Table of Contents

SYSTEMATICS

A new Phemeranthus (Portulacaceae) from the Piedmont of Virginia and North Carolina

Stewart Ware

A new hedge-nettle (Stachys: Lamiaceae) from the Mid-Atlantic Piedmont and Coastal Plain

of the United States

Gary P. Fleming, John B. Nelson, and John F. Townsend

Polemonium villosissimum (Polemoniaceae), an overlooked species in Alaska and Yukon

Territory

David F. Murray and Reidar Elven

California

Roger Raiche and James L. Reveal

Cymopterus spellenbergii (Apiaceae), a new species from north central New Mexico, U.S.A.

Ronald L. FIartman and M>i Larson

A new hybrid of Hibiscus (Malvaceae) from Texas

Carex austrodeflexa (Cyperaceae), a new species of Carex sect. Acrocystis from the Atlantic

Coastal Plain of the southeastern United States

Bruce A. Sorrie, Patrick D. McMillan, Brian van Eerden, Richard J. LeBlond, Philip E. Hyatt,

Phaseolus hygrophilus (Leguminosae-Papilionoideae), a new wild bean species from the

wet forests of Costa Rica, with notes about section Brevilegumeni

J. Sai.cldo-Castano, R. Araya-Villalobos, N. Castaneda-Alvarez, O. Toro-Chica, and D.G. Debouck

Lepechinia yecorana (Lamiaceae), a new dioecious species from the Yecora area of Sonora,

Mexico

James Henrickson, Mark Fishbein, and Thomas R. Van Devender

Columnea bivalvis (Gesneriaceae), a new species from the eastern slopes of the Ecuadorian

Marisol Amaya-MArquez and John L. Clark

John L. Clark

1 species from northwestern Ecuador

f species from northwestern Ecuador

John L. Clark and James F. !

Begonia bernicei (Begoniaceae), a new species for the lowland humid forests of the Venezuelan

Guayana

Gerardo A. Aymard C. and Gustavo A. Romero-GonzAlez

Matelea pakaraimensis (Apocynaceae: Asclepiadoideae), a new species in the Matelea

stenopetala complex from Guyana

Alexander Krings

Lectotypification of Thomas Nuttall’s names applied to North American Polygala (Polygalaceae)

Alina Freire-Fierro and Alicia Landale

101

105

109

Notes on the disintegration of Polygala (Polygalaceae), with four new genera for the flora of

North America

J. Richard Abbott

New combinations in Phoradendron leucarpum (Viscaceae)

J. Richard Abbott and Ralph L. Thompson

Typihcation and synonymy of the species of Euphorbia subgenus Esula (Euphorbiaceae)

native to the United States and Canada

Dmitry V. Geltman, Paul E. Berry, Ricarda Riina, and Jess Peirson

Western Ghats, India

N. Anil Kumar, M.K. Ratheesh Narayanan, P. Sujanapal, R. Meera Raj, K.A. Sujana, and Mithunlal

A phylogenetic assessment of breeding systems and floral morphology of North American

Ipomoea (Convolvulaceae)

Putative morphological synapomorphies of Saxifragales and their major subclades

Spirodela oligorrhiza (Lemnaceae) is the correct name for the lesser greater duckweed

Daniel B. Ward

Thomas Walter’s Orchids

Daniel B. Ward and John Beckner

Billie L. Turner

DEVELOPMENT AND STRUCTURE

Endosperm and cotyledon areole correlation in Leguminosae subfamily Papilionoideae

Micromorfologia de la lemma de los generos Polypogon, xAgropogon y Agrostis (Poaceae)

en Chile

VICTOR L. Finot, Wilson Ulloa, Carlos M. Baeza, Alicia Marticorena y Eduardo Ruiz

ETHNOBOTANY

Caracterizacion y uso de “pimientas” en una comunidad quilombola de la Amazonia Oriental

(Brasil)

Luciano Araujo Pereira, Gloria Estela Barboza, Massimo Giuseppe Bovini,

Mara Zelia De Almeida y Elsie Franklin GuimarAes

CHROMOSOME NUMBERS

Chromosome number for Arcytophyllum fasciculatum (Rubiaceae)

A. Michael Powell and Allison Leavitt

FLORISTICS, ECOLOGY, AND CONSERVATION

Flora endemica de Nuevo Leon, Mexico y estados colindantes

Carlos G. Velazco Macias, Glafiro J. Alanis Flores, Marco A. Alvarado Vazquez,

Liliana Ramirez Freire y Rahim Foroughbakhch Pournavab

Lucio Lozada Perez y Claudia Gallardo HernAndez

125

143

153

159

179

197

205

213

219

237

255

273

275

299

R.W. Pemberton and H. Liu

254, 320, 330, 336, 340, 356, 3

152, 178, 204, 212, 218,

A NEW PHEMERANTHUS (PORTULACACEAE) FROM THE PIEDMONT

OF VIRGINIA AND NORTH CAROLINA

Stewart Ware

Department of Biology

College of William and Mary

Williamsburg, Virginia 23187-8795, US. A.

saware@wm.edu

RESUMEN

INTRODUCTION

Only two species of Phemeranthus Raf. have been previously recognized from east of the Appalachians. They

and all other eastern United States species were first described in the genus Talinum Adans. and have been

treated under that name in all but the most recent literature (Kiger 2003). Phemeranthus teretifolius (Pursh)

Raf., with a style about the same length as the 15-20 stamens, occurs in rock outcrop communities from

the serpentine barrens of Pennsylvania and Maryland southward through the granitic and gneissic outcrops

of the Piedmont from Virginia to Georgia. It also occurs on sandstone outcrops of the Ridge and Valley in

Virginia and Tennessee (Ware & Pinion 1990; Murdy & Carter 2001; Kiger 2003) and on mafic and ultramafic

outcrops in the western Piedmont of Virginia and North Carolina (C. Ludwig & A. Weakley, pers. comm.).

Phemeranthus mengesii (W. Wolf) Kiger, with larger flowers than P. teretifolius and a style noticeably longer

than the very many (50+) stamens, was first described from sandstone in the Ridge and Valley of Alabama

(Wolf 1920, 1939), but also occurs on sandstone in the Ridge and Valley of Tennessee and in the Altamaha

Grit region of Georgia (Montgomery & Blake 1969; Murdy & Carter 2001; Kiger 2003) and occasionally on

granite (Carter & Murdy 1986).

Producing good herbarium specimens of species in this genus is made especially difficult by a com-

bination of very delicate, ephemeral flowers coupled with thick succulent leaves and large, extremely firm

rhizomes (Holzinger 1900; Ware 1967). Species can usually be distinguished by a variety of floral characters

visible in fresh specimens with open flowers (such as petal color, petal shape, style color, stigma shape,

filament color, and pollen color), but most of these characters are obscured in most herbarium specimens.

As a consequence, identifications of herbarium specimens from east of the Appalachians have been based

largely on stamen numbers (few versus many) and on whether the style is longer than or the same length

as the stamens. Herein is described a new species of Phemeranthus that is like P. mengesii in having a style

longer than the numerous stamens, but with a much smaller flower size and various other floral and habit

characters that distinguish it both from that species and from P. teretifolius, as well as from other Phemeranthus

species in the eastern United States.

, sp. nov. (Fig. 1).-

t 2/3 as long ass

f the University of>

(UNC, WILLI) were examined to try to find a

Table 2. Results of crosses between Phemeranthus species.

Pistillate parent

P. piedmontanus

P. mengesii

P. piedmontanus

P. calcaricus

P. piedmontanus

P. calycinus from AR SS

P. piedmontanus

P. calycinus from MO LS

P. piedmontanus

P. calycinus from AR shale

Staminate parent

Number of Number of normal-

producing appearing seeds/

capsules capsules

P. calcaricus

P. piedmontanus

P. calycinus from AR SS

P. piedmontanus

P. calycinus from MO LS

P. calycinus from AR shale

the same level as or slightly below the anthers, allowing pollination as the flower closes, so a vector is not

essential for fruit set, while a vector is needed in P. piedmontanus.

The subcaptitate stigma and usually the upper 2 /5 of the style are white in both P. piedmontanus and P.

mengesii, but both style and stigma are much smaller in the former. In contrast, P. teretifolius, P. calcaricus,

and all populations I have seen of P. calycinus have strongly tripartite stigmas, and usually (but not always)

have pink pigment in the upper style and sometimes even where the three branches of the stigma join.

This genus is renowned for strong synchrony and predictability of daily opening and closing of flowers,

and flowering time has been invoked as a way to separate species (Holzinger 1900). However, daily flower-

ing time may vary among populations of the same species, may coincide in two different species, and at any

location varies with daily cloudiness and portion of the growth season (Ware 1967; Ware 1993), so it must

be used with caution. Plants from Bald Knob at Rocky Mount, Virginia usually open around 1:30-1:45 pm

EDT and close about 5:30. This contrasts with most Virginia populations of P. teretifolius, which flower later

(often around 3:30 EDT) and stay open later. Earlier opening of flowers in P. piedmontanus also occurs where

it and P. teretifolius occur on the same outcrop in North Carolina (A. Weakley, pers. comm.).

Crossing Attempts. — All self-pollinations produced normal fruit filled with normal-appearing seeds.

None of the cross-pollinations could be called successful. Only three pollinations caused capsules to form,

all when P. mengesii was the pistillate parent and P. piedmontanus was the staminate parent (Table 2). In

one of these capsules, no normal seeds were produced, and only one and three normal-looking seeds were

produced in the other two capsules, far below the 25 to 50 seeds produced in manually self-pollinated P.

mengesii plants. No attempt was made to germinate the four normal-appearing seeds, but such a low fruit

set and such a small number of normal-appearing seeds clearly indicate strong barriers to cross-pollination.

Further, accidental self-pollination of P. mengesii during the crossing efforts cannot be ruled out completely.

Herbarium Specimens . — No specimens from additional locations in North Carolina or Virginia could

be surely identified as P. piedmontanus. The floral characters used to distinguish this species in life are

poorly preserved in most collections. Almost no specimens had been pressed during the afternoon hours

when flowers are fully open. Herbarium labels rarely gave information about relative style/stamen length or

whether stamens were few or very many. In a few specimens with a closed flower that had been open on the

counts of stamens, but such dissection destroys the flower, eliminating evidence for future confirmation of

identity. Phemeranthus specimens should be pressed in the afternoon while flowers are fully (not partially)

open, and labels should include notes on the relative length of style and stamens, whether the stamens are

many or relatively few, and preferably an actual count of stamens in at least one flower.

BOOK REVIEWS

st. Texas 5(1): 8. 2011

A NEW HEDGE-NETTLE ( STACHYS : LAMIACEAE) FROM THE

MID-ATLANTIC PIEDMONT AND COASTAL PLAIN OF THE UNITED STATES

Gary P. Fleming

Virginia Dept, of Conservation and

Recreation, Division of Nat. Heritage

r J 1 1 7 Governor St., 3rd Fioor

Richmond, Virginia 232 19, U.S.A.

gary. fleming@dcr. virginia.gov

John B. Nelson

AC. Moore Herbarium

Dept. Biological Sciences

University of South Carolina

Columbia, South Carolina 29208 U.S.A.

nelson@sc.edu

ABSTRACT

John F. Townsend

Virginia Dept, of Conservation and

Recreation, Division of Nat. Heritage

217 Governor St., 3rd Floor

Richmond, Virginia 23219, U.S.A.

john.townsend@dcr.virginia.gov

RESUMEN

INTRODUCTION

Field work and herbarium studies in North Carolina and Virginia has led to the circumscription of a new

species in the genus Stachys, a large group of mints widely recognized for its taxonomic complexity. Changing

taxonomic interpretations in the past have confounded efforts to classify this plant properly, but enough has

now been learned to recognize this species’ unique place within the genus. A history of taxonomic opinions

regarding Stachys matthewsii is presented, as is a discussion of habitat and geography, particularly its primary

range in the Piedmont and its somewhat disjunct Coastal Plain populations in Virginia.

Stachys matthewsii G.P Fleming, J.B. Nelson, &J.F Townsend, sp. i

t, 11 Jun 1979 John B. Nelson 1152 withj. Matthews (i

:: UNITED STATES: E

), NCU, UNCC[2]).

Perennial herbs from vigorous, pale, fragrant rhizomes; flowering stems erect, 0.8-1. 05m tall, sparingly

branched, even within inflorescence, or branched toward base on older plants; the sides glabrous as high as

the lowest verticil, fertile internodes with scattered short glandular hairs, the angles prominently and con-

sistently pubescent with stifhsh, spreading (or retrorse) hairs, these eglandular, to 3mm long; nodes bearded

with hairs as those on the stem angles; the overall pubescence is dense at the base of the plant, becoming

progressively less so upwards; leaves consistently petioled, the petioles 2-5mm long, blades mostly rounded

at base or somewhat cordate, the blades 5-10 cm long, 2-5cm broad, broadly elliptical to lance-ovate, acute

J. Bot. Res. Inst. Texas 5(1): 9-1

available key or description that would account for the particular combination of characters that are diagnostic

of the taxon. In recent decades, interpretation of the genus has become more stable, with circumscriptions

of native southeastern species generally based on combinations of three character-groups: relative petiole

length, shape and relative length of the calyx lobes, and distribution and type of stem pubescence (Nelson

1981; Mulligan & Munro 1989). Stachys matthewsii is characterized by the combination of short petioles,

short triangular-apiculate calyx lobes, and copious pubescence restricted to the stem angles (Figs. 2, 3, and

5),: This taxon was not clearly recognized in any treatment until that by Nelson (1981), who maintained

it within S. nuttallii (= S. cordata) as a short-petioled variant, disjunct from the rest of the range. Current

treatments of the nutallii-cordata group, however, distinguish its members not only by their long-petioled

leaves but also by the presence of hairs on the stem sides and their musky-scented, atomiferous-glandular

foliage (Nelson 2008), all characters absent in S. matthewsii. Contemporary interpretation of S. clingmanii

as a longer-petioled, high-elevation plant of the southern Appalachians has rendered Fernald’s view of the

Surry County hedge-nettles both taxonomically erroneous and phytogeographically improbable (Nelson

1981; Gleason & Cronquist 1991).

The treatment by Mulligan and Munro (1989) would place the collections of this taxon within S. latidens

Small. Annotations of the holotype and one paratype of Stachys matthewsii ( Nelson 6733), supplied by G.A.

Mulligan in 1990, read “Stachys latidens Small, very close to”; the holotype additionally bears Mulligan’s

note “angles long erect hairs.” Like S. matthewsii, S. latidens has short petioles and short deltoid calyx lobes.

However, it is a relatively smooth plant with only scattered, pustulate, short-deflexed hairs on the stem

angles. Additionally, the bearded nodes and abundant, prominently spreading, multicellular hairs of S.

matthewsu will consistently separate it from S. latidens. Moreover, the two species have allopatric distribu-

tions and disparate ecological affiliations, with S. latidens a plant of higher-elevation forests and clearings i^Jr;

the Blue Ridge and S. matthewsii a plant of low-elevation clearings and edges in the Piedmont and Coastal

Plain. This geographic and ecological separation of S. matthewsii and S. latidens would seem to preclude any

biological interaction between the two taxa at present. Other species, such as S. hispida and S. tenuifolia,

are occasionally found sympatrically with S. matthewsii, but no intermediates have been seen to suggest

any current gene exchange among members of the group. The type of multicellular pubescence seen in S.

matthewsu could possibly indicate some past kinship with S. hispida, although the distribution and density

of these pustular-based, multicellular hairs is distinctly different in the two taxa. Specifically, S. hispida is

more or less uniformly hispid with a moderate to somewhat dense concentration of hairs along the stem,

whereas those of S. matthewsii are only moderately dense above but rapidly become very densely concentrated

in the proximal 3-4 nodes and internodes. Without a molecular framework for placing the various species

of Stachys, such morphological and ecological evidence will remain the most practical tools for recognizing

new taxa in this complex genus.

The population at the type locality is near the margin of the Yadkin River, at the narrows just below

(and within sight of) the Falls Dam, east of the town of Badin. The plants grow in considerable shade on

sandy, loamy ground, with the population best developed on a somewhat sandy ridge. The site is prob-

ably best considered an occasionally flooded-scoured river levee complex. Associated species include

Liriodendron tulipijera L., Pirns echinata Mill., Liquidambar styraciflua L., Quercus nigra L., Robinia pseudoacacia

L., Albiziajulibrissin Durazz., Sassafras albidum (Nutt.) Nees, Alnus serrulata (Aiton) Willd., Amorphajruticosa

L., Parthenocissus quinquefolia (L.) Planch., Apios americana Medik., Vitis rotundifolia Michx., and Asplenium

platyneuron (L.) Britton, Sterns, & Poggenb., all of which are locally common and hardly indicating special

or unusual habitats.

Subsequent study of populations and collections from Virginia demonstrate a wider distribution (Fig.

4). Most Virginia populations occur either along streams or in low, seasonally damp swales. Habitats could

generally be considered mundane, except for those of the Surry County populations, which occupy calcareous

open swales and forest edges along shell-laden shores of the James River. Repeated observations of popula-

tions in Halifax and Charlotte Counties indicate that S. matthewsii is quite shade-intolerant and not very

Journal of the Botanical Research Institute of Texas 5(1)

Fig. 1 . Stachys matthewsii habit.

Fig. 2. Middle stem, leaves, and petioles of Stachys matthewsii.

ig. 3. Flowering and fruiting calyces of Stachys matthewsii.

competitive with taller, ruderal perennials. Plants growing in a formerly cleared floodplain now invaded by

young trees have become progressively shade-stressed and do not flower. A periodically mowed powerline

swale with short to medium-height herbaceous vegetation appears to represent optimal habitat for the species,

supporting a population with hundreds of flowering plants. Perhaps the combination of shade-intolerance

and poor competitive abilities, as well as low levels of botanical inventory in the Piedmont, accounts for the

relatively low number of known populations, despite an abundance of seemingly suitable habitat in the region.

Notes on the Distribution of Stachys matthewsii

There is precedence for the somewhat disjunct distribution of populations of Stachys matthewsii between the

North Carolina - Virginia Piedmont and calcareous habitats of the Virginia Coastal Plain. Previous investiga-

tions by Ware and Ware (1992) and MacDonald (2000) point to numerous species of primarily Piedmont

and Mountain distribution with outlying populations in or adjacent to coastal ravine complexes underlain

by the calcareous Yorktown and Eastover formations. These Pliocene deposits, due to their highly erodible

nature, have allowed for the creation of relatively dramatic ravine complexes within the otherwise slightly

undulating and acidic landscape of the Tidewater region. It is widely assumed that the disjuncts present ifi

or near such ravines are relics of early Holocene climates, when species now confined to montane habitats

were more broadly distributed in Virginia (Delcourt & Delcourt 1986). While Stachys matthewsii is not con-

fined to these steep - sided erosional features, the presence of populations immediately adjacent to them

on shelly substrates (and nowhere else in the region) certainly points to relictual distribution as a possible

factor involved in their presence.

Journal of the Botanical Research Institute of Texas 5(1)

POLEMONIUM VILLOSISSIMUM (POLEMONIACEAE),

AN OVERLOOKED SPECIES IN ALASKA AND YUKON TERRITORY

David F. Murray

Reidar Elven

University of Alaska Museum of the North

907 Yukon Dr

Fairbanks, Alaska, 99709, U.S.A.

dfmurray@alaska.edu

Author for correspondence

Natural History Museum

University of Oslo

Oslo, NORWAY

ABSTRACT

The genus Polemonium L. (Polemoniaceae) is predominantly western North American but has several repre-

sentatives in eastern North America, Europe, and Asia. One of the most widespread and also polymorphic

species is P. boreale Adams, described from the mouth of Lena River in northern Siberia, but with a circumpolar

distribution in the Arctic and extensions southwards into northern boreal mountains. Hulten (1948) stated

that he had worked with Davidson on his material and that they were unable to find characters for subdivid-

ing P. boreale. Accordingly, in his monograph of the genus, Davidson (1950) treated the species collectively.

However, years later, Hulten (1967) formalized variation in Alaska as 1) subsp. boreale (implied), 2)

subsp. macranthum (Cham.) Hulten, based on P. humile macranthum of Chamisso (1831), described from St.

Paul Island and reported by Hulten from “along the southern coast of western Alaska,” and 3) var. villosissimum

Hulten, based on a collection from Cathedral Mountain, Denali National Park, in the Alaska Range (holotype:

S!; isotype: ALA!). Whereas Hulten (1968) mapped subsp. macranthum with a range in southwestern Alaska,

he merely mentioned var. villosissimum in passing as a plant of the high mountains of central and southern

Alaska. His diagnosis of var. villosissimum is brief: “Inflorescentia dense capitata longe albo-villosa.”

The tall-growing and large-flowered plants Hulten assigned to subsp. macranthum represent an endpoint

of continuous variation within P. boreale and do not, in our view, merit taxonomic recognition. Variety vil-

losissimum is another matter. This taxon is now documented at ALA from more than 35 sites from the St.

Elias Mountains of southwestern Yukon throughout the Alaska Range and the more southern mountains

westwards to the Alaska Peninsula and the Kilbuck-Kuskokwim mountains (Pig. 1). These specimens show

several differentiating characters in addition to the two emphasized by Hulten. Variety villosissimum is for

the most part sympatric with P. boreale in central and southern Alaska, but it is generally found at higher

elevations and m screes, often irrigated by meltwater from snowbanks. None of the specimens of var. villosis-

simum show features transitional to P. boreale ; thus, we conclude that Hulten’s var. villosissimum represents

a fully independent species and hereby combine it as such with an emended description.

Perennial, with a prolonged, distally richly branched root-stock typical of scree plants; 5-20 cm tall; :

J.Bot. Res. Inst. Texas 5(1): 19 -2

Journal of the Botanical Research Institute of Texas 5(1)

m, 60 65'N, 138 22'W, 22 Jul 1969, D.F. Murray 3015 (ALA). Photo: D.F. Murray.

solitary to several, erect or suberect from dense cluster of basal leaves; basal leaves 2-8 cm, with (9-)

11— 21(— 23) obliquely ovate to obovate leaflets, apices rounded to obtuse, 4-10 x 3-8 mm, decurrent and

confluent with distinct rachis wings, villous; stem leaves 2-3, similar to basal leaves; inflorescence strongly

congested, 5-10 flowers or more, compact and head-like until maturation of the fruits; pedicels, bracts and

calyces long-villous, eglandular; pedicels less than half the length of the calyx, 3-7 mm; proximal bracts

foliar, distally absent; calyx 7-10 mm, campanulate, segments equalling or slightly shorter than the tube,

narrowly triangular-ovate, apices obtuse or rounded; corolla narrowly campanulate, 10-15 mm long, 5-10

Journal of the Botanical Research Institute of Texas 5(1)

Fig. 3. Polemonium boreale Adams. Voucher: USA. Alaska. Harrison Bay Quad.: National Petroleum Reserve-Alaska. Vic. Fish Creek test well 1, ca. 30 m.

7019'N, 151 58'W, 27 Jul 1977, D.F. Murray & AM Johnson 661 9(ALA). Photo: D.F. Murray.

mm broad, lobes narrowly obovate to broadly linear, 7-11x3-6 mm, longer than the tube, cream to purplish

inserted well below the stamens, anthers yellow; flowers with sweet fragrance.

We have examined specimens of P. boreale from arctic Europe, Siberia, Canada, Greenland, Alaska and

Yukon and found none that approached P. villosissimum. Diagnostic characters between these two species

are given in Table 1 and several of these features are evident in Figures 2-3.

A NEW VARIETY OF HOLODISCUS DUMOSUS (ROSACEAE: SPIRAEOIDEAE)

FROM COASTAL NORTHWESTERN CALIFORNIA

Roger Raiche

James L. Reveal

L.H. Bailey Hortorium, Department of Plant Biology

Ithaca, New York 14853, USA.

jlr326@cornell.edu

Forestville, California 95436, USA.

roger@planethorticulture.com

ABSTRACT

RESUMEN

The Cedars of northwestern California comprises about 6,000 acres of ultramafic (high magnesium and

iron) rock and derived soils located in Sonoma Co. These rocks are commonly referred to as serpentine.

Serpentine has a profound effect on vegetation due to an imbalance of magnesium to calcium, and a lack

of essential nutrients for plant growth. Geophysical isolation of The Cedars from other serpentine areas,

dramatic topographic relief, high winter rainfall and unusually hot summer temperatures, are factors that

have fostered an exceptionally high level of endemic taxa on this geologic “island.” They are: Arctostaphylos

bakeri Eastw. ssp. sublaevis P.V. Wells, Calochortus raichei Farwig & Girard, Epipactis gigantea Hook. f. rubrifolia

P.M. Br., Erigeron serpentinus G.L. Nesom, Eriogonoum cedrorum Reveal & Raiche and Streptanthus glandulosus

Hook. ssp. hoffmanii (Kruckeb.) M.S. Mayer & D.W. Taylor. In addition to the endemic taxa, many others

are disjunct or represent a range extreme at The Cedars.

The Cedars Holodiscus was first noted by Raiche in 1981 who introduced the plant into the University

of California Botanical Garden at Berkeley in 1982 where it has persisted in cultivation ever since. He noted

that the plant differed significantly from the common coastal oceanspray, H. discolor (Pursh) Maxim, which

occurs less than a mile away on non-serpentine substrate. Using the most recently published monograph by

Ley (1943), he attempted to ascertain in this complex genus of eight species where this particular expression

fit best. He initially sought to fit The Cedars plant into one of the published expressions of H. discolor only

to conclude that it persistently tended to key out to what Ley termed H. microphyllus Rydb. and specifically

the var. glabrescens (Greenm.) Ley, a plant found in the Sierra Nevada and mountains of central and eastern

Oregon eastward to Wyoming, Utah and northwestern Arizona, well away from The Cedars, a mere seven

air miles east of the Pacific Ocean in Sonoma Co., California.

By studying the plant over the years, and growing the plant in several different locations, noting that it

retained its consistent features, Raiche concluded that the plant was distinct and required formal recogni-

tion. Working with Reveal while they jointly examined another local endemic, Eriogonum cedrorum (Reveal

& Raiche 2009), they began a detailed study of the two species of Holodiscus found in the United States and

J.Bot. Res. Inst. Texas 5(1): 25 -3

Fig.1 . Holodiscus dumosus var. cedrorus. Flowering panicles. Fully open flowers ± 6 mm across, arranged in compound panicles, the secondary panicle

branchlets subtended by leaf-like bracts. All young growth, including inflorescence (including sepals) suffused with red or pink coloration.

Journal of the Botanical Research Institute of Texas 5(1)

, 1. 8-2.1 mm long, 0.9-1 mm

(30-)35-45(-50) prominent, sessile or stalked glands per face (Fig. ft

foliage without magnification. Branch color in transition from bright red leaf base sometimes decurrent along petiole for all or part of length

to cinnamon, typical of first year growth. (lower center leaf).

Distribution . — Restricted to serpentine substrates at The Cedars 200-620 m elev, Sonoma Co., California

(Fig. 6). May-Jul.

May 1987, Raiche 70194 (JEPS); The Cedars, in the canyon above Austin Creek near Layton Mine, on serpentine gravelly slopes at 1350

ft elev., 38°37T6"N, 123°07'37"W, T9N,R12W, sec. 13, 28 Jul 2009, Reveal & Raiche 8991 (BH; UC).

Our recognition of Holodiscus dumosus as distinct from H. discolor, following Hitchcock (1961) and Holmgren

(1997), rather than merging the two as proposed by Lis (1993) who then recognized H. microphyllus at the

species rank rather than including this expression within H. dumosus, is based on a broad knowledge of these

plants in the field and the original type material. The two species recognized here are not known to hybridize

and their distinction in the Pacific Northwest, and especially in Idaho, where their ranges overlap — although

the populations never intermix — are consistently and easily recognized. Even in California the ranges of

H. discolor and H. dumosus do not overlap geographically, and even when the two are in close proximity, as

is the case of the new variety proposed here, they occur in markedly different ecological settings. Contrary

to Lis, a more troublesome distinction is between H. dumosus and H. microphyllus, and so much so in the

Intermountain West that Welsh (2003) joined Hitchcock and Holmgren in not recognizing H. microphyllus

Journal of the Botanical Research Institute of Texas 5(1)

Distribution of Holodiscus dumosus var. cedrorus in NW Sonoma Co., CA

at The Cedars and contiguous ultramafic rock areas.

is Jeps., etc. On talus slopes the var. c

tsvar. sericeus by Ley) s

the bright, ruby red of the stems, branches and petioles is the most <

CYMOPTERUS SPELLENBERGII (API ACE AE),

A NEW SPECIES FROM NORTH CENTRAL NEW MEXICO, U.S.A.

Ronald L. Hartman

Jill E. Lars^j-

Rocky Mountain Herbarium

Department of Botany, Dept. 3 1 65

University of Wyoming

iOOOE. University Ave.

Laramie, Wyoming 82071, U.S.A.

Black Hills National Forest

201 4 North Main St.

Spearfish, South Dakota 57783, US.| |-

jilllarson@fs.fed.us

ABSTRACT

RESUMEN

North American

More than three decades ago, Richard Spellenberg of New Mexico State University f

specimen of umbel from north central New Mexico. He was aware of my ii

members of the family and was curious if it was a novelty.

Initially I assigned it to Aletes sessiliflorus W.L. Theob. & C.C. Tseng, although I knew little regarding

that taxon, as to my knowledge it had been documented only from the type locality 20 miles west of Cuba,

New Mexico (Theobald et al. 1964). Subsequently, I have collected numerous specimens of A. sessiliflorus

and have studied the holdings of regional herbaria. Published concurrently with A. sessiliflorus was Aletes

macdougaliiJ.M. Coult. & Rose subsp. breviradiatus W.L. Theob. & C.C.Tseng. At that time the subspecies

was known only from Montezuma County, Colorado, and adjacent San Juan County, Utah. In recent years

several botanists have accumulated numerous specimens of the two taxa from the northwestern quarter of

New Mexico and adjacent Arizona, Colorado, and Utah that bridged the morphological gap. This is most

apparent with regards to leaf size and shape but it is equally true for other characters provided in the com-

parison chart by Theobald et al. (1964, p. 311). In general, it appears that as one moves west from the type

locality of A. sessiliflorus, there is a decline in leaf size, likely due to increased aridity. There are numerous

exceptions to this gradient, possibly based on local environmental conditions. Both taxa have terete fruit

subsp. breviradiatus was elevated to species rank and transferred to Cymopterus (C. breviradiatus (W.L. Theob.

& C.C. Tseng) R.L. Hartm.). It was later placed in synonymy under Cymopterus sessiliflorus (W.L. Theob. &

C.C. Tseng) R.L. Hartm. (see discussion, Hartman 2006; Goodrich & Hartman, in press).

Returning to Spellenberg’s specimen, first collected on Black Mesa just to the west of the Rio Grande

sessiliflorus in leaf and fruit characters. Furthermore, the taxa are distantly related evolutionarily based on

molecular and combined molecular and morphological studies (see below).

Cymopterus spellenbergii R.L. Hartm. & J.E. Larson, sp. nov. (Fig. 1). '

J. Bot. Res. Inst. Texas 5(1): 33 - 40. 201 1

Journal of the Botanical Research Institute of Texas 5(1)

Fig. 2. Detailed morphological features of Cymopterus spellenbergii. A. Fruit in side view with the commissure faintly visible down the center (Sivinski

7613, RM). The remaining images are based on Hartman 11742 (RM). B. Dorsal view of mericarp. C. Mericarp in partial side view; upper part of the

deciduous carpophore branch attached at apex. D. Cross section of mericarp, oil tubes evident.

piningli 4.7 air mi WSW of Picuris Pueblo, T23N R11E sec. 29 SV2 (36.1945°N, 105.7947°W), 7,300 ft, north-facing

ion-juniper woodland, 16 Jun 2010, fruit, R.C. Sivinski 7613 (RM, UNM).

Etymology . — -This species is named for Richard Spellenberg of New Mexico State University who dis-

covered it and for his studies on legumes and nyctages and of the flora the Sierra Madre Occidental, Mexico.

Distribution and ecology . — Cymopterus spellenbergii has been documented from the vicinity of the

Rio Grande in two general areas (Fig. 3). One is in north central Taos County where it is known from four

sites west of Questa. The other is in the southwest portion of that county at four locations and two just to

the southwest in Rio Arriba County, all northeast of Espanola. Of the last two sites, the one indicated by

the large star marks the type locality. It has been collected several times here and in a radius of one to two

mile. Much of the public lands between these two centers of distribution and immediately adjacent to the

Rio Grande have been searched for additional populations. Additional searches especially on private lands

are warranted.

Cymopterus spellenbergii typically is found growing on basalt of the Servilleta Formation (5.6 to 4.5

mya) or their degradation products. These volcanic flows cover much of the Taos Plateau or form caprock

along canyons rims. The novelty may also occur in sandy draws below the caprock. In one case it has been

documented growing on metamorphic (R.C. Sivinski 7613).

Typically this species is found in plant communities dominated by pinon pine and juniper or by pon-

derosa pine and Douglas fir.

Phenology . — Cymopterus spellenbergii generally flowers from the end of April to late May; fruits from

mid May to early July. As noted above, mature mericarps can be found below plants in mid August if there

is adequate plant material covering the ground to hold them in place.

so recognized by the New Mexico Rare Plant Technical Committee and Natural Heritage New Mexico.

Notes . — The closest known sites for Cymopterus sessiliflorus (C. breviradiatus included) is about 35 air

miles due west of the type locality of C. spellenbergii. It was collected at three locations in the vicinity of

Ghost Ranch near Canjilon Creek, 3 to 4 miles north of Abiquiu Reservoir ( Larson 5023, 5024, 10348; RM).

These locations are about 30 air miles northeast of the type locality of C. sessiliflorus. The habitat was dry,

sandy bottom and rocky sides of wash or riparian areas in shales and sandstones of Chinle and San Rafael

groups. Sandstone and shales and their erosional products are the most common substrate for this species,

especially in dissected landscapes.

A reason for the delay in describing this novelty, and others in the past, is the senior author’s insistence

on understanding the geographic distribution and morphological variation in adequate detail. Similarly as

with Cymopterus constancei R.L. Hartm., a nearly thirty year gestation period occurred. In this latter case the

than doubled (Hartman 2000). Between 2001 and 2006 (Reif et al. 2009), more than 35,000 numbered col-

lections were obtained in inventories of the Valles Caldera National Preserve, the Santa Fe National Forest,

the Carson National Forest, and the Taos District of the Bureau of Land Management, all in north central

New Mexico.

These studies are part of the larger effort through 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).

This inventory of north central New Mexico allowed for detailed searches for Cymopterus spellenbergii in

the area where it is now known to occur, but also in a vastly larger region with associated ecological diversity.

Furthermore, work in south central Colorado (Elliott 2000) had led to the pursuit of the Colorado endemic

Neoparrya lithophila Mathias (Apiaceae) in New Mexico. Thus both were sought simultaneously. Subsequently

it was discovered in New Mexico, near the Rio Grande about Vi mile south of Colorado (Hartman et al. 2006;

O’Kane 1988).

Journal of the Botanical Research Institute of Texas 5(1)

America have been published by Stephen R. Downie’s Lab at the University of Illinois (Downie et al. 2002;

Sun 2003; Sun et al. 2004; Sun and Downie 2004, 2010a, 2010b). The accession Hartman 13954 (RM) was

misidentibed by Hartman as Aletes sessiliflorus (here treated in Cymopterus ) and was not corrected subse-

quently. In fact, it represents C. spellenbergii from the type locality. Furthermore, three accessions labelled

Aletes macdougalii subsp. breviradiatus are here recognized as C. sessiliflorus. All three of these collections are

from the western part of the geographic range.

Most molecular analyses were based on nuclear ribosomal ITS and chloroplast rps 16 intron and trnF-

L-T sequences. Sun and Downie (2010a, Figs. 2, 3) also included morphological characters (proportionally

weighted). As stated, “these trees are largely congruent with those trees derived from molecular data alone

(Sun 2003; Sun and Downie 2010b). In fact these trees are better resolved and, in general, their branches are

more strongly supported than any phylogenetic tree for the group heretofore available. . . .” In these papers,

subclade 3f depicts C. spellenbergii and Harbouria trachypleura (A. Gray) J.M. Coult. & Rose as strongly sup-

ported sister taxa with C. bakeri (J.M. Coult. & Rose) M.E. Jones (as Oreoxis) being sister to this clade followed

by Cymopterus filifolius (Mathias, Constance, & W.L. Theob.) B.L. Turner (as Aletes) and then Cymopterus

longiradiatus (Mathias, Constance, & W.L. Theob.) B.L. Turner (as Pseudocymopterus) as successively basal

branching linages. All five species occur in New Mexico but only C. spellenbergii, C. filifolius, and Harbouria

trachypleura have filiform to linear leaflets. Subclade 3c has one taxon that consistently has linear leaflets

( Cymopterus davidsonii (J.M. Coult. & Rose) R.L. Hart.) and another that occasionally does ( Cymopterus

lemmonii (J.M. Coult. & Rose) Dorn). Thus only five taxa from New Mexico in Clade 3 (41 accessions) has

or may have this feature. Cymopterus sessiliflorus (as Aletes macdougalii subsp. breviradiatus) is in subclade

lg (Clade 1 consists of 60 accessions) and is quite distant phylogenetically from subclades 3f and 3c. The

conclusion is that C. spellenbergii is always associated with the taxa in subclade 3f and does not represent

C. sessiliflorus (subclade lc).

The concluding paragraph of Sun and Downie (2010a) discusses the recent trend of including related

genera in Cymopterus stating that “given this trend and overlapping [morphological] character variation

among genera, it may very well be possible that future studies will indicate that all or most members of the

group [perennial endemic North American apioids] should be combined into one large, polymorphic genus,

an extreme but possibly inevitable action.”

A key is provided for the species of Clade 3 (Sun and Downie 2010a) that have or may have filiform to

linear leaflets and occur at low to mid elevations in New Mexico. Where a combination is available under the

genus Cymopterus, that name is used. Many of these combinations were made by Turner (1998). Cymopterus

sessiliflorus ( Aletes macdougalii subsp. breviradiatus) in Clade 1 is included for comparison. Cymopterus davidsonii

has been collected sparsely, especially in fruit. Thus its dmerentiation from C. lemmonii needs clarification.

1. Plant acaulescent.

2. Peduncles glabrous throughout or slightly scabrous above; fruit with membranous wings ( Aletes

macdougalii subsp. breviradiatus ) C. sessiliflorus

2. Peduncles minutely puberulent (20x) with peg-like hairs at apex; fruit^^"fhick, corky winijS or q ■ f

. Plants caules|j||p^th 1 or 2 reduced Id^es subtending branch(s).

3. Frgft#$|9tely papillate at least between the wings.

4. Fruit subterete (Pteryxia davidsonii)

4. Fruit dorsally flattened ( Pseudocymopterus montanus )

3. Fruit glabrous or densely warty throughout.

5. Fruit smooth {Aletes filifolius)

5. Fruit prominently and densely warty

Cymopterus spellenbergii and C. filifolius are very similar in appearance but may be differentiated by the char-

acters in the key. Furthermore, the latter has more highly dissected leaves and the sepals in fruit elongate and

become indurate, often with one or more hooked inward (versus deltoid to obsolete, papery, not enlarging in

fruit). A generality can be made regarding the substrate on which these three species occur: C. spellenbergii

grows primarily on or adjacent to basalts, C. filifolius grows primarily on limestone but sometimes on rhyolite

or other volcanics, and C. sessiliflorus is found on sandstones and shales. The distribution of C. filifolius is t0

the south of C. spellenbergii by at least 100 miles in eastern Torrance County on the east slope of Manzano

Mountains and in northeastern Socorro County in the Los Pinos Mountains. It has been collected in both

Journal of the Botanical Research Institute of Texas 5(1)

counties on Chupadera Mesa. To the south and southeast it has been documented also in Dona Ana, Eddy,

Lincoln, and Luna counties, New Mexico, and in Brewster, Culberson, Davis, and Hudspeth counties, Texas.

ACKNOWLEDGMENTS

We wish to thank B. “Ernie” Nelson for assistance in the held and in the RM and Larry Schmidt, Dennis

Moser, and Josh Irwin for assistance with the graphics. Curators of the following herbaria are thanked for

loans: ASC, BRY, COLO, UNM, UT. We acknowledge Santa Le National Lorest, Carson National Lorest and

the Taos District of the Bureau of Land Management for providing funding. Stephen R. Downie, Robert C.

REFERENCES

Downie, S.R., R.L. Hartman, F-J. Sun, and D.S. Katz-Downie. 2002. Polyphyly of the spring-parsleys ( Cymopterus ):

molecular and morphological evidence suggests complex relationships among the perennial endemic taxa

of western North American Apiaceae. Canad. J. Bot. 80:1 295-1 324.

EluotI^^OOO. A vascular flora of south-central Colorado. Master's thesis, Univ. of Wyoming, Laramie.

Goodrich, S. and R.L. Hartman. Apiaceae. In: Bolack San Juan Basin Flora. Missouri Bot. Garden Press (In press).

Hartman, R.L. 1992. The Rocky Mountain Herbarium, associated floristic inventory, and the flora of the Rocky

Mountains project. J. Idaho Acad. Sci. 28(2):22-43.

Hartman, R.L. 2000. A new species of Cymopterus (Apiaceae) from the Rocky Mountains, U.S.A. Brittonia 52:1 36-1 41 .

Hartman, R.L. 2006. New combinations in the genus Cymopterus (Apiaceae) of the southwestern United States.

Slda 22:955-957.

Hartman, R.L. and B.E. Nelson. ,2008. General information for floristics proposals [The Boiler Plate], URL: http://www.

rmh.uwyo.edu/research/GenerallnformationforFloristicsProposals.pdf

Hartman, R.L., B.£ Nelson, and B.S. Legler. 2009. Rocky Mountain plant specimen database: URL www.rmh.uwyo.edu).

Hartman, R.L., B. Reif, B.E. Nelson, and B. Jacobs. 2006. New vascular plant records for New Mexico. Sida 22:1 225-1 233.

O'Kane, S.L., Jr., D.H. Wilken, and R.L. Hartman. 1 988. Noteworthy collections: Colorado. Madrono 35:72-74.

Reif, B., J. Larson, B F*ja§$$, B E Nelson, and R.L. Hartman. 2009. Floristic studies in north central New Mexico, U.5.Af}

TheTusas Mountains and the Jemez Mountains. J. Bot. Res. Inst. Texas 3:921-961.

Sun, F-J. 2003. A phylogenetic study of Cymopterus and related genera (Apiaceae). Ph.D. dissertation, Univ. of

,B°i s at Urbana-Champaign, Urbana.

Sun, F-J. and S.R. Downie. 2004. A molecular systematic investigation of Cymopterus and its allies (Apiaceae) based on

phylogenetic analyses of nuclear (ITS) and plastid (rpsl 6 intron) DNA sequences. S. African J. Bot. 70:407-41 6.

.^||j-J., S.R. Downie, and R.L. Hartman. 2004. An ITS-based phylogenetic analysis of the perennial, endemic Apiaceae

subfamily Apioideae of western North America. Syst. Bot. 29:419-431.

Sun, F-J. and S.R. Downie. 201 0a. Phylogenetic analyses of morphological and molecular data reveal major clades

within the perennial, endemic western North American Apiaceae subfamily Apioideae. J. Torrey Bot. Soc.

137:133-156.

Sun, F-J. and S.R. Downie. 2010b. Phylogenetic relationships among the perennial, endemic Apiaceae subfamily

Apioideae of western North America: additional data from the cpDNa^®^^^nT region cor^t^to sup-

port a highly polyphyletic Cymopterus. PI. Div. Evol. (formerly Bot. Jahrb.) 1 28:1 51 -172.

Theobald, W.L., C.C. Tseng, and M.E. Mathias. 1964. A revision of Aletes and Neoparrya (Umbelliferae). Brittonia

16:296-315,

Turner, B.L. 1998 [2003], Cymopteris (Apiaceae) in Trans-Pecos, Texas. Phytologia 85:331-335.

A NEW HYBRID OF HIBISCUS (MALVACEAE) FROM TEXAS

Wendy Weckesser

Journal of the Botanical Research Institute of Texas 5(1)

parent species are sympatric across most of their ranges (Correll & Johnston 1979; Fryxell 1988; Turner et

al. 2003). Hibiscus coulteri is found from California to West Texas and in adjacent Mexico. Hibiscus denudatus

is found from Arizona to West Texas and in adjacent Mexico.

Previously, a suspected hybrid was collected in the vicinity of the Eagle Mountains in Culberson County,

Texas (Warnock, Johnston & Powell 17991 SRSC). Close examination of that specimen revealed characters that

match those of Hibiscus coulteri and not those of the hybrid described here. No other Hibiscus collections in

SRSC share the characteristics of the hybrid. No hybrid specimens were uncovered as a result of a search

of the Hibiscus collections at TEX. Possible occurrences of hybrids in other areas where populations of both

species exist warrant further attention. Further herbaria searches may yield as-yet unrecognized collections

of the hybrid.

Morphology. — The suspected hybrid was first noticed by the color of its flower, creamy white with purple

streaking along the margins of the petals (Fig.l). In the Big Bend area, the observed color for H. coulteri is

a pale to bright yellow, and for H. denudatus lavender to pale purple-pink. When dried, the flowers of the

hybrids tend to turn purple with some white mottling. An exception to this is seen in the collection from

BBNP ( Alex 108 SRSC). The flower dried predominantly yellow with purple mottling. It can be seen in

herbarium specimens (SRSC) that the flowers in each of the parent species tend to retain, with some fad-

ing, their original color when pressed. Purple streaking occurs in some herbarium specimens of H. coulteri

(SRSC). In published descriptions of the parent species, the petals of H. coulteri are variously described as

“whitish to lemon- or sulfur-yellow and commonly reddish- or purple-tinged” (Correll & Johnston 1979),

“usually yellow, perhaps whitish or purple-tinged” (Powell 1998), and “yellow with a small (often striate)

maroon spot at base or spot absent” (Fryxell 1988). In H. denudatus, the petals are “lavender-purple” (Correll

& Johnston 1979), “lavender to purplish” (Powell 1998), or “lavender or white” (Fryxell 1988). Normal petal

color variation in the parent species may explain the petal color in the hybrid and the variation seen in the

pressed hybrid flowers.

In the absence of flowers, the hybrid plants exhibit gross characteristics of H. coulteri: few to several

stems arising from a woody root, some lower leaves entire, ovate to cordate with dentate margins, and upper

leaves deeply three-lobed. Given these similar characters, the white flowers might be seen as pigment vari-

ability within the species. Upon closer examination, however, the hybrids consistently exhibit intermediate

forms for hair, sepal, and bract characters. Because there is a marked difference in the characters of the

parent species, it is possible to identify the intermediate forms of these characters in the hybrid.

The hairs of the parent species are distinct. In H. coulteri the stellate hairs have 4 long rays, up to 1 mm,

sometimes obvious without magnification, “aligned 2-by-2 longitudinally with the stem axis” (Fryxell 1988),

with sufficient overlap on stems and petioles to produce a hirsute appearance. Hibiscus denudatus also has

stellate hairs, but the rays are numerous, arranged radially, and much shorter than those of H. coulteri. The

stellate hairs are so densely distributed that leaves, petioles, and stems appear tomentose. In contrast to the

parent species, the hairs of the hybrid resemble those of H. coulteri in length of the rays, but with 3-6 rays

arranged radially as in H. denudatus.

In H. coulteri, the bracts are nearly as long as the sepals (up to 20 mm). In H. denudatus, the bracts (no

longer than 4 mm) are markedly shorter than the sepals (11-15 mm), less than half their length (Fryxell

1988). In the hybrid (Fig. 2), the bracts (7-10 mm) are intermediate between the parent species and are

usually just over half the length of the sepals (15-17 mm).

The differences in fruit and seed characters between the parent species are narrowly defined. There is a

degree of overlap that precludes using these characters in distinguishing the hybrid plants from the parent

species. The fruits of H. denudatus are globose and glabrous or pubescent at apex. The fruits of H. coulteri

are ovoid and antrorsely hispid apically (Correll & Johnston 1979; Fryxell 1988). The fruits of the hybrids

do show some pubescence apically. The only marked difference seen in the few hybrid fruits available for

observation is that they seem to be laterally compressed, which may indicate incomplete development of

carpels or incomplete seed development. The seeds of both parent species are sericeous, completely covered

H. denudatus ( 2n=22 ) t

CAREXAUSTRODEFLEXA (CYPERACEAE),

A NEW SPECIES OF CAREX SECT. ACROCYSTIS FROM THE

ATLANTIC COASTAL PLAIN OF THE SOUTHEASTERN UNITED STATES

NC Natural Heritage Program

and UNC Herbarium, CB 3280

Chapel Hill, North Carolina 27599, U.S.A.

bruce.sorrie@ncdenr.gov

Richard J. LeBlond

PO Box 787

Richlands, North Carolina 28574, U.SA

richardleblond@charter.net

Patrick D. McMillan

Museum of Natural History

Clemson University

Clemson, South Carolina 29634, U.S.A.

pmcmill@clemson.edu

„ : Hyatt

610 East Sixth Street

Mountain Home, Arkansas 72653, U.SA.

sedgehead@gmail.com

Brian van Eerden

The Nature Conservancy

530 East Main Street

Richmond, Virginia 2321 9-2428, U.SA.

bvaneerden@tnc.org

Loran C. Anderson

R.K. Godfrey Herbarium

Florida State University

Tallahassee, Florida 32306-4370, U.S.A.

lorananderson@bio.fsu.edu

ABSTRACT

Botanical and ecological studies during the 1990s of the fire-maintained vegetation of the Atlantic and Gulf

Coastal Plain of the Southeastern United States uncovered a distinctive species of wetland sedge in the

genus Carex. Remarkably, this entity was independently discovered by the six authors of this paper. This

unfamiliar Carex was clearly a member of the section Acrocystis, due to the combination of pistillate spikes

produced on short, subradical peduncles and an elongate lower pistillate bract. In most characters it keyed

to Carex deflexa Hornem. Closer examination of the material, combined with additional field and herbarium

searches, revealed a number of distinctive features. It became apparent that this entity did not match any of

the species descriptions in the most recent complete treatment of section Acrocystis (Crins & Rettig 2002).

The new species is described as follows:

Plants perennial, moderately to loosely cespitose by means of slender rhizomes (2.5— )6— 9(— 12) cm long,

,&7~1 't .mm thick, covered with red or maroon-red sheaths at time of flowering. Leaves predominantly

basal, much exceeding culms, principal leaves 11-42 cm long, 1.1-2 .4 mm wide, linear, acuminate, sca-

J. Bot. Res. Inst. Texas 5(1): 45 -5

Journal of the Botanical Research Institute of Texas 5(1)

h 0.9-1 .4 mm wide, a

Sorrie et al., A new species of Carex from the eastern United States

z, 0.65 mi NE of jet Rtes 2 and 629 along

spring branch tributary of Turkey Track Creek, 17 May 2006, GP Fleming 15355 with K Patterson (VPI). Isle of Wight Co.: Blackwater

i, IV jtirtJflfil,

Townsend 2562 (WILLI).

Among eastern North American species of the section Acrocystis, the new species keys readily to Carex dejlexa.

Both species produce terminal staminate spikes closely associated with overlapping, subsessile pistillate

spikes on the primary (distal) inflorescence and also produce pistillate spikes on short basal peduncles. Both

species have long lower pistillate bracts on the primary inflorescence. Carex austrodeflexa may be separated

from C. dejlexa by its longer perigynium beak (0.6-1 mm vs. 0.4-0. 8 mm) and papillose perigynia, which

are normally glabrous or glabrate with a relatively sparse pubescence concentrated at the base of the beak

(vs. the rather copious and widespread pubescence of C. dejlexa). The staminate spike of C. austrodejlexa

is longer and thicker than that of C. dejlexa (3-9 mm long and 1-2 mm thick vs. 2-5 mm. long and 0.5-1

mm thick), and the staminate scales are also proportionately larger. The length of the lowest pistillate bract

in C. austrodejlexa is subequal to the staminate spike, although it may vary from shorter than to greatly

exceeding the staminate spike (vs. typically exceeding the staminate spike in C. dejlexa). Finally, the ranges

of C. dejlexa and C. austrodejlexa do not overlap. The former is a plant of arctic-boreal and north temperate

regions, in the east extending southward to southern New England and disjunctly to the high Appalachians

of West Virginia, North Carolina, and Georgia; the latter is confined to the coastal plain from southeastern

Virginia to northwestern Florida, southwestern Alabama, and northwestern Louisiana.

Carex austrodejlexa could be confused with two other coastal plain species of section Acrocystis that

produce basal pistillate spikes, Carex umbellata Schkuhr ex Willd. and Carex tonsa (Fern.) E.P. Bicknell. They

differ by perigynium shape and indument, staminate spike characters, rhizome length, pistillate bracts, and

habitat. Carex austrodejlexa has elliptic to narrowly obovate perigynia, whereas C. umbellata and C. tonsa have

globose to broadly ovoid perigynia. Perigynia of C. austrodejlexa are glabrous to glabrate and with a papillose

surface; those of C. umbellata are densely puberulent, those of C. tonsa sparsely puberulent or sometimes

glabrous; neither has a papillose surface. In its distal inflorescences C. austrodejlexa always has a staminate

spike closely associated with two pistillate spikes, whereas in C. umbellata and C. tonsa the staminate spike

is associated with one pistillate spike or is solitary. Carex austrodeflexa differs from C. umbellata in its long

slender red rhizomes and matted habit (vs. the short reddish to brown rhizomes and distinctively cespitose

habit of C. umbellata). From C. tonsa, C. austrodejlexa may be separated by its slender, red-sheathed rhizomes

(vs. short, thick rhizomes). The lower pistillate bracts of C. austrodejlexa are typically much longer than

those of C. umbellata and C. tonsa, which have bracts usually much shorter than the staminate spike. Both

C. umbellata and C. tonsa inhabit dry to xeric situations, often in full sun, whereas C. austrodejlexa grows in

wet shaded streamheads and margins of small-stream swamps.

Finally, Carex austrodejlexa may be confused with two other members of section Acrocystis, Carex emmonsii

Dewey ex Torr. and C.jloridana Schweinitz. Carex austrodejlexa shares the bright red sheath character with

emmonsii, and the two species occasionally grow together, but they are easily separated by the long-creeping

rhizomes and basal pistillate spikes of C. austrodejlexa. Some plants of Carex jloridana produce short fertile

culms as well as long ones and thus resemble C. austrodejlexa, but the former is a much coarser plant, has

leaves twice as wide as those of C. austrodejlexa, has longer, much thicker rhizomes forming clonal patches,

has perigynia covered with short pubescence but non-papillate (vs. glabrous and papillate), and occupies

dry to dry-mesic habitats.

Sorrie et al., A new species of Carex from the eastern United States

(Chapm. ex Britton) Small, Sarracenia purpurea L. var. venosa (Raf.) Fern., Sarracenia rubra Walt., Solidago

patula Muhl. ex Willd. var. strictula Torr. & A. Gray, and Xyris platylepis Chapm.

Second, Carex austrodejlexa inhabits non-alluvial swamp forest (also called small-stream swamp for-

est) and especially their ecotones with wet, grass-sedge-herb savannas. In the outer coastal plain of North

Carolina, several populations of C. austrodejlexa occur in association with many globally rare and restricted

species, such as Carex lutea LeBlond, Parnassia caroliniana Michx., and Thalictrum cooleyi Ahles. Common

associates include Acer rubrum, Carex atlantica, Carex leptalea Wahlenb., C. lonchocarpa , C. stylojlexa Buckley,

Liriodendron tulipfera, Morelia cerifera (L.) Small, Osmunda regalis L. var. spectabilis (Willd.) A. Gray, and

Taxodium ascendens Brongn. In this habitat C. austrodejlexa may be a patch dominant in the herb layer.

Carex austrodejlexa flowers early, beginning in late February and often peaking in the first two weeks

of March in the Sandhills region of the Carolinas and eastern Georgia. It reaches its peak fruiting period in

early and mid April. The populations on the outer coastal plain of North Carolina and on the Gulf Coastal

Plain reach floral peak one to two weeks earlier than those in the inland portion of its range.

RARITY AND CONSERVATION

Carex austrodejlexa is not a rare plant from a global perspective. In our opinion it is merely overlooked, as

from a short distance flowering plants can appear to be merely vegetative and fruiting culms wither rapidly

following maturation. In North Carolina new populations are found almost annually without directed

searches. We believe that many more populations will be documented in other southeastern states, once

botanists learn its morphology and habitat. That said, however, we believe that chances of finding C. aus-

trodejlexa increase significantly within areas managed with controlled burns.

ACKNOWLEDGMENTS

The authors wish to thanks curators and staff of the following herbaria for access to specimens: CLEMS, FSU,

MICH, NCSC, NCU. Tony Reznicek (MICH) and two reviewers (Theodore S. Cochrane and one anonymous)

significantly improved the manuscript. Alan Weakley (NCU) provided guidance and support throughout.

Guy Nesom (Fort Worth, Texas) kindly provided the Latin diagnosis.

REFERENCES

Crins, WJ. and J.H. Rettig. 2002. Carex sect. Acrocystis. In: Flora of North America North of Mexico. Vol.,23,

Magnoliophyta: Commelinidae (in part), Cyperaceae. Oxford University Press, New York. Pp. 532-545,

Christensen, N.L. 1 988. Vegetation of the Southeastern Coastal Plain. In: Barbour, M.G. and W.D. Billings, eds. North

American terrestrial vegetation. Cambridge Unit^^p’ress; United Kingdom. Pp. 31 7-363.

Peet, R.K. and DJ. Allard. 1 993. Longleaf pine vegetation of the southern Atlantic and eastern Gulf Coast regions:

a preliminary classification. Proc. Tall Timbers Fire Ecol.Conf. No. 18. Tallahassee, Florida.

Platt, WJ. 1 999. Southeastern pine savannas. Iftf^j&erson, R.C., J.S. Fralish, and J.M. Baskin. Savannas, barrens, and

rock outcrop plant communities of North America. Cambridge University Press, United Kingdom. Pp 23-5V'

Weakley, A.S. 2008. Flora of the Carolinas, Virginia, Georgia, northern Florida, and surrounding areas. Draft of April

2008. Unii^l^ffy of North Carolina Herbarium, Chapel Hill, North Carolina.

Werie^^2006. Carex reznicekii, a new widespread species of Carex section Acrocystis (Cyperaceae) from eastern

North America. Sida 22:1 049-1 070.

Journal of the Botanical Research Institute of Texas 5(1)

BOOK NOTICES

Randall T. Schuh and Andrew VZ. Brower. 2009. Biological Systematics: Principles and Applications,

Second Edition. (978-0-8014-4799-0, hbk.). Comstock Publishing Associates, (Orders: www.cornell-

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750 Cascadilla Street, Ithaca, New York 14851-6525, U.S.A.). $59.95, 328 pp„ 65 illustrations, tables,

charts, graphs, maps, b/w line drawings, glfe" x 9 1 A".

MISSOURI BOTANICAL GARDEN PRESS (ST. LOUIS, MISSOURI)

Orchids

CarlA. Luer, 2010. leones Pleurothallidinarum XXXI: Lepanthes of Bolivia. Systematics of Octomeria.

Species North and West of Brazil (Addenda: New species of Brachionidium, Lepanthes, M asdevallia,

pbk.). Monographs in Systematic Botany from the Missouri Botanical Garden, Volume 120. Missouri Botanical

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$65.00, 154 pp., illus., 7" x 10".

BOOK REVIEW

Elray S. Nixon, with illustrations by Bruce Lyndon Cunningham. 2010. Gymnosperms of the United States

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180 County Road 8201, Nacogdoches, Texas 75964, U.S.A. (Orders: www.artplantaebooks.com/).

$70.00 (hbk.), $59.95 (pbk.), 208 pp., color illustrations, 8V2" x 11".

being described.

of Texas, Fort Worth, Texas 76107-3400, U.S.A.

J.Bot. Res. Inst. Texas 5(1): 52. 2011

PHASEOLUS HYGROPHILUS (LEGUMINOSAE-PAPILIONOIDEAE),

A NEW WILD BEAN SPECIES FROM THE WET FORESTS OF COSTA RICA,

WITH NOTES ABOUT SECTION BREVILEGUMENI

i $mm

Journal of the Botanical Research Institute of Texas 5(1)

Phaseolus hygrophilus Debouck, sp. nov. (Figs. 1-6).

Plant a pluriannual, mid-sized, indeterminate vine to 4-6 m long with 8-16 lateral inflorescences (Fig.

1). Seedling 7-8 cm high, from hypogeal germination, epicotyle green with some red (147A) 1 spots, terete,

30-40 mm long, sparsely covered with minute uncinate hairs. Eophylls petiolate, opposite, 27 x 18 mm, dark

green (144A) with variegation along the midvein, covered with minute uncinate hairs, the blade acuminate,

base somewhat rounded, the petioles green, canaliculate, 15-18 mm long, the pulvini markedly reddish, the

stipules 2 mm long, triangular purplish (59A) 4-nerved. First true leaf trifoliolate (Fig. lb). Root herbaceous,

fibrous, up to 60-80 cm long, tan yellowish brown (22C), much branched at the beginning of growth (Fig.

la), the 4-6 main lateral roots becoming tuberous during the second and subsequent years. Adventitious

roots produced at lower nodes of stems (Fig. 2c), after months becoming tuberous, 8-15 cm long, fusiform

pyriform. Stems terete, green (147D), exposed parts turning reddish in sunlight, the nodes purplish (60B),

internodes 9.5-13.4 cm covered with uncinate trichomes; sprawling horizontally from the lower nodes of

the main stem, rooting with help of adventitious roots, climbing and twining from the upper nodes of main

stem; usually 2-4 shoots in the lower part of the plant, 2-4 shoots in the upper part. Stipules green (141C)

triangular lanceolate basihx multiveined 5x3 mm. Leaves trifoliolate, dark green (136A) with bluish tone,

membranaceous, lustrous, with a clear variegation (138C) along central veins (still apparent on dry herbarium

vouchers) (Fig. Id), covered with uncinate trichomes more abundant on the inner surface as compared to

the upper side. Leaflets lanceolate apiculate, the terminal leaflets with a more acute apex as compared to the

lateral ones, the base rounded. ‘i^fpnal leaflet 7.7(-13.1) x 5.0(-8.0) cm. Lateral leaflet 5.3(-10.8) x 3.3(-7.1)

cm. No lobation observed. Petiole terete, canaliculate, green 3. 3-8. 5 cm. Loliar rachis terete, green, 1.2-2. 8

cm. Proximal pulvini purplish (77A), conspicuous. Distal pulvini green (136B). Stipels green (138B), linear,

narrowly triangular, 2-3 x 1-1 jlpaia, two at the base of the pulvinus of the terminal leaflet, one subtending

the pulvinus of each lateral leaflet. Inflorescence a pseudoraceme (Pig. lc, le; Pig. 2b), appearing as a green-

ish white (145B) axillary spherical mass at the beginning of floral development. Peduncle 40-50 mm long,

terete, scandent, curved in lower part, erect in upper part, almost glabrous with sparsely distributed minute

uncinate hairs, light green (143B). Rachis erect, terete, twice as long as the peduncle, 70-120 mm long, light

green (143A). Number of primary bracts subtending the secondary racemes: 9-16 or more. Primary bracts

5x5 mm, rounded, cupped, multi(8-10)nerved, light green (145D) with many brick reddish dots (79D),

sometimes caducous after anthesis. Pedicels terete, glabrous, light green, 12-16 mm to 20 mm with matur-

ing pod. Pedicelar bracts rounded, cupped, 2.5 x 2 mm. Flowers papilionaceous (Fig. If, lg; Fig. 2d) white

sometimes with standard light pink, two for each secondary raceme (Fig. 2b). Bracteoles (Fig. li) pale light

green (149C), ovate cordiform, 3.5 x 3 mm, 6-veined, a few veins purplish (79D). Calyx round, glabrous,

cupped (Fig. lh), green turning tan at insertion of pedicel when pod matures; tube 3.5 mm long, greenish

(142D) with minute reddish dots (79D); lobes conspicuous, convex (prominent during early ontogeny of

floral buds), two upper lobes rounded 4x4 mm, three lower lobes rounded apiculate 3x3 mm narrowing

to 2 mm at insertion on the tube. Corolla fading cream yellow (13A-C) on the day after anthesis. Standard

(Fig. lj) outer face whitish glabrous with light purple (75B) dots at flexure, inner face white sometimes with

a pink (75D) cast towards the upper half and left auricle, broadly rectangular with sinus 3 mm deep, length

from insertion to flexure 9 mm and from flexure to upper margin 12.5 mm, strong and 12 mm at flexure,

width 14 mm; asymmetrical with left auricle 6 mm wide right auricle 8 mm wide, left auricle with margin

o-Castano et al., A new species of Phaseolus from Costa Rica

Journal of the Botanical Research Institute of Texas 5(1)

Fig. 2. a: habitat of P. hygrophilus. N

close-up of flower of P. hygrophilus (DGD-3172). e: close-up of flower of P. oligospermus (X-16694) (photographs a, c taken at the collection

Debouck&RAraya Villalobos 3172; b, d and e taken at CIAT Popayan station in Colombia).

much reflexed; claw whitish with prominent parallel callosities. Wings (Fig. Ik) pure white, glabrous,

spreading obtusely, left wing 20 x 9 mm, right wing 20 x 10 mm, rounded obovate; margins not revolute

but right wing often superseding on the left one; spur rounded adhering to the keel; claws linear 6 mm long.

Keel tubular (Fig. 11) white greenish at insertion, asymmetrical with two close coils in the terminal part

counterclockwise (facing the keel) with a diameter of 4 mm, the final coil smaller with a diameter of 3 mm,

the two petals joined all along, ca 14 mm long, 3 mm wide up to the non prominent pockets adhering to

the wings, 2 mm wide at the tube, tube 4 mm long to the coils. Stamens diadelphous (9+1), staminal tube

(Fig. lm) 12 mm long delicately veined, vexillary stamen (Fig. In) 15 mm long, claw terete 2 mm long with

rounded cupped knob diameter 2.5 mm, anthers oval dithecal dorsihxed. Ovary (Fig. lo) light greenish,

heavily pubescent, laterally compressed, 7.5 mm long, 2 mm wide, with 4-5 ovules inserted on a delicately

veined basal disk 2 mm long. Style (Fig. lo) 12 mm long, coiled with 15 close coil at the tip, terete, glabrous

but with a loose brush of sparsely distributed hairs below and along the stigma; stigma (Fig. Ip) internal

narrowly triangular 1 mm long, not extending beyond the tip of the style. Pollen (Fig. 4a, 4d) triporate,

spherical, diameter 40 microns, exine finely reticulate, tectum finely punctate. Pods laterally compressed

(Fig. lq), usually 3-seeded, rarely 2 or 4-seeded, when mature 52-64 mm long and 9-13 mm wide, green

(145A) turning dark tan or brown at maturity, with sparsely (densely on 4-5 days old pods) distributed

pubescence of straight coffee brown hairs 0.8 mm long. Upper suture rectilinear, lower suture curved boat

shaped. Base of pod stipitate, narrow, 1.5-2 mm diameter. Beak right, short, 1.5-3 mm long. Seeds circular

and lenticular (Fig. lr; Fig. 3a) 6 mm long, 6 mm wide, 2 mm thick, 6.4 g/ 100 seed, grey (199A) with fine

black spots at higher density around the hilum, shiny.

Etymology. — The epithet has been chosen because of the wet habitat in which the population Jpj

has been found (Fig. 2a). According to local informants just outside the ranch of Santa Eduviges about 1

km from the type collection site, the dry season lasts only about 6-8 weeks in the period April-May.

Geographic distribution, ecology and conservation status. — Phaseolus hygrophilus seems to be the first bean

species found in wet/ rain forests with frequent mist, among tree ferns, many climbing lianas and epiphytes

(Fig. 2a). The original location would correspond through the geographic coordinates to the Bp-P or Bp-MB

(for ‘Bosque pluvial premontano’ and ‘Bosque pluvial montano bajo’, or lower montane rain/ wet forest, ac-

cording to Tosi 1969; Bolanos & Watson 1993, or Gargiullo 2008). According to Herrera-Soto and Gomez-

Pignataro (1993), the type locality would be at the limit of biotic unit 24 (Fig. 5), which corresponds to

temperate tropical cool wet habitat with a dry season of 3-4 months. Associated plants are typical of humid

habitats with often thin broad leaves, and belong to the families: Acanthaceae, Balsaminaceae, Begoniaceae,

Bromeliaceae, Dioscoreaceae, Heliconiaceae, Melastomataceae, Passifloraceae, and Piperaceae. It grows in

the understory of natural openings and on path sides, with stems sprawling on the ground first and then

climbing before flowering. The new species seems to be rare, known from only two sites on the southern

slope of Cerro Buena Vista, extending over an area of 20-30 km 2 (Fig. 5). The two sites from which it has

been reported are on the southeastern border of the Reserva Forestal Los Santos. At the time of our visits in

2003 and 2004 there was little disturbance inside this part of the Reserva but severe deforestation around

Santa Eduviges confirming observations by Sanchez-Azofeifa and co-workers (2003) about forest cover loss

Td$ km-buffer zones of protected areas. We have so far not found P. hygrophilus with any other Phaseolus

species, although it might be at the limit of the range of P. tuerckheimii Donnell-Smith (Fig. 6).

Morphology and possible species relationships in section Brevilegumeni. — The overall morphology of P. hygrophi-

lus suggests a relationship with P. oligospermus, namely the size of the plant, the ferruginous and abundant

pubescence, shape and size of pods, the size and morphology of flowers and seeds. At first glance an out-

standing feature of this species is the clear variegation on all leaflets (Fig. lc). Variegation is often present

along the main vein of terminal leaflets of P. oligospermus, while very well marked and apparently constant

on all leaflets in P. hygrophilus. Leaflet variegation seems only matched by that of P. pedicellatus Benth.,

Journal of the Botanical Research Institute of Texas 5(1)

Fig. 3. Close-ups of seeds, scales at bottom in mm. Clockwise: a: P hygrophilus (DGD-31 72; INB, F, GH). b: P. oligospermus (DGD-2091; CR). c

(DGD-1617; BR, K, MICH, SI, US), d: P. vulgaris (DGD-3168; INB).

Fig. 4. Pollen of P. hygrophilus (left: a and d, clockwise; DGD-31 72) and of P. oligospermus (right: b and c; X-16694). Upper row: polar views (4a, 4b); lower

row: equatorial views (4c, 4d).

although the color of the leaflets is less intense in the latter (Freytag & Debouck 2002). The variegation, which

can still be seen on some voucher specimens, is so striking in P. hygrophilus that it could be of ornamental

value. The base of terminal leaflets is often cuneate in P. oligospermus, and that of lateral leaflets slightly or

clearly lobed (Freytag & Debouck 2002; Zamora 2010), while the base of all leaflets is smoothly rounded

in P. hygrophilus. In both species, leaflets are membranaceous, but shrivel very quickly upon drying in P.

hygrophilus, because of the humid habitat. Tuberous roots are usually cylindrical-conical in P. oligospermus,

and spherical-fusiform in P. hygrophilus; adventitious roots can appear at lower nodes of lateral stems when

they spread onto the mulch, and later become tuberous (Fig. 2c). These lateral shoots eventually flower,

while the original main stem may lose vigor and die. Wings and standard are pure white often with a pinkish

DISCUSSION

The afore-mentioned facts about the new taxon elicit the following points for discussion, in relation to its

taxonomic identity, taxonomic position, and ecology. First, the habit of the plant forming a small colony,

structure of the pseudoraceme, size and slow unfolding of the floral parts (the lobation of the calyx is

noteworthy), morphology of the pod, and lack of development of one seed, all suggest a relationship with

the Brevilegumeni, and particularly P. oligospermus. This however remains a working hypothesis, pending

chromosomal study to discard a relationship with the Falcati Freytag (where 2n = 2x= 20 instead of 22

usually found in the genus: Marechal 1970; Lackey 1980; Mercado-Ruaro & Delgado-Salinas 1996), and

molecular evidence, for instance nuclear ribosomal ITS sequencing (Delgado-Salinas et al. 2006; Serrano-

Serrano et al. 2010). On the other hand, although the authors have not had P. campanulatus in cultivation for

a direct comparison with P. hygrophilus, there are enough differences in primary bract, bracteole and flower

(Freytag & Debouck 2002, p. 224) to consider that the latter is not a disjunct group of populations of the

former, as it happens for P. tuerckheimii across its range. P. tuerckheimii is indeed scattered in woodlands of

high mountains of Chiapas, central Guatemala, central Honduras, eastern Costa Rica and western Panama

just as in biogeographic “islands” (Standley & Steyermark 1946; Lackey & D’Arcy 1980; Breedlove 1986;

Araya-Villalobos et al. 2001; Debouck 2010b; Zamora 2010). Second, P. hygrophilus thrives in premontane

wet/ rain forests with 1-3 months of dry season (Fig. 5, in/ around biotic unit 1); in that habitat, one usually

finds legumes of Desmodieae instead those of Phaseolinae given the challenge for pod opening and seed

dispersal. Phaseolus oligospermus in contrast is found in subhumid premontane forests with 3-4 months dry

season (Araya-Villalobos et al. 2001) (Fig. 5, in/ around biotic unit 4; Fig. 6). Phaseolus tuerckheimii and P.

dumosus Macfady. thrive in humid montane forests at higher altitudes (Freytag & Debouck 2002; Zamora

2010; Fig. 6), where the dry season is longer or well-marked (Fig. 6). From the map (Fig. 5), it seems that

a few more areas combining the right altitude, rainfall amount and pattern and other ecological conditions

could exist along the Talamanca mountain range, where a large protected area has been established (i.e.

‘Parque Internacional La Amistad’), but with grazing on its lower Pacific slopes (Gomez-Pignataro 1986) and

in the distant buffer zone (Sanchez-Azofeifa et al. 2003). The total number of populations of P. hygrophilus

for Costa Rica might be higher than two but possibly not more than a dozen (from a study of the ecological

maps: Bolanos & Watson 1993; Herrera-Soto & Gomez-Pignataro 1993), and additional sampling in the

Reserva Forestal Los Santos would be a logical starting point. The case of P. talamancensis Debouck & Torres

calls for caution sure enough in concluding about endemic species: this taxon was initially thought to have

its distribution restricted to the Talamanca mountain range in Costa Rica with two populations (Torres-

Gonzalez et al. 2001), and it currently has four known populations, one of them outside that mountain range

(Debouck 2010a; Zamora 2010). Finally, it is interesting to note that the number of species of Phaseolus

continues to increase with the disclosure of more endemic species, this time towards the southern part of the

range of most species in western Mexico (Delgado-Salinas 1985; Freytag & Debouck 2002; Ramirez-Villegas

et al. 2010). For Costa Rica, the number of wild bean taxa has almost doubled since the early floristic work

of Paul C. Standley (1937); including the new species here described, ten wild species of Phaseolus are now

known from there (Zamora 2010).

ACKNOWLEDGMENTS

The authors warmly thank the Bundesministerium fur Wirtschaftliche Zusammenarbeit und Entwicklung

(BMZ) of Germany for the financial support, and the University of Costa Rica and the Centro Internacional

de Agricultura Tropical for logistical and administrative support. This activity has been part of the BMZ

supported project “Gene Flow Analysis for Environmental Safety in the Neotropics.” The effective help of

5 de Mexico. IV. Flora de C

i, USA. Pp. 395-775.

st. Texas 5(1): 66. 2

LEPECHINIA YECORANA (LAMIACEAE),

A NEW DIOECIOUS SPECIES FROM THE YECORA AREA OF SONORA, MEXICO

James Henrickson MarkFishbein

Plant Resources Center

University of Texas

Austin, Texas 78712, i |§§|

Oklahoma State University

Stillwater, Oklahoma 74078, USAi

Thomas R.Van Devender

RESUMEN

INTRODUCTION

The broad Rio Mayo floral region covers most of southern Sonora from the Sinaloan border to the Rio Yaqui

in the north (Martin et al. 1998). The completion of Mexico Federal Highway 16 (MEX 16) in 1992 provided

improved access to this portion of the Sierra Madre Occidental in easternmost Sonora (Burquez et al. 1992;

Van Devender et al. 2005). The flora of the Municipio de Yecora, in the northern portion of the Rio Mayo

region, is very diverse with over 1770 taxa in 3,300 km 2 . Floristic publications in the area include the flora

of the unique sphagnum spring at the Cienega de Camilo east of Maycoba, 10 km east of the Chihuahua

border on MEX 16 (Van Devender et al. 2003) and a summary of the distribution of the grasses in the

Municipio (Van Devender et al. 2005). The Yecora area has steadily yielded a wealth of range extensions

and new species in the last two decades.

We describe herein a species of Lamiaceae that is restricted to pinyon-juniper-oak woodland on mottled

pink and whitish volcanic ash in the Yecora area. In its small shrub habit and solitary flowers, the taxon

appears most similar to Lepechinia mexicana (S. Shauer) Epling [= Sphacele mexicana S. Shauer], a shrub spe-

cies occurring in Nuevo Leon, Tamaulipas, Veracruz, San Luis Potosi, Jalisco, Hidalgo, Queretaro, Puebla,

and Oaxaca. Both are well-branched shrubs with small, bullate upper leaf surfaces, with forked to dendritic

vestiture, and with solitary, axillary flowers. However, L. mexicana occurs in rather arid habitats and has

thickened, strongly vestitured leaves and moderately large blue-lavender corollas mostly (8— )9— 12 mm long,

while the new species occurs at higher elevations among pinyons and yellow pine and, at least at the type

locality, the leaves are narrower, thinner, more flaccid, less conspicuously bullate, and the corollas are white

with purple spots at the throat and on the abaxial limb and only 5-7 mm in length. The less rigid leaves

could be explained away as the plants occur in a more mesic habitat. Further observations found that both

L. mexicana and the new taxon appear to be functionally dioecious — a feature otherwise known only in

South American species of Lepechinia section Parviflorae Epling. But first a look at the genus as a whole.

J.Bot. Res. Inst. Texas 5(1): 67-7

polyphyletic (p. 156), including both herbs and shrubs, with large or small leaves, and variously branched

inflorescences (as above), or the flowers solitary in the leaves-bracts, and the flowers larger, and hermaph-

roditic (gynodioecious in 2 species, one from Haiti the other from Columbia-Venezuela). Corollas range

from 5-40 mm in length, and range from blue, purple, red, yellow or white in color, and they have a ring of

hairs that form an annulus in the lower tube, though this is sometimes poorly developed. They range from

central Mexico to Argentina. Hart (1985) in his publication on dioecy in Lepechinia, used Epling’s sectional

classification, placing the gynodioecious species outside the section Parviflorae in section Salviifoliae Epling.

It appears, however, that both L. mexicana and the new taxon are dioecious. In the new taxon, from

which one of us (JH) has made mass population collections, both staminate and pistillate flowers have a

fully formed ovary-style-stigma, (the pistillate flowers have a fertile ovary, but four reduced staminoda,

the staminate flowers have 4 fertile stamens and a sterile ovary (Fig. 1 A-B). Only one developing fruit was

found in staminate plant specimens, which would make that plant polygamodioecious. All pistillate plants,

of course, produced fruit. The corollas in the new taxon are white, small, slightly larger in staminate than

pistillate flowers (as in the South American section Parviflorae) (see Fig 2) and both staminate and pistil-

late flowers have a well defined annulus in the lower tube [as in Hart’s (1983) section Lepechinia] but the

hairs are continuous along the corolla tube-throat floor to the area between the abaxial filaments unlike

either section (see Fig. 1 A-B). In contrast, flowers in section Parviflorae have an patch of hairs only

between the abaxial filaments and an annulus is absent. The reproductive condition in L. mexicana is less

resolved. Few flowers or fruit were found in the specimens examined. Staminate flowers were observed, as

were flowers with aborted anthers, and as in the new taxon, both staminate and pistillate flowers had fully

formed styles. But as the specimens had few flowers, it was not possible to make many observations.

The new taxon and L. mexicana appear to represent an independent origin of dioecy in the genus,

^^interesting that, at least in the new taxon, it has conformed to certain features also found in section

Parviflorae (i.e., small white corollas that are slightly larger in staminate flowers, the retention of staminoda

in pistillate flowers and of a style in staminate flowers). But whereas in section Parviflorae the white corollas

have purple spots on the largest abaxial lobe, in the new taxon there are purple spots along the rim of the

throat, with a few on the abaxial lobe. Furthermore, the new taxon has a basal annulus, hairs between the

abaxial filaments, and a continuous patch of hairs between the abaxial filaments and annulus (Fig.l, A-B).

SYSTEMATICS

Lepechinia yecorana Henrickson, Fishbein, & T. Van Devender, sp. nov. (Figs. 1A-E, 2). Type: Mexico. Sonora:

came ash slope, flowers white, leaves aromatic with light anise odor; near 28 o 26'09"N, 108°31‘22*W, 1600 m, 29 Mar 1997, T.R.

Rounded to spreading, oppositely, usually densely branched, suffrutescent, dioecious (rarely polygamodioe-

cious) shrubs to 1.2 m tall. Young stems slender, erect to spreading; internodes 3-15(-24) mm long, initially

4 ribbed, soon rounded, often ± purplish-red, pubescent with ± thick-based, translucent, simple to distally

forked or alternately branched dendritic hairs 0.02-0.2(-0.4) mm long, the lateral hair branches thick to

slender, to 0.05-0. l(-0. 15) mm long, also with scattered clear, sessile glands to 0.08 mm in diameter. Leaves

opposite, decussate, lanceolate, ± hastate, 13-20(-26) mm long, (2.6-)4-6(-9) mm wide, [leaf length/width

ratio 1.9— 3(— 4)] bluntly acute to obtuse at the tip, with a single pair of short, divergent teeth in the lower third

(or these absent in some leaves), truncate to abruptly cuneate below to a typically broadly tapered, winged

Journal of the Botanical Research Institute of Texas 5(1)

d, showing smaller size, note reduced staminodes with short filaments,

smaller anthers. C. Calyx at anthesis showing bract and bractlets. [

calyces, the upper cut open showing nutlet. F. Nutlets are hard, black smooth with a asymmetrical attachment scar. All from liquid-preserved material

of Henrickson 24629. G-K. Lepechinia mexicana. G. Staminate corolla expanded (as in A) showing annulus, local patch of larger hairs in throat, fertile

stamens, and showing ovaries, style and style lobes that do not mature. H. Pistillate corolla expanded (as in B) showing smaller size, note reduced

staminodia, annulus, and patch of long hairs on corolla throat and fertile gynoecium. I. Calyx at anthesis showing bractlets, longer sepal lobes. J. Fruiting

calyx, often with 2-3 nutlets are broader and have stronger development of lateral venation. K. Mature leaves are thicker, more strongly bullate,

more ovate. G-J from Cisneros 241 7 (TEX-from Puebla); H, I, K from Chiang etal. 2413 (TEX-from Oaxaca). Scale: short bars = 1 mm; long bars = 1 cm.

Henrickson et al., Lepechinia yecorana, a new s

D. Staminate flower, face view showing white anthers in upper corolla throat. E. Pistillate flower side view showing a drooping abaxial corolla lobe.

F. Leaf and developing calyx. G. Partially mature calyces revealing single nutlets within, note lack of horizontal vein thickening. H. Same, cut open to

show single nutlet. All from Henrickson 24629.

petiole 3.6-7(-10) mm long (but cordate when the lateral teeth are closer to the base), the margins straight to

the crenulations 1 1-14 per 10 mm of margin, the upper surface bullate-rugose with weak to strongly impressed

veins, the veins distinctly raised beneath, the blades ± thin, (more thickened and then coarsely bullate with

long and with yellowish glands to 0.08 mm wide on both surfaces but often more dense abaxially, the de-

veloping and youngest leaves gray canescent with a closer vestiture. Flowers spreading to ascending, usually

peduncles 0.5-1. 5 mm long; paired bracteoles linear-oblanceolate to subulate, 2-7 mm long, 0.2-0.4(-0.5)

Flowers of the two species are of similar configuration (Fig. 1), but the corollas of L. mexicana are larger

(9-15 mm total length), lavender, purplish, or blue, with small white spots, the filaments are longer, the

divergent anther sacs are longer (0. 8-0.9 mm) and the calyx lobes are longer and more attenuate, (2.5-4

mm long, to 5.6 mm long in fruit), the calyces become widely inflated in fruit around a larger number of

nutlets, and the cross veins typically thicken as is characteristic of Lepechinia. In addition the hairs of the

annulus are not continuous with the hairs at the base of the central lobe in L. mexicana, as they are in L.

yecorana. However, it must be mentioned that a specimen from near the city of San Luis Potosi, some 1000

km to the south-southeast, also has smaller, white, purple-spotted flowers (± 6.9 mm long) but the specimen

has broader, ovate, thickened leaves as in L. mexicana.

Hart (1983, 1985) noted that dioecy is rare in the Lamiaceae, previously known only in the South

African genus Iboza N.E. Br. (Brown et al. 1910), which has been transferred into Tetradenia Benth. (Codd

1983), and in section Parviflorae of Lepechinia. In Lepechinia Hart noted that dioecy evolved from gynodioe-

cism at least two times in section Parviflorae. The situation in Lepechinia yecorana and L. mexicana appears

to be an independent origin of dioecy in the family. In these, flowers of both sexes have styles, but in the

male flowers the styles are not functional, except in rare instances where male plants produce fruit, making

the species rarely polygamodioecious. It is remarkable that the flowers of L. yecorana are similar in size and

coloration to those of Lepechinia section Parviflorae. Ana L. Reina-G. was the first to note that the new taxon

exhibited dioecy “about half of plants with fruit” (A.L. Reina-G. 97-1428- in herbarium).

Other species of Lepechinia present in pine-oak forests in the Rio Mayo region in Sonora include L.

caulescens (Ortega) Epling in Sierra Saguaribo at 1430 m (Martin et al. 1998) and on Mesa del Campanero at

1740-2150 m (ca. 69 km W of the westernmost L. yecorana locality, Martin et al. 1998). Also L. schiedeana

(Schltdl.) Vatke occurs at 1700-2200 m in pine-oak forest near the cascada de Basaseachic, Chihuahua

(about 60 km E on MEX 16 of the eastern L. yecorana locality; Spellenberg et al. 1996; Martin et al. 1998).

Both of these species have much larger leaves: L. caulescens is a white-flowered, colonial herb with flowers ip

dense, elongated terminal heads; L. schiedeana is a blue-flowered, rhizomatous herb with flowers in terminal

verticils.

In the Municipio de Yecora, pine-oak forest occurs from 1200-1700 m elevation with oak woodlands

on drier exposures at 1,000-1275 m (Van Devender et al. 2005). The Sierra Madre Occidental in this area is

formed of rhyolites and andesites of Oligocene and Miocene ages (Cocheme & Dement 1991). Unusual volcanic

substrates support distinctive local plant distributions and associations in many areas in the Municipio de

Yecora. Areas with hydrothermically-altered, acidic volcanic soils termed glossans typically support pines

and oaks at low elevations in the tropical deciduous forest zone (Goldberg 1982). The Lepechinia sites at 1400

and 1600 m elevation present the opposite situation with relatively xeric pinyon-juniper-oak woodland in

local areas with white or white and pink mottled volcanic ashes surrounded by more mesic pine-oak forest.

Vegetation at the Lepechinia localities is a locally unique pinyon-juniper-oak woodland unlikely to be

found elsewhere. Dominants include Pinus discolor D.K. Bailey & Hawksw. and Juniperus durangensis Martinez

with occasional Quercus oblongijolia Torr., Q. toumeyi Sarg., and Q. viminea Trel. Shrubs include Arctostaphylos

pungens H.B.K., Erythrinaflabelliformis Kearney, and Garrya wrightii Torr., while M ortonia sp., Aralia humilis

Cav., Berberis pimana Laferr. & Marroq., and Comarostaphylis polifolia (H.B.K.) Zucc. were also present at the

eastern Lepechinia locality.

ACKNOWLEDGMENTS

We thank Neil Harriman for the Latin diagnosis; Norma B. Valganon for Figure 1 delineation; Adolpho Espejo

Serna for the Spanish translation and comments, Richard Felger, Adolfo Espejo-Serna, and an anonymous

reviewer for comments; and Ana Lilia Reina- Guerrero, who has visited the Yecora area for two decades,

who has helped document this rich and diverse flora, and discovered a number of new species, including

Lepechinia yecorana .

COLUMNEA BIVALVIS (GESNERIACEAE), A NEW SPECIES

FROM THE EASTERN SLOPES OF THE ECUADORIAN ANDES

Marisol Amaya-Marquez*

Apartado 7495, Bogota, COLOMBIA

John L. CllpS 1 •

Department of Biological Sciences

Tuscaloosa, Alabama 35487, USA.

Corresponding author

ABSTRACT

RESUMEN

INTRODUCTION

The genus Columnea belongs to the New World subfamily Gesnerioideae. It is the most diverse genus in

the subfamily with over 200 species (Skog & Boggan 2006; Weber 2004; Burtt &Wiehler 1995). The sub-

division of Columnea sensu lato into sections or into segregate genera has caused much controversy and

taxonomic confusion (Kvist & Skog 1993; Wiehler 1973, 1983). We recognize the classification based on

recent phylogenetic hypotheses that strongly support the monophyly of Columnea (Smith 1994; Smith &

Sytsma 1994; Clark et al. 2006) and non-monophyly for segregate genera. Thus, the sectional classification

outlined in Kvist and Skog (1993) is more desirable as an informal classification until segregate clades can

be evaluated phylogenetically.

The new species described here belongs to Collandra, the most diverse section in Columnea. The following

characters are useful for recognizing the species within this section: dorsiventral shoots with anisophyllous

subsessile leaves and ovoid (non-globose) berries. In this paper we describe a new species of Columnea that

is known from two populations between 1800 and 2350 m from the eastern slopes of the Ecuadorian Andes.

Epiphytic vine, suffrutescent, often branched; stem terete, 0.3-0. 8 cm, reddish villous (trichomes 7-10

celled), internodes 1-4 cm long. Leaves opposite, strongly anisophyllous in a pair, chartaceous; larger leaf

with petioles 0.3-0. 9 cm long, densely reddish villous; blade asymmetrical, narrow oblong to oblanceolate,

9.5-20 x 2. 2-4.2 cm, base oblique, apex acuminate, margin dentate; adaxially green, golden villous (trichomes

TAXONOMIC TREATMENT

Columnea bivalvis J.L. Clark & M. Amaya, sp. nov. (Figs. 1 & 2). Type: Ecuador.

J.Bot. Res. Inst. Texas 5(1): 75 -7

Journal of the Botanical Research Institute of Texas 5(1)

/species from Ecuador

Fig. 2. Columnea bivalvis. A. One of the paired bracts removed to show uniformly yellow tubular corolla. B. Dorsiventral shoot showing pendent bracts

(photos by J.L. Clark; from the live plant from which the holotype was collected, J.L. Clark, E. Narvaez &J. Vargas 5693).

Journal of the Botanical Research Institute of Texas 5(1)

green with red venation, villous (trichomes 5-7 celled), 8-10 pairs of lateral veins; smaller leaf sessile, blade

asymmetrical, oblong, 1.5-2 x 0. 3-0.5 cm, base oblique, apex attenuate, margin dentate, adaxially green,

reddish villous (trichomes 7 celled), abaxially green, reddish villous (trichomes 7 celled). Inflorescence

epedunculate with 1 flower per node, larger bracts persistent and paired, green with red margins, outer

surface reddish villous, asymmetrical, broadly ovate, 4-4.7 x 2.5-3 cm, bracteoles 2-4, unequal in size,

narrowly ovate to lanceolate, 1.2-2. 5 x 0.2-1 cm, pedicel 0. 2-0.4 cm long, densely villous (trichomes 4-5

celled). Calyx pale green; lobes 5, nearly free joined only at the base by 1 mm of their length; narrow oblong,

1.5 x 0.3 cm, margin dentate with three glandular teeth on each side, outside densely villous (trichomes

5-7 celled), inside glabrous. Corolla uniformly yellow, tubular, 3. 7-4.5 x 0.8-1 cm, basally gibbous and

slightly oblique in the calyx, gibbosity 0.4 x 0.5 cm constricted apically 0.3 cm, 0.8-1 cm at the widest

part of the tube and constricted again at the limb to 0.7 cm; limb nearly actinomorphic, slightly ampliate

in mid region, lobes rounded, subequal 0.3 * 0.3 cm, outside apically sericeous, glabrescent toward the

base (trichomes 8-12 celled), inside glabrous. Androecium of 4 stamens, filaments 3.2 cm long, glabrous,

basally connate for 0.5 cm; anthers oblong 2.0 x 1.3 mm, connective rectangular, 1.8 x 1.2 mm. Nectary a

single dorsal trilobed gland. Gynoecium with the ovary ovoid, 0.7 x 0.3 cm, densely sericeous; style 2.5-3

cm long, laminar with glandular trichomes; stigma bilobed. Fruit an ovoid berry, 1.5 x 0.8 cm. Seeds light

brown 1.8 x 0.5 mm, elliptic, and longitudinally striate.

Distribution and habitat. — Columnea bivalvis is only known from two localities in the wet Andean cloud

forests in eastern Ecuador between 1800 and 2350 m.

Phenology. — Flowers and fruits collected in April and December.

Columnea bivalvis is unique among the species of Columnea by having a pair of large pendent bracts that

enclose a single axillary flower (Figs. 1-2). The yellow tubular flowers are almost completely enclosed by the

bracts with only the throat extending beyond the bract margins. The bracts in C. bivalvis are superficially

similar to the Drymonia hoppii and D. affinis. Although large bracts are common in many Columnea species

and especially those belonging to the section Collandra, no species is known to have large pendent bracts

and dorsiventral shoots.

Columnea bivalvis is similar to C. medicinalis, C. albiflora, and C.eubracteata. The latter three species differ

by the presence of congested bracts compared to a pair of large pendent bracts (i.e., non-congested) in C.

bivalvis, and by the prominent bilabiate corolla limbs compared to a nearly actinomorphic corolla limb in

C. bivalvis.

Etymology. — The new species is named in reference to the marine and freshwater mollusca belonging to

the class Bivalvia because of the resemblance to the two large rounded bracts that enclose a single axillary

ACKNOWLEDGMENTS

We thank the Herbario Nacional Colombia (COF), the Smithsonian Institution’s National Museum of Natural

History - Department of Botany (US), and the Herbario Nacional del Ecuador (QCNE) for access to their

collections. We thank Laurence E. Skog (US) for his willingness to collaborate on the revision of Collandra.

Funding for MAM came from The Gesneriad Society’s Elvin McDonald Research Endowment Fund. Funding

for JLC came from the National Science Foundation (DEB-0841958 & DEB-0949169). We also thank Juan

Carlos Pinzon for the illustration and Pedro Ortiz for help in selecting an appropriate specific epithet and

the Latin description. We thank John R. Clark and Harry Luther for providing helpful comments to an early

version of the manuscript.

REFERENCES

Burtt, 6.L and H. Wiehler. 1 995. Classification of the family Gesneriaceae. Gesneriana J

/species from Ecuador

Clark, J.L, P.S. Herendeen, L.E. Skog, and E.A. Zimmer. 2006. Phylogenetic relationships and generic boundaries in

the Episcieae (Gesneriaceae) inferred from nuclear, chloroplast, and morphological data. Taxon 55:313-336.

Kvist, L.P. and L.E. Skog. 1 993. The genus Columnea (Gesneriaceae) in Ecuador. Allertonia 6:327-400.

Skog, L.E. and J.K. Boggan. 2006. A new classification of the Western I femisphere Gesneriaceae. Gesneriads 56:1 2-1 7.

Smith, J.F. 1994. Systematics of Columnea section Pentadenia and section Stygnanthe (Gesneriaceae). Syst. Bot.

Smith, K.J. Sytsma. 1 994. Molecules and morphology: congruence of data in Columnea (Gesneriaceae). PI.

Syst. Evol. 193:37-52.

Weber, A. 2004. Gesneriaceae. In: Kubitzki, K. and J.W. Kadereit, eds.The families and genera of vascular plants.

Vol. 7. Flowering plants, dicotyledons: Lapi^ff^ (except Acanthaceae including Avicenniaceae). Berlin &

Heidelberg, Germany: Springer-Verlag. Pp. 63-1 58.

Wiehler, H. 1 973. One hundred transfers from Alloplectus and Columnea (Gesneriaceae). Phytologia 27:309-329.

Wiehler, H. 1 983. A synopsis of the neotropical Gesneriaceae. Selbyana

Journal of the Botanical Research Institute of Texas 5(1)

BOOK REVIEW

Mahendra Rai, Deepak Acharya, and Jose Luis Rios (bps.). 2011.

Traditional Knowledge of Herbs. (ISBN 978-1-57808-696-2, hbk.). Science Publishers, EO. Box

699, Enfield, New Hampshire 03748, U.S.A.; marketed and distributed by CRC Press, 6000 Broken

Sound Parkway, NW, Suite 300, Boca Raton, Florida 33487, U.S.A. (Orders: http://www.crcpress.com/

ecommerce_product/product_detail.jsf?isbn=9781578086962; www.scipub.net). $129.95, 504 pp., 11

color plates, 6 Vs" x 9W.

Worth, Texas 76107-3400, U.S.A.

J.Bot. Res. Inst. Texas 5(1): 80. 2011

COLUMNEA LUCIFER (GESNERIACEAE),

A NEW SPECIES FROM NORTHWESTERN ECUADOR

John L, Clark

Department of Biological Sciences, Box 870345

The University of Alabama

Tuscaloosa, Alabama 35487, U.S.A.

jlc@ua.edu

ABSTRACT

The genus Columned is primarily epiphytic and belongs to the New World subfamily Gesnerioideae. Columned

is distinguished from other closely related genera by the presence of an indehiscent berry instead of a fleshy

bivalved capsule. Columned is the most diverse genus in the subfamily Gesnerioideae with over 200 spe-

cies (Skog & Boggan 2006; Weber 2004; Burtt & Wiehler 1995), The traditional sectional classification of

Columned does not represent monophyletic lineages (Smith 1994; Smith & Sytsma 1994; Clark et al. 2006)

and a revised classification system based on molecular sequence data is currently in preparation by the

author, James F. Smith, and other collaborators. The species described here is not assigned to any of the

traditional subgenera or sections.

TAXONOMIC TREATMENT

3, 00°57'21"N, 78°33'38"W > 664 m, 3 Jun 2009,

s: QCNE, NY).

Epiphyte, suffrutescent to herbaceous, stem erect (not scandent), 15-30 cm tall, dense reddish indument

throughout, internodes 1.5-4 cm long. Leaves opposite, strongly anisophyllous to nearly isosophyllous

in a pair, chartaceous; larger leaf with petioles 0. 5-1.0 cm long, densely reddish villous indument; blade

narrowly oblong to ovate, 9.0-11 x 3. 0-4.0 cm, base asymmetrical, apex acute, margin serrate and red;

adaxially green, sparsely golden villous; abaxially green with prominent secondary red venation, uniformly

villous, 3-6 pairs of lateral veins; smaller leaf variable, nearly equal in size or greatly reduced and stipule like.

Inflorescence epedunculate with 1-5 flowers per node, bracts narrowly lanceolate, densely villous and red,

5.0 x 0.3 mm, pedicel 1.0-2. 5 cm long, red, densely villous and red. Calyx bright white with reflexed red

lobes; lobes 5, nearly free joined only at the base by 1 mm of their length; 4 lobes equal in size, ovate, 1.0 x

0.5 cm, margin serrate with 4-8 serrations on each side, outside densely villous and white, inside glabrous

at base and densely villous at apex, the fifth dorsal lobe smaller, broadly oblong to lanceolate, 0.8 x 0.2 cm.

J.Bot. Res. Inst. Texas 5(1): 81 -8

Journal of the Botanical Research Institute of Texas 5(1)

1 cm

Corolla mostly yellow with red apex, uniformly tubular, 2. 0-3.0 x 0.5 cm, corolla posture erect in calyx,

apical region red, lobes triangular, ca., 1.5 x 2 mm, nearly actinomorphic, outside densely villous, inside

glabrous. Androecium of 4 stamens, didynamous, included; filaments 1. 5-2.0 cm long, basally connate

for 3.5 mm, adnate to the base of the corolla tube, glabrous; anthers broader than long, ca. 1.5 mm long,

ca. 2.0 mm wide, dehiscing by longitudinal slits; staminode not observed. Nectary a single dorsal truncate

Fig. 2. Columned lucifer. A. Stephen Maciejewski holding shoot of entire plant. B. Immature flower. C. Mature flower, (photo A. by Julie Mavity-Hudson.

Photos B & C by J.L. Clark; from the live plant from which the holotype was collected, J.L. Clark & Gesneriad Research Expedition Participants 1 1 100).

1 (Q, QAP, QCA, and QCNE) for a

i R. Anderson for t

Journal of the Botanical Research Institute of Texas 5(1)

BOOK REVIEW

Ralph W. Tiner, with Abigail Rorer (Drawings). 2009. Field Guide to Tidal Wetland Plants of the

Shores, Marshes, Swamps, and Coastal Ponds. (ISBN 978-1-55849-667-5, pbk.; 978-1-55849-

666-8, hbk.). University of Massachusetts Press, East Experiment Station, 671 North Pleasant Street,

Amherst, Massachusetts 01003, U.S.A. (Orders: http://hfs.jhu.edu; hfscustserv@mail.press.jhu.edu;

toll-free 800-537-5487). $29.95 (pbk.), $98.00 (hbk.), 416 pp„ 570 illustrations, 11 maps, 7" x 10".

J. Bot. Res. Inst. Texas 5(1): 86. 2011

COLUMNEA PYGMAEA (GESNERIACEAE),

A NEW SPECIES FROM NORTHWESTERN ECUADOR

John L. Clark

James F. Smith

Department of Biological Sciences, Box 870345

The University of Alabama

Tuscaloosa, Alabama 35487, U.S.A.

Department of Biology

Boise State University

1910 University Drive

Boise, Idaho 83725, U.S.A.

RESUMEN

INTRODUCTION

The genus Columnea is primarily epiphytic and belongs to the New World subfamily Gesnerioideae. The

genus ranges from Mexico south to Bolivia and is one of the largest genera of the family in the New World

tropics with over 200 species. Columnea is distinguished from other closely related genera by the presence

of an indehiscent berry instead of a fleshy bivalved capsule. Columnea is the most diverse genus in the sub-

family Gesnerioideae with over 200 species (Skog & Boggan 2006; Weber 2004; Burtt & Wiehler 1995).

The traditional sectional classification of Columnea does not represent monophyletic lineages (Smith 1994,

Smith & Sytsma 1994; Clark et al. 2006) and a revised classification system based on molecular sequence

data is currently in preparation by the authors. The species described here belongs to a well-supported clade

(Smith et al., in review) that does not yet have a formal designation. Many of the specimens that resemble

Columnea pygmaea J.L. Clark &J.F. Smith are incorrectly determined in herbaria. Plates of digital images

and a key are provided to help differentiate Columnea pygmaea from the following morphologically similar

species: Columnea parvijlora C.V. Morton, C. herthae Mansf., C. minutijlora L.P. Kvist & L.E. Skog, and C.

lehmannii Mansf.

TAXONOMIC TREgKMf(

Obligate epiphytic herb, roots fibrous, stems erect (not scandent), 7-20(-40) cm tall, sparsely villous

throughout, internodes 1-4 cm long. Leaves opposite, subequal to strongly unequal in a pair; larger leaf with

petioles 0.5-1. 5 cm long, villous; blade elliptic to narrowly ovate, 2. 0-7.0 x 1.5-3. 5 cm, base asymmetrical,

apex acute to obtuse, margin shallowly serrate to crenate; adaxially bright green, sparsely villous; abaxially

J. Bot. Res. Inst. Texas 5(1): 87 -9

Journal of the Botanical Research Institute of Texas 5(1)

Fig. 1. Columnea pygmaea. A & B. Mature flower. C. Mature berry. D. Berry removed to show calyx. E. Habit (Photos by J.L. Clark from the field collection

of the holotype, J.L. Clark & The Gesneriad Research Expedition Participants 1 1 180).

greenish- white, sparsely to densely villous (especially on veins), 3-4 pairs of lateral veins; smaller leaf

similar to larger leaf, but greatly reduced, 0.5-1. 5 x 0.2-0.5 cm. Inflorescence reduced, epedunculate and

appearing fasciculate with 1-3 flowers per node, bracts persistent, 0.2 x 0.1 cm, bright red; pedicel 0.3-0. 5

cm long, villous to densely villous near base of calyx and red. Calyx bright red with reflexed red lobes at

anthesis; lobes 5, nearly free; 4 lobes equal in size, ovate to broadly ovate, ca., 0.5 x 0.5 cm, margin with

3-5 serrations on each side, sparsely villous near apex and densely villous at base, inside sparsely villous,

the bfth dorsal lobe smaller, lanceolate, 1.0 x 0.2 cm. Corolla pale yellow, tubular, * 0.2-0. 3 cm,

corolla posture erect in calyx, lobes erect, white to pale-yellow, limb nearly actinomorphic, outside sparsely

villous near base to densely villous near apex, inside glabrous. Androecium of 4 stamens, didynamous,

included; staminode not observed. Nectary a single dorsal truncate gland. Gynoecium immature, ovary

superior, ovoid. Fruit an indehiscent white berry, appearing dorsally flattened, 0.5 cm in diameter.

Distribution and habitat. — Columnea pygmaea is known from two collections from wet forests along the

western slopes of the Ecuadorian Andes. The brst collection of Columnea pygmaea was by Hans Wiehler and

Calaway Dodson (H. Wiehler & C.H. Dodson 7113 ) from the province of Pichincha (Rio Baba, Hwy Santo

Domingo-Quevedo). Additional collections since the early 1970s are from cultivated material that was

distributed from Cornell University’s L.H. Bailey Hortorium. The cultivated material apparently originated

from Wiehler and Dodson’s 1971 held collection.

The tropical wet and moist coastal forests of western Ecuador below 1000 meters once encompassed

a band from Colombia to Peru that reached a maximum width near the northern limit and was very nar-

row and broken to the south. Dodson and Gentry (1991) estimated that while this forest type once covered

15% of western Ecuador, currently less than 0.8 % of the coastal wet and moist forests remain. It is not

surprising that Columnea pygmaea has not been recollected since 1971 because of the near eradication of

this forest type from western Ecuador. The endemism of the flowering plant family Gesneriaceae in western

Ecuador is estimated at 20%, which is significantly higher than the 12% endemicity reported for western

lowland Ecuador (defined as below 1000 m) as reported by Jorgenson and Leon-Yanez (1999) and Valencia

et al. (2000). The endemicity of the Gesneriaceae in western Ecuador is especially important for evaluat-

ing conservation priorities because at least 20 of the 23+ species were evaluated and considered extinct or

endangered by Kvist et al. (2004) according to the IUCN Red List categories (IUCN 2001).

Fieldwork in Ecuador with participants of the 2009 Gesneriaceae Research Expedition resulted in the

discovery of a second population and extension of the species known range into the northern Ecuadorian

province of Esmeraldas. The type locality of Columnea pygmaea is from a remnant patch of primary forest

along the San Lorenzo-Ibarra highway between the towns of Alto Tambo and Durango at 460 m. It was

noticed by the brst author during extensive fieldwork between 2003 and 2009 that most of the forest along

the San Lorenzo-Ibarra highway had been converted to African Palm plantations. The original habitat from

this area is transitional between lowland and montane wet forest. These forests have been classified as bosque

siempreverde piemontano (Sierra 1999); selva ombrofila noroccidental del pie de cordillera (Acosta Solis 1968);

and bosque lluvioso montano bajo (Harling 1979).

Discussion of circumscription. — Columnea pygmaea is unique among the species of Columnea by the

short corollas that are less than 1 cm in length (Fig. IB). Columnea parviflora consistently has corollas that

are less than 1 cm in length (Fig. 2A), but the habit and calyx margin help differentiate this species from

C. pygmaea. The pendent and climbing habit of C. parviflora (Fig. 2E) helps differentiate it from the erect

stems and non-climbing habit of C. pygmaea (Fig. IE). The calyx lobes of Columnea parviflora are fimbriate

(Fig. 2C-D) in contrast to the shallow serrations in C. pygmaea (Fig. ID). Both Columnea pygmaea and C.

herthae have uniformly pale-yellow tubular corollas with four equal calyx lobes and one (dorsal) lobe that

is reduced and lanceolate (Fig. ID & Fig. 3C). In contrast to four equal calyx lobes with one reduced calyx

lobe, Columnea minutiflora (Fig. 4) and C. parviflora (Fig. 2) have five calyx lobes that are nearly equal in size

and shape.

The above three species are common in northwestern Ecuador and southern Colombia. In contrast,

Journal of the Botanical Research Institute of Texas 5(1)

Fig. 2. Columnea parviflora. A. Mature flower. B. Mature berry. C. Mature flower. D. Berry removed to show calyx. E. Habit (Photos by J.L. Clark, A & B

J.L. Clark, R. Fleiss & I. Salinas 8817, C. J.L. Clark & The University of Alabama in Ecuador Program Participants 9644, D & E. J.L. Clark 8t The University of

Alabama in Ecuador Program Participants 10832).

Fig. 3. Columnea herthae. A & B. Mature flower. C & D. Mature berry. E. Habit (Photos by J.L. Clark, A, B & E, J.L. Clark, G. Zapata & G. Toasa 71 1 3, C & D.

J.L. Clark & The Gesneriad Research Expedition Participants 11193).

Journal of the Botanical Research Institute of Texas 5(1)

Fig. 4. Columned minutiflora. A. Dorsal view of mature berry. B. Mature flower. C. Lateral view of mature berry. D. Inflorescence. E. Habit. F. Inflorescence

(Photos by J.L. Clark, A, J.L. Clark 7092, B. & D. J.L. Clark & The Gesneriad Research Expedition Participants 1 1091, C. & F. J.L. Clark & The Gesneriad Research

Expedition Participants 1 1 132, E. J.L. Clark 7200).

Fig. 5. Columnea lehmannii. A. & B. Habit with erect inflorescence. C. Lectotype of C. lehmannii, F. Lehmann 6063 (K). D. Isolectotype of C. lehmannii, F.

Lehmann 6063 (K). (Photos A & B. by J. Betancur of J. Betancuretal. 12359 from the Parque Nacional Tatama, C. & D. Specimens from the Royal Botanic

Gardens, Kew).

Journal of the Botanical Research Institute of Texas 5(1)

Columnea lehmannii is common in Colombia, but is known from Ecuador by a single collection ( Kvist et al.

48983, AAU), which has not been seen by the authors of this study. Confusion between Columnea lehmannii

and other species discussed here is evident from many misidentihed collections at MO, NY, QCA, QCNE,

SEL, and US. Many herbarium collections previously identified as Columnea lehmannii clearly belong to

Columnea lehmannii is differentiated by the presence of larger corollas (> 2 cm long; Fig. 5 A & B). The pres-

ence of opposite leaves that are nearly equal in size is rare in other species and relatively common in C.

lehmannii (Fig. 5), but occasionally the opposite leaves are strongly anisophyllous (Fig.|^J® . (

Etymology, — The new species is named in reference to the small stature of the plants. The flowers of

Columnea pygmaea (Fig. IB) are similar in shape to Columnea herthae (Fig. 3B), but differ by their significantly

smaller size.

Conservation and IUCN Red List category. — Columnea pygmaea has not been found in any formally pro-

tected area in Ecuador. According to the IUCN Red List criteria (IUCN 2001) for limited geographic range

(B2a, less than 10 km 2 and known to exist at only a single location) and considering the uncertain future

of habitat conservation along the San Lorenzo-Ibarra highway, Columnea pygmaea should be listed in the

category CR (Critically Endangered).

Paratypes. — ECUADOR. Pichincha: Rio Baba (=Rio Quevedo, Rio Palenque), km 7 on road from Santa Domingo to Quevedo, 21 Jul

1971, H. Wiehler & C.H. Dodson 7113 (=Wiehler live accession W-1573) (US); cultivated collection from L.H. Bailey Hortorium of H.

Wiehler & C.H. Dodson 7113, 7 Jan 197% M.H. Stone 1305 (SRP, US).

KEY TO THE COMMON SPECIES OF COLUMNEA

WITH CRENATE-MARGINED OBLONG LEAVES

2. Reduced inflorescence of 1 -3 flowers with well-developed pedicels

3. Corolla less than 1 cm long, stems less than 40 cm tall

3. Corolla 1 .5 - 2.0 cm long, stems more than 40 cm tall

. Corolla 2.0 to 4 cm tong^oorolla not uniformly colored, tube with broad red band on ventral surface

ACKNOWLEDGMENTS

Support for this study was provided by the National Science Foundation (DEB-0841958 & DEB-0949169).

We thank the following herbaria for access to their collections: Smithsonian Institution’s National Museum

of Natural History - Department of Botany (US), Marie Selby Botanical Gardens (SEL), Missouri Botanical

Garden (MO), Herbario Nacional del Ecuador (QCNE), and the Pontihcia Universidad Catolica del Ecuador

(QCA). Christian Feuillet provided the Latin diagnosis and Laura Clavijo provided the Spanish translation of

the abstract. We are grateful to Jeremy Keene and Laurence E. Skog for helpful comments on the manuscript.

Julio Betancur (COL) provided held images of Columnea lehmannii. We thank the Royal Botanic Gardens,

Kew (K) for permission to use photographs of the lectotype and isolectotype of Columnea lehmanii. The first

author would like to thank the participants from the 2009 Gesneriad Research Expedition to Ecuador for

their help in discovering the extant population of Columnea pygmaea.

REFERENCES

Acosta Sous, M. 1 968. Divisiones fitogeograficas y formaciones geobotanicas del Ecuador. Publicaciones Cientfficas

de la Casa de la Cultura Ecuatoriana, Quito.

i i id r. 1 995. Classification of the family Gesneriaceae. Gesneriana lilMM I

Clark, J.L., P.S. Herendeen, LJE. SkOg, and E.A. Zimmer. 2006. Phylogenetic relationships and generic boundaries in

the Episcieae (Gesneriaceae) inferred from nuclear, chlorpplaSt, and morphological data.Taxon 55:31 3-336.

,7 halftones, 6" x 9".

BEGONIA BERNICEI (BEGONIACEAE), A NEW SPECIES FOR THE

LOWLAND HUMID FORESTS OF THE VENEZUELAN GUAYANA

Gerardo A. Aymard C. Gustavo A. Romero-Gonzalez

UNELLEZ-Guanare, Programo de Ciencios

del Agroy del Mar, Herbario Universitario (PORT)

Mesa de Cavacas. Estado Portuguesa

VENEZUELA 3350

gaymard@cantv.net

Orchid Herbarium of Oakes Ames

Harvard University Herbaria

22 Divinity Avenue

Cambridge, Massachusetts 02138, U.S.A.

romero@oeb.harvard.edu

ABSTRACT

. The new

RESUMEN

, \ [ '

The genus Begonia was described by Linnaeus (1753) and later circumscribed by Irmscher (1925), Doorenbos

et al. (1998), Forrest and Hollingsworth (2003), and Tebbitt (2005). Begonia is remarkable because it includes

all the species of the family Begoniaceae, except for one species referred to Hillebrandia sandwicensis Oliver,

which is endemic to Hawaii (Clement et al. 2004). Comprising more than 1500 species (Govaerts 2009),

Begonia is the sixth largest genus of flowering plants, and it is distributed throughout the tropics and sub-

tropics with the exception of Australia (Frodin 2004; Tebbitt 2005).

Begonia, undoubtedly due to its large size, has been divided into numerous sections. The most recent

sectional account is that of Doorenbos et al. (1998), who recognized sixty three sections. Three additional

new sections have been described recently (Shui et al. 2002; Forrest & Hollingsworth 2003; de Wilde &

Plana 2003). The sections of Begonia are recognized based on a combination of morphological features such

as leaf venation, perianth segment number, features of the pistillate and staminate flowers, and the shape

of the fruits (Nguyen & Tebbitt 2006).

Family treatments were prepared for the Flora of Venezuela (Smith & Wasshausen 1989), and the Flora

of the Venezuelan Guayana (Steyermark 1997), who recognized 47 and 11 species, respectively. In addition,

Dorr (1999) reevaluated several taxa from the Venezuela Andes, and recently the family was revised for

the “Catdlogo ilustrado y anotado de las plantas de los Llanos de Venezuela” (Duno et al. 2007) and the “Nuevo

Cat&logo de la Flora vascular de Venezuela” (Hokche et al. 2008). This later contribution increases to forty nine

the number of Begonia species known from Venezuela, 23 of which are considered endemic.

Herbarium material of the new species described here was first pointed out to the auhors in 1991 by

L.B. Smith (1904-1997), a noted bromeliad and Begonia taxonomist. Subsequently, the authors found two

additional collections, which together provide enough material to describe it.

TAXONOMIC DESCRIPTION

J. Bot. Res. Inst. Texas 5(1): 97-1

Journal of the Botanical Research Institute of Texas 5(1)

by Angelina Licata.

Suffrutescent herb, stems 2-5 mm in diameter, green, grooved, glabrous, internodes 3-8 cm long. Stipules

deciduous, ca. 15 x 6 mm, green, ovate to triangulate, glabrous and reticulate on both surfaces, margins entire,

apex with an acumen ca. 2 mm long. Leaf lamina 7-14 x 4-8 cm, asymmetric, delicately membranaceous,

broadly ovate, glabrous on both surfaces, apex acuminate, base obliquely cordate, margins ciliate-denticulate,

venation palmate, main veins 8-9, these red on the lower surface, tertiary veins reticulate, inconspicuous;

petiole 2-4 cm long, green, glabrous, canaliculate at the base joining lamina at an ca. 65° angle. Inflorescence

axillary and terminal, erect, 6-8 cm long, 4-6-flowered, cymose, once dichotomous branched, glabrous,

penduncles pink to light green; bracts caducous, ovate, 2-3 x ca. 2 mm, glabrous on both surfaces, margins

entire, apex acuminate. Staminate flowers white; pedicels 3-5 mm long, glabrous, tepals 2, white, orbicular to

reniform, 3x3 mm, glabrous and reticulate on both surfaces, androphore absent; stamens 25-30, arranged in

an actinomorphic cluster, filaments free, 1-1.5 mm long, glabrous, anthers oblong, ca. 1.4 mm long, glabrous,

connective projecting; pistillate flowers white; pedicel ca. 2 mm long, glabrous; bracteoles caducous, ca. 3 x

ca. 4 mm, orbicular, glabrous on both surfaces, margins ciliate-serrulate, tepals 4, glabrous and reticulate

on both surfaces, apex obtuse, margins entire, outer pair ovate, 1-2 x 1-1.5 mm; inner pair oblanceolate,

1-3 x ca. 1 mm; tepals persistent on young fruit, covering the stigmas, outer pair ovate, 3-3.5 xl-1.5 mm,

inner pair oblanceolate, 2-3 x ca. 1 mm, delicately membranaceous, glabrous, with a conspicuous network

of veins on both surfaces; styles three, free, 1-1.5 mm long, bifid, stigmatic area arranged in a spiral band;

ovary green, ca. 6 mm long, glabrous and reticulate, spherical with three rib-like wings, placentation axile,

one placentae per locule. Fruits pendulous, green, ca.10 x ca. 8 mm, with three wings, slightly unequal, the

longer wing ca. 4 mm wide, shorter wings ca. 3 mm wide, glabrous, with a conspicuous network of veins.

Distribution and ecology. — Begonia bernicei is known from three gatherings collected in the understory

of lowland moist forests in the lower Erebato river, in Bolivar state, Venezuela, all three coincidentally at an

altitude of 360 m.

Because of its cymose inflorescence, ovary with three locules, female flowers with 4 perianth segments,

male flowers with 2 perianth segments and one placenta per locule, Begonia bernicei belongs to the neotropi-

cal section Doratometra (Klotzsch) A. DC. (Doorenbos et al.1998). The annual plants asymmetric leaves,

the few-flowered, one time dichotomous inflorescence, and the slightly unequal fruit wings place this new

species close to B. semiovata Liebm. However, B. bernicei is distinguished by its leaves 7-14 x 4-8 cm (vs.

4.5-7 x 1-2 cm in B. semiovata ), petiole 2-4 cm long (vs. 0.5-2 cm), inflorescence axillary and terminal,

6-8 cm long (vs. inflorescence axillary, 2-3 cm), tepals of the staminate flowers orbicular to reniform, 3x3

mm (vs. ovate to suborbicular, ca 2 x 2 mm), and tepals of the pistillate flowers 4 (vs. tepals of the pistillate

flowers 5).

Etymology. — The epithet honors Bernice G. Schubert (1913-2000), noted authority on Dioscoreaceae,

the genus Desmodium Desv. (Leguminosae-Papilionoideae), and particularly on Begoniaceae.

Paratypes. VENEZUELA. Estado Bolivar: 1

1989 (fl, fr), E. Sanoja2397, 2603 (PORT):

ACKNOWLEDGMENTS

We thank Lisa M. Campbell (NY) for her useful suggestions, to Mark Tebbitt (California University of

Pennsylvania) and Mark Hughes who provided helpful comments, corrections and additional references for

the manuscript. We are also grateful to Bruno Manara (VEN) for revising the Latin description and preparing

the illustration, Angelina Licata for additions to the illustration, and to the New York Botanical Botanical

Garden (NY) for making their facilities available for our research.

n relict. Amer.J.Bot. 91:^05-91 7.

M ATELEA PAKARAIMENSIS (APOCYNACEAE: ASCLEPIADOIDEAE),

A NEW SPECIES IN THE M ATELEA STENOPETALA COMPLEX FROM GUYANA

Alexander Krings

Herbarium, Department of Plant Biology

North Carolina State University

Raleigh, North Carolina 27695-7612, U.S.A.

Alexander_Krings@ncsu.edu

ABSTRACT

M atelea Aubl. (Apocynaceae: Asclepiadoideae: Gonolobinae) is a New World genus estimated to include

20-330 species, depending on its circumscription (Morillo 1997; Stevens 2001). More than 150 species have

been named and at least eighty-four combinations made since the one hundred some combinations made

by Woodson (1941). Because of Woodson’s (1941) mass synonymization and the subsequent description of

numerous new species, Matelea s.l. now represents a morphologically heterogeneous group (Morillo 1997;

Stevens 2001; Krings & Saville 2007; Krings et al. 2008). Unfortunately, despite recent advances in our un-

derstanding of the Gonolobinae (Rapini et al. 2003; Liede-Schumann et al. 2005; Rapini et al. 2006; Krings et

al. 2008), the circumscription of Matelea s.s. and numerous other lineages remains unresolved. Consequently,

in the absence of improved generic level understanding of the Gonolobinae, it remains convenient to de-

scribe new species belonging to the subtribe into one of the three genera recognized by Woodson (1941):

Fisheria DC. (dorsal anther appendages [Cd sensu Kunze 1995] vesicular), Gonolobus Michx. (dorsal anther

appendages laminar), and Matelea s.l. (dorsal anther appendages absent). Ongoing study of the Gonolobinae

in northern South America, as a component of the Smithsonian’s Biological Diversity of the Guiana Shield

Program, has now resulted in the discovery of an additional species of Matelea from northern South America.

The new species belongs to the Matelea stenopetala Sandwith complex, to which five species have been

previously referred: M. grenandii Morillo, M. hildegardiana Morillo, M. quindecimlobata Farinaccio & W.D§g

Stevens, M. stenopetala, and M. sucrensis Morillo (Morillo 1980, 1988, 1991; Farinaccio & Stevens 2009).

Among these, adaxially pubescent corolla lobes are found in the (1) new species described below (corolla

purplish), (2) M. hildegardiana (corolla green or greenish-white), and, very rarely, in (3) M. stenopetala (see

Mori 24547, US; corolla greenish-white to greenish-yellow). All other species in the complex — and usually

M. stenopetala as well — exhibit adaxially glabrous corolla lobes. The recently described M. quindecimlobata

is particularly interesting among these in bearing staminal corona lobes (Cs sensu Liede & Kunze 1993)

trapezoidal in outline with the outer margins deeply irregularly 3-6-lobed-fimbriate and the lateral lob-

ules broader than the central lobule (Farinaccio & Stevens 2009). Matelea grenandii, M. stenopetala, and M.

sucrensis, in contrast, lack this character. They can be distinguished by the shape and size of the calyx and

corolla lobes as noted in the key below.

J. Bot. Res. Inst. Texas 5(1): 101-1

Krings, Matelea pakaraimensis, a new species from Guyana

103

Journal of the Botanical Research Institute of Texas 5(1)

distinctly more distant from the gynostegial center than the base, base somewhat swollen, lobed, and slightly

fimbriate, ca. 1.2 mm tall, ligule an apical ridge, not free, Ci unlobed, not ligulate. Style-head ca. 2.4 mm

caudicles present, pollinia oblong, ca. 0.45 x 0.2 mm. Follicles unknown. Seeds unknown.

Notes. — M ateleapakaraimensis is currently known only from the type, which was collected in tall mesic

ACKNOWLEDGMENTS

of their collections: A, B, BG, BH, BKL, BM, BOLO, BR, BREM, BSC, BUF, C, CGE, COLO, CR, DUKE, E,

F, FI, FLAS, FR, FIG, G, GH, GOET, H, HAC, HAJB, HBG, IA, IJ, ISC, JBSD, JE, K, L, LD, M, MICH, MIN,

MO, MSU, NEU, NSW, NY, O, OXF, P, PH, RSA, U, UC, UPS, US, USF, TRIN, TUR, VEN, WILLI, WU,

Z. I thank David Goyder (K) and Bruce Hansen (USF) for reviewing the manuscript, and Mark Garland for

translating the diagnosis into Latin.

REFERENCES

Endress, M.E. and P.V. Bruyns. 2000. A revised classification of the Apocynaceae, s.l. Bot. Rev. 66:1-56.

Farinaccio, M.A. and W.D. Stevens. 2009. Matelea quindecimlobata (Apocynaceae, Asclepiadoideae), a new species

from Amazonas, Brazil. Novon 19:156-158.

Krings, A. and A.C. Saville. 2007. Two new species and three lectotypifications in the Ibatia-Matelea complex

(Apocynaceae: Asclepiadoideae) from northern South America. Syst. Bot. 32:862-871 .

Krings, A., D.T. Thomas, and Q.-Y. (J.) Xiang. 2008. On the generic circumscription of Gonolobus (Apocynaceae,

Asclepiadoideae): Evidence f^^^^sfifes and morphology. Syst. Bot. 33:403-415.

Kunze, H. 1995. Floral morphology of some Gonolobeae (Asclepiadaceae). Bot. Jahrb. Syst. 1 17:21 1-238.

Liede, S. and H. Kunze. 1993. A descriptive system for corona analysis in Asclepiadaceae and Periplocaceae. PI.

Syst. Evol. 185:275-284.

Liede-Schumann, S., A. Rapini, DJ. Goyder, and M.W. Chase.. 2005. Phylogenetics of the New World subtribes oft

Asclepiadeae (Apocynaceae-Asclepiadoideae): Metastelmatinae, Oxypetalinae, and Gonolobinae. Syst. Bot.

30:184-195.

LivshultzJ. 2003. Systematics of Dischidia (Apocynaceae, Asclepiadoideae). Ph.D. dissertation. Cornell University,

Morillo, G. 1980. Nuevas especies suramericanas en el genero Matelea Aublet (Asclepiadaceae). Mem. Soc. Ci.

Nat. "La Salle" 40:73-82.

Morillo, G. 1 988. Especies, combinaciones y sinonimos nuevos en Fischeria DC., Gonolobus Mich. Y Matelea Aubl.

Comentarios sobre una especie interesante de Gonolobus M|||§fJ|Lis afines. Ernstia 50: ; : )/ ■-.

Morillo, G. 1 991 . Once Asclepiadaceae sudamericanas nuevas para la ciencia. Ernstia 1 1 09-1 20.

Morillo, G. 1 997. Asclepiadaceae. In: P.E. Berry, B.K. Holst, and K. Yatskievych, eds. Flora of the Venezuelan Guyana,

Vol. 3. Missouri Botanical Garden, St. Louis. Pp. 129—1 77.

Rapini, A., M.W. Chase, andT.U.P. Konno. 2006. Phylogenetics of South American Asclepiadoideae (Apocynaceae).

Taxon 55:119-124.

Rapini, A., M.W. Chase, DJ. Goyder, and J. Griffiths. 2003. Asclepiadeae classification: evaluating the phylogenetic

relationships of New World Asclepiadoideae (Apocynaceae). Taxon 52:33-50.

Stevens, W.D. 2001 . Asclepiadaceae. In: W.D. Stevens, C. Ulloa U., A. Pool, and O.M. Montiel, eds. Flora de Nicaragua.

Monogr. Syst. Bot. Missouri Bot. Card. 85:234-270.

Woodson, R.E., Jr. 1941. The North American Asclepiadaceae. Ann. Missouri Bot. Gard. 28:193-244.

A NEW COMBINATION BASED ON

M YRCIANTHES IRREGULARIS (MYRTACEAE)— A NEW GENUS FOR ECUADOR

) (fl, fr), i

LECTOTYPIFICATION OF THOMAS NUTTAFF’S NAMES APPFIED

TO NORTH AMERICAN POLYGALA (POFYGAFACEAE)

Alina Freire-Fierro

Academy of Natural Sciences

Botany Department, PH Herbarium

1 900 Benjamin Franklin Parkway

Philadelphia, Pennsylvania 19103, U.S,A.

freirefierro@ansp.org (author for correspondence)

INTRODUCTION

Since the founding of the Academy of Natural Sciences in Philadelphia in 1812, botanical collections of

many North American species have been deposited at this institution (Freire-Fierro 2008). However, the

type status of many has not been verified. Such is the case for many taxa described by Thomas Nuttall

(1818, 1834), who used the Academy’s herbarium (PH) as his home institution during the early stages of his

career in the United States (Pennell 1936). In publications and on specimen labels, Nuttall generally added

an asterisk before his new names for the plant species he described. Also, many of the PH Nuttall labels

have the abbreviation “Nutt.” in parenthesis. According to Stuckey (1966), this abbreviation had been added

later by Charles Pickering (1805-1878) (Fig. 1). Following standard practice, we could have chosen Nuttall

specimens at PH as the holotypes. We did not to do that, however, because Nuttall generally studied more

than one specimen per species, and he was not consistent in the use of the asterisk (Stuckey 1966). In several

instances, he did not provide much detail about the localities. Additionally, material used by him for the

description of new taxa is also deposited at several herbaria (e.g., BM, CGE, LIV, and MANCH). These facts,

combined with the scarcity of current published literature on North American Polygalaceae (Wendt 1979)

and the familiarity of AFF with Polygalaceae, motivated the initiation of this project.

This paper presents results obtained from a study on the types of North American Polygala (Polygalaceae)

described by Thomas Nuttall (1818, 1834) and deposited at the Academy of Natural Sciences, Philadelphia

(PH). This genus has 1,554 species described, and less than 40% of these have been taxonomically/no-

menclaturally assessed (The Plant List 2011). So, the current total estimate of 300-350 species (Eriksen &

Persson 2006) is very likely an underestimate (R. Abbott, pers. comm., 2011). Although there are various

floristic and taxonomic revisions for the genus that include taxa from the United States (Chodat 1893; Blake

1916, 1924; Smith & Ward 1976; Wendt 1979; Gleason & Cronquist 1991 and Bernardi 2000), these do

J. Bot. Res. Inst. Texas 5(1): 109-1

110

Journal of the Botanical Research Institute of Texas 5(1)

laceae for Flora of >

or the region (R. Ab

i for 53 of New Wor

, Gen. N. Amer. PL 2:87. 1818. (Fig. 2). T

fP. alba, it is very likely that t

i of his Fl. Amer. Sept. (1814: 750), c

,Kp.p. digital image!, MANCH

112

Journal of the Botanical Research Institute of Texas 5(1)

g. 2. Lectotype of Polygala alba Nutt. (PH 1049944 p.p. (rightmost specimen), barcode number 471 58)

114

d right (Fig. 4). With the first (lei

id handwriting/Polygala attenu;

> of its height (2 to 3 feet tall), t

■; This specimen is r

likely that “N.” stands for Nuttall. If so, tl

lia 7(1):86. 1834. (Fig. 6). Type: U

. The first (PH 1049947) has a N

115

Freire-Fierro and Landale, L<

. Lectotype of Polygala attenuata Nutt. (PH 1049951 p.p. (leftmost specimen), barcode number 20518)

116

Journal of the Botanical Research Institute of Texas 5(1)

Lectotype of Polygala balduinii Nutt. (PH 1049950 p.p. (leftmost specimen), t

e number 2051 9)

117

Freire-Fierro and Landale, L<

. Lectotype of Polygala boykinii Uutt. (PH 1049947, barcode number 47170)

at BM, K, and PH, ii

120

Journal of the Botanical Research Institute of Texas 5(1)

Fig. 8. Lectotype of Polygala pubescens Nutt. (PH 1072615 p.p. (leftmost specimen), barcode number 20567)

Freire-Fierro and Landale, I

121

9. Lectotype of Polygala purpurea Nutt. (K p.p. (leftmost specimen), barcode number 555052)

122

Journal of the Botanical Research Institute of Texas 5(1)

Polygalafastigiata, though described by Nuttall as closely related to P. cruciata L., was later synonymized

under P. mariana Mill. (Blake 1924).

Polygala pubescens Nutt., Gen. N. Amer. Pi.

(1815). (Fig. 8). A Georgia: ‘7

For the locality, Nuttall mentions in the protologue: “HAB. Around Savannah in Georgia, &c.” So, it is likely

that he collected it himself in Savannah (GA) while visiting Baldwin in the fall of 1815 (Ewan 1971). While

the label allows for little identification, it does contain Nuttall’s asterisk before the word “pubescens.” Blake

(1924) considered Polygala pubescens Nutt, a synonym of P. grandiflora Walt.

Polygala purpurea Nutt., Gen. N. Amer. PL 2:88. 1818, nom. illeg., later homonym non P purpurea Ait.

f. Hortus Kew. (WT. Aiton), ed. 2, 4:244. 1812. (Fig. 9). Type: U S a. New Jersey: “Common throughout North

digital image!; PH 107^j^^p).!).

When describing Polygala purpurea, Nuttall acknowledged that his specimen along with those described by

Michaux and Pursh under P. sanguinea L. corresponded to a species distinct from the latter. From his descrip-

tion, it is evident that he also studied the Pursh material deposited at K, and also the one at PH (mounted

on the same sheet with specimens identified as Polygala sanguinea ). Although the PH specimen containing

Nuttall’s label ordinarily would have been designated as the lectotype, we chose the sheet at K because the

sheet at PH has four labels and eight specimens that have been intermingled sometime in the past during

the mounting process. Regarding the type locality, despite the fact that the protologue does not mention

a particular location, the label at PH reads “Camden” (New Jersey) and the label at K reads “New Jersey.”

According to Art. 64 from the Code (McNeill et al. 2006), the name Polygala purpurea Nutt, is considered

illegitimate, because of the publication of an earlier homonym (P. purpurea Ait.f.) published in 1812 and

based on another North American collection that had been introduced to Europe in 1791 (Aiton 1812). No

new name is assigned to P. purpurea Nutt, because it was considered a synonym of P. viridescens L. (Blake

1924). Currently, the name Polygala viridescens L. is considered a synonym of Polygala sanguinea L. (Fernald

1950).

ACKNOWLEDGMENTS

The following herbaria and staff members provided information and/or E-loans for this study: N. Kilian (B),

R. Huxley, J. Hunnex, and J. Gregson (BM), I. Bonnie (CM), T. Hogan (COLO), D. Trock (DS), A. Smith (E), L.

Kawasaki (F), R. Clark (K), C. Beans, K. Ghandi, J. Macklin and E. Wood (A, GH), G. Thijsse (L), R. Rabeler

(MICH), A.F. Cholewa (MIN), Jim Solomon (MO), D. Bailey (NMC), B. Thiers and T. Zanoni (NY), C. Loup

(P), S. Boyd (POM), J. Stepanek (PRC), R.H. Hartman (RM), T. Wendt (TEX), A.S. Doran (UC), R. Kennedy

(UMO), R. Russell (US), D. Barrington and K. Mauz (VT), and P. Sweeney (YU). Suggestions from R. Abbott,

B. Eriksen Luteyh, N. Turland (MO), and T. Wendt (TEX) greatly improved the manuscript.

Thanks also to staff and volunteers of the Botany Department (PH), in particular to A.E. Schuyler, Jon Gelhaus,

Rosemary Malh, and Eileen Mathias. Alinia Freire-Fierro is very grateful to A.E. Schuyler for his invaluable

comments regarding the nomenclature and history of the Academy’s botanical collections. Alicia Landale’s

work was supported by a REU Site Program Research Experiences for Undergraduates (REU) grant #0353930

from The National Science Foundation awarded to the Academy of Natural Sciences of Philadelphia, for

their Collections-Based Undergraduate Research at the Academy of Natural Sciences program. Images of

the types at PH were taken under the Global Plants Initiative (GPI) financed by the G.P. Mellon Foundation.

REFERENCES

Aiton, W.T. 1812. Hortus Kewensis; or, a catalogue of the plants cultivated in the Royal Botanic Garden at Kew.

London, 2nd ed.Vol.4:1-522.

124

j Co. Pp. 173-174.

of botany: being outlines of the s

NOTES ON THE DISINTEGRATION OF POLYGALA (POLYGALACEAE),

WITH FOUR NEW GENERA FOR THE FLORA OF NORTH AMERICA

J. Richard Abbott

University of Florida Herbarium (FLAS)

Florida Museum of Natural History

University of Florida

P.O. Box 1 17800

Gainesville, Florida 3261 1-7800, USA.

Research Associate

Missouri Botanical Garden

RO. Box 299

Saint Louis, Missouri 63166-0299, USA.

rabbott@ufl.edu

Raf., Hebecarpa (Chodat) J.R. Abbott, sts

ABSTRACT

RESUMO

Asemeia Raf., Hebecarpa (Chodat) J.R. Abbott, stat nov., Polygaloides Haller e Rhinotropis (S.F. Blake) J.R. Abbott, stat. nov. como

INTRODUCTION

Polygalaceae Hoffmanns, and Link of North America north of Mexico [hereafter referred to as North America]

consist of one species of Monnina Ruiz and Pav. s.l., ca. 52 native species of traditional Polygala L., and four

reportedly naturalized species of Polygala (only one of which, P. vulgaris L., appears to be truly naturalized).

A morphologically diverse assemblage of herbs and shrubs, they are most speciose in the southern United

States and occur in a wide range of habitats, from wetlands to woodlands or deserts, mirroring the global

diversity and range of Polygala, which, as traditionally circumscribed, includes everything from annual herbs

to lianas and trees, occurring from sea-level to high montane regions. Eriksen and Persson (2007) provided

a comprehensive description of the family, as well as a discussion in which they pointed out that there are

discrepancies between published phylogenetic analyses and that there is a lack of clarity with respect to

relationships within the tribe Polygaleae Chodat. Nonetheless, all phylogenetic analyses have indicated that

traditional Polygala s.l. is a polyphyletic assemblage (Eriksen 1993a; Persson 2001; Forest et al. 2007), i.e.,

that several groups of species are more closely related to other genera than they are to other Polygala species.

Traditionally, nearly anything with a falsely papilionoid flower (two petals forming a ‘standard,’ one

conduplicate petal forming a ‘keel,’ and two petaloid lateral sepals forming ‘wings’) and with a bilocular

capsule has been left in Polygala, while groups with conspicuous specialized features, primarily modified

fruits, have been segregated as other genera. Globally, many additional infrageneric groups have been named,

based on vegetative and reproductive differences, but there has been neither consensus on which groups

to recognize nor agreement as to which rank is appropriate. There have been numerous regional floristic

treatments, sometimes even of a monographic nature, which have suggested various morphological features

for taxonomic use (e.g., Blake 1916, 1924; Marques 1979; Meijden 1988; Paiva 1998; Bernardi 2000), but not

J. Bot. Res. Inst. Texas 5(1): 125-1

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Journal of the Botanical Research Institute of Texas 5(1)

since Chodat’s century-old work (1893, 1896) has there been a global treatment of the genus Polygala. Thus,

there is no clear or reasonably comprehensive phylogenetic framework for assessing the level of universality

for putative synapomorphies.

As a result, attempts to stress any morphological feature (or suite of features) as more important than

any other for taxonomic purposes run the risk of being arbitrary or incomplete in focus. In the analyses of

Eriksen (1993a), Persson (2001), and Forest et al. (2007), several of the traditional subgroups within Polygala

have been shown to be non-monophyletic (or, if monophyletic, to make other groups non-monophyletic),

while some subgroups have been supported as monophyletic and many smaller groups have simply not

been adequately addressed. The two most complete phylogenetic analyses of Polygalaceae, both using only

plastid sequence data, included 25 species of Polygala s.l. (Persson 2001), six of which occur natively in

North America (a 7th, P. vulgaris L., is sparingly naturalized and an 8th, P. violacea Aubl. emend Marques,

is part of a widespread complex that may include the North American P. grandiflora Walter), and 49 species

of Polygala s.l. (Forest et al. 2007), seven of which occur in North America. It is clear that some of the North

American species are more closely related to other genera than they are to each other, but a robust global

phylogenetic context for assessing morphology and for generic circumscription has been lacking.

Forty-five species of North American Polygalaceae (a six-fold increase over the most complete published

study), representing 80% of the ca. 53 native species and all of the traditional subgenera, sections, and series

in North America north of Mexico, along with 90 extra-limital Polygala taxa and 44 members of other genera

(including two legumes as the outgroup), were phylogenetically analyzed using nrlTS and the plastid trnL

intron and adjacent spacer trnL-F (2778 total aligned characters). Those analyses were important in shaping

my taxonomic conclusions with respect to the Flora of North America treatment, and they will be discussed

in a forthcoming publication. As per the requirement for new names to have been published prior to their

use in the Flora of North America project, I present here a discussion of the morphologically diagnosable

groups that I will be recognizing as genera of Polygalaceae in North America.

RESULTS

In all analyses, the tribe Polygaleae was resolved as monophyletic, and in no analyses were all species tradi-

tionally recognized as Polygala recovered as a monophyletic group (Fig. 1). Although the exact relationships

between some of the clades within the Polygaleae were not consistently recovered, the following clades were

consistently supported across all, or nearly all, analyses (taxa traditionally included within Polygala s.l. are

indicated with an asterisk): Acanthocladus Klotzsch ex Hassk.*, Asemeia Raf.*, Badiera DC.*, Bredemeyera

Willd. s.str., Comesperma Fabill., Hebecarpa (Chodat) J.R. Abbott, stat. nov.*, Heterosamara Kuntze*, Polygala

subgen. Ligustrina (Chodat) Paiva*, Monnina Ruiz and Pav., Muraltia DC., Phlebotaenia Griseb.*, Polygala L.

s.str. (with an Old World subclade and a New World subclade), Polygaloides Haller*, Rhinotropis (S.F. Blake)

J.R. Abbott, stat. nov.*, Salomonia Four., and Securidaca L. In the few analyses where one of these clades

was not recovered, it was a result of a lack of resolution and not from a well-supported conflicting topology.

Although the addition of more taxa altered some supported relationships, the overall topology of most clades

from the 56 taxa analysis is robust to the addition of taxa across the combined nuclear and plastid analyses,

with robust support for the distinctiveness of the clades recognized or proposed herein as genera, as well

as for several of the subclades within Polygala s.str.

GENERAL DISCUSSION

A representative example of historical attempts to reclassify the North American species of Polygala is

provided by Small (1933), who recognized Asemeia Raf. (for P. grandiflora Walter and some N hemisphere

allies no longer recognized as separate species), Galypola Nieuwland (for P. incamata L.), Pilostaxis Raf. (for

a few species in an endemic southeastern U.S. group also recognized as Polygala ser. Decurrentes Chodat),

and Trichlisperma Raf. (for P. paucifolia Willd.; an erroneous spelling of Triclisperma Raf.), yet chose not to

recognize Anthalogea Raf. (for P. polygama Walt.) nor Senega Spach. (for P. senega L.), even though all of these

Fig. I.Ge

s s-Str. are part of the s

itlllilllf!

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Journal of the Botanical Research Institute of Texas 5(1)

genetically distinct from other traditional Polygala. This is also true for two groups not treated as distinct

by Small (1933), Hebecarpa and R hinotropis, despite their morphological distinctiveness. The remaining

‘genera’ recognized by Small (1933) are subgroups of the New World Polygala clade and none of them can

be maintained as genera without creating a blatantly artificial, non-monophyletic Polygala or overly splitting

to recognize numerous additional genera, most of which could not be morphologically diagnosed as distinct

from Polygala. Morphological distinctiveness has to be understood within a global phylogenetic context, and

which clades to recognize as genera, while always inherently subjective, should nonetheless be constrained

by the phylogenetic framework. Distinctive subclades can always be recognized as subgenera, without any

compulsion to assign all species to a subgenus, i.e., the undesired situation of paraphyletic or polyphyletic

genera can be circumvented while maintaining names for the more distinctive subgroups. Such an approach

allows us to have taxonomy reflect phylogeny without ignoring distinctive subgroups nor obfuscating patterns

of relationship by formally recognizing artificial assemblages of taxa that are not each other’s closest relatives.

More than a century ago (Robinson 1906), botanists were warned about the dangers of describing

artificial genera while focused on a geographically restricted flora and ignoring global patterns of varia-

tion among potentially related species. A regional approach to resolving relationships leads to confusion,

overlapping circumscriptions, a lack of compatibility or comparability across floras, and taxonomic and

nomenclatural instability (Robertson 1906). Higher level ranking is inherently subjective and was histori-

cally often rather arbitrary, primarily since there was no reliable, agreed-upon methodology for assessing

patterns of evolutionary relationship. Now, phylogenetic hypotheses often provide a solid framework for the

hierarchical grouping of species and understanding the interrelationships between groups, even though the

importance of broad geographical sampling remains. Ultimately, as pointed out by Humphreys & Linder

(2009), genera should be stable and predictive, as well as diagnosable and of a workable size, if possible, in

order to be useful.

Over time, there have been shifts in how we conceive of genera and in the kinds of data we use to analyze

them (Humphreys & Linder 2009). There has been much discussion and criticism of the Linnaean system

and traditional type-based ranked names vs. phylogenetic naming systems, e.g., Benton (2000), Withgott

(2000), Queiroz and Cantino (2001), Langer (2001), Berry (2002), Forey (2002), Nixon et al. (2003), Pickett

systems seem desirable. But, as pointed out by Moore (2003), even though our current system does use

Linnaean conventions, it is not strictly bound by traditional ideas and is flexible and able to handle changing

ideas. Names do not exist in a vacuum, they should and do imply something, and they are important for

communication. Advances in our understanding should attempt to minimize taxonomic change when pos-

sible (Entwisle & Weston 2005), in order to minimize confusion and disruption of existing communication

channels. But advances in our understanding do necessitate change for effective transmission of improved

knowledge (Knapp et al. 2004).

Thus, since it is clear from my results (Fig. 1) and the work of others (Eriksen 1993a; Persson 2001;

Forest et al. 2007) that Polygala s.l. is an artificial construct and must be recircumscribed, I have done so

using names in accordance with the International Code of Botanical Nomenclature (McNeill et al. 2006)

that provide historical continuity and minimize disruption. If only clades that are well-supported and

morphologically diagnosable are named, then taxonomic stability can be achieved, although there is still

often some room for interpretation. Nomenclatural codes are independent of taxonomic opinion, and it is

important not to blur the lines between taxonomy and nomenclature (Dubois 2007), keeping in mind that

true, lasting nomenclatural stability will only come from the quality science required to develop robust

hypotheses (Knapp et al. 2004). Therefore, I have avoided renaming any suprageneric clades for now, as

the relationships between the genera proposed herein are not yet robustly supported. Likewise, hypotheses

of relationships of clades closest to the core group of Polygala are also not yet well understood, and much

more work is needed to sort out which infrageneric groupings within Polygala s.str. will best reflect both

morphology and phylogeny.

129

Monophyly is currently considered by most taxonomists to be the most important criterion for taxonomic

changes (e.g., Backhand & Bremer 1998; Entwisle & Weston 2005; Judd et al. 2007). While phylogenetic

studies can yield robust hypotheses of relationship with predictive power, it should be kept in mind that

monophyletic groups are not necessarily predictive of overall similarity (just of shared derived similarity, e.g.,

Stevens 1985). Robustness of a clade, i.e., its persistence in new analyses and with the addition or deletion of

taxa, should be well-documented before making taxonomic changes, as such robustness is an indication of

the strength of the hypothesis that that group actually exists in nature (as a result of genealogical descent with

modification). The results here are robustly supported, i.e., the clades I am recognizing are supported in all

analyses, across all data sets and using double and triple the number of taxa (Fig. 1). These results also largely

mirror those of other published phenetic (Paiva 1998) and phylogenetic analyses (Eriksen 1993a; Persson

2001; Forest et al. 2007), at least with respect to recognition of the clades named as generic groups here.

Some workers dismiss as inappropriate the idea that genera should be easy to recognize (Entwisle &

Weston 2005) or write that the issue of convenience and utility is trivial (Stevens 1985). I agree that dis-

tinctive characters should not be emphasized nomenclaturally if they result in non-monophyletic groups

(e.g., Pfeil & Crisp 2005), i.e., if the characters have been shown to be an artificial human construct not

representative of shared ancestry they may still be useful for identification purposes but should not form the

basis of formal classification. However, taxonomists need to keep in mind that classifications serve a user

community, and considerations of recognizability, utility, and information retrieval, therefore, are important,

at least as secondary criteria (Backlund & Bremer 1998; Judd et al. 2007).

The genera I propose here, Asemeia, Hebecarpa, Polygaloides, and Rhinotropis, are strongly supported in all

the analyses, even when the relationship between the clades is NOT resolved (Fig. 1), they are morphologi-

cally diagnosable, and they make sense biogeographically. When there are alternative interpretations, such

as combining sister clades, the biology of the organisms should be considered to determine which classifica-

tion is most meaningful or useful, or makes the most sense in terms of diagnosable groups. For instance,

Eriksen et al. (2000) circumscribed the genus Badiera to include Polygala subgen. Hebecarpa (Chodat) S.F.

Blake on the basis of unpublished, preliminary data that suggested they were sister taxa, an hypothesis

not yet supported by any other published analyses (e.g., Forest et al. 2007) and one that was formulated

without any morphological context or detailed study of the species. Even though most of my analyses do

support the hypothesis of Badiera and Hebecarpa as sister groups, it makes more sense morphologically td ’

treat them as separate genera. Treating them as separate genera is also a more stable means of addressing

the lack of robustness, i.e,, all analyses support them as clades, but not all analyses support them as sister

are not yet any known morphological synapomorphies for the putative Badiera + Hebecarpa clade, although

each is independently a morphologically distinctive group that has been traditionally recognized.

The generic proposals provided here are not a reflection of personal opinion about which characters I

think most represent the concept of a genus; instead, they reflect a phylogenetic approach to understand-

ing relationships (based on DNA sequences), using morphology as a secondary guide/criterion for naming

clades and ranking them within a taxonomic system (see Backlund & Bremer 1998; Judd et al. 2007), while

also attempting to adhere to traditional names as much as possible. Even though all of the clades here given

recognition as genera are morphologically diagnosable, it is often difficult to determine which features are

synapomorphic for particular generic clades, especially when the characters are homoplasious. I hope in a

future study to develop a detailed morphological matrix, so that morphological features can be explicitly

mapped on to DNA-based as well as total evidence trees.

All the genera proposed here are traditionally recognized taxa (as subgenera or sections), and their

recognition does not result in the paraphyly of any other genera. Conversely, to create a broadly circum-

scribed monophyletic Polygala would necessitate the inclusion of all other genera of Polygaleae! Despite the

arbitrariness of rank, subsuming within Polygala s.l. a dozen or more distinctive generic clades such as the

Asian mycoheterotrophs (E pirixanthes Blume and Salomonia), the pantropical samaroid lianas ( Securidaca ),

130

Journal of the Botanical Research Institute of Texas 5(1)

the primarily neotropical and mostly drupaceous shrubs ( Monnina ), etc., would do absolutely nothing to

clarify our understanding of relationships within the tribe.

The species names transferred here (and the resulting new combinations) reflect my current under-

standing of the North American species, even though some of them are part of species complexes that could

be split apart or treated more broadly. Some of the genetic polymorphisms detected in my phylogenetic

analyses raise awareness of species-delimitation issues, e.g., at what point should distinctive populations

be nomenclaturally recognized, but such issues must await monographic studies of the groups involved.

In addition, all of the genera addressed here contain species outside of North America that are not being

nomenclaturally transferred at this time, because the species have not been studied by me and because the

primary purpose of this publication is to make the formal name changes required prior to using a name in

the Flora North America treatment of Polygalaceae (Abbott, in review.).

Taxonomic Conclusions. — Paiva (1998) re-segregated the Old World genus Heterosamara and recog-

nized 12 subgenera within Polygala s.l. My most complete cladistic analyses (Fig. 1), in congruence with other

published analyses, show that eight of the subgenera of Polygala are more closely related to other genera than

they are to subgenus Polygala and that Polygala subgen. Brachytropis (DC.) Chodat is phylogenetically nested

within the subgen. Polygala s.str. The other two subgenera, Polygala subgen. Chodatia Paiva and P. subgen.

Gymnospora (Chodat) Paiva, have been supported as distinct from Polygala s.str. in other published phylo-

genetic analyses (Forest et al. 2007). Thus, Paiva recognized many distinctive, morphologically defensible,

monophyletic groups but maintained these (except Heterosamara) in an artificial, polyphyletic assemblage,

Polygala s.l.

With respect to the circumscription of Polygala s.str. (as here circumscribed), the two primary concerns

are that the relationships between the primarily Old World groups Polygala subgen. Chodatia , Heterosamara,

and Polygaloides are not well-resolved, and that the New World and Old World clades of Polygala s.str. are

not always supported as sister taxa. Taxon sampling for the group containing Polygala subgen. Chodatia,

Heterosamara, and Polygaloides has been limited, but morphology does indicate that the three groups are

distinctive. It is conceivable that Polygala s.str. could be expanded to include these three lineages, although

results to-date suggest that this is unlikely and depends on the exact placement of the genera Muraltia and

Salomoma.

North American Groups and Brief Biogeographic Discussion. — Monnina is represented by a single

species in the U.S., M. wrightii A. Gray, in Arizona and New Mexico, representing the northern limit of a

wider range in Mexico. Monnina wrightii also occurs disjunctly in Bolivia and Argentina, with single collec-

tions known from Peru and Uruguay (Eriksen 1993b). Eriksen hypothesized that the Mexican-United States

population is likely the result of long-distance dispersal from the region of Bolivia-Argentina. My analyses

show material from Bolivia does form a clade with material from Mexico, although there is some genetic

divergence, suggesting that these populations may have been isolated for some time. My data show that the

Central American species of Monnina do not form a clade, thus it seems clear that there have been multiple

movements between the Andes and Central America. Most species of Monnina are shrubs with drupaceous

fruits in the mountains of Central and South America, but Monnina wrightii is a member of a basal grade

partially characterized by herbaceous habit and dry, flat, samaroid fruits. This group has been recognized

generically as Pteromonnina, but Eriksen and Persson (2007), although citing unpublished, preliminary data

suggesting the group is monophyletic, pointed out that the group is difficult to separate morphologically from

other Monnina species. My cladistic analyses indicate that the species of Pteromonnina form a paraphyletic

grade (M. leptostachya Benth., M. stenophylla A. St.-Hil., M. tristaniana A. St.-Hil., M. wrightii), not a clade,

providing preliminary support for the conclusion that Pteromonnina should be included within Monnina.

Amphitropical distribution patterns that span the neotropical lowlands are well-known in other groups of

plants ( e.g ., Raven 1963; Solbrig 1972a, 1972b), and although many involve California and Chile (e.g., Morrell

et al. 2000), some patterns are similar to that of Monnina wrightii ( e.g ., Holmes et al. 2008). Long-distance

dispersal is accepted as the most likely scenario, but possible vectors and timing are not well understood.

132

Journal of the Botanical Research Institute of Texas 5(1)

to be related to P. violacea Aubl., and it is not clear how to separate it morphologically from some of the

Latin American entities. Many workers, including Blake (1916, 1924) have recognized numerous segregate

species out of P. grandiflora within the U.S. alone, while other workers have lumped all of them, including P.

grandiflora, into P. violacea (e.g., Bernardi 2000). Preliminary molecular analyses (Abbott, unpublished data)

suggest that P. grandiflora and P. violacea may not be each other’s closest relatives. Molecular DNA analyses

and observational field work suggest that the North American segregates can not reliably be segregated

taxonomically and that P. grandiflora can only be included in P. violacea if several other currently recognized

species are also synonymized, including many taxa considered by Brazilian workers to be distinct (e.g.,

Aguiar 2008). Therefore, I recommend that P. grandiflora be maintained apart from P. violacea, at least until

there has been a detailed, robustly supported, comprehensive study of the complex. Restricted to the SE

U.S. within North America, P. grandiflora may have moved in from Mexico during one of the hypsithermal

intervals known to have brought western species into the southeast, or it may have spread into the southeast

through Florida via the Caribbean.

Hebecarpa contains 40-70 species. Most species are Mexican, with a few extending into South America

and nine extending into the SW U.S, five of which were included in my phylogenetic analyses. There are

several species complexes in this group that are in serious need of revision, with overly fine, and perhaps

meaningless, historical splitting (often based on ||||p^single or a few specimens), so that the total number

of species is probably fewer than 50. Chodat (1893) included the genus Badiera DC. as a subsect, in his

sect. Hebecarpa, recognizing 25 species in subsect. E uhebecarpa, but Badiera has since been shown to be a

distinctive clade separate from Hebecarpa, e.g., Forest et al. (2007) and my analyses (Fig. 1; Abbott 2009).

Blake (1916) circumscribed three sections, three subsections, and two series within his subgen. Hebecarpa.

One of the sections, sect. Biloba Blake, is now known to be part of the Rhinotropis group (Wendt 1978;

Abbott 2009) and is discussed there. The monotypic sect. Huateca Blake, with the Mexican P. tehuacana

Brandegee, has not been included in any phylogenetic analyses, so it should be tentatively viewed as incertae

sedis, although morphological descriptions do suggest it belongs in Hebecarpa. Section Hebecarpa, which

includes all of the North American species, was divided by Blake into three subsections, subsect. Microthrix

Blake, subsect. Hebantha Blake, and subsect. Adenophora Blake. Subsection Microthrix contains seven of the

North American species, one in ser. Ovatifoliae Blake (P. ovatifolia Gray), the others in ser. Obscurae Blake.

Subsection Hebantha Blake was circumscribed with 34 species, none of them North American. Two of the

three species of subsect. Adenophora range into North America, P. greggii S. Watson [= P. glandulosa Kunth]

and P. macradenia A. Gray. My preliminary molecular DNA analyses suggest that subsect. Adenophora is

sister to the rest of Hebecarpa. Final circumscription of subgroups must await additional study of more taxa.

Species of Hebecarpa in the U.S. are largely restricted to (semi-)arid regions of the southwest, representing

contiguous northern expansions of their ranges in Mexico, which is the center of diversity for this group. A

1-most species,

suggesting that they have arrived in these regions secondarily.

Polygaloides contains six or seven species globally and corresponds to subsection III of Chodat’s (1893)

Polygala sect. Chamaebuxus, plus one species unplaced by him to subsection, P. paucifolia, disjunct in northern

North America from the remaining five or six species in Europe and northern Africa. This North American

species is isolated and morphologically distinct from its closest relatives in Europe and northern Africa, per-

haps as a vicariant relict of a historical circumboreal flora. While its creeping herbaceous habit with relatively

thin leaves and large showy crest set the North American taxon apart from the Old World taxa [subshrubs or

shrubs to 1 m tall with subcoriaceous leaves and small, fairly inconspicuous crest], the form of the stigma,

the glanduliform disc at the base of the flower, and the few-flowered inflorescences morphologically unite

the taxa (Chodat 1893). No existing descriptions for this Polygaloides group are entirely accurate, as, based

on phylogenetic analyses, they all include elements that are not part of this clade or exclude species that are,

e.g., in Blake (1916) the. only species treated in the Chamaebuxus group were those belonging to the group

here recognized as Rhinotropis, with no reference to the Old World species, nor to P. paucifolia. Blake (1924)

38:151-177.

Schwarz, 0. 1949.

Wendt, T.L. 1978.-

Journal of the Botanical Research Institute of Texas 5(1)

BOOK REVIEW

Michel Poulain, Marianne Meyer, and Jean Bozonnet, Text in French with translation of glossary and keys to

identification into English by Annie Kohn. 2011. Les Myxomycetes: Guide to Identification. (ISBN:

978-209518540-2-4, EAN: 9782951854024, pbk.). La Federation Mycologique et Botanique Dauphine-

Savoie. (Orders: www.fmbds.org; tresorier@fmbds.org). €120.00, 568 pp. (Vol. 1), 544 pp. (Vol. 2),

675 color images, 556 b/w line drawings mostly microscopic structures, 9W x 6 3 A".

The illustrations are grouped by orders: Liceales, Echinosteliales, .

oring countries. 1

1700 University Drive, Fort Worth, Texas 76107-3400, ty'lrg

J. Bot. Res. Inst. Texas 5(1):138.2011

NEW COMBINATIONS IN PHORADENDRON LEUCARPUM (VISCACEAE)

J. Richard Abbott

Ralph L.Thompson

Missouri Botanical Garden Research Associate

St. Louis, Missouri 63166, USA.

University of Florida Flerbarium Courtesy Assistant Scientist

Florida Museum of Natural Flistory

Gainesville, Florida 326 1 1, U.S.A.

rabbott@ufl.edu

Flancock Biological Station, 56 1 Emma Drive

Murray State University

Murray, Kentucky 42071, U.S.A.

Berea College Flerbarium, Department of Biology

Berea College, Berea, Kentucky 40404, USA,

ralph_thompson@berea.edu

ABSTRACT

Reveal and Johnston (1989) established Phoradendron leucarpum (Raf.) Reveal & M.C. Johnst. as the no-

menclaturally correct name for American mistletoe, a species native to the United States and Mexico, based

on a 1817 collection from the Carolinas. Initially, Rafinesque gave two binomials for American mistletoe:

Viscum leucarpum in 1817 and V. serotinum in 1820 (Reveal and Johnston 1989). Johnston (1957) made the

combination Phoradendron serotinum (Raf.) M.C. Johnst. for American mistletoe because he considered the

specific epithets leucarpum and leucocarpum to be orthographic variants. Brummitt (1988) reported a

ruling from the Committee for Spermatophyta that Phoradendron leucarpum, an eastern United States spe-

cies, and P. leucocarpum, a Peruvian species, were not homonyms. Following Brummitt (1988) and the rule

of priority, Johnston’s (1957) combination became a synonym. Distinctive Phoradendron taxa ranging from

Texas to California to Oregon, and into Mexico have variously been proposed as infraspecific entities under

epithets other than “leucarpum,” e.g., “serotinum,” or as distinct species (Wiens 1964; Kuijt 2003). Despite

the conclusions reached in the 1989 article, the most recent monographer of Phoradendron (Kuijt 2003) did

not use the correct binomial P. leucarpum. Consequently, for workers who want to accept Kuijt’s taxonomic

judgment, the correct nomenclatural combinations for infraspecific taxa within P. leucarpum do not exist.

The choice of which name(s) to use for American mistletoe has been the source of much taxonomic and

nomenclatural contention during the last century, as discussed by Kuijt (2003). Prior to 1989, Phoradendron

^tiflso taxonomic judgment. We are not challenging Kuijt’s t;

and ranks as he defined them. Our goal is to provide the corre

(Raf.) Reveal & M.C. Johnst. i

Major post-2003 national and international online databases accept Phoradendron leucarpum (i.e., GRIN,

USDA, ARS, National Genetic Resources Program 2010; The International Plant Names Index 2010; PLANTS

Database, USDA, NRCS 2010), as do numerous publications on a wide variety of themes, e.g., general parasite

J.Bot. Res. Inst. Texas 5(1): 139-1

Reveal, J.L. and M.C. Johnston. 1 989. A new combination in Phoradendron (Viscaceae), Taxon 38:1 1 / ,

The International Plant Names Index (IPNI). 201 0. Online Database, http://www.ipni.org, 1 9 Dec 201 0.

USDA, ARS, National Genetic Resources Program. 2010. Germplasm Resources Information Network - (GRIN)

Online Database. National Germplasm Resources Laboratory, Beltsville, MD. http://www.ars-grin.gov/cgi-bin/

npgs/html/index.pl, 1 9 Dec 201 0.

USDA, NRCS, 2Q1Q. The PLANTS Database (http://plants.usda.gov, 19 Dec 2010). National Plant Data Center,

Baton Rouge, LA 70874-4490 USA.

Wiens, D. 1 964. A revision of the acataphyllous species of Phoradendron. Brittqisij^^^^

142

$125.00 (US), $141.50 (Canada), 352 pp„ 170 c

TYPIFICATION AND SYNONYMY OF THE SPECIES OF EUPHORBIA SUBGENUS

ESULA (EUPHORBIACEAE) NATIVE TO THE UNITED STATES AND CANADA

Dmitry V. Geltman

Paul E. Berry

Komarov Botanical Institute of the

Russian Academy of Sciences

2 Prof. Popov Street

St. Petersburg, 197376, RUSSIA

geltman@mail.ru

Ricarda Riina

University of Michigan Herbarium and

Department of Ecology and Evolutionary Biology

3600 Varsity Drive

Ann Arbor, Michigan 48108-2228, U.S.A.

riina@umich.edu

University of Michigan Herbarium and

Department of Ecology and Evolutionary Biology

3600 Varsity Drive

Ann Arbor, Michigan 48108-2228, U.S.A.

peberry@umich.edu

Jess Peirson

University of Michigan Herbarium and

Department of Ecology and Evolutionary Biology

3600 Varsity Drive

Ann Arbor, Michigan 48108-2228, U.S.A.

peirsonj@umich.edu

ABSTRACT

INTRODUCTION

Euphorbia L. subgenus Esula Pers. contains approximately 470 species, with the majority of the species

occurring in temperate Eurasia. There are also several smaller, disjunct centers of diversity in areas such

as Macaronesia, the East African mountains, South Africa, and the New World. This group was formerly

recognized by some botanists as the separate genus Tithymalus Gaertn. (e.g., Klotzsch 1860; Small 1903;

Chrtek & Krisa 1982, 1992), but it is generally maintained within Euphorbia. Molecular phylogenetic studies

(Steinmann & Porter 2002; Bruyns et al. 2006; Kryukov et al. 2010; Zimmermann et al. 2010) have shown

that the subgenus corresponds to one of four major clades within the genus, and the latest results, using

multiple markers from all three plant genome compartments, confirm that it is the sister group to the other

three subgenera (Horn et al. 2009).

Euphorbia subgenus Esula is represented in the continental United States and Canada (the area covered

by the Flora of North America project) by 17 native species and 15 naturalized Eurasian species. The only

paper dealing specifically with the North American species is Norton (1899), where he described several

new infraspecific taxa and followed the infrageneric classification system of Boissier (1862). There have been

no further attempts since then to systematize the nomenclature of this group, discuss possible relationships

to Eurasian species, or place the North American species in more recent infrageneric classifications, such

as those of Prokhanov (1949, 1964) and Geltman (2007).

This paper is part of a global initiative to update the systematics of Euphorbia worldwide (Berry & Riina

2007; Esser et al. 2009), and it is also a precursor to the treatment of Euphorbia for the Flora of North America

J. Bot. Res. Inst. Texas 5(1): 143-1

Journal of the Botanical Research Institute of Texas 5(1)

(FNA) project. Here we aim to systematize the nomenclature and typification of the native North American

species of subgenus Esula and to provide a discussion of their relationships with Eurasian taxa.

Holdings of several North American (DWC, GH, KSC, MICH, MO, NY, RM, US) and European (G, E I I !

herbaria were examined, as well as available online type collections of other herbaria. Protologues of all

names treated were examined. Norton (1899) published several new species and varieties of Euphorbia from

North America, and although he cited several syntypes for these names, he usually made annotations of

“types” on particular herbarium sheets. Such specimens are designated here as lectotypes. In other cases, if

several duplicates of the type collection were located, the holotype was assumed to come from the herbarium

where the author of the name was working at the time (e.g., MO for G. Engelmann, RM for A. Nelson, and

F for C. Millspaugh).

Native species in the FNA area are arranged alphabetically. For every accepted name or synonym rec-

ognized, typification information is provided. Relationships of North American species or species groups

to Eurasian groups are discussed initially and in the individual species listings.

The 17 native species of Euphorbia subgenus Esula represent a number of different lineages, based on morpho-

logical and molecular data, and they can be separated into the five main groups listed below. Since molecular

phylogenetic data for the subgenus is still preliminary and broader taxon sampling is in progress, we focus

our discussion here on current morphological concepts of sections, which we acknowledge are likely to be

superseded once more molecular data is obtained and analyzed. Our main reference is to the infrageneric

system developed by Geltman (2007), which covers all the temperate Eurasian species of Euphorbia.

1. Euphorbia purpurea (Raf.) Fernald. — this species is unique among the North American esu-

loid Euphorbias, since it is a perennial, shady forest-edge species with warty fruits, and it has an eastern

Appalachian/Alleghenian distribution. In the Geltman (2007, 2008) system, it belongs to E. sect. Chamaebuxus

Lazaro, which consists of about 95 perennial species that have hornless, elliptical cyathial glands, usually

warty fruits, and shiny brown seeds. With the exception of E. purpurea, all other members of the section are

native to the Old World, occurring in Eurasia or Mediterranean North Africa, from the Atlantic Ocean across

the continent to the Pacific, and with a few species extending into the mountains of east tropical Africa.

Two European species that may be closely allied to E. purpurea are E. hyberna L. (Spain, France, Italy, and

the British Isles) and E. squamosa Willd. (Caucasus region), both of which also occur in broadleaf forests.

2. Euphorbia trichotoma Kunth. — this species is also unique among the North American esuloid

Euphorbias, because it is a subtropical Caribbean maritime perennial species, occurring along beaches

in southern Florida. It is similar in habit to the maritime E. paralias L., which is native to the Atlantic and

Mediterranean shores of Europe, and molecular data (Geltman et al. 2010; Kryukov et al. 2010) place it close

to this species, which belongs to E. sect. Paralias Dumort. subsect. Paraliodieae Prokh.

3. Euphorbia brachycera group. — this is a group of five species that occur mainly in the western U.S.

and are perennials with short-horned or irregularly dentate cyathial glands and irregularly and shallowly

pitted to nearly smooth seeds. Based on these characters, they are morphologically most closely allied with

the Eurasian E. sect. Herpetorrhiza (Prokh.) Prokh., which comprises about 15 species from Central Asia

and the Mediterranean region. Besides E. brachycera Engelm., the group includes E. chamaesula Boiss., E.

lurida Engelm., E. schizoloba Engelm., and E. yaquiana Tidestr.

4. Euphorbia commutata group. — this is a group of seven annual species with horned glands, smooth

fruits, and diversely ornamented (or rarely smooth) seeds. In the Eurasian classification system, they belong

to E. sect. Peplus Lazaro, which consists of seven subsections that are probably paraphyletic, based on initial

molecular data (Kryukov et al. 2010). Besides E. commutata Engelm., the group includes E. crenulata Engelm.,

E. helleri Millsp., E. longicruris Scheele, E. peplidion Engelm., E. roemeriana Scheele, and E. tetrapora Engelm.

150

Journal of the Botanical Research Institute of Texas 5(1)

This is a perennial species in the E. brachycera species group that is endemic to Arizona.

ACKNOWLEDGMENTS

The work was supported by a joint grant of the U.S. Civil Research and Development Foundation (project

N RUB1-2916-ST-07) and the Russian Foundation for Basic Research (project N 08-04-91103), as well as

a NSF Planetary Biodiversity Inventory award (DEB 0616533). We are grateful to the curators of herbaria

(DWC, G, GH, KSC, LE, NY, MICH, MO, P, RM, US) for allowing us to examine their holdings, and to

Sharon Bartholomew-Began for providing photographs of the historic Euphorbiaceae specimens at DWC,

Roger Kiger for his expertise on Rafinesque handwriting, and Mark Mayfield for discussions on the North

American euphorbias. Victor W. Steeinmann and an anonymous reviewer provided helpful reviews.

REFERENCES

Berry, P.E. and R. Riina. 2007. A new collaborative research project: a global inventory of the spurges. Euphorbia

World 3(1 ):1 2-1 3.

Boissier, E. 1 862. Euphorbiaceae— Euphorbieae. In: Candolle, A. P. de. Prodromus systematis naturalis regni veg-

etabilis. Sumptibus Victoris Masson et filii, Parisiis. 1 5(2):3-1 88.

B%pi$/Pty, RJ Mapaya, andT Hedderson. 2006. A new subgeneric classification for Euphorbia (Euphorbiaceae) irp

southern Africa based and psbA-trnH sequence data. Taxon 55:397-420.

Chrtek, J. and B. Krisa. 1 982. Euphorbiaceae. In: J. Futak (ed.). Flora Slovenska 3:406-462. Bratislava.

Chrtek, J. and B. KrJsa. 1 992. lithymalus Gaertn. In: Hejny, S. arid B. Slavik, eds. Kvetena Ceske Republiky. Academia,

Prague. 3:321-346.

Correa, M.N. 1988. Euphorbiaceae. In: M.N. Correa, ed. Flora Patagonica. Parte V. INTA, Buenos Aires. Pp. 75-^91.

Esser, FI P.E. Berry, and R. Riina. 2009. Euphorbia: a global inventory of the spurges. Blumea 54:1 ..

Geltman, D.V. 2007. Conspectus of the system of Euphorbia L. subgenus Esula Pers. (Euphorbiaceae) of the non-

tropical Eurasia. Nov. Sist. Vyssh. Rast. 39:224-240. [In Russian],

Geltman, D.V. 2008. Conspectus of Euphorbia L. section Chamaebuxus Lazaro (Euphorbiaceae). Nov. Sist. Vyssh.

Rast. 40:1 09-1 58. [In Russian]

Geltman, D.V., P.E. Berry, C. Fergusson, W. Jin, A. Kryukov, M. Mayfield, R. Riina, and A. Rodionov. 2010. Phylogenetic

relationships between Eurasian and North American species of Euphorbia subgenus Esula (Euphorbiaceae):

, ; y^^^^from morphological and molecular (ITS sequences) data. XII Moscow conference on plant phylogeny,

devoted to 250th anniversary of Georg-Franz Floffmann. Proceedings. KMKScientific press, Moscow. Pp. 79-82.

Govaerts, R., D. Frodin, and A. Radcliffe-Smith. 2000. World checklist and bibliography of Euphorbiaceae (with

Pandaceae). 3 vols. Royal Botanic Gardens, Kew.

Goyne, M.A. 1 991 . A life among the Texas flora. Texas A&M University Press, College Station.

FIorn, J.W., B.W. van Ee, JJ. Morawetz, R. Riina, P.E. Berry, V.W. Steinmann, and K. Wurdack. 2009. Phylogeny and evolu-

tion of growth forms in the giant genus Euphorbia (Euphorbiaceae). Abstract, Botany and Mycology 200|§

meetings, Snowbird, UT, 25-29 July, 2009.

Kearney, T.H. and R:H. Peebles. 1951. Arizona flora. University of California Press, Berkeley.

1860. Linne's naturliche Pflanzenklasse Tricoccae des Berliner Herbarium's im Allgemeinen und"

die naturlicheche Ordnung Euphorbiaceae insbesondere. Abhandl. Konigl. Preuss. Akad. Wiss. Berlin (1859),

Phys. Abhandl. Pp. 1-108.

Kryukov, A.A., D.V. Geltman, E.M. Machs, and A.V. Rodionov. 2010. Phylogeny of Euphorbia subgenus Esula

(Euphorbiaceae) inferred on the sequences of ITS1 -5.8S rDNA-ITS2. Bot. Zhurn. 95:801 -81 9 [in Russian],

Merrill, 1949. Index Rafinesquianus. Arnold Arboretum of Harvard University, Jamaica Plain, MA.

Norton, J.B.S. 1899. North American species of Euphorbia section Tithymalus.^^58, plates 11-52. St. Louis.

Reprint issued 10 July 1899 in advance of Norton, J. B. S. 1900. A revision of the American species of the

Euphorbia of the section Tithymalus occurring north of Mexico. Ann. Rep. Missouri Bot. Gard. 1900. 1 1:85-144,

152

Journal of the Botanical Research Institute of Texas 5(1)

BOOK REVIEWS

Gregory P Cheplick and Stanley H. Faeth. 2009 . Ecology and Evolution of the Grass-Endophyte Symbiosis.

(ISBN 13: 978-0-19-530808-2, ISBN 10: 0195308085, hbk.), Oxford University Press, Inc, 198 Madison

Avenue, New York, New York 10016, U.S.A. (Orders: www.oup.com/us/catalog/general/subject/

LifeSciences/Ecology/?view=usa&ci=9 780195308082; 800-451-7556; custserv.us@oup.com). $75.00,

256 pp., 5 halftones, 24 line illus., 6 Vs" x 9 VC.

Jan Wrede. 2010. Trees, Shrubs, and Vines of the Texas Hill Country: A Field Guide, Second Edition.

(ISBN- 1 3 :978- 1 -60344-188-9, ISBN-10: 1-60344-188-3, Flexbound with flaps). Texas A&M University

Press, John H. Lindsey Building, Lewis Street, 4354 TAMU, College Station, Texas 77843-4354, U.S.A.

aspx; 1-800-826-8911; Toll Free U.S. only 979-847-8752 fax). $24.00, 272 pp., 195 color photos’, map

2 tables, 8 3 A" x 8V2 M .

J. Bot. Res. Inst. Texas 5(1):152. 2011

IMPATIENS VEERAPAZHASII (BALSAMINACEAE), A NEW SCAPIGEROUS BALSAM

FROM WAYANAD, WESTERN GHATS, INDIA

N. Anil Kumar, M.K. Ratheesh Narayanan, P. Sujanapal,

R. Meera Raj, K.A. Sujana, and Mithunlal

MS. Swaminathan Research Foundation

Puthoorvayal - 673 121

Kalpetta, Wayanad, Kerala, INDIA

RESUMEN

INTRODUCTION

The genus Impatiens (Balsaminaceae) is diversified with more than 1000 species distributed mainly in the

wet tropics and in north temperate regions (Mabberley 2008). The western part of south Asia and contiguous

southeast Asian regions are the major centers of diversity and endemism. A few representatives are reported

from America and Europe (Mabberley 2008). In India the genus is represented by 203 taxa mainly distrib-

uted in three major centers of diversity i.e., Western Himalayas, North East India, and the Western Ghats,

each characterized by its own species group (Vivekananthan et al. 1997). In India, scapigerous Impatiens

form a peculiar section ‘Scapigerae’ and are restricted to Western Ghats. So far, among the world distribu-

tion range of Impatiens, this section is reported only from Western Ghats-Sri Lankan phytogeographical

region (Grey-Wilson 1980; Vivekananthan et al. 1997) and is endemic to Western Ghats-Sri Lanka Hotspot

of Biodiversity. Except Impatiens acaulis Arn., which is distributed throughout the wet tropical forests of

Western Ghats and Sri Lanka, all the other species have very narrow distribution patterns in various small

microcenters of endemism in Western Ghats, especially in its southern part. The Wayanad district of Kerala

forms a biodiversity rich area in the Nilgiri Phytogeographical region of Western Ghats. Because of rich

tropical forests and suitable climatic factors at an average altitude of 1000 m asl, the area supports diverse

ephemerals especially a genus such as Impatiens.

The Wayanad district in Kerala is unique and the entire area is along the Western Ghats with altitudes

ranging from 700 m to 2100 m msl. Nineteen species of Impatiens were reported from the Wayanad district

(Ratheesh Narayanan 2009). During our floristic exploration in the tropical evergreen forests of Wayanad,

some specimens of epiphytic scapigerous Impatiens were collected from the Kurichiarmala-Banasura hill

ranges. These scapigerous Impatiens were strikingly different from other species of scapigerous Impatiens,

due to their large leafy bracts, a spurless lower sepal, and tubercled, hairy, golden yellow seeds. A careful

examination revealed that the leaf, bracts, and texture of the scape, size and shape of lateral united petals,

dorsal auricle and seed structure are unique and distinct from all the other described species. Subsequent

detailed study, close observation, and consultation of relevant literature (Hooker 1908; Barnes 1939; Grey-

Wilson 1980 & 1985; Bhaskar 1981; Bhaskar & Razi 1983; Pandurangan & Nair 1995; Vivekananthan et

al. 1997; Viswanathan et al. 2003; Bhaskar 2006) have confirmed the plants are distinct from all the other

described taxa. Hence, it is described and illustrated here as a new species.

J. Bot. Res. Inst. Texas 5(1): 153-1

Single flower, F. Lateral sepal, G. Lower sepal, H. Dorsal petal, I. Lateral united petal, J. Anthers, K. Ovary, L. Fruit, M. Seed.

e, D. Bract, E.

156

Journal of the Botanical Research Institute of Texas 5(1)

Sujanapal, & Meera. Habit and close-up of flowers.

Pedicel 1.3-1. 5 cm

n July and peak time i:

>r ofM.S.

India. Bull. BotSurv. India 45:1-4.

) L.K. Ghara. 1997. Balsaminaceae. In: P.K. Hajra, VJ. Njj$

ay of India, Calcutta. Pp. 95-229.

A PHYLOGENETIC ASSESSMENT OF BREEDING SYSTEMS AND

FLORAL MORPHOLOGY OF NORTH AMERICAN 1POMOEA (CONVOLVULACEAE)

J. Andrew McDonald

Department of Biology

University of Texas - Pan American

1201 W. University Dr.

Edinburg, Texas 78539, U.S.A.

amcdonald@utpa. edu

Joshua R. McDill

Plant Resources Center and

Section of Integrative Biology

1 University Station F0404

The University of Texas at Austin

Austin, Texas 78712, U.S.A.

Plant Resources Center and

Section of Integrative Biology

1 University Station F0404

The University of Texas at Austin

Austin, Texas 7871 2 f USA.

Beryl B. Simpson

Plant Resources Center and

Section of Integrative Biology

1 University Station F0404

The University of Texas at Austin

Austin, Texas 78712, U.S.A.

ABSTRACT

RESUMEN

INTRODUCTION

Ipomoea, the largest genus in the Convolvulaceae is known globally for the sweet potato (I. batatas), the

sixth most important starch crop of the world (faostat.fao.org). In addition to the sweet potato, the genus

contains 600 to 1000 species (Austin & Huaman 1996; Manos et al. 2001), most of which occur in the New

World with approximately 167 (25%) native to temperate and tropical North America (including Mexico).

These species display a wide range of floral morphologies indicative of different pollinators and/or breeding

systems (autogamy versus xenogamy; Fig. 1A-F, 2A-D). While recent molecular phylogenetic studies have

included Ipomoea species (Miller et al. 1999; Manos et al. 2001, 2004; Miller et al. 2004), they have not used

phylogenetic hypotheses to examine changes in breeding system. Similarly, numerous studies have described

the breeding systems of various Ipomoea species (cf., Ennos 1981; Bullock et al. 1987; Devall & Thien 1992;

Chang & Rausher 1998; Chemas & Bullock 2002) but have not set these studies into a comparative frame-

work. Here we combine a phylogenetic study of North America Ipomoea with greenhouse studies of breeding

systems of all taxa included to assess the occurrence of different breeding systems in an evolutionary context

and examine the relationship between breeding systems and floral traits.

An historical and prevailing assumption that outcrossing (xenogamy) in flowering plants is responsible

J. Bot. Res. Inst. Texas 5(1): 159-1

Journal of the Botanical Research Institute of Texas 5(1)

for the remarkable diversity and evolutionary success of the angiosperms (Darwin 1877; Stebbins 1974;

Faegri & van de Pijl 1979) has been tempered in recent years by the recognition of high rates of autogamy

in herbaceous groups (Barrett et al. 1996; Kohn et al. 1996; Schoen et al. 1997; Goodwilliel999; Richman

& Kohn 2000; Lu 2001; Barrett 2002; Allem 2003; Igic et al. 2003) and the frequent occurrence of apo-

mixis in tropical tree groups (Kaur et al. 1978; Allem 2003; Ashman & Majestic 2006). Since 20-30% of all

flowering plant species produce offspring by means of self-fertilization (Barrett 2002; Allem 2003), there

is ample evidence that self-compatibility confers a competitive edge over self-incompatibility under certain

environmental conditions. Nevertheless, cross-fertilization is the favored mode of reproduction in most

angiosperm groups, leading to a general consensus that cross-fertilization promotes heterozygosity and

genetic polymorphisms in plant populations (Grant 1958; Faegri & van der Pijl 1979; Liu et al. 1998; Allem

2003), thereby facilitating adaptability to changing environments over the long course of evolutionary time

(Holsinger 2000).

Evolutionary biologists recognize two different explanations for high rates of self-pollination among

herbaceous angiosperm groups. The ‘automatic selection’ hypothesis states that alleles for self- compatibility

within gene pools of outcrossing individuals will be inherited in larger proportions than self-incompatibility

alleles (3:2) due to increased rates of transmission, this owing to the additive effects of both selhng and

cross-fertilization mechanisms (Fisher 1941; Jain 1976; Schoen et al. 1996; Holsinger 2000). Alternatively,

the ‘reproductive assurance’ hypothesis ascribes high rates of autogamy in plant populations of transitory

and insular habitats (Baker 1955; Webb & Kelly 1993; Bernardello et al. 2001) to a filter mechanism that

favors self-compatible disseminules and their ability to establish viable colonies in the absence of mates or

pollinators after long-distance dispersal events (Baker 1955, 1967; Stebbins 1974; Lloyd 1980; Pannell &

Theoreticians agree that selhng advantages are counter-balanced by genetic benefits that derive from

the process of cross-fertilization by increasing rates of pollen discounting and diminishing the expression

of lethal homozygous deleterious alleles (Lande & Schemske 1985; Charlesworth & Charlesworth 1987;

Harder & Wilson 1998). These predictions are substantiated by studies of natural plant populations, many

of which conclude that perennial plant species are more susceptible to inbreeding depression than annuals

on account of higher genetic loads (Lande & Schemske 1985; Charlesworth & Charlesworth 1987; Lloyd

& Schoen 1992; Barrett et al. 1996; Liu et al. 1998; Holsinger 2000; Barrett 2002). While experimental

crossing studies on both plants and animals support this perspective (Crnokrak & Barrett 2002), Lande

and Schemske (1985) predict plant populations should be able to purge themselves of deleterious alleles

by means of occasional selhng events within outcrossing populations. Populations must apparently pay,

however, an initial competitive price in progeny success rates during the purgation process.

Shifts from outcrossing to inbreeding modes of reproduction are believed to be initiated by the malfunc-

tion of self-incompatibility mechanisms (Barrett 2002). If selhng behaviors prove advantageous, then selective

forces will no longer favor floral characteristics that are associated with cross-pollination syndromes. Hence

Barrett et al. (1996) recognize that outcrossing is usually associated with a number of floral features and

syndromes that are attractive to pollinators, just as self- compatible flowers exhibit dysfunctional vestiges

of these same features. They also note that co-variation of two or more floral characters between taxa can

inform us about the adaptive signihcance of such features if we understand these attributes in the context

of their phylogenetic and functional linkages.

In the present study we generate a phylogenetic hypothesis based on existing and newly-collected ITS ■

sequence data for 70 North American Ipomoea taxa and use this phylogeny to pinpoint changes in breeding

systems and associated changes in floral traits. Breeding systems for the species included were determined

by experimental greenhouse studies. We also use these data to examine the impact of breeding system

changes on floral structures. Most Ipomoea species present large and showy flowers and produce substantial

amounts of nectar and pollen as attractants (Figs. 1, 2). In shifts from xenogamy to autogamy, one might

expect the relaxation of selection pressures that maintain showy floral features and the increased influence

microsepala and /. minutiflora, the sole members of Ipomoea ser. Microsepalae, exhibit strong differences in flower sizes as xenogamous and a

species (respectively). EJpomoeo rfumeforum exhibits typical features of an autogamous species derived from a melitophilous ancestor. Note

small (ca. 2.5 cm long) and stamens are subequal. The stigma maintains close contact with several anthers. F. Ipomoea aristolochiaefolia is an a

species that belongs to a group that is predominantly pollinated by hummingbirds or long-tongued flies (I.Exogonium si). Note the flowers a

of mechanisms that enhance selhng abilities. We point out the relationships between breeding system and

of large flowers and large amounts of pollen is hypothesized to decrease in selhng lineages (Goodwillie et

al. 2009) while the distance between anthers and stigma is hypothesized to decrease in order to facilitate

selhng.

Although many studies have discussed relationships between breeding system and floral evolution

(e.g., Cruden 1977, 2000; Lord & Eckhard 1984; Plitman & Levin 1990; Gallardo et al. 1994), this study

With an aim to examine the multiple origins of inbreeding in North American Ipomoea, we obtained DNA

162

Journal of the Botanical Research Institute of Texas 5(1)

Fig. 2. Contrastive floral features of closely related xenogamous and autogamous sister species of Ipomoea in North America. A. Ipomoea variabilis (/.

sect. Exogonium) is xenogamous and shares a number of synapomorphic characteristics with the sister species, I. meyeri (late leaf primordia sub-bullate;

sepals foliose and hispid, corolla tube basally yellow-pigmented). Like other outcrossing Ipomoea, this species has relatively large and colorful flow-

ers, heteromophic stamens and style (herkogamous flowers), and high pollen-ovule ratios (Table 1). B. Ipomoea meyeri is autogamous and sister to /.

variabilis. Flowers are relatively small (<2.5 cm long), stamens and style subequal, and pollen-ovule ratios low (Table 1). C. Ipomoea tricolor (/. sect.

Tricolores) is xenogamous and shares a number of synapomorphic characteristics with the sister species, /. cardiophylla (sepals small, deltoid-elongate,

papillate, corolla tube yellow). Like other outcrossing Ipomoea, this species has relatively large and colorful flowers, heteromophic stamens and style

(herkogamous flowers), and relatively high pollen-ovule ratios (Table 1 ). D. Ipomoea cardiophylla is autogamous and exhibits flowers that are relatively

small (<3 mm long), the stamens and style subequal, and low pollen-ovule ratios (Table 1).

163

sequences of about 90% of all known autogamous species in this region of the world and sampled xenoga-

mous species that demonstrate phylogenetic affinities with inbreeding taxa on the basis of morphological

characters. We measured several floral characteristics that have long been associated with inbreeding species

in Ipomoea, including small corollas, subequal stamens and styles, and low pollen-ovule ratios, to examine

the distribution of such these characteristics in a phylogenetic context and to discern multiple origins of

these associative characteristics. The multiple and correlative origins of these characteristics are considered

in an evolutionary context.

Taxon Sampling

In order to obtain a broad sample of North American species and their close relatives, published ITS se-

quences of 50 species (47 Ipomoea and 3 outgroups) were acquired from GenBank (Manos et al. 2001; Miller

et al. 2004) and an additional 23 xenogamous species of Ipomoea that demonstrate close relationships with

moreeamg taxa on the basis of morphological synapomorphies, representing 21 species and 2 varieties,

were sequenced. As a whole, these species comprise a substantial complement of taxa that fall within two

subgenera of the genus (I. subg. Eriospermum and Quamoclit), representing 17 different sections and series

(Table 1). The former subgenus comprises a paraphyletic assemblage of clades that include both New and

Old World elements while the latter is monophyletic and consists predominantly of tropical North American

taxa. Voucher specimens for a total of 73 Convolvulaceae are listed in Appendix 1 along with their collection

localities and GenBank accession numbers.

We initially included samples from two or three populations of 25 species in our phylogenetic inves-

tigations, but duplicate accessions did not alter the topologies of trees from those including only a single

population of each taxon. We therefore include only one sequence for each taxon in our final results.

Because various Mexican species of Ipomoea occupy a basal phylogenetic position within the Ipomoeae

tribe (Miller et al. 1999, 2004), we employed three different outgroups of the sister tribe, Merremiaea

(Stefanovic et al. 2002): M erremia dissecta, Merremia tuberosa, and Operculina pteripes (Appendix 1).

For newly sequenced species, DNA was extracted from fresh leaf material grown in greenhouse facilities

at The University of Texas. Seeds were collected by the first author in the held or obtained from the USDA

Southern Regional Plant Introduction Station, Griffin, Georgia.

Whole genomic DNA was extracted by following a modified Doyle and Doyle (1987) protocol using a CTAB

extraction buffer with 0.2% (v:v) beta-mercaptoethanol. The internal transcribed spacer (ITS) region of the

18S-26S nuclear ribosomal repeat was chosen for phylogeny reconstruction. We employed primers ITS 1A

(5’GGA AGG AGA AGT CGT AAC AAG G 3’) and ITS 4^T&;fC4;§&TAT TGA TAT GC 3’) as primers

to amplify the ITS I, the 5.8s region, and ITS II via the polymerase chain reaction (PCR). Reaction volumes

of 25 ul included 2.5pL of lOx Triton-X PCR buffer (final concentration lx), 2pL lOmM dNTP mix (final

concentration 0.8 mM dNTPs), 2 pL 25mM MgCl (final concentration 2mM MgCl), DMSO (final

concentration 5% v:v), 1 unit of Taq enzyme, and 0.25 pL 20pM forward and reverse primers (0.2pM final

concentration of each primer). After initial denaturation at 95° C for 5 min, amplification proceeded at 94°

3 min, 46-52° C for 1 min, &}$$$* 1 min, followed by 35 cycles of 94° C % min, 46-52° C 1 min,

72° C for 45 sec + 3 sec/cycle, with a final 7 min extension at 72° C. Amplification products were visual-

ized on a 1.5% agarose gel stained with ethidium bromide and viewed with UV on a transilluminator. PCR

products were cleaned prior to sequencing with QIAquick spin columns (Qiagen Inc.) according to the

manufacturer s instructions.

Cycle sequencing was performed using Big Dye terminator chemistry and either the forward or reverse

amplification primer. Centri-sep columns were used to remove residual salts and unincorporated nucleo-

tides from the sequencing products. Automated sequencing was performed on a BaseStation sequencer

(MJ GeneWorks, San Francisco, CA). Forward and reverse sequence strands were assembled and edited

167

Ten measurements of flower length and anther-stigma distance (herkogamy) were collected from either

fresh or herbarium materials (TEX, UNAM). Pollen-ovule ratios for 51 of the 68 species were determined

by counting the total number of pollen grains in five anthers of each flower and then dividing this sum by

the total number of ovules (i.e., four in most Ipomoea, six for members of I. series Pharbitis; Table 1). When

live material was available, an average pollen-ovule ratio was measured by directly observing and count-

ing every pollen grain from a total of 10 flowers. Ratios obtained from herbarium specimens were based

on measurements of 1-3 flowers. The herbarium sheet approximations of pollen-ovule ratios should prove

reliable for the present survey, as formerly published measurements indicate pollen number within species

varies from 1-10% (McDonald 1982; Chemas Jaramillo & Bullock 2002) while pollen-ovule ratios between

sister autogamous and xenogamous species usually differ by factors that range from 100-2000% (Table 1).

We also collected data of all 68 species for three floral traits coded as discrete characters. Based on ob-

servation of living and herbarium materials at TEX, flowers with corolla tubes that exceed 2.5 cm in length

were scored as ‘large’ while those that range from 0.5-2. 5 cm in length were classified as ‘small’ (Table 1).

Species were deemed herkogamous if anthers and stigma are separated spatially (usually by several mil-

limeters).

We were not able to use methods currently available for predicting ancestral traits or for rigorous com-

parative analyses because we were able to sample only 41% of the species of North America and our sample

is not truly random. However, we did code the floral and reproductive characters in Table 1 (autogamy

versus xenogamy, herkogamy vs. homomorphic sexual structures, floral length, and pollen:ovule ratios) as

discrete characters and then optimized them on the tree shown in Figure 3 using MacClade 4.08 (Maddison

& Maddison 2005) under a parsimony criterion (Fig. 4A, B, Fig. 5 A, B).

Phylogeny estimation

The aligned matrix of ITS sequence data totaled 676 bp. Across these characters, 355 were variable and 221

were parsimony-informative. For the parsimony analysis, we excluded uninformative characters, leaving

221 characters analyzed. The parsimony analysis generated 120,950 equally most-parsimonious trees. The

consensus of these trees was similar to that obtained from the Bayesian analysis.

For Bayesian analysis, the best-fitting model of sequence evolution for this dataset was a general time-

reversible model with gamma-distributed rate variation and the number of invariant sites estimated. Both

independent runs produced similar consensus topologies, suggesting convergence on the most probable

area of tree space. The average standard deviation of split frequencies was less than 0.01 after the burn-in

phase (30%), also indicating good convergence. The resulting majority-rule consensus of 56,000 trees is

used (Fig. 3) for the assessment of the breeding system and floral morphology across the species.

Breeding System and Floral Trait Data

Greenhouse studies confirmed that Ipomoea species range from completely selling (ca. 100% viable seed set

in non-treated flowers) to completely outcrossing (0% viable seed set in non-treated flowers). Based on our

scoring methods for an ingroup of 87 taxa (Table 1), 59 species were scored as xenogamous, 23 as autoga-

mous, and 5 as having a mixed breeding system (self-compatible but facultatively herkogamous). Flowers of

69 taxa were scored as large and 18 as small. Distances between the anthers and stigma yielded 51 species

being scored as herkogamous and 38 as homogamous.

51-3177:1 (x = 597). High extremes of autogamous pollen-ovule ratios in I. amnicola (604:1) and I clavata

q|||||p:l) occasionally exceed the low extreme of pollen-ovule ratios in various xenogamous species (Table

1), such as those of I bracteata (405:1) and I simulans (591:1). In a similar fashion, several self-compatible,

outcrossing species that produce herkogamous flowers, such as I chamelana (P/O = 108), I. cordatotriloba (P/O

= 476), and I. hederifolia (P/O = 134), exhibit pollen-ovule ratios that fall within limits of many autogamous

species (Table 1).

169

The results of the character optimization are shown in Figures 4 and 5. These optimizations reveal

considerable congruency in character evolution. Herkogamy, long flowers and large pollen:ovule ratios usually

occur on outcrossing lineages while homomorphic sexual structures, short flowers and small pollen:ovule

ratios trace primarily along inbreeding clades.

The present study uses a new phylogenetic reconstruction for North American Ipomoea to explore breeding

systems and their floral traits. Our analyses indicate multiple (potentially 16) independent transitions to

autogamy and possibly reversals (Fig. 3). We found that apparent transitions between xenogamy and autogamy

were strongly associated with changes in a suite of floral traits including flower size, distance between the

anthers and stigma within a flower, and pollen:ovule ratio. Here we discuss the results in the context of

existing studies of breeding system evolution and suggest avenues for future work.

Although a few phylogenetic studies based on morphological data suggest xenogamy may be a derived

feature (Olmstead 1989; Armbruster 1993; Kelly 1997), almost all studies based on molecular evidence

indicate that inbreeding lineages are derived from, and rarely revert back to, outcrossing habits (Barrett et

al. 1996; Goodwills 1999; Weller & Sakai 1999; Vieira & Charlesworth 2002; Igic et al. 2003). Similarly,

our estimates based on a sample of 68 Ipomoea species suggest strong directionality in transitions, with

many shifts from xenogamy to autogamy and a few in the reverse direction (notably Ipomoea sers. Mina and

Pharbitis; Fig. 3).

Although most autogamous species appear as isolated tips in xenogamous clades, in a few cases autoga-

mous species are closely related to other autogamous species, as observed among members of Ipomoea ser.

Pharbitis (I. pubescens is sister to I. purpurea, I. nil is sister to I. hederacea), Ipomoea ser. Batatas (I. lacunosa, I.

cordatotriloba, I. ramosissima), and particularly species in Ipomoea sect. Mina. Apart from these exceptional

examples, however, autogamous lineages in Ipomoea appear to undergo cladogenesis less often than their

most closely related xenogamous lineages, this being consistent with the notion that transitions to selfing may

decrease speciation or increase extinction, leading to an evolutionary dead-end (Stebbins 1957; Takebayashi

and Morrell 2001). Substantiating this pattern across Ipomoea will require, however, a broader sampling of

taxa for statistical analyses, as discussed below.

The lack of species-rich clades in terms of autogamous taxa in Ipomoea compares closely with those

of the Polemoniaceae but contrast significantly with inbreeding lineages of the Solanaceae. Self-compatible

lineages have arisen at least 16 times in the Polemoniaceae and a large majority of these have failed to diver-

sify over the course of time (Barrett et al. 1996). In contrast, an estimated 60% of species in the Solanaceae

(ca. 1200 spp.) are identified as self-compatible, and these autogamous nightshades are often closely related

to one another, comprising clades of exclusively inbreeding taxa (Igic et al. 2003). This distinction is note-

worthy (and as yet unexplained in terms of their genetic architecture and evolutionary histories), as the

Convolvulaceae are sister to Solanaceae and phylogenetically distant to the Polemoniaceae (Savolainen et

al. 2000; Soltis et al. 2000).

Similar to evolutionary trends of autogamy in the Polemoniaceae (Barrett et al. 1996), selfing has arisen

in distantly related groups of Ipomoea clades that exhibit a variety of pollination syndromes: i.e., humming-

bird pollination (I. cholulensis, I. hederifolia, I. neei, I. quamoclit), bee pollination (I. amnicola, I. cardiophylla, I.

costellata, I. fimbriosepala, I. meyeri, I. leptophyla, I. microsepala, I. minutiflora, I. orizabensis, I. pubescens, etc.),

hawk-moth pollination (I. neurocephala, I. muricata), and long-tongued fly pollination (I. aristolochiaefolia,

Fig. 1; McDonald 1982, 1991). Consequently, there is little evidence that specific animal vectors predispose

or preclude a plant population’s ability to become autogamous.

Inbreeding species in Ipomoea commonly possess a suite of floral characteristics that apparently arise

convergently. These characteristics include relatively small (tubes usually < 2.5 cm long) and pale corol-

las, subequal stamens, anthers that make direct contact with the stigma (Fig. 1 E & F, Fig. 2 B & D), and

low pollen:ovule ratios. Similar floral syndromes typify autogamous elements in various genera of the

170

Journal of the Botanical Research Institute of Texas 5(1)

Polemoniaceae (Grant 1981; Barrett et al. 1996), Dalechampia (Armbruster 1993), Scutellaria, Mazus and Hosta

(Ushimaru & Nataka 2002), and often concomitant with a reduction in flower size in Amsinckia, Limnanthes,

and Epilobium (Ornduff & Crovello 1968; Olmstead 1989; Parker et al. 1995; Schoen et al. 1997).

Our results suggest that shifts between autogamy and xenogamy are strongly correlated with simul-

taneous changes in flower size, pollen-ovule ratio, and anther-stigma distance (Table 1, Fig. 4A, B, 5A, B).

Small autogamous flowers with anthers that touch stigmas and relatively low pollen-ovule ratios have arisen

independently in numerous unrelated clades, including those comprising Ipomoea ser. Batatas (I. triloba), I.

sect. Calonyction (I. muricata), I sect. Exogonium s. lat. (I. aristolochiaefolia, b dumetorum, and I meyeri; Fig.

1 F&E, Fig. 2 B), I. sect. M icrosepalae (I. minutiflora; Fig. 1 D), I. sect. Mina (I. hederifolia), I ser. Pharbitis (I.

neurocephala), I. sect. Tricolores (I. cardiophylla; Fig. 2 D), and I. sect. Jalapae (I. amnicola; see Table 1, McDonald

1982, 1991; Chemas Jaramillo & Bullock 2002). Such characteristics contrast with typical floral features

of xenogamous relatives, whose flowers are normally large, brightly pigmented, herkogamous (Fig. 1 A-C,

Fig. 2 A, C), and exhibit high pollen-ovule ratios (Table 1).

Pollen:ovule ratios can serve as a reliable indicator of breeding system. High pollen-ovule ratios are

normally associated with obligate outcrossing behaviors, moderate pollen-ovule ratios with facultatively

xenogamy, and low ratios with obligate autogamy (Cruden 1977, 2000; Lord & Eckhard 1984; Plitman &

Levin 1990). As noted earlier, pollen-ovule ratios in our broad sampling of both autogamous and xenogamous

taxa within Ipomoea varied considerably from 51-3100 grains per ovule (McDonald 1982; Chemas Jaramillo

& Bullock 2002, Erbar & Langlotz 2005), and therefore provide a model system to examine this variation

in a phylogenetic context. Most xenogamous taxa produced more than 400 pollen grains per ovule, roughly

2-14 times their autogamous relatives (Table 1). Lower pollen-ovule ratios (51-223:1) were invariably as-

sociated with autogamous relatives and lineages (Table 1; Fig. 3). While some autogamous species, such as I.

amnicola, I. clavata, and I.fimbriosepala in Ipomoea subg. Eriospermum, exhibited relatively high pollen-ovule

ratios (458-1172:1), these values are still 4-5 times lower than their closest xenogamous relatives [i.e., I.

leptophylla ;217o:l) and I. setifera (2971:1)]. Hence the quantitative relationship between pollen-ovule ratios

of outcrossing and inbreeding taxa is not measurably precise, but predictably disparate in relative terms

between closely related autogamous and xenogamous species.

While this study has focused on the evolution of floral traits associated with autogamy, other traits,

such as life histories, are likely to be associated with shifts in breeding system. For example, a considerable

portion of selling Ipomoea species are annuals and share close ancestry with a xenogamous and (usually)

perennial species (Fig. 3). Such associations are exemplified by the following species-pairs: I. costellata and

I. ternifolia ; I. muricata and I. santillanii; I. purpurea and I. pubescens; I. cardiophylla and I. tricolor (Fig. 2 C

& D); I. minutiflora and I. microsepala (Fig. ID); I. meyeri and I. variabilis (Fig. 2 A & B); I. dumetorum and I.

simulans/I. purga; I.fimbriosepala and I. pes-caprae, and I. amnicola and I. pandurata/I. leptophylla.. Exceptions

to this pattern include several selling perennials, such as I. amnicola, I. clavata, and I. fimbriosepala, which

belong to the woody and hairy seeded Ipomoea subgenus Eriospermum (sensu lato, Fig. 3; Table 1).

Additional research might consider the genetic and evolutionary mechanisms by which these autoga-

mous lineages have arisen from xenogamous ancestors. Self-compatibility probably arises independently

by unique point mutations at the S-locus (Kowyama et al. 2000), and the convergent evolution of floral

syndromes associated with inbreeding habits likely arises secondarily and independently on account of

shifts in selective forces on flower phenotypes. Without positive selection for floral features that attract and

reward animal vectors, selection will favor floral reductions that allow more resource allocation for growth

and seed production (Cruden & Miller-Ward 1981; Charnov 1982; Williams & Rouse 1990; Kirk 1992;

Barrett et al. 1996; Goodwillie 2009). Therefore, autogamous species tend to produce relative smaller flowers

that are less vivid in color (Richards 1997; Tate & Simpson 2004). The repeated evolution of the same suite

of floral characters in association with selfing presents the opportunity to examine how often parallel trait

evolution is caused by parallel genetic mechanisms.

Finally, Ipomoea offers a prime system for testing the effects of breeding system transitions on the fate

171

Fig. 4. Optimizations of breeding system and floral characters on the tree shown in Fig. 3. A. Breeding systems: xenogamy (X), autogamy (A), or mixed

(XA). B. Anther and stigma separation: herkogamy (He), Homogamy (Ho), or mixed (He, Ho).

of lineages. Recently developed models by Maddison et al. (2007) make it possible to test statistically the

effect of a binary character on rates of speciation and extinction. With such methods, one can determine

processes. Although these analyses will require more intensive taxon sampling, they hold the possibility of

HDD

172

Journal of the Botanical Research Institute of Texas 5(1)

to be strongest with respect herkogamy vs. homomorphic styles and stamens, while a relatively weaker cor-

example, a less consistent correspondence between relatively large corollas and inbreeding behaviors in I.

173

coccinea, 1. cholulensis ; Figs. 3, 5) can be attributed to the unique pollination syndromes of these two groups

present extraordinarily long and narrow corolla tubes. Alternatively, relatively less congruency is observed

in pollemovule ratios (Fig. 5) and breeding systems on account of our incomplete data set for pollen counts

and an exceedingly wide range of variation in this floral parameter. Nevertheless, character optimization

analyses generally reveal consistent phylogenetic correlations between autogamy, homomorphic sexual

structures and low pollemovule ratios.

APPENDIX 1

Voucher and DNA sample information on Ipomoea species examined for ITS sequence variation, including (in order): taxon

examined, DNA accession number, plant locality, collector, and GenBank accession number. McDonald vouchers are located

at XAL orTEX; Miller vouchers are maintained at Southeastern Louisiana University (SLU), Hammond, Louisiana. Germplasm

from SRPIS (USDA-Southern Regional Plant Introduction Station, Griffin, Georgia) provided fresh material. GenBank accession

numbers for new sequences are presented in boldface.

Outgroup. Merremia tuberosa (L.) Rendle, REM 1 9, Unknown, Miller 8 (SLE), AF1 1 0909; Merrmia dissecta (Jacq.) Hallier f., REM 384,

Chihuahua, Mexico, Miller 305, GQ388262; Operculina pteripes (G. Don) Odonell, REM 322, Jalisco, Mexico, Miller 365, GQ388263.

Ingroup. I. alba L, REM 1 29, Oaxaca, Mexico, McDonald AF538275; I. amnicola Morong., REM 3,SRPIS-5382.65,Texas,

USA, Miller 3 (SLE), AF1 1 0928; /. ampullacea Fernald, REM 124, Jalisco, Mexico, Lott 2362 (MEXU), AF$38277, / Of'US&ttH. Bouse j

REM 1 ji&a, Mexico, van Devender 97-1 263 (TEX), DQ355304; I. arborescens (H.B.K.) G.Qtoft,lEM 38, SBE Universal Seedbank,

Miller 84 (SLE), AF1 1 0924; I. aristolochiifolia G. Don, JAM 6, Jalisco, Mexico, McDonald 238 (TEX), DQ355309; I. batatas (L.) Lam.

var. batatas, REM 1 , SRPIS- 561558, Mexico, Miller 39 (SLE), AF1 1 0938; I. batatas (L.) Lam. var. apiculata (M. Martens & Galeotti)

J.A. McDonald & D.F. Austin, JAM 33, Veracruz, Mexico, McDonald 1949 (TEX), DQ355319; I. cardiophylla A. Gray, REM 1 34, New

Mexico, USA, McDonald 14 1 (SLE), AY538280; /. carnea Jacq. var. fistulosa (M. Mart, ex Choisy) D.F. Austin, B&T World Seeds - 1 472,

Miller 6 (SLE), AF 1 1 0920; /. chamelana J.A. McDonald, REM 1 53, Jalisco, Mexico, McDonald 1930 (SLE), AY538281 ; /. cholulensis

tfll’ifUAM 2, Veracruz, Mexico, McDonald 1886 (TEX), DQ355305; I. da vata v.Ooststr. ex Macbride, JAM 30, Veracruz, Mexico,

Mt§|rrain s.n. (TEX), DQ355325; I. coccinea L, REM 13, North Carolina, USA, Miller 47 (SLE), AF1 10941, / co/^|ff^reenm.,

REM 44, B&TWorld Seeds-32404, Mi»#fSLE), AF1 1 0927; I. cordatotriloba Dennstedt, REM 52, B&T World Seeds-7493 1 , Miller

73 (SLE), AF1 10939;/. costellataJorr., JAM 1, Sonora, Mexico, van Devender 20001 -859 (TEX), DQ335306; I. crinicalyx S. Moore,

WMC 8329, Jalisco, Mexico, Bullock 2000 (Kew), AF309164; I. dumetorum Willd. ex Roem. & Schult, REM 147, New Mexico,

USA, McDonald 140 (SLE), AF538284; /. expansa J.A. McDonald, REM 1 35, Guerrero, Mexico, McDonald 1910 (SLE), AY538285; /.

fimbriosepala Choisy, JAM 44, Veracruz, Mexico, Calzada 5985 (XAL), DQ35531 5; /. funis Schlecht. & Cham., REM 1 23, Guerrero,

Mexico, McDonald 1895 (SLE), AY538286; I. hastigera H.B.K., REM 1 39, Veracruz, Mexico, McDonald 1998 (SLE), AY538287; I. hart-

wegii Benth., JAM 1 3, Queretaro, Mexico, Carranza & Silva 6355 (TEX), DQ33531 3; I. hederacea Jacq., REM 1 63, Georgia,, USA,.

Miller 330 (SLE), AY538291 ; I. hederifolia L„ REM 1 83, Mexico, Miller 31 8 (SLE), AY538294; I. imperati (Vahl) Griseb., North Carolina,

USA, Miller 99 (SLE), AF1 1 091 7,hrtka(8iirn\.ij Merr., REM 1 68, Madagascar, Miller 256 (SLE), AY538297; l.jalapa (L.) Pursh, JAM

29,Tamaulipas, Mexico, McDonald 1288 (XAL), DQ33531 6; I. lacunosa L, JAM 20, Missouri, USA, Brant 4182 (TEX), DQ335324;

I. lindheimeri A. Gray, REM 25, SPRIS-55301 1 (Texas, USA), AF11Q944, h leptophyllalonM^, SRP1S-303327,

Miller 30 (SLE), AF1 1 0929; I. lobata (Cerv.) Thellung, REM 39, B &T World Seeds - 34004, Miller 7 (SLE), AF1 1 0940; I. lutea Hemsl.,

REM 141 , Chiapas, Mexico, McDonald 1994 (SLE), AF538299; I. matretu Choisy, REM 1 37, Chiapas, Mexico, McDonald 2024 (SLE),

AY538300; /. meyeri (Spreng.) G. Don, JAM 3, Jalisco, Mexico, McDonald 1939 (TEX), DQ33531 1; I. microsepala Benth., JAM 4,

Oaxaca, Mexico, Elorsa2801 (TEX), DQ335307; /. minutiflora (M. Mart. & Galeotti) House, JAM 5, Mexico, Mexico, McDonald

m (TEX), DQ335308; I. muricata Cav. REM 144, Jalisco, Mexico, McDonald 1940 (SLE), AY538332; I. neei (Spr.) O'Donell, REM

1 40, Jalisco, Mexico, Bullock &Martijena 2098 (SLE), AF538302; /. neurocephala Hallier, REM 145, Michoacan, Mexico, McDonald

1963 (SLE), AY538303 ;L nil( L.) Roth, REM 1 95, New Mexico, USA, Miller 2^pj| AY538308, / orizabensis (Pell.) Led. exStdud'L

var. orizabensis, REM 142 (SLE), Veracruz, Mexico, McDonald 2434, AY538309; /. orizabensis (Pell.) Led. ex Steudl. var. collina (F§

H. House) J.A. McDonald, JAM 12, Nuevo Leon, Mexico, Hinton 23668 (TEX), DQ335303; /. pandurata (L.) G. Mey., REM 48,

Durham Co., North Carolina, Miller 98 (SLE), AF1 10930; I. parasitica (H.B.K.) G. Don, REM 1 79, Morelos, Mexico, Miller 333 (SLE),

AY538313; I. pauciflora M. Mart. & Galeotti, JAM 25, Puebla, Mexico, McDonald 2010 (XAL), DQ335314; /. pedicellaris Benth.,

REM 97, Mexico, Miller 185 (SLE), AF309165; I. pes-caprae L, REM 8, Kew Botanical Garden (Mali), Miller 51 (SLE), AF1 10932; /.

pubescens Lam., REM 76, Mexico, Miller 1 13 (SLE), AF538314; /. purga (Wender.) Hayne, REM 1 33, Veracruz, Mexico, Linajes s.n.

(SLE), AY53831 5; I. purpurea (L.) Roth, REM 1 TTi'NOrt^Wna, USA, Miller 254 (SLE), AY538322; /. quamoclitL, REM 1 67, Surinam,

Miller 181 (SLE), AF538323; I. ramosissima Choisy, JAM 56, Guanacaste, Costa Rica, Moraga 604 (MO), DQ335323; I. santillanii

O'Donell, REM 1 38, Veracruz, Mexico, McDonald 1946 (SLE), AY538324; I. seducta House, REM 1 46, Chiapas, Mexico, McDonald

1986{ SLE), AY538325; l. sepacuitensis Donn. Sm„ JAM 61 , Chiapas, Mexico, £ Martinez S. 16636 (TEX), DQ33531 7; /. sescossiana

Bail Ion, REM 1 43, Zacatecas, McDonald s.n., AY538326; I. setosa Ker Gawl., JAM 47, SRPIS 61 62, Griffin 6162, DQ33531 8; I. simulans

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s of plastid atpB and n

PUTATIVE MORPHOLOGICAL SYNAPOMORPHIES OF SAXIFRAGALES

AND THEIR MAJOR SUBCLADES

Barbara S. Carlsward

Department of Biological Sciences

Eastern Illinois University

Charleston, Illinois 61920, USA.

bscarlsward@eiu.edu

Walter S. Judd and Douglas E. Soltis

Department of Biology

University of Florida

Gainesville, Florida, 32611, USA.

wjudd@botany. ufl. edu; dsoltis@botany. ufl. edu

Steven Manchester and Pamela S. Soltis

Florida Museum of Natural History

University of Florida

Gainesville, Florida 32611, USA.

steven@flmnh.ufl.edu; psoltis@flmnh.ufl.edu

RESUMEN

As part of the Angiosperm AToL project, which resulted in a phylogenetic analysis of 640 species representing

330 families based on 25,260 aligned base pairs from 17 genes, representing all three plant genomes (Soltis

et al., 2011), one goal was to assess morphological synapomorphies for each of the major clades of angio-

sperms. We present here a morphological exploration of Saxifragales, which represents the first treatment

in a series of analyses (Judd et al., unpublished data) that will address morphological synapomorphies and

the clades they support. Our data almost exclusively come from original observations taken directly from

herbarium specimens representing the same species used in the molecular analyses and allow us to assess

the morphological diagnosability of an array of large angiosperm clades, such as Saxifragales, based on easily

J. Bot. Res. Inst. Texas 5(1): 179-1

Journal of the Botanical Research Institute of Texas 5(1)

acquired morphological characters. This study, therefore, provides an example and a test for methods that

we will soon apply across our large sampling of angiosperms.

This is the first analysis to explicitly assess morphological traits and potential synapomorphies across

Saxifragales, which only recently has been recognized in its current circumscription, and represents an

ancient and rapid diversification within the angiosperms (Angiosperm Phylogeny Group 2009; Fishbein

et al. 2001; Jian et al. 2008; Kubitzki 2007a; Soltis et al. 2000, 2005, 2007). We note, however, that the

phylogenetic analyses of Hermsen et al. (2006) employed both morphological and molecular characters,

but these were not explicitly mapped onto the tree, since the focus of their paper was on the placement of

fossil taxa.

MATERIALS & METHODS

Morphological characters and character states (Tables 1, 2) were determined from herbarium material at

FLAS and MO, supplemented by reports from the literature. Most observations were made using a Wild

were also made using an Hitachi S-4000 scanning electron microscope (SEM) at a running voltage of 5-7

kV. Pollen and stamens used for SEM were taken directly from herbarium sheets, mounted on aluminum

stubs with adhesive graphite tabs, and coated with a gold-palladium alloy for 60 s. Metadata were input into

TOLKIN (Beaman & Cellinese 2010}!’

Sixty-three morphological features (Table 1) were used to construct a character matrix (Table 2) in

MacClade v.4.08 for Mac OS X (Maddison & Maddison 2000). Characters then were mapped using par-

simony (Brokaw & Hufford 2010; Soltis et al. 2005) onto the cladogram of Saxifragales published by Jian

et al. (2008; see their Fig. 1) using particular species of representative genera as terminals (Fig. 1; Table

3). This topology differs in a few minor ways from that recovered in the 17-gene analysis of angiosperms

(Soltis et al., 2011). We used only Liquidambar L. to represent Altingiaceae because Liquidambar and Altingia

Noronha are not reciprocally monophyletic, and only Peridiscus Benth. represented Peridiscaceae. In addi-

tion, we took into account the morphological character states seen in Crassula (Crassulaceae) and Pterostemon

Schauer (Pterostemonaceae; sister to Iteaceae) using the phylogenetic placements of those two supported

in the analyses of Jian et al. (2008). Vitaceae (represented by Leea Royen ex L. and Vitis L.) were used as an

outgroup, and we note that the polarity of a few characters is equivocal.

Observations of wood anatomy were taken from the Inside Wood online database (http://insidewood.

lib.ncsu.edu/). Vitaceae wood was so autapomorphic, due to its liana habit, that Peridiscus was used as a

functional outgroup for the anatomy of the rest of the Saxifragales.

Potential synapomorphies for Saxifragales include basifixed anthers, with the filament attached at a basal

several groups (Fig. 2A-C; char. #42-1), but reversals to the dorsihxed condition (Fig. 2D-E) have

occurred, and under DELTRAN optimization would be placed on the lineage leading to Ribes L., and at

the common ancestor of the Pterostemon + Iteaceae clade (see also Fig. 1). Additionally, the presence of

violoid to theoid teeth (char. #18-1) is possibly synapomorphic for Saxifragales, but salicoid teeth occur in

Aphanopetalum Endl., and non-glandular teeth occur in Hamamelis L., Itea L., and Haloragis Forst. & Forst.

f., while the condition cannot be assessed in taxa with entire leaves. All but Peridiscaceae are united by fol-

licular fruits (char. #60-2) but this feature shows extensive homoplasy and is correlated with ovaries being

at least distally distinct. All but Peridiscaceae also are united by latrorse anther dehiscence (Fig. 2B; char.

#44-2, when ACCTRAN optimization is assumed), but this character also is quite homoplasious.

The woody clade (i.e., Altingiaceae, Cercidiphyllaceae, Daphniphyllaceae, and Hamamelidaceae; see

Soltis et al. 2005) can be diagnosed by the flattened adaxial surface of the petiole (Fig. 3A; char. #14-1,

but a grooved petiole occurs in Cercidiphyllum Sieb. & Zucc., and adaxially flattened petioles have evolved

in parallel in Choristylis Harv. and Kalanchoe Adans.), polytelic or racemose inflorescences (char. #21-1,

also evolving in parallel in Peridiscus, Ribes L., Itea, Myriophyllum L., and Tetracarpaea Hook.), and the loss

182

Journal of the Botanical Research Institute of Texas 5(1)

Table! continued

49. Pollen aperture distribution and symmetry: triaperturate (0); panaperturate (1); many random pores (2);

one pore at each pole of a bilateral grain (3)

50. Pollen aperture type: colporate (0); colpatp'(l); porate (2)

5! Pollen grain ornamentation ^ jBffll jte (0), smooth/psilate (1); rugulate (2); spinulate, even (3); spinulate,

uneven (4); scabrate (5); striate (6)

52. Tectum organization: puncta^^feontinuous (1); interrupted or incomplete (2); reticulate (3)

53. Aperture membrane: rugose to spinulose (0); smooth (1)

55. Carpel number: two (0); one p| 8w|(2): three (3); four (4)

56. Ovary position: superior (0); inferior (1); partially inferior (2)

57. Stigma: minute, capitate or punctate (0); decurrent along side of style (1); broad and globose-capitate (2)

58. Placentation: axile or basal-axile (0); parietal (1 ); apical-axile (2)

59. per focule two (0); or^^yi|umerous/more than two (2)

60. Fruit type: berry (2); drupe (3); capsule (4); dry-drupaceous schizocarp (5)

’ SI", ’-Fruit attachment: to receptacle (0); to gynotihpf#|T )

62. Persistent styles on fruit: absent (0); present (1 )

63. Winged seeds: absent (0); present (1 )

of floral nectaries (char. #28-2, a feature that also characterizes Haloragis, Myriophyllum, Penthorum, and

Tetracarpaea, while a nectary does occur - possibly representing a reversal - in the Hamamelidaceae, Endress

1989a). Although not included as a formal part of our analysis, sessile flowers are apparently an additional

synapomorphy of the woody clade. Most members of the woody clade share anthers opening by valves/

flaps (Fig. 3B; char. #43-1), including Hamamelidaceae and Cercidiphyllum. Stamen appendages (Fig. 3B;

char. 39-1) unite Hamamelidaceae with Cercidiphyllum and Daphniphyllum (a character also found in the core

Saxifragales). The woody clade is united with Paeonia F. on the basis of stigmas decurrent along the side

of the style (Figs. 3C & 4; char. #57-1), although this stigmatic condition has been lost in Rhodoleia (where

it is apical, minute and truncate). Hamamelidaceae, represented by Corylopsis Sieb. & Zucc., Hamamelis L.,

Exbucklandia , and Rhodoleia, are united by the apomorphy of stipules expanded and surrounding the api-

cal bud (char. #9-1, but character reversed in Rhodoleia) and styles persistent in fruit (Fig. 5A; char. #62-1,

under DELTRAN optimization, and occurring also in Liquidambar and Altingia). Within Hamamelidaceae,

stellate hairs unite Corylopsis and Hamamelis (under DEFTRAN optimization, as they also occur in the related

Rhodoleia), while entire-margined leaves (char. #17-1) are a potential synapomorphy for Exbucklandia and

Rhodoleia. Cercidiphyllum and Daphniphyllum share imperfect flowers (char. #26-1, a condition also present

in Liquidambar and Myriophyllum and reported to be present in other Hamamelidaceae not sampled here but

in Magallon 2007). Bisporangiate anthers (Fig. 3B; char. #45-1) appear to be an autapomorphy of Hamamelis

and of Exbucklandia.

A synapomorphy of Core Saxifragales (i.e., all members of the order except Perdiscaceae and the woody

clade + Paeoniaceae - see Endress 1967) is the presence of a hypanthium in their flowers (Fig. 6; char. #27-

1, under ACCTRAN optimization) and this feature characterizes all representatives of core Saxifragales

included in the analysis except for Crassulaceae. The Saxifragaceae + Ribes + Pterostemon + Iteaceae clade is

supported by the presence of a nectary on the inner surface of the hypanthium (Fig. 7; char. #28-1), while

the Ribes + Saxifragaceae clade is supported by the lack of a pulvinus (char. #13-1, but very homoplasious,

evolving also elsewhere in cladogram, such as in Haloragaceae and relatives) and the petiole merely flattened

above (char. #14-1, very homoplasious, also evolving in Hamamelidaceae and relatives, as outlined above).

Saxifragaceae, as represented by Heuchera L. and Saxifraga L., are united by the herbaceous habit with stems

rooting at the nodes (chars. #1-1 and 2-1, but very homoplasious, evolving several times in parallel, see Fig. II

Journal of the Botanical Research Institute of Texas 5(1)

48-1, 49-1, 50-2, 51-3, 53-1, 56-1, 57-2, 58-2, 59-1; oo = 3-2, 19-2, 22-1, 30-0, 52-0, 55-2, 60-4; pp = 5-1, 6-3, 8-1, 12-1,1

10-1, 19-1, 20-0; rr = 27-1, 30-0, 34-1, 50-1; ss = 18-0, 28-4, 32-4, 44-0, 45-1, 54-1.

,41-1,47-2,

186

Journal of the Botanical Research Institute of Texas 5(1)

Fig. 2. Synapomorphies for Saxifragales. A-C. Stamens showing basifixed anther (char. #42-1) with filament attached at a basal pit. A. Tetracarpaea

tasmanica closed stamen, scale bar = 300 pm. B. Open stamen of Paeonia lactiflora also showing latrorse dehiscence (char. #44-2), scale bar = 857 pm.

C. Androecium of Penthorum sedoides with open and closed stamens, scale bar = 500 pm. D-E. Stamens showing dorsifixed condition (char. #42-0).

D. Open Itea virginiana stamen (foreground) showing unicellular, simple filament hairs (char. #46-1), apically expanded connective (char. #39-1), and

(background) globose-capitate stigma (char. #57-2), scale bar = 500 pm. E. Open stamen of Pterostemon rotundifolius showing stamen appendage

formed from apically expanded connective (pointing down, char. #39-1) and unicellular, simple filament hairs (char. #46-1), scale bar = 1.0 mm.

the outer perianth whorl slightly connate (char. #33-0, but also occurring in Daphniphyllum, Rhodoleia,

Kalanchoe, and Crassula L.), and oblate pollen grains (char. #47-2, but also occurring in Daphniphyllum,

Crassula, Kalanchoe, and Haloragaceae). The Pterostemon + Itea + Choristylis clade is supported by the

presence of stamen appendages (Fig. 2D-E; char. #39-1), which develop from the anther connective, but

appendages have also evolved in Cercidiphyllum, Daphniphyllum, and Hamamelidaceae, and by staminal fila-

187

(char. #57-1), scale bar = 500 (im.

ments with unicellular hairs (Fig. 2D-E; char. #46-1); other possible synapomorphies of this clade are the

capsular fruits (char. #60-4) and expanded, capitate stigmas (Fig. 2D; char. #57-2). Iteaceae (represented

by Itea and Choristylis; probably better treated as a single genus; see Kubitzki, 2007b) share the following

putative apomorphies: simple hairs (char. #19-1, under DELTRAN optimization, but quite homoplasious,

evolving also in Haloragis, Cercidiphyllum, Liquidambar, and Exbucklandia), flowers with open sepals and

valvate petals (char. #32-2), and distinctive pollen grains that are bilateral, with only two apertures, one

at each pole, with smooth aperture membranes (Fig. 8A; chars. #46-1, 47-1, 48-1, 49-1, 53-1). A putative

synapomorphy of the Crassulaceae + Aphanopetalum + Tetracarpaea + Penthorum + Haloragaceae clade is the

lack of stipules (char. #8-1, but only under ACCTRAN optimization, and with reversals in Myriophyllum

and Aphanopetalum ); but we note that stipules have been lost in parallel elsewhere in the cladogram. Other

possible synapomorphies of this large and diverse group are striate pollen (Fig. 8B; char. #51-5, although

the pollen of Tetracarpaea is smooth, and that of Haloragis and Myriophyllum spinulate), four carpels (char.

#55-4, although both Penthorum and Crassula have hve-carpellate flowers), and the stem frequently rooting

at the nodes (char. #2-1, quite homoplasious; see Fig. 1). Crassulaceae are well supported as monophyletic

by their succulent leaves (char. #7-1) and the presence of a nectar gland opposite each carpel (Figs. 7 & 8C;

char. #28-3). The Tetracarpaea + Penthorum + Myriophyllum + Haloragis clade (i.e., Haloragaceae along with

closely related families Penthoraceae and Tetracarpaeaceae, although some have suggested that these three

188 Journal of the Botanical Research Institute of Texas 5(1)

2, a broad and globose-capitate stigma (see Fig. 2D).

= 1.5 mm. B. Flower of Rhodoleia

championii showing nectar disk (char. #28-1 ), scale bar = 1 .

families could be combined into Haloragaceae si; see Soltis et al. 2005, Jian et al. 2008) are supported

by the lack of a pulvinus (char. #13-0) and the absence of nectaries (char. #28-2), but these characters are

quite homoplasious. The Penthorum + Myriophyllum + Haloragis clade is supported by the herbaceous habit

(char. #1-1) and nonglandular teeth (char. #18-0), while Haloragaceae, as represented by Myriophyllum and

Haloragis, are united by their short filaments in relation to the anther length (Fig. 8D; char. #41-1), spinulose

pollen (Figs. 6 & 8E; char. #51-3), globose-capitate stigma (char. #57-2), apical-axile placentation (char.

#58-2), and single ovule/locule (char. #59-1).

DISCUSSION

We have identified potential morphological synapomorphies for Saxifragales, including basihxed anthers,

often with the filament attached at a basal pit, and violoid to theoid leaf teeth. Stevens (2001) also has sug-

gested that basihxed anthers with a basal pit represent a synapomorphy for members of Saxifragales, however,

Endress (1989b; Endress & Stumpf 1991) has noted basal pits only in Aphanopetalum, Saxifragaceae, and

Crassulaceae. Although chemical, seed-anatomical, and micromorphological characters were outside the

scope of our investigation, Stevens also notes the following as potentially apomorphic of the order: presence

of ellagic acid, myricetin, flavonols, epicuticular waxes as clustered tubules, and more or less exotestal seeds.

He considers the presence of free carpels, or at least apically free carpels, as an ordinal synapomorphy, but

our analysis placed this character as a synapomorphy for the clade containing all members of the order

190

Journal of the Botanical Research Institute of Texas 5(1)

(state 0), and black lines represent the presence of a hypanthium (state 1).

Fig. 7. Character state reconstruction for nectary presence and form (char. #28). White lines represent the plesiomorphic state of nectaries present as a

disk (state 0); hatched lines represent nectaries present on or near inner surface of hypanthium (state 1); grey lines represent a loss of nectaries (state

2); black lines represent nectaries present as glands opposite each carpel, pictured in Fig. 8C (state 3); and the striped line represents staminodial

Journal of the Botanical Research Institute of Texas 5(1)

#51-5), scale bar = 4.29 |jm. C. Crassula flower showing nectar glands opposite each carpel (char. #28-3), scale bar = 1 mm. D-E, Synapomorphies of

Haloragaceae. D. Haloragis aspera stamen showing short filaments relative to anther length (char. #41-1), scale bar = 231 pm. E. Close up of exine

showing spinulose pattern, in pollen grains oWyriophyllum exalbescens (char. #51-3), scale bar = 3.0 pm.

193

except Peridiscaceae. Kubitzki (2007a, p. 17) also considered apocarpy derived in Saxifragales, stating that

“it is difficult to imagine that this character expression should be plesiomorphic in these groups.”

Our analyses also elucidate character evolution within the clade, a topic briefly addressed by Soltis et

al. (2005), although their selection of taxa, cladogram topology, as well as characters considered differed

somewhat from ours. Comparisons with our results, therefore, are limited. They hypothesized an ancestral

bicarpellate condition in Saxifragales and suggested that only two increases in carpel number have oc-

curred, while our data suggest one increase: in the common ancestor of the Crassulaceae + Haloragaceae +

Aphnopetalum + Tetracarpaea + Penthorum clade. Hufford and Endress (1989) suggested that valvate anther

dehiscence is plesiomorphic at the level of Hamamelidaceae. Our analysis supports this conclusion, while

Soltis et al. (2005) suggested that valvate anthers evolved within the Hamamelidaceae. While our analyses

only found imperfect flowers in the clade formed by Cercidiphyllum and Daphniphyllum, there are representa-

tives of Hamamelidaceae reported with this condition as well (Endress & Igersheim 1999; Magallon 2007).

It is of interest that Hermsen et al. (2006, p. 141), who included 44 morphological characters (several

similar to ours) in a phylogenetic analysis of Saxifragales, reported that the Saxifragales “as well as most

major infraordinal clades lack unambiguous morphological synapomorphies,” however, we note that even

homoplasious synapomorphies can be informative. Hermsen and associates included no figures in which

morphological characters were mapped onto their tree topologies, but they did mention a few synapomorphies

for infraordinal clades, i.e., the presence of a hypanthial nectary in the Saxifragaceae + Ribes + Pterostemon +

Iteaceae clade (Fig. 7; our character #28-1), and the presence of sessile flowers and palmate venation in the

Hamamelidaceae + Liquidambar + Cercidiphyllum clade. We note that their tree topology differed from ours

in that Daphniphyllum was placed outside this clade. Their hypothesized synapomorphies are in agreement

with ours, except that we interpret palmate venation as a retained ancestral character in these taxa. Finally,

their results agree with ours in hypothesizing 2-aperturate and porate pollen as synapomorphies for the

Itea + Choristylis clade (i.e., Iteaceae).

Not surprisingly, given the confused taxonomic history of Saxifragales, and reconstruction of phylogeny

relying exclusively on molecular characters, clear morphological synapomorphies for major subclades within

the order are few. Nonetheless we did discover likely synapomorphies for both the woody clade and Core

Saxifragales, as circumscribed in Jian et al. (2008) and Soltis and associates (2011), as well as less inclu-

sive groupings. As might be expected, clades traditionally recognized at familial rank, and represented by

more than one taxon in this analysis, usually possess unambiguous shared apomorphies, and these match

traditional diagnostic characters, e.g., capsules with persistent styles in Hamamelidaceae, the succulent

leaves and distinctive nectar glands of Crassulaceae, the stamens with short filaments, spinulose pollen,

and distinctive gynoecium of the flowers of Haloragaceae, or the unusual pollen of Iteaceae. Importantly,

phylogenetically useful morphological characters occur across all plant parts (e.g., vegetative morphology,

floral and fruit morphology, pollen structure) and can often be correlated with ecologically important syn-

dromes, such as pollination by animals in clades with capitate stigmas, e.g., in our Pterostemon + Iteaceae

and in Hamamelidaceae (Endress 1989b) or wind pollination in the woody clade where most members have

at best a poorly differentiated perianth and are either completely or at least partly wind pollinated (Endress

2010 ).

The characteristic of bisporangiate anthers within Hamamelidaceae found in Exbucklandia and Hamamelis

appears to be an autapomorphic feature for each genus, and is thus not homologous. Developmental data from

Magallon (2007) show that the loss of sporangia involves the reduction of dorsal sporangia in Exbucklandia,

while in Hamamelis it involves the reduction of ventral ones. Also within Hamamelidaceae, while the species

of Corylopsis we used (i.e., C. pauciflora) was haplostemous, there are diplostemous examples in the genus

(Endress 1989a). Endress (1989a) also suggested that Rhodoleia is diplostemous, but our observations of R.

championu suggest that it is haplostemous.

The clade united by basifixed anthers without basal pits that includes Altingiaceae, Daphniphyllaceae,

Cercidiphyllaceae , and Hamamelidaceae share a suite of common wood anatomical characters including

195

Table 3. continued

Family

Pterostemonaceae

Saxifragaceae

Tetracarpaeaceae

Vitaceae

Pterostemon rotundifolius Ramirez

Pterostemon rotundifolius

Heuchera micrantha Douglas ex Lindl.

Heuchera micrantha

Saxifraga virginiensis Michx.

Saxifraga virginiensis

Tetracarpaea tasmanica

Leea manillensis Walp.

Vitis aestivalis Michx.

Vitis aestivalis

MO (3417656)

MO (4028404)

’^§ 155609 )

FUS (125070)

29997)

MO (2314048)

MO (4895512)

FLAS (188082)

FLAS (120160)

FLAS (26062)

FLAS (68520)

FLAS (154089)

FLAS (154090)

D. Nava, F. Nava &F. Castillo 43

P. Tenorio 1 1302

LS. Rose 41 289

S. C. Hood 1040

D. Soltis 807

W.C. Brumbach 5 1-32< *

DA. Ratkowsky & A.V. Ratkowsky21

1 Bradford, R. Barnes & K. Bradford 898

J.R. Abbott 7323

G. Joyner s.n.

K.W. Loucks23

iJgpd 3980

L.M. Baltzell 7fjfP

vessels predominately (>90%) solitary, intervascular pitting opposite to scalariform, perforation plates exclu-

sively scalariform usually with more than 20 bars, fibers non-septate and with distinctly bordered pits, and

mostly narrow rays 1-3 seriate (based on information available in the “Inside wood” database and imagery).

These features (except for vessels predominately solitary) are also shared with Peridiscus. Therefore, none of

these wood anatomical features can be considered as synapomorphies of the woody clade. It is noteworthy

that vessels with simple perforations characterize most herbaceous members of Saxifragales, apparently as

a result of parallel evolutionary events (Kubitski 2007a).

Finally, it is significant that even this comparatively straightforward morphological survey (using char-

acters easily recorded from herbarium specimens) generated useful information, allowing morphological

characterization of nearly all clades recovered in molecular analyses. This applies even in a group such as

Saxifragales, which likely represent an ancient, rapid diversification and are thus relatively character poor

(Jian et al. 2008; Hermsen et al. 2006). We anticipate, therefore, useful results upon completion of a com-

parable morphological survey across the breadth of our large angiosperm cladogram, as well as comparable

analyses of Caryophyllales and the Campanulidae (see Cantino et al. 2007).

ACKNOWLEDGMENTS

This study was carried out as part of the Angiosperm Tree of Life Project (NSF EF-0431266). The authors

thank the University of Florida Herbarium and the Missouri Botanical Garden for providing samples used

in this study and Kent Perkins for assistance in processing specimen loans. We thank Peter Endress for his

insightful and very useful comments in reviewing our manuscript and two anonymous reviewers for their

comments. We would also like the thank Karen Kelley, of the Interdisciplinary Center for Biotechnology

Research at the University of Florida for her help with the scanning electron microscope.

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the orders and families of flowering plants: APG J I t 1 ’■

Beaman, R.S., and N.QaLpfeSE. 2010.TOLKIN:TheTree of Life Knowledge and Information Network. Website: http://

www.tolkin.org/.

Brokaw, J.M. and L. Hufford. 2010. Phylogeny, introgression, and character evolution of diploid species in Mentzelia

section Trachyphytum (Loasaceae). Syst. Bot. 35:601 617.

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Cantino, P.D., J.A. Doyle, S.W. Gra *y,W$ Xr* vfljG Olmstead, D.E. Soltis, P.S. Soltis, and M.J. Donoghue. 2007. Towards

a phylogenetic nomenclature of Tracheophyta. Taxon 56:822-846; E1-E44.

Fishbein, M., C. Hibsch-Jetter, D.E. Soltis, and L. Hufford. 2001. Phylogeny of Saxifragales (angiosperms, eudicots):

analysis of a rapid, ancient radiation. Syst. Bot. 50:81 7-847.

Endress, PK. 1967. Systematische Studie ^&|Jie verwandtschaftlichen Beziehungen zwischen den

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Endress, P.K. 1 989a. Aspects of evolutionary differentiation of the Hamamelidaceae and the Lower Hamamelididae.

PI. Syst. Evol. 162:493-211.

Endress, P.K. 1989b. Phylogenetic relationships in the Hamamelidoideae. In: P.R. Crane and S. Blackmore, eds.

, |||j, systematics, and fossil history of the Elamamelidae, 1: Introduction and 'lower' Hamamelidae.

Clarendon Press, Oxford. Pp. 227-248.

Endress, P.K. and A. Igersheim. 1999. Gynoecium diversity and systematics of the basal eudicots. Bot. J. Linn. Soc.

130:305-393.

Endress, P.K. and S. Stumpf. 1 991 .The diversity of stamen structures in Lower Rosidae (Rosales, Fabales, Proteales,

Sapindales). Bot. J. Linn. Soc. 1 07:21 7-293.

Endress, P.K. 201 0. Flower structure and trends of evolution in eudicots and their major subclades. Ann, Missouri

Bot. Gard. 97:541-583.

Hermsen, E.J., K.C. Nixon, and W.L. Crepet. 2006. The impact of extinct taxa on understanding the early evolution

of angiosperm clades: an example incorporating fossil reproductive structures of Saxifragales. PI. Syst. Evol|.

260:141-169.

Hiepko, P. 1 965. Vergleichend-morphologische und entwicklungsgeschichtliche Untersuchungen uber das

Perianth bei den Polycarpae. Bot. Jahrb. Syst. 84:359-508.

AN© P.K. Endress. 1989. The diversity of anther structures and dehiscence patterns among

Hamamelidaceae. Bot. J. Linn. Soc. 99:301 -346.

Jian, S„ P.S. Soltis, M.A. Gitzendanner, MJ. Moore, R. Li,T.A. Hendry, Y.-L. Qiu, A. Dhingra, C.(|||i§ and D.E. Soltis. 2008.

Resolving an ancient, rapid radiation in Saxifragales. Syst. Bio. 57:38-57.

Kubitzki, K. 2007a. Introduction to the groups treated in this volume. In: K. Kubitzki, ed.The families and genera

of vascular plants. Vol. 9. Flowering plants - Eudicots. Berberidopsidales, Buxales, Crossosomatales, Fabales

p.p., Geraniales, Gunnerales, Myrtales p.p., Proteales, Saxifragales, Vitales, Zygophyllales, Clusiaceae alliance,

Passifloraceae alliance, Dilleniaceae, Huaceae, Picramniaceae, Sabiaceae. Springer Verlag, Berlin. Pp. 1-22.

Kubitzki, K. 2007b. Iteaceae. In: K. Kubitzki, ed.The families and genera of vascular plants. Vol. 9. Flowering plants

- Eudicots. Berberidopsidales, Buxales, Crossosomatales, Fabales p.p., Geraniales, Gunnerales, Myrtales p.p.,

Proteales, Saxifragales, Vitales, Zygophyllales, Clusiaceae alliance, Passifloraceae alliance, Dilleniaceae, Huaceae,

Picramniaceae, Sabiaceae. Springer Verlag, Berlin. Pp ’ - , ' 1 ,

Maddison, D.R. and W.P Maddison. 2000. MacClade 4: Analysis of phylogeny and character evolution. Version 4.08

Sinauer Associates, Sunderland, Massachusetts.

Magallon, i. ;||pr. From fossils to motecgles: phylogeny and the core eudicot floral groundplan in

Hamamelidoideae (Hamamelidaceae, Saxifragales). Syst. Bot. 32:31 7-347.

Soltis, D.E., J.W. Clayton, C.C. Davis, M.A. Gitzendanner, M. Cheek, V. Savolainen, A.M. Amorim, and P.S. Soltis. 2007.

Monophylyand relationships of the enigmatic family Peridiscaceae. Taxon 56:65-73.

Soltis, D.E. (and 1 5 others). 2000. Angiosperm phylogeny inferred from 1 8S rDNA, rbcL, and atpB sequences. Bot.

Soltis, D.E., F|§fl||tis, P.K. Endress, and M.W. Chase. 2005. Phylogeny and evolution of the angiosperms. Sinauer

Associates, Sunderland, Massachusetts.

Soltis, D.E. (and 25 others). 201 1 . Angiosperm phylogeny: 1 7-genes, 640 taxa. Amer. J. Bot. 98:704-730.

Stevens, P.F. 2001 onwards. Angiosperm Phylogeny Website. Version 9, June 2008 [and more or less continuously

updated since], http://www.mobot.org/MOBOT/ resea rch/APweb/.

SPIRODELA 0L1G0RRH1ZA (LEMNACEAE) IS THE CORRECT NAME

FOR THE LESSER GREATER DUCKWEED

Daniel B. Ward

Department of Botany

University of Florida

Gainesville, Florida 326 |«pl$

INTRODUCTION

The family Lemnaceae consists of four genera: Lemna, Spirodela, Wolffia, and Wolffiella. (A fifth genus,

Landoltia, has been proposed, and will be discussed here.) All species are diminutive thalloid plantlets,

floating or slightly submersed on quiet waters. Each plantlet of Spirodela produces several roots, while the

genus Lemna is characterized by each plantlet producing only a single root. (The two remaining genera,

Wolffia and Wolffiella, are without roots.) Spirodela, in turn, is formed of 3 species (or 2, if Landoltia is split

off): S. polyrrhiza (L.) Schleiden, whose name is uncontested; a second, known as either S. oligorrhiza (Kurz)

Hegelmaier or S. punctata (Meyer) Thompson (or Landoltia punctata (Meyer) Les & Crawford); and a third,

presently known as S. intermedia W. Koch.

The original ranges of these three species are largely discrete. As based on an extensive compilation of

herbarium records (Landolt 1986), Spirodela polyrrhiza occurs throughout much of North America, Europe,

and temperate Asia. The second species with the conflicted nomenclature (at times cited here as the “Lesser

Greater Duckweed”) is common in eastern Australia, southeastern Asia, and throughout the islands of the

western Pacific. The third species, S. intermedia, is widespread in South America. Each species, by modern

records, may have extended its range beyond its historic homeland. The most notable — and best recorded —

range extension is the introduction of the Australia/Pacific species into the southeastern United States.

A prefatory justification is in order for the vernacular name “Lesser Greater Duckweed,” as used in the

above title and following text. In matters of nomenclature, where the scientific name is in dispute, an unam-

biguous common name has value. Plants of the Lemnaceae have from ancient times been called “duck-weed”

or “duck-meal.” The most familiar genus of the family, Lemna, is now invariably reported as “Duckweed”

(the archaic European “duck-meal” having faded into obscurity). Species of the related genus Spirodela, al-

though similar in form, are generally larger. So that Spirodela may be distinguished from Lemna, yet reflect

their close kinship, botanists have called the native S. polyrrhiza the “Greater Duckweed.” The introduced

S. oligorrhiza — the species to be addressed here — has elsewhere been assigned the common name “Dotted

Duckweed” in transliteration of the epithet of a misapplied synonym. Plants of S. oligorrhiza are smaller

than those of S. polyrrhiza (yet larger than Lemna spp.). In that this little plant is gaining importance as an

J. Bot. Res. Inst. Texas 5(1): 197 - 203. 2011

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Journal of the Botanical Research Institute of Texas 5(1)

unwelcome weed and thus is in need of its own distinctive common name — and to show its congeneric

relation with its larger relative — “Lesser Greater Duckweed” has been volunteered.

The Lesser Greater Duckweed carries three scientific names. Two of these are within the genus

Spirodela, and one as the single species of the newly-formed genus Landoltia. The present task is to explore

the history and legitimacy of these names and to determine which of the proposed names best conforms

to the morphology of the plant and to the rules of nomenclature. As a secondary but worthy by-product of

this nomenclatural mission, a previously unexplained detail of South American plant distribution will be

resolved. The search for these answers will require an examination not only of botanical literature but of a

nearly forgotten moment of glory in the American history of scientific exploration.

The Lesser Greater Duckweed became known to American held botanists — though not by that name — with

a report by Albert Saeger (1934) of its discovery in a city park of Kansas City, Missouri. Whether known as

Spirodela oligorrhiza or Spirodela punctata, this species is not native to the Americas (Landolt 1986). Saeger

included in his report 15 figures showing the plantlets, roots, stamens, and pollen; his plant was unmis-

takably the Lesser Greater Duckweed. Prior to Saeger’s discovery only the Greater Duckweed, S. polyrrhiza

(L.) Schleid., was known in North America. Saeger made the identification by consulting the then-standard

world monograph, that of Friedrich Hegelmaier (1895). Saeger followed Hegelmaier’s nomenclature, and

reported the plant as Spirodela oligorrhiza (Kurz) Hegelm.

Saeger was familiar with an impressive prior study of North American Lemnaceae by Charles H.

Thompson (1898). Thompson, in a supplementary footnote, had reported a South American plant he named

Spirodela punctata (Meyer) Thompson. Thompson had based his description on a collection made in 1838-1842

by an American expedition to the South Pacific. The specimen label indicated (incorrectly; see below) the

material to have been collected at Orange Harbor, Tierra del Fuego. Saeger rejected Thompson’s S. punctata

for his Missouri plant, noting that Thompson’s description and accompanying figure did not correspond.

The Lemnaceae were again given serious attention by an Illinois botanist, Edwin H. Daubs, first (1962)

by reporting the discovery of Spirodela oligorrhiza from a number of additional locations in Florida, Illinois,

and Louisiana, then second (1965) with a world monograph. Daubs also rejected Thompson’s S. punctata ;

he followed Hegelmaier and Saeger in using S. oligorrhiza.

In Europe two centers arose with interest in the Lemnaceae. In the Netherlands, den Hartog and van

der Plas (1970) summarized their investigations into taxonomy of the family by a synopsis of all pertinent

scientific names with full synonymy; they treated the Lesser Greater Duckweed — though again not by

that name — as Spirodela punctata. In Switzerland, Elias Landolt conducted an extensive research program

covering all aspects of the Lemnaceae, culminating (1986) in two magnificently detailed volumes; he too

used S. punctata. A recent survey of Florida duckweeds (Ward & Hall 2010) touched lightly on the correct

name for this species; in agreement with Saeger (1934) and Daubs (1965), they again employed S. oligorrhiza.

Whatever its scientific name, the Lesser Greater Duckweed in Florida has reached such abundance that ifjv

2009 it became the first member of the Lemnaceae to be classified as “Invasive Plant, Category II” (Florida

Exotic Pest Plant Council 2009).

THE EXPEDITION

The probable source of the specimen which Thompson (1898) identified as Spirodela punctata needs deter-

mination. This will require an understanding of the itinerary of the venture on which it was obtained, the

United States South Pacific Exploring Expedition of 1838-1842. The Expedition was often referred to as

the “Wilkes Expedition” in acknowledgment of the contribution to its success of its leader, Capt. Charles

Wilkes. In 1838 the young and self-confident United States sent out what was at that time the world’s largest

“voyage of discovery,” consisting of six sailing ships and 346 men, charged with exploring and reporting

, on 18 August 1838. The s

1 of Capt. Wilkes, U.S.N., 1838-42.” Data on L

i of S. punctata in f

i Goodall], pers. comm., Oct 1989, Nov 1

the Expedition provided many opportunities for collection of the Australia/South Pacific species. The true

source of the supposed Tierra del Fuego specimen is most probably eastern Australia or one of the South

201

Saeger (1934) and Daubs (1965), relying on the descriptions and distributions reported by Hegelmaier

(1895), rejected Thompson’s name for the species they found introduced into the central and southeastern

United States. They may have been influenced by Hegelmaier’s world-wide monographic coverage, while

Thompson was reporting on a single specimen from outside his North American area of expertise.

Landolt, not withstanding his thorough coverage of the biology of the Lemnaceae, did not present

himself as a nomenclaturalist. He was aware that Spirodela punctata was insecurely typified. He (1986: 439)

forthrightly apologized for using Meyer’s name: “[T]he identity of L. punctata G.F.W. Meyer is not quite sure.

The type collection is lost. The description of L. punctata can also fit that of poorly developed plants of S.

intermedia, which occurs near the original type locality (Essequibo River, Guyana).” He even cast doubt

on the status of the few South American collections of S. punctata cited by him, suggesting that “this loose

group was included by mistake.” But he moved on, noting that “Thompson retypified the species name of

S. punctata clearly. His typification has been accepted for a long time.”

Landolt’s doubts regarding the plant that stood behind Meyer’s early name were not given weight by

the numerous subsequent writers who, impressed by his overall academic achievement, accepted without

question Spirodela punctata as the name for the introduced North American species.

Landolt merits praise for his scrupulous detailing of data, even where it encourages reappraisal by other

parties. But he erred in stating Thompson “retypified” the species. Thompson’s text shows he was describing

a specimen he believed to be the same as the type. The modern International Code of Botanical Nomenclature

(McNeill et al. 2006) permits a later person to select a replacement, a neotype. If Thompson had indicated

in any clear way that the specimen he was describing was intended as a replacement for the missing type,

his action would be interpreted as neotypification. But he seemed unaware that Meyer’s type had been lost.

He simply stated, “My description of the species is based on a collection of the plants . . . made by the United

States South Pacific Exploring Expedition.”

If neotypification by Thompson were to be put forward, the choice of a plant essentially absent from

the continent of its basionym does violence to the concept that a neotype should be as similar as possible to

type), invocation of the Code (Art. 9.16) would permit the Expedition specimen to be superseded.

A NEOTYPE FOR SPIRODELA PUNCTATA

In the belief that the plant from Guyana described by Meyer in 1818 is the same as the plant widespread

and native in South America, and to give stability to the nomenclature of this small genus, a neotype is

here selected.

THE GENUS LANDOLTIA

The Lesser Greater Duckweed cannot yet rest secure with Spirodela oligorrhiza as its scientific name. Modern

science, in the form of DNA-based cladistics, has asserted its claim. Les and Crawford (1999) have proposed

that S. oligorrhiza (their S. punctata ) be separated at generic rank from its two close relatives, S. intermedia

and S. polyrrhiza.

Spirodela and Lemna are morphologically quite similar. Species of Spirodela (s.l.) are distinguished from

all species of Lemna by the presence of several roots per plantlet (vs. a single root). Landolt (1986:464) but-

tressed this striking difference with others less apparent: Spirodela has the “frond surrounded at the base

by a small, scale-like leaflet covering the root point of attachment; pigment cells present; crystal cells with

either raphides or druses; external locules of the stamen situated at the same level as or somewhat higher

than the internal ones,” while Lemna has “no leaflet at base of the frond covering the point of attachment of

the root; no pigment cells (red pigmentation present in some species); raphides but no druses present; the

external locules of the stamen situated at the top of the

202

Journal of the Botanical Research Institute of Texas 5(1)

Les and Crawford (1999) employed chloroplast DNA sequences, as well as morphological and allozyme

data, to support their contention that S. oligorrhiza (= S. punctata, sensu Les & Crawford) was sufficiently dif-

ferent from the other two species of Spirodela as to merit generic ranking. They argued that their re-analysis of

Spirodela and Lemna morphology demonstrated S. oligorrhiza (= S. punctata, ibid.) to fall into a clade between

the two recognized genera, and not within either. They stated their allozyme data (part yet unpublished)

also supported an intermediate position. They acknowledged, however, that Spirodela had not previously

been suggested for formation of segregate genera.

Les and Crawford (1999) dismissed the classification employed by Landolt (and others) as being

paraphyletic. They considered S. polyrrhiza and S. intermedia to be sister species, but with S. oligorrhiza (= S.

punctata, ibid.) associating with Lemna. They provided phylogenetic diagrams of the relationships as inferred

by (a) Landolt; (b) themselves, as based on morphology; and (c) their DNA analysis. Each diagram implied

a different evolutionary pathway. They chose the third diagram, based on data obtained from the selected

segment of chloroplast DNA, as best representing the evolutionary relationships of the Lemnaceae.

With their target species considered closer to Lemna than to Spirodela (s.s.) and not a member of either,

Les and Crawford (1999) believed it appropriate to recognize S. oligorrhiza (= S. punctata, ibid.) as a new

genus to better reflect this “revised hypothesis of duckweed relationships.” They selected Landoltia for its

name, in honor of the scholarly work on the family by Elias Landolt.

A response to Les and Crawford (1999) must be made from four directions, some factual and others

philosophical. First, of course, as established by selection (above) of a South American neotype for Meyer’s

Guyana plant, the name Landoltia becomes irretrievably attached to the plant presently known as Spirodela

intermedia, a wholly unwelcome outcome.

Second, the comments by Les and Crawford (1999) regarding morphology are not unchallengeable. As

they observed, no previous author has considered dividing Spirodela into two genera; even the lesser rank

of section has been employed cautiously. Den Hartog and van der Plas (1970: 359) noted that one previous

author had distinguished two “species groups” within Spirodela; but they stated that, in their opinion, these

groups were too insignificant to be regarded even as separate sections. The reason for the failure of previous

workers to divide Spirodela must in large part rest on the conspicuous morphological character found in all

species of the genus — the difference in number of roots on a plantlet. Though less readily observed, other

morphological differences described by Landolt (1986), especially the scale-like leaflet surrounding the root

base, cannot be matched in Lemna.

Third, Les and Crawford’s cpDNA conclusions are based on a “hypothesis” (their word), as is of course

any analysis that has access to no more that a segment of chloroplast DNA and no nuclear DNA. This is not

to fault the technology, but to call attention to the necessarily tentative basis of close DNA-based evolution-

ary pathways.

And fourth — and perhaps most importantly — heavy reliance on DNA data causes a slighting of char-

acters that can actually be observed in every-day taxonomic work. Verifiable observation becomes second-

ary to trust in the conclusions of others. If a plant is to be identified in the held, or even under laboratory

determination. Several roots are present, or they are not; a scale is present, or it is not. Once these unifying

genus of incontestable morphological similarity, especially one of such small (three-species) size, cannot

but confuse those parties whose interest is plant identification. Morphological clarity is lost.

The approach of Les and Crawford (1999) is not to be belittled, for even as a hypothesis their suggested

diagram of evolutionary pathways is of informational value. Yet to use that scant basis as justification for

nomenclatural change is an unproductive over-extension of their work.

CONCLUSION

The Lesser Greater Duckweed, a species native to Australia and the South Pacific and introduced into eastern

BOOK REVIEW

U.S.A.; 314-57-9594; 314-577-9547 f;

THOMAS WALTER’S ORCHIDS

Daniel B. Ward

John Beckner

r Fir, Abies fraseri (his F

is (L.) Lindl. [= Habenaria ciharis (L.) R. Br.], I

i SC. The first two h

210

Journal of the Botanical Research Institute of Texas 5(1)

Modern name: Cypripedium reginae Walt.

Comments: Very rare: in the Carolinas, known only in NC (2 counties). Spm. 39-B of the Fraser/Walter her-

barium is labeled “Cypripedium Reginae” in Fraser’s hand. It has been marked as “type” (by O. Ames?), but

the designation was not published. Since Walter could most likely have seen this species only through the

agency of Fraser, this specimen (or another of the same gathering) may well have used by spm.

39-B has been designated (Ward 2007c) lectotype of Cypripedium reginae Walt.

Walter’s name: Arethusa divaricata (p. 222). Linnaeus, Sp. Pi. 951. 1753.

Walter’s description: radice fibrosa, foliis lato-subulatis, capsula oblonga subtereti sexsulcata, petalis tribus lance-

olatis purpureis suberectis, aliis duobus latis incarnatis, caule unifloro.

Modern name: Cleistes divaricata (L.) Ames

Comments: Frequent on SC coastal plain. Spm. 8-E of the Fraser/Walter herbarium (BM) was labeled “Arethusa

Divaricata” by Fraser. Walter very probably correctly identified Linnaeus’ Arethusa divaricata.

Walter’s name: Arethusa racemosa Walter (p. 222)

Walter’s description: radice tuberosa bipartita, foliis radicahbus ovatis nervosis, scapo longo subnudo racemifloro,

floribus parvis alternis pedicellatis, petalis patulis aequalibus albidis, capsulis ovatis.

Modern name: Ponthieva racemosa (Walt.) Mohr

Comments: Occasional in coastal SC. The label of spm. 8-D (“Arethusa racemosa ”) is in Walter’s hand. But

since the species would have been available to Walter near his home and the label bears a 3 -digit number

(“??9”) which indicates the specimen to be a Fraser collection, it may not have been seen by Walter until

after completion of his manuscript. Blake (1915) correctly called it “an excellent specimen.” It has been

selected (Ward 2007c) as neotype for Arethusa racemosa Walt., basionym of Ponthieva racemosa (Walt.) Mohr.

Walter’s name: Arethusa spicata Walter (p. 222)

Walter’s description: radice tuberosa, caule sesquipedali succulento aphyllo, floribus bracteatis sessilibus alternis

spicatim positis, petalis aequalibus ovatis conniventibusflavescentibus striis purpureis, nectario longitudine petalorum

rugosa subtrilobo, lobis lateralibus brevioribus erectis, colore petalorum, lobo medio purpureo propendente, capsula

columnan angulare.

Modern name: Hexalectris spicata (Walt.) Barnh.

Comments: Infrequent throughout. As Blake (1915) has stated, Walter’s diagnosis is “quite distinctive” of this

species. There is no specimen in the Fraser/Walter herbarium. A specimen from Richland County, South

Carolina, has been selected (Ward 2008) as neotype for Arethusa spicata Walt., basionym of Hexalectris

spicata (Walt.) Barnh.

Walter’s name: Arethusa foliosa Walter (p. 223)

Walter’s description: radice bulbosa, caule folioso unifloro, foliis alternis brevibus nervosis, flore terminali lineari

albido, capsula oblonga hexangula.

Modern name: Probably Triphora trianthophora (Sw.) Rydb.; possibly Isotria verticillata (Muhl. ex

Willd.) Raf.

Comments: The name Arethusa foliosa is not recorded in modern floras, even in synonymy. Walter’s rather

detailed description cannot be wholly matched with any orchid of eastern America. But Walter must have

had something, and if his plant were to be identified, his epithet could be prior to a familiar name. Two species

come to mind, each rare in the Carolinas. Triphora trianthophora is almost restricted to the mountains, yet

has been found in Berkeley Co.; Isotria verticillata is wholly unknown on the coastal plain (possibly reaching

Walter via the agency of Fraser). The flowers of Triphora are white (“albido”), its leaves cauline (“caule folioso”)

and short (“brevibus”); yet its flowers are only rarely single (“unifloro”) and never terminal (“terminali”). Isotria

bears solitary terminal flowers, and its elongate, narrowly lanceolate sepals conform to Walter’s “lineari”;

yet its perianth is purplish to green, and its distinctive whorled leaves could scarcely have been overlooked.

No specimen has been identified in the Fraser/Walter herbarium. Walter’s description is surely an amalgam

212

Journal of the Botanical Research Institute of Texas 5(1)

BOOK REVIEW

Tim Robinson. 2008. William Roxburgh: The Founding Father of Indian Botany. (ISBN-13: 978-186077-

434-8, ISBN 10: 978-1-86077-434-2, hbk.). Joint publication of the Royal Botanic Garden Edinburgh and

Phillimore & Co. Ltd., Phillimore & Co. Ltd., Healey House, Dene Road, Andover, Hampshire, England,

SP10 2AA, U.K.). (Orders: The History Press, Sales (Phillimore & Co. Ltd), The Mill, Brimscombe Port,

Stroud, Gloucestershire, England, GL5 2QG, U.K.; http://www.thehistorypress.co.uk/; 01453 883300

(International +44 1453 883300; 01453 883233 fax). £50.00, 286 pp„ 150 b/w, 16 color, 8 x 11".

University Dr., Fort Worth, Texas 76107-3400, U.S.A.

J. Bot. Res. Inst. Texas 5(1): 212. 2011

BIOLOGICAL STATUS OF CROTON BIGBENDENSIS (EUPHORBI ACE AE) :

SPECIES, VARIETY, OR INFORMAL VARIANT?

214

Journal of the Botanical Research Institute of Texas 5(1)

Example 4 notes that “The holotype of Cephaelis acanthacea Steyerm., Cuatrecasas 16752 (F), consists of a

single specimen mounted on two herbaruim sheets, labeled ‘sheet V and ‘sheet 2.’ Although the two sheets

have separate herbarium numbers, F-1153741 and F-1153742, respectively, the cross-labeling indicates that

they constitute a single specimen.”

The dioecious Croton bigbendensis is only fully represented as a species by at least two plants, which show

the staminate and the pistillate condition. A single plant would be acceptable as the nomenclatural type,

but the two plants more fully define the species. The original typification clearly and explicitly represents

the two plants as part of a single specimen.

Biological Status of Croton bigbendensis

Regarding the procedures of his revisitation, Henrickson informs the reader that “Data were collected from

all specimens at TEX-FF (n=296) with 41 from trans-Pecos Texas identified by Turner as “Croton bigbendensis”

plus six of my specimens that are clearly referable to that taxon, 72 from typical Croton dioicus in trans-Pecos

Texas (west of the River), 98 from C. dioicus from cis-Pecos Texas (east of the Pecos River) and adjacent New

Mexico, and 85 from C. dioicus from Mexico.” All of these plants were reexamined by me in the present

study, except for the six specimens of C. bigbendensis collected by Henrickson, these presumably not as yet

mounted; at least they were not found among the current collections at TEX-LL.

In the current reassessment of Croton bigbendensis and C. dioicus, l have studied again the two taxa

concerned, most all of which had been annotated by me during my initial studies. Not a single sheet, how-

ever, was formally annotated by Henrickson, even before or after his publication. His failure to annotate

specimens of C. dioicus was of no concern, for the taxon is relatively invariant throughout its range, but his

failure to annotate sheets of C. bigbendensis, or the hypothetical intergrades has made it difficult for me to

understand his assessment. It should also be noted that Henrickson did not borrow, or annotate, the large

number of collections of this complex housed at SRSC, the herbarium out of which I was working at the

time of my interest in the taxa concerned.

My restudy of the Croton bigbendensis- C. dioicus complex has not changed my original conclusions. There

are clearly two taxa in the trans-Pecos (Fig. 1), one of these a robust erect, much-branched, perennial herb

to 1 m high, having mostly linear lanceolate upper leaves (C. bigbendensis), the other a low, mostly compact

perennial herb with more nearly ovate leaves, the blades usually obtuse or rounded at the apices (C. dioicus).

The two taxa are readily recognized in the held, where they often exist as large populations. In my original

publication of C. bigbendensis, I was unaware of the extension of C. dioicus into southern Brewster County,

where they approach populations of C. bigbendensis, mainly because at the time of my original publication I

had not reexamined the TEX-LF collections, because of my Alpine residence during that period. But, during

my many held trips in that region at the time, I never found the two taxa growing together or near each other.

Subsequently, however, I discovered a collection at TEX-FF ( Warnock s.n. Persimmon Gap. Chisos

Mountain area, 2 May 1937) that documented their co-occurrence in southern Brewster Co., if the sheet

concerned is not an error in mounting. Henrickson himself, by penciled notation (“= Bigbendensis”), recog-

nized the two specimens on that sheet as distinct “somethings” (having failed to annotate C. dioicus, which

bore my annotation with this name). The only other area or population in which the two taxa have been

noted as growing together has been in collections made by me, called to the fore by Henrickson (p. 299):

Henrickson, in his evaluation of the taxa concerned, attempting to make his study more quantitative than

my own, constructed a graph (his Fig. 1) on which was positioned those plants referable (in his opinion)

Turner, Status of Croton bigbendensis 215

to C. bigbendensis, and those referable to C. dioicus. The data used to construct the graph were said to be

internode length (the longest internode observed on the plants concerned) and leaf blade length/width

ratios. To his credit, most of the measurements were penciled on most of the sheets that he examined.

The measurements, however, were made from herbarium sheets from over a very large area, which merely

showed eclectic variation of the taxa concerned. What would have been more meaningful were graphs for

individuals from populations in the held, especially where the two taxa might occur together, suggesting gene

exchange — in short, following the procedures of Edgar Anderson (1949, 1954), using scatter diagrams, with

care given to the particular measurements recorded (1954). Thus, Anderson (1954) would have demanded

that leaf measurements be made consistently from leaves at a given node, and internodal length, likewise,

say between the 6th and 7th nodes, if such could be determined.

cause they were not populational in nature, but also because the sheets were not formally annotated by him.

This brings me to Henrickson’s Figure 2, “Distribution of Croton dioicus and “bigbendensis” in trans-

Pecos Texas, based on specimens so identified by Turner at TEX-LL).” Henrickson mapped 10 plants and/

216

Journal of the Botanical Research Institute of Texas 5(1)

or populations of “bigbendensis” as occurring in southern Brewster Co., 10 such collections from adjacent

Presidio Co. and 2 from Hudspeth Co., most of these occurring along the Rio Grande. He notes these to be

so identified by myself. My assumption is that he would not have identified the collections concerned as C.

bigbendensis, but rather would have dubbed them C. dioicus. As already noted, he failed to formally annotate

any of the sheets concerned, either at TEX or SRSC. On this same map, Henrickson included a series of open

squares that were said to be “intergrades,” 7 of these in Brewster Co, 1 in Presidio Co. and several others in

more western counties. Unfortunately, he did not list the so-called intergrades in his paper, nor did he an-

notate any of the sheets on hie at TEX-LL, as such, but I could infer some of these by location and penciled

comments on some of the mapped sheets. Thus, on the single so-called intergrade from Presidio Co. (Butterwick

& Strong 829), Henrickson had scribbled in pencil “Closer to” with an arrow pointing to Croton dioicus (my

formal annotation prior to my discovery of C. bigbendensis). I could locate only one other possible, so-called

intergrade among the Brewster Co, sheets of C. bigbendensis on hie at TEX-LL, this being Baker 1512, in which

Henrickson placed, in pencil, the word “No!” next to my inked annotation, “C. bigbendensis B.L. Turner.” In

my opinion, both of these so-called intergrades are typical elements of C. bigbendensis. Among the C. dioicus

specimens on hie at TEX-LL (Nine Point Mesa, Webster & Westford 32586) I found one plant with the pen-

ciled “No!” placed next to an old annotation label of mine inked in as B. bigbendensis. My original annotation

was in error; it should have been C. dioicus, but not an intergrade, as suggested by Henrickson’s notation.

More egregious is Henrickson’s contention that pressed specimens identihable (to him) as “bigbendensis”

were found well within the Mexican range of C. dioicus (e.g., states of Nuevo Leon, Hidalgo and Puebla). I

217

have examined all of the mounted sheets of from these states at TEX-LL (none annotated by Henrickson)

and I would identify these as part of the fabric of C. dioicus, these annotated accordingly. Henrickson did

call attention to a single collection from the Samalayuca dunes of north-central Chihuahua that I had not

examined in my initial studies; he thought this might compare well with “bigbendensis.” I take the sheet

concerned to be closer to C. dioicus, but perhaps an undescribed taxon, having larger stellate hairs that seem

atypical of C. bigbendensis, and larger ovaries and peculiar leaves. While Henrickson called attention to the

plant concerned, he did not annotate or place notes upon the specimen. I annotated the sheet as perhaps

undescribed, but mapped this as C. dioicus in Figure 2.

Biological Status and Taxonomic Rank of Croton bigbendensi

be recognized as a sound biological taxon, or not. In my opinion, yes! Most any experienced field taxonomist

could recognize the populational elements concerned at a glance, surely each deserving of a name. Should

it be called a species or a variety? Lacking adequate populational data in regions of contact, it is difficult to

say. If the populations coexist without clear intermediates, it seems best to threat these as species, as has

been argued elsewhere (Turner and Nesom 2000); if it can be shown that the two taxa intergrade to some

significant extent near regions of contact they are perhaps better treated as infraspecific taxa. Henrickson,

s a differen

point of view; his concluding paragraph reads as follows (p. 300):

Henrickson (2010) has not, in my opinion, as discussed above, presented adequate quantitative populational

data to show that the taxa concerned intergrade near regions of contact. Indeed, to paraphrase his sentence

bolded in the above paragraph, I can state this with certainty. As it stands, Croton bigbendensis is a

very distinctive species, easily recognized over its range by substantial characters. Henrickson’s

final sentence is true, depending upon his definition of experimental data; certainly none is provided in his

quantitative assessments of the taxa concerned here. Indeed, most all of the taxa among higher plants gener-

ally, are lacking in experimental data, but this does not preclude our recognition of a multitude of species,

applied to populations. No doubt, the final arbiter in all this ado will be the application of DNA data. I look

forward to the systematic conclusions drawn from such studies.

ACKNOWLEDGMENTS

I am grateful to my long-time colleagues, Guy Nesom and Mike Powell, for reading the paper and making

helpful suggestions. Distribution maps (Figs. 1, 2) are based upon specimens on file at SRSC and TEX-LL.

One anonymous reviewer’s comments are greatly appreciated.

REFERENCES

Anderson, E. 1949. Introgressive hybridization. John Wiley & Sons, New York

Anderson, E. 1954. Efficient and inefficient methods of measuring specific differences. In: Kempthorne,% : 3|.

Bancroft, J.W. Gowen, and J.L. Lush. Statistics and mathematics in biology. Iowa State College Press, Ames, Iowa.

Henrickson, J.^l4§ Croton bigbendensis Turner (Euphorbiaceae) revisited. J. Bot. Res. Inst. Texas 4:295-301 .

Turner, B.L. 2004. Croton bigbendensis (Euphorbiaceae), a new species from trans Pecos Texas. Sida 21:79-85.

Turner, B.L. and G.L. Nesom. 2000. Use of variety and subspecies and new varietal combinations for Styraxplatani-

folius (Styracaceae). Sida 19:257-262.

218

Journal of the Botanical Research Institute of Texas 5(1)

BOOK REVIEW

Kathryn Mauz. 201 1. An Agreeable Landscape: Historical Botany and Plant Biodiversity of a Sonoran

Desert Bottomland, 1855-1920. (ISBN 13: 978-1-889878-35-5; ISSN 0833-1475; pbk.). SBM #35.

Botanical Research Institute of Texas, 1700 University Dr., Fort Worth, Texas 76107-3400, U.S.A.

(Orders: http://www.brit.org/brit-press/books/sbm-35; orders@brit.org; 817-332-4441 x 33). $30.00,

234 pp., b/w and color figures, 6.5" x 9.5".

J. Bot. Res. Inst. Texas 5(1): 218. 2011

ENDOSPERM AND COTYLEDON AREOLE CORRELATION IN

LEGUMINOSAE SUBFAMILY PAPILIONOIDEAE

James A. Lackey

Smithsonian Institution

Department of Botany, NMNH MRC- 1 66

PO Box 3701 2

Washington, DC 2001 3-7012

lackeyj@si.edu

INTRODUCTION

Beck (1878) discovered a small peculiar characteristic spot “besondere Eigenthmulichkeit . . . Fleck,” which

he named an “Aleuronfleck” (protein-spot), on each abaxial cotyledon surface of dormant mature seeds of all

studied species of legume tribe Vicieae DC. (= Fabeae Rchb.). At this spot, epidermal cells and subtending

mesophyll cells contain chlorophyll-colored protein-containing bodies, and have different size and shape,

surrounding cells. In the soybean and other species of the genus Glycine Willd. (tribe Phaseoleae DC.), the

spot, often subtle and known as a “pit” was rediscovered at least twice and has been studied extensively

J. Bot. Res. Inst. Texas 5(1): 219 -2

220

Journal of the Botanical Research Institute of Texas 5(1)

(Dzikowski 1936, 1937; Miksche 1961; Yaklich et al. 1984, 1986, 1987, 1989, 1992, 1995, 1996, 1998;

Baker & Minor 1987; Ma et al. 2004). All these soybean researchers thought that the “pit” was unique to

the genus Glycine. Endo and Ohashi (1997) again rediscovered this spot in tribes Cicereae Alef., Trifolieae

Endl., and Vicieae and named it a cotyledon areole, and later (Endo and Ohashi 1998a) established that the

soybean “pit” is the same as their cotyledon areole. Endo and Ohashi (1998a, 1998b, 1999a, 1999b), found

cotyledon areoles in 73% of studied species of Leguminosae Juss. subfamily Papilionoideae Giseke but absent

in subfamilies Caesalpinioideae DC. and Mimosoideae DC. Presence in Bobgunnia J.H. Kirkbr. & Wiersema

(tribe Swartzieae DC.), indicates that cotyledon areoles developed early in papilionoid evolution (Lackey

2009). Within the papilionoid legumes, presence or absence, shape, and position seem to have taxonomic

significance for genera, tribes, and groups of tribes (Endo and Ohashi 1999a). Cotyledon areole function

remains unknown (Ma et al. 2004; Lackey 2009).

Despite frequent statements to the contrary (Candolle 1825; Bentham 1865; Carlson & Lersten 2004),

conspicuous endosperm deposits are found in most legume seeds (Schleiden & Vogel 1839b; Nadelmann

1890; Pammel 1899; Kopooshian 1963; Smith, 1983). These deposits almost always occur by lateral local-

ized enlargement of endosperm tissue, which accommodates massive development of galactomannan, also

known as gum or mucilage, secondary cell-wall material (Nadelmann 1890; Buckeridge et al. 2000). The

relatively small lateral endosperm deposits of most modern soybean cultivars have been called an “antipit”

(Ma et al. 2004). Presence or absence and degree of development seem to have taxonomic significance for

species, genera, and tribes (Smith 1983). Studies of model legumes indicate that during germination, galac-

tomannan is hydrolyzed to galactose and mannose, then transformed to sucrose in the endosperm, followed

by transport to the cotyledons (Buckeridge et al. 2000).

Previous studies (Lackey 2007, 2008, 2009, 2010, unpublished results) of some tribe Phaseoleae, tribe

Swartzieae, and other legumes suggested that cotyledon areole presence and position are always positively

correlated with presence and position of conspicuous endosperm, as does an analysis of the results of Beck

(1878) and Ma et al. (2004; etl^ljifra) for tribe Vicieae and Glycine. However, there is no broad survey to

see if this correlation holds for all papilionoid legume tribes. The purpose of the current study is to ex-

amine at least one species from every papilionoid legume tribe in the systems of Polhill and Raven (1981),

Polhill (1994a, 1994b) and Lewis et al. (2005) to verify that the correlation holds for all tribes. In addition,

a substantial review is given of the extensive disconnected literature, particularly rarely-cited significant

literature, of the last 200 years to see if current results shed any light on the function of cotyledon areoles

and endosperm galactomannan, and thus the parallel systematic utility of these characteristics.

MATERIAL AND METHODS

At least one representative of all papilionoid tribes in the classification systems of Polhill and Raven (1981),

Polhill (1994b), and Lewis et al. (2005) was examined. Included are the common commercial sources of

papilionoid galactomannan gum: guar ( Cyamopsis tetragonoloba (L.) Taub.) and fenugreek ( Trigonellafoenum -

graecum L.). Specimens studied and results are given in Table 1.

Seeds were examined and stained by the methods of Lackey (2007, 2008). Lor those species with endo-

sperm or cotyledon areoles, camera lucida outline drawings were made (Lig. 1), except for Glycine max, which

is already well-depicted in the literature (Ma et al. 2004). Endosperm was considered present if seen with 50x

optics and adjacent to any portion of the abaxial cotyledon surface. Minute endosperm residues between the

cotyledon margins and the crease between the cotyledons and radicle, as well as any minutely thin remnants

of the endosperm envelope surrounding the embryo were ignored. In this study, the term endosperm refers to

visible lateral endosperm tissue unless designated otherwise. Approximate endosperm percent in cross section

at the point of maximum occurrence was measured by the method of Anderson (1949). Cotyledon areoles

were considered present if they were manifested by an unambiguously distinct spot of projecting epidermal

cell walls on the cotyledon abaxial surface or a corresponding mirror-image impression on the endosperm,

or if two or more of the following six cotyledon areole characteristics were present on the cotyledon abaxial

side: 1) an area of epidermal cells along midvein of different size or shape than surrounding cells, 2) an area

221

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deposits are indicated by arrows. For each specimen, a left cotyledon and radicle is outlined. Cotyledon areoles are outlined and filled with a grid pattern.

(1) Abrus precatorius L. (2) Adesmia bicolor (Poir.) DC. (3) Ammopiptanthus mongolicus (Maxim, ex Korn.) S.H. Cheng (4) Amorpha fruticosa L.

(5) Aotus ericoides (Vent.) G. Don (6) Apoplanesia paniculata Presl (7) Astragalus sinicus L. (8) Baptisia sp. (9) Bobgunnia madagascariensis (Desv.) J.H.

Kirkbr. & Wiersema (10) Brongniartia canescens (S. Watson) Rydb. (11) Calpurnia aurea (Aiton) Benth. (12) Carmichaelia australis R. Br. (13) Chorizema

cordatum Lindl. (14) Cicer arietinum L. (1 5) Securigera varia (L.) Lassen (1 6) Crotalaria cunninghamii R. Br. (1 7) Cyamopsis tetragonoloba (L.) Taub. (1 8)

(22) Gompholobium scabrum Sm. (23) Goodia medicaginea F. Muell. (24) Hedysarum coronarium L. (25) Hypocalyptus sophoroides (PJ. Bergius) Baill.

(26) Indigoferasuffruticosa Mill. (27) Lotus maritimus L. (28) Lupinus luteus L. (29) Onobrychis conferta (Desf.) Desv. (30) Otholobium candicans (Eckl. &

Zeyh.) C.H. Stirt. (31) Podalyria calyptrata (Retz.) Willd. (32) Pseudoeriosema borianii (Schweinf.) Hauman (33) Robinia pseudoacacia L. (34) Sesbania

cannabina (Retz.) Pers. (35) Stylosanthes guianensis (Aubl.) Sw. (36) Templetonia retusa (Vent.) R. Br. (37) Tephrosia purpurea (L.) Pers. (38) Thermopsis

villosa (Walter) Fernald & B.G. Schub. (39) Trigonellafoenum-graecum L. (40) Viciafaba L. Scale bars = 1 mm

222

Journal of the Botanical Research Institute of Texas 5(1)

of epidermal cells that stain differently from surrounding cells, 3) a distinct green to yellow spot, 4) an area of

epidermal cells lacking stomata, 5) an area with subepidermal cells of different size, shape, or stainability from

surrounding cells, or 6) an area of epidermal cells with anticlinal walls sinuate parallel to the embryo axis.

Of the 52 species examined, 11 lacked cotyledon areoles and discernible endosperm, and 39 showed both

cotyledon areoles and discernible endosperm. Dupuya madagascariensis (R. Vig.) J.H. Kirkbr. and Hypocalyptus

sophoroides (P.J. Bergius) Baill. have abundant endosperm, but cotyledon areoles were either absent (D. mada-

gascariensis) or questionably discernible (H. sophoroides ). In all specimens, the maximum extent of lateral

endosperm was always immediately external to the cotyledon areole (Table 1, Figs. 1-2).

Cotyledon areole position, size, shape, and other attributes varied from species to species (Table 1). Many

were basal to medial. Hedysarum coronarium L. and Tephrosia purpurea (L.) Pers. were apical or subapical. All

were positioned external to the midvein or when branched (Fig. 2s), external to the midvein and portions of

major branch veins. Size and shape varied from minute to perhaps almost a third of the cotyledon surface

(. Amorphafruticosa L., Fig. 2u), from oval to linear, and from unbranched to branched. Within Glycine, the

wild soybean, G. soja Siebold & Zucc., had a much larger cotyledon areole, relative to overall seed size, than

the cultivated soybean, G. max (L.) Merr. Cotyledon areole epidermal cells differed in size and shape from

other cotyledon epidermal cells (Fig. 2h, 2t). In some species and some seeds of some samples, cotyledon

areole epidermal cells were slightly green-yellow or a slightly darker color relative to most other cotyledon

epidermal cells (Fig. 2g, 21). Sometimes the cotyledon areole was in a localized concave area (Fig. 2g, 2n)

filled with corresponding endosperm tissue (Fig. 2g, 2o). Most species had no such concave area. Cotyledon

areole epidermal cells almost always had convex outer periclinal walls, which gave a shagreened or granular

appearance when viewed under low magnification and lateral bright light (Fig. 2e, 2n). The anticlinal walls

parallel to the cotyledon axis were often sinuate in surface view (Fig. 2t). The cotyledon areole epidermal

cells usually stained less readily with surface-applied toluidine blue than other cotyledon epidermal cells

(Fig. 2f, 21, 2r). Cotyledon mesophyll cells between the cotyledon areole epidermal cells and the veins usually

stained more readily with toluidine blue than other mesophyll cells outside the cotyledon areole, and cell size

and position were different from other mesophyll cells (Fig. 2m). In embryos hydrated after removal from

the endosperm envelope, epidermal blistering of epidermal cells outside the cotyledon areole was generally

greater, but sometimes lesser, than the minimal epidermal blistering occurring in embryos hydrated while

still within the endosperm envelope. Cotyledon areole cell size and shape and other epidermal features

could often be seen readily as a mirror-image impression on the interior surface of dry endosperm tissue

(Fig. 2n, 2o). Many times this impression gave a better surface view of the cotyledon areole appearance than

the direct view of the cotyledon.

Endosperm extent varied from unobservable with 50x optics to occupying 60% of seed width (Fig.

2q). When visible, it always formed a complete envelope around the embryo (Fig. 2d) and always adherent

to the testa. It was almost always free from, rarely adherent to, the embryo. The embryo outline (Fig. 2b)

and the most minute details of the embryo epidermis could be seen as impressions on the dry endosperm

interior. Thickness of the endosperm envelope was not uniform, but varied according to a consistent layout.

It was thinnest under the hilum. A slight thickening occurred in the triangular (in cross section) slit formed

where the cotyledon margins touch (Fig. 2i). At a similar, but usually larger, slit between the radicle and

cotyledons a somewhat larger deposit formed (Fig. 2b). The greatest development, if any, occurred adjacent

to the abaxial cotyledon faces (Fig. 2a). The development of this area varied in position to be directly exterior

to the cotyledon areole. It was usually more or less symmetrically positioned about the cotyledon areole,

but sometimes it was slightly asymmetrically positioned towards the apex of the cotyledon, especially if

the cotyledon areole was basal. Hardness of the endosperm to a knife or cutters varied from relatively soft

to fairly hard ( Cyamopsis tetragonoloha), but always softer than the dry Malpighian layer of the testa. It was

usually colorless, sometimes dark to red-purple ( Viciafaba L.), and generally appeared crystalline when

cut. When well-developed, three layers of endosperm could generally be seen. The outer aleurone layer was

5 in the testa (Fig. 2c). Endosperm of Glycine (Fig. 3) was u

225

species. The wild soybean, G. soja, had endosperm larger than average (Anderson 1949; current results) for

papilionoid legumes. The cultivated soybean, G. max, had a much smaller than average endosperm.

Although often subtle, detection of cotyledon areoles and endosperm was usually immediately unambiguous,

but several specimens presented problems. Detection generally seemed easiest for compact or even small coty-

ledon areoles that showed a clear differentiation from remaining cotyledon epidermal cells. Very small seeds,

very small cotyledon areoles, or very large cotyledon areoles made observation difficult. Ammopipthanthus

mongolicus (Maxim, ex Kom.) S.H. Cheng and Stylosanth.es guianensis (Aubl.) Sw. were difficult to dissect, and

had small cotyledon areoles and endosperm, near the limit of detection. Lespedeza bicolor Turcz. likewise was

J^pPtult to dissect, and any cotyledon areole or endosperm were too small to allow unambiguous detection.

Amorphafruticosa had a large endosperm, and the presumptive cotyledon areole appears large, covering much

of the abaxial surface (Fig. 2u); in cross section, the usual disruption of mesophyll cell patterns below the

cotyledon areole is not readily evident, but these mesophyll cells do stain more readily than adjacent ones

(Fig. 2m). Similarly, Hypocalyptus sophoroides had massive white endosperm, but a cotyledon areole was not

readily conspicuous, although a patch of larger cells with slightly protruding periclinal exterior walls over

the middle vein suggests a cotyledon areole, but with no clear distinction and no differential staining from

the remaining smaller epidermal cells (Fig. 2t); in cross section through the cotyledon base, mesophyll is

entirely spongy, with no readily apparent distinction in the cotyledon areole area. Dupuya madagascariensis

was the only species with a conspicuous prominent endosperm and no indication of cotyledon areole.

Cotyledon areole size seemed generally positively correlated with endosperm size. The correlation was

demonstrated by measurement. Cotyledon areole length relative to cotyledon length ranged from 0.06 ( Cicer

arietinum L.) to 0.82 ( Crotalaria cunninghamii R. Br.). In those species with cotyledon areoles, there was a

positive correlation between cotyledon areole relative length and relative endosperm thickness (R 2 =0.35).

Correlation

Results of the current broad sampling of all papilionoid legume tribes, concurs with previous work in specific

groups to show an almost perfect positive correlation between cotyledon areole presence and position and

endosperm presence and position in papilionoid legumes. The only known exception, discovered in the

current work, is the absence of a cotyledon areole in Dupuya madagascariensis, where abundant endosperm

is found. A similar possible exception is Hypocalyptus sophoroides.

Although prior authors did not note any correlation, a review of their observations in specific groups

supports it. In Vicieae, analysis of Beck’s (1878) purely descriptive anatomical work indicates that basal

cotyledon areoles are always present and positioned adjacent to endosperm in all studied species of Vida,

and noted for the genera Pisum, Lens, and Lathyrus, for which he cited no species. Yaklich et al. (1989) ob-

served medial cotyledon areoles and what is now known to be adjacent endosperm in all 185 accessions of

soybean and 45 accessions of seven other species of Glycine (tribe Phaseoleae subtribe Glycininae Benth.).

Additional support is in Phaseoleae subtribe Clitoriinae DC., in which Lackey (2007) found cotyledon areoles

and endosperm in all studied species of Periandra Mart, ex Benth., Centrosema (DC.) Benth., and Barbieria

DC., and two species of Clitoria L., but no cotyledon areoles and no endosperm in four Clitoria species. In

Phaseoleae subtribe Kennediinae Benth., Lackey (2008) found cotyledon areoles and prominent endosperm

in all studied species. Lackey (2009) found cotyledon areoles and endosperm in Bobgunnia madagascariensis,

and no cotyledon areoles or endosperm in two Swartzia species (tribe Swartzieae). Lackey (2010) found both

endosperm and cotyledon areoles in Pueraria montana (Lour.) Merr. and Cyamopsis tetragonoloba.

Across the range of papilionoid legumes with cotyledon areoles studied here, in addition to the above

by length along the midvein, was positively correlated with endosperm size, when adjusted for seed size

differences. Lackey (2010) found a similar positive size correlation across a range of wild, semiwild, and

domesticated soybean samples, which are a single interbreeding gene pool.

June 22

Fig. 2. Cotyledon areoles and endosperm in papilionoid legumes.

{a)Sesbaniacannabina (Retz.) Pers. JAL 1443. (b-f) Robinia pseudoacacia L. JAL 1442. (g) Glycine max (L) Merr. JAL 1598. (h) Indigoferasuffruticosa

Mill. JAL 1615. (i) Goodia medicaginea F. Muell. JAL 1635. (j-k) Vicia faba L. lithograph figures 8, 24 from Beck (1878). (I) Vicia faba L. JAL 1002. (m)

Amorphafruticosa L. JAL 1613. (n-o) Vicia faba L. JAL 1002. (p) Glycine max ( L.) Merr. lithograph figures 76, 80, 81 from Dzikowski (1937). (q) Cyamopsis

tetragonoioba (L.) Taub. JAL 1582. (r-s) Cyamopsis tetragonoloba (L.) Taub. JAL 1585. (t) Hypocalyptus sophoroides (PJ. Bergius) Baill. JAL 1645. (u)

Amorpha fruticosa L. JAL 1 61 3.

(a) Transverse dry seed section. Conspicuous endosperm in two large lens-shaped areas on each abaxial side of cotyledons, (b) Median dry seed

radicle and cotyledon pockets, (c) Exterior hydrated seed, left side. Cracks around hilum and radicle, showing endosperm beneath cracks in testa, (d)

Endosperm thickening at margins of cotyledons, (j) Cotyledon areole ("Aleuronfleck") in cross section (300X), a large granules ("Korner") of the epider-

mis, b smaller granules of the parenchyma, g globoids ("Globoide"). Note large and smaller granules are hand-tinted yellow-green (Pantone 366 U) in

original figure. External periclinal epidermal cell walls slightly convex, (k) Ventral view of hulled seed of Vicia faba, r radicle ("Radicula"), a cotyledon

areole ("Aleuronfleck"). (I) Embryo, dry, stained on one side with toluidine blue. Cotyledon areole is yellow-green, stains less readily than surrounding

cotyledon area, (m) Transverse section of cotyledon through cotyledon areole and midvein, (n) Cotyledon areole, SEM. (o) Endosperm from same seed

227

soybeans are similar with galactoman

US soybean cultivars lack such a well-

. Many other landraces, wild, and semiwild

jcture on each side of the cotyledons. Most

ercial galactomannan gum.

Problem observations

Most observations are unambiguous. However, two contradict previous observations: Indigofera suffruticosa

ambiguous cotyledon areole (Fig. 2h). As discussed in results, detection of the cotyledon areole of Amorpha

in (n), unstained and natural color. Endosperm dark pigmented, (p) 76. Seed coat with lenticular formation ("formation lenticulaire") of the endosperm

and adjacent cotyledon. Transverse section in glycerin, pi palisade tissue ("couche en palissade"), pdp hourglass layer ("cellules auxiliaires"), m paren-

chyma tissue ("couches du parenchyme"), ends endosperm remnants with lens ("lenticule")between aleurone tissue and parenchyma, cot cotyledon.

80. Cotyledon abaxial surface below the endosperm lens ("lenticule"). Transverse section, ep epidermis, m parenchyma. 81 . As figure 80, Longitudinal

section. Note Dzikowski's depiction of cotyledon areole epidermal cells with convex peridinal walls in figure 80. Also note the cotyledon areole epidermal

cells width greater than surrounding cells in figure 80 and smaller in figure 81 . (q) Transverse dry seed section. Massive hard endosperm on each abaxial

side of cotyledons, (r) Embryo, dry, stained with toluidine blue. Staining lighter at cotyledon areole. (s) Cotyledon areole, SEM. Branching at lateral veins,

(t) Cotyledon surface, SEM. Cotyledon areole epidermal cells in lower left show elongated cells with sinuate anticlinal longitudinal walls. Remaining

cotyledon epidermal cells in upper right show more isodiametric cells with no such sinuate walls, (u) Cotyledon surface, SEM. Large cotyledon areole

contrasts with remainder of cotyledon surface on periphery.

c a = cotyledon areole, cot = cotyledon, cot p = cotyledon pocket, en = endosperm, h = hilum, mv = midvein, r = radicle, r p = radicle pocket,

t = testa. Abbreviations and translated description for (j, k, p) follow Beck (1878) and Dzikowski (1937). Scale bars = 100 pm in h-i, m-o, s-u 1 mm in

all others.

228

Journal of the Botanical Research Institute of Texas 5(1)

fruticosa is problematical. As with Hypocalyptus sophoroides, the endosperm in A.fruticosa is very large, and

the presumed cotyledon areole is also very large, covering most of the cotyledon abaxial surface, thus not

presenting a clear distinction between cotyledon areole epidermal cells and surrounding epidermal cells

(Fig. 2u). The difficulty of false negative reports for cotyledon areoles has been noted previously (Lackey

2008; Yaklich et al. 1989) for initial survey studies that were later followed by in-depth anatomical work,

particularly when seeds and cotyledon areoles are small. The same difficulty seems to occur when cotyledon

areoles appear to cover most of the cotyledon surface. Current results confirm the occasional difficulty of

form with this known problem. The most readily distinctive cotyledon areoles seem to be in those specimens

with relatively large seeds, average to small endosperm, and relatively small cotyledon areoles, such as the

Vicieae and Glycine spp., the very species in which the cotyledon areoles were first discovered. Detection of

lateral endosperm deposits was even less ambiguous than cotyledon areoles. However, absent exhaustive

observation, it may sometimes be possible to overlook small endosperm deposits if they are localized and

near the cotyledon base or apex.

To combine current results with past findings, requires a preliminary literature summary that is necessar-

ily lengthy, despite all attempts at brevity. The combination with recent literature (Endo & Ohashi 1998a,

1998b, 1999a, 1999b; Ma et al. 2004; Lackey 2007, 2008, 2009, 2010) and long-overlooked anatomical

findings (Schleiden & Vogel 1839b; Beck 1878), indicate a need to reassess some basic questions regarding

the structure and function of cotyledon areoles and endosperm in papilionoid legumes, and any possible

selective advantage.

Cotyledon areole structure

Three independent sources form the basis for detailed understanding of cotyledon areole anatomy: Beck’s

(1878) comparative anatomy of mature dormant Vicieae seeds, Ma et al.’s (2004) anatomical study of mature

dormant and germinating soybean seeds, and Endo and Ohashi ’s (1997, 1998a, 1998b, 1999a, 1999b) treat-

ments on developing, dormant, and germinating seeds of many papilionoid legumes. The three results have

never been combined. None of these authors knew of each others work at the times of publication (Yasuhiko

Endo & Fengshan Ma, pers. comm. 2009), however, Endo and Ohashi (1998a) and Ma et al. (2004) both

knew about Miksche’s (1961) brief description of the cotyledon areole in soybean. Ma et al. (2004) also ref-

erenced a substantial body of work on soybean cotyledon areoles (Dzikowski 1936, 1937; Yaklich et al. 1984,

1986, 1987, 1989, 1992, 1995, 1996, 1998; Baker & Minor 1987). Despite their varied scope and emphasis,

and lack of awareness of each other’s work, the overlapping areas of these three sources present a generally

consistent description of the cotyledon areole, with variation from taxon to taxon and even from cultivar to

cultivar within a species. In some taxa, some attributes appear to be completely missing. The three sources

combined with Lackey (2007, 2008, 2009, 2010) characterize the cotyledon areole as a slightly dark, greenish

or yellow-green, oval to linear, protein body containing spot on each abaxial cotyledon epidermis, above the

midvein, sometimes branched, and sometimes in a depression. Cotyledon areole epidermal cells are usually

larger, sometimes smaller, but always differ in size and shape from other cotyledon abaxial epidermal cells,

do not form stomata, and have strongly convex outer periclinal walls which give a “granular” appearance

to the cotyledon areole surface. In surface view, the anticlinal walls parallel to the cotyledon axis are often

“wavy” (Ma et al. 2004) or sinuate parallel to the cotyledon midvein. The cotyledon areole epidermal celtV

walls have thicker outer periclinal walls and cuticle, and stain less readily than surrounding epidermal

cells. Each cotyledon areole epidermal cell contains one large proteinaceous, chlorophyll-colored body, or

“Aleuronkorn” (Beck 1878). Mesophyll cells between the cotyledon areole epidermal cells and the midvein

are generally smaller than other mesophyll cells and contain several smaller similar bodies or “Korner”

(Beck 1878), “granules” or “structures” (Endo & Ohashi 1999b). In alcohol, chlorophyll is extracted from

these chloroplasts more slowly than from others in the cotyledon, leaving a yellow-green, and eventually a

red-brown pigment. The cotyledon areole appears during late stages of seed development, reaches its full

229

expression in the mature dormant seed, and the plastids and other internal and external cell structures

degenerate upon germination. In a summary article, Endo and Ohashi (2010) proposed the Japanese name

‘Shiyomon’ for the cotyledon areole.

The current research conforms with the above description and adds differential imbibition damage

confirms prior suggestions (Lackey 2009, 2010) that cotyledon areole presence and position is correlated

with endosperm presence and position.

Most of the literature is ambiguous as to whether the cotyledon areole refers to a cotyledon surface

feature, and therefore merely the epidermal cells, or also includes the internal cotyledon cells between the

epidermis and midvein. It seems more useful to define the cotyledon areole in the broader sense, which

would include both the specialized epidermal cells and specialized cotyledon mesophyll cells, and this

broader definition is used here.

The structure of legume endosperm in the mature dormant seeds has been studied longer and in more

detail than the structure of cotyledon areoles. A huge body of significant 19 th century anatomical work,

beginning with Bronn’s (1822) detailed findings, is almost never cited now, and with trivial exception has

never been compared with recent findings. At least since perhaps Schleiden (1838) and Schleiden and Vogel

(1839a, 1839b) legume endosperm has two well-known functions with corresponding structures. In the

mature dormant seed it acts as a polysaccharide energy reserve for germination (Buckeridge et al. 2000),

but in early developmental stages it supplies nourishment of the developing embryo (Chamberlain et al.

not be reviewed.

In a remarkable, and strangely, seemingly modern work, the anatomical treatment of endosperm, or

albumin, by Schleiden and Vogel (1839b), carried out with the most rudimentary optical and mechanical

instruments (Schleiden 1849), and unfortunately rarely cited now, is an expansion of their parenthetic treat-

ment during their classic study of legume floral development (Schleiden and Vogel 1839a), and remains the

most comprehensive, and in many instances the most detailed, legume endosperm study to date. They estab-

lished, by means of development studies and anatomical examination of almost 300 different legume seeds,

many basic facts. First, the albumin of legumes is endosperm, i.e. of embryo sac origin, and not perisperm,

i.e. of nucellar or maternal origin. Also, the endosperm consists of cells, in accord with Schleiden’s (1838)

cell theory. They found that, contrary to common statements (Candolle 1825), endosperm is widespread

and often well-developed in legumes. Although often well-developed, it can be reduced to a greater or a

lesser degree, which in their opinion is caused by an enlarged embryo, which displaces and compresses the

endosperm. The endosperm development is generally greatest on the sides of the cotyledons, least on the

cotyledon margins and under the hilum in subfamily Papilionoideae, and most persistently present in the

transparent, swells in water, particularly hot water, is tasteless, and is harder in Caesalpinioideae and

Mimosoideae than in Papilionoideae. They described the anatomy of the tissue layers of endosperm in detail,

which when fully developed, consists of three layers. Although the details varied considerably in such a

vast family, exterior is the first tissue layer, usually a single row of regular cells with distinct cell contents,

now called the aleurone layer (Pammel 1899). A second or middle tissue layer consists of a few to many cell

layers, which is the chief constituent of the endosperm. The individual secondary cell walls are filled with

mucilage, to the extent that in some cells, the lumen may almost disappear. The cells are of many shapes, hui i

often elongated with the long axis towards the embryo, and are often arranged in radiating patterns with the

embryo as the center. The mucilage seems to be deposited in the secondary cell walls in layers, the mucilage

layer nearest the lumen sometimes denser, and is often traversed by pore canals to the primary cell wall.

The third or innermost endosperm tissue layer consists of a few layers of cells that are similar to the middle

layer, and sometimes intergrade with it, but are crushed by pressure of the immediately interior cotyledon

230

Journal of the Botanical Research Institute of Texas 5(1)

epidermis, and it is this third layer that is seen when only a thin layer of endosperm remains. Later research

(Sempolowski 1874; Chalon 1875; Beck 1878; Avetta 1884; Nadelmann 1890; Pammel 1899; Smith 1981,

1983; Meier & Reid 1977) has clarified, extended, confirmed and updated Schleiden and Vogel’s (1839b)

above summarized basic results about legume endosperm to conform with advances in basic biological

theory, but never contradicted it to any significant degree, with the exception of their theory of endosperm

reduction caused by embryo expansion.

The complex method of mucilage deposition in the secondary cell walls (Dhugga et al. 2004) was

studied by Nadelmann (1890) by light microscopy for many legume species and Meier and Reid (1977) by

TEM in fenugreek, Trigonellafoenum-graecum. Both confirmed Schleiden and Vogel’s (1839b) observations

that mucilage is deposited in secondary cell walls, i.e. cell walls that are formed after the cells have stopped

enlarging, and both reported deposition from vesicles formed within the cells. Meier and Reid (1977) show in

their figures 2 and 8, TEM photographs of endosperm cells before and after deposition of about 20 microns

broad, about the same as seen in the current study, although under different preparation protocols. Although

without scale indication, Nadelmann (1890) presents cell line drawings in three stages of deposition, which

indicate no apparent cell enlargement. This confirms prior observations that mucilage deposition, or lack

thereof, has no effect on increasing endosperm cell size, but merely is accommodated by the pre-determined

cell size. Both works also confirm centripetal step-wise deposition of mucilage in layers, starting with a posi-

tion adjacent to the middle lamella and proceeding inward, towards the lumen; the space occupied by the

mucilage causes a reduction in lumen size, finally resulting in a very small, sometimes almost non-existent

lumen, and possible cell death, with few remnants of cell contents. However Ma et al. (2004) report these

mature endosperm cells as still living in soybean, as do Tonini et al. (2006) for Sesbania virgata.

Anatomical findings in the current study conform with endosperm structure described above. Endosperm

is always found as a complete, although often very thin, envelope around the embryo, and is always visible

as a minute enlargement at the cotyledon margins and in the slit between radicle and cotyledons. When

prominent, it is always by enlargement on the lateral embryo sides, occasionally quite small and distinct,

as seen here in Vidajaba, and previously observed by Beck (1878) in tribe Vicieae and Bronn (1822) in pea

( Pisum sativum). Sometimes it occupies most of the seed, as in Hypocalyptus sophoroides. The material is dense,

almost crystalline, occasionally dark colored, and dissolves in water.

Endosperm in Glycine was visually unexceptional relative to other legumes, and in the wild and semiwild

soybeans seen here (G. soja and G. gracilis Skvortz., Fig. 3), endosperm was substantial. The descriptive en-

dosperm results of Ma et al. (2004), although describing a small lateral endosperm deposit, given the unique

name “antipit” in the cultivated soybean, are likewise unexceptional relative to other endospermic legumes.

The chemical structure and physiology of the endosperm mucilage has been subject to extensive study because

of its pervasive commercial importance in petroleum extraction, food, cosmetics, pharmaceuticals, textiles,

paper, and a wide variety of industrial uses (Buckeridge et al. 2000). Although potentially available from a

variety of legume species (Tookey & Jones 1965), current primary commercial sources are guar ( Cyamopsis

tetragonoloba). locust ( Ceratonia siliqua L.), and fenugreek ( Trigonellafoenum-graecum ). All of which are sub-

ject to supply and price uncertainties, and alternatives, such as the soybean, are eagerly sought (Anderson

1949; Dhugga et al. 2004; Lackey 2010). Prior to my studies, the parallel presence and absence of cotyledon

areoles and endosperm in papilionoid legumes has not been reported, so any functional relationship of the

two has not been explored.

Nadelmann’s (1890) physiological and anatomical study of legume endosperm mucilage covered 74

legume species. He concluded that the primary function of mucilage in the middle layer was a reserve car-

bohydrate, which is broken down during germination and moves to the cotyledons and becomes transitory

starch. He speculated that the endosperm may have other functions, such as a reservoir for water or other

materials. He found a complicated correlation between the degree of development of endosperm mucilage,

cotyledon storage carbohydrates, and cotyledon proteins and oils, in which the increase in material in one

group was associated with decreases in the others. Although he performed several microchemical tests on the

231

mucilage, and indicated that it is carbohydrate and is transformed into cotyledon starch during germination,

he did not indicate its composition, which is now known to consist of mannose and galactose in the form of

galactomannan. Avetta (1874) had previously demonstrated laboratory degradation of galactomannan into

sugars. Mitchell (1930) indicates authors after Nadelmann (1890) and before 1900 with knowledge of the

composition of galactomannan or “Mannogalaktan.”

Pammel’s (1899) study was primarily anatomical, but he did review current physiological literature,

and concluded that mucilage is the reserve energy source in legumes because it is the most condensed form

compared to starch, proteins, or fats. He also noted the hydrophilic nature of mucilage in endosperm in

his summary of contemporary thought: “When once the water passes the Malpighian cells and reaches the

endosperm the latter has a great affinity for it and additional amounts are taken up readily. . . . After the

endosperm has taken up water the aleurone layer secretes a ferment which dissolves the cell-walls, and the

soluble material is conveyed to the embryo.”

Reid and Bewley (1979) observed in Trigonellafoenum-graecum that removal of the endosperm seven hours

after hydration still allowed germination of the embryo, although with decreased gain in weight. Because

galactomannan breakdown and mobilization does not occur until much later, this observation demonstrated

that any possible energy function of the endosperm galactomannan is not necessary for germination, and that

galactomannan does not differ qualitatively from energy reserves in the cotyledons. Because the endosperm

was never removed prior to seven hours, any necessity for galactomannan during this early period remains

unknown. They confirmed the extreme hydrophilic nature of galactomannan and its role in buffering the

embryo against moisture loss.

Buckeridge et al. (2000) summarized galactomannan literature and intense current interest.

Galactomannans consist of a linear backbone of mannose residues with side branches of galactose residues.

Galactomannans with few side branches (more or less pure mannans) are hard and insoluble in water. As

branching increases, galactomannans are softer and more readily extracted with water, particularly hot water.

The Caesalpinioideae tend to have abundant endosperm with galactomannans bearing few side branches.

There is a trend within the Caesalpinioideae, and from Caesalpinioideae to Papilionoideae or to Mimosoideae

to decreased galactomannan yields and increased branching. Galactomannans are formed within the middle

endosperm cells in the Golgi (Dhugga et al. 2004) and then deposited by vesicles into the secondary cell

walls. In some plants, a decrease in branching, and therefore increased hardness or density, occurs in the

cell wall layer towards the lumen. Although mobilization of galactomannan during germination has been

intensively studied in only a few legumes, it seems to be initiated after radicle protrusion by three enzymes of

the aleurone layer, which hydrolyze galactomannan to mannose and galactose. In some legumes, hydrolytic

enzymes are also stored in protein bodies within the endosperm, in addition to the aleurone layer (Tonini

et al. 2010). Mannose and galactose are converted to sucrose in the endosperm, and it is transported to the

cotyledons, where it forms transient starch. Recent research activity (Reid et al. 2003; Edwards et al. 2004;

Dhugga 2003, 2005; Dhugga et al. 2004) demonstrates that legumes can be genetically transformed to alter

galactomannan formation.

In the context of the above integrated literature summary, current observations combined with recent

literature and a review of unusual galactomannan properties employed for commercial purposes suggest new

functional explanations for the plant as well as variations on the three traditional functional explanations:

energy reserves, water relations, and hardness for protection (Nadelmann 1890; Buckeridge et al. 2000).

Furnish energy reserves for germination

Galactomannans are known to provide energy to the germinating seed. Pammel (1899) concurs with previous

workers who advocated primarily a superior energy function for galactomannan, and concluded: “. . . the

plant stores away its food in the most condensed form to save space.” However, current energy calculations

do not support Pammel’s assertion. Galactomannan has about the same energy per gram as other carbo-

hydrates and protein, and less than half the energy per gram of oil (Lopes & Larkins 1993). As a source of

energy, Reid and Bewley (1979) showed that galactomannan has no obvious advantage over other energy

storage products in the cotyledons, such as oil, protein, or starch. Therefore, if galactomannan is a superior

232

y of supression oficefc

233

surrounding cotyledon epidermis, and thus less likely to be susceptible to bacterial invasion. Also, there

are no literature reports of bacterial invasion in the cotyledon areole area. Additionally, now that cotyledon

areole presence is known to occur widely outside of soybean, there seems no reason to suspect a correlation

between cotyledon areole presence and invasion by rhizobial bacteria.

Yaklich et al. (1996) suggested that the cotyledon areole may be involved with nutrient transport and

unloading to the developing embryo. For several reasons this is not likely. The cotyledon areole is initiated

late in seed development, reaching full expression in the mature dormant seed, then degenerating within

some days after germination (Endo & Ohashi 1999b; Ma et al. 2004). Cotyledon areole development, there-

fore, is not concurrent with the developing embryo. Also, the thick cuticle of the cotyledon areole would

seem to impede nutrient transport, rather than facilitate it. And the peculiar presence of chloroplasts with

pigments different from other cotyledon cells and protein bodies in mature endosperm does not suggest a

transport function during embryo development.

Yaklich et al. (1996) and Ma et al. (2004) suggested that the cotyledon areole may help align the en-

dosperm and embryo or maintain a round seed shape. This is also unlikely. With or without a cotyledon

areole, papilionoid embryos observed here remain consistently positioned within the endosperm envelope

and testa. The radicle pocket alone seems to ensure proper alignment. And in many species, cotyledon ar-

eoles are not in a depression (Endo & Ohashi 1998b; current results), which is contrary to this suggestion.

Current observations suggest that presence or absence of a depression seems to bear no relation to seed

shape or embryo alignment.

Yaklich et al. (1987) and Ma et al. (2004) suggested that the cotyledon areole functions m germination,

possibly for transport of nutrients to the cotyledons. Although not demonstrated, this is consistent with most

anatomy and known physiology. As shown in the current study, cotyledon areole and endosperm presence

or absence and position are almost always positively associated, which immediately suggests a functional

relationship. The position of the cotyledon areole along the midvein, or when the cotyledon areole is branched,

additionally along major branch veins, suggests a transport function, as do the modified mesophyll cells

between the cotyledon areole epidermis cells and the veins. It should be remembered that the veins in the

mature dormant seed are not fully functional and only acquire functionality some days after imbibition, as

seen in soybean (Miksche 1961), about the same time as galactomannan mobilization. The developmental

and degeneration timing of cotyledon areoles corresponds with the fully mature development of endosperm

and its degeneration, and the functional development of cotyledon vascular tissue. In conflict with an ex-

planation of a germination function, the thick cuticle over the cotyledon areole would suggest a barrier to,

not an enabler of transport, and the functional significance of distinctive chloroplasts, pigmentation, and

protein bodies in the cotyledon areole is an enigma. It is also possible that the cotyledon areole functions in

a different, and unknown, fashion during germination, such as a trigger, or as part of Varner et al.’s (1963)

unknown embryo axis factor, which was a requirement for proper cotyledon metabolism.

CONCLUSIONS

At this time, the reasons why most legumes have maintained, for tens of millions of years, galactomannan

bearing endosperm in the mature dormant seed is only a matter of speculation. Whatever the reason, it

must have substantial selective advantage for the plants, because the deposition of the material during seed

development and mobilization during germination are both precise and complicated systems involving

highly specific physical, chemical, and temporal activities. Similarly, the function of the cotyledon areole

is still speculative, but as shown here, the positive correlation of presence or absence, size, position, and

developmental timing with endosperm suggests a functional relationship with endosperm energy reserves

and germination.

Any legume seed structure, such as a cotyledon areole, must be viewed as but one component of a

whole functioning seed, which has many structural, physiological, and ecological requirements for dor-

mancy, germination, protection from predators, etc. Smith (1983) has shown through extensive survey that

legume seeds occur in two basic forms. About 60% (his form 1) have endospermic seeds with foliaceous

234

Journal of the Botanical Research Institute of Texas 5(1)

cotyledons and epigeal germination. About 30% (his form 4) have non-endospermic seeds with enlarged

storage cotyledons and usually hypogeal germination. The remainder (forms 3 and 4) have intermediate and

anomalous forms. Evolution from the presumed primitive form 1 to the derived forms has occurred many

times. The cotyledon areole would thus be a characteristic of form 1 seeds. How the cotyledon areole fits

into the functioning of this basic form 1 is, again, unknown.

It would be instructive to study further the functional role of cotyledon areoles for several reasons.

One is a general interest in seed physiology and ecology, with a possible role in legume germination strate-

gies. A comparison with the non-cotyledon areole and scarcely studied caesalpinioid and mimosoid legume

seeds could help one understand how these plants fulfill the functions of a cotyledon areole in papilionoid

legumes. A second interest would be to understand the reasons for development of a very elaborate system of

galactomannan accumulation and mobilization during the evolution of legumes in the early Eocene, a period

much hotter and more humid than now. And third, germination of agricultural legumes, such as soybean,

peas, and many others, is always a critical stage of the life cycle, and any role that the cotyledon areole may

have has implications of commercial importance. The soybean would seem to be an ideal candidate for a

comparative study of cotyledon areole function because the plant is well-studied, abundant material is readily

available, and the diversity of wild, semiwild, and cultivated plants presents a wide range of cotyledon areole

and endosperm extent from large to almost non-existent, although neither has been completely eliminated

through non-scientihc or scientific plant breeding.

The demonstrated correlation of cotyledon areoles with endosperm indicates that use of cotyledon ar-

eoles in systematic studies cannot be analyzed as an independent character, but must be viewed in relation

to other seed and germination structures and functions, especially the endosperm.

ACKNOWLEDGMENTS

The author thanks D. Anderson, M. Buckeridge, Y. Endo, M. Fleet, J. Kirkbride, F. Ma, R. Palmer, and G.

Reid for helpful advice and discussion. I thank Joseph H. Kirkbride, Jr. and one anonymous reviewer for

their detailed reviews.

REFERENCES

Anderson, E. 1 949. Endosperm mucilages of legumes occurrence and composition. Ind^D^fhem. 41 :2887-2890.

Avetta, C. 1 884. Richerche anatomiche ed istogeniche sugli organi vegetativi della Pueraria Thumbergiana Benth..

Ann. del 1st. Bot. di Roma 1 :201 -222 + 3 ic.

Baker, D.M. Minor. 1987. Frequency and comparative anatomy of the extended endothelium among

soybean plant introductions and weathering resistant genotypes. Crop Sci. 27:1 301 -1 303.

Beck, G. 1878. Vergleichende Anatomie der Samen von Vicia und Ervum. Sitzungberichte der Kaiserlichen

Akademie der Wissenschaften 77:545-579 + ic. 2.

Bentham, G. 1 865. Leguminosae. In: Bentham, G. and J.D. Hooker. 1 865. Genera plantarum. Reeve & Co., London.

Bronn, H.G. 1822. De formis plantarum Leguminosarum primitivis et derivatis. Heidelberg.

Buckeridge, M.S., S.M.C. Dietrich, and DU d Lima. 2000. Galactomannans as the reserve carbohydrate in legume seeds.

In: Gupta, Anil Kumar, and Narinder Kaur. 2000. Carbohydrate reserves in plants - synthesis and regulation.

Elsevier; Amsterdam et alibi. Pp. 283-31 6.

Candolle, A.P.D, 1825. Memoires^UrT'afamille des Legumineuses. A. Belin, Paris.

Carlson, J.B. and N.R. Lersten. 2004. Reproductive morphology. In: Boerma, H. Roger, and James E. Specht. 2004.

Soybeans: improvement, production, and uses. American Society of Agronomy, Madison, Wisconsin. Pp. 59-96.

Chalon, J. 1875. La graine des Legumineuses. Soc. Sci. Arts Lett. Hainaut 10:3-66:

an anatomical and autoradiographic study. Canad. J. Bot. 71:11 53-1 1 68.

Dhugga, K,S. 2003. Genes for galactomannan production in plants and methods of use. US patent application

20040143871.

MICROMORFOLOGIA DE LA LEMMA DE LOS GENEROS POLYPOGON,

xAGROPOGONY AGROSTIS (POACEAE) EN CHILE

Victor L. Finot*

Wilsonfl!^, Carlos M. Baeza,

Alicia Marticorena & Eduardo Ruii;

Universidad de Concepcion

Departamento de Produccion Animal

Facultad deAgronomfa

Casilla 537, Chilian, CHILE

vifinot@udec.cl

*Autor de correspondencia

Casilla 160-C, Concepcion, CHILE

wulloa@udec.cl, cbaeza@udec.d

amartic@udec.cl, eruiz@udec.cl

El genero Polypogon fue descrito por Desfontaines (1798). Pertenece a la familia Poaceae, subfamilia Pooideae,

tribu Poeae, subtribu Agrostidinae (Soreng et al. 2003). El nombre del genero alude a la morfologla de la

mflorescencia, una panlcula densa provista de muchas aristas (del gr. polys = muchos; pogon = barba) (Watson

& Dallwitz 1992). Las especies de Polypogon son conocidas por sus nombres comunes de “cola de zorro o

cola de conejo los que aluden tambien a la morfologla de la mflorescencia.

Polypogon incluye unas 26 especies (Tzvelev 1983; Renvoize 1998; Lu & Phillips 2006) de zonas tem-

pladas y templado-calidas de ambos hemisferios y en zonas tropicales en areas montanosas por sobre los

1.200 m de altitud (Giraldo-Canas 2004; Lu & Phillips 2006). Para America, Soreng et al. (2003) reconocen

11 especies, de las cuales 10 estan presentes en Chile. El hlbrido xAgropogon lutosus (Poir.) P. Fourn. ( Agrostis

stolonifera L. x Polypogon monspeliensis (L.) Desf.) fue tambien citado para el sur Chile (Osorno, Chiloe) por

Rugolo de Agrasar & Molina (1997). Comprende plantas anuales o perennes que habitan sectores humedos

y muchas veces arenosos o salobres, en las margenes de cursos de agua (Glenn 1987). Se caracteriza por sus

espiguillas unifloras que se desprenden a la madurez con una parte o todo el pedicelo, y carecen de exten-

sion de la raquilla; las glumas son mas largas que el antecio, escabrosas y usualmente aristadas; la lemma es

hialina, con 5-nervios y puede ser aristada o mutica; la palea puede ser tan larga como la lemma o menor

que esta, alcanzando aproximadamente la mitad de su longitud (Tzvelev 1983; Nicora & Rugolo de Agrasar

1987).

Polypogon presenta una estrecha afmidad con Chaetotropis (Kunth, 1829) del cual se diferencia por tener

J. Bot. Res. Inst. Texas 5(1): 237 -2

Journal of the Botanical Research Institute of Texas 5(1)

la palea de una longitud menor que la mitad del largo de la lemma (Nicora & Rugolo de Agrasar 1987). El

genero Chaetotropis ha sido aceptado, entre otros autores, por Kunth (1835), Desvaux (1854), Pilger (1920),

Skottsberg (1920), Bjorkman (1960), Nicora (1970, 1978, 1993), Marticorena & Quezada (1985), Nicora

& Rugolo de Agrasar (1987) y Morrone et al. (2005). Otros autores, en cambio, consideran a Chaetotropis

un sinonimo de Polypogon, como Kunth (1816), Trinius (1840), Steudel (1854), Martius & Eichler (1893),

Stuckert (1904), Hitchcock (1931, 1951), Macbride (1936), Rosengurtt et al. (1970), Navas (1973), Gould

(1983), Tzvelev (1983), Muller (1985), Clayton & Renvoize (1986), Tovar (1993), Renvoize (1998), Barkworth

(2007), Zuloaga et al. (2008), Finot et al. (2009).

El tratamiento taxonomico mas completo de las especies de Polypogon s.l. fue realizado por Muller (1985)

quien reconoce 10 especies reunidas en dos secciones: Polypogon sect. Polypogon tipihcada por P. r

con seis especies y Polypogon sect. Polypogonagrostis Asch. & Graeb. tipihcada por P. elongatm

especies. Muller (1985) caracteriza la seccion Polypogonagrostis como plantas perennes, con glumas aristadas

desde el apice, con lemmas de ca. 1.5 mm de largo, palea mas corta que la lemma que alcanza hasta 2/5-1/3

de su largo y epidermis de la lemma provista de una malla de engrosamientos de la pared periclinal de las

celulas epidermicas denominada “trichodium net” (Rugolo de Agrasar & Molina 1997a; Favret et al. 2007).

En la seccion Polypogonagrostis, Muller (1985) incluye a P. chilensis (Kunth) Pilger (= Ch. chilensis Kunth), P.

elongatus Kunth (= Ch. elongata (Kunth) Bjorkman), P. rioplatensis Herter (= Ch. rioplatensis (Herter) Bjorkman)

y P. imberbis (Phil.) Johow (= Ch. imberbis (Phil) Bjorkman). De estas, P. rioplatensis se considera actualmente

un sinonimo de P. imberbis (Soreng et al. 2003).

La seccion Polypogon fue caracterizada por Muller (1985) como aquella que comprende plantas anuales

o perennes, con glumas aristadas desde el apice o con el apice bilobado, aristado desde la incision de los

dos lobulos en que se divide el apice, lemma de hasta 1,5 mm de largo, sin trichodium net y la palea tan larga

como la lemma o poco menor que esta. En esta seccion incluye a P australis Brongn., P interruptus Kunth, P.

linearis Trin., P. maritimus Willd., P. monspeliensis (L.) Desf. y P. viridis (Gouan) Breistr. Con la excepcion de P.

rioplatensis, todas las restantes especies tratadas por Muller (1985) han sido citadas para Chile (Marticorena

& Quezada 1985; Soreng et al. 2003; Zuloaga et al. 2008). A estas especies hay que agregar P exasperatus

(Trin.) Renvoize (= Agrostis exasperata Trin.) y P. hackelii (R.E. Fr.) Renvoize (= Ch. hackelii (R.E. Fr.) Bjorkman)

transferidas a Polypogon desde el genero Agrostis por Renvoize (1998). Ambas especies debieran incluirse en

la seccion Polypogonagrostis por poseer la palea mas corta que la lemma.

util para distinguir las secciones Polypogon y Polypogonagrostis, o bien para distinguir el genero Chaetotropis

de Polypogon. No obstante, especies de otros generos como Agrostis L., Deyeuxia Clarion ex P. Beauv. (D.

parviseta Vickery) y Bromidium Nees & Meyen (B. hygrometricum (Nees) Nees & Meyen, B. ramboi (Parodi)

Rugolo), tambien presentan malla (Favret et al. 2007), asi como dentro de Agrostis carecen de malla las espe-

cies del subgenero Zinagrostis Romero-Garcia, Blanca & Morales-Torres (A. truncatula Park, A. reuteri Boiss.

y A. nebulosa Boiss. & Reuter), lo que hace que el valor taxonomico de la malla sea discutible.

Polypogon tambien presenta una estrecha ahnidad con Agrostis descrito por Linnaeus (1753). Estos

generos se distinguen porque en Agrostis las espiguillas se desarticulan sobre las glumas quedando estas

persistentes en la inflorescencia a la madurez de los frutos luego del desprendimiento del antecio; en Polypogon,

en cambio, las espiguillas se desarticulan con todo o parte del pedicelo, de modo que las glumas caen junto

con el antecio que queda unido al resto de la espiguilla a la madurez de los frutos (Rugolo de Agrasar 1982;

Clayton & Renvoize 1986).

El hibrido xAgropogon muestra caracteres intermedios entre Agrostis y Polypogon. En este genero el antecio

se desarticula y se desprende de las glumas como ocurre en Agrostis, pero las glumas aristadas y escabrosas,

que se desprenden con el pedicelo, recuerdan a Polypogon (Rugolo de Agrasar & Molina 1997b).

Los objetivos del presente trabajo fueron examinar y describir la epidermis abaxial de la lemma de

las especies estudiadas mediante microscopia electronica de barrido (MEB), comparar las especies sobre la

base de los caracteres micro y macro morfologicos de la lemma y discutir la posicion sistematica de algunas

especies en los generos Polypogon, Agrostis y xAgropogon.

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Journal of the Botanical Research Institute of Texas 5(1)

Tabla 2. Matriz basica de datos de los caracteres utilizados para el analisis multivariado. M = malla; AG = aguijones; ON = ondulacion de la pared anticlinal de las celulas largas; EN

= engrosamiento de la pared anticlinal de las celulas largas; AC= ancho de las celulas largas; TEM = tipo de engrosamiento de la malla; FEM = forma de los engrosamientos de la

malla; PC = longitud de los pelos del callus; AR = arista; na = no aplicable; DE= Desprendimiento de la espiguilla; RLPL=

RLPL

Polypogon sect. Polypogon

P. australis

P. interruptus

P. linearis

P. monspeliensis

4. insularis

A kuntzei

A. magellanica

masafuerana

A. stolonifera var. palustris

A. stolonifera var. stolonifera

xAgropogon

A. lutosus

6 1 2

los engrosamientos de la malla, engrosamiento de la pared anticlinal de las celulas largas y ondulacion de

la pared anticlinal de las celulas largas) y 4 a caracteristicas macromorfologicas (pelos del callus, presencia

o ausencia de arista, desarticulacion de la espiguilla y largo de la palea en relacion al largo de la lemma). La

las especies del genero Chaetotropis. La MBD fue analizada mediante analisis de conglomerados utilizando

el algoritmo de Ward, que entrega grupos mutuamente exclusivos, donde cada grupo incluye los miembros

con la mayor similitud otorgando la maxima homogeneidad interna (Ward 1963) y el coeficiente de distancia

de Gower, que permite el uso de variables cualitativas y cuantitativas simultaneamente (Gower 1966), para

la confeccion de un fenograma y un analisis de coordenadas principales; utilizando el programa estadistico

Infostat 2009 (Di Rienzo et al. 2009).

RESULTADOS

3), malla (Figs. 1-6)

y tricomas (aguijones) (Fig. 1A-D). No se observo la presencia de estomas, celulas cortas ni macropelos.

La longitud de celulas largas pudo ser observada solo en Polypogon sect. Polypogon (Fig. 7A-D), xAgropogon

lutosus (Fig. 7E-F), Agrostis kuntzei (Fig. 3A-B), A. magellanica (Fig. 3C-D) yA. philippiana (Fig. 4C-D), pues en

las restantes especies d e Agrostis y en Polypogon sect. Polypogonagrostis la presencia de malla impide distinguir

el largo de las celulas. En la seccion Polypogon las celulas largas miden de 60-160 pm de largo por 3-7 pm

241

Fig. 1 . Fotomicrograffas de MEB de la epidermis de la lemma en especies de Agrostis. A-B. Agrostis brachyathera A. (1 OOOx), B. (3000x) (Finot& Lopez

1442, CONC); C-D. Agrostis breviculmisC. (lOOOx) (Mihocetal.4437, CONC), D. (3000x) (Ruiz & Lopez 91 4, CONC); E-F. Agrostis glabra E. (lOOOx), F. (3000x)

(Dominguez 160, CONC); ag, aguijones; m, malla; pa: pared de celula larga.

1C-D), A. mertensii Trin. (Fig. 4A-B), Polypogon australis (Fig. 7A-B), P. interrupts (Tabla 2) y P. linearis (Fig.

7C-D) u ondulada en forma de U, como en xAgropogon lutosus (Poir.) P. Fourn. (Fig. 7E-F), Agrostis glabra

(J. Presl) Kunth (Fig. 1E-F), A. imberbis Phil. (Fig. 2A-B), A. inconspicua Kunze ex E. Desv. (Fig. 2C-D),

A. insularis Rugolo & Molina (Fig. 2E-F), A. kuntzei Mez (Fig. 3A-B), A. magellanica Lam. (Fig. 3C-D), A.

Journal of the Botanical Research Institute of Texas 5(1)

Fig. 2. Fotomicrografi'as de MEB de la epidermis de la lemma en especies de Agrostis. A-B. Agrostis imberbisA. (1 OOOx), B. (3000x) ( Marticorena & Matthei

898, CONC); C-D .Agrostis insconspicua C. (1 OOOx) (Teneb 577, CONC), D. (3000x) (Soreng & Soreng 7136 CONC); E-F. Agrostis insularis E. (1 OOOx), F. (3000x)

( Villagrdn&Aguila 6063, CONC). ag, aguijones; m, malla; e, engrosamiento de la pared anticlinal; pa, pared anticlinal.

masafuerana Pilg. (Fig. 3E-F), A. philippiana Rugolo & De Paula (Fig. 4C-D), A. scabra Willd. (Fig. 4E-F),

A. stolonifera L. var. palustris (Huds.) Farw. (Fig. 5A-B) y var. stolonifera (Fig. 5C-D), P. chilensis, P elongatus

(Fig. 6A-B), P. exasperatus (Fig. 6C-D), P imberbis (Fig. 6E-F). En A. brachyathera Steud., P. monspeliensis y

P. viridis se observaron celulas largas con paredes rectas y onduladas en la misma epidermis. Engrosamiento

especies de la sect. Polypogonagrostis (Fig. 6B) y en P australis (Fig. 7A-B). Carecen de engrosamiento en la

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Journal of the Botanical Research Institute of Texas 5(1)

stolonifera var. palustris (Fig. 5A-B) y var. stolonifera (Fig. 5C-D) con cerca de 16-25 aguijones por campo

visual (1800 pm 2 ) o muy escasos, como en Agrostis insularis (Fig. 2E-F), A. philippiana (Fig. 4C-D), P. australis

(Fig. 7A-B), xAgropogon lutosus (Fig. 7E-F) con 1-5 aguijones por 1800 pm 2 . Todas las especies de la sect.

Polypogonagrostis carecen de aguijones en la epidermis de la lemma (Figs. 5E-F, 6); en la sect. Polypogon fueron

observados solo en P. australis. En cambio, la mayor parte de las especies de Agrostis presentaron aguijones

FinotetaU

Fig. 5. Fotomicrografi'as de MEB de la epidermis de la lemma en especies de Agrostis y Polypogon sect. Polypogonagrostis. A-B. Agrostis stolonifera vat.

palustris A. (1 OOOx) (Villagrdn & Leiva 7557, CONC), B. (3000x) (Parra & Torres 669, CONC); C-D. Agrostis stolonifera var. stolonifera C. (1 OOOx), D. (3000x)

(Peterson & Soreng 15620, CONC); E-F. Polypogon chilensis E (3000x) ( Villagrdn 4899, CONC), F. (3000x) (Villagrdn etal. 6954, CONC). ag, aguijones; e.

epidermicos, con la exception de A. glabra (Fig. 1E-F), A. kuntzei (Fig. 3A-B) y A. magellanica (Fig. 3C-D).

En todas las especies las aristas de la lemma presentaron aguijones (no mostrado).

La malla fue observada en todas las especies de Agrostis (excepto A. kuntzei, A. magellanica y A. philip-

piana ) (Figs. 3A-D, 4C-D) asi como en la section Polypogonagrostis. Las especies pertenecientes a la section

Polypogon (Lig. 7A-D) y xAgropogon lutosus (Lig. 7E-L) no presentan malla. En general, la malla se presento

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Journal of the Botanical Research Institute of Texas 5(1)

Fig. 6. Fotomicrografi'as de MEB de la epidermis de la lemma en especies de Polypogon sect. Polypogonagrostis. A-B. Polypogon elongatus. A. (lOOOx),

B. (3000x) ( Tedane 522 1, SI); C-D. Polypogon exasperatus. C. (1 OOOx) (Araya 69, CONC), D. (3000x) (Levi 1742, CONC); E-F. Polypogon imberbis E. (lOOOx),

F. (3000x) ( Villagran & Leiva 7280, CONC). ag, aguijones; m, malla; pa, pared anticlinal.

como engrosamientos de contorno eliptico (Fig. 5E-F), subrectangular (Fig. 2F) o irregular (Fig. 4F). El eje

mayor del engrosamiento es siempre perpendicular al eje mayor de la lemma. La malla presento una estructura

compacta en A. breviculmis, A. glabra, A. insularis, A. masafuerana, A. mertensii, A. scabra, A. stolonifera y en

todas las especies de la sect. Polypogonagrostis. Se considero malla de tipo compacto cuando el espacio entre

engrosamientos fue menor que el eje menor del engrosamiento, paralelo al eje mayor de la lemma (Fig. ID).

Finotetal.,

247

Fig. 7. Fotomicrografias de MEB de la epidermis de la lemma en especies de Polypogon sect. Polypogon. A-B. Polypogon australis. A. (1 OOOx) (Pisano & Bravo

200, CONC), B. (3000x) (Gardner & Knees 6974, CONC); C-D. Polypogon linearis. C. (lOOOx), D. (3000x) (Bliss & Lusk 808, CONC); E-F. xAgropogon lutosus.

E. (1 OOOx), F. (3000x) (Rugolo de Agrasar 1218, SI). Ag, aguijones; cl, celulas largas; e, engrosamiento pared anticlinal; m, malla; pa, pared anticlinal.

malla se definio como laxo.

En Agrostis, la presencia de arista en el dorso de la lemma fue observada en A. brachyathera, A. kuntzei,

geniculada inserta en el medio del dorso de la lemma y que excede ampliamente la longitud de las glumas.

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Journal of the Botanical Research Institute of Texas 5(1)

i (Fig. 8),

250

Journal of the Botanical Research Institute of Texas 5(1)

En general, la seccion Polypogonagrostis comparte con Agrostis la presencia de malla. La malla ha sido

considerada un caracter de valor taxonomico importante para distinguir la seccion Polypogonagrostis de

la seccion Polypogon (Muller 1985) o para reconocer a Chaetotropis como un genero distinto de Polypogon

(Nicora & Rugolo de Agrasar 1987), as! como para distinguir Agrostis de Polypogon s.s. (Rugolo de Agrasar &

Molina 1997a). En general, nuestros resultados coinciden con la observation de Rugolo de Agrasar & Molina

(1997a), en que la malla permite la separation de Polypogon s.s. y Agrostis, con la exception de A. magellanica,

A. kuntzeiyA. philippiana, que no muestran malla sino celulas largas no engrosadas con paredes anticlinales

profundamente onduladas.

Nicora & Rugolo de Agrasar (1987) destacan que algunas especies de Agrostis como A. magellanica son

problematicas debido a su similitud con Chaetotropis. La principal diferencia entre Chaetotropis y Agrostis es

la desarticulacion de la raquilla. En Agrostis tipicamente desarticula sobre las glumas persistiendo estas en

la planta despues de desprendidos los antecios, mientras que en Chaetotropis las espiguillas caen completas

llevando una parte o todo el pedicelo (Rugolo de Agrasar 1982). Por otra parte, las especies de Agrostis y

Chaetotropis comparten la presencia de malla en la epidermis de la lemma. Si bien A. magellanica y A. kuntzei

se acercan a Chaetotropis por la desarticulacion de la raquilla, ambas especies muestran ausencia de malla.

La epidermis de la lemma en estas especies presenta celulas largas con paredes anticlinales profundamente

onduladas, caracter compartido unicamente por xAgropogon lutosus. De acuerdo con Rugolo de Agrasar &

Molina (1997b), este caracter acerca xAgropogon a Polypogon, no obstante, en Polypogon s.s. la epidermis de

la lemma posee celulas largas con paredes anticlinales rectas o con tenues ondulaciones, nunca profunda-

mente onduladas, como ocurre en xAgropogon o en Agrostis kuntzei, A. magellanica y A. philippiana. Nuestros

resultados acercan xAgropogon a Agrostis y situan a A. magellanica y A. kuntzei en Chaetotropis (Polypogon sect.

senalar ademas, que celulas largas profundamente onduladas, con una conformation similar a la descrita

para xAgropogon lutosus se han descrito en especies del genero Lachnagrostis Trin. (L. hillardierei (R. Br.) Trinjf-, 1

y Deyeuxia ( D . decipiens (R. Br.) Vickery, D. innominata D.I. Morris, D. scaberula Vickery) (Jacobs 2001).

Soreng et al. (2003) senalan que A. magellanica se encuentra en una position intermedia entre Agrostis y

el “grupo Chaetotropis” de Polypogon. Con respecto a A. kuntzei senalan su posible sinonimia con P. exaspera-

tus. Esta especie, transferida a Polypogon por Renvoize (1998) se acerca tambien al grupo Chaetotropis. Las

Figs. 8 y 9 muestran ambas especies asociadas a Chaetotropis ( =Polypogon sect. Polypogonagrostis), a pesar de

carecer de malla. Otros caracteres que A. magellanica y A. kuntzei comparten con la seccion Polypogonagrostis

son la desarticulacion de la raquilla, la longitud de la palea respecto del largo de la lemma y la ausencia de

aguijones en la epidermis de la lemma.

La clasibcacion infragenerica de las especies chilenas de Polypogon es la siguiente:

POLYPOGON Desf.

1. Polypogon sect. Polypogon:

3 DIUC 210.121.014-1.0 el fi

AS,lfi3V J

A.S. 1951..

s. Contr. U.S.Natl. Herb. 24:384-385.

of the United States (2 nd ed). U.S.D.A.

MacBride, J.F. 1 936. Flora of Peru. Field Mus. Nat. Hist. 1 3(1 ):1 -320.

254

Journal of the Botanical Research Institute of Texas 5(1)

BOOK NOTICES

Ruth Kassinger. 2010. Paradise Under Glass: An Amateur Creates a Conservancy Garden. (ISBN13:

978-0-06-154774-4, ISBN10: 0061547743, hbk.). William Morrow Imprint, HarperCollins Publishers,

10 East 53rd Street, New York, New York 10022, U.S.A. (Orders: http://harpercollins.com/books/

Paradise-Under-Glass-Ruth-Kassinger/?isbn=9 7 8006 1 547744) . $24.99, 368 pp„ 5V2" x Sfe

MISSOURI BOTANICAL GARDEN PRESS (ST. LOUIS, MISSOURI)

Sedges (Cyperaceae)

Robert EC. Naczi and Bruce A. Ford, Eds. 2008. Sedges: Uses, Diversity, and Systematics of the Cyperaceae.

(ISBN 978-1-930723-72-6, hbk.). Monographs in Systematic Botany from the Missouri Botanical Garden,

Volume 108. Missouri Botanical Garden Press, PO. Box 299, St. Louis, Missouri 63199-0299, U.S.A.

(Orders: www.mbgpress.org). $75.00, 298 pp., illus., 7" x 10".

J. Bot. Res. Inst. Texas 5(1): 254.2011

CARACTERIZACION Y USO DE “PIMIENTAS” EN UNA COMUNIDAD

QUILOMBOLA DE LA AMAZONIA ORIENTAL (BRASIL)

Luciano Araujo Pereira

Parte de la Tesis Doctoral del primer autor

Institute de Pesquisas

Jardim Botanico do Rio de Janeiro

Rua Pacheco Leao, 915, Horto, CEP: 22.460-030

Rio de Janeiro, RJ, BRASIL

lpereira@jbrj.gov.br

Massimo Giuseppe Bovini

Instituto de Pesquisas

Rua Pacheco Leao, 91 5

Horto, CEP: 22460-030

Rio de Janeiro, RJ, BRASIL

Gloria Estela Barboza

Instituto Multidisciplinario de Biologia Vegetal

(IMBIV-CONICET)

Facultad de Ciencias Quimicas

Universidad Nacional de Cordoba

Casilla de Correo 495, 5000, Cordoba,

ARGENTINA

gbarboza@imbiv.unc.edu.ar

Mara Zelia De Almeida

Universidade Federal da Bahia (UFBA)

Rua Barao de Geremoabo, S/A/, Ondina

CEP: 401 70-290, Salvador, BA, BRASIL

Elsie Franklin Guimaraes

Becario de Productividad del CNPq

Instituto de Pesquisas Jardim Botanico do Rio de Janeiro

Rua Pacheco Leao, 915, Horto, CEP: 22.460-030, Rio de Janeiro, RJ, BRASIL

eguimar@jbrj.gov. br

IXTRODUCCIOX

La conservation de la biodiversidad necesariamente involucra aspectos biologicos, ademas de sociales y

culturales, que ciertamente, proveen el conocimiento de las culturas locales que son fuertes elementos para

su conservation (Albuquerque 2005).

J.Bot. Res. Inst. Texas 5(1): 255-2

e los servicios de los

259

microscopio estereoscopico Leica MZ75, acoplado con camara clara y procesados a traves del metodo usual

en taxonomia (Fidalgo & Bononi 1984). El material comprobatorio fue depositado en los Herbarios BHCB,

CORD, HAMAB, HB, MBML y RB, siglas segun Thiers (2010); material de referencia de otras localidades

tambien fue analizado, cuando necesario, en el herbario del Museo Botanico de Cordoba (CORD, Argentina).

Ademas, fue elaborada una clave analitica de los taxones estudiados, para su reconocimiento. Para la distri-

bution biogeografica de las especies, se han seguido los conceptos de Cabrera & Willink (1980) y Cavalcante

& Major (2006).

RESULTADOS Y DISCUSION

Diversidad y taxonomia de las “pimientas”

Los resultados indican que, a pesar del gran numero de etnoespecies citado en las entrevistas por los es-

pecialistas (22 etnoespecies), no hubo un aumento considerable en el numero de especies cultivadas en

las quintas (siete especies); solo se observo la adicion de nuevos nombres populares para la mayoria de las

especies (Tabla 1).

Un informante calificado o especialista local aplica claramente distintos nombres a los diferentes

morfotipos que identifican cada etnoespecie, es decir que un unico informante reconoce etnoespecies

pertenecientes a una misma especie botanica. No hubo diferencias sigmhcativas entre los informantes de

una misma comunidad ni entre comunidades distintas ya que en general, los especialistas consultados tienen

un conocimiento arraigado de las “pimientas” y de sus usos.

Para el 75% de los especialistas locales (12), todos los nombres populares por ellos citados aluden a

diferentes morfotipos, cuyas variaciones se centran principalmente en la morfologia de los frutos (tamano,

forma y colores). Asi por ejemplo, para “pimienta” (Capsicum frutescens), la poblacion local, como en otras

regiones de Brasil, utiliza el tamano del fruto como un caracter importante para la determination de las

etnoespecies “pimenta malaguetinha” (fruto pequeno) o “pimenta malaguetao” (fruto grande). En cambio,

las etnoespecies (Tabla 1) que corresponden a C. chinense presentaron morfologia carpologica muy diversa

en relation con las descripciones existentes en la literatura (Reifschneider 2000; Barbosa et al. 2006), si

bien nuevos tipos son obtenidos por hibridacion natural en el campo o por diferentes ambientes de cultivo

(Carvalho et al. 2006). Tales variaciones dihcultaron asociar la etnoespecie recolectada en las quintas con

su nombre cientifico, por lo que se hizo un estudio mas detallado “in situ” en la epoca de floracion. De esta

manera, se logro constatar que a pesar de las diferentes formas, tamanos y colores que presentan estas

etnoespecies, todas corresponden a una misma especie (C. chinense).

La denomination de las etnoespecies acorde a las caracteristicas morfologicas que presentan en deter-

minadas localidades fue conhrmada por Nascimento-Filho et al. (2007) en una encuesta llevada a cabo en el

estado de Roraima (Brasil). Los autores ahrmaron que de 180 “pimientas” en el campo (163 cultivadas y 17

nativas), 78 morfotipos diheren por la forma, color y grado de pungencia. Estos morfotipos fueron atribuidos

a C. annuum, C. baccatum var. pendulum, C. chinense y C. frutescens, siendo los predominantes la “pimenta

malagueta” (C. frutescens), la “pimenta murupi” (C. chinense) y la “pimenta-olho-de-peixe” (C. chinense). Estos

resultados coinciden con los encontrados en este trabajo donde C. frutescens y C. chinense fueron las especies

mayormente citadas por los quilombolas de Curiau (Tabla 2).

En el area estudiada en el Area de Protection Ambiental (APA) del Curiau se registraron siete taxones

(cinco especies y dos variedades) pertenecientes a Piperaceae y Solanaceae (Tabla 1), a saber: Piper marginatum

Jacq. (Fig. 2C), P. tuberculatum Jacq. (Fig. 2D), Capsicum annuum L. var. annuum, C. annuum var. glabriusculum

(Dunal) Heiser & Pickersgill (Fig. 3), C. chinense Jacq. (Fig. 4A-C, Fig. 2 A), C. frutescens L. (Fig. 4 D-G) y

C. baccatum var. umbilicatum (Veil.) Hunz. et Barboza (Fig. 2B). Unicamente las dos especies de Piper crecen

en forma espontanea en los bosques del area estudiada (Pereira et al. 2007); ambas especies fueron citadas

por dos de los 16 especialistas entrevistados y solo como planta medicinal (Tabla 2).

260

Journal of the Botanical Research Institute of Texas 5(1)

Tabla 1 . Lista de las"pimientas"del APA del Rio Curiau (Macapa, Amapa, Brasil) y su lugar de cultivo.

Piper tuberculatum

C. annuum var. glabriusculum

Capsicum frutescens

pimenta malagueta, pimenta malaguetao, X

pimenta malaguetinha

CLAVES PARA LOS GENEROS DE LAS “PIMIENTAS”

1. Hierbas o subarbustos, con (1 — )2— 4 flores axilares en fasciculos; flores diclamideas, carpelos 2 (raro 3-4-

1. PIPERACEAE

Piper L.,Sp. PI. 1:28. 1753.

Subarbustos o arbustos, arboles pequenos, 1-10 m de alt. Tallo con ramas nudosas, profilo generalmente

caduco. Hojas alternas, simples, enteras, sesiles o pecioladas; peciolo provisto de vainas cortas, alargadas o

canaliculadas; nerviacion predominante camptodromo-acrodromo o broquidodromo, nervaduras secundarias

sulcado, liso, papiloso o fimbriado; bracteolas peltadas, redondeadas, triangulares, semi-lunares o calcifor-

mes, de margenes lisas o fimbriadas. Flores sesiles o pediceladas, aperiantadas, aclamideas; estambres 2-6;

ovario supero obovoide a ovoide, vistoso o no, sesil o no, estigmas 3-4. Drupas con pericarpo delgado,

con estigma persistente.

Genero pantropical con especies dioicas o monoicas para el Viejo Mundo y hermafroditas en America

(Tebbs 1993). Con gran distribution en las regiones tropicales y templadas de los dos hemisferios. En Brasil,

habitan cerca de 266 especies (Guimaraes & Giordano 2004).

numerosas variedades para la production de la “pimienta”; se usa ampliamente como condimento ademas

de medicinal (Tebbs 1993; Guimaraes & Monteiro 2006).

CLAVE PARA LA IDENTIFICACION DE LAS ESPECIES DE PIPER L. RECOLECTADAS EN EL AREA DE ESTUDIO

1 . Lamina foliar palmatinervada con base simetrica y cordada, margen densamente ciliado P. marginatum

1 . Lamina foliar penninervada con base asimetrica no cordada, margen eciliada P. tuberculatum

a; B. Caliz; C. Gineceo; D. Flor.

265

peninervadas (8-10 pares de nervios ascendentes), base asimetrica y no cordada, apice agudo, margen

eciliado; peciolos rojos, puberulos y papilosos, de 0, 5-1,0 cm de largo, vaina con alas hasta la lamina.

Espigas erectas, cortas, de 4-7 cm de largo. Estambres 4. Drupas tetragonales, ovoideas o subobovoideas,

lateralmente comprimidas, glabras, con 3 estigmas persistentes sesiles.

Distribucidngeogrdfica . — Por todo el continente americano. En Brasil, ocurre en los estados de Amazonas,

Amapa, Para, Maranhao, Piaui, Ceara, Rio Grande do Norte, Paraiba, Pernambuco, Mato Grosso y Rio de

Janeiro. Habita en areas perturbadas de bosques abiertos (= capoeiras), principalmente, en lugares humedos

de la region biogeografica del Cerrado (Cabrera & Willink 1980); en otras regiones de Brasil, crece en las

pendientes humedas de los bosques abiertos y en lugares pantanosos (= brejos), en altitudes aproximadas a

los 550 m.

En Rio Grande del Norte se emplea como condimento. En Amapa, donde crece espontaneamente, es

conocida como “pimenta-de-macaco” o “pimenta longa” (Tabla 1). En el area estudiada y en algunos otros

lugares del Estado, se usa como planta medicinal, principalmente como antiespasmodica.

2. SOLANACEAE

Capsicum L , Sp. Pi. 1:188. lJFJjdt,

Hierbas perennes, arbustos o subarbustos ca. 0,50-2,5(-3) m alt., glabrescentes a muy pilosos, a veces villosas.

Tallos frecuentemente fistulosos, generalmente con dicotomia evidente. Hojas simples, alternas, aisladas o

falsamente geminadas, pecioladas; lamina entera, ovada a estrechamente lanceolada; base abruptamente

cuneada a rotunda; apice agudo a acuminado. Flores axilares, solitarias o en fasciculos, actinomorfas,

pentameras por lo general, pedicelos erectos, semi-erectos o pendulos, a veces geniculados distalmente.

Caliz persistente, truncado, con (0-)5-10 dientes o apendices lineares, inconspicuos o muy desarrollados,

iguales entre si, que nacen por debajo del margen. Corola 5-partida en diferentes niveles, rotacea, estrellada,

raramente campanulada o urceolada, integramente blanca, amarilla o lila, o bien blanquecina con diferentes

combinaciones de manchas verdes, vinaceas, amarillas y violetas; tubo pequeno, lobulos 5 bien evidentes;

estambres por lo general 5, filamentos glabros, parte basal ensanchada adherida a la corola, formando un

canal nectarifero; anteras amarillas o violaceas; ovario supero, con anillo nectarifero basal, bilocular o

falsamente trilocular, ovulos 2-muchos por loculo; estilo obsubulado; estigma pequeno, aplanado. Frutos

bayas, deciduas o persistentes, conicos, globosos u ovoides, de color amarillo, naranja, rojo, verde o verde-

reniformes aplanadas, oscuras o claras; testa generalmente foveolada.

Capsicum es un genero americano de ca. 32 especies nativas (Barboza et al. 2011). Brasil es el pais con

la mayor concentracion de especies (Barboza & Bianchetti 2005), especialmente en la costa sureste (Est.

Espirito Santo a Rio Grande do Sul), con algunos representantes en el noreste brasilero y en la Caatinga

(Barboza et al. 2010).

Los frutos de la mayoria de sus especies contienen una gran variedad y cantidad de metabolitos (vita-

minas, carotenoides, minerales, proteinas, carbohidratos, grasas, fibras) pero los principales componentes

pungent* m la capsaicina y la dihidrocapscina (Bosland & Zewdie 2001; Manirakiza et al. 2003), los que

se acumulan en la epidermis secretora del septo (Filippa & Bernadello 1992).

1LAVE PARA IDENTIFICACION DE LOS TAXOlS^^^p CAPSICUM L. RE C OLECTAD AS EN EL AREA DE ESTUDIO

. Corola blanca con dos manchas amarillas o verdosas difusas en la base de cada lobulo y parte del limbo.

Gineceo con heteromorfismo estilar. Baya de coloracion rojo-anaranjada, umbonado-umbilicada

C. baccatum var. umbilicatum

. Corola blanca, amarillenta, verdosa o purpura, raramente violeta; si blanca, entonces desprovista de manchas

amarillentas o verdosas en la base de los lobulos: Gineceo sin heteromorfismo estilar. Baya de coloracion

variada, morada, roja, amarilla, amarillo-verdosa o anaranjada, no umbonado-umbilicada.

266

de 4-7 x 2-2,5 cm, \

® PI. 1:189. 1753. (Fig. 4D-G).

1,5-4 cm del

272

Journal of the Botanical Research Institute of Texas 5(1)

Rodrigues, E. 2009. Festas d'O -Curiau. www.ferias.tur.br/informacoes/307/macapa-ap.html.

Rodrigues, E. 201 0. Historico do Curiau e suas manifestagoes culturais. www.navegaramazonia.org.br/2005/09/1 9/

Silva, R.B.L. 2002. A etnobotanica de plantas medicinais da comunidade quilombola de Curiau, Macapa-AP, Brasil.

Dissertagao Mestrado em Agronomia-Universidade Federal Rural da Amazonia, Belem, www.iepa.ap.gov.br/

arquivopdf/etnobotanica_de_plantasmedicinaisdo_Curiau.pdf.

Souza, V.C. & FI. Lorenzi. 2005. Botanica sistematica: Guia ilustrado para identificagao das familias de Angiospermas

da flora brasileira, baseado em APG II. Nova Odessa: Plantarum, Sao Paulo.

Tebbs, M.C. 1 993. Piperaceae. In K. Kubitzki, ed.The familes and genera of vascular plants 2. Springer-Verlag, Berlin.

Tewksburry, J.J. & G.P. Nabhan. 2001 . Seed dispersal. Directed deterrence by capsaicin in chilies. Nature 41 2:403-404.

Thiers, B. [continuously updated]. Index Herbariorum: A global directory of public herbaria and associated staff.

New York Botanical Garden's Virtual Flerbarium. http://sweetgum.nybg.org/ih/. [Consultado el 21 junio 201 0]|

,1. n/fVfJ Paris, FI. Jacquemin, & R.R. Paris. 1 978. Flavonoides de Piper marginatum. PI. Med. (Sttutgart) 33:46-52.

CHROMOSOME NUMBER FOR ARCYTOPHYLLUM FASCICULATUM (RUBIACEAE)

A. Michael Powell

Department of Biology, Herbarium

Sul Ross State University

Alpine, Texas 79832, USA.

ampowell@sulross.edu

Science and Resource Management

Big Bend National Park

Panther Junction, Texas 79834, U.S. A.

allisonleavitt@gmail.com

A meiotic chromosome number of 2 n =

Arcytophyllum fasciculatum (A. Gray) Terrell & H. Rob.: Texas. Brewster Co.: Big Bend National Park, Upper

Burro Mesa Trail, 0.43 mi from Ross Maxwell Scenic Drive, 3875 ft., 16 Jul 2010, A. Leavitt 325, 326 (SRSC).

Buds were bxed in modified Carnoy’s Solution (4:3:1). Meiotic observations followed the use of standard

squash techniques and acetocarmine stain. Interpretation of meiotic configurations was complicated by 1)

small cell size, 2) consistently “clumped” chromosomes at metaphase I, anaphase I, and in meiosis II, and 3)

heteromorphic chromosomes (Fig. 1). Bivalents were dark-stained and often clearly differentiated at diaki-

nesis, prophase I; hundreds of chromosomal spreads were observed at this stage. Diakinetic configurations

were interpreted as consisting of 12 II, approximately (if not exactly) half of them larger “ring” bivalents

and half of them smaller “rod” bivalents.

Based on morphological considerations, Terrell and Robinson (2010) transferred Hedyotis intricata Fosberg,

a species of northern Mexico, Trans-Pecos Texas, and southern New Mexico, to Arcytophyllum Willd. ex

Schult. & Schult. f., a genus of 15 primarily Andean species (Mena 1990), concluding that A. fasciculatum

(= H. intricata ) is most closely related to A. thymifolium (Ruiz & Pavon) Standi, of Columbia, Ecuador, and

Peru. Apparently the only species of Arcytophyllum for which chromosome numbers have been reported pre-

viously is A. thymifolium ; 2 n = ca. 30 and 2 n = 36 (Federov 1969). Chromosome numbers have been useful

in evaluating relationships in tribe Spermacoceae (Groeninckx et al. 2009), which includes Arcytophyllum,

Hedyotis L., Houstonia L., Oldenlandia L., and segregate genera, and involves the aneuploid series x = 6,7, 8,

9, 10 JE 13, and 17 as reviewed by (Lewis 1962; Terrell 1991; 2001; Terrell et al. 1986; Church 2003). The

reported here, previously missing from the series, is relevant fentture phylogenetic considerations in

tribe Spermacoceae. Furthermore, the heteromorphic karyotype of A. fasciculatum suggests that chromosome

morphology might also provide important phylogenetic information in the lineage concerned.

ACKNOWLEDGMENTS

We are grateful to Martin Terry for the Resumen Spanish and to Martin and Ed Terrell for reviewing the

manuscript.

J. Bot. Res. Inst. Texas 5(1): 273 -2

274

Journal of the Botanical Research Institute of Texas 5(1)

?e hand drawing, meiotic chromi

ot Arcytophyllum fasciculatum {Leavitt 325): prophase I, diakinesis; interpreted as2n=12 heteromorphic

REFERENCES

Church, S.A. 2003. Molecular phylogenetics of Houstonia (Rubiaceae): Descending aneuploidy and breeding

system evolution in the radiation of the lineage across America. Molec. Phylogen. Evol. 27:223-238,

Federov, A.A. (ed.). 1 969. Chromosome numbers of flowering plants. Acad Sci. U.S.S.R., Komarov Botanical Institute,

Groeninckx, Ochoterena, C PerssoH, i KArehed, B. Bremer, S. Huysmans, and E. Smets. 2009.

Phylogeny of the herbaceous tribe Spermacoceae (Rubiaceae) based on plastid DNA data. Ann. Missouri

Bot.Gard. 96:1 09-1 32.'

Lewis, W.H. 1 962. Phylogenetic study of Hedyotis (Rubiaceae) in North America. Amer. J. Bot. 49:855-865.

Mena, V., P. 1990. A revision of the genus Arcytophyllum (Rubiaceae: Hedyotideae). Mem. New York Bot. Gard.

60:1-26,

Terrel|SJ|§ 991 . Overview and annotated list of North American species of Hedyotis, Houstonia, Oldenlandia

(Rubiaceae), and related genera. Phytologia 71 :21 2-243.

T rri . I ,. E.E. 2001 . Stenotis (Rubiaceae), a new segregate genus from Baja California, Mexico. Sida 1 9:899-91 1 .

Terrel^^^id H. Robinson. 201 0. Transfer of Hedyotis intricataXo Arcytophyllum (Rubiaceae). J. Bot. Res. Inst. Texas

4:625-626.

Terrell, : E.E # W-H. Lewis, H. Robinson, and J.W. Nowicke. 1 986. Phylogenetic implications of diverse seed types, chro-

mosome numbers, and pollen morphology in Houstonia (Rubiaceae). Amer. J. Bot. 73:1 03-1 1 5.

FLORA ENDEMICA DE NUEVO LEON, MEXICO Y ESTADOS COLINDANTES

Glafiro J. Alanis Flores 1 , Marco A. Alvarado Vazquez 2 ,

a’"’ 11 Ramirez Freire 1 ,

*Carlos G. Velazco Macias 1 Rahim Foroughbakhch Pournavab 2

1 Laboratorio de Manejo de Vida Silvestre

Departamento de Ecologla

Facultad de Ciencias Biologicas

Universidad Autonoma de Nuevo Leon, MEXICO

carlos.velazco@gmail.com

2 Departamento de Botanica

Facultad de Ciencias Biologicas

Universidad Autdnoma de Nuevo Leon

San Nicolas de los Garza, Nuevo Leon, MEXICO

RESUMEN

4-059-SEMARNAT-2001.

1-059-SEMARNAT-2001. Ecosystems with

Mexico es un pais megadiverso, se estima que ocupa el 4° lugar mundial en riqueza floristica (Mittermeier et

al. 1997; Villasenor 2003), la cual incluye casi 23,000 especies de plantas vasculares, con un endemismo di?T , !

52% (Rzedowski 1998). Esta diversidad esta distribuida en 2,804 generos (7.8% endemicos) y 304 familias

(Villasenor 2004), y se atribuye a factores fisicos como la orografia, climas, hidrologia y la ubicacion del pais

entre los reinos biogeograficos neartico y neotropical (Rzedowski 1978).

A pesar de que Mexico cuenta con mas de 2.3 millones de colectas botanicas y existen numerosas floras

regionales. aun no se cuenta con una flora nacional y se estima que un 30% del territorio aun no esta incluido en

algun estudio floristico formal (Sosa y Davila 1994). La necesidad de documentar la flora es fundamental para

respaldar numerosas investigaciones biologicas, tanto basicas como aplicadas (Vazquez- Garcia et al. 2004).

En cuanto a la diversidad del estado de Nuevo Leon se tiene el antecedente de un listado floristico elab-

orado por Rojas-Mendoza (1965), el cual incluye 148 familias de plantas vasculares, con un total de 1,484

especies o categorias subespecificas en 657 generos; esta lista floristica se incremento a 3,175 especies, 1,031

generos y 158 familias de plantas vasculares (Villarreal y Estrada 2008), mientras que otras estimaciones

senalan un total de 2,903 especies de plantas vasculares distribuidas en 157 familias y 910 generos, ademas

de 360 taxa infraespecificos, registrandose 110 especies endemicas para el estado (Velazco 2009).

El estudio de especies endemicas o consideradas como prioritarias para su conservacion presenta

avances en estados como Coahuila donde se estima existen 190 especies en esa categoria (Villarreal y Encina

J. Bot. Res. Inst. Texas 5(1): 275 -2

i 22 m de alto. Se 1<

maticas en la region, la preser

; que destacan el encino de £

282

Journal of the Botanical Research Institute of Texas 5(1)

Vegetacion edafica. — Se presentan en el estado y el norte de Mexico diversos afloramientos minerales entre

los que destacan aquellos de sulfato de calcio (yeso), la importancia de estas areas, asi como un listado de

especies de flora asociadas a este tipo de habitat es presentado por Johnston (1941); este tipo de sitios se

denominan comunmente “suelos yesoso” o “habitat gipsohlo” y se presentan por lo comun en valles inter-

montanos del Altiplano Mexicano y en en los taludes inferiores de la Sierra Madre Oriental, en ellos se puede

presentar casi cualquier tipo de asociacion vegetal, sin embargo destacan entre estas, la vegetacion desertica

ya sea de tipo rosetohla o microhla, las areas de pastizal natural, es comun observar tambien, que conforme

aumenta la elevation se comienzan a presentar elementos de bosque como Pinus y Quercus, asociados a

otros elementos arbustivos o semiarboreos como Arbutus, Buddleja, Mortonia y Krameria, entro otros. Para

el presente estudio, cuando sea necesario a cada tipo de vegetacion se agregara la denomination “gipsohlo”

para denotar la presencia de yeso en el habitat donde crece una determinada especie.

METODO

Se realizaron consultas bibliograhcas, asi como de bases de datos de la Red Mundial de Informacion sobre

Biodiversidad (REMIB) disponibles en la pagina de Internet de la Comision Nacional para el Conocimiento y

Uso de la Biodiversidad (CONABIO - http://www.conabio.gob.mx/remib/doctos/remib_esp.html), se genero

una lista de plantas vasculares cuya distribucion esta restringida a los limites politicos del estado de Nuevo

Leon y estados colindantes. Se obtuvieron datos de distribucion por municipio, asi como del habitat en donde

ocurren, para este ultimo se aplica la abreviatura “ND” en caso de que la informacion no se especihque en

las fuetes de informacion consultadas; su nomenclatura y sinonimia fue revisada en las bases de datos de

Tropicos (Jar din Botanico de Missouri - http:Zwww.tropicos.org) y The Internacional Plant Names Index

(Real Jardin Botanico de Kew, Herbario de la Universidad de Harvard y el Herbario Nacional de Australia -

http:/www.ipni.org). El arreglo seguido para los generos y familias, es el propuesto Mickel y Smith (2004)

para el grupo de las pteridohtas, Martinez (1963) para las gimnospermas y por ultimo para las plantas con

flor se sigue el arreglo del Grupo de Filogenia de las Angiospermas (Stevens 2010).

RESULTADOS

Se presentan en el estado de Nuevo Leon un total de 159 especies restringidas a los limites estatales, las cuales

se distribuyen en 106 generos y 45 familias de plantas vasculares, ademas de cinco categorias infraespecifcas

(Anexo 1). Las familias con mayor numero de especies endemicas al estado son Asteraceae, Cactaceae y

Madre Oriental hacia el sur y centro del estado (Fig. 2), de manera especihca en el municipio de Galeana

que presenta mas del 50% de las especies endemicas para el estado; Los ecosistemas con mayor diversidad

de especies endemicas son los bosques en sus diversas asociaciones (encino, encino - pino, pino - encino,

pino y coniferas) y zonas con sustratos gipsohlos. Se registran cinco generos cuya distribucion geograhca

se restringe a los limites estatales: Aztekium, Digitostigma, Geohintonia (Cactaceae), Strotheria (Asteraceae) y

Jaimehintonia (Themidaceae), donde al menos para la familia cactaceae no se presenta una situation similar

en otro estado de la Republica Mexicana. Solamente 13 especies cuya distribucion se restringe a Nuevo Leon

estan incluidas dentro de la NOM-059-SEMARNAT-2001 (SEMARNAT 2002), una de ellas en la familia

Al incluir taxa cuya distribucion geograhca abarca ademas los estados vecinos de Nuevo Leon

(Tamaulipas, Coahuila, San Luis Potosi y Zacatecas en Mexico y Texas en los Estados Unidos de America),

la cifra de especies endemicas a un nivel regional aumenta a 191, en 146 generos y 53 familias de plantas

vasculares (Anexo 2). Tambien se registra un incremento de 6 especies protegidas por la NOM-059-

SEMARNAT-2001, al ampliar el rango geograhco a los estados vecinos.

Asteraceae 42

Cactaceae 1 1

Lamiaceae 7

Fabaceae 6

Boraginaceae 5

DISCUSION Y ' ,,

Tomando en cuenta las estimaciones hechas por Villarreal y Estrada (2008) y por Velazco (2009) para la

totalidad de las especies de flora presentes en Nuevo Leon, las especies endemicas aqui reportadas, represen-

tan alrededor del 5 % de la flora de Nuevo Leon, la cual se alberga de manera particular en la Sierra Madre

Oriental, de manera especifica en los municipios de Galeana, Aramberri, General Zaragoza y Rayones, donde

tan solo el municipio de Galeana presenta mas de 50 % de los endemismos estatales. Esta riqueza puede

atribuirse en gran medida a la variedad de habitats y los factores climaticos presentes en la zona, los cuales

van desde zonas deserticas con poca precipitation hasta el limite de la vegetacion arborea en montanas con

mas de 3000 metros de altitud, pasando por bosques templados y subtropicales; ademas de que estos tipos

de vegetacion han pasado por una rica historia de cambios climaticos (McDonald 1998) dando tiempo a la

formacion de elementos unicos o sirviendo como refugio de especies.

Por otra parte, y a pesar de los diversos trabajos floristicos que se han realizado en el estado de Nuevo

Leon, los cuales abarcan poco mas de un siglo (Velazco 2009), se ha dado poca importancia al conocimiento

de las especies endemicas o con una distribucion restringida a nivel regional, salvo algunos casos senalados

en la normatividad federal (NOM-059-SEMARNAT-2001). Asi mismo, los botanicos han dedicado pocos

esfuerzos a entender las relaciones ecologicas de estas especies, su biologia basica o dar un seguimiento pos-

terior a su situation taxonomica, teniendo por ejemplo el caso de Viguiera nesomii Turner, la cual fue descrita

a partir de un solo ejemplar (Turner 1989), de manera subsiguiente no se ha publicado nada referente a esta

especie, la cual actualmente esta reconocida por el “Catalogo taxonomico de especies de Mexico” (Sarukhan

et al. 2009); lo anterior nos plantea un claro ejemplo del nivel de desconocimiento de las especies endemicas

y por lo general mas vulnerables a extincion.

Tomando en cuenta que menos de 20 especies de este listado de endemismos estatales y regionales

estan incluidas en la normatividad federal bajo alguna categoria de protection, este tipo de listados, puede

servir tambien como base para la generation de normas estatales que tengan efectos directos en la toma de

decisiones por parte de las autoridades locales. Asi mismo, es importante resaltar la importancia de las areas

naturales protegidas (ANPs), por ejemplo el Cerro del Potosi, declarado desde el ano 2000 como un ANP

de caracter estatal con una extension de 989 has. (Anonimo 2000), lo cual se constituye como una herra-

mienta basica para la protection de especies como Festuca hintoniana (Poaceae), Potentilla leonina (Rosaceae),

y Astranthium beamanii (Asteraceae), entre otras que ocurren en la pradera alpina y subalpina, asi como los

bosques de coniferas. En este mismo rubro, el Parque Nacional Cumbres de Monterrey (de caracter federal),

abarca una mayor superficie (177,395 has.) de la portion norte de la Sierra Madre Oriental, donde se destacan

las siguientes especies endemicas: Notholaena leonina (Pteridaceae), Agave albopilosa (Agavaceae), Mirandea

hua tecen i (Acanthaceae) y Lobelia sublibera (Campanulaceae).

Las listas de especies endemicas y su categorization de acuerdo a su grado de vulnerabilidad siguiendo

estandares nacionales o internacionales, deberia ser una prioridad entre los botanicos de Mexico; pero la

falta de recursos humanos y materiales es uno de los principales factores que limitan esta investigation. El

Journal of the Botanical Research Institute of Texas 5(1)

Notholaena leonma Maxon tontf USNatt Herb 16:58.1912.

43(2):308. 1989.

Distribucion: Allende, Monterrey y Santa Catarina, N.L.

Habitat: matorral submonl

HSbrtat: matorral gipsofilo.

Gibasisgypsophila B.LTurner, Phytologia 75(5):406-407.

Distribudoa Aramberri, Galeana y General Zaragoza, I'

Habitat: matorral gipsofilo y bosques de pino encino.

Distribi

Turner, Phytologia 75(5):407-408.

Doctor Arroyo y General Zaragoza,

GIMNOSPERMAS

lonimo: Trodescantia pexata H.E. Moore, Baileya 4:1 00. 1

stribucion: Guadalupe, Mier y Noriega, Monterrr

Rayones, N.L.

NOM-059-SEMARNAT-200 1 :EmfeIig||gY .

LILIOPSIDA

Agavaceae

Agave albopilosa I.Cabral, Villarreal & A.E. Estra

Mex. 80:52. 2007.

Santa Catannti§|' , t

i nuevoleonensis Matuda, Anales Ir

Nac. Mexico 26:72.1955.

Sinonimo: Tradescantia potos,

181056 - 1976 .

Agave ovati folia G.D.Starr & Villarreal, Sida 20(2):495-499. 2

Distribucion: Bustamante, N.L.

Habitat: bosques de encino - pino.

Eleocharis rzedowski S. Gonzalez, Phytol

Distribucion: Galeana, N.L.

um ownbeyi Traub, PI. Life 24(2-4): 139. 1

oitat: Matorral desertico.

um traubii T.M. Howard, PI. Life 23:62, 91

Amaryllidaceae

Zephyranthes howardii Jraub, PI. Life 1 9(1 ):49. 1963.

‘ 1fet|p imo: Hobranthus howardii (Traub) T.M. Howard, Herbertia

46(1-2) "

Distribucion 4pf|§irrey, N.L.

if/'/Traub&T.M. Howard, PI. Life 26(1 ):62.

Tiy Galeana, Nil •

I B.L. Turner, Phytologia 75(4):277-279,

)erri y General Zaragoza, N.L.

syrinchium microbracteatum G.L. Nesom, Phytologia

76(6):470. 1994.

istribucion: Aramberri, Galeana, General Zaragoza y

Rayones, N.L.

syrinchium novoleonense G.L. Nesom & L. Hern., Phytologia

73(6):430. 1992.

istribucion: Allende, Aramberri, Doctor Arroyo, Galeana y

General Zaragoza, N.L.

iemiphylacus hintoniorum L. Hern., Syst. Bot 20(4):549. 1 995.

)istribucion: Galeana, Iturbide y Rayones, /> , t ,

286

Journal of the Botanical Research Institute of Texas 5(1)

Distribudbn: Galeana, Santiago y San Pedro Garza Garcia, il bosque de encjnb -ipino y coniferas

Habitat: bosque de e§^J||ppconiferas.

Orchidaceae

Brachystele chiangii (hA.C Johnst) Bums-Bal.,0rquidea (Mexico

Sinonimos: Spiranthes chiangii M.C. Johnst., Phytologia 45:449.

1980; Mesadenus chiangii (M.C. Johnst.), Garay Botanical

Museum Leaflets 28(4):336. 1 982.

Galeottiella hmtomorumTodzia, Brittonia 46(4):332. 1994.

Sinonimos: Brachystele hintoniorum (Todzia) Espejo & Lopez-

Ferrari Phytologia 82(2):79. 1 997; Microthelys hintoniorum

(Todzia) Szlach., Rutk. & Mytnik, Fragm. Florist. Geobot.

41(6)>476-477. 2004.

Distribucion: General Zaragoza, N.L.

Mirandea huastecensis T.F. Daniel, Syst. Bot. 3(4):428.

1978(1979]

Distribucion: Santa Catarina, N.L,

322, f. 1. 1995.

Acourtia hintoniorurrm^mner, Phytologia 75:404. 1993.

itl^fj-iucion: Aramberri, N.L.

Sinonimo: Stipa hirticulmis S.L. Hatch, Valdes-Reyna & Morden,

Distribucion: Galeana, N.L.

Distribucion: Galeana, ^

Habitat: bosque de pino.

\Auhlenbergia jaime-hintonii P.M. Peterson & Valdes-Re

Distribucion: Aramberri y General Zaragoza, N.L.

Habitat: matorrales y bosque de pino gipsofilos.

°oa mulleri Swallen, J. Wash. Acad. Sci. 30(5):21 1 . 1 940.

35(4):375,f. 1.1983.

Jaimehintonia gypsophila B.L. Turner, Novon 3(1 ):86. 1993.

Habitat: matorral desertico gipsofilo.

MAGNOLIOPSIDA

Ageratina nesomii B.L. Turner, Phytologia 53:241 . 1 983.

|S§ipfjmo: Ageratina rollinsif^^mrnet, Phytologia 53:2|I|J

1983..

Ageratina potosina B.L. Turner, Phytologia 64:20. 1987.

^^wfeUddrr; Galeana y General Zaragoza, N.L.

Habitat: bosque de e|p|j||' pino.

Ageratina viejoana B.L. Turner, Phytologia 75(2):147. 1993.

Distribucion: Aramberri, N.L.

Astranthium splendens DeJong, Publ. Mus. Michigan State

Univ., Biol. Ser. 2(9):51 1-514. 1965.

Distribucion: Aramberri, Doctor Arroyo, Galeana, General

Habitat: bosque de encino - pino.

Brickellia aramberrana B.LTurner, Phytologia 75(2):1 41 . 1993.

Distribucion: Aramberri, N.L>. , ' ’ ’

Habitat: bosque de pino.

Sinonimo: Erigeron gypsophilus B.L. Turner, Wrightia 5(5):1 18.

oyo, Iturbide y Galeana, N.L.

) gipsofilo.

Distribucion: Rayones, N.L.

Habitat: ND.

Acanthaceae

Justicia hintoniorum G.L. Nessom, Phytologia 73(2):140. 1992

Distribucion: Aramberri e Iturbide, N.L.

Erigeron pattersonii G.L. Nesom, Phytologia 76(1 ):96-99. 1 994.

> Distribucion: Galeana y Rayonds, N.L,

Habitat: matorral gipsofilo.

Flyriella leonensis (B.L. Rob.) R.M. King & H. Rob., Phytol

24:69. 1972.

.Sinbhimos: Eupatorium iconense B.L Rob., Proc. Amer. A

Arts 36:479. 1901; Eupatorium chrysostyloides B.L. F

Distribution: Gale

Grindeliaobovatifc

Senecio powellii B.L. Turner, Phytologia 75(4):326-328. 1993.

Distfibucforc Galeana y Mina, N.L.

Habitat: rmatorral gipsofilo.

Solidago ericamerioides G.L. Nesom, Phytologia 67(2):143.

Distribucion: Galeana, N.L.

Habitat: vegetacion riparia, matorral gipsofilo.

Steviopsis nesomii B.L. Turner, Phytologia 68:410. 1990.

Sinonimos: Brickelliastrum nesomii (B.L. Turner) R.M. King & H.

1 995.; Brickelliastrum villarrealii

'., Phytologia 76(1 ):1 7. 1 994.

ri. Doctor Arroyo y General Zaragoza,

otheria gypsophila B.L. Turn

i, Phytologia 68(4):3 13-315.1 990.

20(2)27-30. 1975.

p||j|n: Galeana, N.L

aitat: pastizales gipsofilos.

Heterotheca gypsophila B.L. Turner, Phytologia 55:206. 1984.

Distribucion: Galeana, N.L.

Isocoma gypsophila 1972

Distribucion: Aramberri y Galeana, luk.

r, Phytologia

', Phytologia

ner, Phytologia

Distribucion: Aramberri, Galeana e Iturbide, N.

Habitat: matorral y bosque de pinof- ,

Perymenium hintoniorum B.L. Turner, Phytolo

1991.

Distribucion: Aramberri, General Zaragoza, Iturb

N.L.

Habitat: matorral y chaparral.

Pinaropappus pattei

r, Phytologia 73(4)261-

r, Phytologia 65(5};373. 1

Phytologia 77:283. 1 994.

Sinonimo: Aster gypsophilus B.L. Turner, Souths I'

1$jgjf&74.

Distribucion: Galeana, N.L.

Habitat: pastizal gispofilo.

Tagetes mulleri S.F. Blake, J. Wash. Acad. Sci. 32:150. 1

EgBL

erff) Melchert, Phytologia

Sinonimo: Bidens muelleri Sherff, Publ. Field Mus. Nat. Hist.,

Bot. Ser. 1 6(2):645-646. 1937.

^^^Galeana, N.L.

Tridax hintoniorum B.L Turner, Phytologia 73(5):350. 1992.

Distribucion: Galeana, N.L.

Habitat: matorrales gipsofilos, bosque de pinos.

Verbesina aramberrana B.LTurner, Phytologia 75:134. 1993.

Distribucion: Aramberri, General Zaragoza, N.L.

Habitat: matorrales gipsofilos.

Verbesina hintoniorum B.L. Turner, Brittonia 37(1 ):96,f.1 .1985.

t: matorral gipsofilo.

no olsenii B.L. Turner, PI. Syst. Evol. 150(3-4)254-255.

'ones y Santiago, N.L.

t: matorrales ybosques.

nazaragosana B.L.Turner, Phytologia 73(4):302. 1992.

i: Montemorelos, N

J illustrations of plants

>:rt92S>

Suninimo: Thelocactus tulensis subsp. buekii (Klein) N PTaylor,

Cactaceae Consensus Init. 5:14. 1 998.

Distribucion: Aramberri y Galeana, N.L.

Habitat: matorral y bosque de pino.

Thelocactus conothelos sobsp. argenteus (Glass & R.A. Foster)

Glass, Guia de Identificion de Cactaceas Amenazadas de

Distribucion: Aramberri y Doctor Arroyo, N.L.

Calcaratolobelia pringlei (B.L. Rob.) Wilbur, Sida 1 7(3)

Sinonimos: Heterotoma pringlei B.L. Rob. Proc. Ai

Arts 44(21 ):61 5. 1909; Lobelia gypsophila TJ. J

Calcaratolobelia rr

nm.) Wilbur, Sida 17(3):564.

& R.A. Foster, Cact. 5

Caryophyllaceae

Arena ria gypsostrata B.LTurner, Phytologia 75(6):481. 1993.

Distribucion: Aramberri, Mina y Rayones, N.L.

Habitat: matorral gipsofilo.

Sinonimo: Turbinicarpus schmiedickeanus subt

& R.A. Foster) Glass, Guia de Identificion de Cactaceas

Amenazadas de Mexico. 1997.

5 Aramberri, N.L.

NOM-059-SEMARNAT-200 1 :En tfeligrcj (?)

Turbinicarpus hoferi Luthy & A.B. Lau, Kakteen Sukk. 42(2):37.

nbooleanum B.L. Turner, Phytologia 79(1):31. 1995.

oucibn: Rayones, N.L.

at: matorral gipsofilo.

nhintoniorum pypftner, Phytologia 78(6):405. 1 995),

NOM-059-SEMARNAT-2001 :Amenazada (A)

Turbinicarpus saueri subsp. septentrionalis K

Kaktusy; Zpravodaj Svazu Ceskych ,

40(3):85-91 . 2004.

^^^iC^'Hidalgo, N.L.

Euphorbiaceae

Euphorbia correlli M.C. Johnst., Wrightia 5(5):1 30. 1 975.

Sinonimo: Tithymalus correllii (M.C. Johnst.) Sojak, Cas. Na

Mus., Odd. Phr. 148:199. 1979

Distribucion: Galeana, N.L.

Habitat: matorral gipsofilo.

Euphorbia neilmulleri M.C. Johnst., Wrightia 5(5):1 25. 1 97^;

Sinonimo: Tithymalus neilmulleri (M.C. Johnst.) Sojak, Cas. Na

„ Odd. F

Distril

:1 99. 1979

i, Galeana, Gem

290

Journal of the Botanical Research Institute of Texas 5(1)

ragalusregiomontanus Barneby, (V

Phytologia 76(4):295. 1 994.

Habitat: bosque de encino, pino y coniferas.

Myrospermum sousonum A. Delgado & M.C. .

Phytologia 79(2)97 19S&TI

Distribucion: Doctor Arroyo, General Zaragoza, flurbtife'y

Santiago, N.L.

Habitat: bosque de encino y bosque de pino.

Salvia jorgehintoniana Ramamoorthy ex B.LTurner, Phytologia

79(2):80. 1995.

Distribucion: Doctor Arroyo, General Zaragoza, fturbrefp ( y

Santiago, N.L.

Habitat: bosque de encino - pino.

Sophorajuanhintoniana B.L. Turner, Phytologia 76:385. 1994.

Distribucion: Aramberri, N.L.

Habitat: matorral gipsofilo.

Frankenia gypsophila I.M. Johnst., J. Arnold Arbor. 20:237. 1 939.

Habitat: pastizal gipsofilo.

Geniostemon gypsophilum B.LTurner, Phytologia 76(iy.l 1 —13* '

1994 '.,

Scutellaria aramberrana B.LTurner, Phytologia 76(5):352. 199-

Distribucidn: Aramberri, N.L.

Scutellaria hintoniorum Henrickson, Aliso 1 2(3)51 7-5 1 9. 1 98‘

Distribucion: Galeana, N.L.

Distribucion: Aramberri, Galeana y General Zaragoza, N.L.

r, Phytologia 80(2):1 1 8. 1 996.

Hedeomaciliolata (Epling) R.S. Irving, Brittonia 22:345. 1970.

Distribucion: Doctor Arroyo, Galeana e Iturbide, N.L.

Hedeoma palmeri var. galeana B.L. Turner, Phytologia

71 (1 ):33— 34. 1991.

Distribucion: Galeana e Iturbide, N.L.

Pinguicula immaculata Zamudio & Lux, Acta Bot. Mex.

20:40-44, f. 1-2. 1992.

Distribucion: Aramberi, Galeana, General Zaragoza y Rayones,

7i gypsogemum G.L. Nesom, Madrono 3U:25

ibucion: Doctor Arroyo, .N.L.

tat: matorral gipsofilo.

)istribucion: Aramberri, Galeana y Rayones,

Monarda pringlei Fernald, Proc. /

Distribucion: Galeana, f

Nyctaginaceae

ii B.LTurner, Phytologia 70(1 ):44. 1

DIODIA SAPONARIIFOLIA (RUBIACEAE: SPERMACOCEAE),

ESPECIE DISYUNTA ENTRE SUDAMERICA Y MEXICO

Lucio Lozada Perez

Plantas Vasculares, Facultadde Ciencias

Universidad Nacional Autonoma de Mexico

Ciudad LJniversitaria A. P. 70-283

MEXICO, D.F. 04510

luciolozada@hotmail.com

Claudia Gallardo Hernandez

Instituto de Ecologta A.C.

Carretera antigua a Coatepec 351, El Haya

Xaiapa 91 070, Veracruz, MEXICO

1 NT RO DU C: Cl O N

Diodia L. s. str. (Rubiaceae: Spermacoceae), comprende cinco especies americanas, distribuidas princi-

palmente en terrenos humedos, anegados, pantanosos o arenosos, bordes de arroyos, lagunas o rios. Son

hierbas postradas, rastreras, de hojas opuestas con vainas estipulares casi siempre bien desarrolladas, sus

flores axilares, 1 a 5, isomorfas, con corolas mfundibuliformes, de lobulos pilosos en la superficie interna,

estilo y ramas estigmaticas filiformes y frutos indehiscentes (Bacigalupo y Cabral 1999).

Diodia saponariifolia (Cham. & Schltdl.) K. Schum., se conocia unicamente de Sudamerica, en particular

de Brasil, Paraguay y recientemente de Argentina en los limites con Brasil. En el presente trabajo se registra

su presencia en los estados de Puebla y Veracruz, Mexico. Varios especimenes colectados desde 1974 en el

estado de Veracruz, y depositados en los herbarios ENCB, MEXU y XAL, se habian identificado erroneamente

como Diodia maritima Thonn. (sinonimo de Diodia serrulata (P. Beauv.) G. Taylor), y mas recientemente

ha sido considerada como Diodella serrulata (P. Beauv.) Borhidi, (Borhidi 2006). Diodia maritima Thonn.

se describio de Africa y su distribucion se ha extendido a algunas playas del Caribe, Belice, Guatemala y

Nicaragua (Burger & Taylor 1993; Standley & Williams 1975; Taylor 2001). Se caracteriza por tener 4 sepalos

y estigmas capitados; a diferencia de los especimenes examinados que poseen 2(3) sepalos y estilos bifidos

con ramas estigmaticas filiformes.

Diodia saponariifolia (Cham. & Schltdl.) K. Schum., in Martius, Fl. Bra. 6(6): 16. 1889. (Fig. 1). Borreria

Hierbas rastreras, suculentas; tallos verdes a rojizos en algunas partes en fresco, oscuros cuando secos, teretes,

de hasta 2 m de largo, de 4-6 mm de diametro, glabros, con entrenudos de 2-7 cm de largo; estipulas unidas

en una vaina estipular, bianco verdosa en fresco y cafe claro en seco, de 6-9 mm de largo, glabras, con 3 setas

rojizas, glandulares en el apice, triangulares, la central de 4-5 mm de largo, las laterales de 0.3-1. 5 mm,

glabras en el dorso, ciliadas en el margen, alternando con coleteres en la base. Hojas opuestas y decusadas,

subsesiles, peciolos de ca. 1 mm de largo; oblanceoladas a elipticas, apice agudo a muy corto acuminado,

margen rojizo, entero, estrigiloso, gradualmente adelgazandose a hacia la base, redondeada en la base, de

J. Bot. Res. Inst. Texas 5(1): 299 - 302. 201 1

300

Journal of the Botanical Research Institute of Texas 5(1)

Fig. 1.0/01

, de 5-5.5 mm de largo y 1.5-2 mm de

de 5-6 mm de largo y ca. 1.2

0 en el ano de 1978 en una \

302

Journal of the Botanical Research Institute of Texas 5(1)

AGRADECIMIENTOS

Los autores agradecen a los curadores de los herbarios ENCB, MEXU y XAL por permitirnos la revision de

los ejemplares citados y a Laura Padilla por la elaboracion del dibujo. Los autores desean agradecer a David

REFERENCIAS

Bacigalupo, Cabral. 1 999. Revision de las especies americanas del genero Diodia. Darwiniana 37:1 53-1 65.

Borhidi, A. 2006. Rubiaceas de Mexico. Akademiai Kiado, Budapest.

^^,W.C&C.M. Taylor. 1993. Flora Costaricensis: Family #202. Rubiaceae. Fieldiana, Bot. n.s. 33:T-333.

Standley, P.C.^^ Williams. 1975. Rubiaceae. In: Flora of Guatemala. Fieldiana, Bot !

Taylor, C.M. 2001 . Rubiaceae. In: W.D. Stevens et al., eds. Flora de Nicaragua 3:2206-2284.

E.W, Lathrop. T9 70. Pilularia americana on the Santa Rosa Plateau of Riverside County, California.

Aliso 7:14f§fi|

CHRYSOTHAMNUS BAILEYI (ASTERACEAE) NUEVO PARA NUEVO LEON, MEXICO

Jose A. Villarreal-Q.

Departamento de Botanica

Universidad Autonoma Agraria Antonio Narro

Buenavista 1 923, Saltillo 253 15

Coahuila, MEXICO

e-mail: javillarrealOO@hotmail.com

George S. Hinton

Rancho Agumlla

San Rafael, Galeana

Nuevo Leon, MEXICO

Eduardo Estrada-C.

Facultad de Ciencias Forestales

Universidad Autonoma de Nuevo Leon

A.P. 41, Linares 67700

Nuevo Leon, MEXICO

RESUMEN

ABSTRACT

del genero Chrysothamnus para la flora del estado, cuya existencia no habia sido registrada previamente

(Henrickson y Johnston 1997; Villarreal 2001). La especie al parecer es C. haileyi Wooton & Stand., la cual

fue descrita en 1913 de un ejemplar colectado por V.O. Bailey 498 en Nuevo Mexico. En 1923 fue transferido al

siguiente nombre: Chrysothamnus pulchelus (A. Gray) Greene ssp. bailey (Wooton & Stand.) Hall & Clements,

posteiormente en 1995 a Ericameria pulchella var. bailey (Wooton & Stand.) L.C. Anderson y recientemente

en el 2005 a Lorandersonia bailey (Wooton & Stand.) Urbatsch, R.P. Roberts & Neubig.

,61 and the Hwy 3, 14.1 mi al S 100°28'24.96"W, 23°30'0.74"N, 26 Oct 1981, Poole, Nixon & Smith 2508 (TEX).

La especie es un arbusto de 30-50 cm de alto que crece en matorrales y bosques de pino con Pinus cembroi-

des, Quercus intricata, Arctostaphylos pungens, Viguiera greggii, Chrysactinia mexicana, Helianthella gypsophila,

Porophyllumfiliforme, Dalea uniflora, Cheilanthes spp., entre otras. Es poco comun en el area y forma colonias

esparcidas, creciendo tambien en suelos yesosos.

Chrysothamnus baileyi es conocido en los Estados Unidos de Arizona, Colorado, Kansas, Nuevo Mexico,

Oklahoma, Utah y de Trans-Pecos en Texas, y en Mexico del noroeste-centro de Coahuila (Nesom in prep.;

Urbatsch et al. 2005). Los nuevos reportes son del sur de Nuevo Leon por lo que la distribution conocida

se extiende ahora hasta el sur de la region del Desierto Chihuahuense.

AGRADECIMIENTOS

Agradecemos la ayuda de Thomas Wendt por proporcionar informacion de ejemplares en la coleccion del

herbario de la Universidad de Texas en Austin, al B.L. Turner por la revision del escrito.

J. Bot. Res. Inst. Texas 5(1): 303 - 304. 201 1

ECOLOGIA DEL BOSQUE DE OSTRYA VIRGINIANA (BETULACEAE)

DE LA SIERRA DE ZAPALINAME, COAHUILA, MEXICO

Juan A. Encina D., Francisco J. Eri^$j£ D. y Efren Mata R.

Departamento Forestal

Universidad Autdnoma Agraria Antonio Norro

Buenavista 253 15 Saltillo, Coah., MEXICO

juanencina@gmail.com

ABSTRACT

Ostrya virginiana K. Koch es un arbol caducifolio de 15 m o mas de altura, con copa de forma piramidal, nativo

de regiones templadas y humedas de Norteamerica hasta zonas tropicales montanosas de Centroamerica

(Islebe et al. 1994; Felger et al. 2001). Su distribution comprende amplios bosques en Canada y el Este

de Estados Unidos de America, donde forma parte del bosque deciduo templado o bosque de latifoliadas

(Nixon et al. 1997). En Mexico se distribuye en el bosque mesofilo de montana, en los estados de Jalisco,

Michoacan, Queretaro, Hidalgo, Veracruz, Guerrero y Chiapas (Rzedowski 1978, 1996), en la Sierra Madre

Occidental se reporta para el suroeste de Durango (Gonzalez et al. 2007), suroeste de Chihuahua y sureste

de Sonora (Felger et al. 2001). En el noreste de Mexico tiene distribution reducida y presenta bosques ais-

lados (Valdez et al. 2003), en Tamaulipas se reporta para la sierra de San Carlos (Briones 1991) y la reserva

de la Biosfera El Cielo (Puig et al. 1983); en Nuevo Leon se presenta en el centro-sur, en los municipios de

Galeana, Santiago y Zaragoza (Hinton y Hinton 1995; Villarreal y Estrada 2008). Para el estado de Coahuila

se tienen reportes de colecta en las sierras del Carmen, Santa Rosa y La Gloria, ubicadas en el centro-norte

de la entidad (Villarreal 2001). Actualmente Ostrya virginiana es una especie listada en estatus de conser-

vation segun la Norma oficial mexicana NOM-059-SEMARNAT-2010 (SEMARNAT 2010), bajo la categoria

de Sujeta a protection especial; sin embargo, su distribution ha sido afectada de manera importante por la

tala inmoderada y la destruction del habitat con fines agricolas y ganaderos (Luna 2003); a pesar de ello

se carece de estudios enfocados a conocer el estado actual de sus poblaciones que permitan establecer las

estrategias mas adecuadas para su conservation a largo plazo en el pais. La Sierra de Zapaliname es un area

natural protegida bajo la categoria de Zona Sujeta a Conservation Ecologica, decretada por el gobierno deb

estado de Coahuila (Anonimo 1996). Se pretende estudiar la estructura y ecologia del bosque de Ostrya

virginiana de la Sierra de Zapaliname y con ello promover su conservation.

La sierra de Zapaliname se localiza en el sureste de Coahuila, en los municipios de Saltillo y Arteaga; es una

zona de transition entre el Desierto Chihuahuense y la Sierra Madre Oriental. Se ubica entre los 25°15'00"

- 25°25'58.35" de latitud Norte y entre 100°47'14.5" - 101°05'3.8" de longitud Oeste (Fig. 1). Pertenece a la

J. Bot. Res. Inst. Texas 5(1): 305 -3

306

Journal of the Botanical Research Institute of Texas 5(1)

)6reas presentes en el bosque de Ostrya virginic

i lo sucesivo IDR)

aS. Marines G. y J.J. C

| 20

Ostrya virginiana

Y/////X Sideroxylon lanugi

insssssi Fraxinus cuspidata

RXXX&a Cercis canadensis

n n

1 ( cm )

S.A. Mexico. D.F.

3 of thmmfa del Carmen, USA and l\

BEE FLORA OF AN INSULAR ECOSYSTEM IN SOUTHERN BRAZIL

Cassia Monica Sakuragui

Departamento de Botanica

Universidade Federal do Rio de Janeiro

Rio de Janeiro - BRASIL

Av. Brigadeiro Trompowsky s.n. Ilha do Fundao

CEP: 21 941-490

cmsakura 12@gmail.com

Emi Rainildes Lorenzetti

Departamento de Fitopatologia

Universidade Federal de Lavras

Caixa Postal 3037

CEP: 37200-000

Lavras -MG -BRASIL

elorenzetti@gmail. com

Rafael Augusto Xavier Borges

Centro Nacional de Conservagao da Flora

Instituto de Pesquisas Jardim Botdnico do RJ

Rio de Janeiro - BRASIL

Rua Pacheco Leao 915

Rio de Janeiro RJ CEP 22460-030

rafaelborges@jbrj.gov. br

Eloi Machado Alves

Departamento deZootecnia

Universidade Estadual de Maringa

Maringa -PR- BRASIL

Av. Colombo, 5790

Maringa - PR, CEP: 87020-900

eloimachadoalves@yahoo.com.br

Angela Maria Janunzzi

Universidade Estadual de Maringa

Av. Colombo, 5790

Maringa - PR, CEP: 87020-900

Maringa -PR- BRASIL

amjcorsi@uem.br

Vagner Arnaut De Toledo

Departamento deZootecnia

Universidade Estadual de Maringa

Av. Colombo r 5790

Maringa - PR, CEP: 87020-900

abelha.vagner@gmail.com

RESUMEN

INTRODUCTION

Insects, especially bees, comprise one of the main groups of angiosperm pollinators, and, as such, they play

a vital role in the biology of plant communities, being mainly responsible for gene flow among individuals

and populations. From this perspective, considerable research is in progress to increase the knowledge

about bee flora and its interaction with the pollinators (Rodarte et al. 2008; Mendonga et al. 2008; Vieira

et al. 2008). The diversity of Brazilian flora, given the territorial extension and climatic variability of the

country, greatly influences the potential of beekeeping (Marchini et al. 2004).

Bee flora is defined by the number of plant species that bees use as a source of nectar and pollen for

J. Bot. Res. Inst. Texas 5(1): 311 -3

312

Journal of the Botanical Research Institute of Texas 5(1)

their survival and production of honey (Pereira 1990). While bee flora in Brazil is rich and varied, there is

still insufficient information on beekeeping and the plants used by honey bees. Most data available come

from studies from temperate zones (Almeida-Anacleto 2007). Identification of plant species visited by bees

is important for beekeepers because such information can optimize honey production through the use of

appropriate sources of nectar and pollen (Alcoforado-Filho 1993). Furthermore, knowing the floral source

of honey enables beekeepers to characterize their products (Almeida-Anacleto 2007). The pollen found in

honey is an indicator of its botanical origin and, through quantitative analysis, it is possible to establish the

relative contribution of each plant to honey production (Iwama & Melhem 1979). Knowledge of the botani-

cal and geographical origin of pollen sources (Almeida-Anacleto 2007) is used in this type of analysis, and

it is also a requirement of the international quality control of honey.

Brazilian apiculture has been steadily growing, and it is internationally renowned for its control methods

of Africanized honey bees, as well as the large diversity and quality of its manufacturing products (centrifuge,

tanks, cylinders to produce molded wax, and bee hives), and the production, mainly organic, of such bee

products as honey, pollen, royal jelly and propolis. For those reasons, Brazil has potential for the production

of organic honey, which, if properly certified, can be competitive in the international market (Buainin &

Batalha 2007). There are several projects for the production of organic honey now underway in the country.

In Parana State, for example, it is estimated that more than nine thousand certified beehives exist, with an

average annual production of 400,000 kg of organic honey by 75 beekeepers. It is estimated that the islands

of the upper and middle Parana River have more than 15,000 productive hives. World production of honey

by the 15 major producing countries grew 30.15% from 1994 to 2005. Over the last ten years, Brazil has

been the country with the largest expansion of honey exports, both in value and quantity. In 2006, the

production reached 38 thousand tons, representing R$ 120 million. (IBGE 2009).

The Laranjeiras and Floresta Islands are located in a protected area of the upper Parana River floodplain.

The southern region of the area can be considered the last remnant of “varzeas” in the Parana River, free

of dams (Campos 2001). Several floristic and phytosociological studies have been carried out in the area,

(e.g., Campos et al. 2000; Campos & Souza 2003). The basis for consolidated beekeeping depends on sound

information about local flora and its phenology, as well as the determination of apiculture potential (Silveira

1983).

This study aimed to identify the plants visited by bees in a protected area of the Parana River, which

is known as the Parque Nacional da Ilha Grande, and characterize them, taking into account the floral

morphology, to provide better information for the management of bee colonies in the studied area.

The Ilha Grande National Park (PNIG) is a Unity of Conservation for Integral Protection. It covers ap-

proximately 78,878 ha of semideciduous forests at the borders of Parana and Mato Grosso do Sul States in

Brazil (IBGE 1992). The study area is located in the PNIG and is bounded by the Laranjeira and Floresta

islands near the municipality of Querencia do Norte (22°55'00"S, 53°35'00"W and 22°48'00"S, 53°25'00"W,

respectively). The region is dominated by lakes, ponds and about 300 islands (Campos 2001). Since this

region contains the Biodiversity Corridor of the Parana River, it is important for the conservation of the

natural resources present in the area. Most of the original vegetation was destroyed, giving way to coffee

plantations and cattle fields. The park protects the last stretch of the Parana River free of dams in Brazil. For

those reasons, the PNIG is internationally recognized as a core area of the Biosphere Reserve of the Atlantic

Forest (MaB Programme of UNESCO). The park protects areas used by migratory birds and rare species,

as well as both endemic and endangered flora and fauna. The area also provides several types of economic

activity for the local population. Particularly important to this work, it is estimated that there are about

10,000 bee boxes within the park, according to beekeepers of Parana. This activity has generated additional

income for local residents, and it has grown in importance in a regional e

MATERIAL AND METHODS

The collection sites at Laranjeira and Floresta islands are indicated in Figure 1. The climate of the region,

313

according to the classification of Koppen, is type Cfa - humid, subtropical (Maak 1968). The rainfall is

distributed throughout the year, with higher precipitations from September to December and a drier season

from June to August (Caviglione et al. 2000).

The samples were collected monthly, between August 2005 and August 2006. Trails (12 for Laranjeira

Island and 10 for Floresta Island) were established throughout the study area, especially near the apiaries.

A plant was considered visited by Apis melifera only when the bee was found within its flowers. All plants

visited by Apis mellifera were collected. In the field, notes were taken to describe the color and symmetry of

the flowers. In the laboratory, the symmetry of all corollas was reanalyzed and classified as actinomorphic,

zygomorphic or asymmetric (Bell 2008).

All fertile material was collected according to the standard botanical procedures (Bononi et al. 1984;

Liesner 2010). Specimens were deposited into HUEM and identified using specific literature and compari-

son with collections held at RB andif§ff§J| (acronyms follow Holmgren et al. 1990). Whenever necessary,

taxonomy specialists were also consulted. Duplicates were sent to RB. The organization of families followed

APG II (2003).

RESULTS AND DISCUSSION

The studied area presents a high degree of deforestation as a consequence of its use for pasture land and fires

that have occurred throughout the last three decades (Campos & Souza 2003). Between 1999 and 2003,

314

Journal of the Botanical Research Institute of Texas 5(1)

52 fires were documented in the IGNP, and most of them reached meadow areas and fragments of seasonal

semideciduous forest (Abreu et al. 2004). During the development of this study, part of Laranjeira Island

was also struck by fire in September, 2005. In spite of both fire damage and deforestation, plant species

visited by bees were found in all habitats: gallery forest, meadow, paludiculous and aquatic (terminology for

riparian formations according to Souza & Kitta 2007). Several species were typical of seasonal semidecidu-

ous submontane vegetation, possibly indicating a reasonable degree of preservation of local flora: Guarea

macrophylla Lam., Trichilia pallida Sw. (Meliaceae), Balfourodendron riedelianum (Engl.) Engl. (Rutaceae),

Crysophyllum lucentifolium Cronq. (Sapotaceae), Nectandra cissiflora Ness, Ocotea langsdorffii (Meiss.) Mez

(Lauraceae), Eugenia punicifolia Kunth (DC) (Myrtaceae) and Inga sessilis (Veil.) Mart. (Fabaceae). On the

other hand, other species were considered invasive plants, evidencing the impact of human activities in the

invasive, but also as apiculous plants. Vernonanthura phosphorica (Veil.) H. Rob. (Asteraceae), Lantana trifolia

(Verbenaceae) and Crotalaria micans (Fabaceae) have all been considered as important bee plants (Lorenzi

2000). V. phosphorica and some species of Mikania were listed in the Directory of Important World Sources,

confirming the importance of the family Asteraceae among the families visited by bees.

Pfaffia glomerata (Amaranthaceae), one of the visited species, is sometimes considered as invasive as

well as an important medicinal plant found in the area. Since it is also one of the first species to appear in

areas affected by fire, it is considered ecologically important as a pioneer species (Taschetto & Pagliarini

This study found 135 bee-visited species distributed among 104 genera and 53 families in the area

(Table 1). Besides the importance for beekeepers, this information is also useful for plant ecology studies,

since bee-visiting can be a good proxy for pollination. For example, Freitas and Sazima (2006) verified that

Apis mellifera was responsible f^p^inating 31 out of 124 bee-visiting species. Using the number of species

as a metric, the most important families were Asteraceae (15), Rubiaceae (10), Fabaceae (7) and Solanaceae

Croton, Passiflora, Gouania and Psycothria presented three species each. Asteraceae, Myrtaceae, Rubiaceae,

Euphorbiaceae and Lamiaceae are the most important botanical families for A. mellifera in the neotropics

(Ramalho et al. 1990). Locatelli and Machado (2001) pointed out the importance of Asteraceae as a particu-

larly rich family of plant species visited by bees. They suggested that the great number of species and the

wide geographical distribution of the family favor this fact.

Comparatively, the Fabaceae, one of the most diverse families in the studied area, was highly represented

(seven species) in the bee flora of a sandbank area in Bahia, northeastern Brazil (Viana et al. 2006). This

family was also among those with the largest number of species (14) visited by bee species in a transition

area among seasonal deciduous forest, savanna and ombrohlous forest in Castro Alves, Bahia (Carvalho &

Marchini 1999).

Although the studied area was near Maringa (about 250 km), the results of the present study are very

different from those of Toledo et al. (2003). These authors found that Anacardiaceae and Sterculiaceae were

the most visited families by Apis mellifera in Maringa, probably reflecting the choice of an urban environment

for their bee flora study.

With respect to pollen, geographical distribution of plants and honey production, the pollen of Allophyllus

(Sapindaceae) and Casearia (Salicaceae) was considered by Ramalho et al. (1991) to be a good geographical

indicator of honey collected in southern Brazil. The genera were considered an important honey source for

that region and important for beekeepers in the tropical Americas.

Only Maranta divaricata was considered asymmetrical, while the remaining visited plants presented

actinomorphic (83%) and zygomorphic flowers (17%). In the study of Freitas and Sazima (2006) involving

124 species, radial symmetry (actinomorphic flowers) occurred in 69% of the flowers versus 26% of flow-

ers with bilateral symmetry (zygomorphic flowers). According to Zoucas et al. (2004), Apis mellifera prefers

actinomorphic flowers. Since our study concluded that the majority of plants in the study area presented

315

Table 1 . Plants visited by Apis mellifera in the Laranjeiras and Floresta Islands. Hab - Habit, Infl - Inflorescence, HUEM (registration number in the Herbarium of the

University of Maringa), HER- Herb, SUB- Subshrub, SBR- shrub, CRE- Climber or Liana, TRE-Tree, ACT- Actinomorphic flower, ZIG- Zygomorphic flower, ASY- Asymmetrical

flower, RED- Red, WHI- White, CRE- Cream, YEL- Yellow, ORA- Orange, PUR- Purple, PIN- Pink, VIN- Vinaceous.

Chamissoo macrocarpa (Kut

Pfaffia glomerata (Spreng.) f

Pfaffia paniculata (Mart.) Ku

Guatteria campestris R.E. Fr

Jabermontana catharinensis A.

Aristolochia cymbifero Marrt. & (

Achyrocline saturejoides (Lam.) [

Ageratum conyzoides L.

Conyza bonariensis (L.) Cronq.

Chromolaena laevigata (Lam) R

Pluchea sagittalis (Lam.) Cabrera

Porophyllum ruderale (Jacq.) Cass.

Schkuhria pinnata (Lam.) Kuntze

Verbesina encelioides (Cav.) H. & B.

Vernonanthura phosphorica (Veil.) H. Rob.

Chrysolaena platensis (Spreng.) H.Rob.

Heliotropium cf. angiospermum Murr;

Heliotropium transalpinum Veil.

Protium heptaphyllum Marchand

Trema micrantha (L.) Blume

Cheiloclinium d.cognatum (Miers.) A

Combretum fruticosum (Loefl.) Stuntz

Commelina erecta L.

Ipomoea cairica (L.) Sweet

Ipomoea tiliacea (Willd.) Choisy

Merremia digitata (Spreng.) Hallier f.

Doliocarpus dentatus (Aubl.) Standi.

Tetracera breyniana Schlech.

Erythroxylum pulchru

di glgndulosu

Croton floribundus Be

Apocynaceae HER

Apocynaceae CRE

Apocynaceae TRE

Aristolochiaceae CRE

Asteraceae SUB

Boraginaceae CRE

Boraginaceae HER

Burserceae TRE

Cannabaceae ,f|l j

Celastraceae CRE

Combretaceae CRE

Commelinaceae HER

Convolvulaceae CRE

Dilleniaceae CRE

Elaeocarpaceae

Erythroxylaceae TRE

Euphorbiaceae SCR

Euphorbiaceae

Euphorbiaceae TRE

Euphorbiaceae CRE

ZIG

ACT

■ACT

ACT

'act,';

ACT

ZIG

ACT

$$$

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316

Journal of the Botanical Research Institute of Texas 5(1)

HUEM Color

Diodea grandiflora Mart, ex Benth.

Ingasessilis (Veil.) Mart, ex Benth

Mimosa micropteris (Burk.) Barneby

Aeschynomene sensitiva Sw.

Heliconia laneana Barreiro:

Hyppericum sp.

Aegiphila sellowiana Char r

Aegiphilla s\

Ocotea langsdorffli (Meiss.) Mez

Cuphea melvilla Linc||^)sp

Cuphea calophylla Cham. & Schltdl.

Mascagnia affinis W.R. Anders. & C.C. Dav

Luehea cf. divaricata Mart.

Clidemia hirta (L.) D. Don

Miconiastaminea (Desr.) DC.

Tibouchina sebastianopolitana (Raddi) Cogn.

Guarea macrophylla Lam.

Trichilia pallida Sw.

Trichillia elegans A. Juss,

Eugenia punid folia (Kunth) DC.

Syzygium jambolanum DC.

Pisonia ambigua Heimeri

Ludwigia hookeri

Ludwigia leptocarpa (Nutt.) H. Hara

Ludwigia octovalvis (Jacq.) Raven

Ludwigia uruguayensis (Cambessa) H. Hara

Passiflora foetida L.

Passiflora misera H.B.K.

Passiflora alata Curtis

Picramniasellowii Planch.

Piper umbellatum L.

Angelonia integerrima Spreng.

Polygonum acuminatum H.B.K.

Polygonum punctatum (Meiss.) Small

Triplaris americana L.

Eichornea crassipes (Mart.) Sol ms.

Pontederia cordata Nutt.

Gouania latifolia Reissek

Borreria cf. capitata (Ruiz & Pavon) DC.

Cephalanthus glabratus Schum.

Diodia gymnocephala (DC) Schum.

Diodia sarmentosa Sw.

Phyllanthaceae SUB

Fabaceae (Pap.) SBR

Fabaceae (Mim.)’ TRE

Fabaceae (Mim.) !

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Fabao i| 'dfe':

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12731

and M.O. Batalha. MAPA/SPA, Brasilia 9:85-140.

NEPTUNIA OLERACEA (FABACEAE) NEW TO THE CONTINENTAL

UNITED STATES, WITH NEW AND NOTEWORTHY RECORDS

OF SEVERAL ANGIOSPERMS IN ARKANSAS

Depart men t of Biology

University of Arkansas at Little Rock

Little Rock, Arkansas 72204, U.S.A.

jhpeck@ualr.edu

Brett E. Serviss

Department of Biology

Henderson State University

Arkadelphia, Arkansas 71999-0001, U.S.A.

servisb@hsu.edu

ABSTRACT

INTRODUCTION

Numerous introductions of non-native species and range expansions of both native and non-native species

have occurred in Arkansas over the past 10 years, with our (and others) held work on ruderals and recent

escapes in urban environments leading to the recognition of many plant species being documented as new to

Arkansas or exhibiting significant range expansion (Nesom 2009; Peck 2003; Peck & Serviss 2006; Serviss

2006; Serviss et al. 2006, 2007a, 2007b; Serviss & Peck 2008; Serviss 2009). Many of these species when

initially encountered were present as a single, spontaneous individual or as a relatively small, isolated popu-

lation; however, subsequent to discovery, some have increased in numbers (Serviss 2006). Spontaneous is

used here to mean the autonomous occurrence through sexual or asexual reproduction of a plant species in

a region or flora to which it is not native. It is equivalent to the term “escaped” as defined by Nesom (2000).

Non-native plant species have frequently not been adequately documented and incorporated into North

American floristic studies (Neves et al. 2009). It is therefore highly imperative that continued documentation

and subsequent investigation be focused on the occurrence and biology of non-native species both in the

state’s flora and nationally. Many of these species are initially represented in our flora as waifs from repeat

accidental introductions (this is typical from ballast or rubbish discards), or as local escapes from cultivated

ornamentals. Subsequent to introduction, they demonstrate an ability to persist or establish, with some

eventually becoming invasive. Therefore, a heightened level of awareness, documentation, and attention

must be afforded to non-native species in floras and herbaria, as supported by Pysek et al. (2004). Instead

of being dismissed and ignored when initially encountered, which over time may lead to the probability of

J. Bot. Res. Inst. Texas 5(1): 321 -3

322

Journal of the Botanical Research Institute of Texas 5(1)

control or eradication being minimized or lost, a heightened level of awareness, documentation, and atten-

tion must be afforded to non-native species in our flora.

The occurrence and distribution data of the taxa listed below was determined from the on-line national

flora database, PLANTS Database (USDA, NRCS 2008).

NEW SPECIES RECORD FOR THE CONTINENTAL UNITED STATES

Neptunia oleracea Lour. [syn. N. prostrata (Lam.). Baill.] (Fabaceae). Garden puff or water mimosa (Figs. 1,

2) is an aggressive and invasive aquatic species, presumably native to India and southeastern Asia (possibly

also tropical Africa), where it is used as an edible food plant; however, its present distribution is pantropi-

cal (Windier 1966). It has previously been reported from Puerto Rico, but no previous records exist for its

spontaneous occurrence in the United States (Isely 1998; USDA, NRCS 2008). It is reported here for the first

time in the continental United States, along the flood plain of the Little Maumelle River in Pulaski County,

Arkansas. At this site, it co-occurred with Hydrilla verticillata. Neptunia oleracea is sometimes sold in the

water-garden trade across the southeastern United States in USDA hardiness zones 9-11. It is called water

sensitive plant or garden mimosa in the trade. Once established, it spreads rapidly across the surface of the

water, aided or buoyed by white, spongy aerenchyma tissue underneath trailing stems. Neptunia oleracea is

extremely similar morphologically to N. plena (water dead and awake), but can be distinguished from it and

the two other species of Neptunia that occur in the continental United States by the following key (adapted

from Windier 1966):

KEY TO SPECIES OF NEPTUNIA IN THE CONTINENTAL UNITED STATES

. Plants of wet habitats {N.plena can also si t es) typically growing at edge of water or

directly within; floating and submergent stems c^^l^it^'-^ith white to yellowish-brown aerenchyma

tissue and rooting vigorously at the nodes; rare, currently only known from southeastern Texas [N. plena) or

Arkansas [N. oleracea).

2. Stems typically erect and often highly branched; leaves with a conspicuous gland at the base to slightly

below the lowermost (proximal) pair of pinnae, pinnae (2— )4— 5 per leaf, sometimes more, typically 20 or

more pairs of leaflets per pinna

. Plants completely terrestrial, possibly of moist sites but not aquatic, stems not inflated; widespread in the US.

3. Leaflets 18-30 per pinna; inflorescence bearing 30-60 flowers, all flowers bearing stamens with func-

tional anthers

3. Leaflets (24-)30-50 per pinna; inflorescence with less than 30 flowers, the lower flowers with the stamens

modified into staminodes N. |

SPECIES NEW OR NOTEWORTHY FOR ARKANSAS

Castanea mollissima Blume. (Fagaceae). Chinese chestnut is a large, deciduous tree to 18 m or more tall

that is native to China and Korea. This is the first documentation of this species in Arkansas, and may be the

first record of it outside of cultivation west of the Mississippi River. Castanea mollissima is also known from

several eastern states, ranging from Illinois and Alabama eastward to Massachusetts and Florida. Castanea

mollissima can be confused with Castanea pumila (chinkapin) in Arkansas, but can be distinguished by the

following key:

KEY TO SPECIES OF CASTANEA IN ARKANSAS

323

r, 24 Oct 2010, Serviss 7410 (HE IStm',

Celosia argentea L. [syn. C. cristata L.] (Amaranthaceae). Silver cock’s comb or crested cock’s comb is a

commonly cultivated, ornamental, annual species native to India. It is naturalized in at least 22 eastern

states, including Arkansas; however, this record represents only the third collection of it in the state out-

side of cultivation, as it was previously known only from Baxter and Cross counties (Sarah Nunn, pers.

comm.)— reported as C. cristata, AVFC (2006). Celosia argentea is probably more common in the state’s

flora than collections indicate, as it is frequently cultivated and readily sets seed, often culminating in the

production of numerous additional plants.

Euphorbia esula L. (Euphorbiaceae). Leafy spurge or wolf’s milk is a highly invasive perennial native tp »

Europe. This is the first record of this species from Arkansas, though outside of the southeastern United

States and the southern plains, it is well established (and invasive). Euphorbia esula can easily be confused

with another, introduced species of spurge naturalized in Arkansas, E. cyparissias (cypress spurge). The two

species can be distinguished using the following key:

324

Journal of the Botanical Research Institute of Texas 5(1)

e of M. <

325

published record of M. cordata from Arkansas, the species was actually previously collected in Washington

County along a roadside south of Rogers by Aileen McWilliam on 9 July 1960. The McWilliam specimen was

sent unidentified from UARK to NCU as part of a massive thinning of old and duplicate specimens, without

retention of a duplicate specimen. The specimen at NCU [NCU 212600] was determined as M. cordata by

R. W. Kiger in 1983. The species was not listed for Arkansas by Smith (1988), nor more recently by Kiger

(1997) or the Arkansas Vascular Flora Committee (2006), but was attributed to the state in the PLANTS

Database (USDA, NRCS 2008).

Pistacia chinensis Bunge. (Anacardiaceae). Chinese pistachio is a large, deciduous tree to 18 m tall that

is native to China. This is the first documentation of this species in Arkansas outside of cultivation. It has

been previously documented from Alabama, California, Georgia, Oklahoma (Serviss 2006), and Texas. This

species was planted in many locations in Little Rock over the past 20 years, and a few adventive plants have

been found peripheral to a staging nursery. Additional spontaneous plants should be expected elsewhere in

Arkansas. Vegetatively, P. chinensis somewhat resembles a few other woody species present in the Arkansas

flora, including Ailanthus altissima (tree-of-heaven), Sapindus drummondii (western soapberry), and the three

arborescent species of Rhus (sumac), but can be distinguished from them by the following key:

{ TO PISTACIA CHINENSIS AND SIMILAR WOODY SPECIES IN ARKANSAS

2. Rachis of leaf without wings or wings very narrow and obscure (sometimes present in Sapindus ).

3. Leaflets entire ( Sapindus occasionally with a few toothed leaflets).

4. Foliage strongly (anacardiaceous) aromatic, upper surfaces of leaflets semi-glossy; twigs with a large,

brown or sometimes red, conical-shaped terminal bud; flowers red to reddish-green; drupes red at

maturity, eventually turning dark blue, not translucent_

flowers yellow

. Leaflets prominently toothed.

5. Rachis and lower surfaces of leaflets glabrous or nearly so _

5. Rachis and lower surfaces of leaflets densely velutinous-pubescent_

Rhus typhina Torner (Anacardiaceae). Staghorn sumac is a deciduous shrub or small tree to 10 m tall that

is native to the eastern United States and southern Canada. This is the first documentation of this species

outside of cultivation in Arkansas. Rhus typhina resembles the other two arborescent species of sumac that

occur in Arkansas, R. glabra (smooth sumac) and R. copallina (winged sumac), but can easily be distinguished

using the key above.

River, Little Rock, Two Rivers Park, 21 Jul 2010, Peck 2010283 (HEND).

We would like to very sincerely thank Ann Willyard (Hendrix College), Kristen Benjamin (Henderson State

University), and one anonymous reviewer for their extremely helpful comments and suggestions regarding

this paper. We would also like to thank Brent Baker (Arkansas Natural Heritage Commission) and Sarah

Nunn (University of Arkansas, Fayetteville) for providing species distribution data for Arkansas. Additionally,

we would like to thank Betty Patton for her assistance in acquisition of some voucher specimens.

FIRST RECORD OF IRIS PSEUDACORIS (IRIDACEAE) FROM COLORADO

Michael W. Denslow and Gabrielle L. Katz

William F. Jennings

Department of Geography and Planning

Appalachian State University

Boone, North Carolina 28608, U.S.A.

md68135@email.appstate.edu

ABSTRACT

INTRODUCTION

Iris pseudacorus L. (Iridaceae) is a non-native obligate wetland perennial that is widely established across

the Ul^pj States and Canada (Henderson 2002; USDA, NRCS 2011). It is native to Eurasia and northern

Africa (Sutherland 1990). The earliest records of this plant in North America appear to be from the north-

eastern United States and Canada. Cody (1961) cites a Fernald & Wiegand collection from 1911 as being

the first record from Canada. The plant was listed as a rare escape from gardens in Connecticut by Graves

et al. (1910). This Iris species is widely cultivated and has been in use in North American gardens since as

early as the 18th century (L.H. Bailey Hortorium 1976; Wells & Brown 2000). Iris pseudacorus is listed on

the Noxious Weed list in 5 states, with some states listing it as banned (i.e., importation, sale, and trade are

prohibited) (USDA, NRCS 2011). However, there have been few studies conducted to quantify its threats

(but see Thomas 1980).

NEW RECORDS FOR COLORADO

Iris pseudacorus has not been previously reported for Colorado in floristic treatments covering the state

(Harrington 1964; Weber & Wittmann 2000; Hartman & Nelson 2001; Snow 2009) or Boulder County

specifically (Hogan 1993; Weber 1995). In addition, no Colorado specimens were located in a search of

.tgg&O and KHD or through an online database search of CS and RM (Southwest Environmental Information

Network ''

DISCUSSION

The three specimens cited represent two distinct populations, each with several individuals of this species.

This species is an established part of the Colorado flora. Extensive surveys for this plant have not been

conducted, so its range in Colorado is not clear at this time. However, additional populations have been

observed along Boulder Creek. It is also unclear how long this species has been established in the wild,

since the vouchered populations do not appear to be newly established. This species has been cultivated on

the campus of The University of Colorado, Boulder since at least the 1970s.

J. Bot. Res. Inst. Texas 5(1): 327 -3

NATURALIZED YELLOW COWHORN ORCHID, CYRTOPODIUM FLAVUM

(ORCHIDACEAE), SPREADING IN FLORIDA

R.W. Pemberton 1

H. Liu

Courtesy Curator, Herbarium

Florida Museum of Natural History

Research Associate

Fairchild Tropical Botanic Garden

V SW28th Terrace

Fort Lauderdale, Florida 333 12, VSJt! \

rpemberton5@gmail.com

Department of Earth and Environment

Florida International University

1 1200 5W8 th Street, Miami, Florida 33199, U.S.A.

Center for Tropical Plant Conservation

Fairchild Tropical Botanic Garden

1 1935 Old Cutler Road, Miami, Florida 33 756 ^^

hliu@fiu.edu

The yellow cowhorn orchid, Cyrtopodium Jlavum Link & Otto ex Rchb. is a terrestrial orchid native to the

eastern coast of Brazil (Romero-Gonzalez et al. 2008). It was first found to be naturalized in Florida during

the early 1970s (Hammer 2001). The earliest specimen that we are aware of was collected by George Avery

( Avery 1155 in the herbarium of the Fairchild Tropical Botanic Garden, FTG) on May 12, 1972, from plants

that volunteered on gravel inside a screened porch in Miami-Dade County (Hammer 2001). Hammer (2001)

and McCartney (2010) discuss Avery’s notes on the cultivation of the orchid during the 1950s and the early

nal 1 l on Luer (1972) included the orchid in his book “The native orchids of Florida” as C. andersonii

(Lamb, ex Andrews) R. Br. from South America and said it occurred in the Everglades. The orchid was also

known as C. polyphyllum (Veil.) Pabst. ex F. Barrios, until Romero-Gonzalez et al. (2008) decided that it was

conspecihc with C. Jlavum.

For many years the orchid appeared to be limited to Kendall, a suburb of Miami in the southwestern

urban area of Miami-Dade County. The primary location of the orchid is a pine rockland preserve known

as “Boystown” at 25 39 28N by 80 25 16W, where we estimated there to be 10,000 mature plants (Liu &

Pemberton 2010). The yellow cowhorn orchid also appeared periodically in residential gardens in Kendall.

We studied the pollination of the orchid (Liu & Pemberton 2010) and found that it is being pollinated

by an alien bee, Centris nitida Smith, a bee native to Central and South America that we had found to be

naturalrzed in Florida (Pemberton & Liu 2008a). Only about 1% of the flowers set fruit, but given the large

J. Bot. Res. Inst. Texas 5(1): 331 -3

332

: Dade Chapter oft!

apart of Everglades National!

336

BOOK NOTICE

it. Res. Inst. Texas 5(1): 336. 2011

NATURALIZATION OF THE NUN’S HOOD ORCHID

(. PHAIUS TANKERVILLEAE: ORCHIDACEAE) IN CENTRAL FLORIDA

Donald J. Robinson

18041 Applejack Ct.

Springhill, Florida, 34610, U.S.A.

djrobinson63@yahoo.com

Carmel VanHoek

Volunteer

Week! Wachee State Park

1 027 W. Berry Avenue

Tampa, Florida 33603, USA.

Elizabeth Gandy

Florida Park Service

Florida Dept, of Environmental Protection

Division of Recreation and Parks, District 4 Administration

1843 S. Tamiami Trail, Osprey, Florida 34229, USA.

Robert W. Pemberton*

Courtesy Curator

Flerbarium, Florida Museum of Natural History

2121 SW 28th Terrace

Fort Lauderdale, Florida 333 12, USA.

rpemberton5@gmail.com

Corresponding author

ABSTRACT

Plants of the nun’s hood orchid, Phaius tankervilleae (Banks ex L’Her.) Blume, were found in Weeki Wachee

State Park in Hernando County (near the western coast of Florida north of Tampa) in April, 2010. This or-

chid population is located in a hydric hammock forest on the south side of the Weeki Wachee River. During

the summer of 2010, the area was surveyed by walking zigzig transects, ca. 3 meters apart, flagging plant

clumps as they were encountered. Using this method, we estimate that there are ca. 500 plants at the site. The

number of pseudobulbs in the marked clumps was noted and they generally ranged from 5-19 per clump,

but some had as few as 1-3, and one clump had more than 45 pseudobulbs. Points along the outer edges of

the population were marked with a GPS unit in order to determine its area. This provided an estimate that

the area containing the population of the Phaius is ca 2.3 hectares. The central location of the population is

N 28 31 04.0 W 82 34 32.4. The average elevation of the site is estimated to be 10 m above sea level.

The hydric hammock where the plants were found has the soil type of Anclote Fine Sand and the most

frequent canopy trees in this native forest are Acer rubrum L.,Juniperus virginiana L., Magnolia virginiana L.,

Nyssa sylvatica var. biflora (Walter) Sarg., Persea palustris (Raf.) Sarg., Quercus laurifolia Michx., Sabal palmetto

(Walter) Lodd. ex Schult. & Schult.f., and Taxodium distichum L., Orchids growing in association with the

nun’s hood orchid are the following native species: Habenariajloribunda Lindl., an unidentified Spiranthes

sp., Ponthieva racemosa (Walter) C. Mohr, and Platantheraflava (L.) Lindl.; and the naturalized Oeceoclades

maculata (Lindl.) Lindl.).

Some plants were in flower in April, although freezes in December, January and February, including

-10C, appeared to have damaged most of the plants.

J. Bot. Res. Inst. Texas 5(1): 337 -3

„ Batty, K. Dixon, J. Koch, andK.S

340

Journal of the Botanical Research Institute of Texas 5(1)

BOOK NOTICES I BOOKS RECEIVED

Trees of Florida

Gil Nelson. 2011. The Trees of Florida: A Reference and Field Guide, Second Edition. (ISBN 978-1-

56164-474-2, pbk.; 978-1-56164-475-9, pbk.). Pineapple Press, Inc., PO. Box 3889, Sarasota, Florida

34230, U.S.A. (Orders: www.pineapplepress.com; 800-746-3275). $24.95 (hbk.), 428 pp., 170 line

drawings, 150 color photos, 6" x 9".

Trees of Panama and Costa Rica

Richard Condit, Rolando Perez, and Nefertaris Daguerre. 2011. Trees of Panama and Costa Rica. (ISBN

978-0-691-14707-9, hbk.; 978-0-691-14710-9, pbk.). Princeton University Press, 41 William Street,

Princeton, New Jersey 08540, U.S.A. (Orders: press.princeton.edu/titles/9289.html). $85.00 (hbk.),

$45.00 (pbk.), 552 pp., 438 color illus., 5 line illus., 482 maps, 6" x 9".

Trees in Patagonia

Bernardo Gut. 2008. Trees in Patagonia. (ISBN 978-3-7643-8837-9, hbk.). Birkhauser Verlag AG, PO. Box

133, CH-4010, Basel, SWITZERLAND. (Orders: www.birdhauser.ch; www.springer.com/birkhauser/

biosciences/book/978-3-7643-8837-9). $89.95, 283 pp., 760 illus. with 600 in color by Bernardo Gut,

7" x lOW.

J. Bot. Res. Inst. Texas 5(1): 340. 2011

GOMPHRENA SERRATA (AMARANTHACEAE) NEW

TO THE FLORA OF LOUISIANA

Charles M. Allen, Rhonda Hampton, and Brian Early

Colorado State University

Fort Polk Station 164523rd St.

Fort Polk, Louisiana 71459, U.S.A.

Charles.m.Allen1@us.army.mil

ABSTRACT

Prostrate globe amaranth, ( Gomphrena serrata L.), an annual species in the Amaranthaceae is reported as

new to the Louisiana flora. This species also has the common names of arrasa con todo and perpetua.

roots, decumbent stems, heads narrower than 12 mm in diameter, and bractlets with narrowly cristate keels

on some flowers as described as key characteristics for this species in the Flora of North America treatment

by Clemants in 2003. Its range in the United States includes FL, GA, HI, MD, TX, and VA (Clemants 2003).

Clemants (2003) also reports it from Puerto Rico, Virgin Islands, Mexico, Central America, and South America.

It is not listed for Louisiana by Thomas and Allen (1996) but is mapped for Louisiana in the plants database

(USDA NRCS 2011). The USDA NRCS listing for Louisiana is based on literature (Rollins 1981), but this is

Therefore our collection is apparently the first for Louisiana. A duplicate of our collection was verified by

Steve Clemants of the Brooklyn Botanic Gardens. The plant was probably introduced at the collection site

which is a disturbed area. Associated species include Croton capitatus Michx., Conyza canadensis (L.) Cronq.,

and Hypericum drummondii (Grev. & Hook.) Torr. & A. Gray.

Voucher specimen: LOUISIANA. Vernon Parish: southern end of Range 40, N of LA 463 ca of Pitkin, 31 Jul 2007, Alien et al

ACKNOWLEDGMENTS

REFERENCES

Clemants, S.E. 2003. Gomphrena. In: Flora of North America Committee, eds. Flora of North America north of

Mexico. Vol. 4. Oxford University Press, New York. Pp. 451-454.

Rollins, R.C. 1 981 . Weeds of the Cruciferae (Brassicaceae) in North America. J. Arnold Arbor. 62:51 7-540.

Thomas, R.D. and C.M. Allen. 1996. Atlas of the vascular flora of Louisiana, Vol. 2: Dicotyledons Acanthaceae-

Euphorbiaceae. Louisiana Department ofWildlife and Fisheries, Baton Rouge.

USDA, NRCS. 201-1. The PLANTS database (http://plants.usda.gov/plants). National Plant Data Center, Baton

Rouge, LA 70874-4490,

Journal of the Botanical Research Institute of Texas 5(1)

2011 APPLICATION PROCESS, DELZIE DEMAREE TRAVEL AWARD

Applications for the 2011 Delzie Demaree Travel Award should include a letter from the applicant telling

how symposium attendance will benefit his/her graduate work and a letter of recommendation sent by the

major professor. The Systematics Symposium (www.mobot.org/MBGSystematicsSymposium) dates for 2011

are October 7-10, 2011. The period for receiving applications will end three weeks prior to the date of the

symposium if a sufficient number of applications are in hand at that time. Anyone wishing to apply after

that date should inquire whether applications are still being accepted before applying.

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.

More information: 1-757-221-2799; Email: ddmware@wm.edu

J. Bot. Res. Inst. Texas 5(1): 342. 2011

> TORRESIANA (THELYPTERIDACEAE) NEW TO KENTUCKY

Courtney E. Gorman MattS. Bruton L. Dwayne Estes

344

Journal of the Botanical Research Institute of Texas 5(1)

PLA

HYPOCHAERIS MICROCEPHALA VAR. ALBIFLORA ( HYPOCHAERIS ALBIFLORA:

ASTERACEAE), NEW FOR THE VASCULAR FLORA OF MISSISSIPPI

AND ITS DISTRIBUTION IN NORTH AMERICA

John F. Pruski

Missouri Botanical Garden

$0. Box 299

St. Louis, Missouri 63166-0299, U.S.A.

ABSTRACT

RESUMEN

A few springs ago while botanizing for Plucheas near Bay St. Louis in Gulf Coastal Mississippi, I collected

the South American native Hypochaeris microcephala var. albiflora (Kuntze) Cabrera (Compositae: Cichorieae).

This taxon is typical of Hypochaeris by its plumose pappus bristles and paleate receptacles but is noteworthy

among Cichorieae by having florets with white corollas. Hypochaeris microcephala var. albiflora was not reported

for Mississippi in the statewide treatments by Lowe (1921) and Temple and Pullen (1968). The treatments

by Cronquist (1980) and Bogler (2006) do not list H. microcephala var. albiflora for Mississippi. Neither

the McCook and Kartesz (2010) nor the USDA-NRCS (2010) websites list H. microcephala var. albiflora for

Mississippi, and it is not listed in recent floras in the southern part of the state (e.g., Alford 2001). Thus, it

appears that H. microcephala var. albiflora is a new report for the vascular flora of Mississippi. The purpose

of this note is to voucher H. microcephala var. albiflora in Mississippi, to give a quick overview of this taxon,

and to track the range expansion of this invasive weed in North America. Hypochaeris microcephala (Sch.

Bip.) Cabrera var. microcephala , a different taxon, is a yellow-flowered South American endemic.

Hypochaeris microcephala var. albiflora (sub the orthographic variant “Hypochoeris”) was reported by

Shinners (1966) as new to North America from material collected within five miles of the Sabine River in

Orange County, Texas. Shortly thereafter, Thieret (1969) reported H. microcephala var. albiflora as new to

Louisiana. The range mffT^JImted States of H. microcephala var. albiflora was given by Cronquist (1980) as

Louisiana and Texas. Correll and Johnston (1970) treated H. microcephala var. albiflora as only in Orange

County and as the only member of the genus in Texas, but soon thereafter Tomb (1974) provided an illustra-

tion of H. microcephala var. albiflora, cited it in four Texan counties, and reported an additional two species of

Hypochaeris for Texas. Because the report by Tomb (1974) so quickly followed Correll and Johnston (1970),

this suggests that species of Hypochaeris are overlooked and/or expanding their ranges.

The dot-map of H. microcephala var. albiflora (sub the binomial H. microcephala) in Texas by Turner et

al. (2003) shows it to occur in 13 counties, all in southeastern Texas. The dot-map of H. microcephala var.

albiflora in Louisiana by Gandhi and Thomas (1989) shows it in eight parishes and seven years later Thomas

and Allen (1996) plot it in 13 parishes. Study at the Tulane University herbarium in May 2010 showed that

H. microcephala var. albiflora is known in Louisiana also from East Baton Rouge and Tangipahoa Parishes.

Both Gandhi and Thomas (1989) and Thomas and Allen (1996) report it in St. Tammany Parish, so the report

herein of this taxon in adjacent Hancock County on the Mississippi Gulf Coast is not fully unexpected.

Carter et al. (2009) report a noteworthy eastward leap of H. microcephala var. albiflora into coastal Georgia,

J. Bot. Res. Inst. Texas 5(1): 345 -3

347

Tomb, A.S. 1974J

348

Journal of the Botanical Research Institute of Texas 5(1)

TuRNE|^^^yoLS, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas, vol. 1. Sida Bot. Misc.

Tyrl, R.J., S.C. Barber, P. Buck, WJ. Elisens, J.R. Estes, P. Folley, L.K. Magrath, C.L. Murray, B.A. Smith, C.E.S. Taylor, R.A.

Oklahoma Inc., Noble.

USDA-NRCS. 201 0. The PLANTS Database, (http://plants.usda.gov, accessed 20 October 201 0).

CRATAEGUS CHRYSOCARPA VAR. DODGEI (ROSACEAE)

NEW TO NORTH CAROLINA

fj^^^jPoindexter

LW, Carpenter, Jr. Herbarium

Appalachian State University

The Balsam Mountain Trust

Senior Naturalist/Land Manager

riance@bmtrust.org

Ron W. Lance

poindexterdb@appstate.edu

ABSTRACT

RESUMEN

In North America, Crataegus chrysocarpa Ashe [series Rotundifoliae (Eggleston ex Sargent) Rehder] is a wide-

spread, but primarily boreal taxon found in northern regions of the continent (USDA, NRCS 2011). When

broadly interpreted as the core species of a complex of closely related hawthorns, the composite range

extends from the Pacific Northwest across southern Canada and adjacent states of the U.S. to the Atlantic,

with southern extensions in the Rocky Mountain Region to central Colorado and recently attributed to New

Mexico (Legler 2010), and also in the Appalachians to West Virginia, Virginia, and now North Carolina. In

congruence with this broad distribution pattern, numerous varieties have been attributed to this polymorphic

species and these varieties have in turn been variously interpreted as independent taxa (e.g., Palmer 1937,

1950, 1952; Kruschke 1965; Smith & Phipps 1987; Phipps & O’Kennon 2004).

dodgei Ashe. We have chosen to follow the early taxonomy of Palmer (1937), relegating this taxon to varietal

status inclusive within the C. chrysocarpa species complex. Rationale behind this opinion is based on ob-

servance of unstable morphological characters used to distinguish dodgei from other eastern entities of the

chrysocarpa complex. Specifically, the range of marginal lobing in the leaves of dodgei, which is one of the

most important characters used in the separation of dodgei (Palmer 1950, 1952; Smith & Phipps 1987) var-

ies from obscure to obtuse (typical) to acute (most similar to other chrysocarpa entities). The North Carolina

dodgei plants exhibit distinctly acute leaf lobes, which puts them on the more extreme end of resemblance to

other eastern varieties of chrysocarpa. Considering dodgei as a variety of a species complex that encompasses

a wide range of intergradation is a simplifying approach, though arguments for specific recognition exist,

as well. It should be noted that Palmer changed his recognition of dodgei from a variety to species level in

1950. It is not the purpose or intent of this paper to define or revise any new taxonomic rank for dodgei, but

merely to document its presence in North Carolina.

member of the chrysocarpa- complex known to reach southeastern U.S. states, where it was previously recorded

only as far south as Grayson County, Virginia (Wieboldt et al. 2011). The significance of our collections is

var. dodgei (Ashe) Palmer (syn. Crataegus dodgei Ashe) — -This is the only

J. Bot. Res. Inst. Texas 5(1): 349 -3

(US DA, NRCS). 2011. The PLANTS

North Carolina Herbarium, North Carolina Botanical Garden, Chapel Hill, NC. (http://www.herbarium.unc.

Wieboldt,T.F., G.P. I

LLAVEA CORDIFOLIA (PTERIDACEAE),

NEW FOR TEXAS AND THE UNITED STATES

Robert J. O'Kennon

Botanical Research Institute of Texas

1700 University Dr.

Ft. Worth, Texas 76107-3400, US. A.

George M. Diggs, Jr.

Department of Biology, 900 N. Grand Ave.

Sherman, Texas 75090, USA. and

Botanical Research Institute of T&a$ 1

gdiggs@austmcollege.edu

Llavea is a morphologically distinctive, monotypic genus previously known to occur primarily in the moun-

tains of Mexico from Coahuila, Nuevo Leon, and Tamaulipas south to Chiapas, but also in Guatemala and

Costa Rica (Stolze 1981; Mickel & Beitel 1988; Lellinger 1989; Jaramillo et al. 2000; Mickel & Smith 2004).

It has not previously been reported from Texas (e.g., Correll & Johnston 1970; Hatch et al. 1990; Jones et

al. 1997; Yarborough & Powell 2002; Diggs et al. 2006) or from the United States (Windham 1993 [Flora

North America]; Kartesz 1999; USDA Plants 2011).

The evolutionary relationships of the genus have been unclear — Copeland (1947) considered it to be

derived from Pellaea, Pichi Sermolli (1963) placed it with Cryptogramma and Onychium, while Tryon and

Tryon (1982) suggested there was a distant relationship to Lygodium. Family placement has ranged from

Adiantaceae to Cryptogrammaceae (e.g., Smith et al. 2006), Llaveaceae, and Pteridaceae. Recent molecular

research (e.g., Schuettpelz et al. 2007; Schuettpelz & Pryer 2007, 2008) has shown it to be in the Pteridaceae

in a cryptogammoid clade consisting of three genera including Cryptogramma (6-1 lspecies of North America,

South America, Europe, and Asia) and Coniogramme (an Old World genus). This cryptogammoid clade is

the most basal of the five clades comprising the Pteridaceae, making Llavea only distantly related to other

members of that family. Schuettpelz et al. (2007) noted regarding this clade, “The morphology is highly

variable in this group, and a clear morphological synapomorphy is lacking. Although all three genera display

sterile-fertile leaf dimorphism, this character is widespread throughout the Pteridaceae.” Llavea has long

been cultivated as an ornamental — Hooker (1860) pointed out that it was “one of the most beautiful, and,

in a state of cultivation, rarest of ferns, native of Mexico, with a very peculiar habit. ... No fern-collection

suitable to a warm greenhouse should be without this charming plant.”

A collection from Presidio County, Texas is apparently the first documented occurrence of this species

for both Texas and the States.

. 11.1 km (6.9 mi) from

While the Texas collection of this species was made in 1992, at that time it was thought to be an aberrant

individual of an Osmunda species. The specimen remained in storage until late 2010 when its identity was

realized. In Mexico, it occurs on rocky slopes of moist woods and cliffs (Mickel & Smith 2004). Tryon and

Tryon (1982) noted, “Llavea grows in mesic canyons, or other rocky places in pine and oak woods, or in

tropical forests. Sometimes it occurs on roadsides, on rock walls or in damp soil. It is principally, perhaps

always, a calciophile.” It superficially somewhat resembles Osmunda regalis, royal fern, but is quite different

in appearance from all other Texas species. The current status of the species in Texas is unknown. However,

J. Bot. Res. Inst. Texas 5(1): 351 -3

352

Journal of the Botanical Research Institute of Texas 5(1)

Fig. 1. Lin

12004, fig. 174).

Fig. 2. Scan of Texas vouche

/iillo, I.R., B. F

;v. Biol. Trap

v S.D., J.K. \N

356

Journal of the Botanical Research Institute of Texas 5(1)

MISSOURI BOTANICAL GARDEN PRESS (ST. LOUIS, MISSOURI)

Moss Flora of Central America

Bruce Allen. Contributons erom Ronald A. Pursell and Robert R. Ireland. 2010. Moss Flora of Central

America. Part 3. Anomodontaceae-Symphyodontaceae. (ISBN 978-930723-95-5, hbk.). Monographs

in Systematic Botany from the Missouri Botanical Garden, Volume 117 (ISSN 0161-1542). Missouri Botanical

Garden Press, RO. Box 299, St. Louis, Missouri 63199-0299, U.S.A. (Orders: www.mbgpress.org).

$110.00, 732 pp„ illus., 7" x 10" .

Flora of Costa Rica

B.E. Hammel, M.H. Grayum, C. Herrera y N. Zamora, eds, S^^||royo, Illustrations. 2010. 1

de Costa Rica. Volumen V. Dicotiledoneas (Clusiaceae-Gunneraceae). (ISBN 978-935641-01-8,

hbk.). Monographs in Systematic Botany from the Missouri Botanical Garden, Volume 119 (ISSN 0161-1542).

Missouri Botanical Garden Press, RO. Box 299, St. Louis, Missouri 63199-0299, U.S.A. (Orders: www.

mbgpress.org). $125.00, 970 pp., 412 line drawings, 30 b/w photos, 8 color plates, 7" x 10".

Symposium Proceedings

Bruce E. Ponman and James S. Miller, eds. 2011. Realizing Nature’s Potential: Proceedings of the William

Systematic Botany from the Missouri Botanical Garden, Volume 1 18 (ISSI^^^-1542). Missouri Botanical

Garden Press, RO. Box 299, St. Louis, Missouri 63199-0299, U.S.A. (Orders: www.mbgpress.org).

$35.00, 196 pp., illus., 7" x 10".

n the fall of 2006 ir

J. Bot. Res. Inst. Texas 5(1): 356. 2011

A FLORISTIC INVENTORY OF GRAND TETON NATIONAL PARK,

PINYON PEAK HIGHLANDS, AND VICINITY, WYOMING, U.S.A.

DaveT. (Scott) Kesonie Ronald L. Hartman

PlnelM* • .

Bailey, Colorado 80421, U.S.A |

dave@kesoniebotanical.com

Rocky Mountain Herbarium

Department of Botany, Dept. 3165

University of Wyoming

1000E. University Ave.

Laramie, Wyoming 82071, U.S.A.

ABSTRACT

INTRODUCTION

This contribution discusses a broad-scale inventory of the vascular plants documented on federal lands of

the southern Greater Yellowstone Ecosystem. The areas are: Grand Teton National Park (Park), the John D.

Rockefeller Jr. Memorial Parkway (Parkway), the northern-most portion of Targhee National Forest (Targhee)

in Wyoming, and a remote segment of the Bridger-Teton National Forest (Highlands). This region is known

for its lack of human developments and its wild character. Inventories had been conducted on lands adjacent

to the current one: the Southwest Absarokas (Snow 1989); the Targhee National Forest (Markow 1994); and

the Gros Ventres and Mount Feidy Highlands (Hartman 1996; Fichvar 1979).

This botanical inventory is 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 specimen, the distributions of its

taxa in relatively fine detail (Hartman 1992; Hartman & Nelson 2008; Hartman et al. 2009; Reif et al. 2009).

To this end, over 64 (48 as Master’s degree projects) major floristic inventories have been conducted during

the past 32 years in Arizona, Colorado, Idaho, Kansas, Montana, Nebraska, New Mexico, Oregon, South

Dakota, Utah, Washington, and Wyoming. Over 600,000 new numbered collections have been obtained

by graduate students, staff, and research associates of the RM. These specimens form the core of the Rocky

Mountain Herbarium plant specimen database (700,000+ specimen record, 25,000+ specimen images)

(Hartman et al. 2009).

Study area. — The lands surveyed lie in the Greater Yellowstone Ecosystem of northwest Wyoming,

wholly within Teton County (Figure 1). They belong to the Middle Rocky Mountain Province (Horberg 1938).

J. Bot. Res. Inst. Texas 5(1): 357 - 388. 201 1

Journal of the Botanical Research Institute of Texas 5(1)

Fig. 1 . Boundaries of the federal lands included in the floristic inventory: Grand Teton National Park, John D. Rockefeller Jr. Memorial Parkway, Bridger-

Teton National Forest (Pinyon Peak Highlands), and a portion of the Targhee National Forest, Wyoming. The area is located in extreme northwest

Wyoming, just south of Yellowstone National Park.

The unique geology and associated topography coupled with three converging floristic regions (Rocky

Mountain, Great Basin, and northern Great Plains) yield a highly diverse flora.

The National Park Service administers the bulk of the lands. Grand Teton National Park and the John D.

Rockefeller Jr. Memorial Parkway total 521 mi 2 (134,938 ha). To the east, the contiguous western portion of

the Teton Wilderness, Bridger-Teton National Forest, consists of 194 mi 2 (50,246 ha). Finally, the Wyoming

section of the Targhee National Forest, immediately west of the Parkway, totals 51 mi 2 (13,209 ha). Thus,

the inventory covered 766 mi 2 (490,240 acres; 198,393 ha) of public lands.

Boundaries. — The western boundary is along the Teton Divide, from near Jackson Hole Mountain

Resort on the south to the Parkway (Figure 1). It continues west to Indian Lake on the Targhee, north to

the southern edge of Yellowstone National Park, and east to Gravel Creek, in the Pinyon Peak Highlands.

south, it extends west (four miles north of Jackson) where it completes the loop at the Resort. The area lies

between approximately N44°07' and N43°32' latitude, north and south, respectively; likewise W110°25'

and WlirOl 1 longitude, east and west.

Topography. — The Teton landscape is nothing short of spectacular. The range rises abruptly from the

valley floor (Jackson Hole) with a maximum relief of 6,772 feet (2,064 m). Glaciers and permanent snow-

fields occur on north and east exposures at high elevations. The Grand Teton rises to an elevation of 13,770

feet (4,197 m) with associated peaks extending to the north and south, along the Teton Divide, for a total

distance of 40 miles (64 km).

In contrast, rolling hills and abundant wetlands characterize the Targhee (west of the Parkway). These

wetlands, perpetuated by groundwater discharge and general runoff, are often extensive and complex in

structure. The floating peat mats and ephemeral ponds provided novel habitats.

The Parkway bridges the northern slope of the Teton Range and the Yellowstone Plateau. It contains

extensive wetlands including thermal springs and pools. A rolling topography dissected by the Snake River

contributes to the landscape.

The Pinyon Peak Highlands lie east of the Parkway and the Park. Some shared features are porous

volcanic soils and rolling terrain. The remote region is indeed a “perched highlands” with its namesake

Pinyon Peak (9,705 feet; 2,958 m) (Whitlock 1993; Love et al. 2003). Although mostly montane, a number

of scattered peaks provide subalpine environments. The Highlands drain to the south and then into Jackson

Hole. Much of the Teton Wilderness was burned by “The Yellowstone Fire” of 1988 (Christensen et al. 1989).

Lowest in elevation is Jackson Hole, a flat valley dominated by sagebrush. It is the downthrown block

of the Teton Fault that gave rise to the Tetons (Love et al. 2003). It ranges in elevation from 6,354 to 6,772

feet (1,937-2,064 m). The Snake River bisects Jackson Hole creating braided channels with oxbow ponds.

Many of the lakes were formed by glaciers descending from the Yellowstone plateau or the Tetons near the

end of the Pleistocene (Love et al. 2003).

Climate. — Reliable long-term weather data are from the Moose, Moran, and Snake River weather

stations (Curtis & Grimes 2004). Average daily high and low temperatures for the Moran station, centrally

located and most representative of the area, are 69 and 37, 79 and 42, and 78 and 40° F for June, July and

August, respectively (21 and 3, 26 and 6, 26 and 4° C) (HPRCC 2010). From 1933 through 2002, the highest

recorded temperature was 97° F (36° C) at Moose while the lowest recorded temperature was -63° F (-53°

C) at Moran. The extreme northwest corner of Wyoming is generally the coldest. The high latitude and high

elevation makes it subject to the passage of cold polar air masses (Dirks & Martner 1982). As a consequence,

cold air becomes trapped in the valleys and radiative heat loss from snow-covered surfaces increases winter

cooling (Dirks & Martner 1982). Average summer frost-free periods are short: 14 days at Moran, 29 days at

Moose (Becker et al. 1961).

Most precipitation falls as snow (Knight 1994). Estimated annual snowfall at higher elevations ranges

from 200 to 500 inches (508-1,270 cm) (Fames 1974). In the Tetons, melt water flows from alpine snow-

packs throughout the summer. Summer brings the dry period with average rains totaling a mere 4.12 inches

cm ) for June, July, and August at Moran (period 1911-2010) (HPRCC 2010). A predominant west to f

southwest flow of air delivers both winds and thunderstorms in the summer. This is also true of frontal

winds in winter (Curtis & Grimes 2004), although they may range from north to south.

Geology and geomorphology. — The greater Teton landscape is the result of three regional mountain-

building phases. First, the Sevier Orogeny of the Cretaceous caused widespread compression of the Rocky

Mountain region (Love et al. 2003). Thus, sedimentary strata were pushed eastward to the southern Tetons

as evidenced by the Teton-Gros Ventre uplift. Second, and partially concurrent, was the Laramide Orogeny

that began in late Cretaceous (70 mya). It led to the first significant uplift of the Tetons, again due to com-

pression. Thrust activity along the Buck Mountain, Cache Creek, and Forellen Peak faults elevated the

basement Precambrian. These include the Mount Owen Granite, Webb Canyon Gneiss, and Rendezvous

Metagabbro, some of the oldest rocks in Wyoming with an estimated age of 2.68 billion years (Love et al.

360

Journal of the Botanical Research Institute of Texas 5(1)

sites was 375; GPS coordinates were recorded for each.

2003). Correspondingly, erosion striped away the overlying sedimentary layers of Lower Paleozoic age.

Third, the largest uplift in the Tetons occurred in recent geologic time (7.5 mya). Movement along the Teton

fault, due to extension or pulling, allowed the downthrusting of the Jackson Hole block and an uplift of the

Teton block (5 to 1 ratio of drop to lift) (Love et al. 2003). The high and steep east face of the Tetons thus is

largely a result of the subsidence of Jackson Hole.

The northern portion of the area has been influenced primarily by the Yellowstone Plateau Volcanic

Field. The subterranean plume of superheated magma has led to repeated volcanic activity during the past

2.1 million years (Pierce & Morgan 2009). Furthermore, as the plume is more buoyant, the elevation of the

Plateau increased. Two recent volcanic eruptions have shaped the geology. The first was when hot gases and

ash poured south into the Highlands, the Parkway, and the Targhee. The resulting tuff (Huckleberry Ridge

formation) is widespread across the northern portion of these areas. A later eruption produced the Lewis

Canyon Rhyolite, evident in the Parkway (Love et al. 1992).

361

The Yellowstone Plateau has also influenced the surficial geology by hosting ice masses during the last

two glacial periods (Bull Lake and Pinedale). The Plateau’s high elevation and cold temperatures produced

orographic precipitation that formed large ice sheets. The sheets moved south due to the elevational gradi-

ent. The penultimate event, Bull Lake glaciations, (maximum 155,000 years ago) covered most of Jackson

Hole and areas to the south (Pierce & Good 1992; Love et al. 2007). Smaller glaciers in the canyons of the

Tetons joined this Yellowstone ice mass. It is likely that only the highest elevations in the Tetons and the

Highlands were exposed.

The Pinedale glacial maximum (25,000 years ago) also influenced the surficial geology of Jackson Hole

(Pierce 2003). The process parallels that of the Bull Lake event but the Yellowstone ice sheet only extended

to just south of Jackson Lake. One result was the scouring of the basin now occupied by Jackson Lake

(maximum depth 800 feet, 244 m; currently 446 feet, 136 m, deep to sediment level) (Brewer & Thompson

1991; Love et al. 2007).

Many lakes formed on the valley bottom due to scouring (the largest being Emma, Matilda, Two Ocean,

Leigh, Jenny). Fine scale effects include numerous pothole ponds (Knight 1994). Finally, when the Yellowstone

ice sheet receded, glacial outwash and rivers redistributed large quantities of cobblestone, gravel, and sand.

METHODS

The procedures largely follow practices employed by students and staff at the Rocky Mountain Herbarium

for inventories in the region (Hartman 1992; Hartman & Nelson 2008). The main objective is to collect the

diversity of vascular plants, in a stratified manner, throughout the growing season. We collected in these

four areas during the summers of 2006 (May 27 to Aug 22) and 2007 (June 12 to August 22).

Some specifics on methodology are as follows. When collecting, a route was chosen to sample the

greatest diversity of plant habitats. GPS coordinates were taken every 0.5 mile (0.8 km) or less. Often a

unique habitat (e.g., bog, neoglacial deposit, alpine communities) was preselected as the destination with

frequent collecting in route. About half of the routes were off-trail. The total number of collecting sites was

375 (Figure 2). For rare plants (as designated by the Wyoming Natural Diversity Database, University of

Wyoming) (WYNDD), vouchers were taken only for populations that consisted of 20 or more individuals.

Specimens were processed at the University of Wyoming/National Park Service Research Station (the AMK

Ranch) near Leeks Marina in the Park.

The dried specimens were identified at the RM using appropriate taxonomic literature and then com-

pared with authenticated material. Data for each collection were entered into the RM database (Hartman

et al. 2009). The original set of vouchers is housed at the RM and a representative set of duplicates are at

the Bridger-Teton National Forest Herbarium in Jackson. The RM vouchers have been mounted and imaged

(16.7 megapixels, Canon Mark II). Each high-resolution image is attached to the corresponding entry in the

database. They may be viewed from University of Wyoming Digital Iniative website (UWDI 2010) along

with the 6,360 images (and database) of specimens housed at the Grand Teton National Park Herbarium

(GTNP) (UWDI 2010).

RESULTS AND DISCUSSION

Vegetation Types

Two systems of classification are commonly used to describe plant communities. The first is community

typing” which is based on “existing” vegetation of an area (Cogan et al. 2005). This classification describes

assemblages of plants where the dominant species is often used as part of its name. The second system,

“habitat typing,” describes a community as it would be at “climax.” It is a theoretical approach that incorpo-

rates succession (Anderson 1986). The following discussion will refer primarily to plant communities of the

area - as they exist today - yet some comments will cover successional processes as used with habitat typing.

This discussion facilitates the listing of relevant vegetation types for each taxon in the Annotated Checklist.

Herblands

Alpine Tundra and Talus . — True alpine conditions promote perennial tufted or mat-forming herbs and

ground-hugging shrubs; they are limited to the Park. Topography and aspect dictate the elevation of these

communities but most occur above 9,500 feet (2,896 m) (Spence & Shaw 1981). Unique are dwarf shrub-

lands dominated by Salix arctica, just above treeline (Cogan et al. 2005). Here and elsewhere are mosaics

of meadows, barren areas, patterned ground formed by cryogenic perturbation of soils, rock outcrop, and

talus with significant areas consisting of shear rock.

Common mat-formers on the driest microsites include Phlox multiflora, Minuartia obtusiloba, and Sibbaldia

procumbens. Representative graminoids include Agrostis scabrajuncus parryii, Carex paysonis, and Poa alpina;

also Deschampsia cespitosa on mesic sites. Species often associated with cliffs, rock crevices, and talus include

Saxifraga bronchialis ssp. austromontana, Smelowskia calycina var. americana, Cryptogramma acrostichoides, and

Telesonix heucheriformis.

Subalpine Meadow and Talus . — Meadows adjacent to the subalpine forests vary greatly in size. About half

of this zone is characterized by these meadows. They may be lush with tall forb assemblages dominated by

Symphyotrichum ascendens or Erigeron peregrinus ssp. callianthemus with a minimal cover of grasses.

The herbaceous perennials Castilleja sulphurea, Epilobium anagallidifolium, and Phleum alpinum are com-

mon. Talus slopes and rocky outcrops including cliffs provide habitat for Leucopoa kingii, Erysimum capitatum

var. capitatum, and Penstemon whippleanus. Mesic sites or stream sides are often inhabited by Carex microptera,

Kalmia microphylla, Mertensia ciliata, and Dodecatheon pulchellum ssp. pulchellum.

Sparsely Vegetated Montane or Subalpine Slope . — Hillsides with the Sparsely Vegetated Montane communi-

ties cover extensive areas of the Highlands. They may intergrade with xeric montane grasslands (Cogan et al.

2005) or with limber pine communities ( Pinus flexilis ) that often include Douglas-hr ( Pseudotsuga menziesii).

Rockslides and talus may be placed under the Subalpine Slope type. Frequent associates on open slopes are

Mahonia repens, Lomatium ambiguum, Crepis spp., Phacelia hastata var. hastata, and Penstemon cyaneus.

Montane Meadow. — A xeric montane meadow community occurs in the Pinyon Peak Highlands (Cogan

et al. 2005). Dry conditions are promoted by well-drained soils or southern exposures; grasses are domi-

nant. More common are the moister meadows with abundant forb cover. Elevations range from 6,500 feet

(1,981 m) in the vicinity of Jackson Hole to about 8,800 feet (2,682 m) where they transition to subalpine

meadow. Frequent forbs are Balsamhoriza sagittata, Linum lewisii var. lewisii, Hieracium albiflorum, Achillea

millefolium var. occidentalis, and Taraxacum spp., whereas graminoids include Melica spectabilis, Poa secunda,

Poa pratensis, Carex hoodii, and Bromus carinatus.

Mesic Montane Meadow. — These diverse communities occur at montane elevations but with wet condi-

tions throughout the growing season. Intergradation into stream and seep communities is frequent and

some are influenced by thermal springs in the Parkway. Associations listed for the Park and the Parkway are

Mertensia ciliata, Heracleum maximum-Rudbeckia occidentalis, Geranium viscosissimum, and Ligusticum filici-

num-Delphinium xoccidentale type which intergrade into subalpine meadows (Cogan et al. 2005). Recurring

species include Gentianopsis thermalis, Deschampsia cespitosa, Senecio integerrimus var. exaltatus, Perideridia

gairdneri ssp. borealis, and Calamagrostis canadensis.

Montane Grassland . — They occur in areas on or near Jackson Hole and are frequently adjacent to

sagebrush shrublands. Pastures or old fields also may be of this type. Burned sagebrush communities, not

classified under “burned area,” usually fall into this category. Representatives include Koeleria macrantha,

Poa pratensis, Bromus inermis ssp. inermis, and Antennaria rosea.

Hydrologically-Influenced Vegetation

Aquatic. — The majority of aquatic habitats are ponds, lakes, and fens occurring below the alpine. Unique

aquatic habitats include thermal pools and streams. Free-floating aquatics include Lemna spp., while common

rooted taxa with floating leaves include Nuphar lutea ssp. polysepala, Sagittaria cuneata, and Potamogeton spp.

Largely submerged species, often overlooked, include Isoetes bolanderi and Utricularia macrorhiza.

Scoured Streambed . — Floristically diverse areas heavily influenced by flood runoff are described below

364

Journal of the Botanical Research Institute of Texas 5(1)

(Reed 1952). They typically occur where streambeds have a low gradient, such as in the Highlands, north-

ern portions of the Park, and the Parkway. Deposits of sand, gravel, and cobble are often in ribbon mosaics

and are typically wet or moist. Sedges and rushes are common, as are species adapted to disturbed areas.

Frequently, plants of higher elevations shed propagules that are washed downstream to low elevations, thus

appearing as oddities. Woody species may include Piceapungens and Populus angustifolia; herbs may consist of

Epilobium spp., Platanthera spp., Heterothecavillosav ar. depressajuncus saximontanus. Trifolium spp., Potentilla

arguta, and Crepis tectorum. A local endemic, Stephanomeriafluminea, is limited to these habitats.

Stream and Seep . — Perennial and ephemeral riparian habitats and springs are here included. Intergradation

with mesic montane meadow, subalpine meadow, scoured streambed, aquatic habitats, willow bottomland,

and even montane mixed-conifer forest are possible and may form mosaics. Thus, these communities are

diverse and may support Equisetum arvense, Equisetum laevigatum, and Botrychium multifidum. Flowering

plants include Mertensia ciliata, Lonicera involucrata, Saxifraga odontoloma, Epilobium spp., Senecio triangularis,

and Mentha arvensis, and a few species of slight stature: Stellaria longipes ssp. longipes, Parnassiafimbriata var.

fimbriata, and Spiranthes romanzoffiana.

Wetland. — Next are habitats that support emergent aquatics. Other communities intergrade: aquatics

on the hydric side; montane mesic meadow, stream and seep, and willow bottomland on the drier flanks.

Sedges are important and pure stands of Carex utriculata or Carex vesicaria are common. Likewise, Poa

palustris, Agrostis scabra, Calamagrostis stricta, and Glyceria spp. provide high canopy cover. Willows are

expected, but are a minor component. Herbs include Eleocharis palustris, Carex aureajuncus ensifolius, and

M enyanthes trifoliata as well as Siam suave and Sparganium angustifolium. Ephemeral ponds may become mud

flats and thus classed as “disturbed area.”

Disturbed

Aspen Serai Forest . — Aspen forests are promoted by fire yet not classified under “burn areas” due to their

successional nature (White et al. 1998). Most are serai, thus the “habitat type” would indicate a climax

dominated by conifer (Youngblood 1979). These forest types are rich in herbs, diverse, and physiognomically

complex. Sixteen named associations occur in the Park and the Parkway (Cogan et al. 2005).

Burned Area. — The greater extent of these fires occurred in 1988 when nearly half of Yellowstone National

Park burned (Christensen et al. 1989). Such areas may be covered by lodgepole pine (Kashian et al. 2004)

whereas those previously occupied by shrublands revert to a serai grass community. Exotic species may be

frequent, especially near roads and trails. Common species are Chamerion angustifolium ssp. circumvagum,

Calamagrostis spp., Phlox longifolia ssp. longifolia, and Viola adunca. Resprouting shrubs such as Symphoricarpos

oreophilus var. utahensis and Prunus virginiana var. melanocarpa are typical while such communities may also

have the fire-obligate Ceanothus velutinus.

Roadside-agricultural . — Human-disturbed sites include gravel quarry, parking lot, roadside, homestead,

and heavily-used trail. Both non-native and native weeds are conspicuous and may include Taraxacum laev-

igatum, Carduus nutans, Cirsium arvense, Tragopogon dubius, Capsella bursa-pastoris, Medicago lupulina, Plantago

major, Bromus tectorum, and Potentilla norvegica.

TAXON NEWLY DOCUMENTED IN WYOMING

Surprisingly, only one “state record” was documented, an ornamental established in numerous states in

the country (2010). Achillea ptarmica (Scott 5641 ) is a double-variant cultivar, bred for its showy white ray

florets and sold as an ornamental (Thornton-Wood 2000). It was collected near a historic ranch that has

seen significant activity for many decades (Daugherty et al. 1999). Irrigation ditches appear to be the cor-

ridor for migration.

TAXA NEWLY DOCUMENTED IN THE PARK AND PARKWAY

A comparison was made between Shaw’s (1992) “Annotated Checklist of the Vascular Plants of Grand Teton

365

total of 112 unique taxa new to these federal lands were obtained in our inventory. Of these, 65 were new

to this area based solely on our inventory. An additional 47 unique taxa were also added based on our col-

lections although these had been documented previously based on vouchers in the Grand Teton National

Park Herbarium (UWDI 2010) and the Rocky Mountain Herbarium (Hartman et al. 2009).

SPECIES. OF SPECIAL CONCERN

Species of conservation concern in Wyoming are designated by the Wyoming Natural Diversity Database

(Heidel 2007). Only “Plant Species of Concern,” the category covering the rarest plants at the state level (SI,

SH) are discussed. They represent 21 species documented at 39 sites. The 21 “Species of Concern,” (S2) are

so indicated in the checklist. No taxon listed under the Endangered Species Act was encountered nor are

any known from northwestern Wyoming.

Achnatherum nevadense (Nevada needlegrass) was collected in a 21 year old burn area with lodge-

pole pine in the Pinyon Peak Highlands. All other Wyoming localities are east of the Continental Divide in

Carbon and Fremont counties. Voucher: Hartman 82299.

Agrostis oregonensis (Oregon bentgrass) was collected in the Pinyon Peak Highlands on the margin

of a pond with scattered individuals of Engelmann spruce. The WYNDD list indicates uncertainity (SI?) as

to its status. Voucher: Scott 3299.

Aquilegia formosa (crimson columbine) has a distribution centered in the Pacific Northwest (2010).

Its status in Wyoming was previously unknown as the first specimen, collected in 1951, is thought to be a

hybrid between Aquilegia formosa and A.flavescens (Fertig 2000). Voucher: Scott 961.

Aspidotis densa (pod-fern) is known from cliff habitats (Shaw 1992). It was collected at montane

elevations in the Park. Vouchers: Nelson 68545, 68589.

Astragalus terminalis (railhead milkvetch) is a regional endemic of northwest Wyoming, central

Idaho, and southwest Montana that grows on steep, often eroded, and bare slopes (Shaw 1992). It was col-

lected in the Park and the Highlands, all in Jackson Hole. Vouchers: Hartman 82399; Nelson 68354, 68503.

Botrychium minganense (Mingan moonwort) is known in Wyoming primarily from sites east of the

Continental Divide. It is widespread across the western United States, yet populations probably go unnoticed

due to its cryptic form. It was collected in willow bottomlands in the Park. Voucher: Scott 5690.

Carex echinata ssp. echinata (little prickly sedge) was collected in all juristictions surveyed. It was

obtained in montane wetland and montane mesic meadow. Vouchers: Scott 1868, 2283; Hartman 86597,

86612, 86685.

Carex proposita (Smoky Mountain sedge) has not been documented in Wyoming since 1899 and

thus had been classified as a “historical species” based in the WYNDD list (Fertig 2001; Heidel 2007). The

collection was obtained from a subalpine wet meadow in the Park. Voucher: Scott 1525.

Cicuta bulbifera (bulb-bearing water-hemlock) was collected in the Targhee on the margin of Tillery

Lake. Voucher: Hartman 86575.

Gentianopsis simplex (hiker’s gentian) was previously known in the state only from Yellowstone

(Heidel 2007). The new collections expand its range southward into the Parkway and the Highlands. This

species was found in wetlands at the margins of both a thermal and cold springs and a pond with scattered

individuals of Engelmann spruce. Vouchers: Scott 2456, 3295a.

Huperzia haleakalae (hr clubmoss) was documented in Wyoming only from a 1932 collection by L.

O. Williams. The historical collection and the new record are from the north exposures of the Cathedral

Group in the Park. The habitat was a cool, moist, shady cliff ledge in the subalpine. Voucher: Scott 2666.

Kelloggia galioides (milk kelloggia) is rare in western Wyoming. It was collected on talus in the

montane in the Park. Voucher: Scott 1889.

Luzula glabrata var. hitchcockii (smooth wood-rush) is locally abundant in the Park (Cogan et al.

2005). It is now recognized as Luzula hitchcockii by Swab (2000) as the typical L. glabrata is European in distri-

bution. The collections were from the montane to the alpine. Vouchers: Scott 850, 3218, 5260; Hartman 82255.

366

Journal of the Botanical Research Institute of Texas 5(1)

Minuartia macrantha (House’s Stitchwort) was previously recognized as Minuartia filiorum, now a

synonym of Minuartia macrantha (Hartman & Rabeler 2008). It was collected on high elevation limestone

in the Park. Voucher: Scott 5711a.

Myriophyllum verticillatum (whorled water-milfoil) is an aquatic documented once in the Park.

Previously it was known in Wyoming only from Yellowstone National Park and the Shoshone National Forest

(Heidel 2007). Voucher: Scott 2543.

Porterella carnosula (western porterella) is a showy annual, locally abundant in drying ponds. It was

collected in the Park. Voucher: Scott 5421.

Scheuchzeria palustris (pod-grass) is known from bogs in Yellowstone (Lemly 2007). The population

in Targhee extends its Wyoming range a bit to the south. Voucher: Hartman 86799.

Spirodela polyrhiza (duckmeat) was collected in the Highlands and in northeastern portions of the

Park. Most small populations do not overwinter, but must be established anew or at other sites through

introductions on the feet of waterfowl. Thus they are difficult to track over time (Landolt 2000). Vouchers:

Scott 2380, 3818, 4098, 4412.

Stellaria crispa (crimped stitchwort) is known from mesic habitats of northwest Wyoming. Thus its

occurrence in this Park is limited. Vouchers: Scott 990, 1954; Hartman 86755.

Viola renifolia (kidney leaf white violet) is new to the Park where it was collected in a willow bog.

Voucher: Nelson 68832.

Xerophyllum tenax (western beargrass) is abundant just south of Yellowstone where it was documented

in the Park, the Parkway, and the Targhee. Vouchers: Scott 2977, 3990, 5436.

INVASIVE PLANTS

Invasive plants of exotic origins pose certain threats to ecosystems and human economies. Because these

threats can be insidious and persistent, noxious weeds are carefully monitored by governmental agencies.

In Wyoming, the state legislation has designated 25 plant species as “noxious” (Wyoming Weed and Pest

Council 2008). This study identified nine such taxa; nearly all were collected in human-disturbed areas

(see The Annotated Checklist). Exceptions were three species obtained from areas seemingly free from

such disturbance: Cirsium arvense, Linaria vulgaris, and Sonchus arvensis. The first two were found in the

backcountry at multiple sites.

Teton County has established a list of “Declared Weeds” that compliments the state list of noxious plants

(Wyoming Weed and Pest Council 2007). The three species, collected along roads or developed sites, are

Berteroa incana, Cirsium vulgare, and Verbascum thapsus.

Roughly 7.5 percent (72 species) of the documented taxa are not native to North America (USDA 2010).

This is a relatively low number as levels of 12 to 14 percent have been typical in other studies through the

RM (Hartman et al. 2009).

SUMMARY OF TAXA

A summary of collection results for the study follows. Numbers in parentheses are the subset for the Park

and Parkway.

List by taxonomic category

Families 86 (82)

Genera 347 (338)

Species, 904 (861)

Infraspecies

Hybrids 6 (6)

Unique taxa 962 (909)

List by special category

Exotic taxa

Percent exotic taxa

'i^fi^xious weeds

Species conservation conc^fe*''

State records

Taxa new to the Park and Parkway

this study

this and other studies

72 (69)

7.5 (7.6)

42 (37)';

(47)

tp||f unique taxa b\|^gi§||f plant giftfl):

CONCLUSIONS

s of 2006 and 2007 were devoted to inventorying the vascular plants of Grand Teton

National Park, the J. D. Rockefeller Jr. Memorial Parkway, a portion of the Targhee National Forest, and the

Pinyon Peak Highlands, Bridger-Teton National Forest. Detailed collecting was done at 375 sites (Fig. 2).

A total of 962 unique taxa (904 species, 52 infraspecies) in 347 genera and 86 families were documented

in the study. Also, six hybrid taxa, as documented in the literature, were collected. For the Park and Parkway

proper, the relevant numbers are 909 unique taxa (861 species, 42 infraspecies, and 6 hybrids).

The following botanists obtained a total of 8,002 numbered voucher specimens: David Scott, 5,752

collections, Ronald L. Hartman, 1,351, and B.E. Nelson, 899.

We documented one Wyoming record, the cultivar Achillea ptarmica, one local endemic, Stephanomeria

fluminea described 12 years ago (Gottlieb 1999), and a number of regional endemics. Species of conservation

totaled 42 vouchered at 89 sites. Finally, 112 unique taxa new to the Park and Parkway have been docu-

mented.

Roughly 7.5 percent (72 species) of the flora is not native to North America. Of these exotics, nine

species are considered “Noxious” in Wyoming.

, IfESg ANNOTATED CHECKLIST

The checklist is divided into major vascular plant groups (ferns and fern allies, gymnosperms, and angio-

sperms) each with alphabetical listings by family and species. Nomenclature follows PLANTS database

(USDA 2010). In cases where names differ from those in Dorn’s Vascular Plants of Wyoming (2001), the

latter name is placed in brackets.

Following is a guide to format and abbreviations associated with individual taxa in the checklist.

Collection data are available online (Hartman et al. 2009).

Taxon with Authority Federal entities [G,W,H,T] elevational range in feet; GEOLOGIC AREA [number

of collections] vegetation type.

[Synonym with Authority] based on Dorn (2001). If the binomial for the synonym is the same for the

accepted name, the authority is omitted]

Entity abbreviations:

Geologic a

Grand Teton National Park JAC Jackson Hole

J.D. Rockefeller Jr. Memorial Park Way PPH Pinyon Peak Highlands

Pinyon Highlands, Bridger-Teton TET Teton Mountains

National Forest YEL Yellowstone Plateau (J. D. Rockefeller Jr.

Targhee National Forest Memorial Parkway and Targhee National Forest)

Habitat type:

alt Alpine tundra

aqu Aquatic

asf Aspen serai forest

elf

Montane shrubland

Sagebrush shrubland

Subalpine meadow

Scoured streambed

Subalpine mixed-conifer forest

Journal of the Botanical Research Institute of Texas 5(1)

dis Disturbed area

mmc Montane mixed-conifer forest

sps Sparsely vegetated montane or subalp. slope

Symbols by category preceeding Taxon

▲ “Noxious,” weeds so designation in Wyoming

◄ Taxon new to Park and Parkway, this study and other collections

> Plant Species of [Conservation] Concern (SI, SH) (WYNDD; Heidel 2007)

< Species of [Conservation] Concern (S2) (WYNDD; Heidel 2007)

x Hybrid

FERN ALLIES

Equisetaceae

Equisetum arvense L [G,W,H ,T] 6527-8885', |^^H,TET, YEL

[23] mmm, scs, str,wet,wib.

Equisetum hyemale L. var. affine (Engelm.) A.A. Eat. [G,H,T]

l"j|te4lp o JAC YEL mmc, sps, wet.

Equisetum laevigatum A. Br. [G,W,H] 6551-7200'; JAC, PPH,

YEL [11] sas, scs, str,wet, wib.

Equisetum variegatum Schleich. ex Weber & Mohr var. var-

[Equisetum variegatum]

► Dryopterisfilix-m^^§)ott [G] 6800-7200';TET[1] mjjfr

< Qymnocarpium disjunctum (Rupr.) Ching [G] 6824'; TET

Polystichum ionchitis (L.) Roth [G] 8285-9400'; TET [4] elf,

Woodsia scopulina D.C. Eat. [G] 7200-9200'; TET [3] sps, tas.

Pteridaceae

> Aspidotis densa (Brack.) Lellinger [G] 7200-8000'; TET [2]

Isoetaceae

Isoetes bolanderi Engelm. [G,T] 7340-9530'; TET, YEL [2] aqu.

Lycopodiaceae

> Huperzia haleakalae (Brack.) Holub [G] 95pjjif [1 ] alt, elf.

Lycopodium annotinum L. [G] 7825'^]^^^^' ' *

Cryptogramma acrostichoides R. Br. [G] 7000-1 0800'; TET [1 7]

alt, elf, mmc, mos, sps, tas.

Pellaea brewed D.C. Eat. [G,H] 6938'; JAC, TET [6] alt, elf, sum,

GYMNOSPERMS

Ophioglossaceae

► > Botrychium minganense Viet. [G] 7350'^M wib.

Botrychium multifidu^^^$jme\i) Trevisan [G,W,H,T] 6333-

7539'; JAqjffff EL [10] mmmg|fffil|tr, wet.

Selaginellaceae

Selaginella densa Rydb. [G,H] 6700-1 0800'; JAC, TET [1 4] alt,

FERNS

Aspleniaceae

< Asplenium trichomanes-ramosum L. [G] 9320-9635'; TET

Juniperus communis L. var. depressa Pursh [G,H] 6640-1 0000';

JAC, PPH, TET [16] asf, elf, sps, mmc, tas.

Juniperus scopulorum Sarg. [G,H] 6860-7367'; JAC, TET [4]

Pinaceae

Abies lasiocarpa (Hook.) Nutt. var. lasiocarpa [G,W,H] 6700- 1

9485'; JAC, PPH, TET, YEL [20] bua, mmc, mos, smc.

Picea engelmannii Parry ex Engelm. [G,H] 6920-9800'; JAC,

PPH, TET [16] mmc, scs, smc, sps.

Picea pungens Engelm. [G,H] 6640-6865'; JAC [5] asf, scs,

'icaulis Engelm. [G,H] 6758-9620'; JAC, PPH, TET [7]

3xo n. [G] 9590-10160';

Athyrium filix-femina (L.) Roth [G] 6868'; JAC [1] str.

Cystopteris fragiiis (L.) Bernh. [G,W] 7000-1 0060'; TET, YEL [27]

7s James [G] 6700-7367'; JAC [3] mmc, sps

f[G,H] 6700-9200'; JAC, TET [2of asf, bu;

Kesonie and Hartman, Flora of Grand Teton National Park

373

, jaj faffi mcstr [Descuramia pmnata var. paysonii (Detl.)

Welsh & Reveal]

• Descurainia sophia (L.) Webb ex Prantl [G,H] 6686-6900';

JAC, PPH [3] mmc, mos, sps.

Draba albertina Greene [G,H] 6607-8275'; JAC, PPH,TET,YEL

[13] mm, mmm, mmc

Draba aurea Vahl ex^§g^tri [G] 8770-1 0g£j$ET [4] alt,

► Draba brewer/ Wats. var. cana (Rydb.) Roll. [G] 8500-9740';

imm, sps [ Draba cana Rydb.]

◄ Draba crassa Rydb. [G,H] 8680-10640'; PPH, TET [6] alt,

◄ Draba fladnizensis Wulf. var. pattersonii (O. E. Schulz) Roll.

[G] 9530'; TET [1] sps, tas. [Draba fladnizensis ]

► Draba incerta Pays. [G] 9800'; TET [1] sps, tas.

Draba lonchocarpa Rydb. var. lonchocarpa [G] 8878-11320';

TET [9] alt, elf, sps, tas.

Draba nemorosa L. [G,H] 6479-8320 JaC, pft I-IF/ '{fij'mirt;*

orippa curvipes Greene var

PPH [1] scs. [Rorippa tru

orippa curvisiliqua (Hook.)

eps.) Rollins [H] 7220';

.) Stuckey]

itt. var. curvisiliqua [H]

x Britt, var. orientalis

Rorippa curvisiliqua (Hook.) Besser

Stuckey [H] 7980'; PPH [1]dis.

Rorippa palustris (L.) Bess. var. fernaldiana (Butters & Abbe)

Stuckey [G] 6563'; JAC [1] aqu.

Rorippa palustris (L.) Bess. ssp. hispida (Desv.) Johnsell. [G,T]

6400'; JAC, YEL [2] mmm, wet. [Rorippa palustris var. elon-

gata Stuckey] [Rorippa palustris var. hispida (D^^^bJ

[2] sps.

• Sisymbrium altissimum L. [G] 6932-6400'; JAC [2] sps, scs.

Smelowskia calycina (Steph. ex Willd.) C. A. Mey. var. ameri-

cana (Regel & Herd.) Drury & Roll. [G] 9300-1 If^fET

[8] alt, sps, tas.

• Thlaspiarvense L. [G,H] 6520-7000'; JAC [1 1 ] dis, mm, mmsr^

Draba oligosperma Hook. [G] 9300-1 0000'; TET [7] alt, sps, tas.

► Draba praealta Greene [G] 6597-9320'; JAC, TET [4] elf,

► Draba reptans (Lam.) Fern. [G] 6560-6852'; JAC [2] bua, dis.

Erysimum capitatum (Dougl. ex Hook.) Greene var. capita-

turn [G] 6800-9640'; TET [14] mm, mmc, mos, sum, sps.

[Erysimum asperum (Nu^^var arkansanum (Nutt)

A. Gray]

• Erysimum cheiranthoides L. [G] 6680'; JAC [1] dis, mmm.

Erysimum inconspicuum (Wats.) MacM. [G,H] 6900-6865';

JAC [2] mos, sps.

• Lepidium campestre (L.) W.T. Aiton [G] 6560-7000'; JAC [4]

Lepidium densiflorum Schrad. var. densiflorum [G] 6760'; JAC

[1] dis.

Lepidium densiflorum Schrad. var. macrocarpum Mulligan [G,H]

6743-7000'; JAC [6] asf, dis, mmc.

Lepidium densiflorum Schrad. var .pubecarpum (A. Nels ) Thefl '

[G] 6560';gfgji bua. [Lepidium densiflorum var. pubicar-

pum (A. Nels.) Thell.]

• Lepidium perfoliatum L. [G] 6400-6560'; JAC [2] bua, scs.

Lepidium ramosissimum A. Nels. [H] 7220'; PPH [1] scs.

◄ Lepidium virginicum L. var. pubescens (Greene) Thell. [G]

6400-7220'; JAC, TET [8] dis, mos, sas, scs, sps.

◄ Lesquerella carinata var. carinata Roll. [G,H] 7000-8400'; jyjjjgs

TET [4] mm, sum, sps.

◄ Lesquerella paysonii Roll. [G] 9800'; TET [1] elf, sps, tas.

• Nasturtium officinale W.T. Aiton [G,W] |plg|90 0'| JAC,

YEL [2] str, wet.

Noccaea montana (L.) F.K. Mey. var. montana [G] 8275-9320';

TET [2] elf, mm, tas. [Thlaspi montanum L. var. montanum]

Physaria didymocarpa (Hook.) A. Gray var. didymocarpa [G]

7367'; JAC [1] sps.

Physaria integrifoiia (Roll.) Lichvar var. integrifolia [G] 6900-

Rorippa alpina (Wats.) Rydb. [G] 101 00-1 ^^ET [2] alt,

mmc. [Rorippa cun/ipes Greene var. alpina (Wats.) Stuckey]

Rorippa curvipes Greene var. curvipes [G] 6680-7000'; JAC [2]

Cactaceae

Opuntia fragilis (Nutt.) Haw. var. fragilis [G] 6400-6800'; JAC

[2] sps.

Callitriche hermaphroditica L. [G] 7200'; JAC [1] aqu.

Callitriche heterophylla Pursh var. heterophylla [T] 7340'; YEL

[1] mmc.

Callitriche palustris L. [G,W] 6680-10100'; JAC, PPH, TET [5]

aqu.

Campanulaceae

Campanula rotundifolia L. [G,W,H,T] 6700-1 0480'; JAC, PPH,

TET, YEL [35] bua, mm, mmc, mog, mos, sum, scs, sps, wib.

► Campanula uniflora L. [G] 1 0255f§g$| alt.

^#ff?re//a carnosulc fp||| ) Torr [G] 6860'; JAC [1 ] wet.

Cannabaceae

Humulus lupulus L. var. neomexicanus Nels. & Ckll. [G] 6760';

JAC [1] dis.

► *x Lonicera xbella Zabel [G] 6760'; JAC [1 ] dis.

Lonicera involucrata (Richards.) Banks ex Spreng. var. involu-

crata [G,W,H,] 6573-8375'; JAC, PPH, TET, YEL [38] mmm,

Sambucus racemosa L. var. racemosa [G,W,H] 65f^^m

JA^S^PS^Jifilmm, mmc, tas. [Sambucus racemosa var.

microbotrys (Rydb.) Kearn. & Peeb.]

Sambucus racemosa L. var. melanocarpa (A. Gray) McMinn [H]

7740'; PPH [1] mm, mmc.

Symphoricarpos occidentalis Hook. [G] 7366'; TET [1] mog.

Symphoricarpos oreophilus A. Gray var. utahensis (Rydb.) A.

Nels. [G,W] 6232-8915'; JAC, PFfflteYEL [18] bua, dis,

Caryophyllaceae

Arenaria congesta Nutt. var. congesta [G,W,H] 6560-10795';

JAC, PPH, TET, YEL [34] alt, elf, dis, mm, mmc, sas, sps.

[Eremogone congesta (Nutt.) Ikonnikov var. congesta]

374

Journal of the Botanical Research Institute of Texas 5(1)

Cerostium arvense L. [G,W,H] 6520-7400'; JAC, PPH, TET, YEL

[18] mm,^^^^|nos sas, scs, wib.

Cerastium beeringianum Cham. & Schlecht. var. earlei (Rydb.)

Hulten [G] 1 0000- ,1 #390'; TET [2] tas. [Cerastium beerin-

gianum. var. capillare Fern. & WiegJ

• Cerastium fontanum Baumg. ssp. vulgare (Hartm.) Greuter

& Burdet [G,W,H,T] 6680-7350'; JAC, PPH, TET, YEL [13]

◄ Minuartia austromontana S. J. Wolf & Packer [G] 10198-

10795'; TET [2] alt.

◄ >Minuartia macrantha (Rydb.) House [G] 91 40';'|^^plt.

[Minuartia filiorum (Maguire) McNeil]

Minuartia nuttalin (Pax) Briq. ssp. nuttallii [G] 81 60-9640'; TET

Minuartia obtus'iloba (Rydb.) House [G] 8600-10800'; TET

[16] alt, elf, str, tas.

◄ Minuartia rubella (Wahlenb.) Hiern [G] 8878-1 «p|gif;

[8]alt,clf,^^nc,tas.

Moehringia lateriflora (L.) Fenzl [G,H,T] 6460-7185'; JAC, TET,

YEL [10] rf|fy|igog,5as str, wib

Saginasaginoides (L.) Karst. [G,W,H,] 6780-10640'; JAC, PPH,

TET, YEL [1 8] alt, elf, dis, sum, scs, wet, wib.

Silene acaulis (L.) Jacq. var. subacaulescens (F. N. Wms.) Fern. &

St John [G] 9400-1 1 320'; TET [8] alt, elf, tas.

► Silene drummondii Hook. var. drummondii [G] 6800-7340';

JA^f$t|2] bua,^^SI^

6400-6840'; JAC, YEL [2] wet. [Silene latifolia]

Silene menziesii Hook. [G,H] 6560-7360'; JAC, PPI

dis, scs. [Silene menziesii var. menziesii] [Silenemei

viscosa (Greene) Hitchc. & Maguire]

Silene oregana Wats [G,H] 7200-8450'; PPH, TET [2] n

Silene parryi (Wats.) Hitchc. & Maguire [G] 7840-9

• Spergularia rubra (L.) J. & K. Presl [G,W,H] 6700-7000'; JAC,

YEL [5] dis, rnmnScSl;

Stellaria calycantha $£$£$ Bong. [W,H] 7020-8320'; PPH,

TET, YEL [3] str, wet.

Stellaria longifolia Muhl. ex Willd. [G,W,H] 6680-7580'; JAC,

PPH, TET, YEL [6] mmm, str, wet, wib.

Stellaria longipes Goldie ssp. longipes [G,W,H] 6760-8320'; JAC,

PPpSRyeL I 25 ! asf ' bi|^^n, scs, sps, str, wet, wib

Stellaria obtusa Engel m. [G,H] 6760-8885',

Stellaria umbellata Ti^^ Karel. & Kir. [G,H] 7580-l064Q i ;

JAC, PPH, TET [13] alt, elf, mm, mmm, sum, tas, wet.

Paxistima myrsinites (Pursh) Raf. [G,H] 6597-8953'; JAC, PPH,

TET [1 7] bua,

Ceratophyllaceae

Ceratophyllum demersum L. [T] 6333-6400'; YEL [1] wet.

Chenopodiaceae

Chenopodium atrovirens Rydb. [G] 6760-9060'; JAC, TET [4]

Chenopodium capitatum (L.) Asch. [G] 7366'; TET [1] bua.

Chenopodium foliosum (Moench) Asch. [G,H] 6597-7220'; JAC,

PPH [3] dis, scs, sps. [Chenopodium capitatum (L.) Asch:,.

Hypericum scouleri Hook. ssp. scouleri [G,W,H] 6900-10060';

PF^&YEL [7] mm, str, tas, wet. [Hypericum formosum

var. formosum] [Hypericum formosum var. scouleri (Hook.)

Coult.]

Cornaceae

fjgHMs sericea L. ssp. sericea [G] 657$^|$f; JAC, itif |f||tr

Rhodiola integrifolia (Raf.) ssp. integrifolia [G] 10000-10480';

TET [2] alt, elf, sum. [Sedum integrifolium (Raf.) A. NelsJ

Rhodiola rhodantha (A. Gray) H. Jacobsen [G] 7840-10305';

[3] alt, sum, str, wib. [Sedum rhodanthum A. Gray]

Sedumdebile Wats. [G] 9240- 10480'; TET [5]alt, ,

Sedum lanceolatum Torr. var. lanceolatum [G,H] 6400-1 0350';

JAC, PPH, TET [21] alt, elf, dis, mmc, sas, sum, tas.

Cyperaceae

Carexaquatilis Wahlenb. var. aquatilis [G,W,H,T] 6460-91 80';

‘ J ' ‘*5^6 PPHSfplL [29] str, wet, wib.

Carex athrostachya Olney [G,T] 6400 6900'; JAC, YEL [5]

aqu, mog, str.

Carex aurea Nutt. [G,W,H] 6820-7780'; JAC, PPH, TET, YEL [1 6]

◄ Carex bebbii Olney ex. Fern. [G] 6680'; JAC [1] mmm.

Carex buxbaumii Wahlenb. [W,T] 6460-7160'; YEL [2] mmm,

' ;^St L [4]str,wet.

Carex disperma Dewey [G,H] 6840-7560'; JAC, PPH [4] str,

Carex douglasii Boott [G] 6398'; JAC [1] scs, wib.

Carexduriuscula C. A. Mey. [G,H] 7000-7550'; JAC, PPH [2] sps,

wet. [Carex stenophyl la Wahlenb.]

>Carex echinata J. A. Murray ssp. echinata [G,W,H,T] 6460-

7539'; JAC|_^&L [5] mmm, str.

Carex elynoides Holm [G] 8600-108^^ [7] alt, tas.

Carex engelmannii LJ|||s|y [G] 9400-10795'; TET [4] alt,

tas. [Carex breweri Boott var. paddoensis (Suksd.) Cronq.]

Carex filifolia Nutt. [G] 6560'; JAC [1] bua.

Carex geyeri Boon [G,W,H] 6920-9267'; JAC, PPH, TET, YEL [36]

asf, bua, mm, mmc, mog, mos, sas, tas.

Carex haydeniana Olney [G,H] 6900-10795'; JAC, PPH, TET

[7] alt, mmm, tas.

Carex hoodii Boott [G,H] 6520-8915'; JAC, PPH, TET, YEL [43]

Carex i I lota Bailey [G,H] 8320-9240'^^^^#imm, str.

*<Carexincurviformis Mack. var. danaensis (Stacey) F. J. Herm.

[G] 10600'; TET [1]alt.

Carex interior Bailey [G] 6816-7840'; JAC, TET [2] mmc, str.

► Carex lachenalii Schkuhr. [G] 10100-1 0390'; TET [3] alt, smc,

Carex /oppon/co O.F.Lang [G,W,H] 6900-8065'; JAC, PPH,\||’

[8] mmm, str, wet. [Carex canescens L.]

Carex lasiocarpa Ehrh. [G] 6lJ®ipj|^t.

Kesonie and Hartman, Flora of Grand Teton National Park

375

< Carex leptalea Wa h len b. [G] 6854'; TET[1] mmc, str.

Carexlimosa L. [W,T] 6460-7340'; YEL [5] mmm, mmc, wet.

► < Carex livida (Wahlenb.) Willd. var. radicaulis Paine [W]

6980'; YEL [1] str. [Carex livida]

arexluzulina Olney var. ablata (Bailey) F. J. Herm. [G,W,T]

► Carex macloviana Urv. [G] 8975~T|pf|ET [6] alt, mm,

arex microptera Mack. [G,W,H] 6816-8680'; JAC, PPH,TET,YEL

[1 6] mm, mmm, scs, str, wib. [Carex microptera var. microp-

tera ] [Carex microptera var. limnophila (F. J. Herm.) Dorn]

arex nardina Fries [G,H] 10000-1 0795'; TET [5] alt.

arex nebrascensis Dewey [G,W,T] 6573-7340'; JAC, YEL [6]

[G,H] 7

10-8320'; PPH/TET [3]

Carex nigricans C. A. Mey. [G] 9240-1 0100'; TET [2]^®$:.

Carex norvegica Retz. ssp. stevenii (Holm) E. Murray [G] 7090';

TET [1] str. [Carex norvegica var. stevenii (Holm) Dorn]

◄ Carex pachystachya Cham, ex Steud. [G,H] 7680-11320';

PPH,TET [9] alt, mmm, scs, tas, wet, wib.

Carex paysonis Clokey [G] 9590-1 1 320'; I El [4] alt, tas.

◄ Carex pelhtaM uhl. ex Willd. [G,W,T] 6460-6845'; JAC, YEL

[5] mmm, wet.. "

Carex pelocarpa FJ. Herm. [G] 9760-1 0795'; TET [2] alt. [Carex

nova Bailey var. pelocarpa (F. J. E^^M)orn]

Carex petasata Dewey [G] 6700-7160'; JAC [4] bua, sas.

Carex phaeocephala Piper [G] 6975-1 1 320'; JAC, TET [5] alt,

► Carex platylepis Mack. [G] 681 6-7680'; JAC, TET [2] str.

Carex praeceptorum Mack. [G,H] 7255-10100'; PPH JET [2]

Carex praticola Rydb. [G] 6880'; JAC jjfef m

► > Carex proposita Mack. [G] 9607^|3^»J^

Carex pyrenaica Wahlenb. [G] 1 0060 10640'; I El [3] alt, elf) tas.

Carex raynoldsii Dewey [G,H] 6930-91 30'; JAC, PPH,TET, YEL

[13] asf, mm, mmc, mos, sas.

Carex rossii Boott [G,H] 6573-9615'; JAC, PPH, TET, YEL [18]

► Carex scopulorum Holm var. scopulorum [G] 9240'; TET

[1] sum.

aqu, wet.

◄ Carex spectabilis Dewey [G,H] 8270-10640'; PPH, TET [3]

alt, mm, tas.

Carex stenoptila F. J. Herm. [H] 7580-8400'; PPH [2] mm, mmm.

Carex utriculata Boott [G,W,H,T] 6400-8400'; JAC, PPH, TET,

YEL [25] aqu, mmm, wet.

Carex vallicola Dewey var. vallicola [G,H] 6398-7367'; JAC

[6] bua, sps, str.

TET, YEL [9] mm^ilf^et

◄ Carexviridula Michx. var. viridula [G,W,T] 6900-7340'; JAC,

YEL [3] str, wet.

Cyperus squarrosus L. [G] 6860'; JAC [1] wet.

Eleocharis acicularis (L.) R. & S. [ W,H,T] 6400-8680'; PPH, YEL [5]

mmm, wet. Eleocharis flavescens (Poir.) Urban var. thermal is

(Rydb.) Cronq. [W] 6840'; YEL [1] str, wet.

Eleocharis palustris (L.) R. & S. [G,W,H,T] 6491 -7550'; JAC, PPH,

YEL [11] aqu, mmm, str, wet.

Eleocharis quinquefiora (Hartmann) O. Schwarz [G,W]

6563-6859'; JAC, YEL [2] mmm, str. [Eleocharis pauciflora

(Lightf.) Link]

Eleocharis rostellata (Torr.) Torn [W] '^Sl^-lfetr

◄ Eriophorum angustifolium Honck. ssp. angustifolium [W,T]

6500-7160'; YEL [3] mmm, str, wet.

< Eriophorum gracile Koch var. gracile [T] 6500-7340'; YEL

[2] n

is [T] 6460-6527'; YEL [2] mmm, wet.

< Schoenoplectus americanus (Pers.) Volkart ex Schinz & R.

Keller [W] 6859-6905'; YEL [2] str, wet.

Schoenoplectus tabernaemontani (C.C. Gmel.) Palla [W]6680' ;

';VJ^C[1]wet.

Droseraceae

< Drosera anglica Huds. [W,T] 6460-7340'; YEL [9] mmm,

Elaeagnaceae

Elaeagnus commutata Bernh. ex Rydb. [G] 6560-8500'; JAC,

TET [5] bua, str, sps.

Shepherdia canadensis (L.) Nutt. [G,W,H ,T] 6500-8400'; JAC,

PPH, TEE YEL [41] mm, mmc, sps, str.

Arctostaphylos uva-ursi (L.) Spreng. [G,W,H] 6865-10000';

JAC, PPH, TET, YEL [5] mmc, sps. [Arctostaphylos uva-ursi

var. stipitata (Packer & Denford) Dorn] [Arctostaphylos

^.flilten [G,W] 6852-7670'; JAC, TE“fe^'|3] mmc

[Chimaphila umbellata var. occidentalis (Rydb.) Blake]

Gaultheria humifusa (Grah.) Rydb. [G,W,H,T] 6980-9240'; PPH,

TET, YEL [7] mmm, mmc.

Kalmia microphylla (Hook.) Heller [G,W,H] 6980-1 0305'; TET,

YEL [8] alt, mmm, mmc, str.

Menziesia ferruginea Sm. [G,H] 6852-9267' JAC PFjf||| f7]

mmc, str. [Menziesia ferruginea var. glabella (A. Gray) Peck]

Moneses uniflora (L.) A. Gray [W] fSfiftaskjf} mmc, str.

Orthiliasecunda (L.) House [G,W,T] 6460-1 0450'; JAC, TET, YEL

[10] mmr^^, str.

Phyllodoce empetriformis (Sw.) D. Don [G] 8975-1 Q795 : -JET

[4] alt, sum, sps.

Phyllodoce glanduliflora (Hook.) Cov. [G] 9680-10305'; TET

[4] alt, sps, wet.

Pterospora andromedea Nutt. [G,W] 6868-7344'; JAC, YEL

Pyrola asarifolia Michx. var. asarifolia [G,W,H,T] 6824-7340';

[10] mfn^Jimc, scs, str, wet, wib.

Pyrola chlorantha Sw. [G] 6852-6854'; TET [3] mmi^/ 1 „

Pyrola minor L. [G,T] 6980-7340'; JAC, YEL [2] mm, mmc.

Vaccinium cespitosum Michx. [G,W,T] 7020-7760'; TET, YEL

[3] mmc.

376

Journal of the Botanical Research Institute of Texas 5(1)

l /actinium membranaceum Dougl. ex Torn fG,HI 6575^9189 *

TET [23] alt, mm, mmc, smc, str.

Vaccinium scoparium Leib. ex Cov. [G,W,H,T] 6600-10293';

JAC, PPH,TET,YEL [32] alt, bua, elf, mm, mmc, smc, sps.

Vaccinium uiiginosum L. [G,W,T] 6491-1 0250'; TET, YEL [5] alt,

mmm, mmc, wet. [ Vaccinium occidentale A. Gray]

Euphorbiaceae

Astragalus agrestis Dougl. ex G. Don [G,H] 6470-7740'; JAC,-

P#6$jfe [9] mos, sas, sps, wib.

Astragalus alpinus L. var. alpinus [G,W,H] 6800-9740'; JAC, PPH,

TET, YEL [23] mm, mmm, mmc, scs, sps, str, wib.

Astragalus argophyllus Nutt. var. argophyllus [G] 6640-7367';

JAC [5] mm, sas, sps.

Astragalus australis (L.) Lam. [G] 9300'; TET [1] alt.

► Astragalus convallarius Greene var. convallarius [G] 6560';

, ■ J^Mjlua

Astragalus eucosmus Robins. [G] 6820'; JAC [1] §|p ^ • " '

Astragalus kentrophyta A. Gray var. tegetarius (Wats.) Djijlp,

[G,H] 72^^5'; PPH, TET [14] alt, sum, tas.

► Astragalus laxmannii Jacq. var. robustior (Hook.) Barneby &

S.L. Welsh [G] 6950-7785'; JAC, TET [2] str, tas, wib.

Astragalus miser Dougl. ex. Hook. var. hylophilus (Rydb.)

Barneby [G,H] 6900-8160'; JAC, PPH, TET [9] mmm,

Astragalus miser Doug I. ex. i

6398-73§fp|fe [8] as

Astragalus miser Dougl. c

Barneby [H] 702^%t

◄ Astragalus molybdenus

is Barneby [G,H]

8760- 10795'; TET [3]

. purshii [G,H] 6398-

Astragalus purshii Dougl. ex Hoc

71 60'; JAC [5] bua, sas, sps.

Astragalus tenellus Pursh [G] 6900-7367'; JAC [3] sps.

> Astragalus terminate Wats. [G,H] 6640-71 80'; JAC [3J bua,

Glycyrrhiza lepidota Pursh [G] 6680'; JAC [1] str. [Glycyrrhiza

lepidota var. glutinosa (Nutt.) Wats.]

Hedysarum alpinum L. [G,H] 6885-8985'; JAC, TET [3] asf,

mm, sum. [ Hedysarum alpinum var. americanum Michx.]

Hedysarum boreale Nutt. var. boreale [G] 6900'; JAC [1] sps.

[Hedysarum boreale var. pabulare (A. Nets:) ,'Dprn]

Hedysarum occidentale Greene [G] ?21Q \(M50JET][73sum, v

Lupinus argenteus Pursh ssp. argenteus [G,H] 7360-9780';. JAC,

P#p|[5] mn4|^, mmcj^s.

[G,H] 6720-8975'; JAC, PPH, TET [16] bua, dis, mm, mmc,

[Lupinus lepidus Dougl. dx^M§3/ar utahensis (Wats.)

ipinus caudatus Kellogg ssp. argophyllus (A. Gray) L. Phillips

[G,W] 6795-7155'; JAC, YEL [2] str. [ Lupinus argenteus

Pursh var. argophyllus (A. Gray) Wats.]

pinusdepressus Rydb. [G,H] 8000-9680'; PPH, TET [3] mm,

jfimc, sps, tas. [ Lupinus argenteus Pursh var. depressus

(Rydt^ft^^&f 1

pinus leucophyllus Dougl. ex Lindl. [G,H] 6520-8000'; JAC,

ipinus prunophilus M.E. Jones [G,W,H] 6400-9530'; JAC,

PPH, TET, YEL [14] bua, mm, mmc, sas, sps, tas. [Lupinus

polyphyllus Lindl. var. prunophilus (Jones) L. Phillips]

ipinus sericeus Pursh [G,H] 6400-8500'; JAC, PPH [11] bua,

• Medicago lupulina L. [G,W,H] 6398-7350'; JAC,TEl^lgp5]

• Medicago sativa L. [G] 6400-6560'; JAC [2] bua, mm.

• Melilotus officinalis (L.) Lam. [G] 6400'; JAC [1 ] mm. Oxytropis

borealis DC. var. viscida (Nutt.) S.L. Welsh [G] 9665'; TET [1 ]

Oxytropis campestris (L.) DC. var. cusickii (Greenm.) Barneby

[G] 9300'; TET [1] alt.

Oxytropis deflexa (Pall|i^^^r. foliolosa (Hook.) Barneby [G]

9640-9840'; TET [4] sps, tas.

Oxytropis deflexa (Pal I.) DC. var. sericea T.&G. [G,H] 6700-7360';

• Trifolium hybridum L. [G,W,H,T] 6560-7340'; JAC, PPH, YEL

[25] bua, dis, mm, mmm, mmc, mog, sas, scs, wib.

Trifolium longipes Nutt. ssp. reflexum (A. Nels.) J.M. Gillett

[G,W,H] 6398-7360'; JAC, PPH, T^ta> [18] mm, mmm,

se L. [G,W,H] 6540-7000'; JAC, F

.. [G,W,H,T] 6460-81 00'; JAC, PPH,

Fraseraspeciosa Dougl. exGriseb. [G,H] 6600-9300'; JAC, PPH,

TET [14] asf, bua, mm, mmc, sas, sps.

Gentiana affinis Griseb. [G] 6790'; JAC [1 ] str.

Gentiana calycosa Griseb. [G] 7860-10795'; TET [10] alt, elf,

Gentianella amarella (L.) Borner [G,W,H] 6980-9060'; JAC, PPH,

( tft YEL [1 0] mm, mmrh/mrnc, str, wib.

► > Gentianopsis simplex (A., Gray) litis [W,H] 6905-8360';

PPH, YEL [2] str, wet.

Gentianopsis thermalis (Kuntze.) litis [G,W,H,T] 6333-8360';

Kesonie and Hartman, Flora of Grand Teton National Park

377

JAC, PPH, YEL £18] mrr|^^wet, wib. [Genticmopsis

detonsa (Rottb.) Ma var. elegans (A. Nels.) N. Holmgren]

Swertia perennis L. [G] 71 85';TET HI wib.

6S?i-&0G5:, JAC, YEL [6] aqu, sps, wet. [Juncus JjaiticdS

Willd. var. montanus Engel mj

Geranium richardsonii Hsch. & Trautv. [G,W,H] 6770-8420';:

* [26] mmm ' scs - str ' wib -

Geranium viscosissimum Fisch. & Mey. ex Mey. var. viscosis-

simum [G,W,H] 6555-^320'; JAC, PPH,TET, YEL [35] asf,

Grossulariaceae

Ribes cereum Dougl. var. pedicellare Brewer & Wats. [G,H]

6595-81 30'; JAC, PPH [7]mmc,sps.

Ribes hudsonianum Richards, var. petiolare (Dougl.) Jancz.

[G,H] 6590-7598'; JAC, PPH, TET [4] mmc, str. [Ribes

hudsonianum ]

Ribes inerme Rydb. var. inerme [G,H] 6790-6920'; JAC [3] asf,

Ribes lacustre (Pers.) Poir. [G,H] 6600-961 5'; JAC, PPH, TET, YEL

[16] bua, mmc, mos, str.

Ribes montigenum McClat. [G,H] 6900-9635'; JAC, Wf, TET

[14] alt, mm, mmc, sum, sps.

Ribes viscosissimum Pursh [G,H] 6760-9200'; JAC, PPH, TET,

YEL [22] bua, '

Haloragaceae

Myriophyllum sibiricum Korn. [G,W,H] 6820-7560'; JAC, PPH,

YEL [3] aqu.

► > Myriophyllum verticillatum L. [G] 6790'; JAC [1] aqu.

Hippuridaceae

Hippuris vulgaris L. [G,H,T] 6400-8140'; JAC, PPH, YEL [3] aqu.

Hydrocharitaceae

Elodea canadensis Michx. [T|.^^»|fj,aqu

Juncus confusus Cov. [G,H,T] 6527-8320'; JAC, PPH, TET, YEL

[8] mm, mmm, mmc, mog, wet.

Juncus drummondii E. Mey. [G] 7680-1 0800'; TET [1 0] alt, elf,

Juncus ensifolius Wikstr. [G,W,H,T] 6460-8360'; JAC, PPH 1^

YEL [33] aqu, mrtj|PRic, str, wet, wib. [. Juncus ensifolius

◄ Juncus filiformis L. [G,T] 6900-7340'; JAC, YEL [2] str.

◄ Juncus interior Wieg var. interior [G,H] 6700-8120'; JAC,

YEL [2] bua, dis.

Juncus longistylis Torr. var. longistylis [G,H] 6816-7250'; JAC,

PPH [4] mmc, scs, str.

mertensianus Bong. [G,W,H] HM UMJ) pi\H, TET, '

YEL [1 1 ] ta

Juncus nevadensis Wats. var. nevadensis [G,W] 6960-7020';

JAC, YEL [2] mmc, str, wet.

Juncus nodosus L. [T] 6500'; YEL [1] wet.

Juncus parryi Engelm. [G,H] 7995-11320'; PPH, TET [14] alt,

Juncus re'gelii Buch. [H] 6930'; PPH [1] str.

Juncus saximontanus A. Nelson [G,W,H,T] 6491-7780'; JAC,

Plf|#EL [1 6] aqu, mmm, scs, str, wet. [Juncus ensifo-

Juncus tweedy! Rydb. [H,T] 6460-7539'; PPH, YEL [3] str, wet.

◄ > Luzuia glabrata (Hoppe ex Rostk.) Desv. var. hitchcockii

(Hamet-Ahti) Dorn [G] 7860- 10600'; TET [4] mmc,tas.

► Luzuia multiflora (Ehrh.) Lej. [G,W] 6920-7360'; JAC, YEL

Luzuia parviflora (Ehrh.) Desv. [G,W,H,T] 6960-10390'; JAC,

PF^^YEL [1 7] alt, mmm, mime, sum, str, tas, wib.

[G] 10640-10800'; TET [2] alt, elf.

[Luzuia wahlenbergii Rupr.]

Luzuia spicata (L.) DC. [G] 8000-1 1 320 f ‘TETtf|i^elf smc, str.

Hydrophyllaceae

y [G,\N,H] 6595-8975'; JAC, PPH, TET, YEL [26] dis, nfe’

Nemophila breviflora A. Gray [G,H] 6600-8000'; JAC, PPH, TET

[28] mm, mmc, mos, sas.

Phacelia franklinii (R. Br.) A. Gray [G] 6900-7000'; JAC [1 ] dis.

Phacelia hastata Dougl. ex Lehm. var. hastata [G,W,H]

6700-10480'; JAC, PPH, TET, YEL [22] mmc, sum, sps, tas.

Phacelia heterophylla Pursh ssp. virgata (Greene) Heckard [G,H]

6700-7200'; JAC, PPH [6] bua, mm, mmc, mos.

Phacelia sericea (Graham) A. Gray ssp. sericea [G,H] 6600-

1 0600'; alt, rrtpT^J^; sum, sps.

Iridaceae

Sisyrinchium idahoense Bickn. var. occidentale (Bickn.) D. M.

Hend. [G,W,H] 6700-7250'; JAC, PPH, YEL [6] scs, str,

Juncaceae

Juncus arcticus Willd. ssp. littoralis (Engelm.) Hulten [G,W,H]

Juncaginaceae

Triglochin maritima L. [G,W] 6790-6905'; JAC, YEL [3] aqu, wet.

[Triglochin maritima var p||||utt) A Gray]

► Triglochin palustris L. [W] $05^ YEL [1 ] wet.

Lamiaceae

Agastache urticifolia (Benth.) Kuntze var. urticifolia [G,H]

6750-7840', PPH, Ilf [6] mm, mmc, mos, str.

Dracocephalum parviflorum Nutt. [G] 6560-6932'; JAC [2]

Mentha arvensis L. [G,W,H,T] 6333-7550 , J^T^CU] 5]

mmm, str, wet. [Mentha arvensisvar canadensis (L.) Kuntze]

Prunella vulgaris L.ssp. lanceolata (Barton)' Hulten [G,W]

6563-6905'; JAC, TET, YEL [3] str, wet. [Prunella vulgaris]

Scutellaria galericulata L. [G,W,H,T] 6491-7550'; JAC, PPH, YEL

[8] mmm, str, wet.

► Lemnagibba L. [W] 6905'; YEL [1 ] aqu. Lemnaminuta Kunth

[G,W] 6905-7390'; JAC, YEL [2] aqu.

Lemna trisulca L. [H] 7440-8320'; PPH [3] aqu.

Lemna turionifera Landolt [G,H] 6563-8320'; JAC, PPH [8] aqu.

Kesonie and Hartman, Flora of Grand Teton National Park

379

I] 6750-7000'; JAC, F

Raven [G,W] 6400-6880'; JAC, YEL [3] dis, str.

Orchidaceae

Calypso bulbosa (L.) Oakes var. americana (R. Br.) Luer [G,W,H]

6594-7080'; JAC, TET, YEL [1 0] m me, str. [Calypso bulbosa]

Corallorrhiza maculata (Raf.) Raf. var. occidentalis (Lindl.) Ames

[G,H] 6800-7960'; JAC, PPH,TET [8] bua, mm, mmc.

Corallorrhiza mertensiana Bong. [G] 6800-8285'; JAC, TET [6]

Corallorrhiza striata Lindl. [G] 6760'; TET [1] mmc.

Corallorrhiza trifida Chat. [G] 75 15'; TET [1] mmc.

Corallorrhiza wisteriana Conrad [G] 6594-8285|^^flp'

[3] mmc|;|I§ 5

Goodyera oblongifolia Raf. [G] 6852-7360; JAC, TET [3] mmc.

Listera borealis Morong [G] 6980'; JAC [1 ] wet.

Listera caurina Piper [G,T] 6852-7340'; TET, YEL [2] mmc.

< Listera convallarioides (Sw.) Nutt, ex Elliott [G] 6854-6975';

TET [2] mmc, str.

Listera cordata (I .) R. Br. [G,W] 6854-9140'; JA(|®.'^J

s (Spreng.) Rydb. [G] 6938-7691 '; JAC, TET

[3] mm, mmc, sps.

Platanthera aquiloms Sheviak [W] 7155' YEL [1]

Platahthera dilatata (Pursh) Lindl. ex Beck var. dilatata [G,H,T]

6680-8420'; JAC, PPH,TET, YEL [11] mmm, scs, str.

Platanthera dilatata (Pursh) Lindl. ex Beck var. albiflora (Cham.)

Ledeb. [G,H] 6816-8400'; JAC, PPH,TET[8] mmm, mmc,

Platanthera huronensis (Nutt.) Lindl. [G,H] 6820-8975'; JAC,

‘ str

Platanthera obtusata (Banks ex Pursh) Lindl. [G] 7390'; JAC

[1] mmm.

◄ Platanthera stricta Lindl. [G,W,H] 6816-8680'; JAC, PPH,

TET, YEL [8] mmm, str.

Spiranthes romanzoffiana Cham. [G,W,H,T] 6500-8320'; JAC,

PPH, YEL [10] mm, mmc, str, wet.

obanche uniflora L. [G,H] 671 0-9240'; JAC, TET, YEL [9] bua,

mm, sas, sps. [Orobanche uniflora var. occidentalis (Greene)

Taylor & MacBryde]

•Piantago major L. [G] 6680-6900'; JAC [6] dis, mmm, str,

Plantago tweedy! A. Gray [H] 7250-7390'; PPH [2] scs.

Poaceae

Achnatherum hymenoides (Roem. & Schult.) Barkworth [G,H]

6400-7367'; JAC [6] sas, sps, str.

Achnatherum lettermanii (Vasey) Barkworth [G,H] 6595-9300';

T ^feA,TET[10]mm,mmm,sas,sps,sum.

[G,W,H] 6900-8280'; JAC^^^M|ua, mmc, sas, sps.

• Agropyron desertorum (Fisch. ex Link) Schult. [G] 6400-6820';

JAC [4] dis, sas, sps. [Agropyron cristatum (L.) Gaertn. var.

► • Agropyron fragile (Roth) P. Candargy [G] 6560'; JAC [1 ] bua.

[Agropyron cristatum (L.) Gaertn. var. fragile (Roth) Dor^-;'!

Agrostis exarataJhn. [G,W,H,T] 6460-8360'; JAC, PPH, TET, YEL

[14] mmm, mmc, scs, str, wib.

Agrostis humilis Vasey [G,H] 7860-ljra^gjj^ [10] alt,

J i ^ mmc. sum. smc. [Agrostis thurberiana Hitch^saj

Agrostis idahoensis Nash [W,H] 7120-8680'; PPH, YEL [4]

► < Agrostis mertensiiJhn. [G] 1 0800-1 1 320'; TET [2] alt, tas.

► Agrostis oregonensis Vasey [H] 8360'; PPH [1 ] wet.

Agrostis scabra Willd. [G,W,H,T] 6491 -1 1 320'; JAC, PPH, TET,

YEL [23] alt, mm, mmm, scs, tas, wet.

• Agrostis stolonifera L. [G,W,T] 6680-7340'; JAC, YEL [8]

-s Rydb. [G,W,H]

[G,W,H,T] 6400-8360';

Alopecurus aequalis Sob<

JAC, PPH, YEL [1 1 ] aqu, mmm, wet.

Alopecurus alpinus J. E. Sm. [G,W] 6845-7400'; JAC, YEL

► Alopecurus pratensis L. [G,H] 6790-8320'; JAC, PPH [7]

aqu, dis, mm, mmc

► Arrhenatherum elatius (L.) Beauv. ex J. & K. Presl var. elatius

[G] 6790'; JAC [2] dis.

Iromus arvensis L. [G] 6400-6938'; JAC [3] dis, sas, sps.

mus carinatus H. & A. [G,W,H] 6700-9060'; JAC, PPH, TET,

YEL [38] bua, mm, mmc, sas, scs, sum.

us L. [G,W,H] 6680-8885'; JAC, PPH, TET [21] mm,

• Bromus inermis Leyss. ssp. inermis [G,W,H] 655 1 -7580'; JAC,

PPH [8] bua, X*

[G] 6800-8000'; JAC,TE#^^nc, wet, wib.

Bromus ported (J.M. Coult) Nash [G,H] 6760-7390'; JAC, PPH,

TET [4] mmc, sas, scs, wib. [Bromus anomalus Rupr. ex

Fourn.]

• Bromus racemosus L. [G] 6750-7000'; JAC [1] mmc, mog.

[Bromus commutatus Schrad ]

• Bromus tectorum L. [G,H] 6551-7160'; JAC [13] bua, cfc

Bromus vulgaris (Hook.) Shear [G] IWfr/mil I [2Jbud, str.

Calamagrostis canadensis (Michx.) Beauv. [G,W,H,T] 6460-

1 G250';JAC, PPFI^pJfL [29] alt, aqu, bua, mm, mmm,

Calamagrostis koelerioidesVasey [G] 6852'; TET [1] mmc.

380

Journal of the Botanical Research Institute of Texas 5(1)

► Calamagrostis montanensis Scribn, ex Vasey [G] 7366';

TET [1 ] bua.

9740-1 0350'; TET [6] alt, sps.

Calamagrostis rubescens Buckl. [G,W,H] 6920-8360'; JAC,

PPH,YEL [4] mmc.

Calamagrostis scopulorum Jones [H] 7740'; PPH [1] scs.

6460-7390'; JAC, ;>;> l I, Y: I . [7] aqu, bua, rrirnm, str, wet.

Calamagrostis stricta ssp. inexpansa (A. Gray)

C.W. Greene [G,H] 7120-8885'; PPH, TET [2] mrll;

[Calamagrostis inexpansa A. Gray]

Cinna latifolia (Trev. ex Goepp.) Griseb. [G,W,H] 7230-7740';

PPH,TET, YEL [5] mmc, scs, str.

• Dactylis glomerata L. [G,W,H] 6400-6930'; JAC, PPH, YEL

[7] dis, mf^^

C^^^i^ifomicaMandi. [G] 6880'; JAC [1] . ,

Danthonia intermedia Vasey [G,W,H] 6700-10305'; JAC, PPH,

TE|f|j9] alt, mm, mmc, sas, sum.

Dant

>un [G] 6573-

0'; JAC [t

Deschampsia cespitosa (L.) Beauv. [G,W,H] 6770-1 1 320'; JAC,

PPH, TET, YEL [33] alt, aqu, mmm, sum, tas, wib.

Deschampsia elongata (Hook.) Munro [G,H] 7580-8320'; Pf»E|-

TET[5]dis,mm,«fcscs,wib.

Dichanthelium acuminatum (Sw.) Gould & Clark var. acumina-

tum [W] 6840-6859'; YEL [2] str, wet.

► Elymus alaskanus (Scribn. & Merr.) A. L6ve ssp. latiglumis

(Scribn. & J. G. Snipgiive [G] 9060-1 0795'; TET [4] alt,

mm, sps. [Not recognized by Dorn]

MElymus albicans (Scribn. & Sm.) Love [G] 6900'; JAC [1] sps.

[Elymus albicans var. griffithsn (Scribn. & Sm. ex Piper) Dorn]

Elymus elymoides (Raf.) Swezey ssp. elymoides [G,W,H]

6700-7920'; JAC, PPH, TET, YEL [10] bua, mm, mmc, sas.

Elymus elymoides (Raf.) Swezey ssp. brevifolius (J. G. Sm.) Barkw.

[G,H] 6400-8600'; JAC PPH II 1^7^^*,

Elymus glaucus Buckl. ssp. glaucus [G,W,H,T] 6795-9400'; JAC,

PPH, TET, YEL [24] alt, m str, sum

Elymus lanceolatusi, S<j$§®®Sm.) Gould ssp. lanceolatus [G,H]

6560-7367'; JAC [4] bua, mmc, sas.

Elymus scribneri (Vasey) Jones [G] 10000-11320'; TET [3]

Elymus trachycaulus (Link) Gould ex Shinners ssp. trachycaulus

[G,W,H] 6760-10450'; JAC, PPH,|m|. [40] alt, dis, mm,

Festuca brachyphylla Schult. ex Schult. & Schult. ssp. colora-

densis Frederiksen [G] 8878-1 0640'; TET [7] alt, mmm, tas.

Festuca idahoensis Elmer ssp. idahoensis [G,H] 6560-8400';

JAC, PPH [8] bua, mm, mmc.

◄ Festuca minutiflora Rydb. [G] 1 1 320'; TET [1] alt, tas.

Festuca saximontana Rydb. var. saximontana [G,W,H]

6700-1 0350'; JAC, PPH.’mgj [1 1 ] alt, dis, mm, sas, tas.

jpjfa borealis (Nash) Batch. [G,H,T] 65 1 5-7550'^^^

YEL[4]acE|ftm, wet.

Qlyceria grandis Wats. [G,W] 6680-7390'; JAC, YE£^‘^

Glyceria striata (Lam.) Hitchc. [G,W,H,T] 6860-7780'; JAC, PPH

[1 3] str, wet. [Qlyceria elata (Nash ex Rydb.) Jones]

Hesperostipa comata (Trin. & Rupr.) Barkworth ssp. comata [G]

6900-7 160'; JAC [2] sps.

dia (Scribn. & Tweedy) Barkw. [G] 6400-6900'; JAC [4]

) Beauv. [G,W,H] 6880-8270'; PPH, TET,

L [5] mm, mmm, str, wib.

Hordeum brachyantherum Nevski [G,H] 6770-7780'; JAC, pgjjjJ

[10] mmr^^^tr, wib.

Hordeum jubatum L. [G] 6845'; JAC [1) dis.

Koeleria macrantha (Ledeb.) Schult. [G,W,H] 6550-9300*; JAC,

PPH, TET, YEL [23] asf, bua, mm, mmc, sps.

Leucopoa kingii (Wats.) W. A. Weber [G] 7200-9530'; TET [12]

alt, elf, mmc, sum, sps.

Melica bulbosa Geyer ex Porter & Coult. [G,H] 7300-8130';

JAC, PPI I [6] mm]- mmc' '

Melica spectabilis Scribn. [G,W,H] 6551 -8720'; JAC;^P^||B;.

YEL [41 ] asf, bua dts, , f

► Melica subulata (Gffijflcribn. va^ pammelu (Scribn ) U ;

Hitchc. [G,H] 6795-7960'; JAC, PPH TET [9] |§ft«m

mmc, str. [Melica subulata]

Muhlenbergia andina (Nutt.) Hitchc. [G,W] 6840-6905'; JAC,

YEL [3] str.

Muhlenbergia asperifolia (Nees & Meyen ex Trin.) Parodi [W]

6859'; YEL [1] str.

Muhlenbergia filiformis (Thqrb. -ex' Wats.) Rydb. [G,W,H,T]

6460-8320'; JAC, PPH, TET, YEL [15] dis, mm, mmm, str,

► Muhlenbergia mexicana (L.) Trin. [W] 6840'; YEL [1] str, wet.

Muhlenbergia richardsonis (Trin.) Rydb. [G,W,H,T]

[1] SCS.

Nassella viridula (Trin.) Barkworth [G] 6700-6900'; JAC [1 ] sas.

Phalaris arundinacea L. [G] 6563-6980'; JAC [4] aqu, str.

Phleum alpinum L. [G,H] 7000-1 0305'; JAC, PPH, TET [22] alt,

Phleum pratense L. [G,H] 6597 r 8915', J/^^^|7]

Piptatherum exiguum (Thurb.) Dorn [G,H] 7560-9^^^1

TET [3] mmc, sps.

Poa abbreviata R. Br. ssp. pattersonii (Vasey) A. Love & D. Love

& Kapoor [G] 8975-10795'; TET [4] alt, mm, sps. [Poa

pattersoniiV asey]

Poa alpina L. [G,H] 6770-10390'; JAC, PPH, TET [29] alt, elf,

• Poa annua L. [G] 6680-8420'; JAC, TET [4] mmm, str.

• Poa bulbosa l. [G] 6520-6570'; JAC [1 ] dis.

• Poa compressa L. [G] 6400-6900'; JAC [3] dis, wet.

Poa cusickii Vasey ssp. epilis (Scribn.) W. A. Weber [G,H]

8915-10600'; PPH, TET [7] alt, elf, sum, tas.

Poa cusickii Vasey ssp. pallida Soreng [G] 9060-9800'; TET

[2] sum, tas.

Poa fendleriana (Steud.) Vasey [G,H] 6758-10305'; JAC, &H'

TET [15] alt, mm, mmm, mmc, mos, sas, tas.

Poa leptocoma Jrin. [G,H] 6/9(^r03SO', iAC, Pft f, ff L [7] aqu,

Poa nemoralis L. ssp. interior (Rydb.) W.A. Weber [G,H,T]

^1^15-10040'; JAC, PPH, TET, YEL [12] alt, mmc, scs, sps,

sum, tas, wib. [Poa interior Rydb.]

Kesonie and Hartman, Flora of Grand Teton National Park

Poa palustris L. [G,W,T] 6400-7785'; JAC, PPH, TET, YEL [16]

• Poa pratensis L. [G,W,H,T] 6400-1 0795'; JAC, PPH, TET, YEL

[40] bua, dis, mm, mmm, mmc, mos, sas, scs, sps, sum, wib.

Poa reflexa Vasey & Scribn. ex Vasey [G,W,H] 6980-1 0640'; JAC,

PPH, TET, YEL [23] alt, aqu, mm, mmm, mmc, mos, scs, str.

Poa secunda Presl [G,W,H] 6520-11320'; JAC, PPH, TET, YEL

n ][Poas

[57] a

Pseudoroegneria spicata (Pursh) A. Lo'

6560-9300'; JAC, TET [14] alt, bua

[Elymus spicatus (Pursh) Gould]

x Pseudelymus xsaxicola (Scrib. & J.G. S

Dewey [G] 1 0480'; TET [1 ] alt. [Elym

• Schedonorus phoen/x (Scop.) Holub [t

[Festua

a Schre

Trisetum spicatum (L.) Richt. [G,W,H] 6780-1 1 320'; JAC, PPH,

TET, YEL [37] alt, mm, mmm, mmc, scs, str, tas.

Trisetum wolfii Vasey [H] 7300-8680'; PPH [3] mmm, str.

8680-10640'; PPH, 'TET [5] alt, elf, smc, str. [Deschampsia

atropurpurea (Wahl.) Scheele]

Polemoniaceae

Collomia linearis Nutt. [G,W,H] 6400-9130'; JAC, PPH, TET,

' Xii [43] asf buay^«A^^^»s sas scs, sum.

► Collomia tenella A. Gray [G,H] 7040-8640'; JAC, PPH, TET

Ipomopsis aggregata (Pursh) V, Grant var. aggregata [G,H]

6551-8880'; JAC, PPH, TET [27] bua, mm, mmc, sas, scs,

Lathrocasis tenerrima (A. Gray) L. A. Johnson [G,H] 6700-7440';

JAC, PPH [4] mmc, mos, sas.

P nuttallii [G] 8300-9635'; Tit IS] mm, sps, tas.

Leptosiphon septentrionalis (Mason) J.M. Porter & L.A Johnson

[G,H] 6551-8500'; JAC, §§|l||T [2*6] bua, mm, mmm,

Polemonium occidental Grr^^§p. occidentale [G] 71^|,

TE l»f b -

Polemonium pulcherrimum Hook. ssp. pulcherrimum [G,H]

6700-8400'; JAC, PPH [3] scs, sum.

Polemonium viscosum Nutt. [G] 8878-10600'; TET [8] alt,

sps, tas.

Polygonaceae

Eriogonum brevicaule Nutt. var. laxifolium (T. & G.) Reveal [H]

10040'; TET [1] alt.

Eriogonum flavum Nutt. var. piped (Greene) Jones [H] 8450';

PPH [1] mm.

Eriogonum ovalifolium Nutt, var .purpureum (Nutt.) Durand [G]

9300-1 08|||% [6] alt, tas

Eriogonum umbellatum Torr, vm, umbellatum [G,H] 6806-

Eriogonum mnbellatumlon.vdr. majus Hook. [G,W,H] 6551 -

1 0480'; JAC, PPH, TET, YEL [44] bua, dis, mmc

Oxyria digyna (L.) Hill [G,H] 7210-10800'; TET, YEL [14] alt,

fell tas.

Polygonum amphibium L. var. stipulaceum Coleman [G,W,H,T]

6400-7549'; JAC, PPH, YEL [1 0] aqu, wet.

Polygonum aviculare L. [G] 6680'; JAC [1] mmm.

Polygonum bistortoides Pursh [G,H] 6607-10795; JAC, PPH,

TET [20] alt, elf, n^ttjifrlc sps, tas.

Polygonum douglasii Greene var. douglasii [G,W,H,T] 6400-

891 5'; JAC, PPH, TET, YEL [26] aqu, mm, mmm, mmc, sas,

Polygonum lapathifolium L. [G] 6845'; JAC [1 ] wet.

Polygonum minimum Wats. [G] 7360'; TET [1] sps.

Polygonum polygaloides Wall, ex Meisn ssp. confertiflorum

(Nutt, ex Piper) J.C. Hickman [G,H] 6900-7980'; JAC, PPH

[3] dis, wet. [ Polygonum kelloggii Greene var. confertiflorum

(Nutt -ex Piper) Dorn]

Polygonum polygaloides Wall, ex Meisn ssp. kelloggii (Greene)

J.C. Hickman [G,H] 6700-8680'; JAC, PPH [4] dis, mmm.

[Polygonum kelloggii Greene var. kelloggii ]

Polygonum viviparum L. [G] 71 85-1 0255'; TET [5] alt, elf, scs,

• Rumex acetosella L. [G] 6551-7000'; JAC [7] sas, sps.

► Rumex aquaticus L. var. fenestratus (Gr]|pf|born [W]

68M|i.i!^^

• Rumex crispus L. [G,W,T] 6400-6980'; JAC, YEL [3] aqu,

Linanthus pungens (Torr.) J. M. Porter & L.A. Johnson [G,H]

6560-8320'; JAC [9] mmc, sas, sps.

Microsteris gracilis (Hook.) Greene var. gracilis [G] 6700'; JAC

[1] sas.

[G] 6520 7000'; JAC [6] dis, sas, sps, str.

Phlox hoodii njggs. [G,H] 6597-8878'|jjp[ET [5] ij|

Phlox longifolia Nutt. ssp. longifolia [G,H] 6397-8640'; JAC,

PPH [11] bua, mmc, sas.

Phlox multiflora A. Nels. [G,W,H] 6760-1 0795'; JAC, PPH, TET,

YEL [13] alt, elf, mmm, sas, sps.

Phlox pulvinata (Wherry) Cronq. [G] 8500-9640'; TET [2]

Rumex maritimus L. [G] 6680-6975'; JAC [3] mmm, wet.

[Rumex maritimus var. fueginus (Phil.) Dusen]

Rumex paucifolius Nutt. [G,W,H] 6397-1 01 00'; JAC, PPH, TET,

YEL [26] asf, bua, mm, mmm, sas, smc, wib.

Rumex salicifolius\Ne\r\m. var. mexicanus (Meisn.)

gulivalvis ||fef^t) C L H ’ ) , T ’ * 1 ,

Portulacaceae

Cistanthe umbellata (Torr.) Hershkovitz var. caudicifera (A.

Gray) Kartesz& Gandhi [H] 9590'; PPH [1] sum. [Cistanthe

umbellata ]

Claytonia lanceolata Pall ex Pursh var. lanceolata [G,W,H]

6594-9640'; JAC, PPH, TET, YEL [25] dis, mm, mmm, mr^'. 1 ,!

382

Journal of the Botanical Research Institute of Texas 5(1)

Cloytonia megarhizo (A. Gray) Parry ex Wats. var. megarhiza

[G] 1132CMgiJ]alt.

Lewisia pygmaea (A. Gray) Robins, var. pygmaea [G,H] 6700-

T JAC, PPH, TET, YEL [11] alt, mm, sas, smc, sps, str.

Lewisia triphylla (Wats.) Robins. [G,H] 7000-9530'; PPH, TET

[7] mm, mmc, sps, sum.

Montia chamissoi (Ledeb. ex Spreng.) Greene [G,W] 6850-

7185'; JACjfgglfeL [4] mmm, str, wib.

Potamogetonaceae

Potamogeton alpinus Balbis [G,W,T] 6680-7340'; JAC, JDR,

YEL [3] aqu, mmm.

Potamogeton epihydrus Raf. [T] 6460'; YEL [1] aqu.

Potamogeton foliosus Raf. [G,W] 6820-6900'; JAC, YEL [4]

Potamogeton

L. [T] 7348'; YEL [1] wet.

son/7 (Benn.) Rydb. [G] 6845-6935'; JAC

Stuckenia pectinata (L.) Boerner |

I 6680'; JAC [1] aqu.

Androsace filiformis Retz. [G,W,H] 7159-8320'; JAC, PPH, YEL

[7] mmm, str, wib.

Robbins [G,W,H] 6607-10600'; JAC, PPH, TET, YEL [35]

alt, asf, bua, dis, mm, mmm, mmc, sas, sps.

Dodecatheon conjugens Greene ssp. conjugens [G] 6889-8000';

JAC, ILL [2] sas, str.

Dodecatheon pulchellum (Raf.) Merr. ssp. pulchejlum [G,H]

6790-10305 JAC PPHl§fe|1 1] alt^tj^^^lsps

Primula parryi A. Gray [G] 9607-1 [6] alt, clffe

Aconitum columbianum NifM| columbianum [G,W,H]

OOOO-IOlWi^^^^^^l] mmm, smc, str, wib.

Actaea rubra (Ait.) Willd. [G,H] 6800-7598$. JAC, TET [14]

Anemone multifida Poir. var. multifida [G,H] 6640-7200'; JAC,

TET [6] asf, mmc, wib.

Anemone multifida Poir. var. tetonensis (Porter ex Britton) C.L.

Hitchc. [G,H] 6760-1 0040'; JAC, TET [1 3] alt, sps, sum, tas.

[Anemone tetonensis Porter ex \

Aquilegia coerulea James var. coerulea [G,H] 6800-1 0350'; JAC,

PPH, TET [1 7] elf ggp^ps, str, sum, tas.

Aquilegia flavescens Wats. var. flavescens [G,H] f '60 <3H&p,

JAC, PPH, TET [8] mmc, str, tas.

► > Aquilegia formosa Fisch. ex DC. [G] 6800'; JAC [1 ] str.

Caltha leptosepala DC. ssp. leptosepala [G,H] 8270-10305';

PPH, TET [5] bua, mm, mmc, sum.

Clematis hirsutissima Pursh var. hirsutissima [G,H] 6806-9635';

[1 1 ] bua, mmm, mmc, sum, tas, sps.

Clematis occidentalis (Hornem.) DC. var. grosseserrata (Rydb.)

J. Pringle [G,H] 6800-7440'; JAC, PPH, TET [10] mmc, sir,

◄ Delphinium bicolor Nutt. [G,H] 6594-8310', JAC PPH' Ibf 1

[14] mm, mmc, sas, sps.

x Delphinium xburkei Greene [G] 6880'; JAC [1] mmm.

[Delphinium burkei]

Delphin

[G,H] 7255-9140'; PPH, TET [7]

tz. ex Walp. [G,W,H] 6397-8300';

6930-7740';

P JAC, PPH, TET, YEL [25] asf, dis, mm, mr

x Delphinium xoccidentale (Wats.) Wats. [

PPH, TET [3] mm. [Delphinium occidentale]

Myosurus minimus L. [H] 7780'; PPH [1] mmm.

Ranunculus acriformis A. Gray var. montanensis (Rydb.) Benson

[G] 6720-7W; JAC [5] wib.

◄ Ranunculus adoneus A. Gray [G] 7598-10324'; TET [6]

alt, mmc.

► Ranunculus alismifolius Geyer ex Benth. var. alismifolius [G]

6880'; JAC [1] mmm. [Ranunculus alismifoliusv ar. hartwegii

(Greene) Jeps.]

Ranunculus alismifolius Geyer ex Benth. var. davisii Benson [H]

i [G] 6lMP#^»jAC [2]

[Ranunculus cymoaiaria var. saximontanus hern.j

Ranunculus eschscholtzii Schlecht. var. eschscholtzii [G,H]

7600-10305'; PPH, TET [10] alt, elf, mmc, str, wet.

Ranunculus eschscholtzii Schlecht. var. trisectus (Eastw.) Benson

[G,H] 8915-10640'; PPH, TET [9] alt, rnm,mmc,$um; '

Ranunculus flammula L. var. filiformis (Michx.) Hook. [G,W]

6860-71 YEL [4] aqu s Avunculus flammula

var. reptans (L.) Meyer.]

Ranunculus glaberrimus Hook. var. ellipticus (Greene) Greene

[G,W] 6900-8400'; JAC, TET, YEL [4] mm, mmm.

Ranunculus gmelinii DC. [G,H] 6980-7550'; JAC, TET [2] aqu,

◄ Ranunculus hyperboreus Rottb. [W] 7186'; YEL [1 ] wet.

Ranunculus inamoenus Greene var. inamoenus [G,W,H]

6790-8270'; JAC,$^ffr, YEL [1 0] mrWp^n, str

Ranunculus jovis A. Nels. [G,W] 7320-974g|^|EL f5] <$gj

Ranunculus macounii Britt. [G,W] 6563-690(jaSpEL [2] wet.

Ranunculus orthorhynchus Hook. var. orthorhynchus [W] 6900';

YEL [1] mmc. [Ranunculus orthorhynchus var. platyphyllus

A. Gray]

► Ranunculus pensylvanicus L. f. [G,T] 6400-6880'; JAC, YEL

[2] mmm, wet.

► Ranunculus pygmaeus\Nah\enb.var.pygmaeus[G] 10480';

TEtj$if, elf.

► • Ranunculus repens L. [G] 6850';; JAC [1] mmm.

[1] wet.

Ranunculus trichophyllus Chaix. var. trichophyllus [G,W]

6563-7200'; JAC, YEL [3] str, wet. [Ranunculus aquatilis L.

var. difusus With.]

Ranunculus uncmatus D. Dok^sf^Dan var. uncinatus [G,H]

Benson [G,H,T] 6400-7785'; JAC, PPH, TET, YEL [16] mmm,

Thalictrum fendleri Engelm. ex A. Gray [G,W,H] 6780-8880';

JAC, PPH, TET [14] mmc, str, sum, wib.

Kesonie and Hartman, Flora of Grand Teton National Park

Thalictrum occidentals A. Gray [G,W,H] 681 6-8450'; JAC, PPH,

TET, YEL [9] mm, m me, str.

num Boivin [T] 6460-7340'; YEL [2] mmm, str.

◄ Thalictrum venulosum Trel. [G,W] 7280-7600'; TET, YEL

[2] mm, mmc.

Trollius laxus Salisb. ssp. albiflorus (A. Gray) A. Love & D. Love &

Kapoor [G,W,H,T] 6900-10100'; [14] i^jnm, mmc,

str, smc. [Trollius albiflorus (A. Gray) Rydb.]

a Hook. <

t [G] 9140-10305'; TET

Ceanothus velutinus Dougl. ex Hook. [G,H] 6800-8280'; JAC,

PfPP YEL [15] mmc, mos, sps, str.

Rhamnus alnifolia L'Her. [G] 6680-6854'; JAC, TET [2] mmm,

Rosaceae

[G,H] 6551-8300'; JAC, [18] asf, bua, mm, mmc,

Amelanchier pumila (Torr. & A. Gray) Nutt, ex Roem. [G,H,T]

6479-8080'; JAC, PPH, TET, YEL|®Jim, mmc, sas, sps,

wet. [Amelanchier alnifolia (Nutt.) Nutt, ex Roem. var.

pumila (T. & G.) A. Nels.]

◄ Amelanchier utahensis Koehne [G,H] 6820-8000'; JAC, PPH,

TET [8] mm, mmc, mos, sas, sps, wib.

Argentina anserina (L.) Rydb. [G,W] 6700-6900'; JAC, YEL

[3] wet.

Comarum palustre L. [W,H,T] 6400-755f|PPH, YEL [1 3] m m m,

wet. [Potentilla palustris (L.) Scop.]

Dasiphora friii^jL) Rydb. [G,W,H] 6400-9300'; JAC, PPH,

TET, YEL [24] asf, mm, mmm, sas, sps, str, wib.

Dryas octopetala L. ssp. hookeriana (Juz.) Hulten [G] 10198-

10795'; TET [2] alt.

Fragaria vesca L. [G,H] 6800-8000'; JAC, PPH, TET [17] mm,

Fragaria virginiana Duchesne [G,W,H] 6607-9635'; JAC, PPH,

TET, YEL [36] asf, bua, mm, mmm, mmc, str.

► Geum aleppicum Jacq. [G,W] 6790-6859'; JAC,- YEL [2] str.

Geum macrophyllum Willd. var. perincisum (Rydb.) Raup

[G,W,H,T] 6680-8320'; [31] JAC, PPH, TET, YEL mrflfcp

Geum rossii( R. Br.) Ser. var. turbinatum (Rydb.) C.L. Hitchc. [G]

8980-1 0800'; TET [4] alt, sum, tas.

6560-7200'; JAC, TET [1 5] asf, bua, mr^rip|r, sas.

Ivesia gordonii (Hook.) T. & G. [G,H] 9640-1 004§g1ff [4]

alt, sps.

Petrophytum caespitosum (Nutt.) Rydb. var. caespitosum [G,H]

»i||20-1 0040'; TET [2] alt, elf [Petrophyton caespitosum ]

◄ • Potentilla argentea L. [G] 6551-6880'; JAC [3] dis, sas.

◄ Potentilla arguta Pursh [G,H]6640-10450' ; JAC, PPH, TET,

YEL [22] mm, mmc, sas, scs, str, wet.

Potentilla brevifolia Nutt. exT. & G. [G] 9590'; TET [1] alt.

Potentilla diversifolia Lehm. var. diversify

10795'; JAC, PP^^YEL [25] alt,

r [G,W,H] 6

[5] mrmfa'Ai,

nentilla glandulosa Lindl. ssp. glabrata (Rydb.) D.D. Keck [H]

7140'; PPH [1] mmc. [Potentilla glandulosa var. intermedia

(Rydb.) C.L. Hitchc]

nentilla glandulosa Lindl. ssp. pseudorupestris (Rydb.) Keck

[G,H] 6800-9485', mm [18] |pjnrn smc, sps,

Watson [G,W,H] 6573-8120'; JAC, PPH, TET, YEL [16] dis,

Potentilla gracilis Dougl. ex Hook. var. flabelliformis (Lehm.)

Nutt. exT. & G. [G] 6551-6960'; JAC [3] mmc, sas.

► Potentilla hookeriana Lehm. [G] 9385'; TET [1] alt, tas.

Potentilla norvegica L. [G,W,T] 6333-7366'; JA0|jf jEL

[11]dfs,n^Ss,wet.

9607'; TET [2] alt, ffT>‘ ^

Potentilla pectinisecta Rydb. [H] 6940'; PPH [1] mm. [Potentilla

6597-8720'; JAC, PPH, TET, YEL [27] bua, dis, mm, mmr

Purshia tridentata (Pu^sh) DC. [G,H] 6397-6970'; JAC [1 7] d

6938';.

l|Spndl. ssp

G [1] sps. [Rosa

s [G]

Rosa nutkana Presl var. hispida Fern. [G] 6806fJACt1§]«pps.

Rosa woodsii Lindl. [G,H] 6400-769] '; JACJEI [3] mmc.

Rubus idaeus L. ssp. strigosus (Michx.) Focke [G,H] 6573-8885'; .

JAC, TET [9] mm, mmc, sps, tas.

Rubus parviflorus Nutt. var. parviflorus [G] 6573-7200'; JAC,

TET [8] mos, str.

Sibbaldiaprocumbens L. [G,H] 8680-1 0795'; PPH, TET [1 8] alt,

Sorbus scopulina Greene var. scopulina [G,H] 6800-8000'; JAC,

PPH, TET [8] mm, mmc, mos, sps.

Spiraea betulifolia Pall. var. lucida (Dougl. ex Hook.) (M:

Hitchc. [G,W,H,T] 6597-8885'; JAC, PPH, TET, YEL [13]

Galium boreale L. [G,W,H,T] 6

f JAC, PPH, TET, YEL

Kesonie and Hartman, Flora of Grand Teton National Park

3 (Wat

Holmgren [G] 6400-7000'; JAC [12] sas, sps.

Castilleja pulchello Rydb. [G] 9600-1 0324'; TET [5] alt, sps, tas.

Castilleja rhexiifolia Rydb. [G,H]8400-1 0305'; PPH,TET [3] alt,

str, sum. [Castilleja rhexifolia Rydb.]

Castilleja sulphurea Rydb. [G,H]7340-1 0600'; TET [11] thfk)

Collin

I. [G,W,H] 6397-8t

3 (L.) K

lea [G] 6

[1] s;

A • Unaria vulgaris Mill. [G,T,Y] 6460-7366'; JAC, TET, YEL [6]

'; JAC, PPH,TET,YEL

H,TET,YEL[14]

imulus guttatus DC. [G,W,H] 6.

[16] rf|l|tr,wet,wib.

'muluslew hirsh [G,W,H] 71 20-10640'; F

Mimulusmoschotjs Dougl. ex Lindl. var. moschatus [G,W,H,T]

6400-8360'; JAC, PPH, YEL [15] mmm, mmc, str, wet.

Mimulus tilingii Regel var. tilingii [H,T] 6333-8320'; PPH, YEL

[6] mmm, scs, str, wet.

◄ Mimulus washingtonensis Gandog. [G] 7000-8953';TET[3]

elf, sps. [Mimulus patulus Penn.]

Orthocarpus luteus Nutt. [G,W,H] 6810-7580'; JAC, PPH, TET,

YEL [6] diS/flFT mmm, wet, wib.

[G,W,H] 6

TET [1]alt.

Retz. [G,H] 6854-10305'; JAC,. PPH, :

- ssp. purpurea (Parry) G.D. Carr [G]

TET [23] mm, mmm,;

Ped leu laris parryi A. Gray

8980';TEfH||!fi

Ped icu laris racemosa DougL ex ssp alba P«

[G,W,H,T] 6460-1 0450'; JAC, PPH, TET, YEL [19] bua, i

(Rydb.) Cronq. [G,H] 7210-10600'; 1^^ [11] alt, mm,

nstemon cyananthus Hook. [G] 7691-9300'; TET [4] elf,

nstemon cyaneus Penn. [G,H] 7200-8400'; JAC, PPH, TET [8]

nstemon deustus Dougl. ex Lindl. var. deustus [G] 6806-9200';

JAC, TET [4] mos, sps, tas.

nstemon montanus Greene var. montanus [G,H] 8878-

10000'; PPH, TET [6] alt, sps, tas.

nstemon procerus Dougl. ex Grah. var. procerus [G,H]

6400-8270', JAC, PPH, TET fl 1 ] asf, Mmm, sas, scs.

nstemon radicosus A. Nels. [G,H] 6560-7367'; JAC [5] bua,

wKeij $ps.

nstemon subglaber Rydb. [G] 6400-6938'; JAC [3] sps

nstemon whippleanus A. Gray [G,H] 6770-1 0350'; JAC, PPH,

TET [10] mmm, mmc, scs, sme, sps, sum, tas.

Scrophularia lanceolata Pursh EG]

• Verbascum thapsus L. [G,W] 6790-6840'; JAC, YEL [2] dis.

Veronica americana Schwein. ex Benth. [G,W,H,T] 6460-

1 01 00'; JAC, PPH, TET, YEL [20] dis, mmm, sme, str, wib.

Veronica anagaiiis-aquatica L. [G] 6680'; JAC [1] str.

◄ • Veronica biloba L. [G,H] 6640-8720'; JAC, PPH [5] dis, sps.

◄ • Veronica officinalis L. [G] 6760'; JAC [1 ] dis.

Veronica peregrina L. ssp. xalapensis (H.B.K.) Pennell [G]

6700-6900'; JAC [2] dis, sas.

Veronica scutellata L. [T] 6400-6527'; YEL [2] wet.

Veronica serpyllifolia L. ssp. humifusa (Dicks.) Syme [G,H]

6700-8680'; JAC, PPH, TET, YEL [11] mmm, mmc, sas,

Veronica wormskjoldii R. & S. [G,W,H] 7580-1 0640'; PPH, TET,

YEL [17] alt, elf, mmm, str, wet.

Sparganiaceae

Sparganium angustifolium Michx. [G,W,H,T] 6500-8065'; JAC,

PPH, YEL [8] aqu, str, wet.

• Sparganium emersum Rehm. [G,T] 6400-6980'; JAC, YEL

[4] aqu.

Typhaceae

Typha latifolia L. [W] 681 O'; YEL [1 ] wet.

Urticaceae

Urticadioica L. ssp. gracilis (/|gp |eland. [G,H] 6520-7350';

JAC, TET [10] mmc, mos, str. [Urtica dioica var. procera

(Muhl. ex Willd.) Wedd.]

Valeriana acutiloba Rydb. var. pubicarpa (Rydb.) (MspLfG]

93j$jJ&rUclf»sps.

Valeriana dioica L. var. sylvatica Wats, [H]'7560' ; PPH [1]

Valeriana edulis Nutt. exT. & G. var. edulis [G,H] 6720-9300';

JAC, PPH [11] mmm, mm, sps, wib.

Valeriana occidentalis Heller [G,W,H] 6520-9140'; JAC, PPH,

TET, YEL [41] asf, bua, mm, mmm, mmc, mos, sas.

Verbena bracteata Cav. ex Lag. & Rodr. [G] 6700-6900'; JAC

[4] dis, sas, wet.

Viola adunca Sm. [G,W,H] 6640-81 00'; JAC, pj|@fr, YEL [21 ]

Viola macloskeyi Lloyd ssp. pallens (Banks ex Ging.) M.S. Baker

[G,H,T] 6460-8975'; JAC, PPH, TET, YEL [10] mm, mmm,

Viola nephrophylla Greene [G,H] 7560-8400'; PPH, TET [2]

Viola orbiculata Geyer ex Holz. [G,H] 6760-8040'; PPH, TET

[2] mm, str.

Viola palustris L. [G,H] 7360-9140'; JAC, PPH, TET [7] mmm,

Viola praemorsa Dougl. ex Lindl. ssp. linguifolia (Nutt.) M.S.

Baker &J.C. Clausen ex M. Peck [G,W,H] 6720-8975'; JAC,

PPH, TET, YEL [20] bua, dis, mm, mmm, mmc, mos, sas,

'and the flora of the Rocky

Journal of the Botanical Research Institute of Texas 5(1)

S Department of Agriculture) 2010, Natural Resources Conservation Services;: tJatabatey

National Plant Center. Baton Rouge, LA. URL: http://plants.usda.gov (15 May 2008).

of Wyoming Digital Initiative). 2010. Grand Teton National Herbarium Database, http://www.rmh.

uwyo.edu/digitalherbaria/about.php

White, C A, C£Ou^^|nd CE Kay. 1998. Aspen, elk, and fire in the Rocky Mountain National Parks of North

America. Wildlife Soc. Bull. 26:449-462.

Whitlock, C. 1993. Postglacial vegetation and climate of Grand Teton and southern Yellowstone National Parks.

Ecol. Monogr. 63:1 73-1 98.

Wyoming Weed and Pest Council. 2007. Teton County declared weed list. Unpubl.

Wyoming Weed and Pest C 'js'n.*2008 Designated noxious weed list. URL: http://www.wyoweed.org/statelist.

html (14 Nov 2008).

Youngblood, A.P. 1 979. Aspen community type classification for the Bridger-Teton National Forest. Master's thesis.

Utah State Univ., Logan.

DONALD E. STONE

10 December 1930-4 March 2011

Jonathan Gilej|

Vice President for Development

Organization for Tropical Studies

Box 90630, Duke University

Durham, North Carolina 27708-0630, U.S.A.

jgiles@duke.edu.

Donald Eugene Stone died from a short bout with cancer on Friday, March 4,2011 in Durham, North Carolina,

Don was born on December 10, 1930 and grew up in Eureka, California. He took his undergraduate work

at Humboldt State College and the University of California at Berkeley, graduating in 1952. He remained at

Berkeley and was awarded his Ph.D. in Botany in 1957. Subsequently, he taught as a Lecturer and Assistant

Professor at Tulane University for six years. In 1963, he joined the Botany Department at Duke University

and taught at Duke for the remainder of his career. In 1969-70, Don took a one-year leave of absence to

serve as the Associate Program Director in Systematic Biology at NSF where he was in charge of all plant

research proposals. In 1976, while teaching as full professor at Duke, he began shepherding the Organization

for Tropical Studies (OTS) as its Executive Director. During his tenure with OTS, Don expanded the con-

sortium from twenty to more than 55 universities, museums, and research institutions. He enhanced the

Organization’s field-based graduate courses and created an on-the-ground training program for policy-makers.

Most importantly, he strengthened OTS’s three biological held stations in Costa Rica, La Selva, Las Cruces,

and Palo Verde, as major research centers, and, in particular, established the La Selva station as one of the

most important sites in the world for research in tropical biology. In the early 1980s, under Don’s guidance,

OTS took a leadership role, along with the John D. and Catherine T. MacArthur Foundation, the World

Wildlife Fund, and The Nature Conservancy, in establishing a protected, 47,000-hectare, forested corridor

from the Braulio Carrillo National Park, located in the central highlands of Costa Rica, to La Selva, more than

/.^Ifhiles away in the Caribbean lowlands. As result of these efforts, in 1985 OTS was the first organization

to be awarded the John and Alice Tyler Prize for Environmental Achievement. When Don retired from OTS

in 1996, he served as the chair of the Botany Department at Duke for three years. In 2000, he joined the

OTS Board of Visitors, which he formed in 1992, and from 2003-2005 he served in a volunteer capacity as

OTS’s Interim Executive Director during an 18-month search for the current CEO. In 1988 Don received

J. Bot. Res. Inst. Texas 5(1): 389 - 392. 201 1

393

BOOKS RECEIVED | BOOK NOTICES

John Leslie Dowe. 2010. Australian Palms: Biogeography, Ecology, and Systematics. (ISBN 978-

0643096158, pbk.). CSIRO Publishing, 150 Oxford Street (PO. Box 1139), Collingwood VIC 3066,

AUSTRALIA. (Orders: www.publish.csiro.au/pid/6164.htm; publishing-sales@csiro.au). AU140.00,

290 pp., b/w and color photos, IOC" x SC".

Howe Island, Norfolk Island, £

Sara Oldfield. 2010. Botanic Gardens: Modern-Day Arks. (ISBN 978-0-262-01516-5, hbk.). The MIT

Press, 55 Hayward Street, Cambridge, Massachusetts 02142-1493, U.S.A. (Orders: mitpress.mit.edu/

catalog/item/default. asp?ttype=2&tid= 12399; special_sales@mitpress.mikt.edu). $29.95, 240 pp., 200

color pho

Juliet C. Stromberg and Barbara Tellman, eds. 2009. Ecology and Conservation of the San Pedro River.

(ISBN 978-0-8165-2752-6, hbk.). The University of Arizona Press, 355 S. Euclid Ave., Suite 103, Tucson,

Arizona 85719, U.S.A. (Orders: www.uapress.arizona.edu/Books/bid2087.htm; 800-426-3797). $85.00,

529 pp., 7" x 10".

Klaus Mehltreter, Lawrence R. Walker, and Joanne M. Sharpe, eds. 2010. Fern Ecology. (ISBN 978-0-521-

89940-6, hbk.; 978-0-521-72820-1, pbk.). Cambridge University Press, The Edinburgh Building,

Cambridge CB2, 8RU, UK. (Orders: www.cambridge.org; www.cambridge.org/aus/catalogue/catalogue.

asp?isbn=978052 1728201 &ss=fro) . AUD$99.95, 460 pp., 93 b/w illus., 14 color illus., 32 tables, 9 3 /4 M

Walter Kingsley Taylor. 2009. A Guide to Florida Grasses. (ISBN 978-0-8130-3319-8, flexbound with

flaps). University Press of Florida, 15 Northwest 25 th Street, Gainesville, Florida 32611-2079, U.S.A.

J. Bot. Res. Inst. Texas 5(1): 393. 2011

Journal of the Botanical Research Institute of Texas 5(1)

Patricia A. Harding. 2008. Huntleyas and Related Orchids. (ISBN 978-0-88192-884-6, hbk.). Timber Press,

The Haseltine Building, 133 S.W Second Avenue, Suite 450, Portland, Oregon 97204-3527, U.S.A.

(Orders: www.timberpress.com/books/huntleyas_related_orchids/harding/9780881928846; www.

timberpress.com). $39.95, 260 pp., 150 color photos, 5 line drawings, 7 3 /s" x lOYs".

Alexander Krings. 2010. Manual of the Vascular Flora of Nags Head Woods, Outer Banks, North

Carolina. (ISBN 978-0-89327-500-6, hbk.). Memoirs of the New York Botanical Garden, Volume 1 \ j

(ISSN 0077-8931). The New York Botanical Garden Press, 200th Street & Southern Boulevard, Bronx,

New York 10458-5126, U.S.A. (Orders: www.nybgshop.org/Manual-of-the-Vascular-Flora-of-Nags-

Head-Woods-Outer-Banks-North-Carolina-p-21584.html; Customer Service 718-817-8869). $62.00,

260 pp., illustrated, 7 1 /2 1 ' x lOVi".

From the Publisher: “The Nags Head Woods complex (Oute

Lane Greer and John M. Dole. 2009. Woody Cut Stems for Growers and Florists: How to Produce and

Use Branches for Flowers, Fruit, and Foliage. (ISBN 978-0-88192-892-1, hbk.). Timber Press, The

Haseltine Building, 133 S.W Second Avenue, Suite 450, Portland, Oregon 97204-3527, U.S.A. (Orders:

www.timberpress.com). $39.95, 512 pp., 137 color photos, 9 line drawings, 6" x 9".

Matthew A. Jenks and Andrew J. Wood. 2007. Plant Desiccation Tolerance. (ISBN 978-0-8138-1263-2,

hbk.). Blackwell Publishing Professional, 2121 State Avenue, Ames, Iowa 50014, U.S.A. (Orders: www.

blackwellprofessional.com; www.wiley.com/WileyCDA/WileyTitle/productCd-0813812631.html; 1-800-

0862-6657). $219.95, 340 pp., 7" x 10".

J. Bot. Res. Inst. Texas 5(1): 394.2011

396

Journal of the Botanical Research Institute of Texas 5(1)

MISSOURI BOTANICAL GARDEN PRESS (ST. LOUIS, MISSOURI)

Peruvian Flora

Rodolfo VAsquez Martinez, Rocio Rojas GonzAles, and Henk van der Werff, eds. 2010. Flora del Rio Cenepa,

Amazonas, Peru. Volumen 1. Introduccion, Pteridophyta, Gymnospermae y Angiospermae

(Acanthaceae-Fabaceae); Volumen 2. Angiospermae (Gentianaceae-Zingiberaceae). (ISBN

978-930723-93-1 978-930723-94-8 [vol. 2], hbk.). Monographs in Systematic Botany from the

Missouri Botanical Garden, Volume 114 (ISSN 0161-1342). Missouri Botanical Garden Press, PO. Box

299, St. Louis, Missouri 63199-0299, U.S.A. (Orders: www.mbgpress.org). $140.00 (2 vols.), 1,568

pp., illus., 7" x 10".

(ISBN 978-930723-88-7, hbk.). Science Press, 16 Donghuangchenggen North Street, Beijing 100717,

CHINA and Missouri Botanical Garden Press, PO. Box 299, St. Louis, Missouri 63199-0299, U.S.A.

(Orders: www.mbgpress.org). $90.00, 520 pp., illus., 8 3 A" x IDA".

Wu Zhengyi and Peter H. Raven, Co-chairs of the editorial committee, Hong Deyuan, Vice co-chair of the edi-

torial committee. 2009. Flora of China: Volume 25, Orchidaceae. (ISBN 978-930723-90-0, hbk.).

Science Press, 16 Donghuangchenggen North Street, Beijing 100717, CHINA and Missouri Botanical

Garden Press, PO. Box 299, St. Louis, Missouri 63199-0299, U.S.A. (Orders: www.mbgpress.org).

$125.00, 570 pp., illus., 8 3 /4 M x IDA".

J. Bot. Res. Inst. Texas 5(1): 396. 2011