louisa Dari leon is om Mississippi (Harrison oe Henderson Point, 18 Mar 1954, Demaree

34767-GH!; amid m "p isa ien of O. allen: (as nee Se Eiten 1963 and as annotated by him in 1965; confirmed as O.

2009 na.

Oxalis recurva Ell. Val. floridana Sin Bhodors 27: 138. 1925. Tyee: U.S.A. Fronma: East Florida, no date, D.C. Eaton s.n. (HOLOTYPE:

Oxalis stricta var. piletocarpa Wieg., Rhodora 27:123. 1925. Tyre: U.S.A. New Hawesuire. [Cheshire Co.]: Alstead, old gravel pit, 1901,

E.E Williams (HoLotyee: GH!).

Plants perennial, caulescent, arising from a ligneous or lignescent rhizome, sometimes appearing taproot-

like. Stems 10—25 cm, proximally ligneous to lignescent, 1.5-2 mm thick, usually 2-8 from the base, erect

initially, often becoming decument to prostrate and rhizome-like, sometimes rooting at nodes, strigillose

to strigose with antrorsely appressed, nonseptate, sharp-pointed hairs. Stipules usually greatly reduced at

least above midstem, margins narrowly flanged or without any free portion, without free apical auricles.

Leaves basal and cauline; leaflets 3, obcordate, lobed 1/5-1/3 length, (4-)6-15(-21) mm, green on both

surfaces, glabrous adaxially, sparsely strigillose abaxially, petioles 1-4 cm. Flowers 1 or 2-3(-4, very rarely

to 8) in umbelliform (rarely irregular) cymes, mostly homostylous; peduncles 1-6(-10) cm; pedicels in fruit

deflexed (to horizontal), usually without bracteoles; petals 5-11 mm, yellow. Capsules angular-columnar,

abruptly tapered at apex, 12—20(-25) mm, densely strigose-pilose with mixture of appressed and spreading

hairs, with a puberulent understory. Seeds brownish, tranverse ridges with strong grayish or white lines.

2n - 18, 20, 22, 20-24.

Flowering Feb—May(—Oct). Pastures, roadsides, lawns, river bottoms, sandy, rocky, or gravelly soils;

Nesom, T f vellow-fl ] | t Oxali 731

5-300 m; N.B.!, N.S.!, Ont.!, P.E.L!, Que.!, Sask.!; Ala.!, Ark.!, Ariz.!, Colo.!, Conn.!, Del.!, D.C.!, Fla.!, Ga.!,

Ill.!, Ind.!, lowa!, Kans.!, Ky.!, La.!, Maine!, Md.!, Mass.!, Mich.!, Minn.!, Miss.!, Mo.!, Nebr.!, N.H.!, N J.!,

N.Mex.!, N-Y.!, N.C.!, N.Dak.!, Ohio!, Okla.!, Pa.!, R.L!, S.C.!, S.Dak.!, Tenn.!, Tex.!, Pus Vt.!, Wash.!, W.Va.!,

Wis.!, Wyo.!; introduced in Bermuda, Europe. The PLANTS Datab ] f Alta, Man., Idaho,

and Oreg.

Decumbent branches Dr Oxalis dillenii often ap toloniform, producing erect pratiches Ha leaves

7] f , small

at the nodes, rarely | lventiti ts. Such pl

corniculata, but they differ in | their overall habit, ligneous stems aud rhizomes, reduced stipules, TAM

cauline vestiture, PESE fruit vestiture, and seed color. In O. dillenii, the stems are consistently strigose,

hile the | ly are pilose-villous, and in a quick scan of a specimen not carefully pressed,

stem — mm be urne

Oxalis dillenii may continue to flower, or begin to reflower, in moist places especially in the fall. Plants

are often abundant in lawns and other mowed areas, and later-flowering plants commonly produce very

small flowers. The species shows much plasticity in habit, some of which may be genetically partitioned, as

dysploid chromosome races apparently exist.

Oxalis dillenii and O. stricta are the two most abundant Species of the eastern United States and both

commonly in disturbed sites. In view of recent l confusion [ see commen ts below, under

O. stricta), a key that separates these two is p here.

ef -— I

1. Stems usually 2-8 from the base, 10-25 cm, erect initi

like,

g at nodes; stem vestit trigose t strigillos from base to peduncl | pedicels

| |, nonseptate, sharp- ted | | t t lucing lig

or rlignescentr rhizomes or stolons newer ] Eres ae 5, rarely to 8) in umbelliform cymes Oxalis dillenii

ally arising singly fron dia o! 20-60(-90) cm, erect or later leaning or fall-

ing o cene and d P t vestiture ed 5palselyt tO sparsely d madera pes or ia Fd non-

E rf

fomathin: P | to lig t rhi ; fl ally (3-)5-7(-15) in 1 regular (rarely irregular)

cymes Oxalis stricta

3. Oxalis priceae Small, Bull. Torrey Bot. Club 25:612. 1898. Xanthoxalis priceae (Small) Small, Fl. S.E. U.S. 668. 1903.

Tyee: U.S.A. Kentucky, Warren Co.: Bowling Green, rocky hillsides, open woods, fields, and roadsides, 6 Nov 1898, S.E Price s.n.

(LECTOTYPE, Eiten 1963: NY, NY-digital image!).

Oxalis caespitosa Raf., New Fl. N. Amer. 2:27. 1838 (non O. caespitosa A. St. Hil. 1825). Tyee: U.S.A. Kentucky: glades of West Kentucky

and Tennessee, May—Jun 1823, C.S. Rafinesque s.n. (LECTOTYPE, Lourteig 1979: P).

Oxalis macrantha (Trel.) Small, Fl. S.E. U.S. 667. 1903. Oxalis corniculata L. var. macrantha Trel., Mem. Boston Soc. Nat. Hist. 4:88, pl.

11, fig. 5. 1888. Oxalis recurva Ell. var. macrantha (Trel.) Wieg., Rhodora 27:138. 1925. Tyre: U.S.A. ALasama. Alabama, no date, P

Hatch s.n. (Lectotyre, Lourteig 1979: GH).

Oxalis hirsuticaulis iene Bul a Pd bs 2 ODE add TYPE: it iege DONOR Co.: Nashville, [no date], A. Gattinger

| O. priceae—adoption of the latter here

Aa Eitens Ba aeie in 1955.

Oxalis recurva Ell. var. macrantha (Trel.) Wieg. forma sericea Wieg., Rhodora 27:138. 1925. Tree: U.S.A. ALapama: no other data, Dr. Cabell

s.n. UE GH)).

X Small, Fl. S.E. U.S. 668, 1333. 1903. Oxali (Small) Eiten, Amer. Midl. Naturalist 69:302. 1963

Tyre: U.S.A. Georaia. DeKalb Co.: On and about Stone ML. 100-1500 ft, 1-16 May 1895 , J-K. Small s.n. (HoLotyre: NY, NY digital

image!; isotypes: E, NY, NY digital image!).

form i offset like) hi M and produci t stems from the nodes. Stems proximally

Tel s]

, caulescent, arising from a ligneous

t taproot, usually with lignescent, stoloni-

lignescent, Euer 2—8 from the base, 5—20(-40) cm, erect or ul becoming decumbent, villous-hirsute

with nonseptate hairs spreading or deflexed or spreading in ia orientations. ene with margins

obcordate, lobed 1/5-1/3 length, 3.5-12 mm, green on both Mrd strigose-hirsute on both sores less

commonly glabrate, petioles 2-7 cm. Flowers 1 or (2-)3-8 in umbelliform cymes, les ly irregular

cymes, produced well above the level of the leaves, distylous; peduncles (3-)5- 10(-15) cm; pedicels in fruit

narrowly flanged or without any free portion, without free apical auricles. Lea cauline; leaflets 3,

732 | I tanical Institute of Texas 3(2)

deflexed to horizontal, often bracteolate; petals (1314-20 mm, yellow, with prominent red stripes at the

base (corolla throat). Capsules angular-cylindric, abruptly tapered at apex, 10-15 mm, sparsely to densely

villous with long, deflexed, nonseptate hairs, less commonly puberulent with short, straight, deflexed hairs.

Seeds usually with white transverse ridges. 2n = unknown.

Flowering Mar—May. Dry limestone areas, glades, cedar barrens, chalk prairies, limestone bluffs and

outcrops, sandstone cliffs, rocky slopes, talus, sandy hedgerows, oak-pine, longleaf pine; 5-300 m; Ala.!,

Fla.!, ie | Ey Miss.!, n s | S.C?, Tenn.!; Mexico SEND León».

tive species of tl theastern USA, recognized by its villous to villous-hirsute

stems, lowers in umbelliform cymes, and large yellow to yellow-orange corollas with red lines in the throat.

The lines in the throat remain visible after drying and usually can be seen on herbarium specimens even

from the outside of ne Omer A similar pattern also occurs in O. texana, O. illinoensis, and O. grandis.

Plants of Oxali Mississippi and eastern Louisiana grow in pineland and have slightly smaller

corollas than those "et Vouchers for Lousiana records are cited here. Jefferson Par.: at Gretna, opposite

New Orleans, 6 May 1899, Ball 352 (GH); Rapides Par.: Red River valley, near Zimmerman RR Sta., frequent

in dryer spots beneath tall pines on crest of hills, 24 Apr 1948, Ewan 17608 (MO); St. Tammany Par.: 5 mi

S of Pearl River, US 90, hwy embankment, 30 Apr 1953, Ewan 18546 (GH).

Two collections from a single area in North Carolina are typical Oxalis priceae, at the northeastern

extremity of its main range. Stanly Co.: rocky slopes below bluffs of Yadkin River, near Charlotte, 20 Apr

1932, Palmer 39985 (GH); steep moist banks above the Yadkin River, just above the second or lower power

dam east of Badin, 10 May 1963, Wilbur 6826 (GH). Similarly, a collection of Oxalis priceae from eastern

Tennessee appears to be disjunct. Cocke Co.: near Del Rio, sili bluffs along Newport Rd., 18 Apr 1963,

Sharp et al. 17255 (TENN digital image!). Further study may fill in the known distribution, especially in

Georgia and South Carolina, so that the Cocke Co. and Stanly Co. localities no longer appear disjunct.

Two collections from montane Nuevo León, Mexico, are unmistakably Oxalis priceae, disjunct from

the closest localities in the southeastern USA by about 900 kilometers. Nuevo León. Mpio. Santiago, along

the Cola de Caballo-Laguna Sanchez road into high sierra SE of Monterrey, between Puerto Genovevo and

La Cienega, ca. 1.5-2.5 air km NW of Puerto Genovevo, temperate pine-oak-hickory dominated by Pinus,

Carya ovata, 1500 m, 15 Mar 1994, Mayfield 1887 (TEX); Mpio. Bustamante, along switchbacks of road be-

low the Grutas de Bustamante, ravines and bluffs along road from opening of cave to ca. 4 road km below

cave, Quercus-Cercis-Brahea-Ungnadia-Fraxinus greggii, dark, rich, rocky soil in talus slopes, 1000 m, fairly

common in moist ravine to E of cave opening, 18 Mar 1994, Mayfield 1908 (TEX).

4. Oxalis texana (Small) Fedde, Justs Bot. Jahresber. 32:410. 1905. . Aan texana Small, pu: E U.S. 667, 1332.

1903. Oxalis recurva Ell. var. texana (Small) Wieg., Rhodora 27: 138. 1925. Oxali r. Midl

Naturalist 69:301. 1963. Tyre: U.S.A. Texas. Brazoria Co.: Alvin, 20 Apr 1894, E. N. Plank s.n. (notLorvee: NY, ? ¡Y digital i

Plants perennial, caulescent, DN: tromi a woody taproot, caespitose with stems arising from a caudex or

with rhizome-like offsets g at the nodes. Stems 5-15 cm, proximally ligneous to lignescent,

erect to ascending, strigose to strigillose with antrorsely appressed to ascending nonseptate hairs. Stipules

usually with margins very narrowly flanged, usually with rounded and slightly free apical auricles. Leaves

basal and cauline, leaflets 3, cordate, lobed 1/5-1/3 the length, (426—-12(-18) mm long, green to purple

on both surfaces, glabrous to sparsely strigose adaxially, sparsely strigose abaxially; petioles 2-6 cm long.

Flowers (2-)3-5(-8) in umbelliform cymes, very rarely irregular cymes, distylous, peduncles 4-10 cm;

pedicels horizontal to deflexed in fruit, without bracteoles; petals (10-)12-16-17) mm long, yellow, with

prominent red lines at the base (corolla throat). Capsules angular-columnar, abruptly tapered at apex,

8-15 mm long, moderately to densely puberulent to puberulent-villous. Seeds brownish, transverse ridges

distinctly whitish. 2n = unknown.

Flowering Mar-May(Jun). Commonly in undisturbed habitats and usually in deep, loose sand, but

also fields, roadsides, and edges and openings in woods, pine, pine-oak, and mixed hardwoods; 10-200 m;

Ark.!, La.!, Tex.!

Nesom, T. f vellow-fl | caul t Oxali 733

vi PRY

Oxalis texana is very similar to O. dillenii—differing from O. dillenii primarily in its more numerous

flowers per inflorescence and larger, distylous flowers with red-lined corolla throats. The distinctive red

striping in the corolla throat remains visible after drying and usually can be seen on herbarium specimens

even from the outside of the flower. Plants of O. texana also are distinct in their relatively larger taproots and

habit either caespitose or with short stolon-like offsets. Plants of O. dillenii with larger flowers on elevated

peduncles might be mistaken for O. texana, yet the two taxa exist sympatrically in the range of O. texana

and it seems clear that they are separate species. Oxalis priceae and O. texana are separate in geography and

morphology.

Lourteig (1979) used the name Oxalis lyonii Pursh for the species identified here as O. texana, and her

illustrations of O. lyonii were drawn from a Texas collection identified here as O. texana (Lourteig 1979, Fig.

7, a-e). Turner et al. (2003) followed Lourteig’s nomenclature and mapped the same species as O. lyonii.

Lourteig neotypified O. lyonii to place it within her concept of O. texana (as identified here) and included

O. [Xanthoxalis] texana in synonymy—her neotype, however, is from southern Mississippi, outside of the

range of O. texana, and is instead a collection of O. dillenii (as noted by Eiten 1963 and annotated by him in

1965; confirmed here as O. dillenii). Further comments regarding O. lyonii are in the synonymy of O. dillenii

in the present paper.

5. Oxalis florida Salisb., Prodr. Stirp. Chap. Allerton, 322. 1796. Terr: f ls f hC bably collected

by John Fraser, aem Hortus Chapel Allerton, 1789, Salisbury s.n. (nororvre: BM; isorvee: G-DC). Fide Lourtéig 1979.

Oxalis cabs Small, N. Amer. Fl. 25:52. 1907. Type: U.S.A. New York. Richmond Co.: Giffords, Staten Island, 19 May 1889, N.L.

n. (Horory: NY, NY digital image!).

Oxalis fine Small i in Britton & Brown, Ill. Fl. N. U.S. 2:346. 1897. Oxalis florida Salisb. var. filipes (Small) Ahles, J. Elisha Mitchell Sci.

1964. Xanthoxalis filipes (Small) Small, Fl. S.E. U.S. 667. 1903. Oxalis dillenii Jacq. subsp. filipes (Small) Eiten, Amer.

Midl. Naturalist 69:301. 1963. Type: U.S.A. NORTH ian Stanley Co.: Falls of the Yadkin River, 100-200 ft, 23 Aug 1894, J.K.

Small 61 (Lectotype, Fiten 1963: NY, NY-digi THE LECTOTYPE: F MO!, MSC, NY, NY-digital image!).

aa prostrata Haworth, Misc. Natur. 183. 1803. Oxalis anida Salisb. subsp. prostrata ay RE Mii 42:156. 1979.

yee: USA, cultivated in England, Haworth s.n. (destroyed). Tyre: U.S.A. Veemi. Henrico t W of

2 Station, 21 Oct 1938, M.L. Fernald and B. Long 9355 (Neortvre, Lourteig 1979:US, US- digital: image!; ISONEOTYPE: GH!). Lourteig

cited the collection date as 21 Sep 1928.

Oxalis recurva Ell., Sketch Bot. S.C. 1:526. 1821. Oxalis florida Salisb. var. recurva (ElL) Ahles, J. Elisha Mitchell Sci. Soc. 80:173. 1964.

Oxalis dillenii Jacq. subsp. recurva (Ell.) C.F Reed, A PILAM 63:411. 1987. Tyee: U.S.A. SouTH Canouma: In cultis at pecus circa

Charleston, Apr, Elliot s.n. (CHARL, photo-GH!). Eiten (1 | , lacking flowers, of O

and stem vestiture that would place it with equal probabili as either O. e.” Teibss. priceae) or O. florida. Geography places

it with the ui nnd Eiten chose to keep it as a “nomen dubium." Elliott's protologue noted "Very common near Charleston,

intermingled stricta, with which it has been confounded."

Oxalis inp dida Raf., New Fl. N. Amer. 2:26. 1838 (non O. rupestris A. St. Hil. 1825). Type: U.S.A. K Kentucky, 1823?, Rafinesque

LECTOTYPE, Lourteig 1979:P).

Plants perennial, caulescent, arising from slender lignescent stolons, youngest plants from short, slender

taproots. Stems usually single from the base, (5—)8—30(—35) cm, erect or rarely leaning and decumbent,

subglabrous to sparsely to moderately pilose-villous with very fine hairs spreading in irregular orientation,

sometimes strigose distally or just beneath the flowers and pilose-villous on proximal portions. Stipules

obsolescent, without free margins or apical auricles. Leaves basal and cauline, leaflets 3, obcordate, lobed

1/5-1/3 length, 4-11 mm long, green on both surfaces, sparsely strigose abaxially; petioles 2-5 cm. Flow-

ers | or 2(-3, rarely 4-6) in umbelliform cymes at level of the leaves or slightly above, tristylous; peduncles

(Q-)3-8 cm; pedicels reflexing to ascending, often bracteolate; petals 5-9(-11) mm, yellow. Capsules

angular-cylindric, 8-12(-15) mm, glabrous to sparsely puberulent. Seeds brownish, including transverse

ridges. 2n - 16

Flowering Mar-May(-Aug). Low woods, swamp forests, rich woods, pine woods, sandy sites, burned-

over woods, ditches, roadside banks, floodplains, low fields, lake edges, creek banks, pastures, disturbed

sites, bluffs, rocky slopes; 10-350 m; Ala.!, Ark.!, Conn.!, D.C.!, Fla.!, Ga.!, Ind. (hide Kay Yatskievych), Ky.!,

La.!, Maine!, Md.!, Mass.!, Miss.!, Mo.!, NJ.!, N.Y.!, N.C.!, Pa.!, S.C.!, Tenn.!, Vt.!, Va.!, WVa.!

I [| fal Dos A In LI ga de ET ía

734

Oxalis florida is recognized by its mostly erect stems, sparse and spreading cauline vestiture without

multicellular hairs, obsolescent stipules, relatively small, and yellow flowers without red lines in the throat;

the distal stems and peduncles are thin compared to other species. It is a species primarily of the Atlantic

states and Gulf coast, much more Ty represented in more inland regions. It is known from a cluster of

counties in southeastern Missouri (Butler, Carter, Ripley, and Wayne cos., MO!), and from southern Arkansas

(Ashley, Bradley, and Ouachita cos.; BRIT!, NLU!) northward to apparently isolated localities in Arkansas

(e.g., Yell Co.; MOD. The voucher for the West Virginia record perhaps was a waif—Tucker Co.: Otter Creek

Lumber Co., near Hendriks, dry grounds along railroad track, 10 Sep 1904, Greenman 400 (GH). A collection

annotated by Eiten as O. dillenii var. filipes is interpreted here as O. dillenii with slightly reduced vestiture,

perhaps resulting from damaged stems: Polk Co: 12 Apr 1941, Tharp s.n. (GH).

Lourteig (1979) also treated Oxalis florida at specific rank, and for Florida, Ward (2004, p. 35) noted

that “the differences between O. dillenii subsp. dillenii and subsp. filipes (= O. florida) “are appreciable and

intermediates seem few.” In contrast, Eiten (1963, p. 268) observed that Oxalis dillenii subsp. filipes “is dis-

tinctive in its most epee form but intergrades with [subsp. dillenii], both in forming intermediate

g I o, in disturbed ground, variable pe swarms. The most distinctive por-

tion of this variable sul ies is ntrated in the northern Appalachi The morphological evidence

is conflicting as to sneer it originated from [subsp. dillenii] or directly from [O.] corniculata.”

Wiegand (1925, p. 124), preceding Eiten, also observed a measure of intermediacy in Oxalis florida,

noting that “O. florida and O. filipes have much the appearance of hybrids between [O. stricta and either O.

dillenii or O. corniculata], as no new cl are found in either species. The frequency of their occurrence

in the east and absence in the west, where the possible parents both occur is against this hypothesis.”

Ward (2004, p. 35) noted that Eiten was incorrect in claiming that the name Oxalis florida Salisb. is

illegitimate. “The name [phrase-name] cited in synonymy by Salisbury (1796) was pre-Linnaean which,

since not available for his use, in not disturb BE Mer Ad of O. florida."

In Eiten's view (1963), th bet calis dillenii subsp. filipes and O. pri bsp. colorea was

primarily in flower size. aA filipes keyed under *Flowers 13 mm dona or less,” while subsp. colorea keyed

under "Flowers 10 mm long or more." The seemingly arbitrary difference in size apparently is reflected in

his comment that he identified a ee ©) of the pel in Xanthoxalis d as uS ni dd or ee

pkg | _ [have enna

)

—

In the

between O. florida and O. priceae, and the reality of subsp. cola remains sone ats in dubi. even though

itis placed here as a synonym of O. priceae. Mulcahy (1964, p. 1048) found that both O. priceae sensu stricto

and O. priceae subsp. colorea exhibit a high degree of self-fertility, the former strictly distylous, the latter

tristylous. "A further difference between the 2 subspecies is n pne ed forms very dense and

extensive clones, some of which extend for several meters, while subspecies rather diminutive

clones, very few of which contain more than 3-4 flowering stalks.” Ward did not find significant variation

within O. priceae, listing Xanthoxalis colorea as a synonym.

6. Oxalis stricta I. , SP. Pl. 1:435. 1753. Fre: Morison, Pl. Hist. Univ. 2:184, s. 2, t. 17, fig. 3. 1680 (LecrorvrE. Eiten 1955).

The ps an an patei in ia rad

; ; To fant TONER pl F +] China, | ppli ] 1) O. stricta to the

species identified by Eiten (1955, 1963) and ie as o. dillenii. T basis for the diff lies in selecti f1 for O. stricta

Lourteig (1979), and earlier Robinson oe opined e HEBR pe with Gronovius and was most likely to have been

familiar. with a John Cl yt 4, BM, ee bik. BM- Aa M cn A

11 0) dilleni Fiten (1955), foll 1 by Jarvis (2007)

of O. stricta best cl t lized and weedy in Pope and probably known first-hand by Linnaeus. The

1 1 1I +1 31 1r ] fA AKA,

SUIT ar zr Dy Ward AVUT).

Oxalis mie m s Torrey Bot. Club 25:611. 1898. Oxalis europaea var. bushii (Small) Wieg., Rhodora 27:135. 1925. Oxalis stricta

var ll, Mich. Acad. Sci. Rep. 20:183. 1918. Oxalis fontana var. bushii (Small) H. Hara, J. Jap. Bot. 24:106. 1949.

Tyre: U.S. A Missouri. Jackson Co.: NW Jackson Co., dry ground, 28 May 1893, B.E Bush 30 (HoLotyre: NY, NY-digital image!;

ISOTYPE: GH!).

Oxalis coloradensis Rydb., Bull. Torrey Bot. Club 29:243. 1902. Tyre: U.S.A. COLORADO. Larimer Co.: Sangre de Christo Creek, 2400-2700

Nesom, T f vellow-fl | caul t Oxali 735

m, 2 Jul 1900, PA. Rydberg 5920 with EK. Vreeland ( NY, NY-digital i NY, NY-digital image

Oxalis cymosa Small, Bull. Torrey Bot. Club 23:267. 1896. Oxalis europaea TM rosa a (Small) WB: HAN ii 135. 1925. Tyee:

A, NEw ee oe Co.]: Riverdale, 26 Aug 1894, E.P Bicknell s.n. (p E g

AT L]: Arch El F m Allemagne 309 1854 Ter: A ] t 1 for thi qum reckoned

t t] i the American ones identi fied. as Oxalis stricta L. and because Linnaeus had

that (is ES plants

indicated that O. stricta i pie rd nm he diode a new name for i Maur nese ones. nah noted en a specie “est si

z t

n la trouve si l

JA lm i 2 ali + > BLA mT] “fe : J]: : [a

+ +

this. P Wiegand (1925), Fiten (1955), I teig (1979) p ided inf ti the tvvification of O europaea

ee (1927, p. 136) noted that “O. europaea is with little doubt ive of A ica, but j luced into Europe about 1658

A Adit A f +1 ] 1

It seems to MAE a inda d Morison in 1660 and was figured by him in his Historia y history of this

149],

Oxalis europaea Jordán forma ii Wieg., Rhodora 27:135. 1925. Tyre: U.S.A. Missouri. Jackson Co.: Courtney [Greenwood], 20 May

1912, B.E Bush 6701 (HoLorYPE: GH!; tsotyres: NY, US, US-digital image!).

Oxalis A forma eine Wieg., Rhodora 27:136. 1925. Tree: USA. Iowa. [Story Co.]: Ames, Pammel and Ball 4 (HOLOTYPE:

GH!; isorYPE: MO!, MO-digital image!, US, US-digital image!).

Oxalis penne forma vestita Wieg., Rhodora 27:136. 1925. Tre: U.S.A. MassacnuseTts. [Middlesex Co.]: Cambridge, 1904, Miss

I erson s.n. (HOLOTYPE: GH!).

Oxalis sonora forma villicauiis Wieg., Rhodora 27:135. 1925. Oxalis stricta var. villicaulis (Wieg.) Farwell, Amer. Midl. Naturalist

2. 1928. Tyre: USA. Michigan. [St. Clair Co.]: Port Huron, 1914, C.K. Dodge 41 (HoLoTYPE: GH!

ou pa Bunge, Enum. Pl. China Bor. 13. 1833. Tyre: CHINA: "Borealis, Pan-Sham, hab. ad fontam,” 1831, A.A. Bunge s.n. (HOLOTYPE:

P; isotypes: E, P). Fide Lourteig (1979).

Oxalis interior (Small) Fedde, Just's Bot. Jahresber. 32:410. 1905. Xanthoxalis interi ana de S.E. U.S., 668. 1903. Tyre: U. on ARKANSAS.

Benton Co.: no other locality data, 1899, E.N. Plank s.n. (LecrorY?E, Eiten 1955:NY, NY-

Plants annual to short-lived perennial, caulescent, arising from a thin, short rhizome. Stems usually arising

singly from the base cay = begets), erect Or ind leaning or falling over and decumbent, 20—60(-90)

cm, villous with ton stems and petioles, commonly concen-

trated at nodes, sometimes gale on paneles Stipules PM without free margins or apical auricles.

Leaves basal and cauline; leaflets 3, obcordate, lobed 1/5-1/3 length, (8210—-20(-30) mm, light green to

yellowish-green on both surfaces, petioles 2-8 cm. Flowers rarely 1 vo (3-)5-7(-15) in regular cymes,

less commonly in irregular cymes, usually within level of the leaves ghtly above, homostylous or slightly

to strongly heterostylous; peduncles 3-9(-11) cm; pedicels in fruit "in to ascending, often bracteolate;

petals (62)8-11 mm, yellow. Capsules columnar, nearly terete, abruptly tapering toward apex, 8-15 mm,

villous with septate hairs to glabrate. Seeds brown, transverse ridges rarely whitish. 2n = 18, 24.

Flowering (Apr-)Jul-Oct. Prairie ravines, river and stream banks, sand bars, low woods, floodplains,

roadsides, fields, lawns, gardens; 20-1200 m; B.C., Man., N.B.!, Nfld., N.S.!, Ont.!, P.E.L!, Que.!; Ala.!, Ark.!,

Colo.!, Conn.!, Del.!, D.C.!, Ga.!, Idaho, Ill.!, Ind.!, lowa!, Kans.!, Ky.!, La.!, Maine!, Mass.!, Md.!, Mich.!,

Minn.!, Miss.!, Mo.!, Nebr.!, N.H.!, NJ.!, NY.!, N.C.!, N.Dak.!, Ohio!, Okla.!, Pa.!, R.1.!, S.C.!, S.Dak.!, Tenn.!,

Va.!, Vt.!, WVa.!, Wis.!; introduced in Europe, Asia, Africa, Pacific Islands (New Zealand), Australia. The

PLANTS Database shows records from Nfld., Sask., Ariz., Mont., N.Mex., Wash., and Wyo.

Oxalis stricta is recognized by its tall, erect stems from a short, simple rhizome, presence of septate

hairs, cymose inflorescence, and relatively small flowers. Septate hairs on the stems and petioles are easily

crosswalls, but they vary greatly in density, as do the

* |

recognized (lens), especially because of their |]

nonseptate hairs. In “villicaulis” and “pilosella” forms, as well as “var.” bushii, the septate hairs are dense and

evenly distributed on the stems, but more often they are localized around the nodes and intermixed with

nonseptate hairs. Often they are few in number, and, in rare cases, plants with greatly reduced vestiture

= Du a PER za

ts, of Oxalis strict ditscl lati 0) grandis and O. illinoensis

is dine and a basic biological difference: The bases of Oxalis plants often are incon lee. collected,

however, especially in the other species, which commonly develop stoloniform or rhizomiform branches

above the taproot.

Oxalis stricta in western states is uncommon and probably adventive. Reports of the species from Texas

736 t tani titute of Texas 3(2)

(e.g., USDA, NRCS 2009) have not been confirmed in the present study and it is unlikely that such reports

are accurate. The early establishment of O. stricta in Europe is noted above with the type of O. europaea.

Eiten (1963) appeared to suggest that the species may be native in eastern Asia as well as North America.

Eiter's comments (1963, p. 304-305) on variability in t ful and repeated here: “Very

variable, both as between east Asia and North America, and within each felon. I have not seen enough

material from east Asia to determine if it is useful to set up subspecies. In North America all the variation

that is regionally based is too intergrading to set up useful infraspecific categories. The most distinctive

geographically-based character is the presence of hairs on the upper surface of the leaflets. This is most

frequent in the Midwest, but even there less than half the collections in each state have this feature, and

sometimes it varies even within single populations. This and other characters that have been used to set up

infraspecific groups (presence of septate hairs on stems or on pedicels) often vary in the same population

or even on the same plant, and present all intergrades of expression. In addition, the extreme expressions

of characters that vary regionally center in different areas, i.e., they do not vary together. The introduced

European plants represent a small part of the North American variation, in general being less pubescent.

They are not at all like the plants of east Asia."

7. Oxalis grandis Small, Bull. Torrey Bot. Club 21:475. 1894. Xanthoxalis grandis (Small) Small, Fl. S.E. U.S. 668. 1903. Tyee:

U.S.A. PENNSYLVANIA. [Bradford Co.]: Wysox, Jul 1836, J. Carey s.n. (LECTOTYPE, Fiten 1963: NY, NY. digital image!; isoLectoTyPE: GH!).

Oxalis recurva Trel., Mem. Boston Soc. Nat. Hist. 4:89. 1888 (not Elliott 1821). Fide Wiegand (1925).

Plants perennial, caulescent, arising from slender, lignescent, stoloniform rhizomes without tubers. Stems

erect, usually single from the base (rarely 23 together), (10—25-60(-100) cm before branching, nearly

glabrous to sparsely or densely pilose or villous with a mixture of septate and nonseptate hairs. Stipules

absent. Leaves cauline, mostly on the distal half of the stem; leaflets 3, obcordate, lobed 1/5 length, 5 ——

mm, upper shoulders of lobes usually rounded, rarely flattened, green on both surfaces,

narrowly brownish-purple, ciliate, petioles 5-7.5 cm. Flowers 1 or 2—4(—8) in regular or irregular cymes or

umbelliform cymes produced above the level of the leaves, tristylous; peduncles 7-12 cm; pedicels in fruit

erect to ascending; petals 10—14 mm, yellow, throat yellow to faintly or weakly red-lined within. Capsules

ovoid to ovoid-oblong, 6-10 mm long. Seeds brown, traverse ridges rarely whitish. 2n = 28 (hide Weller &

Denton 1976, reporting unpublished counts by Orndulf).

Flowering May-Aug. Sandy woods and alluvial soils; 100—1100 m. Ala.!, D.C.!, Ga.!, Ind.!, Ky.!, Md.!,

N.C.!, Ohio.!, Penn.!, S.C.!, Tenn.!, Va.!, WVa.!

A report of Oxalis grandis from southern Mississippi (Forrest Co.; Carter & Jones 1968) is far out of

range and habitat for the species and probably is based on a misidentification, perhaps of O. priceae, which

also is large-tlowered.

8. Oxalis illi is Schwegm., Phytologia 50:467. 1982. Tre: U.S.A. Imre Macon En by Tenn. [Hwy] 10, S side of

and id P de Fd deep hardwood ravine, a! loam, 5 n 1975 155 , desig

BY. T ft 71 4 show e hi 1 infl Alas, d lecti

from Illinois, but th icall i ter of tl is in eastern EE e the Lan e occurs in A

(comments below). The holosgpew Was — at SIU but cannot now

Illinois. Pope Co.: Wooded floodplain forest along Little Lusk Creek at Marthas Woods, 4 mile ENE of Eddyville, 11 May 1968,

J-E. Schwegman 1661.

Plants perennial, caulescent, arising from slender, herbaceous rhizomes at intervals producing white fusi-

form tubers or tuberlike thickenings. Stems erect, usually single from the base (rarely 2-3 together), 15-40

cm, nearly glabrous or sparsely to densely villous with a mixture of septate and nonseptate hairs. Stipules

obsolescent, without free margins or apical auricles. Leaves EE leaflets 3, obcordate, lobed 1/5 length,

(12—)20-30(-35) mm, upr f lobes flat, green on botl , ciliate, petioles 4—7.5

cm long. Flowers 1—3(—6) in regular or irregular cymes produced — at the level of the leaves, tristylous;

peduncles 3-10 cm, pedicels in fruit erect to ascending; petals 12-18 mm, yellow, throat strongly red-lined

within. Capsules oblong-ovoid, 7-10 mm. Seeds brown, traverse ridges rarely whitish. 2n = unknown.

T. g Il W a [| [| a PF. E 737

Nesom

C a +l Bs E E Pu

Flowering Apr—Sep. Slopes, bluffs, ravines, floodplains, mesic forests,

ground cover, commonly on limestone, shale, or calcareous loess substrate; 200-500 m; s TL l, s Ind.!, w

Ky.!, and c Tenn.!

Differences between Oxalis illinoensis and O. grandis are subile but mi Are to be correlated with

I 3

nmonly CALAS LACAL

hi ara

geography; the tuberous portions of the O. illinoensis

off ms collection: Oxalis illi d oe M western n part of es range of e grandis sensu

lato; the to be closely arely ove n range in Ten-

nessee (Univ. Bi Tennessee Herbarium 2008; persone P by "a of many collections at BRIT/VDB

and MO). Michael Homoya (pers. comm.), of the Indiana Dept. of Natural Resources, observes that the two

species overlap in range in southern Indiana, where they are mostly separated by habitat, with “Oxalis grandis

on the drier, more acidic slopes, and O. illinoensis in the mesic, alkaline environments.” Oxalis illinoensis in

Illinois is known from only three counties immediately bordering Indiana and Kentucky, and is state-listed

as endangered (S1, Illinois Endangered Species Protection Board 1999). John Schwegman (pers. comm.)

observes that he has never seen O. grandis in Illinois, nor have I seen any collections (MO). County-level

distribution records in Indiana and Kentucky shown by USDA, NRCS (2008) need to be reexamined.

Medley (1993) has observed that Oxalis illinoensis and O. grandis intergrade, and a hybrid population is

said to exist in Indiana (Heikens 2003, citing unpublished and undocumented observations by S. Olson).

ACKNOWLEDGMENTS

I am grateful to the staffs at BRIT-SMU-VDB, GH-NEBC, MO, NLU, and TEX-LL for help during visits to

those herbaria and ASU for a loan of specimens of sect. Corniculatae to MO. Kanchi Gandhi and Jim Reveal

gave advice on typification of Oxalis dillenii, Nancy Elder (Librarian, Univ. of Texas Life Science Library)

provided literature relevant to the typification (or lack of it) of O. europaea. Michael Homoya (Indiana Dept.

of Natural Resources), John Schwegman, Steve Hill, and Bob Mohlenbrock provided information on O. il-

linoensis, Gene Wofford provided information and an image of the collection of O. priceae from Cocke Co.,

Tennessee, and Carol McCormick for help with selecting a neotype for Oxalis lyonii. George Yatskievych,

Kay Yatskievych, Gordon Tucker, Bruce Sorrie, Alan Weakley, and Dan Ward reviewed the key and manu-

scripts and gave helpful comments. Comments and critique by Jim Henrickson have sharpened the focus

of definitions of species in this group. Steve Hill and an anonymous reviewer provided useful critiques for

the journal review. This study was done as part of the work under contract for the Flora of North America

Association in conjunction with preparation of the FNA treatment of Oxalis.

REFERENCES

ABRAMS, L. 1951. Illustrated flora of the Pacific States, Washington, Oregon, California. Vol. III. Stanford Univ. Press,

Stanford, Calif.

ASCHERSON, P. AND P. GRAEBNER. 1913. Synopsis der mitteleuropaischen Flora, Vol 7. Verlag von Wilhelm Engelmann, Leipzig.

CARTER, J.W., JR, AND S.B. Jones. 1968. The vascular flora of Johnson State Park, Mississippi. Castanea 33:194-205.

Druce, G.C. AND S.H. Vines. 1907. The Dillenian herbaria. Clarendon Press, Oxford.

Ermen, G. 1955. Typification of the names “Oxalis corniculata L” and "Oxalis stricta L'Taxon 4:99-105.

EITEN, G. 1963. Taxonomy and regional variation of Oxalis section Corniculatae, |. Introduction, keys and synopsis

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CLARIFICATION OF THE TYPIFICATION OF MICHAUX NAMES

IN EASTERN NORTH AMERICAN VITIS (VITACEAE)

Connie Fisk Alexander Krings

Department of em 7 cience mese dll of Plant Biology

North Carolina State University North Carolina State University

Raleigh, North Carolina 27695- pue U.S.A. Raleigh, North Carolina 27695-7612, U.S.A.

Connie Fiskancsu.edu Alexander. Krings&ncsu.edu

ABSTRACT

Fl [. hazel : de AT 1 A , 17:4 1 SIT ] : +1 f.

Tk e Pu

V. aestivalis, V. cordifolia, V. riparia, and V. rotundifolia.

RESUMEN

T E k PEA a og J 1 1 - Va 1] «AE 1 "T Pa | AT my A el 1 :f pa pepe | 1 f.

en la bibliografía: V. aestivalis, V. cordifolia, V. riparia, y V. rotundifolia.

Uttal (1984) typified a large number of Michaux names for North American plants, including several in

eastern North American Vitis L. (Vitaceae). Uttal provided a citation for each name that indicated the card

number in the IDC microfiche set of the Michaux herbarium (IDC 1967) and the photo number on the

card in which the selected element appeared. Uttal noted that: “[...] each such designation is equivalent

to the selection of the specimen photographed as the holotype, if single, or lectotype if one of a series” (p.

5). However, he referred to the specimens only as “Type.” Unfortunately, this leaves readers to speculate

as to whether Uttal considered a given cited specimen as a holotype or a lectotype. Fortunately, Art. 9.8 of

the International Code of Botanical Nomenclature (ICBN; McNeill et al. 2006) allows for the correction of

the mistaken use of a term for type, thus Uttal’s use of the term “Type,” abbreviated throughout his paper

to “T; is an error correctable to “Lectotype.” In 1991, Moore indicated lectotypes for the same Michaux

names in Vitis, incorrectly labeling each as “here designated.” It remains unclear whether Moore (1991) was

unaware of Uttal’s work (it was not cited) or whether he felt the ambiguity between the categories holo- and

lectotype needed correction. In either case, unless demonstrably incorrect, Uttal’s typifications have prior-

ity and must be followed according to Art. 9.17 of the ICBN (McNeill et al. 2006). However, because the

typifications of both Uttal and Moore do not coincide in all cases and as errors (regardless whether made

by printer or author) are found in each, we seek to clarify the typification ascriptions of the four Michaux

names in eastern North American Vitis. The typification statements of both Uttal and Moore are shown.

The accepted typification is bolded.

Vitis aestivalis Michx., Fl. Bor.-Amer. 2: 2 E

Uttal pud. p. is Lectotype ES ee I li ; È IDC Michaux, microfiche no. 122, photo 17!

T Teea] / 13600943

hj me

Vitis cordifolia Michx., Fl. Bor.-Amer. - oe a a de Tun L.]

Pa eee P- es Lectotype vs p P IDC Michaux, microfiche no. 123, photo 3!

(iO AN

Vitis les Michx., Fl. Bor.-Amer. 2:231. 1803.

Uttal (1984): “Type”: “ad ripas et in inusulis fluviorum Ohio, Misisipi, etc.": P, IDC Michaux, microfiche no. 122, photo 20!

Moore (1991, p. 361): Lectotype (as “Holotype”): "ad ripas et in insulis fluviorum Ohio, Mississippi, etc.”: P, IDC microfiche no. 122,

photo 19!, bottom specimen!

Photo 20 of fiche 122 contains no specimens, only labels and descriptive inf ion for the speci in pl 19, including a label

J. Bot. Res. Inst. Texas 3(2): 739 — 740. 2009

dated 1897 (“the two upper specimens seem to be Vitis rupestris Scheele”) and four labels for “Vitis riparia.” Uttal's type citation for

V riparia (i.e., fiche 122, photo 20) includes only the label of the corresponding specimen cited by Moore as the holotype (here

corrected to lectotype). The label reads: “ad ripas et insulas, Ohio, Misissipi.”

Vitis rotundifolia Michx., Fl. Bor.-Amer. 2:231. 1803.

Uttal (1984, p. 62): Lectotype (as “Type”): [Hab. a Virginia ad Floridam] P, IDC Michaux, microfiche no. 122, photo 21!

Moore (1991): “Lectotype”: a Virginia ad Floridum: P, IDC Michaux, microfiche no. 122, photo

Uttal designated the element in photo 21 of fiche 122 as the type of V rotundifolia. B

set, we reasonably assume that Uttal was aware that V. rotundifolia was also represented by another specimen (fiche 123, photo 1).

His selection of the element on un 122, pun 21, as the ' aod ndn E to dd a was a Ape choice as it is a better

learly sl orth Americ ie

J qe +l +i Els

"m v owl

ecimen,

interpret Moore's An cf photo “20” as an mane — error, since fis

only labels for elements mounted on another sheet (as evidenced by his careful discrimination among elements in nthe case of Y

riparia, see above). He accepted photo 1, fiche 123 as a syntype.

ACKNOWLEDGMENTS

We thank Fred Barrie, Eric Wada, and an anonymous reviewer for constructive comments improving this

manuscript.

REFERENCES

INTER DOCUMENTATION Company, 1967. Microfiche set of the Herbarium of A. Michaux, Flora boreali-americana. Zug,

Switzerland. 6211:122-123.

McNett, J., ER. Barrie, H.M. Bunper, V. DEMOULIN, D.L. HAWKSWORTH, K. ee D.H. Mee p Prano, P.C. SiL vA, J.E. SKOG,

J.H. WIERSEMA, AND N.J. TURLAND. 2006. Interna tional code of bot ture ( code). Regnum Veg.

146, A.R.G. Ganter Verlag KG.

MicHAUX, A. 1803. Vitis. Fl. Bor.-Amer. 2:230-231.

Moore, M.O. 1991. Classification and systematics of eastern North American Vitis L. (Vitaceae) north of Mexico.

Sida 14:339-367.

Utra, LJ. 1984. The type localities of the Flora Boreali-Americana of André Michaux. Rhodora 86:1—65.

ADDITIONAL MORPHOLOGICAL MEASUREMENTS OF

ARCEUTHOBIUM SISKIYOUENSE AND A. MONTICOLA (VISCACEAE)

Robert L. Mathiasen Carolyn M. Daugherty

School of Forestry Department of Geography, Planning, and Recreation

Northern Arizona University Northern Arizona University

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

Robert.Mathiasen@nau.edu Carolyn.Daugherty@nau.edu

ABSTRACT

auo A 21 Lt eee PE f. Exe [1z] es : 1

JP. jeffreyi, and P. contorta

Es e e Jo

emi 4 ] : -1 11 +1 1 E 1 to Has 1 $ D ff G | n

var. contorta. , E g jeffres contorta var.

contorta A1 35,5 1 1 1 11 au E. A monticola, +l J = A I PN ET fil wI el

iu e

Siskiyou Mountains. Results d trated that A. siskiy i pholog Uy similar to A. eee: and that A. monticola is

morphologically similar to A. californicum. O for fl ] A. siskiyouense and A

onticola.

-—

TE 1

Key Worps: Arceut

Pinus monticola

, Arceuthobium siskiyouense, morphology, Pinus attenuata, Pinus contorta var. contorta, Pinus jeffreyi,

RESUMEN

] r-.1 Jd Á LL 1 ] 1 4 VE As 1 t A CO

E o ES

California. 1 lici hici bre plant linas y f i iendo sobre Pi tt i Perey, yP. contorta var.

T Js. E = -1 1 1 4 1 J T 1 J Te = T

contorta I E , P I

P. jeffreyi y más pequeño sobre P. contorta var. contorta. Tambié t lici licional bre A. monticola, otro pierda se

enano endémico de las wi eue Add Los resultados muestran que A. QU ence es aie sania ae a A.

i di sla y que A ti g A. californicum. ?

son | tan tant A. siski A. monticola

I 5 I r z 1

PALABRAS CLAVE: A thobi ticola, A thobi ishiy , morfología, Pinus attenuata, Pi torta var. contorta, Pinus jeffreyi,

Pinus monticola

Arceuthobium siskiyouense Hawksw., Wiens & Nickrent (Knobcone pine dwarf mistletoe, Viscaceae) and A.

monticola Hawksw., Wiens & Nickrent (western white pine dwarf mistletoe) are endemic to the Klamath/

Siskiyou Mountains of northwestern California and southwestern Oregon (Hickman 1993; Hawksworth &

Wiens 1996). Prior to 1992, A. siskiyouense was classified as A. campylopodum Engelm. and A. montiolca was

grouped with A. californicum Hawksw. & Wiens. Hawksworth et al. (1992) separated A. siskiyouense from A.

campylopodum based on differences in plant and fruit size, flowering period, host affinities, and the induce-

ment of witches' brooms or not on infected principal hosts (see their Table 1). They separated A. monticola

from A. californicum based on plant and fruit size, plant color, seed dispersal period, and host affinities (see

their Table 2). An electrophoretic analysis of isozymes also supported the separation of A. siskiyouense from

A. pui e ape and A. monticola from A. californicum (Nickrent & Butler 1991). However, recent apad

work has ted that these taxa are closely related and that further study of their I ical and

I e

1 e

vas needed (Nickrent et al. 2004). Because A. siskiyouense commonly parastizes

Pinus ated Lemm., P. jeffreyi Grev. & Balf., and P. contorta Dougl. ex Pond var. contorta within its geo-

graphic range (Matiniasen & Daugherty 2009), we collected ac morphological data for this dwarf

mistletoe from each of these hosts. We also collected additional logical data for A. monticola because

little was known regarding its flower and fruit morphology (Hawkeworth & Wiens 1996). This allowed us

to compare morphological characteristics of A. siskiyouense and A. monticola with A. campylopodum and A.

californicum, respectively, using plant and flower characteristics for all four species.

] Rot Rec Inct Tawas 32): 741 749. 2009

I 1 fal Dos . in LI "LP fot

742 t f Texas 314)

MATERIALS AND METHODS

Cl 1 1 . | E an EL: bh A E 7 : 1 x

In order to make a comparison ol

we sampled 15 pawalanens of the miie (Fig. 1). Plants were collected from Pinus attenuata only Q loca-

tions); from P. jeffreyi only (2 locations); from both P. penu ane P pe. © ae Rus from us A

attenuata and P. contorta var. contorta (4 locations). Fi

2 in southwestern Oregon and 3 in northwestern Coma (Fig. 1). Plants were measured from ie type

localities for both dwarf mistletoes (Fig. 1; locations 5 and 6; Hawksworth et al. 1992). Identification of host

trees was based on Hickman (1993) and Critchfield po er B contorta var. elias

From each population, 10-20 male and 10-20

from each infection was used for morphological measurements. The dwarf mistletoe plant characters mea-

sured were those used by Hawksworth and Wiens (1996) for taxonomic classification of Arceuthobium spp.

The following morphological characters were measured: 1) height, basal diameter, third internode length

and width, and color of male and pd ps 2 mature d oS rus and color; 3) seed length,

width and color; 4) length and width nate spikes; 5) for 3- and 4-merous

flowers; 6) length and width of staminate flower peram Idee and 7) anther diameter and anther distance

from the perianth lobe tip. Plants were measured within 24 hours after collection and were measured us-

ing a digital saan a juri microscope with a micrometer, or diio a 7x Hue eus Fs ein with a

ed t shoot

micrometer. Stamin pike and flower g ie d fruit and

cee D QN were made ous the pes of seed dispersal

ispersal for both dwarf mistletoes were poo (Hawks

— nn! 1996), additi del f the phenology of these taxa we ing the summer

and fall of 1998, 2003, 2004, 2007, and 2008.

RESULTS AND DISCUSSION

Arceuthobium siskiyouense

Plant heights for male and female plants on the pine hosts of Arceuthobium siskiyouense were similar, but

plants were slightly smaller from Pinus contorta and slightly larger from P. jeffreyi when compared to those

measured on P. attenuata (Table 1). The means and ranges for basal diameters and third internode dimen-

sions were also similar for male and female plants from the different pine hosts. Flower characteristics were

also similar, but the diameter of 3-merous flowers was smaller by 0.2-0.3 mm from male plants growing

on P. contorta (Table 1). Fruit and seed sizes followed the same pattern as plant heights; fruits and seeds

were largest on average from P. jeffreyi and smallest from P. contorta. Plant color of both male and female

plants ranged from dark brown to red-brown to brown-green, independent of the host from which they were

sampled. The dark brown to red-brown color of plants was the principal characteristic used to identify A.

siskiyouense at two locations where there were no infected P. attenuata growing near infected P. jeffreyi (Fig.

1, locations 7 and 19).

The mean size of plants reported by Hawksworth et al. (1992) when they described A. siskiyouense was

similar to what we obtained, but we measured plants on P. attenuata that were larger (15 cm) than they

reported (10 cm). We also found that the basal diameters and third internode dimensions of A. siskiyouense

plants were slightly larger than those reported by Hawksworth el al. (1992). Results also indicate that the

size of fruits of A. siskiyouense was larger (5.1 x 3.3 mm) than reported by Hawksworth et al. (1992) for this

species (4.0 x 2.5 mm) or by Hawksworth and Wiens (1996) (3.6 x 2.1 mm).

T lec af Ingi MONET

cies is morphologically very similar to A. campylopodum (Table 2). The latter t iginall; i dA

O ais sapien et al. ee separated the dwarf mistletoe populations parasitizing Pinus at

Oregon at t] ific level. Based on

of A. siksiyouense, the size of plants, flowers, and fruits are only slightly larger than those of A. campylopodum,

the primary morphological difference between these species being plant color. Arceuthobium siskiyouense is

RA athe : T^ 1 4 A 2L L’ e. NP Ba

monticola 743

, L

Curry Co. Josephine Co. ;

O NA

1 A?

X Cave Junction

Brooki 8 GA

rookings

¿qlo 3 OREGON

l Li /

Wie VA J CALIFORNIA

A 12 16 O

PACIFIC 13 15 M 18

14 O 19

OCEAN

Crescent City 20

Del Norte Co. Siskiyou Co.

0 10. — 20

ilb.

(

Fc 1. Approximate location of collection sites in California and Oregon. Open triang

tarada Iskiy lected and | ly Pj ffreyi. Of l | t Ineati | A tien a 5

d on A monticola. Numb pond to the following locati s: 1-Snowcamp Mountain; 2-Chrome Ridge; 3-Lookout Gap; 4-Rock Creek

7-Oregon M in; 8 Bain Station; 9 T thwest of Bain Stati ; 10-Smith River Bridge; 11-Sourdough Junction; 12— Pine Flat Mountain;

13- —High Divide; 14-01d Gasquet Toll Road, near Danger Point; 15—Old Gasquet Toll Road, Elevenmile Creek; 16—0id Gasquet Toll Road, Twelvemile

Creek; 17-5 km north of Black Butte; 18—Black Butte; 19—Bear Gulch; 20-Ship Mounta

ABLE | q y

(range) [n]. Letters in brackets designat ple sizes already listed in ti | Plant heights i

Character Pinus attenuata Pinus jeffreyi Pinus contorta

var. contorta

Plant Height

Male 8.1 (5.1-15.2) [120a] 8.3 (5.1-14.7) [100a] 7.7 (5.0-13.9) [60a]

Female 8.9 (5.3-14.8) [160b] 9.3 (5.8-17.3) [120b] 8.7 (5.7-14.1)[a]

Basal Diameter

Male 3.0 (1.8-4.3) [a] 3.1 (24-6.1) [a] 2.9 (2.1—5.1) [a]

Female 2.9 (1.9—5.7) [b] 3.2 (2.4-5.9) [b] 3.0 (1.9-4.9) [a]

Length of Third Internode

Male 11.7 (7.1-22.3) [a] 12.8 (7.5-22.9) [a] 11.3 (6.3-21.2) [a]

Female 12.5 (7.4-22.8) [b] 14.7 (7.5-25.4) [b] 12.7 (5.8-20.7) [a]

Width of Third Internode

Male 2.0 (1.5-2.8) [a] 2.2 (1.6-3.0) [a] 1.9 (1.2-2.8) fa]

Female 1.9 (1.3-3.0) [b] 2.1 (1.6-3.0) [b] 1.9 (1.4-2.8) [a]

Staminate Spike Length 11.1 (6.0-17.3) [200c] 124 (5.2-19.1) [160c] 10.5 (6.2-15.8) [100b]

Staminate Spike Width 1.9 (1.5-2.4) [C] 2.0 (1.6-2.6) [c] 1.9 (1.4-2.3) [b]

Mean Flower Diameter

3-merous 3.2 (2.6-3.9) [100d] 3.3 (2.7-3.8) [50d] 3.0 (2.5-3.7) [50c]

4-merous 44 (3.7—5.6) [d] 4.5 (3.7—5.9) [d] 4.4 (3.6-5.5) [c]

Perianth Lobe Length 1.5 (1.2-1.9) [c] 1.5 (1.1—2.0) [a] 1.5 (1.1-1.8) [b]

Perianth Lobe Width 1.5 (1.2-2.1) [c] 14 (1.1-1.9) [a] 1.4 (1.0-2.0) [b]

Anther Diameter 0.8 (0.5-1.1) [c] 0.7 (0.4-1.1) [a] 0.8 (0.5-1.2) [b]

Anther Distance from Tip 0.8 (0.5-1.1) [c] 0.8 (0.4-1.1) [a] 0.7 (0.4-1.0) [b]

Mean Fruit Length (mm) 5.1 (4.1-6.1) [110e] 5.3 (4.0—6.0) [80e] 4.9 (3.8-5.7) [c]

Mean Fruit Width (mm) 3.3 (2.6-4.1) [el 5 (2.8-4.3) fe] 3.1 (2.5-3.8) [c]

Seed Length 3.0 (2.1-4.0) [e] 3.1 (2.0-3.9) [e] 2.9 (2.1-2.8) [c]

Seed Width 1.3 (1.0-1.8) [e] 3 (0.9-1.9) le] 1.2 (0.8-1.8) [c]

consistently dark brown, red-brown, or brown-green, and we did not observe any male or female plants of

this mistletoe that we considered yellow, a plant characteristic described for A. campylopodum. However, we

did observe a few male peu of - iic adie that were brown-green and it is difficult to determine if our

use of brown t is different from the olive-green designation used by Hawksworth

and Wiens (1972, 1996) for A. campylopodum. It was also difficult to compare plant dimensions between

these species because Hawksworth et al. (1992, Table 1) and Hawksworth and Wiens (1996, Table 16. 6)

indicate the height of plants of A. campylopodum as averaging 12 cm with a range from 10-14 cm. However,

they describe A. campylopodum plants as averaging about 8 cm in height with a maximum height of 13 cm

in their written descriptions of the species (Hawksworth & Wiens 1972, 1996). We chose to adopt their

written descriptions of A. campylopodum for our compari with A. siskiyouense in Table 2

The mean diameter of 3-merous flowers for A. siskiyouense and A. campylopodum was similar also (3.2

and 3.0 mm, respectively), but Hawksworth et al. (1992) and Hawksworth and Wiens (1996) did not pro-

vide a range for flower diameters of either species. Our flowet ts indicate the width of perianth

lobes and anther diameters were smaller for A. campylopodum when compared to A. siskiyouense (Table 2),

but other flower characteristics were similar for these taxa.

The mean dimensions of fruits of A. siskiyouense we obtained (5.1 x 3.3 mm) were approximately the

same size as the fruits of A. campylopodum when information for this character reported in the descriptions

of A. campylopodum in Hawksworth and Wiens (1972, 1996) is used (5.0 x 3.0 mm). But fruit dimensions

monticola 745

pu» ] A l J RA E. : al id

A siskiyouense

U Pr, i

L: I£ IIL te Mata E A ] J £, El I +.) Mar "Fara Tal naan TOU Ct I L : I

t fd ~ A m

was available for A campylopdum P|

Character Arceuthobium siskiyouense Arceuthobium campylopodum

Plant ini dil 8.7 (5-17) 8.0 (Maximum - 13)

Basal Diameter? 3.0 (1.8-6.1) 3.0 (1.5-5.0)

Length of mad Internode ? 12.9 (6-25) 11.3 (7-22)

Width of Third Internode * 2.0(1.2-3,0) 2.0 (1.5-2.5

Plant Color Dark brown, brown-green, Olive-green, yellow

red-brown

Mean Flower Diameter ^ B 30

Perianth Lobe Length 1.5 14

Perianth Lobe Width 1.5 1.1

Anther Diameter 0.8 0.5

Anther Distance from Tip 0.8 0.8

Mean Fruit Length 3.1 5.0

Mean Fruit Width 3.3 30

— Male and female plants combined.

— 3-merous flowers.

of A. campylopodum have also been reported as large as 6.0 x 3.5 mm in Hawksworth et al. (1992, Table

1) and 5.6 x 3.0 mm in Hawksworth and Wiens (1996, Table 16.6). Therefore, it was difficult to compare

not only plant dimensions for these dwarf mistletoes, but fruit dimensions as well. Nevertheless, when our

measurements of characters as i siskiyouense are one with the written descriptions, it is evident

these dwarf mistl phologically more similar than reported by Haw] th et al. (1992). A more

detailed l lanalysis i led for A. campylopodum, as well as for other closely related California

taxa such as A. occ dentale Engelm. and A. littorum Hawksw., Wiens & Nickrent. The latter species are also

morphologically similar to A. campylopodum as are their hosts, phenology, and molecular characteristics

(Nickrent & Butler 1990: Hawksworth et al. 1992; Hawksworth & gs pu Mien et al. 2004).

Although Hawksworth et al. (1992) reported that A. siski id noti tion of witches’

brooms on infected Pinus attenuata, our observations el that it frequently does form brooms on this

host (Mathiasen and Daugherty 2009). Host P mid was also used by Hawksworth et al. (1992) to

ly infecting both P. jeffreyi and P. ponderosa

separate A. siskiyouense from A.

Laws., both of which are principal hosts of A. pow However, A. campylopodum commonly infects

P. attenuata, the principal host of A. siskiyouense, and the latter species has now been shown to commonly

parasitize P. jeffreyi (Mathiasen & Daugherty 2009).

Anthesis of Arceuthobium siskiyouense begins in late-July, peaks in mid-August, and continues into

early-September. In every year we observed flowering for this mistletoe, it had completed flowering by late-

September. Seed dispersal of A. siskiyouense begins in late-September and peaks in mid- to late-October.

It continues into early-November, but we were not able to determine when seed dispersal is completed for

this species. Our observations of seed dispersal for Arceuthobium siskiyouense agree with those reported by

Hawksworth et al. (1992), but our observations of flowering indicate its peak flowering period is in August

rather than in SeptemBens.

tl icaldat presented here, the ] | Ic] teristi f Arceutho

Based on

bium siskiyouense siti A. cumpyllopodum (phenology and host alfinities; m the molecular data available

thus far (Nickrent et al. 2004), the classification of the dwarf mistletoe populations parasitizing P. attenu-

ata in the Klamath/Siskiyou Mountains as a subspecies of A. campylopodum may be more representative

746 F [| al 4 Li [| [| H de fTexas 3(2)

of the phylogenetic relationships of these mistletoes. Our data indicate that there are a few morphological

discontinuities between A. siskiyouense and A. campylopodum and Nickrent and Butler (1991) found genetic

differences between these taxa based on their isozyme analysis. However, until we collect additional mor-

phological, host infection, phenology, and distribution data for A. campylopodum throughout its geographic

range, we feel it would be inappropriate to recombine A. siskiyouense as a subspecies of A. campylopodum

here. Furthermore, subspecies of Arceuthobium are usually geographically isolated (Hawksworth and Wiens

1996) and thus far we are uncertain to what degree the geographic range of these two mistletoes does or

does not overlap, even tl listributi maps in Haw] th and Wiens (1996) clearly indicate these

species are sympatric. Although Nickrent et al. (2004) considered A. campylopodum and A. siskiyouense to

be conspecific, this conclusion is based solely on molecular data and does not consider the morphological

differences (plant color), phenology differences, or host susceptibility discontinuities between these popu-

lations. Further research is needed to clarify the host susceptibility of P. ponderosa to A. siskiyouense and P.

attenuata to A. campylopodum discussed by Hawksworth et al. (1992) and Hawksworth and Wiens (1996).

In addition, a detailed morphometric analysis of A. campylopodum is needed and this work should include

analysis of the other mistletoes closely aligned with A. campylopodum in California: A. occidentale and A.

littorum.

Arceuthobium monticola

Our morphological measurements of Arceuthobium monticola are summarized in Table 3. The plant dimen-

sions we obtained for A. monticola were similar to those reported by Hawksworth et al. (1992), except that

we measured plants that were larger (13 cm) than what they reported (10 cm). We also measured larger third

internode and fruit dimensions than what they reported. The color of male and female plants of A. monticola

was consistently dark brown, red-brown, or brown-green. Rarely male plants were brown-yellow to almost

completely yellow, as are those of A. californicum. Our data include the first measurements of flower charac-

teristics. Previous descriptions (Hawksworth et al. 1992) indicated that A. monticola only forms 3-merous

flowers. However, we found that 3-merous flowers were usually on the same plant, but this

character varied. The species produces both 3- and 4-merous flowers, and rarely 5-merous flowers, and we

observed a om male uu unn pouce bas AH 4-merous flowers.

J J 41 eee |

in Hasicewoith and Wiens (1972, 1996) for A. cdi fomteum is aene] in Table 4. Plant dimensions were ap-

proximately the same for both species, but plant color for A. monticola was usually dark brown or brown-green.

However, some male plants were brown-yellow to yellow, as are those of A. californicum. Flowers and fruits

of A. monticola were slightly larger on average than those of A. californicum, but seeds were smaller in length.

Although Hawksworth et al. (1992) and Hawksworth and Wiens (1996) reported that Arceuthobium

californicum rarely parasitizes Pinus monticola, this is evidently based on one report of this mistletoe/host

combination from Castle Lake Campground, west of Shasta City, CA (see Hawksworth and Wiens 1996,

p. 334). Our field observations at Castle Lake, and our comparison of specimens of A. californicum and A.

cyanocarpum (A. Nelson ex Rydb.) Coulter & Nelson, indicate the mistletoe infecting P. monticola at Castle

Lake is A. cyanocarpum. Therefore, we consider there to be no confi 1 reports of A. californicum on P. mon-

ticola at this time. We have observed A. monticola infecting P. lambertiana Dougl. at two locations in northern

California, but this pine is clearly a rare host of A. monticola at both locations. In contrast, P. lambertiana is

the principal host of A. californicum.

Anthesis of A. monticola begins in mid-July, peaks in early- to mid-August, and continues into early-

September. Seed dispersal of A. monticola starts in early September, peaks in late September, and is finished

by late October. While our observations of flowering for A. monticola agree with those reported by Hawk-

sworth et al. (1992), our observations of seed dispersal indicate that peak seed dispersal for this species is

in September, and not October to November. We consistently observed that A. monticola had completed

seed dispersal by late October and did not extend this aspect of its life cycle into November as reported by

Hawksworth et al. (1992).

RA papa ND trn L de A

Taste 3, Morphological ts for A t} ticol Pj

Character A. monticola

Plant Height

Male 7.8 (4.4—-12.6) [50a]

Female 8.5 (6.1—13.4) [a]

Basal Diameter

Male 2.8 (2.0-3.5) [a]

Female 2.9 (2.1-3.8) [a]

Length of Third Internode

Male 9.9 (4.8-17.1) [a]

Female 11.5 (6.9-23.4) [a]

Width of Third Internode

Male 1.7 (1.3-2.4) [a]

Female 1.7 (1.3-2.5) [a]

Staminate Spike Length 8.6 (5.0-14.2)[100b]

Staminate Spike Width

Mean Flower Diameter

erous

14 (1.1-1.8) [b]

2.5 (2.0-3.1) [a]

4-merous 3.6 (3.0—4.6) [a]

Perianth Lobe Length 1.3 (0.8-1.7) [b]

Perianth Lobe Width 1.1 (0.7-1.3) [b]

Anther Diameter

Anther Distance from Tip

Mean Fruit Length

0.5 (0.4—0.8) [b]

0.6 (0.4—0.9) [b]

4.7 (4.0-5.6) [a]

Mean Fruit Width 3.0 (2.4-3.5) [a]

Seed Length 2.5 (1.8-3.2) [a]

Seed Width 1.2 (0.9-1.4) [a]

monticola

Our morphological data for Arceuthobium monticola further demonstrates its similarity to A. californi-

cum, but these two species do have some morphological differences that can be used to distinguish them.

While male and female plant sizes are similar and their ranges are nearly identical, the mean flower diam-

eter of 3-merous flowers of A. monticola is smaller than that of A. californicum. In addition, the fruits of A.

monticola are much larger and they are distinctly glaucous which causes them to appear blue, in contrast to

the lightly-glaucous, green fruits of A. californicum. Plant color also distinguishes A. monticola (dark brown

to brown-green) from A. californicum (yellow to green). Furthermore, A. monticola rarely parasitizes Pinus

lambertiana, the principal host of A. californicum and thus far, A. californicum has not been confirmed to

parasitize P. monticola, although this pine is common in the geographic range of A. californicum in the Sierra

Nevada Mountains (Griffin & Critchfield 1972; Hawksworth and Wiens 1996). Although we have now col-

lected A. californicum from the Klamath Mountains (near Etna and Orleans), its distribution still remains

south of that for A. monticola, so thus S it Bee not we n the GM ranges of A. monticola and A.

californicum overlap. Therefore, the cl of A ubspecies of A. californicum

could be given consideration given the morphological, physiological, and molecular differences between

them and their apparent geographic isolation (Nickrent & Butler 1991; Hawksworth et al. 1992; Nickrent et

al. 2004). However, until we complete a detailed morphometric analysis with data for A. californicum from

throughout its geographic distribution, we do not feel it would be appropriate to recombine A. monticola as

a subspecies of A. californicum here.

Although the dimensions and color of male and female plants of A. siskiyouense and A. monticola were

|| i £ ai n.a a In LI ri 2 >

748 f Texas 3(2)

TABLE 4. A f hol ld for A thobi ticola and A californicum Data is listed (range). Data for A californicum

is from Hawksworth eral 155 (abl 2) and Hawksworth dba (1996, page 197). Singl | ge informati ilable for A

californicum | all otl

Character A. monticola A. californicum

Plant Height ° 82 (4.4-13.4) 8.0 (6.0-14.0)

Basal Diameter ? 2.8 (2.0-3.8) 2.0 (1.5-4.0)

Length of Third Internode * 107 (4.8-23.4) 10.5 (5.0-16.0)

Width of Third Internode ? 1.7 (1.3-2.5) 1.5 (1.0-2.0)

Plant Color ? Dark brown, red-brown, brown-green Yellow to greenish

or rarely yellow-brown or yellow

Mean Flower Diameter ^ 25 3.3

Perianth Lobe Length 1.3 ies)

Perianth Lobe Width 1.1 1.1

Anther Diamete 05 0.5

Anther Distance ann Tip 0.6 0.7

Mean Fruit Length 4.7 4.0

Mean Fruit Width 3.0 2.5

Fruit Color Blue (highly glaucous) Green (slightly glaucous)

Seed Length 2.5 3,2

Seed Width 1,2 12

— Male and female plants combined.

— 3-merous flowers,

also similar based on our findings, they differed in their flower characteristics for both 3- and 4-merous

flowers, and fruit and seed sizes (compare Tables 1 and 3). In addition, Pinus attenuata is considered to be

immune to infection by A. monticola (Hawksworth and Wiens 1996), a conclusion our field observations in

several severely A. monticola-infested mixed stands of Pinus monticola and P. attenuata supports. Our field

observati d those of Hawksworth and Wiens (1996) also indicate that P. monticola is immune to infec-

tion by A. siskiyouense. Furthermore, Nickrent and Butler (1991) found that these species were genetically

different based on their UD rb "i idi LA

A comparison of th lum in Hawl th and Wiens (1972, 1996)

with our popolo edi and vhysiclogical data for A. P and A. monticola could be interpreted as

support for the classification of these species as bun dE with A. campylpodum as suggested by Nickrent

et al. d unt could i Support their as subspecies of A. campylopodum. ne or not

Es Pn

t Spe les, subspecies, Or are conspecitc With

thece A 143 ac

A. campylopodum will continue to be debated by plant systematists, as well as forest managers mä wildlife

biologists whose interests in Arceuthobium spp. have economic and ecological implications (Hawksworth

& Wiens 1996; Nickrent et al. 2004; Mathiasen et al. 2008). Clearly, further research is warranted on the

morphological, physiological, and molecular relationships of the species discussed here as well as all the

taxa included in Section Campylopoda by Hawksworth and Wiens (1996).

ACKNOWLEDGMENTS

The assistance of M. Socorro González Elizondo with an early review of the manuscript and the Spanish

translation for the Spanish resumen is greatly appreciated. Robert Scharpf kindly reviewed the paper as

well.

Math! in harty A thahi icki 1A monticola 749

REFERENCES

GRIFFIN, J.R. AND W.B. CritcHFIELD. 1972. The distribution of forest trees in California. Research Paper PSW-82, USDA

Forest Service, Pacific Southwest Forest and Range Experiment Station, Berkeley, CA

CRITCHFIELD, W.B. 1980. Genetics of lodgepole pine. Research Paper WO-37. USDA Forest Service, Washington,

€

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

Handbook 401, USDA Forest Service, Malón DC.

HAWKSWORTH, F.G. AND D. Wiens. 1996. Dwarf mistlet iology, pathology, and systematics. Agriculture Handbook

709, USDA Forest Service, Washington, DC

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

Novon 2:204-211.

Hickman, J.C. (ed.). 1993. The Jepson manual: pine plants ona AI o Sal roma Press, Berkeley

MATHIASEN, R.L. AND C.M. DAUGHERTY. 2009. Com} y | warf mistletoe.

W. N. Amer. Naturalist 69:42—48.

MATHIASEN, R.L, D.L. Nickrent, D.C. SHaw, AND D.M. Watson. 2008. Mistletoes: pathology, systematics, ecology, and

management. Pl, Dis, 92:988- 10006.

NickRENT, D.L. AND T.L. Butter. 1990. Allozyme relationships of Arceuthobi pylopodum and allies in California.

Biochem. Syst. Ecol. 18:253-265.

NiCKRENT, D.L, AND T.L, BUTLER. 1991. Genetic relations! Arceuthobi ticol | A. siskiyouense (Viscaceae):

New dwarf mistletoe species from California ~ Oregon. Biochem. Syst. Ecol. 19:305-313.

NickRENT, D.L., M.A. García, M.P. Martin, AND R.L. MATHIASEN. 2004. A phylogeny of all species of Arceuthobium

(Viscaceae) using nuclear and chloroplast DNA sequences. Amer. J. Bot. 91:125-138

750 [| I Fal Dat FR” im

Jourt i f Texas 3(2)

BOOK REVIEW

GINA DONOVAN, WITH STEPHEN D. LANGE AND ADRIAN E VAN DELLEN. 2009. Neches River User Guide. (ISBN

978-1603441384, pbk.). Texas A&M University Press, John H. Lindsey Building, 4354 TAMU, College

Station, Texas 77843-4354, U.S.A. (Orders: www.tamu.edu/upress, 1-800-826-8911). $17.95, 83 pp

72 color Posen 40 color maps, 5 b&w photos, 18 tables, 5 1/2" x 8 1/2".

^1 KT D I-11

ti f Texas' publi 1s, both ] future, and privately

owned land. ihe MC River ad Guide isa project c j the T C Alli d ed d he myriad ways thi

river can be UA ed. The gui Lake Palestine Dam to the Downt Dock & 1 Beaumont

itf t, the guide i ttol d spi principally to facilitat f tł tion, thus users can

L l?*l1 i P i *

flip through it ud easily view té maps illustrations, and a GPS li ] B i ps that “reveal the towns,

roads, E pun bridges, Ert lands, pu a Poil PM aoe pies. 360 miles of the river’s course.” One map offers a

] J aa s +

quick overview g I I The larger and

] vi ] 1 [71 1-2 I. x 1 "n y y 15 E 11 J r, Vies "DC s +

i r L r O

I r

‘for points of interest" along the river.

aes B is a ‘Duet won tion ` e SIE and x of human id iss nia river, as e as pheeaes of

pl Se ae eta nT

feeling of st ]ship for the ri tl y be enjoyed Ley al for da to come.

Printed on waterproof paper, mM Neches River User G ; 1st dy Pg ide f j ] t

peri he Neches Ri first-] el paddlers, birders, Ps hunters, and ] g l Recommended

for all public lil 1 collecti that ] , ecology, and lei tiviti Texas

à tive director of the Houston Audubon Societ; Co- founder of the East Texas Forest & Wildlife Hips

she also served on the Texas Parks ane Wildlife Der 's Ri C tion Advisory Board. Stephen D. Lange is a Regional G

ements for the Texas a an Der 's Wildlife Divisi Tyler, Texas. Aden F. Van pum PE is a retired ui

d ] pl g pl Gary Jennings, Librarian, B f Texa

Fort Worth, Texas 76102-4025, U.S.A.

J. Bot. Res. Inst. Texas 3(2): 750. 2009

ADDITIONAL OBSERVATIONS ABOUT PHASEOLUS ROTUNDATUS (FABACEAE),

AN ENDEMIC BEAN SPECIES FROM WESTERN MEXICO

Jesus Salcedo C. Rogelio Lépiz |. Nora Castaneda A.

Bioversity International University of Guadalajara Bioversity International

Office for the Americas Centro Universitario de Ciencias Office for the Americas

c/o CIAT, COLOMBIA Biológicas y Agropecuarias c/o CIAT, COLOMBIA

jm.salcedogcgiar.org Jalisco, MÉXICO

rlepiz@cucba.udg.mx

César Ocampo N. and Renal G. Debouck

International Center for Tropical iculture (CIAT)

o —_

AA 6713

Cali, COLOMBIA

d.debouck@cgiar.org

ABSTRACT

H +l E r : D ] 1 + DI 1 " E

es bla for western Mexico, where it appears to be endemic and

endangered, with very few popul je 1 to date, and al I ee ee BENE B disce

AD 1 es rl +l FD I s D H 124 L ^ +1 1 +4 ^e]

the validity of the former taxon.

RESUMEN

d (roe ee Ere E 1 1:1 TD 1 " Ta, - £ - . "n na EN

Se presentan ltados q d : p d y en peligro d ]

dente de México, con muy p pobl das a la fecha y ubicad y f de 4 tegidas. S gnan |

At : y = 1 D 1 1 r 1 Jj 1 D 3 E J] DL T A 1 1 xz

i i po e o r rE

confundirse, para confirmar la validez de P. rotundatus.

INLRODUCTLION

Anew wild bean species, P} lus rotundatus F Del , was described for western Mexico (Freytag

& Debouck 2002). It is a tall woody abne vine wib rounded rhombic, densely pubescent, trifoliolate

bs and stout panicles with lilac-purple flowers. The double racemic structure of its inflorescences with

] t of the lary raceme axis (Debouck 1983) clearly places it in the section Paniculati Freytag

of the genus. The species was first collected b Rogelio Lépiz in October 1978, and shown later in that year

as a herbarium voucher to tl thor (DGD), who could not identify it, but confirmed it as a ‘Phaseolus

sensu stricto' as the genus was understood by then (Maréchal et al. 1978). Although the voucher was slightly

overburnt during drying, Robert Maréchal was able to obtain a living specimen in the glass-house in Gem-

bloux, Belgium (as NI 1046, also G40693 at CIAT). George F. Freytag obtained later seed of that plant, and

on the indication by the senior author (DGD) aptly described P. rotundatus as grown in a growth chamber

at the Tropical Agriculture Research Station of USDA, Mayagüez, Puerto Rico (Freytag & Debouck 2002).

Conditions were not ideal for the preparation of type specimens (currently the holotype is US3386751, and

isotypes are kept at CS, F, and MICH, none with complete reproductive parts). Additional material was col-

lected in the field by R. Lépiz and D. Debouck in recent years. These latest collections enabled additional

observations on blastogeny and palynology, which are reported below. Because of a possible confusion with

a closely related species, viz Phaseolus marechalii Delgado (Delgado 2000), attention is drawn to differences

between these two taxa.

|| Rat Rac Inct Tayac 3(2): 741 — 762. 2009

752 j tl tanical hl exas 3(2)

RESULTS

Morphological observations

The following complements the description given elsewhere (Freytag & Debouck 2002) on plants grown

for several years in CIAT Palmira, Colombia, and seen in Rio Verde, Jalisco, Mexico in November 2003.

The small amount of variation observed for many morphological traits between plants grown artificially in

Mayagtiez (18°12'28" lat N, 67°08'18" long W !) and in the field in Palmira G°30'06" lat N, 76°21'21"long

W) is noteworthy.

Seedling 15-18 cm high, from hypogeal germination; epicotyl green, terete, 80-100 mm long, covered

with white uncinate trichomes (Fig. 1a, 2e left). Eophylls simple, deltoid, green (146B Yellow — green group;

Anonymous 1966), opposite, with minute white uncinate trichomes on adaxial and abaxial surfaces, 30 x

25 mm, apex acute to acuminate, base slightly cordate; petioles green, 15 mm long, minute stipels, pulvini

dark green, pubescent; stipules triangular, basifixed, 1-2 mm long, highly pubescent, 3- nerved, green.

First true leaf trifoliolate, leaflets ovate to elliptic. Root diameter 8-12 cm or more, 50 cm long or more,

pluriannual thick fleshy and woody, with 3-6 conical subdivisions 15-30 cm long of 1-4 cm diameter,

with prominent whitish nodules and dark brown corky cortex. Stems terete stout erect in lower parts

and climbing in upper parts, woody and enlarged at base, total length 2-4-8 m (Fig. 2b), diameter 2-5-8

mm, internodes 12-14 cm green (144A) densely covered with white short strigose pubescence. Stipules

triangular acute 4 mm long 2 mm wide covered by whitish pubescence. Leaves trifoliolate 13-18 cm long,

terminal leaflets rhombic rounded to orbicular (57 x 55 mm to 67 x 62 mm) greenish (147A) because of

whitish dense pubescence, lateral leaflets similar but slightly inequilateral (53 x 45 mm to 65 x 64 mm);

petioles 3—6—10 cm long, petiolules 9-20-30 mm long; lower pulvini 4-6 mm long, upper pulvini 2-3

mm long; stipels ligulate 2-3 mm long. Inflorescence a panicle with 20-30 or more secondary racemes,

some of them developed at base, 35-75 cm long (Fig. 2a); the peduncle 12-22 cm long stout slightly curved

and reddish at base densely covered with white short strigose pubescence and minute uncinate hairs; the

rachis 18-51 cm long usually erect green with 15-20-30 secondary racemes irregularly spaced, often with

development of axis beyond second flower, subtended by triangular aciculate primary bracts 2.53 x 1.2

mm strongly 3-nerved covered by whitish pubescence. Pedicels terete straight or slightly recurved 4-6 mm

long covered with white short strigose pubescence and minute uncinate hairs. Pedicelar bracts linear 1 mm

long 1-nerved densely covered by whitish pubescence. Bracteoles green triangular narrow hirsute Imm

long or less strongly 1-nerved densely covered by whitish pul nce (Fig. le). Flowers papilionaceous (Fig.

lc, 1d, 2d left) purple pink (72B red-purple) fading violet blue (85A violet), two for each secondary raceme

(Fig. 1b); calyx round cupped green turning to tan at insertion of pedicel, hirsute, margin shortly ciliate,

tube 3.5 mm long, two upper calyx lobes fused unconspicuous, three lower lobes triangular apiculate 1 x 1

mm long (Fig. le); standard outer face pale lilac glabrous with a few straight short hairs on top, inner face

purple pink intense with auricles 4-5 veined, squarish with a deep central sinus, 11 x 8.5 mm, thickened at

flexure, auricles rounded slightly reflexed, the right auricl e developed, claw green reddish 1.5 mm long

with two parallel rounded green pinkish callosities (Fig. 1f); wings spreading horizontally, intense purple

pink, obovate rounded subequal, the margins often revolute particularly on the left wing, the blade 8 x 6

mm glabrous, the squarish spur slightly adhering to the keel, claw linear 4 mm long (Fig. 1g); keel tubular

in close asymmetrical spiral spreading to the left white greenish at base, pink purple in the ascending part,

greenish at top, 1 34 counterclockwise closed coils, ca 12 mm long, two claws divided 2 mm long whitish

with a few pink spots, a convex rounded pocket 1 mm long above each claw (Fig. 1h); stamens diadelphous

(9+1): vexillary stamen free 11 mm long claw terete 2 mm long to cupped knob 1 mm diameter (Fig. lj);

Medus Hue "s mm ue py veined with two small rounded callosities flanking the knob (Fig. 1i);

al dorsifixed. Ovary green glabrous 5 x 1 mm laterally compressed margins slightly

sranounced inserted on a smooth basal disk 1 mm diameter, 4 ovules; style spiraled ca 11 mm long terete

glabrous, a brush of straight 1 mm long hairs unilaterally extending below the stigma not beyond, stigma

narrowly triangular white 1 mm long not extending beyond style tip (Fig. 1k). Pods falcate small 28-45

Salcedo C. et al : Ol

Em uw

un Wy

> H *

Fic 1 FE a FI F mL

k a Seedling b. Habit of

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(eh): e. Pedicel, bracteoles and calyx. f. Standard. g. Wings. h. Keel. i. Staminal tube. j. Vexillary stamen. k. Ovary and style. I. Pod. m. Seed.. n. Root

system and lower part of oneyear plant. Scale: a

~b, |, n in cm; d-m in m

iew). Floral parts

Li T

i Tepatitlán, Jalisco, Mexico

,C olombia, of seed from

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Salcedo C. et al., 0l ti hout PI | tundat 755

mm long, 8-10 mm wide, 2-4 seeded, flattened, slightly constricted between the seeds; valves glaucous

green (144B) drying tan (164B), twisting 1 Y at dehiscence, the sutures finely thickened, the beak curved

downwards 1-2 mm long. Seed round ovale oblong 6-8 mm long 5 mm wide 2-3 mm thick (Fig. 3a), 100-

seed weight 7—9 g, cream tan background covered by a dense fine black speckle with a black ring around

the hilum; hilum irregularly elliptic 1.2 mm long 0.6 mm wide; lens small and slightly raised. Pollen (Fig.

4a and 4d) round (polar view), subprolate, tricolporate with 6 pseudocolpi, medium-sized, diameter 25-27

um, almost spherical; margins surrounding the colpi prominent and smooth 3 um wide; endoaperture el-

liptic (longer axis 7 pm); endoaperture membrane smooth, ectoaperture membrane finely punctate; tectum

cdd punctuate.

S 1: For P. rotundatus: ¡State ee bu dd i jaan de pcd) er al 21°00'07"N. Long. 102?48'

10"WI. 1,200 m, 22 Oct 1978, R Lépiz s.n. ( Palmira station, and hereby

distributed to BRIT, COL, em F, PHO; de GH, K, IBI IG. IEB, MEXU, MICH. NA. NY, SI, TEX, UC; in addition, small amounts of seeds,

accessioned as G40693,

For P. aman Merco: Villa del Carbón, 12 n SE de Villa del Carbón, 19°41'N, 99°25'W, 2520 m, 6 Nov 1987, DG Debouck &

JS M , Cuernavaca, [1901'N. 99?15"W], 2000 m, 7 Nov 1969,

Hinton 17462 (K; callestéil by James Binion 12); Tepoztlán 18°59'N, 99°06'W, 1900 m, Leroi Mex-130 (COL, GH, K, IBUG); in pedregal

barranca just below 62 Km marker, which is about 14 ! th of Cuernavaca, [19°04'N. 99°15'W], 27 Oct 1947, OW Norvell HM 215

(UC933928; los E E glaucocarpus newer Pe nov. ined); "No d'introduction 402. Faculté des Sciences figs de l'Etat

Gembloux, Chai les. Herbier no. 2386. Phaseolus sp. P M' Schubert,

Morelos 623, Mexique. [19°02'N. 9e"17WIE Rec: ed 1971 serre Gembloux” (K). PUEBLA: Acajete, Cerro 4 im NE de Pueblo Acajete,

19%08'N, 97°54'W, 2400 m, 11 Nov 1987, D j g 2389 (BR; MICH1179986; US3168489).

On the basis of the revision of herbarium specimens mentioned above and ag EON of peus En

out side-by-side in the experimental station of CIAT in Palmira,

P. rotundatus and P. marechalii in Table 1.

Ecology

From the two Pala Sua so far (at Río Verde, with a difference in altitude of 200 m), P. rotundatus

seems dist; ] d the upper reaches of the valley of Río Grande de Santiago, while

P. marechalii thrives in ue oak forests of the Trans-Mexican Volcanic Belt (Fig. 6).

Electrophoretic evidence and species relationships

A quicl f seed st in one-dimension SDS-PAGE electrophoresis, using 1596 polyacrylam-

ide slab sels: as in Brown et al (1981) modified by Gepts et al. (1986) (Fig. 5), indicates very similar patterns

of banding for P. rotundatus (lane 3), P. marechalii (lane 4) and P. jaliscanus Piper (lane 5). P. novoleonensis

Debouck (lane 6) of the Coriacei section (Salcedo et al. 2006) is then close, although with much less in

globulins around 50-52 kD. P. lunatus L. (wild Mesoamerican oe lane Da and : plabeltus d (lane "

show almost no globulins in the range of 50-52 kD, but ] igni

35 kD. The section Phaseoli is represented by P. costaricensis Freytag & Debouck (lane 9) EE P. vulgaris L.

(wild Mescamerican - lane 10) with a significant amount of phaseolin (50—52 kD), a possible arcelin in wild

P. vulgaris (35 kD) and a possible lectin-like protein in P. costaricensis (24—25 kD). P. chiapasanus Piper (lane

8) would have a significant amount of globulin too (50 kD).

DISCUSSION

] P T 125 pi | A 1

The data above were collated to show the differences between P.

to the ones reported elsewhere (Freytag & Debouck 2002). They elicit the following points for discussion.

First, P. rotundatus is a taller, aggressively climbing vine as compared to P. marechalii; plants topping trees

5-6 m high (Fig. 2b) are frequent, thus with stems up to 8-10 m long. Racemes and flowers are larger as

compared to those of P. marechalii (Fig. 2a, c, and d). Other differences exist in seedlings (Fig. 2e), seeds

(Fig. 3a and 3b), and pollen (Fig. 4a,d and 4b,c). Terminal leaflets of fully developed leaves are 67 x 62 mm

in P. rotundatus, while 50 x 32 mm in P. marechalii.

[| I £ al Dos H In LI POP "hi

756 f Texas 3(2)

En 2 Fl r J log [| Li y...” rJ Le fi I . h DER dd n " | A rg m PARAFRASI L. Pi i Pa ta h P marerhalir

dues o diei id curd i * LÀ r r E f FI

£n H £P. ri L 775. FAAPFASL. hI Amm LE Je £ Lites Morelos Mexica r i K L FAg D "EI ran ros £, BL

A f f i a Li F P P e i F ; 3

of Gainesville, Alachua County. Florida. USA. d. P lunatus (DGD-527: 625741) from CI Hecelchakan, Campeche. Mexico. TI le bar i

f af F V e yews J ¥ r f J,

for each division ] | lightly f hooting t ti

r ,£ - , a

Salcedo ( at al ' Ri "11 L z ni I 4 Jag 757

0089 20KU 2,560 10ym WI 0003 28KU az ,388 10Yn

0808 20KY — 542,700 | 0902 28KU — 42,588

Fic. 4. Pollen types of P. rotundatus (left, 4a and 4d) and P. marechalii (right, 4b and 4c). Polar (4a, 4b) and equatorial (4c, 4d) views. Scale bar: 10

microns

Tape 1 C y Lal , rm L d. n j ) n L T

Trait P. rotundatus P. marechalii

Seedling Larger eophylls (30 x 25 mm); leaflets of Smaller eophylls (22 x 15 mm); leaflets of

1st true leaf bluntly acuminate 1st true leaf acuminate

Plant Stout and puri citing vine, usually Stout climbing vine, usually less than 3 m high

more than 4 m

Leaves Large 13-18 cm (d leaflets rhombic rounded Small 7-11 cm long, leaflets ovate acuminate

Racemes Large panicles, more than 20 cm long Small panicles, less than 20 cm long

Inner standard Veining ie Clearly vertically veined

ee vale flattened ound ovoid thicker

Habitat Oak md of N-NE Jalisco, Mexico Pine-oak forests of Trans-Mexican Volcanic Belt,

central Mexico

Journal of the Botanical Research Institute of Texas 3(2)

Fic. 5. One dimensional SDS-PAGE gel of pal rotundatus and selected taxa. Lane 1: molecular weights in kiloDaltons (range 14.3 lower band to

66 upper band). Lane 2: P. Junatus wild oe om Jalisco, Mexico; 625704). Lane 3: P rotundatus (Lépiz s.n. from Jalisco, Mexico; G40728). Lane 4:

P marechalii (B Schubert 623; G40506). Lane 5: : pela (R Buhrow M21 from Jalisco, ipn PI 535313). Lane 6: P novoleonensis andi from

Nuevo León, Mexico; G40590). Lane 7: "pru, 5 (DGD-2043 from San Luís Potosí, Mexico; G40585). Lane 8: P. chiapasanus () Muruaga Mart

from Oaxaca, Mexico; 640676). Lane 9: P. costaricensis (DGD-2119 from Cartago, Costa Rica; 640604). Lane 10: P vulgaris wild (HSG-22202 an en

Mexico; 612866).

Salcedo C. et al., Ol ti bout P! | tundat 759

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98 GO'NW

89 ('ONV

100 SOW

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pero

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in =

BACONA or

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Lts al! In L ger i 15.1 Ets al JA t J

Fic. 6. Distribut

central Mexico. 1 = Wildlife refuge“La Primavera"; 2 = Wildlife refuge"Sierra de Quila": 3- Biosphere reserve “Sierra de Manantlán”; 4- Wildlife refuge

“El Jabali”; 5 = National park"Nevado de Colima"; 6 = Biosphere reserve “Mariposa Monarca”; 7 = pariona! park Nevado de Toluca’; 8= Nation al park

“Desierto del Carmen”; 9, 10, M SIME euge Cenagas de Lema 12 = National park"Insurg Hid illa”; 13 = National park

“Desierto de Los Leones”; 14 = National "C National park “I le Z la^; 16 = -Wildlife S DIDA.

17 = National d ium 18 = National park“Iztaccihuatl- Popocatepeti" 19- National park“La Malinche”.

760

Second, P. rotundatus seems to be known from only one locality to date, distributed on the upper slopes

of the high valley of Rio Grande de Santiago and possibly surrounding mountains or volcanoes. Study of

over fifty herbaria holding specimens of the section Paniculati (Debouck 2009) has not revealed additional

populations. Delgado (2000) reported a distribution around the Mexican state of Morelos for P. marechalii;

from the novel records it is still a rare species as he indicated, restricted to the Trans-Mexican Volcanic

Belt. Both species were not reported in “Flora Novo-Galiciana” (McVaugh 1987), perhaps another indica-

tion of their scarcity in and around Jalisco. A hypothesis worth further testing is that P. rotundatus had a

wider distribution in the past than at present, namely in the valley of Guadalajara, before encroachment by

agriculture and urbanization. The putative reduction in its distribution could be attributed to its plant type:

it is a perennial vine reaching considerable development, and unlikely to survive under intense grazing

pressure, plowing and fire, all characteristic of post-Columbian land use practices in central Jalisco. Today

it only survives where humans, cattle and goats have no access, although places with little deep soil like

cliffs are not its primary habitat.

Currently, as it can be judged from few sources (Toledo & Ordoñez 1993; ee et » 1996; Toledo et

al. 1997), few populations of P. marechalii and none of P. rotund luded in protected areas However,

it would still be timely to include populations of these two species into ten schemes, by slightly

expanding current protected areas (for example, to the east the National Park ‘La Malinche”, or to the north

west the National Park “El Tepozteco”), or creating an area S of Río Verde bridge for P. rotundatus (where the

slopes of road sides of federal road Mex 25 are a wasteland anyway). An additional justification for further

in Det conservation efforts stems from the fact that the upper Santiago watershed might be involved in the

an (Chacón et al. 2005; Kwak et al. 2009), because of the presence of wild forms

(our germplasm collections of 1978; Zizumbo et al. 2009).

On the other hand, the unicate specimen collected by Howard Scott Gentry (HS Gentry 22509) in

Telixtlahuaca, Oaxaca, Mexico, and kept at the US National Arboretum (NA0027957) has been identified

by GF Freytag on July 26, 1989, as Phaseolus MURS PEE & diua (ined.). We have seen this

specimen on July 18, 2006, and can discard this i has two short racemes

(12 and 9 cm) with 5—6 primary bracts and the leaflets are almost glabrous This means that there would be

no disjunct distribution for P. rotundatus between Jalisco and Oaxaca, of limited distribution in the upper

valley of Río Grande de Santiago in Jalisco, and that HS Gentry 22509 belongs to another taxon.

Third, on the basis of electrophoretic evidence of seed storage proteins, P. rotundatus seems to share

phylogenetic affinities with species such as P. marechalii, P. jaliscanus, perhaps even with P. novoleonensis.

Since Lima bean (lane 2) is the type species of the section Paniculati (Freytag & Debouck 2002), one would

expect more similarities with other members of Paniculati (viz. P. rotundatus, P. marechalii, P. jaliscanus).

This result confirms those of a study on an intergenic section of cpDNA using PCR-RFLP (Fofana et al.

1999), where P. rotundatus (= NI 1046) is shown to be close to P. ritensis Jones and P. reticulatus Freytag &

Debouck (= NI 702), both of which are related to P. marechalii. Preliminary results of ITS sequencing (Gaitan

et al. 2000) indicated a close proximity between P. rotundatus and P. jaliscanus and P. acinaciformis Freytag

& Debouck. In a combined analysis of the nuclear ribosomal DNA ITS region and of the chloroplast DNA

intron trnK, Delgado et al. (2006) have shown the close phylogenetic proximity between the Paniculati and

the Coriacei.

As recognized by Freytag and Debouck (2002) and Delgado et al. (2006), the Paniculati seems to be the

section of the genus with the largest number of species, and thus is of special interest to breeders of Lima

bean given the pe for wide crossing o 1990). The base for further progress lies first in the

proper botanical identification of this biologi set and its conservation. From the review of the Paniculati

carried out so far, clearly not all taxonomical probl solved, but we hope to have herein advanced one

step further for one of them, P. rotundatus.

Salcedo C. et al., Ob ti bout PI | tundat 761

ACKNOWLEDGMENTS

This work initiated in the fall of 1978 has been made possible thanks to generous grants from CIAT, INIA

of Mexico, the University of Guadalajara (the latter for continuing field work in western Mexico), BMZ of

Germany, and the World Bank for the visit of one ofus to the U.S. National Arboretum. The continuing help

of Orlando Toro for keeping track of all data and specimens over years has been immensely valuable to our

research. The help of José Alejandro Arroyave and Camilo Oliveros for the scanning electron micrographs

and the setting of the figures, respectively, is particularly appreciated. The authors desire to warmly thank

Dr George Freytag for his continuing support and interest on all these jeu pl ds observations

on the poorly known bean species. The y appreciated. One

of us (DGD) would like to express his deepest gratitude n the Curators of the (alles nena (that keep

specimens of section Paniculati; see Debouck 2009): A, AGUAT, AHUC, BAA, BAFC, BM, BR, BRIT, CAS,

CHAPA, CICY, COL, CORD, CPUN, CR, CUZ, DAV, DES, DS, EBUM, ENCB, F, Fl, G, GH, HAO, HNMN,

HUT, IBUG, IEB, INB, K, L, LL, LIL, LPB, M, MA, MEXU, MICH, MO, MOL, MSC, NA, NEBC, NY, O, OXF,

P, PH, PRG, QCA, SI, TEX, UC, US, USJ, USM, WIS, who allowed him to study their collections ‘in sitw’.

REPERENCES

ANONYMOUS. 1966. Royal Horticultural Society Colour Chart. The Royal Horticultural Society, London. Fans 2, 3,

and 4.

BAUDONN, J.P. 1990. l'amélioration de Phaseolus lunatus L. en zones tropicales. 6. Réservoirs génétiques, travaux

de sélection, conclusions et perspectives. Bull. Rech. Agron. Gembloux 25:395-442

BROWN, J.W.S., Y. Ma, E.A. Buss, AND T.C. HALL. 1981. Genetic variation in the subunits of globulin-1 storage protein

of French bean. Theor. Appl. Genet. 59:83-88.

CASTILLEJA, G., D. EVANS, AND S. EGGEN-MicInTOSH. 1996. Mexico. In: C.S. Harcourt and J.A. Sayer, eds. The conservation

atlas of tropical forests. The Americas. International Union for Conservation of Nature and Natural Resources,

Simon & Schuster, New York, New York, USA. Pp. Iun la

CHACÓN, M.I., B. PICKERSGILL, AND D.G. Desouck. 2005. D | patterns in common bean (Phasec Igaris L.)

and the origin of the Mesoamerica and Andean cultivated races. Theor. Appl. Genet. 1 10432444.

Desouck, D.G. 1983. Observations sur les processus de floraison et de ramification chez quelques Phaseolus.

Comp. Rend. Séances Acad. Sci., Sér. 3, Sci ue 296:425-428.

DeBouck, D.G. 2009. Cahiers d ion Paniculati. International Center for Tropical Agriculture (CIAT),

Cali, Colombia. Accessed on n line on 7 ner 2009 at http://www.ciat.cgiar.org/urg. 261 p.

DELGADO SALINAS, A.O. 2000. New Species of Mexican Phaseolus (Fabaceae). Syst. Bot. 25:414-436.

DELGADO SALINAS, A., R. BIBLER, AND M. Lavin. 2006. Phylogeny of the genus Phaseolus (Leguminosae): a recent diver-

sification in an ancient landscape. Syst. Bot. 31:779-791.

Forana B., JP BAUDOIN, X Vekemans, DG Degouck, AND P. DU Jaro. 1999, Molecular evidence for an Andean origin and

a secondary gene poo! for the Lima bean (Phaseolus lunatus L.) using chloroplast DNA. Theor. Appl. Genet.

98:202-212.

FREYTAG G.F. AND D.G. Desouck. 2002. Taxonomy, distribution, and ecology of the genus Phaseolus (Leguminosae-

Papilionoideae) in North America, Mexico and Central America. Sida Bot. Misc. 23:1-300.

GAITÁN, E., O. Toro, J. TOHME, AND D.G. Desouck. 2000. Molecular taxonomy of the genus Phaseolus through ITS se-

quencing. Annual report, Centro Internacional de Agricultura Tropical, Cali, Colombia. Pp. 56-60

Gerts, P, T.C. OSBORN, K. RASHKA, AND F.A. Buss. 1986. Phaseolin protein variability in wild forms and landraces

of the common bean (Phaseolus vulgaris L.): evidence for multiple centers of domestication. Econ. Bot.

40:451-468

Kwak, M. J.A. Kami, AND P. Gerts. 2009. The putative Mesoamerican domestication center of Phaseolus vulgaris is

located in the Lerma-Santiago basin of Mexico. Crop Sci. id 554—563.

d'i Im mirmirimae raranlava Alacnarcac rare marnrac

MARÉCHAL, R., J. MASCHERPA, AND F. STAINIER. 1978. Etude t | un groupe complexe d'espèces des genres

762 j tl tanical Insti Texas 3(2)

Phaseolus et Vigna (Papili )surla! le d : phologiq t polliniques, traitées par l'analyse

informatique. Boissiera 28:1-273.

McVauGH, R. 1987. Flora Novo-Galiciana—A descripti tofthe vascular plants of western Mexico— Volume

5 Leguminosae. The University of Michigan Press, Ann Arbor, Michigan, U.S.A.

SALCEDO, J., J.A. ARROYAVE, O. Tono, AND D.G. Desouck. 2006. Phaseolus novoleonensis, a new species (Leguminosae,

Phaseolinae) from the Sierra Madre Oriental, Nuevo León, Mexico. Novon 16:105-11 1.

ToLepo, V.M. AND M.J. OgpoREz. 1993. The biodiversity scenario of Mexico: a review of terrestrial habitats. In: T.P.

Ramamoorthy, R. Bye, A. Lot, and J. Fa, eds. Biological diversity of Mexico: origins and distribution. Oxford

University Press, New York, New York, U.S.A. Pp. 757—777.

ToLepo, V.M., J. Rzepowsk1, AND J. VitLA-LoBos. 1997. Regional overview: Middle America, Mexico. In: S.D. Davis, V.H.

Heywood, O. Herrera-Macbryde, J. Villa-Lobos, and A.C. Hamilton, eds. Centres of plant diversity—A guide

and strategy for their conservation, vol. 3 The Americas. World Wide Fund for Nature and International Union

for Conservation of Nature and Natural Resources, Cambridge, United Kingdom. Pp. 97-107.

Zizumso, D., R. Papa, M. HUFFORD, S. REPiNski, AND P. Gers. 2009. Identification of new wild populations of Phaseolus

vulgaris in western Jalisco, Mexico, near the Mesoamerican domestication center of common bean. Ann.

Rept. Bean Improv. Coop. (U.S.A.) 52:24-25.

DISCOVERY OF A LEWIS AND CLARK GRASS COLLECTION,

POA SECUNDA (POACEAE) SENSU LATO, AT THE HERBARIUM

OF THE ROYAL BOTANIC GARDENS, KEW

feque EON James L. Reveal

National P. Biodiversity L.H. Bailey Hortorium Herbarium

Arce and Agri-Food Canada Department of Plant Biology

unders Building (#49) Cornell University

a on Farm Ithaca, New York 14853-4301, U.S.A.

Ottawa, Ontario, CANADA K1A 0C6 jir326@cornell.edu

jacques.cayouetteGagr.gc.ca

Gary E. Moulton

University of Nebraska

Lincoln, Nebraska 68588, U.S.A.

gmoulton@unl.edu

ABSTRACT

zr AAG 1: I 7 As y) a T TF Prea unda J D 15 1 n 1D Ce an J LL ‘yy 11 4 ] J L +1

J i Bue [a]

jr. i " 3-1 1 i 11 1 T E 171 1- LT 1 : +1 A

ana _TaNnAT ua ir] T

l Sci in Philadelphia (PH-LC) i ixt f a June 1806 collection: made i in Idaho and a late July 1806 collection par

for the LE of A brevifo lia to the

in Montana. Accordingly, a second step" lectotypification is propos

element Pursh had at hand in London, namely plants collected die Lewis un near the mouth E: Crow Coulee, Chouteau County,

Montana, on 28 Jul 1806.

RESUMEN

Ela Brimi ANZ A Es dJa Aq 1 ‘Cys y ly. (Dee bei A TOD 15 1D 1n r 1 17 ‘wy 1 3

J M T d Pi MA COME LANLA

inn la e MM de Lewisy Clarke: en Edd 1806 d tró que el lectotipo designad l Herbario Lewi 1 Clark en la Acad

1 3 | 1 aed -l G L q 1enx 1 1 THal ] pa | ps 1 A

julio de 1806 de Mont D d to, se eon una lectotipificación “segundo pue pos el pliego PH-LC n enaa la

aplicación de A. brevifoli lel t P t Led

cerca de la boca de Gee cales Bond de Chouteau, Montana, el 28 de M de 1806.

INTRODUCTION

In September 2005 while visiting the herbarium of the Royal Botanic Gardens at Kew (K), the first author

discovered a collection labelled Aira brevifolia Pursh (Fig. 1) in a Deschampsia cespitosa folder (Merritt et al.

2006). It bears labels and annotations congruent with previously detected Lewis and Clark specimens at K

by Ewan (1979) and Moulton (1999), but had not been seen by Ewan, Moulton, or botanists investigating

the Lewis and Clark collections examined by Frederick Traugott Pursh (1774-1820) in the preparation of

his flora (Pursh 1813). Only about ten or eleven Lewis and Clark specimens were known to be at K (Ewan

1979; Moulton 1999; Reveal et al. 1999; McCourt & Spamer 2004a), these having been obtained indirectly

at auction by William J. Hooker (1785-1865) in 1842 (Miller 1970).

RESULTS AND DISCUSSION

Features of the Kew and PH Specimens, and Historical Background

The Kew Aira brevifolia collection has four features typical of Lewis and Clark specimens found at K: 1) a

label in Pursh’s hand with the locality and the date: “Doubtfull River Missouri plains Jul. 28. 1806” (Fig.

1A), thus indicating a Meriwether Lewis collection; 2) annotations by Aylmer Bourke Lambert (1761-1842),

J. Bot. Res. Inst. Texas 3(2): 763 — 769. 2009

£T ai”

284.

= fhe tecismtn ds C Aii bt sof

= : 5]

boat fría ce Puro? AT dinie

liti The plant, P,

Cayouette et al., Lewi i Clari g llection, Poa secunda 765

protector and patron of Pursh, one on the front of the sheet: “Aira brevifolia Pursh Fl: Amer.” and one on the

back: *[N.] America Fred. Pursh" (Fig. 1B); e an annotation by Kup Pao ai (1806-1899), a bookseller

and editor: “Herb. Pursh proper”; and 4), an Hb Hool An tation by George Vasey (Fig.

1) 1822-1893), an American grass family specialist who eE the specimen as “Deschampsia caespitosa”

(sic), is one of the reasons why the specimen was filed in Deschampsia and not detected previously (Merritt

et al. 2006). Aira and Deschampsia were once considered to be related genera and some of the species were

placed in Aira by some, and in Deschampsia by others (see for example, Hitchcock 1920).

Nevertheless, the sad is correctly identified as ME Poa secunda J. is ou (annotated by J.

Cayouette and SJ. Darbyshire in 2005). In fact, as the specimen has most of it ttered, it was not

recognized easily as a Poa species. qe shattered state of the spikelets of the P. — specimen is congru-

ent with its having been collected late in the season, on July 28, 1806. On that day, a Monday, Lewis and

his men were in western Montana near today's Fort Benton. In the morning they were in Chouteau County,

close to the Missouri River, after retreating from the Blackfeet Indians for three days (they killed two in the

only “battle” of the expedition; see Moulton 1993). They were fatigued but determined to reach the river.

When they heard rifle fire, they assumed it was Sergeant Patrick Gass and his party bringing the canoes

down river. They reached the Missouri River, still in Chouteau County, and the two groups joined up that

afternoon. From near the mouth of the Marias River they journeyed downstream about fifteen miles and

camped.

It is difficult to determine exactly where Lewis collected the plant. Also uncertain is the “Doubtful

River" reported on the label (Fig. 1A), probably transcribed by Pursh from an annotation by Lewis that is

now lost. This location is not mentioned in the journals nor is it shown on Clark's maps, but might refer to

the area near the mouth of Crow Coulee (Chouteau County, see Merritt et al. 2006) which would have been

a “Doubtful River.”

The main Lewis and Clark plant collections are at the herbarium of the Academy of Natural Sciences

of Philadelphia (PH-LC) and have been fully documented (Moulton 1999; Reveal et al. 1999; McCourt &

Spamer 2004a, b; Spamer & Meet 2000 When Pursh went to London in 1811, he brought some of

Lewis's plants, most probably tl ht were new to science (Reveal et al. 1999; McCourt & Spamer

2004b) to prepare his Flora Americae CHEM that was published in late December 1813. The num-

bers of Lewis and Clark specimens he brought with him are not known but probably there were about 50,

representing approximately 2596 of the entire Lewis and Clark collection (McCourt & Spamer 2003).

After the publication of his flora, Pursh returned to North America (Canada) and left his specimens

with A.B. Lambert who kept them in his poser) M, in London. After Lambert's death in 1842, his

herbarium was auctioned off, with Pursh' laced in at least two lots (Miller 1970; Moulton 1999).

The one acquired 2 the botanist Kawana Tuckerman (1817-1886) contained may of Lewis’s specimens,

and they e ually found their | Philadelphi with the other coll lat the American

philosophical ps that were recovered in 1897. An uian number of Lewis’s plants were mixed with

the Pursh herbarium in a lot that was acquired by Pamphlin. The Pursh plants came later to WJ. Hooker

and became the property of Kew in 1867 when Hooker’s herbarium was purchased by Kew. Joseph Ewan

discovered the Lewis specimens at Kew (Ewan 1979).

Most of the ten or eleven collections E seis to Lewis are ss ceca öl al

retained in Philadelphia and now part of PH-LC. The Philadelphia bed

by Pursh in his flora. Most of the Kew duplicates bear a Pursh label and share the same bieen locality. but

this is not the case with the PH-LC counterpart of Aira brevifolia (Fig. 2). Until recently only one collection

from the Lewis and Clark expedition could be associated with Aira brevifolia, PH-LC 170. On that sheet the

label written by Pursh reads: “The most common g tl h the plains of Columbia & near the Kooskoo-

skee [Clearwater] R. Jun: 10th 1806.” Pursh a de: “Poa trivialis L. var.” The name of Aira brevifolia was not

written by Pursh on the sheet but added by Thomas Meehan (1826-1901, fide Moulton 1999, fig. 133 and

transcription of the annotations) probably in 1897. That specimen was reported to have been collected in

766

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Anericaa prilosepiical Soler 3

ug.

Aira Iehetes Perch

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breegets

LEWIS AND CLARK HERBARIUM

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Ei J Se yl EAr L LF dt TA L (PH LC 170) | a: gP Ej Let li J ty, Idaho, ł AA all || m$ Le

label enlarged at the top left of the sheet.

= Pi "

FL Amu. Sept: 76. Bos CC 1815, 90

Cayouette et al., Lewi 1 Clark llection, Poa secunda 767

Idaho County, Idaho (Moulton 1999; Reveal et al. 1999) and was identified as belonging to the Poa secunda

group (Hitchcock 1935; Cutright 1969; Ewan 1979; Moulton 1999; Reveal et al. 1999; Soreng 2003).

Clarification of the Lectotypification

It seems likely that the PH-LC collection consists of portions of both the Idaho and Montana collections

(Fig. 2) for the second left hand specimen (with a bent panicle) most closely resembles the specimens on

the K sheet, while the large clump to the right has roots and mature (not shattered) spikelets, and seems to

correspond to collections made earlier June 10). The K panicles, with mostly brown and shattered spike-

lets, clearly correspond to a later collection July 28). A fragment of the PH-LC lectotype is present in the

US herbarium (US-76299) and was reported in Soreng (2003: 90). A.S. Hitchcock and A. Chase used to

Sota M ps of RE material of North American grasses and they were deposited in US as reference

935). According to RJ. Soreng (pers. comm. 2006), the fragment of the panicle of the

US -76299 leues — spikelets. An examination of the spikelets of the two supposed groups of specimens

on the PH-LC sheet did not reveal a difference in color (A.E. Schuyler, pers. comm. 2006). At this time, it

is not known from which group of panicles of the PH-LC sheet the panicle fragment at US originated and

therefore its type status is in question.

Aira brevifolia Pursh was described from specimens Frederick Pursh obtained from Meriwether Lewis

who collected them during the Lewis and Clark expedition of 1804-1806 (Pursh, Fl. Amer. Sept.: 76, sero

1813). Following his description, Pursh indicated: "In the plains of Missouri. M. Lewis. June, July. v.s. in

Herb. Lewis. This grass is the most common in those plains." Unaware of the existence of the K specimen

of Aira brevifolia, Cutright (1969: 401) selected PH-LC 170 as a lectotype (as “type”) for Aira brevifolia and it

was considered as such by Reveal et al. (1999: 39).

The PH-LC and the K specimens represent the same taxon, Poa secunda J. Presl, so Pursh's description

(protologue) applies to both specimens. It seems evident that Pursh saw the two specimens and transcribed

Lewis' annotations of the labels; this is why June and July appear in the protologue. The PH-LC specimen

remained in Philadelphia while the K sheet was brought by Pursh to London in 1811 where he prepared his

flora (Moulton 1999). So Pursh had the K specimen before him when he described Aira brevifolia but not

PH-LC 170. Pursh personally wrote Aira brevifolia on the K label (Fig. 1A), and the specimen has fewer and

shorter leaves (brevifolia) than the PH-LC specimens. Pursh did not inscribe “Aira brevifolia” on the PH-LC

specimen but rather “Poa trivialis L. var." Clearly, his decision to recognize a new species was made after he

left Philadelphia in 1809. While the two collections from Idaho and Montana taxonomically represent the

same taxon, PH-LC 170 remains the lectotype of Aira brevifolia even if the K specimen would have made a

more suitable choice.

Nonetheless, as noted here, the PH-LC sheet is a mixture of two collections, one made in Idaho in June

of 1806 and a second obtained in Montana in July of 1806. Although the designation of the PH-LC as the

lectotype cannot be altered, it is necessary to restrict pH of the name to one of the two discordant

elements. Accordingly, we ide restrict the | ype to just a portion of the PH-LC specimen and designate

here (hoc loc.) as the lectot ing el t: Lewis s.n., "Doubtfull River Missouri plains," probably

near the mouth of Crow Coulee Chouteau County Montana, 28 Jul 1806, PH-LC-170, the second left hand

specimen with a bent panicle. A duplicate of the lectotype is at K.

CONCLUSION

Numerous grasses were seen by Lewis and Clark in the BUR areas of tiig pane habitats and some were

actually collected by Lewis because of their obvious or potent & Reveal 2003)

It is not clear why Lewis would have collected such a common and a grass as Sandberg’s blue-

grass (Poa secunda), but it probably was because the seeds were readily available, and he needed a voucher

specimen to go with the seeds. There might be at least another reason for this collection. By late July 1806

Lewis knew that all of his 1805 plant collections cached in Montana before crossing the Rocky Mountains

(save one, a Ribes) were lost, and obtaining a grass may have been an attempt to acquire a more diverse col-

I dal Daj . In LJ di dion de ai”

768 J f Texas 3(2)

lection. Lewis continued to collect, albeit rarely, going down river until he was accidentally wounded on

11 August (Moulton 1993).

As previously recorded, some of the K specimens of the Lewis and Clark expedition are duplicates of

the PH-LC ones (Ewan 1979; Moulton 1999; Reveal et al. 1999). Interestingly, according to Moulton (1999),

there are seven specimens found at K that have no counterpart at PH-LC. A search for other potential Lewis

and Clark authentic specimens has been intensive at these herbaria and elsewhere, and most of the pos-

sibilities have been documented, discussed, and resolved (Moulton 1999; Reveal et al. 1999).

On the other hand, despite a careful search at K by Moulton, Ewan and others investigating the Lewis

and Clark collections examined by Pursh, a grass collection of Poa secunda sensu lato was still there and

unrevealed. Is it possible that other Lewis and Clark collections brought by Pursh to London in 1811 are

still undetected within the seven million specimens of the K herbarium, one of the largest of its kind in the

world? Even if browsing the North American folders at K may represent a monumental task, it may lead to

tracing lost or unknown Lewis and Clark specimens. With the eventual data basing of the K herbarium,

such a search would be facilitated, but the largest herbaria are likely to be the last ones to have such docu-

mentation.

ACKNOWLEDGMENTS

We wish to thank J. Marsden, T. Cope, ana 5 oe (K) Ir the loan oi the Kew specimen, C. Kiel and L.A.

McDade (PH) f respectively tl the PH-LC specimen, R.J. Soreng

(US) for Mic examination of the US A ALE. les (PH) hor examination of the spikelets of the PH-

LC specimens, G. Mitrow (DAO) for the scanning of the K specimen, and J. McCarthy and E. Johnson for

preparation of the figures. EE K specimen was reproduced with kind permission of the Director and Board

of Trustees, Royal Botani dens, Kew. We thank Bernard R. Baum, Venice Bayrd, Stephen J. Darbyshire,

Ernest Small, and an anonymous reviewer for their constructive comments on earlier drafts.

REFERENCES

CutricHT, PR. 1969. Lewis and Clark: pioneering naturalists. University of Illinois Press, Urbana.

EARLE, A.S. AND J.L. ReveaL. 2003. Lewis and Clark's green world. The expedition and its plants. Farcountry Press,

Helena, MT.

Ewan, J.A. 1979. Introduction to b seine reprint i Frederick Pursh's Flora Americae Septentrionalis (1814).

In: F.T. Pursh, Flora Americae S lis (Facsimile reprint, J. Cramer, Vaduz). Pp. 7-117.

HitcHcock, A.S. 1920. The genera ases in the United States, with special reference to the ic species

U.S.D.A., Bull. 772.

HitcHcock, A.S. 1935. Manual of the grasses of the United States. U.S.D.A., Misc. Publ. 200.

McCourt, R.M. AND E.E. Spamer. 2003. The botanical legacy of Lewis and Clark: the most famous collection you

never heard of. PI. Sci. Bull. 49:126-131.

McCounr, R.M. AND E.E. Spamer. 2004a. On the paper trail in the Lewis and Clark herbarium. Bartonia 62:1—24.

McCounr, R.M. AND E.E. Spamer. 2004b. Jefferson's botanists: Lewis and Clark discover the plants of the West. Acad-

emy of Natural Sciences, Philadelphia.

Mennrrr, J.L., J. Cavouerre, J.L. REVEAL, AND G.E. MourroN. 2006, Lewis & Clark plant specimen found in London. We

Proceeded On (Lewis and Clark Trail Heritage Foundation) 32(1):12.

Mier, H.S. 1970. The herbarium of Aylmer Bourke Lambert: notes on its acquisition, dispersal, and present

whereabouts. Taxon 19:492-497.

Moutton, G.E. (ed.). 1993. The journals of the Lewis and Clark expedition. June 10-September 26, 1806. Vol. 8.

University of Nebraska Press, Lincoln.

MouttTon, G.E. (ed.). 1999. The journals of the Lewis and Clark expedition. The herbarium of the Lewis and Clark

expedition. Vol. 12. University of Nebraska Press, Lincoln.

PunsH, F.T. 1813. Flora americae septentrionalis. 2 vols. White, Cochrane, and Co., London.

Cayouette et al., Lewi 1 Clark flection, Poa secunda 769

REVEAL, J.L., G.E. MOULTON, AND A.E. SCHUYLER 1999. The Lewis and Clark collection of vascular plants: names, types,

and comments. Proc, Acad. Natl. Hist. Philadelphia 149:1-64.

SORENG, R.J. 2003. Aira L. In: Soreng, R.J., PM. Peterson, G. Davidse, E.J. Judziewicz, FO. Zuloaga, T.S. Filgueiras, and

O. Morrone. Catalogue of New World grasses (Poaceae): IV. Subfamily Pooideae. Edited by RJ. Soreng. Contr.

U.S. Natl. Herb. 48:89—96.

SPAMER, E.E. AND R.M. McCourt. 2004. The second two hundred years are better: the endearing, enduring plants

of Lewis and Clark. Bartonia 62:95-102.

[| Hn ls F dg A!

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770

BOOK REVIEW

RICHARD A. Minnicu, 2008. California's Fading Wildflowers: Lost Legacy and Biological Invasions. (ISBN

978-0-520253537, hbk.). University of California Press, 2120 Berkeley Way, Berkeley, California 94704-

1012, U.S.A. (Orders: www.ucpress.edu or California-Princeton Fulfillment Services, 1445 Lower Ferry

Road, Ewing, New Jersey 08618, U.S.A., orders@cpfsinc.com, 1-800-777-4726). $49.95, 360 pp., 55

illustrations (23 b&w photographs, 13 line illustrations, 19 tables), 6" x 9".

Contents:

1. The Golden State

2. Pre-Hispanic Herbaceous Vegetation

3. Invasion of Franciscan qu Lines: pe and California Pasture i inet y

4 AC WEIL f B 5 Wildflowers

5. Lessons “oe the Rose Parade

Appendix 1. 1 ti f Franciscan campsites, F i ] ] 1 l

E + I 4 t

Appendix 2. S ish pl for California vegetation

PP I I Califo getatio

Annendix 5 Cz] "| Li + 1 + = 1 11 > r xm 1 M : E EE ud dux

Ir I

Appendix 4. References to wildflowers in the Los Angeles Times, The Desert Magazine, 1 the Riverside Press Enterpri

ae s Rome cial by ewes A. Minnich is a derful ple of how | levelor ti , or in this case, the

FI T1 1 1-7 1 11 1 j -11 4 " 1 473.1 t TE

San Diego to San Francisco. It accurately displays how | gricultural practi d f livestock on the land—in a time when

only 200 en area—still greatly ff 1 tł ti g | f ti land to tl tral alley.

Pu book goes aa to get t information on va California from the j ls of early expl d then on into the mid 1800s

1 J f. 1 "E 11£1 1 1 1 foala

g y from the native w Rush

3 r 1 4 zT] L:f 1 1 ` 1 1 1£ J 1 1 Fo

ae he state. T I l EI

x paste +l [1 fel

e a al the book are dedicated to o ihe Bess and now posce. UE poppy nee dn There

are ih Spec ee to the cemeg bear, now extinct in Sa hich y illustrate how imp

j d as d ] ulated as California. Altogether this

[==] I P oe "I 4 e

hl | 1; 1 1 1 Hh tr] ] 1 g ] intl ] t 300 years — Reherra Swadek, Botanical

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

J. Bot. Res. Inst. Texas 3(2): 770. 2009

PINUS PINCEANA (PINACEAE), NUEVO REGISTRO PARA

EL ESTADO DE NUEVO LEON, MEXICO

Susana Favela Lara Carlos G. Velazco Macias

Laboratorio de Vida Silvestre Parques y a see de nuevo León

Facultad de Ciencias Biológicas Av. Eugen 6604

Ave dda AANO ma de Nuevo León Col. i del Bosque

Nuevo León, MEXICO Monterrey, Nuevo León, MEXICO

Glafiro J. Alanis Flores

Laboratorio de Vida Silvestre

Facultad de Ciencias mae

AVEG POROM de Nuevo León

, Nuevo León, n

RESUMEN

Se registra por primera vez a Pinus pinceana para el estado de Nuevo León, México, las poblaci tán ubicad la Sierra Madre

Oriental, en el municipio de Santa Catarina.

ABSTRACT

Tks d +1 fi 4 jr D: i] : 41 nd F AT T z : +1

Sierra Madre, i t t icipality

México ha sido reconocido como el centro con mayor riqueza de especies para el género Pinus con un total

de 43 (Farjon 1996); Pinus pinceana Gordon es uno de estos elementos endémicos dentro del género para

México, fue descrita para el centro del país en el siglo XIX (Gordon 1858), hoy en día su distribución cono-

cida abarca los estados de Hidalgo, Querétaro, Zacatecas, San Luis Potosí y Coahuila en centro y noreste de

México (Perry 1991; Villarreal 2001). Las poblaci de dicha especie se encuentran dispersas y no forman

un continuo a través de su rango de distribución, aunque son consistentes en cuanto a las características

físicas y biológicas en las que se desarrollan (Farjon y Styles 1997; Ledig et al. 2001). La especie no se en-

cuentra reportada para la flora de Nuevo León (Standley 1920; Martínez 1948; Rojas-Mendoza 1965; Capó

1972; Hinton y Hinton 1995).

La especie fue encontrada en Nuevo León por las exploraciones botánicas del Dr. Jorge Armando Ver-

duzco Martínez, alrededor del ario 2000, pero su registro no fue publicado y no se colectaron ejemplares de

herbario, se dio seguimiento a dichos trabajos de exploración en la parte norte de la Sierra Madre Oriental

con la finalidad de documentar f Imente la existencia de dicha especie para el estado, por lo que se

realizaron salidas a diversos EISE RS cañones Ubicados en la zona en cuestión, específicamente en el

municipio de Santa Catarina para ul e la especie. Una vez ubicadas, se georeferenciaron

y delimitaron las poblaciones, se realizaron anotaciones sobre el tipo de vegetación y las especies de flora

asociada, así como del medio físico en el que se desarrollan; se colectaron ejemplares para ser depositados

en herbarios. Por medio de herramientas geoespaciales ArcView y Google Earth, se elaboró un mapa de

distribución para la especie en Nuevo León.

Las poblaciones de Pinus pinceana se localizan en el cañón El Montoso, en el municipio de Santa Ca-

tarina, a 8 kilómetros en linea recta al suroeste de la ceases uu y la zona urbana de la ciudad de

O ición norte dentro

Monterrey Los grupos d

del cañón, se presenta una altitud inicial de 1 100 msnm y eee hasta los 1,700 msnm, las poblaciones

se extienden cerca de 10 km en dirección oeste de manera intermitente en cada una de las cañadas que

J. Bot. Res, Inst. Texas 3(2): 771 — 774. 2009

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Favela et al., Pinus pinceana, nuevo para Nuevo Leon 773

integran el cañón El Montoso partiendo de la localidad denominada Loma Alta el inicio del cañón (Fig.

1). Se presentan suelos de tipo litosol más rendzina con una clase textural media en los primeros 30 cm

del suelo (CETENAL 1977a); la geología se compone de rocas sedimentarias de tipo caliza y calizas más

lutitas, así como brecha sedimentaria (CETENAL 1976); el clima es de tipo seco semicálido con lluvias en

verano (BS hw), la precipitación es de entre 325 y 400 mm durante el verano y de 50 a 75 mm durante el

invierno, pueden presentarse heladas durante los meses de noviembre a marzo (Secretaría de Programaci-

ón y Presupuesto 1981). P. pinceana se integra a una vegetación de tipo matorral submontano subinerme y

matorral desértico rosetófilo (CETENAL 1977b), donde las especies más comunes son Acacia greggii A. Gray

(guajillo), Helietta parvifolia (A. Gray) Benth. (barreta), Cordia boissieri C. DC. (anacahuita), Acacia rigidula

Benth. (chaparro prieto), Gochnatia hypoleuca (DC.) A. Gray (ocotillo), Mortonia greggii A. Gray (afinador),

Calia secundiflora (Gomez-Ortega) Yakovlev (colorín), Decatropis bicolor (Zucc.) Radlk. (hoja dorada), Agave

lecheguilla Torr. (lechuguilla), Hechtia glomerata Zucc. (guapilla), Dasylirion texanum Scheele (sotol), Opuntia

stenopetala Engelm. (nopal) y Echinocereus stramineus add DA e de =

TG: 1 ] s BAC WIS re

Nuevo León

25*3746. 70"N-100?31'34.55"W, 1160 msnm, 17 Feb 2009. Velazco-Macías y Favela-Lara s.n. (ANSM, UNT).

Con anterioridad se había ya sugerido la presencia de esta especie en Nuevo León (Martínez 1948), y la

ocurrencia de poblaciones de esta especie en la Sierra de Zapalinamé y la Sierra San José de los Nuncios en

el estado de Coahuila hacen más factible la ocurrencia de otras poblaciones en el complejo orográfico de la

Sierra Madre Oriental entre los límites de Nuevo León y Coahuila. Pinus pinceana se encuentra incluido en

la norma oficial mexicana NOM-059-SEMARNAT-2001, bajo la categoría de “protección especial” (Pr), y

había sido mencionado como parte de la flora en riesgo para el estado de Nuevo León (Alanís et al. 2004); a

pesar de su cercanía con la zona metropolitana de Monterrey y la cabecera municipal de Santa Catarina, no

se observan amenazas a corto o mediano plazo como la ganadería o el impacto por turismo, sin embargo,

el desarrollo urbano y habitacional en la zona si representa una amenaza a lago plazo, cabe resaltar que

la especie se encuentra dentro de los límites del Parque Nacional Cumbres de Monterrey, lo cual podría

favorecer su conservación y protección.

AGRADECIMIENTOS

Al Biol. Héctor M. Villalón 2 u d te los trabajos de campo; Al Ing. Jorge Martínez Pineda

por la generación del mapa de distribución y el procesamiento dé la información geográfica; y a la M. en C.

Liliana Ramírez Freire por la revisión al manuscrito.

REFERENCIAS

ALANIS FLORES, G.J, C.G. VeLazco Macías, R. FOROUGHBAKHCH Porunavav, V. VALDEZ Tamez y M.A. ALVARADO VAZQUEZ. 2004.

Diversidad florística de Nuevo León: Eee en el de riesgo. Ciencia UANL. 7:209-218.

CAPÓ ARTEAGA, M. 1972. Observaci axonomía y distribución de las coníferas de Nuevo León, México

Tesis de Licenciatura, U.A.N.L. F.C.B. México

CETENAL. 1976. Carta geológica. Garza García G14C25. Escala 1:50,000. Primera edición. México.

CETENAL. 1977a. Carta edafológica. Garza García G14C25. Escala 1:50,000. Primera edición. México

CETENAL. 1977b. Carta de uso de suelo. Garza García G14C25. Escala 1:50,000. Primera edición. México

FARJON, A. 1996. Biodiversity of Pinus (Pinaceae) in Mexico: speciation and palaeo-endemism. Bot. J. Linn. Soc.

121:365-384.

FARJON, A. v B.T. Styles. 1997. Pinus (Pinaceae). Fl. Neotropica Monogr. 75. The New York Botanical Garden, Bronx.

GORDON, G. 1858. The Pinetum: being a sinopsis of all coniferous plants at present known, with descriptions,

populations and synonyms. Henry G. Bohn, London, UK.

HINTON, J. Y G.S. Hinton. 1995. Checklist of Hinton's collection of the flora of south tral N

Coahuila. Acta Bot. Mex. 30:41—1 12.

| fal n.a . in LI bi bak fT Tio A!

77A

Lepi, ET. M.A. Capo AnrEAGA, PD. Hopaskiss, H. SaAv, C. Flores Lopez, M.T. CONKLE, AND B. BERMEJO VELAZQUEZ. 2001. Ge-

netic diversity and the mating system of a rare Mexican piñon, Pinus pinceana, and a compari with Pinus

maximartinezii (Pinaceae). Amer. J. Bot. 88:1977-1987.

Martinez, M. 1948. Los Pinos Mexicanos. Segunda edición. Universidad Autonóma de México. México.

Perry, J.P. 1991. The pines of Mexico and Central America. Timber Press, Portland, Oregon.

SECRETARÍA DE PROGRAMACIÓN Y PRESUPUESTO. 1981. Carta estatal, climas. Estado de Nuevo León. México.

SrANDLEv, PC. 1920. Tress and shrubs of Mexico (Gleicheniaceae — Betulaceae). Contr. U.S. Natl. Herb. 23(1):

1-170.

Rojas-MENDOZA, P. 1965. Generalidades sobre la vegetación del estado de Nuevo León y datos acerca de su flora.

Tesis doctoral. Facultad de Ciencias, ell nia Daci: Ali bs México.

VILLARREAL QUINTANILLA, J.Á. 2001. Flora de Coahuila. Listad xico. Instituto de Biología. Universidad

Nacional Autónoma de México. México D.F.

DIVERSIDAD Y DISTRIBUCION DE LAS GRAMINEAS (POACEAE)

EN EL ESTADO DE ZACATECAS

Yolanda Herrera Arrieta y Armando Cortés Ortiz

Instituto Politécnico Nacional

CHDIR Unidad Durango

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PALABRAS CLAVE: Gramíneas, Poaceae, pat de distribución, Zacatecas, México

ABSTRACT

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I p g 4 ; Zacatecas ) y of the

family in the state. The distribution of species was studied within four g l types of tati d altitudi The state was

Rad in ques of 0.5 x 0.5 REEL I I ber of collecti | species were placed. We apres the Diva- GIS program (Hi-

JANAN + -l /T m E f QM L : +1 AXI

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INTRODUCCIÓN

Las gramíneas son elementos importantes en la composición florística de varios tipos de vegetación y en

ocasiones forman comunidades vegetales denominadas pastizales, donde juegan un papel predominante

(Rzedowski 1978). México cuenta con una importante superficie cubierta por pastizales de un 18 a 20 %

del territorio (Rzedowski 1978); en Zacatecas un porcentaje similar al del pais está cubierto por pastizales

naturales, lo que ha permitido que la producción ganadera sea una de las actividades económicas de mayor

relevancia en el estado (INEGI 2006). Los pastizales de Zacatecas, además de su reconocida capacidad de

producción de ganado vacuno para carne en terrenos de pastoreo extensivo, revisten una gran importan-

cia ecológica por formar parte del continuo de pastizales del SW de Estados Unidos y el NW de México,

tratándose de comunidades vegetales cuyo origen y formación se remontan a cerca de 20 millones de años

atrás (Bailey 2000). Las actividades humanas de los últimos siglos están causando la desertificación de los

pastizales con el sobrepastoreo y la roturación para terrenos de cultivo, además de las sequías prolongadas,

fuegos e invasión de especies exóticas que incrementan la vulnerabilidad de estas comunidades.

De las plantas vasculares, la familia de las gramíneas es una de las más grandes y más distribuidas

en el mundo, se encuentra desde los círculos polares hasta el ecuador, en las cumbres de las montañas y

al nivel del mar (Hitchcock 1951). Por su número de especies se estima que ocupa el cuarto lugar a nivel

mundial después de las compuestas, las leguminosas y las orquideas (Lawrence 1951); en México ocupan el

tercer lugar superando en número a las orquídeas (Rzedowski 1978). No obstante, ocupan el primer lugar en

cuanto a su importancia económica, a que pertenecen a esta familia los cereales como el arroz, trigo, maíz

y caña de azúcar, base de la ali 5n humana. Es la familia de plantas con mayor importancia ecológica

] Bat Rac Inet Tayas 3(2): 77^ 792. 2009

776 t tani titute of Texas 3(2)

por su diversidad, por su característica capacidad de formar suelos y porque la mayoría de sus especies son

elementos naturales de vegetaciones primarias, aunque también un porcentaje no muy alto de ellas (<5%

estimado) han desarrollado la capacidad de dispersarse por el mundo y establecerse como adventicias en

hábitats diversos.

Se estima que existen alrededor de 700 géneros y 10,000 especies de gramíneas en el mundo (Clayton

Sr Renvoize 1986), de ellos 204 géneros y 1182 especies se encuentran en México (Dávila et al. 2006). El

de: DUE HE los Bee y ni de las especies distribuidas en México se encuentran en el estado de Zacatecas

91 géneros y 284 especies, 6 subespecies, 30 variedades y 4 formas (Herrera

et al. 2009).

La diversidad biológica o od es la “variedad de vida” y se refiere de manera general a la varia-

ción en todos los nive les de organización biológica. Existen has definiciones, DeLong (1996) registra 85

definiciones de la diversidad biológica o biodiversidad. La biodiversidad puede dividirse en tres grupos: 1)

diversidad genética, 2) diversidad de Mns vivos y 3) diversidad ecológica PURA & peers 20 ua La

AA £100 mila actri1r4+11 nal ES

diversidad genética abarca | d go genético que (nucledti

dos, genes, SINGERS y la variación de estructura genética entre individuos en da seno ids una población

y entre poblaciones. La diversidad de organismos abarca la jerarquía taxonómica y sus componentes, desde

los individuos hasta categorías taxonómicas de orden superior. La diversidad ecológica abarca las escalas

de diferencias ecológicas desde las poblaciones pasando por nichos y hábitats hasta más allá de los biomas

(Gaston & Spicer 2007). De manera que la biodiversidad puede ser medida de muchas maneras, en la prác-

tica tiende a ser medida más frecuentemente en términos de riqueza de especies, esto es, por el numero

de especies. Esta medida de biodiversidad de organismos es muy comün debido a su aplicación práctica,

información existente, sustitución, aplicación amplia (Gaston & Spicer 2007). La familia de las gramíneas

parece ser interesante como un indicador de biodiversidad, por ser un grupo importante en la riqueza

florística mexicana, especialmente en la región norte de México (Challenger 1998). Con base en lo anterior

se propone llevar a cabo un análisis de la diversidad relativa de organismos y la distribución geográfica de

las gramíneas encontradas en 4 tipos de vegetación principales del estado de Zacatecas, con base en datos

georreferidos de colectas y del conocimiento actualizado de los taxa que conforman esta familia (Herrera et

al. 2009

MATERIALES Y MÉTODOS

Área de estudio

El estado de Zacatecas se localiza en la porción centro-norte de la Repüblica Mexicana. con coordenadas

extremas 25.12? a 21.04? N y -104.33? a -100.74? O, rodeado por los estados de Durango, Coahuila, Nuevo

León, San Luis Potosí, Jalisco, Aguascalientes y Nayarit, segün el Mapa Geoestadístico de México elaborado

por INEGI, 1996. El estado de Zacatecas posee un territorio muy irregular en cuanto a su topografía, ex-

isten sitios que tienen una altitud de 600 a 700 metros sobre el nivel del mar hasta 3192 metros. Cerca del

7596 de su superficie se encuentra en una altitud entre 1800 y 2400 metros, con un máximo en el intervalo

de 2000 a 2200, segün los datos de altitud (Modelo Digital de Elevación) obtenidos por la misión SRTM

(Shuttle Radar Topography Mission) de la NASA (2000). A pesar de las diferencias en altitud, el terreno de

Zacatecas tiene pendientes poco pronunciadas, cerca del 7596 presenta pendientes menores de 8 grados,

según los cálculos realizados con el MDE del SRTM. Las zonas que presentan mayor frecuencia de fuertes

pendientes se localizan al sur, suroeste y en el complejo montañoso del norte del estado. Por su fisiografía

Zacatecas se divide en las provincias: Mesa Central (44.896), Sierra Madre Occidental al sur (38.98), Sierra

Madre Oriental al norte (15.3496) y Eje Neovolcánico al suroeste (0.8596). Los procesos geohidrológicos

han creado en el terreno zacatecano una serie de formas del terreno, llamados sistemas de topoformas en

el diccionario de datos fisiográficos, escala 1: 1000,000 (INEGI 1998b), conocidos como llanuras, bajadas,

lomeríos, sierras, cafiones, mesetas. Siendo las bajadas y las sierras los más comunes en la entidad, segui-

dos de las llanuras y lomeríos. Por su ubicación geográfica y características topográficas el territorio de

Li r E FK iH [| al m | 24-7

5 1 + + 117

FISTING y

Zacatecas presenta un clima del grupo B [BS, seco (30.21%) y BS, semiseco (48.34%)], seguido por el grupo

C(w) templado subhúmedo (21.16%) y (A)C(w) cálido subhúmedo (0.29%) (INEGI 19802). Debido a las

características climáticas, edáficas y de relieve, la cobertura vegetal del estado está dominado por matorral

xerófilo (40%), seguido por pastizal (16%), bosque templado (14%) constituido por latifoliadas y coníferas,

bosque tropical (2.5%) y áreas agrícolas (27.5%), esta ultima representa la cuarta parte de la superficie del

estado (INEGI 2003).

Materiales

Se utilizaron: a) los registros digitales de los ejemplares de gramíneas colectados en el estado de Zacatecas

y existentes en diversos herbarios; b) el marco geoestadístico de la República Mexicana en formato digital

elaborado y distribuido por INEGI (1996); c) el conjunto de datos de uso del suelo y vegetación a escala

1:250 000 elaborado por INEGI (2003); y d) el Modelo Digital de Elevación (MDE) generado por la misión

SRTM; y e) los Programas de cómputo: Diva-SIG.

Material botánico

Se recopilaron los datos de las colectas realizadas en el estado y dep en i (CHA-

PA, CIIDIR, ENCB, HUAZ, IEB, INEGI, MEXU y SLPM) y de Estados Unidos de America (GH, MO, TAES,

TEX y US), donde se concentran los ejemplares colectados en el estado durante los últimos 100 años. Estos

datos fueron capturados en el sistema Biótica (Base de datos de la CONABIO), con la que se realizó un

estudio florístico del estado de Zacatecas (Herrera et al. 2009). Entre los datos de las colectas se incluyó

las coordenadas geográficas del sitio, que fueron establecidas a partir de los mapas topográficos de INEGI

(1998a) escala 1:250 000.

Métodos

Se realizó un análisis de número de colectas, especies e índice de intercambio mediante la ejecución de los

siguientes pasos: 1. Elaboración del mapa estatal con colectas; utilizando el marco geo-estadístico nacional

de INEGI (1996), dicho mapa se manejó en coordenadas geográficas. Los sitios de colecta de las especies se

sobrepusieron al mapa estatal, por medio de las coordenadas en grados decimales. 2. Elaboración de una

cuadrícula; se construyó una cuadrícula n en e mapa, formada por cuadros de 0.5 por 0.5 grados.

Dicha cuadrícula se utilizó para realizar diversidad, como se indica en las etapas siguientes.

3. w ii número de gn y de especies en la cuadrícula; se determinó e nümero D mapas de las

o, así como el número t i | idad de especies)

Esto se realizo con ayuda del programa Diva-GIS. 4. Determinación de la relación EN número de colectas y

número de especies. La relación de un número de muestras y el número de especies se realizó por medio de

una regresión lineal y una regresión logarítmica, por medio de algoritmos incorporados en el programa de

aplicación Diva-GIS. 5. Cálculo del índice de intercambio de especies, se estimó un indice de intercambio

para conocer cuales cuadros tienen mayor posibilidad de intercambio de especies, para lo que se aplicó el

indice de Whitaker B, = (S/a)- 1, donde S = número total de especies en las celdas consideradas, a = número

promedio de especies en las celdas consideradas. El índice fue calculado con una de las funciones integradas

en el programa Diva-SIG. Con la opción *Queen case" que calcula el índice para cada celda considerando

sus ocho vecinos (2 horizontales, 2 verticales y 4 diagonales).

El intercambio, también llamado Beta-diversidad, es una medida de intercambio de especies, indica

cuan diferentes o similares son un nümero de áreas cercanas. Dos grandes áreas con nümero de especies

similares, la primera con especies diferentes en todas las celdas de la cuadrícula, y la otra con la misma

especie en todos sus cuadros. La primera tendría una elevada diversidad beta, y la segunda tendría una baja

diversidad beta (Hijmans et al. 2004).

RESULTADOS

Para llevar a cabo el análisis de diversidad se generó un mapa estatal con una cuadrícula de 30 x 30 minutos

(0.5 grados), el número de cuadros fue de 50 como se aprecia en la Figura 1.

778 J laf the Ratanical R h Inctituta afT

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Fc1M A 1. n^ ro r J i 4 * d J z d J

El námero de registros (ejemplares de herbario) considerados en este trabajo tue de 2408, cuyos sitios

georreferidos de muestreo se indican en la Figura 1. En cada cuadro existe un numero distinto de colectas

o registros, variando desde 0 a más de 200.

El número (riqueza) de taxa trad cada cuadro también varía desde 0 a más de 100. Es notorio

que la cercanía a las ciudades y a las vías de infl en el numero de especies colectad

(Fig. 2).

Al observar los mapas de las Figuras 1 y 2, se puede apreciar cierta relación entre el número de colec-

tas y el número de especies presentes en cada cuadro, para confirmar esa relación se realizó una regresión

lineal entre las dos variables (número de colectas y diversidad), se confirmó que existe esa relación como se

puede ver en el gráfico de la Figura 3, donde la regresión lineal tiene un Coeficiente de correlación (^) de

0.9498.

y pr esentes

En la regresión logarítmica muestra poca relación entre ambas variables (nümero de colectas y diversi-

dad), esto se puede apreciar fácilmente por la distribución de los puntos en relación a línea de la gráfica de

la Figura 4, donde la regresión logarítmica tiene un Coeficiente de correlación (r^) de 0.7627.

Plantas colectadas

En »] na & E L - ( = ) J F || ri I

E] resultado del análisis de intercambio fue un mapa que indica las celdas con mayor posibilidad de

intercambio de especies (Fig. 5). En el mapa se aprecia que el valor de intercambio es mayor en la parte

suroeste del estado, en donde se presenta mayor variación ambiental. Mientras que en la parte noreste

presenta los valores de intercambio más bajos, debido a que la variación climática y del medio físico son

relativamente más uniformes. Esta división coincide en términos generales con la distribución reportada

por el INEGI: de la cubierta vegetal (2003), los rasgos climáticos (19803), fisiográficos (19805) y geológicos

del estado (19800).

En cuanto a la distribución altitudinal de las especies, la frecuencia es mayor en el intervalo altitudinal

de 2020 a 2440 m, como puede verse en la gráfica de la Figura 8. Donde el 6896 de las gramineas prefieren

este intervalo altitudinal.

Otras especies se favorecen por el sustrato donde se desarrollan (Tabla 1) como: I) especies adaptadas a

suelos salinos, alcalinos o yesosos como oe curvifolia, Boutel , Eragrostis obtusiflora, Erioneuron

pilosum, Paspalum crinitum y Pleurt das en suelos alcalinos; Bouteloua Lamina Distich-

lis spicata, Sporobolus airoides y S. iaa en suelos salinos y Bouteloua Fu B. karwinskii, B. ramosa,

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acuáticos 6 subacuáticos para su ets como CI

E. crusgalli, E. cruspavonis, E. holciformis, E. oplismenoides, E obtusiflora, Enola acuminata, Leersia

hexandra, Leptochloa aquatica, L. fusca, Paspalum arsenei y P. denticulatum, P. distichum que crecen en ambien-

tes acuáticos. een lid Guadua angustifolia, Panicum decolorans, P. elephantipes, Pappophorum

vaginatum, Polypog 1 , Setaria adhaerens, Sporobolus pyramidatus y Urochloa texana de ambientes

3 y tol tes a vivir bajo el agua en alguna época como Arundo donax, Chloris rufes-

cens, Digitaria ternata, Eleusine indica, Leptochl , Muhlenbergia diversiglumis, M. polycaulis, M. rigens,

Peyritschya deyeuxioides, P. pringlei, Polypogon ONZAS, P. viridis, Tuum duds y Urochloa plantaginea.

DISCUSIÓN Y CONCLUSIONES

La familia Poaceae parece ser la segunda más diversa en el estado de Zacatecas, con sus 91 géneros, 284

especies y 40 infraespecies, es superada con mucho por la familia Asteraceae que cuenta en esta enti

141 géneros, 456 especies y 119 infraespecies (Balleza y Villasefior 2005). Existe una distribución de los taxa

de acuerdo a las características ecológicas del estado que concuerdan con lo encontrado y presentado por

Balleza y Villasefior (2005), aun con las mismas limitaciones muestrales existentes en ambos estudios.

Los tipos de cobertura vegetal que resultaron tener mayor diversidad de gramíneas son los bosques

templados de encino-pino con 172 taxa (3296), le sigue el pastizal con 157 taxa (29.596), el matorral con 139

taxa (26.196) y el bosque tropical con sólo 64 (1296) (Tabla 1 y Fig. 7). Datos que coinciden con las propor-

ciones de endemismos reportados por Rzedowski (1995) para la flora de diferentes tipos de vegetación de

México. Comparando la riqueza relativa (Fig. 5) con el área cubierta por los tipos de vegetación (Fig. 6),

puede notarse que los cuadros del noreste y este correspondientes al área de los matorrales de la región

más árida del estado, son los menos diversos con 69 taxa (1596), donde los matorrales son comunidades de

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tipo micrófilo y rosetófilo, en un gradiente altitudinal de 1640 a 2300 m. El área media del estado (cuadros

naranja en Fig. 5) que corresponde a la región de pastizales, muchos l ora abiertos a la agricultura,

con algo de bosque en la a oeste y a entreverados, presenta la diversidad más alta con 230

taxa (49%), con el gradiente altitudinal lo (1970 a 2440 m). Mientras que el área de los cuadros

del suroeste (en e Fig. 5) se asocia a climas templados en las partes altas de la sierra, semiáridos en las

partes medias y semicálidos en las partes bajas, tienen una riqueza media de 169 taxa (3696) en relación a

los anteriores. Está región alberga los 4 tipos de vegetación aquí considerados de forma mezclada; bosques

templados primordialmente de encino y encino-pino en las partes altas, pastizales y matorral xerófilo con

elementos crasicaules y arbustos espinosos en las partes medias, y matorral subcaducifolio en las tierras

bajas; en un gradiente altitudinal de 1100 a 2720 m

Los 295 taxa (91 géneros y 284 especies 6 subespecies, 30 variedades y 4 formas en Anexo Tabla 1)

encontrados en Zacatecas por Herrera et al. a forman parte de ii de las 10 e ptadas para

México por Davila cot oe La mayor di se encuentra en la subfamilia Chloridoideae que resultó

ser la mejor en 125 taxa (42.37%) de la diversidad total, por ser este estado parte

de la región de alta diversidad lada para las Chloridoideae por Valdés y Cabral (1993). Esta subfamilia

incluye en su haber a los 3 géneros con mayor número de taxa: Muhlenbergia (43), Bouteloua (25), Eragrostis

(16), llegando a concentrar estos 3 aaa casi 1/3 de la diversidad de la familia (84 taxa = 28.47%). Le sigue

en importancia la subfamilia Panicoideae con 93 taxa (31.52%) de la diversidad total, lo que se exp

influencia que la Sierra Madre Oriental tiene sobre Zacatecas (Valdés y Cabral 1993). La subfamilia Pooideae

le sigue con 53 taxa (17.97%) de la riqueza total. Las subfamilias Arundinoideae (2 taxa), Bambusoideae (3

taxa), Danthonioideae (1 taxa) y Ehrhartoideae (2 taxa), se encuentran poco representadas en la entidad,

lo que se explica por ser subfamilias de origen y distribución en ecosistemas tropicales húmedos hasta ser

pol la

especies acuáticas.

782 tani titute of Texas 3(

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kilometers

En 5 ARA I || 3. ed je J a F1 L? MA |

P finidades geográficas de las 284 ies de gramíneas de Zacatecas, 189 (66-5%) son nativas,

40 especies (14%) son introducidas (o exóticas) y 55 (19.5 96) son endémicas de México. De]

son nativas de Zacatecas se encuentran 82 especies de afinidad Holártica (43.2%), 28 especies de afinidad

D dn oe a 79 epee de toda na s Mad) debido a la ubicación de esta entidad en la región

Si Madre Occidental, Sierra Madre

Oriental y A Neovolcánico) y Neotropical (en is: provadis: Mesa Central y Sierra Madre Oriental). De

particular importancia son las 55 especies presentes en Zacatecas que son endémicas de México (19.5%),

de las cuales sólo 7 especies son endémicas de Zacatecas o de Zacatecas con algún estado vecino (2.48%):

Agrostis rosei, Blepharoneuron shepherdii, Bouteloua reederorum, Festuca roblensis, Muhlenbergia durangensis,

M. flaviseta y M. pubigluma. Dicho porcentaje de endemismos de Gramineas en Zacatecas se aproxima, sin

llegar al estimado por Rzedowski (1991) para esta familia en México (25%), o al 30% citado por Valdés y

Cabral (1993). La calidad de los pastizales de Zacatecas para la producción pecuaria, se debe a que los taxa

(especies e infraespecies) de gramíneas de esta región son por su origen principalmente de tipo nativo, y en

escasa proporción exóticas como puede apreciarse en la Figura 9.

| |

BON QU

783

Tasta 1. Relación de grami le Z bfamilia, t q

Formación vegetal Origen

Bos- Selva Mato- Past- Otros Nativa End- Exótica Sub-

Especie Que rral izal émica familia

Achnatherum editorum X X M Poo

Achnatherum eminens X X X Na Poo

Achnatherum multimode X M Poo

Aegopogon cenchroides X Sa Chl

Aegopogon tenellus X x Na Chl

Agrostis hyemalis X A Poo

Agrostis rosel X D,Z Poo

Andropogon gerardii x A Pan

Andropogon glomeratus X VS A Pan

var. pumilus

Andropogon pringlei X M Pan

Andropogon virginicus x Na Pan

Aristida adscensionis X X x X C Ari

Aristida appressa X X M Ari

Aristida arizonica X Na Ari

Aristida curvifolia X X SA M Ari

Aristida divaricata X X X X Na Ari

Aristida havardii x X x Na Ari

Aristida laxa var. laxa X X Sa Ari

Aristida pansa fo. dissita MDM Na Ari

Aristida pansa fo. pansa X X Na Ari

Aristida purpurea var. nealleyi X X Na Ari

Aristida purpurea var. purpurea X X Na Ari

Aristida purpurea var. wrightii X X Na Ari

Aristida schiedeana X X X Na Ari

var. schiedeana

Aristida scribneriana X X M Ari

Aristida ternipes var. gentilis X X Na Ari

Aristida ternipes var. ternipes X X X X A Ari

Arundo donax fo. donax X X VR, T X Aru

Arundo donax fo. versicolor x x VR, T X Aru

Avena fatua X AC X Poo

Avena sativa AC, AD x Poo

Blepharoneuron shepherdii X Ch,D,So,Z Chl

Bothriochloa alta X Ca Pan

Bothriochloa barbinodis X X X A Pan

Bothriochloa hirtifolia x X Ca Pan

Bothriochloa hybrida X X Na Pan

Bothriochloa ischaemum X X Pan

Bothriochloa laguroides X X AC Sa Pan

ar. torreyana

Bothriochloa perforata X X X Na Pan

Bothriochloa wrightii X Na Pan

Bouteloua aristidoides X x A Chl

Bouteloua barbata var. barbata X X A Chl

Bouteloua chasei MDM SY M Chl

Bouteloua chondrosioides X X X Na Chl

Bouteloua curtipendula X K X X Sa Chl

var. caespitosa

Bouteloua curtipendula MDM x BG Na Chl

var. curtipendula

784

TagLa 1. continuación

ia Y de A!

f Texas 3(2)

Formación vegetal Origen

Bos- Selva Mato- Past- Otros Nativa End- Exótica Sub-

Especie rral izal émica familia

Bouteloua curtipendula MDM x M Chl

var. tenuis

Bouteloua dactyloides X X Na Chl

Bouteloua diversispicula X X Ca Chl

Bouteloua erecta X Na Chl

Bouteloua eriopoda X Na Chl

Bouteloua gracilis X X X Na Chl

Bouteloua hirsuta X X Na Chl

var. glandulosa

Bouteloua hirsuta var. hirsuta X X X Na Chl

Bouteloua karwinskii MDM x SH,SY M Chl

Bouteloua parryi var. parryi BG Na Chl

Bouteloua polymorpha x X M Chl

Bouteloua dod X x Na Chl

Bouteloua ramosa MDM SA,SY Na Chl

Bouteloua reederorum X x Ag,D,SL,Z Chl

Bouteloua repens X x X Na Chl

Bouteloua scorpioides X X X M Chl

Bouteloua simplex x X X A Chl

Bouteloua uniflora X X M Chl

var. coahuilensis

Bouteloua williamsii X X X Ca Chl

Brachypodium mexicanum X X X Sa Poo

Bromus anomalus X X X Na Poo

Bromus carinatus X X X Na Poo

var. californicus

Bromus en X AD Sa Poo

var. catharticu

Calamagrostis oe X M Poo

Cenchrus brownii X AD X Pan

Cenchrus ciliaris X X AD, VR X Pan

Cenchrus echinatus X X x Pan

Cenchrus incertus X X X AD A Pan

Cenchrus multiflorus X M Pan

Cenchrus pilosus MDM Sa Pan

Chaboissaea ligulata X X VÀ M Chl

Chascolytrum subaristatum X Sa Poo

Chloris gayana X X X Chl

Chloris rufescens X X Ti Ca Chl

Chloris submutica X X X AD A Chl

Chloris virgata X X X X ADVR C Chl

Cortaderia selloana Or Sa Dan

Cottea pappophoroides X X A Chl

Ctenium planifolium X M Chl

Cynodon dactylo X X X X VR x Chl

Dactyloctenium aegyptium X X Chl

Dasyochloa pulchella x X VR Na Chl

Dichanthelium sphaerocarpon x A Pan

Digitaria califérnica X X X VR Na Pan

Digitaria ciliaris X X X VR E Pan

Digitaria cognata MDM x Na Pan

d. | ll An T

TABLA 1. continuación

785

Formación vegetal Origen

Bos- Selva Mato- Past- Otros Nativa End- Exótica Sub-

Especie rral izal émica familia

Digitaria filiformis X X Na Pan

igitaria panicea X X M Pan

Digitaria sanguinalis X X Pan

Digitaria ternata x X T X Pan

Distichlis spicata var. mexicana X SH,VA A Chl

Echinochloa colona X X VA, VR X Pan

Echinochloa crusgalli X X X VA,VR A Pan

var. crusgalli

Echinochloa crusgalli X X X VA VR A Pan

var. zelayensis

Echinochloa crus-pavonis X VA A Pan

var. crus-pavonis

Echinochloa holciformis VA, VR Ca Pan

Echinochloa jaliscana X X X M Pan

Echinochloa oplismenoides X X VA Na Pan

Eleusine indica X X X T X Chl

Eleusine multiflora X X Chl

Elionurus barbiculmis X X Na Pan

Elionurus tripsacoides X Sa Pan

Elymus arizonicus X Na Poo

Elymus elymoides x Na Poo

subsp. brevifolius

Elymus repens subsp. repens x AD x Poo

Enneapogon desvauxii X VR A Chl

Enteropogon chlorideus X AD Na Chl

Fragrostis barrelieri VR X Chl

Eragrostis cilianensis X VR X Chl

Fragrostis X x Chl

Eragrostis erosa Na Chl

Eragrostis hirta X Ca ChI

Eragrostis intermedia Na Chl

var. appressa

Eragrostis lehmanniana X X Chl

Eragrostis lugens X X A Chl

Eragrostis mexicana X x X VR A Chl

subsp. mexica

Eragrostis obtusiflora x SA,VA Na Chl

Eragrostis palmeri X X Na Chi

Eragrostis pectinacea X X X A Chl

var. miserrima

Eragrostis pectinacea X X X A Chl

var. pectinacea

Eragrostis plumbea X X M Chl

Eragrostis pringlei M Chl

Fragrostis superba X X Chl

Friochloa acuminata X X VR,VA Na Chl

var. acuminata

Erioneuron avenaceum X A Chl

var. avenaceum

Erioneuron nealleyi X Na Chl

Erioneuron pilosum SA A Chi

Festuca arizonica x Na Poo

786

Tasta 1. continuación

Formación vegetal

Selva Mato- P

Origen J—. .

Bos- ast- Otros Nativa End- Exótica Sub-

Especie rral izal émi familia

Festuca pringlei X M Poo

Festuca roblensis x X Poo

Festuca rosei X M Poo

Gouinia virgata var. robusta Sa Ch!

Guadua angustifolia X VS Ca Bam

Heteropogon contortus xX X X A Pan

Heteropogon melanocarpus x X A Pan

Hilaria cenchroides X X X Ca Chl

Hilaria swallenii X Na Chl

Hopia obtu X X X VR Na Pan

Hordeum jubatum AD A Poo

Hordeum vulgare AC Poo

Koeleria pyramidata X E Poo

Lasiacis nigra Sa Pan

Leersia hexandra VA, VR Na Ehr

Leptochloa aquatica VA M Chl

Leptochloa dubia x X X A Chl

Leptochloa fusca

subsp. fascicularis X VA,VR A Chl

subsp. uninervia VA A Chl

Leptochloa panicea X VR, T A Chl

subsp. brachiata

Lolium multiflorum X X Poo

Lolium perenn X A Poo

Lolium temulentum AD A Poo

Luziola fluitans X VA NA Ehr

Lycurus phalaroides X X x A Chl

Lycurus phleoides X x X Na Chl

Melinis repens X X X Pan

Metcalfia mexicana X Poo

Microchloa kunthii X X X A Chl

Mnesithea granularis X Pan

Mnesithea ramosa VR SA Pan

Muhlenbergia alamosae X x M Chl

Muhlenbergia arenacea X x X Na Chl

Muhlenbergia arenicola X X A Chi

Muhlenbergia brevis X Na Chl

Muhlenbergia brevivaginata X M Chl

Muhlenbergia ciliata X x Sa Chl

Muhlenbergia crispiseta X BG Na Chl

Muhlenbergia depauperata X X Na Chl

Muhlenbergia diversiglumis X X T Sa Chl

Muhlenbergia dubia X Na Ch!

Muhlenbergia dumosa MDM Na Chl

Muhlenbergia durangensis x Ch,D Chi

Muhlenbergia emersleyi X X X VR Na Chl

Muhlenbergia eriophylla x M Chi

Muhlenbergia flaviseta X X Ch,D Chl

Muhlenbergia fragilis X Na Chl

Muhlenbergia glauca X Na Chl

Muhlenbergia implicata X Sa Chl

Muhlenbergia longiglumis X M Chl

Y | pa | A, T

TABLA T. continuación

Formación vegetal Orige

Bos- Selva Mato- Past- Otros Nativa End- Exótica Sub-

Especie e izal émic amilia

Muhlenbergia microsperma X X A Chl

Muhlenbergia minutissima X Na Chl

Muhlenbergia montana x Na Chl

Muhlenbergia mucronata X M Chl

Muhlenbergia pectinata » Na Chl

Muhlenbergia peruviana X X BG A Chl

Muhlenbergia plumbea X Ca Chl

Muhlenbergia polycaulis x x X VR, T Na Chl

Muhlenbergia porteri X VR Na Chl

Muhlenbergia pubescens X X X M Chl

Muhlenbergia pubigluma X X X Co,NL,Z Chl

Muhlenbergia quadridentata X M Chl

Muhlenbergia repens X Na Chl

Muhlenbergia rigens X X VR, T Na Chi

Muhlenbergia rigida X X X A Chl

Muhlenbergia robusta X X Ca Chl

Muhlenbergia scoparia X M Chl

Muhlenbergia speciosa x M Chl

Muhlenbergia stricta X M Chl

Muhlenbergia tenuifolia X X X AD A Chl

Muhlenbergia texana X X Na Chl

Muhlenbergia utilis X Na Chl

jede villiflora X X M Chl

var. viiliflor

Muhle Eno virescens X Na Chl

ns leucotri x Na Poo

Nassella Oo X Sa Poo

Nassella tenuissima X A Poo

Oplismenus burmannii BG Sa Pan

var. burmannii

Otatea acuminata X M Bam

subsp. aztecorum

Panicum antidotale AC X Pan

Panicum decolorans VSVR M Pan

Panicum elephantipes VSVR A Pan

Panicum hirticaule X X Na Pan

var. verrucosum

Panicum lepidulum X X Ca Pan

Panicum vaseyanum X M Pan

Panicum virgatum X Na Pan

Ra M al vaginatum X VS A Chl

Paspalum arsenei VA M Pan

Paspalum convexum X X Sa Pan

Paspalum crinitum SA,V.S M Pan

Paspalum denticulatum VAVR A Pan

Paspalum distichum X X x VA A Pan

Paspalum humboldtianum X X SA Pan

Paspalum jaliscanum X X CA Pan

Paspalum notatum X A Pan

Paspalum paniculatum SA Pan

Paspalum plicatulum X X A Pan

Paspalum tenellum X X SA Pan

788

TABLA 1. continuación

Formación vegetal Origen

Bos- Selva Mato- Past- Otros Nativa End- Exótica Sub-

Especie rral izal émica familia

Pennisetum clandestinum Or X Pan

Pennisteum crinitum X X X M Pan

Pennisetum villosum X X X x Pan

Pereilema ciliatum x Ca Chl

Pereilema crinitum x X X Sa Chl

Peyritschia deyeuxioides X X VR, T Sa Poo

Peyritschia pringlei X VR, T Sa Poo

Phalaris canariensis X X Poo

Phyllostachys aurea X AC X Bam

Piptochaetium fimbriatum X X X Na Poo

Piptochaetium virescens X Ca Poo

var. virescens

Pleurhaphis mutica MDM Xx SA,SY Na Chl

Poa annua X X X A Poo

Poa strictira X Na Poo

Polypogon ner x X T A Poo

Polypogon monspeliensis x VS A Poo

Polypogon viridis X x E A Poo

Saccharum officinarum AC X Pan

Schedonorus arundinaceus AC X Poo

Schizachyrium brevifolium X X SA Pan

Schizachyrium cirratu X X A Pan

hizachyrium sanguineum X X X BG A Pan

Schizachyrium tenerum X X A Pan

da uM orevifolus X X A Chl

Setaria adhaerens x VR, VS X Pan

Setaria pel x X X VR A Pan

Setaria leucopila X X X A Pan

etaria macrostachya X X A Pan

Setaria parviflora X X X X AD A Pan

Setariopsis auriculata X X Sa Pan

Setariopsis latiglumis X M Pan

Sorghastrum incompletum X X X Sa Pan

Sorghastrum nutans X X A Pan

Sorghum bicolor X x Pan

Sorghum halepense X X X X Pan

Sporobolus airoides X PH Na Poo

var. airoides

Sporobolus atrovirens X X X M Poo

Sporobolus cryptandrus X A Poo

Sporobolus indicus var. indicus x X X X VR A Poo

Sporobolus macrospermus MDM Ca Poo

Sporobolus palmeri X SASY M Poo

Sporobolus pyramidatus X PH SA,VR, A Poo

Sporobolus trichodes X X M Poo

Sporobolus wrightii X X SA Na Poo

Steinchisma cuprea X X M Pan

Trachypogon spicatus X X X A Pan

Tragus berteronianus X X VR X Chl

Tridens muticus X VR Na Chl

Tripogon spicatus X A Chl

Tasa 1. continuación

Formación vegetal Origen

Bos- Selva Mato- Past- Otros Nativa End- Exótica Sub-

Especie Que rral izal émica familia

Tripsacum dactyloides X X X jji Na Pan

ar. dactyloides

scum lanceolatum X x Na Pan

Tripsacum pilosum X a Pan

Tripsacum zopilotense X Ca Pan

Trisetum viride X X Ca Poo

Triticum aestivum X x E Poo

Urochloa fusca X X VR A Pan

Urochloa meziana x M Pan

Urochloa panicoides X X X Pan

Urochloa plantaginea X X X Tm A Pan

Urochloa texana X x X VS Na Pan

Vulpia myuros var. hirsuta X Sa Poo

s subsp. mays AC M Pan

Zuloagaea bulbosa X X X BG M Pan

Por su M cae en: A= América*, C= Cosmopolita, Ca= Centroamérica**, M= México, Na= Norteamérica***, Sa=

Sudamérica**

an sa en f todo el continente americano

en ee O

Norteamérica***= pd Endo O Estados ccs EE:

Sudamérica****z desde CO UTE E

Endémicas en los Estados de: Ag= Ag lientes, Ch= Chihuahua, D= Durango, J= Jalisco, N= Nayarit, Mi= Michoacán, Si=

Sinaloa, SLP= San Luis Potosi, So= Sonora, Z= Zacatecas

Otros: AC= área de cultivo, AD= área de disturbio, BG= bosque de galería, Or= Ornamental, PH= pastizal halófilo, SA= sue-

los alcalinos, ii jonas Tio SY= ee yesosos, VA= vegetación acuática, VR= vegetación riparia, VS= vegetación

subacuática, T= tolerante (subsiste en medios secos y acuosos por igual).

Ub millas: ee Aru=Arundinoideae, Bam=Bambusoideae, Chl=Chloridoideae, Dan=Danthonioideae,

Ehr= =Panicoideae, Poo-Pooideae

SUGERENCIAS

Los muestreos realizados con propósitos taxonómicos o floristicos se llevan a cabo tradicionalmente a lo

largo de las principales vías de comunicación, lo que ocasiona que los estudios de biodiversidad con base

en dichos muestreos, aplicando técnicas cuantitativas, arrojan resultados sesgados que dan pie a factibles

malas interpretaciones. Sin embargo, los resultados del análisis de estos datos, aportan o contribuyen a una

primera aproximación al conocimiento de la distribución y riqueza de especies (gramíneas en este caso), la

cual podría mejorarse significati te si el muestreo se realizara bajo un diseño apropiado para fines de

conocer más ner la distribución de riqueza o diversidad.

AGRADECIMIENTOS

Se reconoce y agradece el financiamiento recibido de la Comisión Nacional para el Uso y Manejo de la Biodi-

versidad (CONABIO-EEO14), del ON 53142 y del Instituto Politécnico Nacional (SIP-20060623 y SIP

20070429) al proyecto “Florística de las gramíneas de Zacatecas” y “Distribución y Diversidad de Gramíneas

de Zacatecas” de los que se baaa resultados para el análisis del presente trabajo. A las autoridades

del CIIDIR IPN Unidad Durango, les agradecemos las facilidades recibidas para realizar las tareas que nos

permitieron llevar a cabo y concluir el presente estudio. A los curadores de los herbarios consultados, se les

agradece sus atenciones. Finalmente agradecemos las sugerencias de 2 revisores anónimos.

I lafal n.a L In Ls des im

790 f Texas 3(2)

Estado de Zacatecas

COBERTURA DEL SUELO

osque

Aatorral Xerófilo

selva

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oblado

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50 0 50 100 Kilometers yy Veg. halófila

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E A Cnhort La sula dal actada da Tarcatarac Cuanta: f'nniunta da datne da Uca dal cuala y Venetación escala

q; f

REFERENCIAS

BALLEZA C., J.J. Y J.L. VILLASEÑOR 2005. Regi lización bi ‘fica de Zacatecas, México, con base en los patrones

de distribución de la familia Asteraceae. Rev. Mex Biodivers 76:71-78.

CHALLENGER, A. 1998. Utilización y Conservación de los Ecosistemas sa de México. Pasado, presente y

futuro. Comisión Nacional para el Conocimiento y Uso de la Biodi | de Biología, Universidad

Nacional Autónoma de México; y Agrupación Sierra Madre, S.C.

CLAYTON, W.D. v S.A. Renvoize. 1986. Genera graminum grasses of the World. Kew Bull. Addit. Ser. XIII. Royal Botanic

Gardens, Kew

Davia, P, MT. Meta-Saués, M. Gomez-SANCHEZ, J. VALDÉS-REYNA, J.J. Ortiz, C. MORÍN, J. CASTREJÓN y A, Ocampo, 2006. Ca-

tálogo de las gramíneas de México. UNAM-CONABIO, México.

DeLong, D.C. Ja. 1996. Defining biodiversity. Wildlife Soc. Bull. 24:738—749.

Gaston, KJ. AND J.l. Spicer. 2007. Biodiversidad introducción. Ed. Acribia, S.A., Zaragoza, España.

HERRERA A., Y, PM. Peterson v A. Cortés Ortiz. 2009, Gramíneas de Zacatecas, México. Sida, Bot. Misc.. In Press.

Humans, RJ., L. Guarino, C. Bussink, P. MatHur, M. Cruz, |. BARRANTES, Y E. Rosas. 2004. Diva-GlS version 5, Sistema de

información Geográfica para el análisis de Datos de Distribución de Especies. Manual. Internacional Plant

Genetic Resources Institute (IPGRI) y UC Berkeley Museum of Vertebrate Zoology. http//www.diva-gis.org/

Materials.htm

% de especies

Matorral

26%

191

nm "

9 8 s B

= = =

v = Te T

iud

Fic. 8. Número de especies según la altitud.

£ al Dos a In LI da den die ET as 0/70

192

m Nativa

mEndémica

Exótica

Fic. 9 P taj | 1 7

HicHcock, A.S. 1951. Manual of the grasses of the United States. Ed. 2, revised by Agnes Chase. U.S. Dept. Agric.

Washington, D.C.

INEGI. 1980a. Carta de climas, escala 1:1 000 000. Instituto Nacional de Estadística, Geografía e Informática,

México.

INEGI. 1980b. Carta fisiográfica, Escala 1: 1 000 000. Instituto Nacional de Estadística, Geografía e Informática,

México.

Inea. 1980c. Carta geológica, Escala 1: 1 000 000. Instituto Nacional de Estadística, Geografía e Informática,

México.

INEGi. 1996. Mapa digital Del Marco Geoestadistico 1995. Instituto Nacional de Estadística, Geografía e Informá-

tica, México.

INEGi. 1998a. Carta topográfica, escala 1: 250 000. f1303, f1305, f1306, f1309, f1404, f1409, 91309, g1312 y g1410.

Instituto Nacional BE ESEAMISHGA, eoa e Informática, México.

INEGI. 1998b. Dicci o de datos de uso del suelo y vegetación (vectorial), escala 1:1 000 000. Sistema Nacional

de Información Geográfica. net Nacional de Estadística, Geografía e Informatica, Méxic

Inga! 2003. Conjunto o de datos de uso del su lo y vegetación serie Il, escala 1: 250 000. r1303, r1305, F1306, F1309,

F1404, £1409, 61309, 61312 y61410, Instituto Nacional de Estadística, Geografía e Informática, México.

INEGI. 2006. Anuario Estadístico de zacate cas. Instituto Nacional de Estadística, Geografía e Informática, México

Lawrence, H.M. 1951. Taxonomy of vascular plants. Macmillan Publishing Co., New York.

Nasa. 2000. http://www2,jpl.nasa.gov/srtm/. (consultado 15 Enero 2009)

Rzepowskl, J. 1978. Vegetación de Mexico. Limusa.

Rzepowski, J. 1991. El Endemismo en la Flora Fanerogámica Mexicana: una apreciación analítica preliminar. Acta

Bot. Mex. 15:47-64.

VALDES Reyna, J. AND |, CABRA Cabral. 1993. Chorology of Mexican grasses. In: Ramamoorthy, T.P, R Bye, A. Lot & J. Fa.

Biological diversity of Mexico. Pp. 439-458

LIPOCARPHA MICRANTHA (CYPERACEAE), NUEVA CITA PARA LA FLORA

DE ARGENTINA: MICROMORFOLOGIA DEL FRUTO

Priscila R. Van de Velde y M. Gabriela Lopez

Instituto de Botdnica del Nordeste

argento Cabral 2131

Corrientes 3400, ARGENTINA

Ff a: - y "eun -

ABSTRACT

4: ] J f +l fs EE f. +5 i] TA q A J t4 Ados

Lipocarpha micrantha aan G.C. Tucker (C

area in South America. It was Suevi ME on Africa, North America (from the South of Canada), Central Ámerica and South

d icd nar ra the Antilles, Min to A ane PEE ue new iin Eo Sarre. ae is based ina

g with a study of

E - Lan] Mido Locus Jal y far the identifies: frthet 1 in Argentina

r oy i [e]

Key Worbs: Lipocarpha micrantha, new record, Argentinian flora, mi phology of the fruit

RESUMEN

Ro a Da Tuc! er (Cyr ) se ci pri la flora de Argentina) li área de distril

— África, América del Norte (desde el Sur de Canadá), América C ls América del Sur

ic oo y ES Antillas hasta Paraguay y aM A La nueva especie de ne ic rra se basa en una n hecha

i 4

] la identifi in deal q 1 Aroentina

E F e

P Ü Ii ] 7 +} : nueva cita, flara de Argentina,

INTRODUCCIÓN

Lipocarpha R. Br. Pan poles a ie SUA E a, tribu Cypereae (Goetghebeur 1998) e

incluye alrededor de 35 esq Van den Borre 1989) y de regiones templado

cálidas. Son plantas aps rara vez — llos condes o curvados, Bare Hojas a

más cortas que los tallos, aplanadas o rara vez algo revolutas, glabras. Infl en cabezuela terminal o

pseudolateral, por la posición erguida de la bráctea involucral inferior, constituida por una o varias espigas

ovoides o globosas; brácteas desiguales foliiformes, la inferior de mayor longitud, expandida o encerrando

parcialmente la base de la inflorescencia. Espigas (20—)50-350+ floras, brácteas glumiformes, densamente

dispuestas en espiral a lo largo de un eje, caedizas; cada una de ellas tectriz de una espiguilla uniflora,

generalmente encerrada por un profilo acne in una iid abaxial o al eee a la madurez. En

algunas especies tanto el profilo como la gluma pueden estar en parte o totalmente reducidos. Perianto ausente;

1-3 estambres; estilo filiforme, 2-3 estigmas, Feitio obovoide o dron de sección subtrigono, eae

aplanado dorsiventralmente o subterete, base sésil o estipitada; ápice con la base del estilo persistente, no

engrosada; superficie finamente papilosa.

De Argentina se conocía solo L. humboldtiana Nees en las provincias de Chaco, Misiones, Entre Ríos y

Corrientes (Barros 1947; Guaglianone 1996). En este tra bajo se comunica la presencia de L. micrantha (Vahl)

G.C. Tucker, encontrada en la provincia de Corrientes.

MATERIALES Y METODOS

Para la preparación de este trabajo se estudiaron ejemplares del Instituto de Botánica del Nordeste (CTES) y

se realizaron observaciones de plantas vivas de L. humboldtiana y de L. micrantha a campo. Se extrajeron 10

J. Bot. Res. inst. Texas 3(2): 793 — 798. 2009

q || Eal n.a H In LI di dian ndh Sry

P94 SUT a f Texas 3(2)

frutos de cada ejemplar de herbario, a un grupo no se le realizó ningún tratamiento, al resto se les extrajo la

pared periclinal externa (Schyuler 1971), por último los frutos se colocaron a centrífuga durante 5 min en 2

o más repeticiones (según necesidad), para realizar la remoción total de las impurezas. 5e montaron en una

platina de cobre y se metalizaron con oro paladio. Para su observación se utilizó el Microscopio Electrónico

de Barrido (SEM), JOEL 5800 LV de La Universidad Nacional del Nordeste-Secretaría General de Ciencia y

Técnica (UNNE-SECyT). La observación de los frutos se realizó a 15 kv y se tomaron fotografías digitales.

La descripción de L. micrantha se basa en los materiales conocidos para Argentina.

RESULTADOS

Lipocarpha micrantha (Vahl) G.C. Tucker, J. Arnold Arbor. 68: 410. oe oo D. i indi micranthus Vahl,

num. Pl. 2:254. 1805. Tiro: L.C. Richard s.n. (moLoriro: P! , Nat. Pflanzenfam

2(2):105. 1888

Planta de 2-10 cm alt.; tallos 0.3-0.5 mm diám., densamente agrupados, rectos o curvados, cilíndricos,

estriados. Hojas Memes de 5-60 mm long. x 0.3-0.5 mm lat. oda pseudolateral; brácteas in-

volucrales 1-2(3), la inferior siguiendo la dirección del tallo, | ás cortas. Espigas compuestas

liad ovoides o globosas, constituidas por bracteas glumiformes Hs 0.7-1 x 0.3-0.5 mm, densamente

li espiral, amarillas, rojizas en los bordes, 2 trales desde la base hasta cerca

dii ápice, o sólo hasta a la mitad de su longitud, verdes; cada una tectriz de una espiguilla reducida a 1 flor.

Raquis persistente, patente, cilíndrico, con cicatrices romboidales. Flores bisexuales. Estambres 1 (raro 2),

anteras 0,2-0,4 mm, elipsoides, apiculadas, filamentos capilares. Estilo 0,2 mm long., filiforme, estigmas

2. Aquenio 0.5-0.7 mm long. x 0.2-0.3 mm lat., obovoide, sección subterete, cortamente apiculado, sésil,

conspicuamente papiloso, amarillento o iridiscente, marrón claro cuando maduro.

Fenologia.—Florece y fructifica todo el año (Kral 2001)

Ecología.—Se la encuentra en campos bajos temporalmente inundados, terrenos alterados, abiertos,

bordes de ríos, suelos arenosos, pedregosos o bua: 0— Pide m.s.n.m.

Distribución geográfica (Fig. 2).—Lig a especie del género que habita en ambos

hemisferios, en los continentes de África y America (Goetghebeur & Van den porie 1989), en regiones tro-

picales y templadas. En América se distribuye desde el sur de Canadá en América del Norte hasta América

del Sur llegando hasta Paraguay, Noreste de Argentina y Noreste de Uruguay. Su límite de distribución es

33° lat. Sur.

M diado: ARGENTINA. CORRIENTES. Dpto. It R io Cué, 15 km S de Ituzaingó, 23 May 1982, R. Carnevalli 5374

(CTES). Dpto. Empedrado: Camping de Empedrado, 14 Mar 2006, M.G. a et al. 385 (CTES). Dpto. Capital: Laguna Soto, Playa

Bounganville, 10 Dic 2006, M.G.López & R. O. Vanni 386 (CTES)

CLAVE PARA SEPARAR LAS ESPECIES DE ARGENTINA

1. Hierbas con talios de 2-10 cm long. Hojas con : filif: de 0.3-0.5 mm lat. Inflorescencia seudola-

teral con 1-2(-3) espigas; espiguillas Si sin profilo; estigmas 2. Aquenio de 0,5-0,7 mm long. x

at. L micrantha

1. Hierbas con tallos de 30- £ cm long. Hojas con lámi filif redondeadas acanaladas en el e

de 1 mm iat. | i 3-5(-8) espigas; espiguill ! ; estig 3. Aq eses 1,3

mm long. x 0,4-0,5 mm lat. L. humboldtiana

MICROMORFOLOGÍA DEL FRUTO

La ornamentación de los frutos tiene valor taxonómico en muchos de los géneros de Ciperáceas: Schuyler

(1971) realizó el estudio de la superficie de los aquenios con MEB en los géneros Scirpus L. y Eriophorum L.;

más tarde lo siguieron otros autores: Walter (1975), Toivonen & Timonen (1976), Standley (1987 y 1990)

en el género Carex L.; Guaglianone (1979 y 1981) y Ragonese et al. (1984), en Rhynchospora Vahl, López &

Matthei (1995) y Araújo & p oM e SUE en ie i L.; Goetghebeur & Coudijzer (1984) han

publicado fotos de la superficie de frutos de las especies e Bulbostylis de África Central; López (2006, 2007)

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a | ou " flay J on | || L Fa H] mn B E a | J L "Am m i || H £1 . Li

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cara adaxial. I. cara abaxial. J. Inflorescencia

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K. Espiga. (A, C—F, H-K de López et al. 385. B, G de López et al. 386).

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Fie 2. Distrihució sf j A ag

y Lopez et al. (2007) en algunas especies de Bulbostylis de Sudamérica. Lye (2000) estudio detalladamente

con MEB los frutos de alrededor de 600 especies de Cyperáceas de todos los continentes Goetghebeur &

Van den Borre (1989) realizaron el estudio de 16 especies de las 35 del género haciendo referencia a la falta

de datos suficientes sobre la variabilidad intraespecifica. Entre las especies estudiadas no incluyeron a L.

micrantha.

En L. micrantha se distinguen en vista superficial células en su mayoria hexagonales y algunas pen-

tagonales con protuberancias centrales, redondeadas (Fig. 3; D, E); Al remover la pared periclinal externa

del exocarpo, se observa que la pared anticlinal es muy levemente ondulada y delgada, la plataforma basal

es cóncava y lisa. En frutos no maduros el cuerpo central en el ápice es peltado, liso, con una leve concavi-

dad y a en su extremo distal ve 3 E E En irutós IBaduios el cuerpo central es cónico, con

co alas de Cyperus reflexus Vahl

Var. pm López & Matthei 1995), (Fig. 3; a Lā cara aan de la plataforma de los cuerpos de silice, es

ligeramente excavada y cubierta por pequeñas 1 siliceas en toda la superficie (Fig. 3; H, D.

En los frutos de L. humboldtiana en vista meram no se puede definir la forma de las células ya que

el limite es oscuro y se observan protuberancias truncadas (Fig. 3; A, B). Sin la pared periclinal externa

las células son hexagonales, la pared anticlinal es recta y delgada, la plataforma basal es convexa y lisa, el

cuerpo central cilíndrico, truncado, sin ondulaciones y plano (Fig. 3, C).

—

AGRADECIMIENTOS

Y^ 1: ] E 1 i Jad -ala

t d lecimiento a la Prof. R

w Laura Simón por n an de la lámina, Srita. Valeria Manibiti pot su on en la -—"

de los frutos y Lic. Cristina Salgado por su atención en el MEB

797

Fic. 3. Microfot fi MEB d ha R.: L. humboldtiana (A-0): A. Fruto. B. Vist ficial d linal C

Detall | ed linal : : le silicio: £ micrantha (D-J): D Fruto. E, Vist imerfigal del | periclinal ext F

Detalle de la ] linal y Je silicio G-J Cuerpos de silicio. G. Cuerpo cenia H. Vista inferior de plataf basal

|. Vista inferior de la plataforma basal J. Detalle de cuerpo central cóni ificaciones apicales. (A- C de Ahumada 2139. F-H de

López et al 385. |, ) de R. Carnevalli 5374).

798 J t tanical h Institute of Texas 3(2)

REFERENCIAS

Araújo, A.C. AND H.M. LONGHI-WAGNER. 1997. Anatomia foliar e micromofologia do fruto na taxonomia de Cyperus

L. (Cyperaceae). Iheringia, Bot. 48:103-120.

Barros, M. 1947. Cyperaceae - Scirpoideae. En: H. Descole, ed. Gen. et Sp. Pl. Argent. 4(1):9-13.

GOETGHEBEUR, P. AND J. COUDIUZER 1984. Studies in Cyperaceae 3. Fimbristylis and Abildgaardia in Central Africa. Bull.

Jard. Bot. Belg. 54:65-89,

GOETGHEBEUR, P. AND A. VAN DEN Borre. 1989. Studies in Cyperaceae. 8. A revision of Lipocarpha, including Hemicarpha

and Rikliella. Wageninger Agric. Univ. Pap. 89:1-87.

GOETGHEBEUR, P. 1998. Cyperaceae. En: K. Kubitzki y collaborators, eds. The families and genera of vascular plants

4:141-190.

GUAGLIANONE, E.R. 1979. Sobre Rhynchosj gosa (Vahl) Gale (Cyperaceae) y alg pecies afines. Darwiniana

22:255-311

GUAGLIANONE, E.R. 1981. Contribución al estudio del género Rhynchospora Vahl (Cyperaceae) Ill. Darwiniana

23:489-506

GUAGLIANONE, E.R. 1996. Cyperaceae. En: Zuloaga F. and O. Morrone, eds. Catálogo de las plantas vasculares de la

República Argentina |. Monogr. Syst. Bot., Missouri Bot. Gard, 60:175.

KraL, R. 2001. Lipocarpha (Cyperaceae). En: Stevens, W. D, C. Ulloa Ulloa, A. Pool, O. M. Montiel, A. L. Arbeláez, and

D. M. Cutaia, eds. eds. Flora de Nicaragua (Tomo I) Monogr. Syst. Bot., Missouri Bot. Gard. 85:773-774.

Lopez, M.G. 2006. Nuevas citas de Bulbostylis (Cyperaceae) para las floras de Bolivia y Paraguay. Bonplandia

15(3-4):149-160.

Lopez, M.G. 2007. Tres Nuevas Especies en el Género Bulbostylis (Cyperaceae) de Sudamérica. Novon 17:

497—502

LOPEZ, M.G., A.P. PRATA, AND W.W. Thomas. 2007. New synonymy and new distributional Is in Bulbostylis (Cyper-

aceae) from South America. Brittonia 59:88—96.

Lopez, PS. AND OJ, Mamme. 1995. Micromorfología del Aquenio en especies del género Cyperus L. (Cyperaceae),

Chile. Gayana Bot. 52:67-75.

Lye, K. 2000. Achene structure and function of structure in Cyperaceae. En: Wilson K. L. and D.A. Morrison, eds.

Monocots: systematics and evolution. CSIRO, Melbourne. Pp. 615-628

Raconese A.M., E.R. GUAGLIANONE, AND C. DIZEO DE STRITTMATTER. 1984. Desarrollo del pericarpio con cuerpos de sílice

de dos especies de Rhynchospora Vahl (- y pEracede. panei 25:27-41.

SCHUYLER, A. 1971. Scanning electron microscopy of achene epidermis in species of Scirpeae and related genera.

Proc. Acad. Nat. Sci. Philadelphia. 123:29—52.

STANDLEY, L. 1987. Anatomical studies of Carex cuchumatensis, C. decidua, and C. hermanii (Cyperaceae) and

comparisons with North American taxa of the C acuta complex. Brittonia 39:11-19.

STANDLEY, L.1990. Anatomical aspects of the taxonomy of sedges (Carex Cyperaceae). Canad. J. Bot. 68:

1449-1456.

Toivonen, H. AND T. Timonen. 1976. Perigynium and achene epidermis in some species of Carex, subg. Vignea

(Cyperaceae) studied by scaning electron microscopy. Ann. Bot. Fenn. 13:49-59.

Water, K.S. 1975. A preliminary study of the achene epidermis of certain Carex (Cyperaceae), using scaning

electron microscopy. Michigan Bot. 14:67—72.

CHECKLIST OF THE PLANTS OF THE GUIANA SHIELD

1. AN UPDATE TO THE ANGIOSPERMS

Christian Feuillet

Department of Botany, MRC-166

Smithsonian Institution

PO. Box 37012

Washington, D.C. 20013-7012, U.S.A.

feuillec@si.edu

ABSTRACT

1 Ce A A ihe Coa Shield i l. 1t i tl 700 additions, corrections d

+ J L i 1 e g P Ed

for ile Angiosperms. Two nev binati le in the Rubi Appunia longiped lat d Appunia ped l

RESUME

Une mise à jour de la Checklist of the Plants of the Guiana Shield est offerte. Ell d plus de 700 additions, corrections et synony-

mies pour les Angiospermes. D binai 11 t faites dans les Rublaceas: Appunia longipedunculata et Appunia

peduncularis.

The flora of the Guiana Shield is the object of strong interest that fostered various international programs

as evidenced and enhanced by the publication of the Flora of the Guianas (Gorts-van Rijn & Janssen-Jacobs

1985—present; in progress), the Flora of the Venezuelan Guayana (Berry et al. 1995-2005), and the Guide to

the vascular plants of central French Guiana (Mori et al. 1997 and 2002), as well as by the large specimen-

based database at the Department of Botany of the Smithsonian Institution. There was a need to compare

results and therefore for conformity of the taxonomy. The Checklist of the Plants of the Guiana Shield (Funk et

al. 2007) was the tool to fill that need. The phase of data gathering for the Checklist was completed in 2005.

Since then, many papers have been published on the flora of the region adding several species, changing

their generic placement, or refuting their presence in the Guiana Shield. Furthermore, due to the widely

scattered literature, some taxa were overlooked. Hence many additions, deletions, and corrections must be

made before an updated version of the Checklist can be offered for consultation on-line. This paper presents

719 modifications needed for the Angiosperms and gathers in one place the references for the new species.

The results are presented in the Appendix. For the Angiosperms, 211 names are added to the checklist and

159 are placed in synonymy or rejected. Therefore, there is a gain of 52 and the total number of accepted

subgeneric names is now 14,784.

A later pos will UD ane Hs and allied groups. The Checklist (Funk et al. 2007) can be down-

loaded from http:;//l ts.html along with this update and relevant pdf files.

If you know of taxa that should have been added, edited, or deleted, please let me know.

APPENDIX: MODIFICATIONS

The famili ted in alphabetical order under the family | in the Checklist (Funk et al. 2007). The names of

¿l c A E. i | el pag l Ll ) I £ ] xil Checklict c " TES | E NE "NE

added are in bold, followed | by the symbols of t! ions (AM = Amazonas BO = Bolivar; DA = Delta Amacuro; FG = French

Guiana; GU = Guyana; 2 = Surinam). Names of species ir in 1 italics are to obe edited Or deleted. As in the checklist, the family

patas ificat “and “References.” Occasionally a

chapters GH POV OU Y LM Playa 113 J

note clarifies a problem.

Reference (p. 15-16)

Add: Cuarx, H., R. Liesner, PE. Berry, A. FERNÁNDEZ, G. AYMARD, AND P. MAQUIRINO, 2000. Catálogo anotado de la flora del

área de San Carlos de Río Negro, Venezuela. Sci. Guaianae 11:101-333 + 21 color photos.

| Rat Rac Inct Tavac 23/91 740 — 814. 2009

800

ANTHOPHYTA: MONOCOTYLEDONAE

ARACEAE (p. 59-67)

Monstera barrieri Croat, Moonen & Poncy—FG

Philodendron scottmorianum Croat & Moonen—-FG

References

Croat, T.B. AND J. Moonen. 2007. Philodendron scottmo-

rianum, a new species of P sect. Philodendron (Ar-

aceae) from the Guianas. Willdenowia 37:535—539.

Croat, T.B., J. Moonen, AND O. Poncy. 2005. New species of

Monstera (Araceae) from French Guiana. Rodriguésia

56(88):61—64.

ARECACEAE (p. 67-73)

Bactris nancibaensis J.J. de Granville—FG

Reference

GRANVILLE, J.-J. DE. 2007. A new species of Bactris (Palmae)

from French Guiana. Brittonia 59:354-356

BROMELIACEAE (p. 73-83)

Pitcairnia cremers Couga SU. ra

lata

Pitcairnia Jeprieuri Bal ker=shouig be deleted

carract aithor acd

eet PRL CILALI TMT CANA "ud t

Dit 1 a (B | ) I] B S

spelling of species pina

Pitcairnia saxosa Gou da —FG

Pitcairnia semijuncta Baker—GU, SU, FG

Synonyms and misidentifications

Hepetis amazonica (Baker) Mez—should read "—Pitcairnia

ubiginosa var. amazonica (Baker) L.B. S

io incarnata (Baker) G.S. Varad. & Gilman — Pitcairnia

icifolia Mart. ex Schult. f.

Biene amazonica ES reag '=Pitcairnia rubigi-

r. amazonic B. Sm

Pitcairnia incarnata dd Es

Schult. f.

Pitcairnia leprieurii Baker — Pitcairnia rubiginosa Baker

picem caricifolia Mart. ex

Reference

GOUDA, E.J. 2009. Studi the flora ofthe Guianas: the

genus Pitcairnia (B li ) kelba: 30:80-88.

COSTACEAE (p. 86-87)

Chamaecostus congestiflorus (Rich. ex L.F. Gagnep.) C.

Specht & D.W. Stev U. SU, FG

T Li ,

Chamaecostus curcumoides (Maas) C. Specht & D.W.

Stev.—FG

Chamaecostus lanceolatus (Petersen) C. Specht & D.W.

Stev.—SU, FG

Chamaecostus lanceolatus subsp. pulchriflorus (Ducke)

| & DW. Stev.—

Costus congestiflorus Rich. ex L.F. Gagnep.—should be

deleted

," d AA h lA kh

Fred Lea eee (lg) A ere

C Pe Ne Na LI wd PRADA Pul No Nel nun et

be deleted

Synonyms and misidentifications

Costus congestiflorus Rich. ex L.F. Gagnep. = Chamaecostus

congestiflorus (Rich. ex L.F. Gagnep.) C. Specht & DW.

Stev.

Costus curcumoides om = Chamaecostus curcumoides (Maas)

echt & DW

Eostiis Wis o = Chamaecostus lanceolatus

(Petersen) C. Specht & DW, Stev.

is NEENA subsp. PAORS (LUCRE) middi =

I ^

YAA A Pi wea

sii & DW. Stev.

Reference

SPECHT, C. AND D.W. STEVENSON. 2006. A new phylogeny-

based generic classification of Costaceae (Zingiber-

ales). Taxon 55:153-163.

CYPERACEAE (p. 88-103)

Bulbostylis medusae Prata, Reynders & Goetgh.—edit author

and delete “sp. nov. ined’

Reference

PRATA, A.P, K. CAMELBEKE, M. REYNDERS, I.C. FeDON, P. Go-

ETGHEBEUR, AND O. Huser. 2007. Bulbostylis medusae

(Cyperaceae), a new species from Venezuela. Novon

17:67-71

ORCHIDACEAE " 118-149)

d armata Rchb. f —add S

Habenar | i que comment in the

square Sn ds BO

Habenaria hamata Barb. R

Note: The only collection of " seticauda Lindl. ex Benth.

in CAY, Cremers & Hoff 10717, has been identified as H.

hamata by Batista et al. 2008.

p" d

I J l | AA bem

Habenaria pratensis (Salzm. ex Lindl) Rchb. f —should be

deleted

Habenaria pratensis var. parviflora Cogn.—should be de-

leted

Habenaria schomburgkii Lindl.—BO, GU, SU

Habenaria schwackei Barb. Rodr.—GU, SU, FG

Habenaria secundiflora Barb. Rodr.—BO, GU, FG

dote seticauda Lindl. ex Benth.—delete FG; it should

bla spathulifera Cogn.—BO, GU?, SU, FG

Habenaria subfiliformis Cogn.—BO, SU

Heterotaxis proboscidea (Rchb. f.) F. Barros—should be

deleted

Ligeophila clavigera var. dd ri Ormerod—GU

Ligeophila unicornis Orm —

Maxillaria nasuta Rchb. eode be deleted

Feuillet, U m E! | m En £ H r Al F..* CL 5.1.

Microchilus campanulatus Ormerod—AM, BO, GU

Microchilus guianensis Ormerod—

Nitidobulbon nasutum (Rchb. f) Ojeda & Carnevali AM,

BO, GU

Nitidobulbon proboscidea (Rchb. f) Ojeda € Carnevali—

Ornithidium elianae Carnevali & M.A. Blanco—AM, GU,

U, FG

Specklinia feuilletii Luer—FG

Note: in the Checklist, the species of Specklinia Lindl. were

placed la the cy of POROU R. d one in

4 (p. 237)

as well as Specklinia feuilletii (Luer) Luer 2004 v n are

mala penso they MEE dd published in the same

The only name

available is Specklinia feuilletii Luer validated in 2005

Synonyms and misidentifications

m platydactyla Kraenzl. = Habenaria schwackei Barb.

mun proboscidea (Rchb. f) F. Barros = = Nitidobulbon

pr OO G (Rchb, 1 Ojeda & Carneva

M & C. Schw S read

= Nitidobulbon nasutum (Rchb. f.) dues & Carnevali”

Maxillaria nasuta Rchb. f. = Nitidobuloon nasutum (Rchb. f)

Carnevali

Maxillaria proboscidea Rchb. f. = Nitidobulbon proboscidea

(Rchb. f.) Ojeda €: Carnevali

Pleurothallis feuilletii Luer (invalid)

= Specklinia feuilletii Luer

References

BATISTA, J.A.N., J.B.F. DA SiLVA, AND L. DE Bem BiaNcHETTI. 2008.

The genus Habenaria (Orchidaceae) in the Brazilian

Amazon. Rev. Brasil. Bot. 31:105-134

BLANCO, M.A., G. CARNEVALI, D. BOGARÍN, AND R.B. SINGER,

2008. Further AO ofa tonom puzzle:

RÁ "tj

and anew

species, O. elena (Orchidaceae) Harvard Pap. Bot.

13:135-154.

Luer, C.A. 2004. Pleurothallis subgen. Acianthera and

three allied subgenera; A second century of new

species of Stelis of Ecuador; Epibator, Ophidion,

Zootrophion. Monogr. Syst. Bot. Missouri Bot. Gard.

95:1-265

Luer, C.A. 2005. Validation of names cited in Monogr.

Syst. Bot. Missouri Bot. Gard. 95. 2004 Monogr. Syst

Bot. Missouri Bot. Gard. 103:310-31 1.

OJEDA l, G. CARNEVALI, AND G.A. RoMERO-GONZALEz. 2009.

Nitidobulbon, a new genus of Maxillariinae (Orchi-

daceae). Novon 19:96-101.

Ormeroo, P. 2008. Studies of neotropical Goodyerinae

(Orchidaceae) 3. Harvard Pap. Bot. 13:55-87.

POACEAE (p. 149-172)

A Ld L iT, nz | g A mee eee AM, BO, GU

Anachias alinulara (NE Br.) Fiilnana 8, Marrana "Pa!

F = M M

A mmelslm [Daviden "Ti ile ater b e NAarranga BO

Apochloa sipapoensis (Swallen) Zuloaga & Morrone—

Apochloa steyermarkii (Swallen) Zuloaga & Morrone—

+l re H H | E T e E N I eus AM

Panicum inode Trin. ould be deleted

Panicum eligulatum N.E. Br—should be deleted

n y s Ty FEN -— Led lee ny PRA |

I Jl LEO V ee he Ju La

D H H C li edil x A alata

n s PAN a I] P E ess OR oe S A |

Raddiella vanessiae Judziewicz & Sepsenwol—FG

Synonyms and misidentifications

Panicum auyanense Swallen—should read “= Apochloa eligu-

lata (N.E. Br) Zuloaga €: Morrone”

Panicum chnoodes Trin. = Apochloa chnoodes (Trin.) Zuloaga

one

Panicum churunense seis Fiala read “= Apochloa

chnoodes (Trin.) Zuloaga & Morro

Panicum cowanii Swalle d one E “= Apochloa chnoodes

(Trin.) Zuloaga & Morron

Panicum curvifolium T ahedd read “= Apochloa

chnoodes (Trin.) Zuloaga €: Morrone”

Panicum eligulatum N.E. Br. = Apochloa eligulata (N.E. Br.)

a & Morrone

Panicum fontanale Swallen—should read*= Apochloa steyer-

markii (Swallen) Zuloaga & Morrone”

Panicum inversum Swallen—should read “= Apochloa

chnoodes (Trin.) Zuloaga & Morrone”

Panicum jauanum Davidse = Apochloa jauana (Davidse)

Zuloaga & Morrone

Panicum kavanayense MN Um: read “= Apochloa

chnoodes (Trin.) Zuloaga & Mor

Panicum maguirei yale ull read “= Apochloa

chnoodes (Trin.) Zuloaga & uiid ne"

Zuloaga & Morrone

n A Ll " Lo fe I|

len) Zuloaga & Morrone

Panicum tatei Swallen—should read “= Apochloa chnoodes

(Trin.) Zuloaga & Morrone”

Panicum tropinoblephare Tutin—should read “= Apochloa

chnoodes (Trin.) Zuloaga & Morrone”

Panicum vannum dos Id read"— Apochloa chnoodes

(Trin) Zuloaga & Mor

Panicum wurdackii Suallen should ie “= Apochloa eligu-

lata (N.E. Br) Zuloaga 8: Morrone

References

Juoziewicz, E.J. AND G. DaviDsE. 2008. Arthrostylidium berryi

(Poaceae, Bambusoideae, Bambuseae, Arthrostylidii-

nae), anew species from white sand shrublands in

Venezuela and Colombia. Novon 18:361-365.

JUDZIEWICZ, E.J. AND S. SEPSENWOL. 2007. The world's small-

D 5 n

est hamhan:

a Fm

vanesside \rOacede: Bambu-

802

soideae: Olyreae), a new species from French Guiana.

Jj. Bot. Res. Inst. Texas 1:1-7.

Sepe, S.M., O. Morrone, L.M. Giussani, AND F.O. ZULOAGA.

2008. Phylogenetic studies in the Paniceae (Poace-

ae): a realignment of section Lorea of Panicum. Syst.

Bot. 33:284-300.

ANTHOPHYTA: DICOTYLEDONAE

ACANTHACEAE (p. 183-187)

Asystasia gangetica (L) T. Anderson—add F

ote: Common weed around villages. i specimens, cf.

C. Feuillet 26 (CAY, P, US)

Synonyms and misidentifications

Ruellia fulgens (Bremek.) E. Tripp = Polylychnis fulgens

Note: The merge of Polylychnis in Ruellia might be the right

thing to do, but the other species of Polylychnis in the

Checklist ales has r Mel been transferred. It seems better

tl Ho regio! 1al VW IX.

1 tog in

Reference

Tripp, E.A. 2007. Evolutionary relashionships within the

species-rich genus Ruellia (Acanthaceae). Syst. Bot.

32:628-649

ANNONACEAE (p. 190-195)

Guatteria alticola Scharf & Maas—GU

Guatteria anteridifera Scharf & Maas—FG

Guatteria anthracina Scharf € Maas—GU, SU, FG

Guatteria ayangannae Scharf & Maas—GU

Guatteria elegans Scharf —FG

Guatteria intermedia Scharf —SU, FG

Guatteria RUcOUICNA ean à Maas—FG

/aas—QGU, SU

Guatteria montis-trinitatis Scharf—FG

Soin pannosa Scharf & Maas—FG

uatteria partangensis Scharf & Maas—GU

Guat richardii R.E. Fr—add $ 3H

fO NAanac

Oa DA el 8 RAN LA

"- Wa

GENTES EIA VALITA VICIS e MEM

Eolsib mel

Synonyms and misidentifications

References

Maas, PJ.M., L.Y.T. Westra, AND M. Vermeer. 2007. Revision

of the neotropical genera Bocageopsis, Onycho-

petalum, and Unonopsis (Annonaceae). Blumea

52:413—554.

ScHaRE, U., PJ.M. Maas, AND W. Monawtrz. 2006. Guat-

teria richardii (Annonaceae) rediscovered along

with two new species from French Guiana. Blumea

51:541-552.

1 fal n. b In LI bibih £Taw

ac 2/0

Ad» HL]

SCHARF, U., PJ.M. Maas, ed Morawetz. 2005. Five new

species Ol GUCGLLGCTIG the Pakara-

ima Mountains, Guyane ote 50: 563- 573.

ScHaRF, U., PJ.M. Maas, ANDAN. NOMEA 2006. Five new

~Cranch (71

POUT Gui

species of Guatteria (, laceae)

ana, Guyana and came Blumea 51:117-130.

Scharr, U., PJ.M. Maas, AND M.-F. Prévost. 2008. An un-

usual new species of Guatteria (Annonaceae) from

French Guiana and adjacent Brazil (Amapá). Blumea

53:515—523.

ARISTOLOCHIACEAE (p. 207-208)

A TI | l | £4| : DA Aris

larkhir e Laltnid LI | d Art] I i n

t Li HU ELE LA

FGUINGL Upe the alignment E the checklist text and

] J -Tl Erll

should read as follows.

iie a Feuillet—delete“ined,, correct author

elling—

eee a Duch.—GU, FG

Aristolochia melgueiroi Barringer & Guánchez—AM, BO

Aristolochia mossii S. Moore—AM, BO, FG

Aristolochia nummularifolia Kunth—AM, BO

Aristolochia pannosoides Hoehne—BO

Aristolochia paramaribensis Duch.—GU, SU, FG

Reference

FeuiLLET, C. 2007. Folia taxonomica 1. Validation of two

taxa from northern South Arnerica. J. Bot. Res. Inst.

Texas 1:143-144.

BORAGINACEAE (p. 224-226)

Cordia bullata (L) Roem. & Schult.—should be deleted

Cordia bullata subsp. humilis (Jacq) Gaviria—should be

delete

ii Feuillet—should be deleted

Cordia curassavica (Jacq.) Roem. & Schult.—should be

deleted

—

mins multispicata Cham.—should be deleted

Cordia polycephala (Lam) I.M. Johnst.—should be deleted

€ " l " 1 Fa +i l LA Tha Halata A

Cordia roraimae |. M. Johnst.—should be deleted

Sai schomburgkii DC. cop Pe deleted:

FATE mp pun

Id kh Halatan

Cordi kii G Gaviria

Cordia tomentosa cn Roem: 2 Schult.—should be de-

leted

Euploca filiformis (Lehm.) J..M. Melo & Semir—AM, BO,

GU, SU, FG

i Iger—BO, SU

Euploca polyphylla (Lehm JJ. LM. Melo 8 Semir—BO

ploca procumbens (Mill) Diane € Hilger—AM, BO,

E^ GU

* NAS ATV e Ll:

Feuillet, Uj lated checklist of gi I for the Gui Shield

Euploca ternata (Vahl) J..M. Melo & Semir—BO, GU

(ay poe IIA i CRL I | lw adem Le Lm pee |

ie ees : E i SLA X foe ol A A eel coal tie ced

Ip 5 alus = hH * rM SET Pdl IA Ll mE

Lint + de $ Me cl bee | IMA

q E | [| TRA" ^ [7 A

LI Li wk

, GU

Myrlopus maculatus Jaca) Feulllet—BO, GU, SU, FG

Myniopus paniculatus (Cham; pee au: SU, FG

A. DC.) Feuillet—

LI -—

Myriopus volubilis (L.) Smali—BO, GU, F

Tournefortia candidula (Miers) I.M. can —should be

deleted

Tournefortia maculata x —should be deleted

Tournefortia paniculat m.—should be deleted

Tournefortia ipn var. elo (A. DC) LM. Johnst.—

shoul

Tournefortia vo! us L. mieu i deleted

Varronia bullata L.—BO, DA, G

Varronia bullata subsp. unii - Feuillet—BO, DA,

GU?

Varronia cremersii (Feuillet) Feuillet—FG

Varronia curassavica Jacq.—AM, BO, DA, GU, SU, FG

A, GU

Varronia grandifiora | Desv.—AM, BO, D

"Jj EV C "Jl " GU I

Varronis polycephala Lam. —BO, Gu, SU, FG

hii AM

Varronia ı roraimae ( M. p i & Mil. —BO

Varronia schomburgkii (DC) Borhidi—BO, GU, SU, FG

Varronia stenostachya (Gaviria) J.S. Mill. —AM, GU

Varronia steyermarkii (Gaviria) J.S. Mill.—BO

Varronia tomentosa Lam.—SU, FG

Synonyms and misidentifications

Carla eu tL) Roem. & & Schi ilt. = Marena HUdig -

C j a

H ;

subsp. humilis (Jacq) Feuillet -

Cordia cremersii Feuillet = Varronia cremersii (Feuillet)

Feuillet

Cordia curassavica (Jacq.) Roem. & Schult. 2 Varronia curas-

savica Jac a

'H l j (Dri 9, Da; Y Baam

& Schult. [misiden-

tin cation for Varroni a schomburgki (DC) Borhidi]

Varronia schomburgki oc ) —

Cordia | marioniae Feuillet = = ‘Vanonia marioniae (Feuillet)

Feuillet

Cordia multispicata Cham. [not in our area]

m eiue (Lam.) ILM. Johnst. = Varronia polycephala

s Pasada Kunth = Varronia polystachya (Kunth)

Borhidi

Cordia roraimae (LM. Johnst.) J.S. Mill. = Varronia roraimae (I.M.

ohns ill

3. iVII

Cordia schomburgkii DC. = Varronia schomburgkii (DC.)

Borhidi

803

Cordia spinescens (L.) Borhidi [misidentification for Varronia

schomburgkii (DC) Borhidi]

Cordia stenostachya Killip ex Gaviria = Varronia stenostachya

(Gaviria) J.S. Mill.

Cordia steyermarkii Gaviria = Varronia steyermarkii (Gaviria)

Mill.

Cordia tomentosa (Lam.) Roem. & Schult. = Varronia tomen-

tosa Lam.

Heliotropium filiforme Lehm. = Euploca filiformis (Lehm) J.J.M.

Melo & Semir

Heliotropium foliosum Roem. & a anoula read “= Eu-

oca POPMA ) J.M o & Sem

| S dr “= Euploca

ternata (Vahl) J.l. M. Melo & Semir"

e lagoense (Warm.) Gürke = Euploca lagoensis

( Diane & Hilger

TT dum ottonis Lehm.—should read “= Euploca ternata

(Vahl.) J.1.M. Melo & Semir“

Heliotropium polyphyllum Lehm. = Euploca ternata (Vahl.)

J..M. Melo & Semir

Heliotropium procumbens Mill. = Euploca procumbens (Mill)

Diane & Hi

Heliotropium ternatum Vahl =

Melo & Semir

Heliotropium trinitensis Urb. SEUA read “= Euploca lagoensis

(War mt) Di ane & Hilger

—

= Euploca ternata (Vahl) J.1.M.

IJ: 34. .]

-4

Feuillet

Schleidenia fumana Fresen "iu read “= Euploca ternata

Vahl) J.M. Melo & Sem

Schleidenia lagoensis Warm.—s inedia read “= Euploca lagoensis

war m.) Dia ane & lige

Ca polyphylla (Lehm ) M. M. Melo & Semir'

Tournefortia candidula (Miers) I.M. Johnst. = Myriopus candi-

ulus (Miers) Feuillet

Tournefortia maculata Jacq. = Myriopus maculatus (Jacq)

Feuillet

ia inr paniculata Cham. = Myriopus paniculatus (Cham.)

euillet

rounetort paniculata var. spigeliiflora (A. DC.) I.M. Johnst. =

yriopus paniculatus var. spigeliiflorus (A. DC.) Feuillet

Tournefortia spigeliaeflora A. DC. = Myriopus paniculatus var.

spigeliiflorus (A. DC.) Feuillet

Tournefortia volubilis L. = Myriopus volubilis (L.) Small

References

FeuiLLET, C. 2008. Folia taxonomica 4. Conspectus of

Myriopus (Heliotropiaceae, Boraginales) in the Gui-

ana Shield. J. Bot. Res. Inst. Texas 2:263-265.

FeuiLLET, C. 2008. Folia taxonomica 11. Conspectus of

Varronia (Cordiaceae, Boraginales) in the Guiana

Shield with three new combinations. J. Bot. Res.

Inst. Texas 2:83/-842.

MeLo, J.I.M. DE. 2009. Two new Brazilian species and

new combinations in Euploca (Heliotropiuaceae).

Kew Bull. 64:285-289,

804

BURSERACEAE (p. 227-231)

Protium calendulinum pia FG

Protium retusum Daly—

Protium urophyllidium ae

Reference

Daty, D.C. 2007. A new section of Protium from the

Neotropics. Studies in neotropical Burseraceae XIII.

Brittonia 59:1-24.

CAPPARACEAE (p. 234-235)

Capparis flexuosa subsp. polyantha (Triana & Planch.) H.H

IItis—should be deleted

rr 1

D BS I LA |

EAM muco HH. Iltis, L. Cumaná, R. Delgado €: Aymard—

should be deleted

nas polyantha (Triana & Planch.) X. Cornejo € H.H.

Iltis O, SU

| "1 [| * es y +

+) LJ LJ Iltis—GU, SU, FG

ist) X. Cornejo & H.H.

i $ l

Ai | | 4 || Li r] in

tis —BO, GU, SU,

Neocalyptrocalyx morii X. Cornejo & H.H. IItis—FG

(H.H. Iltis, L. Cumaná, R. Delgado &

Aymara) X. Cornejo & H. H. tis BO

Synonyms and misidentifications

Capparis flexuosa subsp. antha (Triana & Planch.) H.H

Iltis = fecal polyantha (Triana & Planch.) X. Cornejo

& H Pe tis

“nr de A! i " j J ; su y s +)

AH H. tis

(Benoist) X. Cornejo & H.H. Iltis

Capparis muco H.H. Iltis, L. Cumaná, R. Delgado & Aymard =

Neocalyptrocalyx muco (H.H. Iltis, L. Cumaná, R. Delgado

& Aymard) X. Cornejo & H.H. Iltis

Capparis polyantha Triana & Planch. = Cynophalla polyantha

(Triana & Planch.) X. Cornejo & H.H. Iltis

References

Cornejo, X. AND H.H. ILris. 2008. Two new genera of Cap-

paraceae: S j | M pparis stat. nov.,

andtl t of Neocalyptrocalyx. Harvard

Pap. Bot. 13:103-116.

Cornejo, X. AND H.H. Iris. 2008. New combinations in

South American Capparaceae. Harvard Pap. Bot.

13:117-120.

Cornejo, X. AND H.H. Ins. 2008. Neocalyptrocalyx morii

(Capparaceae), a new species from Central French

Guiana. J. Bot. Res. Inst. Texas 2:807-810.

CLUSIACEAE (p. 247-253)

Tovomita gazelii Poncy & Offroy—FG

Reference

Poncy, O. AND B. Orrroy. 2006. Une nouvelle espèce de

Tovomita Aubl. (Clusiaceae) de Guyane francaise.

Adansonia, sér. 3, 28(1):113-117.

COMBRETACEAE (p. 254-255)

The references are misplaced on the bottom of page 253

after those of the Clusiaceae.

COMPOS

Galinsoga parviflora Cab.—GU

p. 255-269)

DROSERACEAE (p. 282)

Drosera solaris A. Fleischm., Wistuba & S. McPherson—GU

Reference

FLEISCHMANN, Å., A. WiSTUBA, AND S. McPherson. 2007. Dros-

era solaris (Droseraceae), a new sundew from the

Guayana Highlands. Willdenowia 37:551-555.

EBENACEAE (p. 282-283)

Diospyros gallo B. Walln.—BO

Diospyros ottohuberi B. Walln.—BO

Diospyros tepu B. Walln.—BO

Reference

WALLNOFER, B. 2000. Neue Diospyros-Arten (Ebenaceae)

aus Südamerika—ll. Ann. Naturhist. Mus. Wien, ser.

B, 102:419-432.

EUPHORBIACEAE (p. 290-300)

Paul Berry

Astraea lobata (L.) Klotzsch—BO, DA, GU, SU, FG

ai lobatus Enea DE Seele

i. ia. | pp

gyropnynoides KUN — shou

: "AS ty dei | "M L om

J i

Croton cajucara Benth. —add FG

Croton essequiboensis Klotzsch—remove*[=Croton populifolius

P. Mill.?]"

Sd Kaieteuri Jabl. Odio uu"

tar

ene aff. nepetifolius Baill should replace Croton nepe-

tifolius

Croton niveus Jacq.—BO

o polypleurus Croizat—add e

I esl

Coton foranmersi var .roraimensis —should | be deleted

ae sacaquinha Crolzat—GU

Croton sampatik Muell. Arg.—AM

Croton sp. A—should be deleted

Croton sp. B—should be deleted

Croton sp. G—should be deleted

Croton sp. H—should be deleted

Croton sp. X [= Croton nervosus Klotzsch, nom. illeg.] —AM,

BO, GU

Croton tricolor Klotzsch—FG

Euphorbia comosa Vell.—erase" SU?

Euphorbia aff. laurifolia Juss.—should be deleted

t ll d ad A al E £ H £ al T cL:.

Feuille ;

Euphorbia pulcherrima Willd. Ex Klotsch—correct epithet

Euphorbia tirucalli L—add FG

Euphorbia tithymaloides L.—BO, GU, SU, FG

Euphorbia tithymaloides subsp. tithymaloides [native

to GU, SU; cultivated in all; collected from]—BO, GU,

U, FG

Manihot baccata Allem—FG

Micrandra sp. A—should be deleted

Micrandra spruceana (Baill.) R.E. Schult.—remove the mention

"Irecord probably misident.]’

Note: Next species in the Checklist, M sprucei (Muell. Arg.

R.E. Schult, is a different present in the area

covered by the checklist.

Synonyms and misidentifications

{| E Í IL A

should be deleted

Note: They are two different species and C. adenoptera is

n r

Croton argyrophylloides Kunth = Croton tricolor Klotzsch

Croton barlettii Lanj.—should be deleted

Croton glandulosus subsp. hirtus (L'Hérit.) Croizat, “Bull*—

should be deleted

Note: n the Checklist it is a repeat of the entry 2 nes anor:

“Croton glandulosus subsp. hirtus (L'Hérit.) Pu

Croton ic Jabl. = Croton cuneatus Klotzs

Croton kavanayensis Steyerm. = Croton aes Aubl.

Croton lobatus L. = Astraea lobata (L.) Klotzsch

Croton micans Sw. = Croton argyrophyllus—instead of "=" it

should read “misidentification for”

Croton nervosus Klotzsch [nom. illeg.] = Croton argyrophyllus—

zou read “= Croton D x"

iAH | ^E

Croton rhamnifolius Kunth (illeg.) = Croton conduplicatus

unt

Croton sp. A = Croton parodianus Croizat

Croton sp. B = Croton sampatik Muell. Arg.

Croton sp. G = Croton polypleurus Croizat

Croton sp n niveus Jacq.

Fuphorbia aff. laurifolia Juss. [misidentification; Andean

species)

Euphorbia tithymaloides L.—should be deleted

Euphorbia tithymaloides subsp. tithymaloides—should be

deleted

Micrandra sp. A e

Rene tithymaloides (L.) Poit. — Euphorbia tithymaloides

Al = doo at ne p OLAS L

"m tithymaloides subsp. tithymaloides = Euphorbia

tithymaloides subsp. tithymaloides

Reference

Auem, A.C. 1999. A new species of Manihot (Euphor-

biaceae) from the Brazilian Amazon. Int. J. Plant Sci.

160:181-187.

GENTIANACEAE (p. 304-309)

Curtia ayangannae L. Cobb & Jans.-Jac.—GU

805

Reference

Coss, L., M.J. JANSEN-JACOBS, AND R. VAN DER HAM. 2007.

Curtia ayangannae, a new species of Gentianaceae

from the Pakaraima Mountains, Guyana. Blumea

52:5-10.

GESNERIACEAE (p. 309-312)

C. Feuillet and L.E. Skog

Alloplectus cristatus (L) Mart.—should be deleted

Alloplectus cristatus var. epirotes Leeuwenb.—should be

deleted

Alloplectus savannarum CN. Morton—should be deleted

Besleria neblinae Feuillet—AM

Besleria so. A—should be deleted

Besleria sp. B—should be deleted

Besleria sp. C—should be deleted

Besleria yatuana Feuillet—

Crantzia epirotes (Leeuwenb.) J.L. Clark—GU

Episcia duidae Feuillet—A

Episcia rubra Feuillet —AM

Episcia sp. A—should be deleted

Episcia sp. B—should be deleted

Sloxinla erinoides (DC) E. la Roa SoN ded K. ac cal wees

M MA

Koellikeria erinoides (DC) ange ME be deleted

euillet—AM

Nautilocalyx roseus Feuillet—

Nautilocalyx ruber Feuillet— an

aut ilocalyx Sp. À

Nautilocalyx sp. D—should be deleted

Nautilocalyx sp. E—should be deleted

Nautilocalyx sp. F—should be deleted

Nautilocalyx sp. G—should be deleted

Nautilocalyx vestitus Feuillet—BO

Nematanthus savannarum (C.V. Morton) J.L. Clark—BO,

GU, SU

Paradrymonia glandulosa a

Paradrymonia hamata Feuillet

Paradrymonia maculata (Hook. f. T" uic DA

Paradrymonia lutea Feuillet ——AM

Paradrymonia maguirei Feuillet—AM

Paradrymonia sp. A—should be deleted

Paradrymonia sp. B—should be deleted

Paradrymonia sp. C—should be deleted

Paradrymonia sp. D—should be deleted

Paradrymonia sp. E—should be deleted

Paradrymonia tepui Feuillet—AM

Paradrymonia yatua Feuillet —AM

Seemannia purpurascens Rusby—GU, FG

Synonyms ane Ds

SAIA n l^

Crantzia epirotes

(Leeuwenb) J.L. Clark

806

Alloplectus savannarum CV. Morton = Nematanthus savan-

arum (CV. Morton) J.L. Clark

Besleria sp. A = Besleria neblinae Feuillet

Besleria sp. B — Besleria neblinae Feuillet

Episcia sp. B = Episcia duidae Feuillet

~l os £n | 3 AAA AL]

cens Rusby

Koellikeria erinoides (DC.) Mansf. = Gloxinia erinoides (DC.) E.H

Roalson & J.K. Boggan

Nautilocalyx sp. A = Nautilocalyx vestitus Feuillet

Nautilocalyx sp. B = Nautilocalyx orinocensis Feuillet

Nautilocalyx sp. C = Nautilocalyx roseus Feuillet

Nautilocalyx sp. D = Nautilocalyx paujiensis Feuillet

Nautilocalyx sp. E = Nautilocalyx ruber Feuillet

Nautilocalyx sp. F = Nautilocalyx crenatus Feuillet

Nautilocalyx sp. G = i ] i

Paradrymonia sp. B = Paradrymonia hamata Feuillet

Paradrymonia sp. C = Paradrymonia glandulosa Feuillet

Paradrymonia sp. D = Paradrymonia tepui Feuillet

Paradrymonia sp. E = Paradrymonia maguirei Feuillet

References

Clark, J.L. 2005. A Monograph of Alloplectus (Gesneri-

aceae). Selbyana 25:182-209.

FeuiLLET, C. 2008. Folia Taxonomica 6. Two new species

of Besleria (Gesneriaceae) from the Venezuelan

Guayana J. Bot. Res. Inst. Texas 2:269-273.

FeuiLLET, C. 2008. Folia Taxonomica 7. Two new species

and a new section in Episcia (Gesneriaceae) from

the Venezuelan Guayana. J. Bot. Res. Inst. Texas

2:275-280.

FeuiLLET, C. 2008. Folia Taxonomica 10. New species of

ezuelan Guayana. J. Bot. Res. Inst. Texas 2:825-836.

FeuiLLET, C. 2009. 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.

J. Bot. Res. Inst. Texas 3:133-138.

FEuiLLET, C. 2009. FOODS 15. Five new ERES

of P Paradrymonia (Gesne

aceae, ide from the Venezuelan Guayana. )

Bot. Res. Inst. Texas 3:583-592,

Roatson, E.H., J.K. BOGGAN, AND L.E. Skos. 2005. Reorga-

nization of tribal and generic boundaries in the

Gloxinieae (Gesneriaceae: Gesnerioideae) and the

description of a new tribe in the Gesnerioideae,

Sphaerorrhizeae. Selbyana 25:225—-238.

Skoc, L.E. anp C. FeuiLLET. 2008. Gesneriaceae. In: M.J.

Jansen-Jacobs, ed. Flora ofthe Guianas ser. A, 26. 136

pages. Royal Botanic Gardens Kew, Richmond.

HIPPOCRATEACEAE (p. 313-314)

In the family authors, after Mennega, delete ^t"

LEGUMINOSAE-CAESALP

Paloue sandwithii Redden—GU

(p. 333-346)

Reference

Reppen, K.M. 2008. A new species of Paloue (Legumi-

nosae: Caesalpinioideae: Detarieae) from Guyana,

South America. Brittonia 60:25 7-260.

LEGUMINOSAE-FABOIDEAE (p. 346-365)

A la Cer

wr Y wt

P s J 3 : fc J tl

& Azevedo—GU

Swartzia anomala R.S. Cowan—add GU

Swartzia benthamiana var. yatuensis R.S. Cowan—should

be deleted

Lora D ps

It. LJ IA | Halata

Swartzia brachyrachis var. glabrata R.S. Cowan—should be

Swartzia canescens Torke—

Swartzia glabrata (R.S. Cowan) Torke—BO, GU, SU

Swartzia kaieteurensis Torke—delete "sp. nov. ined"

Swartzia ramiflora Torke—AM

Swartzia rosea Mart. ex Benth.—AM

Swartzia stipellata R.S. Cowan should be deleted

Trischidium alternum (Benth.) H.E. ireland —GU

Synonyms and misidentifications

Bocoa alterna (Benth.) R.S. Cowan = Trischidium alternum

(Benth.) H.E. Ireland

Swartzia alterna Benth. = Trischidium alternum (Benth.) H.E.

Ireland

Swartzia benthamiana var. yatuensis R.S. Cowan = Swartzia

rosea Mart. ex Benth.

Swartzia brachyrachis var. glabrata R.S. Cowan = Swartzia

glabrata (R.S. Cowan) Torke

Swartzia stipellata R.S. Cowan = Swartzia anomala R.S.

References

IRELAND, H.E. 2007. Taxonomic changes in the South

American genus Bocoa (Leguminosae-Swartzieae):

reinstatement of the name Trischidium, and a syn-

opsis of both genera. Kew Bull. 62:333-350.

Scuürz RODRIGUES, R. AND A.M.G. Azevedo. 2006. Guianoden-

dron, a new genus of Leguminosae (Papilionoideae)

from South America. Novon 16:129- 132.

SrERGIOS, B. AND G.A. AYMARD. 2008. A striking new species

of Aldina (Fabaceae-Swartzieae-Aldinae) from the

Venezuelan Guayana highlands. Harvard Pap. Bot.

13(1):29-33.

Feuillet, Uj lated checklist of

Tonke, B.M. 2007. New combinations and species-level

synonyms in Swartzia (Leguminosae: Papilionoide-

ae). Novon 17:110-119.

Torke, B.M. 2007. Three new species of Swartzia (Le-

guminosae-Papilionoideae) from northern South

America. Bot. J. Linn Soc. 153:343-355.

LEGUMINOSAE-MIMOSOIDEAE (p. 365-376)

Inga loubryana Poncy—GU, FG

Reference

Poncy, O. 2007. A new species of inga Mill. (Fabaceae,

Mimosoideae) from the Guianas. Adansonia sér. 3,

29(2):249—254.

LISSOCARPACEAE (p. 379)

i YA/-Il LI i J 5$ "| ri

A A II

o of tl

belong here.

LORANTHACEAE (p. 381-384)

Oryctina atrolineata Kuijt—GU

Reference

Kuur, J. 2003. A new species of Oryctina (Loranthaceae)

from Guyana. Brittonia 55:169-172

MALPIGHIACEAE (p. 386-393)

Alicia macrodisca (Triana & Planch.) W.R. Anderson—AM,

Carolus sinemariensis (Aub!) W.R. Anderson—AM, BO,

G F

Christianella glandulifera (Cuatrec.) W.R. Anderson—AM

Christianella surinamensis (Kosterm.) W.R. Anderson—

AM, GU, SU

Aalpighiod is (W.R. Anderson) WR. Anderson—

GU, SU, FG

Malpighiodes liesneri (W.R. Anderson) W.R. Anderson—

Mascagnia castanea (Cuatrec) W.R. Anderson—should be

elete

Mascagnia glandulifera Cuatrec.—should be deleted

Mascagnia guianensis W.R. lel he A be deleted

Mascagnia liesneri W.R. Anderson—should be deleted

Mascagnia macrodisca ne a Planch.) Nied.—should be

deleted

Mascagnia microcarpa (Sandwith) W.R. Anderson—should

be deleted

LIX” eme bed A A |

Mascagnia surinamensis (Kosterm) WR Aleron

e deleted

Niedenzuella acutifolia (Cav.) W.R. Anderson—GuU, SU,

Niedenzuella castanea (Cuatrec.) W.R. Anderson—AM

Niedenzuella poeppigiana (A. Juss.) W.R. Anderson—AM,

Tetrapterys acutifolia Cav.—should be deleted

807

Tetrapterys poeppigiana (A. Juss.) Griseb.—should be

deleted

Synonyms and misidentifications

Banisteria sinemariensis Aubl.—should read “=

mariensis (Aubl.) W.R. Anderson"

Diplopterys microcarpa (Sandwith) W.R. Anderson—should

ad “= Carolus sinemariensis (Aubl) W.R. Anderson”

Heteropterys castanea (Cuatrec.) W.R. Anderson—s ii read

"= Niedenzuella castanea (Cuatrec.) W.R. Anderson"

Hiraea macrodisca (Triana & Planch.) Nied. ed read “=

Alicia macrodisca (Triana & Planch.) W.R. Anderson"

Hiraea poeppigiana A. Juss.—should ial = Niedenzuella

oeppigiana (A. Juss.) W.R. Anderso

Hiraea schizoptera Turcz.—should read “= ‘cies sinemariensis

(Aubl.) W.R. Anderson”

Carolus sine-

Malpighia volubilis Sims—should read “= Carolus sinemariensis

(Aubl.) W.R. Anderson"

Mascagnia castaneaa (Cuatrec.) W.R. Anderson = Niedenzuella

castanea (Cuatrec) W.R. Anderson

Mascagnia anisopetala var. macrodisca (Triana & Planch.)

Kosterm.—should read “= Alicia macrodisca (Triana &

Planch.) W.R. Anderson"

Mascagnia elles fera Cuatrec.—should read “= Christianella

glandulifera E rec.) W.R. Anderson"

Mascagnia guianensis W.R. Anderson—should read “= Mal-

pighiodes guianensts (WAR. DAD MM Anderson"

Mascagnia hondensis (Sandwith) W. Anderson—should read

= = Carolus sinemariensis (Au bl. ) W. R. Anderson"

Mascagnia liesner erson—should read “=

pighiodes liesneri (WR. Anderson) W.R. Anderson"

Mascagnia macrodisca (Trian a & Planch.) Nied. = Alicia mac-

rodisca (Triana & Planch) WR Anderson

Mascagnia microcarpa (Sandwith) W.R. Anderson = Carolus

sinemariensis (Aubl.) W.R. Anderson

Mal-

= Chris-

Lam

tianella surinamensis (Kosterm.) W.R. Anderson

Mascagnia poeppigiana (A. Juss.) W.R. Anderson—should

be deleted

Mascagnia sinemariensis (Aubl.) Griseb. = Carolus sinemariensis

(Aubl) W.R. Anderson

Mascagnia subsericea Cuatrec.—should read “= Niedenzuella

poeppigiana (A. Juss.) Griseb”

Mascagnia surinamensis Vice WA. Dr = Chris-

ianella surinamensis (Kosterm . Ander

Tetrapterys acutifolia Cav. = ere Minn (Cav.) W.R.

Tetrapterys poeppigiana (A. Juss.) Griseb.—should read “=

Niedenzuella poeppigiana (A. Juss.) Griseb?

Reference

ANDERSON, W.R. 2006. Eight segragates from the neo-

tropical genus Mascagnia (Malpighiaceae). Novon

16:168-204.

MORACEAE (p. 423-428)

Ficus duartei C.C. Berg & Carauta—FG

Ficus duckeana C.C. Berg & Ribeiro—FG

808

Ficus popenoei subsp. malacocarpa (Standl) CC. Berg—

AM, BO, DA, GU, SU, FG

Synonyms and misidentifications

Ficus malacocarpa Standl. = Ficus popenoei subsp. malaco-

carpa (Standl) C.C. Berg

References

Bera, C.C. AND J.PP. CARAUTA. 2002 [2003]. New species of

Ficus (Moraceae) from Brazil. Brittonia 54:236-250.

Bera, C.C. 2007. New species of Ficus (Moraceae) from

South America. Blumea 52:569-594.

MYRSINACEAE (p. 430-434)

Cybianthus p tiae Pipoly | pelling

Reference

PiroLy, JJ. 1999. Two new species of Myrsinaceae from

French Guiana. Brittonia 51:128-133.

MYRTACEAE (p. 434-442)

B. Holst and C. Feuillet

Calycolpus cochleatus McVaugh—Huber 9375 (MO, VEN)—

add BO

Calycorectes batavorum Mevaulg a Be pa

Á —

Can t

-t 4

Ai ee OK

Cal yptranthes forsteri O. Berg— —Oldeman B-2825 (CAY)—add

Calyptranthes pulchella DC.—Cremers 8213 (CAY)—add FG

ia od sp.—was in the Checklist as C. lepida

ugh

pe a L r a

LA AAR SHE Cil

Eugenia enormis (McVaugh) Mattos—BO,

Eugenia exaltata O. Berg—Acevedo et al. 5750 Coe SU

Eugenia gomesiana O. Berg—Fleury 1012 (CAY)—add FG

Eugenia maxima Mevaugn): Ma

SAFE KI

LAT A 2s IN,

Amshoff—Service Foretie -6259 (CAY)—add

ana MA

E Aad VA

H T

Myrcia gonini Maio is Granville 899 (CAY aun FG

)—add FG

A sd 1-45: 4(C

Eugenia galbaoensis Mattos—F

Synonyms and misidentifications

Calycorectes batavorum McVaugh = Eugenia batavorum

(McVaugh) Mattos

Mattos

Cal

attos

Calyptranthes lepida McVaugh—misidentification for an

undescribed species

Eugenia aubletiana Mattos (nom. illeg.) =

ubl.

Eugenia yatuae (McVaugh) Mattos (superfluous) —

yatuae (McVaugh) Holst

Eugenia latifolia

Eugenia

Z£' Tons ST

References

is sr, B.K. 2002 bii species and notes on Myrtaceae

th America Selbyana 23:137-180.

Marros, J.R. 1996. N

XII. Loefgrénia 108:1—2.

Mattos, J.R. 2005. Consideragóes sobre Calycorectes

Berg. Loefgrénia 120:1—24.

A Às rer 703m

OCHNACEAE (p. 445-449)

Ouratea candelabra Sastre —GU

Ouratea jansen-jacobsiae Sastre—GU, SU

Ouratea miniguianensis Sastre—F

CI]

Ouratea retrorsa Sastre—FG

Ouratea sipaliwiniensis Sastre—SU

Ouratea superimpressa Sastre—GU

Ouratea takutuensis Sastre—GU

References

SASTRE C. 2006. Deux nouvelles espéces d'Ouratea

(Ochnaceae) des Guyanes. Adansonia, sér. 3,

28(1):119-1 E

SasTRE C. 2007.5 I

ceae) des Guyanes. dansa sér. 3, 29(1):77-91,

WALLNOFER B. 1998: A revision of Perissocarpa Steyerm.

& Maguire (Ochnaceae). Ann. Naturhist. Mus. Wien,

ser. B, 100:683-70/.

J'Ouratea (Ochna-

PASSIFLORACEAE (p. 454-456)

Passiflora angusta Feuillet & J.M. MacDougal—BO, GU

Passiflora arta Feuillet—remove "ined”-—GU

Passiflora compar Feuillet—remove "ined^—GU, FG

Passiflora curva Feuillet —FG

Passiflora davidii Feuillet—FG

Passiflora foetida var. orinocensis (Killip) Feuillet—remove

“ined”

Passiflora rufa Feuillet & J.M. MacDougal—GU, SU, FG

Passiflora sp. D—should be deleted

Passiflora sp. G—should be deleted

Passiflora sp. H—should be deleted

Passiflora tecta Feuillet —BO, GU, SU?

dn ang misidentifications

Passiflor ill

afi pa es Passiflora davidii Feuillet

Passiflora sp. H = Passiflora rufa Feuillet & J.M. MacDougal

t&J.M. MacDougal

References

FeuiLLET C. 2007. Folia taxonomica 1. Validation of two

taxa from northern South America. J. Bot. Res. Inst.

Texas 1:143-144.

Feuitet C. 2007. Folia taxonomica 2. New species of

Passiflora subgenus Passiflora (Passiftoraceae) from

the Guianas. J. Bot. Res. Inst. Texas 1:819-825.

Feuer C. 2007. Folia taxonomica 3. Passiflora davidii

1.15 a. £r

Feuillet, U lated cl

and a key to the sections of superecon Stipulata.

J. Bot. Res. Inst. Texas 1:895—898.

FEUILLET C, 2008. Folia taxonomica 8. Passiflora tecta, a

new species in subgenus Passiflora from the Gui-

anas. J. Bot. Res. Inst. Texas 2:281-283.

FeuiLLEr C. 2009. Folia taxonomica 13. Passiflora curva

(Passifloraceae), a new species in subg. Passiflora

supersect. Coccinea from French Guiana. J. Bot. Res.

Inst. Texas 3:577-579.

FEUiLLET C. AND J.M. MacDoucat 2008. Folia taxonomica

9. New species of Passiflora subg. Decaloba (Passi-

floraceae) from northern South America. J. Bot. Res.

Inst. Texas 2:81 7-824.

Passiflora

PIPERACEAE (p. 458-466)

Piper aulacospermum Callejas—

Piper ciliomarginatum Górts & Christenh.—GU

Piper remotinervium Górts—GU, FG

Reference

GORTS-VAN Run, A.R.A. AND R. CALLEIAS. 2005. Three new

species of Piper (Piperaceae) from the Guianas.

Blumea 50:367-373.

RUBIACEAE (p. 479-507)

P.G. Delprete

et al. (2009) support the separation of Appunia Hook. f. from

Morinda L., therefore two n mbinations in Appunia

from the Guayana Shield are necessary. The phylogenetic

studies of Dessein (2003), Dessein et al. (20068), Groeninckx

et da (2009a, 20090) zd | ens etal mee confirm the wide

lin incl. Borreria G. Mey),

Aen | B 1993), Boom

& De Mm (2002), Delprete et al. (2005), Delprete & Cortés

("2006" 2007), Delprete (2007), De Vré (2000), Dessein et al.

(2003, 20063), and Vaes et al. (2006). Arecenti re- -delimitation

of Gi (“2004"

AA fi D

[2005], and the necessary synonymies are listed Delon Ad-

ditional synonyms and rearrangements here presented are

based on direct observations and ongoing revisionary and

floristic treatments by the present author.

3U, SU, FG

A . a? fL? Lf itc

e e

fT»

Appunia calycina (Benth ) mw GU

Appunia debilis Sandwith—GU

diis a longipedunculata (Steyerm.) Delprete, comb.

— mund Morinda longipedunculata Steyerm., Ann

Missouri Bot. Gard. 75:340. 1988; Tyre: VENEZUELA: Siérta

de la Neblina, 22-23 Jul 1984, Davidse 27651 (HOLOTYPE:

MO, isoTYPE: VEN).

809

Appunia peduncularis (Kunth) Delprete, comb. nov.—

AM, GU

Al J LY LJ I p Bonpl,

Nov. Gen. Sp. 3(12, quarto): a Todes TUE VENEZUELA

Rio Orinoco, San Borja

P- Bonpl).

SU [i

Appunia tenuiflora Hook. f—AM, BO, GU, SU, FG

nia venezuelensis Steyerm

PONeId alata Aubl) Dc AMB be oe deleted

—

Borreria capitata (Ruiz & Pav) DC.—should be lc

Borreria capitata var. capitata—should be delet

Borreria capitata var. suaveolens (G. Mey.) etoile

deleted

Borreria capitata var. tenella (Kunth) Steyerm.—should be

deleted

C4 ba IA | d led

D 1 £ pp qe C4

Borreria densiflora DC. ~ should be deleted

Borreria hispida Spruce ex K. Schum.—should be deleted

chniild ha Halatan

R chruvila ha Halatan

Borreria latifolia (Aubl) K. Schum.—should be deleted

PONE Inge de Standl, e AT e sou ok Boek

Asartanc

D i " FA chari les wm

Borreria | pygmaea Spruce ex K. Schum.—should þe deleted

Borreria remota (Lam.) Bacigalupo & E.L. Cabral—should be

deleted

Borreria repens DC.—should be deleted

Borreria verticillata (L.) G. Mey.—should be deleted

i l^ IA! xl ex J

Calycophyllum intonsum | Steyerm. —GU

GU ||

Calycophyllum spectabile Steyerm. —GU

Calepiaies guianensis AUD. ~aU, FG

Inrate—__(Gl]

Chimarrhis cymosa Jaca. —should be deleted [endemic

to Lesser Antilles; cf. Delprete, 1999, Flora Neotropica

77:152-155)

fe fa? j Late C4 Ll Id kh ^ eed

Chiococca nitida var. amazonica Muell. Arg.—should be

deleted

Chiococca nitida var. chimantensis Steyerm.—should be

deleted

Chiococca nitida var. nitida.—should be deleted

rete—FG

Diodella apiculata (Willd. ex Roem. & Schult) Delprete—

BO, GU, SU, FG

Note: Delprete 2004. in A. Reis, Fl. llutrada Catar., Rubiaceas

lall Baala AAA Roem & Sel li Mal

ex BO

Note: Delprete 2004. in A. Reis, Fl. llutrada Cater, Bupiacess

810

Diodella sarmentosa (Sw. Bacigalupo & E.L. Cabral ex Bo-

rhidi— GU, SU, FG

Note: somal ace para de México. 186.

em. & Schult.) K. Schum.—should

deleted -

Diodia Od ae RN ex Roem. & Schult) Cham. &

S —should be del

Diodia mutton DG shi M deleted

be deleted.

i D LA] | aa | Ay

tula(Willd &H anne: ex Mon & Schult.) Cham.

& Schltdl. —should be deleted

Diodia radula subsp. venezuelensis Steyerm.—should be

ny: fe

Diodia sarmentosa Sw.—should be deleted

Diodia spicata Mig.—should be deleted

Diodia teres Walter Solo be deleted

lata t U, FG

Faramea sessiliflora Aubl. var, pedunculata peer —should

be deleted

Faramea sessiliflora var. Ea OU! ild be ge elo

L if

A J m V MAMAS MM

D I | Lal ka Halatan

PIMLVAVILTVEE TA

Kutchubaea surinamensis (Bremek.) C.H. Perss.—SU, FG

Note: C.H. Perss. 2005. Revista Biol. Neotrop. 2:67.

LA Dad pl

m . J iD “¿LA Os l ld BA Halorar]

Pal ki f

AA H d jt. E fC J Hn

inda | (Si St should be deleted

iiis longipedunculata Steyerm. [Sierra de la Neblina]—

leted

hould be de

rn EP +l l LAI l lE

j mal {Ca | C4 karil H ha dalaotar

Morinda surinamensis (Bremek) Steyerm.—should be

deleted

j " aff iD tl.) C4 | LA |

j I ul WW

be deleted

Palicourea lemoniana C.M. Taylor—AM

pou gulanensis Aubl.—ada SU

A Ce l Ie ha Halata

Psychotria ula var. ligularis should be deleted

Semaphyllanthe intonsa (Steyerm.) L. Andersson—should

deleted

LL LE d dg

) | Andersson—should

be deleted

Semaphyllanthe spectabilis (Steyerm.) L. Andersson—should

e deleted

Spermacoce alata Au b.—AM, GU, su, FG

+ RO)

or d

Spermacoce capitata Ruiz & Pav —AM, Bo, „GU, ue: y

(C———— ÁN U

Spermacoce confertifolia (Steven) Govaerts—AM, BO,

Spermacoce densiflora (DC.) Liogier—AM, BO, GU

Spermacoce exilis (L.O. Williams) C.D. Adams—AM, BO,

DA, GU, SU, FG

3/79)

f Texas 3(2)

Spermacoce hexangularis Aubl.—FG

Ee L of fe AFIL ay PR

l /i4030^

nem

^ fi

Kunth (1819)—AM, BO, DA, GU, SU, FG

Spermacoce latifolia Aubl —AM, BO, DA, GU, SU, FG

nal. & Steyerm.) Govaerts—

Spermacoce multiflora (DC) Delprete—DA

Spermacoce neohispida iuga BO, GU, SU, FG

illd. ex Roem. €: Schult.—AM, BO,

DA, GU, SU, FG

Ba SKA Martane Mal AH GU 1)

Spermacoce prostrata Aubl —AM, BO, ps GU, SU, FG

GU , SU

Spermacoce remota Lam.—AM, BO, GU. su, FG

Spermacoce spicata (Miq.) Delprete—AM, GU, SU, FG

Spermacoce spruceana Kuntze—AM, BO

Spermacoce suaveolens (G. Mey.) Kuntze—GU, SU

Spermacoce tenella Kunth—AM, BO, GU, SU, FG

Spermacoce verticliata L- TAM Po, DA, Gu, ee FG

ee an

MA Ca A ua ue TU A AA MA P, M MU V UAM

L ||| ro

Synonyms and misidentifications

Alibertia surinamensis (Bremek.) a = Kutchubaea suri-

namensis (Bremek.) C.H

Appunia longiloba Steyerm: er tenuiflora Hook. f.

Appunia sparsiflora Steyerm. = Appunia tenuiflora Hook. f.

Appunia tenuiflora var. leiophylla Steyerm. = Appunia tenuiflora

ook. f.

d n Bremek. = Appunia brachycalyx (Bre-

er. calycina (Benth.) Bremek. = Appunia calycina

(Benth.) Sandwith

Bellynkxia surinamensis Bremek. = Appunia surinamensis

(Bremek.) Steyerm

Bigelovia sphaerica DC. = Spermacoce remota Lam

Bigelowia verticillata (L) Spreng. = Spermacoce verticillata n

Borreria alata (Aubl) DC. = Spermacoce alata Aubl.

Borreria doc Bello 2 Spermacoce ocymifolia Willd. ex

ult

lis Mes DC. = Spermacoce ocymifolia Willd. ex Roem.

Borreria assurgens (Ruiz & Pav.) Griseb. = Spermacoce remota

Borreria nU oo = Spermacoce bolivarensis

teyerm.) Gov

Borreria capitata n E Pav.) DC. = Spermacoce capitata Ruiz

& Pav.

Borreria capitata var. capitata f. ferruginea (A. St.-Hil.) Steyerm.

= Spe ermacoce capitata e i a

mien

m. = Spermacoce

capitata Ruiz & Pav.

Borreria capitata var. suaveolens - a Steyerm. = Sperma-

coce suaveolens (G. Mey.) Ku

Borreria capitata var. tenella (Ku a Steyerm. = Spermacoce

tenella Kunth

t lindatad ct impe E H E al fet a *.1.

Feuille :

Borreria cataractarum Steyerm. = Spermacoce cataractarum

(S aerts

Borreria confertifolia Steyerm. = Spermacoce confertifolia

teyerm.) Govaerts

Borreria densiflora DC. 2 Spermacoce densiflora (DC.) Liogier

Borreria elongata DC. = Spermacoce capitata Ruiz

Borreria ferruginea (A. St.-Hil.) DC. 2 Spermacoce capitata Ruiz

& Pav.

Borreria fockeana Miq. = Spermacoce latifolia Aubl.

Borreria hispida Spruce ex K. Schum. = Spermacoce neohispida

Govaerts

Borreria intricata Steyerm. = Spermacoce intricata (Steyerm.

ovaerts

Borreria jangouxii Steyerm. = Spermacoce jangouxii (Steyerm.)

Borreria kappleriana Miq. = Spermacoce capitata Ruiz & Pav.

Borreria laevis (Lam.) Griseb. = Spermacoce remota Lam

Borreria latifolia (Aubl.) K. Schum. = Spermacoce latifolia

Aubl.

Borreria latifolia var. fockeana (Miq.) Bremek. = Spermacoce

latifolia Aubl.

Borreria linifolia DC. = Spermacoce remota Lam.

Borreria linoi

orreria macrocephala Standl. & S . = Spermacoce

macrocephala (

Borreria minima DC. = Spermacoce vertici

Borreria multiflora (DC.) Bacigalupo €: Cabral

multiflora (DC) Delpiete e

FC loci M

vaerts

lata L.

— Spermacoce

= 0

44D o Da;

Borreria ocymifolia (Willd. ex x Roem. & Schult) Bacigalupo

& E.L. Cabral = Spermacoce ocymifolia Willd. ex Roem.

& Sc hu i

A DUO Te ppl ices Aa L.

Aartanc fialantti

folia (M. Martens € Galeotti) Hemsl.

Borreria penicillata Miq. = Spermacoce latifolia Aubl.

Borreria perrottetii DC. = Spermacoce latifolia Aubl.

iud pound a pe uu ue — L.

Cabral = Sper-

macoce pulchisioula (Bremek) o

Borreria pygmaea Spruce ex K. Schum. = Spermacoce spru-

ceana Kuntze

Cabral = Spermacoce

remota Lam.

Borreria repens DC. = Spermacoce exilis (L.O. Williams) C.D.

Borreria spicata (Miq.) oe & Cabral = Spermacoce

spicata (Miq.) Delpret

Borreria splitgerberi Beds Spermacoce latifolia Aubl.

Borreria stricta G. Mey. = Spermacoce verticillata L.

Borreria suaveolens G. Mey. = Spermacoce suaveolens (G. Mey.)

untze

Borreria suaveolens var. mE (DC) Standl. 2 Spermacoce

suaveolens (G. Me l Kun

Aav war pia tis

ave

phylla (K. Sct MA

= Spermacoce tenella: Ku nth

811

Borreria tenella (Kunth) Cham. & Schltdl. = Spermacoce tenella

Kunth

Borreria tenella var. angustifolia DC. = Spermacoce tenella

unt

Borreria tenella var. genuina K. Schum. = Spermacoce tenella

unt

Borreria tenella var. linoides K. Schum. = Spermacoce tenella

Kunt

Borreria tenella var. platyphylla K. Schum. = Spermacoce

tenella Kunth

Borreria tenella var. suaveolens (G. Mey.) K. Schum. = Sperma-

coce suaveolens (G. Mey.) Kuntze

Borreria tetraptera Miq. = Spermacoce latifolia Aubl.

Borreria trichantha Miq. = Spermacoce ocymifolia Willd. €x

chult

Borreria verticillata (L) G. Mey. = Spermacoce verticillata L.

Borreria virgata Cham. & Schltdl. = Spermacoce ocymifolia Willd.

ex Roem jit

Borreria wurdackii Steyerm, = Spermacoce wurdackii (Steyerm.)

Govaer

vaerts

Borreria xanthophylla Bremek. = Spermacoce xanthophylla

(Bremek) Govaerts

Callicocca guianensis (Aubl.) Gmel. = Carapichea guianensis

Aubl.

Carapichea aubletii DC. = Carapichea guianensis Aubl.

Cephaelis guianensis (Aubl.) Standl. = Carapichea guianensis

Aubl.

m is calycina Benth. = Appunia calycina (Benth) Sandwith

Chiococca cor Ben

| = = Chiococca nitida Benth.

Chiococca pcia pra = Chiococca nitida

Benth

Chiococca nitida var. amazonica Muell. Arg. = Chiococca

nitida Bent

Chiococca nitida var. ani f. cordata (Cowan) Steyerm.

= Chiococca nitida

Chiococca nitida var. anne Steyerm. = Chiococca nitida

entn.

ae tenuiflora Benth. = Appunia tenuiflora Hook. f.

Cordiera surinamensis Miq. ex Bremek. = Appunia surinamensis

(Bremek.) Steyerm

Dasycephala spicata (Miq.) Hook. f. = Spermacoce spicata

(Mig.) Delprete

Diodia apiculata (Willd. ex Roem. & Schult.) K. Schum. =

Min dall icula (Willd ex Roam & Schult) Del ;

Diodia articulata Pohl ex DC. 2 Spermacoce hyssopifolia Willd.

cnult

Diodia Sopon len ex SES & Schult.) Cham. & Schltdl.

|. ex Roem. & Schult

Diodia hyssoplola var. articulata (Pohl ex DC.) Steyerm. =

Spermacoce hyssopifolia Willd. ex Roem. & Schult

Diodia Nonis var. a f. psiloclada Steyerm. =

Spermacoce Respond Wilg Roem. & Schult

D fil Roem. 8: Schult.) Stey-

erm. = Spermacoce onc Willd. ex Roem. & Schult.

Diodia hyssopifolia var. linearis f. ala iS

Spermacoce hyssopifolia Willd. ex Roem. & Sch

lieomric

812

Diodia multiflora DC. = Spermacoce multiflora (DC.)

Delprete

Diodia a ll (Willd. ex Roem. & Schult.) Bremek. = Sper-

coce ocymifolia Willd. ex Roem. & Schult.

boda e Bremek. = Spermacoce pulchristipula

(Bremek) ore

Dj Hoffmanns. ex Roem. & Schult.) Cham.

& Schltdl. = o. radula (Willd. ex Roem. & Schult.)

Delprete

Diodia radula subsp. venezuelensis Steyerm. = Diodella radula

(Willd. ex Roem & Schult) Delprete

IN it ^ D I

f Texas 3(2)

C L Tl LL H CA

) L. Andersson = Calycophyl-

lum intonsum Steyerm.

| L i}

) L. Andersson = Caly-

cophyllum merumense Steyerm

E spectabilis Steyerm) L. Andersson = Calyco-

phyllum tabile Steyer

Sas ae Ruiz h Pav. — Spermacoce remota

TIED

Spermacoce n Kunth = Spermacoce hy Willd.

Schult

Spermacoce cal ae ón. tenella Kunth

em. € Schult. = Spermacoce

E.L. Cabral ex Borhidi

Diodia spicata Miq. = Spermacoce spicata (Miq.) Delprete

iodia surinamensis Mig. = Spermacoce hyssopifolia Willd. ex

E & Schult.

LAPUA LU Ve

k. = Faramea

r sport A Lal? tio j A p]

pedunculata (Bremek) Delprete

Hemidiodia ocymifolia (Willd. ex Roem. & Schult.) K. Schum.

(as"ocimifolia" fort. var.]) = Spermacoce ocymifolia Willd. ex

Roem. & Schult

Hoffmannia megistophylla Standl. = Patima guianensis Aubl.

Ibetralia surinamensis Bremek. = Kutchubaea surinamensis

(Bremek) C.H. P

Vorndaa aurantiaca K Kauso Steyerm. = Appunia aurantiaca

e) San

one bay Bree ) Steyerm. = Appunia brachycalyx

emek.) Ste l

Morinda calycina ue Steyerm. = Appunia calycina (Benth.)

Morinda debilis (Sandwith) Steyerm.=Appunia debilis Sandwith

Morinda longiloba (Steyerm.) Steyerm. — Appunia tenuiflora

Hook. f.

Morinda longipedunculata Steyerm. = Appunia longipeduncu-

lata (Steyerm.) Delprete

j J fs FL +h}

Delpr F Cr

Moin mum (Steyerm.) Steyerm. = Appunia tenuiflora

ook. f.

Te surinamensis (Bremek.) Steyerm. = Appunia surina-

mensis (Bremek.

gin tenuiflora (Benth.) Pieve — Appunia tenuiflora

ook. f.

st oo

mas tenuiflora var. leiophylla (Steyerm.) Steyerm. = Ap-

punia tenuiflora Hook. f.

Morinda venezuelensis (Steyerm.) Steyerm. — Appunia ven-

ezuelensis Ste

Psychotria a (Steyerm.) Delprete = Carapichea

guianensis Aubl.

Psychotria galbaoensis Steyerm. = Carapichea guianensis

Aubl,

scho ligularis — Steyerm. = Carapichea ligularis

Delpret

anchor llas v var. carapichea Steyerm. = Carapichea

gui Aubl.

ee [es Rudge - Carapichea ligularis (Rudge)

Delprete

remota Lam.

Spermacoce coerulea Pohl ex DC. = Spermacoce ocymifolia Willd.

ex Roe Schult

Spermacoce coerulescens Aubl.

Spermacoce ernstii R. Fosberg & D. ii

ovalifolia (M. Martens & Galeotti) H

Spermacoce ferruginea A. St.-Hil — m capitata Ruiz

& Pav.

— Spermacoce latifolia Aubl.

ell = Spermacoce

Spermacoce abi Bremek. = Spermacoce remota Lam.

Spermacoce laevis Lam. = Spermacoce remota Lam.

Spermacoce linearis will ex Roem. & Schult. = Spermacoce

ifolia Willd oem. & Schult

Spermacoce linoides e ex DC. = Spermacoce suaveolens (G.

Mey.

Spermacoce longifolia Aubl. = Mitracarpus sp. ?

Spermacoce longifolia f. glabrata Miq. = Spermacoce ocymifolia

Willd. ex Roem. & Schult.

Spermacoce oligodonta (Steyerm.) Govaerts = Spermacoce

rticillata L.

Spermacoce orinocensis Roem. & Schult. = Spermacoce capitata

Ruiz

Spermacoce pode Balb. ex DC. = Spermacoceocymifolia

Willd. ex Roem. & Schult.

Spermacoce pringlei S. Watson = Spermacoce ovalifolia (M.

& Galeotti) Hemsl.

Spermacoce radicans Aubl. = Spermacoce prostrata Aubl.

lens (G. Mey.) Kuntze

Spermacoce suaveolens f. pubescens Kuntze — 5permacoce

suaveolens (G. Mey.) Kuntze

Spermacoce suaveolens var. tenella (Kunth) Kuntze = Sperma-

coce tenella Kunt

Tardavel tenella (Kunth) Standl. = Spermacoce tenella Kunth

Uragoga guianensis (Aubl.) Pulle = Carapichea guianensis

Aubl

References

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Rhin

RUTACEAE us 507-510)

Fr». L I] J

"(Ducke) Kallunki”

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Rutaceae) and description of two new species from

Eastern Brazil. Brittonia 6:28-34. Author as “Emmer-

ich ex Kallunki"

SABIACEAE (p. 510-511)

Ay mard & Daly

P^ | IL

—AM

Reference

AYMARD, G.A. AND D.C. Daty. 2006. Two new species of

Ophiocaryon (Sabiaceae) from South America. Brit-

tonia 58:2/0-2/6.

VIOLACEAE (p. 547-549)

Ll Lo: SHE Rallard & Mimnzinrar EG

Reference

MUNZINGER, J.K. AND H.E. BALLARD. 2003. Hekkingia (Vi-

olaceae), a new arborescent violet genus from

French Guiana, with a key to genera in the family.

Syst. Bot. 28:345-351

814 J | of the Botanical R h Institute of Texas 3(2)

VITACEAE (p. 552-553) d'Amérique tropicale. |!I—Cissus spinosa Cambess.,

Cissus descoingsii J.A. Lombardi—should be deleted C. subrhomboidea (Baker) Planchon, C. flavens et C.

Cissus duarteana Cambess.—should be deleted kawensis, spp. nov. J. Bot. Soc. Bot. France 44:3-17.

Cissus guyanensis Descoings—add BO, GU

ne Kawelsis ae ie FG VOCHYSIACEAE (p. 553-555)

(Baker) Planch.—add FG Qualea marioniae Marcano-Berti (as “marionii")—-GU

Synonyms and misidentificatio fieforanca

Cissus descoingsii J.A. scd ue and illegitimate) — :

GisusGipanenss Deco a eet L. 2002. studies on the flora or the

Cissus duarteana Cambess. d cation for Cissus sub- Guianas n? 101. New Vochysiaceae: Qualea johan-

rhomboidea (Baker) Planch] nabakkeri and Qualea marionii. Acta Bot. Venez.

Reference 25:111-118.

Descoincs, B. 2008. Contribution à l'étude des Vitacées

ACKNOWLEDGMENTS

I am grateful to Vicki Funk who encouraged me to publish this update. 1 appreciate the input from the

reviewers, Howard L. Clark (Quito, Ecuador, 127 Aug 2009, cf. p. 996) and Paul Hiepko (B). I am thankful

for the help provided at the Smit! ian Institution by the Department of Botany librarian Robin Everly and

database specialist Sara Alexander; their patience through multiple requests was endless. This is number

149 in the Smithsonian's Biological Diversity of the Guiana Shield Program publication series.

REFERENCES

Berry, PE., K. YATSKIEVYCH, AND B.K. Horst (eds.). 1995-2005. Flora of the Venezuelan Guayana, 9 volumes. Missouri

Botanical Garden, Saint Louis.

Funk, V., T. Houtoweut, PE. Berry, C. KELLOFF, AND S.N. ALEXANDER. 2007. Checklist of the plants of the Guiana Shield

(Venezuela: Amazonas, Bolivar, Delta Amacuro; Guyana, Surinam, French Guiana). Contr. U.S. Natl. Herb.

Gorts-vAN Run, A.R.A. AND M. JaNsEN-JacoBs (eds.). 1985-2008. Flora of the Guianas. Ser. A, fasc. 1—26 (in progress);

Ser. B, fasc. 3, 4 & 6 (in progress). Koeltz, Koenigstein and Kew Royal Botanical Garden, Richmond.

Mori, S.A, G. Cremers, C.A. Gracie, J.-J. pe GRANVILLE, M. Horr, AND J.D. MitcHeLt (eds). 1997. Guide to the plants of

Central French Guiana. Part 1. Pteridophytes, Gymnosperms, and Monocotyledons. Mem. New York Bot.

Gard. 76(1):1-422.

Mori, S.A., G. Cremers, C.A, Gracie, J.-J. DE GRANVILLE, S.V. HEAD, M. Horr, AND J.D. MircheL (eds.). 2002. Guide to the plants

of Central French Guiana. Part 1. Dicotyledons. Mem. New York Bot. Gard. 76(2):1-776.

NOTES ON THE TAXONOMIC HISTORY, REDISCOVERY AND CONSERVATION

STATUS OF TWO ENDANGERED SPECIES OF CEROPEGIA (ASCLEPIADACEAE)

FROM SIKKIM HIMALAYA

D.C. manele 2 K. SUUM dis i i tangle

Cent IUG i j

Academic Research Centre Building

University of Delhi, Delhi - 1 10007, INDIA

ABSTRACT

Ceropegia hookeri C.B. Clarke ex Hook.f. and C. lucida Wall. have been reported to be endangered or possibly extinct from India. This

contribution marks Ps d art bi e i from Sikkim after a gap of nearly 133 years. We could locate a single population of C.

hookeri 8-10 i one nu ba ea oe a pea ee in eu one square

^lcorve n

kilometer area. Like ewis

have low population sizes and are faced with degradation a loss B Pour ~ studies confirm bn ay ici status of

these species and we report pu larg; pol Yg wild. Int pL dn we pa

collection history, taxonomy, tat cl isti d ti tat fel t pecies in view of the t

it * IE 1t 4 r 1 :

liic a peccas.

ooo e

Key Wonps: Ceropegia hookeri, C. lucida, endangered, habitat deg tion, g hi ge shift, Sikkim Himalaya

RESUMEN

Ceropegia hookeri C.B. Clarke ex Hook.f. y C. lucida Wall. se han citado como amenazadas o posiblemente extintas de la India. Esta

contribución marca el redescubrimiento de C. lucida de Sikkim después de unos 133 afios. Pudimos localizar una sola población de

C ] 1 : oO Jf: dal hla de T h E tm EEEE 1 E 3 C 1 A Aa

i ar]

cerca de un kilómet drado. Así mismo, ol jlo dos poblaci de C. lucid ingidas al á le North Sikkim. Ambas

: : = ] 1.1 1A E ru P J 1 22 Eu 15.1 Admd£shsene* MI 4 1: LL ] Poe

Li Ir I L 4 o Jo E

is : LES S P c M EUM Cu c acc] NN E blaci ———— — PI (€

r E

en la naturaleza. E Í ticulo discutimos la historia de 1 lecci , taxonomía, hábitat ísti y lo d nservación

pa | t 1 E 1 k| 1 2 : x 1 1 : 3i A

F pang n 7 * = S

ex situ e in situ para estas especies.

Ceropegia L. is represented by about 44—45 species, 1 subspecies and 2-3 varieties in India, of which 28

species are endemic (Ansari 1984; Jagtap & Singh 1999). Ceropegia has greater representation in Peninsular

India with 23 species (Kavade & Diwakar 2004). There are 9 species of this genus known from Himalaya

of which five are present in Eastern Himalaya. Sikkim harbors 3 out of 5 known species from the Eastern

Himalayan region. East Himalayan species of Ceropegia have been studied ud wae (1831), King (1874),

Hooker (1885), Kanjilal and Bor (1939) and Hara et al. (1982). However, based o | tion from Indian

herbaria as well as published literature, these species. are highly a a eo in collections (Nayar &

Sastry 1988). Ceropegia is represented by a disprop tely higher number of endangered species with at

least 12, out of 44 species, reported to be endangered or possibly extinct from the Indian ep ean

The Red Data Book of India (Nayar & Sastry 1987—90) was the first serious attempt at thenti

assessment of the status of Indian plants. This publication paved way for further irene studies which

unfortunately could not be followed up by action. Nearly 158 Himalayan plant species were reported to face

various degrees of threat. In Sikkim, Eastern Himalaya, more than 40 species of higher plants are reported

to be facing survival risks. Being a rich zone of biodiversity, it is crucial to understand and reassess the

Corresponding author, e-mail: mkpanditecismhe.org

J, Bot. Res. Inst. Texas 3(2): 815 — 822. 2009

816 tani Institute of Texas 3(2)

status of various end l/rare species in Sikkim. The habitats in cer are d severe pressure from

increasing human sopu laton; ]i icul , and large-scale d tivities. Recent stud-

ies have one Mn a number E NA are e likely to be consigned to extinction in Himalaya as a result of

and land use changes (Pandit et al. 2007). A number of proposed hydro-power

Du ects along Teesta River and its tril ies pose a serious threat to habitats which may further exacerbate

species population losses in these habitats (CISMHE 2007). This contribution is a part of the studies aimed

at reviewing the status of endangered species in Sikkim. In this paper we have attempted to assess the

magnitude of loss of species populations of Ceropegia and appropriate steps for their effective conservation

are indicated.

MATERIAL AND METHODS

During plant explorations carried out during 2002 to 2006 in Sikkim, covering various habitats from sub-

tropical to alpine zones, we collected a few specimens of Ceropegia L. from North Sikkim (Fig. 1). After a

thorough consultation of herbaria at Central National Herbarium (CAL), Kolkata and Botani-

cal Survey of india, Sikkim Himalaya Circle, Gangtok (SHC), followed by review of literature, these species

were identified as Ceropegia hookeri C.B. Clarke ex Hook.f. and C. lucida Wall. The voucher specimens are

deposited at the Herbarium, CISMHE, University of Delhi, Delhi, India. During the survey we also recorded

the habitat conditions, associates of these species as well as biotic and abiotic pressures so as to asses vari-

ous threats to their populations.

RESULTS

The taxonomic description of Ceropegia hookeri and C. lucida is given below. Line diagrams of the collected

specimens are given in Figures 2 and 3, respectively. The photographs of the flowers of C. hooheri and C.

lucida in Figures 4 and 5, respectively.

Ceropegia hookeri C.B. Clarke ex Hook.f., Fl. Brit. India 4:73. 1885; Ansari, Fasc. Fl. India 16:17. 1984.

Erect or decumbent twining, hairy herbs, with a strong tuberous rootstock. Stem very slender, 20-60 cm

long and 0.2—0.3 cm wide, slightly pubescent. Leaves membranous, ovate, acuminate, 2-4 cm long and

0.5-3 cm wide, hairy on both surfaces as well as on margins; peus PR about 0.5 cm, hairy. Peduncles

hairy, longer than petioles, about 1 cm long. Flowers in axill led cymes, 1-2 cm long, dark purple;

pedicels glabrous. Calyx minute, 0.2-0.3 cm long; corolla tübe 1-1.2 cm long; narrow from base to the

mouth; upper lobes short, linear-lanceolate, one third the length of the tube, gently curved and forms a short

crown. Outer corona entire, hairy; inner erect, linear. Follicles cylindrical, terete, 4-6 cm long.

Habitat.—A single population of the species on temperate grassland slopes and scrub near Nyakha-

Lachen village (North Sikkim) was located at an altitude of 2700 m. It grew with other herbaceous species

like Anaphalis contorta (D. pi Hook.f., Anemone obtusiloba E pi. Drosera peltata Thunb., Eulalia mollis

(Griseb.) O. Kuntze., Euph llichii Hook.f., Polygonatum ci (Wall) Royle, Pteris nepalensis H. Ito,

Roscoea purpurea Sm. and Vincetoxicum hirundinaria Medik. A hrubs, the associates of the species were

Eurya acuminata DC., Gaultheria nummularioides D. Don, Pieris omen (Wall) D. Don, Rhododendron arboreum

Sm., etc. This locality has shallow to moderately deep coarse loamy soil with sandy loam surface texture.

The area receives high rainfall bee: pue to es with an average rainfall of ca. 2500 mm. The average

minimum/ maximum t 4.2°C-17.1°C ane 13.9°C-23.9°C in this region. In winter

this region receives high mówi "E remains under snow int ttently from late December to February.

Distribution.— India: North Sikkim; Bhutan; sae Tibet.

Status.—During the explorations only a si tion of 8-10 individuals (Fig. 1) could be located.

The majority of the idas in this T were adul and in flowering/fruiting stages. Juvenile stages

of the species were not encountered. Attempts to relocate the species from localities like Zemu area, from

where it was recorded earlier, were unsuccessful.

Nautival et al

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BAY OF BENGAL

817

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re qur DiekritDoundan "m eae L = E jud kinn Kn adqearer

Ba oda c mmmC cmi

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NORTH DISTRICT

Kilometers

40 30 12.0 16.0 20.0

Eal

Fic. 1. Map

MAT al n.a *. Inm LI PI fT af

Fic. 2. Line di f (, ia hookeri C. B. Clarke ex Hook. f Fl

—

Nautiyal et al ' i I 819

Fic. 3. Line diagram of Ceropegia lucida Wall.

peak d ee —tThe species is threatened by habitat loss due to a number of human activities.

ral expansion, road building and grazing by d ti imals. A number of tourists

also visit besoa areas and owing to its showy flowers, these plants become targets of removal, which is also

poses a serious threat to the survival of this species.

Ceropegia lucida Wall. Pl. As. Rar. 2:33, t. 139. 1831; Hook.f. Fl. Brit. India 4:73.1885; Kanjilal & Bor. Fl.

Assam 3:309. 1939; Ansari, Fasc. Fl. India 16:22. 1984.

820 J tl tanical h Institute of Texas 3(2)

Fic. 4. A flower of Ceropegia hookeri. Fic. 5. A plant of C. lucida in its natural habitat. Note its showy

the genus derives its name.

Long twining, Sparen EE or he nee herbs, with tuberous roots. Stem solid, long creeping, up to 2 m

long. I or lanceolate, petiolate, 6.5—10.5 cm long and 1.5-2.5 cm wide; petiole short,

0.5—1.1cm long. peduncles short, 1.5-2 cm long, hairy in line. Flowers few (5), in axillary peduncled cymes;

pedicels glabrous. Calyx small, about 0.7 cm long; corolla curved, 3.5—4.5 cm long, sparsely dialated at the

base, green spotted with purple; upper corolla lobes linear, about 2 cm long, ciliate within the spathulate

apex. Outer corona of 5 bifid-deltoid ciliate lobes; inner erect, linear-clavate.

Habitat.—Two populations of C. lucida were located near Lachung village (North Sikkim) at an altitude

of 1750 m ina damp grassland/scrub (Fig. 4). The scrub was comprised of scattered trees such as Alnus nep-

alensis D. Don, Carpinus viminea Wall. ex Lindl, Erythrina arborescens Roxb., Lyonia ovalifolia (Wall.) Drude,

Rhus hookeri Sahni & Bahadur and Salix disperma Roxb. ex D. Don. Ceropegia lucida often twines on other

herbaceous species like Artemisia parviflora D. Don, Chrysopogon gryllus (L.) Trin., Capillipedium assimile

(Steud.) A. Camus, Colocasia affinis Schott., Commelina benghalensis L., Cynoglossum glochidiatum Wall. ex

Benth., Didymocarpus pedicellatus R. Br., Heracleum nepalense D. Don, Persicaria capitata (D. Don) H. Gross

and Stephania elegans Hook.f.& Thomson. This locality has shallow, loamy skeletal soil with gravelly sandy

loam surface texture. The area is humid almost round the year and receives A d Bue june to

August with an annual average rainfall of ca 3000 mm with the average mini I

ranging between 5—16?C and 14.7-24.8?C.

Distribution.—India: dad na Lo Burma, China, PORE Kong, Malaysia, Thailand.

Status.—During the survey 10 individual ld be located

(Fig. 1). All the individuals in this pea were seats and in opere amd de Res We undertook

f Ceropegia in Indi 821

Nautival et aj End

4 A

field exploration to various localities in Sikkim from where the occurrence of this species was reported

earlier. We could not relocate this species at the known localities like Lower and Upper Rayang.

Conservation threat.—The populations suffer from habitat loss due to agriculture expansion, overgraz-

ing and road construction activities, particularly in Rayang valley from where it was recorded earlier. These

habitats have undergone major land use changes due to on-going human activities.

DISCUSSION

Ceropegia hookeri C.B. Clarke ex Hook.f. is one of the poorly known endangered species of Himalaya. It was

first reported from its type locality, Lachen valley in North Sikkim (Clarke 1879). In the nineteenth-century,

the species was collected from Zemu valley and the adjacent Tibet Himalaya and its specimens deposited in

the Central National Herbarium, Kolkata (see Nayar & Sastry 1988). Apart from the e only a few speci-

mens from Zemu valley are available at CAL (July, 1909; 2 sheets). Since then, this species has remained

elusive, though the area of its occurrence has been extensively surveyed by different research groups from

India and abroad. Grierson and Long (1999) reported the species from Lachen and Chumbi valleys. A few

sheets—reportedly of this species—from Lingthem in upper Zongu, Rang Rang, Bichu and Chungthang

areas, are in the Botanical Survey of India herbarium, Sikkim Himalaya Circle (SHC), Gangtok. These sheets

neither matched Hooker's description of the species (Hooker 1885) nor with the few specimens at CAL. It

seems that the sheets at SHC may have been wrongly ascribed to C. hooheri. In view of the wrong identity

of the specimens at SHC, Gangtok, our paper places on record the occurrence of C. hooheri from its type

locality, Lachen, North Sikkim. We one E bad from Lachen valley at an altitude of 2700 m during

2003 and SE to ol tl gh 2004-2006. The specimen of C. hookeri from Lachen

agrees with tl deseription(Hoolter 1885) and ul: its type locality. It is apparently endemic to

Lachen and possibly to the adi oining Nepal cule qu WHETE it a been aa earlier (Hooker 1885).

Ceropegia lucida Wall. was first reported f lhet in Bangladesh (Wallich 1831). In India, its collection

history dates back to nineteenth-century when Look collecied it from Khasia Hills (Meghalaya), followed

by the collection of Keenam from Cachar (Assam) and King (1874) from the banks of the Rayang River,

in Sikkim. Since there was no fresh collection of this species after 1874, it was considered as endangered

or possibly extinct (Nayar & Sastry 1988). Collection of this species from Lachung Valley is therefore of

significant interest to conservationists and taxonomists and marks its rediscovery after a gap of over hun-

dred years. Also, this species is reported for the first time from North Sikkim, thus extending its range of

distribution from previously reported tropical zone to the temperate de een It may be too early

to suggest, but likely, that the species may have shifted its geographic range upv under the influence of

piona, warming (see Davis & Shaw 2001). The occurrence d the species in dia valles (1750 m) and its

from its known localities like Lower and Upper Rayang, do suggest the possibility of a range

shift Howeret, the other likely cause of species extinction of Ceropegia lucida from its known localities ap-

pears to be habitat destruction due to agricultural expansion, overgrazing, and road construction activities

in Rayang Valley. These habitats have P major ana use changes due to ongoing human activities

(Pandit et al. 2007). Additionally, the tubers of some C ies are known to be consumed locally by

humans during periods of scarcity (see Nayar & Sastry 1987, 1988). Enquiries among the local population,

however, revealed that the tubers of Ceropegia are not consumed at present. However, the respondents did

not rule out the possibility of consumption of tubers in the past, in these inaccessible terrains, during the

times of food shortage. This biotic pressure on the species in the past might have accentuated the problem

of dwindling numbers of this species in nature. All these pressures seem to be responsible for pushing the

species to the brink of extinction.

Both C. hookeri and C. lucida have low population sizes and are threatened by habitat loss due to a

number of human activities. It is well known that many Himalayan plant species with low population sizes

face survival risks (Pandit 1998, 2003). Humid conditions and adequate amount of water in the soil seem

822 | lof the Botanical R h Institute of Texas 3(2)

to be essential for their growth in the natural habitat. Due to these specific habitat requirements, the spe-

cies are difficult to cultivate at other places. To save these species from extinction efforts are on to multiply

them through seeds and tubers and reintroduce them in their natural habitats. Besides, there is an urgent

need for carrying out detailed biological and ecological studies on these species and their habitats need to

be brought under some kind of protective regulation.

Ex-situ conservation strategies

Propagating by seeds in habitats with similar ecological conditions or in nearby botanic gardens will be

useful in ex-situ conservation of these species. Micropropagation using tissue and organ culture needs to

be explored for mass multiplication and reintroduction into the wild. Since the population sizes of these

species are small, harvesting of plants or plant parts from the wild should be strictly prohibited. In addition,

there is a need to understand the reproductive biology, genetic architecture, and evolutionary relationship

of these species to ensure their reestablishment in nature and multiplication in botanic gardens, herbal

gardens, and other protected areas.

ACKNOWLEDGMENTS

The authors are thankful to H.M. Mukherjee, Central National Herbarium, Kolkata (CAL) for permission

and help with the herbarium. The tinangal suppor: n the research project sanctioned to M.K. Pandit vide

project no. J.12011/11/99-IA.I. from t t, Forests and Wildlife, Govermmeni of India

and National Hydro-electric Power ooo: Faridabad, is eratefully acknowledged. Two

Santhosk Kumar and one anonymous, provided helpful reviews.

REFERENCES

Ansari, M.Y. 1984 Fascicles of flora of India, fascicle 16, Asclepiadaceae, genus Ceropegia. Botanical Survey of

India, Kolkatta. Pp 1-34.

CISMHE. 2007. Carrying capacity studies of Teesta basin in Sikkim, Vol. IV, Ministry of Environment & Forests,

Govt. of India, New Delhi.

Davis, M.B. AND R.G. SHAW. 2001. Range shifts and adaptive responses to Quaternary climate change. Science

292:673-679.

GRIERSON, A.J.C. AND D.G. Lona. 1999, Flora of Bhutan. Vol. 2, Part 2. Royal Botanic Garden, Edinburgh. Pp

728-730

Hara, H., A.C. CHATER, AND L.H.J. Wizuiams. 1982. An enumeration of flowering plants of Nepal. Vol. IIl, London.

Hooker, J.D. 1885. Flora British India. Vol. IV. London. Repr. Bishen Singh Mahendra Pal Singh, Dehra Dun.

JAGTAP, A.P. AND N. P. SINGH. 1999. Fascicles of flora India 24, Botanical Survey of India, Kolkata. Pp 211—241.

KANJILAL, U.N. AND N.L. Bor. 1939. Flora of Assam. Vol. Ill. New Delhi.

KAVADE, S.P. AND PG. Diwakar. 2004. Ceropegia pubescens Wall. (Asclepiadaceae) — A new distributional record for

Peninsular India. J. Econ. Taxon. Bot. 28:113-116.

KiNG, G. 1874. Cited from CAL herbarium, Botanical Survey of India, Kolkata.

Nayar, M.P. AND A.R.K. Sastry. 1987. Red data book of Indian Plants. Vol. |. Botanical Survey of India, Kolkata.

Nayar, M.P AND A.R.K. Sastry. 1988. Red data book of Indian Plants. Vol. Il. Botanical Survey of India, Kolkata.

Nayar, M.P. AND A.R.K Sastry. 1990. Red data book of Indian Plants. Vol. Ill. Botanical Survey of Moa ivi

PanorT, M.K. AND C.R. Basu. 1998. Biology and conservation of Coptis teeta - an endemic licinal

herb of Eastern Himalaya. Environm. Conservation 25:262-272

Panor, M.K. AND C.R. Basu. 2003. The effects of loss of sex in clonal populations of an endangered perennial Coptis

teeta (R | ). Bot. J. Linn. Soc. 143:47-54

Panor, M.K., N. Sophi, L.P. Koh, A. Bhaskar, AND B.W. Brook. 2007. Unreported yet massive deforestation driving loss

of endemic biodiversity in Indian Himalaya. Biodivers. €: Conservation 16:153-163.

WALLICH, N. 1831. Plantae Asiaticae rariores 2:33. t. 139

NOTES ON THE TAXONOMIC HISTORY OF TWO RARE SPECIES OF BEGONIA

(BEGONIACEAE) FROM SIKKIM HIMALAYA AND THEIR CONSERVATION

DG cum Gan Bis cuela ais M.K. Panai

D IE nd

cade emic - Research Centre Building

Tian of Delhi, Delhi -110007, INDIA

ABSTRACT

Begonia satrapis C.B. Clarke and B. scutata Wall. are two pecies 1 from Eastern Himalaya. Begonia satrapis i endemic of

amis and eee drin while B. scutata is rey ] in Sikki d the adjacent Nepal Himalaya. In thi tril

to: : Hu 1 E d:

WE ICVICVV I ,and identify e

1] s 1 Bu p a a zl eL A | ono s | 1.1 1 3 TI =

found that many of the species I PE and g y Begonia

" +1 ] ] : I IE im Sans LEG 1 AQ En ] ] 1 1 E

JUL apis L 2 only t I J e H 5 Like-

wise, B ] 1 by tl ] fal 10-15 individuals each gg g ] ti

1 [- t

e Lr

KEY Wi Begonia satrapis, B. scutata, endemic, rare, habitat loss, Sikkim Himalaya

RESUMEN

Begonia satrapis C.B. Clarke y B. scutata Wall. Son dos especi onocidas del este del Himalaya. Begonia satrapi demica de

PE 1 | la Cil IN 13. E Eh " iE i l4 1 pa |

Sikkim y Darj g, mientras que B I I

recolecciones, f d id tual, e identificamos vari I l ]

LI t -l 1 11 14] L ] 1 1 1 J J J 1.1

Li ri Eu a

hos pequefios. B i trapi å la úni blaci 1 qoo d

Pu L o L Iu r [ r L

40—50 individ ] D d jante, B. scutat á rer tada [ poblaci d 10-15 individuos cada una.

C a j: gJ m d + EL E args ] JA 1 r3 3:5.

c k i

INI RODUC LION

Sikkim, with a geographical area of 7096 sq km, is a small state of India located in the Eastern Himalaya.

Owing to its varied altitudinal gradient (240—8,598 m), diverse habitats and high rainfall, the area supports

a rich flora in its tropical, sub-tropical, temperate and alpine ecosystems. This region has been identified

as one of the global hot spots of biological diversity (Myers et al. 2000). This rich diversity has of late come

under severe pressure and as a result a large number of plant taxa are threatened in this region (Nayar &

Sastry 1987, 1988, 1990). Recent studies suggest probability of large scale extinctions of Himalayan plant

species following habitat loss (Pandit et al. 2007).

The first assessment of conservation status of Indian plant species about two decades ago (Nayar &

Sastry 1987, 1988, 1990), could not be followed by intensive studies on taxa faced with survival risks. With

MS Dieu B e and habitat D several species have been restricted to only a few

Easte laya (Pandit & Babu 1998, 2003). Sikkim Himalaya, particularly,

i one of the richest areas of za dive sys where a number of plant taxa are reported to be facing risk of

survival. It is estimated that at least 45 species of flowering plants are known to be facing varying degrees

of threat in Sikkim Himalaya (Nayar & Sastry 1987, 1988, 1990; Singh & Chauhan 1998). The habitats are

under threat from increasing human population, expanding agriculture and a plethora of developmental

activities. In Sikkim, proposed large-scale hydro-power Poen did Teesta River Ada on its tributaries, pose

a serious threat to habitats, which are likely to cause tion losses if proper care

is not taken to rescue them (CISMHE 2007). In view of ihe impending yao DORE developmental activities

‘Corresponding author, e-mail: mkpandit@cismhe.org

] Rat Bac Incet Tayac 3(2): 092 _ 830. 2009

3 Eau Doa H In Y ] y a a

824 J f Texas 3(

in Sikkim Himalaya, a survey and assessment of the rare and threatened plant biodiversity in the area was

taken up so as to understand the magnitude of their population loss and also to suggest corrective measures.

MATERIALS AND METHODS

During a series of recent plant explorations (2002-2006) in Sikkim Himalaya, we collected specimens of

ns m Tm the tropical and sub-tropical regions in South Sikkim (Fig. 1). After critical study, review of

tation of specimens at Central National Herbarium (CAL), Kolkata and Botanical Survey

of India, Sikkim Circle, Gangtok, (SHC), our Begonia collections were identified as B. satrapis C.B. Clarke

and B. scutata Wall. The locations of the various populations of the two species in Sikkim were mapped

using Leica — GS5 GPS (global positioning system). The voucher specimens were deposited in the CISMHE

Herbarium, University of Delhi, Delhi, India with duplicates being deposited at the Herbarium of Botanical

Survey of India Sikkim Himalaya Circle, Gangtok (SHC) after publication of this study. During the surveys,

the habitat conditions, plant associates as well as biotic and abiotic pressures, were recorded in order to

asses the threat the Begonia populations.

RESULTS

Notes on the taxonomic description, habitat, distribution, population status in the study area, and conser-

vation methods are described below. Illustrations of Begonia satrapis and B. scutata are provided in Figures

2 and 3, respectively.

Begonia satrapis C.B. Clarke, in Hooker, Fl. Brit. India 2:638. 1879.

Tall pubescent herbs, with a tuberous rootstock. Stem erect or shortly decumbent, 24-28 cm long, usually

elongate above the radical leaves. Leaves reniform or ovate-acuminate, 4-9 x 4.5—12cm, cordate at the base,

minutely denticulate, ciliate, closely pubescent above, rufous-villous on the nerves beneath; stipules lan-

ceolate, pubescent. Peduncle elongate, up to 7 cm, densely pubescent; bracts persistent oblong-lanceolate,

hairy. Male flower: pedicel 1.5-2 cm, pubescent; tepals 2+2, Aa outer larger, about 1cm long, pubescent; the

inner smaller, 0.5 cm long, glabrous; stamens many, anthers obovoid, 0.1cm long, connec-

tive absent. Female flower: pedicel long, about 3 cm, pen pals 2+3, the outer larger, about 0.6 cm

long, pubescent, inner smaller, glabrous; styles 3, separate. Fruit: capsule with 3 equal wings, about 1.5 cm

long, pubescent. Seeds ellipsoid.

Habitat.—The populations were located aranna ee and Kitam, in aa vay dd MSIE at

altitudes of 550-1100 m. Begoni pical and sub-tropical m

ing of trees like Callicarpa arborea Roxb., Gmelin arborea Roxb., Ostodes paniculata Blume, Phoenix sylvestris

(L.) Roxb., Schima wallichii Choisy, Shorea robusta C.F. Gaertn., Tectona grandis L.f. and Terminalia myriocarpa

Van Heurck & Mull-Arg. The species is usually associated with other herbaceous species such as Ageratina

adenophora (Spreng.) R.M. King & H. ee dec dai haat assimile (Steud.) A. Camus and idi cdi flexuosus

(Nees ex Steud.) Will Watson. B led

areas along road cuttings. The species is found restricted to a narrow area in the Majitar Forest Block in South

Sikkim. The habitat is exploited for | [ timber trees, fodder, We wood, and other human activities.

Distribution.—Sikkim (Rangit Valley), West Bengal (Darjeeling district).

Population mapping and status.—Only three isolated populations of B. satrapis around Sumbuk (lat.

27°6'19"N, long. 88?22'07"E at alt. 550 m; lat. 27?06'06"N, long. 88°23'07"E at alt. 555 m) and Kitam (27°

07'42"N and 88?21'04'E at 1100 m) in South Sikkim (Fig. 1) were encountered during the study. Each of

these populations comprised 40—50 individuals and the majority of them were in flowering and fruiting

stages. However, no seedlings of the species could be located.

Threat. —We observed that the natural habitat of B. satrapis is und bi The number

and geographic area of the type localities of the species have been considerably rediised owing to activities

like agricultural expansion, road-widening, and fodder collection by the local population.

rocky forest floors near the forest fringes and dthe

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Collection history.—Begonia satrapis has not been collected from the Rangit hills since 1910. There are

only two herbarium sheets deposited in the Central National Herbarium (CAL) which were collected by

Ribu and Rhomo (Griffith Herbarium), from Rangit, dated October 7, 1910 and these are without flowers.

Biswas (1966) reported the occurrence of B. satrapis from the adjoining (Darjeeling district), West Bengal,

however, no voucher specimens were found at CAL and SHC herbaria for reference.

Begonia scutata Wall., Prodr. 15(1):328.1864; Clarke, in Fl. Brit. India 2:642. 1879.

Small, glabrous herbs with a very short tuberous rootstock. Stem erect, filiform, 7-14 cm high, leafless in

fruiting stage. Leaves ovate-acute, cordate at base, irregularly serrate margins, minutely pubescent on the

nerves beneath; petiole filiform, 2—4 cm long in cauline leaf, longer up to 7 cm in radical leaves. Peduncles

5—7cm long, 2-3 flowered; bracts caducous. Male flower: pedicel about 1cm long; tepals 2+2, the outer tepals

ovate, 1 x 0. i cm, DUAE tie inner smaller, 0.5 x 0.4 cm, glabrous; stamens about 20, shortly monoa-

delphous; B fl tepals 2+1, the outer tepals rounded, 0.8 cm in diameter, the inner

one seules 0.4 cm wide; styles 3, distinct, shortly bifid above. Fruit: capsules, unequally 3-winged, the

larger wing 0.9 cm wide, smaller wing about 0.2 cm wide. Seeds ellipsoid.

Habitat. —Three populations of B. scutata were located near Yangyang in Teesta Valley and Legship in

Rangit Valley (South Sikkim) at altitudes of 300—1200 m. These populations inhabited open, tropical, and

sub-tropical mixed deciduous secondary forests. The surrounding vegetation in the habitat comprised of

trees like Bombax ceiba L., Canarium strictum Roxb., Engelhardtia spicata Blume, Erythrina arborescens Roxb.,

Garuga pinnata Roxb., Gynocardia odorata R. Br., Schima wallichii Choisy and Toona ciliata M. Roem. The main

herbaceous associates of B. scutata include Begonia picta Sm., Cissus repens Lam., Didymocarpus pedicellatus

R. Br., Pilea scripta (D. Don) Wedd., Persicaria capitata (D. Don) H. Gross, Urtica dioica L. and Selaginella

aitchisonii Hieron. Begonia scutata grows on sandy, wet soils, and rocky strata.

Distribution.—Himalaya: Sikkim, West Bengal (Darjeeling); Nepal; Peninsular India.

Population mapping and status.—Only three populations of B. scutata around Legship (lat. 2/?1625"N,

long. 88°16'49"E at 300 m alt.; lat. 27916'28.27"N, long. 88°16'48"E at 400 m alt.) and Yangyang (lat. 27°16!

49"N, long. 88°24'45"E at 1100 m alt.) could be observed in South Sikkim (Fig. 1). Each of these populations

comprised 10—15 individuals and the majority of these were in flowering/fruiting stages.

Threat.—The vast forested tracts in the distribution range of the species have already been cleared for

agriculture and road construction purposes. The habitats are being continuously disturbed and degraded

owing to various activities being undertaken in those areas such as road construction, sand mining, and

agricultural expansion.

Collection history.—This species was first collected by Wallich in 1821 and its type specimen is avail-

able in CAL. It was later reported from Eastern Himalaya (Sikkim and Darjeeling) by Biswas (1966) and

Grierson and Long (1991). Apart from the type (Wallich 1821) only a few other specimens without flowers

are available in CAL.

DISCUSSION

trapi dB tat listed ies in the Red Data Book of Indian Plants (Nayar & Sastry

199 0). Begora satrapis is a very rare plant oe was reported to occur only in Rangit valley in Sikkim Hima-

laya (Hooker 1879). The CAL specimens of B. satrapis—collected by Ribu and Rhomo, G.H. Cave and 1.H.

Burkil during 1875 to 1914 from the hills of Rangit—are fairly old and do not represent adequate material

for its proper identity. Though, Biswas (1966) reported the occurrence of the species from Darjeeling, no

specimen of his collections was found at either CAL or SHC. Reports of Grierson and Long (1991) suggest

presence of the species in Darjeeling and Sikkim Himalaya. Apart from the type and a few specimens of B.

satrapis by Ribu and Rhomo (1910), available at CAL, no other specimen of this species has been preserved

anywhere in the Indian herbaria. Therefore, the recent collection of these species by the aut! fter a long

gap is noteworthy.

Nautiyal etal., R i fB ia in Indi 829

LI -

Begonia scutata was first collected by Wallich in 1821 and described by De Candole in 1864 from Nepal.

This species was later reported from Sikkim and Darjeeling Himalaya (Biswas 1966) and Nepal Himalaya

(Hara 1966). A collection of tl ecies from locality in Peninsular India by Kumar and Banerjee (1989)

could not be verified as the specimen of their collection was not found at CAL. Grierson and Long (1991)

have mecoraed this species near Beater in Darjeeling Himalaya and described it as B. rubella. However, our

evaluation of the d descriptions reveal that B. rubella and B. scutata are two distinct

species (see also Nayar & Bastiy 1990). Apart from the type specimen of Wallich (CAL), no other authentic

specimen of B. scutata was located in other Indian herbaria.

The populations of B. satrapis and B. scutata in South Sikkim were mapped and various threats faced

by these species were assessed. It was observed that the population sizes of the two species are small and

their habitats face severe pressure from various anthropogenic activities including road-widening, tree fell-

ing, expansion of agricultural fields, grazing, etc. In a recent work, Pandit et al. (2007) have reported that

large scale species extinction is likely to occur in Himalaya following deforestation and continuing land

use changes. Studies have shown that reduction in the population size occurs due to the human activities

in their habitats which lead to the reduction of genetic diversity and rarity of the species (Godt et al. 1998;

Di Battista 2007). The present nd uds that Begonia satrapis is represented by merely three scattered

populations of about 40-50 individuals dispersed over an area of 2-3 sq km each. Additionally, regular col-

lection of fodder and fuel ao) Eon the forest floors by humans seems to be a serious threat to the survival

of this species in its natural habitat.

Begonia scutata is represented by only three populations comprising about 10-15 individuals each.

These populations survive in the degraded forests along river banks, which are threatened by activities

like road construction, sand mining, and agricultural expansion. These habitats are the last bastions of its

survival and the species may be consigned to extinction once these habitats are lost. The two species of

Begonia discussed here are faced with probable extinction. To save these species from being lost efforts are

under way to locate more populations of these taxa, multiply them, and re-introduce them in their natural

habitats. As such, there is an urgent need for carrying out detailed biological and ecological studies on these

species for their proper conservation. To begin with the habitats of these species need to be brought under

protective regulation before these species are ae forever. We also recommend that if activities such as road

extension, agricultural expansion in these habitats, the last remaining populations of these

species must be rescued and rehabilitated to safer habitats within the region.

Ex-situ conservation strategies

Re-establishing wild populations of B. satrapis and B. scutata in suitable habitats that do not have severe

habitat pressure can prove to he an dun aid means dh ex-situ conservation. Since both the O es of Begonia

AS ti g these

CQ4 7200

have limited number ofi

species in nearby botanic gardens and parks shouid be aadorin. Fa a long-term conservation program,

establishing seed banks will be essential.

ACKNOWLEDGMENTS

The authors are thankful to H.M. Mukherjee, Central National Herbarium, Kolkata (CAL) for permission

and help with the herbarium. The financial support for the research project sanctioned to M.K. Pandit vide

project no. J.12011/11/99-IA.I. from the Ministry of Environment, Forests and Wildlife, Government of India

and National Hydro-electric Power Corporation, Faridabad, is gratefully acknowledged. Two anonymous

reviewers kindly reviewed and thus improved the manuscript.

REFERENCES

Biswas, K.P. 1966. Plants of Darjeeling and Sikkim Himalaya, Vol. |. Kolkata. P. 377,

CismHE. 2007. Carrying capacity studies of Teesta basin in Sikkim, Vol. IV, Ministry of Environment & Forests, Govt.

of India, New Delhi.

£ al Ds LI Im | I] d len die e F da h

830 J | f Texas 3(2)

De CanpoLte, A. 1864, Reg. Veg. Prod. 153:328.

Di Barısta, J.D. 2007. Patterns of genetic variation in anthropogenically impacted populations. Cons. Genet.

(online first, April 12, 2007).

Grierson, A.J.C. AND D.G. Lone. 1991. Flora of Bhutan, Vol. 2, Part 1. Royal Botanic Garden, Edinburgh. Pp.

241-242.

Gopr, M.J.W., B.R. JOHNSON, AND J.L. FLOURICH. 1998. Genetic diversity and population size in four rare Southern Ap-

palachian plant species. Conservation Biol. 10:796-805.

Hara, H. 1966. The flora of Eastern Himalaya: results of botanical expedition of eastern Himalaya. Tokyo. Pp.

1-215.

Hooker, J.D. 1879. Flora British India. Vol. Il. L Reeve & Co. London. Pp. 638-642.

Kumar, K.D. AND R.N. Banerjee 1989. A note on Begonia scutata Wall. ex DC. (Begoniaceae). J. Econ. Tax. Bot.

13:247-248.

Myers, N., R.A. Mitrermeler, C.G. MirreRMEIER, G.A.B. DA FONSECA, AND J. KENT. 2000. Biodiversity hotspots for conservation

priorities. Nature 403:853-858.

Nayar, M.P. AND A.R.K. Sastry. 1987. Red Data Book of Indian Plants. Vol. |. Botanical Survey of India, Kolkata.

Nayar, M.P AND A.R.K. Sastry. 1988. Red Data Book of Indian Plants. Vol. Il. Botanical Survey of India. Kolkata.

Nayar, M.P. AND A.R.K. Sastry. 1990. Red Data Book of Indian Plants. Vol. Ill. Botanical Survey of India, Kolkata.

Pano, M.K. AND C.R. Basu. 1998. Biology and Conservation of Coptis teeta—an endemic and endangered medicinal

herb of Eastern Himalaya. Environm. Conservation 25:262-272.

PanpiT, M.K. AND C.R. Basu. 2003. The effects of loss of sex in clonal populations of an endangered | ial Coptis

teeta (Ranunculaceae). Bot. J. Linn. Soc. 143:47-54.

PANDIT, M.K., N.S. Sons, L.P. Koh, A. Bhaskar, AND B.W. Brook. 2007. Unreported yet massive deforestation driving loss

of endemic biodiversity in Indian Himalaya. Biodivers. Conservation 16:153-163.

SINGH, P. AND A.S. CHAUHAN. 1998. An overview of plant diversity of Sikkim. In: S.C. Rai, R.C. Sundriyal, and E. Sharma,

eds. Sikkim Perspectives for Planning €: Development. Sikkim Science Society, Sikkim. Pp. 219-233,

STATUS ASSESSMENT OF ELEOCHARIS WOLFII (CYPERACEAE)

IN THE UNITED STATES

Paul M. McKenzie C. Theo Witsell Loy R. Phillippe

U.S. Fish and Wildlife Service’ Arkansas Natural Heritage Commission Illinois Natural History Survey

101 Park DeVille Dr; Suite A 1500 Tower Building, 323 Center St. 607 East Pea

Columbia, Missouri 65203, U.S.A. Little Rock, Arkansas 72201, U.S.A. Champaign, Illinois 18820, U.S.A.

paul_mckenzie@fws.gov theo@arkansasheritage.org rickp@inhs.uiuc.edu

Christopher S. Reid Michael A. Homoya

Louisiana Natural Heritage Program Division of Nature Preserves

Louisiana Department of Wildlife & Fisheries Indiana Department of Natural Resources

2000 Quail Dr. 402 West Washington St; Room 267

Baton Rouge, Louisiana 70898, U.S.A. Indianapolis, Indiana 46204, U.S.A.

creidawlf.louisiana.gov mhomoyaqQgd4ánr.IN.gov

Steven B. Rolfsmeier Caleb A. Morse

Kansas State University Herbarium R.L. McGregor Herbarium

Manhattan, Kansas 66506, U.S.A. University of Kansas Campus West

srolfsm2@yahoo.com 2045 Constant Ave

Lawrence, Kansas 66047, U.S.A.

@ku.edu

ABSTRACT

r1 1 . VPF AV 21 aa es 1 by AT [eH S37 A pa | cl ( re I D p -1 d), S2 (imperiled), SNR SU ( 1 ]

or currently imer review). or SH à istorical) t states wl the sr has | documen ted. R t field studies in Arkansas,

Illinois, Kansas, Louisiana, Missouri, and Nebrasl ggest that the sy t li 11 ked

Pre-1978, tl tesin 43 i K ishesin 17 states and MEN two atic Post-1978, Wolf S pe ud

mari tt 16 states. Since 2000, 135 new sites distributed among

has been documented at 222 sites from 104

64 counties/parishes in 15 states have Dee discovered an ii is € po in the field by the combination of tiny

rhizomes; purple-red culm bases; blue-green, f Í tly iin achenes « " E mm long with 9-19

longitudinal ridges and 30— de transverse becas T pecies is | f wide va I g wet depressions

J a : : Beak al 1 1 1 =

s = nd p p I pen grasslands, limestone and saline barrens, I I

haliow i dstone, granite, tzit t El i H r ii 1 1 "m

E L in J I

is often found with a wide var iety f pl t lat d p di g il d t ] ity type. Significant threats to Eleocharis

fi: 1 Ls : 1 E: J J k| mi r 1 1 l Er 1 q S 1 " Ec ] - i s] ]

I le] a 4 F

T J ] 1l [ + : Te ] ra 1 € 113 ] «L1 1 FN +] i |

F , f 8 ps, Į y There are

ffici lat 1 j inr l i to p id y mea e EE species. The potential = of lanes

1 n] J: E 1 1

ey negatively A

climate uns. on E.| fii i

ditional a are needed in E oe n Great Plains states, and states bordering or adjacent to the Great Lakes. The species

s ura an mn ice sped Mond a pemet dues

would b

. sf. . E 1 J

L , F F ee F E I & b

Lie a E a E :] 1. 1 1 -1 JE 1 J at 1 x c ] dias 1 1

benefit P L ’ yg 45 [e] i E u

] ] ru 335 T + 1 1 r1 m n ——

and low g l growth

r1 1 fee 1 3 IAT C Panis Fan C1 f a PP Pa 2X c (amenazada) CNIT CT 7 f. tal E

H + pi V ey S Za X b

" " P CTT 7? 21.1] i . : "HS J 1 4 ] pj A 1 Š 3 v]. —€————— e el

1 y 1 AN ii h dad da. Re-

EF E

i tudios d E Arkansas, Illinois, Kansas, Louisiana, Missouri, y Nebrasl gi jue la especi t

1 : Fd A sal Az me

I I p Antes de 1978, I lugares en “To cof ndados/

1] re , Eal He PLL unite. E

J. Bot. Res. Inst. Texas 3(2): 831 — 854. 2009

832 tani titute of Texas 3(2)

parroquias de 17 estados y posiblemente otros dos. Después de1978, se ha documentado en 222 sitios de 104 condados/parroquias

distribuidos por 16 estados. Desde 2000, se han ld MUN 135 nuevas localidades distribuid 64 condados/parroquias en 15

stados. Eleocha If o por la a de rizomas finos; bases b bn oe inp PORUM

culmos verde- azulados, plastados y idos espiral yaq < 1.0 le 1

1 3 J 1; cee uo: es | q A : 1 J J x Ls

y 30 I H P L Pi I

das de fondo de valle y mésicas de ti ltas, cl les el prad biertas, baldi ] y salinos, y n" temporales

afloramientos de areniscas, , granitos, o cuarcitas. Eleocharis wolfii pued trarse casi en cultivos puros

| + 1.3.1 1 DES | 3 1

Į Existen

| bre el hábitat de Eleocharis wolfii en gran poe de su distribución que Belen la tala de bosques; destruc-

ción del habitat por el ae ans y residencial; "s conversión de praderas ind: en tierras de is pastos o prados; y

la owe i M ion es. Hay g la mayoría d para proteg especie. El impacto

TA tod 1 E Wes ceux. T 1 ] x 4 1 J nA 2] " ]

I I 4 + r i

pueden af gati te a la especie. 5 it tudios adicional l Medio Oeste, estados de las grandes llanuras, y

1 1 3] l4 ] x^ ] ] 1 r. J EL : Joas 1 : 1 fay E pa | em |

o L L +

vital y ecología; viabilidad de las semillas; "s tica de poblaci imientos para la germinación;

cambios en el uso del territorio; y los impactos potenciales de especies invasoras. Las acciones de gestión que pueden bene a E.

bd p

wolfii ae la prohibición del fuego, siega o p R leñosa en los hábitats de pradera, y

1 : à 1 1. Lx As] 1 f, 1 : x A 11 Asl

i a +

INTRODUCTION

1 M: ERA eee E

Eleocharis R. Br genus of Cy consis y 200 species worldwide.

Smith (2002) eae 67 species of CMM in North American noh of Mies No recent compre-

hensive worldwide taxonomic treatment is available. González-Elizondo and Peterson (1997) recognized

four subgenera in a worldwide ific classification of Eleocharis: subgenus Eleocharis, subgenus Lim-

nochloa (P. Beauv. ex T. Lestib.) Torr, — Scirpidium (Nees) Kukkonen and subgenus Zinserlingia TV.

Egorova. Most North American treatments of Eleocharis rely heavily on the extensive studies by Svenson

(1929, 1932, 1934, 1937, 1939, 1957) and they include difficult species complexes that need taxonomic

revision (Smith 2002).

Eleocharis wolfii (A. Gray) A. Gray ex Britton in Patt. (Wolf's spike-rush) is native but scattered and

local from North Dakota south to Texas and east to Georgia, Virginia (Godfrey & Wooten 1979; Kolstad

1986; Robertson & Phillippe 1992; Yatskievych 1999; Smith 2002) and New York where it is considered by

some to be adventive (e. B. Svenson 1957). Galen Smith could not locate substantiate previous

reports of the species from Alberta, Saskatcl , and Colorado (Smith 2002). E We wolfii is a member

of subgenus Scirpidium. Members of this bubaenis are M MEME oe having 3-fid ae and diagnostic

cuol cellate cells or fine horizontal

trigonous to nearly terete achenes with distinct t

ridges (trabeculae) between prominent longitudinal ridges or achane angles (Smith 2002). Smith (2002)

recognized seven species within subgenus Scirpidium north of Mexico including E. wolfti. Wolf's spike-rush

is named for John Wolf who collected the type specimen in Fulton County, Illinois (Gray 1874; Mohlenbrock

1976; Phillippe 2005).

Botanical nomenclature listed herein follows Yastkievych (1999) or Yatskievych and Turner (1990),

except for Carex aureolensis, C. opaca, Crataegus brachyacantha, Dichanthelium dichotomum ssp. dichotomum,

D. scoparium, Isolepis pseudosetacea, Juncus coriaceus, Ludwigia linearis, L. palustris, Quercus similis, Ranunculus

laxicaulis, Rhynchospora globularis var. globularis, R. recognita and Steinchisma hians which follow Arkansas

Vascular Flora Committee (2006); Eleocharis coloradoensis, E. elliptica, E. montevidensis and E. tenuis which

follow Smith (2002); Hierochloé odorata, Polygonum ao m Rhamnus frangula which follow Swink and

Wilhelm (1994); Bidens tricl , Boltonia diffusa, Euth ides, E. leptocephala, Doellingeria

ampenata, ia il la annua, gana Symphyotrichum loncedlatum which follow Yatskievych ads

and Eriopl which follow Kolstad (1986). Botanical

the mosses imac and Daum follows United States Department of Agriculture, National Resources

Conservation Service (2009). Author citations and abbreviations follow Brummitt and Powell (1992).

McKenzie et al., St i £ Elanrchar "TP -

Plant description (from Smith 2002)

Plants perennial, rhizomatous, may grow in small circular clumps (tussocks) or often coalescing to form

large mats (Fig. la—d); light brown to white rhizomes 0.25-0.6 mm thick (Fig. 2a), internodes 1-4 cm,

the stems in erect or decumbent tufts or clumps along the rhizome (Fig. 2a), scales 2 mm. Culms erect

(sometimes decumbent when dry), sides variably smooth or with 1 to few acute ridges (often nearly smooth

or with 1 ridge on 1 side and several ridges on the other), greatly compressed (Fig. le-f € Fig. 2b), usually

inrolled when dry, rectangular in cross section, 8 to 50 cm x 0.3-1.5 mm, 0.2-0.5 mm thick, firm, margins

often sharply acute, margins and often 1 or more ridges minutely serrulate at 20-30 x. Leaves: leaf sheaths

persistent, dark red, brown, or straw-colored proximally (Fig 2a), terminating at the acute apex as a thin,

hyaline tooth that may be split and free from the culm (slightly inflated). Spikelets ovoid or lanceoloid,

3-9 x 1.5-2.5 mm, apex acute (Fig. 2b); 15-30 imbricate, ovate floral scales with acute apexes (Fig. 2b,d),

scales dark red, orange-brown, stramineous, or colorless, and generally with broad hyaline margins and

tip, the prominent midrib stramineous , brown, or greenish, ovate-lanceolate, (2.2—)2.7-3.2 x 1.5 mm (Fig.

2d). The 2 lower basal scales usually larger. Flowers: perianth bristles absent; anthers 3, 1.1-1.75 mm, style

trifid, wind pollinated. Achenes brown, gray, or nearly white, lustrous, compressed-trigonous, obovoid,

mostly 2 times longer than wide, 0.7-0.9(—1.1) x (C0.4)0.5 mm, with angles and 9-19 longitudinal ridges

and 30—60 transverse trabeculae (Fig. 2c). Tubercles brownish, pyramidal, 0.1-0.15 x 0.2-0.25 mm (Fig

3a, 3b). Fruiting late spring-early summer (May-June). Although not mentioned in most ref outside

of McKenzie and Jacobs (2000), Robertson and Phillippe (1992), and Phillippe (2005), the culms of E. wolfii

are usually bluish-green and conspicuously twisted in a diagnostic spiral (Fig. lef). Based on personal

experience of the authors, the bluish-green and spirally twisted culms of Wolf's spike-rush provides for

immediate recognition in the field, especially PASE aeoea with other species of Eleocharis spp.

Eleocharis wolfii most closely bl e fused with Eleocharis acicularis (L.) Roem.

& Schult., especially var. porcata S.G. Sm. Smith (2001a) described variety porcata based on material that

was collected mostly from the Great Plains. Due to the need to revise the taxonomy of subgenus Scirpidium

on a global scale, and the variability associated with E. acicularis sensu lato, Smith (2002) did not formally

recognize var. porcata in his Flora of North America treatment. Smith (20012) provided the specific epithet

porcata due to the presence of 6-12 prominent ridges on the culms, but also noted that this variety has

flattened stems, and is therefore similar to, and possibly confused with, E. wolfii. Eleocharis wolfti can be

distinguished from E. acicularis var. porcata by the combination of twisted, blue-green culms (Fig. le—f)

with fewer (1-3) sharp ridges and usually longer spikelet scales [(2.2—)2.7-3.2 mm] (Smith 2002). Eleocharis

acicularis var. porcata has dark green culms with 6-12 blunt ridges and shorter spikelet scales [(1.52—2.5

mm] (Smith 2001a, 2002). Eleocharis wolfti also usually has wider culms (0.3-1.5 mm) than E. acicularis

var. porcata (0.2—0.5 mm). The generally narrower culms of E. acicularis give this species a wiry appear-

ance (pers. obs.). Additionally, E. acicularis usually has shorter culms that form dense, low growing, often

sterile mats that superficially resemble sterile clumps of E. parvula (Roem. & Schult.) Link ex Bluffet al.

(Mohlenbrock 1976), E. coloradoensis (Britton) Gilly, or the small fern Pilularia americana A. Braun, species

with which it times associates (pers. obs.). Smith (2001a, 2002) reported that E. acicularis var. porcata

has been misidentified as E. wolfii at some localities. Although confusion between the two taxa could occur

where the two taxa are sympatric (Smith 20012), the characters identified above should readily separate E.

wolfii from E. acicularis var. porcata, especially for culms of E. wolfii that are > 0.8 mm wide. Finally, E. wolfii

is typically not found in the same habitat as E. acicularis. Eleocharis acicularis is often found on the shores of

ponds, lakes, vernal pools, sloughs, ditches and frequently on disturbed habitats (Godfrey & Wooten 1979;

Yatskievych 1999; Smith 2002). As noted by Robertson and Phillippe (1992), Swink and Wilhelm (1994),

Smith (2002), Phillippe (2005), and others, E. wolfii is usually associated with native, mostly undisturbed

habitats. Eleocharis wolfii was cited as a weed in Asia and Pacific Islands by Holm et al. (1979) and Moody

(1989), respectively. There are, however, no vouchers to verify the identity of these claims (Charles Bryson,

USDA, pers. comm. 2009).

Journal of the Botanical Research Institute of Texas 3(2)

3 F

Fic. 1. ad: Clumped habit of Eleocharis wolfii (a, b, & c photos by Theo Witsell, Saline County, AR; d photo by John Pelton, Prairie County, AR); e-f:

flattened and sprially twisted culms of Eleocharis wolfi (photos by Christopher Reid, Morehouse Parish, LA).

McKenzie et al., St t £ Flanchari Sor ae ee ree m

rhizomes

uu OL

WLU G

Fic. 2. a: Purple-red culm | | thin white to tan rhi f Eleocharis wolfii (photo by Theo Witsell from Witsell #07-24 [ANHC]; Hempstead

T P

f tw AP hoeh el PT P a

county, nnj, 9 p [ i achenes,

836 J I tanical h Institute of Texas 3(2)

Based on field observations El the Rue E. wolfii produces flower and fruit for about a 3-4 week

period. After fruiting, spikes often become lated from the stem and the stems turn brown or wither

(McKenzie & Jacobs 2000). Populations of Wolf's spike-rush occurring in flatwoods under shaded conditions

may contain mostly sterile culms that lie prostrate or decumbent on the forest floor (Fig. la—c) (Robertson

& Phillippe 1992, McKenzie & Jacobs 2000, Phillippe 2005). Specimens of E. wolfii in such areas of reduced

light intensity tend to be etiolated, and produce fewer di ipud id fewer achenes per spikelet

(Robertson & Phillippe 1992) and achenes are often i ie & Jacobs 2000). Fertile

and sterile culms of E. wolfii can be identified by the combination of very narrow rhizomes, purple-red culm

bases, and conspicuously flattened and spirally twisted culms (Fig. le—f; 2a).

The habit of E. wolfii is variable depending on soil type and natural community type (see below). At

many sites, the species is often hidden among taller vegetation where it can be difficult to detect, especially

late in the season when spikelets are absent and the culms no longer exhibit their conspicuous blue-green

color (Fig. le—f). At some sites, however, the species can form large colonies that are in nearly pure stands

(Fig. la—d).

Previous evaluations by the United States Fish and Wildlife Service (USFWS)

Eleocharis wolfiii was listed as a Category 2 (C2) candidate species in the USFWS's 1993 Plant Candidate

Review for Listing as Endangered or Threatened Species (U.S. Fish & Wildlife Service 1993). Category 2

candidate species comprised taxa for which information indicated that a proposal to list as endangered or

threatened was possibly appropriate, but for which conclusive data on biological vulnerability and threats

were not currently available to support proposed rules.

On 19 Jul 1994, the Director of the USFWS issued a memorandum establishing new policy on the

definition of candidate species (formerly Category 1 or Cl) and how the USFWS would consider species

for which they remained concerned (formerly C2). Under the new policy, candidates are defined as those

species for which the USFWS has on file sufficient information on biological vulnerability and threats to

support issuance of a proposed rule to list as endangered or threatened, but issuance of the proposed rule

is precluded by other listing actions.

Former C2 species for which the USFWS lacks sufficient information to classify as candidate species

will no longer be enumerated on an official list. Nonetheless, other agencies that have developed extensive

databases on former C2 species [e.g., The Nature Conservancy (TNC) & state Natural Heritage programs]

will continue to monitor these species and maintain communication with the USFWS to further assess the

status of these species in their respective states. This information will be useful in determining when there

is sufficient information to warrant their addition to the USFWS's list of candidate pes

Eleocharis wolfti is a species the USFWS conti | 1 the lack of a ive summar

of the species’ rangewide status and distribution, coupled with the large number of a coa!

in the central E S. since ane is Pau ine si as i indes the current assessment.

In 2003,t TE. licited information f

-l

botanists throughout the ange of the species. a received a such requests is providedi in this

report. Although we could not find any published results of a rang, of ies conducted in

North America, S. Galen Smith compiled a distribution map of E. wolfti in his 2002 Flora of North America

treatment (Smith 2002). Martin (1993) drafted a status report of E. wolfii for the USFWS but the document

was never finalized (Karen Kreil, USFWS, pers. comm. 2006). Phillippe (2005) prepared a conservation

assessment for the U.S. Forest Service on E. wolfii and provided information on the species’ taxonomy, life

history, habitat requirements, ecology, range-wide distribution and abundance, conservation status, popu-

lation biology and viability and known or suspected threats. NatureServe (2009) provided the projected

current distribution of E. wolfti for North America, listed global and state ranks outlining the species global

rarity and conservation status in each state in the United States and included information on ecology and

life history, habitat requirements, and threats to the species.

Recent field surveys conducted primarily in Arkansas, Kansas, Louisiana, Missouri, and Nebraska

"Om lille elo Inita d Cond 837

McKenzie etal Stat

between 2000 and 2008 and in Indiana between 1983 and 1995 have yielded numerous new localities

for the species (Phillippe 2005). The new discoveries suggest that the species may be more common than

previously believed and simply overlooked. To obtain a more complete depiction of the current range of the

species, including recently collected data, we referenced recently published and unpublished survey results,

published accounts (Godfrey & Wooten 1979; Kolstad 1986; Robertson & Phillippe 1992; McKenzie &

Jacobs 2000; Smith 2002; Phillippe 2005; Diggs et al. 2006), as well as accounts provided by personnel of

various state Natural Heritage programs and Academia.

Distribution

A summary of historical and extant documented occurrences of E. wolfti in the United States is provided in

Table 1 and Figure 3. We define any oe eocumentcs prior to 1978 to be historical and those recorded

after that date to be extant. Historically G that have extant populations), Wolf’s spike-

rush was known from imately 59 sit ttered across 43 counties of 20 states (Table 1). Of these, 31

sites are known solely fon peni iae (Table 2; Fig. 3). Within the last 30 years, E. wolfii has been

collected at 222 sites in 104 counties of 16 states (Table 1) and over 61% of these sites (135) and 62% of the

]

counties (64) have been documented since 2000 (Table 3). Although the species has disappeared from some

historical locations due to habitat destruction, the large number of extant sit that the species was

historically widespread. The smaller number of historical records (Table 1, Table 2) is undoubtedly due, at

least in part, to the lack of surveys for E. wolfti prior to 1978 or due to the lack of field experience of many

botanists with the species. This conclusion is supported given: a) the large number of discoveries of this

species since 2000 (Table 1, Table 3), b) plants are apparently long lived and individual populations may

exist for hundreds of years (Phillippe 2005), c) the species occurs in a wide variety of habitats (see below),

and d) there is an abundance of unsurveyed habitat, it is likely that additional populations will be found

with further survey efforts.

Habitat requirements and soil types

Eleocharis wolfii has been documented from a wide NEUE of habitats including: *wet depressions of bot-

wet, open sites;" “wet river and lake margins;” aene and seeps;"

vr) tt

tomland and mesic upland prairies;

“ephemeral pools in open grasslands;

wet depressions, pond and river margins; wet sand prairies; wet meadows and other moist areas (Kolstad

1986; Godfrey & Wooten 1979; Hedge et al. 1994; Robertson & Phillippe 1992; Eilers & Roosa 1994; Swink

& Wilhelm 1994; Yatskievych 1999; Smith. 2002; Diggs et al. 2006). Eleocharis wolfii has recently been

discovered in wet areas on sandstone glades in Arkansas and Missouri and in wet saline barrens and in wet

areas within shale glades in Arkansas (Witsell & McKenzie, pers. obs., 2000—2008). In Georgia, E. wolfii is

known from ephermal pools on granite outcrops [Tom Patrick, GA Nae Heritage eee (NHP), pers.

comm. 2006] and from shallow depressions on sandstone, gneiss granite , and Si ite in Minnesota

[Fred Harris, Minnesota Department of Natural Besos (DNR), pers. comm. 2007].

We could find limited information on soil types associated with the natural communities where E.

wolfii is currently extant. Prairies in northern Illinois are open, wet and are on poorly to somewhat poorly

drained soils that are nearly level or depressional on outwash plains. These soils are often ponded in spring

and are dark gray and friable fine sand or sandy loam (Phillippe 2005). At prairie sites in southern Illinois,

the soil type is categorized as deep gray silt loam or gray silt loam on tight clay (Robertson & Phillippe

1992). River terraces of southern Illinois flatwoods are on poorly drained silt loam and silty clay loam soils

(Miles 1988; Phillippe 2005). In northern Indiana, most populations of E. wolfti occur on wet, acidic sandy

loam (Hedge et al. 1994). In southern Indiana, the species is found on slowly p ble silt loam (Aldrich

& Homoya 1984; Homoya pers. obs. 1983-1993). In Kansas, prairie soils where Wolf's spike-rush occurs

range from hard-pan clay, to shallow calcareous soils over limestone, to sandy soils over sandstone (Morse,

pers. obs.). Poorly drained sodic and non-sodic silt loam soils are characteristic of Louisiana E. wolfii sta-

tions (Edwards et al. 1991; Martin et al. 1981). A site in Juneau County, Wisconsin is described as “...moist,

limestone barrens;” “oak flatwoods or woodlands on river terraces;”

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TABLE 1, | f (

State No. of historical No. of historical No. of extant No. of extant

counties/parishes sites counties/parishes sites

AL 1 1 0 0

AR 2 2 22 54

CO 1? 1? 0 0

GA 0 0 2 4

IA 4 4 ] ]

IL 9 21 16 4/

IN 1 2 8 15

KS 1 1 6 10

LA 2 3 9 14

MN 3 3 4 5

MS 1 1 1 1

MO 2 2 18 30

NE ] ] 7 29

NY 1 1 0 0

ND 1 2 0 0

OH 2 2 2 1

OK 2 2 1 1

TN 4 6 2 2

TX 4? 27 3 6

WI 1 1 2 2

TOTAL 43 59 104 222

TABLE 2. C t /Pari

State Historical counties

AL Lauderdale

CO El Paso?

IL Fulton, LaSalle, Peoria, Perry, Stark

lA Cedar, Emmett, Union

KS erokee

LA East Baton Rouge, Ouachita

MN Nicollet, Norman, Traverse

MO Linn

MS Tishomingo

NY Nassau

OH Crawford, Ross

OK Latimer, McCurtain

TN Coffee, Franklin, Grundy, Warren

Bowie, Jefferson, Morris

r ri I . pep J: n Ll tz ] Cons A ^"Inwn

TABLE 3 | |

State No. of new counties/ No. ofnew

arishes sites

AR 19 53

GA 1 1

IA 1 1

IL 3 7

KS 5 E

LA 7 11

MN 4 5

MO 16 29

MS ] 1

NE 3 10

OH 1 1

OK 1 1

TN 0 1

TX [ 4

WI 1 1

TOTAL 64 135

slightly peaty, dark gray fine sandy loam, prob, slightly acid” (http://www. botany.wisc.edu/cgi-bin/st

cgi?Accession-v0008579WIS)

Associated species

Due to the wide range of habitats recorded for E. wolfii, the list of plant species iated with the species is

extensive and there are data from most states a its Ens At as puse Muenchh. and Q. stellata

Wan genh flat ds sites in southern I tens (E.P. Bicknell) E.P.

Bicknell var. xanthocarpa (E.P. Bicknell) Wiegand, C. bicknellii Britton, C. caroliniana Schwein: C. ae

L., Cinna arundinacea L., Eleocharis verrucosa (Svenson) L.J. Harms, Galium obtusum Bigel.,

Gay € Dur. and Q. bicolor Willd. (Phillippe 2005).

In the wet sand prairies of northern Illinois, E. wolfti is associated with Agalinis purpurea (L.) Pennell,

Andropogon gerardii Vitman, Carex buxbaumii Wahl, C. haydenii Dewey, C. stricta Lam., C. suberecta (Olney)

Britt., Eleocharis verrucosa, Eupatorium perfoliatum L., Hypericum adpressum Bart., Hypericum sphaerocarpum

Michx., Leersia oryzoides (L.) Swartz, Osmunda Eu A "dad laeve nca ls paa > ) Timok,

var. herbiola (R. Br.) Luer, Polygonum careyi Olney, P hx., )

Vahl, R hneideri Bailey, Sisyrinchium atlanticum ME P. Bicknell, Verbena hastata L., Viola lanceolata L., and

Viola His i L. (Phillippe 2005).

in northern Indiana, associates listed include Alisma sp., Carex lasiocarpa Ehrhart, Cype-

rus un Torr., C. strigosus L., Drosera sp., Echinochloa crus-galli (L.) P. Beauv., Eleocharis elliptica Kunth,

E. ovata (Roth.) Roem. & Schult., E. palustris L., Galium obtusum, Juncus acuminatus Michx., J. brachycarpus

Engelm., J. dudleyi Wiegand, Lysimachia quadrifolia Sims, Onoclea sensibilis L., Panicum rigidulum Bosc ex

Nees, Poa pratensis L., Polygala sanguinea L., Rotala ramosior (L.) Koehne, Spartina pectinata Link, Spirea alba

Duroi, and Viola lanceolata (Robertson & Phillippe 1992: Appendix 1; Phillippe 2005: Appendix 2).

Swink and Wilhelm (1994 also listed the evens associates for sites in northeastern Illinois and

| Indiana: Agrostis gigantea Roth., Bid (Michx.) Britton, Calamagrostis canadensis

(Michx.) P. Beauv., Carex (ORIG Schkuhr ex x Willd., C. pellita Muhl. ex Willd., Cornus amomum Mill. ssp.

obliqua (Raf.) J.S. Wilson, Doellingeria umbellata (Mill.) Nees, Eleocharis erythropoda Steud., Erigeron philadel-

phicus L., Eryngium yuccifolium Michx., Euthamia gymnospermoides Greene, Hierochloé odorata (L.) P. Beauv.,

"——"——— 841

McKenzie et al l Stat

Hedyotis caerulea (L.) Hook., Hypericum mutilum L., Hypoxis hirsuta (L.) Cov., Iris virginica L. var. shrevei

(Small) E.S. Anderson, Juncus marginatus Rostkov, J. tenuis T Lechea tenuifolia Michx., Lobelia ho

Lam., Lycopus rubellus Moench, Panicum virgatum L., Populus deltoides Bartram ex Marshall, R

L., Rubus flagellaris Willd., Salix humilis Marshall. Saxifraga pensylvanica L., Thelypteris palustris Scher var.

M MM (G. Lawson) Fernald and Veronica eue L.

In flatwoods in sout? Indiana,

Eleocharis tenuis (Willd.) Schult., Isoëtes melanopoda, Ludwigia palustris (L.) Elliott, Quercus bicolor, Q. —

Raf., Q. palustris Q. stellata, and Ranunculus pusillus Poiret (Aldrich & Homoya 1994).

In Kansas, E. wolfti has been observed with Carex annectens, C. festucacea Schkuhr ex Willd., C. frankii

Kunth, C. granularis Muhl. ex Willd., C. missouriensis P. Rothr. & Reznicek, C. scoparia Schkuhr ex Willd.,

Eleocharis erythropoda, E. macrostachya Britton, Glyceria striata (Lam.) Hitchc., Juncus acuminatus, J. interior

Wiegand, and Elymus smithii (Rydb.) Gould in swales of native tall grass prairie and in disturbed wet prairies.

In the floodplain of the Marais des Cygnes River in Linn County, the species was observed along the edge

ofa d Bn to a pin oak-hickory-pecan forest remnant (Morse, pers. obs.).

ies at Missouri flat 1 sites include Carex scoparia, C. triangularis Boeck., C. tribuloides

Wahlenb. “Eleocharis verrucosa, Isoétes melanopoda, Juncus spp., Ludwipis peploides (Kunth) Raven, Steinchisma

hians (Elliott) Nash, and Polygonum spp. (McKenzie & Jacobs 2000). At Missouri prairie swale sites, E. wolfi

is often associated with Carex annectens, Carex bicknellii, C. oklahomensis Mack., C. tribuloides, Dichanthelium

scoparium (Lam.) Gould, Eleocharis palustris, E. verrucosa, Glyceria striata, Juncus spp., Muhlenbergia spp.,

Penstemon tubiflorus Nutt., Physostegia virginiana (L.) Benth., Scirpus atrovirens Willd., S. pendulus Muhl. ex

EM Beaman nion (^. BM A. Gray and Tripsacum dactyloides (L.) L. (McKenzie & Jacobs 2000).

ther in Missouri include Carex conoidea, C. granularis, C. trichocarpa Muhl.

ex Willd. , Carex vesicaria L var. monile (Tucker.) Fernald, Carex vulpinoidea Michx., Lysimachia sp., Lythrum

alatum push: Platanthera lacera (Michx.) G. Don, Polygonum amphibium L., Scutellaria galericulata L., and

Spartina pectinata (pers. obs.).

Nebraska, associates recorded for E. wolfti include Agrostis gigantea, Alisma triviale Pursh, Apocynum

cannabinum L., Carex atherodes Sprengel, C. buxbaumii, C. emoryi Dewey, C. interior L.H. Bailey, C. pellita, C.

sartwellii Dewey, C. scoparia, Dichanthelium acuminatum (Sw.) Gould & C.A. Clark, Dulichium arundinaceum

(L) Britton, Eleocharis acicularis, E. elliptica, E. erythropoda, E. palustris, Equisetum arvense L., Eriophorum

polystachion L., Euthamia gymnospermoides, Juncus marginatus, J. torreyi Cov., Lysimachia hybrida Michx., L.

thyrsiflora L., Lycopus americanus Muhl., Phalaris arundinacea L., Polygonum amphibium, Pycnantheum virgin-

ianum (L.) T. Durand & B.D. Jackson ex Rob & Fernald, n eriocephala M MUR, ¿> ave Diis = hee

de Carex annectens, C. muskingumensis Schwein.,

S. petiolaris Smith, Sorghastrum nutans (L.) Nash, S

Nesom, Spiranthes cernua (L.) Rich., Thelypteris wm Schott var. pubescens (G. Lawson) Fernald, pm

maritima L. var. elata (Nutt.) A. Gray, Iypha angustifolia L., and Verbena hastata L. (Robert Steinauer, pers.

comm. 2007).

In Arkansas, E. wolfii occurs with E. verrucosa, Isoétes melanopoda, Juncus spp., Rhynchospora spp.,

Steinchisma hians on saline barrens; Carex albolutescens Schwein., C. oklahomensis, C. opaca (FJ. Herm.) P.E.

Rothr. & Reznicek, Dichanthelium dichotomum (L.) Gould subsp. roanokense (Ashe) Freckmann & Lelong,

and Ranunculus laxicaulis Darby in prairie swales; Allium canadense L. var. mobilense (Regel) Ownbey, E.

lanceolota Fernald, E. verrucosa, Isoétes butleri Engelm., I. melanopoda, Juncus secundus P. Beauv. ex Poir., Lud-

wigia palustris, Sporobolus ozarkanus Fernald, and Steinchisma hians on shale Dartens, Beochan nes E

verrucosa, Eriocaulon koernickianum Van Heurck & Müll. Arg. in Van Heurck, Isol

Gand., Juncus secundus, Rhynchospora recognita (Gale) Kral and Utricularia subulata L. on sandstone sine

Carex albolutescens, C. complanta Torr. & Hook., C. debilis Michx. var. debilis, C. flaccosperma Dewey, Fraxinus

pennsylvanica Marshall, Juncus coriaceus Mack., Leersia virginica Me a styraciflua L., Quercus

lyrata Walter, and Q. phellos L. in flatwoods; and Cyperus spp., El pp, and Fimbrisylis autumnalis (L.)

Roem. & Schult. on a wet salt lick (Witsell, pers. obs.).

842

Associates listed at recently discovered sites in Minnesota include Alopecurus carolinianus Walter, Eleo-

charis acicularis, E. compressa Sullivant, E. ovata, Hordeum jubatum L., and Juncus interior (Fred Harris, MN

DNR, pers. comm. 2007).

Associates of E. wolfii in small depressions of sodic flat d 1 prairies in Louisiana i

similis Ashe and Crataegus brachyacantha Sarg. & Engelm. in the flatwoods overstory, with herbaceous as-

sociates consisting of Carex annectens, C. aureolensis Steud., C. complanata, C. reniformis (L.H. Bailey) Small,

Cyperus pseudovegetus Steud., Dichanthelium scoparium, Eleocharis verrucosa, paii ec Juncus spp.,

Iva annua L., Ludwigia linearis Walter, L. glandulosa Walter, Pani lata (Lam.)

A. Gray, R. globularis (Chapm.) Small var. globularis, Steinchisma TM and eee cines (Nutt.) Nash. Non-

sodic poorly drained flatwoods supporting E. wolfii have Quercus phellos and Fraxinus pennsylvanica in the

overstory with an understory supporting Boltonia diffusa Elliott, Campsis radicans (L.) Bureau, Carex compla-

nata, C. lurida Wahlenb., C. triangularis, Dichanthelium dichotomum var. dichotomum, D. scoparium, Euthamia

leptocephala (Tor. & A. Gray) Greene ex Porter & Britton, Helenium flexuosum Raf., Juncus coriaceus Mack.,

and Steinchisma hians. The moss genus Climacium F. Weber & D. Mohr is often characteristic of shaded sites

supporting E. wolfii in Louisiana and Reid has used the presence of this moss with success to locate E. wolfi.

Sphagnum L. is also an occasional associate in Louisiana.

taAa yiceeepc

Population estimates

Other than very rough estimates, there are limited detailed demographic data available on Wolf's spike-

rush at most sites in North America. Robertson and Phillippe (1992), however, closely examined two large

populations of E. wolfii in southern Illinois (one from a prairie swale habitat and the second from a Quercus

spp. flatwoods) and counted the number of tufts or clumps, the number of stems/tuft, and fruiting stems

within eight randomly selected 4 m? circular plots. They also counted spikelet scales and achenes from 50

randomly selected spikelets and obtained an estimate on fruit set. The total area of the prairie population

was 321.4 m? and that of the flatwoods population was about 3,380 m". Measurements from the prairie

population plots yielded 1,108 tufts and 5,059 stems, of which 1,807 (3696) were fruiting. By extrapola-

tion, this population contained 178,078 tufts, 813,082 stems and 290,421 fruiting culms. The 50 spikelets

produced an average of 22 scales and 10 achenes per spikelet. This yielded a total estimate of 6,389,262

scales, about 2,904,210 achenes, and a fruit set of approximately 4596. Similar procedures were performed

for the flatwoods site except that 100 spikelets were collected to calculate fruit set data. The 100 spikelets

produced an average of 25 scales and 14 achenes per spikelet. This yielded a total estimate of 13,308,750

scales, about 7,452,900 achenes, and a fruit set of approximately 5696.

In a second population examined in Washington County (Robertson & Phillippe 1992: collection-

Phillippe & Gehlhausen 20311), results were P different. This population was located at an

ephemeral pool surrounded by 9 l T ds. The Q. stellata crowns nearly covered

the ephemeral pool such that the E. wolfii bosulstibn had low available light. The population was in an area

of 122.6 m? and 3396 of the site was surveyed. Robertson and Phillippe (1992) extrapolated that the total

area had 1,167 tufts with a total of 5,400 stems, of which only 228 (496) of the stems were fruiting. Rough

estimates at other localities have been reported as few as 25 culms and as high as hundreds of thousands of

plants (Robertson & Phillippe 1992; McKenzie & Jacobs 2000; Fred Harris, MN DNR, pers. comm. 2006).

We could not find any gray or published literature on seed viability, predation, or germination requirements

for Wolf's spike-rush.

Conservation status

Phillippe (2005) conducted a conservation assessment on tl ies for the Eastern Region of the U.S. Forest

Service but his report only covered Illinois and Indiana. NatureServe (2009) listed global and state ranks

outlining the global rarity and conservation status of E. wolfii in each state in the United States. Eleocharis

wolfii currently has a global rank of G3G4 (NatureServe 2009). A G3 ranking indicates that a species is

vulnerable globally due to a limited distribution and potential threats to its continued existence. A species

McKenzie et al., St i fElanrhari ao E SR E TA -

ranked G4 is one that is generally 1 widespread but apparently secure. The GRANK for E.wolfii

was reviewed in 2006 in light of recent discoveries outlined in this report (Leah Oliver, NatureServe, pers.

comm. 2006). NatureServe (2006) SRANK categories included SH (historical with the expectation that it

still may be extant and possibly rediscovered), S1 (critically imperiled), S2 (imperiled), S3 (vulnerable), 54

(apparently secure), and SNR/SU (not ranked/under review). States and NatureServe (2009) SRANKs listed

for Wolf's spike-rush are: Alabama (S1), Arkansas (S3), Georgia (SD, Illinois (SD), Indiana (S2), Iowa (S1),

Kansas (S2), Louisiana (S3), Minnesota (S1), Missouri (S354), Mississippi (SU/NR), Nebraska ($2), North

Dakota (SH), Ohio (SD, Oklahoma (SU/NR), Tennessee (S1), Texas (SI), Virginia (SD), and Wisconsin (S1)

(Louisiana Natural Heritage Program 2009; NatureServe 2006, 2009; Tim Smith, Missouri Department of

Conservation, pers. comm. 2005).

In some states, E. wo dn is given special designations separate from the NatureServe ranking. Wolf's

spike-rush is as an endangered species in Minnesota, Ohio, Tennessee, and Wisconsin, rare in Indiana,

a species of pane concern in Arkansas, Georgia, and lowa, and a species of uncertain status in Virginia

(Appendix 1). Separate state d tions are not provided for Eleocharis wolfii in Alabama, Illinois, Kansas,

Louisiana, Mississippi, Misses Neth Dakota, Oklahoma, or Texas (Appendix 1).

In Nebraska, E. wolfii is listed as a Tier 1 At-Risk Species as part of the Nebraska Game and Parks Com-

mission’s Natural Legacy Project (Nebraska Game & Parks Commission 2009a). This program is part of the

state’s development of a Comprehensive Wildlife Strategy that has been initiated in all 50 states (Nebraska

Game & Parks Commission 2009b). Eleocharis wolfii is included on the U.S. Forest Service's regional forester

sensitive plant list (Phillippe 2005; U.S. Department of Agriculture 2006).

Wilhelm (1977, 1978) developed a system to assess how conservative various plant taxa were in Kane

County, Illinois by giving each species a “coefficient of conservatism.” Species with the least conservative

value were ranked as O while those with the highest conservative rating were graded as 10. This system was

a d and modified by Swink and Wilhelm (1979), and further refined by Wilhelm and Ladd (1988). The

logy was summarized by Swink and Wilhelm (1994) and Taft et al. (1997). Swink and Wilhelm

(1994) -— coefficients of conservatism for each plant species in the Chicago Region. Eleocharis wolfi

was given the highest conservative rank of 10 and was listed as a representative of high quality wet prairies

(Swink and Wilhelm 1994). In other states in the Midwest, however, the coefficient of conservatism ranges

from 5-9 for Wolf's spike-rush.

State accounts

Alabama.—Smith (2002) listed Alabama within the range of E. wolfti and the species is known from a sole

historical record from Lauderdale County (Al Schotz, AL NHP, pers. comm. 2009)

Arkansas.—Historically, Wolf's spike-rush was only known from two specimens with vague local-

ity data from Arkansas and Little River counties (Table 1, Table 2). Recent surveys by Witsell, McKenzie,

and others in Arkansas from 2002 to 2008 have yielded 54 new sites that are widely distributed among 22

counties, including the two historical counties (Table 1, Table 3, Fig. 3). In Arkansas E. wolfii is found in a

wide variety of natural, historically open, wet to seasonally wet habitats. These habitats include unplowed

tallgrass prairies, saline barrens, sandstone glades, shale barrens, open hydric pine flatwoods dominated

by Pinus taeda, open hydric oak-dominated flatwoods, forested channel scar wetlands on abandoned stream

terraces in the Ouachita Mountains, and a seasonally wet deer-maintained salt lick. Eleocharis wolfii may

also occasionally occur in ditches along roads that cut through these habitats.

Colorado.—Although Smith (2002) stated that he had not seen specimens of E. wolfti to verily litera-

ture reports of the species in Colorado, Martin (1993) listed a record from El Paso County and noted that

the specimen had been cited by Svenson. However, Svenson was unaware of E. acicularis var. porcata and

misidentified specimens of this taxon for E. wolfii (Smith 2001a, 2002). Surveys in Colorado will be needed

to assess the current status of E. wolfii in the state.

Georgia.—Wolf's spike-rush is currently known from three sites in Walton County and one site in

DeKalb County (Jim Allison, and Tom Patrick, GÀ NHP, pers. comm. 2006).

[| [| £ all Dos H LN T L. | "EFE ET "3 £083

A F

Illinois.—The type locality for Wolf’s spike-rush is Fulton County, Illinois (Gray 1874; Mohlenbrock

1976; Phillippe 2005) and the species has undoubtedly received more attention in this state than any other

within its range. Robertson and Phillippe (1992) conducted an extensive survey for E. wolfi in Illinois and

documented 26 populations in 11 counties. Of these, nine populations were documented from four coun-

ties and the species was collected from ephemeral pools of remnant native prairie habitat. An additional

17 populations were recorded across seven counties and were found in ephemeral pools within open oak

flatwoods on river terraces (Robertson and Phillippe 1992). In Illinois E. wolfii often occurs in large colonies

that range from “several ad to many thousands of culms,” and these populations may cover a “few

hundred m? to a few thousand m?area” (Phillippe 2005; Phillippe, pers. obs.). Since 1978, E. wolfii has been

documented from approximately 47 sites in 16 counties in Illinois (Table 1, Fig. 3).

Indiana.—Eleocharis wolfii was first collected in Indiana in 1935 by FJ. Hermann and C.C. Deam at

apparently two separate sites (Phillippe 2005). There are currently 15 extant occurrences scattered across

Lake, Pulaski, and White counties in northern Indiana and from Jefferson, Posey, Ripley, Spencer, and

Warrick counties in southern Indiana (Swink and Wilhelm 1994; Phillippe 2005; Table 1, Fig. 3). Habitat

for the species in Indiana is identical to that is for Illinois (Phillippe 20055 Indiana Natural Heritage

Data Center 2009). Eleocharis wolfti occurs al l pools in 9 tris flatwoods in southern

Indiana and is associated with wet, sandy penes in the northern portions of the state.

lowa.—There are four historical records for the state: one each for Cedar, Emmet, Johnson and Union

counties (Eilers & Roosa 1994). The only known extant site occurs on the Williams Prairie State Preserve

in Johnson County (Table 1, Fig. 3). The species was found during a floristic inventory of Williams Prairie

in 2002 (John Pearson, Iowa DNR, pers. comm. 2009).

Kansas.—Wolf’s spike-rush was collected as early as 1897 in Cherokee County, but not again in the

state until 1986, when a specimen was taken in Crawford County (C. Freeman, pers. comm.). Since 2000,

E. wolfii has been documented to occur at nine aceon sites in dide Douglas, Franklin, Linn, and

Greenwood counties. The species is known in Kansas only from the eastern one-quarter of the state, where it

has bas o in a in ae Fa over both limestone and sandstone soils, in roadside ditches

uality sites, and in a disturbed, cool-season grassland adjacent

to remnant ponon aai pin o cs forest (Table 1, Fig. 3).

Louisiana.—Historically, E. wolfti was collected from three localities at one site each in East Baton

Rouge, Franklin, and Ouachita Parishes (= counties) between 1931 and 1970. Surveys conducted in 2007 and

2008 by Reid led to the discovery of 14 new sites in nine parishes that included seven new parish records.

Previous reports recorded for Natchitoches, Richland, St. Martin, and St. Tammany Parishes proved to be

misidentifications of other species, namely Eleocharis acicularis and E. montevidensis Kunth. In Louisiana,

Wolf's spike-rush is found in depressions in flatwoods and saline prairies, broader wet flatwoods, and in

ditches within these habitats. Several flatwoods records are from utility corridors where some portion of

the native herbaceous component persists. Much of this herbaceous layer is shaded out by dense stocking

of adjacent forests, due to fire exclusion or by aia types of manag en (e. 8- Pinus taeda - i pentations)

Minnesota eee is wolfit i tly known from fi (Pip , Ren-

ville, Rock, Scott) and the species historically known f tl dditional sitesin th te counties

(Nicollet, TE nia = P is T in e swales in edi is along Bus of

shallow d g the Minnesota River, and i in prairi alesan ] pools

of — Sioux UM outcrops (Fred Harris and Welby Smith, Manesa DNR, "s comm. 2006,

2007). Population estimates at extant sites range from 50 to 10,000s of plants (Harris, pers. comm. 2007).

Mississippi.—Eleocharis wolfii was historically known from one site in Tishomingo County and was

considered likely extirpated in the state until it was rediscovered in Alcorn County in 2008 by Charles

Bryson (Bryson, pers. comm. 2008

Missouri.—Wolf's spike rush was previously known in Missouri solely from collections taken in

Linn and Callaway counties (Steyermark 1963; Yatskievych 1999). Even with repeated searches to docu-

ment its occurrence in Missouri, the species had not been observed in recent years and was believed to be

McKenzie et al., St i f Clanchari IG: in tha IInitad Ctat dls

possibly extirpated from the state (Yatskievych 1999). The species was rediscovered in 1999 along the edge

of a seasonally-flooded bottomland forest in Stoddard County, in the Missouri Bootheel (McKenzie and

Jacobs 2000). Subsequent to this discovery, the species was documented in wet swales of five high quality

prairies in southwest Missouri, in early June 1999 (McKenzie and Jacobs 2000). In 2000, a second location

was documented in seasonally-flooded bottomland in Stoddard County (Smith 2001b). Between 2000 and

2008, Wolf's spike-rush was discovered at 21 additional prairie sites at scattered localities in northern and

southern Missouri and was rediscovered at its historical location in Callaway County. Eleocharis wolfti was

discovered on a sandstone glade in Henry County in 2003 (Missouri Natural Heritage Database 2009).

Currently Wolf’s spike-rush is known from 30 localities scattered throughout 18 counties (Fig. 3). Most

localities occur within the Unglaciated Plains Division (Thom & Wilson 1980, 1983; Yatskievych 1999) of

southwestern Missouri but there are a few records from the Glaciated Plains (Thom & Wilson 1980, 1983;

Yatskievych 1999) of central and northern Missouri and the RA Lowlands (Thom & Wilson 1980,

Te, ile gale in ext theastern Missouri. Duet t discoveries, the spe-

les of tion concern (Missouri ¡ Natural Heritage Prog 2009)

Nebraska. aha eee was first collected in Brown County in 1890 by J. Bates. Based on misiden-

tification of E. acicularis, Wolf’s spike-rush was deleted from the Nebraska list (Rolfsmeier 1995), but was

subsequently reconfirmed after a collection by Rolfsmeier in 1996 in Cherry County (Rolfsmeier & Steinauer

1999). Rolfsmeier and Robert Steinauer collected the species in Garfield and Rock counties, respectively, in

1999. pai deci aa! os E. wolfii from " sites between 2000 and 2005 while conducting

surveys for Platantl 1 Bowles and 5 lectus hallii (A. Gray) S.G. Sm. in the eastern

Sandhills of Nebraska, dud discovered an additional 14 new "m on the Valentine National Wildlife Refuge

in Cherry County in 2006 (Steinauer, pers. comm. 2007). Currently, E. wolfti is extant at 29 sites scattered

across seven counties in the northern one-half of Nebraska (Table 1; Fig. 3). Because it has been estimated

that there are 19,300 square miles (Knue 1997) of sandy habitat within the sandhills region of the state,

ongoing surveys are likely to yield additional new populations of this species.

New York.—Eleocharis wolfii is known solely from historical collections taken in 1927 from an open,

wet meadow adjacent to a train station on Long Island (Nassau County) (Troy Weldy, NY NHP, pers. comm.

2006). The population is now extirpated and is considered by most to be adventive (Svenson 1957; Troy

Weldy, pers. comm.).

North Dakota.—Martin (1993) listed two sites from Cass County, North Dakota. Kolstad (1986) and

Smith (2002) included the state within the range of E. wolfti. One site was known from a roadside ditch and

another site has since been oe by lige aa (Justin Parks, ND NHP, pers. comm. 2006). Wolf's

spike-rush is considered possibly ted from North Dakota (Parks, pers. comm. 2006; NatureServe 2009).

Ohio.—There is one extant site for E. wolfii in Ohio on a TNC preserve in Jackson County and adjacent

Pike County de was oy discovered in 1993 (Rick Gardner, Ohio DNR, pers. comm. 2009). There are

also two hist ken from remnant prairies in Ross County (1945) and Crawford County west

of Monnett (1953). The site in Ross County has been largely destroyed and apparently no longer supports E.

wolfii. Although the exact location of the historical collection in Crawford County can not be determined,

it is possible that it was taken from a high quality prairie that was converted to cropland in 1963 (Gardner,

pers. comm. 2009). The single extant population on the TNC preserve is apparently increasing due to good

management (Gardner, pers. comm. 2009).

Oklahoma.—In Oklahoma, E. wolfti was historically known from two collections: one taken by F.H.

Means, Jr. in Latimer Co., 1968 (KANU collection) and one collected in 1969 by U.T. Waterfall from a wet

prairie in McCurtain County (Amy Buthod, OK NHP, pers. comm. 2006). There have been no subsequent

surveys for the species in the state (Bruce Hoagland, University of ala pers. comm. 2000 BONEN,

in 2008, Jason Singhurst collected E. wolfii in Atoka County from TNC's B

where it was growing in a sandy hillside seepage bog (Singhurst, pers. comm. 20 09). This record apparently

represents the only known extant population in the state.

[| l at al Ds a Inm LI dM. dn adis Sia)

846 f Texas 3(2)

Tennessee.—Wolf’s spike-rush is known historically from seven sites in five counties: (Coffee, Frank-

lin, Grundy, Rutherford, Waren) (Table 1, Fig. 3) but currently only extant at one site each in Marshall and

Rutherford counties (Roger McCoy, Tennessee Division of Natural Areas, pers. comm. 2007). A new site

was discovered in 2003 in Rutherford County, but the site is threatened by construction and a new housing

subdivision. Construction activities possibly altered the hydrology of the site and the E. wolfii was being

displaced by species more typical of upland localities when visited in 2005 (McCoy, pers. comm. 2007).

Texas.—The status of reports - E. cl ME for Texas is unclear. Diggs et al. (2006) listed the

species for Bowie, Burnet, Jefferson and Morris counties , but Barney Lipscomb was only able to locate speci-

mens ork Bowie County (omne Research Institute of Texas [BRIT], pers. comm. 2007). S.Galen Smith has

seen specimens of Wolf’s spike-rush from Burnett and Jefferson counties (Smith pers comm. 2007). David

Rosen of Lee College in Baytown, Texas has collected the species from four separate sites 1 n Brazoria County

between 2004 and 2005 (Lipscomb, pers. comm. March 2007; Rosen 2007; Rosen. pers. comm. 2007). Smith

noted a specimen taken by Crockett in 1944 from Beaumont, Texas, but was unsure from what county the

specimen was taken (Smith pers. comm. 2007). We suspect that it is from Jefferson County. Martin (1993)

listed a specimen from Harris County but neither Smith nor Lipscomb has confirmed the identification

(Lipscomb and Smith pers. comm. 2007). A recent specimen collected by Jason Singhurst confirms the

presence of E. wolfii in Harris County (pers. comm. 2009). Recent collections in Brazoria County by Rosen

strongly suggest that additional survey efforts in this state are warranted. Several saline prairies/barrens

were recently identified in Harrison, Marion, and Panola Counties in northeast Texas, and have received

some preliminary botanical exploration (M. MacRoberts, pers. comm. 2009 and J. Singhurst, pers. comm.

2009). Given that E. wolfii occurs in this habitat in adjacent Caddo and De Soto Parishes, Louisiana, it will

likely be found in this part of Texas.

Wisconsin.—Eleocharis wolfti was recently rediscovered in Wisconsin (Smith ZOIL and the only extant

records of tl ies in the state are from one site each in Juneau and Marinette counties (Craig Anderson, WI

NHP, pers. comm. 2007; Galen Smith, pers. comm. 2007). There is also one historical collection taken from

nae Louny. Habitat sarees SOME isa Eanes and Lei dominated a mieado W a road side

Y AS AS O MA W t dou T4

Mosen prr YY YY Pi e E^

desa Dess comm. .. 2007).

Summary of threats

A. The present or threatened destruction, modification, or curtailment of the species' habitat or

range.—Although numerous new populations of E. wolfii have been discovered in the United States since

2000, some historical locations have been destroyed and extant populations are subject to various threats.

Some historical sites have been eliminated due to the conversion of native prairie and wetland habitat to

agricultural crops, the conversion of native prairie to hayfields and pasture, the cutting of many bottomland

forests, or the loss of habitat due to residential and commercial development (Turner 1934; Martin 1993;

Greenberg 2002; Nelson 2005; Phillippe 2005). Swink and Wilhelm (1994) and Greenberg (2002) sum-

marized the widespread destruction of native prairies and oak savannas near Chicago. Given the extent of

native prairie prior to European settlement, the historical range of E. wolfii was undoubtedly much more

extensive even than recent discoveries in the Midwest and south-central Gulf Coast would suggest.

In Illinois, Robertson and Phillippe (1992) noted that herbicide application at one site in Effingham

County killed ca. 9096 of the culms of a population discovered five days earlier by Phillippe. The alteration

of hydrologic cycles associated with overland flooding was identified as a threat to Wolf’s spike-rush in the

state by Phillippe (2005).

Due to the immediacy and magnitude of threats to Wolf’s spike-rush habitat in Illinois, especially on

private property, Robertson and Phillippe (1992) recommended that E. wolfii be listed as a threatened spe-

cies in that state. Despite the large number of extant sites in Illinois (Table 2, Table 3), species experts in

that state have recommended that the SRANK for the species be retained as S1.

MrKanzia at al fan £ El L a MESS all A Y T"

B. Over-utilization for commercial, recreational, scientific, or educational purposes.—To our

knowledge there is no evidence that Wolf's spike-rush is being negatively impacted due to over-utilization

for commercial, recreational, scientific, or educational purposes

C. Disease or predation.—Phillippe (2005) included livestock grazing as a potential threat to popu-

lations of E. wolfii. Other than possible negative impacts from grazing, we are unaware of any additional

threats to the species from disease or predation.

D. The inadequacy of existing regulatory mechanisms.—Although the NatureServe global and

state rankings for E. wolfii do not provide any regulatory protection for E. wolfii, separate designations in

some states afford limited regulatory protection for the species.

Nebraska's listing of E. wolfii as a Tier 1 At-Risk Species does not provide any regulatory protection

status to the species, but the designation has heightened awareness of its management needs and the NE

Department of Roads Morne consults with the NE Game and Parks Commission for highway projects

that may impact the species (R. Schneider, NE NHP, pers. comm. 2006). Eleocharis wolfti can not be collected

in the state without a permit (Schneider, pers. comm.).

A state listing as endangered provides little protection for E. wolfii in Wisconsin under state statutes

29.604 and NR (Natural Resources) 27.03-NR27.07 (Wisconsin Legislature 2007a, 2007b; Michigan State

University 2007). Under Wisconsin law, the taking of E. wolfti is prohibited add a pene under section

27.05. Permits are not required, however, for persons who want to take this or pecies: 1) on

property which they own or lease or for which they have been granted landowner permission, except if the

plants or their progeny are sold or processed, 2) on property that is being used for agriculture, construction,

or forestry practices, or 3) on property that is being operated or maintained as a utility facility (Michigan

State University 2007; Wisconsin Legislature 2007b).

Little protection is afforded E. wolfti under Indiana law, but personnel with the Indiana DNR have an

opportunity to provide input on state-funded projects that could negatively impact the species. In Georgia,

E. wolfii is listed as a species of conservation concern appendix 1) m it is not listed as a protected species

under Georgia state law (http:// ildlife.dnr.state.ga Iplants.asp). Region 9 of the

U.S. Forest Service includes: Es wo Ifi on their regional forester venetie plant list (Phillippe 2005; U.S. De-

partment of Agriculture 2006) but the designation does not provide any regulatory protection (Dave Moore,

USFS, pers. comm., Mar 2009).

E. Other natural or manmade factors affecting its pace existence. Non AIDE ns

climate change could contribute to loss of wetland habitat Peu by this species by g be

more extensive and persistent, especially in the Midwest w drought liti | isted for several

years, and are predicted to continue with i ae na d in the future (Hansen 1989; Rosenzweig et

al. 2000). Climate models indicate that high d an increase in pests (Rosenzweig et al. 2000)

and other invasive species (Vitousek 1994) will POER the increased frequency of droughts and other

extreme events (Dai et al 1996). Global warming and climate change have increasingly been identified as

factors which may contribute to the loss of biodiversity and extinction of imperiled species (Wilcove et al.

1998; Thomas et al. 2004; Maschinski et al. 2006). Habitat for E. wolfi is likely further threatened due to

projected changes in agricultural development, especially corn production that is needed for increasing

demands for ethanol production (Keeney and Mueller 2006; U.S. Department of Agriculture 2007). Keeney

and Mueller (2006) estimated a 254% increase in volume of water used in ethanol production from 1998

to 2008 for only one state in the Midwest. Corn production for ethanol plants is projected to increase in

2007 by 14.2% in Illinois (U.S. Department of Agriculture 2007), the state with the second largest number

of extant sites for Wolf’s spike-rush. Nebraska has large areas of a habitat in its sandhills region

(Knue 1997), but Keeney and Mueller (2006) postulated that i 1 for ethanol production may

result in competing water uses in that state. Increased demands for valer from above ground sources and

underground aquifers could prevent the formation and maintanence of important wetland habitat needed

by E. wolfti throughout the Midwest.

[| A £ sl n e Inm L ] FECI iT ara)

848 I lexas 3(2)

Research needs

Additional surveys are warranted, especially in states where the species is known only from historical loca-

tions believed to have been extirpated. Based on recent discoveries in Arkansas, Louisiana, Missouri, and

Nebraska, surveys should be conducted between March and May in the southern U.S. and between May

and June in the northern limits of the eem ines M da iue ug on the e ned and Mire is

requirements would be helpful in ident d. Recent

discoveries of E. wolfti in previously ntes ed habitats (e.g., sadermi glades in AR and MO, shale glades

and saline barrens in AR and LA, shallow depressions in gneiss bedrock in MN) suggests that the species

occupies a broader range of habitats than reported by McKenzie and Jacobs (2000), Smith (2002), and Phil-

lippe (2005). Natural community type, soil type, plant associates, and phenology should be recorded at

each site. Other tnan estimates taken at different sites in ins by eid and s (1992), we are

JA; ] ] xn Lea lata A

y additional report Lo F WELE MIAU LU

117^ auxr7TaTo0

Eon "-— E MA ; CHE ity of this species E

the long-term ee and conservation status mer this species in the future.

RESCATE on seed viability, predation, Į population genetics and germ ination i for the species

is | (Phillippe 2005). Phillippe (2005) postulated that gravity and flood water were likely mechanisms

for seed dispersal but additional investigations are needed, especially for upland sites. Studies should be

initiated to further examine competition and responses by E. wolfii to various levels of management and

disturbance, especia grazing Į [ native and non-native | Analyses of land use changes

would help threats to the species, especially related to the conversion of native habitat for

agriculture, silviculture, and residential development. Further evaluations are needed to assess differences

in habit, light requirements and reproductive success of individuals of E. wolfii in shaded sites vs. localities

in full sunlight.

Oak flatwood sites in AR, IL, IN, and MO are dependent upon flood waters or water accumulated from

precipitation. The importance of seasonal flooding to the dispersal of E. wolfii and its germination require-

ments should be further studied. What role the increase and spread of invasive species has on Eleocharis

wolfii should be investigated, particularly in light of possible scenarios associated with predicted impacts

from projected changes in climate.

Results of searches conducted by authors of this report suggest that additional populations of Wolf's

spike-rush are likely to be found if ys are conducted in appropriate habitats during the proper season.

Prairie swales, including those associated with remnant railroad prairies in IA, IN, MI, MN, ND, OH, OK,

and TX should be targeted for surveys. Oak flatwoods in MS, western TN, and southwestern KY may yield

additional populations.

Management needs

We can find little information on management recommendations to benefit Wolf's spike-rush. Phillippe

(2005) suggested that the maintenance of openings in flatwood habitats, the monitoring of potential im-

pacts of exotic species, and the use of prescribed fire to prevent the encroachment of woody vegetation were

treatments that could be useful in maintaining populations of E. wolfii. The import of using prescribed

fire to maintain prairie, savannah, and glade habitats has been extensively reported on in the published

and gray literature (e.g., Swink & Wilhelm 1994; Davit 1999; Greenberg 2002; Nowacki & Abrams 2008).

Prescribed fires not only maintain open habitats and halt the spread of trees and shrubs, but the repeated

practice prevents the establishment of fire sensitive eA pene. rue m CRUS reviewed in

Nowacki & Abrams 2008). Mowing and haying may also | hel trol woody

vegetation, by opening the canopy, and by providing a low level of distance necessity [ot colonization

of rhizomes or achene germination.

Phillippe (2005) advised against actions that alter soils where Wolf's spike-rush occurs but the rela-

tionship of soil disturbance in maintaining the species' habitat is not fully understood. Bowles et al. (1990)

EE, p o Mv: d Vie ec 849

McKenzie et al. Stat

included E. wolfti on a list of species that positively responded to ant! ic influences of humans at the

Indiana Dunes National Lakeshore but Phillippe (2005) believed that the response there was likely due to

increased light levels that resulted from the disturbance. Some level of disturbance from natural processes

(e.g., fire, flooding) is likely important for achene germination and plant growth of E. wolfii but we are un-

aware of the initiation of any experiments to test this hypothesis. The importance of maintaining natural

hydrologic regimes was noted by Phillippe (2005) and NatureServe (2009).

CONCLUSION

Although E. wolfii has disappeared from some historical sites across the range of the species and is threat-

ened with various land use changes, the large number of discoveries in the central U.S. between 2000 and

2008 strongly suggest that the species is not as rare as Previously ene but has been simply overlooked

by botanists who were unfamiliar with the species’ habitat 1 phenology or confused it with

other members of the genus. Significant threats to the habitat of E. wolfti still persist and, due to potential

negative effects associated with climate change, the increased demand for agriculture and ethanol produc-

tion, and commercial and residential develop t associated with population growth, we recommend

that botanists continue to monitor the status of this species throughout its range. The current NatureServe

global rank for this species is G3G4. Although 222 extant sites and populations at some localities number-

ing in the thousands of individuals may suggest a G4 ranking, Eleocharis wolfii remains threatened in many

areas. Due to these threats, the range-wide population status is likely somewhere between a G3 and G4.

Consequently, we recommend the retention of the current global rank of G3G4. If additional populations

are found, however, and/or threats to extant populations are significantly reduced, the global rank of this

species should be reassessed at a future date.

APPENDIX 1

tion for listings.

Alabama-No status; the species is E ios en ida list of species of special ai (Alabama Peparniment ui

Conservation and Natural Racniircoc e ol dye

ORT | ES NC frenlanta | i. Accessed 7 Mar 2009.

4 4 Ld F

Arlancac_ € 1 f [ TELE '

Ai Raisds lal |

(Arkansas Vascular Flora Committee 2006). Eleocharis wolfii is not listed tected species unde: OT state law.

Colorado Al hom LAG. [fe "J kK: gs | J J 4l Mies de s | | tfield guide ( (C | la Kl at it g Program

1999). Available at: http:// hi lostate.edu/ lants/ terlist.t Accessed Mar 2009.

Georgia-Species of special concern: An Ceo die the species i liste as a APAN species of special c concern e De-

ao of Natural h Resources 2008). £ offi is not | pecies under g /ailable

ri T i H I | " Load ff E = JPE J do f

cies asp; http:// georgiawildlif 'd tdetail.aspx?docid=89&pageid=7&category conservation

Accessed 7 Mar 2009.

Minels-Ne status: although Robertson aña Phillippe ee) a aed that E wolfi be included in the Illinois’ l er

2007). Available at: http:// Inr.state.il.us/espb/datelist.ht Accessed 7 Mar 2009.

ment of Natural R 2007). Available at: http:// gov/dnr/nature| /files/np spencer.pdf. Accessed 7 Mar

2009,

lowa-Species of special concern: Eleocharis wolfii is included on Lone B of Fragile Flora" (lowa Department of Natural

Resources 2007). Available at: http://fm.i i |71 | | Accessed 7 Mar 2

Kansas Al de de ri i . ¡AA hs I J | al Lh ry | IVA 54 IE r 1 fu 4 " £VAFIJInr1L

ING P, I A | VV IATA

and Darke 2005) A HM EM +. hp / f. i E: Los ESPECIES; | ttp:// | J + I yl ipl S l j ices/

Threatened-and-Endangered-Species/Threatened -and- Endangered-Species/Statewid List: http://www. ios state.ks.us/

í t/ h?SearchText=endangered. Accessed 7 Mar 2009

a s

FS liSsilana

dial is no acai status other t Natural Heritage state rank of 53 (Louisiana De] f Wildlife and Fist 2009)

C A A | / LE. H f " IL Hn / | " I L "Aa Ta LE: Io. h++ if

VE Lege f ics p » " j Il m ge/20099620Rare9620Plant9620T Lo 99620! x t] E Accessed 7 TN 2009.

Minnesota t langered: the Minneso ta DNR lists Wolf's spil h as ah endangered species on their Rare Species Guide

fl IR 2009). Available at: http:// Inr.stat Irsg/a-z_search.htmi?action=a-z

Searchelenermcecolunne SCIENTIFIC_NAME. Accessed 7 Mar 2009.

MEUS status: due to recent discoveries ir in Missouri, E. wolfii is no longer tracked on the states list of species and

i Natural Her

communities of conservation concern ( eritage Program 2009).

Ne braska-N No status: in LINGBIas au E. wolfii is listed Tier 1 At- à Br as part of the asus Game ang Pane Com:

Ni | Game Q Parks Commiss sio

"i a Comprehensive Wildlife Strategy that has Deen initiated in i 39 states (Nebraska Game & Parks commission 20090)

HER IE

+. bee

uc pdf. Accessed Mar 2009.

no status: this Spese. is nat tracked by the ING TOI Natural Heritage Program (New York Natural Heritage

| wild esi ML n Accessed 14 Mar 2009.

| +h hi] tl, T-] + larriral Llbaritgtann errar FA rs

Al Adda Fuel 4 Al es " | +

fs.fed.us/ hehe ep Tenis api rios 1998. pdf.

— e

Ohio-Endangered: Eleocharis wolfii is a on ES tales. 2008- dd is S: rare lis (Ohio Division of ios Areas and

iioi 2008). died’ a ttp// /3 2009%200hi0%20R OPlant%20Li

odf http ‘2018/default.asox#Anchor-46443. Accessed 7 Mar 2009.

í | roe M fete lab MAI LI

lahoma-No status: the Oklahoma Natural Heritage | tory does not include Wolf's spil h on their list of rare and

+ f, | " Il T4-

= Fa »

GAT BSFRR EGS

— Á L]

Ll I + A

OU. edu/plants rare _vulnerable.htm. Accessed 7 Mar 2009.

Tennessee-Endangered: the species is included on Tennessee's pate) Heritage Program: rare plant i MENESES

Department of Environment and Conservation. 2008). Available at: | t_list.

pdf. Accessed 7 Mar 2009.

Texas-No status: Wolf's spikerush is not tracked by the Wildlife Diversity Program of Texas Parks and Wildlife Department

as a State tarea teneo or r endangered species (Texas Parks ae ve DEBA du Available at: http U RONN, Roui

a em IE A

Ji th

ecc _ Accessed 7 Mar 2009.

J a e L+ L 4 H A AF Tea I-A mm emm mm

a CORseniatión and Recreation 2006). Available at http://www. EM gov/natural duel dou petet os

pdf. Accessed 7 Mar 2009

Wisconsin-Endangered: Wolf's P is included on the states list 9r threatened and endangered sj (Wisconsin

07) ere ee CR ] 5 | Aantal 2 orrn—o

Y Je i L Pp IRA ERAT p ae

Accessed 7 Mar 2009.

ACKNOWLEDGMENTS

We are grateful to the numerous botanists, data managers, and species experts who provided information

on E. wolfti: Jim Allison and Tom Patrick, GA NHP; Craig Anderson, WI DNR; Marlin Bowles, Morton

Arboretum; Brent Baker, UARK Herbarium,; Charles Bryson, USDA; Amy Buthod, OK NHP; Bill Carr,

TX Parks and Wildlife Department; Craig Freeman, McGregor Herbarium, University of Kansas (KANU);

Rick Gardner, OH DNR; Fred Harris and Welby Smith, MN DNR; Bruce Hoagland, Oklahoma University;

Tara Gibbs Kieninger, IL DNR; Karen Kreil USFWS; Barney Lipscomb, BRIT; Roger McCoy, TN Division of

Natural Areas; Dave Moore, USFS; Leah Oliver, NatureServe; Justin Parks, ND NHP; John Pearson, lowa

DNR: David Rosen, Lee College, Baytown, TX; Al Schotz, AL NHP; Rick Schneider, NE Game and Parks

Commission; Jason Singhurst, Ls a and Wildlife; Tim na me Troy Weldy, NY ra and O

Yatskievych, Missouri Botanica rden (MO). S.G. Smith ificant input on

the manuscript. We thank John Pelton and George Yatskievych for "ur photographs of MEM achenes

and floral scales. Barney Lipscomb of BRIT helped clarify the status of the species in Texas. Robert Steinauer

McKenzie et al., St t f Flanrhari Mi on tha linitard Chat -

supplied us with detailed information on his survey results for the species in Nebraska. Charles Bryson of

the USDA, Mike Penskar of the Michigan Natural Features Inventory and an anonymous reviewer provided

helpful and constructive comments on an earlier draft of the manuscript. Elderine Milligan of the USFWS

edited multiple drafts of the document.

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SALVINIA MINIMA AND S. OBLONGIFOLIA (SALVINIACEAE)

NEW TO CALIFORNIA, WITH NOTES ON THE S. AURICULATA COMPLEX

Richard E. Riefner, Jr. Alan R. Smith

Research Botanist

University Herbarium

University of California

1001 Valley Life Science Building #2465

Berkeley, California 94720-2

Research Associate

Claremont, California 91711-3157, USA.

rriefner@earthlink.net 65, U.S.A.

arsmith@berkeley.edu

ABSTRACT

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fram rtecidential ar rammoerrial ] I ] : water-gardens, zx 1 +1 | PC | Calo; tahkl-al 1;

sout hern Calif , but S. oblongifolia is likel persistent waif. We review the enon, a the Sana Sed ae and

] f 1 ] molesta

I [^]

in California ] I : | 1 y 1 J r. Casos I 1 p a EA 1 C

1 ] n4 1 Ma -l J s 1 1 1 1

i o I D

Key W juatic plants, Salvinia minima, Salvinia molesta, Salvinia obl gif lia, ti pl , wetlands, wildland-urban interface

RESUMEN

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4 oy i r r

J; A + tal : EF + ] t E + pa | Tis T Co Tos 15 Il

ki L J , e

l Sur de Calif S. oblongifolia es posibl planta abandonad persi También revisamos la taxonomía del

a ae F ein auriculata” y | las d inaci previas de esp tériles utilizando l isticas d

a l ia de S. molesta en California. Tambié j ] asia

A LI Lr r r

3.3 -J Ja Caleta? 5 J 1 A; A 1 tac ro -

I ] | I I I alifornia y resumiremos el

inka: A A aetna 1A + 1; Led ] ge ] E 7 J A dr : ` .

1 F > la» LEBAH LIC S Cll $ E

INTRODUCTION

Species of Salvinia are often difficult to identify, have a long history of use in cultivation, are known to es-

cape and colonize indigenous wetlands, and often become invasive in new and remote regions around the

world (Mitchell & Thomas 1972; de la Sota 1976; Forno 1983; Jacono & Pitman 2001). In this paper, we

provide the first documented records of Salvinia minima Baker and Salvinia oblongifolia Mart. for California,

and review their range, ecology, reproduction, potentially invasive behavior, mode of introduction, and

likely pathways of dispersal.

Salvinia molesta D.S. Mitch. (giant salvinia or kariba-weed) has been reported previously for California

ok et al. 2002; DiTomaso & Healy 2003; Riefner & Boyd 2005; Riefner & Boyd 2007). It is one of four

ically similar species frequently referred to as the “Salvinia auriculata complex" (i.e., S. auriculata

Aubl. p qM E la Sota, S. biloba Raddi, and S. molesta), Mese are Dun euet to Separate when plants

are infertile (Mitchell & Thomas 1972). To our] ledge, fertile I

oe outside of cultivation in Mora, making previous e problematic. We have deter-

rile herl of S. molesta by examining leaf primary venation patterns

and comparing them with the iligstratibns depicted by Forno (1983). As a result, we confirm the presence

of S. molesta in California, but cannot positively exclude other species in the S. auriculata complex. We also

provide a key to the identification of Salvinia species known or expected to occur in the State, review the

current naturalized status of urban and wildland habitats occupied by Salvinia species, and speculate on

potential regions in California where each species might become invasive.

| Rat Bac Inct Tavac 3(2): Le _ 866. 2009

856 J tl tanical h Insti exas 3(2)

SALVINIA SPECIES NEW TO CALIFORNIA

Salvinia minima (water spangles, common salvinia) and S. oblongifolia (giant water velvet) have not been re-

ported previously from California in treatments of the Salviniaceae that address nonnative species growing

outside of cultivation (Hickman 1993; Nauman 1993; Bossard et al. 2000; Hrusa et al. 2002; Dilomaso &

Healy 2003; Roberts et al. 2004; Rebman & Simpson 2006; Bossard & Randall 2007; Clarke et al. 2007;

DiTomaso & Healy 2007; Riefner & Boyd 2007; Dean et al. 2008; Roberts 2008; Jepson Flora Project 2009;

USDA 2009a).

Salvinia minima

Salvinia minima is a floating aquatic fern widely recognized as an invasive species in the southern United

States (Madeira et al. 2003; ISSG 2006; USDA 2009a). Like many aquatic plants, the natural range of S.

minima is cre to determine because it is grown in ue and oe ania and used in the

water and tl de, where it frequent! n (de la Sota 1976; Tryon

& Tryon 1982). Salvinia minima is aa to Florida and Georgia in the caen United States, southern

and eastern Mexico, Bermuda, the Greater and Lesser Antilles, Central America, and South America from

Venezuela to northern Argentina (Mickel & Smith 2004). Tryon & Tryon (1982) and DiTomaso & Healy

(2003) also report S. minima as native to the southern United States or Florida. Others, however, believe it

has been introduced to Florida and elsewhere in North America (Weatherby 1937; Crow & Hellquist 2000;

Jacono et al. 2001; Morgan 2009). The potential native occurrence of water-lettuce (Pistia stratiotes L.) in

Florida has also stimulated a similar debate (Schmitz et al. 1988).

In the United States, S. minima is now established outside of cultivation in Alabama, Arkansas, Florida,

Georgia, Louisiana, Mississippi, South Carolina, Texas, and Puerto Rico (Jacono et al. 2001; Jacono 2003;

Morgan 2009). The USDA (2009a) maps these and other occurrences, including New Mexico and several

mid-Atlantic and northeastern states. In New Mexico, S. minima is believed to be a non-persistent waif, and

reports from Maryland, Massachusetts, New York, and Oklahoma have not been verified (Morgan 2009). A

report from Minnesota also has not been confirmed (Nauman 1993).

Salvinia minima grows on still or stagnant waters of lakes and ponds, canals, slow-moving streams,

ditches, shallow backwaters of bayous, oxbows, cypress swamps, marshes, and on mud (Lellinger 1985;

Nauman 1993; Jacono et al. iin Much iie S. molesta, the distribution of S. minima is, in part, limited by

its low tolerance to freezing tem line waters (Jacono et al. 2001; Jacono 2003; Morgan 2009).

edad minima is BEEN to be eee uas and He ) molesta, noes by continuous

egetative ghter plants (Schneller

1980; jacomo et al 2001; Morgan 2009). Neuman (1993) recorded three appren fertile specimens, all

from Florida, and we have documented plants having a fertile axis growing in coastal San Diego County.

Jacono (2003) reported that sporocarps are common on large plants in the southeastern United States.

Salvinia minima can reproduce at remarkably high rates and completely cover the surface of waterways

if optimum growing conditions are present (Jacono et al. dn. acme Min. ISSG Martii In the southern

United States, particularly in Louisiana and Mississippi, 5. mi dinvasive

aquatic weed (Jacono et al. 2001). Dense infestations can block sunlight, decrease oxygen concentration

to the detriment of other aquatic species, replace native plants that provide food and habitat for wildlife,

clog irrigation systems, and block water intake screens and pipes of power plant facilities (Madeira et al.

2003: ISSG 2006). Salvinia minima, however, is prohibited only in Louisiana and Texas, and has not been

included on the Federal Noxious Weed List or the State of California list of pest plants (Plant Protection &

Quarantine 2002; California Department of Food & Agriculture 2008; Morgan 2009).

In southern California, S. minima is established at Be wildland-urban interface (WUD where it grows

kwat mt native a

on mud, in algal mats, ane nid waters IE urban draina ges, and in stagnant |

Tas J 11 HI 1 A 4 1

with üibanized watersheds (i.e., el flooding, üucrmanons in water devel and e water quality), S.

TP + me M a eee EE Ac Ll TA LL H PIS a 857

minima could superficially resemble small patches of Lemna and therefore may be easily overlooked in the

field (Fig. 1). Accordingly, S. minima may be more widespread in California than our records indicate.

Salvinia minima can overwinter in southern California stream environments or perennial wetlands

when fragments of rhizomes are dispersed, get tangled in debris racks, and/or are sheltered among stems or

roots of wetland plants, and then lie dormant until the return of favorable growing conditions in summer.

Along Alberhill Creek, western Riverside County, S. minima grows in a large slough infested with Eichhornia

erase Man) Solms- Laupach (water hyacinth). In this population, discovered in February, 2009, most

t f leaf dieback, and the pattern of browning is due presumably

to cold- Pa embolism. However. at his site San pans also pees new leaf growth and eaten

plants at or just below the water surface, which aj ool |

In the mild Mediterranean climate of rea, in coastal San eee ay e S. minima DOBULAON

discovered in April, 2009, overwinters without obvious col damage. At this site, and based on initial

observations, S. minima can compete with but apparently cannot TM Lemna in stagnant waters of a

culvert outflow basin (Fig. 2).

REINO the ability of Salvinia minima t i iods of reduced moisture or cool temperatures,

and its potential for rapid tati luction have conelbuted to its ongoing invasive spread across the

southern United States (Jacono 2002). ‘Salvinia minima may not become a nuisance when growing in urban

streams, but it could become invasive in sloughs, backwaters, ponds, and irrigation canals in California’s

mild Mediterranean coastal zones, possibly San Diego County, and the Lower Colorado River watershed in

the Sonoran Desert region.

Salvinia minima is easily separated from other species of Salvinia, including species of the S. auriculata

complex or S. oblongifolia, and other small floating aquatic plants using morphological features. Salvinia

minima produces hairs on top of papillae that divide into four thin, spreading branches that remain free at

the tips. Salvinia minima and S. molesta co-occur in drainages throughout the southeastern United States

(Jacono 2003). In Brazil, S. minima can grow mixed with species of the S. auriculata complex (Forno 1983).

Flooding, local environmental conditions, seasonal growth patterns, herbivory, and competition for space

and nutrients with other floating plants mae affect me local apne of S. minima (Dickinson & Miller

1998). Asa result, variable growth responses t nditions and micro-habitat preferences in

AMORE oul ae detection of S. minima difficult in the field. In the Lower Colorado River, few Salvinia

collected, and confusion between S. minima and small forms of S. molesta may

be prolematie (WAPMS News 1999). Therefore, small Salvinia plants growing with S. molesta populations

or rafting with Azolla and Lemna should be examined carefully in order to avoid misidentifications.

As noted by Forno (1983) and Nauman (1993), botanists should make every effort to locate and col-

lect fertile material to avoid misidentifications of sterile plants. In San Diego County, the fertile branches

of S. minima plants bearing sporangia appear to be more-or-less subumbellate, i.e., originating at or near a

common point on the axis, or divaricately branched (Fig. 3). However, the arrangement of the sporangia

in the California specimens conflicts with the illustration of the fertile axis—elongate with the sporangia

pinnately and alternately arranged—depicted by de la Sota and Jankowski (1996). The California material

is somewhat similar to the fertile axis shown in the photograph of southeastern plants provided by Jacono

(2003), but less elongate. Forno (1983) illustrated similar variation (short and long forms) of the fertile axes

in S. auriculata and S. herzogii, and this may be an indication that minor variations of the fertile axis are not

unusual in Salvinia. Accordingly, additional study and documentation is needed to accurately characterize

the S. minima populations growing in California.

1: U.S.A. CALIFORNIA Orange a City of Laguna Beach aaa! of MES St. and Pacific Coast Highway, UTM

(NAD 83) 115 0430540E 3707490N, elev. 4 1 shall basin, 9 Jul 2008, Riefner 08-219

(RSA, UC), same locality, 22 Oct 2008, foie 08 307 (RSA). Riverside Co.: Cathedral City, Perez Rd. at Date Palm Dr., urban creek

tributary to the Whitewater River, UTM (NAD 83) 115 0550217E 3737981N, elev. ca. 82 m, uncommon but ay ttered

, growin g

TTT NA Gp fT 4T Tl 1:1b r7 1

1 , W of

in algae, on mud, and in slow-movi ters, 5 Jul 2008, Riefner 08-214 (RSA, U

Hwy. 74, along Baker St. vicinity o ley Áve. at Bennett St., UTM (NAD 83) 1 Js 0467394E 3728133N, elev. ca. 380 m, uncommon

858 rnal of t tani i Texas 3(2)

Exc 1 Salvini aa ee 1

but scattered in o slough infested with Eichhornia crassipes, 25 Feb 2009, Riefner 09-17 (RSA, UC). iin Diego Co.: iod 2 bro

side, North Santa Fe Ave. at Mesa Dr., UTM (NAD 83) 115 0474261E 3678151N, elev. ca. 32m

Md E growing with Lemna and Wolffia in culvert outflow basin, 9 Apr 2009, Riefner 09 26 (RSA, UC).

Salvinia oblongifolia

Salvinia oblongifolia is native to Brazil where it can form dense, nearly pure colonies in lagoon habitats;

occasionally, it can also grow mixed with species of the S. auriculata complex (de la Sota 1962; Richerson

& Jacono 2005; Forno 1983). Salvinia oblongifolia is also grown for the horticultural industry, but is not

known to grow in natural environments outside of cultivation in the United States (Kartesz & Meacham

2005; Richerson & Jacono 2005). Sexual reproduction is expected in S. oblongifolia, but it also reproduces

primarily by vegetative propagation (Richerson & Jacono 2005).

Salvinia oblongifolia is a distinctive aquatic fern, and generally is easily identified (Fig. 4). Salvinia

oblongifolia has somewhat rectangular floating leaves approximately three times as long as wide and short

blunt papillae arranged in double rows on the upper leaf surface (Tryon & Tyron 1982; Richerson & Jacono

2005). Based on field observations, its prominent lower leaf keel and spongy aerenchyma tissue provide

extra buoyancy and, like S. molesta, may increase its ability to compete for space on crowded waters by

oye eee small m peu d leaf debris.

lt to grow when temperatures drop below 50°F and its growth may

"s bg vi ds by Pii dm in pH ene & Pitman 2001; Bed & Jacono 20 s dinde dd

A lel

was emd T the Bixby Slough population, its presence may represent: a SN waif dey

dispersed from cultivated sources to wildland habitats. Salvinia oblongifolia, however, could naturalize in

ssfully thern California

Fic. 2. Salvini

raz a FA " a £ al LJ | | FA CA || F1 L H aL! L "1 L £ PE | nL 5 Ll se] H A H |

2009, City of Oceanside, San Diego County, California.

subtropical and tropical climate zones elsewhere in the southern United States, especially Florida (Jacono

& Pitman 2001; Richerson & Jacono 2005).

Fund LI 1

St d: U.S.A. CALIFORNIA. Los Angel City, Bixby Slough, ity of Dodge St. at 255" St., UTM (NAD 83)

11S 0380730E 3740064N, elev. ca. 11 m, uncommon, floating with Lemna sp. and Ludwigia peploides subsp. montevidensis in sluggish

t d clustered t roots along bank of riparian woodland, 28 Jul 2008, Riefner 08-251 (RSA, UC).

O e

The Salvinia auriculata complex

Species of the S. auriculata complex share the defining character of "egg-beater" hairs (four hairs that join

at the tip), which grow from the top of papillae on the upper surface of floating leaves (Mitchell & Thomas

1972). The floating leaves of all four species are orbicular to ovate, but vary widely in size according to

environmental conditions and growth stage. Therefore, species of the S. auriculata complex are difficult to

separate when fertile plants bearing sporocarps are absent (Tryon & Tryon 1982; Forno 1983; DiTomaso

& Healy 2003; Mickel & Smith 2004).

Sporocarp-bearing plants are in the southern United States, and to our knowledge, fertile S.

molesta has not been documented by herbarium specimens or reported in the literature for California (Hrusa

et al. 2002; DiTomaso & Healy 2003; Riefner & Boyd 2005; Riefner & Boyd 2007). In Brazil, more than

one species of the S. auriculata complex can grow together in the wild (Forno 1983). We cannot, therefore,

860 tani titute of Texas 3(2)

ll d. I £ ] XL H LI Fr * J c n: P" de L H a L ^n

Lolas Ie (nal. La . mm safal 12 005 aba AO a : peg £. 4l | ta ninnataly

a ft [ 4

completely rule out the posso that other species of the S. ub dex LA could occur in California.

Accordingly, rey t sterile herbarium st as S. molesta or identified as

the S. auriculata complex representing a wide range of row innu were selected for study using illustra-

tions of leaf venation patterns and the methodology outlined by Forno (1983), which follows:

Methodology: Dried and pressed leaves were placed in boiling water with a few drops of household

detergent for a few minutes and allowed to cool. The leaves were then scraped of surface hairs and cleared

using a 596 NaOH solution (adapted here using 2096 Clorox solution) to aid viewing the leaf venation pat-

terns under a dissecting microscope. The venation pattern of each specimen was then compared to the

species-specific illustrations depicted in Forno (1983). (Clearings made by A. Smith and H. Driscoll, with

permanent slide mounts archived at UC.) Cleared leaves of specimens cited by CCH (2009) and deposited

at CDA (Herbarium of California Department of Food and Agriculture) include: Imperial Co. (Cason & Mi-

zumoto s.n., Herod & Pilman s.n), Mendocino Co. (Harrie s.n.), Riverside Co. (O'Connell s.n.), San Diego Co.

(Riveria s.n., Stevens s.n.), and San cii ue Co. (Knowles s.n., Stoltz s.n.).

As a result of this study, we of S. molesta in California, but cannot positively exclude

S. biloba, which has somewhat similar — However, S. auriculata, which is morphologically similar to

S. molesta, is the other species of the S. auriculata complex most likely to occur in southern California.

Fic. 4. Salvini LI CF ae CS | Los " J J z "EN NE PE] 2

J FI E J J £.

hilitv t Il flaat laavec and roots

taken in July, 2008, at Bixby Slough, Harbor City, Los Angeles County, California.

Salvinia molesta

Salvinia molesta is well known for its phenomenal growth rate and worldwide invasive spread, and is there-

fore widely regarded as one of the world’s worst aquatic weeds (Holm et al. 1977; Thomas & Room 1986;

Jacono & Pitman 2001). In the course of just a few decades, S. molesta spread from its native Brazil to aquatic

habitats in many tropical and subtropical regions around the world, including the southern United States

and northern Mexico (Jacono & Pitman 2001; McFarland et al. 2004; LCR 2009). It may be expected to

co-occur wherever Fichhornia crassipes is established (McFarland et al. 2004), although Salvinia species ap-

pear to be more cold-sensitive.

Even the name of this Salvinia has been a source of contention. De la Sota (1995) replaced S. molesta

with S. adnata Desv., an older name. However, because the Desvaux type is sterile and cannot be positively

identified with confidence, and is also of uncertain provenance, the use of S. adnata should be considered

a “name of uncertain application” (Moran & Smith 1999). Most authors have continued to use the name S.

molesta, as recommended by Moran and Smith (1999).

In the United States, S. molesta was first documented in 1995 growing outside of cultivation in South

Carolina; by 1999, the first collections from California were made from the Imperial National Wildlife Ref-

uge, Imperial Co., from near Blythe, Riverside Co., in the Lower Colorado River drainage, from a pond near

Fallbrook, San Diego Co., and from the San Diego River floodplain (without voucher), also in San Diego Co.,

southwestern California (Jacono & Pitman 2001; Hrusa et al. 2002). It has since been documented from

|| £ sl nm.a b IP L I "X B ET * £3

862

canals near El Centro and Holtville in Imperial County, and from man-made ponds and native wetland

habitats from coastal San Diego and Orange counties north to Fort Bragg, Mendocino County, in coastal

northern California (CCH 2009). Salvini lesta has been reported recently from Humboldt County, which

is also in coastal northern California (Calflora 2009). Accordingly, the distribution of S. molesta in California

is more widespread than previously mapped by McFarland et al. (2004) and Jacono & Richerson (2005).

Though benign within its native range, S. molesta is an aggressive aquatic menace that can suppress

growth of native vegetation, degrade water quality and fish and wildlife habitats, clog een canals,

interfere with recreational activities and utilization of water resources, and impact wetland

values and functions. Salvinia molesta and other species of the S. auriculata complex are included on the

Federal Noxious Weed List, and therefore prohibited from importation to the United States and from trans-

port across state lines (Plant Protection & Quarantine 2002). Although species of the S. auriculata complex

are included on the State of California weed list (“A List”) of pest plants, S. molesta has become an invasive

pest in several national wildlife refuges located along the Lower Colorado River (California Department of

Food & Agriculture 2008; LCR 2009). Salvinia molesta has also been identified as a “High Alert” plant by

the California Invasive Plant Council (Bossard et al. 2006).

Salvinia molesta is distinguished from other species in the S. auriculata complex by the arrangement of

the fertile axis, which bears a large number of sessile sporangia arranged in a dorsiventrally arranged cyme

(Forno 1983). Sporocarps are small, ovoid, apiculate, and contain a high number of empty sporangia or

abortive spores (Mitchell & Thomas 1972; Tyron & Tyron 1982). However, no fertile material of any Salvinia

in California has been collected.

The leaf venation pattern of S. molesta was used by Forno (1983) to distinguish it from other members

of the S. auriculata complex; however, this pattern is somewhat similar to that found in S. biloba. These

two species can be carefully separated by examining the number and shape of the primary areoles situated

between the keel and the margin of a tertiary leaf; i.e., 5 to 10 areoles in S. molesta, and 8 to 12 areoles in S.

biloba, with the latter generally having 10 or more present. Because of the similarity of leaf size, shape, and

patterns of venation, populations of S. molesta in California and Arizona should be carefully examined for

the presence of fertile plants to ensure that S. biloba is not present in California. Salvinia auriculata, possibly

to be expected in California, greatly resembles and can be confused with S. molesta. The leaf venation of

S. auriculata is reticulate and similar to the venation of the outer third of a S. ‘ies we Illustrations of the

primary leaf venation patterns of S. auriculata, S. biloba, and S. molesta, and | bearing

e L r r

plants are depicted in Forno (1983).

Salvinia auriculata has been reported previously idis Mexico and Puerto Rico, Bu not from California

(Mickel & Smith 2004; USDA 2009c). Therefore, S i luded in the foll g key that will serve

to identify the sterile species of Salvinia known or ene to occur in California.

A KEY TO STERILE MATERIAL OF SPECIES OF SALVINIA IN CALIFORNIA

(ADAPTED, IN PART, FROM FORNO 1983)

+ L H Il y Et L

1. Floating leaf blades oblong, generally 2-3 times longer ; par ( ) SUP} g hai pl

blade surfaces « 0.1 mm long S. oblongifolia

1. Floating leaf ds round, or if slightly (suborbicular), less than 1.5 times a wide; papillae

BONO! hairs an upper mage EEES af wall developed leaves to 1 du. lon

2 | pal ill f uppel surface TI | 8 | | Y lly 0.5 1.0 (-1.5)

cm long S. minima

2. Hairs on tips of papillae of upper surface of floating leaves joined into a darkened knot at tips ('egg-

cdd Palisa, leaf aS n z= .5 cm long.

al | ù I ral xb. un De dali : | 2/3

EL M \ J

£l gi L H H Lt | I] í Lo y H'WI L + J | L J +]!

| ; g gans) y;

known from CA, to be expected S. auriculata

3. Leaf venation of areoles of unequal length (the outer much shorter than the inner) in marginal 2/3 of

ar Dl

in coastal and southern CA S. molesta

J 1: J J A A PS J n eI : hal n LC pocos H I dul d:

TABLE 1 y of distributi tna! tat , and | p

in California,

Species Significance in California Naturalized Status Potential Invasive Behavior

and Habitats

Salvinia auriculata Not currently known, but NCI Possibly Sonoran Desert region

expected in southern counties

Salvinia minimc New to State; expected N — urban creeks NW - ip coastal southern California,

elsewhere in NOM California slough specially San Diego Co., and

ES Desert region

Salvinia molesta Identification of infertile N - backyard ponds, Sonoran Desert region, central a

molesta confirmed. agricultural ponds, southern coastal regions, died

Similar species in the S. auriculata drains, and canals San Diego Co, but unlikely

complex cannot be excluded NW - river channels, elsewhere

backwaters, an

floodplain ponds

Salvinia oblongifolia New to State; firs d growing C-sloug Unlikely in California

outside of eed in US.

DISCUSSION

Urbanization in arid and semiarid regions often brings unintended changes to fluvial ecosystem processes

and threats to native biodiversity, including replacement of ephemeral washes with year-round waters and

the introduction of new exotic species (Schwartz et al. 2006; White & Greer 2006; Roach et al. 2008).

Seasonal stream flows and summer-dry conditions of southern California’s arid climate have in the past

acted as a batrier to establishment by many nonnative plants dispersed at the WUI (Brigham 2007). The

availability of year-round sources of water, however, generated by man-made activities has inadvertently

created numerous new wetland and riparian habitats that have significantly increased the likelihood that

nonnative facultative wetland plants escaping cultivation could migrate from benign urban environments

outward along the WUI to establish in native wetland habitats (Riefner & Columbus 2008).

In addition to intentional introductions for water-gardens, federally listed noxious plants, including

S. molesta, have also been inadvertently introduced as contaminants with non-invasive wetland plants sold

over the internet (Kay & Hoyle 2001; Maki & Galotowitsch 2003). The continuing growth of internet sales

and demand for the water-garden industry will increase the likelihood that species of Salvinia will continue

to be introduced by cultivation and escape to California’s urban and wildland wetland habitats.

As a result of field experiences in southern California and a review of pertinent literature, the new

distributiona! records reported here, current urban and wildland habitats occupied by Salvinia species, and

potential regions where Salvinia species might become invasive are summarized in Table 1. Naturalization

categories shown in Table 1, which are somewhat subjective, follow Hrusa et al. (2002) in order to allow

consistency in data compilation of new introductions, which include: naturalized in wildlands (NW); natu-

ralized outside of wildlands, including WUI habitats (N); casual or waif populations (C); and no current

information (NCI).

ACKNOWLEDGMENTS

We greatly appreciate the efforts of Harvey Brenneise and Irene Holiman (Library of Rancho Santa Ana

Botanic Garden) for assistance with document retrieval, Fred Hrusa (California Department of Food and

Agriculture, CDA) for a loan of specimens, and Heather Driscoll (UC Herbarium) for some slide prepara-

tions. Two anonymous reviewers provided helpful comments that greatly improved this manuscript. We

also thank our friends and colleagues who provided locations and directions to ponds, basins, and sloughs

that might support nonnative aquatic plants.

864 J loft tanical h Institute of Texas 3(2)

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vegetation of Los Peñasquitos Creek, California. Landscape Urban Planning 74:125-138

SAGINA CAESPITOSA (ALSINACEAE) IN COLORADO, U.S.A.

A.E. Porsildt W.A. Weber and R.C. Wittmann

National Museum of Canada University of Colorado Museum

Ottawa, Ontario, CANADA UCB 265, Boulder, Colorado 80309, U.S.A

bill weber@colorado.edu

ABSTRACT

Sagina caespitosa (J. Vahl) Lange in HJ MD: 1, : p ] f. +1 ES PS » 21 TT ] cua r. +l D 1 RA aes To »

RESUMEN

Sagina caespitosa (J. Vahl) Lange in HJ. Rin! tra por la pri vez en Los Estados Unidos en los Montañas Rocosas de Colo-

In this note, we report Sagina caespitosa (J. Vahl) Lange in HJ. Rink for Colorado and the United States.

Otherwise, S. caespitosa occurs in North America in Greenland, Manitoba, Newfoundland, Labrador, Nu-

navut, and Quebec (Crow 2005).

According to the Flora of North America (Crow 2005), two species of Sagina occur in Colorado, namely

S. saginoides and S. procumbens. For many years, we assumed that our only native species was the common

S. saginoides. Erling Porsild visited in 1950 and the senior author took him, as he has Eilif Dahl, Eric Hultén

and other European field botanists, on excursions to 14000-foot Mount Evans, the site of many significant

alpine or arctic disjuncts (Weber 1965, 1991). Since then, the authors have repeatedly collected Sagina there

and at similar sites. Recently, Wendy Dathan, Porsild's biographer (manuscript in preparation), sent me a

copy of the page in his field notebook where he mentioned collecting Sagina caespitosa and how thrilled

he was, since he had not seen it since Hudson Bay and Greenland. To Porsild should go the credit for the

discovery.

Sagina caespitosa has been considered an Amphiatlantic species; however, in Colorado it is locally

abundant on high mountain summits from 12000-13500 ft (3700-4100 m) altitude, on frost boils and

well-developed gravelly moss tundra. Throughout the remainder of its range, S. caespitosa is not known to

grow at elevations greater than 1500 ft (450 m).

A report of S. nivalis (Lindblom) Fries from Colorado by Rydberg (1906, p. 130) was overlooked until

recently. Rydberg's report was erroneous, based on a Harry Patterson collection which is here assigned to

S. caespitosa. S. nivalis, being a species of the American arctic, might also be expected in Colorado, but thus

far it has not been detected; however, S. nivalis does occur in a few localities in Montana.

In the Flora of dis America dM is ció scene (L.) H. Karsten was described as “Plants

perennial, tufted or ! habitats.” This is incorrect and all collections of S. caespitosa

at Herbarium COLO were anmotated as S. NEM In the field, S.caespitosa is easily distinguished by its

tightly massed, button-like form. Plants are also characterized by short shoots lacking internodes, by stiffly

erect pedicels, by plump, purplish sepals, and by petals equaling or slightly exceeding the calyx. It occurs

on mature gravelly tundra usually at very high altitudes. The shoots can hardly be separated to make less

bulky herbarium specimens.

Sagina nivalis (Lindblom) Fries might be expected to occur in Colorado, since it is a western American

arctic species that reaches northern Montana. It differs from S. caespitosa by being 4-merous with petals

shorter than the sepals, and by not being densely caespitose because of the lack of secondary shoots.

Sagina saginoides (L.) Fenzl isa very weak, often somewhat procumbent plant with obvious internodes,

filiform pedicels and pale white sepals. Its habitat is not special, occurring on wet sand or mud, and it gener-

ally occurs at lower altitudes.

| Rat Bac Inet Tavac 3(2): Qr7 _ 868. 2009

868 I | fal Dos a In LI dt f Texas 3(2)

Voucher collections of S. caespitosa, Herbarium COLO: COLORADO. Boulder Co.: sandy seepage area between the two uppermost

Green Lakes, TIN R754W Sec. 13, Green Lakes Valley N of Kiowa Peak, 11,500-12000 ft, Boulder watershed, 21 Jul 1953, Weber &

Dahl; N slopes of Kiowa Peak Green Lakes Valley, 11,400-1200 ft, 1 Jul 1972, Komarkova s.n.; summit of South Arapahoe Peak on its

southeast ridge, 13300 ft, 10 Jul 1972, Komarkova s.n. Clear Creek Co.: high mountains, vicinity of Gray's Peak, 23 Aug 1892 (as S.

nivalis), H.N. Patterson s.n. [the collection cited by Rydberg (1906)] [on the specimen at F, an inked note says "halfway up"]; Summit Lake,

Mount Evans, 12,800 ft, 5 Aug 1950, Weber 5927, 15 Aug 1950, Weber, Porsild, & Holmen 5927, 11135, 22868, "e Tue 1954, Weber & Dahl

8937; 16 Aug 2006, Weber 19505; 16 Aug 2006, Weber 19505; 25 Aug 2007, Weber & Wittmann 19514 t Mts. Evans

and Epaulet, 13590 ft, 6 Aug 1998, Weber & Wittmann 19403. El Paso Co.: Pikes Peak, along auto a ca. 13,000 ft, 7 Aug 1932,

Penland, Wershing, & Hartwell 789 (COCO). Pant Co.: M a Nat. ivit Trail URGES on terrace $ of oe Lake, a ft, on

bare wet humus around sis scars, MM 196 g

hummocks, pide 1960, 1 f Hagues Peak, 13300 ft, i ls of a very ilite Snow lati , 7 Aug 1961,

Willard 6 Park C fosquito Range, along st N ofI ick Tarn, SSE of Peerless Mt. summit, 12320 ft, 30 Jul 1991, Tonnessen

& Roy be Mosquito Pass, saturated below snow-bed beside the jeer 1, 30 Jul 2007, Weber & Wittmann19527.

ACKNOWLEDGMENTS

We thank Garrett Crow and Wendy Dathan for assistance. Del Wiens kindly reviewed the ms.

REFERENCES

Crow, G.E. 2005. Sagina. In: Flora of North America Editorial Committee, eds. Flora of North America North of

Mexico. 5:140-147.

DATHAN, PW. 2010 estimated. The Reindeer Botanist: AIf Erling Porsild. Manuscript in preparation.

Weser, W.A. 1965. Mount Evans, concentration point for Pleistocene relict plants. Guidebook for one-day field

conferences, Boulder Area, Colorado. International Association for Quaternary Research, VIIth Congress. Pp.

Weser, W.A. 1991. The alpine flora of Summit Lake, Mount Evans, Colorado. 15(4):3-15.

ERIGERON BELLIOIDES (ASTERACEAE), NEW TO FLORIDA AND

THE CONTINENTAL UNITED STATES

Randy L. Mears

10852 SW 88th Street, Unit #108

Miami, Florida 33176, U.S.A.

ABSTRACT

TI 1 [:.1.1 1 Li È m JE 1 11: .: J 1 1 ] y rn 1 A

[e] A O J i F

Dade County, Florida. A description, localities, d ji g } luded

RESUMEN

4 " T E A 1 4 | A = — m —Lraelf Lael T ER | 1 ] J J 1; J

r * oO

en el condado de Miami Dade, Florida. Se incluy descripción, localidades, especi iadasy, fotografías y una ilustración.

Erigeron bellioides DC. grows outside of cultivation in southern Florida (Fig. 1). The collections cited below

are apparently the first records from Florida as well as the continental USA. The species was not included

or mentioned in any of the following major floristic summaries: Correll and Johnston (1970), Cronquist

(1980), Cronquist et al. (1994), Hickman (1993), Long and Lakela (1976), Wiggins (1980), Wunderlin and

Hansen AE Nee ae,

tive tot ter Antilles in the southern Caribbean region (Wagner et al. 1999).

It occurs on Cuba, Hi pne aici Puerto Rico (Loigier et al. 1982). It also occurs in , the Baharia Archipelago

(Correll & Correll 1996). The species has become established on some of the Hawaiian Islands as well as

other Pacific islands (PIER 2009). I personally observed the species naturalized on the Hawaiian islands

of Kauai and Oahu in May of 2004. Erigeron bellivides is referred to as “bellorita” in parts of its native range.

Members of the genus often are referred to as “fleabanes.”

Voucher specimens: Florida. Miami-Dade Co.: 10 May 2000, Mears 4725 (DOV, FIG, USF); 11 Dec 2005, Mears 4986 (DOV, FTG,

USF); 4 Nov 2003, Mears 4836 (DOV, FTG, USF); 20 Nov 2005, Mears 4971 (DOV, FTG, USF); 11 Dec 2005, Mears 4988 (DOV, FIG,

USF); 2 Dec 2005, Mears 4979 (DOV, FTG, USF).

The plants of Erigeron bellioides have been observed flowering throughout the year in south Florida and

were growing on grassy roadsides and medians of parking areas, in damp to wet soil at all of the sites. The

following species were associated: Kyllinga brevifolia Rottb., Paspalum setaceum Michx., Fleusine indica (L.)

Gaertn., Dichondra caroliniensis Michx., Pilea microphylla (L.) Liebm., Coronopus didymus (L.) Sm., Drymaria

cordata (L.) Willd. ex Schult., Calyptocarpus vialis Less., Synadrella nodiflora (L.) Gaertn., Youngia japonica (L.)

DE, 3 didi qd iiim L., and Enna opu de

f Erigeron bell q on (Wagner et al. 1999; Correll & Correll 1996)

and on personal abina oe made of the species as it occurs within Miami-Dade County. Plants with an

oblique rootstock 1-3 cm long, sparingly and loosely villous throughout. Stems simple or few-branched,

ascending or stoloniform, 4-10 cm long. Basal leaves rosulate, with a conspicuously margined petiole

usually longer than the blade, blades suborbicular to broadly ovate, to ca. 3 cm long and wide, entire to

obscurely few-toothed; cauline leaves few, much reduced, and mostly obovate to oblong. Heads solitary or

2-3: involucres 3-3.5 mm high, ca. 5 mm broad; phyllaries in ca. 2 series, linear, acute, strigillose. Rays

white, barely as long as the phyllaries, tightly coiling at maturity. Achenes elliptic, flat, about 1 mm long,

puberulent. Pappus brownish, about 1.5 mm long.

This species is distinctive among Erigeron species in the USA in its combination of annual duration,

very small size, spatulate leaves arranged in a basal rosette, and much reduced cauline leaves.

J. Bot. Res. Inst. Texas 3(2): 869 — 871. 2009

870 t tani titute of Texas 3(2)

Fic. tographs of Eri bellioides, Miami-Dade County, Florida. Habit, Mears 4725 (left); close up of flower (right, 19 Oct 2009). Photographs

by Randy L. Mears.

ACKNOWLEDGMENTS

Guy L. Nesom provided helpful review comments that greatly improved an earlier version of this manuscript.

I would also like to thank Robert F.C. Naczi of the New York Botanical Garden for so graciously critiquing

and providing several helpful suggestions pertaining to the format and content of this paper.

REFERENCES

Correll, D.S. AND M.C. JOHNSTON. 1970. Manual of the vascular plants of Texas. The University of Texas Printing

Division, Dallas, Texas.

CorreLL, D.S. AND H.B. Correct. 1996. Flora of the Bahama Archipelago. A.R. Gantner Verlag KG., FL-9490 Vaduz.

Cronauist, A. 1980. Vascular flora of the southeastern United States Vol. 1, Asteraceae. Univ. of North Carolina

Press, Chapel Hill.

CRONQUIST, A. et al. 1994. Int

New York Botanical G

Hickman J.C. 1993. The Jepson manual: higher plants of California. Univ. of California Press, Berkeley and Los

Angeles.

LogiER, H.A. AND L.F. MarTORELL. 1982. Flora of Puerto Rico and adjacent islands: a systematic synopsis. Editorial

Univ. Puerto Rico.

Lone, R.W. AND O. LaxeLa. 1976. A flora of tropical Florida: a manual of the seed plants and ferns of Southern

Peninsular Florida. Banyan Books, Miami, Florida.

vascular plants of the Int tain West, U.S.A. Volume 5 Asterales.

, Bronx

Mears, Erigeron bellioides, new to Flordia 871

Pier. 2009. Pacific Island ecosystems at risk (PIER). Institute of Pacific Islands Forestry. «http//www.hear.org/

pier/index.html»

USDA-NRCS. 2009. The PLANTS Database. National Plant Data Center, Baton Rouge, La. <http://plants.usda.

gov»

WAGNER, W.L., D.R. Her8sT, AND S.H. SouwrR. 1999, Manual of the flowering plants of Hawai'i revised edition Volume

1. Univ. of Hawaii Press Bishop Museum Press, Honolulu.

Wicains, 1.L. 1980. Flora of Baja California. Stanford Univ. Press, Stanford, California.

WUNDERLIN, R.P AND B.F. HANSEN, 2003. Guide to the vascular plants of Florida, Second Edition. Univ. Press of Florida,

Gainesville.

|| || f a nm a * if Lo) E e ET ia

lexas 312)

872

BOOK REVIEW

LYTTON JOHN MUSSELMAN, WITH FOREWARD BY GARRISON KEILLOR. 2007. Figs, Dates, Laurel, and Myrrh: Plants

of the Bible and the Quran. (ISBN 978-0-88192-855-6, hbk.). Timber Press, Inc., 133 S.W. Second

Avenue, Suite 450, Portland, Oregon 97204-3527, U.S.A. (Orders: www.timberpress.com, 1-800-327-

5680, 1-503-227-3070 fax). $24.95, 336 pp., 255 color photos, 1 map, general index and index to

verses in the Bible and Quran, 5 1/2" x 8".

From the publisher: “This book Ba the ie of the Old Testament, New Testament, and Quran. From acacia, the wood of the

A to Mero absinthe, fl , 81 chapters cover the more than 100 oe. in s

holy l coi Aleman tories of the fruits, grains, grasses, trees, flowers, and frag

ill J Bei ones. anlar this Aa lightful emmimeninral botany

med Fa end uses, and ovde de context. n illustrated f

11 LE »

VIII Lilt

]

1 Tus 1 T3] fl DAI J at nm ] 4 1 + 1 1 3 1 1 1

Figs, Dat 9, I

pon to in rtir the Bible or dn Quran. T plant listed alphabetically by thei but the author does list many

+ Fila baal Far ` ] +4 difiere cli ghtly,

g ariou I but the ius will A

find a brief d ipti f the plant (e.g sd onion, saffron, wheat) that includ lor imag habitat,

and uses. Mussel Iso includes tl l scripture or simply the specific chapter or verse from the Bible or Quran asa A

for the reader.

T 3 eee 4s 4 + «lI + +1 1 ] LR Eos ral = —:L71 Ary 1 : +l 4 n

O IL acids x Bod r

duction of this book. In s plant is listed i ly f the religi (e.g IE Times re of the wee versus

the New Living Translation). F ling this book, it appears that M ] trul ]

der to determi f tl icular pl that ioned withi h text This PORR was mde to understand and would be

1 1f lag ] 1 tl that ira Keri Mr New

5 + E e a LAT A LYRLILNYLVE,

MS Biology, Prog Manager, Botanical R h Institut UAE Fort t Worth, Texas 76102-4025, U.S.A.

T.L BA. J 1 4 ] ] D:L. 1 4 2 +1 : PRA 1 eus E 1 q q Jt td 1 7 . Jnl

AE the ndi I I

bolis: j : +1 1 1 e IIn in AA IT T E. fo ] «I C Fil

f Biological Sci Uni in Norfolk Virginia. He iso wrote te Jordan in Bloom (2000), ok was s commissioned

t, has held

by Queen Rania Al-Abdullah. With

three AUDE a and has na asa visting soles at a American University o of Beirut. Also a field naturalist, he co-edits

l Preserve.”

Haustorium, I

J. Bot. Res, Inst. Texas 3(2): 872. 2009

A FLORISTIC INVENTORY OF MARCO ISLAND (COLLIER COUNTY), FLORIDA

GJ. Wilder B.J. Roche

eee ea Garden Conservancy of South se mE

re Drive 1450 Merrihue Driv

Naples, vies Bs ie 7336, U.S.A. Naples, ada 34102- ein U.S.A.

GWilder@naplesgarden.org jeanr@conservancy.org

ABSTRACT

Marco Island, situated within the Gulf of Mexico th thern tip of peninsu lar Florida, exhibits six main kinds of natural

habitats together with ruderal land and other disturbed ar areas. oon daa e re Mens 324 genera, 461 species, and 470 in-

c dpi taxa Ee e» eins varieties, and a hy vascul pa grow ild land, and 325 of the infrageneric taxa

habit f to six of each e six main kinds of natural habitats present. Fourteen

of the infrageneric taxa are rare in Florida. Trichostigma octandrum is abundant on the Island, but might grow wild at solely two other

localities in the United States.

Key Worps: Flora, Marco Island, vascular plants

RESUMEN

Marco Island, situada en el MM e México yc cerca = t peninsular de Florida, presenta seis a e de hábitats

naturales, junto das. En la isla crecen ciento once familias, 324 géneros, 461 especies, y 470

taxa ON ee subespecies mn y un niu de plantis Y RS y Vii be a taxa infagenencos son nativos

de Florida. pos ] j j tes. Catorce de

los taxa infragenéricos son raros en Florida. Trichostig tand bundant la Isla, pero pudiera vivir únicamente en otras dos

localidades en los Estados Unidos.

INTRODUCTION

Thisis the 1 paper of a series ing the flora of southwestern Florida (Wilder & McCombs 2006).

Herein, we present the results of a study of the native and nonnative taxa of vascular plants growing wild

on Marco Island, Collier Co., Florida.

We undertook this study for five reasons. First, no published flora existed for Marco Island, overall.

Second, the potential for additional residential development had created pressing need for a floristic inven-

tory of the Island that would serve as a “baseline” for future floristic studies. Third, the Island flora appeared

unusual compared to the floras of adjacent portions of mainland Florida, and included rare taxa. Fourth,

we wished to document the flora by vouchering inventoried species. Previous investigators had produced

species lists or accounts of species for parts of the Island, but had not cited herbarium specimens (Beriault

1980; Bradley 1999; James N. Burch [list reproduced in Collier County Facilities Management Department

2007]; Johnson and Muller 1993; Matt Finn and Eileen Ward [lists reproduced in Tsandoulas @ Ward

2008]; Natural Areas Management Co., no date; Small 1924, 1929). In contrast, Gann et al. (2002) had cited

specimens of rare taxa previously collected on the Island. Fifth, Marco Island, being subtropical, warranted

study, because of the diversity and uniqueness of the Floridian subtropical flora.

We undertook a tripartite approach to this investigation. (1) We inventoried the wild native and non-

native taxa (Appendix I). (2) We intensively studied selected sites within the main kinds of natural habitats

on Marco Island, in order to determine which native taxa occupied each kind of habitats and, for certain

habitats, to gauge the frequency of these taxa (Appendix II). (3) We tabulated additional taxa of vascular

plants collected by previous investigators on the Island but not encountered during the present study (Ap-

pendix I)

Marco Island has long appealed to floristic botanists. In addition to workers, aforementioned, the fol-

lowing noteworthy investigators made botanical visits to the Island: F. Almeda, George N. Avery, Leonard

J. Bot, Res. Inst. Texas 3(2): 873 — 899. 2009

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J. Brass, Walter M. Buswell, Frank C. Craighead, Bruce Hansen, Olga Lakela, Robert W. Long, Harold N.

Moldenke, Ruben P. Sauleda, Joseph H. Simpson, Daniel B. Ward, Richard P. Wunderlin, and Alvan W.

Chapman (Chapman 1878; Wunderlin & Hansen 2008; Appendix I). John K. Small admired the Island flora

at the Caxambas region (Austin 1987; Small 1922, 1923, 1929).

Geography, Climate, and Soils

Marco Island, centered at N 25°56'13.5" and W 81°42'57", is situated by the southern border of Collier County,

in southwestern Florida. Encircled by the Gulf of Mexico, the Island is the largest of the Ten Thousand

Islands group. Beriault (1980) cited measurements of 7.3 km along a north-south axis and 8.75 km along

an east-west axis. Elevations range from sea level to 16 m (Beriault 1280)

Marco Island occurs within an area “...overlapped by a humid subt l climate and a tropical nna

climate, in which temperatures are nia by winds from the Gulf al Mexico and the Atlantic Ocean”

(Collier County Facilities Management Department 2007). Yearly, there occur a well defined rainy season

and dry season. Liudahl et al. (1998) reported seven soil types on Marco Island: urban land; urban land-

Immokalee-Oldsmar, limestone substratum, complex; urban land-Aquents complex, organic substratum;

udorthents, shaped; Durbin and Wulfert mucks, frequently flooded; Canaveral-Beaches complex; and, Paola

fine sand, gently rolling. Wilder and McCombs (2006) reviewed the climate, geology, and soils of Collier

County and of adjacent Lee Co.

Historical Sketch

Paleo Indians were the initial inhabitants of Florida, living there 12,000 years ago (Milanich 1994). Beriault

(1980), summarizing previous literature, cited a pottery sherd from Marco Island that had been C-14 dated

(by association with charcoal) as ca. 5150 years old.

In south Florida, The Calusa Indians succeeded the Paleo Indians. A nonagricultural people, the Calusa

Indians hunted, fished, harvested mollusks, and gathered wild plant foods (Gilliland 1989, 1989a; Tebeau

1966). They modified Marco Island, creating canals, a sea wall, reservoirs, and th hell mounds

plus many maler shell mounds. Mounds, composed largely of conch, oyster, and clam shells, were multi-

functional, fuse heaps and platforms. One mound, once located at the present site of Goodland

on Marco Island, eied 1400 by 1700 feet (Beriault 1980; Tebeau 1965).

Marco Island ranks among the most important of archaeological sites in North America because of

Indian artifacts. During 1895-1896 the archaeologist Frank Hamilton Cushing led the famous Pepper-

Hearst Expedition to Key Marco (at the northwestern corner of Marco Island). From muck which Cushing

interpreted as a courtyard, his team excavated superbly preserved Calusa remains: painted wooden masks,

carved figurines, fishing nets, hafted tools, and many other objects (Gilliland 1989, 1989a).

Calusa Indians were in South Florida when Spaniards arrived during the early 1500s, but their civiliza-

tion plummeted afterward because of warfare, disease, abduction by Europeans, and apparently migration.

By 1800 almost all Calusa Indians were gone from the region (Tebeau 1966).

The William T. Collier family was the first permanent non-Indian settlers of Marco Island. Arriving in

1871, they encountered a few “squatters,” but found the Island otherwise cient re deos 2006;

Tebeau 1966). Additional pioneers followed, undertaking farming, construction, and ot ties. Pioneers

mined the shell mounds to build roads.

Three settlements arose on the Island: Key Marco (to the north; this is indicated as Marco Village, in

Fig. 1), Caxambas (to the south), and Goodland (to the east). In 1926 the entire island was incorporated

as Collier City (Tebeau 1966), but that municipality was discontinued. In 1998 the City of Marco Island

was incorporated and now includes Key Marco, Caxambas, and additional land, but not Goodland (which

remains an unincorporated portion of Collier County).

In 1922 Barron G. Collier (not related to W. T. Collier) purchased extensive property on Marco Island.

Barron Collier died in 1939 and his descendants sold their Marco-Island holdings to the Deltona Corpora-

tion (Waitley 1999).

In 1964-1965 the Deltona Corporation initiated a

1 at

y scheme for developing Marco Island,

Wilder and Roche, Marco Island flora 875

North

Marco inn Marco Village

Y) Factory O 0

Pass Old Railroad

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eros 2 Site of Burnham Clam Ten Thousand

Caxambas Factory (Ruins) Islands

Fic. 1. Map of Marco Island as it appeared in 1962. A thick li

of a design and scope likely rivaled in few places elsewhere (cf. Figs. 1, 2). Catastrophic, from an ecologi-

cal standpoint, the scheme included land clearing; road building; the construction of residential housing,

of commercial buildings, of dikes, and of sea walls; dredging, filling, and the concomitant creation of an

extensive matrix of many miles of canals (Waitley 1999). The acreage of the Island was increased from ca.

6,800 to 8,374 acres (Ms. Nancy Richie [City of Marco Island City Hall], personal communication of Dec. 8,

2008; Tebeau 1966). Construction obliterated natural areas, undoubtedly diminished taxonomic diversity,

and probably rendered one species of plants extinct (Lechea lahelae). In the early 1980s, under pressure

from government agencies and environmentalists, the Deltona Corporation ceased activities on the Island

(Waitley 1999).

Seven main kinds of natural habitats occurred on Marco Island prior to development: mangrove

swamps, saltwater marshes, freshwater marshes, pinelands (once covering much of the flat, sandy terrain

in the center of the island), dense xeric scrub vegetation, tropical hammock, and unaltered shell mounds

(Beriault 1980). Quoting from work of J. Miller and M. Fryman, of 1978, Beriault (1980) also indicated *a

dynamic sand beach, shore and lagoon strand."

METHODS

One hundred and eleven field trips were conducted at Marco Island from 2003-2009, with most visits in

2008. The majority of field work was undertaken during the rainy season (lasting from June through Sep-

tember or October) and early and late in the dry season.

We investigiteo ide proad portions of Marco Island, including publicly owned and privately owned

areas. Pri spe luable to this study were Hideaway Beach Golf Club and the Key Marco

ALE EA EE See e

development, ES manifesting cone taxa absent elsewhere on the Island.

Except for noting Sonchus oleraceus, a European weed broadly dispersed within the United States, we

did not inventory plant species at Otter Mound Preserve, a disturbed 2.46 acre shell mound and recently-

established Collier County park situated southward on ithe Island. Ü isi site in a Eu 1900s, this

property was planted repeatedly with native and species planting today. Repeated

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canals contiguous with the Gulf of Mexico.

Wilder and Roche, Marco Island flora 877

plantings have made it questionable which plants grow there naturally, thus making this area untenable for

the present study. James N. Burch—botanist of the Big Cypress National Preserve—had inventoried plants

of Otter Mound Preserve during 2007, recording 127 plant species (Collier County Facilities Management

Department 2007). Burch listed certain species excluded from our own inventory of Marco Island.

In addition to compiling an island-wide inventory, we determined which native taxa occupied each of

the main kinds of natural habitats remaining on the Island (Appendix 2). We did so, by i PES Ee

edly, selected sites of each kind of habitats (Table 1). For coastal tropical hardwood |

we surveyed sites that were fairly uniform in size—each site ca. the size of a housing lot (the sole exception

being a larger hammock; Table 1, hammock no. 3). For these two kinds of habitats, the comparable sizes of

sites studied enabled us to compare, quantitatively, the frequencies of individual taxa present within each

kind of habitats. We surveyed coastal strand vegetation within large areas of Crescent Beach, Hideaway

Beach, Sand Dollar Island (a sand spit protruding into the Gulf of Mexico), and Tiger Tail Beach. We sur-

veyed mangrove habitat along the eastern and western sides of the Island. Pine flatwoods was represented

by a parcel of second growth, described below.

We documented taxa with dried herbarium specimens (Appendix 1). Specimens have been deposited

at the Herbarium of Southwestern Florida (SWF), housed at the Naples Botanical Garden.

We also compiled a list of taxa that previous workers had collected on Marco Island, but that we did

not observe there (Appendix 1), based on sources cited in that Appendix. The list is probably incomplete

because the sources are still being developed (Virtual Herbarium 2008; New York Botanical Garden 2008;

Wunderlin & Hansen 2008).

We characterize taxa as native, alien, and endemic to Florida, according to Wunderlin and Hansen

(2003). Mostly, our nomenclature for species and infraspecific taxa follows Wunderlin and Hansen (2003);

however, Appendix 1 (footnote 1) specifies nomenclatural differences between that work and the present

paper.

Brenda Thomas (of Florida Gulf Coast University [FGCU]) provided GPS readings, using a Garmin 60

CSx GPS unit. Brian Bovard (of FGCU) measured the salinity of a water sample from Marco Lake, using an

MRIOOATC refractometer at room temperature (Milwaukee Instruments, Rocky Mount, NC). Dr. Bovard

undertook two measurements: the first with microorganisms in place, the second with microorganisms

removed by centrifugation (3,700 rpm for 10 minutes).

RESULTS AND DISCUSSION

Natural Habitats Remaining on Marco Island

Mangrove las and one strand Vegetation are the most abundant kinds of remaining natural habitats.

M est tly along/near the shoreline and lagoons. Coastal strand vegetation exists

bris by thes western shore of the Island: at Crescent Beach, Tiger Tail Beach County Park, Sand Dollar

Island, and Hideaway Beach Golf Club.

Previous workers classified coastal strand vegetation in different ways Johnson & Barbour 1990). For

Marco Island, for purposes of simplicity we define two well-delimited zones of beach habitat, collectively,

as coastal strand vegetation: an herbaceous zone and a woody-plant zone. Generally, the herbaceous zone

is seaward of the woody-plant zone, and the two zones typically border and interdigitate with one another.

Based on terminology of the Florida Natural Areas Inventory (1990), the herbaceous zone includes “beach

dune” and “coastal grassland,” whereas the woody-plant 1s to “coastal strand.” The herbaceous

zone exhibits primarily herbaceous species, but also includes opa woody species and sporadic

seedlings, saplings, and stunted individuals of woody species that normally grow larger in other habitats

(e.g., species of mangroves). In the ia eae zone, = and trees are most d

Coastal tropical hardwood hammock and scrubland are rare Hammock diem

the Island and westward at Beene) ae Gan Club. Scrubland ts persist more-or-less medially also.

hwat hes, unaltered shell mounds,

Eliminated totally from Marco I

878 | I £ al n Fi a Um L 3 Pas ri f Texas 3Q)

Tape 1 List of certain of the localities cited in thi ladies — "E zs

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east; N = orth S = south; W = west.

SELECTED REMNANTS OF COASTAL TROPICAL HARDWOOD HAMMOCK

1. N 25° 56.198, W 81° 40.936, + 18 ft; Sheffield Ave. (N)

2. N 25° 56.227, W 81? 41.359, + 16 ft; Sheffield Ave. (S)

3. N 25° 56.303, W 81° 41.138, + 16 ft; San Marco Dr. (N)

4. N 25° 56.133, W 81? 41.616, + 20 ft; Sheffield Ave. (S)

5. N 25? 56.228! W 81° 41.156! + 20 ft; Sheffield Ave. (S)

6. N 25° 55.804, W 81° 40.636, + 16 ft; Whiskey Creek Dr. (W)

7. N 25° 57.766, W 81° 44.545, + 17 ft; Royal Marco Way (E)

SELECTED REMNANT OF PINE FLATWOODS

N 25° 57.089, W 81? 43.428, + 19 ft - Bald Eagle Dr. (W)

SELECTED REMNANTS OF SCRUBLAND

1. N 25° 55.048, W 81? 42.012) + 14 ft; S. Barfield Dr. (E)

2. N 25° 56.282; W 81° 41.299, + 14 ft; San Marco Dr. (S)

3. N 25° 56.253, W 81? 41.407", + 13 ft; Sheffield Dr. (N)

4. N 25° 54.899, W 81? 41.735, + 14 ft; Caxambas Dr. (E)

5. N 25° 55.016, W 81? 41.949) + 14 ft; Inlet Dr. (W)

6. N 25° 56.819, W 81° 43.212) + 15ft; Bald Eagle Dr. (E)

LOCATIONS OF WILD hs OF TRICHOSTIGMA oo TANDRUM

1. N 25° 54.753, W 81° 41.9335 + 16 ft; Addison Ct. (N); a | ls exhibit Trichostigma

2. N 25° 55,044, W 81° 42.001) + 15ft; S Barfield Dr. (E); by disturbed scrubland -

3. N 25° 54.943, W 81° 41.776; 11 ft; Caxambas Dr. (E) hind bland

4. N 25° 54.027, W 81? 41.720" + 19 ft; Caxambas Dr. (E); disturbed land

5. N 25° 54.780, W 81? 41.586'; + 19 ft; Caxambas Dr. (E); c potion O torest

6. N 25° 54.651, W B1? 41.636'; + 18 ft; Caxambas Dr. (N of the Ji t | large field

7.N 25° 55.373, W 81° 41.891'; + 16 ft; Inlet Dr. (E); disturbed la

8. N 25° 54.682, W 81? 41.830'; + 13 ft; Inlet Dr. (E); tree island aah field

9. N 25° 55.169), W 81? 41.945'; + 16 ft; Inlet Dr. (W); disturbed land

10. N 25° 54.913, W 81? 42.147; + 17 ft; Ludlow Rd. (N); disturbed land

11. N 25° 54.947, W 81° 42.274; + 17 ft; Ludlow Rd. (N); disturbed scrubland

12. N 25° 54.786, W 81° 41.734; + 17 ft; Scott Dr. (E); disturbed land

13. N 25° 56.133, W 81? 41.620; + 23 ft; Sheffield Ave. (S); coastal tropical hardwood hammock

14. N 25° 56203, W 81° 41.503 + 22 ft; Sheffield Ave. (S); coastal tropical hardwood hammock

15. N 25° 56.103, W 81? 41.693, + 22 ft; Sheffield Ave. (S); coastal tropical hardwood hammock

and original pine flatwoods. A small parcel of flatwoods exists, as do a few highly disturbed pineland frag-

ments, but the flatwoods is misleading. The parcel was stripped of vegetation and used as a borrow pit.

Vegetation reappeared naturally, as second growth. In 1984, following re-growth of the vegetation, the parcel

was dedicated as a nature park, but the park was abandoned (Natural Areas Management Co., no date; The

Marco Island Eagle 1984).

A few artificial ponds exist. At least one pond (Marco Lake) exhibits fresh water (Brian Bovard; salinity

measurement of Dec. 4, 2008). We noted species at the periphery of Marco Lake that we did not observe

elsewhere on the Island (e.g., Eleocharis interstincta, Ipomoea sagittata, Lygodium microphyllum, Melaleuca

quinquenervia, and Thelypteris interrupta).

Taxonomic Analysis of Present Data

The flora of Marco Island includes 111 families, 324 genera, 461 species, and 470 infrageneric taxa (spe-

cies, varieties, subspecies, and a hybrid; appendix I). Between parentheses, the numbers of families, genera,

and infrageneric taxa are indicated, respectively, for each of the following major groups of vascular plants:

Wilder and Roche, Marco Island flora 879

pteridophytes (10, 13, 19), gymnosperms (1, 1, 1), angiosperms (100, 310, 450), monocotyledons (17, 67,

134), and dicotyledons (83, 243, 316). For infrageneric taxa of each of these major groups, their percent-

age of all 470 infrageneric taxa is listed: pteridophytes, 4.0%; gymnosperms, 0.2%; angiosperms, 95.7%;

monocotyledons, 28.5%; and dicotyledons, 67.2%.

The eight largest families of monocotyledons on Marco Island, as gauged by the numbers of infrage-

neric taxa present, are Poaceae (64), Cyperaceae (34), Bromeliaceae (8), Agavaceae (4), Commelinaceae (4),

Amaryllidaceae (3), Arecaceae (3), and Orchidaceae (3) (for each family, the number of infrageneric taxa is

listed between parentheses). The families Poaceae and Cyperaceae, collectively, exhibited 20.9% of all 470

infrageneric taxa listed (i.e., 98 infrageneric taxa).

The E bacs families of Ola on ne Island, as gauged by the numbers of infrageneric taxa

present, are Fal 42). E j (22), Rubiaceae (17), Amaranthaceae (8), Malvaceae

(8), Myrtaceae (8), abd Convelvulasede (7. The families bae and Asteraceae, collectively, exhibited

18.1% of all 470 infrageneric taxa listed (i.e., 83 infrageneric taxa).

Native a Joram Ta Inventoried During the Present Study

Sixty t of all 470 inf tive to Florida (i.e., 328 taxa; Wunderlin

and Hansen 2003). The centre of native truces taxa within each major group of vascular plants is

listed: pteridophytes (73.796), gymnosperms (10096), angiosperms (69.696), monocotyledons (68.796), and

dicotyledons (69. i

Ten inf tly inventoried are endemic to Florida (Wunderlin and Hansen 2003): Agave

1..T

decipiens, Asimina reticulata, Chamaesyce uo oe lance Croton gl var. floridanus,

hy R} yn hosia cinerea, and Stipulicida

setacea var. lacerat.

Native Taxa and Natural Habitats

Appendix II represents the 221 to 226 native infrageneric taxa that we inventores at rd sites within

the six main kinds of natural habitats on Marco Island (the number of taxa is approxi we could

not assign species names to certain sterile or otherwise unidentifiable specimens).

Each infrageneric taxon inhabited from one to six of these main kinds of natural habitats (Appendix

ID. Crea of Pus Durs. are presented; within each group s ciii d aide sequentially: (a) a

I of (written in script), and (b, c { tage of the 226 taxa which

occupied that number of habitats (indicated with ] d between tl ): one (108, 47.8%); two

(66, 29.2%): PDIEE (29, 12.8%); jour (15, 6.696); five (5, 2.296); and six (3, 1.396). These data manifest pro-

nounced tive correlation between the number of habitats occupied and the number of taxa represented.

Proceedings stepwise through the data, from one to four habitats, each step entails, roughly, an exponential

(ca. two-fold) decrease in the number of taxa.

We designate as especially facultative those 10.2% of taxa that occured in four to six of the main kinds

of habitats surveyed. Chrysobalanus icaco, Sabal palmetto, and Toxicodendron radicans inhabited all six kinds.

Twenty additional taxa occupied sou or five kinds: idi iid: flavescens, Bursera simaruba, Caesalpinea

bonduc, Coccoloba uvifera, Cy igul Dalbergia phyllum, Ficus sp., Fimbristylis cymosa, Iresine diffusa,

Parthenocissus quinquefolia, Piscidia violen: Quercus virginiana, R tata, S , Sideroxylon

celastrinum, Smilax auriculata, Tillandsia fasciculata, Tillandsia usneoides, Vitis sotundiolia: Bud Yucca aloifolia

(Appendix ID.

For coastal tropical hardwood |

of relative frequencies of taxa. For each fraction, the numerator indicates the number of sites of a given

habitat that exhibited a particular taxon, the denominator signifies the total number of sites of that habitat

evaluated. Thus, a fraction equaling the number one indicates a taxon that occupied all sites studied of a

habitat, whereas, fractions of less than one represent taxa present at fewer sites.

Within coastal tropical hardwood hammock, we noted twenty-seven taxa at five to seven (71% to 100%)

of the seven sites investigated: Bursera simaruba, Callicarpa americana, Chiococca alba, Chrysobalanus icaco,

] d scrubland, tl f ti | Appendix Il as measures

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880 fTexas 3(2)

Chrysophyllum oliviforme, Eugenia axillaris, Ficus aurea, Part i quinquefolia, P borbonia, Phlebodium

aureum, Pi idia piscipula, Pleopeltis polypodioides, Psychotria nervosa, Quercus virginiana, Randia nena Rapa-

nea punctata, Rivina humilis. Sabal palmett Shoe chrysophylloides, S , Sideroxylon celastrinum,

Sideroxylon foetidissimum, Tillandsia psal ata, Tillandsia setacea, Tillandsia e Toxicodendion radicans,

and Vitis rotundifolia. Based on our criterion for relative frequency, we rank these taxa as the most frequent

hammock taxa within the sites surveyed. Remaining taxa occupied fewer sites.

Within scrubland, we noted 27 taxa at four to six (67% to 100%) of the six sites investigated: Bulbostylis

ciliatifolia, Bursera lisa dL di K din. dni pes lolas Aa id cumu-

licola, Cyperu humifusa, Palafe i, Polygonella

sola var. brachystadhya ener nervosa, 1, Quercus geminata, Q yrtifolia, Rhus opallinum, Serenoa

repens, Vitis rotundifolia, Callicarpa americana borbonia, Polanisia t f lia, DTE

tenax, Smilax auriculata, Tillandsia MER Tillandsia imiia, ana Ximenia americana.

Applying the same criterion as before, k tl t frequent of scrubland taxa within

the sites surveyed. Remaining taxa —! fewer sites.

We did not observe all native taxa of Marco Island at the

grew within portions of natural habitats outside of these sites Additonal native taxa grew in ruderal habitat

]

in Appendix II. Certain taxa

or in other kinds of disturbed areas.

FLEPPC Species

The Florida Exotic Pest Plant Council (FLEPPC) recognizes two categories of plant species alien to Florida,

that pose especial threats to the ecology of the State, overall, i.e., Category I and Category II (these categories

indicate decreasing degree of threat; Florida Exotic Pest Plant Council 2008).

Alien species abound on Marco Island. During the present study, we inventoried 27 Category I species

and 19 Category II species (i.e., 40.3 96 and 27.196 of all 67 and 70 species in these categories, respectively).

Indicated, in Appendix I are all Category I and Category II species that we inventoried on Marco Island.

Taxa Collected by Previous Workers

Previous investigators collected at least 40 infrageneric taxa on Marco Island that we did not observe (ap-

pendix I, infrageneric taxa listed with bold font). Eighty five percent of these taxa were native (34 taxa).

Four taxa were endemic to Florida: Lechea cernua, Lechea divaricata, Lechea lakelae, and Stylisma abdita. The

40 taxa constitute 7.896 of all 510 infrageneric taxa reported by us and by previous workers.

Small (1922) also reported Pinus clausa and Persea palustris (Raf.) Sarg. and Chapman (1878) indicated

Aristida gyrans n. sp. (at “Robert’s Key”) and Pectis ciliaris L. (at "Colliers Key"); however, neither worker

cited voucher specimens.

Rare Taxa Reported Presently and by Previous Investigators

Rare taxa that we (and in some cases, previous investigators) collected include: Acanthocereus tetragonus,

Acrostichum aureum, Chamaescyce cumulicola, Chrysophyllum oliviforme, Clitoria mariana, Myrcianthes fragrans,

Scaevola plumieri, Scleria ciliata var. curtissii, ess id T. Ga T. flexuosa, T. utriculata, Tricho-

l rare taxa (Table 2, Appendix

stigma octandrum, and Triplasis americana. Previous

I). Table 2 provides assessments of the rarity of these taxa | (Coile & Garland 2003; Gann et al. 2002).

Lechea lakelae is the rarest of species previously reported from Marco Island, and is known with cer-

tainty solely from there. Olga Lakela first collected L. lakelae on the Island in 1964 and participated in most

subsequent collecting of that species. The species was last collected on the Island in 1987 (Gann et al. 2002).

We failed to locate L. lakelae, despite considerable searching, and we suspect that the species is extinct —a

victim of development. Probably, other taxa listed in Table 2 have been extirpated from the Island, also.

The presence of wild Trichostigma octandrum on Marco Island is noteworthy. Previously unreported

from Marco Island, this species of high-climbing vines now grows in at least 15 discrete locations (Table 1).

Plants of T. octandrum also grow at Otter Mound Preserve and we suspect that at least some were formerly

cultivated, based on the regular spacing between them. The wild populations compose two groups on the

Wilder and Roche, Marco Island flora 881

A + H 7 NS T +l | £ f Fl Ir J l

Table 2. List of lant f^ Island. Included,

2003) and for south Florida (Gann et al. 2002). Crit. imp. = critically imperiled; End. — endangered; Hist. — historical; Threat. = threatened.

Taxon Coile and Garland (2003) Gann et al. (2002)

jp pie ua Threat.

Acrostichum um Threat.

Amorpha aes var. herbacea Hist.

Asclepias tomentosa! Hist.

Cenchrus myosuroides Crit. imp.

hamaesyce cumulicola End.

Chrysoph yllum oliviforme Threat.

Clitoria m Crit. imp.

Eleocharis albida Crit. imp.

Gymnopogon chapmanianus Crit. imp.

indigofera trita End. Crit. imp.

Lechea cernua Threat

Lechea divaricata End

Lechea lak En Hist.

let fragrans Threat.

aevola plumieri Threat.

pa ciliata var. curtissii Crit. imp.

Solanum donianum Threat.

Stylisma abdita End. Crit. imp.

Tillandsia balbisiana Threat.

Tillandsia fasciculata nd.

Tillandsia fle Threat

Tillandsia utriculata End.

Trichostigma octandrum End. Crit. imp.

Triplasis americana Crit. imp.

Viola primulifolia Crit. imp.

| s papel | ed one individual of Asclepias t ee eee ee NE

(Wilder & McCombs 29783, a portion of the pia

Island: a southern group situated more-or-less near the Preserve, and a medially located group associated with

Sheffield Ave. These populations grow in remnants of coastal tropical hardwood hammock and scrubland,

on tree island(s) within fields, and within disturbed land (most populations; Table 1).

Trichostigma octandrum exists in Florida at the northern limit of its range, and the species is unknown

outside of Florida in the United States. Southward it ranges from Cuba to Venezuela and Peru (Austin 2004).

Wunderlin and Hansen (2003) considered T. octandrum native to Florida, and Gann et al (2002) ranked this

species as Critically Imperiled in south Florida. Austin (2004), however, concluded that “we do not know’

whether T. octandrum is native or alien in Florida.

Previous workers had reported T. octandrum from five sites in Florida. In 1891 Joseph H. Simpson first

discovered the species on Chokoloskee Island (Collier Co.), a location ca. 23 miles from Marco Island (as

measured from the Caxambas region of Marco Island). Investigators subsequently located it on Pumpkin

Key (Monroe Co.), in Miami (Miami-Dade Co.), and in the Big Cypress National Preserve (county unspeci-

fied). In 2000, Michael Anderson discovered the species in Water Conservation Area 3A (Broward Co.;

Gann et al. 2002). Gann et al. (2002), citing habitat destruction at various locations, considered the plants

at Conservation Area 3A as “... apparently the only extant occurrence in Florida.”

An unpublished discovery of one plant of T. octandrum was made on a shell mound at Everglades Na-

tional Park, by Jesse Hoffmann (Jimi Sadle, pers. comm., 28 Jan 2009; Sadle vouchered this individual [/.

Sadle 553, ENPS]).

[| A £ sl Ds . ..» LA a4

882000 JT tute of Texas 3(2)

We do not know whether Trichostigma octandrum is native to Marco Island. Perhaps, the wild popula-

tions there are escapes from the apparently cultivated individuals at Otter Mound Preserve. The absence of

previous reports/collections of T. octandrum from Marco Island suggests that the wi Id pl t original

In contrast, the proximity of Marco Island to Chokoloskee Island suggests that the Dedes might be native

to Marco Island. If T. octandrum is native on Marco Island, then plants growing at the Preserve may have

been acquired from nature on Marco Island, or they may be the eHsprinig of wild plants there.

Chamaesyce cumulicola, Myrcianthes fragrans, and S la plumieri are fairly common on Marco Island,

yet, Coile and Garland (2003), the Florida Natural Areas inventory (2008), and Gann et al. (2002), collec-

tively, considered these species rare in South Florida.

Many native species are rare on Marco Island that vary from rare to common in Florida. Native species

with one to ca. Seve] individuals observed on the Island during this study include Acanthocereus tetrago-

nus, Acer rubrum, Andropogon ternarius, Aristida condensata, Asimina reticulata, Baccharis angustifolia, Bucida

buceras, Capparis mapraloprera onal ae pe gaa annuum, Carex longii, Chrysopogon pauciflorus,

Chrysopsis scabrella, Cli nashii, Heliotropium curassavicum, Heris-

santia crispa, ewe ao mara — ¡ee sessiliflora, Liatris tenuifolia, as deiade

Osmunda regalis, P Pentalinon luteum, d Pisonia aculeata, Pluchea carolinensis, 5 y

creta, Psilotum nudum, leria E var. curtissii, Se tinata, Smilax bona-nox, Sorghastrum secundum,

Thelypteris kunthii, Tournefortia volubilis, and Vaccinium Sd

CONCLUSIONS

Marco Island exhibits fewer wild taxa than one might expect on the basis of its considerable area and sub-

tropical character. We attribute the apparent deficit of taxa to the extensive development of the Island and

to the elimination of ne water Sica ae Due and original pine flatwoods.

The Island flora it the floras of adjacent portions of mainland

Collier County. (1) PET li is frequent o on Marco Island, yet, we have never observed it in mainland

Collier County. (2) Sideroxy ton Marco Island but is absent or uncommon within main-

land Collier County. (3) eee a reclinatum is frequent in mainland Collier County, but is absent from

Marco Island. (4) P disjunct, albeit ] lation on Marco Island, located

within scrubland. We do not know whether the Population is native Within eastern Florida, the nearest

populations of P. caroliniana occur three counties northward (in De Soto Co. and Sarasota Co.; Wunderlin

and Hansen, 2008). (5) Since its discovery in 1964, Lechea lakelae has been known unequivocally, solely,

from Marco Island.

APPENDIX I !

List of taxa donee for Metco Island (plus coros dela L., which was reconocen Dut not cocumenteg during Uis

study). Non! f ybrid, subspeci ies, and

m no

il 4 * [| J " L j 1 H "M de de J

investigation

listed with Pol font. Solely one voucher Spe en Ue each ieee nel taxon. For nfrageneri taxa documented

f he any rid Prackets)

common name; the Wilder and McComi llecti ber? of a voucher the habitat

collected; and the ae of the taxon, if any, by the Florida Exotic Plant Pest Council For infrageneric tz taxa cientes

solely by previous investigators, data are provide the Latin name of a taxon: relevant synonym, if any (be-

tween brackets); common name; collector: collection number; collection date; and acronym of the herbarium where the

specimen is on deposit? After the name of each family and suprafamilial taxon, between parentheses are Included two or

four numbers; the two numbers not in italics—if present Especial me P To presently of genera

within that Ton e] suprafamilial taxon; it ie two n ics—if present—signify, respectively,

the sums of vorkers, of eue genera and Al taxa. * = alien to Florida;

g = endemic to Florida; FLEPPC land FLEPPC | ¡e = ~~ aecognizsd as Category | or Category li species by the Florida Exotic

Plant Pest Council (2008).

Wilder and Roche, Marco Island flora

PTERIDOPHYTES (13, 19)

BLECHNACEAE (2, 2)

Blechnum serrulatum Rich., Swamp Fern—28501; pine

flatwoods

Woodwardia virginica (L) Sm., Virginia Chain Fern—28505;

pine flatwcods

DENNSTAEDTIACEAE (1, 2)

Pteridium aquilinum var. caudatum (L) Sadeb., Lacy Brack-

445; edge of scrubla

ec aqulinum var. Lice ed € ute) Clute ex de

r, fa lar Bracke JOC HR

mod

LYGODIACEAE (1, 1)

ks cedi ae um (Cav.) R. Br, Old World Climbing

Fer ; climbing on other vegetation rooted in

don Ware Marco Lake; FLEPPC I

PEET ROEE EE (1, a

760127 tul

ofo ak-Sabal ha amm ock

a cordifolia if C. fae Tuberous Sword Fern—

; scruoland; FLEPP

A exaltata (i VU Sword Fern— 27875; coastal

tropical hardwood hammock

*Nephrolepis mulio (Roxb.) FM. Jarrett ex CV. Morton,

Asian Sword Fern— 28752; coastal tropical hardweod

hammock; ae |

OSMUNDACEAE (1, 1)

Osmunda regalis L., Royal Fern—-28489; pine flatwoods

POLYPODIACEAE (2, 2)

Phlebodium aureum (L.) J. Sm., Golden Polypody—28502;

ine flatwoods

Pleopeltis polypodioides (L.) E.G. Andrews €: Windham,

Resurrection Fern—27599; coastal tropical hardwood

hammock

PSILOTACEAE (1, 1)

Psilotum nudum (L.) P Beauv, WHISK Ee 207a along

woody I

vegetation

PTERIDACEAE (2, 3)

Acrostichum aureum L., Golden Leather Fern—28407; along

boardwalk traversing mangrove habitat

Acrostichum danaeifolium Langsd. Ro Fisch., Giant Leaner

ern—28408; at/near periphery o

*Pteris vittata L., Chinese Ladder ces periphery of

coastal tropical hardwood hammock; FLEPPC Il

THELYPTERIDACEAE a, 2)

illd.) K. iwats., Hottentot Fern—28931;

by shore of Marco Lake

Thelypteris kunthii (Desv.) C.V. Morton, Southern Shield

rn—28551; disturbed land

VIETARINCERE (1,1 in

V fet XJ |

cal hardwood ae

883

GYMNOSPERM (1, 1)

PINACEAE (1, 1)

Pinus elliottii Engelm., Slash Pine—28479; pine flatwoods

MONOCOTYLEDONS (67, 134; 73, 147)

AGAVACEAE (2, 4)

Agave decipiens Baker, False Sisal— 28594; disturbed land

*Agave sisalana Perrine, Sisal Hemp—28561; coastal strand

vegetation; FLEPPC Il

*Agave vivipara L., Century Plant—27994; scrubland

Yucca aloifolia L., Spanish Bayonet—28726; coastal strand

vegetation

ALISMATACEAE (1, 1)

Sagittaria lancifolia L., Bulltongue Arrowhead—28943; by

O rco Lake

AMARYLLIDACEAE (2, 3)

Crinum americanum in String-Lily—28742; coastal tropical

hardw ae hammock

*Crinum asiaticum » a eee scrubland

Hymenocalis a (Mill) M. Roem., Mangrove spider-

MMe Se LOPLI I

ARACEAE (2, 2)

*Epipremnum pinnatum (L) Engl, Golden Pothos—29172;

coastal tropical hardwood hammock; FLEPPC II

*Syngonium podophyllum Schott, American Ever-

green—28944; disturbed land; FLEPPC |

ARECACEAE (3, 3)

*Cocos nucifera L, Coconut Palm — coconut with attached

SESION on be eac

S ter) Lodd. ex Schult coo t Cabbage

Palm—29212: list

Serenoa repens (W. Bartram) Small, Saw Palmetto—28625;

pine flatwoods

ASPARAGACEAE (1, ae

‘coastal strand vegetation; FLEPPC I

BROMELIACEAE (1, 8)

Lir odd balbisiana schuit. & Schult. f, Northern esac:

Tillandsia la Sw., Cardinal Airplant—29213; coastal

t ropical parecer ham ock

Ji tar Airplant (2766 AR talt

: LOAF L

cal hardwood hamm ock

Tillandsia paucifolia es Potbelly Airplant—28005; dis-

ed lan

Tillandsia recurvata (L) L., Ballmoss—28006

Tillandsia setacea Sw. Southern Needleleaf—27766; dis-

Tillandsia usneoides (L.) L., Spanish Moss—28271; scrubland

Tillandsia utriculata L, Giant Airplant—29214; found lying

P [| P

CANNACEAE (7, 7)

E T ad ff... H

Olaa Lakela

-—

29067 with D. Laker; 20 Aug 1965 (USF)

884

EAE (2, 4)

*Commelina diffusa Burm. f, Dayflower—28419; disturbed

land

Commelina erecta L., Whitemouth Dayflower—28597; dis-

turbed lan

*Tradescantia pallida (Rose) D.R. Hunt, Purplequeen—28420;

distur be d la nd

RT, hr AES AAA

tropical hardwood hammock

CYMODOCEACEAE (1, 1)

Halodule wrightii Asch., Shoalweed—28814; shallow water

of bay

CYPERACEAE (11, 34; 11, 37)

*Bulbostylis barbata (Rottb.) C.B. Clarke, Watergrass— 28805;

highly insolated, cleared land

Bulbostylis ciliatifolia (Elliott) Fernald, Capillary

Hairsedge— 28320; scrubland

Bulbostylis stenophylla hae C.B. Clarke, Sandyfield

Hairsedge—28938; s

Carex longii Mack., qn E highly insolated

lawn

H c Ç 920712:

40) 123,

Crantz, J g

pine flatwoods

Cyperus compressus L, Poorland Flatsedge—29103; highly

insolated lawn

Cyperus croceus Vahl,

land

*Cyperus esculentus L, Yellow Nutgrass—28819; highly inso-

lated, cleared la

Cyperus Aavescent E Yellow Flatsedge—28900; wet portion

of highly insolated tree lawn

SOYA TE jtag | =

E AANA KAONA

Du ES a l

J ^01 777,.: las Jt

Cyr 1 ' 3

of Marco Lake

Cyperus ligularis L, Swamp bos RE herbaceous

zone of coastal strand vegetatio

Cyperus odoratus L., Fragrant reese Sree oak-Sabal

mmock

Cyperus polystachyos Rottb., Manyspike Flatsedge— 29106;

highly insolated lawn

bos pumilus L; Low Flatsedge—29168; boundary be-

n lawn and brackish wetlan

opens retrorsus Chapm., Pinebarren Flatsedge—29081;

rubland

Or rotundus L., Nutgrass—28499; tree lawn

*Cyperus sphacelatus Rottb., Roadside Flatsedge—28994;

highly insolated lawn

Cyperus squarrosus L., Bearded Flatsedge, Olga Lakela

29238, 1965 (NY, USF)

pen surinamensis HOLD, Tropical pened ee in

d +l

SI | i

zem: eragonis Elliott, Poiana Flatsedge—28854;

sha ergrown scrubland

Eleocharis ‘bids Torr., White Spikerush, Olga Lakela

30960, 196/10 SF)

"m )Chap J Ine A | 1

Olga Lakela 31680, 21 Si 1968 (USF)

Eleocharis cellulosa Torr., Gulf Coast DE inso-

lated portion of mangrove habita

Eleocharis geniculata (L.) Roem. e Schult.,

Spikerush—28626; highly aig lawn

Fleocharis interstincta (Vahl) Roem. & Schult., Knotted

Splkerush— PU o by le of Man Lake

rolina Fimbry—2881 1;

herbaceous ale of coastal ein WO n

U, erica rennracc

Canada

zone of coastal i vegetati lo

Fimbristylis spadicea (L.) Vahl, Marsh m 3; herba-

ous zone of coastal strand vegetation

. | A At Al C +l | l |] J 28939:

hu] fi

shore of Marco Lake

*Kyllinga brevifolia Rottb., Shortleaf Spikesedge—28903;

Fu y) O Ld At cC "J l

e — [i 7 LI -

lawn

*Lipocarpha aristulata (Coville) G.C. Tucker, Awned Halfchaff

Penge e— 28857, wet POR of lawn

alfc haff

Sedge— — 29201; woody- plant zone of ee strand

Hitchc.], S 28514:

coastal strand vegetation

Rhynchospora megalocarpa A. Gray, Sandyfield

pe - E wie nd

Nutrus bes 28942; scrubland

Scleria triglomerata Michx, Tall Nutgrass—28500; scrubland

DIOSCOREACEAE (1, 1)

*Dioscorea bulbifera L., Air-Potato—28821; on mangrove

vegetation by parking lot; FLEPPC |

IRIDACEAE (7, 7)

Sisyrinchium angustifolium Mill., Narrowleaf Blue-Eyed

Grass, F.C. Craighead sn, 24 Mar 1964 (FTG)

JUNCACEAE (1, 1; 1, 2)

Juncus marginatus Rostk, Shore Rush—28948; by shore of

all) JW, Kessler, Curtiss’

Juncus repens Michx., Lesser Creeping Rush, Olga Lakela

0847, date unspecified (USF)

ORCHIDACEAE (3, 3

Encyclia tampensis (Lindl.) Small, Florida pea Or-

chid—29109; coastal tropical hardwood ham

to maculata (Lindl) Lindl., lle ae

of tropical hardwood ham

Bie concreta Jaca) aay & H j^ AEN eae! d

lowspike Orchi

ESAS

POACEAE (33, 64; 36, 70)

Amphi hlenbergi , Blue Maid-

encane—29165; disturbed land

Andropogon glomeratus (Walter) Britton et al., Bushy

Bluestem—29069; edge of scrubland

Andropogon ternarius Michx, Splitbeard Bluestem—29070;

edge of scrubland

Wilder and Roche, Marco Island flora

BROOROQOR CUR i sa Bluestem—28980;

Andropogon virginicus var. glaucus Hack., Chalky

Bluestem, Olga Lakela 31192; 22 Oct 1967 (USF)

Aristida condensata Chapm., Big Threeawn—28933; edge

of scrubland

Aristida patula Chapm. ex Nash, Tall Threeawn—28410; highly

insolated, disturbed land

Aristida a bu var. purpurascens, Arrowfeather

Thre —28889; Scrubland

Axonopus ees hi Kuhlm., Common Carpet-

Olga Lakela 31373; 7 Apr 1968 (USF)

ee pertusa (L) A. Camus, Pitted Beardgrass—28552;

highly insolated, disturbed land

Il OFC. J

between mangrove habitat and parking lot

Cenchrus myosuroides Kunth, Big Sandbur, Joseph H.

2 impson 281, ne 1 (US)

| i | A. Curtis], Coastal

Sandur 2904 herbaceous zone of coastal strand

veget

Chysapagn PUES (Chapm.) Benth. ex Vasey, Florida

d eardgras 5—29043; SIBLE? ana peldcnne

"Cynder cn (L) Pers., Bermudagrass 26447, dis-

rb vergrown portion of bea

ie iU aegyptium (L.) Willd. ex Asch. € Schweinf.,

Durban Crowfootgrass— 28657; disturbed, overgrown

portion of beac

Dichanthelium aciculare (Desv. ex Poir) Gould & C.A. Clark,

M dM cA E d scrubland

Dic ult) G id, Variable witch

rass—28890: ed F 1 | |

"Digitaria bicornis (Lam) Roem. & Senats Asia Crab-

of beach

PUL FAFSA AIS

Digitaria ciliaris (Retz) Koeler: Mar r. ciliaris SOS Crab-

rass—28283 3, hig

Digitaria filiformis (L.) Koeler v var. sid (Walter) Fernald, Sr

der

rahniracc 28564

Lal d»» Z0322

*Digitaria longiflora (Retz.) e m Crabgrass— 28556;

highly insolated, cleared land

Distichlis spicata (L.) Greene, Saltgrass—28981; edge of

mangrove habitat

Echinochloa walteri (Pursh) A. Heller, Coast Cockspur— 28507;

highly insolated, cleared land

is indica (L.) Gaertn., Indian Mods

overgrown portion of bea

E oma (L.) Wight & Arn. ex uk & Arn., Feather

Lov —28714; crack in pavement of roa

“Eragrostis o atrovirens (Desf.) Trin. ex Steud., Thalia Loveg-

27658; disturbed land

eer: ciliaris (L.) R. Br, Gophertail Lovegrass—28284;

ruderal land

Fragrostis pid S. Watson, Elliott's m her-

b e of coastal strand veget

SA MA sandra Presi, Red | n cL field

*Fre a ophiuroides (Munro) Hack., Centipede-

Mika ruderal land

885

Eustachys petraea (Sw.) Desv, Pinewoods Fingergrass—28553;

tree lawn

Gymnopogon chapmanianus Hitchc., Chapman's Skel-

etongrass, Olga Lakela 27527, 1967 (FLAS, USF)

*Hemarthria altissima (Poir. Stapf & C.E. Hubb., Lim-

pograss—28717; highly insolated tree lawn; FLEPPC il

Heteropogon contortus (L) P. Beauv. ex Roem. €: Schult.,

Tanglehead— 28509; field

gi o fusca subsp. fascicularis (Lam. N. Snow, Bearded

angletop—287 18; highly insolated ruderal land

Muhlenbergia gue var. HDS (M E Curtis) Chap ex Beal,

and vegetation

ose hirtellus (L.) P. Beauv., Woodsgrass—28754; coastal

tropical hardwood hammock

A amarum Elliott, Be uu E herba-

eous zone of coastal strand vegetat

*Panicum maximum Jacq., Cures eed 00; highly inso-

lated, d ue) land; FLEPPC H

*Panicum ala L. Au M c. tree lawn; FLEPPC |

Panicum witchgrass— 29166; shaded, disturbed

land

: 7 nicorae Parodi, oe oe field

m, Bahiagrass—28628;

overgrown tree lawn

Paspalum notatum var. saurae Parodi, Bahiagrass—28491;

edge of scrublan

Paspalum setaceum var. ciliatifolium (Michx.) Vasey, Frin-

geleaf Paspalum; nonstereotypical material—28268;

scrubland

Paspalum setaceum var. longipedunculatum (Leconte)

A.W. Wood, Barestem Paspalum, Olga Lakela 31083,

8 Sep 1967 (USF)

Paspalum setaceum var. rigidifolium (Nash) DJ. Banks, Stiff

Paspalum—29016; scrublan

Paspalum setaceum var. stramineum (Nash) DJ. Banks, Yellow

and Paspalum; 29017; scrubland

Paspalum setaceum var. villosissimum (Nash)

is aol: aa SUBANG

|, Va seygrass;

D.J. Banks, Hairy

^"On(107. kh! Ll] $ | " J

law

RUM vaginatum Sw.,

Nu zone of coastal Tang vegetation

C | rn |

20723; herba-

, Olga

Lakela 29244, 6 Oct 1965 (USF)

i n australis (Cav.) Trin. ex Steud., Common Reed—

ruderal land

Mie nd repens (Willd.) C.E. Hubb., Rose Natal-

gra NOU ; Scrublanci MEME UM

Caf, ict] 20077:

EIN E

a i

edge of scrubland

Setaria macrosperma (Scribn. €: Merr.) K. Schum., Coral Bris-

tlegrass—28936; disturbed land

Setaria parviflora (Poir.) Kerguelén, Yellow Bristlegrass—29102;

highly insolated, disturbed land

Sorghastrum secundum (Elliott) Nash, Lopsided Indian-

grass—29078; edge of scrubland

886

Spartina patens (Aiton) Muhl., Marshhay oe 99;

he s zone of coastal strand vegetati

*Sporobolus diandrus (Retz.) P. Beauv., D ve ssibly, S.

jacquemontii)— —28511; ; highly; nsolated bd land

— 28988;

herbaceous zc zone of coastal stand wegen

)Kunth,S 28416:

herbaceous zone of coastal strand vegetation

e secundatum (Walter) Kuntze, St. Augustine-

grass—29019; insolated ruderal land

TN d americana P. Beauv., Perennial Sandgrass— 28937;

scrublan

THp purpurea (aten chapmi Purple sandgrass—28989;

coastal strand vegetation

Uniola paniculata L., Seacats—28559; herbaceous zone of

coastal strand vegetation

Urochloa adspersa (Trin.) R.D. Webster, Dominican Signal-

gras ; periphery of

*Urochloa fusca (Sw.) B. : ~ & Wunderlin, Browntop

Signalgrass—28804

oe subquachpara ‘in R.D. Webster, AAO TaS, Mi

AOorny

RUSCACEAE (1, 1)

*Sansevieria hyacinthoides (L.) Druce, Bowstring Hemp—

28995; disturbed land; FLEPPC H

SMILACACEAE (1, 2)

r | ET | T. 70707 II "m

Smilax bona-nox L. Saw Greenbrier —28822: edge of coastal

tropical hardwood hammock

TYPHACEAE (1, 1)

Typha domingensis Pers., Southern Cattail—29320; edge of

large pond at Mackle Park

XYRIDACEAE (7, 7)

Xyris sible Michx., Shortleaf Yelloweyed Grass, Olga

Lakela 31371, 7 Apr 1968 (FTG)

DICOTYLEDONS (243, 316; 260, 343)

ACANTHACEAE (2, 2)

28451;

lawn; FLEPPC I

ue brittoniana Leonard, Mexican Bluebell—28782; dis-

urbed land; FLEPPC I

ACERACEAE (1, 1)

Acer rubrum L., Red Maple—28516; pine flatwoods

AIZOACEAE (1, 1)

Sesuvium portulacastrum (L.) L., Shoreline Seapurslane

—28074; mangrove habitat

AMARANTHACEAE (5, 8)

Alternanthera flavescens Kunth, Yellow Joyweed—27661;

peripneny ene? moun

A. Gray) J

8598; dried- bb water retention pond

D blitum L., Purple pe a 28904; cultivated

land

€ ri Amaranth

PF da A!

f Texas 3(2)

*Amaranthus hybridus L., Slim Amaranth—28517; highly

insolated, cleared lan

*Amaranthus viridis L, Slender Amaranth—28566; highly

insolated, ruderal land

Blutaparon vermiculare (L.) Mears, Samphire—28461; dis-

turbed, overgrown portion of beach

Froelichia floridana (Nutt.) Moq., Cottonweed—28518;

scrubland

Iresine diffusa Humb. & Bonpl ex x Willd, Juba’s n EE

P A LAD ULAT AAI LA

ANACARDIACEAE (4, 4)

*Mangifera indica L, Mango—27878; coastal tropical hard-

wood hammock

Rhus copallinum L., Winged Sumac—28785; scrubland

*Schinus terebinthifolius Raddi, Brazilian Pepper—28567; edge

stal tropical hardwood hammock; FLEPPC I

Toxicodendron radicans (L) Kuntze, Eastern Poison lvy—2 7879;

coastal tropical hardwood hammock

ANNONACEAE (1, 1)

nAsimina reticulata Shuttlew. ex Chapm., Netted Paw-

pa 517; road-facing edge of scrubland

APIACEAE (1, 1)

Ptilimnium capillaceum (Michx.) Raf., Mock Bishop-

sweed— 29111; in ditch situated within highly insolated

ruderal land

APOCYNACEAE (3, 3)

*Catharanthus roseus (L) G. Don, Madagascar Periwinkle

— both white-flowered forma and pink-flowered

forma-— 29140; disturbed land

Pentalinon luteum (L) B.F. Hansen & Wunderlin, Wild Alla-

a—28692; coastal strand vegetation

Rhabdadenia biflora (Jacq) Mull. Arg., Rubbervine—27880;

mangrove habitat

AQUIFOLIACEAE (7, 7)

Hex dile (L.) A. Gray, Inkberry, Olga Lakela 30795 with

Almeda, 20 Apr 1967 (USF)

ARALIACEAE (3, 3)

Centella asiatica (L.) Urb., Spadeleaf—28452; overgrown

ee lawn

Hydrocotyle umbellata L., Manyflower Marshpenny-

wort—28786; highly insolated, overgrown tree lawn

*Schefflera actinophylla (Endl.) Harms, Australian Umbrella

Tree—28860; coastal tropical hardwood hammock;

FLEPPC I

ASCLEPIADACEAE (4, ^ 4, 5».

land

A E Fad | Lo xs Caaf ILI | pP" |

N. Moldenke 5204, 1930 (NY)

, Madagascar

a A

Pio SASAS | mda

Rubber VIN ie

ated at roadside; FLEPPC I

Cynanchum angustifolium Pers., Gulf Coast swallow

vegetatior 1 situ-

“00731

Ant.

VJI C

Wilder and Roche, Marco Island flora

S | )R & Schult., White Twinev-

ine—28453; highly insolated, cleared land

ASTERACEAE (35, 42; 36, 44

Ageratina jucunda (Greene) Clewell & Wooten, Hammock

Snakeroot—291 13; scrubland

Ambrosia artemisiifolia L., Pa Ragweed— 28454; dis-

rb vergrown portio

aoe ae rss Michx, Saltwater A mre ice

Bacchus glonenifor Pers, Silverling—29050; pine

flatwoo

Baccharis siete. suet a woody-plant

f coastal strand ve

Bidens alba (L.) DC, cuc e lawn

Borrichia frutescens (L) DC., Bushy Seaside Oxeye—29204;

ove habitat

*Calyptocarpus vialis Less., Straggler Daisy—28906; lawn

Chrysopsis scabrella Torr. & A. Gray, Coastalplain Goldenas-

ter—291 15; scrubland

Cirsium nuttallii DC., Nuttall's Thistle —291 16; highly insolated

lawn

Conyza canadensis (L.)) Cronquist, Canadian Horseweed

—29141; ruderal land

nCoreopsis dise thii Torr. & A. Gray, Leavenworth's Tick-

seed 3; highly insolated, cleared lan

EE cinereum (Lò H. Rob, Little Ironweed— 28862;

field

se dl (L) L, False Daisy—291 17; in ditch situated

in highly insolated ruderal lan

ea fase Nicolson, Florida Tasselflower—28455;

developed lan

*Emilia senha (L) DC, Lilac Tasselflower—28569; tree

law

Erechtites ha eed (L.) Raf. ex DC., Fireweed, Olga

9441, 10 Jan 1966 (USF)

A a Poir, Oakleaf Fleabane—28456; over-

grown tree lawn

ai ii capillifolium (Lam.) Small ex Porter & Britton,

Dogfennel—28865; disturbed land

Eupatorium SCION Micnx., Lateflowering Thorough:

wort

VOL ZU Y

abitat

Flaveria linearis Lag., c me woody-

plant zone of coastal strand veget

Gale pulchella Foug., rene! 5205 herbaceous

zone of coastal strand vegeta

"a o ah Cabrera, Pennsylvania

erlasting— 29518;

Non debilis Nutt. Md debilis, East Coast Dune Sun-

flower—28424; scrublan

Heterotheca subaxillaris au Britton & Rusby, Camphor-

weed ;

iva imbricata Walter, ous oe M herba-

us zone of coastal strand veget

Liatris tenuifolia Nutt., Shortleaf nO edge

of scrubland

(L) Small, S S

of overgrown lawn

Mikania scandens (L.) Willd., Climbing Hempvine—28323;

disturbed land

&Palafoxia feayi A. Gray, Feay's Palafox—28973; scrubland

Pectis prostrata Cav., Spreading Cinchweed—28788; tree

lawn

Pityopsis a (Michx.) Nutt., Narrowleaf Silkgrass

—28696; highly insolated lan

Pluchea BA (Jacq.) G. Don, Cure-For-All—29143;

oak-Sabal hammoc

Pluchea quud (L) Cass., Sweetscent—29118; highly inso-

lated law

Pluchea rosea 2n K. Godfrey, Rosy Camphorweed—28951;

shore of Marco Lake

Solidago odora Aiton, Goldenrod—28697; scrubland

la nd

Solidago delega Elliott, debes eb Alvan

middle to late 18 US)

T an " dob PES 28599; shell

mound

me trilobata (L.) Pruski [Wedelia triiobata [L]

c.], Creeping Oxeye—291 19; highly insolated lawn;

Symphyotrichum subulatum (Michx.) G.L. Nesom, Annual

tmarsh Aster—291 14; in ditch situated within highly

insolated ruderal land

*Tridax procumbens L., Coatbuttons— 28457; highly insolated,

ruderal land

Verbesina virginica L., White Crownbeard—28525; bound-

ary between mangrove habitat and highly insolated

cleared land

IRAN:

COUUVU,

) DC, Ori " Ir. ry | L J

lawn

AVICENNIACEAE (1, 1)

Avicennia germinans (L.) L., Black Mangrove—28630; dis-

turbed, overgrown portion of beach

BASELLACEAE (1, 1)

E T RA Ir

elirmkhina Aan el f +l &-l-

har] AS MI vegetation ARS GHG DIRT

BATACEAE (1, 1)

Batis maritima L., Saltwort—28438; periphery of mangrove

habitat

BIGNONIACEAE (2, 3)

*Pyrostegia venusta (Ker Gawl.) Miers, Flamevine—29232;

as scuba

-T Lx] Ji LI LI

E 28458

coastal tropical hardwood hammo ck

*Tecoma stans (L.) Juss. ex Kunth, Yellow Elder—27669; Indian

Hill (a dune)

BORAGINACEAE (2, 4)

Heliotropium angiospermum Murray, Scorpionstail—291 44;

oak-Sabal hammock

Heliotropium curassavicum L., Seaside Heliotrope—28823;

boundary I [ M | bitat | ki lot

888

Heliotropium polyphyllum Lehm., Pineland Heliotrope— 28526;

land

Tournefortia volubilis L, Twining Soldierbush—28825; edge

of coastal tropical hardwood hammock

BRASSICACEAE (1, 1)

Lepidium virginicum L., Virginia Pepperweed—28572; tree

lawn

BURSERACEAE (1, 1)

a simaruba (L) Sarg., Gumbo-Limbo— 27691; coastal

tropical hardwood hammock

CACTACEAE (2, 3; 2, 4)

ip ena tetragonus (L.) neues Barbed-Wire

Cac 9218; disturbed lan

ies ei (L) Mill, ie Cactus—29223;

disturbed land

Opuntia humifusa (Raf. : Raf., Pricklypear—28291; scrubland

paris stricta (Haw.) Haw., Erect Pricklypear, Olga Lakela

540, ae (USF)

CAPPARACEAE (2, 3)

Capparis leal L. damal an eo

eripher

a ledot o i Bayleaf Capertree—28864; coastal

opical hardwood hammock

ms tenuifolia Torr. & "n Gray, Slenderleaf Clammy-

weed— 28601; scrubland

CARICACEAE (1, 1)

*Carica papaya L., Papaya—27965; coastal tropical hardwood

A (2, 2

UNUTLJ | Frenzl ex Walp., American Nail-

—28602; scrubland

"adm setacea var. lacerata CW. James, Pineland Scaly-

pink— 28952; scrubland

CASUARINACEAE (1, 1)

*Casuarina equisetifolia L., Australian-Pine— 28528; disturbed

land; FLEPPC I

CELASTRACEAE (1, 1)

Hippocratea volubilis L., Medicine Vine— 27884; coastal tropi-

cal hardwood hammock

CELTIDACEAE (1, 1; 2, 2)

Celtis laevigata Willd., Hackberry, Olga Lakela 28519A;

1 May 1965 (USF)

Trema micranthum (L) Blume, Nettle Tree— 28548; disturbed

land

CHENOPODIACEAE (4, 5)

Atriplex cristata Humb. & Bonpl. ex Willd., Crested Salt-

bus n" m herbaceous zone of coastal strand

veget

panes ambrosioides L., Mexican Tea—28292; highly

insolated, cleared land

Chenopodium berlandieri Moq, Pitseed Goosefoot—28531;

highly insolated, cleared lan

Sarcocornia perennis (Mill) AJ. Scott, Perennial Glass-

aaa along boardwalk through mangrove

habita

Suaeda -— (Elliott) Moq., Sea Blite—28462; disturbed,

overgrown portion of beach

ILLE (2, 2)

+] E |] La E, La

and the forma with white fruits), Coco Plum—27671;

coastal tropical hardwood hammoc

Licania michauxii Prance, Gopher Apple—28273; scrubland

CISTACEAE (2, 2; 2, 3)

nHelianthemum nashii Britton, Florida Scrub Frost-

weed—28797; highly insolated sand

nLechea cernua Small, Nodding Pinweed; Olga Lakela

31974, 30 Dec 1969 (USF)

»Lechea divaricata Shuttlew. ex Britton, Drysand Pin-

eed, Don Richardson sn, 6 May 1987 (USF)

ees lakelae Wilbur, Lakela's Pinweed, Olga Lakela

7 Aug 1967 (USF

Lechea sessiliflora Raf., Pineland Pinweed—27586; scrubland

COMBRETACEAE (3, 4)

Bucida buceras Lu Black Olive—28632; boundary between

mangrove | t and ruderal land

Conocarpus erectus L., Buttonwood— 29002; mangrove

abitat

Conocarpus erectus var. sericeus DC., Silver Button-

wood—28799; edge of mangrove habitat

Laguncularia racemosa (L.) C.F. Gaertn, White Mangrove

— 28832; mangrove habitat

CONVOLVULACEAE (1, 7; 2, 8)

Ipomoea alba L, q Aare edge of coastal tropi-

cal hardwood hammock

Ipomoea indica (Burm.) m Oceanblue Morning-Glory

— 28463; disturbed, overgrown portion of bea

Ipomoea pes-caprae (L.) R. Br., Railroad EC 28603; herba-

ceous zone of coastal strand vegeta

*Inomoea quamoclit L, Cypressvine— p boundary be-

tween forest and ruderal land

Ipomoea o Poir, Saltmarsh Morning-Glory—28953;

of Marco Lake

*Ipomoed ae L., Littlebell—291 20; highly insolated lawn

ge i violacea L, deal aa Morning-Glory—28870;

of mangrove habita

"Simo iig Myint, Showy Dawnflower, Jim. Burch

iain

CRASSULACEAE (1, 3)

Hamer & H. Perrier, Devil's

_ Backbone—29628 disturbed land

Eckl. & Zeyh , Chandelier Plant— 29427;

disturbed land

*Kalanchoe pinnata (Lam.) Pers., Cathedral Bells—29522;

boundary between disturbed scrubland and ruderal

land; FLEPPC Il

CUCURBITACEAE n e 3,3)

*C ; aloupe , Olgal kela 28539, 1 May,

. 1965 (USF)

Wilder and Roche, Marco Island flora

Melothria pendula L., Creeping oo by board-

walk through mangrove habita

*Momordica charantia L., Balsampea -— coastal tropical

hardwood hammock

EMPETRACEAE (1, 7)

27526; 27 Sep 1964 (USF)

ERICACEAE (2, 2)

Lyonia fruticosa (Michx.) G.S. Torr., Coastalplain Stagger-

bush—28077; scrubland

EUPHORBIACEAE (9, 22)

*Acalypha arvensis Poepp., Threeseed Mercury—29221;

. distur be d lan d

Bic | IRENA: hy

LANAS T, eal A

mangrove habitat, FLEPPC |

Lit L AC

ll, Limestone

Sandma 1—28274; scrubla nd

Chamaesyce bombensis Vaca) SR Pixie SERE

SOLNE

aChamaesyce cumulicola Small Coastal ue ande

mat—281 92; disturbed land

Chamaesyce hirta (L.) Millsp., Pillpod Sandmat—28669;

ruderal land

L oc ft js

r EIC | z

(L) Millsp., 29122:

highly insolated law

Chamaesyce propia (L.) Small, Hyssopleaf Sandmat

—-28670; law

gaa soca (Klotzsch) Arthur, Roadside Sand-

; in ditch situated within highly insolated

ee Bins

Chamaesyce maculata (L.) Small, Spotted Sandmat—28606;

highly insolated gravel

Chamaesyce inns e ie (Jacq.) Dugand, Coastal

each Sa bells 426; herbaceous zone of coastal

stran

ER REM (Pers.) D.G. Burch, Florida Hammock

Sandmat— 28532; ruderal land

Chamaesyce prostrata (Aiton) Small, prostrate Sandmat

9; tree lawn

dccus udis osus (Michx.) Engelm. & A. Gray, Tread

Softly— 289 O; field

| (A.M. Ferguson) R.W. Long,

Vente Conmigo 28758; herbaceous zone of coastal

strand vegetatio

Croton pop var. glandulosus L., Vente Conmigo

A highly ikea disturbed land

OR}. ; hia

AAN P IM 311

insolated gravel

a amarus Schumach. & Thonn., Gale-Of-

Wind—28579; disturbed land

Pha e Roxb., Mascarene Island Leafflower

mes MOM (Murray) Bartl, Paintedleaf—28275;

scrubland

hulla (A VUlatzerh 2, | V Lea

h, Fid-

dler’s Sourge—28580; disturbed land

889

*Ricinus communis L., Castorbean—28834; cleared lana;

FLEPPC Il

FABACEAE (30, 43; 33, 48)

*Abrus precatorius L., Rosary Pea—29053; disturbed land;

FLEPPC I

*Acacia auriculiformis A. EU ex Benth., Farleaf Acacia

—28295; scrubland; FLE

Aeschynomene americana L. ere highly insolated,

overgrown tree lawn

Aeschynomene viscidula Michx., Sticky Jointvetch—28957;

lawn

Alysi lifolius (Sct h. & Thonn.) J. Leonard, False

29092 insolated, cleared land

ad AAIFrT.

Moneywort

Amorpha herbacea Walter var. herbacea, erica

False Indigobush; John K. Small 10487, 1922 (NY)

Caesalpinia bonduc (L) Roxb., Gray Nicker— 29006; herba-

s zone of coastal strand vegetation

Canavalia rosea (Sw.) DC., Baybean—29054; herbaceous zone

of coastal strand vegetation

Centrosema virginianum (L) Benth., Spurred Butterfly Pea—

28328; disturbed land

Chamaecrista fasciculata (Michx.) Greene, Partridge Pea—

27693; disturbed land

Chamaecrista nictitans var. aspera (Muhl. ex Elliott) H.S. Irwin

& Barneby, Sensitive Pea—28533; cleared land

Clitoria mariana L., Atlantic Pigeonwings—28866; edge of

scrubland

*Crotalaria incana L., Shakeshake—27 561; disturbed land

*Crotalaria pallida Aiton, Smooth Rattlebox—28908; dis-

turbed land

, Rattleweed, Ol

Long, F.C. Craighead; 5 Dec 1964 (USF

mun rotundifolia J.F. Gmel., Rabbitbells—28276;

scrubland

Dalbergia ecastaphyllum (L) Taub., Coinvine— 28581; edge

of natural vegetation

Lakela 28682, 29 May, 1965 (USF)

Desmanthus virgatus (L) Willd., Wild Tantan—28700; boundary

n mangrove habitat and field

*Desmodium incanum DC., Zarzabacoa Comun— 28277;

disturbed land

dla tortuosum (Sw.) DC, Dixie NÓ

dist d, overgrown portion

E triflorum (L) DC, Tete erie eae

highly insolated sand

Erythrina herbacea L., AS coastal tropical

hardwood hamm

Galactia Beda a. ie Milkpea—28909; upland

Olaa

, Po

veget

Galactia ice (L) Britton et al., Eastern Milkpea—28837;

scrubland

y^ xvid Les al AC i LARA 28872:

coastal tropical hardwood hammock

Galactia volubilis (L.) Britton, Downy Milkpea— 28873; edge

f scrublan

of scru

*Indigofera hirsuta L., Hairy Indigo— 28296; highly insolated

cleared land

890

*Indigofera spicata Forssk,, Trailing Indigo—28467; lawn

*Indigofera tinctoria L., True Indigo—27632; scrubland

Indigofera trita L. f. [Indigofera mucronata Spreng. ex

DC.], Florida Keys Indigo, Alvan W. Chapman s.n.,

1800s (NY)

EX i i

LAMIACEAE (3, 3)

Callicarpa americana L., American Beautyberry—28339;

scrubland

*Hyptis pectinata (L.) Poit, Comb Bushmint—29057; edge

of lawn

ay get + = pa j i | LE | I pl |

) de Wit, White Leadt 29124;

highly insolated, disturbed land: FLEPPC It

Lupinus diffusus Nutt., Skyblue Lupine—28078; disturbed

land

*Macroptilium lathyroides (L.) Urb., Wild Bushbean—28838;

ighly insolated tree lawn

*Melilotus albus Medik., White Sweetclover—28536; highly

insolated cleared land

Neptunia dl Benth., Tropical Puff—28537; highly

insolated law

Piscidia iE T Sarg., Florida Fishpoison Tree— 28593;

for

pthecellobium unguis cati (L ) Benth Catclaw— Le

and manele habitat

aRhynchosia cinerea Nash, Brownhair Snoutbean—27587;

insolated, disturbed land

Rhynchosia minima (L) DC., Least Snoutbean—28766; field

*Senna obtusifolia (L.) H.S. Irwin & Barneby, Coffee-

weed—28959; insolated, disturbed land

Sesbania herbacea (Mill) McVaugh, Danglepod—28703;

highly insolated tree lawn

*Sophora tomentosa var. occidentalis (L) Isely, Necklacepod

— 27589; scrubland

Sophora tomentosa var. truncata Torr. & A. Gray, Necklacepod

—29009; landward portion of beac

E hamata (L.) Taub., Cheesytoes— 28439; tree

law

ads indica L., Tamarind, Olga Lakela 27763, 23

Vicia acutifolia Elliott, Fourleaf Vetch—29449; tree lawn

Vigna luteola (Jacq.) Benth., Hairypod Cowpea—28468;

disturbed, overgrown portion of beach

FAGACEAE (1, 4)

Quercus chapmanii Sarg., Chapman's Oak—28297; scrubland

Quercus geminata Small, Sand Live Oak— 27676; scrubland

Quercus myrtifolia Willd., Myrtle Oak— 27677; scrubland

Quercus virginiana Mill, Live Oak—28081; edge of coastal

tropical hardwood hammock

GENTIANACEAE (1, 1)

Fustoma exaltatum (L ) Salisb. ex G. Don, Marshgentian

370107.

GOODENIACEAE (1, 2)

Scaevola plumieri (L.) Vahl, daa o herbaceous

zone of coastal strand vegetatio

*Scaevola taccada var. taccada n Roxb., Beach

Naupaka—28610; woody-plant zone of coastal strand

vegetation; FLEPPC I

HYPERICACEAE (1, 1)

| , Fourpetal St John's-Wort,

- Olga Lakela 29521 A, 16 Mar 1966 (USF)

l 28611; woody-

plant zone of coastal strand vegetation

LAURACEAE (2, 2)

Cassytha filiformis L, Love Vine—28298; scrubland

Persea borbonia (L} Spreng, Bay—27636; scrubland

LENTIBULARIACEAE (7, 1)

t Olga Lakela

30952, 7 Aug 1967 (USF)

LOASACEAE (1, 1)

Mentzelia floridana Nutt. ex Torr. & A. Gray, Poorman's

Patch— 28840; field

LOGANIACEAE (1, 1)

Mueola nea (J.F. Gmel) TIE & A, Gray, Lax Hornpoc

| j

astal strand vegetatior L]

LYTHRACEAE (2, 2)

Ammannia latifolia L., Pink Redstem-—291 25; in ditch situated

within highly insolated ruderal land

Ts alatum Pursh, Winged Loosestrife—28538; highly

insolated, cleared land

MARIA EEE (6, 8; di 9)

rizi icky, B

Hill (a hune ie

dicio pentacarpos (L.) Ledeb. [Kosteletzkya

virgini . Presl ex A. Gray], Virginia Saltmarsh

Mallow, Olga Lakela 30793 with F. Almeda, 20 Apr

1807 (USF)

L E abr fT» ADe.e4r- C

Il, False Mallow

—29153; disturbed land (grassy area)

Sida acuta Burm. f, Common Wireweed—28301; insolated

disturbed land

*Sida cordifolia L., Llima—-29058; edge of lawn

Sida rhombifolia L., Cuban Jute—28707; field

*Talipariti tiliaceum var. oo (L.) Fryxell, Mahoe—28930;

natural land; FLEPPC

“o populnea (L.) d ex Correa, Portia Tree—28627;

urbed, overgrown portion of beach; FLEPPC

e lobata L., Caesarweed—28539; periphery of pine

flatwoods; FLEPPC II

MELIACEAE (1, 1)

bes azedarach L., Chinaberrytree— 27892; edge of coastal

opical hardwood hammock; FLEPPC Il

MOLLUGINACEAE (1, 1)

ri verticillata L., Indian Chickweed—28769; highly

solated, cleared land

MORACEAE (1, 2; 1, 3)

Ficus aurea Nutt., Strangler Fig—29209; coastal tropical

hardwood hamm

Ficus elastica Roxb., Rubber-Plant, H.R. Loconte 954 with

J.B. Walker, 12 Mar 1994 (NY)

Wilder and Roche, Marco Island flora

*Ficus microcarpa L. f, Indian Laurel—29059; disturbed land;

FLEPPC I

MUNTINGIACEAE (1, 1)

*Muntingia calabura L., Strawberrytree—29138; oak-Sabal

hammock

50570 + Li ~|

MYRICACEAE (1, 1)

-Ndowhee zv m x | CrArpitharn RPavhorry

MYRSINACEAE (2, 2)

Ardisia escallonioides schiede & Deppe ex a & ENS

Marlb SITEAR

EUER puntata (am) Lundell Myrsine—27699; coastal

tropical hardwood hammock

MYRTACEAE (6, 8)

Eugenia axillaris (Sw. Willd, a Stopper—27727; coastal

tropical hardwood hammock

Eugenia foetida Pers., Lae Stopper—28915; edge of

coastal tropical hardwood hammoc

usd uniflora L., Surinam Cherry— 28471; coastal tropical

hardwood hammock; FLEPPC

*Melaleuca quinquenervia (Cav.) S.T. Biake, Punktree— 29096;

cleared land by Marco Lake; FLEPPC I

Myrcianthes fragrans (Sw. McVaugh, Twinberry— 28135;

coastal tropical hardwood hammock

*Psidium guajava L., Guava—27894; coastal tropical hard-

wood hammock; FLEPPC 1

pine flatwoods; FLEPPC I

*Syzygium cumini a Skeels, Java Plum—29157; disturbed

land; FLEPPC

k., Rose Myrtle-—28540;

NYCTAGINACEAE (2, 2)

Boerhavia diffusa L., Red Spiderling—28472; field

Pisonia aculeata L., Devil's Claws—28920; coastal tropical

hardwood hammock

OLACACEAE (2, 2)

Schoepfia chrysophylloides (A. Rich.) Planch, Graytwig—27728;

coastal tropical hardwood hammock

Ximenia americana L., Tallow code 27683; scrubland

OLEACEAE (1, 1)

Forestiera segregata (Jacq.) Krug & Urb., Florida Swamp-

privet—27684; disturbed land

ONAGRACEAE (3, 5)

Gaura angustifolia Michx., Southern Beeblossom—28473;

disturbed, overgrown portion of beach

Ludwigia E (Jacq.) Raven, Mexican Primrosewil-

low— 28474; highly insolated, cleared an

Es de peruviana (L.) H. Hara, Peruvian Primrosewil-

low— 28963; by shore d Mee Lake; FLEPPC I

Oenothera DUNS Nutt., seabeach ponla

No ADM

EE etation

Oenothera laciniata Hill, Cutleaf enine c

highly insolated sand

OROBANCHACEAE (2, 2)

Buchnera americana L, American Bluehearts (blue-flowered

891

eub irn Ate) Ha + it! H EE les 3 lask j

Hy VISTA LEA

forma) within hig|

ruderal lan d

Seymeria pectinata Pursh, Piedmont Blacksenna—28480;

scrubland

OXALIDACEAE (1, 1)

Oxalis dillenii Jacq., Southern Yellow Woodsorrel —28585;

tree lawn

PASSIFLORACEAE (1, 1)

Passiflora suberosa L., Corkystem Passionflower—28541; pine

flatwoods

PETIVERIACEAE (3, 3)

Petiveria alliacea L., Guinea Hen Weed—28875; edge of tree

island within field

Rivina humilis L, Rougeplant—27729; coastal tropical hard-

wood hammoc

Trichostigma octandrum (L.) H. Walter, Hoopvine—28588;

coastal tropical hardwood hammock

PHYTOLACCACEAE (1

hytolacca americana L, American Pokeweed—28586; coastal

tropical hardwood hammock

PLUMBAGINAEERE (2, UR

[eee a [a || J

—29215; shaded edge of mangrove habita

*Plumbago auriculata Lam., Cape puede edge

of fores

Plumbago scandens L., Doctorbush—27730; edge of coastal

tropical hardwood hammock

O AT (1, 1:

“plant: zone of coastal strand vedere

POLYGONACEAE (4, 4)

*Antigonon leptopus Hook. & Arn., Coral Vine— 28679; dis-

turbed land; FLEPPC li

Coccoloba uvifera (L) L, Seagrape—27900; coastal strand

tanion.

D Ji L I TRA E LiAd I:

October Flower—28542 scrubland

p kL, "| aan | I

VALLI situated VVILIIIHL TL TA higl ly i

PORTULACACEAE (1, 2)

Portulaca oleracea L., Little Hogweed— 28686; highly inso-

ed, ruderal lan

Portulaca pilosa L., Pink Purslane— 28543; highly insolated,

cleared land

PRIMULACEAE (1, 1)

Samolus ebracteatus Kunth, Water Pimpernel—28544; herba-

us zone of coastal strand vegetation

RHAMNACEAE (1, 1)

NE asiatica (L) Brongn., Asian Nakedwood— 29013;

e of mangrove habitat; FLEPPC

RHIZOPHORACEAE (1, 1)

Rhizophora mangle L., Red Mangrove— 28635; by shore of

Fast Marco Bay

892

ROSACEAE (1, 1)

Ail Y Aiton Carolina | aurelcherny

t 4

28710;

scrubland

RUBIACEAE (13, 17)

Chiococca alba (L.) Hitchc., Snowberry—28966; edge of

coastal tropical hardwood hammoc

Diodia teres Walter, Poor Joe—28922; edge of scrubland

Blog HOO" Ls ome POE A in ditch

Enodea littoralis Sw., Beach Creeper—28442; woody-plant

e of coastal strand vegetation

Cum hispidulum Michx., Coastal Bedstraw— 28967;

crubland

Hamelia patens Jacq., Firebush—28878; edge of coastal

tropical hardwood hammo ck

H S il. In

junction of lawn and remnant of ped fone e

wood hammock

*Mitracarpus hirtus (L.) DC., Tropical Girdlepod—28879;

ruderal lan

NA royoc L, bnc NEN edge of coastal tropical

hardwood ham

*Oldenlandia pee ee L., Flattop Mille Graines—28616;

highly insolated gravel

Oldenlandia uniflora L., Clustered Mille Graines—28969;

highly insolated, disturbed land

Psychotria nervosa Sw., Wild Coffee— 27734; coastal tropical

hardwood pein aes

| VAL j; L

SFCEA dm] 3

cal hardwood hammo ck

*Richardia da Gomes, Tropical Mexican Clover

; Shaded area within yar

*Richardia geno (Cham. € Schltdl) Schult. & Schult. f.

geflower Mexican Clover—28303; scrubland

on assurgens Ruiz & Pav., Woodland False Button-

weed—28881; disturbed land

Pi i

} 907 20):

427150

DE L* LI E

V¥ell Hil EL IL d

RUTACEAE (3, 3)

*Citrus x aurantium L., grapefruit group—29219; natural

"Muraya ponit (LJ Jack, Orange Jessamine—28545;

k; FLEPPC H

T E fagara (L) Sarg., Wild Lime—28687; woody-

piant zone of coastal strand vegetation

SALICACEAE (1,

Salix caroliniana MEE Carolina Willow—29160; edge of

mangrove habitat

SAPINDACEAE (1, 1; 4, 4)

Cardiospermum microcarpum Kunth., Heartseed, Olga

Lakela 29241; 6 Oct 1965 (USF

*Cupaniopsis anacardioides (A. Rich.) Radlk., Carrot-

oa coastal tropical hardwood hammock;

FLEP Ci

x T. Durand, Inkwood,

Olga Lakela 29507, 15 Mar 1966 (USF)

[| lafal n.a a mm L I di dg pde nia

f Texas 3(2)

Sapindus saponaria L., Soapberry, Olga Lakela 28514, 1

May 1965 (USF)

SAPOTACEAE (2, 4)

Chrysophyllum di L, Satinleaf—27904; coastal tropical

hardwood hammock

ola a (Kunth) T.D. Penn. Saffron Plum—

8546; bal hammock

Sideroxylon foetidissimum Jacq. False Mastic— 27555; coastal

tropical har mm

Sideroxylon tenax L, Tough Bully—27736; coastal tropical

hardwood hammock

SOLANACEAE bs 5; 4, 6)

Capsicum annu , Bird Pepper— 28713; scrubland

Lycium fiin Wa Iter, Christmasberry—28443; periph-

ery of mangrove habita

Physalis angustifolia Nutt., Coastal Groundcherry—29098;

periphery of scrubland

Solanum americanum Mill, American Black Nightshade

—28433; En to boardwalk through mangrove

habita

Solanum PP”. m Walp., Mu ein Nightshade, Olga

Lakela 29834, 22 May 1966 (US

*Solanum lycopersicum L. (= A esculentum Mill.),

Garden Tomato—29133; edge of food-concession

building

STERCULIACEAE (1, 1; 2, 2)

Melochia spicata (L.) Fryxell, Bretonica Peluda, Olga

Lakela 27744, 22 Oct 1964 (USF)

Rai

d TU tu SA MAy

cleared land

SURIANACEAE (1, 1)

Suriana maritima L., Bay Cedar—28444; woody plant zone of

coastal strand vegetation

TETRACHONDRACEAE (1, 1)

Polypremum procumbens L., Rustweed— 28459; lawn

TURNERACEAE (2, 2)

Piriqueta cistoides (L.) Griseb, Pitted a

MI OVAS LA SUALINA m

*Turnera ulmifolia L., Yellow Alder—28434; disturbed land

URTICACEAE (5, 5)

Boehmeria em (L.) Sw., False Nettle—28971; by shore

of Marco lake

ej " 4 i} \ CH LAL 4) J: LAT J fae | SONED.

ZZ,

L Kits

Parietaria floridana Nutt., Florida Pellitory—29135; in shade

under building

Pilea microphylla (L.) Liebm., Artillery Plant— 28435; by cul-

tivated shrubs

*Pouzolzia zeylanica (L.) Benn., Pouzolz's Bush—28926;

woodland

VERBENACEAE (3, 4)

*Lantana camara L., Lantana— 28340; scrubland; FLEPPC I

Lantana involucrata L., Buttonsag oundary

between the herbaceous zone of, and the eed. plant

zone of coastal strand vegetation

Wilder and Roche, Marco Island flora

Phyla nodiflora (L) Greene, Turkey bis Fogfruit —28477;

J tuned, overgrown portion of bea

insolated, ruderal land; FLEPPC ll

VERONICACEAE (6, 6) :

ipn monnieri l) Pennell, Herb- One i in ditch

VALET bal hl

Capraria biflora Ls Gootweec 29363, boundary between

angrove habitat and tree law

"Conobea lias (Michx.) Penne en Paleseed

2; bare, disturbed land situated by w

ma crustacea (L.) F. Muell., Malaysian ES p

—28925; shrub bed

*Russelia EQUITIS SEDULO & e ais Fountain-

bush SET uc A A

Scoparia dulcis L., E 2; field

893

VIOLACEAE (1, 7)

Viola primulifolia L., Primroseleaf Violet, Olga Lakela

(USF)

VITACEAE (4, 5)

Ampelopsis arborea (L.) Koehne, Peppervine—28972; bound-

ary be RES TES s m j J ] I] J

ticillat colson & CE. Jarvis, Seasonvine— 28549;

edge of bn od hardwood hamm

Parthenocissus quinquefolia (L.) Planch., "d Creeper

— 28550; disturbed lan

Vitis aestivalis Michx., Summer Grape— 28436; scrubland

Vitis rotundifolia Michx., Muscadine— 28280; disturbed land

ZYGOPHYLLACEAE (2, 2)

Kallstroemia maxima (L) Hook. €: Arn., Big Caltrop—28619;

awn

*Tribulus cistoides L., Burrnut—28731; overgrown disturbed

portion of beach; FLEPPC II

E darna md lihan innl mms nrn SAAD 4 £ gf Le £.

Fl z El E

n £ rJ E. Ag Anna. A fe e PA

2 LI b Pi A i ,

Fr iD A = E E £i rc X F |

Li £T LE V

es - P a in r 3 r * e ÍAnNOA| [7 * hn I. N | | :* ] EET I|

s | 4 P

FT. inm m PL Fa we A! Dual LE JTL FARRS unana A Ulsredavlin ard llnn £553 an fh.

Li * d LI

H YT noa Er n ik n In

corniculata |

me - -L

H h In fanart YA L£ nha NM SRP oe a ek ales Lahore wl ie ad Lemon AAAS AL eat oe or CPV TA eros

ICcnx

2 i J V 1 2 Fl

tr p ^^AASY A YY A A aad nnn SARASI ALA n elos LAT | PPA H J

f T

itl ID P FT O A A A E PA O Y O A A on A ee «ac Y PR de T T 1 A | J i LY. Fak}

and Wofford and

i 3 i af

m. RATE n Im n A m. e n de zel, al

CP I - E E n £

varieties)

Ej de TL En L 1

LÀ - LÀ

2 Me P CUIE, L L £ CUIT y P

d - La

^L E

nm £ IE E mnnojr El mp z P" f" rn fawn nbus AASI Vl O amd Liancann [SAAS 4

and Mr

a V PH Y Pr

Inna

Indiaofera trita. Juncus renens

, F 7

comm., 1 Dec 2008). 6 Gann et al. (

*, a

lia. Gann et al. (

APPENDIX II

Tabulation of inventories compiled during this study, of native taxa growing in selected site(s) within each of the six main

kinds of natural habitats remaining on Marco Island. An X signifies the presence of a taxon within a kind of habitats when

all selected site(s) of that kind are represented, collectively (each of HZ, WZ, MANG, PF). A fraction alse eigiallles the occur-

rence of a taxon in a kind of | of CTHH, SCR); the numerator indicates the number of selected | sites exhibiting

that taxon, the denominator signifi es the total uber of selected sites investigated (7 sites from CTHH; b Sites Hom SCR.

Asimina sp. Ficus sp., Galactia sp., Mikania sp., Rhynchosia sp., pang Smilax sp. represent individuals not i S

other ae ie Se herein. HZ

V OE

. J: + +l L

E the PEMA PICU Ir PAÑAL LPI PT A of

KATL L> BRO |

ity ot Li pe

| T | " BAAS

| | al strand vegeta nio! ly we OQ MALIS AWI VI CN LO DIO!

TAXON HZ WZ MANG CTHH PF SCR

Acer rubrum X

Acrostichum aureum X

Acrostichum danaeifolium X 1/7

Alternanthera flavescens X X X 1/6

mbrosia artemisiifolia X X

Ampelopsis arborea X

Andropogon glomeratus X X 1/6

1/6

Andropogon ternarius

Sry)

894 f Texas 3(2)

TAXON HZ WZ MANG CTHH PF SCR

Andropogon virginicus X X 1/6

Ardisia escallonioides 4/7 1/6

Aristida cond 1/6

Aristida patula X

Aristida purpurascens X 3/6

imina S 2/6

Atriplex cristata X

Avicennia germina X X

Baccharis Semen X

Baccharis halimifolia X X 147

copa monnieri X

Batis maritima X

Bidens alba X X 2/6

Blechnum serrulatum X 2/f X

Blutaparon vermiculare X X

Borrichia frutescens X

Buchnera americana X

Bulbostylis ciliatifolia 6/6

Bulbostylis stenophylla 2/6

Bursera simaruba X X 7/7 X 4/6

Caesalpinea bonduc X X X 1/7

Callicarpa americana n X 5/6

Canavalia rosea X X

Cassytha filiformis X 6/6

Cenchrus spinifex X X 6/6

Centrosema virginianum X 2/6

Chamaecrista fasciculata X X 6/6

hamaecrista nictitans var. aspera X

Chamaesyce blodgettii X

Chamaesyce bombensis X

Chamaesyce cumulicola 4/6

Chamaesyce mesembrianthemifolia X

Chiococca alba X X 55

hr a anus X X X 6/7 X 3/6

Chrysophyllum oliviforme 5/7 X

Chrysopsis scabn 1/6

Cissus verticil X 1/7 X

ladium jamaicense X

Coccoloba uvifera X X X 4/7 1/6

Commelina erecta X X 1/6

Conocarpus e b X X

Conocarpus erectus var. sericeus X

Conyza canadensis X 1/6

Coreopsis leavenworthii X

rinum americanum 4/7

Crotalaria rotundifolia X

Croton glandulosus var. floridanus X X 3/6

Croton glandulosus var. glandulosus X

Cynanchum angustifolium X

Cyperus croceus 1/6

Cyperus ligularis X X 1/7 1/6

in dpi E X

Cyper X X 6/6

Cype oa 1747

o ean X X X 3/7

Wilder and Roche, Marco Island flora

895

TAXON

MANG

CTHH

SCR

Di chanthelium aciculare

Ernodea littoralis

Eupatorium serotinum

Eustachys petraea

Ficus aurea

Ficus sp.

Fimbristylis ie

Fimbristylis osa

Fimbristylis spadicea

Galactia regularis

Galactia sp.

Galactia striata

Galactia volubilis

Galium hispidulum

Gaura angustifolia

Helia nthus debilis su ih debilis

Hippocratea volubilis

Laguncularia racemosa

Lantana involucrata

Liatris tenuifolia

Lyonia fruticosa

Melothria pendula

Mikania scandens

Mikania sp.

ed eel

Morinda royoc

Mea Aii var. filipes

lali s fragra

AN erifera

NOIRS exaltata

Oenothera humifusa

Oplismenus hirtellus

KKK X

2X OD& DX X

5/7

52.

2/7

1/7

3/6

1/6

2/6

1/6

2/6

4/6

2/6

1/6

5/6

4/6

1/6

896 J Bot f Texas 3(2)

TAXON HZ WZ MANG CTHH PF SCR

Opuntia humifusa X X 6/6

Osmunda regalis X

Palafoxia feayi 6/6

anicum amaru X

Paronychia americana 1/6

Parthenocissus quinquefoli X X X 5/7 2/6

Paspalum setaceum var. ciliatifolium X

Paspalum setaceum var. rigidifolium 2/6

Paspalum setaceum var. stramineum X 1/6

Paspalum setaceum var. villosissimum 1/6

Paspalum vaginatum X

Passiflora suberosa 3/7 X

Pentalinon luteum X

Persea borboni 6/7 5/6

Phlebodium aureum 5/7 X

Phyla nodiflor X

Physalis angustifolia X X 2/6

Phytolacca american 2/7

Pinus ellio X 1/6

Piriqueta cistoides X

Piscidia piscipula X X X 6/7

ithecellobium unguis-cati 4/7

Pityopsis graminifolia 1/6

Pleopeltis polypodioides 5/7

P dorat X

scandens 2/7 1/6

Poinsettia cyathophora X X 2/6

Polanisia tenuifoli 5/6

olygala grandiflora X X

Polygonella polygama var. brachystachya 4/6

olypremum EE X

Prunus iip) 1/6

Psilotu um 3/7

ea nervosa UT X 4/6

Pteridium aquilinum var. caudat 1/7

Pteridium aquilinum var. inc 2/7 X

Quercus chapmanii 1/7 3/6

Quercus geminata 6/6

Quercus myrtifolia 6/6

Quercus virginiana X 6/7 X 2/6

Randia aculeata X X 1/1

anea p X X LU X 2/6

Rhabdadenia biflora X 3/7

Rhizophora mangle X X

Rhus copallinum X 1/7 6/6

Rhynchosia cinerea X

Rhynchosia sp. 1/6

Rhynchospora colorata X

Rhynchospora megalocarpa 1/6

ving humilis 5/7 2/6

abal palmett X X X A X 1/6

Salix caroliniana X

Samolus ebracteatus X X

Sarcocornia perennis X

Sarcostemma clausum X X

Wilder and Roche, Marco Island flora

TAXON HZ WZ MANG CTHH PF SCR

Scaevola plumieri X X

Schoepfia chrysophylloides 5/7

Scleria ciliata var. curtissii 1/6

Scleria triglomerata 3/6

Serenoa re, X 5/7 X 6/6

Sesuvium portulacastrum X X

Setaria corrugata 3/6

Seymeria pectinata 1/6

Sida acuta X 1/6

Sideroxylon celastrinum X X X 5/7

Sideroxylon foetidissimum 517

Si 1/7 5/6

Smilax auriculata X X X 5/6

Smilax bona-nox 1/7

Smilax s 4/7 1/6

Solanum americanum X

Solidago odora 2/6

Solidago sempervirens X X X

ophora tomentosa var. truncata X X

Sorghastrum secund 1/6

Spartina pat X

Spermacoce assurgens X

Sporobolus domingensis X X

Sporobolus virginicus X

Stipulicida setacea var. lacerata 1/6

Suaeda linearis X X

Suriana maritima X X

Tillandsia balbisiana 4/7

illandsia fasciculata X X 5/7 3/6

Tillandsia flexuosa X 1/7 3/6

Tillandsia paucifolia X 177 1/6

Tillandsia recurvata X 4/7 5/6

Tillandsia setacea X 6/7

Tillandsia usneoides X X 5/7 4/6

Tillandsia utriculata X 3/7 3/6

Tournefortia volubilis 1/7

oxicodendron radicans X X X 7/7 X 1/6

Trichostema dichotomum X X

Trichostigma octandrum 1/7

Triplasis americana 1/6

Triplasis purpurea X 1/6

iola paniculata X X

Vaccinium myrsinites 1/6

Verbesina virginica X

Vigna luteola X X

Vitis aestivali 3/7 1/6

Vitis rotundifolia X X 7/7 X 6/6

Vittaria lineata 4/7

dai se indica X X

oodwardia virginica X

imenia americana 3/7 5/6

Yucca aloifolia X X 1/7 X

4/7

Zanthoxylum fagara

DEDICATION AND ACKNOWLEDGMENTS

We dedicate this paper to environmentally concerned citizens of Marco Island, with the hope that it will

foster increased appreciation and understanding of their botanical heritage.

We express deep appreciation to Brian Holley and the Naples Botanical Garden for providing laboratory

space for the present study and for housing the Herbarium of Southwestern Florida. We are grateful to Kate

Kintz, a graphic designer who modified Figs. 1 and 2. We also thank the following individuals and institu-

tions for their assistance: Brian Bovard; Chris Brighton; Ronald Echols; Susan Gallagher; Bruce Hansen,

John Kunzer, Richard Wunderlin, and the Institute for Systematic Botany (University of South Florida);

Melissa Hennig: Brandon Jones; Jeannette Kling; John Mickel; William Marquardt; Jacob Mitchell; Elizabeth

Perdichizzi and the Marco Island Historical Society; Nancy Richie, Julie Ross, and the City of Marco Island;

Pamela Riggs; Tom Ryan and Hideaway Beach Golf Club: Eva Schliesser; Brenda Thomas; Gerry Tsandoulas,

Marie Tsandoulas, and the Key Marco Community Association, Douglas Waitley; Laura Wewerka; Joseph

Wipff; and Lynka Woodbury.

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BOOK NOTICE

NEAL K. VAN ALFEN, GEORGE BRUENING, AND JAN E. Leach (eps). 2009. Annual Review of Phytopathology, Volume

47. (ISSN 0066-4286; ISBN 978-0-8243-1347-0, hbk). Annual REVIENS, Inc., Ns E] Camino Way,

PO. Box 10139, Palo Alto, California 94303-0139, U.S.A. (Orders: ww org, science?

annualreviews.org, 1-800-523-8635, 1-650-493-4400). $84.00 TM 445 pp., 7 5/8" x 9 3/8".

Contents of Volume 47:

l. OE Before You lain a a a ne in ird Plant Pathologist— Michele C. Heath

2. —Sally A. Miller, Fen D. Beed, & Carrie Lapaire Harmon

3. Diversity, pathogenicity: a Management de ia Species—Steven J. Klosterman, Zahi K. Atallah, Gary E. Vallad, & Krishna

V. Su diri

4 Bact /F g l Interactions: F

5. Community Ecology of Fungal Pathogens Causing Wheat Head Blight—Xiangming Xu & Paul Nicholson

6. The Biology of Viroid-Host Interactions—Biao Ding

7. Recent Evolution of Bacterial Pathogens: The Gall-F

8. Fatty Acid-Derived Signals in Plant Defense—Aardra Rachmoo & radar Kachroo

9. Salicylic Acid, a Me ee to amiga pee = Corina Vlot, D’Maris Amick Dempsey, & Daniel F. Klessig

RE d E bionts— Donald Y. Kobayashi & Jo Anne Crouch

L uc oy | 1 e RÁ T CA ones

T p

OL, LIAI LLOPL NS

NÀi and F M.N. Rosso, J.T. Jones, & P. Abad

11. Fungal Eff Prot I los & Pie Wit

12. Durability of Resistance in Tomato ad AM to Fes mien" Causing Bacterial Spot—Robert E. Stall, Jeffrey B. Jones, & Gerald

V. Minsavage

13. Seed Pathology i din in oo na NE. ary P. Munkvold

ts and Divergence—Maurice Moens & Roland N. Perry

f Root-Knot N d David McK. Bird, Valerie M. Williamson, Pierre Abad, James McCarter, Etienne G.J. Danchin,

" Migra ator P

pe Castagnone-Sereno, & Charles H. Opperma

16. Viruses of Plant me dan AM Á. Gari e pi Suzuki

7. 1 ] wO. JE i ci Pi Lim Me iai dd Bragg, l Uma Ganesan, & Mi Yeon Lee

, Armin Djamei, &

M ovement,

18. Ustilago ms as a Pathogen—-Thomas Brefort, a Doehlemann, A

egine Kahmann

J. Bot. Res. Inst. Texas 3(2): 900, 2009

THYMELAEA PASSERINA (THYMELAEACEAB) IN SOUTH DAKOTA

Grace Kostel

Black Hills State University

1200 University Street, Unit 9003

erbarium, J159

Spearfish, South Dakota 57799-9003, U.S.A.

GraceKostelabhsu.edu

ABSTRACT

Thymelaea passerina (L.) Cosson & J. G in (Thymel ) is reported ddition to the fl f South Dakota. A description of

+1 E IG a f. 1

1 d Les | ]

1 L Eu

Key Wonps: Thymelaea passerina, Thymelaeaceae, naturalized, Conata Basin, South Dakota, invasive species

RESUMEN

Thymelaea passerina (L.) Cosson & J. Germain (Thymelaeaceae) se presenta como adición a la flora de Dakota del Sur. Se describe la

E 4

INIRODUCTION

In the summer of 2008, the author discovered a population of Thymelaea passerina (common name, spurge

flax), a species thought to be unknown from the state, to be well etalon ona a distursed site in Conata

Basin, Pennington County, South Dakota. The population of ca. 1000 i ttered throughout

several acres of Conata Basin clearly indicates a potential for naturalization. Although no published record

of T. passerina in South Dakota exists, a later search through herbarium records revealed a collection made

in 1989 by Craig Freeman and Ralph E. Brooks in Tripp County. A formal description of the species and

new information about its known distribution in the state follow.

i ad cam Lov U.S.A. SOUTH rinm SET Co.: Midi R76E $4 NW p 7 mi W of Colome, level to slightly rolling

11 Jul 1989, R.E. Brooks 19582 & C. Freeman

(KANU, KS, U.S. 298531).

R i U.S.A P gton Co.: T3S R16E $23. 43.7707000 N; -102.1685000 W (Datum = NAD83),

Buffalo e National sini nae nia ca. 14 mi Me S of we and 0.5-1 mi E » Conata Basin Road, elev. 2570 ft, prairie dog

d heavy use, ca.

town, soils overlain

1000 plants Seded over several acres, A Sep 2008, Grace M. Kostel 12666 (BHSC, SD, U.S. 41571).

MORPHOLOGY

Thymelaea passerina is a taprooted herbaceous annual to 55-60 cm tall and characteristically slender, wiry

and erect. Spurge flax grows as one main stem or, more commonly, branches from the upper portion of the

main stem. Leaves are alternate and sessile, leathery, and jointed at a yellow cartilaginous base; blades are

small, narrow and linear, 8-14 mm long, progressively smaller upward along the stems. The flowers are

perfect, greenish, tubular and axillary, 2-3 mm long, with 4 sepals, no petals, and 8 stamens (in 2 whorls

of 4). Below each flower, two very small bracts arise from a tuft of tiny white hairs. The fruit is one-seeded,

the enclosed seed round, brown to black and 2-3 mm long (McGregor 1986; Kaul et al. 2006).

DISCUSSION

Thymel ina was first reported in the United States by Pohl in 1955. Thymelaea passerina is native to

Northern iim Europe and Asia where it is considered a common weed of dry soils and grain fields (Pohl

J. Bot. Res. Inst. Texas 3(2): 901 — 903. 2009

902 | | tanical hl Texas 3(2)

1955; Wofford & De Selm 1988). It is also found in central Russia, Australia and North America. In the

U.S., it is recorded from Iowa and Nebraska (Pohl 1955), Illinois (Mohlenbrock & Ladd 1978), Wisconsin

(Harriman 1979), Kansas (McGregor 1986), Ohio (Vincent & Thieret 1987), Mississippi (Wofford & De

Selm 1988), Alabama (Webb et al. 1997), and Texas (Holmes et al. 2000). In Washington state T. passerina

was identified in 1996 in Okanogan County and was subsequently listed as a Class A noxious weed (USDA

NRCS 1999).

Introduction to Tripp County may have been facilitated through agricultural activity associated with

grazing and feeding cattle, e.g., introduced as seed in hay. We can only speculate as to how T. passerina may

have been introduced to Conata Basin. Several years of activity associated with the recreational killing of

prairie dogs (Cynomys ludoviciana) and the actions iated with the reintroduction of black footed ferrets

(Mustela nigripes) by the U.S. Fish and Wildlife Service (USFWS) and U.S. Forest 5ervice (USFS) in Conata

ectahlickhend in a reeion

Basin may have played a role. K ledge of how long an introduced si

is important. Assessments of a species' invasiveness can be made too soon, for example, thus increasing the

likelihood of labeling a potentially invasive species in its lag phase as “safe.” Such mistakes are common

(IUCN 2002). Lag phases of up to 180 years have been found for herbaceous species (PySek & Prach 1993).

In Washington State, Class A plants, such as T. passerina, are required by law to be given highest priority.

Such effort is consistent with the state's purpose that nonnative species whose distribution is still limited

are best dealt with by preventing new infestations and eradicating existing ones as soon as possible.

Thymelaea passerina can be difficult to identify. In the field, the plant closely resembles some Polygonum

species but upon close examination, no ocrea (a papery sheath) is found at the leaf nodes. Spurge flax is

the only species in Thymelaea and the only herbaceous annual Thymelaeaceae, thus it is absent from many

taxonomic keys. Officials in Washington have observed that animals will not graze this plant. Alkaline

rangeland and disturbed places are typical of sites where T. passerina has been found (USDA NRCS 1999),

and such habitats are common to the complex and highly altered Conata Basin ecosystem. Spurge flax is

reported to be difficult to control by herbicides due to the lack of surface area of the small, leathery leaves.

A KANU specimen label for T. passerina (KANU 298425, and many duplicates at NEB, collected in 1950)

allude to its weediness by indicating the plant had *almost completely taken over a pasture" in the northern

part of Cedar County, Nebraska.

CONCLUSION

Thymelaea passerina is clearly established in South Dakota and has the potential for wider naturalization.

The 1989 collection introduced a new family, Thymelaeaceae, to the flora of the state. Thymelaea passerina,

or spurge flax, has proven to be aggressive and difficult to control in other regions. Livestock do not graze

the plant, and at Conata Basin, none of the ca. 1000 plants observed were noted to have been grazed by

prairie dogs. An assertive control program at Conata Basin makes management, and possibly eradication,

of spurge flax feasible at this time. Given the relatively isolated geology of Conata Basin, it may be possible

to contain the alien and prevent it from spreading. In addition, monitoring by the USFS of the Conata Basin

flora would greatly aid in identifying new invasions. A return to the site of the initial 1989 find in Tripp

County could also provide valuable information about its persistence and invasive pattern.

ACKNOWLEDGMENTS

Biodiversity occurrence data provided by: University of Kansas Natural History Museum, R.L. McGregor

oh the efforts of S lf j 1] Caleb Morse. Si iation is

Appreciation is extended to Craig Freeman for verifying the data of his initial collection in South Dakota

in 1989 with Ralph Brooks. The author is grateful for the work of M.L. Gabel and Peggy Buckwalter on the

Spanish translation.

Kostel, Thymelaea passerina in South Dakota 903

REFERENCES

GREAT PLAINS FLORA ASSOCIATION. 1986, Flora of the Great Plains. University Press of Kansas, Lawrence.

Harriman, N.A. 1979. Four additions to the Wisconsin flora. Michigan Bot. 18:143-145.

Hormes, W.C., J.F. Pruski, AND J.R. SINGHURST. 2000. Thymelaea passerina (Thymelaeaceae) new to Texas. Sida

19:403-406.

IUCN. 2002. Turning the tide: the eradication of invasive species. Veitch, C.R. and M.N. Cloud, eds. Occasional

Paper of the IUCN Species Survival Commission No. 27. Available at: http//www.hear.org/articles/turning-

thetide/turningthetide.pdf.

KAUL, R.B., D.M. SUTHERLAND, AND S.B. RoLrsmelER 2006. The flora of Nebraska. University of Nebraska, Lincoln.

McGrecor, R.L. 1986. Thymelaeaceae. In: Flora of the Great Plains University Press of Kansas, Lawrence. P 498.

MOHLENBROCK, R.H. AND D.M. Lapp. 1978. Distribution of Illinois vascular plants. Southern Illinois University Press,

Carbondale.

POHL, RW, 1955. Thymelaea passerina, a new weed in the United States. lowa Acad. Sci. 62:152-154.

Pyšek, P. AND K. Prach. 1993. Plant invasions and the role of riparian habitats - a comparison of four species alien

to central Europe. J. Biogeogr. 20:413-420.

USDA, NRCS. 1999. The PLANTS database (http://plants.ussda.gov/plants) accessed 4/19/2009. National Plant

Data Center, Baton Rouge, LA 70874-4490 USA.

VINCENT, M.A. AND J.W. THiereT. 1987. Thymelaea passerina (Thymelaeaceae) in Ohio. Sida 12:75-78.

Weep, D.H., H.R. DeSeLM, AND W.M. Dennis. 1997. Studies of prairie barrens of northwestern Alabama. Castanea

62:1 73-184.

WOFFORD, B.E. AND H.R. De Sem. 1988. Distribution of and first report of Thymelaea passerina from the southeastern

United States. Castanea 53:305-306.

904

BOOK NOTICE

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 Garden Press, 4344 Shaw Boulevard, St. Louis, Missouri 63110-

2291, U.S.A. (Orders: www.mbgpress.org, mbgpress@mobot.org, 1-314-577-9534, 1-314-577-9594

fax). $375.00 set, Vol. 1: 984 pp., Vol. 2: 1302 pp., Vol. 3: 1602 pp., 8 1/2" x 11 1/4".

Vol. 1. Pteridophyta, Gymnospermae, Monocotyledoneae

Vol. 2. Dicotyledoneae: Acanthaceae—Fabaceae (Abarema-Schizolobium)

Vol. 3. NN Fabaceae buda dd i aaa

1 1 Cal r +1 1 £ 1 1 tot ld_temnerate

E e I L NET ek prie Ex tenter

areas of ed Aa E Cono mi Or pde ODE hich includ tl 4,700,000 sq kil t The Introduction (in

1 LE YA. - 1 . E z1

EE od I O [e] i [e]

pw of 17, 697 pted d, al tl that in the United Stat hl i More

11 ther t luded f tl ] d dubi citations and names. At least one

than 42,000 synonyms are indicar ed,

Mene ee each or the dd names. The de ed is y n alphabetically 5 anys eta and aaa and documented

, habit, elevation,

and MU CMS synonymy.

From the publisher: “TI i | i i , 3 I

+1 + 1 de

GLE VERSE LLLA M

ao pa]

tion, overharvesting, or global warming.”

J. Bot. Res. Inst. Texas 3(2): 904. 2009

ANNOTATED LIST OF MARYLAND GRASSES (POACEAE)

Edward E. Terrell and Pau! M. Peterson

Department of Botany

National Museum of Natural History

mithsonian Institution

Washington, D.C. 20013-7012, U.S.A.

terr60@msn.com; peterson@si.edu

ABSTRACT

oo names, common names, and widely = sy ae for 347 species of g found in Maryland are reported. We provide

| y, distribution,

RESUMEN

Se divulgan | ] ptados, ] , y los sinónimos ampliamente utilizados para 347 especies de pastos encontradas

en Maryland. Proporci inf ión d istribución, y estado de la conservación

The grass family (Poaceae or Gramineae) is the fourth largest flowering plant family in the world and con-

tains about 11,000 species in 800 genera worldwide. Twenty-three genera contain 100 or more species or

about half of all grass species, and almost half of the 800 genera are monotypic or diatypic, i.e., with only

one or two species (Watson & Dallwitz 1992, 1999). Specializations for open habitats and grazing toler-

ance, highly reduced floral structure, and wind pollination in the grasses have enabled the family to be

extremely successful at planet-wide radiation and colonization. One notable feature of grasses and other

monocots are intercalary meristems that allow individual culms to resprout once they have been removed.

Grasses are well adapted to open, marginal, and frequently disturbed habitats, and can be found on every

continent, including Antarctica.

With the infusion of molecular data our present concept and classification of the grasses is changing at

a rapid rate. In Maryland we currently eight subfamilies: i El leae, Pooideae,

Arundinoideae, Danthonoideae, mom Ca and Panicoideae Cas Pieces Working

Group 2001; Soreng et al. 2009).

Over the last 120 years or so the flora of the eastern United States has been well documented. In Mary-

land, local treatments such as the flora of the District of Columbia and vicinity (Hitchcock 1919), Maryland

grasses (Norton 1930), herbaceous plants of Maryland (Brown & Brown 1984), a checklist of plants in the

Washington-Baltimore area (Hermann 1946), an annotated checklist of vascular plants of Rock Creek Park

(Fleming & Canal 1995), an annotated list of the flora of Beltsville Agricultural Research Center (Terrell

et al. 2000), and an annotated list of vascular plants of the Washington-Baltimore area (Shetler & Stone

Orli 2002) have vastly improved our distribution records. Regional treatments have also contributed to an

environmental awakening making the identification of grasses more accessible (Fernald 1950; Brown &

Brown 1984; Gleason & Cronquist 1991).

The natural flora is always in a tenuous position since its survival is often in the hands of bureaucrats,

politicians, and ultimately developers. PUEDE of our ever changing flora is critical to the conservation

of our plant biodiversity, including tl ing of native, introduced, and invasive species. We hope this

annotated checklist will be useful to everyone who is concerned with maintaining our natural heritage and

that it will serve as a useful step toward subsequent lists.

sas

J. Bot. Res. Inst. Texas 3(2): 905 — 919. 2009

906 | t tani fTexas 3(

METHODS

We recognize four zones in Maryland: the coastal plain, piedmont, upper midland, and mountain, The

coastal plain is east of Montgomery, Howard, Baltimore, Harford, and Cecil counties (Fig. 1) extending to

Delaware and the Atlantic coast and including the lands surrounding the Chesapeake Bay. The piedmont,

termed the lower midland zone in Brown & Brown (1984), includes all of the above-named five counties

and the eastern part of Carroll County. The upper midland zone extends west through Washington County

and a small part of Allegany County. This zone includes the northern extension of the Blue Ridge province

and part of the Ridge and Valley province. The mountain zone includes most of Allegany County and all of

Garrett County. This zone often has elevations above 1500 feet and its highest point is 3360 ft.

This annotated list is based on reports taken from publications and from collections at the University

of Maryland (MARY) and the United States National Herbarium (US) at the National Museum of Natural

History, Smithsonian Institution. The main source of distribution data was taken from Brown and Brown

(1984) and often the first data listed here for a species have been taken from this source. In addition we

consulted the Flora of North America, volumes 24 and 25 (Barkworth et al. 2003, 2007), a comprehensive

treatment of the grasses. We cite Reed (1964) frequently because he investigated plants found on chrome,

manganese, or other ore piles in the Canton area of the port district of Baltimore. Many of these species

were first reports for North America or new to the DC area. Shetler and Stone Orli (2002) who produced a

list for the DC area was also extensively consulted.

We include data from the Maryland Department of Natural Resources (list downloaded on 13 Nov

2007) concerning rare, threatened, and endangered species. There are no Maryland grasses federally listed

as Endangered or Threatened. Seventy-three grass species are listed, including mostly species that are

state-rare, regardless of how common they are in the other sections of their ranges, e.g., Zizaniopsis miliacea

is very rare in Maryland, but frequent to common in the remainder of its range, Maryland to Florida and

Texas and north to southeastern Missouri. Many other Maryland grasses have similar ranges, and are rated

G5 for “demonstrably secure globally.” In the present list we are concerned with species that are “globally

rare”. Consequently, we list here and in the text only 14 species that are designated G3 (very rare and local

or with a restricted range) or G4 (secure globally put oy mare): i lial indi amphicarpon (G4), Bromus

nottowayanus (G3-G5), Calamagrostis porteri (G4), D bsp. aciculare (G4-G5), D. aciculare

subsp. angustifolium (G4-G5), D. acuminatum subsp. da ae D. n ous D. Mi sea

(G4), Muhlenbergia glabriflora (G4), M. t (G3), P lp} hiladelphicum (G3-G5),

Poa alsodes (G4-G5), P. saltuensis subsp. lancia (G3- G4), nl pensylvanica (64.

The following abbreviations are used in the list: BARC- Beltsville Agricultural Research Center; DC-

District of Columbia; DOV- Phillips Herbarium, Delaware State College, Dover, Delaware; MD- Maryland,

MARY- University of Maryland herbarium; TAWES- Herbarium of the Maryland Department of Natural

Resources, Annapolis, Maryland; and US- United States National Herbarium. Accepted names appear in

bold, synonyms appear in regular type, and an asterisk * designates introduced species.

ANNOTATED LIST OF GRASSES OF MARYLAND

AEGILOPS ; SOLES Saner, somerset, Wicomico a (all ocr oe

*Aegilops. triuncialis L. - barbed goat grass. Baltimore, chrome Harvey

"m piles maar 1964). nd (Te Il collecti ) ¡probably

ery and g

occurs more widely in MD.

(AGROPYRON Gaertn.) A is elliotti hult — Elliott b d

Agropyron repens (L) P. Beauv. = Elymus repens grostis elliottiana Schult. — Elliott's entgrass. Reporte

from Charles m (Shetler & Stone Orli 2002). No col-

AGROSTIS L. bentgrass. lections at MARY

Agrostis alba of authors = Agrostis gigantea *Agrostis gigantea aie redtop. (A. alba L., misapplied).

*Agrostis canina - — velvet bent. A lawn grass reported Common and widespread.

from Frederick C Agrostis hyemalis (Walter) Britton, Sterns, & Poggenb. —

*Agrostis capillas L - Rhode Isiand bentgrass, colonial ticklegrass. Common

bentgrass. Collections at US: Montgomery, Prince

907

Allegany

‘ae

E» T ol i

£ ~ Washington E

E zi A 7

phe

Fic. 1. Location of counties in Maryland.

Agrostis mertensii Trin. — northern bent. Reported by Reed

(1986).

Agrostis dan (Walter) Tuck. - upland bentgrass.

Frequent in s and coastal plain.

Agrostis unr Willd. — rough bentgrass. (A. hyemalis var.

scabra (Wilid.) Bloma. n widespread.

Agrostis spica-venti L. — Apera spica-venti

o is 5 stolonifera 7 - E bentgrass. Widespread

VIL Paik ist ha

eun tenuis Sibth. = A. canion

iia L. nalrgrass,

T :

canillaric

pry ar. Caplllaris

“Aira a caryophyllea L

Dj Koch) Bh Œ 9, Ei

cate hairgrass. Rare, Worcester Co. and south.

var. caryophyllea - silver hairgrass. Frequent on coastal

E in. Common at BARC, Prince George's Co. (Terrell et al.

00).

Aira pu Willd. ex Roem. & Schult. = A. caryophyllea

var. capillaris

*Aira praecox L. — spike hairgrass. Prince George's, Calvert

Worcester cos. Rare on southern coastal plain.

ALLOTEROPSIS J.P esi

*Alloteropsis cimicina (L.) Stapf - Baltimore, ore piles (Reed

1964). (Coridochloa cimicina (L.) Nees ex B.D. Jacks.)

ALOPECURUS L.

Alopecurus aequalis Sobol, var. aequalis — xir Lead foxtail.

Rare, Cecil Co. Prince dd Co, two collections

BARC (Terrell et ai. 200 Itural

ment on Delmarva, ke adventive Frye).

tufted fox

an agricultural weed of field edges (Frye).

*Alopecurus geniculatus L. — marsh foxtail. Rare. Dorchester

tail. Infrequent,

Co. and DC area. Prince George's Co., Oxon Run (Shetler

& Stone Orli 2002

*Alopecurus myosuroides Huds. — slender foxtail. Rare,

DC area. Baltimore, chrome ore piles (Reed x Prince

George's Co, Beltsville (Shetler & Stone Orli 2

"Alopecurus id A haa Dorchester

COP

ema"

gomery Co, Terrell lesión

AMMOPHILA Host Beachgrass.

Ammophila arenaria (L.) Link — European beachgrass. Oc-

casionally introduced. Reported by Kartesz & Meacham

(2008).

nal

mMmarcan

Ocean and beaches and dunes. ie s to

prevent erosion

AMAN Kunth

cane. Locally abundant coastal eli G4.

F T

mphicarpon

ANDROPOGON L. lom

Andropogon elliottii Chapm. = A. gyra

Andropogon gerardii Vitman - big e Infrequent

throughout. Widely planted in warm season grass mixes

Andropogon glomeratus (Wa!ter) Britton, Sterns, &

Poggenb.

var. glaucopsis (Elliott) C. Mohr - Reported by Kartesz €:

Meacham (20

08).

var. glomeratus — bushy bluestem. Common, coastal

plain.

var. hirsutior (Hack) C. Mohr - Reported by Kartesz &

Meacham (2008).

908

Andropogon gyrans Ashe var. gyrans - Elliott's bluestem.

Coastal plain.

AK At. | cL? L

- F -

n |] > har! ,

Coastal plain, infrequent.

Andropogon vi rginicus L.

war VIFAInRIrI Ig

abundant.

A LE Ll Las l.i TI IBI

A glomeratus

ANTHOXANTHUM L

i sui hirtum (Schrank) Y. schouten &Veldkamp

Reported by Kartesz & Meacha 8).

*Anthoxanthum nitens (Weber) Y. Praed & Veldkamp

- vanilla sweetgrass. Hierochloé odorata (L.) Wahlenb.

treated as synonym in Allred & Barkworth (2007: 762).

Rare. Baltimore Co, Soldiers Delight, 4 May 1980, Reveal

& Broome 5133 (MARY).

Ant nie : — sweet vernal grass. Abun-

APERA Adans.

*Apera spica-venti (L) P. Beauv. - Weed in grass plots, College

Park, Prince George's Co

APLUDA L.

*Apluda mutica L. — Baltimore, chrome ore piles (Reed

ARISTIDA L. Three-awn grass

Aristida adscensionis L. — Baltimore, chrome ore piles

eed 1964).

Aristida curtissii (A. Gray) Nash = A. dichotoma var. curtissii

Aristida dichotoma Michx.

var. curtissii A. Gray — Curtiss's three-awn grass. Uncom-

mon, coastal plain and piedmont. Prince George's Co,

BARC (Terrell et al. 2000).

var. dichotoma - Common, coastal plain and piedmont.

n lanosa Muhl. ex Elliott — woolly three-awn grass.

, eastern shore

mh longespic

var. geniculata pir Fernald — Reported by Kartesz &

Meacham (2008).

var. longespica — Frequent, coastal plain and lower

piedmont.

Aristida oligantha Michx. - pe m -awn grass. Fairly

mmon, coastal plain and DC a

Aristida purpurascens Poir.

var. purpurascens — purple three-awn grass. Frequent,

coastal plain

var. virgata (Trin ) Allred — Rare, eastern shore. Five collec-

tions from Wicomico and Worcester cos. in TAWES (Frye).

Aristida tuberculosa Nutt. — seasice three-awn grass. Rare,

Assateague Island, Worcester Co.

ARRHENATHERUM P Beau

*Arrhenatherum elatius (L.) P. Beauv. ex J. p & C. Pres|

var. elatius — tall oatgrass. Naturalized, sometimes planted.

Patner Intrequent, mostly in the Pippo DE

oa IM C +l

Ma”

eastern "n

fay

f Texas 3(2)

ARTHRAXON P. Beauv.

*Arthraxon hispidus (Thunb.) Makino - arthraxon, small

carp grass. Common and widespread wee

UNDINARIA Michx. Cane

Arundinaria tecta (Walter) Muhl. — switch cane. Rare,

coastal plain

ARUNDO L. Reed

*Arundo donax L. - giant reed. Grown as ornamental. Estab-

lished in Wicomico Co.

AVENA L. Oats

*Avena fatua L. - wild oats. Rare. Baltimore, chrome ore

piles (Reed 1964).

*Avena sativa L. — Oats. Cultivated, sometimes escaped and

established

AVENELLA (Bluff & Fingerh.) Drejer

Avenella flexuosa (L) Drejer — crinkled hairgrass. Frequent,

northern coastal plain and mountains.

AXONOPUS P. Beauv.

Axonopus furcatus (Flüggé) Hitchc. - big carpetgrass.

mon, coastal plain. None in MARY, US. Locally

abundant in Somerset Co. (Frye).

BOTHRIOCHLOA Kuntze

*Bothriochloa pertusa (L) A. Camus - Baltimore, chrome

ore piles (Reed 1964).

DOES Lag: rama m

*n altimnara chrome

ore piles (Reed 1964).

*Bouteloua barbata Lag. - Baltimore, chrome ore piles

eed 1964).

Bouteloua curtipendula (Michx.) Torr. — side-oats grama.

Rare, ridge and valley (Frye). Montgomery, Baltimore,

Washington, Allegany cos. (Brown & Brown 1984)

Bouteloua hirsuta Lag. — Reported by Kartesz & Meacham

(2008)

BRACHIARIA (Trin.) Griseb. = UROCHLOA, in part

Brachiaria distachya (L.) Stapf = Urochloa distachya

BRACHYELYTRUM. P. Beauv.

Brachyelytrum aristosum (Michx.) P. Beauv. ex Branner &

Coville — Garrett Co. (Saarela et al. 2003).

l Iré n. (Cr kRiIDPp Eairlu ramman

Ud y VELIT TAT M,

piedmont and mountains.

BRIZA L. Quaking grass.

*Briza media L. — Rare. No data (Brown & Brown 1984).

*Briza minor - little quaking grass. Rarely established; As-

sateague Island, Worcester Co. (Brown & Brown 1984).

MUS L. Brome grass

*Bromus arvensis L. — field bromegrass. Rare, Prince Georges,

orcester, and Talbot cos.

*Bromus briziformis Fisch. & C.A.Mey. - awnless brome. Rare,

Baltimore and Dorchester cos.

*Bromus catharticus Vahl — rescue grass. Uncommon. Prince

George's Co., BARC (Terrell et al. 2000

e ciliatus L.

ciliatus — fringed bromegrass. Reported from Mary-

End needing taxonomic study (Frye).

Bromus ciliatus var. laeviglumis Scribn. = B. pubescens.

*Bromus commutatus Schrad. — d chess, meadow

brome. Common and widespread w

*Bromus diandrus Roth - ripgut grass. E he rare, inner

coastal plain. (B. rigidus in Brown & Brown

Bromus dudleyi Fernald- B. ciliatus

*Bromus hordeaceus L. subsp. hordeaceus - soft chess,

soft brome. Known from several counties (B. mollis in

Brown & Brown 1984). Prince George's Co., BARC (Terrell

et al.

*Bromus inermis Leyss. - smooth brome. Sometimes grown,

but established widely in a variety of habitats

*Bromus japonicus Thunb. - Japanese brome, Japanese

Ss Known from four counties but c common at in

BARC, Pr ince George's Co. (Terrell et al, 2

more widely distribute d.

*Bromus kalmii A. Gray — Rare, Garrett Co.

Bromus latiglumis (Shear) Hitchc. — Rare, coastal plain and

along Potomac River.

*Bromus madritensis L. Cultivated as ornamental. Listed by

(2o

7 Dp

Bromus nottowayanus cun Rare, Baltimore and Mont-

gomery cos. (Brown & Brown 1984). In calcareous habitats

throughout Maryland (Frye). G3—G5

Bromus pubescens Muhi ex la Canada brome. Com-

tt | Allegany Cos, jaiten

n trace -iliatiicwar laavials imis (B.

& Brown 1984).

Bromus purgans L. = B. pubescens

romus racemosus L. - Uncommon, reported from Anne

Arundel, Baltimore, HONO Kent, Prince Georges, Mont-

gomery and Queen Anne

Bromus rigidus Roth — B. dia ndra

*Bromus rubens L. — foxtail " Rare, reported from

orcester Co.

*Bromussecalinus L.—chess, cheat. Emos cua eae:

*Bromus sterilis |. — brome Uncom-

Weed in Prince Georges Co, BARC (Terrell et al.

20. In 2008 abundant in Frederick Co. and in parts of

Montgomery Co. (seen by Terrell).

*Bromus tectorum L. — downy chess, downy brome. Com-

mon we

Bromus erm Kunth = B. a

Bromus villosus Forssk. = B. diandru

CALAMAGROSTIS Adans. Reed Bentgrass.

- bluejoint. Infrequent. Garrett, Montgomery, and Prince

George’s cos. (Brown & Brown 1984)

Calamagrostis cinnoides (Muhl.) W.P.C.Bart.) — small

reedgrass. A proposal is planned to preserve this name

over the name €. coarctata Torr. ex Eaton (Marr et al, 2007:

a 7). AIOE ERA:

i A. Gray subsp. porteri — Reported by

Kartesz & Meacham (2008). G4.

909

CENCHRUS L. Sandbur.

aco echinatus L. — Baltimore, chrome ore piles (Reed

iud

SE LI

fio tel candhi If Infra-

quent. Prince Georges Co, BARC (Terrell et al. 2000).

Cenchrus spinifex C astal sandbur. Reported by

Kartesz & escam ien

Cenchrus tribuloides |. - dune sandbur. Common in

coastal plain.

(CHAETOCHLOA Scribn.) = SETARIA

CHASMANTHIUM Link

Chasmanthium latifolium (Michx.) H. O. Yates - Indian

wood oats. (Uniola latifolia Michx.) Frequent in Potomac

drainage.

Chasmanthium laxum (L.) H. O. Yates — slender wood

oats. (Uniola laxa (L.) BSP). Chiefly in coastal plain where

frequent.

HLORIS Sw. Finger Grass.

"Chloris "Ps Nutt. - windmill grass. Established

n field adjoining library in BARC, Prince George's Co.

Gel et al. 2000).

LEVAS of an

| 1964).

L HI ry

CINNA L. Woodreed.

Cinna arundinacea L. - Woodreed. Common throughout.

Cinna latifolia (Trevir. ex Gópp.) Griseb. - drooping wood-

reed. Infrequent in mountain zone

COELORACHIS Brongn.

= E L" fee

b i

+h, + |

Brown 1984). No specimens from MD in MARY.

Rare

CYNODON Rich.

*Cynodon dactylon (l) Pers. - Bermuda grass. Planted in

lawns and widely established as a weed.

CYNOSURUS L. ica Grass.

*Cynosurus cristatus L. — crested dog's-tail grass. Rare

weed. E Park, ADS George's Co. (Shetler & Stone

Orli 2002).

ricus echinatus L. - Rare weed reported from MD.

DACTYLIS |. Orchard Grass.

*Dactylis glomerata L. — orchard gr

spread. Sometimes cultivated.

DACTYLOCTENIUM Willd.

*Dactyloctenium aegyptium (L.) Willd. — Rare or absent.

One collection Pond s.n. (MARY) as a weed in farm plots,

rk.

| wide-

enium m Hack. - Baltimore, chrome

ore es (Reed 1964).

ANTHONIA Lam.& DC. Wild Oatgrass.

bunten compressa Aust. - mountain oatgrass. Com-

mon in western MD, also Frederick, Montgomery, Prince

George’s, and Calvert cos.

Danthonia sericea Nutt. - downy oatgrass. Common in

Dorchester, Wicomico, and Worcester cos. and in coastal

plain.

910

f! YD Baarns

ex Roem. & Schult. — poverty

grass. Common and widespread.

DASYOCHLOA Willd. ex Rydb.

*Dasyochloa pulchella (Kunth) Willd. ex Rydb. — Baltimore,

chrome ore piles (Reed 1964).

DESCHAMPSIA P. Beauv. Hairgrass.

Deschampsia caespitosa (L.) P. Beauv. — tufted hairgrass.

Reported from Baltimore Co. No collections in MARY, US.

Several collections at TAWES (Frye)

Deschampsia flexuosa (L.) Trin. Avenella flexuosa

DIARRHENA P Beauv.

l I " lc AA I LO J

65), recent coll t | llb D obovata

Diarrhena obovata (Gleason) Brandenburg - obovate

orasan Collected during ds D 20 from Mont-

C Lea, (DOV. US). RJ. S tl || | tl

American Legion Bridge and at Bear Island, T

Co, personal collections.

Gene nets (Hitchc. & a Saar

CI Ibspecie

(1984), Some species listed by Guala & Becca (2004) are

rece M Lelong determined some taxa collected in

Prince George's Co.

acia aciculare (Desv. ex Poir) Gould €: C.A.Clark

- narrow-leaved panicgrass.

subsp. aciculare. — Rare in coastal plain. G4—G5.

subsp. angustifolium (Elion) rreckmann . od m

var aciculare in Brown & Brown (1 984) NG data. G4— (35.

hantheli inat (S \G Id&CA Clark - hairy

panicgrass.

subsp. acuminatum (var. acuminatum) — Infrequent in

coastal plain.

subsp. columbianum (Scribn.) Freckmann & Lelong. As

: ica var. ad OS 8 MA Goulia a c A.

us fasciculatum (Torr) Freckmann & ene

Widesprea

subsp. implicatum (Scribn) Freckmann & Lelong. Pied-

mont and coastal plain.

subsp. leucothrix (Nash) Freckmann & Lelong. As D.

acuminatum var. implicatum pu ) Gould €: C.A. Clark

in Brown & Brown i 984). No dat

subsp. | lash) Frecl & Lelong. Frequent

subsp. longiligulatum (Nash) Freckmann & Lelong.

Coastal plain.

subsp. spretum (Schult) ae is Leong AS D.

darncifl Tarn

KA Sa F

f Texas 3(2)

Dichanthelium aes (L) Gould - deer-tongue

grass. Common and widespread.

Dichanthelium ON pom Gould - variable

panicgrass.

subsp. ashei (T.G. Pearson ex Ashe) Freckmann & Lelong.

No data

subsp. commutatum - Frequent.

Dichanthelium depauperatum (Muhl.) Gould — dei

panic grass. Frequent in coastal plain and piedmon

Dichanthelium dichotomum (L.) Gould - forked panic em

bushy panic grass.

subsp. dichotomum - commons

(A & Lelong. No data.

FRA Ll ril t ee

subsp lucidum

subsp

- Com

en pr AA (Ashe) Freckmann & Lelong. No

dat

elas ensifolium (Baldwin ex Elliott) Gould —

sword-leaf panic grass. No data.

Dichanthelium latifolium (L.) M — broad-leaved panic

9 rass. cl n piedmon

fd aM E E +16

Pile LI

icgrass Rare in MD

| m. H L FN I I EI ga b

En: ic 21 T g |]

panicgrass, low panic grass. No data.

Dichanthelium meridionale (Ashe) Freckmann. Reported by

Kartesz € Meacham (2008). = D. acuminatum subsp.

implicatum

ge | H són | IC ml iey rn

X J

panicgrass.

subsp. oligosanthes — euer coasta p

subsp ik i Freckmann

frequent, piedmont.

PC le (Elliott) Gould & C.A.Clark — stiff-leaved

panicgrass.

subsp. ovale - Rare, coastal dunes.

subsp 2 lalana

i f 2

hare;

hE a " SAT LAr | Do! |

g.No data

Dichanthelium polyanthes (Schult) Mohlenbr. - any-

flowered panic grass. As D. sphaerocarpon var. da

La Gould & C.A. Clark in Brown & Brown 1984).

Com

Siena portoricense (Desv. ex Ham.) B.F. Hansen

& Wunderlin

subsp. patulum (Scribn. € Merr) Freckmann & Lelong —

_ blunt-giumed panicgrass. is data,

Anarri Eml Ravenel's

LATE

_ panic grass. pale in » coastal eb

mJ Gould & C.A. Clark

var. Ries D. portoricense subsp. patulum

var. thinium = D. acuminatum subsp. columbianum

è Clarki in Brown and Brown Pee pastal | piein.

Dichanthelium scabriusculum (Ell) Gould & C.A. Clark —

Uncommon in coasta! plain. Wicomico Co. G4.

Dichanthelium scoparium ano e E - ey pan-

fy my rmxm a a!

us boreale (Nash) Freckman — iiri panic

grass.

enel basal (Poir) Gould € C. A. Clark — Bosc's

panicgrass. Common.

ctal nlain

Dichanthelium sphaerocarpon (Elliott) Gould - round-

fruited panicgrass. Comm

E tenue (Muhl.) Freckmann & Lelong - Re-

Kartesz & Meacham (2008).

ohana villosissimum = D. ovale subsp. villosis-

pichanthelium wrightianum (Scribn.) Freckmann — Re-

orted by Kartesz € Meacham (2008). G4.

DIGITARIA Haller Crab Grass, Finger Grass.

Digitaria ciliaris (Retz) Koeler — southern crabgrass. Prob-

ably common.

a cognata (Schult, Pilg. — fall witchgrass. (Leptoloma

cognata (Schult.) Chase). Anne Arundel Co. (Shetler &

Stone Orli 2002), Prince George's Co., BARC (Terrell et al.

2000). Frequent on coastal plain (Frye).

—,

Digitaria filiformis (L) Koeler — slender cra Frequent,

Stal plain

*Digitaria ischaemum (Schreb.) Muhl. - smooth crabgrass.

mmon and widespread weed.

Digitaria longiflora (Retz) Pers. — Baltimore, chrome ore

piles (Reed 1964)

*Digitaria sanguinalis (L.) Scop. — hairy crabgrass. Common

and widespread weed.

DINEBRA Jaca.

*Dinebra retroflexa (Vahl) Panz. — Baltimore, chrome ore

piles (Reed 1964).

DISTICHLIS Raf. Saltgrass.

Bo spicata (L) Greene - seashore saltgrass. Common

n salt marshes, coastal plain

CHINOCHLOA P Beauv.

tana corona (L) Link — awnless Deity ae grass.

Balti 1964).

Acre Co., Riefner “ 721 (MARY). Prince Georges Co,

BARC (Terrell et al. 200

*Echinochloa crusgalli " P. Beauv. - barnyard grass. Com-

mon and widespread.

*Echinochloa crus-pavonis (Kunth) Hitchc. - Baltimore,

chrome ore piles (Reed 1964

Echinochloa frumentacea bes Reported by Kartesz &

Meacham (2008).

D D YC IA Da ra in AAD

Lo. Le xt MA GITS HIY

var. mi Wiegand Prince Georges Co, BARC

Terrell et al. 2000).

var. muricata — Prince George's Co,BARC (Terrell et al.

2000).

Echinochloa walteri (Pursh) A. Heller - Walter's barnyard

rass. Common, coastal plain.

ELEUSINE Gaertn. Goose Grass.

*Eleusine indica (L.) Gaertn. — goose grass. Common weed

especially on sidewalks and paved places.

LYMUS L. Wildrye.

Elymus canadensis L. - Canada wildrye. Limestone regions,

northern counties, Prince rine Talbot cos.

mus hystrix L. — bottlebrus

ar nystu- - Widespread. (sr patula Meench).

LIO

911

El ii R. Brooks & JJ.N. Campb. - early wildrye.

- Reported. by! Kartesz & Meacham (2008).

*Elymus repens (L.) Gould - quackgrass. Common and

widespread weed.

Elymus biais Min — riverbank wildrye. Frequent,

piedmont and Worcester Co.

Elymus trachycaulus tin Gould

subsp (Link) A. Love & D. Lóve — Reported

by Kartesz & Meacham (2008).

subsp. trachycaulus — Reported by Kartesz & Meacham

2008).

Elymus villosus Muhl. ex Willd. — slender wildrye. Infrequent,

piedmont and coastal plain.

Elymus virginicus

var. halophilus (E P. t Wiegand. - Reported by

Kartesz & Meacham 1 (20

var virniniet Ig \

VOl. Wil we Ms i"

a S | r 4 EA | J

a Des) -ELYMUS

Elytrigia repens (L.) Des —Elymus repens

ENNEAPOGON Desv. ex P. Beau

*Enneapogon desvauxii P. Beauv. — moe chrome ore

piles (Reed 1964).

ERAGROSTIS P. Beauv. Lovegrass

“Eragrostis il Daveau - Baltimore, manganese ore

piles (Reed 1964).

a pd (L.) Nees - lacegrass. Common, pied-

infrequent in coastal plain.

ibn cilianensis (All) Vignolo ex Janch. - stinkgrass.

mmon weed in gardens and waste places through-

"Eragrostis curvum onrad) Nees - weeping ie ieee

ecomation often escaped and established (Peterson

3a: 76).

iis cylindriflora Hochst. - Baltimore, manganese

re piles d 1964).

gs ies frankii C. A. n ex Steud. — Frank's lovegrass.

ont, infrequen

Enos hirsuta ao Nees — big-top lovegrass Com-

mon on coastal plain (Frye).

is hypnoides n Britton, Sterns, & Poggenb. -

us megastachya a Link = E. cilianensis

*Eragrostis mexicana ven) Link

subsp. mexicana

Prince George’s Co.

subsp. virescens (J. Presl) S.D. iret & Sánchez Vega. -

Reported by Kartesz & Meacham (2002).

- Infrequent, Piedmont

Rare, known from

Eragrostis minor cw

lovegrass — Common and widespread weed.

ipic pilosa (L -) EE Ec val. pipes India lovegrass.

quent coastal plain F non

east of Chesapeake Bay.

m. & Schult. = E. m

& Meacham (2008).

Eragrostis refracta (Muhl.) a — Infrequent, oie:

from Wicomico and Worcester cos.

912

Eragrostis spectabilis (Pursh) Steud. — purple lovegrass.

ommon and widesprea

*Eragrostis suaveolens A x Claus — lovegrass.

Prince George’s Co. (Shetler €: Stone Orli 2

a poca Coss. €: Durieu — ieee Chrome

re piles (Reed 1964).

ics ida (Retz.) Nees ex Steud. — Baltimore,

chrome ore piles (Reed 1964).

(ERIANTHUS Michx.) = SACCHARUM

Erianthus alopecuroides (L L) Ell. = Saccharum alopecuroides

S. brevibarbe

Erian tus F Id— S. coarctatum

chia contortus Ell. = S. brevibarbe var. contortum

Frianthus giganteus (Walt.) Muhl. = S. giganteum

Erianthus ravennae (L.) Beauv= S. ravennae

ERIOCHLOA Kunth

*Eriochloa acuminata (J. Presl) Kunth - Baltimore, chrome

ore piles (Reed 1964).

var. minor (Vasey) R.B. Shaw - Reported by Kartesz &

Meac ).

oia So (L.) Desv. ex Ham. — Baltimore, chrome

ore piles (Reed 1964).

(EULALIA Kunth)

Eulalia vimineum (Trin.) A. Camus = Microstegium vimineum

STUCA L. Fescue.

Festuca arundinacea Schreb. = Schedonorus arundinaceus

Festuca elatior L. = Schedonorus arundinaceus

*Festuca filiformis Pourr. — hair fescue. Howard Co. Escaped

rom cultivation.

Festuca longifolia Thuill. =F. trachyphylla

Festuca obtusa Biehler = F. subverticellata

Festuca ovina L. - sheep fescue. Planted as ornamental.

Festuca paradoxa Desv. - cluster fescue. Rare, Garrett Co. and

usquehanna Valley (Brown & Brown 1984). Apparently

re with few records (Frye).

Festuca pratensis Huds. = Schedonorus pratensis

*Festuca rubra L. — red fescue.

subsp. commutata Gaudin — Chewings fescue. Planted

for DNUS ana Eataa ENGO B

subsp Nyman - Planted.

subsp. rubra. Plant

Festuca bo (Pers) E. B. Alexeev - nodding

mmon throughout (Frye

Festuca tenuifolia Sibth. = F. filiformis

*Festuca trachyphylla (Hack.) Krajina — hard fescue. Natu-

ralized. Established in BARC, Prince George's Co. (Terrell

t al. 2000)

GLYCERIA R. Br. Manna Grass.

Glyceria acutiflora Torr. — Rare on coastal plain and Blue

Ridge (Frye).

Glyceria canadensis ede ) zu

a cm Remi sd um eus un

piedmont and mountain zones (Brown & Brown 1984).

*Glyceria fluitans (L.) R. Br. - water manna grass Reported

from Baltimore

Glyceria grandis S. Watson — Reported by Kartesz & Mea-

cham (2008).

Glyceria laxa (Scribn.) Scribn. = G is var. laxa

Glyceria melicaria (Michx.) F.T. Hubb, - melic manna grass.

Infrequent, mountain and piedmo

Glyceria ide (Muhl.) Trin. — a manna grass. Com-

mon in coastal plain.

Glyceria ogi Hitchc. -.northern manna grass.

Frequent in coastal plain

Glyceria striata (Lam.) Hitchc. — fowl manna grass. Wide-

spread and common.

IN Oe em edu vid A

tarne E Poggenb.

- Frequent i in coastal plain.

Gymnopogon brevifolius Trin. - Rare, Worcester and

Wicomico cos.

HACKELOCHLOA Kuntze

*Hackelochloa granularis (L.) Kuntze — Baltimore, chrome

ore piles (Reed 1964).

R OER Br) = a

ANT OKAT

Hierochloe odorata |. Wahlenbos Anthoxanthum nitens

HOLCUS L. Velvet Grass.

*Holcus lanatus L. — velvet grass. Common weed.

HORDEUM L. Bar

Hordeum brachyantherum Nevski —

& Meacham (2008

*Hordeum jubatum L. — foxtail barley. Infrequent mostly

on Pid plain. Prince George's Co. BARC, (Terrell et

al. 2000).

NA marinum Huds. - Baltimore, chrome ore piles

(Reed 1964).

*Hordeum murinum L.

subsp. leporinum (Link) Arcang. — Reported by Kartesz &

Meacham (2008).

subsp. murinum - Baltimore, chrome ore piles (Reed

rey.

Reported by Kartesz

1964).

Hordeum pusillum Nutt. - little barley. Frequent in coastal

plain

“Hordeum vulgare L. - barley. Cultivated, escaping, not

stin

persisting,

HYSTRIX Moench

Hystrix patula Moench = Elymus hystrix

ISCHAEMUM L.

*Ischaemum indicum (Houtt.) Merr. — Baltimore, chrome

ore piles (Reed 1964).

*Ischaemum rugosum Salisb. — Baltimore, chrome ore piles

Reed 1964).

KOELERIA Pers.

Koeleria macrantha (Ledeb) 2e RE NER

uala & Bogler (2004), but no records found

"s phleoides (Vill) Pers. = Rostraria cristats

LEERSIA w

Rare. Data lacking

Leersia lenticularis Michx. — - catch-fly g grass. Rare, Worcester

Co.

Leersia oryzoides (L.) Sw. — rice cutgrass. Widespread.

Leersia virginica Willd. — white cutgrass. Widespread.

LEPTOCHLOA P Beauv.

*Leptochloa dubia (Kunth) Nees — Baltimore, chrome ore

piles (Reed 1964

Leptochloa fusca (L.) Kunth subsp. fascicularis (Lam.) N.

Snow — feather grass. a dandi a var. maritima

(E.P. Bicknell) G coastal plain, Calvert

Co., Cove Point ae B a

Leptochloa pauicea (Reiz) ad subsp bles id

N. SNOW Repot ted by K eacha

LEPTOLOMA Chase

Leptoloma cognata (Schult.) Chase = Digitaria cognata

LOLIUM i. Ryegras

*Lolium multiflorum Lam. - annual ane Italian ryegrass.

Often planted. Widespread weed.

*Lolium perenne L. - erem] ryegrass. Often planted.

Widespread weed

*Lolium temulentum L. - darnel. Uncommon weed. Balti-

more, chrome ore piles (Reed 1964).

MANISURIS L.

MELICA |. Melic Grass.

Melica mutica Walter — two-flowered melic. Rare, southern

D and piedmont (Frye).

Melica nitens (Scribn.) Nutt. ex Piper — Rare, ridg

(Frye).

| valley

MELINIS P. Beauv.

*Melinis repens (Willd) Zizka — Natal grass. (Rhynchelytrum

repens (Willd.) C.E.Hubb.). Treated as synonym by Zizka

(1988). Baltimore, chrome and manganese ore piles

(Reed 1964).

MIC ees

*Microstegium vimineum (Trin.) A. Camus — cover-all grass,

stilt grass, Nepalese browntop. Widespread noxious

weed

MILIUM L.

Milium effusum L. var. cisatlanticum Fernald - tall millet

grass. Locally abundant in Garrett Co. (Frye). Collections

from MD in MARY.

MISCANTHUS Andersson

Mica ys sinensis Andersson — miscanthus. Widely

plan | ly escaped.

E aid. Pan

ARIS

granularis

Mnesithea rugosa (Nutt.) de Koning & Sosef. = Coelorachis

rugosa

MOOROCHLOA Veldkamp.

MEAN ON A MA K

913

*Moorochloa eruciformis (Sm.) Veldkamp — Baltimore,

chrome ore piles (Reed 1964). Formerly placed under

Brachiaria

MEL ela

*Muhl i Aeyen ex Trin.) Parodi

-F rederick Co. (Reed 1 964).

Muhlenbergia capillaris (Lam.) Trin. - DC a

Muhlenbergia frondosa (Poir) Fernald - wirestem muhly.

Baltimore, Frederick, Howard, and Montgomery cos.

Muhlenbergia glabriflora Scribn. — Hyattsville, Prince

George's Co., Scribner s.n. at US (Shetler & Stone Orli

2002). G4

Muhlenbergia glomerata (Willd.) Trin. - Reported by Kartesz

& Me 200

Muhlenbergia mexicana (L.) Trin. — Infrequent in coastal

plain and piedmont. Plants with lemmatal awns 3-10 mm

long have been referred to as M. mexicana var. filiformis

(Torr) Scribn. (Peterson 2003b: 154).

rsen racemosa (Michx.) Britton, Sterns, and

Poggenb. - Reported from Garrett Co.

Muhlenbergia schreberi J. FGmel. — nimblewill. Common

Trin. - branched

Muhlenbergia sonnets (Mun. ex TIS

muhly. Freq

Muhlenbergia te Du Torr.ex A, Gray — woodland

ign Rare, mostly pean

A L No

, Sterns, & Poggenb.

- slender- flowered muhly. Common in western MD

rye).

(Fr

Muhlenbergia torreyana (Schult) Hitchc. — Rare, Caroline

Co. G3.

George's í Co, Suitland (Shetler. 8 sana Orli 2002),

OPLISMENUS P. Beauv.

*Oplismenus hirtellus (L.) P. Beauv. subsp. undulatifolius

Ard.) U. Scholz - Baltimore Co, Patapsco Valley State Park,

collected by Edward Uebel in 1996, the first record of this

taxon for North America (Peterson et al. 1999), Spreading

as noxious weed in 2008—2009

ORYZOPSIS Michx.

Oryzopsis asperifolia Michx. — Rare, mountains (Frye).

ANICUM L

Some subspecies were tested as 5 varieties in dioi and

Brown 1984. M. Lel d | certain collec

Panicum agrostoides Spreng. = P. rigidulum subsp. cipe

Panicum amarum Elliott — bitter beachgrass.

subsp. amarulum (Hitchc. & Chase) Freckmann & Lelong

- Infrequent in coastal plain.

subsp. amarum - Common in coastal plain.

Panicum anceps Michx. — beaked panicgrass.

subsp. anceps - frequent.

einen wh a (Hitcher 2

Fa MES | oO]

— Reported from MD.

: AN] Dada

Cnm eanillara

1 LI

witch grass Freq lent

throughout.

914

Panicum dichotomiflorum Micnx. — fall panicum. Com-

mon weed.

Panicum flexile (Gatt.) Scribn. — wiry witchgrass. Rare in

piedmont

Daricoons mnnttimmaor! D

gattingeri

Panem: hernitomon [s = gr A Infrequent i in

coastal plain (Frye)

Panicum longifolium Torr. = P. rigidulum subsp. pubescens

*Panicum m room-corn millet, proso millet.

Occasionally cultivated and escaped

*Panicum paludosum Roxb. - Baitimore, chrome ore piles

(Reed 1964).

Panicum philadelphicum Bernh. ex Trin. — Philadelphia

witchgrass

subsp. philadelphicum. Rare, Washington-Baltimore area.

RE | Rarnartarl

NST LO

D E alann

subsp. g

by Kartesz & Meacham (2008 ).

Panicum m Trin. — Baltimore, chrome ore piles

(Reed 1964).

subsp. combsii (Scribn. & C.R. Ball) Eeckaanin & dora

- No aata on distribution..

so]

elong

nd AA tt p

Pied-

and mountain zones. Prince George’s Co, BC

us et al. 2000

subsp. pubescens (Vasey) Freckmann & Lelong —

quent in piedmont and coastai plain.

subsp. rigidulum (P agrostoides) - Common in piedmont

and coastal n

| | [p cite PEN | Ius

Goll

Infre-

gatum

[d LI

m 2, Cehil

+ D.

Eee ore n Meca DEA.

Panicum verrucosum | Muhl. - warty panicgrass. Frequent

throughout.

Panicum virgatuniy i L= Nd aes ERR in coastal

plain

PARAPHOLIS C.F. Hubb.

*Parapholis incurva (L.) C.E. Hubb. — curved sickle grass

— Baltimore, chrome ore piles (Reed 1964). Calvert Co.,

Cove Point dunes (Steury 1997). Rarely adventive from

PASPALUM L.

Paspalum boscianum Flüggé - bull paspalum. Uncom-

*p. Humb. & Bonpl. ex Flüggé - Baltimore,

chrome ore piles (Reed 1964).

*Paspalum dilatatum Poir. — dallis grass. Baltimore Co.,

Prince Georges Co.

Paspalum dissectum (L) L. - Infrequent in coastal plain.

Paspalum distichum L. — Listed by Guala & Bogler (2004).

Data lacking

Paspalum floridanum Michx. — Florida paspalum. Frequent

near and in coastal plain

Paspalum fluitans (Ell) Kunth = P. repens

[| [| fal Das a In Li "FA ra

f Texas 3(2)

Paspalum laeve Michx. — smooth paspalum. Common and

widespread

*Paspalum orbiculatum Poir. — Baltimore, chrome ore piles

(Reed 1964

Paspalum pubiflorum Ruprex E. Fourn. - Reported from

Anne Arundel Co

Paspalum repens P J. Bergius — water paspalum. Uncommon.

*Paspalum scrobiculatum L. — Baltimore, chrome ore piles

(Reed 1964)

Paspalum setaceum Michx. - slender paspalum. No data for

these varieties in Brown & Brown (1984).

var. ciliatifolium (Michx.) Vasey

var. muhlenbergii (Nash) DJ. Banks

var. psammophilum it D.J. Banks - Occurs in DC.

var. setaceum - Freque

var. stramineum eue Al Banks

PENNISETUM Rich.

*Pennisetum glaucum (L.) R. Br. - pearl millet. Rarely

escaped from cultivation. Anne Arundel, Brooklyn Park

(Shetler & Stone Orli 2002). Baltimore, chrome ore piles

(Reed 1964)

ALARIS L.

Pda deidad i - ae canary grass. Common in

niarnraiant

here.

*Phalaris canariensis L - canary grass. Baltimore, chrome

ore piles (Reed 1964).

Phalaris caroliniana Walter — maygrass. Rare in coastal plain.

ds Co, probably extirpated (Shetler & Stone Orli

002).

"al minor Retz. - Reported by Kartesz & Meacham

ues pandora L. — Baltimore, chrome ore piles (Reed

1964).

PHLEUM L.

*Phleum pratense |. - timothy. Planted and naturalized.

Widesprea

*Phleum subulatum (Savi) Asch. € Graebn. - Baltimore,

chrome ore piles (Reed 1964).

PHRAGMITES Adans. Reed.

en australis (Cav) Trin. ex Steud. - common reed

and widespread. Two lineages are recognized

in Vsus us by Saltonstall et al. (2004), P australis subsp.

americanus Saltonstall, PM. Peterson, & Soreng, the native

Phragmites communis Trin. = P. australis

PHYLLOSTACHYS Siebold 2 LUGE

xp] Ri Chis Listed

_ by Guala & Bogier (2004).

Siebold & Zucc. - Cultivated

and sometimes spreading. Hyattsville, Prince George's

Co.

Li aes flexuosa Riviére & C. Riviére - Cultivated and

metimes spreading. Frederick Co.

PIPTATHERUM P Beauv

*Piptatherum miliaceum (L.) Coss. — Baltimore, pumice

piles (Reed 1964)

rat dis om dL rc X Ea +=

- (Oryzopsis racemosa (Sm) Ricker), Rare, mountain z zone

and Washington Co. Three MD collections at MAR

PIPTOCHAETIUM J. Pres!

Piptochaetium Se la Parodi — blackseed spear-

grass. (Sti oastal plain Montgomery Co.

(seen by Terrell along A : Q. canal). MARY has vouch-

ers from Anne Arundel, Caroline, Howard, and Prince

George’s cos.

POA L. Bluegrass.

Poa alsodes A. Gray - Uncommon, Garrett and Allegany

*Poa annua L. - annual bluegrass. Common and wide-

spread.

Poa autumnalis Muhl. ex Elliott - autumn bluegrass. Infre-

uent piedmont and coastal plain.

*Poa bulbosa L. - Infrequent. Montgomery Co, seen by RJ.

Soreng and P. Peterson

Poa chapmaniana serib: - Uncommon. Reported by

Kartesz & Meacham (2008

*Poa pc L. — Canada bluegrass. Common and

wides

Poa ae Nutt. — short-leaved bluegrass. Common

throughout (Fr a

*p oods bluegrass. Prince George’s (Shetler

& Orli 200 and Montgomery cos. (RJ. Soreng,

per

*Poa Susa hs fow! meadow a li in Howard

and Washington cos. and D

*Poa pratensis L

subsp. cuele (L) Lej. - narrow-leaved Kentucky

bluegrass. Prince Georges Co., BARC (det. by RJ. Soreng),

Montgomery Co. (RJ. Soreng collection)

subsp. lamas — Kentucky bluegrass. Very common

and widesprea

Poa saltuensis Fernald lada

(Hitchc) A. Haines. Mountains. No data.

= en

subsp. saltuensis — Rare, western MD. Garrett Co. collec-

tion at TAWES (Frye).

Poa PA A. Gray — sylvan bluegrass. ens in pied-

ont and mountains, rare on coastal p

os ade L.- - rough- -stemmed bluegrass. Common and

invasive, pieamont and northern coastal

POLYPOGON Desf.

*Polypogon monspeliensis (L.) Desf. — rabbitsfoot grass.

Frequent in coastal plain. St. Mary’s Co., Point Lookout

(Shetler & Stone Orli 2002).

PSEUDOSASA Makino ex Nakai

and sometimes spreading. DC area.

UCCINELLIA Parl. Alkali Grass.

*Puccinellia distans (L.) Parl. — rare to Milo in coastal

pla ain. Baltimore Co. (Shetler & Stone Orli 2

Puccinellia fasciculata (Torr) Bickn. — a 2 ain

is pallida (Torr) Clausen 2 Torreyochloa pallida

915

(RHYNCHELYTRUM Nees)

Rhynchelytrum repens (Willd.) C.E. Hubb. = Melinus repens

ROSTRARIA Trin.

*Rostraria cristata (L.) Tzvelev — annual June grass. [Koeleria

phleoides (Vill) Pers.]. Baltimore, chrome ore piles (Reed

1964)

SACCHARUM L. Plumegrass. (These species formerly in

Erianthus).

Saccharum alopecuroides (L.) Nutt. — silver plumegrass. Rare

in coastal plain. Calvert Co., Cove Point (Steury 2002).

Saccharum baldwinii Spreng. — Rare. Reported by Kartesz

& Meacham (2008).

MEE brevibarbe (Michx.) Pers. - shortbeard plume-

gra

var, — Somerset Co.

var. contortum rid R.D. Webster —

oe plain. -

Common in

corm nraccarl

VILI d d MIA NA

LAN MT Al 1

plumegrass. Frequent tal plain (Frye). Listed in Brown

& Brown (1984) without locations.

Saccharum giganteum (Walter) Pers. — sugarcane plume-

ommon in coastal plain.

“Saccharum ravennae (L.) L.— ravenna grass. Cultivated and

sometimes escaped to roadsides.

SACCIOLEPIS Nash

Sacciolepis striata (L.) Nash — American cupscale. Rare, coast-

al plain. Calvert Co. (Shetler & Stone Orli 2002). Worcester

Co.: Assateague Island, Aug 1984, Hill 15576 (MARY, US);

Shad Landing State Park, Sep 2005, Terrell 6311 (US).

SCHEDONORUS P. BE UN

tall facri io

Lai iar O Lus

E | [|

(Festuca arundinacea Schreb) Planted on roadsides and

elsewhere; an abundant

I-ino £

(Festuca pratensis Huds). Occasional meed dp Dres

ably planted. Believed to be much less frequent than

formerly.

SCHIZACHNE Hack.

Schizachne quibiis (Torr) Swallen —false melic. Rare,

mountain zone. Cited as occurring in MD (Cayouette &

BuU de m 103). Garrett Co., Jun 1940, Kearney,

Norton et dl. s.n. (MARY,

ee Nees

Schizachyrum littorale (Nash) E.PBicknell - Reported in

Brown & Brown (1984). MARY voucher reported by Kartesz

& Meacham

8).

sehizaenyrum scoparium (MIENK) NaS little bluestem.

in n coastal plain.

LEROCHLOA P Beauv.

ecerccnon! dura (L) P. Beauv. - ede Co., cornfield, Hill

in 1 li 2002). Frederick Co., location

unknown, sent in by uiii Extension Service, 8 May

1986 (MARY); Howard Co., Soreng 59290 (US).

—

916

SECALE L. Rye.

*Secale cereale L.—rye. Cultivated and occasionally escaped.

SETARIA P. Beauv. Foxtail, Bristlegrass.

*Setaria faberi RAW. Herrm. — nodding foxtail. Common

throughout

Setaria geniculata (Lam.) P. Beauv. = S. parviflora

Setaria glauca (L) P. Beauv. = Pennisetum glaucum. Yellow

oxtail was formerly widely known as Setaria glauca; see

Mar

S. pumila.

Setaria grisebachii F. Fourn. — Baltimore, chrome ore piles

(Reed 1964).

*Setaria italica (L) P. Beauv. — foxtail millet. Cultivated and

escaped to roadsides

Setaria magna Griseb. — giant foxtail, saltmarsh foxtail. In-

frequent in coastal plain. Blackwater Refuge, Dorchester

Co. (seen oy Ten re e

) K

AL! uéle rf

knotroot foxtail. Infre-

quenti in a coastal Bl Prince George's Co., BARC (Terrell

et al. 2

*Setaria Soci (Poir) Roem. & Schult. - yellow foxtail. Com-

on throu

*Setaria verticillata A P. Beauv. - hooked bristlegrass.

Infrequent throughou

*Setaria verticilliformis bumom - Reported by Kartesz &

Meacham (2008).

*Setaria viridis (L.) P Beauv. — green foxtail, green bristlegrass.

Common throughout.

SORGHASTRUM Nash. Indian Grass.

Sorghastrum elliottii (C. Mohr) Nash — long-bristled Indian

rass. Rare, coastal plain. Wicomico, Dorchester, and

Worcester cos. (Frye)

Sorghastrum nutans (L.) Nash —

throughout MD.

SORGHUM Adans. Sorghum.

*Sorghum bicolor (L) Moench - sorghum. Cultivated and

occasionally escaped.

*Sorghum halepense (L.) Pers. - Johnson grass. Common

and widespread weed.

SPARTINA Schreb. Cordgrass.

ids iria Loisel - smooth cordgrass. Common

al plain.

Eos o ode A. Eaton - tufted cordgrass. Puta-

tive cross of S. patens and S pectinata. Calvert Co (Shetler

& Stone Orli

a cynosuroides (L) Roth - big cordgrass. Common

al plain.

Seine pim (Aiton) Muhl. — salt-meadow cordgrass.

on coastal plain.

Spartina pectinata Link — prairie cordgrass. Infrequent in a

variety of habitats.

SPHENOPHOLIS Scribn. Wedgegrass.

Sphenopholis intermedia (Rydb.) Rydb. - slender

wedgegrass. Rather widespread.

Sphenopholis nitida (Biehler) Scribn. — Frequent throughout.

Sphenopholis obtusata (Michx, Scribn. — prairie wedgegrass.

Rather widespread.

Indian grass. Common

—

3/93

Sphenopholis xpallens (Biehler) Scribn. = $. obtusata x

S. pensylvanica as reported from MARY by Kartesz &

Meacham (2008).

Sphenopholis pensylvanica (L.) Hitchc. - swamp wedge-

d Uem pen NOH cup (L.) i: Beal ex Roem. &

chult.). | t in. Finotet

al. (2005) also placed this species in A T G4.

SPOROBOLUS Be BE pues

im D

ompositus

dne — Rare, eastern

Ma”

Sporobolus clandestinus (Biehler

Sporobolus compositus (Poir.) Merr. - Reported from

Queen Anne and Talbot cos. as S. asper in Brown &

Brown (1984)

*Sporobolus coromandelianus (Retz) Kunth — Baltimore:

chrome ore piles (Reed 1964).

Sporobolus heterolepis (A. Gray) A. Gray — Rare in piedmont.

Reported by Kartesz & aah (2008).

*Sporobolus indicus (L.) R. Br. - smut grass. Calvert Co, Cove

Point Park, Steury mim ient Co, two collections

as roadside weed, 7 mi ENE of Pocomoke City and As-

sateague Island, Terrell 6309 and 6314 (MARY). Probably

freq uent on n pan

c n " Es

MD (Peterson et al. 2003: 121). NE reports a | collection

in Baltimore County

Y Lian! Dale chrama

y 1

ore piles (Reed 1964). Prince George's Co, near Laurel, as

an abundant roadside weed, Terrell 6286, in 2005; det.

by P. M. Peterson). A southern and southwestern species.

Sporobolus vaginiiords (Torr. ex B Gray) Alph. Wood -

poverty d t we

Sporobolus virginicus (L) Kun - “seashore dropseed.

Baltimore, manganese ore piles (Reed 1964).

STIPA L. Needle grass.

Stipa avenacea L. = Piptochaetium avenaceum

THEMEDA Forssk.

*Themeda arguens (L.) Hack. - Baltimore, chrome ore piles

(Reed 1964).

TORREYOCHLOA G.L. Church

Torreyochloa pallida (Torr) G.L.Church - Frequent in coastal

plain. [Puccinellia pallida (Torr.) Clausen].

RAGUS Haller

*Tragus racemosus (L.) All. — Baltimore, chrome ore piles

(Reed 1964)

TRIDENS Roem. & Schult.

Tridens flavus (L) Hitchc. — purpletop. See key to separate

Wis vars. in Valdés nS (2003: 39).

Report Li A | by Kal tesz X

Meacham (2008).

var. flavus — Common and widespread.

(TRIODIA R. Br.) = Tridens

TRIPLASIS P. Beauv. Sand Grass

Triplasis purpurea (Walter) Chapm. - atie sandgrass.

Frequent in coastal piain.

TRIPSACUM L. Gama Grass.

Tripsacum dactyloides (L) L. - eastern gama grass. Frequent.

Trisetum pensylvanicum (L.) P. Beauv. = Sphenopholis pen-

sylvanica (L.) Hitchc

TRITICUM L. Wheat.

"Triticum aestivum L. — wheat. Rarely escaped from

cultivation.

UNIOLA L.

Uniola latifolia Michx. 2 Chasmanthium latifolium

Uniola laxa (L.) BSP = Chasmanthium laxum

Uniola paniculata L. — sea oats. Planted on beaches as ero-

sion preventive. Common in southeastern U.S.

UROCHLOA P. Beau

*Urochloa distachya (L.) T.Q. Nguyen — p by Kartesz

Meacham (2008).

*Urochloa mutica (Forssk.) T.Q. Nguyen - (Brachiaria mutica

(Forssk.) Stapf). Baltimore, chrome ore piles, reported

as Panicum purpurascens (Reed 1964; Shetler & Stone

Orli 2002).

l^ M

Baltimore,

(Reed 1964)

*Urochloa plantaginea (Link) R.D. Webster - plantain sign-

algrass. [Brachiaria plantaginea (Link) Hitchc.]. Wicomico

Co. (Terrell & Reveal 1996).

*Urochloa platyphylla (Munro ex C. Wright) R.D. Webster

— broadleaf signalgrass. (Brachiaria platyphylla Munro

ex C. Wright). Wicomico and Somerset cos. (Terrell &

Reveal 1996).

*Urochloa ramosa (L) T.Q. Nguyen - Baltimore, chrome ore

piles, reported as Panicum ramosum (Reed 1964; Shetler

& Stone Orli 2002).

917

*Urochloa reptans (L.) Stapf - Baltimore, chrome ore piles,

a as Panicum reptans (Reed 1964; Shetler & Stone

Orli 2

02).

3g | L. E r1 iD Imi ADISA I 4 T, l

[Brachiaria texana (Buckley) S.T dm Dorchester and

Wicomico cos. (Terrell & Reveal 1996).

*Urochloa villosa (Lam.) T.O. qu Reported by Kartesz

& Meacham (2008).

LPIA C.C.Gmel.

Vulpia elliotea (Raf.) Fernald = V. sciure

C. Gmel. - rat-tail fescue. Common.

Vulpia octoflora (Walter) Rydb, — six-weeks fescue. Com-

LV EE | = z fN h A L 5 | + AE A

e Freg quent

on coastal plain. Maritasme " ane Prince George's

cos. (Shetler & Stone Orli 2002); Calvert Co., Cove Point

(Steury 1997).

ZEA L.

*Zea mays L. - corn, maize. Rarely escapes from cultivation.

ZANIA L. Wild-rice.

Zizania aquatica L. - southern wild-rice. Frequent in coastal

plain.

Zizania palustris |. Reported by Guala & Bolger (2004) but

occurs only north and west of MD (Terrell 2003: 48; Ter-

rell et al. 1997).

ZIZANIOPSIS Doell & Aschers.

Zizaniopsis miliacea (Michx.) Doell. & Aschers. — giant

cutgrass. Rare in coastal plain. Calvert Co., Cove Point

(Steury 1

Willd.

*Zoysia japonica Steud. — zoysia. Lawn grass, sometimes

spreading.

ACKNOWLEDGMENTS

We are grateful to Christopher Frye for his review which added a number of additional records based on

his extensive knowledge of the Maryland flora. We thank Robert J. Soreng for information on Diarrhena

and Poa, and for suggestions regarding the format of the list. An anonymous reviewer contributed helpful

suggestions.

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Ls fil weeds nl s

I I

920 Journal of t tani i Texas 3(2)

BOOK NOTICE

STEPHEN SHENNAN (EDS). 2009. Pattern and Process in Cultural Evolution. (ISBN 978-0-520-25599-9, hbk.).

University of California Press, 2120 Berkeley Way, Berkeley, California 94704-1012, U.S.A. (Orders:

www.ucpress.edu or California-Princeton Fulfillment Services, 1445 Lower Ferry Road, Ewing, New

Jersey 08618, U.S.A., orders@cpfsinc.com, 1-800-777-4726). $60.00, 341 pp., b&w photographs, line

drawings, maps, tables, 7" x 10".

Contents:

Acknowledgments

Pattern and P in Cultural Evolution: An Introduction—Stephen Shennan

I TIT TTITO AT CUITCECTMAI

2. Placing Archaeology within a Unified Sci f Cultural Evolution—Alex Mesoudi & Michael J. O'Brien

3. H C icati Niche Construction—Robert Aunger

4. Modes of Transmission and Material Culture Patt in Craft Skills—Robert Hosfield

pipe me. Dats D : a] 15 1 T E AA a as | T^; Á AT 1 171 CWT +l r3

de o o + E

Siberia—Peter Jordan

6. Tangled Trees: Model

T. ae Poko of Material d nian d LM Iranian er Populations Jane Tehrani & Mark Collard

8. E R g y it Theory to Explain

= 1 | II a LI + A "T 1 i Beli+ Ri

A o Ried

Le e Record—Ethan E. Cochrane

9. Identifying Iron Production Lineages: A Case Study in Northwest Wales—Michael F. Charlton

Il. TESTING EVOLUTIONARY HYPOTHESES

10. Q itative Analysis of M luti y Patterning in Technological Evolution: Bicy le Design f 1800 to 2000—Mark

W. Lake & Jay Venti

11. Innovation Diffusion and Traveling Waves—James Steele

12. Explaining Global Patterns in Lower Paleolithic Technology: Simulations of Hominin Dispersal and Cultural Transmission

Using Stepping ucc pos Dm ionis & Steven Mithen

e E +h À A AS | T; } ] 1 [a n € Ai 1 ^4 Fai }

13. ee History A, a = FA ce

1T Pe SEC lanin D Ao Ao aes Cte Test lnoical Fvoluti f Kodiak. Alaska—

Ben Fitzhugh & A. Kate Trusler

Ill. SOCIAL EVOLUTION

15. Mind the Bonding G 5 t the Best fH Societ R.LM. vds

"T Tz lL 1 AT er” Y hs]. 137, "Es. A WA o6 xr 11313 +] i t Á alszeie af Indra Enranean

16 g ; Cultural y y European

MA . D +2 7 E " n ED Marx

&e IILIVLULI! CL L OLI LE A AVULLILE LULA

1 DAA ALT. ` oo T I: - L. C ‘10 Toos nw = d wo Te A 1 T SILT :eT5 1

Ladiclit H 5 5 o J

A Wiessner

t ithe F tion of Political Hierarchy: A Costly Signaling Model—Aimée M. Plourde

19. online and Warfare: A Test of the Turchin Model in Pueblo Societies—Timothy A. Kohler, Sarah Cole, & Stanca Ciupe

20. An Ecological Model for the Emergence of Institutionalized Social Hierarchies on California’s Northern Channel Islands—

Douglas J. Kennett, Bruce ME id Aa o aide

AT Tithi VH Tan Eu üj dp

21. Population, S t Cult i viliagco IUTL IX

List of Contributors

J. Bot, Res. Inst. Texas 3(2): 920. 2009

FLORISTIC STUDIES IN NORTH CENTRAL NEW MEXICO, U.S.A.

THE TUSAS MOUNTAINS AND THE JEMEZ MOUNTAINS

Brian Reif Jill Larson

_ ale University de MAN National Forest

15018 4 North Main St,

nein p 86001, U.S.A. Spearfi E a Dakota 57783, U.S.A.

rian. reif@nau.edu jilllarsonefs.fed.us

Brian F. Jacobs B.E. Nelson and Ronald L. Hartman

Bandelier National Monumen Rocky Mountain Herbarium

Los Alamos, New Mexico 87544, i SA, Department of Botany, Dept. 3165

brian_jacobs @nps.gov University oFWyomin

1000 E. University Ave

Laramie, Wyoming 82071, U.S.A.

bnelsonn or rhartman@uwyo.edu

ABSTRACT

This pay I ts the first of t tributions that the floristic diversity i tl tral New Mexico. The area encompasses

T: AA J¿1 RA f. 3 1; C RA y r.1 Fani AT r 1T lc Ta MT s 1T

[ar]

Also included is Bandelier National Monument, Valles Caldera National P ,B f Land Manag (Taos District), and other

federal, state, and west of the Rio Grande. The second paper will di the floristics of the portions of the two f

and A to ilie east of € Rio Grande, thus primarily bn MR de Cristo Range. The goal of this two-part series is to enumer-

y ever 1 in New Mexico. Here, we sna ae on 19,929 one collections of

1] T fel E +l 3s i +l

3.7 million acres is 35,857 n 11 ). A total of 1,384 unique taxa,

including 93 infraspecics and 10 E dh are e documented om 107 nes Of Mes 154 are exotics ota are Pp as noxious

in New Mexico

p22 pecies of conservation concern, 28 represent first reports or their r New Mexico, and finally

17 are endemics to New Mexico. Based on verified nr at ~~ herbaria, 64 Iditional uni t j Annotated

Checklist; thus the grand total is 1,445.

RESUMEN

ese +7 1 1 : m | J Pa | 7 1 1; s J £1 H d 1 B E 1. AT MAG

te área rodea

las iode Tus » las noua Tene incluyendo la ba Nd del Carson National x y in Fe iue Forest.

I Preserve, B i

Taos), y otras tierras federales, estatales y privadas al oeste sal Rio = El segundo artículo discutirá la florística de las porci ]

los dos a ae a este gel Rio Grande. P llera Sangre de Cristo. El objetivo de est ie de dos p

y istico int ivo ll ] b Nuevo México. Aqui, citamos 19,929 colecciones

las de pl ] (1 la pleta q I ás de 3.7 millones de acres es de 35,857 nuevas colecciones).

"i documentan un total de 1 3841 taxa únicos, que incluyen 93 infraespecíficos y 10 híbridos, de 107 familias. De estas, 154 son exóticas

^ 7 on

iu I I

fi: DE Į NT. "E pa Ja = 1. MT AAA nn ] sT C ED 1 + 1 1

ok incluy Li 64 t lici ] j i b Í ] 3 ll e 1 i total d 1,445.

INTRODUCTION

We report on botanical inventories in the western portions of the Carson National Forest (CNF) and the

Santa Fe National Forest (SFNF) by Jill Larson (2008) and Brian Reif (2006), respectively. Included are sur-

rounding public lands administered by the State of New Mexico as well as the Bureau of Land Management,

and some private lands (Fig. 1). These segments of the two Forests are defined as the portions occurring

west of the Rio Grande. Thus they include the Tusas Mountains to the north and the Jemez Mountains and

Sierra Nacimiento to the south. Results of two additional studies in the Jemez Mountains are included:

| Bat Rac Incet Tayac 3(2): ón. 961, 2009

922

I | fall Dos Li In LI dde S e F da $)

Journal 0 f Texas 3(2)

MÍ Santa Fe NF

| dui Carson NF

gu Bandelier Nat Mon

| —— vcnp

[| BLM

— Rio Grande

ais

' Velarde

Truchas

Los Alamos

Santa Fe

Fic. 1.1

4 <= al. PRA "1 J H e En, "1 IB | AS a TL LH fa | LR L | A yj: re | a: 4 2L 4. AUD al

Rio Grande. |

1 E laj: aol f ye ls | a oe |

f J

Reif et al., FI f the Ti ij tains, New Mexico 923

Bandelier National Monument (BNM; Jacobs 1989) and the Valles Caldera National Preserve (VCNP). Data

from the latter, although reported separately (Hartman & Nelson 2005), were incorporated into the SFNF

thesis (Reif 2006)

This is the first of two papers on federal and adjoining lands in north central New Mexico. The second

will focus on the Sangre de Cristo Mountains to the east of the Rio Grande (largely CNF and SFNF). The

Jicarilla Ranger District of the Carson National Forest in northwestern Rio Arriba County is excluded, al-

though inventoried by Larson (2008).

The geographic area covered in this paper is over 2.4 million acres (the area as a whole inventoried in

the two thesis projects and BNM was over 3.7 million acres). Included are all of Los Alamos, eastern Rio

Arriba excluding the extreme southeast corner, northeastern Sandoval, northwest Santa Fe, and western

Taos counties. Data on the Tusas Mountains are limited to the portion under the jurisdiction of the Carson

National Forest (see Heil & O’Kane 2007 for partial coverage of the western slope). The area ranges in eleva-

tion from about 5,500 feet to just over 10,900 feet

These botanical inventories are part of the larger effort by the Rocky Mountain Herbarium (RM) to

produce a critical flora of the Rocky Mountains and to map, based on vouchered specimens, the distributions

of its taxa in relatively fine detail (Hartman 1992; Hartman & Nelson 2008; Hartman et al. 2009). To this

end, over 64 major floristic inventories have been conducted since 1978 from Oregon, Washington, Idaho,

Montana, and South Dakota south to north central New Mexico. Over 575,000 new numbered collections

have been obtained. They form the core of the 700,000 record Rocky Mountain Herbarium plant specimen

database (Hartman et al. 2009).

Topography.—A majority of the area is within the Southern Rocky Mountain Region (Fenneman

1931). To the north, in southern Colorado, this region is divided by the headwaters of the Rio Grande, thus

forming the broad, treeless San Luis Valley. Here the San Juan Mountains are to the west and the Sangre

de Cristo Mountains are to the east. This division extends into New Mexico (the CNF), where the San Juan

Mountains that extend southeast from Colorado are known locally as the Tusas Mountains. The San Juan

or Tusas Mountains are separated to the south from the Jemez Mountains and the Sierra Nacimiento by the

Rio Chama (Fenneman 1931). The surficial geology is largely volcanic. On the east side of the Rio Grande,

the Sangre de Cristo Mountains form the southernmost range of the Rockies. At the southern terminus,

near Santa Fe, they dissolve into a landscape of mesas and plains.

Unless stated to the contrary, the following discussion is limited to the Tusas Mountains, Jemez Moun-

tains, Sierra Nacimiento, and vicinity. The region may be divided broadly into three geomorphic units: the

Rio Grande Rift (including Taos Plateau), the Tusas Mountains, and the Jemez Mountains (including the

Sierra Nacimiento).

The Taos Plateau is a broad basin and the southern extension of the San Luis Valley of Colorado in

the northern portion of the valley of the Rio Grande. lt is peppered with volcanic cones some of which rise

3,000 feet or more above the surrounding desert grass- and shrublands. The Rio Grande has cut a deep,

narrow canyon through the basin and in places gorge exceeds 800 feet in depth. To the south, the Rio

Grande Rift is the portion constricted by | and formed by the Embudo Fault (Muehlberger 1978;

Muehlberger & Muehlberger 1982).

To the west, the Tusas Mountain unit consists of low mountains and expansive valleys, descending

south to the Rio Chama. Thus, a portion of the southern Chama Basin is included. The highest point in the

Tusas Mountains under jurisdiction of the CNF is Canjilon Mountain at 10,903 feet.

To tbe south of the Rio Chama lies the Jemez Mountain unit, the location of Bandelier National Monu-

ment, Valles Caldera National Preserve (Anschueta & Merlan 2007; Martin 2003; Muldavin & Tonne 2003),

and the town of Los Alamos. Here, the highest point in the area is Redondo Peak at 11,255 feet. This unit

includes the Sierra Nacimiento which merges on its east flank with Meu ME the Tusas and the Jemez

Mountains, the Sierra Nacimiento was part of the Laramide orogeny as was the Sangre de Cristo Mountains

to the east of the Rio Grande.

[| £ sl Dak Li in LJ ri. ds mi

924 J f Texas 3(2)

Based on Fenneman (1931), the Southern Rocky Mountain Physi hic Province includes the Tusas

and the Jemez mountains, as well as the Sangre de Cristos to the cast PT the intervening Rio Grande Rift

including the Taos Plateau. The lowlands to the west of the Tusas and Jemez mountains are in the Colorado

Plateau Province, while those to the south of Santa Fe are in the Basin and Range Province.

Climate.—The climate of north central New Mexico is arid in the lowlands, increasing in moisture

as one ascends in elevation. Precipitation is significantly affected by the North American monsoon. The

monsoon results in a seasonal pattern of precipitation with a primary maximum during July, August, and

Seen ace onis loua cover and precipitation accompany this season. Pa corresponds with

letal 1

y leading to frequent afternoon tl 1 (Shepp 1999).

The mountains exert ona E effects on precipitation in direct zelatiomshin to elevation while

temperature is inversely related. New Mexico receives between 30 and 40 percent of its annual precipitation

during the summer months.

Winter tends to be drier, with a secondary maximum Duor precipitation peraman November through

March (Sheppard etal. 1999). 5 Winter pre ipitat ion is | ) E lly i import tant as E ] recharges

surface and ground water (Redmond 2002).

In the Rio Grande Rift, annual precipitation is about 10 inches at Española (elevation 5,640 feet) and

14 inches at Santa Fe (6,720); in the Tusas Mountains, 10 inches at Abiquiu Dam (6,300) and 14 inches at

Tres Piedras (8,140); while in the Jemez Mountains, 17 Hu at Jemez hend (5, Pd he inches at Los

Alamos (7,300), and 23 inches at Wolf Canyon (8,220). R hadows may explain d Abiquiu

Dam, located in intermontane basins, receive the least annual pon In a ACRES differential

pam: of east anes west exposures and localized cold air drainag e also produce tremendous differences in

1 1979). There is no long-term climate data for higher elevations in the

mountains. The al ipitation dat | dona 30 year ecco] (National Climatic Data Center 2009).

Drought during a porno of this study had an impact on collecting. Dry conditions were particularly

severe in 2002 and 2003 and again in 2006 until the monsoon developed in late July and early August.

Current drought conditions and climate change information for New Mexico can be obtained from the

Governor’s Drought Task Force (2008).

Geology and geomorphology.—The current landscape of the Southern Rocky Mountain Region is

mainly a product of the Laramide Orogeny (late Cretaceous into the Tertiary (Eocene epoch), 70 to 55 million

years ago (mya)) that produced the Sangre de Cristos and the Sierra Nacimiento. Subsequently, volcanism,

erosion, infill, and subsidence of the mountains and basins occurred ae et al. ee un older rock

types of ig igins | | lifted and expose d as a result of the istory. Thus, the

summits of the Pane de Cristo modi T tai ist of Precambrian jode to I. 78 billion vais in age.

The Tusas Mountains underwent extensive volcanic activity in the Tertiary (Bingler 1968). The northern

portion received lava and ash flows from volcanic fields in the San Juan Mountains to the northwest. This

portion of the range is now covered in ash-flow tuffs and volcaniclastic rocks of the Conejos Formation

(New Mexico Bureau of Geology and Mineral Resources 2003).

The southern portion of the Tusas Mountains includes the Brazos uplift. This area is largely comprised

of Precambrian granite, quartzite, schist, and gneiss (Bingler 1968). The ling valleys and slopes are

covered in volcanic detritus eroded from the San Juan and northern Sangre de Cristo Mountains (Muehl-

berger & Muehlberger 1982).

During the Pleistocene, the Tusas Mountains were glaciated above 8,000 feet. Glacial outwash over

Cretaceous strata has created large landslide deposits on its west slopes (Chronic 1987).

Separating the Tusas from the Jemez Mountains is the southern Chama Basin. It is filled with sand-

stones and shale of Jurassic and Triassic age (Smith et al. 1961). Present are fossils of the Upper Triassic

that include conifers and cycads (Ash 1974). Of particular importance for the extant flora are outcrops of

Todilto gypsum which harbor edaphic endemics such as Abronia bigelovii and Astragalus micromerius (New

Mexico Rare Plant Technical Councii 2008).

Reif et al., Fl f the Ti iJ tains, New Mexico 925

Volcanic activity intensified (10 mya) in the western Española Basin al the Rio Grande Rift. Repeated

volcanic episodes gave rise to the Jemez Mountains, emerging directly out e the Rio Grande Basin. The

earliest eruptions (9-11 mya) were basalt flows (e.g., caprock on Borrego Mesa) over arkosic sediments of

the Santa Fe Group (Bailey & Smith 1978). Volcanic activity a culminated in a series of large pyroclastic

eruptions at1.6 and again 1.2 mya that left a crater 16 miles in | 1 deposited many hundreds of feet

of volcanic tuff (Goff et al. 1996). The center of the Jemez Mountains is ne by this crater that consists

of broad valleys interrupted by resurgent domes. It is the site of the Valles Caldera National Preserve. The

most recent eruption was 40,000—50,000 years ago os the El Cajete crater wl outfall blanketed

Bandelier National Monument. N more detail "e on of the volcanic history

and the resultant geologic structure of the Jemez (Goff et al. 1996; Kues et al. 2006; Smith et al. 1970).

The Rio Grande Rift was formed in the Cenozoic (70—0 mya). To the north it forms a broad, relatively

flat surface of basaltic lava flows] as the Taos Plateau volcanic field. These flows issued from hundreds

1] prominent and include San

of vents during the Pliocene (Chronic 1987). The larger shield

Antonio and Ute mountains. Rocks from some vents are dated from 2.2 to 1.8 mya (Lipman 1978).

The Rio Grande Rift popa isa mare trough formed in two phases. Eden separate "subsided blocks

or grabens with high topog f and bound by widely spaced, high l faults” formed (Kelly

1990). T d ts of arkosic sediments, or quartzite derived. ane. were deposited primarily from

the Sangre de Cristo Mountains into the grabens. This led to the characteristic Santa Fe Group fill found

within the Española Basin (Muehlberger & Muehlberger 1982).

The Rio Grande became a free-flowing river during the Pliocene, having overcome containment by a

succession of closed basins within the Rift (Chronic 1987). On the Taos Plateau, it is now confined to the

Rio Grande gorge.

METHODS

Field work on the Carson National Forest, the Santa Fe National Forest, and Valles Caldera National Preserve

was conducted during the summers of 2001 through 2006 whereas the P QE National Monument was

inventoried by Brian Jacobs from 1986 through 1988. These stu dies ined represent the most extensive

and exhaustive floristic survey ever conducted in New Mexico.

Botanists have been roaming the country sides throughout the world for centuries, documenting the

riches of floristic diversity. In keeping with the tradition of field botanists, collecting sites were selected and

searched based on tl her’s judgment. Thus, the “meander search" strategy was employed (Goff et al.

1982; Ristau 1998; Hartman 1992; and Hartman & Nelson 2008). As sites were not selected at random, the

result was the exploration of a much greater diversity of plant communities, soil types, geologic substrates,

and topography leading to the documentation of a substantially greater diversity of taxa.

A total of 697 waypoints (each a geographic coordinate determined using a GPS unit) are represented

in this paper (for the two thesis projects as a whole, the total was 1,542). Each waypoint represents a general

location for collecting plant specimens, usually to within one-half mile (either as a radius from a point or a

trail segment defined by two successive points). Details concerning collecting as followed in ali RM studies

are found in Hartman (1992) and Hartman and Nelson (2008).

Nomenclature in the Annotated Checklist follows Allred (2008) with some exceptions as treated in

Hartman and Nelson (2001) or Flora of North America Editorial Committee, eds. (1993+).

This paper is based on 19,929 collections from the Tusas Mountains, Jemez Mountains, and vicinity

(total number of collections obtained for the two thesis projects and the BNM inventory was 35,857 collec-

tions). A complete set of vouchers from CNF and SFNF are housed at RM. Bandelier specimens are housed

at the monument herbarium with a partial set of duplicates at DUKE and miscellaneous vouchers at RM

and UNM. All authors of this paper have made major contributions to the collection, identification, and

verification of specimens as well as the writing of this paper.

The databases of the University of New Mexico Herbarium (UNM) and the Rocky Mountain Herbarium

926 I tani titute of Texas 3(2)

(RM, USFS) were searched for species not encountered during our field studies. Their identifications were

verified, and the institution and voucher is so indicated in the Annotated Checklist.

RESULTS AND DISCUSSION

The followi ti ill hasize tl Its of research on the Tusas and Jemez mountains and vicinity

with some on on various topics. Furt! f the will await the second paper on the

Carson and Santa Fe National Forests to the east of the Rio Grande, thus data from the entire area.

Vegetation Types

New Mexico's dias has been mido and described in a number of studies at various spatial scales.

The most relevant to thi | by Brown (19942, b), Daubenmire (1943), Dick-Peddie

(1995), and Whittaker (1978). (econ types reported here are based on the application of the above-

mentioned classifications as confirmed or modified by our observations. We report 14 vegetation types

within five broad physiognomic and zonal categories. These may foster an understanding of the ecological

amplitude, common associates, and environmental requirements of the taxa documented by these floristic

inventories.

Forests and Woodlands

Spruce-fir forest.—These forests occur in subalpine habitats above 9,500 feet or on cooler and moister

slopes at somewhat lower elevations. Picea engelmannii and Abies arizonica are codominants, although Picea

pungens may occur as a seral species on moist sites (Moir 1993; Pase & Brown 1994a; U. S. Forest Service

1997). Scattered stands of Populus tremuloides may be present in disturbed areas. Conifers of lower eleva-

tions, such as P Abies concolor, and Pinus flexilis, may be present in low numbers. In the

Tusas Mountains, STE fir forevis were observed primarily around the Canjilon Lakes and Cruces Basin

while in the Jemez Mountains they were common at higher elevations. The shrub and herb layer can be

quite sparse due to the closed canopy.

Associated shrubs and pe include Arctostaphylos uva-ursi, Juniperus communis, Paxistima

myrsinites, Ribes wolfii, Sali | Sambucus racemosa, and Vaccinium myrtillus. The herb layer is

quite sparse yet may includes the forbs oo rubra, Castilleja sulphurea, Cymopterus lemmonnii, Ligus-

ticum porteri, Pedicularis racemosa, Penstemon whippleanus, Solidago simplex, and Viola canadensis and

such graminoids as Bromus ciliatus, B. richardsonii, Luzula parviflora, Poa leptocoma, Trisetum spicatum

and T. montanum.

Mixed conifer forest.—This forest type covers large areas in the Tusas and Jemez mountains (8,000—

10,000 feet). Small islands of mixed conifer forest are also found on the summits of the highest volcanic

cinder cones on the Taos Plateau, including San Antonio Mountain, Ute Mountain, and Cerro de la Olla.

Pseudotsuga menziesii and Abies concolor are the most widespread and important conifers. They occur with

Pinus strobiformis in cooler sites while Pinus flexilis is found in dry, exposed areas, and Abies arizonica in the

ecotones. Picea pungens is restricted to locations subject to cold-air inversions including lower slopes and

drainages while Pinus ponderosa is found in xeric sites at lower elevations (DeVelice et al. 1986; Moir 1993).

The mixed conifer forests are heterogeneous thus reflecting the landscape.

Common understory shrubs and subshrubs include Acer glabrum, Arctostaphylos uva-ursi, Berberis

repens, Holodiscus dumosus, Jamesia americana, Juniperus communis, J. scopulorum, Paxistima myrsin-

ites, Physocarpus monogynus, Rosa arkansana, R. woodsii, Symphoricarpos rotundifolius, and Vaccinium

myrtillus. Forbs may be oe by Artemisia franserioides, ae miniata, Cymopterus lemmonii,

Fragaria vesca, Lat! multiflorum, Potenti la gracil is, Thalictrum fendleri, and

Thermopsis homes while m graminoides eve aches n a a O

Carex siccata, C. geophila, C. occidentalis, El nd A

asperifolia.

Fire has created a natural mosaic within this vegetation type, with burns ranging from patchy, low

Reif et al., Fl f theT IJ tains, New Mexico 927

intensity to stand replacing fires (U. S. Forest Service 1997; Moir 1993). Insect outbreaks and dwarf mistletoe

also add to the patchwork.

Ponderosa pine Tort ms armer aus drier conditions from 6,700—8,500 feet support this forest type

that often f n hout the Tusas and Jemez mountains. At its upper limit, Pinus

ponderosa and eed conifer forests da while at its lower limit it merges into piñon-juniper wood-

land. However, ponderosa pine typically is the sole aora conifer. Where trees are large and scattered

the forests may be open and park-like, with a predominately grass y understor y formed under pre-settlement

fire conditions (Allen 2001; Moir 1993; Pase & Brown 19945). D ponderosa pine forest

include frequent, low intensity understory fires (DeVelice et al. 1986; Moir 1993: U.S. Forest Service 1997).

Associated deciduous shrubs in the understory include Berberis repens, Ceanothus fendleri, Cercocarpus

montanus, Failugia paradoxa, Juniperus communis, Ribes cereum, R. inerme, R. leptanthus, Rosa acicularis, R.

woodsii, and Symphoricarpos rotundifolius. Relictual individuals of Juniperus deppeana are here documented

for the Jemez Mountains. These are of Ma an AR with a center of ela in Mexico (Peet 1988).

Representative forbs include Achill mauriopsis dissecta, Á vifolia, Cymopterus lem-

monii, Enden diti dis E. pacers, Heierotheca villosa, PEN NETS Lithospermum ead es

4 .o.

nin , and Vi rin ^11 1m

wu Carex inco ssp. keliorhila, Elymus elymoides var. brevifolius, oe arizonica, TEN macrantha,

Muhlenbergia montana, and Poa fendleriana.

Piñon-juniper woodland.—Pinus edulis and Juniperus monosperma are dominant in the foothills and

form a discontinuous transitional belt (6,000 to 8,500 feet). Piñon-juniper woodlands represents the low-

est elevation, forest type. Pifion forms closed woodlands at the upper elevational range, whereas juniper

occurs in savanna-like communities at the lower elevational range and interface with grasslands. Juniperus

scopulorum ly occurs with piñon in i d has recently encroached into Pinus ponderosa

understories in absence of fire disturbance (U.S. Forest Cie 1997). Understory conditions are dynamic

and vary with canopy cover, soil conditions, and land use and fire history (West 1999).

Shrub cover is variable dno MG events ini and Q. undulata, out also Artemisia tridentata,

Cercocarpus montanus, Guti Ribes cereum, and Rhus trilobata. Forbs include

Castilleja integra, Chaetopappa etipoldes amarse fendl eri, Cryptantha cinerea, Eriogonum jamesii, Gaillardia

pinnatifida, Hymenoxys richardsonii, Lappula occidentalis, Oenothera suffrutescens, Sphaeralcea coccinea, and

Tetraneuris argentea with several succulents including Cylindropuntia imbricata, Opuntia phaeacantha, O.

polyacantha, and Yucca spp. Common graminoids are Achnatherum Alla: nud i am Bouteloua

curtipendula, B. gracilis, Carex geophila, Elymus elymoides, Hesperostipa comata, j

ee PONE RM mierdas and Poa fendi: eriana.

linclude fuel harvest and g j g (] Toi 1993). Degraded

conditions due t to historic grazing are often characterized by high densities: of Juniperus monosperma and

intercanopy spaces d p aie gracilis and Gutierrezia sarothrae with exposed soils and active

erosion. Bark beetle outbreaks o iated with drought stress can result in high levels of pifion

mortality with the most recent rento —— in 2002 (Breshears et al. 2005).

Shrublands

Montane Shrubland.—These shrublands are found throughout the Tusas and Jemez mountains. They are

distributed in a patchy manner throughout ponderosa pine forest or pifion-juniper woodland. Areas of

montane shrubland are often too limited in extent for mapping because they are produced by site-specific

factors: past disturbance, local substrate, and patterns of moisture. Comparably drier and more rocky sites

are typical of montane shrubland, although some associates such as Prunus virginiana and Ptelea trifoliata

are found in areas of increased moisture such as small catchments (Dick-Peddie 1993).

Included in this classification of montane shrubland are thick stands of Quercus gambelii and Robinia

neomexicana. These species occur on soils that are poorly developed and xeric (Brown 19943) and have also

I [| fal Dos H Im Lt g*. ai

928 f Texas 3(2)

been regarded as a seral association produced by past disturbance (Dick-Peddie 1993). Also included here

is the “climax” shrubland of Dick-Peddie (1993) or Cercocarpus montanus in association with Amelanchier

alnifolia, Philadelphus microphyllus, Quercus ungulata, and Rhus Pe oaae Other shrubs include Ceanothus

fendleri, Holodiscus dumosus, Jamesia americana, Juniperus communis, P gynus, Ribes cernum, R

inerme, Rubus parviflorus, Sambucus racemosa, and iaa POE NUS

Desert Shrubland.—There are three subtypes that qualify as desert shrubland. One is found within a

large area of the Taos Plateau, especially west of the Rio Grande. This subtype is characterized by the absence

of Artemisia tridentata var. tridentata and the presence of a number of other shrubs and subshrubs. Included

are Artemisia nova, Atriplex canescens, Chrysothamnus greenei, Ericameria nauseosa, Gutierrezia sarothrae, and

Tetradymia canescens.

The second subtype, the desert scrub, is dominated by Artemisia tridentata var. tridentata. It most often

occurs in openings or glades within pifion-juniper woodland. Atriplex canescens, Cylindropuntia imbricata,

and Ericameria nauseosa are common woody associates.

The third Ee is Hx sorub d in SÉ areas on Po anal of ue Rio uama ane Rio

Grande. Some 1 A Z

and Rhus trilobata.

The herbaceous cover is often sparse in desert shrublands but may include the forbs Castilleja integra,

Pa aa Croton texensis, Eriogonum annuum, E. pl OACI Euler, ane

the grasses A hymenoides, Aristida purpurea, Boute lo hl

torreyi, and Sporobolus airoides.

Grasslands

Montane meadow and grassland.—This vegetation type occurs Pom REM 8,500 feet to i ee sum-

mits throughout the Tusas and Jemez mountains. Upper-elevation si

whereas lower sites are often surround by coniferous forests. Blegharonturon dan Danthonia nam

Festuca idahoensis, F. HAAS S ih mann. dini au montana, and Poa fendleriana dominate at

higher elevations, while f may outnumber grasses in lower elevation meadows. These

grasslands are frequently associated vit areas where poor drainage or other edaphic conditions inhibit

tree establishment (Brown 1994b). While species composition varies with elevation and moisture levels,

montane meadows are characterized by a diverse array of grasses and forbs. Some frequent representatives

of the latter group include Achillea millefolium, Campania FOtuniaond, ae LTODIENG, hi ale fendleri,

Erigeron flagellaris, Erysimum capitatum, F } , » 11) ) H. hoopesii,

Ipomopsis aggregata, I iensis, Lupinus argenteus, Orthocarpus luteus, Potentilla hippiana, P gracilis, and

Thermopsis rhombifolia.

Generally, this vegetation type has experienced a high level or disturbance by humans. Loss of natural

fire cycles often leads to encroachment by oe plants. Overg g and changes in climate also may have

an effect (Merola-Zwartjes 2004; Raish 2004; Dick-Peddie 1993).

Plains-desert grassland.—This vegetation type is ecotonal to pifion-juniper woodlands or juniper

savannas. It may be interspersed with, or sometimes indistinguishable from, Great Basin desert scrub

(Dick-Peddie 1993). Desert grassland occurs primarily along the Rio Grande Rift, on the edge of, or within,

surrounding pifion-juniper woodlands, and to the west of the Tusas same Jemez mountains where it is transi-

tional to desert. It has a significant shrub and forb cover. Cl Achnatherum hymenoides,

Aristida pupa Bouteloua gracilis, B. EU ERIS a E comata, and Hilari jamesii,

hile f ntered forbs include A ylla, A. rosea, Castilleja integra, Cryptantha spp.,

Clanáuana ada Grindelia squarrosa, Oenothera coroncpifolia, O. suffrutescens, Opuntia polyacantha, O.

phaeacantha, Ratibida tagetes, Sphaeralcea coccinea, Teucrium laciniatum, and Zinnia grandiflora.

Wetlands

Montane Riparian.— Margins of perennial and intermittent st pport variable species assemblages.

Montane riparian vegetation is found in moist areas within spruce-fir and mixed conifer forests. The above

Reif et al., Fl f the T IJ tains, New Mexico 929

are especially well developed in the Jemez Mountains but may also be encountered in the northern Tusas

Mountains. As with montane meadows, the species composition follows an elevation gradient (Dick-Peddie

1993). Obligate and facultative riparian species of trees and shrubs can be arranged along a descending

gradient: Picea pungens, Salix bebbiana, Alnus incana, Acer glabrum, sali sericea, eons angustifolia, and

Acer negundo. A facultative i riparian trees and shrul P. , Prunus virginiana,

bini ich diversity of forbs are Aconitum Cou bum. Caltha

leatasepalis Cardaiine cordifolia, Equisetum arvense, Geum macrophyllum, Heracleum maximum, Hypericum

scouleri, Mertensia fransiscana, Mimulus guttatus, Oxypolis fendleri, Pedicularis groenlandica, Saxifraga odonto-

loma, Veronica americana, and species of Epilobium, Potamogeton, Ranunculus and Salix. Graminoids include

=—

Alopecurus aequalis, Deschampsia cespitosa, Elymus canadensis, Glyceria grandis, G. striata, Juncus arcticus, Tor-

reyochloa pallida, and species of Agrostis, Carex, and Eleocharis.

Floodplain-arroyo riparian.—This type of vegetation occurs at lower elevations on floodplains of the

major rivers, the Chama, Jemez, and Rio Grande, or in arroyos which drain bajadas and mesas. Many spe-

cies that thrive here are well adapted to disturbance and dry conditions with periodic flooding. The most

common dominate is Populus deltoides, with the understory shrubs Baccharis salicina, Ericameria nauseosa,

Fallugia paradoxa, Forestiera pubescens, Rhus trilobata, and Salix exigua. The exotics, Elaeagnus angustifolia

and Tamarix chinensis have proliferated and may persist in a subclimax state (Minckley & Brown 1994;

Dick-Peddie 1993). Arroyo riparian is typically adjacent to desert shrubland and may grade into montane

riparian.

Marsh-lacustrine.—The marsh-lacustrine riparian habitat is found around ponds, springs, and stock

tanks in otherwise arid habitats such as pifion-juniper woodland or desert grassland where the water table

remains sufficiently high, or in various montane vegetation types. Along the shoreline Limosella aquatica,

Potentilla anserina, P. norvegica, Ranunculus cymbalaria, Rorippa spp., and R may be encountered.

Emergents ind aL bli: aliada emersum, n, Typha Jijona and species of oa Eleocharis,

Juncus, and S pp., Lemna

minor, Elodea — sis, Myriophyllum sibiricum, Ranunculus aquatilis, and Zannichellia pdlisine

Disturbed

Aspen seral forest.—Populus tremuloides, an important seral species and post-fire increaser, is widely dis-

tributed in the Rocky Mountains (Peet 1988). Pure stands of this shade-intolerant species arise through

root sprouting following disturbances (Pase & Brown 1994b). Scattered individuals are also found in late-

succession or near cli tages in mixed conifer forest and lower subalpine spruce-fir forests (Moir 1993).

However, like many successional communities, aspen forms a distinct assemblage that may persist for

long periods of time. Aspen are found in various coniferous forest types throughout the Tusas and Jemas

mountains.

Forbs include Campanula rotundifolia, Castilleja miniata, Chamerion angustifolia, Geranium richardsonii,

Pseudostellaria jamesiana, and Thalictrum fendleri while frequently encountered grasses included Poa fendle-

riana, P. pratensis, Bromus carinatus, B. richardsonii, and Festuca arizonica.

Burn Areas.—Natural and on oe DE "es pave been MUS in most of the vegetation types, as

mentioned above. In this regard the J M ively researched. Over 5,200 fires have

been documented between 1906 and 1996, 75 percent of which were miaied by lightning (Allen 2001).

Allen also described near continuous fire activity during the last 9,000 years based in large part on fire

scars on trees and the charcoal record at Alamo Bog (VCNP). Recent extensive fires in the Jemez Mountains

include the Dome (16,000 acres, 1996), the Cerro Grande (45,000 acres, 2000), and the Borrego (12,700

acres, 2002).

Roadside-agricultural.—Here, native vegetation may be largely replaced by exotic or agricultural spe-

cies. Noxious weeds collected along roads include Aegilops cylindrica, Cirsium vulgare, Convolvulus arvensis,

Lepidium latifolium, Linaria dalmatica, and Ulmus pumila. Weedy or agricultural plants include Avena sativa,

Helianthus annuus, Salsola tragus, Sisymbrium altissimum, Tragopogon dubius, and species of Ambrosia, Bromus,

930 Journal of t tani itute of Texas 3(2)

Chamaesyce, Chenopodium, Elymus, Lappula, Lepidium, Medicago, Melilotus, Plantago, Polygonum, and Trifolium.

Roadsides often act as corridors for exotics and thus warrant monitoring.

NOXIOUS WEEDS

Invasive exotic plants are a eres cause for ie loss of madisen oes 1998; MENÉ et al. 2000).

In natural areas, i defined which d plant composi-

tion, structure, and Ecos sem functions (Cronk & Fuller 1995). iayacive plant species that are particularly

damaging or prolific are regulated as noxious weeds (Natural Resources Conservation Service 2008). A

specific goal of our floristic inventories was to document noxious weeds for the purpose of assisting in

monitoring and control efforts. Of the 32 taxa listed in New Mexico, 14 were encountered at 166 sites.

They are A il Aegilops cylindrica, Carduus nutans, Cirsium arvense, C. vulgare, Conium maculatum,

Convolvulus arvensis, Fldeapmis PA Hyoscyamus niger, Lepidium latifolium, Linaria dalmatica, Linaria

vulgaris, Tamarix chinensis, and Ulmus pumila. Noxious weeds and other exotics are indicated by a * or an *,

respectively, in the Annotated Checklist.

TAXA OF CONSERVATION CONCERN

fol E i : à ] ++] A A |

A primary goal of a

cies of conservation concern were d ted from 49 sites in the Tusas and Jemez mountains and vicinity;

four additional records were verified from accessions at UNM. The status is based on plants so designated by

Natural Heritage New Mexico (2008) and New Mexico Rare Plant Technical Committee (2008). These taxa

are listed alphabetically and each is followed by family name, counties of occurrence (apparent new county

records in bold), and voucherís). They also are indicated by a € in the Annotated Che iust

lemic t A total of 22 spe-

Abronia bigelovii (Nyctaginaceae) Rio Arriba, Sandoval, Hackelia hirsuta (Boraginaceae) Colfax, Los Alamos, Mora,

Santa Fe: Larson 4113; Hartman 80977. Rio Arriba, See San Miguel, Santa Fe, Taos, Union:

Astragalus cyaneus (Fabaceae) Rio Arriba, Santa Fe, Taos: pees 636.

Larson 10438, 10480; Hartman 81069; Nelson 67934. Lilium philadelphicum (Liliaceae) Sandoval, San Miguel,

A Meum, (Fabaceae) Rio Arriba, Sandoval, Sani Fe: Jacobs 36

I \ Ni Arriba, Taos:

Nc micromerius (Fabaceae) McKinley, Rio Arriba, Pun 3626; Hartman 83194, 83298.

San Juan: Larson 4112, Mentzelia ) Rio Arriba: Reif 1973.

Astragalus ae (Fabaceae) Rio Arriba, Taos: Larson 2023; Mentzelia springeri (Loasaceae) Los Alamos, Sandoval,

Hartman 81215 x s.n; Hartman 75084; Jacobs 3202, 3296,

Cornus canadensis (Cornaceae) Sandoval, Taos: Hartman 3904, 5124.

18 RA LI |) H

75528, 75600, 75667; Jacobs 5183. g i (P ) Sandoval, Santa Fe: Hart-

Corydalis caseana ssp. brandegeei (Fumariaceae) Rio Ar- man 75187; Jacobs 3094, 3209, 4536.

riba: Lar 4533. Podist t li (A; i ) M , Rio Arriba, Santa

ie sessllifiorus (Apiaceae) Cibola, McKinley, Rio Fe, San Miguel: Hartman 75028.

pipe San Juan, Sandoval, Taos, Valencia: Larson 5024, | Ranunculus abortivus (Ranunculaceae) Rio Arriba, San-

1034 doval, Taos: aroba date cane, 6543, 6678.

aires parviflorum var. pubescens (Orchidaceae) S a, Rio Arriba, Taos: Hartman

Catron, Grant, Los Alamos, San Miguel, Santa Fe: Jacobs 75017.

3621,5 ] 12. Spiranthes romanzoffiana (Orchidaceae) Rio Arriba, San-

Epi, tea (Orchidaceae) Catron, Eddy, Grant, Gua- doval, Santa Fe, Taos: Hartman 78326.

dalupe, Harding, Los Alamos, Otero, Rio Arriba, PEHGOVah The following, also of conservation concern, were encoun-

Hartman 74454; Jacobs 3885; Larson 5027. tered in searches of the University of New Mexico database;

Bib copa lachnogynum var. colobum (Polygonaceae) identifications eye been venned TA puniceus

cKinley, Taos: Hartman 81819, ms verset r. grayi

ma decumbens var. s cisa (Asteraceae) Hidalgo, sa. T a ws PUTA

Grant, Rio Arriba, oa DOR 8648, 9233; Nelson — .. 4 NL cota ge betta /D MERE o

61956. mM

State Records for New Mexico

Species from the Tusas and Jemez mountains and vicinity that appear to be new to the flora of New Mexico,

Raif at al ; Fl f the Ti 1 | ; New Mexico 931

or that have had reports verified, are listed below (based on Allred 2008). A total of 28 apparent state records

based on 78 collections are included along with three additional records verified from accessions at RM and

UNM. The 18 taxa preceded by an * were documented in Hartman et al. (2006). Of these, Carex deweyana

var. deweyana previously had been reported by McGrath (2005). These taxa are listed alphabetically and

each is followed by the family name, the number of collections, and the counties of occurrence. They are

indicated by an ! in the Annotated Checklist.

13 ,Rio Arriba, Sandoval

IA L || .. mm

) 7, Rio Arriba,

ndoval

Astragalus cerussatus (Fabaceae) 3, Rio Arriba, Taos

ID ata | ar ¿Pr laa Ca doval

‘Cardamine cordifolia var. incana (Brassicaceae) 1, Rio

Arriba

C | (Cyperaceae) 2, Los Alamos,

Rio Arriba

aes fases ai Cyperaceae y A

opodium capitatu ) 1, Sandoval

eei echinatus (Poaceeae) 1 Sandoval

Descurainia pinnata var. paysonii (Brassicaceae) 1, Rio

Arriba

Epilobium campestre (Onagraceae) 1, Rio Arriba

Epilobium ciliatum var. glandulosum (Onagraceae) 4, Rio

Arriba, Sandoval

‘Equisetum xnelsonii (Equisetaceae) 1, Sandoval

‘Erigeron nivalis (Asteraceae) 3, Sandoval

‘Geum triflorum var. triflorum (Rosaceae) 1, Sandoval

+l + | l +

TI Eb +

“Juglans nigra (Juglandaceae) 2 Sandoval

'Lactuca biennis (Asteraceae) 2, Sandoval

Lepidium densiflorum Schrad. var. macrocarpum G. A.

Mulligan (Brassicaceae) 1, Rio Arriba

'Lepidium ramosissimum var. bourgeauanum (Brassica-

ceae) 25, Rio Arriba, Sandoval

Mimulus suksdorfii (Plantaginaceae) 1, Rio Arriba

'Neoparrya lithophila (Apiaceae) 2, Taos

T glaber var. alpinus (Plantaginaceae) 1, Los

Alam

‘Piptat ache cain pungens Pran Sandoval

runus persica (Rosaceae) 2, Sandoval

‘Ranunculus a var. la (Ranunculaceae

5, Rio Arri

‘Ranunculus ens (Ranunculaceae) 1, Sandoval

Silene drummondii var. striata (Caryophyllaceae) 2, Rio

Arriba

Sium suave eee 1, Rio Arriba

-- TI

| RA a | ere] sl att Et L L Hal

CELIN MZ

Ul INM 1,2 collections), Lomatium

grayi var. grayi (Apiaceae; UNM), and L. triternatum var. platycarpum (Apiaceae; RM).

SUMMARY OF TAXA

The first number represents results based on our fieldwork. Parenthetical numbers following most of the

former include those verified from other herbaria. The two adjacent numbers below “Exotic taxa” represent

the percent of exotics when compared to total Unique taxa.

List by taxonomic category

List by special category

Families 107 Exotic taxa 154 (158)

Genera 519 (533) Percent exotic taxa (11.1) (9.1)

Species 1,279 (1,334) NM Noxiou 14

Hybrids 10 Species conservator concern 22 (26)

Infraspecies 93 (98) NM Endem 17 (18)

State aa 28 (31)

Unigue taxa 1,384 (1,445)

List of unique taxa by major plant group

Fern Allies 7 (8)

Ferns 20 (23)

Gymnosperms 16

Angiosperms 1,341 (1,398)

Total 1,384 (1,445)

a I £ sl Dd = ID LI d dhea ake af

932 f Texas 3(2)

CONCLUSIONS

This paper represents the first of two contributions that cover the floristic diversity in north central New

Mexico. The area here covered encompasses the Tusas Mountains and the Jemez Mountains (including

Sierra Nacimiento) of the Carson National Forest and Santa Fe National Forest. Also included is Bandelier

National Monument, Valles Caldera National Preserve, and other federal, state, and private lands west of

the Rio Grande. We report on results of 19,929 numbered collections of vascular plants (total for the two-

part series covering more than 3.7 million acres is 35,857 new collections). A total of 1,384 unique taxa,

including 93 infraspecies as well as 10 hybrids, are documented from 107 families. With the addition of

60 taxa (totals within parentheses) verified from four herbaria, the total for unique taxa rise to 1,445. Of

these, 154 (158) are exotic taxa of which 14 are designated as noxious in New Mexico, 22 (26) are species

of conservation concern, 28 (31) represent first reports or their confirmation for New Mexico, and 17 (18)

are endemic to the state.

THE ANNOTATED CHECKLIST

The checklist is divided into major plant groups (f d fern allies, gj and angi ) each

with alphabetical listings by family and species. N lature follows Allred ( 2008) pi some uet dons as

treated in Hartman and Nelson (2001) or Flora of North America Editorial Committee, eds. (1993+). Where

these deviations occur, Allred's scientific name appears in square brackets below the accepted name.

Following is a guide to format and abbreviations associated with individual taxa in the checklist. Ex-

cept for records based on specimens at RM, or UNM, the citation of vouchers are omitted. This is justified

as collection data are available online (Hartman et al. 2009).

Taxon Authority [£, & -INITIALS, +, +] COUNTY; elevational range in feet; GEOLOGIC AREA; habitat

type.

[Taxon Authority, name accepted by Allred]

The number of collections for each project is based on the position of the pound sign (#) within square

brackets: [#,-,-,-] Jemez Mountains and vicinity, |-,#,-,-] Valles Caldera National Preserve, |-,-,#,-] Bandelier

Nasional Monument, and bs ; ^s duds Moun: and vicinity.

h specific ited, e.g., [UNM-Lowrey 1645] (RM,

Rocky Mountain Herbarium; UNM, Universi of New Mexico cea. USFS, National Herbarium of

the U. S. Forest Service at RM) and associated initials, e.g., [-,C,-,-] or [-,#+N,-,-] representing contributor of

VCNP data (C, Jonathan Coop; H, Natural Heritage New Mexico; N, Neal Osborne). Vouchers by Jonathan

Coop are at the University of Wisconsin (verified by Andrew Hipp) whereas those by the Natural Heritage

New Mexico or Neal Osborn (1966) are at the UNM (verified by Hartman).

County abbreviations: Geologic area:

L Los Alamos J Jemez Mountains, s. of Chama River

R Rio Arriba G Rio Grande Rift, s. of Black Mesa

D SanDoval P Taos Plateau, n. of Black Mesa

S SantaFe U TUsas Mountains, n. of Chama River

T Taos

Habitat type:

af dnd seral forest ml Marsh-lacustrine pj Pihon-juniper woodland

br Bur mm Montane meadow and grassland pp Ponderosa pine forest

ds ad shrubland mr Montane riparian ra Roadside-agricultural

fr Foodplain-arroyo riparian ms Montane shrubland sf Spruce-fir forest

mc Mixed conifer forest pg Plains-desert grassland

Symbols by category preceding Taxon:

* Exotic species to New Mexico 9 Species of conservation concern

* Noxious weed in New Mexico ! State record for New Mexico

+ Endemic to New Mexico x Hybrid

Reif at al Fl f the Ti ij ins. New Meyico

FERNS AND FERN ALLIES

Pspieniacede

A le (L) Hoffm. [1,3,1,1] D, R; 8300-9840’,

, U; mm, mc, mr.

Asplenium trichomanes L. [1,-,-,1] D, R; 8200-8460”; J, U;

Dennstaedtiaceae

Pteridium Ae (L) Kuhn var. pubescens Underw. [12,13,1,-]

350-9920’: J, U; af, br, mc, mm, mr, ms, ra.

Athyrium filix-femina (L.) Roth

54,10; E 6700- 10000’; J, U; mc, mm, mr, ms, sf.

r lis (L) Bernh. [9,7,3,3] D, L, R; 6600-10600’; J,

y; al, mc, mm, mr, ms, pp. : sf.

,20,1,-] D, L, R; 7450-10330;

J, U; af, mc, mm, mr, ms,

Dryopteris filix-mas (L.) Schott no 9,4,3] D, L, R; 6150-9325;

J, U; af, mc, mm, mr, ms,

Woodsia neornexicana aan A 1,1,1] D, R; 6150-8940"

J, U; mm, mr,

Woodsia oregana D.C. Eaton var. cathcartiana (B.L. Rob.)

Morton [-,3,1,2] D, R, T; 6660-10300’, J, P, U; ds, mm,

: PJ, PP.

Woodsia plummerae Lemmon [1,-,-,1] D, T; 7150-8460" J,

P; mm, pj.

Equisetaceae

Fguiset L. [31,8,6,10] D, L, R; 5500-10020’; J, G, P,

U; fr, m

r, ra.

x Equisetum xferrissii Clute [3,1,-,-] D, R; 5825-8600"; J; fr,

y A.A. Eaton [8,-,2,1]

L I fr: fT |

D, L, R; 6000- a J, "i fn mm,

un [19,4,2, prm L, R: 5550-8950’;

G, U; fr, ml, mm, mr, r

Ix Equisetum xnelsonii Hi Eaton) J.H. Schaffn. [-,-,1,-] D;

5300'J; mr.

Ophioglossacea

Botrychium SUR rn Gmel.) Ángstr. var. lanceo

[UNM-McGrath 250] R

Botrychium virginianum (L = [-,-,1,-] L; 71505; J; mr.

—

atum

Polypadiaceas

FHINAAGA/ ndham 357

VV I PRAT ICA B] R; U.

Polypodium saximontanum Wind ham [UNM-Stanley 11195]

Pteridac

eset stn fendleri O Windham [2,-,2,1] D, 5, T;

5400-7020". G, P; ds, fr, m

m eatonii Baker [-,1, i ,2] D, R, T; 6600-9050’; J, P,

ai de T Moore [2,-,1,3] D, R; 5500-7200; J, U;

ms, pj.

Cheilanthes fendleri Hook. [-,1,2,2] D, R, T; 5500-8670"; J, P,

7 F i

Cheilanthes wootonii Maxon [1,-,-,-] L; 7500-8900; J; mc.

933

IN Y LUN d

R. Br. [-,2,-,1] D, R; 8300-11150’;

Cry t he

J, U; mm, pp, sf.

lothol i ileyi M T ,1,-] D; 6000” J; pj

Pellaea truncata Goodding [-, A2, 1] D; 5400-5500" J, P; pj.

Pellaea wrightiana Hook. ier -] D; 61007; J; pj.

Selaginaceae

Selaginella densa Rydb. [-,-,-,81 R, T; 7150-10200"; P, U; mm,

Selaginelia mutica D.C. Eaton ex ld var. mutica [3,-,1,-]

oo 5200, J, G; ds, fr, ms,

Selag UNN Clark 9628] D; J.

~o F

GYMNOSPERMS

Cupressaceae

Juniperus communis L, var. depressa Pursh [37,33,2 eS D, L, R,

T: 7500-11150"; J, P, U; af, mc, mm, mr, ms

Juniperus pal Steud. var. deppeana [4,- E A D; 6600-

Juniperus monosperma (Engelm.) Sarg. [32,-,4, a D, R, T;

70-8460’; J, G, P, U; ds, fr, mr, ms, pg, pj, pp, r

iiis scopulorum Sarg. [24,4,3,48] D, R, T; SA 9400" J,

mc, mr, ms, pg, pj, PP, ra.

Edda

Ephedra cutleri Peebles [-,-,5

Ephedra torreyana Torr. ex S. Watson

R; 6100-6340"; J, U; ds, fr, pj.

Ephedra trifurca Torr. ex S. Watson [2,-,-,-] L, S; 5470-6100;

G; ds, pg.

-] D, 5600-5900 J; pj.

VAT torreyana [-,-,1,3] D,

Pinaceae

Abies arizonica Merriam [9,17,1,5] D, L R, T; 8340-111505 J,

U; af, mc, mr, sf.

Abies concolor (Gord. & Glend.) Hildebr. [49,42,2,23] D, L, R;

6800-10650’; J, P, U; af, br, mc, mm, mr, pp,

Picea engelmannii Parry ex Engelm. var. engelmannii

[9,19,1,12] D, L R; 8375-11150’: J, P, U; af, mc, mm, mr, sf.

Picea porns Engelm. [25,17,1,11] D, R; 7450-10200’; J, P,

, br, mc, mm, mr, pp, ra, sf.

Pinus edulis Engelm. [41,3,4 id D, L, R, T; 5860-9850’; J, G, P,

; ds, fr, mc, ms, pg, pj, Pp, r

Pinus flexilis E. James [10,6,2,2] Ü L, R; 7450-10225’; J, U; af,

r,mc, pp, $

Pinus ponderosa Douglas ex P. Lawson & C. Lawson var. sco-

pulorum Engelm. [66,27,1,36] D, L, R, T; 6150-10650’; J,

,B.U;br , mr, ms, pj, p

Pinus strobiformis Engeln: [18,21,1 b D L R; 8000-10650"; J;

mc, mr, ms, pp, S

Pseudotsuga menziesii (Mirb.) Franco var. glauca (Beissn.)

Franco [47,29,5,29] D, L, R, T; 6000-11150’; J, P, U; af, br,

mc, mm, mr, pp, sf.

ANGIOSPERMS

Aceraceae [as Sapindaceae]

Acer glabrum Torr. var. glabrum [37,21,3,5] D, L, R; 6400-

10900’: J, U; af, br, mc, mm, mr, ms, pp, sf.

[Acer glabrum Torr. var. neomexicanum (Greene) Kearney

& Peebles]

934

Acer negundo L. var. interius ai Sarg. [4,-,4,1] D, L, R;

5600-8280’; J, G, U; fr, m

Adoxaceae

Adoxa moschatellina L. 4] D; 8350- B473; Jı mm, mr.

-,-] D, R; 7200-10430' J; af, mc, mm, mr, , Sf.

Sameie racemosa L. var. microbotrys Rydb) Kearney &

Peebles [20,11,2,2] D, L, R; 7780-11254’; J, U; af, br, mc,

mm, mr, S

Agavaceae sonia ae

Yucca baccata T: baccata [ [2 711 ,7] D, R, 3 it 5860- 8630;

J, G, P, U; ds, mm, ms, pj,

+ Yucca intermedia McKelvey [9,- -1, ln D, R, T; 5500-8750" J,

, P, U; ds, fr, mc, ms, Pg, pj, PP, r

[Y. baileyi Wooton & Standl. var. o” (McKelvey)

Reveal; see Sivinski 2008]

Alismataceae

Alisma subcordatum Raf. [UNM-Atwood 21196] R;

Sagittaria cuneata E. Sheld. [1,1,-,1] D, R; 6500- Y J, U;

ml, mr.

Alliaceae bei in pian

Allium bisceptrum S. Watson [1,-,

Allium LU Roth [15,27,1 21] D, LRT 7300- dm up

P, U; af, br, ds, mc, mm, mr, ms, pj, pp.

id His Watson var. geyeri [3, 17, 1,5] D, R; 7550-9625".

; af, mc, mm, mr, ms, pp.

ms gon > Watson var. tenerum M.E. Jones [-,-,1,2] D, R;

0 J, U; m

i beide "t f. ,1,-] D, R; 6250-7360"; J; ds,

ms textile A. Nelson € J. F. Macbr. [-,-,-,5] R, T; 6200-7150";

P, U; fr, pj.

Amaranthac

Amaranthus s L. [2,-,2,3] D, L, R, T; 5450-8740"; J, G, P, U;

fr, ml, mm, pj, r

Amaranthus Li I.M. Johnst. [-,-,2,-] D; 5800-6000";

J; pp.

* Amaranthus blitoides S. Watson [-,-,1,1] L, T; 6500-8500’;

; mi, pj.

saan hybridus bbs 5,] D; 6000-6450’; J; pj.

[-,-,1,-] D; 5300’; J; fr.

Amaranthus powellii S. Watson [-,1,-,7] D, R, T; 6975-8400": J,

> U; ds, mr, pj, pp, ra.

* Amaranthus retroflexus L. [4,5,4,-] D, L, R; 5400-9220"; J, G,

P; iu. 2 pj, Fe ra.

A. Gray) Benth. ex S. Watson [UNM-Yarnell

D; J.

Amaranthus wrightii S. Watson [-,1,-,-] D; 8200-9054’; J; ms.

Anacardiaceae

Rhus glabra L. [2,-,3,-] D, L; 5500-7250"; J; mr, ra.

Rhus trilobata Nutt. var. trilobata [35,2,8,15] D, L, R, S, T;

aa P80; J, G, P, U; ds, UR mc, mr, ms, pj, iiA ra.

2,-] D, R

6300- 3500’: J; B mc, mm, mr, m pi, pp.

|| [| £ al n.a a in LJ a 4 de

f Texas 3(2)

Apiaceae

Aletes acaulis (Torr) J.M. Coult. & Rose [8,-,3,-] D; 5850-7500;

, G; ms, pj, pp.

Angelica pinnata S. Watson [2,-,-,3] D, R; 7100-10020;5 J, U;

D j (Hire Cov ille var incisa (Torr) C KAT I ict 79 _ 2 .1

D, R; 5300- 7350: J; fr mr.

Cicuta maculata L. var. angustifolia Hook. [16,4,3,-] D, L, R;

5825-8375": J: fr, ml, mm, mr, ra

(A Gray) JM. Coult. & Rose [3,17,1,5]

D, R; 6950- 11000’; J, U; af, mc, ml, mm, mr, ra.

*eConium maculatum L. [1,-,-,-] D; 74905; J; m

! Cymopterus spec. nov. [-,-,-,4] R, T; 6200- el P, U; ds, fr,

Pri I 1 Lær pp d

Cymopterus alpinus A. Gray [-,3,-,-] D; 8600-8700"; J; mc,

mm, sf.

Cymopterus bulbosus A. Nelson [1,-,1,3] L, R; 6400-8500’; J,

U; ms, pg. pj

Cymopterus constancei R.L. Hartm. [1,-,1,6] D, R, 5, T; 5550-

8500’: G, P, U; ds, fr,

Cymopterus glomeratus Mus A.P. de Candolle var. fendleri

(A. Gray) R.L. Hartm. [2,-, 1,1] D, S, T; 5600-7150" J, G,

7 OS, DJ.

Cymopterus lemmonii (J.M. Coult. & Rose) Dorn [55,54,3,45]

D, L R, T; 7300-11150% J, P, U; af, br, mc, mm, mr, ms,

PP, r

Gymopten monens teme: Gray [1,-,-,-] D, 7360"; J; ds

4 L.Theob. & C.C. Tseng) R.L. Hartm

I j 42] R; 6300- 7250 U; fr.

/ Bartram [16,4,1,5] D, R; 6950- 11000;

J, U; af, mc, ml, mm, mr, sf.

I I.M. Coult. & Rose [9,

+5] D, R 8180-10900;

^L U; af mc, mm, mr, sf.

1 Lomatium grayi (J.M. Coult.) J.M. Coult. & Rose var. grayi

[UNM-Hei/ 8814] R; U

| Lomatium triternatum (Pursh) J.M. Coult. & Rose var. platycar-

Torr.) B. Boivin [UNM-Heil 8793] R; U.

| Neoparrya lithophila Mathias [-,-,-,2] T; 7900’; P; pj.

Osmorhiza chilensis Hook & Arn. [1,-,-,1] D, R; 7370-9150;

Osmorhiza radica Phil. [29,17,5,3] D, L, R; 6600-10600';

; af, br, mc, mm, mr, ms, pp, sf.

Oxypolis ae (A. Gray) A. Heller ipd 13,1,10] D, L, R;

7740-10900"; J, U; af, br, mc, ml, mm,

$»

4. Coult. & R ans

& Const

[1,-,-,-] R; 9300 10135':J

Sanicula marilandica L. [-,-, ne b: 7200- 7800"; J; mr.

! Sium suave Walter [-,-,-,1] R; 9150% J; mr.

Apocynaceae

Apocynum androsaemifolium L. [8,2,2,1] D, L, R; 7250-9100;

F , F i Sy pp, ra.

Apocynum cannabinum L. [7,2,2,-] D, L; 5500-8300"; J, G; fr,

5, PO, ra.

:L f y rg

Greene [5,-,-,-] D, R; 6875-8125;

J; mr, ra.

[Apocynum medium Greene var. floribundum (Greene)

Woodson]

Reif et al., Fl f the Ti 1 | tains. New Mexico

Araliaceae

Aralia racemosa L. ssp. bicrenata (Wooton & Standl.) S.L

Welsh & N.D. Atwood [3,-,4,-] D, L, R; 5850-8550; J, U;

mc, mm, mr

Asparagaceae [traditionally in Liliaceae]

* Asparagus officinalis L. [2,-,1,-1 D; 5825-7800’; J; fr, mm, ra

Asclepiadaceae.

L S 5600- 6320’ G, P; ds. ft, B.

Asclepias engelmanniana Woodson [-,-,2,-] D; 5300-6000";

J; fr, pj.

Asclepias involucrata Engelmann ex Torr. [-,-,-,3] R, T; 6400-

7150"; P, U; fr,

Asclepias macrosperma Eastw. [-,-,-,3] R; 6100-6990’; P, U; pj.

Asclepias macrotis Torr. [1,-,4,-] L, D; 5300-6600’; G; ds, fr, pj.

Asclepias speciosa Torr. [9,-,1,-] D, L, R; 5400-8500'; J, G, P, U;

r, mm, mr, ra.

Asclepias subverticillata (A. Gray) en [7,1,6,3] D, L, R; 5300-

8450; J, G, P, U; fr, mm, mr,

Asclepias tuberosa L. var. interior edo Shinners [2,-,4,-]

; 590 50’; J; mr, pp, ra.

Asclepias viridiflora Raf. [-,-,3,-] D, L; 6700-7300% J; pj, pp.

Asteraceae

Achillea millefolium L. [46,63,6,38] D, L, R; 6400-111505 J, P,

U; af, br, mc, mm, mr, ms, pp, ra, 5

*eAcroptilon repens (L) DC. [3,-,1,-] L, R, T; 5550-8200; J, G,

; fr, pp, ra.

hert (A. Gray) R.M. King & H Rob. [-,2,3,1] D, R;

. 8625- 9220" J, U; mc, ms, pp, ra.

Agoseris aurantiaca (Hook) Greene var. purpurea (A. Gray)

Cron m t [4,29,2,17] D, R; 8000-11150; J, U; br, mc

Ages glauca (Pursh) Raf. var. glauca [-,9,1,4] D, R; 7550-

U; mm, mr,

— pesos (Nutt) a [2,-,-,-] D, R; 8000-8750;

NUS puc (Nutt) A. Lave & D. Lóve [1,-,-,-] D;

632 40’; J; ml.

a dissecta (A. Gray) Rydb. [2,8,4,3] D, L, R, T;

7020-95807; J, P, U; mm, mc, pg, pj, pp,

Ambrosia E Hook. [-,-,3,2] R, T; Pm 9580; J, P,

Ambrosia artemisiifolia L. [1,-,-,-] L; 63005; G; ra.

Ambrosia psilostachya DC. [2,1,-,-] D, R; 7675-9100"; J, U;

Ambrosia tomentosa Nutt. [2,-,-,-] D, R; 8050-9200; J, U;

Ambrosia trifida L. var. trifida [-,-,1,-] D; 5400"; J; fr.

Anaphalis margaritacea (L.) Benth. & Hook. [-,-,2

;mr.

Antennaria corymbosa E. Nelson [1,7,-,-] R; 9300-101355; J,

r] R; 7150;

ta G [15,7,4,7] D, L, R; 6400-10300’;

; af, mc, mm, mr, pp, sf.

Antennaria oo ie [-,-,-,22] R; 7450-10900"; P, U;

af,

935

Antennaria parvifolia Nutt. [17,11,3,13] D, L, R, T;6400-11150';

3 P, U; af, Dr, mr mc, mm, mr, ms, E

0] D, L, R; 6500- 10600’; J, U;

af, mc, mm, mr, pp

[Antennaria rosea subspecies]

Antennaria rosulata Rydb. [-,4,-,13] D, R; 7780-10940’; J, U;

, mm, ms,

Aphanostephus ramosissimus DC. var. humilis (Benth.

Turner & Birdsong [-,-,1,-] D; 5400’; s fr.

* Arctium minus (Hill) Bernh. [1,-,-,-] R; 6010-70007; J; mr.

Arnica pla Less. var. foliosa (Nutt.) Maguire [2,1,1,5]

400-9950’; J, U; ml, mm, mr, fr.

Amica cordifolia Hook. [1,-,-,4] R; 9300-10150; J, U; mc,

B.L.

—

mm, mr.

* Artemisia biennis Willd. var. biennis [-,-,-,1] R; 8720-8740;

U; ml.

Artemisia OGEI A. ay [1,- ‘2r 7 D; Due 7400; 5 ds, pj.

[-,-,-,2] R, T 6820-7000’, P, U; fr pj.

Artemisia - L. var. pacifica (Nutt) M. Peck [6,9,1,5] D,

RT; a Y: ,P,U; mc, mm, mr, pj, pp, ra.

A ri ici ij ood ex Carruth [3,22,2,12] D, L, RE

6300-9980" J, G, bu ds, mc, mm, ms, pj, pp, ra.

Artemisia dracunculus L. [-,6,2, D, R, T; 6200-10200’; J, P,

; ds, fr, mc, mm, mr, pj, p

Artemisia filifolia Torr. [-,-,3,2] E E 5300- 7280"; J, P, U; ds, pj.

Artemisia franserioides Greene [1,13,2,-] D, L; 7200-9980"; J;

Artemisia frigida Willd. [-,5,4,9] D, R, T; 5850-9850’; J, P, U; ds,

mc, mm, mr, PS, pj, ra

illd. ssp. parryi (A. Gray) W.A. Weber [-,1,-,-]

D; 10340-10875’; J; sf.

Artemisia ludoviciana Nutt. var. mend (Wooton) Shinners [-,-

,3,2] D, T; 5600-7736'; J, P; ds,

Artemisia ludoviciana Nutt. var. ludovciana [2,14,3,4] D, R;

6300-10300; J, U; mc D, ra

Artemisia ludoviciana en var. mexicana 1 (Willd. ex Spreng.

A. Gray [1,2,5,4] D, R; 5850-9350’; J, U; mc, mm, mr, ms,

pj, pp, ra.

Artemisia nova A. Nelson [-,-,-,4] R, T; 8300-9020"; P, U; ms,

pj, p

mera tridentata Nutt. var. ddl [1,-,3,19] D, R, T;

5500-8950"; J, P, U; ds, fr, mm,

Artemisia tridentata Nutt. var. vano Fab) B. Boivin [1,-,

00-8700’; J, U; m

a a Nutt. var. wyomingensis (Beetle € A.M.

ou L. Welsh [-,-,-,3] R, T; 5400-7350’; P, U; fl, pj.

Nh ee Torr. & A. Gray [3,-,1,-] D, L; 5400-5900;

Baccharis wrightii A. Gray [1,-,-,-] 5; 6320"; G; fr.

Baileya multiradiata Harvey € A. Gray ex A. Gray [-,-,1,-] D;

5 Js TE

Berlandiera lyrata Benth. [1,-,4,-] L; 5400-5800" G; fr, ds, pj.

Bidens cernua L. [-,2,1,-] D, R; 5360-8600"; J, U; mm, mr, fr.

Bidens comosa (A. Gray) Weigand [-,-,1,-] D; 5460"; J; fr.

[Bidens tripartita L.]

Bidens frondosa L. [-,-,2,-] D; 5300-5400; J; fr.

936

i tenuisecta A. Gray [-,7,-,1] D, R; 8025-9200’; J, U; mm,

5, PP, ra

sca bcp A. Gray [1,-,5,1] D, L, R; 6100-7120",

U; pj, D

side cofomic (Torr. & A. Gray) A. Gray [1,1,1,-] D;

Q- ; ; Ims,

"ues pda (L) Shinners var. chlorolepis (Wooton

& Stand) B.L. Turner [-,1,3,1] D, R; 6300-8800’; J, U; fr,

mm, ms, pp.

k.) Nutt. [1,10,1,-] D, L, R; 5850-9960":

J, U mom

Brickellia merphyo Am A. Gray var. scabra A. Gray [-,-,-,1]

6300-6600’; U; pj.

Brickellia ase olay Eo r] D, R; 2005 J, U; ms.

Ki ob. [4,3,4,3] D, R;

7950-8700; J, U; mc, mm, mr, ms, pp, ra.

*@Carduus nutans L. [2,2,2,2] D, R; 7540-9700’: J, U; br, mc,

mm, pp, ra.

Chaenactis douglasii (Hook) Hook. €: Arn. [1,-,-,-] R; 7100’;

; p).

ers ericoides (Torr.) G.L. Nesom [26,-,3,25] D, L, R, S, T;

060"; J, G, P, U; dg, ds, fr, mm, ms, pg, pj, pp, ra.

Chatham aH Nutt. [-,-,-,4] R, T; 7185-8655’; P,

mm,

Chosothamnas greeneii (A. Gray) Greene [-,-,-,9] R, T; 8050-

rf " 5, pg, pj,

Chnsorhamnis vaseyi (A. Gray) Greene [-,-,-,4] R, T; 7800-

0 ; ms, pj

Ghaysothamnus viscidiflorus (Hook.) Nutt. var. oo

(Nutt) Greene [-,-,-,2] R; 8720-9500"; U; m

Chrysothamnus viscidiflorus (Hook.) Nutt. var. ey ae [1,-

7,2] R, 16120-90205 P, U; ds, ms, pj.

Cirsium arizonicum (A. Gray) Petrack var. bipinnat

Keil [2,-,-,-] D, R; 6250-7330"; J, U; ms, pj.

*eCirsium arvense (L) Scop. [-,9,1,7] D, R, T; 5400-11000’; J,

U; fr, mc, ml, mm, mr, ra.

Cirsium neomexicanum A. Gray [4,3,3,6] D, R, T; 5860-9000’;

, B, U; fr, mm, ms, pj, pp, ra

Cirsium ochrocnmum A. Gray var. ochrocentrum [1,-,-,-] R;

(Eastw.)

Cirsium paroj iu Gray) Petrak [13,32,1,2] D, L, R;6950-10300';

c, mm, mr, ra, S

Cirsium Ge ean (Nutt.) Soreng. var. undulatum [4,4,1,-] D,

R; 6875-9580": J, U; fr.

*eCirsium vulgare (Savi) Ten. [2,10,3,-] D, L, R; 6013-9800"; J,

U;a m, mr, ra.

" oryza E (L) Cronquist [4,7, 6, 1] D, L, R, S;

6090-9200’; J, G, P, U; fr, mm, mr, ms, pp, ra

[Conyza canadensis varieties]

Cosmos Mp ids (Jacq.) Persoon [-,-,

Jra.

2,-] L; 7050-7450;

} lie Ml arcidonrstic [-, iol ] R; Q^5nn edu aT au

f Qa y

U; ms.

Crepis runcinata (E. J an mi 5 » 7? var. mE (Greene)

B.L. furner [1 0; J; m

Mes runcinata (E. es Torr. 8 A Gry var. obi (Nutt.) B.

oivin [UNM-Halley 20] D; J.

Crepis runcinata (E. James) Torr. & A. Gray var. runcinata [-,6,-,-

—8710'; J; ml, mm, mr

Cyclachaena xanthifolia (Nutt) Fresen. [-,-

3,1] D, R; 5400-

Dieteria asteroides Torr. var. asteroides [-,-,3,-] D, L; 6400-7800";

pj, pp.

Dieteria bigelovii (A. Gray) Morgan €: R.L. Hartm. var. bigelovii

[ 9] D, L, R, T; 7675-10700": J, P, U; mc, mm, mr,

pp, ra.

Dieteria canescens (Pursh) Nutt. var. ambigua (B.L. Turner)

g R.L. Hartm. [2,-,-,-] L; 5800-6300"; J, G; pp, ra

Dieteria canescens (Pursh) Nutt. var. aristata (Eastw.) Morgan

R.L. Hartm. [-,-,-,9] R, T; 7300-9550" P, U; mr, ms, pj,

pp, ra.

Dieteria canescens (Pursh) Nutt. var. glabra (A. Gray) Morgan

& R.L. Hartm. [7,-,-,1] D, L, R; 6000-9100; J, G, P, U; ds,

, pp, mi, ra.

Dyssodia papposa (Vent.) Hitchc. [-,-,1,-] D; 6000’; J; pp.

Ericameria nauseosa (Pall. ex Pursh) G.L. Nesom & GJ. Baird

var. bigelovii (A. Gray) G.L. Nesom & G.l. Baird [1,-,-,2] L, R,

T; 6300-8150"; G, P, U; ds, pp, ra

Ericameria nauseosa (Pall. ex Pursh) G.L. Nesom & G.I. Baird

var. d (Nutt.) Reveal €: Schuyler [-,1,2,7] D, R, T;

-90 ; ds, mm, ms, pg, pj, pp, ra

Bene nauseosa ; (Pall ex Pursh) G.L. Nesom & G.I. Baird

var. holo PUE (A. Gray) G.L. Nesom & G.l. Baird [-,-,-,1] T;

x pj.

Ericameria nauseosa (Pall. ex Pursh) G.L. Nesom & G.I. Baird

var. nitida (L.C. Anderson) G.L. Nesom € G.I. Baird [-,-,-,1]

R; 300-6606 U; fr

Ericameria nauseosa (Pall. ex Pursh) G.L. Nesom & Gl. Baird var.

oreophila (A. Nelson) G.L. Nesom & Gl. Baird [-,-,-, 11] R, T;

6200-9200; P, U; ds, fr, mm, pg, pj.

Mee parryi (A. Gray) G.L. Nesom var. attenuata (M.E.

s) G.L. Nesom & GL. Baird [-,-,2,2] D, R, 5; 6650-9200";

i pj,

Ericameria parryi (A. Gray) G.L. Nesom var. howardii (Parry ex

A. Gray) G.L. Nesom € Gl. Baird [1,-,-,-] D; 7000-7400"

J; pp.

Fricameria parryi (A. — G.L. Nesom var. parryi [-,-,-,4] R;

- ^U;afm

Auc pedi ac. var. bellidiastrum [1,-,-,4] R, T;

55 J, P U; ds, fr,

NM canus A. Gray [-,-,1,2] L, R, T; 5500-8700’; J, P, U;

Erigeron concinnus (Hook. & Arn.) Torr. & A. Gray var. concinnus

Pre Mo F , FS *

geram COUMEN I "oner De i »2]' R; a 11000"; U; mc, mr.

9,31] D, L R, S, T; 5300-

10600; J, G P, U; br, fr mc, mm, mr, ms, pj, op, ra, sf.

d elatior (A. Gray) Greene [-,-,-,1] R; 9980-10020"; U;

mr, sf.

Erigeron ager A. Gray [62,52,3,48] D, R, T; 6500-11150";

r, mc, mm, mr, ms, pg, pj, pp, ra, sf.

Erigeron us Greene var. E [6,48,4,8]

D, R; 7550-111505 J, U; mc, mm, mr, ra, sf.

[4,25,1,4] D, L, R; 8060-10600"; J, U;

Reif et al ,H f the T; ij ins, New Mexico

Erigeron formosissimus Greene var. viscidus (Rydb.) Cronquist

[4, "i D, L, R; 7800-10940*; J, U; mc, mm, mr, m

pp, ra, sf.

ird stanton Hook. [1,-,-,1] R; 9300-10135’; J, U;

inn e Hook. [-,2,-,-] D; 8620-9850"; J; mr, sf.

! Erigeron nivalis Nutt. [-,3,-,-] D; 10160-10360"; J; mm.

Erigeron ecc L. var. philadelphicus [1,-,1,-] L;

5400-5800’; G; fr.

Erigeron pulcherrimus A. Heller [-,-,-,6] R; 6050-10900’; P, U;

ds, pj, pp, mc, mm.

Erigeron pumilis Nutt. var. pumilis [-,-,-,2] R; 6800-7000’; U;

s, ms.

Erigeron speciosus (Lindl.) DC. [3,7,3,3] D, L, R; 7150-9400"; J,

; br, mc, mm, mr, ms, pj, pp, r

Erigeron strigosus Muhl. ex Willd. E 7&5] R, T; 6240-7300’; P,

; ds, pj.

Erigeron subtrinervis Rydb. ex Porter & Britton [14,20,2,20]

D, ih 7 d 10900’; J, P, U; af, br, ds, mc, mm, mr, ms,

pj, p

d er Greene [11,1,3 D, L, R, S, T; 5440-9000” J, G,

; r, ds, fr, mm, mr, pg, pj, pp.

Ene vetensis Rydb. [-,-, ges R: 7500-10600"; U; mc, mm,

ms, pp.

Erigeron vreelandii Greene [1,1,1,1] D, L, R; 7020-9500’; J,

C, p

in: f fA r^ A f

E.E. Lamont [UNM-lvey s sn] D4

Gaillardia aristata Pursh [2,1,-,-] D; 7330- 8500" J; ms, pp, ra.

canada a daTorr. idi -,4,3] D, L, R, S; 5500-8500"; J, G,

7 t PO, pj, pp,

Gaillardia pulchell ES Ds

hrimoridA Gray)

5,-] D, L; 6000-6300’; J, G;

m, pp.

uid exilifolium A, Nelson [-,14,1,4] D, R; 7550-9980’;

ml, mm, mr, ra

Gnaphalium palustre Nutt. [-,-,2,-] D; 5360’; J; fr.

Grindelia arizonica A. Gray [-,-,-,3] D, R; 8250-8550’; J, U; br,

[Grindelia arizonica varieties]

+ pénis decumbens Greene var. e E Iu

Steyerm. [1,-,-,2] D, R; 7200-9150’; J, U; ds,

Gmel ce (Pursh) Dunal [8,-,3,15] D, E A T; 5760-

,G, P, U; ds, fr, me, mm, ms, 5, Pg, pj, pp, ra.

sby 16, 8, 8,41] DL, R,

a 5400- ad: J, G, P, U; ds, fr mm, ms, , P9; pj, pp, ra.

L F F'"rF dJ

D, L; 5400"; J; fr.

iei parryi A. Gray [1,-,-, 10] R; 8280-10940": J, U; af,

c, mm, mr, pp.

To Mies quinquenervis (Hook.) A. Gray [-,-,1,2] D, R;

7850-10000; J, U; br, mm

Helianthus annuus L. [3,-,2, 5] D, L, R, T; 6090-8400"; J, PU;

ds, ml, p

Helianthus CU Yon & A. Gray [1,-,-,-] D; 7675; J; ml.

Helianthus pauciflorus Nutt. var. subrhomboides (Rydb.) Cron-

quist [2, i 2, u D, R nue Bascal J, u; Dr mr, pp.

L. Turner [1,-,-,1]

L; 6300’: G; mm, ra.

937

bcn cr Nutt. var. aerea [2,-,3,1] D, L, R, S;

-—6975', G, P, U; ds, fr,

Tun multiflora Nutt. var. dim [2,16,5,15] D, R;

a E J, P, U; af, br, mc, mm, dd PP, ra, sf

Li EEN WAIL Vat

[3,-,-,3] D, R, T; 7600-9800’; J, G, P, U; mc, ms, B pp, r

Herrickia glauca (Nutt) Brouillet [-,-;-,1] R; 6300- 6600; U; :

[19,19,-,17] D, L R, Ü 5825-10225". J, G, P, U; af, br, fr, me,

mm, mr, ms, pg, pj, PP, ra, sf.

(A. Gray) nis

[12,12,-,23] D, L R, T 5760- 9980" i G, P, U; af, mc, mm

ms, PG, pj, pp, ra

H Semple [-,- ,2] R; 9850 10500; Ui mm.

M LF

IN CL " X ]

* 1

-L Hil fnr

L,-,-,1] R; 8200-8350* U; op.

mucus villosa (Pursh) Shinners var. villosa [19,8,7,15] D,

L, R, 5, T; 5470-10600"; J, G, P, U; ds, fr, mc, mm, mr, ms,

pg, Pj, PP, ra

Migraceum Cameum meresne [-,-,1,-] L; 7200; J; pp.

h. Bip. [7,10,2,6] D, L, R; 7250-11150’; J,

U; br, mc, ms, pp, ra, sf.

Ix Hieraceum xfloribundum Wimm. & Grab. [-,-,1,2] R;

9000- aun , U; mm, Tm

ty Z

mm ILAIA

4. Johnst

116, 10,21] D, L R, $, T; 5500-9450’: J, G, P, U; ds, fr, mc,

mm, ms, 2: pj pp, re ra.

j pap) k. var. pauciflorus (I.M. Johnst.) B.I

Turner [-,-,-,4] R; 6300-6600’; U; fr.

bd bid pd A. Gray var. canotomentosus A. Gray

0-6150'; J, G, P; ds, fr, pg, pj.

o e (A. Gray) I.M. s [18,17,-,-] D,

R; 7330-10330’; J; af, br, mc, mm, mr, ms, pp, ra.

Hymenopappus tenuifolius Pursh [2,-,-,-] R, S; 6320-6880’;

Hymenoxys hoopesii (A. ml Bierner n4 21,3,5] D, R;

8000-11150; J, U; af, m

Hymenoxys richardsonii Hook) deed var. floribunda (A (A.

Gray) Parker [31,17,5,51] D, L,R,T;5710-9750” J, G, P, U;

br, ds, ue mc, mm, mi ms, pj, pp, ra.

,2] R; 6100-6200’; U; fr, pj.

11D, R, T; 5400-8200"; J, G, P; fr, mm,

iva ollas Pursh [1, 1-

mr, ra.

Krigia biflora (Walt) S.F. Blake [-,2,1,-] D, L; 7200-8710; J;

ml, mm, mr.

! Lactuca biennis (Moench) Fernald [-,-,2,-] D; 6000-6500":

J; mr.

Lactuca canadensis L. [UNM-Clark s.n] D, L; J

Lactuca amino Michx. var. arizonica McVaugh [4,4,2,-]

300-9140"; J, P; fr, mm, mr, ms, ra.

lactuca pulchella (Pursh) DC. [2,-,2,-] D, R; 6000-7100’; J, U;

s jgena serili L. [3,2,3,3] D, L, R, T; 5960-8080"; J, G, B. U;

br, fr, mm, pj, pp, ra.

Laennecia schiedeana (Less.) G.L. Nesom [1,6,2,-] D, L:

8040-9980"; J; br, mc, mm, pp, ra.

938

* Leucanthemum vulgare Lam. [4,6,-,-] D; 7860-9150"; J;

mm, mr, ra.

Liatris punctata Hook. var. punctata [1,-,4,-] D, L; 6500-82007,

+ L at AW. VI + O C4 ll^ l.l +

Neubig [2,-, 42] R, T; 6000-6700’; a P; ds, fr,

Lorandersonia linifolia (Greene) Urbatsch, Roberts, & Neubig

ee R; 5760- siis J, G; PS, pj, ra.

= ist) Urbatsch, Roberts,

& Neubig [-,-,-,3] R; 8100- 8600" U; op.

Lygodesmia juncea a D. Don ex Hook. [3,-,-,-] D, R;

6500-7000’; J; fr, pj, r

dd rice raro (Kunth) Nees [-,-,2,4] LR, S, T;

6500-7280’; J, P, U; d

Had gomet cee E a 410] D, R, T; 8450-10230’; J, U;

h, Roberts, &

Malacothrix eae A. Gray [1,-,2,-] D, L; 5550-6600"; G; fr,

Malach sonchoides (Nutt.) Torr. & A. Gray [-,-,-,1] R;

6200’; U; ds.

ü vous s DC. [4,-,-,-] D, R; 7490-8850’; J; mm,

mr, ms,

T e leucanthum Torr. & A. Gray [1,-,3,-] D; 5300-

6460"; J, G; fr, m

Oreochrysum parryi "i ea Rydb. [5,31,2,6] D, L, R; 6150-

11000’; J, U; af, mc, mm, mr, pp, ra,

irs cana (Hook) W.A. Weber & A. e [-,1,-,-] D; 8500’;

J; pg.

acie crocata (Rydb.) W.A. Weber & A Lóve [-,1,-,-] D, R;

a dimophopy la (Greene) WA. Weber & A Lóve var.

dimorphophylla [1,-,-,-] R; 9300-10135% J; mc

a fede ^ e Weber & A Lave [14, 23, 3,-] Dt,

G; af, mc, mm, mr, ms, pj, pp, ra, sf.

Packera multiobat m & A. Gray ex A. Gray) W.A. Weber

& A Lóve [4,-,1,6] D, R, T; 6300-7700’; J, P, U; ds, fr, pg,

pj, PP, ra.

Packera neomexicana (A. Gray) W.A. Weber & A Lóve var.

mutabilis (Greene) W.A. Weber € A Lóve [21,26,4,18]

D, L R, T; 6050-9850", J, P, U; af, br, ds, mc, mm, mr, ms,

pj, pp, ra, sf.

Packera Lamesa (Rydb.) W.A. Weber & A Lóve var. flavula

(Greene) D. Trock €: T.M. Barkley [3,2,2,2] D, L, R; 7000—

mc, mm, mr, sf.

Packera streptanthifolia (Greene) WA. popa Lóve [6,-,-,3]

, R, T; 6875-10180; J, P, U; af, ds, m por

Packera thurberi (A. Gray) B.L. Turner I2. - R; D 8720";

Palafoxia sphacelata Es ex ui d n -- rel ii 5955% J; fr.

Pectis Veo d fore var. ar ET 1] a R; 5400-

r, pj.

Pericome caudata A. ae 2,3,1] D, L, R, T; 5850-8580’; J, P,

Petradoria pale mm Greene var. pumila [1,-,-,6] D, R, T;

7000-8700’; J, P, U; pj, pp.

Pseudognaphalium canescens (DC.) W.A. Weber [-,-,1,-] D;

6500"; J; pj.

O m A LÁ +

5; 7260- 10300’; J, U; mc, mm, ms, p

ET stramineum Kn n Weber [-,-,1,-]

[1,10,2,-] D, R,

LAM MM (A. Gray) A. Nelson [UNM-Castetter

8997] Lx.

ide en deis Greene [5,-,4,3] D, R, 5; 5300-

G, U; ds, fr

ios ao io Don) DC. [-,-,1,-] L; 5500% J; fr.

Pyrrocoma crocea (A. Gray) Greene var. crocea [1,2,2,3] D, R;

8 7 100005 J, U; af, im. mr

& Standl. [6,1,1,1] D, R,T;

6125-8375; J, P, U; ds, fr mm, po, pj, pp, ra.

[Ratibida columnifera formas]

t [1,-,3,1] D, L; 5400-6100";

, G;fr, p

Rudbeckia [m L. var. pulcherrima Farw. [14,12,2,5] D, L, R;

7100-10200’; J, U; af, mc, mm, mr, ms, ra.

Rudbeckia laciniata L. var. ampla (A. Nelson) Cronquist

[22,15,2,2] D, L, R; 5850-9625” J, U; af, mc, ml, mm,

mr, sf.

Sanvitalia abertii A. Gray [-,-,1,-] D; 58005 J; pj.

Schkuhria multiflora Hook. & Arn. [-,3,1,2] D, T; 6420-8625

J, P; ms, pg, pj, pp, ra.

* Scorzonera laciniata L. [9,1,-,3] D, R, S; 6200-8750; J, G, U;

g, pj, pp, ra.

a amprectens a Gray var. amplectens [-,7,-,1] R;

0700-11000"; U; s

mes atratus RA [1,8,1,1] D, L, R; 8200-11150' J, U;

r, mc, sf.

us ee A. Gray var. hallii A. Gray [-,8,3,2] D, R;

000; J, U; af, br, mc, mm, mr, ra, sf.

ms us Richardson var. kingii (Rydb.) Greenm.

[3,38,1,5] D, L R, T; 7150-10300% J, P, U; af, br, mc, mm,

mr, ms, pj, pp, ra, sf.

[Senecio eremophilus var. macdougalii (A. Heller) Cron-

quist

es Poa Less. var. flaccidus [6,-,2,5] D, L, R, 5, T;

6500’; J, G, P, U; fr, mr, pg, pj, ra

E en & A. Gray [1,-,1,7] D, R, T; 6000-8655;

J, P, U; ds, fr, pj, DB:

Senecio triangularis Hook. [2,-,-,2}] D, R; 8120-110005; J, U;

mm, mr.

Senecio wootonii Greene [14,13,3 Pa D, L, R; 6500-11150; J,

; af, mc, mm, mr, ms, pp, Y

Solidago altissima L. ssp. a (Rydb.) Semple [-,-

4,4] D, R, T; 5400-8655’; J, P, U; fr, pj, pp.

Solidago lepida DC. var. salebrosa (Piper) Semple [1,-,3,-] D, L;

5400-7100’; J, G; fr,

Solidago missouriensis Nutt var. fasciculata Holzinger [-,-,-,1]

Solidago missouriensis Nutt. var. missouriensis [2,10,1,2] D, L,

7550-10200’; J, P, U; af, mc, mm, mr, pp, ra

C. Taylor & J. Taylor [1,-,-,-] R; 9100"; J; mm.

Solidago mollis Bartl. [-,-,1,21 D, R; 6700-8500"; J, U; br, ms,

pp.

Solidago nana Nutt. [1,-,-,1] L, R; 7800-9800; J, U; mc, pp.

Reifetal Fl f the Ti a ins. New Mexico

Solidago nemoralis Aiton var. longipetiolata (Mack. & Bush)

dell & Steyerm. [-,2,-,1] D, R; 8080-8625’; J, U; mm,

pp, Y

[So ee nemoralis var. decemflora (DC.) Fernald]

der s Kunth var. simplex [4,16,3,4] D, R; 7260-

U; af, mc, mm, mr, ms, pp, ra

sho speciosa Nutt. var. pallida Porter -- p] L; 7000’;

a dn velutina DC. ssp. Vinili (A. Gray) Semple [1,1,1,-]

7200-81607; J; mc, pp, r

Solidago wrightii A. Gray var. zdenophoras F. Blake [2,1,2,-] D,

dE 6025; 5 ii mr, r

arvensis [1,-,-,-] L; 56005 G: fr.

* Sonchus asper (L.) Hil [2,1,1,-] D, L 5825- 8900’; J, G; fr,

Ce + NE | Nore f

4] R; 6860-8075”; P, U; ds, pj.

7,31 R, 1,6450-6780';

C i

eovimi unt

Sephanomeri pauciflora (Torr.) A. Nelson [8,-,4,7] D, L, R, T;

00-8655’; J, G, P, U; ds, fr, pg, pj, ra

ynehendun ascendens (Lindl.) G.L. Nesom [1,21,-,8] D,

, U; mc, mm, mr, pp, ra.

Smphyorichum f ee (Lindl.) G.L. Nesom var. com-

atum r. & A. Gray) G.L. Nesom [3,-,1,-] D, L, R;

5090- 72000 i, G, P; ui mr, ra.

ll. ex DC. var. apricum (A. Gray)

m [-,-,-,1] R; 10230’; U; mm.

EN UE foliaceum (DC) G.L. Nesom var. canbyi (A.

Gray) G.L. Nesom [1,4,-,1] D, R; 8500-10020’; J, U; mm,

iu ra.

ff fps 1!

phy ex DC) G

(D.C. Eaton) G.L. Nesom [UNM-Castetter 7891] R; U

Symphyotrichum laeve (L) A. Love & D. Love var. geyeri (A.

Gray) G.L. Nesom [3,6,3,-] D, L, R; 6400-8625" J, U; br,

ms, pp, ra.

Symphyotrichum lanceolatum (Willd.) G.L. Nesom var. hespe-

rium (A. Gray) G.L. Nesom [-,2,1,1] D, L, R; 5400-8740” J,

U; mm, mr, ms, ra.

di sia Sell (Lindl.) G.L. Nesom var. spathu-

lat A] R; 8350-10230’; U; mm, mr.

* Tana Bilis at B 741] R, 7300’; U; ra.

ae a (Ledeb.) DC. [-,-,1,-] L; 7400; J; mr.

Andrz. ex Besser [8,5,-,2] D, R;

7360- 10230; J, U; mc, mm, mr, ms, DD, ra.

* Taraxacum officinale Weber ex FH. Wigg. [21,19,1,26] D, R;

6600-10900"; J, U; af, br, mc, mm, mr, ms, pp, ra, sf.

Tetradymia canescens DC. [-,-,2,6] D, R, T; 6100-9200"; J, P,

AIPA TALITY

Tetraneuris aca (Pursh) Greene var. acaulis [1,-,-,10] R, T;

6300-8 P, U; br, ds, fr, ms,

Tetraneuris agent (A. Gray) Ea [13-,10,4] D, L, R, S, T;

5300- ee ds

4] D, R, 1; 7060-9025” J, P,

; fr, ms, mr, po, pj, pp.

Thelesperma filifolium (Hook, A. iind [2,1,4,2] D, L, R, S, T;

900-8800"; J, G, U; pj, p

Thelesperma alan un pa Kuntze [20,-,6,10]

939

D, L, E 5, T; 5470-8100, J, G, P, U; ds, fr, mr, ms, pg, pj,

Strother [1,-,-,-] L; 5600-6600";

Los acerosa (DC.

G; ds.

Townsendia annua Beaman [-,-,-,11] R, T: 6240-7600". P, U;

cali eximia A. a HM D, L, R, S; 6140-8625’; J;

rose exscapa chron T. Porter [1,-,2,4] D, L, R;

500°; J, U; m

dA fendleri A. B Ta R, S, T; 6000-7237"; J, G,

TONSA incana SUE [-,7,7,5] R; 6700-8075’; P, U; ds, pj.

A Gray) Osterh. [2,-,-,1] D, 1; 6900-7700";

J, P; ds, pi, pp.

* Tragopogon dubius Scop. [32,24,3,21] D, L, R, S, T; 5300-

10900’; J, G, P, U; af, br, ds, fr, mc, mm, mr, ms, pg, pj,

Op, ra, sf.

* Tragopogon porrifolius L. [2,-,-,1] D, T; 7150-8400"; J, P, U;

* Tragopogon pratensis L. [5,-,-,1] D, R, T; 5760-9100’; J, G, P,

Verbesina encelioides (Cav.) Benth. & Hook. f. ex A. Gray [1,-

,2,10] D, R, 5, T; 5400-8655"; G, U; ds, fr, ml, pj, ra.

[Verbesina encelioides var. exauriculata B.L. Rob.]

Wyethia arizonica A. Gray [2,-,-,2] R; 7800-8750"; J, U; mm,

if | & R.L. Hartm. [2,-,2,3] D, L, R,

T: 5700- 8600: J, G, P, U; ds, fr, mm, pg, pj, pp, ra.

Xanthisma grindelioides (Nutt. Morgan & R.L. Hartm. var.

Xanthisma spinulosum (Pursh) Morgan & R.L. Hartm. var. spi-

nulosum [14,-,5,28] D, L, R, S, T; 5500-10200’; J, G, P, U;

ds, fr, mm, pg, pj, pp, ra

Xanthium strumarium L. vat. congaense dae Torr. & A. Gray

[2,- 12,31 D, L, Ñ 6000-7100 J,

Zi lutt. [-,-,1,-] D; 6 EIU

Berberidaceae

Berberis fendleri A. Gray [30,5,5,14] D, L, R, T; 6100-9025’; J, P,

; at, br, mc, mm, «DI pp, ra.

Berberis repens Lindl. [30,12,2,11] D, L, R; 6800-9625’; J, U; af,

br, mc, mm, mr, ms, pp.

etulaceae

Alnus incana (L.) Moench var. occidentalis (Dippel) C.L. Hitchc.

[24,9,2,12] D, L, R; 6875-10900"; J, U; ml, mm, mr, ra.

[Alnus incana ssp. tenuifolia (Nutt.) Breitung]

a caen eater Nat B 6200-7675"; J; ml, mr.

8460-9200’: J: ml, mr.

Betula occidentalis Hook. [3,4,5,1] D, L, R; 6700- 8075' J, U;

r,mm, mr, ra

e Ruin

YA UY ILI lect irr rinoranT13 yl ia; 18]

DL R S T. 5860- 8080* J, G, P, U; ads mia ms, pg, pj,

Cryptantha cinerea (Greene) Cronquist var. jamesii Cronquist

[9,-,5,-] D, L, R; 5800-8250’; J, G, P; br, fr, pj, pp, ra

LIMIT PE PLA

940

l. M. cus [1,-,1,9] D, L, R, T; 5460-9200’; G, P, U; ds, fr,

r, pj,

Saas ae (A. Gray) Greene [1,-,-,10] L, R, T; 6300%

G, P, U; PP, ra.

JG ms pj.

fC VÀ £o.

[2,-,-,-] D, R; 5860-7660;

- cens [-,-,-,6] R, T; 6050- 7050"; J, P, U; ds, fr r, pj.

ane minima Rydb. [2,-,-,3] R, S, T; 6320-7200; G, P,

s, fr, pj.

Hackelia besseyi (Rydb.) J.L. Gentry [-,6,1,-] D; 8100-9580"; J;

ml, mm, mr

Hackelia floribunda (Lehm.) I.M. Johnst. [9,6,4,2] D, L, R;

—9800’; J, Us mc, mm, mr, ra

+ @Hackelia hirsuta (Wooton & Stand.) 1.M. Johnst. [-,-,1,-]

Q5 Jz mc

Lappula occidentalis (S. Watson) Greene var. cupulata (A.

Gray) L.C. Higgins [3,-,1,2] D, R, S, T; 5860-71007; J, G, P,

U; ds, fr, pg, pj

Lappula occidentalis (S. Watson) Greene var. occidentalis

[31,20,4,48] D, L, R, S, T; 5550-10050’; J, G, P, U; af, br, fr,

mc, ml, mm, ms j, PP, ra.

Lithospermum cobrense Greene [UNM-Castetter 10732] D; J.

Lithospermum incisum Lehm. [7,-,3,17] D, L, R, T; 6100-8655;

J, G, P, U; mm, pj, pp, ra.

Lithospermum multiflorum Torr. ex A. Gray [35,19,6,8] D, L, R,T;

6300- 2280) M P, U; ale DUE mc, am mr, ms, pj, pp, Y

2] R; a2nn 1 adem

A A of

U; mr.

Mort (1 fi A Heller [28,8,4,5] D, L, R; 6400-10900"

J, U; af, mc, mm, mr, ra, sf.

Mertensia l lata (Pursh) DC. [30,15,3,13] D, L, R; 6300-

ie U; af, br, mc, mm, mr, ms, pj, PP, s

[Mertensia la néeolatà varieties]

agebat scouleri (Hook. & Arn.) LM. Johnst. var. hispi-

us (Greene) Dorn [-,2,2,5] D, R; 7550-9540’; J, U; ml,

mm, mr.

Brassicaceae

* Alyssum simplex Rudolphi [4,-,-,17] D, R, T; 6850-8500; J, P,

, mm, mr, ms, pj, pp, ra

Arabis hirsuta (L.) Scop. var. pycnocarpa (M. O Rollins

[4,3,1,-] D, R; 8060-9140"; J; mc, mm, m

bod vulgaris R. Br. [-,-,1,1] D, R; o HM J, U; mr,

DE 8200-8400" J; mm, ra.

son) À Love & D. Love [3,4,-,13]

D, R, T; 7150- 10600*: J, ben br, mc, mm, mr, ms, pp, ra.

pl plants are an unnamed hybrid 8. fendleri x B.

is Berteroa EARN (L) Be on

ta.]

mn fendleri (S. Watson) W.A. Weber S 5,3,12] D, L, R;

5600-191505; J, P, U; mc, mm, ms, pj, p

-o ur (A. Nelson) W.A. Weber 553] R; 6500-

socie 'palidifola (Rollins) W.A. Weber [-,-,-,7] R, T; 6050-

Boechera perennans (S. Watson) W.A. Weber [1,2,-,

000-10650"; J, U; mc, pp, ra.

-] D, R;

le /^Y

dis E a (Graham) Å. Lave & D. Love [-,-,-,2] R;

x j, pp.

oechera holboelli (Hornem.) A. Lóve & D. Lóvel

Boechera biis (Rydb.) Mii & "à Shehbaz [-,-,-,12]

0-9500; P, U; ds, m

Boechera Aaa (Graham) Al- EUM UM [14,49,1 i D, R;

—-10300; J, U; af, mc, mm, mr, ms, pp, ra, sf.

"mein atari, Andrz. ex DC. [3,-,-,4] D, a S; 5440-

G, U; ds, fr,

“cas burps (L) ea Pan 19,2 i D, R; 7310-

0” J, U; af, br, mc, mm, mr, ms,

Caran cordifolia A. Gray var. la I8, 5,1,3] D, R;

000’; J, U; af, mc, m

Ie cordifolia A. Gray var. incana A. Gray ex M.E. Jones

[1,-,-,-] R; 000’; J;

* Chorispora tenella (Pall.) DC. a =,1,-] D, L, R; 6000-9000’; J,

br, mr, ra

* Conringia orientalis (L) Dumort. [1,-,-,-] D; 73605 J; ra

Descurainia californica (A. Gray) O.E. Schulz [-,-,-,5] R; 8760-

9540’; U; af. :

Descurainia incana (Bernh. ex Fisch. & C.A. Mey.) Dorn ssp.

incisa (Engelm.) Kartesz & Gandhi [2,1,3,6] D, R, T;

5400-10700", J, U; br, fr, mm, mr,

[Descurainia incisa (Engelm. ex A. Gray) Britton ssp. inci-

(Bernh. ex Fisch. & C.A. Mey.) Dorn var. ma-

crosperma (O.E. Schulz) Dorn [1,13,1,1] D, R; 8625-11000;

m, mr, ra, sf.

[Descurainia incana]

Descurainia incana (Bernh. ex Fisch. & CA. Mey.) Dorn var.

viscosa (Rydb) Dorn [1,9,1,4] D, L, R, T; 6200-9980; J, P,

; af, br, fr, mc, mm, mr, pj in ra, TN

rr.

(Rydb) Rollins]

Descurainia pinnata (Walter) Britton var. filipes (A. Gray) Peck

D2] R; 77 700- 1010; U; br, m

ex A cai ssp filipes (A Gray)

| viscosa

Rollins

Mii ^ Dritt nerniririt

m (Cockerell)

Shlhinéts [6,-,2, 10] D, R, S, T; 5860-10100’; J, G, P, U; ds,

fr, mc, pg, pj, ra.

[Descurainia pinnata ssp. halictorum (Cockerell) Detling]

! Descurainia pinnata (Walter) Britton var. paysonii Detling

Er ,", 1] R; 6050-9200* U; ds, mm.

inia incisa (E Im. ex A.G

` Drt

Davsonii

(Detling) Rollins]

* Descurainia sophia (L.) Webb ex Prantl [16,9,2,17] D, L,R, T;

er 9500’; J, G, P, U; ; br, fr me, mm, mr, pg p in ra.

5470- 7220’ J, G, P, U; ds, fr, pa, pj, ra.

Draba aurea Vahl ex Hornem. [1,17,-,-] D, R; 8060-10300' J,

: mc, mm, mr, ra, sf.

Draba helleriana Greene var. helleriana [9,33,4,2] D, L, R;

a us iJ U; br, mc, mm, mr, pp, s

(Hell JOE Schulz [5,-,=,-]

D, R: 6800- 9625" qa

Draba rectifructa C.L. T [1,-,-,3] R; 8840-10150; J, U;

Reif et al ' Fl ftheT {| ins. New Meyico

Draba reptans (Lam.) Fernald [-,-,1,2] R; 6250-7740"; J, U; pj.

Draba Pd A. Gray [-,-,-,5] R; 7050-9200"; U; mc,

mi, ms, pj.

eps TM (Douglas ex Hook) Greene var. capita-

tum 3,2] D, L, R, T; 6400-10225” J, P, U; br, fr, mm,

p , fa.

Erysimum capitatum (Douglas ex Hook.) Greene var. elatum

Nuttall) Torr. [18,11,-,30] D, L, R, S, T; 5860-11150"; J, G,

P, U; br, ds, fr, mc, mm, mr, ms, pg, pj, pp, ra, sf.

* Erysimum cheiranthoides L. [UNM-Yarnell 34] R; U.

Erysimum inconspicuum (S. Watson) MacMillan [-,-,-,3] R, T;

/300-8560' j

—,

; 5, Dj.

* EUST TPAR. x [1,-,-,-] D; 7360" J; ra.

p ia (A Gray) Rydb. [21,2,3,24] D, | RE

5860- 8940’; J, G, P, U; br, fr, mc, mm, mr, ms, pj, pp, ra.

Lepidium alyssoides A. Gray var. ayssodes [3,-,-,11] D, R, T;

6200-8400", J, P, U; ds, fr, pg, pj, r

Lepidium o ^ pa Var. Beads (Wooton) Rollins

3,-,^,-] R; 650 55 J; ft, pj, ra.

» on A ERU P Shehbaz [-,-,-,1] R; 8000-8400’;

id "isum chalapense L [2 sri] D, R, 6000- a : ur he

_ 5550- ony J, G, U; ft pur ms, pg, pj, DD, ra ra.

GA Mulligan

941

[10,-,1,12] D, L, R, S, T; 5600-8500’; J, G, P, U; ds, fr, ms,

Pg, pj, p

Rorippa curvipes Greene var. curvipes [1,-,-,4] D, R; 8300-

10230" J, U; mr

Rorippa curvipes Greene var. truncata (Jeps.) Rollins [-,-,1,-]

D; 5400-5900’, J; fr.

idi cae (L.) Besser var.

key [2,-,2,-] D, R; 5360-8980"; J; fr, m

s ob .) Besser var. hispida eo jane [-,-,1,-]

£, Lr.

E & Abbe)

60 J;

“fr

Rorippa sinuata (Nutt) Hitchc. [-,-,1,-] D; 5360"; J; fr.

Rorippa ee ee (A. Slips Britton [1,16,-,-] D, R; 8450-

,U; mm, mr, m

* Sisymbrium altissimum ^ m, 7 a 10] D, L R, S, T; 5470-8500;

G, P, U; ds, fr, ml,

* Sisymbrium loeselii L. El. A D, R; 8000-8400’; J, P; mm, ra.

Stanleya pinnata (Pursh) Britton var. pinnata [-,-,-,1] R; 8250";

U; pj.

Streptanthella longirostris (S. Watson) Rydb. [1,-

00-6500’; J, U; fr, ra

Streptanthus cordatus Nutt. var. cordatus [1,

6400-8075’; J, P, U;

Thelypodiopsis purpusii decos Rollins [1,-,7, -] S;

d 9500.9; G; f p.

"r1 I a4. 4 1]

71,3] R;

-,1,2] D, R;

ex Walp sar nrarilinoe

prd r

11] R; 9300-9500"; Un mc.

ie are Schrad. var. ramosum (A. Nelson) Thell.

R, S; 6140-9625” J; br, mm, mr, pp, ra

* Lepidium pen L. [-,-,3,-] D, L; 6000-7600’; J; ra.

"eLepidium latifolium L. [2,- -1 0 D, L, n 5400-5600" G; fr ra.

Nlale

Rollins [-,20,-,5] D, R, T; 7950-9850" J, P. U; af, mE Bm.

r, ms, pj, pp, ra.

Eie ramosissimum A. Nelson var. ramosissimum [-,-,-,1]

R; 9250", U; m

ou virginicum L. var. medium (Greene) C.L. Hitchc. [2,-

2,5] D, R, ST (20007 2900, G, U; br, fr, pj, ra

) Thell. [4,21,-,4]

DD R,T; 7550-10230’ J, P, U; mc, ml, mm, mr, pp, ra

* Nasturtium officinale R. Br. [9,2,1,-] D, L, R; 5360- 8625’ J, G,

r, ra.

Noccaea fendleri (A. Gray) Holub ssp. glauca (A. Nelson) Al-

Shebaz & M. Koch [18,11,2,13] D, L, R; 6700-10600’; J,

U; af, mG mm, mr, pp, ra, sf.

th.) Rollins [2,10,-,-] D, L; 7120-10300’;

J; me ms, s PP, ri ra.

A. Gray) Nieuw. [-,1,-,1] D, R; 8300-9000";

J, U; af, p

Praia enden (A. Gray) O'Kane & Al-Shehbaz [2,-,1,6] D, R;

05 J, G, U; ds, fr, pj.

s Penes Rydb. var. floribunda IUNM-Atwood

221] R; U.

Physaria intermedia (S. Watson) A. Helier [UNM-Tonne s.n.]

Phy : (A. Gray) G [1,-,-,10] R, T; 6400-8655";

J, P, U; fr, ms. "m

iyvsana rectipes ( m & Standl) OKane & Al-Shehbaz

Rob. EM -] D; 9500- 9660": J; m

Telpodun wrightii A. Gray subsp. dim [3,-,1,-] D, L, R;

205 J, G, U; pj, pp, ra.

NO arvense L. [-,-,-,1] R; 7540-7740"; U; ms.

Turritis glabra L. [8,8,5,-] D, L, R; 6400-9450 J, U; af, br, mm,

mr, ms, pp, ra.

Cactaceae

Coryphantha vivipara (Nutt) Britton € Rose [1,-,1,9] D, R, T;

3600- diei J, P, U; fr, me, pp.

Ll ] L ]

ta[7,-,1,15]

DR S,T; 5900- “£000 3, G, P, U; fr, ms pj aa

J, F 5, pj, p JJ,

Echinocereus ender aa F. Seitz [3,-,1-] D, L, S; 5440-

r, pj.

conoce da Engelm. [4,-,1,16] D, R, T;

800-9450"; J, G, P, U; ds, fr, pj, pp.

m ania Engelm. [1,-,1,12] L, R; 6500-9000

,'r F C, Pg, pj, pp.

+ Grusonia a (Engelm.) H. Rob. [1,-,-,-] S; 6320"; G; ds.

Opuntia fragilis Nutt. [UNM-Reenea 2146] R;

Opuntia macrorhiza Engelm. [RM-Nelson 228] D; J.

Opuntia padeacanind Engelm. [10,1,2,13] D, L, R, T; 5600-

867 1G, P, U; ds, fr, mc, pj, PP, ra.

B. 7 Parfitt [1, al rd D, R; 6000*: de pj.

O} tha [8,-,1,28] D, L, R,

E T 5440- 8700; A G, P, U; ds, fr, ms, PS, pj, pp

MI Castetter

ALLET

& Bigelow)

1002 A-J] R; U.

Sclerocactus papyracanthus (Engelm.) N.P. Taylor [-,-,1,-] L;

6300" J; pj.

942

Campanulaceae

Campanula parryi A. Gray var. parryi [5,14,2,5] D, R; 8000-

102305 J, U; mm, mr.

Mais rotundifolia L. [27,40,7 a D, L, R; 5850-11150’;

U; af, c, mm, mr, ms, pp,

label anal L. [1,-,2,-] D, L; c Pm J; ml, mr.

Cannabaceae

Celtis reticulata Torr. [3,-,3,-] D, L, S; 5400-6000’; J, G; fr, mr, pj.

mulus lupulus L. var. neomexicanus A. Nelson & Cockerell

[7,3,2,-] D, R; 5550-8750" J; mc, mm, mr, ms, ra

Caprifoliaceae (includes Valerianaceae)

Linnaea borealis L. var. longiflora Torr. [3,3,1,-] D, L, R; 8475-

yD |

g. [19,5,1,2]

D, L, R; 7500- 10260" J, U; af, mc, mm, mr,

parc rotundifolius A. Gray [18,3, p ds Dp ER,

10900’; J, P, U; af, mc, mm, mr, ms

mi acutiloba Rydb. var. ~ [11, 21 E DLR

8000-9300"; J, P, U; af, mc, m

Valeriana arizonica A. Gray "A 5 T 8600; J; af, br,

mr, pp.

aa edulis Nutt. ex Torr. & A. Gray [17,3,2,-] D, R; 7600—

; mc, mm, mr, ms,

Caryophyllaceae

fR At -I 1AM-_l | eee

saxosa (A Gray) /aruc-

chi, R.I qn 2 Rabeler[10, 36,3,6] D, L R: 7850-111505

J, U; al, br, mc, Pam mr, ms, pp. S

ssp. saxosum (A. Gray)

w. A. Weber]

rm J: +L h [14,20,5,3] D

L, R; 6850-10600’; J, U; af, br, mc, mm, mr, ms, r

Cerastium brachypodum (Engelm. ex A. Gray) B.L in M

D; 8570-9725"; J

TLIRIAA Cha LL: co Faa E Dp. E |

L Fiy T,

A 77

*( pj f l B ever l

& Burdet [3,17,1,2] D, R; 6100-10230; J, G, U; mc, ml,

SUE

m 5.

* Dianthus armeria L. subsp. armeria [-,2,-,-] D; 8075-8625;

Du ke [-,-,-,2]

J; pp, ra.

Drymaria effusa A. Gray var. depressa (Greene

R; 8200-8600"; U; pp.

ia nlandil Rartli ,1] R; 6990";

alandi anca T

pi | E. ORE AS ak ee UE

U; pj.

H I PS 1) qi (Cham 0o 0 LL Hip r rs L | {

; PP.

2,1] D, R;6500-8670’:

leptophylla Er R; el -8350; U

J, U; br, pj, rz ra.

Eremogone eastwoodiae (Rydb.) Ikonn. var. adenophora

(Kearney & Peebles) R.L. Hartm. & Rabeler [-,-,-,2] R;

6200-6500” U; fr.

eee eastwoodiae iiw: 2 var. eastwoodiae [1,-,

,3] R; 6050-6500* P, U; ds, f

Ceregore pe (A. Gray) om, E 38,1,9] D, R; 7860-

50 , mm, mr, pp

ve enio (L.) Fenzl e i E R; 92505 U; m

Moe d macrophylla (Hook) Fenzl [1,-,2,-] D, , 6300-

le F da |

f Texas 3(2)

|| fal Dos Li Im LJ Po ow i

Pseudostellaria jamesiana (Torr) W.A. Weber € R.L. Hartm.

3,6,-,5] D, R; 7300-10150’; J, U; af, mc, mm, mr, ms,

H. Karst. [2,-,-,1] D, R; 8460-9720 J,

pp, ra, sf.

Sagina saginoides (L.

; mc, mm, mr.

* Saponaria officinalis L. [-,-,1,-] D; 60005 J; mr.

Silene antirrhina L. [1,-,4,-] D; 5700-6020 J, G; fr.

Silene ia Hook. var. aa [5, bi d 1] D, E

0900"; J, U; af, mc , Pp,

! Silene ae Hook. var. esida (Rydb) m 2 7,4] R;

9300-10940; U; mm, mr.

Silene menziesii Hook. [6,1,-,2] D, R; 7950-9000* J, U; mc,

mr, ms, pp.

Silene scouleri Hook. var. pringlei (S. Watson) C.L. Hitchc. &

Maguire ex Kartesz & Gandhi [1,20,2,2] D, R; 7025-10325';

, U; af, mc, mm, mr, pp, ra.

Er id rubra (L) J. Presl & C. Presl [1,-,-,-] R; 9200-9400";

ED salina J. Pres! €: C. Pres! [-,-,1,-] D; 53605 J; mr.

Stellaria MIS (Ledeb) Bong. [-,1,-,-] D; 9680-10340;

Jim

[St ele calycantha var. simcoei (Howell) Fernald]

Stellaria iip Muhl. ex Willd. [3,3,-,1] D, R; 7740-9100;

, U; mm, mr, pp

stellar pace Goldie var. longipes [3,9,-,5] D, R; 8525-

, U; af, mc, mm, mr, s

à en media (L) Villars [-,-,1,-] L; 7100; J; m

Stellaria umbellata Turcz. ex Karel. € Kir. [-,1,-,

J; mr.

-] R; E 0300’;

Celastrac

Paxistima e (Pursh) Raf. [31,33,3,14] D, L, R, T; 6200-

10700"; J, P, U; af, br, mc, mm, mr, ms, pp, s

Chenopodiaceae

Atriplex argentea Nutt. var. argentea [UNM-Castetter 3929]

Fanacrona [1 A

mm L3 A) ir

1,-,5,24] D, L,

R, S, T; 5470- 8560" J, G, P, U; ds, fr, mr, ms, pg, pj, ra.

Atriblex confertifolia (Torr. € Frem.) S. Watson [2,-,-,-] D, R;

7000-7100’; J; ds, pj.

"5 [-,-,-,4] D, R, T; 8440-10700’;

k Ff c FF

, U; ds, mr, mm.

Chenopodium atrovirens Rydb. [-,9,-,5] D, R; 6975-10100"; J,

, mr, ms, pp, ra.

Chenopodium berlandieri Moq. var. zschackei (Murr) Murr ex

Asc 1,-] D, R; 5400-9625’; J; mc, mm, ms, pp, ra, sf.

j Pde nl capitatum (L.) Asch. [-,1,-,-] D; 9500- o

; mc

EE A desiccatum A. Nelson [-,-,-,3] D, T; 8440-84507;

* ml.

Chenopodium fremontii S. Watson [3,17,5,12] D, L, R, T;

6000-9800’; J, P, U; mc, mm, mr, pp, r

* Chenopodium glaucum L. var. Mum [-,-,1,-] T; 5400;

Lir

Chenopodium hians Standl. [-,-,-,2] R, T; 6600-86557; P, U; pj.

Chenopodium incanum (S.Watson) A. Heller var. incanum [3,-

,2,3] D, L, S, T; 5550-8667’; J, G; P; ds, fr, mm, ms, pj.

Reif et al.. El ftheT I] ins. New Mexico

CH | a

D, R: 6480- 8400" f U; ms, p,

Chenopodium neomexicanum Stand! [UNM-Flecks.n.] R; J.

* Chenopodium overi Aellen [8,15,2,2] D, R; 7450-9800"; J, U;

af, br, mc, ml, mm, mr, ms, ra, sf.

X [Chenopodium capitatum var. pum oT Masi

tt ex S Watson [-,2,1,3]

o Js Ja Jal” il! al ES 7g

y 6, R.U f, mm t, po, ra.

" rr) Wahl]

* Chenopodium rubrum L. var. rubrum [-,-,1,-] D; 5400"; J; fr.

ir rd watsonii A. Nelson [-,-,-,2] R; 6550-8150"; P,

* Doni ambrosioides (L) Mosyakin & Clemants [-,-,1,-]

“Jfr.

Tf Dhan botrys (L.) Mosyakin €: Clemants [-,-,1,-] D;

opi

I3 & Clemants [ ,6,6,13]

DR ST 6500- 9340": J, P. U; br, ds, fr, mc, ms, pg, pj, PP,

“ocha ea - Schrad. [1,1,1,5] D, R, T; 5200-8900", J,

U; fr, pg, p

KARA

& Smit [2,-,1,1 1] D,

R, T; 6100-8320" J, P, U; rs BOUE

io nuttalliana dpi te [1,8,-,7] D, R, T;

900-9600’; J, P, U; mm

n tragus L. [3,-,3,16] D à i 5760- 6150’: J, G, P. U; ds,

fr, pg, pj, ra.

E Edi vermiculatus (Hook.) Torr. [2,-,-,-] D, R: 6000-7000";

J; ds, fr.

nec suckleyana (Torr.) Rydb. [1,-,-,6] D, R, T; 7050-9020’;

J, B. U; ml.

Cleomaceae [Capparaceae]

Cleome serrulata Pursh [3,-,7,-] D, L, R; 5300-8370"; J, G;

inan a

1 tra (L) DC tract (T

& A. Gray)

H.H. Iltis [3,- TUD: L; 5300- 6140’: J, G; fr, pj, ra .

Wislizenia refracta Engelm. var. refracta [UNM-Talley 25] D; J.

Commelinaceae

lina dianthifolia Delile [-,5,3,2] D, R; 6200-9200’: J, U;

I pj, pp

iid occidentals (Britton) Smyth var. . occiden talis [3,-

77] R, T; 5800-72807 J, P, U; ds, fr, p

m d Mv rd En Smyth v bu. ESCONDE

Innean

Convallariaceae [traditionally in Liliaceae]

Maianthemum mosum (L.) Link var. amplexicaule (Nutt.)

Dorn [25,5,2,4] D, R; 6800-10900": J, U; af, br, mc, mm,

mr, ms, pp, sf

AA » HI "PEN E T

(L) Link [25,3,1,5] D,R:6880-10150':

J, U; af, mc, mm, mr, ms, pp, sf.

Convolvulaceae

EC age (L.) R. Br. Ae i rm (Brummitt) N.H.

n [-,-,2,-] L; 5400-5500"; J; m

* Pulls arvensis L. [20,-,-,4] D, L, R, úl 5550-8360’; J, G,

P, U; fr, mm, mr, pg, pj, pp, ra.

943

Cuscuta aplanata Engelm. [-,-,3,-] D, L; 5400-6500, J; ra.

Cuscuta umbellata Kunth [-,-,-,51 R, T; 6750-7185% P, U; ds,

fr, p

Beh Medie Roemer & Schult. [2,-,1,3] D, R, T:

720’; J, P, U; fr, p

n Ene H. Hall [- se 4] D, R; 6000-8350’; J, U; fr,

pp.

* Ipomoea purpurea (L.) Roth [UNM-Dixon A-348] D; U.

Cornaceae

+ Cornus canadensis L. [4,-,-,-] D; 83300-8475"; J: mm, mr

Cornus sericea L. var. sericea [24,4,1,2] D, L, R; 6800-8850" J,

br, mc, mm, mr, ms

Crassulaceae

Sedum cockerellii Britton [-,2,1,-] D, L; 7200-9050” J; ms, pj,

pp.

Sedum lanceolatum Torr. ssp. lanceolatum [-,-,-,1] R; 8600’;

Aa gale

Sedum rhodanthum A. Gray [-,-,-,2] R; 9980-11000; U; mr.

Cucurbitaceae

Cucurbita foetidissima Kunth [2,-,1,-] D, R; 5710-6500’: J; fr,

ml, ra.

Cyperaceae

Carex aquatilis Wahlenb. var. aquatilis [6,6,-,9] D, R; 8350-

11000"; J, U; mc, mm, mr.

a ‘Spreng. rl i di J; ml.

; 8350-9200" U; mr.

a aurea Nutt. 2, 2,- a] D, R: - 8000- 10230"; J, U; mc, mm,

C mu L.H. Bailey) Olney ex Fernald [UNM- Worthington

32790] R; U.

Carex bella L.H. Bailey [1,6,-,1] D, R; 8900-111505 J, U; mm,

bolanderi Olney [-,-,1,-] D; 6800"; J; mr.

Carex brevior (Dewey) Mack. ex. Lunell [-,-,1,-] D; 5600’; J; fr.

Carex brunnescens (Pers.) Poir. var. brunnescens [-,1+C,-,-1 D;

8660"; J; ml.

Carex canescens L. var. canescens [4,5,-,4] D, L, R; 7500-109005

Carex conoidea Willd. [-,C,-,-] D; 8750; J; mm.

! Carex nie Schwein. var. deweyana [1,C,-,1] L, R;

700 l

ey [3,2,-,-] D, L, R; 7740-10000"; J; mr.

Carex douolasi n 8,2,5] D, L, R; 7050-9250; J, U; mm,

mr, pp, ra.

la C.A. Mey. [1,1,-,7] D, R, T; 6850-10330’; J, P,

Carex bene Rydb. m j jn "s 10230; ba mm.

i 1] D, R: 9300 —10020’:

Carex aa Dewey [2,-,-,-] D, L; 5600-7675’; J, G; fr, ml.

Carex filifolia Nutt. var. filifolia [1,-,-,4] D, R; 6300-8500’; J, U;

MS, pj, pp.

Carex geophila Mack. [3,10,1 a, D, R, T; 6850-11000"; J, P, U;

, mm, mr, ms, pj,

Carex geyeri Boott [1,-, dd B R ore 10900"; J, U; mc, mm.

944

Carex Allee Muhl. ex Willd. [1,-,2,-] D, L, R; 5400-7000;

J; fr, m

a LH. Bailey [-,-,-,1] Ri Hed nen U; mr.

Carex inops j Crins [5,4,-,16]

D, R, T; 7130- 9450" J, P, U; br, mc, mm, mr, pj, pp.

Carex interior L.H. Bailey [1,-,-,-] D; 8000-8100’; J; mm.

ah del LH. Paley b F de - E 9800’; U; mr.

M 9800’; U; mr.

n microptera Mack. [25, 41, 2 pe D, L, R; 6600-11150; J,

; af, br, mc, mm, mr, ms, sf.

a o Dewey [3,-,-,3] D, R; 7330-101505; J, G,

H Bailey s S| Ibsp

ial obtusata Lil d peri G5] D; J

Bailey [39,41,3,12] D, L, R; 5860- 10900"

J, G, U; br, mc, mm, mr, ms, pj, pp, ra

Carex pellita Muhl. ex Willd. [8,4,3,5] D, E is 7000-9800"; J,

; ml, mm, mr, ra.

Carex petasata Dewey [-,1,-,-] D; 8810-89257; J; mm

Carex pityophila Mack. [2,6,-,5] D, R; 7100-11000; n U; af,

[Cures eophila M ck]

Mii id is de [1,-,1,4] D, R; 8000-10600'; J, U; mc,

EE Rydb. [-,-,-,1] R; 10175-10600; U; mm

! Carex rosea Schkuhr ex Willd. [2,-,-,-] D; 7025- 7750’ J;

mm, pp.

Carex rossii Boott [5,8,-,1] D, R; 8300-10675; J, U; mc, mm,

mr, pp, sf.

Carex scoparia Schkuhr ex Willd. var. scoparia [1,-,-,-] D;

—7940" J; mr.

Carex siccata Dewey [3,10,-,11] D, R; 8075-11150% J, U; mc,

m, mr, ra, sf.

[Carex foenea Willd. var. foenea]

Carex simulata Mack. [1,7,-,-] D, R; 8350-9180" J, U; mc, ml,

mm, mr.

Carex sprengelii Dewey ex Spreng. [2,-,-,-] D; 7275-8475;

J; mr.

Carex stevenii (Holm) Kalela [1,1,-,-] D, R; 8700-9625"; J; mr.

Carex stipata Muhl. ex Willd. var. stipata [10,7,3,-] D, R;

6600-9150’; J, U; af, mc, ml, mm, mr.

Carex ie Boott [4,-,-,2] D, L, R; 6850-10230’; J, U; mc,

! Carex tahoensis Smiley [UNM-Gierisch 3146] R; U.

Carex utriculata Boott [7,13,-,8] D, L, R; 7920-10900"; J, U;

mc, mm, mr.

Carex vallicola Dewey [-,-,-,5] R, T; 7200-10900’; P, U; ds,

, MI, pp.

Carex vupinoide Michx. [2,-,-,-] D, L; 5550-7490", J, G; mm,

Exe — Mack. [1,5,-,-] D; 8380-9300"; J; mc, mm, mr.

Cyperus bipartitus Torr. [-,-,1,-] D; 5360 J; fr.

[C. rivularis Kunth]

Cyperus esculentus L. var. leptostachyus Boeck. [-,-,

5360; J; ft

2,-] D;

Cyperus fendlerianus Boeck. [-,12,2,4] D, L, R; 6650-9200; J,

, t

Cyperus schweinitzii Torr. [3,-,-,1] R; 5900-6975’; J, P, U; fr,

pg.

[| 1 fall Dos a Inm Lj a es rn

f Texas 3(2)

Cyperus squarrosus L. [-,-,1,1] D, R; 5360-8350; J, U; fr, pp.

Eleocharis acicularis (L) Roem. & Schult. [1,1,-,3] D, R; 8460-

50’; J, U; ml, mm, mr.

Eleocharis ect (L) Roem. & Schult. [5,13,1,10] D, L, R;

0’; J, G, U; fr, mc, ml, mm, mr, pp, ra.

Eleochris enirn Britton [UNM-Worthington 32762] R; U.

Eleocharis onquera UA Hartmann) O. Schwarz [-,1,-,1] D,

Eleocharis rostellata oni Tor [-,-,-,2] R; 6500-7100” U; fr,

mi.

Schoenoplectus acutus (Muhl. ex Bigelow) A. Love & D. Lóve

var. occidentalis (S. Watson) S.G. Sm. [1,-,-,-] R; 6010-7000

ml.

C t I + I} 4 t Ls

de j “2] " 6500- 2100, U; fr, ml

& R. Keller

{Otte M

S.G. Sm. Duel D, L, R: 5600- J, G; fr, ml, mr.

[Schoenoplectus pungens var. pungens]

"o tabernaemontani ce earn Palla [-,-,-,1]

00’; U; fr.

sims Dno Presl €: C. Pres! [8,2,1,3] D, R; 7080-

0020"; J, U; mm, mr.

Scirpus pallidus Britton) Fernald [3,-,-,-] D; 6400-7330", J;

Elaeagnaceae

*eFlaeagnus angustifolia L. [9,-,1,4] D, L, R, S; 5470-7300 J,

U; fr, m

Shepherdia argentea (Pursh) Nutt. [2,-,-,-] R; 6000-7480; J;

fr, ra.

Shepherdia canadensis (L) Nutt. [6,1,-,1] D, L, R; 8340-9180;

J, U; mc, mm, mr, pp.

Elatinaceae

Elatine rubella Rydb. [-,-,1,-] D; 89005 J; mr.

Eric

Arctostaphyos uva-ursi (L.) Spreng. [17,17,2,8] D, L, R; 7350-

1022 , mc, mm, mr, ms, PP, ra, a

Chi hil bellata (L) WPC. Bart talis (Rydb.)

SF Blake [3,-,-,-] L; 7000-8000"; J; mc,

[Chimaphila umbellata var. acuta id) a F. Blake]

Moneses uniflora (L) A. Gray [1,2,-,-] D, R; 9050-10000"; J;

c, mr, sf.

ipei hypopithys L. [-,6,3,-] D, R; 6350-10230' J; af,

c, mm, mr.

RUE secunda (L) House [4,6,1,-] D, R; 6950-10600’; J; af,

F F mr, S

Pterospora andromedea Nutt. [1,4,3,1] D, R; 6300-9800" J,

Pyrola asarifolia Michx. var. asarifolia [-,-,1,-] L; 7450; J; mc.

Pyrola chlorantha Sw. [2,3,4,-] D, L, R; 7450-9650"; J; mc,

Dp, ST.

Pyrola minor L. [2,-,-,-] D, L; 8000-8650"; J; mc, mr

Pyrola picta Sm. [2,-,1,-] D, L; 8020-9200” J; af, m

Vaccinium cespitosum Michx. [1,---,1] R; e J, U;

mc.

Vaccinium myrtillus L. var. oreophilum (Rydb.) Dorn [8,12,1,-]

LR; 88340-11150"; J; af, mc, mm, mr, sf.

Reif et al Fl f tha T 1] taine New Mexico

Vaccinium scoparium Leiberg ex Coville [3,-,-,-] D, L, R;

5% J; mc, mm, pp.

uphorbiaceae

Chamaesyce fendleri (Torr. & A. Gray) Small var. chaetocalyx

(Boiss) Shinners [14,-,2,8] D, L, R, S, T; 5600-7660"; J, G,

' U; br, ds, fr, ms, pj, pp, ra.

Chamaesyce fendleri (Torr. & A. Gray) Small var. fendleri [5,-

,3,17] D, L, R, T; 5300-8500’; J, G, P, U; fr, ms, pj, pp, ra.

Chamaesyce geyeri (Engelm.) Small [1,-,-,-] L; 5500-5600",

G; fr.

Chamaesyce glyptosperma (Engelm.) Small [-,-,1,6] D, R, T;

5300- 7620.4; P, U; ds, fr, pj! ra.

Il [-,-,1,-] L, 6500"; J; pj.

Chamaesyce serpyllifolia. (Pers, Small i4, 12,5,17] D, R, T;

5300-9200’: J, P, U; ds, mc, mm, mr, ms, pg, pp, ra, sf.

Chamaesyce serrula (Engelm.) Wooton € Standl. [1,-,-,-1 D;

5710; Js ra.

Croton texensis (Klotzsch) Muell. Arg. ~ = 1] L, D, R, S;

5300-6300"; J, G, P, U; ds, fr, pg, pj, p

gis. — Engelm. [3,2,2,-] S E 6450-8250; J;

ee a a bils [1,-,2,-]D, L; 6000-74007; J; ra.

Tragia nepetifolia Cav. [2,-,2,-] D, L; 5600-6600’; G; fr, pj.

Fabaceae

Amorpha canescens Pursh [-,-,3,-] D, L; 6500-6600"; J; pj.

Amorpha fruticosa L. [3,-,1,-] D, L, S; 5300-5800’; J, G; ds, fr.

Amorpha nana Nutt. [1,-,-,-] L; P : mr.

Ast | tis D | -,5] R, T; 8200-8700;

P, U; ds, mm, ms, pp.

Astragalus albulus Wooton & Standl. [1,-,-,2] R; 6010-70007

, U; fr, pj.

Astragalus allochrous A. Gray var. dne (M.E. Jones) Isley

[-,-,-,3] R, T; 6900-8600"; P; ds, m

on amphioxys A. Gray var. Bir [1,-,-,1] D, R;

0-6880'; G, U; ms,

oh bisulcatus (Hook) A. Gray var. haydenianus (A.

Gray) Barneby [6,-,1,2] D, R; 6900-8260’; J, U; ds, fr, ms,

Pg, pp, ra.

Astragalus ceramicus E. Sheld. var. ceramicus [-,-,-,31 R, T;

6450-7150’; P, U; fr, pj.

! Astragalus cerussatus E. Sheld. [-,-,-,3] R, T; 8200-8328; P;

+ Astragalus cyaneus A. Gray [-,-,-,4] R; 6700-7600"; U; pj.

Astragalus drummondii Douglas ex Hook. [-,-,-,10] R, T;

50880" ind P, U; idi mm, ms, pj, Pp! ra.

(Hook.) Dou G. Don var. aid

Tt , “Al R, T; EUR 9450; ih P, U; mm, n. p, op, f

Don var. e (A.

Gra ay) Barneby [1,- 42, 1] D, R, S 5700-6600’; J, G, U; fr, pj.

Astragalus gilensis Greene [2,3,-,-] D; 8025-8375"; J; br, mc,

ms, pp.

Astragalus hallii A. Gray var. hallii [-,-,-, 1] R; 8250; U; mm.

Astragalus humistratus A. Gray var. humistratus [3,-,-,-] R;

7310-8120"; J; ds, mr, pp.

945

* Astragalus iodopetalus (Rydb.) Barneby [1,-,-,-] R; 7400-

8600"; J; pp.

+ dei kentrophyta A. Gray var. neomexicanus (Barneby)

neby [1,-,-,-] R; 5900-7100’; P; pj.

mess laxmannii Jacq. var. robustior (Hook.) S.L. Welsh &

Astragalus lentiginosus Douglas ex Hook. var. albiflorus (A.

Gray) Schoener [1,-,2,3] D, R, S, T; 5700-8350'; G, J, P,

Astragalus lonchocarpus Torr. [10,1,1,5] D, R; 5825-9025’; J,

; Os, fr, ms, pj, PP, ra.

Astragalus lotiflorus Hook. [-,-,-,1] R; 6400-6500'; U; fr.

+ Astragalus micromerius Barneby [-,-,-,1] R; 6550-7100;

; pj.

Astragalus missouriensis Nutt. var. missouriensis [2,-,2,6] D, 5,

R, T; 5600-8260; G, J, P, U; ds, fr, pj, pp.

+ d al Torr. var. matthewsii (S. Watson)

Ba -,-] D; 7640-76805 J; ms.

le aca us bus A. Gray [5,-,-,7] D, R; 6900-8350;

; ms, pj, PP, ra.

Astragalus newberryi A. Gray var. newberryi [-,-,-,2] R; 6200-

005 U; pj.

Astragalus nuttallianus A. DC. var. . austrinus (Small) Barneby

[2 : 5700-6020’; G; fr,

ere nuttallianus DC. var. _ micranthiformis Barneby [-,-

4,1] R; 5400-6300" J; U; p

Astragalus pattersonii A. Gray M S; 5470"; G fr.

As eel sea us E. one var. praelongus [2,-,-,2] D, R;

250" J, U; fr, m

mM proximus (Rydb) Wooton & Standl. [1,-,-,-] R;

7400-8600’; J; pp.

+ | ¡A oe 1 n l

* A

[UNM-Spellenberg et al. 1231] R; P.

I o 1 r

BF FF Fos

€ A tit

Astragalus Copla Porter [2,-,-,-] D; 5600-7150’; J; ds, fr

uev humilis Benth. var. humilis [1,-,-,-] D; 6950-7100";

d arboles cens L. [UNM-Hutchins 8472] D; J.

* * Coronilla y varia ls d -] D; 6000"; J; ra.

P Willd. var. oligophylla (Torr.) Shinners

1877,10, L, R,T. 5300- 7400’; J, G, BU; fr, ms, pj, PP, ra.

y E,-,3,-1D; 5450-5700’; J; fr.

Dalea filiformis A. Gray [2,-,-,-] D; 7600-8710’; J; br, mc.

Dalea formosa Torr. [4,-,3,-] D, R, S; 5600-6400" J, G; fr, ms, pj.

Dalea jamesii (Torr) Torr. & A. Gray [4,-,-,2] R, 5, T; 5600-6450"

J, G, P, U; ds, fr, pj

Dalea lanata Spreng. var. terminalis (M.E. Jones) Barneby [2,-

2,1] D, R; 5700-6300’; J, G, U; ds, fr, ra.

Dalea leporina (Aiton) Bullock [-,-,3,-] D, L; 6000-7000; J; ra.

Dalea nana Torr. & A. Gray var. nana [-,-,2,-] D; 5300-6350;

J; fr, ra.

Dalea polygonoides A. Gray [-,-,1,-] D; 6500” J

ae purpurea Vent. var. purpurea [1,-,2,-] “i cane 71007

pj, pp.

a Pa triacanthos L. [1,-,1,-] D, R; 5800’; P; fr, ra

Glycyrrhiza lepidota Nutt. ex Pursh [8,-,3,-] D, L, R, 5; 5300-

7310" J, G, P; ds, fr, mr, ms, ra.

946

h : le Nutt boreale [1,-,-,2] D, R;6100-7100’;

J, U; fr, pj, pp.

2H St John [4,-,-,-] R; 6875-8100

; fr, pp, ra.

* Lathyrus latifolius L. [1,-,1,-] D; 5745-5900", J; fr, ra.

Lathyrus leucanthus Rydb. d 14,2,17] D, L, R; 6500-10675’;

J, U; af, mc, mm, mr, ms, pp, sf.

HUE corniculatus L. DET E 7400- 8600’; J; ra.

by [4,-,-,-] D, R; 7500-8750;

d (use ms, pp.

Lotus "e (A. Gray) Greene [10,5,4,5] D, L, R; 6100-8860;

G, U; br, mc, mm, ms, pj, pp, ra.

Lupinus CUM Pursh var. argenteus [-,

10900"; P, U; mc, mm, ms, pj, pp, ra.

Lupinus o Pursh var. argophyllus (A. Gray) S. Watson

[6,6,6,25] D, L, R, T; 6300-10150% J, G, P, U; mc, mm, mr,

ms, pj, pp, ra.

[Lupinus caudatus Kellogg var. argophyllus (A. Gray) S.L.

Welsh]

-,2,20] R, T; 7150-

la, 1,- 25] D, R; 7950- 10200’ J, bé af, mc, mm, mr, ra.

| ursh x Hook. & Arn)

"Cede "ES 9300-10135": J, P, U; m

uc pin Pursh var. rubricaulis (Greene) a L. Welsh

M-Nelso D; J.

Lupinus area S. Watson [-,-

11,1] D, R; 6250-72377; P,

U; pg, pj.

Lupinus kingii S. Watson [3,2,5,5] D, R; 6240-9020; J, U; mi,

ms, pj, Pp.

if | E II LT ff 10 L

I | £a n.a . in L | £545.24 ET Sf)

1/7

Phaseolus e A. Gray var. latus M.E. Jones [1,-,-,-

Psaralidiut: m (Pursh) Rydb. cl 11,210.57

5300-7130"; J, G, P, U; ds, fr, mr, pj, r

Psoralidium tenuiflorum (Pursh) Rydb. ri -,4,-] D, L, R; 5800-

7660; J; ms, pj,

Robinia neomexicana A. Gray var. neomexicana [41,9,4,4] D, L,

R, S; 9560" J, G, U; af, br, fr, mc, mm mr, ms, pp, ra.

Sophora nuttalliana B.L. Turner [1,-.-,-] S; 5600; G; pj.

Sophora stenophylla A. Gray [UNM-Spellenberg € Spellenberg

4992] R; G.

* Sphaerophysa salsula (Pall) DC. [1,-,-,-] R; 5760; G; ra.

Thermopsis doni as ex i Nutt. ex Richardson

var. divaric ly [7,-,-,-] D, R; 6600-9900’;

J; mc, mr, ms, B ra.

¿Diirehi KB tt Dir] J

ly [29,24,3, 24]D, L,R; 6600-11150! n

; af, br, mcr mm, mr, Ms, pp,

m attenuatum Greene [-,-, a n 9300-10150’; U;

mm.

Trifolium gymnocarpon Nutt. [1,-,-,5] R; 7100-8750", J, U;

mm, PJ, pp.

* Trifolium hybridum L. [-,-,1,1] D,R; 6000-10020; J, U; mr, ra.

us Marion Nutt. var. reflexum A. Nelson [1,1,-,5] D, R,

a 10600"; J, U; mm, mr, ra.

» dm pratense L. ee D, R; 6500-9800"; J, U; fr, mm,

mr, pp, ra.

* Trifolium repens L. [27,37,2,7] D, L, R; 6500-10300’; J, U; af,

c, ml, mim; m na oP ee sf.

"Cl-L.l: YD l

Sl D, R; 7675-8000"; J; pj, pp, ra.

[Lupinus ammophilus Greene]

ue polyphyllus Lindl. var. s si E. Jones) L.

Phillips [-,-,-,4] R; 8200-9680" U; mc

Lupinus pusillus Pursh var. pusillus [1,- 2, i E 5300- 5800’;

, G; fr.

Lupinus sericeus Pursh [14,-,-,3] D, R; 7675-9100"; J, U; af, mc,

¿ Mr, ms, pp, ra.

* Medicago lupulina L. [22,8,2,10] D, R; 5300-9980'; J, U; af,

, MC, , mr, ms, pp, ra.

* Medicago sativa L. [17,2,2,4] D, R; 5710-9640” J, P, U; fr,

mm, mr, pj, pp, ra.

* Melilotus albus Medik. [15,7,3,4] D, L, R; 5710-9200’; J, G, P,

U; fr, mm, mr, pj, pp, ra.

* Melilotus officinalis (L) Pall. [40,6,2,14] D, L, R, 5, T; 5710-

625 U; br, ds, fr, mc, mm, mr, ms, pj, pp, ra.

a ei (Pall.) A. DC. var. pulcherrima S.L. Welsh & A.

r [UNM-Sivinski 1929] R

a al Pursh var. beca Gray [4,-,1,2] D, L, R;

5860-9350’; J, P, U; ds, fr, mc, pi, r

me a sericea Nutt. var. sericea L8] R, T; 7237-109007;

c, mm, pp.

sane ‘ilfoli Torr. € A. Gray ex A. Gray [2,-,3,-] L, R;

re ites aeum p ex Torr. & A. Gray) Rydb. var.

hypo m [-,-,2,-] D; 5460" J; fr.

mois angus us A. s var. angustissimus [-,-,2,-] D,

0-6

5005 J; r

[-,9,-,-] D; 7950- 9200’: J; mm

Vicia americana Muhl. ex 2, e 7,22] D, L, R; A

O0"; J, G, U; af, br, ds, mc, mm, mr, ms, pg, pp, ra, S

Vicia o Nutt ex Torr. & A Ga var. ludoviciana [-, a

D; 5500-6300’; J; ds, pj.

Fagaceae

Quercus gambelii Nutt. [83,30,9,48] D, L, R, T; 5460-10900"; J,

, B, U; af, br, mc, mm, mr, ms, pj, pp, ra

n ee — [7,-,3,3] D, L, R; 5400- 84505 J, P, U;

eu ers CENE [6,-,14,-] D, L; 5400-8200"; J, G;

, MS, pj, pp.

x Quercus xundulata Torr. [4,-,12,9] D, L, R, S; 5400-8450" J,

P, U; ms, pj, pp

Fumariaceae

uoc aurea Willd. var. aurea [14,10,-,6] D, L, R, S, T;

—10900’; J, G, P, U; af, br, mc, mm, mr, ra, sf.

condi aurea Willd. var. occidentalis Engelm. ex A. Gray

,7,2,2] D, R; 5500-9600’; J, U; af, br, fr, mc, mm, mr, pj,

o5 ra.

+ Corydalis caseana S. Watson ssp. brandegeei (S. Watson) G.

Ownbey [-,-,-,1] R; 10700-11000 U; mr

Gentianaceae

0 Eustoma exaltatum (L) c ex G. Don [UNM-Spellen-

berg & Spurrier 8249] R;

Reif et al. Fl ftheT i} New Mexico

Frasera speciosa ea ex Griseb. [12,5,1,1] D, L, R; 7450-

10675

75% 3, U; af, br, mc, mm, ms, pp.

Gentiana bigelovii A. o [-,7,1,-] D, L, R; 7560-10300’; J, U;

mc, mm, mr, pp.

[Gentiana affinis Griseb.]

Gentiana parryi Engelm. [-,-,-,2] R; 10230-10600"; U; mm.

Gentiana prostrata Haenke ex Jacq. [-,1,-,-] D; 8440"; J; mr.

Gentianella amarella (L) Bórner var. aie (Michx) Herder

19,9, 1] D, R; 8460-112755 J, U; mc, mm, mr.

Gentianella amarella (L.) Borner var. heterosepala (Engelm.)

Dorn [UNM-Fleck s.n.] R; J.

PSOE thermalis (Kuntze) H.H. Iltis [-,2,-,1] D, R; 8610-

Swertia perennis L. [-,1,-,1] D, R; 9190-10940; J, U; mm, mr.

Zeltnera arizonica (A. Gray) Mansion [UNM-Tierney & Fox

563] D; J,

Geraniaceae

* yc ien (L) L'Her. ex Aiton [15,9,2,7] D, R, S, T;

0': J, G, P, U; ds, fr, mc, mm, mr, pg, pj, pp, ra.

EL caespitosum E. James [29,30,5,15] D, L, R; 5825-

10200"; J, G, U; af, br, ds, mc, mm, mr, ms, pj, pp, ra.

Geranium dea Fisch. & Trautv. [41,44,4,15] D, L, R;

7750-11000’; J, U; af, br, mc, mm, mr, ms, pp, ra, sf.

Grossulariaceae

Ribes cereum Douglas [28,5,7,35] D, L, R, S, T; 5600-9580"; J,

, P, U; ds, fr, mc, mm, mr, ms, pj, pp, ra.

Ribes inerme Rydb. var. inerme [33,5,3,-] D, L, R; 6950-9075";

; c, mm, mr, ms, pp, ra

Ribes leptanthum A. Gray [11, 25, 6, Nur D, R, T; 9500-10230'; J,

/ c, mm, mr, ms, pj, pp,

Ribes s montigenum n [3,3,- el D. L R; 7550-11150; J,

EM on riley T 0,2,-] D, L, R; 83200-10700’; J; af, mc,

mm, mr, sf.

HAIOIBOBCEAE

hi fei K r 1127: 8200 Q2C fy: Lhe mr

n, OJU

Ltrs! p Mygilll.

ar cea

Fendlera ria in “ay var. falcata (Thornber) Rehder [1,+,---

D; J; fr.

a o A Gray var. wrightii A. Gray [2,-,1,-] L, R;

5500- sJ; fr,

Jamesia americana Torr. & A. Gray var. emericana [25,23,3,2]

D, L, R; 6300-10300’; J, U; af, br, mc, mm, mr, ms, sf.

Philadelphus microphyllus A. Gray [9,6,7,-] D, L, S; 5860-9200;

J; Or, mc, mr, ms, pj, pp, ra

[Philadelphus microphyllus subspecies]

Hydrocharitaceae

Elodea bifoliata H. St. John [-,2,-,-] D; 8560-9570’; J; ml, mr.

odea canadensis Michx. [2,3,-,2] D, R; 7960-10020": J, U;

Hydrophyllaceae

Hydrophyllum fendleri (A. Gray) A. Heller var. ER [12,7,3,3]

D, L, R; 6500-10900'; J, U; af, mc, m

Nama dichotomum (Ruiz & Pavon) E » 2 73] D, R, T;

6990-8328’: J, P, U; br, pg, pj.

947

Nama hispidum A. Gray [-,-,1,1] D, R; 5530-6200"; J, U; ds, fr.

Nama retrorsum J.T. Howell [3,-,-,-] R; 5760-6090"; J, G; ds,

r, ra.

Phacelia alba Rydb. [5,1,-,6] D, R, T; 7185-9850"; J, P, U; mc,

mm, mr, ms, pg, pp.

Phacelia caerulea Greene [1,-,2,-] D, L; 5500-5600" J, G; fr,

pj.

dis crenulata Torr. ex. 5 Watson [-,-,3,3] D, R, S, T;

0-6760'; J, P, U; ds,

Pu ier On Pursh var. heterophylla [15,15,1,10] D,

t Ree 7 , y, / , gi

Phacelia regi Torr. var. E [5,-,2, 11] Di RS

5470-6980"; J, G, P, U; ds, fr, pj, r

us pinkavae N.D. Atwood [-,-, 53 D, S; 5500-6600", J;

pj.

Hypericaceae

Hypericum scouleri Hook. [8,12,1,1] D, R; 7100-9300’; J, U;

Iridaceae

Iris missouriensis Nutt. [41,40,3,25] D, - R; 7100-11150"; J, U:

, Dr, ml, mm, mr, ms, pp, ra,

Sornchum dennis Greene [2, = E D, R; 5300-7490"; J,

al p E.P. Bicknell var. occidentale (E.P. Bick-

nell) D.M. Hend. [2,-,-,-] R; 7060-7200’; J; fr, mr.

Sisyrinchium montanum Greene var. montanum [1,5,1,3] D,

R; 8400-10900’; J, U; ml, mm, mr.

hi glandaceae

M" nigra L. [-,-,3,-] D; 6000; J; mr.

nca

s a us Willd. var. balticus (Willd.) Trautv. [25,27,2,24]

R 5825 9830; J, U; fr, me, m , mm, mr, ms, pp ra.

ti illc]. var (Will Roem. & Schult.)

Balslev [5,7] D, R; 6500- 8700" J; mc, mm, mr, ra.

Juncus bufonius L. [1,1,1,-] D, R; 5360-8920"; J, U; mc, mr.

Juncus confusus Cov. [-,-,1,4] L, R; 6550-10600% J, U; fr, mr,

mm.

Juncus drummondii E. Mey. [-,-,-,1] R; 10350-10940", U; mm.

Juncus dudleyi Wiegand [8,5,1,1] D, L, R; 5600-10100’; J, G,

Juncus ensifolius Wikstr. var. montanus (Engelm.) C.L. Hitchc.

[17,20,2,12] D, L, R; 6500-10900; J, U; mc, ml, mm, mr,

ra, sf.

Juncus interior Wiegand [3,10,-,1] D, L, R; 6700-9700’; J, U; fr,

mc, mi, mm, mr, pp, ra

Juncus pel ini is Torr. [12,14,1,4] D, R; 5360-10600'; J, U; fr,

ml, mm, mr, pp, ra,

Juncus mene NUS "rie -—- R; 9300-101355 J; mc.

Juncus nevadensis S. Watson [-,4,-,2] D, R; 8500-10600’; J, U;

ml, mm, mr.

Juncus nodosus L. [2,-,-,1] D, R; 6010-7100"; J, U; fr, pj.

Juncus torreyi Covilie [-,1,1,-] D; 5400-8500"; J; mr.

Juncus xiphioides E. Mey. [-,-,4,-] D, L; 5400-9700” J; mr.

Luzula comosa E. Mey. [2,2,-,-] D, R; 9200-9720"; J; af, mc,

mr.

948

Luzula parviflora (Ehrh.) Desv. [6,6,1,3] D, L, R; 8350-11000’;

mm, mr, S

Juncagin

OG nm L. [2,-,-,-] D; 6330’; J; ml.

ae palustre L. [-,2,-,-] D; 8610-8720"; J; mm, mr

Lam

+ e pallidiflora (A. Heller) Rydb. var. neomexicana

(Briq) RW. sanders Dol Di 8000, Jr ms.

n.coLcn AFAL

f F

J, G, U; af br, ds, fr, mc, mm, mr, ms, pp, ra , Sf.

Hedeoma drummondii Benth. [4,-,5,9] D, R, S, T 5400-7900";

J, P, U; ds, fr, mr, ms, pj, pp.

Hedeoma nana (Torr) Bria. [3,-,-,-] D, L; 5600-6100’; G; fr, pj.

Hedeoma oblongifolia (A. Gray) A. Heller [-,-,1,-] D; 5500;

fr.

Lycopus americanus Muhl. ex W.P. Barton [-,-,2,-] D, L; 5400;

J; ft, mr.

* Marrubium vulgare L. [4,1,1,5] D, R, T; 5550-87505 J, P, U;

, ms, pj, ra.

Mentha arvensis L. [6,11,1,7] D, L, R; 5825-10020' J, U; fr,

is Menina spicata - [-,-,1,-] D; 5700;. Ji mr.

IPIE E E 2E FA

] D, L, R; 5850-9580’; J, U; br, mc, mm, mr, ms pp.

Monarda pectinata Vu n ,5,3] D, L, R, S, T; 5550- 2280) J,

* Nepeta pai L. eee 1165] D; J.

Poliomintha incana (Torr) A. Gray [1,-,-,-] R; 5955", J; fr.

a ot L. var. lanceolata (W.P.C. Barton) Fernald

a D, L, R; 5825-10900’; J, U; af, fr, mc, ml, mm,

se , Sf.

Salvia farinacea Benth. [UNM-Hutchins 6648] R; U.

d D, L; 8200-9050’; J; br, r

,3,-] D, S; 6000-6540"; J; " pj, ra.

Salvia subincisa Benth. sd 715; 65007; J; pj.

Satureja vulgaris (L) Fritsch [1,-,2,-] D; 5900-8280’; J; mm,

mr.

y] D; 7490-9450'; J; mm,

——

* Scutellaria galericulata L. [1,1,-

UM pilosa Nutt. var. pilosa [-,-,-,1] R; 9000-92007; U; mr.

Teucrium laciniatum Torr. [1,-,-,-] S; 6660-6720”; G; pj.

Lemnaceae

Lemna minor L. [2,5,-,4] D, L R; 5600-10900’; J, G, U; fr, ml,

mr.

Lemna valdiviana Phil. [-,-,1,-1 D; 8000’; J; mr.

Lentibulariace

Utricularia ere LeConte [1,-,-,-] R; 89005 J; ml.

Lili ( Iso Alliaceae, Asparagaceae, Convallariaceae,

Melanthiaceae)

Calochortus gunnisonii S. Watson var. gunnisonii [2,10,2,6] D,

R; —10900’; J, U; af, mc, mm, ms, pp.

Calochortus nuttallii Torr. 8: A. Gray [2,-,1,6] D, R, T; 6200-8600’;

J, B. U; ds, ms, pj

! E J: pp.

Erythronium grandiflorum Pursh var. grandiflorum [UNM-

odsen 9. ]

% Lilium philadelphicum L. [-,-,1,-] D; 68007; J; mr, mc.

£T 3/51

ICAGS HL]

RANN-SO Im

HD LORI.

db LR 8350-9920;

Streptopus amplexifolius (L) DC. [7,3,-,

J; af, mc, mr, sf.

Linaceae

Linum aristatum Engelm. [2,-,-,-] R; 5800-6300’; P; fr, ra.

Linum australe A. ps var. australe [4,-,-,2] R, T; 7200-8600’;

J, P, U; mm, pj, pp, r

Linum osea A. ee [-,-,2,-] D; 6000’; J; pj, ra.

Linum lewisii Pursh var. lewisii [7, 3,1,2] D, R, 5; 6660-11150';

, G, U; ds, mc, mm, mr, pj, pp, ra, sf.

Linum neomexicanum Greene [-, 3, -] D, L; 7150-8200"; J;

)S ] R; 6700-6980"; U; pj.

ar puberulum Engem) À. Heller a -,1,4] D, R, T; 6300—

05 J, P, U; ds,

poasateae:

icaulis (D c Hook.) Douglas ex Torr. & A,

Gray [2,-,3,3] D, L R 3 T 5400- 71505 J, G, P, U; fr, pj.

+ 9 Mentzelia conspicua Todsen [1,-,-,-] R; 7900-8160" J; ra.

Mentzelia E x Gray) J. Darl. var. humilis [-,-,-,1] T;

8320; P

Mentzelia a ud Darl. [2,-,-,3] D, R, T; 6100-8560";

7 fr, pj, pp, ra

Mentzelia dle (Nut A. Gray var. multiflora [15,-,1,10] D,

¿R,S, T; 5" J, G, P, U; ds, fr, ms, pj, pp, ra.

Mentzelia m unis [5,-,-,-] D, R; 7675-8350"; J; ds,

+ * Mentzelia springeri (Standl) Tidestr. [1,-,5,-] D; 5600-7250;

J; pj, pp, ra

Malvaceae

* Alcea rosea L. [-,-,2,-] D, L; 6000-8900; J; ra.

* Malva Mr Wallr. [3,1,1,-] D, R; 8000-8350’; J; fr, mm,

Sea condi A. Gray var. candida [16,14,5,8] D, L, R;

0020"; J, U; mc, ml, mm, mr.

seen neomexicana À. pla var. neomexicana [1,-,-,1] D, R;

7675-8350’; J, U; uU

Spl | ) Rydb. var. coccinea [14,-,3,20] D,

L, R, T; 5300- 8740’ J, G, P U; ds, fr, mm, pj, pp, ra.

Sphaeralcea fendleri A. Gray var. fendleri [11,2,5,4] D, R, S;

-81007 J, G, U; ds, fr, mm, ms, pj, pp, ra.

Sphaeralcea grossulariifolia (Hook. & Arn.) Rydb. var. pedata

(Torr. ex A. — Kearney [5,-,-,-] D, R; 5825-8475’; J; fr,

mm, ms,

Sphaeralcea incana Torr. ex A. Gray var. cuneata Kearney [3,-,

E 5550-7130"; J, G, P, U; ds, fr, pj, ra.

Sphaeralcea i incana Torr. ex A. Gray var. incana [1,-,-,-1 D;

20/5:

T paio A. Nelson [4,-,-,3] R; 5800-6975’; G, P,

U; ft, pg, pj,

Melanthiaceae [traditionally in Liliaceae]

Veratrum californicum T. Durand var. californicum [2,8,-,7] D,

L, R: 8625-10900’; J, U; mc, ml, mm, mr.

Zigadenus elegans Pursh [2,10,3,3] D, L, R; 7500-11150"; J,

mr, sf.

[Anticlea elegans (Pursh) Rydb.]

Reif et al. El f the T y) tains. New Mexico

Morace

* Morus P L. [2,-,1,-] D, L; 5400-5800"; J, G; fr.

bie disi

l [-,-,-,2] R: 6550-7100; U; pj.

Abronia fragrans Nutt. ex Hook. [6,- r6] D, R, S, T; 5470-7220’;

, G, P U; ds, fr, pj.

Boerhavia spicata Choisy [UNM-Yarnel! 189] D; J.

Mirabilis albida (Walter) Heimer! [-,20,-,-] D; 7950-9580"; J;

, mm, mr, ms, pp, ra.

Mirabilis glabra (S. Watson) Standl. [2,-,-,3] R, T; 6000-69755

U; fr, pj, ra

Mirabilis ae (Pursh) Heimerl var. decipiens (Standl.) S.L

,",3,8] D, R; 8300-8475"; J, U; mm, mr.

abi pda (Pursh) Heimerl var. linearis [9,2,1,10] D, L, R;

8900’; J, G, P, U; br, ds, mm, mr, pj, pp, ra

Mic va (Standl.) Spellenb. [2,-,-,-] D; 8000-

05 J; mm, pp.

is multiflora (Torr) A. Gray var. dics (Standl.) J.

7,-] D, L; 6000-6500’; J;

Mirabilis muito (Torr.) A. Gray var. muliftora Sii 510] D, L,

0; J, G, B. U; br, ds, fr, p

Mirai pra (A. "e A. Ene I. 31 D, RT

0-8328; J, P, U; mr, pg, p

Oleaceae

Forestiera | t. [12,1,8,3] D, L, R, S; 5470-8940", J,

G, U; fr, mc, mr, pi ra ra.

M7 l llr

1,-] D: 5400": J; fr.

Bp jJ —, STV,

Menodora scabra A. Gray [-,-,1,-] D: 5700' J; fr.

Onagraceae

Chamerion in (L) Holub var. canescens (Wood) N.

| n & P. Holmgren [8,5,1,3] D, R; 7330-11000% J,

P, U; mc, mm, mr, ra,

1 ,

| RA s |

Circaea alpina L. var. alpina [1,-,2, ] L; 8000", km

ig e IMG Lam. [1,-,-,1] R; ae 10900’; J,

[Epi ae alpinum L.]

Epilobium oo C. Presl [-,2,1,5] D, R; 7200-10180’;

br, mm,

! Epilobium Mes end Hoch & W.L. Wagner [-,-,-,1] R;

Rode 8/40; U; ml.

) Hoch & PH. Raven

Epilobium ciliatum Raf. var. ne [11,19,3,8] D, : 5400-

005 J, U; fr, ml, mm, mr, ra.

[Epilobium ciliatum]

T pE e | Al L 3 fA r2 241

D, R; 8900-9800"; J, U; mm, mr, ra.

[Epilobium ciliatum]

Epilobium halleanum Haussk. [1,4,-,1] D, R; 8600-11000% J,

; mm, mr, ra.

Epilobium hornemanii Rchb. var. hornemanii [UNM-Fleck

E, “Todos . " L] I [-,3,-,2] D, R; 8650-101 50":

a diffusum Torr. & A. Gray var. strictipes (Hook ) Dorn

,91 D, R; 7900-9500’; J, U; mc, mm, ms,

949

Gayophytum ramosissimum Torr. & A. Gray [-,-,-,71 R, T;

7150-9250"; P, U; mm, pj, pp.

Oenothera albicaulis Posh e ¡4,61 D, R, S, T; 5500-7900"; J,

G, P U; ds, fr, Pl. PP:

4" D, oDcr. II

Oenothera brachycarpa A. Gray L-,1 E D; 6400" J;

Oenothera cespitosa Nutt. var. macroglottis (Rydb.) Crenauist

4,-,-,9] D, L, R, T; 6100-8475"; J, G, P, U; ds, ms, pj, pp, ra.

Oenothera cespitosa Nutt. var. marginata (Nutt. ex Hook. &

Arn.) Munz [10,-,1,3] D, L, R, S, T; 5440-8375"; J, G, P, U;

r, ms, pj, pp.

Oenothera cespitosa Nutt. var. i is (W.L. Wagner, Sto-

ckh., & W.M. Klein) Cronquist [1,-,-,1] D, R; 6200-7700;;

r Y; DJ, PP.

das coronopifolia Torr. & A. Gray ~ 3,1,18] D, R, T;

00’; J, P, U; ds, mm, ms, pj, pp,

Moi curtiflora W.L. Wagner & Hoch A -,1,-] D, L, R;

5710-6330’; J, G, P; fr, pj, ra

Oenothera elata Kunth var. hirsutissima (A. Gray ex S. Watson)

Cronquist [14,-,6,-] D, L, R, S; 5850-8500"; J, U; br, mc,

mm, mr, pp, ra.

I I yr

tt [3,16,1,3] D, R; 7550-9600’;

J, U; mc, c ml, mm, mr, , pp; ra

O ssp xis (A Gray) W.L. Wagner

och [-,-,4 E D; 5600-6500" J; fr, pj.

Did pallida Lindley var. latifolia Rydb. [-,-,

5450-6250’; J; fr.

Oenothera pallida Lindl. var. pallida [1,-,-,-] R; 6000-6300’;

J; fr

4,-] D;

P. [I Ge gen iC

Cronquist [7,-

-/61 D, L, R, S, T; 5600-7500"; J, G, BU; ds, Fr ms, pj, pp.

Oenothera serrulata E D, R e oa ji ic

Par A a a $ SEPE AAA lA A O AO PARA = A ES

] D, L, R, S; 5550- 8475" 1,G, U; ds, fr, pa, pj. pp, ra

Oenothera villosa Thunb. var. strigosa (Rydb.) Dorn [1, 4,- 710;

8400" J; mm, mr, ms, pp.

rchidaceae

Calypso bulbosa (L.) Oakes var. americana (R. Br.) Luer [4,-,1,-]

D, R; 7500- ^J; mc.

Corallorrhiza maculata (Raf.) Raf. var. maculata [-,-,1,-] D;

600; J; mr.

I Id da A TÉ

occidentalis (Lindl) Ames

[8,9,3,3] D, L, R; 6600- 10900’: J, U; af, mc, mm, mr, sf.

Corallorhiza striata Lindl. var. vreeranai (Rydb) L.O. Williams

F rer

i t I f 1

Corallorrhiza wisteriana Conrad 12] D, R; 9100-10050’;

* Cypripedium parviflorum Salisb. var. pubescens O.W. Knight

-,7,2,-] L; 7200-7700"; J; ms

* Fpipactis gigantea sino. ex Hook. [1,-,1,1] D, L, R;

7250"; G, U; fr, ml, m

oe oblongifolia Raf. [2, te 1,2] D, R; 8600-10230’; J, U;

,Imc, mm, my, s

eae. repens (L.) R. Br. ex W.T. Aiton [1,3,1,-] D, L; 7950-

^J; mc, mr.

Malaxis porphyrea (Ridley) Kuntze [UNM-Castetter 3498] D; J.

950

Malaxis soulei L.O. Williams [-,5,1,-] D; 8025-8980’; J; mc,

mm, ms, pp

pj lonis Sheviak [-,1,-,-] D; 9050’: J; mr

ee i a A

Elta feta Buranensis (Nutt) Lindl. [2,1,-,4] R; 9500-11000"

Ú (R ) Sheviak &W.F. Jenn. [2,5,-,1]

D, R: 7100- 10900: J, U; mcm

Platanthera sparsiflora (S. om dum -,-,5,-] D; 6500-

000"; J; ml, mr

+ Spiranthes romanzoffiana Cham. [-,1,-,-] D; 8620; J; ml.

F^ L |)

a angustifolia (Nutt) G Don: var. dubia A. Nelson -,7,7,2

R; 7620- ; U; ms, pj.

ms exilis A. Nelson [2,-,1,-] D, L, R; 5400-6560’; G, J; fr.

Castilleja minor (A. Gray) A. Gray]

Castilleja integra A. Gray [28,1,5,39] D, L, R, S, T; 5600-9200’,

J, eu br, ds, A mm, mr, ms, pg, pj, pp, Y

th. [10,-,2,5] D, R;6150- "9680, .U;

br, ds, fr mc, mm, pj, p

Mem R; 7100-10150"; J, U; mc,

mm, mr, pp, ra. ra.

1 | Hook. var. miniata [29,21,4,18]

D, L L, R; 6975- 10900: J, U; af, br, mc, mm, mr, ms, pp, ra, sf.

a occidentalis Torr. [-,-,-,2] R; 9500-10940"; U; mr,

Baie: rhexifolia Rydb. [1,-,-,-] D; 10000’; J; m

Castilleja suiphurea Rydb. [1,3,-,11] D, R; 8350- ine J, U;

f WW MM

Haynes [2, -,2,-] D; 8150-8860" J; mc, ms, pj, pp, ra.

iur wrightii A. id ssp. wrightii [-,-,2 p o LR;

7280; J, U; fr, pj, r

pets fasciculata Nutt. a -,2,71 D, R, T; 5500-8600’; J, P,

var mexicana (A Gray ex S. ll

Orobanche we Nutt. var. ludoviciana [-,-,-,1] T; 7185";

P. R; On oe J, ee pa inp

ib R, T; 7550-10230*; J, P,

c, mm, mr, ms, pi,

Orthocarpus purpureoalbus a dem i Watson [1,-,4,5] D, L,

; 00-8000’; J, P, U; br, ds, pj, p

Pedicularis centranthera A. Gray [5,-, el D, R; 6700-8700’; J,

mr, ms, pj, pp, ra

aig ed Retz. [-,-,-,4] R; 9300-11000’; U;

MAN parryi A. Gray [UNM-McGrath 158] R; U.

Pedicularis procera A. Gray [7,5,1,4] D, R; 8180-10900; J, U;

Mc, mm, mr, 5

Pedicularis racemosa Douglas ex Benth. var. alba (Pennell)

Cronquist [2,2,-,3] D, R; 9300-11150"; J, U; mc, mm,

Oxalidaceae

Oxalis corniculata L. var. wrightii (A. Gray) B.L. Turner [UNM-

McGrath 111] R; U.

Oxalis metcalfei (Small) Knuth sensu G.L. Nesom manuscript

[4,6,2,-] D; 7950-9725'; J; mm, mr, pp, ra.

ET ary)

lexas 3(2)

FP L r . 1 . C Tal 7

* Parnassia fimbriata Konig [UNM-McGrath 1533 R; U.

Parnassia palustris L. var. montanensis (Fernald & Rydb. ex

Rydb.) C.L. Hitchc. [-,1,-,1] D, R; 8610-93507, J, U; mr.

ml I J C L

Callitrichaceae,

in part]

Ca sai Aa L. [-,1,-,1] D, R; 8400-10020’; J;

calice heterophyla P Pursh var. heterophylla [-,-,1,1] D, R;

m s rein i [- i -,2] D, R; 8450-11000"; J, U; ml,

Collinsia parviflora Lindl. [-,-,-,1] R; 7800-7900"; U; pj.

Limosella aquatica L. [2,2,2,2] D, R; 5360-8980’; J, U; ml, mr.

*elinaria dalmatica (L) Mill. subsp. dalmatica [2,-,-,-] D;

5710-8375’; J; fr, ra.

igaris Mill. [1,-,2,5] D, L, R, S; 5400-9200’; J, G, U;

linden dubia is al [-,-,1,-] D; 5360'; J; fr.

mulus glabratus Kunth var. jamesii (Torr. & A. Gray ex

jue. A. Gray [6,3,4,-] D, L R; 5500-8700", J, G, U; fr,

ml, mm, mr.

Mimulus guttatus DC. [17,10,4,8] D, L, R; 5600-11000’; J, U;

Mimulus rubellus A. Gray [-,-,1,1] D, R; 5460-85007; J, U; fr ms.

iie tilingii Regel [4,-,-,-] D, R; 8000-9075"; J; mm, mr

Penstemon ambiguus Torr. var. ambiguus [-,-,1,-] D, 5900;

J; ds.

Penstemon angustifolius Nutt. ex Pursh var. caudatus (A. Heller)

Rydb. [2,-,-,3] D, R, T; 6500-7500’; J, P, U; ds, fr, ms, pp.

Penstemon barbatus (Cav) Roth var. torreyi (Benth.) A. Gray

[39,27,4,9] D, L, R, 5 T; 5500-9850" J, G, P, U; af, br, ds, fr,

j, Pp, ra

Penstemon barbatus (Cav) Roth var. ae A. Gray [10,-,-

300-9830’; J, U; af, mc, ms, pp, r

Penstemor crandallii A. Nelson var. iio (Pennell) G.T.

et & RC. d [7,-,-,17] D, R; 7310-9200", J, P, U; ds,

, 1

+ Pede S uum A. Nelson var. taosensis (Keck) G.T.

Nisbet & R.C. Jacks. [-,-,-,8] D, R; 6850-9450"; P, U; mc,

pj, pp.

! ipods Pl Pursh var. alpinus (Torr.) A. Gray [1,-,-,-] L;

7500-8 ; DE

aa 00 inii A. Nelson [1,-,-,2] R; 8000-9670’; J, U;

S BJ.

"eds inflatus Crosswhite [1,1,-,-] D; 8000-8280"; J;

, MS.

Penstemon jamesii Benth. [3,-,1,-] D, L, S; 5850-6660"; G; ds,

r, pj

A. Gray subsp is (A. Nelson)

EB Maas ^l a R; ye S000: J, U; ds, mr, . pj, pp.

il. [1,-,-,-] D; 8240-8260";

-t

J; pp.

Penstemon palmeri A. Gray [1,-,-,-] 5; 5980-6300; J; ra.

Penstemon rydbergii A. Nelson var. rydbergii [-,6,1,-] D;

7550-9075’; J, U; mm,

Reif et al. FI New Mexico

Penstemon secundiflorus Benth. [3,-,3,5] D, L, R; 6100-8475";

J, U; fr, pj pp.

Penstemon strictiformis Rydb. [-,-,1,-] 5; 6200"; J; ra.

Penstemon strictus Benth. [8,1,-,15] D, R; 6775-10900; J, U;

at, mc, mm, ms, pp, ra.

ydb. [-,-,3,-] D, L; 6600-6800’; J

[Penstemon virgatus ssp. asa- -grayii Crosswhite

Penstemon virgatus A. Gray [3,-,-,7] D, R, T; 7025-9200; J, G,

; pj, pp.

P, U; ds, mm, ms, pj, pp.

Penstemon whippleanus A. Gray [9,8,-,3] D, L, R; 8180-11000";

; mc, mm, mr, sf.

Plantago argyraea Morris [6,-,-,-] D, R; 7125-8000"; J; mm,

j, pp, ra

dicis lanceolata: [12,1,-,1] D, L, R; 5710-9200; J, G, P,

* Plantago major L. [13,17,2,5] D, R; 6000-102305; J, U; mc,

ml, mm, mr, ra

Plant i ica Jaca. [4,-,5,9] D, L, R, S, T; 5400- 81005J,

G, P, U; ds, fr, pj, pp, ra

M plantaginea (E. james) Benth. [5,2,2,3] D, R; 7900-

10300" J, P, U; af, mc, mm, mr, pp.

ipn americana Schwein. ex nii nd 14,4,7] D, L, R;

000-10900'; J, U; af, br, m ?

T s anagallis-aquatica L. "e -43, a B L, R, S; 5360-6500;

Veronica eka hl L. var. xalapensis (Kunth.) St. John & FW.

n [4,28,3,4] D, L, R; 7200-10230’; J, U; mc, ml, mm,

mr, pp, ra.

am "TD" VahlI9.7.2.31

Lave? pp

D, L, R; 6850- 10600"; J, U; mc, mm,

Veronica AA Roem. & Schult. a j ES R; 9980-11000’;

Poaceae

Achnatherum hymenoides (Roem. & Schult) Barkworth [38,-

2,37] D, R, S, T; 5300-8850’; J, G, P, U; br, ds, fr, mc, mm,

rns, v P9; pj, PP, ra.

i (V B th [2,28,3,14] D, R;

8050-11000’; J, U; "m mc, a mr, ms, Pp, ra, sf.

Achnatherum lobatum (Swailen) Barkworth [-,-,-,31 R;

6300-7600; U; ds, pj.

lA ) Bad tl | fi [2,-,-,5]

D, R: 7850-10150": J, Us mc, mm, ms, pp.

ien perplexum Hoge & Barkworth [1,-,-,1] D, R;

205J, U;

uc re vase) 5 edo ud 7,5] D, R T;

7000-815 P, U; ds, m jp

Achnatherum a MM peta [UN M-Sivinski

e cylindrica Host [3,-,1,1] D, R, 5; 5440-7675"; J, G,

U; fr, pj, ra.

* Agropyron cristatum (L.) Gaertn. var. cristatum [6,2,3,10] D,

R, T; 6500-10000; J, P, U; mc, mm, mr, ms, pp, ra

[Agropyron cristatum]

* Agropyron cristatum (L) Gaertn. var. desertorum (Fisch. ex

Link) Dorn [11,-,-,9] D, R, T; 7100-8600"; J, P, U; ds, ms,

Pg, pj, PP, ra.

[Agropyron cristatum]

951

Agrostis exarata Trin. var. minor ls [4,2,1,3] D, R; 5760-

5J, U; mc, ml, m

i gloann Roth E i al 5 R 7200-9200’; J, U; mm,

ds a Willd. [7,43,2,11] D, L, R; 7200-11150 J, U; af,

, mm, mr, ms, pp, ra, sf.

* Agrostis stolonifera L. [10,11,5,4] D, L, R; 5300-9950" J, U;

Alopecurus aequalis Sobol. var. aeua [6,23,2,12] D, R;

7080-10900’; J, U; mc, ml, mm, mr, ra.

* Alopecurus pratensis L. [3,-,-,2] D, R; 7900-10900’; J, U; br,

mr, ra.

Andropogon gerardii Vitman ssp. gerardii [-,-,3,-] D, R;

-7250 J; pp.

Aristida adscensionis L. [2,-,1,-] L, R; 6090-6500; J, P; pg,

pj, ra.

Aristida arizonica Vasey [-,-,2,1] D, L, R; 6500-7300"; J, U; pp.

Aristida divaricata Humb. €: Bonpl. ex Willd. [-,-,1,-] D; 6900’;

J; pj

HN ELEM AI 4-4. ig li a CS IBS M

[1,-,-,13]

R,T; 6450-8300; J, P, U; ds, pj, pp, ra.

yLi™Me

DL, R, S, T: 5825— 9475; J, ic P, U; ds BS pj, Bp

Aristida purpurea Nutt. var. nealleyi (Vasey) Allred [3,-,1,1] D,

:6100—7060* J, G, U; fr, pj.

* Arrhenatherum elatius (L) P. Beauv. ex J. Presl & C. Pres! [1,-

rr] R; 7890; J; ra.

Beckmannia syzigachne d Fernald [-,1,1,2] D, R;

5360-9200"; J, U; fr, ml, m

L * } fd

)T. Koyama

Kuwano]

Blepharoneuron K (Torr.) Nash [3,49,2,10] D, L, R;

6300-11150; J, G, , mc, mm, Mr, MS, pp, ra, sf.

Bothriochloa barbinodis Lào) Herter [-,-,1,-] D; 5950"; J; pj.

Bothriochloa laguroides (DC.) Herter ssp. torreyana (Steud.)

Allred & Gould l2 , ul D, R; a a el " a U; fr, pj, ra.

n --L : El z ff ANA 65005

J; ds, pj. |

Bouteloua barbata Lag. var. barbata [-,-,1,1] R, T; 6200’; P,

ft.

a curtipendula (Michx.) Torr. var. caespitosa Gould &

adia [1,-,-,5] R, S, T; 5760-7350’; G, P, U; fr, pj

P curtipendula (Michx.) Torr. var. curtipendula [8,-

4,12] D, L, R, S, T; 5400-7600" J, G, P, U; br, fr, pg, pj,

, fa.

Bouteloua eriopoda (Torr) Torr. [-,-,2,9] D, L, R, T; 6100-7600;

y Fs U, OS, IM, D ; DJ.

Bouteloua gracilis (Kunth) Lag. [10,17,3,46] D, L, R, T; 5400-

; J, G, P, U; br, ds, fr, mm, ms, pg, pj, pp, ra.

Bouteloua hirsuta Lag. [1,-,2,1] R; 5450-6200; J, P, U; pj, ra.

Bouteloua simplex Lag. [-,2,-,5] D, R, T; 7185-8670"; J, P, U;

j, pp, ra.

Bromus carinatus Hook. & Arn. [3,3,-,5] D, R, 5; 6140-10900';

, U; br, mc, mm, mr, ra.

* Bromus catharticus Vahl [3,-,1,-] D, L, S; 5410-6330” J, G,

; fr, ra.

Bromus ciliatus L. [10,37,3,2] D, R;¡6750-11150';J, P, U; af, mc,

mm, mr, ms, pp, ra, s

952

-6350; 5 i ra.

s. [57,12,3,26] D, | à d 1; 5825-10600";

J, P, U; af, br, ds, fr, mc, mm, mr, m 5

Eoo Thunb. ex Murray 112, i 2 31D, LA/S

5500-8750’; J, G, U; fr, mm, mr, pg, pp, ra

,",2,3] D, L, R: 5700-9200"; J,

Bromus frondosus (Shear) Wooton & Standl. [-,-,3,-] D, L;

5600

Bromus dnd (Shear) Rydb. [-

; fr, mc, mm, mr, pj.

Bromus porteri (J.M. Coult) Nash [2,11,-,10] D, L, R; 8075-

10200; J, U; br, mc, mm, mr, ms, pp.

[Bro malus Rupr. ex E. Fourn.]

nd pu impelli: Scribn. [1,-,=,-] D; 9050"; J; m

Bromus richardsonii Link [-,-,-,8] R; 7960-11000'; A af, mc,

mm, mr.

* Bromus tectorum L. [39,8,4,30] D, L, R, S, T; 5550-10200'; J,

P. U; af, br, ds, fr, mc, mm, mr, ms, pg, pj, pp, ra.

Calamagrostis canadensis (Michx.) P. Beauv. var. canadensis

3,15,1,7] D, L, R; 7950-11000’; J, U; mc, mm, mr, sf.

Ca p n stricta (Timm) Koeler ssp. inexpansa (A. Gray)

CW. eene [-,2,1,-] D, R; 8620- oe en

"m rly] D, R; 5400-5955;

-— f

Chloris ala Nutt. [-,-,-,2] R; 6300-6600’; U; ra.

* Chloris virgata Sw. [2,-,-,1] R; 5760-6600'; G, P, U; ra.

Cinna aoia ee ex Goepp.) Griseb. [1,2,-,1] D, R; 8550-

9 ml, mm

=I meu pad L [^j r] D; 6100; J; mr.

idi glomerata L. [45,15,2,9] D, L, R; 6000-10900", J, U;

» PP, ld, sf

canon intermedia Vasey [-,4,-,2] D, R; 8690-10600"; J,

mm.

Shs eee Scribn. [11,48,4,8] D, L, R; 7550-11150"; J,

af, br, mc, mm, mr, ms, pp, s

Danthonia spicata (L.) P. Beauv. ex Roem. & Schult. [3,8,5,1]

D, L, R; 6200-8910" J, U; mc, mm, mr, pp, ra.

Deschampsia cespitosa (L.) P. Beauv. var. cespitosa [4,32,-,15]

D K ue 11000 4y U; mie mm: mr, pps

fp MIA

pr more

(Nash) Gould [ -,^,3,-] D; 5600- 6200’; J; mr.

* Digitaria sanguinalis (L) Scop. [1,-,-,-] D; 83605 J; ra.

Distichlis spicata (L) Greene var. stricta (Torr) Scribn. [1,-,1,-]

, R; 5300-6000’; J; fr, ra

* Echinochloa crus-galli (L.) P. Beauv. [-,-,1,-] D; 5460", J; fr.

* Echinochloa muricata (P. PCM Fernald var. microstachya

Wiegand [1,-,-,-] R; 5760*; G; ra.

as pde L. var. ok [4,-,3,1] D, L, R; 5400-

7100"; J, G, U; fr, mc, mr, pp, ra

* Elymus on (Host) Runem. var. elongatus [3,1,-,-

S; 6150-83505 J, U; mm, pp, ra

Elymus elymoides (Raf) Swezey var. brevifolius (J.G. Sm.) Dorn

[78,58,6,63] D, L, R, S,T; 5500-11150'; J, G, P, U; af, ds, fr,

mc, mm, mr, ms, pg, pj, pp, ra, sf.

[Elymus longifolius (Smith) Gould]

Flymus glaucus Buckley var. glaucus [8,5,2,3] D, L, R; 7000-

]D,R,

3, U; af, br, mc, mm, mr, pp.

* cos iid (Opiz) pues var. hispidus [10,9,-,3] D, L,

OS J, U; br, me, mm, mr,

z P ee (Opiz) ee var. iene (Griseb.)

Dorn [7,2,2,3] D, L, R; 6900-9980"; J, U; mc, mm, mr, ms

Xx ee ee ee ee |

l Elymus junceus Fisch. [6,-,-,4] D, R; 6050- 8900’ J, U; fr, mm,

pj; pp, ra

[Psathyrostachys juncea (Fisch.) Nevski]

Elymus lanceolatus (Scribn. & J.G. Sm.) Gould [1,-,-,1] L, T;

6300-8560"; G, P; pg, pp.

xmi xpseudorepens (Scribn. € J. G. Sm.) Barkworth €

wey [1,-,-,-] R; 8400-8700’; J; m

* usd repens (L) Gould [5,-,-,-] D; Y 8350’; J; mm,

x Elymus xsaxicolus Scribn. & J.G. Sm. ex Scribn. [-,1,-,-] D;

00 “di

_ [Unstabilizea hybrid]

ld [21,7,3,27] D, R, S, T; 5860-9500’;

J, G, P, U; br “ds, f mc, mm, mr, ms, pg, pj,

Elymus trachycaulus (Link) Gould ex Shinners ssp. nas

(Link) A. Love & D. Love [1,6,1,1] D, R; 8290-10180’; J, U;

af, me uu pp, ri ra.

Ol LA r7 LJ Elis

trachycaulus

[124447111 C D, L R; 5600- 10330; J, P, U; af, br, ds, mc,

mm, mr, ms

Elymus cule un Gould ex Shinners ssp. violaceus

m.) A. Love & D. Love [-,12,-,2] D, R; 8300-1 1000";

, Sf.

* Eragrostis cilianensis (All) Lutati ex Janch. [-,-,2,-] D;

i P oe (Schrad) Nees conferta Stapf [-,-,2,-] D;

Eragrostis ds (Lam.) Britton, Sterns, & Poggenb. [-,-,1,-]

D; 53605 J; fr.

Eragrostis mexicana (Hornem.) Link ssp. mexicana [1,-,-,-] R;

760’; G; ra.

Fj ri (Michx) N Steud. var. miserrima (E

Fourn.) Reeder [RM-Castetter 587] D; J

Eragrostis pectinacea {Michx.) Nees ex Steud. var. pectinacea

1,-,2,-] D, R; 5450-6000’; J, G; fr, ra.

Eragrostis trichodes (Nutt.) A.W. Wood. [-,-,1,-] D; 6300 J; ra.

Festuca arizonica Vasey [11,58,6,35] L, D, R; 6000-109005 J,

RU af, Dr, mc, mm, mr, ms, pp, ra.

hreb. [18,8,1,1] D, L, R; 5600-9250";

J, G, U; fr, uu mi Epa

It. ex Schult. & Schult. f. ssp. colora-

densis F red. [- ¿Cr 1D; 8550"; J; m

Festuca calligera (Piper) Rydb. [-,-,-,6] R; 8450-10600"; U;

m, pp.

e idahoensis Elmer [10,8,-,5] D, R; 6900-9720’; J, U; af,

mc, mm, mr, ms, pp, ra, sf.

* Festuca pratensis Huds. [3,2,2,1] D, R; 5600-8460"; J, U; ml,

Festuca rubra L. ssp. rubra [1,-,-,-] R; 9475-9840’; J; mm.

Festuca saximontana Rydb. var. saximontana [1,4,-,2] D, R;

7940-11000’; J, U; mc, mm, ms, ra, sf.

Festuca sororia Piper [-,10,2,-] D, L, : 8000-10150’ J, U; af,

mc, mm, mr.

Festuca thurberi Vasey [5,33,5,15] D, L, R; 7950-11150; J, U;

r, mc, mm, mr, PP, ra, S

ins. New Mexico

hylla (Hack, Krajina [-,-,1,2] L, R; 8200-9600";

J r T or f + f

introduced in Jemez area after human mediated fire,

etc.

Glyceria borealis (Nash) Batch. [1,5,3,5] D, R; 7950-10020’: J,

F l, f i

Glyceria elata (Nash ex jig i E. Jones [2,1

7100-10900’; J, U; ml, m

Glyceria grandis S. Watson a 17, a 4] D, L, R; 7860-10020’; J,

U; ml, mm, mr.

74] D, R;

\ Lore,

[29,10,4,11] D, L,R:6500-10000"

af, br, mm, mr.

Hesperostipa comata (Trin. & Rupr.) Barkworth var. comata

[8,-,2,17] D, R, S, T; 5860-10900’; J, G, P, U; ds, fr, mm,

rns, pg, pj, pa ra.

0D in

rin r. intermedia

(Scribn. & Tweedy) Dorn [B, 2,- -,3] D, R; 7200- 92005 J, U;

2 0

esperostipa neomexicana (Thurb, ex J.M. Coult) Barkworth

[1,-,2,2] D, L, R; 5600-6600"; J, G, U; fr, pj.

Hierochloé odorata (L.) P. Beauv. [2,3,-,-] D, R; 8760-9600; J;

Hilaria jamesii (Torr.) Benth. [14,-,4,27] D, L, R, S, T; 5530-8000’:

U; ds, fr, ms, pg, pj, pp, ra

[Pleuraphis jamesii Torr.]

Hopia obtusa (Kuntz) Zuloaga & Morrone [-,-,3,-] D; 6000’;

J; ra.

Hordeum brachyantherum Nevski [2,14,1,6] D, R; 8450-10900";

F F f l, ti

A jubatum L. ssp. intermedium Bowden [5,11,-,7] D,

500-9580’; J, U; mG mi mm, 1 PP, fa ra.

Hordeum jubaturn | ssp. jut 8,2,4] D, L, R, $; 5300-

9570"; J, G, U; br, ds, fr, mc, ml, i mm, mr, ms, pj, p

* Hordeum murinum L. ssp. glaucum a a. Pl

, R, S; 5470-8260"; J, G, U; br, fr, pj, ra

Hordeum pusillum Nutt. [1,-,-,1] R, S; 6300- 6720'; G, U; pg,

pj.

Koeleria macrantha (Ledeb.) Schult. [29,58,6,43] D, L, R T;

7 1150’; J, G, P, U; af, br, ds, fr, mc, mm, mr, ms, pj,

pp, ra, sf.

Leersia oryzoides (L.) Sw. [-,-,3,-] D, L; 5400-5500; J; fr.

* Lolium perenne L. var. E Willc. [4,-,-,-] D, L; 7525-

8250* J, U; br, mm, mr, ra.

* Lolium perenne L. var. perenne [3,-,1,1] D, R, S; 5760-9800’;

, G, U; fr, mc, mm, po, ra.

Lycurus setosus (Nutt.) C. Reeder [-,2,2,4] D, L, R, T; 6300-8625";

Melica porteri Scribn. var. porteri [9,8,2,-] D, L, R; 6975-9580’:

; ài, Mc, mm, mr, ms.

Muhlenbergia andina (Nutt) Hitchc. [1,-,-,-] D; 8000-8100’;

J; mm.

* Muhlenbergia arsenei Hitchc. [1,-,3,-] D, L; 5800-6600";

J; pj, ra.

Muhlenbergia asperifolia (Nees & Meyen ex Trin.) Parodi [2,-,-

117 D, R; 6550-7550" J, U; fr, mr, ra

E. Fourn. ex Hemsl. [-,-,1,-] D; 6750’; J; br.

(Thurb. ex S. Watson) Rydb. [-,-,-,6] R,

T, 8300- 10200 P, U; mc, mm, pj.

T MEN UNE REINES] à doi:

A LI CPL

953

icon Mu Ud (L.) Trin. var. mexicana [1,-,1,-] D, R;

; J, U;

muh ls ima (Steud.) Swallen [1,1,1,1] D;

6920-8300" J; pj, pp, t

"esos montana Tm Hitchc. [4,46,4,17] D, L, R, T;

6300-10300’; J, G, P, U; af, mc, mm, mr, ms, pj, pp, ra.

Muhlenbergia pauciflora Buckley [-,1,2,3] D, L, R; 6300-9050’;

ms, pj, pp

Muhlenbergia porteri Scribn. ex Beal [-,-,1,-] D; 5800’; J; fr.

Muhlenbergia pungens Thurb.[3,-,-,9] R, T; 6000-7100" J, P,

U; ds, fr, pg, pj, ra.

iig racemosa (Michx.) Britton, Sterns, & Poggenb.

-,-,1,-] D

nus E (Kunth) Swallen [-,7,1,1] D, R;

00" J; mr.

— pp, ra.

Muhlenbergia repens (J. Presl) Hitchc. [-,1,-,-] D; 8700"; J; mc.

idc richardsonis (Trin.) pini E En ,6] D, R, T;

00’; J, P, U; mc, ml, mm, mr,

Ve. (gens (Benth.) Hitchc. UN Jacobs 4276]

Muhlenbergia sinuosa Swallen [-,1,-,-] D; 7950-8100’; J; pp.

Muhlenbergia thurberi Rydb. [-,-,-,8] R; 6200-7600”; G, U;

"o a. Tuas Hitchc. ex Bush [-,-,2,11] D, R,T;

P, U; ds, p .

o in id " Gu [3,14,1 ip D, L, R,T;

6300-9220’; J, G, P, U; ds, mc, mm, mr, ms, pp, r

Munroa squarrosa lll [2,-,1,12] D, L, RT; P 8720"

J, G, P, U; ds, fr, pj, r

Nassella viridula ae —— [6,-,-,1] D, R; 7100-8250;

J, U; ds, pg, Ppp; fa.

O [20,9,3,1] D, L, R; 6600-10200": J,

; : af br, mc, mm, mr, pp, sf.

Panicum bulbosum Kunth [1,-,2,-] D; 5850-7400": J; pp.

Panicum capillare L. var. brevifolium (Rydb.) Shear [-,-,1,1] D,

R; 5460-8350’; J, U; fr, pp.

Panicum hallii ird var. hallii [-,-,1,-] D; 5600’; J; ds.

[-.-.1 -] D; 6000; J; ra

Fer d

Phalaris arundinacea L Var. Miri dE [2,-,2,1] D, R;

5400-9150" J, U; fr, ml, m

Phleum alpinum L. var. alpinum c ii 3 11]D,L,R;5850-11150';

U; af, mc, mm, mr, ra, sf.

liis pratense L. var. pratense [19,16,2 de 2 F R; 5700-

0180’; J, U; af, fr, mc, ml, mm, mr, ms, pp,

Prog australis (Cav.) Trin. ex Steud, ssp. americanus Sal-

nstall, Peterson, Sog d rl, 7l L; cana J; TE

Pi 19]

D, R, S, T; 5725- 8700": J, G, P, U; n d mm, mr, ms,

pg, pj, PP, ra.

FT. X rr» I "RI

[-,1,-,-] D;8540-8740';

J; pp.

* Poa annua L. [5,9,1,-] D, L, R; 5550-10230; J; af, mc, mm,

mr, ra, S

Poa arctica R. Br. var. grayana (Vasey) À. Lóve & D. Lóve &

Kapoor [-,-,-,3] R; 8300-10150% U; mc, mm.

Poa arida Vasey [2,-,-,-] D; 83400-9900’; J; af, mm.

954

ioi ard ia & Scribn. [UNM- Goodrow 430] R; U.

L. [-,^,-,2] R; 6950-7900" U; pj.

ü pe een L. [4,1 7,2.8] D, L, R; 7800-10250’; J, U; af, br,

me, mm, mi ms, pp. ra.

i ey [16,13,5,32] D, L R, S, T; 6000-

10675’; J, P, U; af, ds, mc, mm, mr, pg, pj, pp. ra.

[Poa fendleriana subspecies]

Poa glauca Vahl var. Mee (Nas ex Rydb.) B. Boivin [-,1,-

D; 10340-10875’; J; sf.

Poa interior Rydb. [-,1,-,2] D, R; 8700-10600" J, U; mc, mm,

sf.

Poa und Trin. [3,-,-,1] R; 9300-11000; J, U; mc, mm,

mr, sf.

Poa wi Vasey [-,6,-,2] D, R; 8100-10900’; J, U; af, mc,

mm, mr, pp.

Poa palustris L. [4,6,2,5] D, R; 7950-9950" J, U; mm, mr

* Poa pratensis L. [59,39,11,40] D, L, R, T; 5600-111505 J, G, P,

U; af, br, ds, fr, mc, mm, mr, ms, pg, pp, ra, sf.

[Poa pratensis subspecies]

Poa tracyi Vasey [UNM-Fleck Rida

Poa wheeleri Vasey [-,-,-,1] R; 7560-7740" U;

* Polypogon monspeliensis ie ES a 2, cs 5300- 5400’;

J; fr.

* "uu viridis (Gouan) Breistr. [2,-,-,-] L, S; 5600-57607;

G; fr.

Puccinellia nuttalliana (Schult) Hitchc. [UNM-Sivinski 2469]

Bde m rdus eah (Nutt.) Trel. [2,-,1,-] L, R; 5650-

200';

; pj, PP,

an ee (Torr) Swallen [-,1,2,-] D, L; 7450-

ED neomexicanum Nash [2,-,3,-] D, L, R; 5760-

sur J, G; pj, Pp, ra.

ran £

,5] D, L, R; 5950- 8450’: J, G, P, U; br, fr, mr, r ms, pj, pp. r ra.

* Secale cereale L. [1,-,1,-] L; 6650-8900'; J; br, pj.

Setaria grisebachii E. Fourn. [-,-,1,-] D; 6000 J; ra.

Setaria Jeneapna (Scribn. € Merr.) K. Schum. [1,-,1,-] L;

5500-5700" J, G; fr.

* Setaria pumila (Poir.) Roem. & Schult. [-,-,1,-] L; 66507; J; ra.

* Setaria viridis (L.) P. Beauv. [1,-,4,-] R; 5400-60007; J, G; mr, ra.

Sorghastrum nutans (L.) Nash [1,-,2,-] D; 6000-7350"; J; mr,

Pp.

* Sorghum bicolor (L) Moench ssp. bicolor [1,-,--] L; 5450';

G; fr.

Sporobolus airoides (Torr) Torr. [2,-,1,4] D, R, T; 6000-8200"; J,

ms, pj, ra

liada ol (Poir) Merr. var. compositus [-,-,1,-]

Sporobolus RM Hitchc. [-,-,3,7] D, R, T; 6000-6975; J,

r, pj. Y

Sonic E T (Torr.) A. Gray [12,2,3 a DER, SI

500-8900’; U; br, ds, fr, mr, pg, pj, p

Sobol flexuosus EE ex Vasey) Rydb. e R; 6200

eee pallida (Torr) G.L. Church var. pauciflora (J. wen

Davis [9,12,1,-] D, L R; 6300-9750’; J, U; mc, ml, m

mr, pp, ra.

I [| E aL Dos a Im LI FE FM £T. Y be Y!

Trisetum montanum Vasey [4,23,2,1] D, L, R; 7950-11150; J,

c, mm, mr, ms, pp, S

Trisetum spicatum (L.) K. Richt. L, 6,-,1] D, R; 7950-10600’; J,

J; mc, mm, mr, S

Trisetum wolfii Vasey [-,1,1,2] L, R; 8000-11000’; J, U; af, mm,

mr, sf.

x Ro d I L Brrr] pan i T; 6000-7600; J, P; fr, ra.

| Ib. [2,-,5,12] D, L, R, T; 5300-8550"

J, P, U; ds, fr, ms, pg, pj, pp.

[Vulpia octoflora varieties]

Polem

dis inna edi ex A Gray) J.M. Porter [2,-,1,-] L 5;

= Inc

mes lineari Nutt. 4 fi ï D, R; 7550-10900; J, U; mc,

ais ALE, TA (A. Gray) H Mason l- =-,7] RT 6050-7450';

P, U; ds, L pj.

A lalenan ccn A setratie (A D firamtr RAE Grant)

A.D. Day & VE. Grant [-,-,2,-] D; 5300- 5550’; J; ds, fr.

Gilia ophthalmoides Brand [-,-,1,3] D, R; 5400-6850"; J, U;

r, pj.

Gilia sinuata Douglas ex Benth. [1,--,1] R, S; 6320-8075”; G,

F r, pj.

Giliastrum acerosum (A. Gray) Rydb. [1,-,-,4] R; 5820-6600;

G, U; fr, pj.

a aggregata (Pursh) V.E. Grant ssp. candida J E.

Gra A.D. Grant [-,-,-,1] R; 8840-9200'; U; m

Pd aggregata (Pursh) V.E. Grant ssp. collina p

Wilken € Allard [1,-,-,-] D; 8400-9340; J; mc

Ipomopsis aggregata (Pursh) V.E. Grant ssp. formosissima

(Greene) Wherry [36,28,2,28] D, L, R, T; run

, B. U; af, br, ds, mc, mm, mr, ms, pj, p

Pa laxiflora (J.M. Coult.) V.E. Grant E j 2 S, T; 6320-

9 ds, pj

pomopss nine (Torr) V.E. Grant ssp. longiflora [-,-,-,14]

RT;6 7300’; P, U; ds, fr, pj

Ipomopsis a En (Torr) V.E. Grant ssp

[6,-,6,-] D, L, R, S; 5300-7000; J, G; ds, fr, pj, pp, ra.

p piis (Nutt.) V.E. Ri [4,-,-,15] D, R, T;

05 J, P, U; br, ds, fr, pj, pp, r

bomopss munis (Nutt.) V.E. Grant UNNE T Tierney P-52175-

Wilken

lepiosiphon nuttallii (A. Gray) ia d ssp. tenuilobus (R.

Patterson) ined. [1,-,-,-] D; 10000; J; m

Linanthus pungens (Torr.) J.M. ei & L.A. Jose [1,-,-,4]

, 1; 7500-9250’; J, P, U; mm, mc, ms, pp.

Microsteris gracilis (Hook) Greene [1,-,-,7] R; 7100-8500; J,

U; mr, pg, pj, pp.

DI t f, Covilla cen dH (A n Wl

H; 6775;

[Phlox austromontanal

Phlox longifolia sp. longifolia [4,-,-,3] D, R; 7300-8350";

J, U; ds, pg, pj, pp, ra

Polemonium foliosissimum A. Gray [14,8,2,4] D, R; 5400-

10900’; J, U; af, mc, mm, mr, pp.

Polygalac

Polygala alba ne var. alba [1,-,-,-] D; 7300-7600; J; pp.

Raif at a] : ri £ al T | ! L] Naw Mavirn

Polygonaceae

Bistorta bistortoides (Pursh) Small [1,-,2,1] D, R; 8960-10600":

F t

Bistorta vivipara (L.) S.F. Gray [UNM-Flecks.n.] R; J, U

riogonum abertianum Torr. var. abertianum [-,-,1 E D; 5500;

J; ds.

Eriogonum alatum Torr. var. alatum [14,-,2,13] D, L, R, T;

450-9

350°; J, P, U; br, ds, mc, mm, ms, pj, pp.

i aed annuum Nutt. [2,-,1,2] D, R; 5950-7280’; J, P, U;

fr, pj, 1

Eregonum a Greene var. arcuatum [1,-,-,-] D;

renum jamesii Benth. var. flavescens S. Watson]

Eriogonum cernuum Nutt. [-,-,3,5] D, R, T; 5300-8328"; J, P;

ds, fr, pg, pi.

Eriogonum corymbosum Benth. var. velutinum Reveal [-,-,-,1]

R; 6818-6900’; U; fr.

id a Benth. var. jamesii [8,-,8,35] D, L, R, T;

5300-9200; J, G, P, U; ds, fr, mr, pg, pj, pp, ra.

M oum lachnogynum Torr. ex Benth. var. colobum

Reveal & A. Clifford [-,-,-,2] T; 6818-6900’; P; ds.

Eriogonum leptophyllum (Torr. & A. Gray) Wooton & Standl.

[-,-,-,1] R; 8250; U;

Eriogonum lonchophyllam Torr. & A. Gray var. lonchophyllum

[1,-,-,1] D, R; 6990-7400"; J, U; ds, pj.

T E ipi iL Torr. & A. Gray var. aia (Torr)

Reveal [1,-,-,-] R; 7100’; J;

Eriogonum PU Nutt. var. simpsonii (Benth.) Reveal

De ,2,20] D, L, R, T; 4004 8700"; J, P, U; ds, ms, o pg, pp.

g t. [9,3,2,32] D, R, T 2000 967054,

P, U; br, mc, mm, ms, pj, p

* Fallopia convoivulus L. [1,1,1 J b 6000-8750: J; ds, mm, m

Persicaria amphibia (L) S.F. Gray [1,1,2,2] D, R; 5300- Bo

, U; fr, ml, mr.

* Persicaria lapathifolia (L) A. Gray [1,5,1,11 D, R; 5360-8740’;

, U; fr, ml, mm, mr.

* Persicaria maculosa A. Gray [6,-,4-] D, R, S; 5300-7350"; J,

; fr, ml, mm, mr

» irren aviculare L. [4,30,4,11] D, L, R, T; 5400-10330’;

J, B, U; af, fr, ml, mm,

Polygonum douglasii Greene " 4,1 ,7] D, L, R; 7300-10230;

J, U; af, br, mm, mr.

Polygonum lis L M] D; 7600’; J; ra.

contortiflarn inn

(Nutt. ex Piper) Hickman [-,-,-,2] R; 9380-9680"; U; mr.

Polygonum ramosissimum Michx. var. ramosissimum [2,-,-,-

RN 5 ees G, P; d

Il [-,25,1,51 D, R, T; 5900-10300’;

J, P, U; af, fr, mc, mm, mr, ms,

ia are L.[14,47,3,2] D, L R: poe 11300" J, U; af,

c mm, mr, ms, pp, ra, sf.

ü uns res L. ssp. ee [9,12,4,-] D, R; 5400-9700"; J;

ZTlD.ZGz6onm an Ir. I. ft

rei y Vee

lic VWlatcan [

* Rumex patientia L. [-,-,1,-] D: 695 0'J; m

Rumex triangulivalvis (Danser) Rech. f. [5, 23 ,A3] D, b, B. S; T;

5400-10020’; J, G, P, U: fr, ml, mm, mr, ra.

955

Stenogonum salsuginosum Nutt. [1,-,-,-] D; 7000’; J; ds.

Portulacaceae

Lewisia pygmaea (A. Gray) B.L. Rob. [-,-,-,2] R; 9850-10600’;

; mm, sf.

Montia chamissoi (Ledeb. ex Spreng.) Greene [2,3,-,-] D;

8350-10000’; J; mm, mr.

Phemeranthus parviflorus (Nutt) Kiger [1,-,1,2] D, R; 6990-

JON ^ U; Or, Ses Ph Pp.

ui E onila e L. [1- 2, 171 D, R i 5460-8350° P, U; ds,

fr, pa, a.

E e" subspecies]

Potamogetonac

Potamogeton dn Balb. [-,2,-,1] D, R; 8590-9950"; J, U;

Ty us + Enf: rr Ccp r5

1 L 1

,11 D, pD.ocnn 9150"

] D, R; 7950-8625’; J, U; mr.

; :ml

-,-] D; 7960- 95705 J; ml, mr.

-,1] D, R;

TOU On edid L. [-,2,=,-

Potamedeton nodosus Poir. [-,6,

Potamogeton richardsonii (A. Benn.) Rydb. [-,1,-

8500-10500"; J, U; mr

Primulaceae

A lis L. [24,38,1,28] D, L, R,T; 5860-1 1000":

J, G, Pu; af, Dr, ft, me, ı mm, mr, ms, pj, pp, ra, sf.

Ichellum [11,8,2,4]

D, R; 7550-1 0600’; J, U; me, mm, mr.

Ranunculaceae

Aconitum columbianum Nutt. var

L, R; 8000-11000; J, U; af, mc, m

Actaea rubra (Aiton) Willd. [37,9,3 ait D, L, R: 6950- 10200’; J,

U; af, br, mc, mm, mr, ra, sf

[Actaea rubra ssp. arguta (Nutt.) Hultén]

Anemone cylindrica A. Gray [1,-,3,-] D, R; 6850-8900’; J, U;

l lr

[5,7,1,5] D,

m, mr.

Neue coerulea E. James var. coerulea [4,4,2,2] D, R, S;

8180-10900"; J, U; mc, mm, mr, sf.

Aquilegia elegantula Greene [22,8,2,1] D, L, R; 6850-10200’;

; c, mm, mr, pp, ra

Caltha leptosepala DC. [4,3,-,2] D, a 8440-11000’; J, U; mc,

ml, mm, mr.

Clematis bigelovii Torr. [1,-,-,-] D; 7600’; J; m

Clematis columbiana (Nutt.) Torr. & A. a var. columbiana

[31 de D, L, R; 6300-10900'; J, P, U; af, br, mc, mr,

ms, PP,

Clematis Ria Pursh var. hirsutissima [1,-,-,] R; 8070-

Clematis hirsutissima Pursh var. scottii (Porter) E.O. Erickson

[-, 1 R; soii 500; U; mr.

Cl t. [10,-,3,1] D, R;

; 3300-8030" J, G, P;

Delphinium M E Huth [3,-,-,2] D, R; 8830-11000’;

, U; af, mc, mr, sf.

Delphinium nuttallianum Pritz. [2,-,-,9] R; 8125-10900’; J, U;

i 1 J f

Delphinium ramosum Ryd b. [-,-,-,1] R; 9980-10020'; U; mr.

956

Delphinium wootonii Rydb. [1,-,2,-] D, S; 6320-6500; J, G;

s, fr, pj.

Myosurus minimus L. [1,-,-,-1D; 7080'; J; r

Pulsatilla patens (LJ Mill. ssp. multifida e Zámels [1,-,1,4]

D, L, R; 7540-9000’; J, U; mc, ms, pp, ra

% Ranunculus abortivus L. [3,-,1,-] D, R; 6850-8550” J; mm,

mr.

! Ranunculus alismifolius Geyer ex Benth. var. montanus 5.

Ranunculus eT L. var. diffusa With. [2,2,2 a D, R;

6500

Ranunculus scons Hook. [1,12,5,-] D, R; 7550-9150;

y, F |, i t .

Ranunculus DD Pursh [9,3,2,-] D, L, R; 5300-8580; J;

me mm, mr, ra

in1mnendsc [14 15.2.51 D

L, R; 6700- 9750’; al U; ana mc, mm, mr, ra, sf.

3,1] D, L, R; 6000- 10020";

ml, mm, mr, ms, P

ines ‘ranunculinus (Nutt.) UE [1,-,-,-] L; 8960" J;

mc.

I* Ranunculus repens L. [1,-,-,-] D; 7490’; J; m

Ranunculus sceleratus L. var. multifidus “ee [1,-,2,-] D, L;

5400-5800"; J, G; fr.

AT R: 53260-9770":

Pa AO FEN

Don ex G. Don [5,-,

J, U; af, mc, mr.

[Ranunculus uncinatus var. earlei (Greene) L.D. Benson]

Thalictrum fendleri Engelm. ex A. Gray [26,13,2,8] D, L, R;

6975-9950’; J, U; af, br, mc, mm, mr, ms, pp, ra, sf.

Trautvetteria caroliniensis (Walter) Vail [9,2,2,1] D, L, R;

8400-10000’; J, U; af, mc, mm, mr, sf.

Rhamnaceae

Ceanothus fendleri A. Gray [21,9,2,4] D, L, R; 6900-9600; J,

U; br, mc, ms, pp, ra

Rosaceae

Agrimonia gryposepala Wallr. [4,-,-,-] D, R; 7050-81507 J; mc,

mm, mr.

Agrimonia striata Michx. [4,3,3,-] D, L, R; 7960-8900 J; mm,

mr.

Amelanchier alnifolia (Nutt) Nutt. ex Roem. var. alnifolia

[23,2,2,3] D, L, R; 6600-10150"; J, U; af, mc, mm, mr, ms,

pp, Sf.

o Roeane [lets dA 8050; J; ra.

7,37] D, LAS,

T 5600- 9450’: J, G, P, U; br, ds, fr, mc, mr, ms, pj, pp.

Crataegus erythropoda Ashe [-,-,4,2] D, L R, S; 6500-8350;

;mr,ms.

Crataegus macracantha Lodd. ex Loud. var. occidentalis (Brit-

ton) Eggles ston [1,-,-,-] D; 7100; J; m

(L) Rydb. [17,16,1 pum R; 7475-10900’;

U; af, mc, mm, mr.

E P (D. Don) Endl. ex Torr. [18,-,6,20] D, L, R, 5,

T; 5550-8475"; J, G, P, U; ds, fr, ms, pj, pp, ra

Fragaria vesca L. [25,31,4,3] D, L, R; 6450- 10150; J, U; af, br,

mc, mm, mr, ms, s PP, ra, sf.

|. [27,21,2,6] D, L, R;6975-10300;; J, U;

af br, mc, mm, mr, ms, pp, ra, st.

Geum pde ig [6,3,2,-] D, L, R; 7800-8900; J; mc,

ml, mm

d macrophyllum Willd. var. perincisum (Rydb.) Raup

,5,6] D, L, R; 6600-10900’; J, U; d mc, mm, mr.

e e L. [-,N,1,-1 D, L; 8000-9600"; J; m

Geum triflorum Pursh var. ciliatum (Pursh) Rut » a 2,21 D, R;

8530-9850; J, U; mc, mm, ms, ra

! Geum triflorum Pursh var. triflorum [1,-,-,-] D; 8500"; J; mr.

Holodiscus dumosus (Nutt. ex Hook.) A. Heller [13,16,4,10] D,

L, R, T: 6000-10230’; J, P, U; af, br, mc, mm, mr, ms, pj, sf.

* je pee Mill. [2,1,1,-] D, R; 6000-8300; J; fr, mm,

pocas irs de (Torr) J.M. Coult. d dd D, L, R;

6400- 0 J, U; af, br, mc, mm, mr, m

Potentilla anserina L. [9,12,2,7] D, L, R; a 9200; J, U; fr,

ml, mm, mr

Mas pU +a D I H " mm

Ib) T.F Wolf [2,1,1,-]

D; 6500-8050"; J; br, me, mr, ra

Potentill Ri ta (Rydb.) S.L. Welsh

& B.C. Johnst. [-,-,-,6] R; 7050-8550"; U; mr, + Dp, ra.

[Potentilla concinna]

ice concinna id var. concinna [-,2,-,-] D;

10340-111505 J; m

Potentilla crinita A. a var. crinita [1,-,2,-] D, L, R; 7310; J;

S, PP.

Potentilla a PA Lehm. var. diversifolia [-,3,-,3] D, R;

8625 ; af, mm

Potentilla un a ex Hook var. glabrata (Lehm) C

Hitchc. [1,-,-,-] R; 8060’; J; mr.

Potentilla gracilis Douglas ex Hook. var. pulcherrima (Lehm.)

Fernald [23,19,5,10] D, L, R; 7675-11150; J, U; af, mc, ml,

1507 J, Deal ne uu. pP, ra, sf.

var

i 31D.L ,R; 8240- 10900:

pms, T

Potentilla iuda Lehm. var. hippiana [29,62,3,24] D, L, R;

6975

al a fi l AO IA FA

DE — mr, ra.

rid ue lg ssp liensis (L) Asch. & Graebn.

a D, L, R; 5700- 10230: J, U; af, br, mc, mm, mr,

ms, pp „ra.

[Poten dia norvegica]

Potentilla paradoxa Nutt. [-,-,2,-] D; 5360-5400’; J; fr.

Potentilla jv L. var. pensylvanica [7,18,-,8] D, R, T;

7300-10230"; J, P, U; br, ds, mc, mm, mr, ms, pp, ra.

Potentilla rivalis Nutt. {1,-,-,1] L, T; 5400-7100’; G, P; fr, ml.

Prunus americana Marshall [-,-,2,2] D, R; 6100-8000” J, U;

mr, ra.

I* Prunus persica (L) Batsch [1,1,-,-] D; 7250-8080’; J; ml, ra.

Prunus virginiana L. var. melanocarpa (A. Nelson) Sarg.

[41,8,6,9] D, L, R, T; 5860-9600’; J, G, P, U; af, br, ds, fr, mc,

, IT, , PP, Ta.

Purshia stansburiana (Torr) Henr. [1,-,-,-] D; 7000-7400; J;

pJ.

Purshia tridentata (Pursh) DC. [1,-,-,-] R; 7400-8600’; J; pp.

Rosa acicularis Lindley var. sayana Erlanson [10,2,-,10] D, L, R;

940-9500”; J, G, U; mc, mm, mr, ms, pp, ra.

Rosa arkansana Porter var. arkansana [24,-,-,-] D, R; 6320-

8900’; J; br, ds, mm, mr, ms, pp, ra.

Reif et al. Fl f the Ti 1] New Mexico

Rosa arkansana Porter var. suffulta (Greene) Cockerell [1,-,-,-

D; Pe ms.

Rosa nutkana C. Presl [6,7,-,4] D, R; 8025-9400"; J, U; af. mc,

, MI, MS, PP, ra.

[Rosa nutkana var. hispida Fernald]

Rosa woodsii Lindl. var. ultramontana (S. Watson) Jeps.

[10,3,6,1] D, L, R; 6000-9425": J, U; fr, mc, mm, mr, ms,

lo TI Fh "mi le ee

pp, ra.

Rubus idaeus L. var

L, R; 6000-10940*; J, U; af, br, mc, mm, mr, ms, DD, ra, TE

[Rubus idaeus var. strigosus (Michx.) Maxim.]

is o Nutt. var. belie [27,13,3,4] D, L, R;

0’; J, U; af, br, mc, mm, mr, ms, pp, ra, sf.

* da minor Scop. ssp. a Brig. [1,-,-,-] D;

6200'

Sanguserba minor aa balearica (Bourg. ex Nyman) M.

Garmendia

proa da procumbens i > rr 1] R; 10350-10940", U; mm.

D; 8400-9200" J; af, mr.

cw LF f F 2 —r

Rubiaceae

+ de aparine L. var. echinospermum (Wallr.) Farw. [-,-,6,-]

D; 6000— mae

Galium aparine]

Galium ndn L. nh 10,3,14] D, L, R; 7200-10600’; J, U: af,

Galium fendleri A. n. [4,-,1,-] D, L; 6150-8400"; J; br, mc,

mr, ms, pp.

Galium mexicanum Kunth var. asperrimum (A. Gray) Higgins

& S.L. Welsh [23,29,1,-] D, L, R; 6500-10330’; J; br, mc,

m, mr, ra, sf.

Galium trifidum L. var. subbiflorum CUm [3,5,-,2] D, R;

9 940" J, U; mc, mr, pp, ra

Galium Mona Michx. [8,-,2,2] D, [^ 6955- 9150"; J, U; af,

, mm, mr.

Houstonia rubra Cav. [3,-,-,5] R; 5800-6800’; J, P, U; ds, fr,

Pg,

Rutaceae

Ptelea trifoliata L. [10,-,4,1] D, L, R, S, T; 5600-8475": J, G, P;

F i Li F Pal =

[Ptelea trifoliata infrataxa]

Salicaceae

x Populus xacuminata Rydb. [1,-,1,-] D, R; 6000-7400; J;

mr.

* Populus alba L. [-,-,1,-] D; 6000'; J; ra. [Individual plant

eradi

Populus o E. James [17,1,2,21] D, L, R; 5600-9000";

U; br, fr, mr, ms, ra.

Populus delióides W. Bartram ex Marshall var. wislizenii (S.

Watson) Dorn [16,-,1,4] D, L, R, S; 5470-7475; J, G, P.

U; fr mr, 5

Poj les Michx. [48,27,2,26] D, L, R; 6975—10700;;

J, P, U; af, br, mc, mm, mr, pp, ra, sf.

* Salix arizonica Dorn [1,-,-,-] R; 9300-10140": J; mc.

Salix bebbiana Sarg. [16,8,2,1] D, L, R; 6975-9725": J, U; mc,

mm, mr.

Salix boothii Dorn [RM-Dorn 8831] R: J.

957

u^ Sio Barratt ex Hook. [4,-,-,2] D, L, R; 8275-

ef F |

Salix see ant et Mich. var. igi (C.R. Ball) Dorn [5,4,-,-

D, R; 7950-9190"; J; mc, m

Salix exigua Nutt. var. exiqua [22, 1 de 6] D, L, R, 5; 5400-8375’;

, G U; fr, mm, mr, ra.

Salix gooddingii C.R. Ball [2,-,-,-] L, S; 5400-5800; G; fr.

Salix irrorata Andersson [6,1,4,-] D, L; 5400-7950’; J, G; fr,

Salix lasiandra Benth. var. caudata (Nutt) Sudw. [-,-,-,8] R;

7050- i

* a

! z |,

[Sali 2 hirida AAAI ssp —— pr Rin Mt urray ia]

Sa lacada Benth. var. lasiandra [4,1,- ,-] D, R; 7100- 8000;

mm, mr.

[Salix lucida Muhl. ssp. lasiandra (Benth.) Murray]

Salix Ae Bebb [3,-,-,2] D, R; 8350-9830"; J, U: mm,

Salix caia Pursh var. planifolia [-,-,-,1] R; 10350-10940’:

U; mr.

k. [7,7,-,5] D, R; 7950-10250’.

A d N

: af, mc, mm, mr, ms,

E un Bebb var. woffii IRM-Dorn 8847 R; U.

Santalaceae (Viscaceae)

Arceuthobium divaricatum Engelm. [2,-,-,4] D, R, T; 6400-7750":

J, B. U; pj, pp.

AUR louglasii Engelm. [-,6,-,2] D, R, T; 8450-10650’;

pP M

—

J, B U; mc.

Arceuthobium vaginatum (Willd.) J. Presl var. cryptopodium

(Engelm.) Cronquist [6,3,1,7] D, L, R; 7300-9340", J, U;

mc, pj, pp.

Comandra umbellata (L) Nutt. var. pallida (A. DC.) M.E. Jones

sa ,10] D, R, T; 5500-9600"; J, P, U; ds, fr, mc, ms, pj,

Pore Pur Engelm. var. UAM [4,-,3,6]

D, R, T; 5500-7950" J, G, P, U; fr, pj, p

aururacea

panei Eom (Nutt.) Hook. €: Arn. [-,-,3,-] D, S; 5350;

ml.

Saxifragac

Heuchera e Nutt. ex Torr. & A. Gray [6,3,2,12] D, R, T;

000-9720" J, P, U; br, mc, mm, mr, ms, pj

Heuchera rubescens Torr. [1,-, 0 J; mr.

Lithophragma tenellum Nutt. IUNM-Heil 8812] R; U.

Mitella pentandra Hook. [UNM-Feck s.n.] R; J

Saxifraga bronchialis L. var. austromontana (Wiegand) Piper

ex G.N. Jones [3,8,2,1] D, L, R; 7960-111505 J, U; mc, mr,

J Y, DOJU 7

S, pp, sf.

Saxifraga odontoloma Piper [2,-,-,1] R; 9300-11000’; J, U;

mc, mr, sf.

Saxifraga rhomboidea G

U m

da SR Ja

Scrophulariaceae (see also Orobanchaceae, Plantag-

inaceae)

Scrophularia lanceolata Pursh [1,-,-,2] R; 8200-9680’; J, U;

958

* Verbascum thapsus L. [7,11,2,4] D, L, R; 6300-9800" J, G, U;

af, br, fr, mm, mr, ms, pp, ra

Simaroubaceae

G; fr, ra.

Solanaceae

a coronopus (Dunal) A. Gray [2,-

00—630 o J, G, P, U; n ps

Datura wrightii Regel [1,-,2,- L; 5500-6000 l f G; fr, ra.

*eHyoscyamus niger L. [-,7,7,4] R, T; 7600-8740’; P, U; ds, ml,

pj, ra.

Lycium ipsi Miers Er D, L R, S, T; 5760-8150"; J, G,

m, pj, Y

eee a es Watson [UNM-Bel! s.n.] D; J.

Pha esis A. Gray v var. comata (Rydb.) Waterf. [-,-,-,2

1 50-73005 P, U; .

Physalis hederifolia A. eae var. -— (A. Gray) Cronquist [3,-

STR, ae 6880’; J, G, P, U; ds, pj, pp, ra

P longifolia [1,-,2,-] D, L; -5400-6100'

JG G; fr.

Physalis subulata Rydb. var. neomexicana (Rydb.) Waterf. ex

Kartesz & Gandhi [1,-,1,-] L, T; 6300-6900’; G, P; pp, ra.

[Physalis foetens Poir. var. neomexicana (Rydb.) Waterf. ex

esz & Gandh

Solanum americanum Mill. [-,-,1,-] D; 6500’; J; fr.

[Solanum nigrum L.]

* Solanum dulcamara L. [1,-,-,-] L; 65255 J; t

Solanum ee M Cav. [4,-,1,2] D, L, R 5550-7000’; J,

—

, U; fr, pj, y

a a on [-,-,3,4] D, L, R, T; 6200-8150"; J, P, U;

S, DJ, Pp.

* solanum ae Rusby var. nitidibaccatum (Bitter)

Edmonds [-,-,1,-} L; 6650; J; ra.

So Pc sarrachoides Sendtner]

* Solanum ptycanthum Dunal ex DC. [1,2,-,-] D; 8050-8900;

J; mm, mr.

x RUE nigrum LJ

Nutt. [1,4,1,6] D, R, T; 5500-8750” J, P, U;

br, ds, fr, mm, pp, ra.

Sparganiaceae

[1,2,-,1] D, R; 8400-10020';

F -t

mr.

Sparganium emersum Rehmann [-,1,-,1] D, R; 8625-9150;

J, U; mr

gle [2,-,2,-] D, L, R; 5400-60007,

,1,3] R;

n Io fal n.a a Inm Li idee dh fT "f£

vem J

Tamaricaceae

*eTamarix chinensis Lour. [12,-,3,3] D, L, R, S; 5300-7200"; J,

, r,

Typhaceae

Typha latifolia L. [6,1,2,1] D, L, R; 5400-9150; J, G, U; fr, ml,

mr.

Ulmaceae

*e lmus pumila L. dnd 1,1,1] D, L, R, S; 5470-8080"; J, G, P, U;

fr, mm, mr, pj, pp, r

Hrtcaceae

Urtica dioica M var. procera (Muhl. ex Willd.) Wedd. 123, 30, 1 12]

D, R; 6600-10100’; J, U; af, br, mc, ml, mm, mr, pp, ra, sf.

[Urtica dioica ssp. gracilis (Aiton) Selander]

Verbenaeeqe

EGG FR y ] Lc ee to Te "* 7]

[DLE RS; 5500- 6900’; 3 G; ds, fr, pj, pp, ra

e pracena Lag. & Rodr. [22,4,2,13] D, L, R, T; 5500-

P, U; br, ds, fr, mc, mm, mr, pg, pj, pp, ra.

ae as L. [1,-,-,-] L; 5650; J; m

ib macdougalii A. Heller [14,1,1 P D, L, R,1;6655-8400";

P, U; af, ds, mm, mr, pj, pp, ra.

Violaceae

Viola adunca J.E. Sm. [10,13,3,6] D, L, R; 6700-10675’; J, U;

af, mc, , mr, sf

Viola canadensis L. qa ls 1,3] D, R; 6300-10330"; J, U; af, br,

mc, mm, mr, pp, r

Viola ee sa [-,-,1,2] D, R; 6600-9800; J, U;

Viola ee da G. Don [-,1,-,-] D; 8500"; J; mr.

[Viola palmata L.]

Vitaceae

Parthenocissus vitacea (Knerr) Hitchc. [5,-,1,1] D, R, T; 5900—

7475’: J, P; fr, mr, ra

Vitis arizonica Engelm. [6,-,2,-] D, L; 5460-6875"; J, G; fr, mr,

Zannichelliaceae

Zannichellia palustris L. [2,2,-,

ml, mr.

Zygophyllaceae

-] D, L; 5600-8620", J, G; fr,

r] D; 5600" J; pj

* Tribulus terrestris L. [5,- d 3] L R S T 5550-6300’; J, G, B U;

ds, fr, pj, ra.

ACKNOWLEDGMENTS

Staff and associates of Bandelier National Monument, Bureau of Land Management (Taos District), Carson

National Forest, Santa Fe National Forest, and Valles Caldera National Preserve are thanked for funding of

the projects, the use of facilities, and assistance in numerous other ways. Key individuals include Craig D.

Allen, Chirre Keckler, Randy McKee, Linus Meyer, and Bob Parmenter.

Assistance in the field was provided by Jonathan Coop, James Fowler, Andrew Hipp, Mark Larson, and

Robert Peters.

T] [T1 s E 3 1 +1 1

Ne TOLOWIIS

] for the identification or verification of specimens: Jennifer Acker-

Reif et al ,H f the T jj tains, New Mexico 959

field, Kelly Allred, Duane Atwood, Mary Barkworth, Karen Clary, Robert D. Dorn, Bill Jennings, Bernadette

Kuhn, Tim Lowrey, Stewart Markow, Bob Sivinski, and Debra Trock. We also thank Tim Hogan and Bob

Sivinski for reviewing the manuscript.

We acknowledge the valuable contribution by the graduate committees that included Greg Brown,

Dave McDonald, and Steve Prager. Alex Buerkle, Larry Schmidt, and Jeff Lockhart also provided assistance.

Graduate students in the MLS. floristics program and other botany students provided invaluable assistance,

support, and inspiration.

Finally, Tim Lowrey, Director of the University of New Mexico Herbarium, and Jane Mygatt, Senior

Collection Manager, provided access to the collection and to the database.

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I 1 .£ al n L In LI Bee S ls F da A!

962 t t f Texas 3(2)

BOOK REVIEW

PauL E. Roturock. 2009. Sedges of Indiana and the Adjacent States: The Non-Carex Species. (ISBN

978-1-883362-14-0, hbk: alk. paper). NE DEINER of Beides, His N one Ave., Indianapolis,

Indiana 46204-2207, U.S.A. (Orders: www.i /special/, 1-317-

205-5440, iaspublications@indy.rr.com). $45.00, 270 pp. 228 color als 3 "n maps, 4 b&w

figures, color range maps for 88 MK 3 appendices, glossary, 6 1/4" x 9 1/4".

1] f ] for 10 minutes or 10 hours,

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WOW! Written by El 1 +1 A I š 1 Ta jc ; thi S iS ea ily "E i351] + + + E

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will be in high demand.

This publication follows a long list of li tural history treatments E Indiana as by the anes Academy of

1 1:54 Lid Ly T: 1 KAZ] LI TOI GrelaA trol Atl na , Dr agonflies

of Indiana by James Curry (2001), Amphibia 1 Reptil fl Indiana by sue! man Minton, a oon. and 1 101 m of iiaia by Marion

Jackson (2004). In the foreword, Indi Acad f Sci Publicati With this visually rich book,

Paul Rothrock, who has studied Cyperaceae for: many years, has provided a tool that will be e assistance to curious amateur

naturalists, as well as to gardeners and natural area and restoration li a This is, essentially, the overriding objective of all our

dones to be attractive, rau ds accurate, "s b co to t ama pas A in as e of study." park

j I I tion. Tl I ] I y commenting, “

beauty, 1 1 ] «I1 T cur E 1 fel [n r1 ] Li] a] Ta 1 n A la | dis and

o Y I D 4 r E

Bau Haus functional desig My clear hope is that the text and pl hat follow adequately capt f thi ld and point

] : mi + ] tono +1 E -l 1 El : [31:1 yl 1.1 11 ra + a I

1 . 11 1 p +1 + +] j > 1 111 Eo 11 ] d ur a ly rl El 1 1 1 e DX D 1 I -l

emphatically p ishes be fully realized. Certainl

i ] i 1 botanists as D re Reznicek, Dr Charles Bs son, Dr. S. Galen Smith, Dr. Gordon Tucker, Dr.

Ci rata and aad AA Another aa is that the author referenced no less than 130 separate publications in providing

"Tl 1 1 1 11 1 i

é r D e

CT ] f. +l 1 1 “11 a ^ JJ f. P" 1 1 ] jp J

n r ES a a Os I QJ! [e] m uo

habitats and ain epeta and aaa of sedge achenes; evolution of sedges; and even short sections on the economic

dditinn) tri t tal 1 ] 1 hi T hi ] f adantation of sedoes to

? r i o! X a

EE soils, iud Es pe C, metabolism

Certainly f the highlig book is that Dr. Rothrocl 1 tl is of conservatism" mim dd de E a &

1070 /nDl " f +1 / s: n : A Cee, Sen AA 11; NU

G. Wilhelm in 1 j go Region, 3rd. ed., tum, Lisle, g

Indiana D ] +l + 1 ee 1 L +1 ] I = 11 I 3 f. * 1 i + For

e I r 4 I

J 1 = 1 1 T CEPR PS A J P ] ] 1 = 1 1 "m nA

e e e O

" mi + att +t aca 1 S x 1 z] ] f. > 1

Species with coefficients of ism values of 8-10 receive Em Highest eine cae those of 0—1 receive the lowest. it should be

alarming and a concern to all read f this book that of the 88 Hu pee idi by HE Aue puc 5096 have coefficients

Pelea 8 and 10. I Hopefully by higl lig! ti E the | igl g p which

the ey e pens the author will g furti pr tt tect , maintain, 1 +1 laah] 1

provided in the introduction is a family description, a key to the genera a Cyperaceae in Indiana, and black and white il-

lustrati f tati h of each genus as well as the achenes of all known species of Fleocharis, Rhynchospora, and Scleria

documented for ithe state. Not onl illustrati but they are drawn to parati le and col gt d

further cad iun a cross section.

173 1] + T 13 + -l hh 1; " * g

bid o eii Wu, Ae E Re liL.1X71 ? 5 > 5

E Re genera and 88 AS E — tl tand heart of the book. To increase the utility of the book, the author

r7 1 f. 1: 1 l 1 ] 4 Y: +l T: E Stat

ALIAS LIE iefly NA IA LA ut o Mir

of dti à Mn Ohio, MEM and/or Wisconsin:

cl Ll a ant Klasse | à 4 A 11 1 1 ea |

Taxononr p | y I

that t ity into the followi tegories: 1) occasional to common and verified by herbarium voucher, 2)

(continued on page 968)

J. Bot. Res. Inst. Texas 3(2): 962. 2009

BERBERIS REPENS (BERBERIDACEAE) NEW AGAIN TO THE

FLORA OF PENNSYLVANIA

Jerry G. Chmielewski

Paes Rock University

Dep f Biology

didi EL HA i 16057, U.S.A.

erry.chmielewski@sru.edu

ABSTRACT

Berberis de a pres iously documented species from Pennsylvania, thougl ly an oo pue from the state record,

This species ] from t] gi dlot in Butler County in western

Peris Ivana.

RESUMEN

Berberis repens, una especi te d tada de Pennsylvania, reyó una especie excluida del estado,

se cita de una población ia Esta especie se colectó en los márgenes de un bosquete en el condado de Butler en el oeste E

Pennsylvania.

Berberi Lindl ing bar! ,isanative, evergreen, low growing, perennial shrub of western North

America that is moy cultivated. though has become naturalized in parts of eastern North America.

Typically upright to decumbent, the solitary stems which arise from rhizomes form colonies that are less

than 40 cm tall. The inner bark is distinctly yellow. The alternate, holly-like leaves are petiolate and pin-

nately compound with 3—7 oval, spiny-toothed (6—14 on each side), bluish-green, leathery, leaflets which

are minutely pappilose on the underside. These turn purplish in winter. Deep yellow, fragrant flowers occur

in terminal, drooping racemes from May-June. These in turn are followed by grape-like, 1-4 seeded, dark

bluish-purple berries which ripen from June-July (Bailey 1949; Whittemore 1997; Harris 2009; USDA For-

est Service 2009). Berberis repens differs from Berberis aquifolium Pursh, with which it hybridizes when the

two grow in proximity to each other, by the latter's greater height (to 2+ m), smooth underside of leaflets,

number of leaflets (5-9) per compound leaf, and the number of teeth (10+ on each side) on leaflets (Bailey

1949; Whittemore 1997).

The native range of B. repens is the western U.S. east to Texas and Minnesota inn British SOUS and

Alberta in western Canada (Whittemore 1997). Further, the species in

Ontario, Indiana, and Pennsylvania in eastern North Acer (USDA, NRCS 2009). Although commonly

cited from a site in Northampton Co., PA based on a 1946 collection (Wherry et al. 1973; Rhoads & Klein

1993) this specimen likely represents a planting that persisted for a period of time after being cultivated,

but never truly became established as part of the Pennsylvania flora (Grund, pers. comm.). For this reason

the species was excluded (Block pers. comm.) from the most recent publication of the Pennsylvania flora

(Rhoads and Block 2000).

A naturalized population of B. repens was observed in a woodlot adjacent to the water tower on the

grounds of the Slippery Rock University campus during the summer of 2008. The population consists of

four clustered, relatively young shoots, two of which were in flower, the others vegetative only. No fruits

were observed later in the growing season. This said, immature fruits were present in mid-June of the cur-

rent year. Based on the number o: leaves alone the plants are presumed to be about 5-6 years old. Although

I am unaware of a nearby cultivated seed source, presumably these shoots are a consequence of a bird

introduction from a more or less locally cultivated specimen. The most commonly associated taxa included

Impatiens capensis Meerb., Liriodendron tulipifera L., Pilea pumila (L.) A. Gray, Rubus allegheniensis Porter, and

J. Bot. Res. Inst. Texas 3(2): 963 — 964. 2009

964 | t tanical h Institute of Texas 3(2)

Toxicodendron radicans (L.) Kuntze. Because the plants are still small only a single leaf was harvested in late

winter.

Voucher specimen: PENNSYLVANIA. Butler Co.: Slippery Rock University campus, woodlot adjacent to water tower, 3 Mar 2009,

Chmielewski 3205 (SLRO).

ACKNOWLEDGMENTS

Both Steve Grund and Timothy Block are thanked for their insights into the Pennsylvania history of the

Berberis repens collection from Northampton Co. Allison Cusick and Timothy Block are thanked for their

helpful reviews of the manuscript.

REFERENCES

Batey, L.H. 1949. Manual of cultivated plant t lyg in the continental United Stat | Canada.

The MacMillan Co., New York.

Harrison, S. 2009. Mahonia repens. (http:// k.ca/agriculture/plantsci/classes, ‘range/berberis.html). Ac-

cessed 30 January 2009.

RHOADS, A.F. AND T.A. Bock. 2000. The plants of Pennsylvania: an illustrated manual. University of Pennsylvania

Press, Philadelphia, Pennsylvania.

RHOADS, A.F. AND W.M. KLEIN, JR. 1993. The vascular flora of P ylvani tated checklist and atlas. American

Philosophical Society, Philadelphia, Pennsylvania.

USDA, FOREST SERVICE. 2009. Mahonia repens (Lindl.) G. Don. (httl:// fs.fed.us/global/iitf/pdf/shrut

honia9620repens.pdf). Accessed 30 January 2009.

USDA, NRCS. 2009. The PLANTS database. National Plant Data Center, Baton Rouge, LA. (http://plants.usda.gov/).

Accessed 30 January 2009.

Wherry, E.T., J.M. Foce, Jr, ano H.E. Wahu. 1979. Atlas ofthe flora of Pennsylvania. Morris Arboretum of the University

of Pennsylvania, Philadelphia, Pennsylvania.

WHITTEMORE, AT. 1997. Berberis. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America

North of Mexico. 12+ vols. New York and Oxford. 3:276-286.

BOUTELOUA CURTIPENDULA VAR. TENUIS (POACEAE: CYNODONTEAE),

NEW TO TEXAS AND THE UNITED STATES AND

A KEY TO B. CURTIPENDULA VARIETIES

Stephan L. Hatch Maria Andrea Tomas

aM . Tracy melanie ( m EE Rafaela, INTA

| lanag t Ruta 34, km 2

as A&M University CC 22, 2300, Rafaela, Sante Fe

College " wn Texas 77843-2138, USA. ARGENTINA

s-hatch@tamu.edu matomas@rafaela.inta.gov.ar

Steven R. Archer

School of Natural Resources

niversity of Arizona

Tuscon, Arizona 85721-0043, U.S.A.

sarcher@ag.arizona.edu

ABSTRACT

Th f Bouteloua curtipendula (Michx.) Torr. var. tenuis oes 2 ens is di new to mee and the Med States. A

key to B tit lul ieties in the U.S.A. is provided, al ng g Į g

RESUMEN

Se reporta la ccurrencia de Bouteloua curtipendula (Michx.) Torr. var. tenuis Gould & Kapadia, nueva para Texas y los Estados Unidos.

Se provee una clave para las variedades de Boutel tij l E.E.U.U. y fotografías digital trando caracteristicas que las

distinguen.

Prior to this report the occurrence of Bouteloua curtipendula (Michx.) Torr. var. tenuis Gould & Kapadia

(CREEPING SIDEOATS GRAMA) in naturalized populations has not been recorded in Texas and the United States

(Gould & Kapadia 1964; Gould 1979; Hatch et al. 1990; Allred 1993; Powell 1994; Jones at al. 1997; Wipff

2003; Hatch 2008; Flora of Texas Database 2008). Specimens of this taxon were collected in July 2001 but

were not identified for a period of time because var. tenuis had not been described in botanical treatments

of Texas or U.S.A. grasses.

This grass was identified using Gould and Kapadia (1964) and verified using specimens in the Tracy

Herbarium (TAES) from Mexico i included de ope Moe 200 D

mex inmmtn e] Aaral lly Specimens

The three varieties of B

with intact bases showing either a caespitose ones habit. or rhizomes, or stolons are critical to the

identification process. The rhizomes must e more than short oe phort to that pm acturi Decamiitlg an

erect stem. The varietal diff ted in the key below and di logical features are

depicted in Figs. 1, 2, and 3. It is likely that var. tenuis in Texas established iom. seed diepasd by livestock

(e.g., cattle, sheep, or goats) from populations in northern Mexico. A review of autecology of B. curtipendula

can be found in Tomas (2004).

KEY TO BOUTELOUA CURTIPENDULA VARIETIES IN THE UNITED STATES

1. Plants caespitose or rarely with short knotty rhizomes B tipendula var. caespitosa

1. Plant stoloniferous or with long rhizomes.

2. Plants stclonif _ thi | t or absent B i lula var. tenuis

2. Plants with long rhizomes B. curtipendula var. curtipendula

J. Bot. Res. Inst. Texas 3(2): 965 — 967. 2009

966 J t tani i Texas 3(2)

Fic. 1. Dig g

var. caespitosa,

Fic. 2. Digital i ftl lant |

curtipendula. Note the rhizomatous specimen base.

I/D} al | a I £

Fi 3 ee TN B fin VE

d 2 X f

n E ri

one curtipendula (Michx.) Torr. var. tenuis Gould & Kapadia, Gould & Kapadia, Brittonia 16:201-203.

1964. CREEPING SIDEOATS GRAMA.

Stoloniferous perennials. Culms ascending, slender rhizomes absent from Texas specimens (see discussion

in Gould & Kapadia 1964). Leaves with long, thin but broad (3-5.5 mm wide) leaf blades. Panicles of 16-35

spicate primary branches; branches 1-3.5 cm long with 4-9 widely spaced spikelets. Spikelets 3.4-8 mm

long, florets 2-4, apical ones rudimentary, rachilla not extending beyond upper floret, rudiment 3-awned;

glumes 2, l-veined, unequal, shorter or longer than first floret; lowermost lemmas 2.9-6.5 mm long, 3-veined,

acute or trifid, mucro minute; lowermost paleas present. Flowering June to October.

I | ined: UNITED STATES: TEXAS. Sutton Co.: TAES Sonora pi Mad Vans zone TX G1 E N; pi :

WIN LLE? SEL

28W ith B. curti ipendi ila var. caespitosa, B curtipendula Var curtipendula, N

Tomas s.n. (TAES). MEXICO. Zacatecas: 10 mi NW di Eonibreiere: 30 Oct 1959, Gould 9000 (holotype) (TAES).

REFERENCES

ALLRED, K.W. 1993. A field guide to the grasses of New Mexico. New Mexico State University. New Mexico Agri-

cultural Experiment Station. Las Cruces.

FLORA OF Texas DATABASE. 2008. University of Texas Herbarium, Plant Resources Center, Austin. < http://www.biosci.

utexas.edu/prc/Tex.html»

Goutp, FW. 1979. The genus Bouteloua (Poaceae). Ann. Missouri Bot. Gard. 66:348-416.

GouLo, FW. AND Z.J. KaPADIA. 1964. Biosystematic studies in the Bouteloua curtipendula complex Il. Taxonomy.

Brittonia 16:182-207.

HarcH, S.L., KN. GANDHI, AND L.E. Brown. 1990. Checklist of the vascular plants of Texas. Texas Agricultural Experi-

ment Station, College Station. MP 1655.

HarcH, S.L. 2008. Checklist of Texas grasses. <http://botany.csdl.tamu.edu/FLORA/taes/tracy/610/Checklist2008.

pdf»

Jones, S.D., J.K. Wierr, AND PM. Montaomery. 1997. Vascular plants of Texas - a comprehensive checklist including

synonymy, bibliography and index. University of Texas Press, Austin.

PoweLL, A.M. 1994. Grasses of the Trans-Pecos and adjacent areas. University of Texas Press, Austin.

Hatch et al., Boutel til lul tenuis, new to Texas 067

Tomas, A. 2004. Ecological implicati f genetic variation in Boutel tipendula (Michx.) Torr. Ph.D. Disserta-

tion, Texas A&M University.

Wir, J.K. 2003. Bouteloua. In: Barkworth, M.E., K.M. Capels, S. Long, and M.B. Piep eds. Magnoliophyta: Com-

melinidae (inpart): Poaceae, part 2. Flora of North America North of Mexico, volume 25. Oxford University

Press, New York. Pp 250-269.

968 J laf tha Ratanical R Tn

ET Siar

lexas 3(2)

BOOK REVIEW

(continued from page 962)

reported by Deam (1940: Flora of Indiana, Indiana Department of C tion, Division of Forestry, Indianapolis),

not ds as state rare, Mo Or Hence and USE state A rare, threatened or penu Additionally, dis over status

j t sd is a fantastic approach because it

TInt A

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p E 1 a is ] FECE nil 1 t SEA E 1 ES 1:57 1

ates; wetness s category DAN Lie U.S. Fish and Wildlife Service's designations Reed, P., Jr. 1988. National List of Plant Species That

Occur in W 3) Biol. Rep. 88 (26.3), U.S. Fish and Aida Service, Washington, D.C.]; a bar graph E

] fruiti iod d ol t natural history, etc. O

I J or , L 1 E ,

4. f, 1 ] 1 1 1 1 1 f: 1 s el L r2

1 6 e

: el; A E | ] l1; J fi] 11 1 " > a +l AE ] f, 1 1

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genera. Whil hard t ist y be ee to this approach, it iw be welcomed by amateurs or those who are not as

"esa | 1 i 7 orm 4] ] AA : 1 pa | “+l

familiar Į PE 1-4 sharp photo-

graphs of habit, rhizomes, infloresence, and/or achenes

r sedge enthusiasts who have ever struggled with pairs of species that are often difficult to distinguish, the author provides

exhaustive and refreshing accounts helpful in separating the following: Cyperus bipartitus vs. C. diandrus, Eleocharis flavescens vs. E.

geniculata, Eleocharis erythropoda vs. E. palustris, Lipocarpha drummondii vs. L. micrantha, Schoenoplectus purshianus vs. S. smithii, Scirpus

cyperinus vs. S. pce alts an nee muehlenbergia vs. S. reticularis.

] One in particular

is Pu favorite: in his account on Rs ! irpoides, Dr. Rothrocl 1 that f I ist John Torrey completed his 500-page

Trg: 14711 "EI f 1 Tiatia J Cig ge

Ther re are e ee appendices A A is heck of dim non- "m ical — known from Papam, each with its

t ?

pp or perennial t, , and its gory. Apt g species

A ] q 1 T 15 1 1 f. 1 1 ] T J]: +l

rr Qo 4 L Pl O D e

While this Lila yl li i ] ili J for Sta DALE, d d dg E t 3 it will likely | ful f ] ] | [ gi

on sedge identification.

e with every book review yen should og find thing to quibble about? This book is so well written, it is extremely

Apps f. 1 T 1 "m On pace 21 el Fa 1.5 " le : £l

L i=] D + “ff

laris have incorrect endings of “a “a” and * ta, respectively It would have | helpful ld I ] hyma, ec

karyotype, monophyletic, and phytolith, etc. to the glossary. hnical ill be i liatel izable by tl

J a E Rn e UR

or untrained Bonus Had IM |i SANGRE time and ADI some extra RC AC of the 40 non-Carex species known con

ode: states would h l g get enough. But a book can only cover so

1 S G 1 11 +1 "E £ A Pe | Era Je Jl Ad a =

y even g. A copy ges of i The Non-Carex

1 14 1 +1 DATE A | YE + A 411 A] nthi act A] r

g 1 , reg Į Given the quality of

this book, we anxiously look forward to the author’s i of the Indian ies of Ca Paul M. McKenzie, Ph.D.,

Va L AUL

Endangered Species Coordinator, U.S. Fish and Wildlife ROM 101 Park DeVille Dr.; Suite A, DE Missouri 65203-0057, U.S.A.

J. Bot. Res. Inst. Texas 3(2): 968. 2009

POLYGALA CYMOSA (POLYGONACEAE) NEW TO TEXAS

Michelle Pollard Jason R. Singhurst

Department of Biology Wildlife n Program

aylor University Texas Parks and Wildlife Department

Waco, Texas 76798-7388, U.S.A. 4200 Smith ed Road

S.A.

jason.singhurstetpwd.state.tx

Walter C. Holmes

Department of Biolog

walter - holmesabaylor.edu

ABSTRACT

Polygala cymosa Walter is reported as new to Texas.

RESUMEN

Se cita Polygala cymosa Walter como nueva para Texas.

The genus Polygala, commonly known as milkwort, includes about 550 species of worldwide distribution.

1 1 ] x nm Ih] : ] ] - el. f] 4 : "EC +1 1 1

I 8 ) E

Of the 26 species reported in Texas, ten species occur in the Pineywoods of East Texas.

Herbarium studies have resulted in the discovery of a specimen of Polygala cymosa Walter from the

Pineywoods, which is herein reported as new to the state.

Voucher specimen: TEXAS. Jasper Co.: 7 mi S of Kirbyville, ca. 2 mi E of Hwy 96, ca. 0.2 mi N of end of unpaved road near Nichols

Creek, wooded uplands, 17 Oct 1977, Ajilvsgi 5505 (SMU).

Polygala cymosa, commonly known as tall pinebarren milkwort, is a plant of 40-120 cm in height. The flow-

ers are lemon-yellow when fresh, and arise from branches emanating from a solitary stem. The linear basal

leaves narrow distally and are 4-7 cm long. The flowers turn dark green upon drying. The species is known

to occur in Alabama, Delaware, Florida, Georgia, Louisiana, Maryland, Mississippi, North Carolina, and

South Carolina (USDA, NRCS 2009). Because this is the only known site of P. cymosa in Texas, and there is

only one known site in western Louisiana (Thomas and Allen 1998), the species is considered to be a rare

peripheral in the West Gulf Coastal Plain. The species is typically found in wet pinelands, swamps and

ditches. The Texas site contains longleaf pine wetland savannas, slightly drier sandy clay upland longleaf

pine savannas, and seepage bogs with Sarracenia alata.

Polygala nana and P. ramosa are the two other species of Polygala in East Texas with yellow perianth.

The following key may be used to distinguish among the three species.

KEY TO THE YELLOW FLOWERED SPECIES OF POLYGALA OF THE PINEYWOODS OF TEXAS

[ADAPTED FROM CORRELL AND JOHNSTON (1970) AND RADFORD ET AL. (1968)]

—À

. Plants 3-15 cm tall, o to semiscapose; racemes solitary at tips of stems, capitate to thick-cylindric,

about as long as wide (9-2 P. nana

1. Plants 10-120 cm tall, st | ie leafy, numerous, cymose, each usually about 2x longer than

wide, in ag te the infl (20-)50-150 mm wide.

2. Plants usually Greater than 30 cm tall basal leaves linear to linear-lanceolate, 3-7 cm long, stem leaves

linear to filiform, gradually reduced above P. cymosa

J. Bot. Res. Inst. Texas 3(2): 969 — 970. 2009

970 Jou the Botanical h Inst Texas 3(2)

2. Plants to 30 Mb T Mii tillate to 25cm lona. st

L I T J wwf

. Fidi UW a sy ¿>|

not reduced above P. ramosa

Following are brief remarks on the distribution and characteristics of the species mentioned in the key.

Information on Polygala cymosa is presented above.

Polygala nana (Michx.) DC. Southeast Texas (Angelina, Hardin, Jasper, Jefferson, Newton, Sabine,

Tyler cos.); United States: Southern North Carolina to Arkansas, south to Florida to Texas. The species is

well-marked by the single head-like raceme at the tip of scapelike stems and the licorice odor of the roots

(earning it the common name of candy-root).

Polygala ramosa Elliott. Southeast Texas (Angelina, Chambers, Hardin, Houston, Jasper, Jefferson,

Liberty, Newton, Orange, San Augustine, and Tyler cos.); United States: Coastal plain from Cape May Co.,

New Jersey, south to Florida, west to Texas. The species is characterized by the amply leaved stems, wide

basal leaves, and height to about 30 cm.

ACKNOWLEDGMENTS

We wish to thank the curator of BRIT for access to the specimens that made this study possible.

REFERENCES

Corretl, D.S. anp M.C. JoHNston. 1970. Manual of the vascular plants of Texas. Texas Research Foundation,

Renner.

Reororo, A.E, H.E. Ahues, AND C.R. Bett. 1964, Manual of the vascular flora of the Carolinas. University of North

Carolina Press, Chapel Hill.

Tuomas, RD. AND C.A. ALLEN. 1998. Atlas of the vascular flora of Louisiana. Vol. Ill. Dicotyledons. Fabaceae-Zygo-

phyllaceae. Louisiana Department of Wildlife and Fisheries, Baton Rouge.

USDA, NRCS. 2009. The PLANTS database (http://plants.usda.gov/plants). National Plant Data Center, Baton

Rouge, LA 78074-4490 USA.

ASSESSMENT OF INVASIVENESS AND ECOLOGICAL IMPACT

IN NON-NATIVE PLANTS OF TEXAS

Guy L. Nesom

2925 Hartwood Drive

Fort Worth, Texas 76109, U.S.A,

guynesom@sbcglobal.net

ABSTRACT

itside of cultivation in Texas has been developed. About 300 of

nl t+nf Qld el

An dnm

since the 1970 vublication ol the Maral of the Vascular duin sid m and n of the latter taxa ES m

these have ]

oe by at leat a literature renee 2 unen cde In

The Index is based on knowledge of the Puedes

from field, herbarium, "E literature, ing ing criteri

d and natural habitats, negatively affecting native species tural biodiversity ll arene native

S ora : À . . KAST e

11 As

F1: Invasive in both

J1 3 ROA "

1 o i

E J , O

O 4 Ir o 4 [m]

cant economic damage; including woody, herbaceous, and aquatic species.

Abundant in number and widespread. ie invasive in disturbed habitats, much less commonly in natural habitats:

F2:

subdivided into woody, herbaceous, and aquatic spec

F3: Relatively few in number, known from ne ely few localities, usually in disturbed habitats; subdivided into woody and

herbaceous species.

F4: Status unknown.

of species per category are Fl-Woody, Herbaceous, and Aquatic (51), F2-Woody (13), F2-Herbaceous (228), F2-

T pin pude (76), F3-Herbaceous (348), and F4 (80). A Watch List includes 52 woody, herbaceous, and aquatic species

most likely to warrant F1 ranking.

RESUMEN

S li inf d tado de 812 esp tivas que crecen fuera de cultivo en Texas. Unas 300 de ellas se han citado

A del HI; 14^ 10760 ol M 1 Aida WU, D A 4 f ray Pa 1 J En 1

I V af Tess

una referencia bibliográfica. Un ' Fundamental I Invasiveness s Index" f para | luación y ord ión de cad de |

a PE ] ] Ki i cl A; +A I 4 ] H : + Ao] 1 ]

I E i C

campo, herbario, y bibliografía, d do a los siguient

EI T : ls. flee 1:f: J 1 E 4 2: s Bs 1 Ts J: LER | ] " 1

4 L [e] Fu Ir

13. 1 1 1 1. :1 1 L i 1 1 z 1 1 lc E

j im] ; incluyend I lef herbáceas, y acuáticas

F2: Abundantes en nümero y extendidas, g i I lterad I habitat ] I

]

divididas en especies lefiosas, herbáceas, y acuáticas.

BR. Dalat; : z - 3

en especies lenosas y herbáceas.

: Estatus desconocido.

Los números de especies por categoría son F1-leñosas, ma, y acuáticas pel e > (13), F2-herbaceas (228),

erbáceas, y acuáticas

F2-acuáticas (16) oe ~ F3- ADNE (348), y F4 (80). 1 y Į

1 1.1

About 516 non-native species were known 40 years ago to occur outside of cultivation in Texas (as counted

from Correll & Johnston 1970)— about 11 percent of the total flora of the state at that time. Since that time

about 300 additional introductions to the state have been reported in various publications (summarized

in the PLANTS Database: USDA, NRCS 2009; the PLANTS Database has drawn its information from a

variety of sources, including major publications on the Texas flora). The present paper provides a summary

account of the currently known non-native Texas flora, with documentation for those species included in

J. Bot. Res. Inst. Texas 3{2): 971 — 991. 2009

|l £a n.a a In LI dh dion ck ET ae 20/70

972

the account since 1970 and a ranking of invasiveness and ecological impact for each species included. The

system presented here is simple enough to allow a quick assessment of a large number of species but still to

provide a fundamental overview of each species.

Various lists of invasive inh have been developed for Texas, but documentation for these lists is not

readily available. The most list is presented by TexasInvasives.org (2009), which includes 139

species, with a photo for each nd a listing of the family, common name, duration, habit, and information

on morphology, biology, and distribution, including links to detailed Texas maps. Two lists have state-wide

legal standing: the Noxious on List of the Texas Department of Agriculture (TDA 2009) with 29 species

and a list of Prohibited Exoti ies from the Texas Parks and Wildlife Department (TPWD 2009) with 19

aquatic species. The list of 13 Bolle species from the North Texas Water Garden Society (NTWGS 2009)

appears to be a subset of the TDA list. The City of Austin lists 26 species (some listed simply as the genus)

of Invasive Plants to Avoid (Austin City Connection 2009).

The Invaders of Texas (2009) program organizes teams of local citizen scientists who seek out and

report occurrences of invasive species. This is a very useful program, both practically and scientifically—as

of May 2009, more than 7400 detailed field observations have been reported and are available online. Each

field report includes precise locality information (with coordinates automatically mapped), an indication of

abundance at the locality, and usually a photo of the plant or populations taken at the site of observation.

The target species sought by the Invaders of Texas participants are the 139 on the TexasInvasives.org list

(2009). For each species, individual field observations are mapped in aggregate to show their total distribu-

tion in the state.

Non-native species in Texas—development of a list and documentation.

A preliminary list of non-native species in Texas was developed by the author from the PLANTS Database

(USDA, NRCS 2009), using an Advanced Search for Texas taxa introduced to the Lower 48 states. Journals

and other literature were reviewed for possible additions, and a number of species have been excluded.

Identifications in the Texas list presented here often are not made to infraspecific rank (in contrast to the

PLANTS list), because vouchers commonly have not been examined or because the validity of the infraspe-

cific taxonomy is not clear. Only plants that are naturalized have been included (vs. cultivated or persisting

from cultivation), though some on the list probably would best be characterized as waifs (Nesom 2000) if

follow-up observations were done at sites where they were reported to occur. Whether or not a species is

native sometimes may be difficult to judge—generally such instances are discussed in the documentation

for the full list (Nesom 2009a).

Documentation for the occurrence of these species in Texas (Nesom 2009a) begins with literature and

other records that have been published mostly after 1970, the date of publication of Correll and Johnston's

“Manual of the Vascular Plants of Texas,” which remains the primary floristic resource for the state. In the

course of assigning invasiveness rankings, however, the rationale for inclusion of each species on the full

list has been examined. For species included in the documentation, at least one full literature reference is

provided, with reported Texas counties and other brief notes on the status of the species, as appropriate

or as known. Many taxa on the PLANTS list or from elsewhere have been excluded from the Texas flora

because of mistaken or faulty documentation (Nesom 2009b for examples from the Asteraceae), and these

are specifically noted in the commentaries (Nesom 2009a). A number of species have been reported simply

by listing (Johnston 1990; Hatch et al. 1990) or by mapping (Turner et al. 2005); vouchers or other docu-

mentation for these are being provided.

Documentation for the full list of non-native taxa is an essential and critical part of both this process

and gus report. ‘aus the information is complex and n is and NL en dE are ongoing, it

(Nesom 20092). Addition of ] tation for

species first reported in lists by Johnston (1990) and Hatch et al. (1990) and elsewhere will be a significant

advance.

Nesom, pP J ani’ 2 a [| £ "E . ET 973

Systems for assessment and ranking of invasiveness.

In developing a protocol for evaluating Texas non-native species for invasiveness and ecological impact, a

representative set of other systems toward the same end has been considered and studied.

* Alien Plants Ranking System (Ver. 5.1) (APRS Implementation Team 2000)

* Ranking Invasive Exotic Plant Species in Virginia (Heffernan et al. 2

* Criteria tor Categorizing Invasive Non-Native Plants that Threaten Wildlands (Warner et al. 2003)

* An Invasive Species Assessment Protocol: Evaluating Non-native Plants for Their Impact on Biodi-

versity (Morse et al. 2004)

* Invasiveness Ranking System for Non-native Plants of Alaska (Carlson et al. 2008)

i New b State Plant Ranking System for Evaluating Non-Native Plant Species for Invasiveness

RAFT) (Jordan et al. 2009).

Perhaps the most widely applied (and adapted) of these is Morse et al. (2004), developed by NatureServe,

in collaboration with The Nature Conservancy and the U.S. National Park Service. It is a “protocol for as-

sessing and categorizing non-native plants according to their impacts on native biodiversity.” NatureServe

Explorer now includes an impact rank CI-rank"—high, medium, low, or insignificant) for many non-native

plant species of the USA.

tha a + I + ] Jus f H Ja : : 1 , Faras A

In each of s of detailed

— grouped into a set of topics. usually inc utis. some or all of Mee

* ecological impact and potential for future impact;

* biological attributes, including dispersal ability and invasiveness potential;

* ecological amplitude;

* geographic j. of invasion; and

* control/management feasibility and cost.

A significant amount of the information required for the ranking criteria is drawn from literature pertinent

to the species under consideration. After scoring is complete for the whole set of questions, the species may

be ranked numerically by its additive score, or the additive score may translate to a broader category.

The ranking protocol for the Tennessee Invasive Exotic Plant List (Tennessee EPPC 2001) is consider-

ably different from those above. In the Tennessee system, a species is assigned by a committee of biologists

to one of the following categories (the number of Tennessee "peers in each category is danos

1 + EN LEER AAA A

Rank 1—Severe Threat: Exotic piant species that p i i I y

into native plant communities and displace native vegetation. e speci

Rank 2—Significant Threat: Exotic plant species that possess dc A ofi invasive species but are not

mo A to spread as easily into native plant communities as those species listed as Rank

1. (49 s

Rank 3 Lesser Iet Exotic plant species that spread in or near disturbed areas; and are not presently

considered a threat to native plant communities. (28 species)

Watch List A—Exotic plants that naturalize and may become a v dm in the future; includes species that

are or could become widespread in Tennessee. At this time more information is needed, and there is

no consensus about their status. (24 species)

Watch List B—Exotic plant species that I j ding states but have not been reported

in Tennessee. (6 species)

The protocol proposed here for Texas non-native ies is very similar to the one developed for Tennessee.

Invasiveness and ecological impact of Texas non-native species.

The system outlined here for use in Texas emphasizes simplicity, allowing assessment of the large number

of non-native species (all that are known to occur in the state). Because essential elements of biology and

geography are included in the assessments, the system is termed the Fundamental Invasiveness Index. After

becoming familiar with the categories of the index and the characteristics of the species under consideration,

the species is assigned to one of the categories. The first category (F1) applies to species that have invasive

biological characteristics as well as a strong and adverse impact on natural systems and biodiversity (or an

adverse economic impact). The other two main categories (F2, F3) include species that are less significant

[| [| £ al D.s . in LI LF Y ds A!

974 Jo f Texas 3(2)

in ecological or economic impact, with the division between species of F2 and F3 made on the basis of

their geographic distribution, abundance, and potential for spread. Within F2 and within F3, slightly dif-

ferent characterizations are given for woody and herl us species, as the two growth forms are generally

distinct in biology.

The biology of aquatic species is distinct from terrestrial ones, in part because of their potential for

extremely rapid dispersal in local aquatic systems. Emergents that grow in open water—sometimes per-

sisting in mud flats or in periodically wet habitats (i.e., Alternanthera, Ludwigia, Nasturtium)—are included

as aquatics. Each species has the potential to cause major ecosystem damage, and many of them already

are recognized as noxious over wide regions; others are ecologically damaging at least locally in the lake

or stream where they occur. Somewhat arbitrarily, aquatic species are recognized here in the category of

maxinum negative effect if they occur in 10 or more counties (as recorded by herbarium voucher specimens,

expert sightings or photos, or other reliable documentation). Hydrilla verticillata is vouchered by specimens

at TEX for only about six Texas counties but clearly occurs more widely (as do some of the other species).

Otherwise, aquatic species known to occur in 9 or fewer counties all are placed in the same category.

A ranking system such as that from NatureServe provides assessments based on array of explicit and

detailed criteria. For Texas and its large number of non-native species, contemplation of such an intricate

system and the massive amount of time required for its completion perhaps has inhibited even the begin-

ning of one. The system here, however, appears to provide a realistic overview of the situation in Texas,

even though many assessments remain to be clarified for species currently included. Further, the current

account at least provides a basis from which more detailed assessments may be developed.

For accurate evaluation of an individual species, knowledge is required of the following.

* Nativity. Is the species native or non-native?

* Approximate date of introduction in Texas (e.g., pre-1970, 1970s, 1980s, 1990s). Documentation

in the current system does not provide specific information for species introduced before 1970 (those in-

cluded in Correll & Johnston’s Manual of the Vascular Plants of Texas), but for species recorded since that

time, dates are evident, as much as possible, in a review posted and periodically updated on the internet

(Nesom 2009a).

* Current geographic distribution. Based on distribution maps in Turner et al. (2003), distribution

maps generated by data from Invaders of Texas (2009), records from herbaria (primarily those in Texas),

and literature.

* Ecological/reproductive behavior in Texas and in other regions. Based on field experience of the

año ana others, alain literature, information from herbarium collections.

(aquatic or terrestrial, herbaceous or woody). Based on field experi-

ence, published literature, information from herbarium collections.

FUNDAMENTAL INVASIVENESS INDEX

Fl—Invasive in both disturbed and natural habitats. Negatively affecti ti i natura l biodiversity

e 4 e Pu

by alteri ti tation and habitats or by out peti hybridizing w ith native species; or, invasive

into ciel as and causing P n economic damage. Aquatic species lotown to occur in 10 or

more counties. Woody, herbaceous, d i poles Arundo donax, Bot! hl

songarica, Centaurea melitensis, Fichhornia ene Ligustrum sinense, Lonicera a Lygodium japonicum,

Nandina domestica, Salvinia molesta, Sorghum halepense, Triadica (Sapium) sebifera, Ulmus pumila.

F2—Abundant in number and widespread, commonly invasive in disturbed habitats, much less commonly

in anes habitats (Table 1).

Woody—Trees, shrubs, subshrubs, and woody vines. Abundant i | 1 widespread, 5 invasive i bed habitat

such as roadsides, fencerows, woods edges, and others, i potenti lly or incipientl y | ] tural habitats (as Fl).

mples: Albizia julibrissin, Morus alba, Nicotiana glauca, Ponci 1 if liat pu stad? cla bns agnus- -castus.

F2-Herbaceous—Annual and ial herbs and grasses. Known invasi y expanding in geograt ge, primarily in

k| j J 11 1 1 4 fe A

lawns, roadsides, and other. open, disturbed babies. i i I , Wi p

Nesom, Fundamental invasi index f ti ies of Texas 975

TasLE 1. Numbers of non-native species in Texas ranked as F1, F2, F3, and F4. The total is 812 species.

Woody Herbaceous Aquatic TOTAL

F1 17 25 9 51

F2 15 228 16 258

F3 76 348 424

F4 0 80 80

ing TANER ae biomass. Not significantly affecti lve species or natural biodiversity or otherwise strongly ua: native

xamples: Ávena sativa ‘Bellardia trixago, Coronilla varia, Daucus carota, Duchesnia indica, E

Galium aparine, Gamochaeta coarctata, Hypoch Lamium amplexicaule, Medicago lupulina, Stachys floridana, Stellaria

media, Md Te Torilis dea Trifolium repens, Verne arvensis, Vicia sativa.

F2-Aquatic 1 to 9 counties

Bode DX AAA. E

odd few in number, known from ic ED lew pocas, usually it in disturbed naitala,

‘oody—Trees, shrubs, subshrubs, and woody vi

gcn repeatedly Med Or perhaps merely | gt isti g t localities t sl g agg y tend E

peeps pid s Examples: Ardisia crenata, Buddl ja lindleyana, Cinna phora, Hibiscus syriacus, Koelreuteria

cea h Photini tifolia, Pistacia chinensis, Py tha koidz i, Pyrus communis, R i offi inali

E2_Herharenne A ] J 14) ] J 1 rn uo f Je: - 1 Fr. 1 E 1.

AE e A A E r ue Pi Is E 3

usually in disturbed Habita p showing ae or no increase of abundance or g hi "RE

Sas pales mS , Phy frat , Plantag Ori in some ae sporadically appears

" 1 DM » a

, Ipomoea batatas, Luffa aegyptiaca, Narcis-

sus pseuicnateissus Tagetes erecta.

F4—Status unknown.

Watch list: terrestrial non-native species in Texas potentially ranked as F1.

Many of the non-native species in Texas occur only in small areas. Among these are a significant number

that have been relatively recently recorded for the state and that are ae to be iu b invasive ean

ecologically destructive in other regions of the United States. These featur

on the Texas “Watch List” (Appendix 5). These species may be expected to sputum in Texas and become

ecologically problematic. Some of the woody species, like Cinnamomum camphora tifolia, Pista-

cia chinensis, and Vitex donus: kas are widely planted and seeds often are nai ubiquitous around urban

areas—each of these species

y appears to be spreading into natural habitats. In contrast, species such

as Albizia julibrissin and Morus alba are invasive and occur in large numbers but appear to be more restricted

to disturbed habitats or to edges of woods.

An interesting point regarding pantropical

tly invasive in Texas

has been raised by Tom Patterson (South Texas College, Rio Gane City). “How to Glaser these new arrivals?

For the most part, they have not escaped from cultivation. But they potentially are a threat and I believe

they need their own category. After tropical storms South Padre EH receives sea beans, red mangrove

fruits, and coconuts. Some of tl les in the past | 1

1 plants and have been infrequently

documented by CONSEDRS, ony later to Vice to a severe freeze. With global warming more of these

pantropical will persist and 1 lly adventive to Texas." In this category Patterson

includes Caesalpina bondie Canavalia rosea, Cassytha filiformis, Conocarpus erectus, Laguncularia racemosa,

Rhizophora mangle, and Scaveola plumieri. These are added as a special category in the Watch List (Appendix

Roadside flora.

The emphasis here on ecological impact to natural habitats is especially significant in Texas, where such a

relatively small portion of the land is unaltered and it is critical to protect remaining natural areas. Following

n.a » inf LJ dd de y

fTexas 3(2)

976 J

E L2 11 eC +] ae significant por-

significant A a s a wti y hr

that thought, however, roadsides in Te

tion of public land in the state and in some since. are important in harboring elements Br the native flora.

“It’s hard to know what to do with roadsides. They're not entirely natural, and

stops at the fence. Don't you wish you had a time machine and could go back 1000 years (or 100) and see

Texas without all of the exotics? I wonder what spring would look like without all the non-native mustards,

mints, beans, grasses, etc. What might we have had that is no longer here at all? All that vetch and clover

and henbit and shepherd's purse had to replace *something*" (Monique Reed, pers. comm...

pa he

VCLCLTLIIIC SS

APPENDIX 1

All non-native species known to occur in Texas, with "Fundamental Invasiveness Index" rankings. Species on the"WATCH List

(Appendix 4) are indicated.

ACANTHACEAE

Hygrophila polysperma (Roxb.) T. Anders.—F2-Woody

Nomaphila stricta (Vahl) Nees—F2-Woody

Ruellia caerulea Morong—F3-Herbaceous

syn= Ruellia brittoniana Leonard

Thunbergia alata Bojer ex Sims—F3-Herbaceous

CEAE

Aloe vera (L) Burm. f—F3-Herbaceous

AMARANTHACEAE

Achyranthes aspera NS Hea

2-Herbaceous

Alt tl phil ides (M D Griseb. F1-Woody

ak Lu +l

E23. Llaris arcanis

rF5-ricrDavccuu»

Amate tenella oe Hemmace cus:

Amaranthus blitum L—F2- Herbaceous

syn= Amaranthus virid non |

Amaranthus cruentus L.—F3-Herbaceous

syn= Amaranthus hybridus var. cruentus (L) Moq.

Amaranthus hypochondriacus | —F3-Herbaceous

maranthus sessilis (L) DC —F4- e

ibn argentea L.—F3-Herbaceo

Gomphrena globosa L.—F2- He beret

AMARYLLIDACEAE

Amaryllis belladonna L.—F3-Herbaceous

nee tubispathus Mea en Fa Herbaceous

Narcissus jonquilla L —F3- Herbaceous

Narcissus pseudonarcissus L.—F3-Herbaceous

Narcissus tazetta L.—F3-Herbaceous

ANACARDIACEAE

Pistacia chinensis Bunge—F3-Woody WATCH

Schinus longifolius (Lindl.) Speg.—F3-Woody

Schinus molle L.—F3-Woody' WAT CH

F3 Dinarly WATCH

"TM. Y WWF RE Wu

APIACEAE

Ammi majus L.—F2-Herbaceous

jd visnaga (L) Lam.—F3-Herbaceous

neum graveco ens. L. Pd neo

"IP Ss Wael oh th]

Bupleurum rotundifolium L.—F2-Herbaceous

Conium maculatum L.—F2-Herbaceous

Coriandrum sativum L.—F3-Herbaceous

Cuminum cyminum L.—F3-Herbaceou

po oe m (Pers) Sprague ex Britt. &

Wilso 2-Herbace

SYN— má B (Pers) F. Muell. ex Benth

Daucus carota L.—F2-Herbaceous

Roeniculum vulgare P js bd enc

nri arant Ig

¡e "ae E PA haha P. a a

Pastinaca sativa L—F3- Herbaceou

Petroselinum crispum (P. Mill.) ace ex AW. Hill—F3-

Herbaceous

syN= Apium petroselinum L.

Scandix pecten-veneris L—F3-Herbace

Torilis arvensis (Huds.) Link—F2- Herbaceous WATCH

Torilis nodosa (L.) Gaertn.—F2-Herbaceous

APOCYNACEAE

Catharanthus roseus (L) G. Don—F3-Woody

syN= Vinca rosea L.

Nerium oleander L.—F3-Wood

Vinca major L.—F2-Herbaceous

Vinca minor L.—F4-Herbaceous

ARACEAE

icr: esculenta (L) Schott —F1-Woody

— Colocasia antiquorum Schott

es ocoryne a ex Trimen—F2-Woody

Pistia stratiodes L.—F1-Woody

Xanthosoma sagittifolium (L.) Schott—F2-Woody

ARALIACEAE

Hedera helix L.—F2-Herbaceous

ASCLEPIADACEAE

Asclepias curassavica L.—F3-Herbaceous

Cryptostegia grandiflora (Roxb. ex R. Br.) R. Br.—F3-

Herbaceous WATCH

Periploca graeca |.—F3-Herbaceous

ASTERACEAE Aster Family

" EX Vie F4 Larharaniic

"es E A "es us P A aa

Acroptilon repens (L) DC.—F4-Herbaceous

sYN= Centaurea repens L.

Anthemis cotula L.—F2-Herbaceous

Bidens pilosa L.—F3-Herbaceous

sYN= Bidens odorata Cav.

Carduus acanthoides L.—F3- Herbaceous

Herbaceous

r [| "I E H H 1 E gt

Nesom

syN= Carduus macrocephalus Desf.

syN= Carduus nutans var. macroceohalus (Desf) Boivin

Carduus pycnocephalus L.—F2-Herbaceous WATCH

Carthamus lanatus L.—F3-Herbac

Carthamus tinctorius Iii e cala

Centaurea cyanus L—F2-Herbaceous

Centaurea melitensis _—F1-Herbaceous

Centaurea solstitialis L.—F4-Herbaceous

Cichorium intybus L.—F2-Herbaceous

Cirsium vulgare (Savi) Ten.—F2-Herbaceous WATCH

sYN— Carduus lanceolatus L.

Cnicus benedictus L.—F4-Herbaceous

syn= Centaurea benedicta (L.) L.

Conyza bonariensis (L) Cronq.—F2-Herbaceous

syN= Erigeron bonariensis L.

Cosmos bipinnatus Cav.—F3-Herbaceous

Cosmos sulphureus Cav.—F3-Herbaceous

Cotula australis (Sieber) Hook. f.—F3-Herbaceous

Crepis capillaris (L.) Wallr—F4- oo

Crepis pulchra | —F3-Herbac

Crepis setosa Haller f—F4-Her Hus eou

Crepis zacintha (L.) Babcock—F4- sE

Emilia fosbergii D.H. Nicols.—F4-Herbaceous

Facelis retusa (Lam. Schultz-Bip.—F2-Herbaceous

Galinsoga parviflora Cav.—F2-Herbaceous

sYN= Galinsoga parviflora var. semicalva Gra

Gamochaeta antillana (Urb) Anderb.—F2-Herbaceous

Gamochaeta calviceps (Fern) Cabrera—F2-Herbaceous

Gamochaeta coarctata (Willd.) Kerg.—F2-Herbaceous

Gamochaeta pensylvanica (Willd.) Cabrera—F2-

Herbaceous

Hedypnois cretica (L.) Dum.-Cours.—F2-Herbaceous

Hypochaeris brasiliensis var. tweediei (Hook. & Arn.)

Baker—F2-Herbaceous

syN= Hypochaeris tweediei Hook. & Arn.

Hypochaeris glabra L.—F2-Herbaceous

Hypochaeris microcephala var. albiflora (Kuntze) Cabre-

ra—F2-Herbaceous

Hypochaeris radicata L.—F2-Herbaceous

Lactuca saligna L.—F3-Herbaceous

Lactuca serriola _—F2-Herbaceous

SYN— Lactuca scariola L.

Lapsana communis L.—F3-Herbaceous

Leontodon hispidus | PU Ipae

EA [u pum raro LT «4

TAI valevus

[a pm me

NERAL UI

[| e | anm E9

LA Iia i de

SYN= Chivsanthehm leucanthemum L.

Matricaria recutita L. Wild Chamomile—F4-Herbaceous

SYN= Psi. [EUH (L) Rauschert

la var. coronata (J. Gay) Coss. &

sYN- Matricar

syN= Matricaria suaveolens L.

Onopordum acanthium L.— rbaceous

dira luteoalbum (L) Hilliard €: Burtt—F2-

Herbac

Sanvitalia tia A. Gray—F3-Herbaceous

Scorzonera laciniata L.—F2-Herbaceous

Senecio vulgaris L.—F2-Herbaceous

f Texas 977

Silybum m marlanum (L) obs .—F3-Herbaceous

LI Ar har Pa Ta EE

F2 HerDaceous

one FNNA niusi Kunth

SYN= Gymnostyles stolonifera (Brot ) Tutin—

sYN— Gymnostyles nasturtiifolia Juss.

Sonchus asper ae Hill—F2 Miei

Sonchus oleraceus L.—F2-Herbaceo

Sphagneticola sri n ) ic es Herbaceous

syN= Wedelia trilobata L.

sl a acaba squamatum (Spreng.) Nesom—F4-

Herbac

SYN = i subse vat. bic A. Gray

Tagetes erecta |.— r

Taraxacum lsevigitum (Willd.) ac

SYN= Taraxacum ti Andrz. ex Bess

A L|

fficinale L. subsp. officinale—F2-Herbaceous

Tragopogon dubius Scop.—F2-Herbaceous

Tragopogon porrifolius |.—F2-Herbaceous

Tridax procumbens |.—F3-Herbaceous

Xanthium spinosum L.—F2-Herbaceous

sYN= Xanthium spinosum var. inerme Bel

Youngia japonica (L.) DC.—F2-Herbaceous

Zinnia violacea Cav—F3-Herbaceous

BASELLACEAE

Anredera cordifolia (Ten.) Steenis—F3-Herbaceous

syN= Boussingaultia gracilis Miers

BERBERIDACEAE

Nandina domestica Thunb.—F1-Woody

BIGNONIACEAE

Macfadyena unguis-cati (L.) A.H. Gentry—F3-Woody

WATCH

(TI IY Siah & Ziice ay Srariel F3

Woody

BORAGINACEAE

Anchusa azurea P. Mill. —F3-Herbaceous

SYN— Anchusa italica Retz.

Buglossoides arvensis (L.) I.M. jonnston—F2-Herbaceous

SyN= Lithospermum arvense L.

Cynoglossum zeylanicum (Vahl) Thunb. ex Lehm.—F3-

Herbaceous

Ecnium vulgare L. mia Herbaceous

AJ F2 [a pow "FS IEA

Heliotropium europaeum L. —F3- jieu ous

Hellotropium nicum T Herbaceous

arhsranie

TF TISDALE

BRASSICACEAE Mustard Family

Arabidopsis thaliana (L.) Heynh.—F2- rd

Brassica juncea (L.) Czern.—F3-Herbaceous

Brassica nigra (L.) W.D.). eon

Brassica oleracea | —F3- de

Brassica rapa L.—F2-Herbaceou

SYN— Brassica campestris var. sd (L.) Hartman

978

SYN= Brassica rapa Ya var. campestris (L.) W.DJ. Koch

n E? Lvl Pra Ta B

FIT

Cakile maritima Scop. —F4- ll

Camelina microcarpa DC.— erbac

SYN= Meno saliva ee microcarpa ie E. Schmid

Harl T

Capsella bursa- pastoris (L.) Hs oo

Cardamine debilis D. Don—F2-Herbac

Cardamine hirsuta quc mcn

Cardaria draba (L) Desv.—F3-Herbaceous

Chorispora tenella (Pallas) DC —F2-Herbaceous

ce orientalis (L.) Dumort.—F2-Herbaceous

Coronopus didymus (L.) 5m.—F2-Herbac

Das sophia (L.) Webb ex ici: “Herbaceous

Diplotaxis muralis (L) DC.—F2-Herbaceou

Diplotaxis ten uifolia (L) DC.—F3-Herbace

Eruca vesicaria subsp. sativa (P. Mill.) Thellung—F2-

Herbaceous

SYN— Brassica eruca L.

SYN— Eruca sativa P. M

syN= Raphanus eruca (L.) C

Erucastrum gallicum (Willd.) O.E. Schulz—F2-Herbaceous

Erysimum repandum L.—F2-Herbaceous

Lepidium campestre (L.) R. Br. —F3-Herbaceous

Lepidium latifolium |.—F3-Herbaceous

Lepidium ruderale | —F3-Herbaceous

Lobularia maritima (L) Desv.—F3-Herbaceous

n= Alyssum maritimum (L.) Lam.

Matthiola incana (L.) Ait. f—F3-Herbace

Matinios longipetala Eno E Pd Herbaceous

PW. Ball

Myagrum perfoliatum L.—F3-Herbaceous

Nasturtium officinale W.T. Ait. —F1-Woody

syn= Rorippa nasturtium-aquaticum (L.) Hayek

Raphanus rapistrum L.—F3- bici

Raphanus sativus L.—F3-Herbaceo

sYN= Raphanus raphanistrum var. sativus (L.) G. Beck

Rapistrum rugosum (L.) All—F1-Herbaceous

Sinapis alba L—F4-Herbaceous

Sinapis arvensis L.—F2-Herbaceous

syN= Brassica kaber (DC.) L.C. Wheeler

Sisymbrium altissimum L.—F2-Herbaceous

Sisymbrium irio L.—F2-Herbaceou

Sisymbrium officinale (L.) Scop.—F2-Herbaceous

Sisymbrium orientale |.—F3-Herbaceous

SYN— Brassica kaber var. orientalis (L) Scoggan

Sisymbrium polyceratium L.—F3-Herbaceous

hlaspi arvense L.—F2-Herbaceous

CACTACEAE

onu ficus-indica (L.) P. Mill. —F3-Woody

ntia compressa J.F. Macbr.

paresis aculeata P. Mill —F3-Woody

CAMPANULACEAE

Campanula rapunculoides | —F3-Herbaceous

Wahlenbergia marginata (Thunb.) A. DC.—F3-Herbaceous

“a

SY ds A!

f Texas 3(2)

CANNABACEAE

Cannabis sativa L. var. sativa—F3-Woody

CANNACEAE

Bail indica] —F3-Herbaceous

SYN= Canna xorchiodes Bailey

Canna indica |.—F3-Herbaceous

sYN= Canna coccinea P. Mill.

CAPPARACEAE

leome gynandra L.—F3-Herbaceous

syN= Cleome pentaphylla L.

Cleome hassleriana Chod.—F4-Herbaceous

syn= Cleome pungens auct. non Willd.

syN= Cleome spinosa auct. non Jacq.

CAPRIFOLIACEAE

Lonicera fragrantissima Lind. & Paxton—F3-Woody

"i ni icera Japonica TAUNO; —F1-Woody

! E33. Ware r WATCH

tw ee € PEF Le Ce

CARYOPHYLLACEAE

grostemma githago L.—F3-Herbaceous

Arenaria serpyllifolia " —F2-Herbaceous

SYN= ii anl holosteoides auct. non abo.

_ "N ri,

= ) Hyl.

an glomeratum Thuill. —F2- -Herbaceous

syn= Cerastium viscosum auct. n

Dianthus barbatus |. —F4-Herbaceous

Gypsophila elegans M. Bieb—F3-Herbaceous

Holosteum umbellatum |.—F2-Herbaceous

Pen organia cuni (Raf) da & M Es eles

pabcamois oy (L) L. —F2- Herbaceous

Sagina procumbens |.—F4-Herbaceous

Saponaria officinalis L.—F3-Herbaceous

zen coneiees A beats

Ls "m———

1eFDaceous

Silene gallica L.—F2- Herbae eous

ape ergula s arvensis L.—F3- Herpacepus

Celak.) Accherc & (Graahn —F3-

Herbaceous—

Spergularia platensis (Camb.) Fenzi—F3-Herbaceous

Stellaria media (L.) Vill. —F2-Herbaceous

Stellaria pallida (Dumort.) Crépin—F2-Herbaceous

syn= Stellaria media subsp. pallida (Dumort) Aschers. &

Graebn

Dar ri ES Larkseaniic

A A I! B n? E dC Ae P Rd

9 Larkaraniic

E Y Ach ee ee iod

4 ff. H LI Mm RASI’ D

OE a ] Ham pur tendu E Nu

CASUARINACEAE

Casuarina equisetifolia | —F3-Woody WATCH

CELASTRACEAE

CHENOPODIACEAE

Atriplex holocarpa F. Muell.—F3-Woody

E»5?.Harharnanmic

MEN itr ae oe

F3 Harharnanmiriec

PR RARE AIDA ha hes RP RR ad

H Ti iros \ Wany RAa>>

rF [| al * a H J £

Nesom

979

Atriplex rosea L.—F3-Herbaceous

AUIpIEX samibaccata ji Rd -Herbaceous

r3 Larkaranis

r3-nerbaceous

= Kochia hyssopifolia Calas Schrad.

Pests scoparia (L) AJ. Scott —F1-Herbaceous

syN= Kochia scoparia (L.) ned

Beta vulgaris | —F3-Herbaceous

syn= Beta vulgaris subsp. maritima (L) n5

Chenopodium album L.—F2-Herbaceo

Chenopodium ambrosioides | —F2- a”

Chenopodium botrys | —F3-Herbaceous

Chenopodium glaucum L.—F3-Herbaceous

Chenopodium murale L.—F2-Herbaceous

Chenopodium opulifolium Schrad. ex Koch & Ziz—F4-

Herbaceous

syN= Chenopodium album var. viride (L) Moq.

sYN— Chenopodium viride

Chenopodium pumilio R. Br.—F3-Herbaceous

syN= Chenopodium carinatum auct. non R. Br.

enopodium vulvaria |. FE

Salsola colma Pallas—F4-Herbac

Salso L. subsp. pontica a Mosyakin—F1-

SYN= Salsola kali var. pontica Pallas

SYN= Salsola pontica (Pallas) A. Degen

sau ategus L Mid Herbace eous

Sal R 1) Botsch. ex Czerepanov

Salsola kali subsp na (L.) Celak.

Salsola pestifer A

Spinacia oleracea L. —F3- em

SYN—

CLUSIACEAE

Hypericum perforatum L.—F2-Woody WATCH

COMBRETACEAE

Conocarpus erectus L.—F3-Woody

Laguncularia racemosa (L.) Gaertn.—F3-Woody

COMMELINACEAE

mmelina communis L.—F2-Herbaceous

Gibasis pellucida (Mart. & Gal.) D.R. Hunt—F2- —

Murdannia nudiflora (L) Brenan—F2-Herbaceous

syN= Aneilema nudicaule (Burm. f) G. Don

syn= Aneilema nudiflorum (L.) Sweet

CONVOLVULACEAE

Convolvulus arvensis L.—F2-Herbaceous

Ipomoea alba |.—F3-Herbaceous

Ipomoea batatas (L.) Lam.—F4- idis

lpomoea cairica (L) Sweet—F3-Herbaceous

lpomoea capillacea (Kunth) p - siis Herbaceous

Ipomoea carnea Jacq.—F2- aceous

omoea crassicaulis diis B.L. Robins.

SYN— pomorai G pp ex Choisy

YN= |

Ipomoea hederifolia L—F4- Herbae

= |pomoea coccinea var. acp (L.) Gray

ponoc: indica (Burm. f) Merr.—F3-Herbaceous

moea acuminata pm Roemer & J.A. Schultes

Ipomoea nil (L) Roth—-F3-Herbaceous

Ipomoea quamoelt L—F3-Herbaceous

Kar Gaw |

"———

SYN= lpomoea muricata (L) Jacq.

Ipomoea violacea L.—F4-Herbaceous

jaa wrightii Gray—F4-Herbac

N= Ipomoea heptaphylla Do oe

CRASSULACEAE

Kalanchoe daigremontana Raym.-Hamet & Perrier—F3-

rbaceous

Kalanchoe delagoensis Ecklon & emi ae

N= Kalanchoe tubiflora (Harvey) Ham

Kalanchoe verticilllata Ell.

Kandis oo Raym.-Hamet & H. Perrier—F4-

Herbac

CUCURBITACEAE

Citrullus colocynthis (L.) Schrad.—F4-Herbaceous

Citrullus lanatus var. citroides (Bailey) Mansf—F4-

Herbaceous

syn= Citrullus vulgaris var. citroides Bailey

Citrullus lanatus (Thunb.) Mansf. var. lanatus—F4-

Herbaceous

syn= Citrullus vulgaris Schrad.

Coccinia grandis (L) Voigt —F4-Herbaceous

syn= Coccinia cordifolia sensu Correll & Johnston

o anguria L- mis ion poe oe

E> Layla SY PASEA

e a Il

Ne Fel "aue P E S

e melo L.—F2- posto

Cucumis melo var. sandalia Naud —F4 Herbaceous

a Liarkaraniic

Eo S INE AIDA e VP Ml

-F

I EE +3 DA AANI C2 WUarkaroniic

tow fF ESE Daceous

SYN- Luffa cylindrica (L) M. Roem

Momordica balsamina Sic PPM

Momordica charantia |.—F4-Herbaceous

CUSCUTACEAE

eua) Japonica ney a -Herbaceous

F3-Herbaceous

CYP

ERACEAE

Bulbostylis barbata (Rottb.) C.B. Clarke—F3-Herbaceous

Cyperus difformis E: msi PTOA ESMS

Herba arnie

Casu a el a

Cyperus eragrostis Lam.—F3- Herbaceous

SYN= Cyperus vegetus ue

E» Lark S22" ei

W dm B Re AIDA es P Ru

Cyperus i iria L.—F2-Herbaceo

Cyperus phaeolepis Cherm o

F3 Herba A Fanlic

Cyperus rotundus | x re Heraeus

is Gale—F3-Herbaceous

DIOSCOREACEAE

Dioscorea bulbifera L—F4-Herbaceous

DIPSACACEAE

Scabiosa atropurpurea L.—F2-Herbaceous

Dipsacus fullonum L.—F3-Herbaceous

980 t tani titute of Texas 3(2)

DRYOPTERIDACEAE Medicago lupulina L—F2-Herbaceous

Cyrtomium falcatum (L. f) K. Pres|—F3-Herbaceous Medicago minima (L.) |. —F2-Herbaceous

syN= Polystichum falcatum (L. f) Diels Medicago orbicularis (L.) Bartalini—F2-Herbaceous

ELAEAGNACEAE e M MR nolis ceous

Plaeagnus angustifolla E pe Woody WATCH medicago Sativa m erbaceous

Sn das syN= Medicago xvaria Martyn

Psu y A "

Elaeagnus pungens THUNB: —F3-Woody Melilotus albus Medik.—F1-Herbaceous

Melilotus indicus (L.) All. —F1-Herbaceous

EUPHORBIACEAE Melilotus officinalis (L.) Lam.—F1-Herbaceous

Croton argenteus L.—F2-Herbaceous Neptunia plena (L.) Benth.—F3-Herbaceous

Chamaesyce hirta (L.) Millsp.—F3-Herbaceous Oxyrhynchus volubilis Brandeg.—F4-Herbaceous

Euphorbia dann L.—F4-Herbaceous Prosopis laevigata (Humb. & Bonpl. ex Wil—ld.) M.C.

Euphorbia exigua L.—F3-Herbaceous Jo hns ton—F3-Woo dy

Euphorbia la L.—F3-Herbaceous P ta ( Willd.) M 2 Almeida

Euphorbia lathyris |.—F3-Herbaceous F2-Woody WATCH

Manihot esculenta Crantz—F3-Woody svNz Pueraria lobata (Willd.) Ohwi

Phyllanthus fraternus G. L. Webster—F3-Herbaceous Senna corymbosa (Lam.) Irwin & Barneby—F3-Woody

F3-Herbaceous syn= Cassia corymbosa Lam

Phyllanthus urinaria L.—F2-Herbaceous Senna occidentalis (L.) Link—F2-Herbaceous

Ricinus communis [.—F2-Herbaceous syn Cassia occidentalis L

Triadica sebifera (L) Small —F1-Woody Carhañta A Urb. —F2-Herbaceous

= Sapium sebiferum (L.) Roxb. Sesbania punicea (Cav) Benth —F3- aos

FABACEAE Sesbania sericea eee RÍA Herbaceous

Spartium junc

nomene evenia C. Wright—F3-Herbaceous oay

ody coria: Rub mi DC.—F3-Herbaceous

Albizia julibrissin Durazz.—F2-Wco

Albizia lebbeck (L) Be Benth. th. —F3-Woody Trifonum arvense n -Fo Herbaceous

F3 Harharaniie

Trifanl: 7l is Cil E? Harla = m mi e

Trifolium incarnatum L.—F2-Herbaceous

Tnronum lappaceum Ap Herbaceous

SYN— N= Alhagi camelorum Fisch.

Alysicarpus vaginalis (L.) DC.—F3-Herbaceous

Arachis hypogaea L.—F3-Herbaceous

Caesalpinia gilliesii (Hook.) Wallich ex D. Dietr.—F3-

Wood

SYN= Trifolium amphanthum Torr. & Gray

Trifolium pratense L.—F3-Herbaceous

Trifolium repens L.—F2-Herbaceous

Trifolium resupinatum L.—F2-Herbaceous

ody

sYN— Poinciana gilliesii Wallich ex Hook.

Caesalpinia mexicana Gray—F3-Woody

SYN= Poinciana mexicana (Gray Britt. & Rose

Caesalpinia sink (L) Sw.—F3-Woody

sYN- Poinciana pulcherrima L.

Clitoria ane L.—F3-Herbaceous

Coronilla varia |. —F2-Herbaceous

caus retusa | —F3- Herbaceous Vicia lathyroides | —F3-Herbaceous

Herbaceous Vicia lutea L.—F3-Herbaceous

SYN= Crotalaria retzi À. S. Hitchc. Vicia sativa subsp. nigra (L.) Ehrh.—F2-Herbaceous

( ki F3-Herbaceous syN= Vicia angustifolia L.

SYN= Lespedeza stipulacea Maxiin SYN= Vicia sativa var. angustifolia (L.) Ser.

Kummerowia striata (Thunb) Schindl.—F2-Herbaceous SYN= Vicia sativa var. nigra

A Chun) Hoo! & Arn. SYN Vicia sativa var. segetalis (Thuill) Ser.

Vicia sativa L. subsp. sativa—F3-Herbaceous

syn= Vicia sativa var. linearis Lan

Vicia tetrasperma (L.) Schreb.—F2-Herbac

Vicia villosa Radzhisu x varia (Host) Corb. E e —

= Vicia dasycarpa Ti

vids villosa Radzhi o villosa— F2-Herbaceous

Vigna unguiculata (L oui -Herbaceous

syN= Vigna sinensis (L.) Sa

Wires sinensis (Sims) ar -Woody

ai aphaca L—F3- Herbaceous

Lathyrus hirsutus L—F2-Herbaceous

Lathyrus latifolius |—F3-Herbaceous

Lespedeza bicolor Turcz.—F3-Herbaceous

Lespedeza cuneata (Dum.-Cours.) G. Don—F2-Herbaceous

WATCH

Leucaena leucocephala (Lam.) de Wit—F3-Woody

WATCH

Lotus corniculatus L.—F3-Herbaceous

Macroptilium gibbosifolium (Ortega) A. Delgado—F4- FUMARIACEAE

aceou Fumaria densiflora DC.—F3-Herbaceous

Medicago arabica (L.) Huds —F2-Herbaceous

r [| awh = H - 3 £ gt

Nacom

Fumana officinalis L —F3-Herbaceous

ANI SE

Centaurium muhlenbergeii (Griseb.) Piper—F2-

Herbaceous

GERANIACEAE

Erodium botrys (Cav.) Bertol —F3-Herbaceous

Erodium malacoides (L.) Willd —F3-Herbaceous

Erodium cicutarium (L.) |'Hér. ext Ait.—F2-Herbaceous

Geranium dissectum L.—F2-Herbaceous

GOODENIACEAE

Scaveola plumieri (L.) Vahl—F3-Herbaceous

LOR Gn Eee

EC> Harhararmt S

E m |; Q*E BG Ee SS

FEPTEM ESAS el

)VOTac.—F i a

SYN— - Myriophyllum brasiliense Camb.

syN= Myriophyllum proserpinacoides Gillies ex Hook. &

Arn.

Myriophyllum spicatum L.—F1-Woody

HYDROCHARITACEAE

Egeria densa Planch.— ody

syN= Elodea densa (Planch.) Caspary

Hydrilla verticillata (L. f) Royle—F1-Woody

Ottelia alismoides (L.) Pers.—F2-Woody

IRIDACEAE

Belamcanda chinensis (L) DC —F3-Herbaceous

Crocosmia xcrocosmiiflora (V. Lemoine) N.E. Br.—F4-

erbaceous

Iris pallida Lam.—F4-Herbaceous

Iris pseudacorus L.—F3-Herbaceous

Romulea rosea (L.) Eckl.—F3-Herbaceous

Tritonia crocosmifolia Lem.—F3-Herbaceous

JUNCACEAE

no.

Yrer! lA "and io

LAMIACEAE

Ajuga reptans L.—F3-Herbaceous

Glechoma er eraros ee Herbaceous

ich.) B F3-Herbaceous

Lamium amplexicaule | —F2- Herbaceous

Lamium purpureum L.—F2-Herbaceous

Leonotis nepetifolia (L) Ait. f—F2-Herbaceous

Leonurus cardiaca L.—F3-Herbaceous

Leonurus sibiricus L.—F3-Herbaceous

Mentha xpiperita L.—F3-Herbac

Mentha xrotundifolia (L.) Huds. a a

Mentha spicata E ml Herbaceous

dee laevis L.—F3- Puen

Nepeta cataria | —F3-Herbac

Perilla fie a: (L) Ad iaa WATCH

Prunella vulgaris L.—F2-Herbaceous

Rosmarinus officinalis _—F3-Woody

Salvia hispanica L.—F4-Herbaceous

f Texas 981

— tiliifolia Vahl—F3-Herbaceous

tellaria minor nue on—F3-Herbaceous

Sideritis lanata L.—F3-Herbaceous

Stachys floridana Shuttlew. ex Benth.—F2-Herbaceous

WATCH

LAURACEAE

Cinnamomum camphora (L.) J. Presl —F3-Woody WATCH

LEMNACEAE

Landoltia punctata (G. Mey.) Les & Crawford —F2-Woody

SYN= Spirodela punctata (G. Mey.) C.H. Thompson

LILIACEAE

Allium ampeloprasum L.—F3- om

Allium porrum L.—F3-Herbac

ous

Alstroemeria pulchella L.f. jipe Herbaceous

Asparagus officinalis | —-F3-Herbaceous

Asphodelus fistulosus _—F3-Herbaceous

Crinum bulbispermum (Burm.f) Milne-R. & Schweick.—F3-

Herbaceous

= Crinum longifolium (L.) Thunb.

Hemerocallis fulva (L) L —F3-Herbaceous

Hemerocallis lilioasphodelus _—F3-Herbaceous

syN= Hemerocallis flava (L) L.

Hyacinthus orientalis L.—F3-Herbaceous

Lilium longiflorum Thunb.—F3-Herbaceous

Muscari comosum (L.) Mill.—F3-Herbaceous

Muscari neglectum Guss. ex Ten.—F3-Herbaceous

syN= Muscari racemosum (L.) Lam. & DC.

rnithogalum umbellatum L.—F4-Herbaceous

Msaga uniflorum (Lindl.) Traub—F3-Herbaceous

= ipheion uniflorum (Lindl) Ra

Zephyranthes candida (Lindl.) -Herbaceousert—F3-

Herbaceous

SYN= iind candida (Lindl) Small

E2_UWarharaniic

PIERDO ECU

LIMNOCHARITACEAE

Hydrocleys nymphoides (H. €: B. ex Willd.) Buch.—F2-

Woody

AERE

Macet

Linum usitatissimum L.—F3- Herbaceous

LOGANIACEAE

Buddleja lindleyana Fort —F3-Woody

LYGODIACEAE

LYTHRACEAE

Cuphea carthagenensis (Jacq) J.F. Macbr.—F2-Herbaceous

Cuphea glutinosa Cham. & Schlecht. —F3-Herbaceous

Cuphea viscosissima Jacq.—F3-Herbaceous

Lagerstroemia indica |. —F2-Woody

Lythrum salicaria |. —F3-Herbaceous

MAGNOLIACEAE

Liriodendron tulipifera _—F3-Woody

982

MALVACEAE

Abutilon theophrasti Medik.—F3-Herbaceous

Alcea rosea |.—F3-Herbaceous

sYN— Althaea rosea (L.) Cav.

Hibiscus esculentus |.—F3-Herbaceous

Hibiscus syriacus L_—F3-Woody

SIIDISCHS tonum Es Pd Herbaceous

E2?_Warkaraniic

Peo -mernvatevuus

1 Presl) ! Fryxell

Lavatera trimestris L—W2

Malva neglecta Wallr.—F2-Herbaceous

Malva parviflora | —F2-Herbaceous

Malva rotundifolia |.—F3-Herbaceous

syN?= Malva pusilla Sm

Malva sylvestris L. —F3-Herbaceous

igi —— (L.) Garcke—F2-

Herbac

VE haaa nenne DC.—F4-Herbac

sYN— Malvaviscus aboreus var. pendulous Le Schery

pavonis hastata Cav.—F3-Herbace

MELIACEAE

Melia azedarach L—F1-Woody

MENYANTHACEAE

Numpnelges indica » RUNES ee lo

F2 VWinndy

ww wn,

MOLLUGINACEAE

Glinus lotoides L.—F3-Herbaceous

Mollugo cerviana (L.) Ser. —F2-Herbaceous

MORACEAE

Broussonete isabel (L) L'Hér. ex Vent.—F2- e

Fatoua hunb.) Nakai—F2-Herbac

Ficus carica e oody

Ficus pumila L.—F3-Woody

Morus alba L.—F2-Woody

MYRSINACEAE

Ardisia crenata Sims—F3-Woody WATCH

MYRTACEAE

Myrtus communis L.—F3-Woody

NYCTAGINACEAE

Bougainvillea glabra Choisy—F3-Herbaceous

Mirabilis jalapa L.—F3-Herbaceous

OLEACEAE

Jasminum mesnyi Hance—F3-Woody

Ligustrum j japonicum Thunb.—F3-Woody

oody

igustrum sinense Lour.—F1-Woody

ONAGRACEAE

Ludwigia erecta (L.) H. Hara—F3-Herbaceous

Ludwigia grandiflora (M. Micheli) Greuter €: Burdet subsp.

grandiflora—F2-Woody

syN= Ludwigia uruguayensis (Camb.) Hara

E graneitiore

la (Hook. & Arn.) Nesom & Kartesz—F2-

as)

Ans

swN- Ludwig tala (Hook. & Arn.) Zardini et al.

Oenothera cordata J. W. Loudon—F2- Herbaceous

ORCHIDACEAE

Zeuxine strateumatica (L.) Schlechter—F3-Herbaceous

OROBANCHACEAE

Orobanche minor Sm.—F3-Herbaceous

Orobanche ramosa L.—F1-Herbaceous

OXALIDACEAE

Oxalis corniculata L.—F3-Herbaceous

Oxalis debilis Kunth—F2-Herbaceous

SYN= Oxalis ESTOS E,

El h

PTa4-hicivatlevus

SYN= walle riara C e"

PAPAVERACEAE

Eschscholzia californica Cham.—F3-Herbaceous

Glaucium corniculatum (L.) J.H. Rudolph—F3-Herbaceous

Papaver rhoeas |.—F3-Herbaceous

Papaver somniferum _—F3-Herbaceous

PARKERIACEAE

Ceratopteris thalictroides (L) Brongn.—F2-Woody

PEDALIACEAE

esamum orientale L—F4-Herbaceous

sYN= Sesamum indicum L.

PINACEAE

Pinus elliottii Engelm.—F3-Woody

PIPERACEAE

Peperomia pellucida (L) Kunth—F3-Herbaceous

PLANTAGINACEAE

Plantago coronopus L.—F3-Herbaceous

Plantago lanceolata L.—F2-Herbaceous

Plantago major L—F2-Herbaceous

POACEAE Grass Family

Achnatherum clandestinum (Hack.) Barkworth—F3-

Herbaceous

SYN= Stipa clandestina Hack.

Aegilops cylindrica Host—F2-Herbaceous

SYN= Triticum cylindricum (Host) Ces., pa & Gib

Agropyron cristatum (L.) Gaertn.—F3- Her baceous

syN= Agropyron subsp. pectinatum (Bieb.) Tzvelev

syYN= Agropyron cristatiforme Sarkar

SYN Agropyron pectiniforme Roemer & Schultes

sYN= Agropyron desertorum (Fisch. ex Link) J.A. Schultes

Agrostis avenacea J.F. Gmel.—F3-Herbaceous

Agrostis stolonifera |.—F3-Herbaceous

SYN— Agrostis alba auct. non L.

SYN= ae Gigantes M

SYN=

+ Drehi Cel L. |

A A

=> 22

Aira caryophyllea | —F2-Herbaceous

syN= Aira elegans Willd. ex Kunth

SYN= ag a capas Host

SYN=

HT fh Fo ae ar pant

Alopecurus geniculatus L. —F4-Herbaceous

ark Poor oe TO

H Be E PA ha as P. rd a

983

Nesom t i i ti

f Lf

me Alopecurus agrestis l E

s.—F3-Herbaceous

SYN— E odoratum var. puelii (Lecoq & La-

motte) Coss. & Dur

Anthoxanthum odor ium —F2-Herbaceous

Arthraxon hispidus (Thunb) Makino—F3-Herbaceous

syn= Arthraxon hispidus var. cryptatherus (Hack) Honda

Arundo donax L.—F1-Woo

; erbac

Bothriochloa d (Retz) a ia Herbaceous

syN= Andropogon caucasicus Trin

SYN= e caucasica (Trin) C.E. Hubbard

Bothriochloa ischaemum var. nia (Rupr. ex & M)

Celarier & Harlan —F1-Herbaceous

Andropogon hue var. songaricus Rupr. ex

SYN—

Fisch. & Mey.

Bothriochloa pertusa (L) E Camus—F3- Herbaceous

Rrarhkiaria ( | F3 Herb araniic

Brachypodium distachyon (L) Beauv. —F3- Herbaceous

Bromus arvensis L.—F2-Herba

SYN= i japonicus uus ex Murr.—

syN= Bromus unioloides Kunth

syn= Bromus willdenowii Kunth

Brom tatus Schrad.—F3-Herb

Bromus diandrus Roth—F3-Herbaceous

syn= Bromus rigidus Rot

Bromus hordeaceus L—F2-Herbaceous

syn= Brornus hordeaceus L. subsp. molliformis (Lloyd)

Maire & Weiller.

syN= Bromus hordeaceus subsp. divaricatus (Bonnier &

Layens) Kerg.

Bromus inermis Leyss.—F3-Herbaceous

syN= Bromopsis inermis (Leyss.) Holub

Promus) japonicus TOUR ex Murr.—F2-Herbaceous

F3-Herbaceous

Bromus rubens L.—F2-Herbaceous

Bromus secalinus L.—F2-Herbaceous

Bromus sterilis |. —F3-Herbaceous

Bromus tectorum L.—F2-Herbaceous

Chloris canterae Arechav.—F2-Herbaceous

Chloris divaricata R. Br. —F3-Herbaceous

syN= Chloris cynodontioides Balansa

Chloris gayana Kunth—F2-Herbaceous

Coix lacryma-jobi L.—F3-Herbaceous

Moda: selloana (J.A. & JH. Schultes) Aschers. &

ody

Cyno ste n aethiopicus Clayton €: pines ms

cynodon dactylon (L) Pers ni meee

C rl | £ ie \/ J + E2.Harharantc

CA_Uarhaecaniic

E A Dil Pa occ ee a id

Cynosurus echinatus | —F3-Herbaceous

Dactylis glomerata |.—F2-Herbaceous

Dactyloctenium aegyptium (L.) Willd.—F2-Herbaceous

ICAd>

SYN= Cynosurus aegyptius L.

Desmazeria rigida (L.) Tutin—F2-Herbaceous

pen gia (L) iuebarg ex Dony

fCaeel TAS

syN= Andropogon annulatus For: sk.

Dichanthium aristatum (Poir.) C.E. Hubbard—F1-

Herbaceous

Dichanthium sericeum (R. n "ard -Herbaceous

sYN— Andropogon sericeu

Digitaria ischaemum cii. Schreb. ex Muhl.—F2-

Herbaceous

Di F4-Herbaceous

Digitaria sanguinalis (L.) Scop. —F2- Herbaceous

Digitaria violascens Link—F2-Herbaceous

syn= Digitaria ischaemum var. violascens di Radford

Echinochloa colona (L.) Link—F2-Herbace

Echinochloa crus-galli (L.) e de

syN= Echinochloa frumentacea Link

syN= Echinochloa crus-galli var. frumentacea (Link) W.

Wight

DO elei E ev

I VH | EA LI arkarani ac

por E Pe AF Ge Se P RAE au

Ehrharta calycina Sm. —F3- Her bac eous

Eleusine indica (LJ Caern Mia Hee

FP Le . J rharaniie

Lia

Oe AA Me ee S el uH

VJ IP

EI Hr? Meldris El _Darharasnioe

MIA Fu MINA IE ll» lib

Elymus repens L) Gould—F3-Herbaceous

SYN— Agropyron repens ai Beauv.

Eragrostis amabilis (L.) Wight & Arnolt ex Nees—F3-

Herbaceous

syN= Eragrostis tenella (L.) Beauv. ex Roemer & Schultes

Eragrostis curvula (Schrad) pe Herbaceous

Ar rhararo

rbaceous

H FATIAS

Eragrostis minor Host—F3- Herbie

Eragrostis a var. perplexa ey S. Koch —F3-

Herbaceo

Eragrostis pilosa nd P ips. var. pilosa—F2-Herbaceous

Pad y eTa -Ta TEE

-— Ed

E ll HP | J ] Lark

IE DA

Eriochloa pseudoacrotricha (Stapf ex a Hubbard

ex Blake - Herbace

+ I Herbaceoi Ic

Eustachys retusa (Lag) Kunth—F2- Herbae

Gastridium p (Nees & Meyen) CE. ee mee

He (pace

| e RES ee a eet eee AARIS 7 mo

c | T

riciDacecuu»

SYN— Rottboellia cylindrica Willd.

syN= Monerma cylindrica (Willd.) Coss, & Dur.

Hemarthria altissima (Poir.) Stapf & Hubbard—F3-

erbaceous

syN= Manisuris altissima (Poir.) A.S. Hitchc.

lat [| (PH R TI F3 Harharariic

a Ms A hace RP Ml a

Holcus lanatus L.—F3-Herbaceous

nO [| a | EM ibsp.g ||

INT. | cA

F. AUT

Herbace

SYN= diliges glaucum Steud.

984

Hordeum murinum subsp. leporinum (Link) Arcang.—F2-

n subsp. teporinum (Link) A. Lave

=— FIFAaAcify

SYN= ae leporinum Link

Hordeum vulgare |.—F3-Herbace

imperata cylindrica (L.) Beauv.—F3- fo WATCH

Ixophorus unisetus (J. Pres!) Schitdl.—F3- a

Lamarckia aurea (L.) Moench—F3-Herbaceous

Leptochloa chloridiformis (Hack. ex wn Parodi—F3-

Herbaceous

Lolium arundinaceum (Schreb.) S.J. Darbyshire—F2-

Herbaceous

sYN= Festuca arundinacea Schreb.

Lolium perenne L.—F1-Herbaceous

syn= Lolium multiflorum Lam

syN= Lolium perenne subsp. multiflorum (Lam.) Husnot

syN= Lolium perenne var. multiflorum (Lam.) Parnell

syN= Lolium perenne subsp. italicum (A. Braun) Husnot

Lolium pratense (Huds.) SJ. Darbyshire—F3-Herbaceous

SYN— Festuca elatior L. p.p.

SYN— a parena Puoi

—À

V CL VASTE GT

SYN— Lolium perenne ae rigidum (Gaudin) Love &

Lolium temulentum L.,—F2-Herbace

syN= Lolium temulentum var. arvense is Lilja

Luziola peruviana Juss. ex J. F. Gmel.—F3-Herbaceous

a repens (Willd.) Zizka—F2-Herbaceous

= Rhynchelytrum repens (Willd.) C.E. Hubbard

Mnene granularis (L)—F4-Herbaceous

syN= Hackelochloa granularis (Lj Kuntze

Microstegium vimineum (Trin.) A. Camus—F3-Herbaceous

ATCH

Oryza sativa L. Muss dile en

Panicum bergii Arech.—F2- Hebeis

Panicum coloratum L.—F2-Herbaceous

Panicum miliaceum _.—F3-Herbaceous

Panicum repens hee jaon

Parapholis incurva (L.) C. 3 Hubbard—F2- Herbaceous

syN= Pholiurus incurvus (L.) Schinz & Thellung

Paspalum almum —F3-Herbaceous

jj ie convexum Humb. & Bonpl. ex Flueggé—F3-

Herba

Paspalum oiseau Rd ian iiid

Paspalum malacophyllum ama —F3- Herbaceous

syN= Paspalum notatum var. laetum Dóil

SYN= Paspalum notatum var. saurae Parodi

Paspalum scrobiculatum L.—F4-Herbaceous

Paspalum urvillei Steud.—F2-Herbaceous

Pennisetum ciliare (L.) Link—F1-Herbaceous

SYN— Cenchrus ciliaris L.

Pennisetum glaucum (L.) R. B. —F3-Herbaceous

syN= Pennisetum americanum (L) Lee

sín= Setaria glauca (L.) Beauv.

[| J £ al n.a H anm LJ M den ndis ray

f Texas 3(2)

Pennisetum nervosum (Nees) Trin.—F3-Herbaceous

SYN= ceneng nervosus s (NBBS) See

P F3-Herbaceous

Pennisetum villosum R. Br. ex Fresen.—F2-Herbaceous

syN= Cenchrus ae M.C. Johnston

Phalaris arundinacea L.—F3-Herbaceous

E dea pid a

alaris canariensis rbaceous

clas minor Retz. —F3- Herbaceous

Phleum pratense L.—F3-Herbaceo

Phyllostachys aurea Carr. ex A.& C. aes F2-Woody

Poa annua L.—F2-Herbaceous

Poa bulbosa L.—F4-Herbaceous

Poa compressa L.—F3-Herbaceous

Poa trivialis _—F4-Herbaceous

Polypogon monspeliensis (L.) Desf —F2-Herbaceous

Polypogon viridis (ouan) Breistr.—F2-Herbaceous

Fisch.) Nevski—F4-Herbaceous

N= Elymus junceus Fis Fis ch.

feces cristata (L) Tzvelev—-F3-Herbaceous

syN= Koeleria gerardii (Vill) Shinners

sYN— Koeleria phleoides (Vill.) Pers.

Rottboellia cochinchinensis (Lour.) W.D. Clayton—F3-

Herbaceous

syN= Aegilops exaltat

syN= Manisuris sce a Kuntze

syn= Rottboellia exaltata (L.) L. f.

Saccharum officinarum L.—F3-Herbaceous

Sacciolepis indica (L.) Chase—F3-Herbaceous

Schismus barbatus (Loefl. ex pce ai aan

Sclerochloa dura (1.) Beauv.—F2-Herbac

Secale cereale | .—F3 Fabrak

SYN— Triticum (L.) Salisb.

Setaria italica (L) Beauv.—F2-Herbaceous

Setaria pumila Poin) ill & Schult.—F2-Herbaceous

taria uv.—F2-Herbaceous

Setaria 2 JP a Herbaceous

= Sella viridis var. major (Gaudin) Pospichal

Daca FA Herba araoamric

Sorghum bicolor (L) Moench—F2- Herbaceous

SYN= Sorghum saccharatum (L.) salen

Sorghum halepense (L.) Pers —F1-Herbac

Stenotaphrum secundatum (Walter) A-

Herbaceous

} la triandra Forssk —F3-Herbaceous

NE Dy T Da rl rh Q Dewey F4

Herbaceous

syN= Agropyron intermedium (Host) Beauv.

SYN= OR inten ia (Host) pees

Baikworth & Dewey

b a) M Pa Ty | X] * OVA

syN= Agropyron elongatum (Host) Beauv.

syN= Elymus elongatus (Host) Runemark

SYN— Elytrigia elongata (Host) Nevski

syn= Elytrigia pontica (Podp.) Holub

Thinopyrum pycnanthum (Godr.) Barkworth—F4-

Herbaceous

r Ji &al* 2 | £ PE)

f Texas 985

Nesom

sYN= Agropyron pycnanthum (Godr) Godr. & Gren.

SYN— Elymus pungens auct. non (Pers.) Melderis

syN= Elymus pycnanthus (Godr.) Melderis

sya= Elytrigia pycnanthes (Godr.) A. Love

Triraphis mollis —F3-Herbaceous

Triticum aestivum _—F3-Herbaceous

LO MAI + ES

Woody

syN= Brachiaria brizantha (Hochstetter ex A. Rich.) Stapf

Erochlos maxima (acq) b Webster) -Herbaceous

F3-Woody

Urochloa mutica (Forsk) T.Q. Nguyen—F3- -Herbaceous

syN= Brachiaria mutica (Forsk.) Stapf

SYN= PORCH EIE Forsk.

Urochloa plantaginea (Link) R. id PU Herbaceous

syN= Brachiaria plantaginea (Link) A.S. Hitchc.

Urochloa ramosa (L) Nguyen—F3- as

sYN= Brachiaria ramosa (L.) Stap

Urochloa reptans (L.) Stapf.—F2-Herbaceous

SYN— Panicum reptans

= Brachiaria reptans (L ) Gard. & Hubb.

Vus bromoides (L) S.F. Gray—F3-Herbaceous

sYN= Festuca bromoides L.

syN= Vulpia dertonensis (All) Gola

Vulpia myuros (L.) K.C. Gmel.—F2-Herbaceous

SYN— Festuca myuros

SyN= Vulpia myuros var Areata Hack

Zea mays L.—F3-Herbaceous

Zea perennis (Hitchc.) Reeves & Manglesdorf—F3-

erbaceous

SYN Euchlaena perennis Hitchc.

POLYGONACEAE

Antigonon leptopus Hook. & Arn.—F3-Herbaceous

Emex spinosa (L.) Campd.—F3-Herbaceous

SYN— BAMIA Apot p

| m

Warkaraniic

Polygonum arenastrum Jord. ex Borea u—F2- ecean

WATCH

syN= Polygonum aviculare var. arenastrum (Jord. ex

Porra Rouy

arlan mrmr s

¿Kiinza F3 LJ

23-HnerDaceous

Polygonum aviculare L ia Herbaceous

EJ3 Lark Ta Ta Ad

oligonum caespitosum var. longisetum (Bruijn) A.N.

Steward——F3-Herbaceou

Polygonum convolvulus en Herbaceous

opiper L.—F3- Herbace eous

—F3-Herbaceous

SYN= ie imd (L.) Spac

dics onum eisnerianum var. beyrichianum (C. & S.)

us Herbaceous

enn scandens L.

Polygonum persicaria L.—F2-Herbaceous WATCH

Rumex acetosella L. mae Herbaceous

HWarkaraniic

SPM RO A ud

Rumex crispus L.—F2-Herbaceous

Rumex obtusifolius L. —F3- ise

Lark L- T Se

F3 E Run ARA a P VE

Rumex pulcher L —F2- Herbaceous

POLYPODIACEAE

Nephrolepis exaltata (1.) Schott—F3-Herbaceous

PONTEDERIACEAE

(Mart) Solms—F1-Woody

POTAMOGETONACEAE

Potamogeton crispus L.—F2-Woody

PRIMULACEAE

Anagallis arvensis |.—F2-Herbaceous

PSILOTACEAE

Psilotum nudum (L) P. Beauv.—F3-Herbaceous

PTERIDACEAE

LAH +l |

3-Herbaceous

Pteris multifida Poir.—F3-Herbaceous

Pteris vittata _—F3-Herbaceous

RANUNCULACEAE

Adonis annua L.—F3-Herbaceous

Clematis terniflora DC.—F2-Herbaceous WATCH

syN= Clematis dioscoreifolia Levl. & Vaniot

Consolida ajacis (L.) Schur—F2-Herbaceous

sw Delphinium ajacis L.

syn= Delphinium ambiguum L.

r lio H +alle {| Z VC el J FA Herb AFA

"es E oll "ee a P di ud

sYN= Delphinium orientale J. Gay

Ranuncus ficaria |. —F3-Herbaceous

Ranunculus Het lira var. trachycarpus (Fisch. & Mey.)

Arn Ni eroac

Es L

Ranünculus muricatus | —F2- Herbaceous

Harum parviflorus L.—F2-Herbaceous

E? He rha

Liza Daceous

Ranunculus repens L. —F2- Herbaceous

Ranunculus sardous Crantz—F2-Herbaceous

SYN= Ranunculus parvulus

E?_Warharaniic

PITA Vales

RHAMNACEAE

Hosen. Bud: Thunb.—F3-Woody

Paliurus spina-christi P. Mil. —F3-Woody

Ziziphus zizyphus (L.) Karst —F3-Woody

syN= Ziziphus jujuba P. Mill.

ROSACEAE

ne Alchemilla microcarpa Bolss, & ies

nr E, | F2 He arhkaraniic

cta | —F3-Herbaceous

i ody

Prunus persica (L.) Batsch—F3-Wood

Pyracantha coccinea M. Roemer—F3-Woody

Pyracantha fortuneana (Maxim.) H.L. Li-——F3-Woody

986

ya TO NS ee i y

Pyrus calleryana Dcne. ap -Woody

Pyrus communis L—F3-Woody

Rosa bracteata J.C. Wendl.—F1-Woody

Rosa eglanteria L.—F3-Wood

Rosa laevigata Michx.—F2-Woody

Rosa micrantha Borrer ex Sm.—F3-Woody

Rosa multiflora Thunb. ex Murr.—F1-Woody

Rosa tomentosa Sm.—F3-Woody

Rubus bifrons Vest ex Tratt. —F3-Herbaceous

RUBIACEAE

Asperula arvensis L.—F3-Herbaceous

Cruciat | tana (Bellardi) Ehrend.—F2-Herbaceous

SYN— Sam Pe dd (Bellardi) All.

Galium aparine L = Heacsous

Paederia foetida | —

syN= Paederia EN. Sur Merr.

Richardia scabra L.—F2-Herbaceous

Sherardia arvensis L.—F2-Herbaceous

Spermacoce verticillata |.—F3-Herbaceous

syn= Borreria verticillata (L) G.F.W. Mey.

ACEAE

Citrus aurantium L—F3-Woody

Poncirus trifoliata (L.) Ra.—F2-Woody

Ruta eint nsis L.—F3- e

Ruta gr s L.—F3-Herbaceou

Triphasia afolia (Burm. f.) P. Med

SALICACEAE

Populus alba L.—F3-Woody

Populus nigra L—F3-Woody

SYN= Populus ales Du on jon

Woodv

WE CF CAE P

SYN= - Salix babylonicoia auct. non iE p.p.

SALVINIACEAE

Salvinia ism Mitchell —F1-Woody

Salvinia minima Baker—F2-Woody

SAPINDACEAE

Cardiospermum halicacabum L.—F2-Herbaceous

Cupaniopsis anacardioides (A. Rich.) Radlk.—F3-Woody

Koelreuteria elegans (Seem.) A.C. Smith—F3-Woody

syn= Koelreuteria paniculata Laxm.

TERORIS LARIACEAE

Hor E2_Warkharaniic

Wettst. Goat FTL RG ee ded

Bellardia trixago (L) All—F2-Herbaceo

Chaenorhinum minus (L.) Lan A”

Kickxia elatine LL Dumort. MA Pina eous

Li hil Wadw

ius TW

Linaria vulgaris ?. Mill.—F3-Herbace

Lindernia crustacea (L) F. Mel baaa

Mazus pumilus (Burm. f.) Steenis—F2-Herbaceous

Mazus japonicus (Thunb) Kuntze

Parentucellia viscosa (L.) Caruel —F2-Herbaceous

Verbascum blattaria |. —F2-Herbaceous

Verbascum thapsus L.—F2-Herbaceous

I £ al Dos * IP LJ st, A

f Texas 3(2)

Verbascum virgatum Stokes—F3-Herbaceous

Veronica agrestis |.—F4-Herbaceous

Veronica arvensis L.—F2-Herbaceous

Veronica persica Poir.—F2-Herbaceous

Veronica polita Fries—F2-Herbaceous

SIMAROUBACEAE

Ailanthus ideis (P. Mill) Swingle—F1-Woody

SOLANACEAE

Cestrum diurnum L.—F3-Woody

Cestrum nocturnum L.—F3-Woody

Cestrum parqui |'Hér—F3-Wood

Datura inoxia P. Mill. —F3- o

Datura stramonium L.—F2-Herbac

Lycianthes asarifolia (Kunth €: Saa Bitter—F3-

Herbaceous

Lycium barbarum L.—F3-Woody

Nicotiana glauca Graham—F2-Woody

Nicotiana longiflora Cav.—F4-Herbaceous

Nicotiana tabacum L. —F3- He naera

Petunia integrifolia (Hook.) Schinz: à Thellung— F3-

Herbaceou

SYN-- zetunia Violacea na

Phy F4-Herbaceous

SYN= Physalis ixocarpa Horn

ages origanifolia dam) al = alesis

cunt ai P + Dir EA. Da Pa

r^a-nerJdavccuu

Canl + : Cel E2 Harhacani<

Solanum capsicoides All. —F3-Herbaceous

Solanum diphyllum L.-—F3-Herbaceous

Solanum erianthum D. Don—F3-Herbaceous

Solanum ly var. cerasiforme (Dunal) Spooner

et al. -Fa Herbac

syn= Lycopersicon pim var. cerasiforme (Dunal)

lef.

Solanum lycopersicum Lam. var. lycopersicum—F3-

erbaceous

n= Lycopersicon esculentum P. Mill.

Solanum pseudocapsicum a Herbaceous

ES Herbaceous

3-ricrDaccuu

LWarkharantric

FS . L.utt£ E- | Ema

a ARAS E]

Solanum viarum Dunal—F1-Herbaceous

SPHENOCLEACEAE

Spl | lani a F3-Herbaceous

STERCULIACEAE

Firmiana simplex (L) W. Wight —F3-Woody

Melochia corchorifolia |.—F2-Herbaceous

TAMARICACEAE

Tamarix aphylla (L.) Karst —F1-Woody

marix chinensis Lour.—F1-Woody

including T. pentandra Pallas, T. ramosissima Ledeb.

Tamarix gallica L.—F1-Woody

including T. canariensis Willd., T. africana Poir.

Tamarix parviflora DC.—F1-Woody

r M | dial * H Li J £ at

Nesom

THELYPTERIDACEAE

othelypteris torresiana (Gaud.) Ching —F3-

Herbaceous

syN= Dryopteris uliginosa (Kunze) C. Christens.

sYN= Thelypteris torresiana (Gaud.) Alston

THYMELAEACEAE

Thymelaea passerina (L.) Coss. & Germ.—F3-Herbaceous

ULMACEAE

Ulmus parvifolia Jacq.—F3-Woody WATCH

Ulmus pumila L —F1-Woody

URTICACEAE

Boehmeria nivea (L) Gaud.—F4-Herbaceous

Parietaria judaica L.—F4-Herbaceous

Pilea microphylla (L.) Lieb.—F3-Herbaceous

Urtica urens L.—F3-Herbaceous

VALERIANACEAE

Valerianella locusta (L.) Betcke—F3-Herbaceous

1 REF NAC EPAD

Clar | F3 Waocw

woody

indui indicum j E y

Duranta ere 3-Woody

SYN= pierre repens Ly var. alba (Mast.) Bailey

sYN= Duranta repens var. repens

Glandularia xhybrida (Grónland & Rümpler) Nesom &

Pruski—F4-Herbaceous

ICAd5

987

syN= Verbena hybrida Grönland & Rümpler

Glandularia pulchella (Sweet) Troncoso —F2-Herbaceous

sYN= Verbena pulchella Swee

SYN= Verbena tenuisecta Briq.

Lantana camara L—F2 "WD ody —

sYN= Lantana laa Link & Otto

= Lippia montevidensis Spreng.

Verbena sonanensi; L. es eo

AFA TS"

Verbena litoralis Ku AFINE DAENS

Verbena rigida Spreng.—F2-Herbaceous

Vitex agnus-castus | —F2-Woody WATCH

Vitex negundo L.—F3-Woody

OLACEAE

Viola tricolor L.—F3-Herbaceous

VITACEAE

Cayratia japonica (Thunb.) Gagnepain—F3-Herbaceous

ZYGOPHYLLACEAE

Kallstroemia maxima (L.) Hook. & Arn.—F3-Herbaceous

Peganum harmala L.—F2-Herbaceous

Tribulus cistoides |.—F4-Herbaceous

Tribulus terrestris |.—F2-Herbaceous

Zygophyllum fabago | —F3-Herbaceous

APPENDIX 2

Non-native species in Texas ranked as F1. For aquatic species, the number of counties of occurrence is indicated in Square

ackets

Ailanthus altissima

Tamarix aphylla

Lonicera japonica

T. ramosissima

F1- HEPBRACEOI Ic FA

Tamarix chinensis (incl. 7. pentandra,

)

F1-WOODY (Trees, shrubs, subshrubs, and woody vines)

Tamarix gallica (incl. 7. canariensis, T.

africana)

Tamarix parviflora

Triadica sebifera

Ulmus pumila

Wisteria sinensis

J pE LL L J 4

Atriplex semibaccata

Bassia scoparia

Bothriochloa ischaemum var.

Carduus nutans subsp. macroce- Lolium perenne

phalus

Centaurea melitensis

Cynodon dactylon

Cyperus entrerianus

Melilotus albus

Alternanthera philoxeroides [20+]

Colocasia esculenta [11

Eichhornia crassipes [25]

Dichanthium annulatum

Dichanthium aristatum

Dichanthium sericeum

songarica Eragrostis lehmanniana

Lygodium japonicum

Marrubium vulga

Melilotus indicus

F1-AQUATIC

Hydrilla verticillata [6]

Myriophyllum aquaticum [18]

Myriophyllum spicatum [10]

Melilotus officinalis

Salsola kali subsp. pontica

Salsola tragu

re Solanum viarum

Sorghum halepense

Urochloa maxima

Nasturtium officinale [35]

Pistia stratiotes [10 or more]

Salvinia molesta [10]

988

m.a b In LI PEPA af

APPENDIX 3

f Texas E]

Non-native species in Texas ranked as F2. For aquatic species, the number of counties of known occurrence is indicated in

square brackets.

Albizia julibrissin

Broussonetia papyrifera

Hypericum perforatum

Lagerstroemia indica

Lantana camara

Aegilops cylindrica

Aira caryophyllea

Alternanthera caracasana

Amaranthus itum

jus

Pole arvensis

peas cotula

aie iin

Aon. tha

Arenaria batte

Buglossoides arvensis

upleurum rotundifolium

Camelina microcar

Cardamine hirsut

Cardiospermum halicacabum

Carduus amd alus

Centaurea cyanus

a rium muhlenbergeii

ellum

opodium murale

erae

Ch ga

Chorispora tenella

Cichorium intyb

Cirsium vu

i Igare

Clematis terniflora

Commelina communis

Conium maculatum

F2-WOODY (Trees, shrubs, subshrubs, and woody vines)

orus alba

Nicotiana glauca

Phyllostachys aurea

Poncirus trifoliata

Pueraria montana var. lobata

F2-HERBAC (A

Pyrus calleryana

Rosa laevigata

Vitex agnus-castus

Conringia orientalis

Consolida ajacis

roton argenteus

Cruciata One

Cucumis m

melo

Cuphea carthagenensis

Cyclospermum leptophyllum

Cyperus difformis

Cyperus involucratus

Datura stramonium

in ischaemum

Echinochloa colon

Saba crus-galli

Eleusine in

Elymus hispi ids

Eragrostis barrelieri

Eragrostis e

Eragrostis curvula

Eragrostis plos var. p

Erodium cicutar

Eruca vesicaria subsp sativa

Gamochaeta coarctata

Gamochaeta ina

Geranium diss

Gibasis pellu a

Glandularia pulchella

Gomphrena globosa

Hedera helix

Hedypnois cretic

Heliotropium dl

Heliotropium indicum

latum

Hordeum murinum subsp. leporinum

jaa ir brasiliensis var.

iei

ined glabra

Hypochaeris microcephala var.

Ibiflora

Hypochaeris radicata

a carnea

Lathyrus hi hirs sutu us

didis parviflor

Melinis

ipii acanthium

Oryza s

| mi all bal :

Nesom

Oxalis articulata

Panicum coloratum

Parapholis incurva

Parentucellia viscosa

Paspalum dilatatum

Paspalum notatum

Pennisetum villosum

Perilla frutescens

Petrorhagia dubi

Phyllanthus urinaria

Plantago lanceolata

Plantago major

Poa annu

Polycarpon tetraphyllum

Polygonum arenastrum

Polygonum aviculare

Polygonum piel

Polygonum persic

Polypogon onodi

Polypogon viridis

Prunella vulgaris

Pseudognaphalium luteoalbum

Ranunculus muricatus

Ranunculus parviflorus

Ranunculus repens

Ranunculus sardous

Ceratopteris thalictroides [3]

Cryptocoryne beckettii [1]

Egeria densa [3]

Hydrocleys nymphoides [2]

Hygrophila polysperma [2]

Landoltia punctata [6]

Scabiosa atropurpurea

Sclerochloa dura

Scorzonera laciniata

Senecio vulgaris

dien eio

Setari

porra arvensis

Solivia anthemiifolia

Soliva sessilis

Soliva stolonifera

Sonchus asper

Stellaria pallida

Taraxacum laevigatum

Taraxacum officinale

F2-AQUATIC

Limnophila sessiliflora [2]

diflor

Ludwigia grandiflora subsp. gran-

a [3]

Ludwigia grandiflora subsp. hexa-

etala [9]

Nomaphila stricta [1]

APPENDIX 4

Thiaspi arvense

Torilis arvensis

Torilis nodosa

Tribulus terrestris

Trifolium dubium

Trifolium incarnatum

Trifolium em

Trifolium repens

Trifolium resupinatum

Trifolium vesiculosum

Verbascum thapsus

Verbena bonariensis

Verbena brasiliensis

Verbena rigida

Vicia sativa nigra

Vicia tetrasper

Vicia villosa a villosa

Xanthium spinosum

Youngia japonica

Nymphoides indica [1]

Potamogeton crispus [8]

Salvinia minima [2]

Xanthosoma sagittifolium [1]

Watch List: non-native species in Texas potentially ranked as F1. All F2 aquatic species are included.

Hypericum perforatu

Pueraria montana var. pre

Ardisia crenata

pedal equisetifolia

nam

os angustifolia

sind hae iced

Cirsiu

mata tern jee

Lespedeza cuneata

From F2-WOODY:

Pyrus calleryana

F F3-WOODY:

Leucaena leucocephala

Lonicera maackii

Macfadyena unguis-cati

Photinia serratifolia

From F2-HERBACEOUS:

onum arenastrum

num persicaria

e floridana

Perilla frutescens

dee =

Vitex agnus-castus

Pistacia chinensis

Schinus molle

Schinus terebinthifolius

Ulmus parvifolia

Torilis arvensis

Verbena brasiliensis

989

990

Cryptostegia grandiflora

Ceratopteris thalictroides

Cryptocoryne beckettii

geri

Hydrocleys nymphoides

Hygrophila polysperma

Landoltia punctata

Dos * in LE d 5 dion ade

From F3-HERBACEOUS:

Imperata cylindrica

from F2-AQUATIC:

Limnophila sessiliflora

Ludwigia grandiflora subsp. gran-

diflora

Ludwigia grandiflora subsp. hexa-

Nomaphila stricta

[| eal E L I

Microstegium vimineum

Nymphoides indica

Xanthosoma sagittifolium

af E |

Texas:

Caesalpina bonduc

Canavalia rosea

` Conocarpus erectu us

Laguncularia racemosa

Rhizophora mangle

Scaveola plumieri

Cassytha filiformis

ACKNOWLEDGMENTS

The authenticity of this report, especially in the F1 species and Watch List but also in discussion of concepts

and other topics, owes to contributions from Bill Carr, Laura Hansen, Stephan Hatch, Eric Keith, Barney

Lipscomb, Andy McDonald, Michael McRoberts, Tom Patterson, Jackie Poole, Mike Powell, Nelson Rich,

Monique Reed, David Rosen, Jason Singhurst, Bruce Sorrie, Damon Waitt, and Justin Williams. Detailed

journal reviews by Jackie Poole, Damon Waitt, and Bruce Hoagland were valuable in refinement of the

ranking protocols—and the detailed commentary by Jackie Poole on everything from A to Z is appreciated

beyond expression.

REFERENCES

APRS IMPLEMENTATION Team. 2000. Alien plants ranking system (Version 5.1): A cooperative effort of the National

Park Service, Ripon College, University of Minnesota, and the U.S. Geological Survey. eid pelle Aa E

Research Center Online, Jamestown, North Dakota. <http:// | gs.g |

index.htm> (Version 30SEP2002).

Austin City CONNECTION, 2009. Invasive plants to avoid. City of Austin. <http:// int growgre

invasive.htm»

CARLSON, M.L., I.V. Lapina, M. SHEPHARD, J.S. CONN, R. Densmore, P. SPENCER, J. Heys, J. RILEY, AND J, NIELSEN 2008. |

ing system for non-native plants of Alaska. U.S.D.A, Forest Service Alaska Region R10-TP-143. Anchorage, Alaska.

Corrett, D.S. AND M.C. JOHNSTON. 1970. Manual of the vascular plants of Texas. Texas Research Foundation, Renner,

Texas.

Hatch, S.L., K-N. GANDHI, AND L.E. Brown. 1990. Checklist of the vascular plants of Texas. MP-1665, Texas Agric. Expt.

Station, Texas A&M Univ., College Station.

HEFFERNAN, K.E., BP. COULUNG, J.F. TOWNSEND, AND CJ. Hutto, 2001.R

Heritage Technical Report 01-13. Virginia Department of Conservation and Recreation, Division of Natural

Heritage, Richmond, Virginia. <http://www.dcrvirginia.gov/natural_heritage/docu ments/rankinv.pdf>

INVADERS OF TEXAS. 2009. Invaders of Texas: a citizen science program to detect and report invasive species. <http://

Lina inves tic plant species in Virginia. Natural

JOHNSTON, M.C. 1990. The vascular plants of Texas: a list, up-dating the ‘Manual of the vascular plants of Texas:

Second edition. Published by the author.

JoRDAN, M.J. G. Moore, AND T.W. WeLoy. 2009. New Mo eis Pant ranking ll for evaluating non-native plant

species for invasiveness (DRAFT). <http://nyis tAssess /Assets/NY_Invasive_Plant_Ranking_

Intro.pdf»

Morse, L.E., J.M. RANDALL, N. BENTON, R. HiEBERT, AND S. Lu, 2004. An invasive species assessment protocol: evaluating

non-native pana for their ee on BOE Version 1. NatureServe, Arlington, Virginia. «http://www.

SS SO E f 23V OOTP Iriel Protocol. pdf»

Nesom, Fund tal i i index f ti ies of Texas 991

Nesom, G.L. 2000. Which non-native plants are included in floristic accounts? Sida 19:189-193.

NESOM, G. " 2009a. Non- "Rae species rc in a reported since 1970. Documentation for the full list.

is G E: 2009b. Notes on non-native mo in A Texas Phytologia 91:325-332.

NTWGS. 2009. Prohibited plant species. North Texas Water Garden Society. «http//www.ntwgs.org/articles/

illegalAquatics.htmlitthe9620list9620»

TENNESSEE EPPC, 2001. Tennessee invasive exotic plant list. Research Committee of the Tennessee Exotic Plant Pest

Council. «http://www.tneppc.org/Invasive Exotic Plant List/Invasive9620List9620Introduction.htm»

TDA. 2009. Noxious plant list. Texas Department of Agriculture. «http;//info.sos.state.tx.us/fids/04. 0019 0300-1.

html»

TEXASINVASIVES.ORG. 2009. Invasive Plant Database. <http:// ‘invasives_database/>

TPWD cad Pronibited ids ipeeles Jus TOME and Wildlife Department. Austin: Texas. <http://www.tpwd.

qM BL, A Meda G. DENE AND ipo Donon: 2003, Atlas of the vascular plants of Texas. Vol. |, Dicots; Vol. II,

Monocots. Sida, Bot. Misc. 24, 1 and 2.

USDA, NRCS. 2009. The PLANTS Database. National Plant Data Center, Baton Rouge, LA 70874-4490 USA. <http://

plants.usda.gov/»

WARNER PJ., C.C. Bossanp, M.L. Brooks, J.M. DiTomaso, J.A. Hatt, A.M. HowaLb, D.W. JOHNSON, J.M. RANDALL, C.L. Rove, M.M.

RYAN, AND A.E. STANTON. 2003. Criteria for categorizing invasive non-native plants that threaten wildlands. Cali-

fornia Exotic Pest Plant Council and Southwest Vegetation Management Association. <http//www.cal-ipc.

org/ip/inventory/pdf/Criteria.pdf>

Postscript.—Since preparation of this report, various minor changes have been made to the lists. The

whole set of information on Texas non-native plants, including these changes and additional commentary,

is posted on the internet—see “Non-natives” at <http://www. euro som: ea o nen

The internet information is frequently updated as new data

In order to reflect the input of Texas botanists to the genesis and development of the information, the set

of online data is cited as below.

Nesom, G.L., J.L. APLACA, W.R. CARR, N.L. Fow er, L.L. HANSEN, S.L. Hatch, B.W. HoacLano, W.C. HoLmes, E.L. Kerry, B.L. LIPSCOMB,

B.R. MacRoserts, M.H. MacRoserts, J.A. McDonaLo, T.F. PATTERSON, J.M. Poore, A.M. Powe, N. Rich, M.D. Reep, DJ. ROSEN,

J.R. SincHursT, B.A. Sorrie, B.L. Turner, D.E. Warrr, AND J.K. WiLLiAMs. 2009. Non- mats oan dias Overview of

occurrence and invasiveness assessments. <http:// guy tiveMENUweb.htm

The total number of non-native plants species in Texas now is 820 and a relatively small number of rankings

have been modified. The Watch List has been expanded with additions of these: Acroptilon repens, Carduus

acanthoides, Cayratia japonica, Colocasia esculenta, Cortaderia selloana, Dioscorea bulbifera, Elymus repens,

Gibasis pellucida, Lespedeza bicolor, Lolium li , and Pyracantha koidzumii. As with many of the F1

and F2 species, some of the Watch List species already are so widespread that it is unlikely that they can

be eradicated or even controlled except by sustained efforts on local levels. From among the Watch List

species, a subset is indicated as a "Super Watch List,” those species that perhaps can be removed from the

Texas landscape before they become impossible to control. A newly added Expected List records species

not currently known with certainty in Texas but that occur in nearby areas—each of these is aggressively

invasive and Dac damaging, and there is a high probability that each soon will reach Texas: Alliaria

petiolata, Ampelopsis 1 ta, Berberis thunbergii, dini nui als stoebe ssp. micran-

thos, Centaurea solstitialis, Cirsium arvense, Dioscorea alata, E bellata, Euonymus

alatus, Euonymus fortunei, Euphorbia esula, Halogeton glbmeratus, Kona morrowii, coda micron lium:

Melaleuca quinquenervia, Miscanthus sinensis, Phalaris aquatica, Pennisetum setaceum, Polygonum cuspidatum,

Schismus arabicus, Setaria faberi, Typha angustifolia, and Wisteria floribunda.

992 t i exas 3(2)

BOOK REVIEW

Tl Doa a Ph | RT

Tatyana A. LOBOVA, CULLEN K. GEISELMAN, AND SCOTT A. MORI 2009. Seed Di

(ISBN 978-089327-501-3, hbk.). Memoirs of The New York Pta En vol. 101. The New ork

Botanical Garden Press, 200th Street and Kazimiroff Boulevard, Bronx, New York 10458-5126, U.S.A.

(Orders: www.nybgpress.org, nybgpress@nybg.org, 1-718-817-8721, 1-718-817-8842 fax). $70.00,

492 pp., 24 b&w illustrations + 32 color plates with more than 200 photographs, 7" x 10".

A OM HRS | :11 s 1 nA.) 1D 1 Cal 11 í mL 11 ^ F1 + r

el [^] r ES aa] + i

fT ils : LEY +l I 1 i| a} 1 1 1 1 4A xl T ` 1 J1 + ] + AL

VT VV y LES pees a 1

] JA 1 1 1 1 1 Eod Dl " f. ] +1 1 pa |

& I 4 Ji pn ed ie

à tüersupb ee well-cited literature ees EN m topi, as well detailed t of field work and tl Its of tl l that

follow. I g y interested i imal-plant interactions, ME asan dam ion to

1 : g Tf ake : f thi RAO 1 + = +1 A +

1:1

the a Aes

There : are five m main sections, consisting o of an introduction, conclusion, materials pe rid. and the two largest sections:

, and

a [94

id I hdi ion of each frugi I i ized by subfamily), including not “Diet Specificity, Seasonality,

and Selectivity”, “Foraging Strategy”, and “Results for French Guiana”. The Plants dispersed by bats section contains detailed plant

treatments (sorted by family) divided into “Bat popesen Hune in Pili Puede d " e Posto Bat bano AKA dern

in del both d contain ERE desc Bptions. Į I

E 1 14 1 [- 1 k : m 1 1 a E 1 FA -l

EE: I _ t E el VE

TIT’ +l 1 ga 11 1 FA S. TT A Je TT 4] f 1 1 in a | ixl 32

LL? a Io

the Literature Cited s section. There is a : page glossary that assists tl ] ith the botanical and dispersal inology referred to

T J :] 11 £l 1 4 iy 1 1 s 1 +l 11 s

[m] IL LI 4 2

housed.

The mcum aod an excellent job n surveyin a existing literature on the topic of bat dispersal, with easy to interpret

"ET g E graphic [ f this sort. I terials and methods section, we find

that the study is supp | by the long time botanical work at the study site of hor Scott Mori, among others. There are beautifully

ead ns lat — hs of infruct , fruits, and ds of y of the families d ibed in the text, which

will likely be of articulo aid in identification. T ] Jude both nat d cul ] pl tl y, indicating

this status in the description f the di iesi ing the literat ER differen Mises ies were pd

[e]

paar were ca or dees taxonomic concepts were applied. 1 Perhaps tl i EI

e [ul 1 a J A 1 1 x1

r E 3). H mum th d 4 rod

ainia

The authors analyzed fleshy fruited species that letermined suitable f imal consumption, and included the following

] t fi lvsi l life fi d habit i l stat fruit type. inf iti frui ti d ition

Pi I , > fr z i Ooo o i

hats teed ] f tl dible part part digested. diaspore. diaspore size 11 1 lt Otl 1 hat

> r d I 1[ e 2 I 1 I E L 2r

discussed, | 1 1 lud t t t ] trit ] tent

"Tl 1 Ve 7 +l El 1 J - 1.:1 NN ADAD UA fl 1 1 gl 1

El " FL e

f. lis: A Jd nd leche 1: E. 1 1 " r : 1+] £41 +1

1 ng, there ALR A IC yy HEIL VILE Lik vv 1 b Pp X F J UCLU thing for

students seeking projects in this field). TI lt pared to tl found at ot] ly sit tl tropics, gh tl

dup] ` FAL uf E + » pP ri H 1 4 ` ¿l m p ne

study with which they could provide any I ( ying g data)

Barro Colorado Island in Panama

lem 1 1 ex 1 A ca E fe D 1 E . A 2 FL. 1 1 = 1 a FR TX

4 E L

(Pepper Family y), {DL :1 1 À Cc 11578 Cy 1 4l iT Trhu ery :1.. 0,1 mm TT 13. x AA 'à Ww smi

inm) ss Clusiaceae (Mangosteen que It ggested that ol d diff 1} he rel of

J I 1 11 + 4-. They g 1 I 1 de 1 I ID 1 by bats to

1 ] 1 1 quc ud rom 1 f.1 1 " 1 EH :1: 1 - 4 EL 1 p. + 1 J I1

e i z i

year, are prtasrtly n iin al bats, ! fruit that ft and defense-l itl Itip] ds or | ingle fruits with

single seeds, d tl that y to swallow and pass.

This book i ite logical in its ] t and tl to t] f it is. but the conclusions the study reaches elucidate

L o P P > y

man y for furt! ly anc critical thought, 1 scientifi ject ought. Iti y marriage ale eee zo ogy.

ki f al ical ti 1 fiar an EM M a necessary reference a y ying plant-anima ]

interactions, Tiana Franklin, Botanical Recani Institute of Texas, Fort Worth, Texas 76102-4025, U.S.A.

J. Bot. Res. Inst. Texas 3(2): 992. 2009

ADDITIONAL FAMILY ACRONYMS FOR VASCULAR PLANTS

BASED ON APG III

Neil Snow

Herbarium Pacificum

Bi useum

1525 Bernice Street,

Honolulu, Hawaii 96817, U.S.A.

neil snow@bishopmuseum.org

ABSTRACT

Six additional 3-letter famil y y pror 1 for vascular plants.

RESUMEN

D Em T 1 -. F ER 3 1 4 1

The recent publication of additional 3-letter family bbreviations for families of vascular plants

(Snow 2009) occurred before the author was aware of ^ additions and suggested changes that appeared

in APG III (2009). The six acronyms are necessary given earlier omissions (e.g., Alford 2006) and the desire

by certain curators to use other family names for which such acronyms are not yet available. The proposed

acronyms are:

Anacampserotaceae (ACM)

Boweniaceae (BOW)

Gerrardinaceae (GRR)

Maundiaceae (MAU)

Montiaceae (MON)

Veronicaceae (VER)

Future authors of new families of vascular plants are strongly encouraged to propose new 3- and 4-letter

acronyms (see Kiger & Reveal 2000, 2009) for each new family along with the family itself, so that future

updates of the present type are unnecessary.

ACKNOWLEDGMENTS

I thank Kent Perkins (FLAS) and Dieter Wilken (SBBG) for their comments. As previously, the Colorado

mannan o E 1. PAE 1

Native Plant Society has kindly rodea to post on-line e | y by family and (sepa-

rately) by 3-letter acronym (see: http://ww ps.org/pdi/Plant%20Key /ACROS pdf)

—

REFERENCES

ALFORD, M.H. 2006. Gerrardinaceae: a new family of African flowering plants unresolved among Brassicales,

Huerteales, Malvales, and Sapindales. Taxon 55:959-964

APG (AuaiosPERM PHyLoGEeNy GROUP). 2009. An update of the Angiosperm Phylogeny Group classification for the

orders and families of flowering plants: APG lli. Bot. J. Linnean Soc. 161:105-121.

KIGER, R.W. AND J.L. ReveaL. 2000. A comprehensive scheme for standardized abbreviation of usable plant-family

names and type-based suprafamilial names. Huntia 11:55—84.

Kicer, R.W. AND J.L. ReveaL. 2009. A comprehensive eene for iei as A vi ids pars

names and type-based suprafamilial names. http:// | y g/reveal/pbio/fam/famabbr.

html [Accessed 4 November 2009].

Snow, N. 2009 Twenty-eight new three-letter family acronyms for vascular plants (with comprehensive listings

on-line). J. Bot. Res. Inst. Texas 3:273-277.

J. Bot. Res. Inst. Texas 3(2): 993. 2009

nm a n L ] Md. den nde Si”

994 Joumal of tl t l f Texas 3(2)

ANNOUNCEMENT

APPLICATIONS FOR THE 2010 DELZIE DEMAREE TRAVEL AWARD

The Delzie Demaree Travel Award provides financial support for graduate students to travel to the Annual

Systematics Symposium held every year at the Missouri Botanical Gardens in St. Louis, Missouri. Typically

one $300 award is granted per year, but value and number of awards may vary depending on availability

of funds and number of total applicants. BDDICASOHS for the 2000 Delzie Demaree Travel Award should

include a letter from the applicant telling how sy will benefit his/her graduate work and

a letter of recommendation sent by the major K Please send letters of application to: Dr. Donna M.E.

Ware, P.O. Box 8795, Herbarium, Biology Department, The College of William and Mary, Williamsburg, VA

23185-8795, U.S.A. 1-757-221-2799; Email: ddmware@wm.edu. The period for receiving applications will

end three weeks prior to the date of the symposium if a sufficient number of applications are in hand at that

time. Anyone wishing to apply after that dace should inquire whether applications are still being accepted

before applying. The Systematics Symposium dates for 2010 are 8—9 October.

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 nid he m yet most as a botanist was assisting students with their field botany research.

d in making a contribution to Delzie Demaree Endowment Fund, which supports the

travel nnd 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: barney@brit.org. Thank you.

J. Bot. Res. Inst. Texas 3(2): 994, 2009

IN MEMORIAM

BRUCE DALE PARFITT, 7 NOVEMBER 1952-3 SEPTEMBER 2009

Neil A. Harriman

iology Department

University of Wisconsin-Oshkosh

Oshkosh, Wisconsin 54901, U.S.A.

harriman@uwosh.edu

Botanist, biologist, birder Bruce Dale Parfitt of Johnson, Vermont, died

at Vermont Respite House in Williston, 3 September 2009. He was 56.

Bruce was born 7 November 1952 in Oshkosh, Wisconsin, the son of

Dale and Joan (neé Barth) Parfitt. From age four during family vaca-

tions, he fished, hunted, and camped in Oconto County in the land of

the jack pines and sweet ferns “up north.” He graduated from Oshkosh

High School in 1970, and earned a bachelor of science degree from

the University of Wisconsin-Oshkosh in 1977. He earned his masters

and doctoral degrees from Arizona State University in 1980 and 1991,

where he was also herbarium curator and teaching associate. For both

degrees, he worked closely with Professor Donald Pinkava, who was

instrumental in encouraging Bruce’s interest in cacti.

At the University of Michigan-Flint he was a valued faculty member

for 14 years, chair of the biology department from 2004-2007, and

director of the university’s herbarium, whose collection grew in size

and value under his care. He kept high standards, edited papers relentlessly, challenged assumptions, but

lavished his time toward students’ success in the classroom, in the lab, in the field, and in their personal

lives. As a result, many became first—generation graduates who found confidence and success in biology—

related careers or entered Master’s and Ph.D. programs at top universities.

Prior to joining the faculty of UM-Flint, he was a research botanist for the Desert Botanical Garden in Phoe-

nix, and a scientific editor of “Flora of North America” head tered at the Missouri Botanical Garden. He was

a significant contributor to the Ranunculaceae, volume 3, 1997, and the Cactaceae in volume 4, 2003. He also

prepared Opuntia for the Jepson Manual, 1993. He authored some 40 papers in the literature, beginning with

a paper on Allenrolfea (Chenopodiaceae) in Rhodora, 1977, while he was an undergraduate at UW-Oshkosh.

Among his friends, he was an avid birder, moose watcher, canoeist, Mr. Fix-it, and hunter and planter

of trees. He was as tenacious in his friendships as in his opinions. He seemed to apply a scientific approach

whether decorating Christmas cookies or designing birdhouses.

And since 1997, he came to love Vermont, first visiting, then buying a place and staying as often as he

could at his “Mooseberry Camp” on the Lamoille River in Johnson.

It was while working as a field biologist in 1979 for the U.S. Bureau of Land Manag t—helicoptered

in to hike the remote Hualapai Mountains and to collect, identify, and preserve rare species—that he dis-

covered a new species, named by Barbara di in 2009 as penal demotica. It was also in the Hualapai

that Bruce developed a blood vessel malf that is spinal column. An unfortunate

reaction to diagnostic tests cost him the use of his legs. Undaunted in in 1980, he learned to walk with canes.

Similarly, throughout his life he cften took on projects of considerable magnitude— building a house, plant-

ing trees, restoring riverbanks, and the like.

In late July, 2009, Bruce and hi ther invited my wife and me to lunch at an Oshkosh restaurant. Dur-

Courtesy, The University of Michigan-Flint

J, Bot. Res. Inst. Texas 3(2): 995 — 996. 2009

996 || || FN FI H | | L I "m FI f Texas 3(2)

ing lunch, he told us that he had come to say good-bye. The doctors had exhausted their armamentarium of

chemotherapies, and surgery would only delay the inevitable end. Bruce and I went that afternoon to watch

the American White Pelicans at a tiny lake nearby; the species was not present in Wisconsin when Bruce

was such an avid local birdwatcher. The pelican in myth and folklore represents loyalty and self-sacrifice,

a most fitting ae ane to Bruce Dale en

CTh mlam n m n n pe a Al ines ado Y e Y

Editors Note. S DI Z¿31Z).

HOWARD L. CLARK, 1941-2009

Kate Clark, wife of Howard L. Clark (Quito, Ecuador), emailed me with sad news that Howard died

on 27 August 2009. Howard was a friend and colleague always willing to help with reviewing uos

“I always will be glad to review papers or anything else for the exSida, as I received many of the first issues

as an undergraduate (student of John W. Thieret, then at Univ. SW Louisiana, Lafayette) for "free" after

making a deal with Lloyd Shinners to trade my plant collections for Sida—1 probably never sent enough to

be adequate. My first botanical publication was in Sida. Yes, that was a few years ago, long before I moved

to the amphiscian realm (as John Thieret phrased it)."

I contacted Howard on June 25, 2009, seeking a review of a manuscript submitted to J. Bot. Res. Inst.

Texas. Of course Howard was more than willing to help and to give of his time (“exceptionally glad” were

his words). The resulting correspondence was like rediscovering a close friend from the past. We had a

good time catching up and reminiscing about our mutual friend John Thieret (his "professor, mentor, and

friend". ..and, 1 might add personally, my mentor and friend). We happily shared stories about Lloyd Shin-

ners of whom neither one of us met in person. Our friend John Thieret worked closely with Lloyd and Sida

from the 1960s to early 70s until Lloyd passed away in 1971. Howard said his old copies of Sida were safely

stored in Atlanta; he even remembered Professor Joe Ewan's poem to Lloyd Shinners in response to the first

issue of Sida (22 Nov 1962).

John W. Thieret (1926-2005) served as Howard’s major professor for his master's degree in Biology at

University of Southwestern Louisiana, Lafayette. Howard studied the Lemnaceae of Louisiana for his M.S.,

a project Howard started as an undergraduate. Howard and John published together in 1968 The duckweeds

[Lemnaceae] of Minnesota (Michigan Bot. 7:67—76

Over the last several years Howard had been living and working in Quito, Ecuador. He was working

mostly as an ecological consultant (increasingly, industrial stuff" as Howard put it), but his love was botany.

He had set aside for me a copy of a recent botanical publication of his (a Venezuela florula) that he was most

AJA

proud of.

CLank, H.[L.], R. Liesner, PE. Berry, A. FERNANDEZ, G. AYMARD, AND P. MAQUIRINO. 2000. Catálogo anotado de la flora del

área de San Carlos de Rio Negro, Venezuela. Scientia Guaianae (Caracas) 11:101-333 + 21 color photos. (103

illustrations by B. Manara).

“For 4.5 years (79—'81), we (my wife is an ichthyologist, but also a good botanist who worked for Joe Ewan

for a year at the Tulane herbarium) lived in San Carlos de Rio Negro, Venezuela, where the soil had a lot

of outwash quartz sands from the tepuis and from the Shield, and the flora had a lot of connection to the

strange stuff above us on the shield and tepuis. San Carlos is across the Rio Negro from Colombia, about 15

km south of the Rio Casiquiare, and about 60 km north of Brasil. It took many years, but I finally published

a florula/checklist (in a pub. more obscure than Sida) for there.” This florula is surely a testament to the

botanist Howard L. Clark.

I'm so sorry and my heart is saddened to lose a friend and colleague. Thanks, Howard, for all of your

contributions to science! —Barney L. Lipscomb, Botanical R h institute of Texas, Fort Worth, Texas 76102-

4025, U.S.A.

REVIEWERS:

_ VOLUME 3 (2009) _

188 reviewers:

Thank you for A of the Botanical Research Institute of Texas.

Alford, Mac H.

Allen, Charles M.

Allred, Kelly W.

Anderson, Gregory J.

Anderson, Loran C.

Aymard, Gerardo C.

Barrie, Fred R.

Bayrd, Venice

Bean, A.R.

Bell, Dennis

Block, Timothy

Bogler, David J.

Bohs, Lynn J.

Brown, Elizabeth

Brown, Larry E.

Bryson, Charles T.

Butler, Ivan Maureira

Butterworth, Charles A.

Carr, Gerald D.

Carr, William R

Carranza-Pérez, Miguel A.

rter, Richard

Cerros-Tlatilpa, Rosa

ambers, Kenton L.

Chandler, Gregory T.

Chautems, Alain

Christensen, Knud Ib

avi

Colin, Rafael Torres

Cowie, Ian D.

Cox, George W.

Crespo, Manuel B.

Cusick, Allison W.

Darbyshire, Stephen J.

Daskin, Ruziye

Davenport, L.J.

Davison, Paul

DiTomaso, Joseph

Doust, Andrew

Dressler, Stefan

Dusen, Olcay Dinc

Dutton, Bryan E.

Edwards, Christine E.

stes, James

Estrada-Castillón, Eduardo

Ezcurra, Cecilia

Ferguson, Carolyn J.

Frye, Christopher T.

Gandhi, Kanchi

ndhi, Thomas

Gelli de Faria, Ana Paula

Gepts, Paul

Mg NUN Farrokh

nchez, Maricela

Spr NN M. Socorro

Guaglianone, Encarnación R.

Gutierrez, Raul

Hansen, Bruce F.

Hartman, Ronald L.

Haston, Elspeth

Hatch, Stephan L.

Henrickson, James

Herrera Arrieta, Yolanda

Hiepko, Paul

Hill, L.M.

Hill, Steven R.

Hoagland, Bruce

Hogan, Tim

Holmes, Walter C.

Judd, Walter S.

Kaul, Robert B.

Kelly, Lawrence M.

KefSler, PJ.A.

Kirkbride, Jr., Joseph H.

Knapp, Sandra

Krosnick, Shawn

Kumar, E.S. Santhosh

Kumar, Muktesh

| Ret Rac Inet Tavac 3(2): 007 — 1008. 2009

Lammers, Thomas G.

Lorence, Lucas

MacDougal, John

MacRoberts, Barbara R.

MacRoberts, Michael H.

Morawetz x

Morgan, David

Morton, aie M.

Mueller, Gregory M.

Miller, J.

Naczi, Robert F.C.

Nath, Virendra

Nesom, Guy L.

Nilsson, Henrik

Olmstead, Richard G.

Oteng-Yeboah, Alfred

Pandit, M

Panero, "P L.

Patterson, Robert W.

Pennington, Terry

Penskar, Michael R.

Perez-Cortez, Silvia

Perkins, Kent D.

Pilz, George E

Plunkett, Gregory M.

Poole, Jackie

Post, Angela R.

Powell, A. Michael

Ramirez-Morillo, Ivón M.

andle, Christopher P.

Ranker, Tom A.

Reed, Monique D.

Reid, Christopher S.

Reveal, James L.

Robertson, Kenneth R.

Robinson, Harold

Rolfsmeier, Steven B.

Rosen, Davi

Saarela, Jeffery M.

adle, Jimi

Salazar, Jackeline

Sanchez-Ken, J. Gabriel

Santelises, mea Aguilar

Scharpf, Robert

Schilling, Edwar

Schneider, Angelo -

Schutzman, Bart

Serna, Adolfo Espejo

Sivinski, Robert C.

Smith, Latimore

Snow, Neil

Soriano, Sara Fuentes

Spaulding, Dan

Spooner, David M.

Stahl, Berti

Taylor, Alberto Sidney

Taylor, Charlotte M.

Telfair II, Ra

Terrell, Edward E.

Tiehm, Arnold

Tillett, Stephen S.

Tucker, Arthur O.

Urban, Lillis

Valdés-Reyna, Jesús

Vanderpool, Staria S.

Wada, E

Mera Warren L.

Waines, J.G.

Waitt, Damon

Ward, Daniel B.

Weber, William A.

Weedon, Ronald R.

Wege, Juliet

Wiens, Del

Wipff, J.

Wunderlin, Richard P.

998

[| [| £ al n.a a nm LI PE he all nia

f Texas 3(2)

INDEX TO 111 TITLES WITH 175 AUTHORS:

VOLUME 3 (2009

We know you have a choice. Thank you for choosing and supporting Journal of the Botanical Research Institute of Texas.

A Pad of geographic elements and analysis of

the flora of Big Bend region cf Texas by ANNA SAGHAT-

ELYAN— 1

A floristic iiie of Marco Island A County), Florida

R AND B.J. RocHE—3(2):87

A new cliff pibus e of s (Zamiaceae) from

Belize by MichagL CaLoNJE—3(1):23

A new combination in North American Tephrosia (Fabaceae)

y . NESOM AND JAMES L. ZARUCCHI—3(1):157

A new genus, Mexotis, for five Mexican species of Hedy-

otideae (Rubiaceae) by EDwARD E. TERRELL AND HAROLD

ROSE US

À i

)

MLN YY oP PRA TN Mv EF i A

l +

K | Arkansas

and taxonomic synopsis of the genus by AARON " FLODEN,

Mark H. MENDES a CAROLYN " EIGUSON cS cde

A |

_ Andes by L.J. PORA AND Bası Srensos—3(2) 605

tains ‘of Belize by MICHAEL CALONIE, JAN MEERMAN, PATRICK

GRIFFITH, AND GEOFFREY HoEse—3(1):3

A Allande of Arctostaphylos (Pricaceae) inferred from

lea A.W

—

THOMAS Panen a AND Mica C. Vase —3(2) 673

FRA |

aceae) by Wenny B. ZOMLEFER AND WALTER S. Juoo—3(1): 159

A revision to the vascular Le of Calcasieu Parish, Louisiana

oy Ray NETANE 10:37

A I bol | ADM

vasculal |

n by Ner Snow—3(2): 993

| LA ti la (Vi ) by ROBERT L.

MATHASE ae nt ih Diuciem—3Q): edi

rw" prr Ba

C hean f. + a ae

hac aaah an endem

by Jesús SALCEDO C, E Lenz l, NoRa CASTAÑEDA A. CESAR

CAMPO N. quo DA ANIEL us cd 751

J | f]

Par-

ish, Louisi by BARBARA R. o SUE H. MacRos-

ERTS, CHRISTOPHER S. REID, AND PATRICIA L. FAULKNER—3(1):379

Again: taxonomy of yellow-flowered caulescent Oxalis

s in eastern North America by Guy L. Ne-

SOM—3(2

Amorpna confusa, a new name fer an old Amorpha (Fa-

C.K. STRAUB, BRUCE A. SORRIE,

AND RUAN a Weno - (1): bud

—

L ] FA A nt

en.

by Curtis R. BJÖRK AND MARK Darraci—3(1): 231

An overview of the Diospyros campechiana complex

(Ebenaceae) and description of three new species by

MITCHELL C. PROVANCE AND ANDREW C. SANDERS—3(1):85

Anatomia de la epidermis foliar en las especies Mexicanas

del género de iuda by m ROSALES

CARRILLO AND YOLANDA HERRERA ARRIETA —3(1):28

Annotated a of ce cae flora of T Menard

Creek Corridor Unit of the Big Thicket National Preserve,

Polk, Liberty, and Hardin counties, Texas by Larry E. Brown,

BARBARA R. MACROBERTS, MICHAEL H. MAcRosERTS, AND WARREN W.

Pruess—3(1):443

Annotated list of Maryland grasses s by EbwarD E.

TERRELL AND PAUL M. PETERSON— 3(2):90

Assessment of invasiveness and oed impact in non-

native species of Texas by Guy L. Nesom—3(2):971

Berberis repens (Berberidaceae) new again to the flora of

Pennsylvania by Jerry G. CHMIELEWSKI—3(2):963

Bouteloua curtipendula var. tenuis (Poaceae: Cynodon-

teae), new to Texas and the United States and a key to B.

curtipendula varieties by STEPHAN L. HaTcH, MARIA ANDREA

TOMAS, AND STEVEN R. ARcHER—3(2):9

Bristlecone pine dwarf mistletoe: Arceuthobium micro-

rpum subsp. aristatae (Viscaceae), a new subspecies

x | A at r +f Ariana

by Jaren M. Scott AND RoBERT L. MATHIASEN—3(1):13

Bruce Dale Parfitt, 7 November 1952-53 September 2009

by Ner A. HARRIMAN—3(2):995

Capsicophysalis: a new genus of Solanaceae (Physaleae)

from Mexico and Central America by John E. AVERETT AND

MAHINDA Martinez —3(1):71

Castilleja angustifolia var. dubia (Scrophulariaceae), a

new record for South Dakota by Grace KosTEL AND LYNN

Her.gr—3(1):391

Checklist of the plants of the Guiana Shield 1. An update to

the Angiosperms by Christian FEUILLET—3(2):799

Chromosome numbers of miscellaneous angiosperm taxa

by Marc Baker, JON REBMAN, Bruce Panrirr, DONALD PINKAVA,

CHARLOTTE CHRISTY, ANDREW SALYWON, RAUL PUENTE-MARTINEZ, ALLAN

ZIMMERMAN, AND J. Huco CorA—3(1):279

Clarification of the typification of Michaux names in eastern

North American Vitis (Vitaceae) by Connie Fisk AND ALEXANDER

eile 3(2):739

Clarifications concerning the nomenclature and tax-

Oxypolis ternata (Apiaceae) by Mary ANN E.

Feist —3(2):661

C + t t gi E v We Pa T

JE u L zT E n

ICAGA Vy

Guy L. Nesom—3(1):46

Cyperus pilosus eco new to the flora of Texas by

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

Lewis—3(1):457

Diplycosia indica (Ericaceae): a new species and a new

generic record for India by M.R. Desta AND H.J. CHowoH-

—3(1):147

Mirem unr Ao m ican

Ar 1 I] +

,Poasecunda

Index

Gardens, Kew by Jacques CAYOUETTE, JAMES L. REVEAL, AND GARY

E. Moutton—3(2):763

Diversidad y distribución de las gramíneas (Poaceae) en el

estado de Zacatecas > YOLANDA HERRERA ÁRRIETA Y ARMANDO

Cortés Ortiz—3(2):77

Erigeron bellioides (Asteraceae), new to Florida and the

continental United States by Ranpy L. Mears—3(2):869

Eriogonum cedrorum (Polygonaceae: Eriogonoideae), a

California by J L. REVEAI

AND TM RACHE SNA]: da

a new species endemic to Acker Rock, Oregon i JAMES L

REvEAL, DANA YORK, AND RICHARD HELLIWELL—3(2):63

Estudios en las Apocynaceae mes XXXVII:

monografía del género Rhabdadenia (Apocynoideae:

Echiteae) by J. Francisco MoRALES—3(2):541

Estudios en las ds ynaceae neotropicales XXXVIII: tres

vas especies de Mandevilla (Apocynoideae:

Mes para Colombia y Venezuela by J. Francisco

MoRALES—3(2):565

m P. | pm | p» | H £T) Lain , two

new species from Hawai'i by PILAR CATALAN, ROBERT J. SORENG,

AND Paur M. PeTERSON—3(1):51

Five new species of Myrtaceae from Ecuador by Maria Lucia

een AND one i HoLst—3(1): ra

undance f

+ | * +1 rr” |

M pon PATRICK K. Eo AND JOHN Y. AMPONSAH—3(1): 309

Floristic studies in north central New Mexico, U.S.A. The

usas Mountains and the Jemez Mountains by Brian Reir,

Jit. Larson, BRIAN F. Jacoss, B.E. NELSCN, AND RONALD L. HART-

n A

cl i "1 C E / I LI d 4l

western Louisiana id BARBARA R. Mac m" AND olaa, H.

MacRoserts—3(1

Folia taxonomica a — (Gesneriaceae: Epi-

. X a bl fn. at Cli. P8 iral a LAS O

necies

from Amazonas, and distribution and floral morphology

of P. lata by Chrisman Feuitter-—3(1):1 33

Folia taxonomica 13. Passiflora curva (Passifloraceae), a

supersection Coccinea by CHRISTIAN FeuiLLer—-3(2):577

Folia RENON a 14. More on Fuss MOTA supersection Coc-

ield by CHRISTIAN

ie AND JOHN VANDERPLANK—=3(2): 581

Folia none 15. Five new cda of Paradrymonia

the Venezuelan Guayana by CHRISTIAN Feu —3(2): 583

Folia taxonomica 16. Dilkea (Passifloraceae) 1. Epkia, a new

subgenus and five new species from western Amazonia

and the Guianas by Christian FeuiLLET—3(2):593

Glyceria Pe (Poaceae) new o ne ie of Texas by

ES M. ALLEN, PATRICIA LEWIS, 3(1):393

3( 15222

Howard L. Clark, 1941-2009 by B L. Lirscoms—-3(2):996

Infrageneric taxonomy of Astrophytum (Cactaceae), with

999

remarks on the status of Digitostigma by RICHARD R.

MONTANO C= I yep)

(A lus fi , Ranuncu-

H ^ ] ta . I! E s +l | +

morphometric analysis by ANGELA R. Post, ALEXANDER KRINGS,

WADE A. WALL, AND JOSEPH C. NEAL—3(1):193

Learning about other species: an updated component

of a liberal arts education by Guy L. Nesom AND ALAN S.

cua ie: "

I p J L £i fae) + |

ner R. Post, ALEXANDER KRINGS, edid ÜENNY) XIANG, BON

2n O NN, AND ID JOSEPH - NEAL—3(1):22

j eraceae), it la fl

da A j fol del fruto by PRISCILA R. VAN

DE ve Y M. GABRIELA Lopez—3(2): 793

Medicago rigidula and M. truncatula (Fabaceae): new

to the California flora by RicHARD E. RiEFNER, JR AND ERNEST

SMALL—3(1):323

j xoa den aeuo DEUS IE Tes Nlareh ae

by MicHAEL W. DENSLOW AN D Derick B. PoiNpExrER—3(1): 383

Miscellaneous typifications, one new series, ánd one new

varietal SE in Crataegus (Rosaceae) by J.B.

Puipps—3(1):23

Muhlenbergia nma (Poaceae: Chloridoideae:

Cynodonteae: Muhlenbergiinae), a new species from

Chihuahua, ie by PauL M. PETERSON AND J. Travis Co-

LUMBUS—3(2):52

Mycological erature on Texas fleshy Basidiomycota, two

Ww new fungal al for Texas by

David P. LEWIS AND Eur L. Ovreso—3(1):25

Al "D 4 CR ue H A

deae) by CALEB A. Morse AND JAMES L. Reveal —3(2): 667

New fie noteworthy plants from Florida by Jonn M. KUNZER,

CHARD P. WUNDERLIN, LORAN C. ANDERSON, AND James R. Bur-

KHALTER—3(1):331

New species of Baccharis (Asteraceae: Astereae) from Rio

de Janeiro state, southeastern Brazil by G. Henen, J.F.A.

te ne PEE Esteves —3(1 Jal "d

by MARIA

] al plar +l D I

Mountain region b. em F. Bu DONALD L. HAZLETT, AND

E a M

hlox (Pol Nith

updated circumscription of P. caespitosa, P. douces

P. missoulensis, and P. richardsonii by James H. Lock-

LEAR—3(2):645

Notes on the taxonomic history of two rare species of

Begonia (Begoniaceae) from Sikkim Himalaya ana

their conservation by D.C. NaunivaL, S.K. SHARMA, AND M.K.

Panor—-3(2):823

Notes on the taxonomic history, rediscovery and conserva-

tion status of two endangered species of Ceropegia

(Asclepiadaceae) from Sikkim Himalaya by D.C. NautivaL,

S.K. SHARMA, AND M.K. PANDIT—3(2):815

1000

Novedades y notas misceláneas en las Bromeliaceae de

Mesoamérica by J. Francisco MoraLes—3(1):113

Orobanche riparia (Orobanchaceae), a new species from

the American Midwest by L. Turner CoLtins, Alison E.L.

OLWELL, ANU GEORGE YATSKIEV YI I—3 1 3

Pinus pinceana Winiaee ae); nuevo os para el estado de

Nuevo León, M ELA LARA, CARLOS G. VELAZCO

Macias Y GLAFIRO J, ALANÍS FLORES—3(2):77 1

Polygala cymosa (Polygonaceae) new to Texas by MICHELLE

POLLARD, JASON R. SINGHURST, AND WALTER C. Hot MEs—3(2):969

Preliminary insights into the phylogeny and speciation of

Scalesia (Asteraceae), Galápagos Ed by Jeremy D,

BLASCHKE AND ROGER W. SANDERS—3(1):17

Primer reporte de Cystopteris reevesiana (Aspleniaceae)

para México by José A. VILLARREAL-Q, AND CELESTINO FLORES-

EDT

yt BRETT E. Serviss—3(1): 319

zia (Marcgraviaceae: complejo Norantea) by DiEGO

GIRALDO- pla eas E

da m

DENN O section Pauletia

riaria (Cercideae: C | | Fal J|

RICHARD P. Wurosun—3(2) cam

Revision of B g tion Schnella

oe | Cal Xl n FILAS

» PY GIN

(Cercideae: C Ipini ) by RICHARD |

DERLIN AND RICHARD M. FILERS—3(2):619

Sagina caespitosa (Alsinaceae) in Colorado, U.S.A. by A.E.

dbi WA. egl R.C. ANTIMA =a) 867

California, with notes on the S. auriculata complex E

RICHARD E. RIEFNER, JR. AND ALAN R. SMiTH—3(2):855

Status pc m "i aan et wolfii (Cyperaceae) in

the uL M. McKenzie, C. THEO WirsELL, Lov

R. PHILLIPPE, eae E on MicHAEL A, Homoya, STEVEN B.

POMEN aie aes fa Moses) BT

tatar

cies from Sonora, Mexico by David H. Lorence—3(2): 52 7

Studies of neotropical Compositae-ll. A new combination in

Chromolaena upareni aa by JOHN F. PRUsci—3(2): 573

Synopsis of G

in Trinidad and il by AUSRNDER KaiNGs—3(1): 77

nu cpm

Astereae) by Guv L. Nesom—3(1): 16

The phylogeny of Selenia os inferred from

chloroplast and nuclear sequence data by James B.

oa: 169

to the genera of neotropical Capparaceae with variously

stellate or peltate indumenta by Xavier CORNEJO AND HUGH

H. lLris—3(2):683

The types of Astragalus section Diphysi (Fabaceae), a

complex endemic to Western North America, Part k lec-

totypification

several taxa by J - ANDREW ALEXANDER 3(1):2

The typification of Crotalaria rotundifolia a Crotalaria

maritima (Fabaceae) by DanieL B. Wagp—3(1):219

Three new species of Solanum section Herpystichum

(Solanaceae) from Ecuador by Eric J. TEPE AND LYNN

Bo s (2):51 P

fT [| Y: ad EL. Tr. | pu |

__ Grace Koste: —3(2): 901

it ^

of Texas by ALFRED RICHARDSON Al AND W. Ken KNc—3(1) 465

Twenty-eight new three-letter family acronyms for vascular

plants (with E listings on-line) by Nei

Snow-—3(1):27

Two new i AM in Tryphane (Alsinaceae) by W.A.

WEBER AND R.C., WiTTHANN—3(2):501

VYA M_ YY I

and UYY AT S.S

c I L IN

Dy STEPHEN R. STERN AND LYNN Bous—3(2): 503

na nueva e en el | género

opanax reine ad en Centro América byl FRANCISCO

MORALES AND ALVARO IDARRAGA—3(1):1 17

Una nueva especie y novedades nomenclaturales en

el género Meliosma (Sabiaceae) by J. Francisco Mo-

RALES—3(2):535

Validation of the name Symplocos pentandra (Symplo-

caceae) by Jo&o Luiz M. ARANHA FILHO, PETER W. FRITSCH, FRANK

LMEDA, AND ANGELA B. ManriNs—3(2):659

Vascular flora of a saline prairie in Winn Parish, Louisiana by

BARBARA R. MACROBERTS, MICHAEL H. eo CHRISTOPHER S.

REID, AND PATRICIA L. FAULKNER—3(1):34

cd ren pane bs County, Florida by

LAS

LAN

Vascular flora of Morse Clay prairies: in northwestern Louisiana

by BARBARA R. MacRoserts, MicHAEL H. MACROBERTS, CHRISTOPHER

S. REID, AND PATRICIA L, FAULKNER—3(1):355

Vascular plant species richness of the Big Thicket, East

as by MicuaEL H. MaAcRoserTs AND BARBARA R. MacRos-

ERTS—3(1):401

Vegetation areas of Ti pt and

C. TELFAIR II—3(1):395

What is Ranunculus rid (Ranunculaceae)? by ALAN T.

WHITTEMOR E—3 1):24

What is Z ina (Z

CALONJE AND JAN AN MEERMAN—3(1) :43

L D

J My EAI

A. ry n. Ju L INF | A a L, +l Deuk

* LI vox. f D

Dolomite li t f Ala) } p

LAJI EE Hdd tid Uy ROBER

J. Mincy MOFFETT, Ja—3(2): :469

T KRAL AND

Index

INDEX OF 175 AUTHORS:

VOLUME 3 (2009)

1001

Thank you for choosing Journal of the Botanical Research Institute of Texas

aaa J. Andrew—3(1):211

n, Charles M.—3(1):393, 457

dls Frank—3(2):659

2):965

Yolanda Herrera—3(1):285;

3(2):

Asase, Alex—3(1):309

Averett, John E.—3(1):71

Baker, Marc—3(1):279

Blaschke, Jeremy D.—3(1):177

Bohs, Lynn—3(2):503, 511

Brown, mi dcm 3(1):443

Burkhalter, James R.—3(1):331

Calonje, Michael—3(1):23, 31, 43

Carrillo, Octavio Rosales—3(1):285

Carter, Richard-—3(1):457

Chowdhery, HJ—3 1

Cota, J. Hugo—3(1):279

Darrach, Mark—3(1):231

Daugherty, Carolyn M.—3(2):741

Debouck, Daniel Em

Debta, MR.—3(1):14

Denslow, Michael vam 383

Dorr, L.].—3(2):60

Eiler rs, Richard RA —3(2):6

Ekpe, Patrick K—3(1): md

Esteves, R.L.—3(1):139

dean bin: |.—3(1):349, 355, 379

Fac: 661

Aars Ann

E ja rii 2):48

eile a Ta 133: - 577, 581,

,799

ae Luiz M. Aranha—3(2):659

in Connie—3(2): 739

Floden, Aaron J.—3(2):48

Flores, a Neon 771

Flores-L., Celestino—3(1):307

Franck, Alan R—3(1):339

Fritsch, Peter W—3(2):659

raldo-Cafias, a 691

ee Patrick—3(1):3

Harriman, Neil A.—3(2):995

Hartman, Ronald L—3(2):921

Hatch, Stephan L.—3(2):965

Hazlett, Donald L.—3(1):317

Heiden, G—3(1):139

d, Michael A

bs Bruce K.—3(1):123; 3(2):609

Homov )

Idarraga, Alvaro—3(1):117

Iltis, Hugh H.—3(2):683

Jacobs, Brian F—-3(2):921

Judd, Walter S. —3(1):159

Kawasaki, Maria Lúcia—3(1):123; 3(2):609

King, W. Ken—3(1):46

Kostel, cece 391; 3(2):901

Kral, Robert—3(2):

Krings, FOE ALME 77, 193, 227;

3(2):

Kunzer, John M.—3(1):331

Lara, Susana Favela—3(2):771

Larson, Jill —3(2):921

Lé

Locklear, James pus ):645

López, M. Gabriela—3(2):793

Lorence, David H.—3(2):521

Macías, sa G. Velazco—3(2):771

MacRo —3(1):349, 355, 367,

Rarhara Q

"cu Mice H.—-3(1):349, 355, 367,

379, 401,

Martínez, Mahi hn

Martins, Angela B.— E

Mathiasen, Robert | ~am: 13; 3(2):741

3(2):48

Mark H —

Morales, J. Ni ME 118, 117

3(2):535, 541, 565

Morse, nw uet ae 831

Moulton, Gary E.—3(2):76

Nautiyal, D.C.—3(2):815, 823

Neal, Joseph C.—3(1):193, 227

Nelson, B.E.—3(2):921

Nesom, - 2e 157, 161, 461;

3(2):727

Neyland, es

mpo N., César—3(2):75

e Armando Conan 725

Ovrebo, Clark L—3(1):25

“o.

Pandit, M.K.—3(2):815, 823

doe eames 279

ho —3(2):673

mas

Patterson Thomas F—

(1):46

Peterson, a a 527, 905

):83

i Angela R.—3(1):193, 227

ovance, Mitchell C.—3(1):85

Puente- Martinez, Raul—3(1):279

Raiche, Roger—3(2):479

ebman, Jon—3(1):279

Reid, Christopher S.—3(1):349, 355, 379;

3(2):831

Reif, Brian—3(2):921

ie James L.—3(2) 639, 667, 763

ichardson, Alfred— 46

toan Richard ER 323; 3(2):855

Robinson, Harold—3(1):59

Roche, B.J.—3(2):873

Rolfsmeier, Steven B.—3(2):831

Saghatelyan, Anna—3(1):407

Salcedo C, Jesús—3(2):751

Salywon, Andrew—3(1):279

Sanders, Andrew C.—3(1):85

Sanders, Roger W.—3(1):177

Stergios ae 605

—3(2):503

lb Shannon CK—3(1)151

Telfair Il, denis 395

Tepe, Eric J.—3(2):5

Terrell, Edward Es 3(2):905

Tomas, Maria Andrea—3(2):965

Van de Velde, Priscila R.—3(2):793

Vanderplank, John—3(2):581

Vasey, Michael C.—3(2):673

Villarreal-Q,, José A.—3(1):307

Wahlert, Gregory A.—3(2):673

Wail, Wade A.—3(1):193

Ward, Daniel B.—3(1):219

1002

Weakley, Alan S.—3(1):1, 151

Weber, W.A.—3(2):501, 867

Whittemore, Alan T.—3(1):245

Wilder, G.J.—3(2):873

Witsell, C. Theo—3(2):831

BOTANICAL NAMES AND SUBJECT

Vol

pa family—3(1):273; 3(2):993

Aechm

angustifola—3(h 113

pia clade—

Africa—3(1):309

Aira Worm 763

Alabama, USA—3(2):469

Alsinaceae—3(2):501, 867

Amaranthus californicus—3(1):317

Amazonas, Venezuela—3(1):13

3(2):583

Amazonia—3(2):593

Amorpha

confusa—3(1):151, 154

aided 151

confusa—3(1):151

Amo heme Fabaceae 30 151

Andes—3(2):60

Adonem, legem Group—

3(2):99

fete CMT 279

Anthemideae (Asteraceae)—3(2):667

Anticlea elegans var. glaucus—

Apiaceae—3(2):661

Apocynaceae—3(1):77, 461; 3(2):541,

565

Apocynoideae (Apocynaceae)—3(2):

541,565

Appunialongipedunculata—3(2):799,

809

Appunia peduncularis—3(2):799,

809

Aquatic plants—3(2):855

Mp a d 117; 3(2):605

Arceuthobiu

Mis subsp. aristatae

74]

Arctic-montane buttercup—3(1):245

Arctostaphylos—3(2):673

glandulosa—3(2):673

Ll "Er ET o Fia h

Wittmann, R.C.—3(2):501, 867

Wunderlin, Richard P.—3(1):331, 339;

3(2):619, 629

Xiang, Qui-Yun (Jenny) —3(1):227

ume 3 (2009

New names (95) in bold face

Mme 673

patula—3(2):67

—3(2):6

Argentina—3(2): 793

Arizona, USA—3(1):13

pene USA—3(1):319; 3(2):485

Asclepiadaceae—3(2):815

Asclepiadoideae (Apocynaceae)—

:77, 461

Asia—3(1):147, 245; 3(2):815, 823

Aspleniaceae—3(1):307

Asteraceae—3(1):139, 161, 177, 231;

:573, 667, 869

Astereae (Asteraceae)—3(1):139, 161,

231

Astragalus

sect. Diphysi—3(1):2

is vat. rone Sil 211,

n N, 1,213

r. albiflorus—3(1):211

var. bryantii—3(1):211,215

var.diphysus—3(1):211,214

var. iodanthus—3(1):211,

var. micans—3(1):211, 216

var. oropedii—3(1):211,216

var. pseudiodanthus—

3(1):211, 216

ii ou o. 251

subg. Astrophytum—3(1):251

sect. cu m3 251

subg. Fuastrophytum—3(1):251

subg. Neoastrophytum—3(1):251

sect. Septentriastrophytum—

DP t y di Y

Yatskievych, George—3(1):3

York, Dana—3(2):639

Zarucchi, James L—3(1):157

Zimmerman, Allan—3(1):279

Zomlefer, Wendy B.—3(1):159

INDEX:

subg. CN

Athenaea cernua—3(1):7

Azolla pinnata subsp. ids 331

Baccharis—3(1):139

altimontana—3(1):139, 140

sect. Caulopterae—3(1):139

friburgensis—3(1):139, 143

genistelloides complex—3(1):139

macrophylla—3(1):139

sect. Oblongifoliae—3(1):139

Bandelier ps Monument, USA—

3(2):9

Exo m 257

Bauhinia

subg. Bauhinia sect. Pauletia ser.

Ariaria—3(2):629

sole

):6

lilacina—3(2):619, 621

maximiliani—3(2):621

microstachya—3(2):623

multinervia—3(2):633

Phanera sect. Schnella—

)

pchinchensis— 30 =

poiteauana—3(2 sea

radiata—3(2):62

vin 326

tarapotensis—3(2):6

tr TE s 9,627

Beautempsia—3(2):68

Ica En 685

Begonia

satrapis—3(2):823, 824

scutata—3(2):823, 828

Begoniaceae—3(2):823

Belize—3(1):23, 31, 43

Berberidaceae—3(2):963

Berberis repens—3(2):963

Big Bend region, Texas, USA—3(1):407

Index

Big Thicket National Preserve, USA—

3(1):443

Big Thicket, Texas, USA—3(1):401

Biological invasions—3(1):323

(2):965

var. tenuis—3(2):965, 966

Brassicaceae—3(1):169, 227

Brazil—3(1):139; 3(2):619, 659

Bristlecone pine—3(1):13

Bromeliaceae—3(1):113

Buffalo Gap National Grassland, USA—

Cactaceae—3(1):25

Caesalpinioideae c 619,

629

California, USA—3(1):323; 3(2):479,

639, 673, 741, 855

Calyptranthes

compactiflora—3(1):123

os 0123, 125

arsiflora—3(1):123, 125

eae Pee Idaho, AMA 231

Capparaceae—3(2):68

Capparis

i R

ovalifolia—3(2):68

doe cona n "a

otosina—3(1):7

Cardamine "iso Ds

Carson National Forest, USA—3(2):921

Caryophyllaceae/Alsinaceae—3(2):

501, 867

Castilleja

angustifolia var. oo 391

chromosa—

Gene -at 2 a 43,711,117;

2):535

n (Caesalpinioideae: Fabaceae)

3( 9

Ceropegia

hookeri—3(2): 815,816

tos

Chihuahua, Mexico—3

Chloridoideae De uA 527

Chloroplas ata—3(1): =

esa AtS duci

Chromosome sae nie ):

279

Circumscription—3(2):645

Clark, Howard L.—3(2):996

Coastal dry forests—3(2):683

Colombia— 3(2):565

Colorado, USA—3(1):317; 3(2):867

Compositae: Eupatorieae—3(2):573

Conata Basin, South Dakota, USA—3(2):

901

Corolla morphology—3(1):13

Costa NE 85,113,117; 305 595

Cracc

AS 157

intermedia—3(1):157

Crataegus—3(1):239

acutiloba—3(1):239

ser. Altaicae—3(1):239, 243

brainerdii—3(1):239

chrysocarpa var. oo

—3(1):239, 2

demissa—3(1): a

fernaldii—3(1):239

gravis—3(1):239

laurentiana—3(1):23

var. demi 239

pastorum—3(1):239

pentandra—3(1):239

populnea—3(1):239

roanensis—3(1):239

subrotundifolia—3(1):239

Creeping sideoats grama—3(2):965

Critically oo 521

Crotalaria—

mE 219,

rotundifolia—3(1): A m

Cryptostegia

grandiflora—3(1):461

da oo.

Cycadaies—3(1):23

Cynodonteae jm 527,965

Cyperaceae—3(1):457; 3(2):793, 831

Cyperus

bipartitus—3(1):331

eragrostis—3(1):331

pilosus—

Cystopteris reevesiana—3(1):307

Miro Mec

caput Medusae a(t): :251

Dilkea-—3(2):59

eu ies 595

subg. Epkia—3(2):593, 594

tillettii—3(2):593, 599

vanessae— 3(2):593, 602

i eme ):85

campechiana—3(1):85, 89

1003

campechiana oo 85

camposii—3(1):

crotalaria—3(1) :85, Es

haberi—3(1):85, 9

hartmanniana—3(1):101

indica—3(1):147

juruensis—3(1):104

oe ee

Dirca—3(2):48

neu :485, 494

mexicana—3(2):485, 496

A :485, 496

palustris—3(2):485, 497

Dwarf B a 3

Ebenaceae—3(1):85

Echiteae (Apocynaceae, Apocynoideae)

—3(2):541

Echites—3(2):541

biflorus—3(2):541

ehrenbergii—3(2): B

madidus—3(2):54

bal impact—3(2 -

Ecuador—3(1):123; o 511, 609,

629, uc

Education—

Eleocharis n am 831

Endangered—3(2):8

Endemic—3(2):485, ER 823

Engelmann spruce—3(1):13

Epidermal anatomy—3(1):285

Episcieae (Gesneriaceae)—3(1):133;

3(2):583

Epitype—3(1):211

Ericaceae—3(1):147; 3(2):673

Erigeron bellioides—3(2):869

ue (Polygonaceae)— 3(2):

—

vU 639

villosissimum—3(2):63

ade

aequatoriensis—3(2):609

castaneiflora—3(2):609, 611

crassimarginata—3(2):609, 61 1

i E 609, 614

silliflora—3(2):609, 616

ia (Asteraceae) —3(2):573

upatorium clematitis—3(2):573

Euphorbia extipulata—3(1):317

Eurybia

o 161,166

glauca—3(1):161

sect. Herrickia—3(1):161

sect. Horrida—3(1):161

1004

horrida—3(1):161

kingii—3(1):161, 166

var.barnebyana—3(1):161,

166

pulchra—3(1):161

wasatchensis—3(1):161

Fabaceae—3(1):151, 157,211,219, 323;

3(2):619, 629, 751

Festuca—3(1):51

aloha—3(1):51

molokaiensis—3(1):51, 54

Flora Londonensis—3(1):227

Florida, USA—3(1):331, 339; 3(2):869,

873

Floristic diversity—3(2):921

ores

upland prohi pacer

upland nec im 367

Formosa firethorn—3(1):

French Guiana—3(2):577

Galapagos elias UN 177

Generation time— 73

Geographic range shift—3(2):81 5

Geranium maculatum—3(1):331

Gesneriaceae—3(1):1 33; 3(2):583

Ghana—3(1):309

Glyceria declinata—3(1):393

Gonolobinae (Apocynaceae, Asclepia-

doideae) —3(1):77

Gonolobus—3(1):77

rostratus—3(1):77, 78

tobagensis—3(1):77, 82

GuianaShield, Checklistofthe Plants ofthe

—3(2):/99

Guiana Shield —3(1):133; 3(2):581

Guianas—3(2):593

Guinea savanna—3(1):309

Gulf o oo 873

Guzm

OET 13

mitis—3(1):113

Gylceria

arkansana—3(1):393

septentrionalis—3(1):393

striata—3(1):393

=- oe >

"M pm ):5

Hedyotideae PU RM 521

Hedyotis—3(1):59

galeottii—3(1):59

kingii—3(1):59

latifolia—3(1):59

terrellii—3(1):59

Heliantheae (Asteraceae)—3(1):177

Herrickia horrida— 1

Honduras—3(1):113

Hygrocybe

chamaeleon—-3(1):257, 264

mississippiensis—3(1): 257, 264

Idaho, USA—3(1):231

India—3(1):147; 3(2):815, 823

Interspecific hybridization—3(2):673

Invasive species—3(1): 323; 3(2):901

Invasiveness 2. ed

Isonym—3(2):661

IUCN Red List—3(2):521

Jemez Mountains, USA—3(2):921

Kansas, USA—3(2):485

Kendrick Peak, Arizona, USA—3(1):13

Kew, The Royal Botanic Gardens—3(2):

763

Lamiaceae—3(1):383

Lamium purpureum—3(1):331

Leaf epidermis—3(1):285

pa 43,211,227

(1):193

| J"

SET Ce ld hallo

wis and Clark—3(2):763

us arts—3(1):1

Lipocarpha T 793, 794

Lolinnae (Poaceae)—3(1):5

Louisiana, USA—3(1):349, m 367,

375,379

du a (Asteraceae)—

3(1):16

ec ):56

albi ap. 565

caquetana—3(2):56

sanctae- nc = 569

Nags ee —3(2):691

Marco Island im 873

Maryland, USA—3(

Maya Mountains, ir 31,43

Medic—3(1):323

Medicago

rigidula—3(1):323

trunculata—3(1):323

dede Od 159

Meliosma— 535

hartshornii—3(2): 535

irazuensis—3(2):535, 537

isthmensis—3(2):535, 537

Melochia dS

Memoriam—3(2):995, 996

Mentha

longifolia—3(1):383

xrotundifolia—3(1):383

suaveolens—3(1):383

Mesechiteae (Apocynaceae, Apocynoi-

deae) —3(2):565

Mesoamerica—3(1):85

Mexico—3(1):59, 71, 85, a 307; 3(2):

521, 527,751,771, 775

Mexotis—3(1):59, 60

galeottii—3(1):59, 61

kingii—3(1):59, 64

latifolia—3(1):59, 61

lorencei—3(1):59, 67

terrellii—3(1):59, i

Michaux names—

Micromorphology, PN 793

Molecular phylogeny—3(2):673

Muhlenbergia—3(1):285; 3(2):527

subg. oo 285;

3(2):52

toda sie 7,528

subg. Trichochloa—3(1):285

Mycology—3(1):257

Myrcia

aequatoriensis—3(1):1 23, 128

verticillata—3(1):1 23, 130

Myrtaceae—3(1):123; 3(2):609

Naturalized—3(2): s

Neotropics—3(2):541,

Neoype/necypiicaion 30 219;

3(2):667

New Mexico, uu t 317; 3(2):921

Nicaragua—

Nonnative Kam 461; 3(2):

North America) 3,13,59, a

, 211, 219, 231,

s 257, a 307, 317, ae a

331, 339, 349, 355, 367, 375, 379,

383, 391, 393, 395, 401, 407, 443,

457, 461, 465; 3(2):469, 479, 485,

521, 639, 727, 739, 741, 751, 771,

de n 855, 867, 869, 873, 901,

, 963, 965, 969, 971

North Carola M T 151,383

Nuclear ribosomal ITS—3(2):673

Nuclear sequence docu 69

Nuevo León, Mexico—3(2):771

Ochna serrulata—3(1):33

Oregon zc m 639, 741

Oreopah

aerate al 117

nicaraguensis—3(1):117

nubigenus—3(1):117

Index

del da 17,118

eltatus—3(1):1

mini

Orobanche

d 1):3

No othaphyllon—3(1): 3

le mies 729

poo 727

ies 727,

subsp. pesas 727

florida—3(2):72

permi i A

wee mae 727,736

lyonii—3(2):7

ias a 7,731

ue 127

ee 727

MMC i cs

Oxypolis

denticulata—3(2):661

rigidior—3(2):661, 662

ternata—3(2):661, 662

Palouse grasslands, Washington, USA—

Panama—3(1):113, 117; 3(2):535

Pappobolus—3(1):177

aradrymonia

glandulosa—3(2):583

subg. Pagothyra—3(1):134

subg. i a 583

To ):58

—3(2): m Es

s Brira Mala 2):995

fon at 331; 3(2):577, 581,593

Supersect. o cen 581

compar—3(2):58

uda

longicuspis—3(2):581

subg. Passiflora supersect. Coccinea

—3(2):577

tecta—3(2):581, 582

vitifolia—3(1):331

Passifloraceae—3(2):577, 581, 593

nsylvania, USA—3(2):963

Peru—3(2):683

lll ternatum—3(2):661

Phaseo

marechalii—3(2):75

sect. ERR 75]

Phlox—3(2):645

austromontana

subsp. jonesii—3(2):645

subsp. lutescens—3(2):645,

646

caespitosa—3(2):645

condensata subsp. covillei—3(2):

douglasii—3(2):645

missoulensis—3(2):645

pilosa subsp longipilosa—

3(2):

5,647

icursont 3a

UPR i (2): 645,

647

subulata d setacea—3(2):

villósissima ise latisepala—

649

Phylogeny—3(2):485

Physaleae (Solanaceae) —3(1):71

Physalineae (Physaleae: Solanaceae) —

3(1):71

Physalis—3(1):71

Picea

engelmannii—3(1):13

ngens—3(1):13

Pinaceae—3(2):771

Pinu

aristata—3(1):13

ontorta var. contorta—3(2):741

nan (2):741

monticola—3(2):741

a esci

Poa secunda—3(2):76

peat i 393; 3(2):527,

763,7 6

eee 645

Polygala cymosa—3(2):969

Polygalaceae—3(2):969

Polygonaceae—3(2):479, 639

Polyploidy—3(2):673

Principle components analysis—3(1):

Priority—3(1):43

Pyr ll koidzumii—3(1):319

Pyrroc

——

scaberula—3(1):231

lir ER 93, 245

Ranunculus

—3(1):245

sect. Epirotes—3(1): 245

ficaria—3(1):193

subsp. bulbilifer—3(1):195

1005

subsp. calthifolius—3(1):

195

subsp. chrysocephalus—

subsp. ficaria—3(1):195

subsp. ficariiformis—3(1):

gelidus—3(1):245

grayi—3(1):245

karelinii—3(1):245

Rare—3(2):823

Rhabdadenia—3(2):541, 547

biflora—3(2):541, 548

lindeniana—3(2):541

macrostoma—3(2):541, 552

adida—3(2):541, 552

pohlii—3(2):541

Var. acd 54]

ragonesei—3(2):54

Rio de Janeiro, mean a

Rocky Mountains, USA—3(1):317

Rosaceae—3(1):239, 319

Rubiaceae—3(1):59; 3(2):521, 799

Sabiaceae—3(2):535

Sagina caespitosa—3(2):867

Shis Toms 349

Salvi

prm complex—3(2):855

minima—3(2):8 a

molesta—3(2):85

mee re 855

Salviniaceae—3(2):855

San Francisco Peaks, Arizona, USA—3(1):

San Luis Potosi, Mexico—3(1):71

Sangre de Cristo Range, USA—3(2):

921

Santa Fe National Forest, USA—3(2):

921

Scalesia—3(1):177

Schefflera

pittieri—3(2):605

eoe 605

ander pl 605

ofa e

strana a 7696

andina—3(2):69

ON 700

brasiliensis—3(2):702

magnifica—3(2):712

Norantea complex—3(2):691

parrae—3(2):712

pterosara—3(2):715

1006

renvoizei—3(2):717

spiciflora—3(2):719

weddelliana—3(2):719

Scrophulariaceae—3(1):391

Selenia—3(1):169

grandis—3(1):169

pns ):169

Sida urens—3(1):33

Sikkim ina a :815,823

Simsia—3(1):1

Singsaglebini = Reserve, Ghana—

1):309

Sium denticulatum—3(2):661

Snake se a Idaho, USA—

3(1):2

ccm B 2 503, 511

Solanum—3(2):5

circinatum oe ramosum—

3(2):509

corymbiflorum subsp. mortonia-

num—3(2):5

crassinervium—3(2):511, 514

ee Med 511

deflex

arate subsp chloranthum

3(2):50

MOS cM 511

endopogon subsp. guianense—

3(2):50

evolvulifolium—3(2):511

io i E 503

huilense—3(2):50

ire ol 511,516

manabiense—3(2):503, 504

obliquum—3(2):503

pacificum—3(2):511, 512

sect. boa E 503

turneroides—3(2):50

zumben go xs 506

Sonora, Mexico—3(2):52

South America—3(1):77, E 123, 133,

139; 3(2):503, 511, 565, 573, 577,

581, 583, 593, 605, 609, 629, 659,

83, 691, 793, 799

South Carolina, USA—3(1):151

South Dakota, USA—-3(1):391; 3(2):

Species richness—3(1):401

—

Spermacoceae E EE Rubiaceae

—3(2):52

Sphaeropteris eT 33]

Stenaria—3(2):521

sanchezii—3(2):521

Superimposed nucleotide S

patterns—3(2):673

Symplocaceae—3(2):659

Symplocos

itatiaiae var. pentandra—3(2):659

pentandra—3(2):659, 6

Synthesis of the North America Flora—

Tephrosia

o ):157

florida—3(1):15

orien cat 157

media—3(1):15

Texas, aes 257,393, 395,401,407,

443, 457,461,465; 3(2):965, 969,97 1

Three-letter family acronyms—3(1):273;

3(2):993

Thymelaea passerina —3(2):901

Thymelaeaceae—3(2):485, 901

so romeliacead—3U):

Torilis ON

Tournefortia hirsutissima—3(1):465

Trans-Pecos Vegetational Area, Texas,

A—3

Trichostigma octandrum—3(2):873

Trifolium rid d anemophilum

1:3

Trin idad íi iotago—30 77

Triniteurybia—3(1):16

Tryphane—3(2):501

rubella—3(2):501

subsp. propinqua—3(2):

501, 502

Tusas Mountains, USA—3(2):921

Lore 211, 219, 227,

239; 3(2):667, 739

USA—3(1):3, 13,51, 151, 193, 219, 257,

319, 323, 331, 339, 349, 355, 367,

375, 379, 383, 391, 393, 395, 401,

407, 443, 457, 461, 465; 3(2):469,

479, 485, 639, 741, 831, 867, 869

873, 901, 905, 921, 963, 965,

969, 971

ET as SEN

Valles Caldera National Preserve, USA—

3(2):921

Vascular flora—3(2):873

Vegetational areas—3(1):39

c 133; 3(2): m 583,

Tem 1217

Viscaceae—3(1): A 3(2):741

Vitaceae—3(2):739

Vitis

aestivalis—3(2):739

cordifolia—3(2):739

riparia—3(2):739

rotundifolia—3(2):739

Washington, USA—3(1):231

Werauhia

brunei—3(1):113

camptoclada—3(1):113

luctuosa—3(1):113, 114

pittieri—3(1):113

West E Coastal Plain, USA—3(1):349,

Wildland-urban nei 855

Wolf's spike-rush—3(2):83 1

Woody plants—3(1):309

Xanthoxalis—3(2):727

Xyridaceae—3(2):469

ris

spathifolia—3(2):469, 472

tennesseensis—3(2):469

— Mexico—-3(2):775

"bussell/onan- -reyesii—3(1):31

tuerckheimii—3(1):31

Zamiaceae—3(1):23, 31, 4

Zigadenus elegans var. glaucus—3(1):

159

Index

1007

95 New NAMES AND NEw COMBINATIONS:

VoLUME 3 (2009)

Amorpha confusa (Wilbur) S. - K. Straub, Sorrie & Weakley,

comb. et stat. nov.—3(1):15

Anticlea elegans var. ee (Nutt) Zomlefer & Judd,

comb. nov.—3(1):159

Appunia eS (Steyerm.) Delprete, comb.

nov.—3(2):80

Appunia unción (Kunth) Delprete, comb. nov.—

3(2):809

A de | BAL e I

| aristatae JM Scott &

Mathiasen, subsp. nov.—3(1):14

Astragalus lentiginosus var. bryantii (Barneby) J.A

Alexander, comb. nov.—3(1):215

Astragalus lentiginosus var. iodanthus (S. Watson) J.A.

Alexander, comb. nov.—3(1):215

iun lentiginosus var. Bo e (Barneby)

A. Alexander, comb. nov.—3(1):2

"eda altimontana Heiden, RUE & Esteves, sp.

nov.—3(1):140

Baccharis friburgensis Heiden, Baumgratz & Esteves, sp.

nov.—3(1):143

pauhinla esmeraldasensis do a nov.—3(2): d

—3(2):62

Bauhinia apes ine (L.P. oa nde & o

comb. nov.—3(2):6

Calyptranthes emo M.L. Kawasaki & B.K. Holst,

sp. nov.—3(1):123

unite fusca M.L. Kawasaki € B.K. Holst, sp.

nov.—3(1):

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

nov.—3(1):125

Capsicophysalis (Bitter) Averett & M. Martínez, stat.

nov.—3(1):72

kp abla ¿eden (B.L. s & Greenm.) Averett &

nez, comb. no 1

drena mais (DC) n comb. nov.—3(2):573

Crataegus series Altaicae J.B. Phipps, ser. nov.—3(1):243

Crataegus chrysocarpa var. subrotundifolia (Sarg.) J.B.

Phipps, comb. et stat. nov.—3(1):240

Dilkea cuneata Feuillet, sp. nov.—3(2):595

Dilkea subg. Epkia Feuillet, subg. nov.—3(2):594

Dilkea lecta Feuillet, sp. nov.—3(2):597

Dilkea ovalis Feuillet, sp. nov.—3(2):599

Dilkea tillettii Feuillet, sp. nov.—3(2):599

Dilkea vanessae Feuillet, sp. nov.—3(2):602

Diospyros camposii M.C. Provance & A.C. Sanders, sp.

nov.—3(1):93

Diospyros crotalaria M.C. Provance & AC. Sanders, sp.

nov.—3(1):95

Diospyros haberi M.C. Provance & A.C. Sanders, sp.

nov.—3(1):98

Diplycosia dein M.R. Debta €: H.J. Chowdhery, sp.

nov.—3(1):1

PIS decipiens A PIOS sp. nO e :494

che, sp. nov.—3(2):479

Eriogonum villosissimum Reveal & D.A. York, sp. nov.—

3(2):639

Eugenia aequatoriensis M.L. Kawasaki & B.K. Holst, sp.

nov.—3(2):6

Eugenia castaneiflora M.L. Kawasaki & B.K. Holst, sp.

nov.—3(2):611

Eugenia crassimarginata M.L. Kawasaki & B.K. Holst, sp.

nov.—3(2):61 1

Eugenia drid M.L. Kawasaki 8: B.K. Holst, sp.

nov.—3(2):6

Eugenia Susa M.L. Kawasaki & B.K. Holst, sp.

n 2):616

bi [ A Nels.) Gl Nesom, comh no 3(1 y:16

E ia | ii (D CFOM] Nesom, comb. n (1

Eurybia kingii var. barnebyana (Welsh & a G.L.

Nesom, comb. nov.—3(1):166

Festuca aloha Catalan, Soreng & P.M. Peterson, sp.

nov.—3(1):51

Festuca molokaiensis Soreng, PM. Peterson & Catalán, sp.

nov.—3(1):54

Hygrocybe chamaeleon (Cibula) Lewis & Ovrebo, comb.

1):264

nov.—3(

i | is (Cibula) Lewis & Ovrebo, comb.

“nov.—3(1):26

da albifoiia JE ae sp. nov.—3(2):565

andevilla

rales, sp. nov.—3(2):56

alpni sancta- doni. A Morales, sp. Mix 569

Meliosma isthmensis J.F. Morales, sp. nov.—3(2):537

Mexotis Terrell & H. Rob., gen. nov.—3(1):60

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

nov.—3(1):61

Mexotis kingii (Terrell) Terrell & H. Rob. comb. nov.—3(1):64

Mexotis latifolia (M. Martens & Galeotti Terrell & H. Rob.,

comb. nov.— 6]

Mexouls: lorencei Terrell &H Rob, sp ne ee

mb eae

Muhlenbergia tarahumara PM. Peterson & Columbus, sp.

nov.—3(2):528

Myrcia aequatoriensis M.L. Kawasaki & B.K. Holst, sp.

nov.— Ases

L Kawasakié:B.K.Holst, sp. nov.—3(1):130

Oreopanax paramicolus J.F. Morales & A. Idárraga, sp.

nov.—3(1):118

Orobanche riparia LT. Collins, sp. nov.—3(1):7

Paradrymonia glandulosa Feuillet, sp. nov.—3(2):583

Paradrymonia hamata Feuillet, sp. nov.—3(2):585

Paradrymonia lutea Feuillet, sp. nov.—3(2):585

Faracrymonia maguinei Penne’ sp. ovs: M04

rn nn I^

et stat. nov—3(1):1 34

Paradrymonia tepui Feuillet, sp. nov.—3(2):588

Paradrymonia yatua Feuillet, sp. nov.—3(2):588

Passiflora curva Feuillet, sp. nov.—3(2):577

Phlox austromontana subsp. jonesii (Wherry) Locklear,

comb. et stat. nov.—3(2):645

1008

Phi t t bsp. lutescens (S.I Welsh) Lock-

lear, comb. et stat. nov.—3(2):646

Phlox condensata subsp. Pa (E.E. Nelson) Locklear,

comb. et stat. nov.—3(2):64

Phlox pilosa subsp. Longi (Waterf.) Locklear, comb.

El stat ova ):64

[| £ aL m.a » in LI dis ding nodi £T, ke F £3

Nal

Solanum huilense Bohs, nom. nov.—3(2):509

Solanum limoncochaense Tepe, sp. nov.—3(2):516

Solanum manabiense S. Stern, sp. asd 504

Stenaria anene upon sp. nov.—3(2):522

eR Anas A j H A | ‚comb

l L Fia AAAF Al en da

ME i

nov.—3(2):647

Phlox subulata subsp. setacea (L.) Locklear, comb. et stat.

nov.—3(2):64

Phlox villosissima subsp. latisepala (Wherry) Locklear,

comb. nov.—3(2):649

Stergios, sp nov.—3(2):605

‘Dak B 5 mb

nov.—3(2):509

Solanum crassinervium Tepe. sp. nov.—3(2):514

comb. nov.—3(2):509

oa | I L mm! * DL pM

LI --

nov.—3(2):509

et stat. nov—3(2 ):660

Tephrosia s (Small) G.L. Nesom & Zarucchi,

mb. nov.—3(1):15

Mini rubella ater) A. €: D. Love ex W.A. Weber,

mb 3(2):50

Dv hane rubella d propinqua M A. & D.

ber, comb. nov.—3(2):5

Werauhia luctuosa J.F. Morales, sp. Map 14

Xyris spathifolia Kral & Moffett, sp. nov.—3(2):472

Zamia decumbens Calonje, Meerman, M.P. Griff. & Hoese,

sp. nov.— 31

Zamia meermanii Calonje, sp. nov.—3(1):23

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

SBM 09 | The Genus Mikania (Compositae: Eupatorieae) in the Greater Antilles

SBM 10 | Checklist of the Vascular Plants of Tennessee

SBM 11 | Text Annotations & Identification Notes for Manual of the Vascular Flora of the Carolinas

SBM 12 | The “El Cielo” Biosphere Reserve, Tamaulipas, México

SBM 13 | Flora de Manantlan

SBM 14 | Niebla & Vermilacinia (Ramalinaceae) from California & Baja California

SBM 15 | Monograph of Northern Mexican Crataegus (Rosaceae, subfam. Maloideae)

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SBM 19 | Emanuel D. Rudolph’s Studies in the History of North American Botany

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

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SBM 28 | Wild Flowers of Mombacho (Nicaragua)

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Keys to the Vascular Plants of the Black Gap Wildlife Management Area, Brewster County

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Eriocaulaceae of Continental oe 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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JOURNAL OF THE BOTANICAL RESEARCH INSTITUTE OF TEXAS

The Journal of the Botanical Research Institute of Texas, formerly -

called Sida, Contributions to Botany, publishes research in classical |

and modern systematic botany—including studies of anatomy, _

biogeography, chemotaxonomy, ecology, evolution, floristics,

genetics, paleobotany, palynology, and phylogenetic systematics.

Geographic coverage is global. Articles are published in either

English or Spanish; an abstract is provided in two languages. All

contributions are peer reviewed and frequently illustrated with

maps, line drawings, and full color photographs.

b R I T The journal is published twice yearly in a multiple topic format

and provides information to scientists, professionals, and the general public for research,

education, and cultural enhancement. All articles are indexed and abstracted in print

and/or electronic form by the following: AGRICOLA Database (National Agricultural

Library); Applied Botany Abstracts; Biosciences Information Service of Biological

Sciences (BIOSIS); Current Awareness in Biological Sciences (CABS); The Kew Record

of Taxonomic Literature; Natural Products Alert (NAPRALERT); and Referativnyi

Zhurnal (Abstracts Journal of the Institute of Paena c e of the Republic of

Russia). ho

Lloyd H. Shinners, whose herbarium and library were the basis of the Southern

Methodist University (SMU) legacy that became BRIT in 1987, founded the journal in

1962. William E Mahler succeeded Shinners as SMU herbarium director in 1971 and

inherited both ownership and editorship of the journal. Since 1993, it has been published

by the Botanical Research Institute of Texas.

The current issue of the Journal of the Botanical Research Institute of Texas is available online

at http://www. britpress.org.

ISSN 1934-5259