Journal of the Botanical Research Institute of Texas

J. Bot Res. Inst Texas ISSN 1934-5259

History and Dedication

1962—Lloyd H. Shinners

(left), a member of the

Southern Methodist University

(SMU) faculty and a prolific

researcher and writer, published the first issues of Sida,

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

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

botany at SMU and director emeritus of BRIT,

inherited editorship and copyright.

1993—BRIT becomes publisher/copyright holder.

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

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The BRIT Press seeks innovation and excellence in

preparation, manufacture, and distribution

of botanical research and scientific discoveries

for the twenty-first century.

The BRIT Press—bringing out the best in botanical

science for plant conservation and education.

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The BRIT Press considers original research papers

concerned with classical and modern systematic

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

All submissions are peer-reviewed.

Guidelines for submissions are available

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

Bibliographical

Citation abbreviation for the

Journal of the Botanical Research Institute of Texas is

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

of B.P.H. (informally

International Standard Serial No. (ISSN 1934-5259)

Frequency of Publication

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

(summer/fall) as one volume

by the Botanical Research Institute of Texas.

VOLUME 8 NUMBER 1 9JUL2014

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Table of Contents

SYSTEMATICS

New species and combinations in Solanum section Androceras (Solanaceae)

Stephen R. Stern, Lynn Bohs, and Jeffrey Keeling 1

Sedum kiersteadiae (Crassulaceae), a newly described species from the Klamath Region

of California, U.S.A.

Barbara L. Wilson, Richard E. Brainerd, and Nick Otting 9

Hexasepalum teres (Rubiaceae), a new combination

Joseph H. Kirkbride, Jr. 17

Passiflora soliana, una especie nueva de Passiflora (Passifloraceae) del Paclhco

Sur de Costa Rica

Armando Estrada Ch. & Gerardo Rivera 19

Cyperus stewartii (Cyperaceae), a new species from Cocos Island, Costa Rica

Gordon C. Tucker 25

Calathea gordonii (Marantaceae), a new endemic Panamanian species

Helen Kennedy 31

Calathea cofaniorum and C. shishicoensis, new endemic species of Marantaceae from Ecuador

Helen Kennedy 37

Glossoloma velutinum (Gesneriaceae), a new species from the Cordillera Central of the

Colombian Andes

Larri A. Rodas and John L. Clark 43

Synopsis of Galvezia (Plantaginaceae: Antirrhineae), including a new cryptic species

from southern Peru

Michael O. Dillon and Victor Quipuscoa Silvestre 47

A new species of Cremosperma (Gesneriaceae) from northeastern Peru

Brian R. Keener and John L. Clark 57

Three new species of Senegalia (Fabaceae) from Brazil

David S. Seigler, John E. Ebinger, Petala Gomes Ribeiro, and Luciano Paganucci De Queiroz 61

A new variety of Phanera glauca subsp. tenuiflora (Fabaceae: Caesalpinioideae) from India

Rajib Gogoi and Subir Bandyopadhyay 71

Taxonomic status of Kobresia curvata and Kobresia fragilis (Cyperaceae)

Bikash Jana and R.C. Srivastava 77

First valid place of publication of Duchesnea indica (Rosaceae: Potentilleae)

James L. Reveal and Barbara Ertter 83

Errata: Four new annual species of Euphorbia section Tithymalus (Euphorbiaceae)

from North America

Mark H. Mayfield 85

DEVELOPMENT AND STRUCTURE

The genus Echinacea (Asteraceae): Floral, stem, and petiole morphology

Harold W. Keller 87

The floral structure of three weedy species of Sida (Malvaceae)

Junior Cezar Muneratto, Luiz Antonio de Souza, and OdairJose Garcia de Almeida 127

PALEOBOTANY

A gasteroid fungus, Palaeogaster micromorpha gen. & sp. nov. (Boletales)

in Cretaceous Myanmar amber

George O. Poinar, Jr., Donis da Silva Alfredo, and Iuri Goulart Baseia 139

Xylaria antiqua sp. nov. (Ascomycota: Xylariaceae) in Dominican amber

George O. Poinar, Jr. 145

FLORISTICS, ECOLOGY, AND CONSERVATION

Flora of the halophytic grasslands in the Valle de Janos, Chihuahua, Mexico

Jose Humberto Vega-Mares, Andres Eduardo Estrada-Castillon, Jose Angel

Villarreal-Quintanilla, and Gustavo Quintana Martinez 151

Primer registro de Astrocasia peltata (Euphorbiaceae) en Costa Rica

Irene Calderon Sanou 165

Pleurisanthes flava (Icacinaceae): A new record for Brazil

Bruno S. Amorim, Rodrigo Duno de Stefano, and Marccus Alves 169

Ecology and conservation of Acacia and Prosopis (Fabaceae) woodlands of the

Mojave Desert, U.S.A.

Scott R. Abella and Kenneth L. Chittick 175

Noteworthy vascular plant collections from the Red River of Arkansas and Louisiana, U.S.A.

Christopher Reid and M. Jerome Lewis 197

Vegetation and vascular flora of tallgrass prairie and wetlands, Black Squirrel Creek drainage,

south-central Colorado: Perspectives from the 1940s and 2011

Sylvia Kelso, Leah Fugere, Miroslav Kummel, and Sebastian Tsocanos 203

A floristic inventory of Dagny Johnson Key Largo Hammock Botanical State Park and

immediately adjacent lands (Monroe County), Florida, U.S.A.

George J. Wilder, Susan V. Sprunt, Janice A. Duquesnel, and Susan F. Kolterman 227

A quantitative study of the vegetation surrounding populations of Zigadenus densus

(Melianthiaceae) at Fort Polk in west central Louisiana, U.S.A.

Jacob Delahoussaye, Charles Allen, Stacy Huskins, and Ariel Dauzart 253

A quantitative study of the vegetation surrounding populations of Uvularia sessilifolia

(Colchicaceae) at Fort Polk in west central Louisiana, U.S.A.

Ariel Dauzart, Charles Allen, Stacy Huskins, and Jacob Delahoussaye 261

Blyxa aubertii (Hydrocharitaceae) new to Mississippi, U.S.A.

Daniel M. McNair and Mac H. Alford 267

Floristic studies in north central New Mexico, U.S.A. The Sangre de Cristo Mountains

Jill Larson, Brian Reif, B.E. Nelson, and Ronald L. Hartman 271

Low genetic diversity and poor dispersal, but not conservation status rank, are linked

to climate change vulnerability

Cynthia M. Morton and Matthew D. Schlesinger 305

The vascular flora of Fort Sumter and Fort Moultrie, South Carolina, U.S.A.

Richard Stalter, Brent A. Berger, Eric E. Lamont, and John Nelson 319

Expanded distribution of Gratiola quartermaniae (Plantaginaceae) in Texas, U.S.A.

Kimberly Norton Taylor and Robert J. O’Kennon

333

The vascular flora of Galveston Island State Park, Galveston County, Texas, U.S.A.

David J. Rosen, Shiron K. Lawrence, and Andrew Sipocz

An annotated flora of Reed Plateau and adjacent areas, Brewster County, Texas, U.S.A.

Wendy Weckesser and Martin Terry

Addendum to the vascular flora of Nash Prairie, Texas, U.S.A.

David J. Rosen

Book Reviews, Notices, and Announcements 8 , 16 , 30 , 36 , 46 , 56 , 70 , 76 , 138 , 144 , 150 , 164 ,

174 , 196 , 202 , 226 , 252 , 260 , 304 , 332 , 338,380

INDEX to new names and new combinations in J. Bot. Res. Inst Texas 8(1), 2014

Calathea cofaniorum H. Kenn., sp. nov. —37

Calathea gordonii H. Kenn., sp. nov. —31

Calathea shishicoensis H. Kenn., sp. nov. —41

Cremosperma inversum B.R. Keener & J.L. Clark, sp. nov. —57

Cyperus stewartii G.C. Tucker, sp. nov. —25

Galvezia elisensii M.O. Dillon & Quipuscoa, sp. nov.— 49

Glossoloma velutinum J.L. Clark & L.A. Rodas, sp. nov. —43

Hexasepalum teres (Walter) J.H. Kirkbr., comb. nov. —17

Palaeogaster Poinar, Alfredo, & Baseia, gen. nov. —140

Palaeogaster micromorpha Poinar, Alfredo, & Baseia, sp. nov. —140

Passiflora soliana A. Estrada & G. Rivera, sp. nov. —19

Phanera glauca subsp. tenuiflora var. gandhiana Gogoi & Bandyop., var. nov. —71

Sedum kiersteadiae B.L. Wilson & R.E. Brainerd, sp. nov. —9

Senegalia harleyi Seigler, Ebinger, & P.G. Ribeiro, sp. nov. —64

Senegalia hatschbachii Seigler, Ebinger, & P.G. Ribeiro, sp. nov. —66

Senegalia irwinii Seigler, Ebinger, & P.G. Ribeiro, sp. nov. —62

Solanum cordicitum S. Stern, sp. nov. —2

Solanum knoblochii (Whalen) S. Stern, comb. & stat. nov. —5

Solanum novomexicanum (Bartlett) S. Stern, comb. & stat. nov. —6

Xylaria antiqua Poinar, sp. nov. —146

NEW SPECIES AND COMBINATIONS IN SOLANUM

SECTION ANDROCERAS (SOLANACEAE)

Stephen R. Stern

Department of Biological Sciences

Colorado Mesa University

1260 Kennedy Ave

Grand Junction, Colorado 81501, U.S. A.

sstern@coloradomesa.edu

Lynn Bohs

Department of Biology

University of Utah

257 South 1400 East

Salt Lake City, Utah 84112, U.S. A.

Jeffrey Keeling

Department of Biology

Sul Ross State University

Alpine, Texas 79832, U.S.A.

ABSTRACT

A new species of Solanum from Texas is described here. Solanum cordicitum S. Stern is a member of Solanum section Androceras. It is simi¬

lar to S. citrullifolium and S. davisense but differs from both in having white corollas, and differs from the latter in having inflorescences with

a significantly longer axis and larger flowers. In addition to the new species, three new combinations are proposed for species in Solanum

section Androceras, Solanum setigeroides (Whalen) S. Stern, Solanum novomexicanum (Bartlett) S. Stern, and Solanum knoblochii

(Whalen) S. Stern.

RESUMEN

Se describe aqui una especie nueva de Solanum de Texas. Solanum cordicitum S. Stern es un miembro de Solanum seccion Androceras. Esta

especie es similar a S. citrullifolium y S. davisense, pero se diferencia de ambas por tener flores con corola blanca, y de la ultima por sus inflo¬

rescences mas largas y flores mas grandes. Tambien se proponen tres combinaciones nuevas de Solanum seccion Androceras, Solanum se¬

tigeroides (Whalen) S. Stern, Solanum novomexicanum (Bartlett) S. Stern, y Solanum knoblochii (Whalen) S. Stern.

INTRODUCTION

A comprehensive project to complete species-level taxonomic treatments and resolve phylogenetic relation¬

ships within the genus Solanum L., supported by the National Science Foundation Planetary Biodiversity In¬

ventory program, has facilitated detailed systematic study of many little known groups within this giant genus

(Knapp et al. 2004; http://www.nhm.ac.uk/solanaceaesource). One of the largest groups within Solanum, the

Leptostemonum clade, includes approximately 350-400 species (Bohs 2005; Levin et al. 2006; Weese & Bohs

2007; Stern et al. 2011) and is commonly known as the “spiny solanums” due to the presence of prickles. Phy¬

logenetic work on this clade has identified a number of monophyletic groups, one of which corresponds to So¬

lanum section Androceras (Nutt.) Whalen (Stern et al. 2010).

Solanum section Androceras is an unusual group within the spiny solanums due to its floral characteris¬

tics, namely its bilaterally symmetrical, heterantherous, and enantiostylous flowers. Most Solanum species

have fleshy berry fruits, but those of section Androceras are dry with a prickly, accrescent calyx. The section has

flavenoids that are unique among known flavenoids in other Solanum groups and has an atypical north tem¬

perate geographic distribution. Whalen (1979) provided a detailed revision of the section and included 12 spe¬

cies and 10 varieties. Stern et al. (2010) used molecular phylogenetic techniques to examine the relationships

among these taxa. Results from that study, and further taxonomic work on Solanum section Androceras, have

uncovered an undescribed Solanum species from Texas and indicates that some taxa described as varieties

should be recognized as distinct species. These new names and combinations are validated here. A revision of

Whalen’s (1979) species key, including the 16 currently recognized species, is also provided.

J.Bot. Res. Inst. Texas 8(1): 1-7.2014

Journal of the Botanical Research Institute of Texas 8(1)

Solanum cordicitum S. Stern, sp. nov. (Figs. 1—2). Type: UNITED STATES. Texas. Jeff Davis Co.: Valentine, Blk 76, Bell 2nd

Add., 23 Sep 1990 (fl, fr) H. Elder 46 (holotype: TEX [00402851]).

Similar to Solanum citrullifolium and S. davisense; differs from both in having white corollas; differs from S. davisense in having inflores¬

cences with a significantly longer axis and larger flowers.

Herb to 35 cm. Stems armed with acicular prickles to 5 mm in length, tan to brown, the base to 1 x 0.5 mm,

sparsely pubescent with simple, uniseriate, multicellular hairs 1-2 mm long, moderately pubescent with sim¬

ple, uniseriate, multicellular, gland-tipped hairs 0.5-1 mm long. Flowering portions of stem of difoliate sym-

podial units, the leaves usually geminate, those of a pair often slightly unequal. Leaves simple, the blades 3-8

x 1.5-4 cm, deeply lobed to pinnatihd or pinnatisect with 3-4 lobes per side, chartaceous, green on both sur¬

faces, the adaxial surface nearly glabrous with occasional simple, uniseriate, unicellular to multicellular hairs

0.5-1 mm long, the abaxial surface sparsely pubescent with stellate hairs, the stalks absent to 0.5 mm, multise-

riate, the rays 2-4, 0.5-1 mm long, unicellular to multicellular, the midpoints 0.5-1 mm long; venation pin¬

nate, the secondary veins 3-4 on each side of the midvein and one per lobe, the midrib and larger secondary

veins occasionally with a few prickles like those of the stem; base obtuse, often asymmetrical; margin deeply

lobed and the lobes with irregularly undulate margins; apex rounded to obtuse; petioles 0.5-3 cm, moderately

pubescent with hairs like those of the stem, sparsely armed with prickles like those of the stem. Inflorescence

8-12 cm, extra-axillary, unbranched, with 5-8 flowers, the axes moderately pubescent with hairs like those of

the stem, moderately armed with prickles like those of the stem; peduncle 2-3 cm; rachis 6-12 mm; pedicels

4-10 mm in flower, 10-18 mm in fruit, spaced 6-12 mm apart, articulated at the base. Flowers 5-merous, zygo-

morphic, enantiostylous. Calyx 4-6 mm long, the tube 1.5-3 mm, the lobes 2-4 x 0.5-1.2 mm, narrowly trian¬

gular, moderately pubescent abaxially with hairs like those of the stem, moderately armed with prickles like

those of the stem; fruiting calyx 9-13 mm, strongly accrescent, completely covering the fruit, densely armed

with prickles like those of the stem. Corolla 2-2.5 cm in diameter, chartaceous, white, rotate-stellate, with

abundant interpetalar tissue, shallowly lobed, the lobes 2-3 x 0.5-1.5 mm, narrowly triangular, sparsely pu¬

bescent on abaxial midveins with hairs like those of the stem, adaxially glabrous. Stamens dimorphic, the

lowermost one 10-12 mm, the upper four 5-8 mm; filaments 1-2 mm long, glabrous; lowermost anthers 9-11

x 1-2 mm, opposite the style in alternating right and left-handed flowers, distally curved upward, yellow; up¬

per anthers 5-6 x 1-2 mm, straight, yellow, all anthers linear-lanceolate, tapering, the base cordate, the apex

acute, the pores directed slightly introrsely, not opening into longitudinal slits. Ovary glabrous; style 10-14 x

0.5-1 mm, cylindrical, glabrous, opposite the lowermost anther and alternating between right and left-handed

flowers; stigma to 1 mm wide. Fruit 10-12 mm in diameter, globose, tightly invested in the prickly accrescent

calyx creating a burr-like fruit, green, turning black, drying and tearing apart at maturity, glabrous. Seeds

30-40 per fruit, ca. 1.5 x 1 mm, chocolate-brown, reniform, the surface with raised ridges.

Distribution and Phenology. —Known only from Jeff Davis Co., Texas from 1350-1820 m in elevation. The

specimens were flowering in September-November and fruiting in September-November.

Etymology.—Solanum cordicitum is taken from the Latin “cordicitus“ for “from the heart” referring to the

type locality of Valentine, Texas.

Conservation Status. —The conservation status of S. cordicitum, according to the IUCN Red List Categories

(IUCN 2010) is Data Deficient due to the low number of collections. Despite searching herbaria, including

BRIT, NY, TAES, TAMU, TEX, US, and UTEP, and collection efforts near Valentine, Texas in 2010 and 2013,

only three collections of S. cordicitum are known. It is our hope that this species description will encourage

further collecting in Jeff Davis County so that the status of S. cordicitum can be determined.

Solanum cordicitum has an overlapping geographic distribution with various members of section Androc-

eras, including S. rostratum, S. davisense, both varieties of S. tenuipes, and S. citrullifolium vars. citrullifolium and

setigerum. Unlike any of these species, S. cordicitum has white corollas. Additionally, S. cordicitum differs from

S. rostratum in that the latter has stellate hairs on the stem while those of S. cordicitum are simple. Solanum

tenuipes is typically found further south in the Big Bend area of Texas and is a perennial whereas S. cordicitum

is an annual. The leaves of S. cordicitum are not thrice pinnatihd as they are in S. davisense and the lowermost

Stern et al., New species and combinations in Solanum

Fig. 1 . Solanum cordicitum S. Stern, A. Habit. Note zygomorphic flower buds. B, C, D. Flowers. E. Dissected corolla. Note elongate lower anther. F. Upper

stamens. G. Lower stamen from a left- and right-handed flower. All from H. Elder 46 (TEX).

Journal of the Botanical Research Institute of Texas 8(1)

PLANTS OF TEXAS

#46 SOLANACEAE

Solanum heterodoxum Dunal

JEFF DAVIS CO. Valentine, TX. BLK 76 BELL 2ND

ADD. in open areas under Prosopis, Ephedra,

with Cucurbita, Bouteloua, Hilaria.

Medium annual herb up to 35cra tall;

herbage and fruit prickly; petals white,

anthers long and yellow; fruit a spiny

capsule.

1YU&C V

granaijiorum Whalen

June 2006

The University of Texas Herbarium <LL, TEX)

Solanum davisense M. D. Whalen

Del: B. L. Turner, 3 May 1997

Siiiltf

Coll: H. Elder 23 September 1990

k< — ————

Fig. 2. Holotype of Solanum cordicitum S. Stern [Elder 46 (TEX)].

Stern et al., New species and combinations in Solanum

anthers of the latter are shorter (6-8 mm in S. davisense versus 9-11 mm in S. cordicitum). Solanum cordicitum

also has longer inflorescences than S. davisense (8-12 cm in S. cordicitum versus 4-7 cm in S. davisense) and

larger flowers (2-2.5 cm in diameter in S. cordicitum versus 1.3-2 cm in S. davisense). The prickles of S. cordici¬

tum are more widely spaced (<20 per cm) than those of S. citrullifolium var. setigerum (>30 per cm). The flowers

of S. cordicitum are generally smaller (2-2.5 cm in diameter) than those of S. citrullifolium var. setigerum

(2.5-3.5 cm in diameter).

Paratypes: UNITED STATES. Texas. Jeff Davis Co.: 30 mi W of Ft. Davis on US 166, 6000 ft, 17 Oct 1974 (fr), Heller 24 (UTEP); town of

Valentine, Elder property, 30°35 , 40"N, 104°39 , 32"W, 1350 m, 7 Nov 2013 (fl, fr),J. Keeling 445 (MESA).

The three combinations below represent taxa previously recognized as varieties by Whalen (1979). These spe¬

cies each have distinctive morphological characteristics and geographical ranges that distinguish them from

other species in sect. Androceras. Additionally, molecular phylogenetic analyses (Stern et al. 2010) have shown

that they do not form monophyletic groups with the other varieties of their respective species.

Solanum knoblochii (Whalen) S. Stern, comb. & stat. nov. Solanum citrullifolium var. knoblochii Whalen, Wrightia 5:237.

1976. Type: MEXICO. Chihuahua: Mojarachic, 16 Aug 1940, Knobloch 8006 (holotype: US [00027510]).

Whalen (1976,1979) designated the variety S. citrullifolium var. knoblochii and distinguished it from S. citrulli¬

folium var. setigerum by the much denser pubescence of the latter (each cm of stem with over 25 prickles versus

scattered prickles < 20 per cm of stem in the former). It is also differentiated from S. citrullifolium var. citrullifo¬

lium by the short (<1.2 mm), uniseriate hairs of the latter versus the presence of much longer (> 2 mm), unise-

riate hairs in var. knoblochii. The easternmost distribution of var. setigerum overlaps with the westernmost

distribution of var. citrullifolium in west Texas. However, S. knoblochii has a disjunct distribution from both of

these varieties and is only known from two populations in the Tarahumara country of western Chihuahua,

Mexico (Whalen 1976).

These distinct morphological characters, isolated geographical range, and evidence from molecular

data (Stern et al. 2010) indicate that S. citrullifolium var. knoblochii should be recognized as a distinct species.

The name Solanum knoblochii was retained, honoring Irving Knobloch, the first collector of this species in

Chihuahua.

The remaining varieties of S. citrullifolium, vars. citrullifolium and setigerum, did not form a monophyletic

group in Stern et al. (2010). Instead, they formed a strongly supported clade with S. davisense and S. hctcro-

doxum var. setigeroides. These species have overlapping distributions and similar morphologies with the main

difference being small flowers in S. hcterodoxum var. setigeroides (corolla <1.5 cm across, large anther < 5 mm),

medium-sized flowers in S. davisense (corolla ca. 1.7 cm across, large anther 6-8 mm), and large flowers in both

varieties of S. citrullifolium (corolla > 2 cm, large anther > 10 mm). There is also reported hybridization between

S. citrullifolium var. setigerum and S. hcterodoxum var. setigeroides (Whalen 1979). Solanum heterodoxum var.

setigeroides is recognized as a distinct species below because it is not monophyletic with the other varieties of

S. heterodoxum in the molecular phylogenetic analyses of Stern et al. (2010). Although S. citrullifolium vars.

citrullifolium and setigerum also did not form a clade in Stern et al. (2010), resolution and support is low in this

part of the tree. Further morphological and molecular data are needed to assess the taxonomic status of S.

citrullifolium vars. citrullifolium and setigerum as well as that of S. davisense.

Conservation Status. —The status of S. knoblochii using the IUCN Red List Categories (IUCN 2010) is Data

Deficient given the few collections and lack of long term monitoring. Recent collections, in addition to the

original collections by Knobloch, show that the range of this species is approximately 34,000 km 2 . This range

is sufficient to place the species out of the Threatened category, but the limited number of collections (only five

are known from two populations) is concerning and warrants Near Threatened status.

Two varieties of S. heterodoxum are designated as distinct species:

Solanum setigeroides (Whalen) S. Stern, comb. & stat. nov. Solanum heterodoxum var. setigeroides Whalen, Wrightia 5:237.

1976. Type: UNITED STATES. New Mexico. Grant Co.: 12 mi W of Silver City, 4 Aug 1975, Whalen 201 (holotype: TT [TT-00372877];

isotypes: MO [MO-503667], WIS).

Journal of the Botanical Research Institute of Texas 8(1)

Solanum novomexicanum (Bartlett) S. Stern, comb. (Srstat. nov. Solanum heterodoxum var. novomexicanum Bartlett, Proc.

Amer. Acad. Arts 44:628. 1909. Androcera novomexicana (Bartlett) Woot. & Standi., Contr. U.S. Natl. Herb. 16:170. 1913. Type:

UNITED STATES. New Mexico: Santa Fe, creek valley, foot of mountains, sunny side, 1847, Fendler 673 (holotype: GH [GH-

00077421]; isotypes: F, GH [GH-00077422], MO [MO-503664]).

Solanum heterodoxum var. setigeroides was recognized as distinct from S. heterodoxum vars. heterodoxum and

novomexicanum due to the dense, narrow prickles on its stems (> 30 prickles per cm, mostly less than 0.5 mm

in diameter versus < 20 prickles per cm, often to 1 mm wide in the latter varieties). Solanum heterodoxum var.

novomexicanum was differentiated from var. heterodoxum by its pentagonal-stellate corolla with narrow deltoid

lobes versus pentagonal corollas with ample interpetalar tissue in the latter. The distributions of these varieties

are also disjunct, with var. heterodoxum occurring in central Mexico from Veracruz to San Luis Potosl, var. se¬

tigeroides in northern Chihuahua, Mexico and westernmost Texas to central Arizona and New Mexico, and

var. novomexicanum in northern New Mexico. Additionally, molecular phylogenetic data in Stern et al. (2010)

found that these varieties do not form a monophyletic group and belong in distinct clades.

Etymology .—The name S. setigeroides was retained from the Latin word “setigermeaning “bristly,” refer¬

ring to the bristly prickles of the stem. Solanum novomexicanum was originally recognized at the species level

as Androcera novomexicanum, so the epithet was transferred and refers to fact that this species is endemic to

New Mexico.

Conservation Status .—The conservation status of S. setigeroides according to the IUCN Red List Catego¬

ries (IUCN 2010) is Least Concern. This is a widespread, weedy species with no obvious threats. Although S.

novomexicanum is not as widespread or as frequently collected as S. setigeroides, its designation according to the

IUCN Red List Categories (IUCN 2010) is also Least Concern due to its relatively weedy nature. Additionally,

S. novomexicanum is afforded some protection with populations in Pecos Ruin National Historic Park and the

Cibola National Forest.

KEY TO THE SPECIES OF SOLANUM SECTION ANDROCERAS

(REVISED FROM WHALEN 1979)

1. Cauline hairs stellate or multiangulate; corollas mostly yellow, rarely pale blue or white.

2. Corollas pale blue or white_ S. tribulosum S. Schauer

2. Corollas yellow.

3. Plants perennial, woody-based; cauline stellae echinoid, some of them with 15 or more rays; distribution in eastern

Durango, Mexico_ S.johnstonii Whalen

3. Plants annual, taprooted; cauline stellae usually with 12 or fewer rays.

4. Large anther essentially glabrous.

5. Hilum of seed sunken in a deep notch; large anther hardly distinct, less than 6 mm long; corollas less than 2

cm across; cauline prickles broad-based, flattened and often recurved_ S. fructo-tecto Cav.

5. Hilum of seed not sunken in a deep notch; large anther very distinct from shorter ones, over 9 mm long; corol¬

las more than 2 cm across; cauline prickles seldom much flattened or recurved_ S. rostratum Dunal

4. Large anther bearded on the proximal portion of the ventral surface_ S. angustifolium Mill.

1. Cauline hairs simple, often glandular, occasionally absent; corollas usually violet, blue, or white, seldom yellow.

6. Corollas violet or blue.

7. Large anther 6 mm long or more; corollas 1.4 cm or more across; stigma unexpanded or only weakly capitate

8. Plants perennial, woody- or corky-based; seeds plump, 2.8 mm long or more_ S. tenuipes Bartlett

8. Plants annual, taprooted; seeds lenticular, shorter than 3 mm

9. Large anther 6-8 mm long; corollas ca. 1.7 cm across; buds obovoid, more or less radially symmetrical; large

leaves often thrice pinnatifid, with acute ultimate lobes_ S. davisense Whalen

9. Large anther 10 mm long or more; corollas 2 cm or more across; buds noticeably curved, bilaterally sym¬

metrical; large leaves usually only twice pinnatifid, with obtuse to acute ultimate lobes.

10. Cauline hairs mostly <0.3 mm with occasional hairs reaching to 1 mm; distributed from central to west

Texas to western Coahuila and eastern Chihuahua, Mexico_ S. citrullifolium A. Braun

10. Cauline hairs often reaching 2.5 mm in length; known only from western Chihuahua, Mexico_ S. knoblochii

(Whalen) S. Stern

7. Large anther 5 mm long or less; corollas 1.5 cm or less across; stigma capitate, twice as thick as the style.

11. Stems sparsely pubescent with simple, glandular hairs ca. 0.2 mm long; stems densely prickly, each cm with 30

or more acicular prickles, the bases less than 0.5 mm in diameter_ S. setigeroides (Whalen) S. Stern

11. Stems densely pubescent with simple, glandular hairs 0.2-0.4 mm long; stems sparsely prickly, each cm with

20 or fewer acicular prickles, the base up to 1 mm in diameter.

Stern et al., New species and combinations in Solanum

12. Corolla pentagonal, with ample, plicate interpetalar tissue; distribution in Mexico, from Veracruz to San Luis

Potosi_ S. heterodoxum Dunal

12. Corolla pentagonal-stellate, with narrowly deltoid lobes; distribution in U.S.A. (northern New Mexico)

_S. novomexicanum (Bartlett) S. Stern

6. Corollas white or yellow.

13. Stems with well-spaced, acicular prickles; corollas pentagonal-stellate with ample, plicate interpetalar tissue, 2 cm

or more across; seeds reticulately ridged.

14. Corolla yellow, 2.5 cm or more across; large anther bearded on the proximal portion of the ventral surface;

distribution from tropical Mexico east to Honduras_ S. angustifolium Mill.

14. Corolla white, 2-2.5 cm across; large anther glabrous; distribution in western Texas, USA_ S. cordidtum S. Stern

13. Stems with dense, filiform, bristle-like prickles; corollas distinctly stellate with little interpetalar tissue, 2.5 cm or

less across; seeds radially ridged at margin.

15. Anthers of three sizes, the longest one flanked by two of intermediate length; mature seeds large, 3 mm or

longer, with the hilum sunken in a deep notch; distribution from southern Arizona, USA to northern Sinaloa,

Mexico_ S. lumholtzianum Bartlett

15. Stamens dimorphic, one large, the other four smaller and essentially equal; mature seeds small, 3 mm long or

less, not deeply notched; distribution from southern Sonora to Puebla and Guerrero, Mexico.

16. Leaves with simple and/or glandular hairs above, or glabrous; plants widespread along the Pacific slope of

Mexico_ S. grayi Rose

16. Leaves with stellate hairs above; endemic near Matamoros, Puebla, Mexico_ S. leucandrum Whalen

ACKNOWLEDGMENTS

We thank G.J. Anderson and one anonymous reviewer for suggestions; herbaria for their loans, particularly

BRIT, LL, TAES, TAMU, TEX, UTEP, and US; Rebecca Ely for assistance in the held; and Bobbi Angell for the

drawing. This work would not have been possible without the exceptional revision of section Androceras by

M.D. Whalen. This work was supported by NSF grant DEB-0316614 (PBI Solanum: A Worldwide Treatment) to

LB.

REFERENCES

Bohs, L. 2005. Major clades in Solanum based on ndhF sequence data. In: R.C. Keating, V.C. Hollowell, andT.B. Croat, eds.

A festschrift for William G. D'Arcy:The legacy of a taxonomist. Mongr. Syst. Bot. Missouri Bot. Gard.Vol. 104. Missouri

Botanical Press, St. Louis, Missouri, U.S.A. Pp. 27-49.

IUCN Standards and Petitions Subcommittee. 2010. Guidelines for using the IUCN Red List Categories and Criteria. Version

8.0. Prepared by the Standards and Petitions Subcommittee in March 2010. Downloadable from http://intranet.iucn.

org/webfiles/doc/SSC/ RedList/RedListGuidelines.pdf. [accessed 5 October 2010].

Knapp, S., L. Bohs, M. Nee, & D.M. Spooner. 2004. Solanaceae - a model for linking genomics with biodiversity. Comp. Funct.

Genomics 5:285-291.

Levin, R.A., N.R. Myers, & L. Bohs. 2006. Phylogenetic relationships among the "spiny solanums" [Solanum subgenus Lept-

ostemonum, Solanaceae). Amer. J. Bot. 93:157-169.

Stern, S.R., T. Weese, & L. Bohs. 2010. Phylogenetic relationships in Solanum section Androceras (Solanaceae). Syst. Bot.

34:885-893.

Stern, S.R., M.F. Agra, & L. Bohs. 2011. Molecular delimitation of clades within New World species of the "spiny solanums"

[Solanum subg. Leptostemonum). Taxon 60:1429-1441.

Weese, T.L. & L. Bohs. 2007. A three-gene phylogeny of the genus Solanum (Solanaceae). Syst. Bot. 32:445-463.

Whalen, M.D. 1976. New taxa of Solanum sect. Androceras. Wrightia 5:228-239.

Whalen, M.D. 1979. Taxonomy of Solanum section Androceras. Gentes Flerb. 11:359-426.

Journal of the Botanical Research Institute of Texas 8(1)

BOOK REVIEW

Tracey Parker. 2014. Arboles Comunes de Nicaragua/Common Trees of Nicaragua. (ISBN-13: 978-0-

9718739-1-9, pbk). The Tree Press, Suite 650, #125, 3300 Bee Cave Road, Austin, Texas 78746, U.S.A.

(Orders: www.thetreepress.com, 1-800-247-6553). $44.95, 404 pp., 170+ line drawings, 1200+ color

photos, map, glossary, index, 6" x 9". Fully bilingual text (Spanish and English).

A fully bilingual guide to the trees and tree-like plants of Nicaragua, this book is packed to the gills with infor¬

mation. Each species is presented over two pages: one for Spanish and English descriptions and one for color

photographs. Each description page includes family name, Spanish and English common names, distribution,

morphological description, habitat, ethnobotanical or commercial uses, and notes (a section for interesting

facts and phenology). In addition, every description page also includes a line drawing and, where appropriate,

taxonomic synonyms. The photo page accompanying each species includes anywhere from 5 to 10 color pho¬

tographs of leaves, fruits, flowers, seeds, bark, habitat, and depiction of usage (e.g., a woman at a market balanc¬

ing a tray of mango slices on her head).

Organized alphabetically by family, genus, and species (with monocots, dicots, and gymnosperms inter¬

mixed), the book also includes an index of both scientific and common names as well as an English and Span¬

ish glossary of botanical terms, making it more novice-friendly. The front matter, however, is bare bones—-just

a map of Nicaragua and a brief introductory note from the author. Truly, this book is all about the trees, devot¬

ing the majority of its 400+-page heft to those wonderfully concise descriptions that manage to clearly depict

the essentials of each species and do so in two languages. Personally, I admire the commitment to full dual¬

language text; by doing so, the author must have had to sacrifice some content, but the resulting text is not only

streamlined and elegant but also accessible to a much wider audience.

Though I cannot speak to the quality of the taxonomic presentation, I believe Common Trees of Nicaragua

to be a well-executed work relevant to the held guide genre. The inclusion of so many color photographs as well

as the line drawings really give this book an edge over other held guides, though I would have liked to see the

line drawings expanded in size where page content allowed. (Some description pages are completely filled

while others have some usable white space remaining.... Why not use that space to increase hgure size?) Other

than this, however, the layout is clean and consistent, the organization logical, pleasing, and easy to navigate.

Dr. Parker also published Trees of Guatemala in 2008, and if it is anything like her current work, I will most

assuredly be making a second purchase. —Brooke Byerley Best, PhD, Botanical Research Institute of Texas, Fort

Worth, Texas, U.S.A.

From the publisher: TRACEY PARKER, PhD, forest ecologist, environmental consultant, professor and photog¬

rapher, holds a bachelor’s degree in forestry from Colorado State University, and masters and doctorate from

the University of Idaho. Dr. Parker moved with her family from Nepal to Guatemala in 1994, and began teach¬

ing dendrology in the Forestry Department at the Universidad del Valle. She held the position of Regional En¬

vironmental Advisor at the U.S. Agency for International Development (USAID) where her work took her to all

the countries of Central America, expanding her knowledge of the vegetation throughout the isthmus.

J. Bot. Res. Inst. Texas 8(1): 8.2014

SEDUM KIERSTEADIAE (CRASSULACEAE), A NEWLY DESCRIBED SPECIES

FROM THE KLAMATH REGION OF CALIFORNIA, U.S.A.

Barbara L. Wilson, Richard E. Brainerd, NickOtting

Carex Working Group

1377Alta Vista Drive

Corvallis, Oregon 97330, U.S.A.

bwiison@peak.org

ABSTRACT

Sedum kiersteadiae is a locally common succulent of rocky, often ultramafic substrates at mid to high elevations, narrowly endemic to the

western Klamath region of northern California. It is recognized by its widely spreading yellow petals and its relatively loose rosettes with

visible internodes.

RESUMEN

Sedum kiersteadiae en una suculenta localmente comun de sustratos rocosos, a menudo ultramaficos en elevaciones de medianas a altas,

endemica del oeste de la region Klamath del norte de California. Se reconoce por sus petalos amarillos ampliamente extendidos y sus rosetas

relativamente flojas con internudos visibles.

INTRODUCTION

In the Klamath Region of northern California and southwest Oregon, Sedum section Gormania has diversified

into six morphologically similar species. Two of the species, S. obtusatum A. Gray and S. laxum (Britton) Berger

(Boyd & Denton 2012) have been treated as having four subspecies each (Denton 1982). The section was stud¬

ied by Clausen (1942,1975), and its classification was further refined by Denton (1982), who had access to ad¬

ditional specimens. This research was made readily accessible by Sedum treatments in the Jepson Manual

(Denton 1993; Boyd & Denton 2012). However, as held botanists worked with populations unknown to previ¬

ous researchers, they found problems with those treatments: similar plants from the same area were often

identified as belonging to two or more species, certain supposedly rare plants were encountered frequently,

and many plants had combinations of traits that did not fully match any of the published descriptions.

In 2011-2013, we studied variation and classification in this group. Full results of the study, including an

identification key, will be published elsewhere. Here we raise to the rank of species one of the more distinctive

elements in Sedum section Gormania. Although the type specimen of S. obtusatum subsp. boreale R.T. Clausen

is an example of this taxon, we find that Clausen’s descriptions do not fully match the plant as we understand

it, so a full description is provided here. The plant cannot be treated as a species under its original epithet be¬

cause that is preempted by Sedum boreale Hort. ex Guenthart (Guenthart 1902).

Sedum kiersteadiae B.L. Wilson & R.E. Brainerd, sp. nov. (Figs. 1-2). Type: U.S.A. California. Shasta Co.: NE of Slate

Mountain, W of USFS Road 37N08Y; T36N R6W S2, SW A of NW V4,41.01159°, -122.53148°, 4500 ft, 15 Jun 2012, Lindstrand III &

VanSusterenNSR-17 (holotype: OSC; isotypes: CAS, CHSC, DAV, HSC, MO, NY, RSA, UC, WTU).

Sedum obtusatum A. Gray subsp. boreale R.T. Clausen, Bull. Torrey Bot. Club 69:32. 1942. Sedum obtusatum A. Gray var. boreale (R.T.

Clausen) H. Ohba, J. Bot. Res. Inst. Texas 1:889. 2007. Type: U.S.A. California. Siskiyou Co.: Mt. Shasta, E side Mud Creek Canyon,

5600 ft, 26 Jul 1940, R.T. Clausen , W.B. Cooke & H. Trapido 4952 (holotype: BH!; isotype: NY!). Not Sedum boreale Hort. ex Guenthart

(1902).

Sterile rosettes usually loose, with visible internodes; leaves obovate and usually notched; sepals less than half as long as the petals; petals

yellow with midribs usually pink to red, whole petals senescing red, the upper half spreading about 90° from the flower axis.

Description. —Plants succulent, herbaceous, perennial. Rhizomes and stolons to 15 cm long, 3-6 mm diame¬

ter; sterile rosettes often numerous. Rosette leaves often loosely arranged with visible internodes, glaucous,

J. Bot. Res. Inst. Texas 8(1): 9 -15.2014

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 1. Type plant of Sedum obtusatum subsp. boreale from Mud Creek Canyon, Mt. Shasta, Siskiyou Co., California, cultivated in greenhouse, Ithaca New

York, May 1943. A. Habit. B. Flower from above (xl .7). C. Flower from side (xl .7). D. Petal and two stamens (x 2.6). E. Carpels (x 2.6). F. Single carpel

and nectary (x3.4). G. Nectary (x4.3). Drawing by Elfriede Abbe, from Clausen (1975) p. 374. Used with permission from Cornell University Press.

Wilson et al., A new species of Sedum from northern California

Fig. 2. Sedum kiersteadiae. Note widely spreading yellow petals often marked with red on the dorsal surface, and the open spacing of leaves on the sterile

shoot. Photo credits: Whole plant and top flower, Julie Kierstead Nelson. Center and lower flower, and sterile shoot, Peter F. Zika.

Journal of the Botanical Research Institute of Texas 8(1)

flattened, broadly obovate to oblanceolate, cuneate at base, 12-32 x 5-18 mm, 1.2-5.3 times as long as wide,

apex usually notched, sometimes obtuse or truncate. Flowering shoots often reddish, 7.5-28.5 cm, nodding in

bud, erect in flower and fruit. Leaves on flowering shoots usually pink to green, (4-) 11-19 x (2-)5-9 mm, 1-3.7

times as long as wide, narrowly obovate, truncate at base, apex usually obtuse. Inflorescences panicle-like

cymes 4-13.6 x 2-3(-4) cm, usually cylindrical, proximal branches solitary at nodes, ascending. Flowers 10-

30. Calyx greenish, (8-)15-45% as long as the petals, free calyx lobes (0.5-)2-4(-6) mm long, acute. Petals

6—8(—12) mm long, light yellow with midribs usually pink or red dorsally, sometimes white ventrally (some¬

times also dorsally) when young, lower half sometimes red, entire petal eventually senescing red, becoming

whitish or pale tan when dead. Upper half of petal blade narrow, spreading (70-)80-90(130)° from the flower

axis at anthesis, apex acute, usually with subterminal mucro. Stamens 10, shorter than or equaling petals; fila¬

ments white or yellow-green, aging red; anthers usually red, rusty, or dark orange aging purple or black, some¬

times yellow aging light brown, becoming tan when dead and dried. Follicles erect, green aging red, fused

0.5-1 mm at base.

Chromosome number. — 2n = 30 (Clausen 1942, based on count for Clausen et al. 4952).

Habitat and Range. —Rock outcrops and rocky open forests, usually on serpentine substrates, NE Trinity

County and adjacent NW Shasta and S Siskiyou counties, CA, in the Klamath Range, also disjunct on the

southeast side of Mt. Shasta (Fig. 3), 1000-2500 m elevation, more common at higher elevations.

Etymology. —The name S. kiersteadiae honors California botanist Julie Kierstead Nelson, who was in¬

spired to study the natural world by her father, Robert William Kierstead. Conveniently, the Dutch “kier stead”

refers to a place or homestead with cracks; this plant generally grows in cracks in its rocky home.

Classification. —This species belongs in Sedum subgenus Gormania section Gormania (Clausen 1975).

Specimens examined: (KNF = herbarium of the Klamath National Forest; other herbarium acronyms follow Thiers (continuously updat¬

ed)). CALIFORNIA: Shasta Co.: Ridge top N of Sanford Pass; T36N R6W S15, 4600 ft, 15 Jun 2012, Lindstrand & Van Susteren NSR-15

(CHSC, DAV, HSC, KNF, STNF, UBC); The Incline, Upper Slate Creek watershed; N of USFS Road 36N44; T36N R6W S12,4200 ft, 15 Jun

2012, Lindstrand & Van Susteren NSR-18 (CAS, MICH, MO, NY OSC, RSA, WTU, UC). Siskiyou Co.: Mount Shasta, Mud Creek, 5716 ft, 7

Aug 2011, Colberg MEC-1 (MO, OSC, RSA, UC, WTU); on the banks of Mud Creek Canyon, 6500 ft, 22 Aug 2013, Wm. Bridge Cooke 15459

(OSC); Mount Bradley, in rocks 10-50 m W of lookout, also E of lookout in rocky openings, 5556 ft, 12 Jul 2011, Lindstrand & Van Susteren

NSR-09 (BH, F, GH, NY, US); Gray Rocks Lake, downslope of trail to lake near trailhead, T39N R5W S21, 5830 ft, 20 Jul 2011, Lindstrand &

Van Susteren NSR-11 (CAS, CHSC, DAV, HSC, MO, NY, RSA, UBC, UC); Rattlesnake Hill, on ridge N of USFS road 38N21, T37N R5W S20,

5580 ft, 20 Jul 2011, Lindstrand & Van Susteren NSR-12 (BH, GH, KANU); along Mount Eddy Trail, ca. 1.75 mi from trailhead, 7000 ft, 29 Jul

2011, Van Susteren JVS 1 (CAS, OSC, UC); off trail between Heart Lake & Castle Lake below Little Castle, 5950 ft, 30 Jul 2011, Van Susteren

JVS 2 (CHSC, DAV, HSC, UBC); Rail Creek road (road 41N08, to Kanagaroo Lake), 4913 ft, 9 Aug 2011, Wilson et al CWG-18 (F, RENO, UC,

US); ridge line W of Kangaroo Lake, near loop at end of road, 6604 ft, 9 Aug 2011, Wilson et al. CWG-19 (CAS, MO, NY, OSC, WTU); 0.9 mi

by road E of summit on Highway 3 at Scott Mountain, 5076 ft, 31 Jul 2011, Zika & Wilson 25677 (CAS, CHSC, OSC, WTU); Shasta-Trinity

National Forest; 10.1 mi NW of Highway 3 on road 42N17,5777 ft, 31 Jul 2011, Zika & Wilson 25680 (F, RENO, UBC, US, WTU); near Pacific

Crest Trail, ca. 0.5 km NW of Parks Creek Pass and junction with FS road 42N17C and Route 17, 2.5 air mi NW of the NW Deadfall Lake,

6890 ft, 31 Jul 2011, Zika & Wilson 25683 (CAS, OSC, UC, US, WTU); South Fork Willow Creek; N end road 42N19, along South Fork, 3691

ft, 21 Jun 2012, Zika et al. 25901 (DAV, MO, NY, RSA, WTU). Siskiyou/Trinity Co. border: Mount Eddy, N side of ridgeline; T40N R5W

Section 20,6600 ft, 7 Aug 2011, Colberg MEC-2 (CHSC, STNF, WS); Pacific Crest Trail W of Toad Lake, to ridge between Toad and Porcupine

lakes, T40N R6W Section 36, ft, 31 Jul 2011, NelsonJKN-1 (GH, KANU). Trinity Co.: ca. 6 mi N on trail to Canyon Creek Lakes, ca. 10 mi N

of Dedrick, 4400 ft, 24 Jun 1976, Denton 3967 (OSC); Little Boulder Lake Trail #8N11, ca. 0.1 to 0.25 mi up from lake, or 0.1-0.25 mi down

from intersection with main Boulder Lake Trail, 6200 ft, 28 Aug 2011, Erwin SE-1 (KNF, MO, NY, RSA, STNF); Parks Creek Summit, Pacific

Crest Trail, ca. Vs to V4 mi W of trailhead on Forest Road 17, Shasta Trinity National Forest, 6897 ft, 28 Jun 2013, Nelson JKN-2013-1 (GH, WS);

3.7 mi up Canyon Creek Trail, 100 ft E of trail on rock outcrop, accessed by Canyon Creek Road off Hwy 299,4699 ft, 23 Jun 2011, Stubbs et

al. RS-27 (MO, NY); between Cedar and Bear Creeks, Forest Service Road 40N45,2.7 road mi S of junction with FS Road 17,1.7 air mi WSW

of SE Deadfall Lakes, 6496 ft, 31 Jul 2011, Zika & Wilson 25687 (US, WTU); Forest Service Road 40N45, 9.1 road mi S of junction with FS

Road 17, 2.7 air mi NNW of Picayune Lake, 31 Jul 2012, Zika 25689 (OSC, UC, WTU), Forest Service road 25 at Horse Heaven Meadows, 3

air km NNW of Grey Rocks Peak, 28 Jun 2013, Zika 26294 (MO, OSC, RSA, UC, WTU).

DISCUSSION

Most populations of Sedum kiersteadiae occur in the western Klamath Region, including Mount Eddy, Scott

Mountain, and the Trinity Mountains (Fig. 3). Clausen (1975) referred these plants to S. obtusatum subsp. re-

Wilson et al., A new species of Sedum from northern California

Trin ity

AN _

C a if f o r n i a

50 mi

□ NV

” Orego

Jackson

Klamath

TMeS&nL

iv' +

50 km

Fig. 3. Distribution of Sedum kiersteadiae in northern California.

Journal of the Botanical Research Institute of Texas 8(1)

tusum (Rose) Clausen. Denton recognized that they did not belong with that taxon and treated them as a dis¬

junct population of yellow-flowered S. obtusatum subsp. obtusatum. We find that they differ from both these

taxa in their narrower, more spreading petals and more open rosettes. We feel this combination of traits is

convincing taxonomic evidence and unique among Sedum section Gormania, to treat S. kiersteadiae at the spe¬

cies level.

The taxon most similar to S. kiersteadiae is the recently described S. citrinum Zika, endemic to a small area

of Del Norte County, California (Zika 2014). Both species have narrow, widely spreading, yellow petals and

open rosettes. Sedum kiersteadiae differs from S. citrinum in having a panicle-like cyme with elongated lower

branches (Fig. 2), rather than a flat-topped cyme, flowering shoots that never branch from the base, and an¬

thers that are usually rusty brown aging black, rather than yellow aging brown. The two do not overlap in

range.

The very open rosettes of S. kiersteadiae resemble those of S. oregonense (S. Watson) M.E. Peck. Sedum ki¬

ersteadiae differs from S. oregonense in having narrower, widely spreading petals that are yellow, often marked

with red. Such petals are unusual in section Gormania and resemble those of S. lanceolatum Torr., in Sedum

subgenus Sedum section Lanceolata. The range of S. lanceolatum overlaps S. kiersteadiae near Mount Eddy,

where the two species are occasionally found on the same slope. We speculate that perhaps S. oregonense and

S. lanceolatum, or their ancestors, hybridized to produce S. kiersteadiae. The range of S. kiersteadiae approaches

that of S. oregonense in the Canyon Creek watershed, at the west edge of the range of S. kiersteadiae and the

south edge of the range of S. oregonense (Figure 3). In that drainage grow plants that resemble S. oregonense in

having yellow anthers and relatively wide, obtuse petals that lack red, and resemble S. kiersteadiae in having

yellow petals. Petals of these plants spread to about 40°, less than S. kiersteadiae and more than is typical of S.

oregonense. The existence of some intermediate plants in small areas where species ranges overlap is not su¬

prising; all tested pairs of taxa in Sedum section Gormania can produce FI hybrids, many of which are fertile

(Denton 1979). The presence of potential hybrids in a limited area does not argue against recognizing the eas¬

ily distinguished S. oregonense and S. kiersteadiae as distinct species.

The only other taxon of Sedum section Gormania with a range approaching that of S. kiersteadiae is S. ob¬

tusatum subsp. paradisum Denton. It grows in the Canyon Creek watershed at the west edge of the S. kier¬

steadiae range, where both S. oregonense and S. kiersteadiae also grow. Sedum paradisum and S. kiersteadiae also

grow near each other at the south edge of the range of S. kiersteadiae. There, S. kiersteadiae can occasionally be

found on the north or northwest side of a ridge and S. paradisum on the south side; the two were not observed

together and morphologically intermediate plants were not observed during this study (Len Lindstrand III,

pers. comm.).

The affinities of the plants represented by Clausen et al. 4952, the type specimen of S. obtusatum subsp.

boreale, have been interpreted in diverse ways. Originally, S. obtusatum subsp. boreale was described as having

pale to deep yellow, somewhat spreading petals, and the type locality on Mount Shasta, in the southern Cas¬

cade Range, was considered the eastern-most population of the taxon (Clausen 1942). We now interpret the

type specimen as S. kiersteadiae but other specimens cited by Clausen [Siskiyou Co.: Caribou Basin, Salmon-

Trinity Alps, 25 July 1937, J. T. Howell 13450 (CAS); Clausen 1942] are better treated as S. oregonense with white

to pale yellow petals.

Later, Clausen (1975) and Denton (1982) treated the type specimen from Mount Shasta as the northwest¬

ern-most, isolated population of a taxon that was more common in the northern Sierra Nevada. This taxon was

characterized by relatively large mats of dense rosettes and by wide, apically obtuse, white petals that are pink

at the base and senesce pink throughout. We agree that this is an accurate description of the northern Sierra

Nevada plants, but not of Clausen et al. 4952.

Plants collected for this study at the type locality, Mud Creek on Mount Shasta ( Colberg MEC-1, Aug 201 1),

resemble S. kiersteadiae of Mount Eddy and elsewhere in the western Klamath Region in having loose rosettes

and yellow, strongly spreading petals that are marked with red. We believe these are representative of the same

population as Clausen et al. 4952 because the taxon was originally described as having yellow petals (Clausen

Wilson et al., A new species of Sedum from northern California

1942), and strongly spreading petals can be observed on both the holotype specimen and the drawing of a live

clone of the holotype (Clausen 1975, p. 374; Fig. 2). Colberg MEC-1, Aug 2011 and the holotype ( Clausen et al.

4952 ) both have relatively loose rosettes. A single location is usually home to a single species of Sedum section

Gormania (Clausen 1975; pers. obs.), and we have not seen plants with white petals or other evidence that more

than one species occurs at Mud Creek.

We suspect that the confusion in interpreting the S. obtusatum subsp. boreale type specimen was caused

in part by the necessary reliance on cultivated plants in previous studies. Variation in plant height, rosette

density, leaf thickness, and glaucescence has been observed in cultivated plants depending on substrate and

vernalization (Denton 1982). Denton (1982) considered the variations in cultivations too minor to affect taxo¬

nomic decisions. We have also observed differences in petal color and degree of spreading (Steven Darington,

pers. comm.). We have observed some individual Sedum that produce pink or yellow flowers in the wild may

produce white flowers in cultivation. Also, each taxon in Sedum section Gormania has a characteristic degree

to which petals spread, but petals may spread excessively in cultivated plants, obscuring the differences be¬

tween taxa. All these variations add a level of confusion to this already difficult group of plants.

ACKNOWLEDGMENTS

This study was funded by the Klamath, Lassen, Mendocino, Shasta-Trinity, and Tahoe National Forests. We

thank the following for providing specimens and information, Caitlin Coberly, Mary Ellen Colberg, Steve

Darington, Talitha Derkson, Ryan Elliot, Tom Engstrom, Susan Erwin, Lawrence P. Janeway, A. Jefferson, Bob

Korfhage, Jenna Lee, Martin J. Lenz, Len Lindstrand III, Andrew Maguire, Julie K. Nelson, Lusetta Nelson,

Rhonda Posey, Cindy Roche, Sierra Spooner, Rebecca Stubbs, Dean W. Taylor, Jane Van Susteren, Sharon Voll-

mann, Jennifer Wheeler, Margaret Widdowson, Gabe Youngblood, and Peter F. Zika. We especially thank

Steven Darington and Peter F. Zika for their insights. Mark E. Mort, Kanchi Gandhi, and an anonymous re¬

viewer provided helpful reviews.

REFERENCES

Boyd, S. & M.F. Denton. 2012. Sedum. In: B.G. Baldwin, D.H. Goldman, DJ. Keil, R. Patterson, TJ. Rosatti, and D.H. Wilken,

eds. The Jepson manual: vascular plants of California, 2nd edition. University of California Press, Berkeley, California,

U.S.A. Pp. 674-676.

Clausen, R.T. 1942. Studies in the Crassulaceae - III. Sedum, subgenus Gormania, section Eugormania. Bull. Torrey Bot.

Club 69:27-40.

Clausen, R.T. 1975. Sedum of North America north of the Mexican Plateau. Cornell University Press, Cornell, New York,

U.S.A.

Denton, M.F. 1982. Revision of Sedum section Gormania (Crassulaceae). Brittonia 34:48-77. www.jstor.org/sta-

ble/2806401.

Denton, M.F. 1993. Sedum. ln:J.C. Flickman, ed. The Jepson manual: higher plants of California. University of California

Press, Berkeley, California, U.S.A. Pp. 531-534.

Gonthart, A. 1902. Beitrage zur Bluthenbiologie der Cruciferen, Crassulaceen und der Gattung Saxifraga. Erwin Naegele,

Stuttgart, Germany.

Thiers, B. [continuously updated]. Index herbariorum: A global directory of public herbaria and associated staff. New

York Botanical Garden's Virtual Herbarium, http://sweetgum.nybg.org/ih/

Zika, P.F. 2014. A new species of stonecrop ( Sedum section Gormania, Crassulaceae) from northern California. Phytotaxa

159:111-121.

Journal of the Botanical Research Institute of Texas 8(1)

BOOK REVIEW

Michael W. Beug, Alan E. Bessette, and Arleen R. Bessette. 2014. Ascomycete Fungi of North America: A

Mushroom Reference Guide. The Corrie Herring Hooks Series, Number 69. (ISBN-13: 978-0-292-

75452-2, cloth, alk. 128 gsm matte paper, paper permanence: ANSI/NISO Z39-48-1992 (R1997)). Uni¬

versity of Texas Press, PO Box 7819, Austin, Texas 78713-7819, U.S.A. (Orders: www.utexaspress.com,

1-800-252-3206). $85.00, 502 pp., 843 color photos, glossary, references, index, 3.9 lbs, 7" x 10" x 1.5".

This is a heavy, bigger book format that far exceeds the coverage of anything previously published for the asco¬

mycete fungi. Beautiful color images! Easy to read and understand text! Stunning macroscopic color habit

photographs! Easy to use picture keys to different ascomycete groups! A bargain basement price! A special

book that falls in the category of “once in a lifetime”! A much needed book that fills a fungal niche vacant for a

long time! These are some of the platitudes that highlight the exceptional features of this book.

The durable book binding (round back Smyth Sewn) should stand the test of time, but this book is best

used at home, in the laboratory, or at the end of the day at display tables to identify specimens rather than as a

held book carried in a backpack. The authors have focused their content on continental United States of

America and Canada. The species described and illustrated are macroscopic in size and likely to be seen by

mushroom hunters in the held.

There are 12 chapters beginning with an Introduction that dehnes the sexual ascus and ascospore stages,

highlighted by Jim Murray’s close-up color image of the rare Texas ascomycete Chorioactis geaster (Devils’s

Cigar) caught in the act of pufhng a cloud of ascospores. This ascomycete species is represented by habit pho¬

tographs that were taken associated with dead, decomposing stumps of Ulmus crassifolia (Cedar Elm) at River

Legacy Parks, Arlington, Texas; the only known locations outside of Japan are in the state of Texas.

Classihcation of the Ascomycetes would have benehted from a taxonomic rank ordering of the different

groups assembled as part of a phylogenetic tree and thus a roadmap for the different families that follow. Chap¬

ter 2 contains the keys that include a combined, mostly dichotomous key with leads interspersed with habit

pictures of different species, beginning with the above- and below-ground Ascomycetes. These picture images

start with the truffles and morels, some of the choice fungal edibles, and include nearly 550 species in total.

This is a lengthy key, extending from page 16 to 69 with 73 leads, so it requires staying power to identify the 12

picture images on the last page of the key.

Chapter 3 treats the hypogeous Ascomycetes with a key to 26 genera. These taxa are represented by pic¬

ture images along with species descriptions. The color images usually occupy about a half-page, either on the

same page as the species description or next to it on the facing page. This conveniently locates both species

description and color image visually in the same space but also unfortunately creates blank white spaces that

break up the flow of the narrative. Each species narrative includes the topical headings Macroscopic Features,

Microscopic Features, Occurrence, and Comments, but unfortunately these topical headings are difficult to

see because they lack contrast formatting. The genus Tuber ; highly prized for its aroma and high quality culi¬

nary delicacy, is included here with five species.

Chapter 4 covers the Pezizomycetes (pp. 117-272, the largest chapter), including the Chorioactidaceae

with Chorioactis and the epigeous (above-ground) Morchellaceae that includes the popular and choice edible

morels. Morels are highly prized edible fungi often hunted, collected, and eaten as a culinary delicacy by my-

cophiles during the spring months in North America and March in Texas. Members of this group are typically

operculate (they have asci with lids) and often forcibly eject spores into the air.

Chapter 5, the Sodariomycetes (pp. 272-352), are sometimes referred to as the pyrenomycetes or stone

fungi, characterized by a sexual stage, the perithecium, and formation of asexual conidia. Certain

(continued on p. 30)

J.Bot. Res. Inst. Texas 8(1): 16.2014

HEXASEPALUM TERES (RUBIACEAE), A NEW COMBINATION

Joseph H. Kirkbride, Jr.

USDA-ARS, US. National Arboretum

Floral & Nursery Plants Research Unit

3501 New York Avenue NE

Washington, DC 20002-1958, US.A.

joseph.kirkbride@ars.usda.gov

ABSTRACT

The new combination Hexasepalum teres (Rubiaceae) is made for a widespread, New World species distributed from the eastern U.S.A. to

Bolivia and Paraguay.

RESUMEN

La nueva combinacion Hexasepalum teres (Rubiaceae) esta hecha para una especie que esta ampliamente difundida en el Nuevo Mundo, y

distribuida desde el este de los EE.UU. hasta Bolivia y Paraguay.

Small (1913) described the genus Diodella Small as a segregate from the genus Diodia L. and transferred a single

species name into the genus as Diodella rigida (Cham. & Schltdl.) Small. Later in the year (Small & Carter

1913) he transferred a second species, as Diodella teres (Walter) Small. For the next 85 years, Diodella was

treated as a junior synonym of the genus Diodia. In 1999 Bacigalupo and Cabral revised the genus Diodia for

the Americas. They treated Diodia in a very narrow sense, reducing it to five species, and suggested that 16 spe¬

cies then included in Diodia belonged to the genus Diodella without making the necessary new combinations.

Over the last 14 years most of the new combinations in the genus Diodella have been made by various authors.

Recently Cabana Fader et al. (2012) discovered that Diodella and Hexasepalum Bartl. ex DC. (1830) are

generic synonyms. They proposed that Hexasepalum and its single species, H. angustifolium Bartl ex DC., be

rejected. They argued that since Hexasepalum is older than Diodella, the 16 species of Diodella would have to be

transferred to Hexasepalum, which would cause significant nomenclatural disruption. The Nomenclature

Committee for Vascular Plants (Applequist 2013) did not recommend rejection of Hexasepalum and H. angusti¬

folium. The Committee felt that since most usage of the name Diodella has only been in the last decade, adop¬

tion of the name Hexasepalum would not cause excessive nomenclatural disruption.

The taxon known as Diodia teres Walter or Diodella teres occurs in North America, north of Mexico. For

the Flora of North America, I will treat it in a genus distinct from Diodia, thus it is necessary to transfer the spe¬

cific epithet of Diodia teres into the genus Hexasepalum.

RESULTS

Hexasepalum teres (Walter) J.H. Kirkbr., comb. nov. Basionym: Diodia teres Walter, FI. carol. 87.1788. Type: united states.

South Carolina: Georgetown Co.: Georgetown, old field, 24 Aug 1939, R.K. Godfrey & R.M. Tryon 1682 (neotype, designated by Ward

2008:476; GH00277018; isoneotypes: NY1163926! US1838313!).

Distribution. —Native: United States (Alabama, Arizona, Arkansas, California, Colorado, Connecticut, District

of Columbia, Delaware, Florida, Georgia, Illinois, Indiana, Iowa, Kansas, Kentucky, Louisiana, Massachusetts,

Maryland, Michigan, Mississippi, Missouri, North Carolina, New Jersey, New Mexico, New York, Ohio, Okla¬

homa, Pennsylvania, Rhode Island, South Carolina, Tennessee, Texas, Virginia, Vermont, West Virginia,

Wisconsin), Mexico, Cuba, Hispaniola, Jamaica, Margarita, Belize, Guatemala, Honduras, El Salvador, Nicara¬

gua, Costa Rica, Panama, Colombia, Venezuela, Guyana, Surinam, French Guiana, Ecuador, Peru, Bolivia,

Brazil, Paraguay; introduced: Netherlands, Cape Verde, Gambia, Guinea-Bissau, Senegal, Angola, southeast

China, Japan, Korea.

J. Bot. Res. Inst. Texas 8(1): 17 -18.2014

Journal of the Botanical Research Institute of Texas 8(1)

Fernald and Griscom (1937) discussed Diodia teres in the eastern United States and published three new

varieties in the species. They presented the following, “Mr. C.A. Weatherby, upon looking for Walter’s type,

reports that there is no Walter material of it in his herbarium at the British Museum; but he found in Paris that

the type of Spermacoce diodina Michx., commonly referred to it, is the common and well known weed with

fruits 2.9-3.6 mm long, covered with short appressed hairs ... and greatly exceeded by the stipules, and the

leaves without prominently setiform tips. Since the latter plant is common all the way from Florida to New

Jersey we are selecting it to stand as typical of Walter’s species.” Various authors (Steyermark 1972:798; Del-

prete 2010:371) have interpreted Fernald and Griscom’s text as a neotypihcation of Diodia teres on the Michaux

specimen of Spermacoce diodina in the Michaux herbarium at P. This is based on a misapplication of Art. 7.10

of the ICN (McNeill et al. 2012), which states “designation of a type is achieved only ... if the type element is

clearly indicated by direct citation including the term “type” (typus) or an equivalent...” The word “typical” in

the last sentence has been interpreted as an equivalent of “type” in the nomenclatural sense, but it is actually

used in the sense of the species S. diodina being representative of the species Diodia teres. Since “typical” was

not used in the sense of the word “type,” i.e. a single specimen, this was not a neotypihcation of Diodia teres.

Steyermark (1972:798) cited the collection locality of Michaux’s specimen as the type of Diodia teres, but

as he did not clearly indicate the type element, as required by Art. 7.10 (McNeill et al. 2012) for an existing

name, he did not achieve typihcation. This interpretation is consistent with Art. 40, Note 2 for new names,

where citation of a collection locality without the collector’s name, collection number, or date would not be

adequate for typihcation. In citing the collectors’ name and number, the collection locality and date, and the

herbarium in which the specimen is deposited, Ward (2008) effectively neotypihed the species.

ACKNOWLEDGMENTS

I thank John Wiersema, USDA-ARS National Germplasm Resources Laboratory, Beltsville, Maryland for his

nomenclatural help and review of the manuscript. Steven P. Darwin (NO) and David H. Lorence (PTBG) pro¬

vided helpful reviews.

REFERENCES

Applequist, W.L. 2013. Report of the nomenclature committee for vascular plants: 65. Taxon 62:1315-1326.

Bacigalupo, N.M. & E.L. Cabral. 1999. Revision de las especies americanas del genero Diodia (Rubiaceae, Spermacoceae).

Darwiniana 37:153-165.

Cabana Fader, A.A., R.M. Salas, & E.L. Cabral. 2012. (2114-2115) Proposals to reject the names Hexasepalum and H. angus-

tifolium (Rubiaceae).Taxon 61:1333-1334.

Candolle, A.P. de. 1830. CLXXVI. Hexasepalum Bartl. in H. Haenke. Prodr. 4:561.Treuttel et Wurtz, Paris. France.

Delprete, P.G. 2010. Rubiaceae, parte 1: Introdugao, generos A-H. In: J.A. Rizzo, ed. Flora dos Estados de Goias eTocantins.

Universidade Federal de Goias, Goiania, Brazil. 40:1 -580.

Fernald, M.L. & L. Griscom. 1937. III. Notes on Diodia. Rhodora 39:306-308.

McNeill, J., F.R. Barrie, W.R. Buck, V. Demoulin, W. Greuter, D.L. Hawksworth, P.S. Herendeen, S. Knapp, K. Marhold, J. Prado, W.F.

Prud'homme van Reine, G.F. Smith, J.H. Wiersema, & NJ. Turland. 2012. International code of nomenclature for algae, fungi,

and plants (Melbourne Code). Koetlz Scientific Books, Konigstein.

Small, J.K. 1913. Flora of Miami. J.K. Small, New York, U.S.A.

Small, J.K. & J.J. Carter. 1913. Flora of Lancaster County. J.K. Small and JJ. Carter, New York, U.S.A.

Steyermark, J.A. 1972. 38. Diodia (Gronovius) ex Linnaeus. Mem. New York Bot. Gard. 23:788-805.

Ward, D.B. 2008. Thomas Walter typification project, V: neotypes and epitypesfor 63 Walter names of genera D through

Z. J. Bot. Res. Inst. Texas 2:475-486.

PASSIFLORA SOLIANA, UNA ESPECIE NUEVA DE PASSIFLORA

(PASSIFLORACEAE) DEL PACIFICO SUR DE COSTA RICA

Armando Estrada Ch.

Museo Nacional de Costa Rica

Apartado Postal 749-1000

San Jose, COSTA RICA

aestrada@museocostarica.go.cr

Gerardo Rivera

grivera@biogenesiscr.com

www.biogenesiscr.com

Biogenesis, COSTA RICA

RESUMEN

Passiflora soliana (subg. Passiflora), una nueva especie procedente del canton de Osa, en el Pactfico Sur de Costa Rica, es descrita e ilus-

trada. Esta especie nueva es afin a Passiflora brevifila Killip de elevaciones mayores, de la cual se distingue por sus flores mas pequenas, sin

pedicelos o estos muy cortos, y principalmente por la morfologia y menor tamano de sus bracteas florales.

ABSTRACT

Passiflora soliana (subg. Passiflora), a new species from the canton of Osa in the South Pacific of Costa Rica is described and illustrated.

This new species is closely related to Passiflora brevifila Killip of higher elevations, but P. soliana is distinguished by its smaller flowers,

without or nearly without pedicel, and mainly by morphology and smaller size of the floral bracts.

Passiflora L. con mas de 560 especies, es por mucho el genero mas grande de Passifloraceae; compuesto desde

enredaderas o bejucos herbaceos principalmente en vegetacion secundaria hasta lianas y algunos arbustos o

arboles pequenos de bosques primarios (Hansen et al. 2006; Krosnick et al. 2013). Es un genero casi exclusivo

del Nuevo Mundo, su distribucion se extiende desde el sur de los Estados Unidos hasta el norte de Argentina,

incluyendo Las Antillas. Solo 24 especies son endemicas de las regiones tropicales y subtropicales del sureste

de Asia, Australia y Nueva Zelanda (Krosnick et al. 2013). En Costa Rica el genero esta representado por 51

especies (Rodriguez y Estrada 2007).

A nivel infragenerico, Killip (1938) dividido Passiflora en 22 subgeneros y mas recientemente Feuillet y

MacDougal (2003) propusieron reducir la cantidad de subgeneros a cuatro en un nuevo sistema de clasih-

cacion: Astrophea (DC.) Mast., Deidamioides (Harms) Killip, Decaloba (DC.) Rchb. y Passiflora. Un quinto

subgenero llamado Tetrapathea (DC.) RS. Green, que incluye tres especies del viejo mundo ha sido reconocido

por Krosnick et al. (2009, 2013). El subgenero Passiflora, al cual pertenece esta especie nueva, incluye cerca de

250 especies y se caracteriza por la presencia de zarcillos, hojas enteras o lobadas, glandulas sobre peciolos,

estlpulas y margenes de hojas, flores grandes con multiples hlamentos de la corona, 3 bracteas usualmente fo-

liaceas, operculo tubular o hlamentoso y limen membranaceo (MacDougal y Feuillet 2004; Hansen et al. 2006;

Krosnick et al. 2013).

Passiflora soliana A. Estrada & G. Rivera, sp. nov. (Figs. 1, 2). Tipo: COSTA RICA: Puntarenas: Canton de Osa, Distrito

Piedras Blancas, La Florida, finca Bellavista, a orilla de rio Bellavista, 8°46.872'N, 83°12.483'W, 297 m, 17 Feb 2012 (fls.), G. Rivera

& M. Nunez 4884 (holotipo: CR; isotipo: MO)

Species nova haec Passiflora brevifila Killip propinqua, sed taxon novum per sequentes floribus characteres aberratur: Flores breviores (4-

4.3 cm diametro), sine pedicello, bracteae florales breviores (2-2.3 x 2-2.5 cm diametro), ovatae usque ad suborbiculatas, cum marginibus

revolutis et in parti proximali connatae, superficie adaxiali papillosa, sine tomento marginali, corona filamentosa biseriata, filamenta ex¬

terna alba, breviora (7-9 mm longa).

Bejuco lenoso, glabro; tallos cillndricos, estriados; estlpulas de 5-8 x 0.7-1 mm, lineares a subfalcadas, enteras,

uno o dos glandular denticuladas en un lado y tambien con una glandula apicalmente, coriaceas. Hojas con

peciolos de 1-2.8 cm, con (4-)6(-7) glandulas, corto-estipitadas en la mitad distal; lamina de 12-17.5(-23) x

4.8-6.5(-8) cm, entera, lanceolada, oblongo-lanceolada a ellptica, acuminada en el apice, cortamente aristada,

J. Bot. Res. Inst. Texas 8(1): 19-24.2014

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 1. A-H. Passiflorasoliana. A. Habito; B. Estipulasy glandulas peciolares; C. Flor, vista frontal; D. Flory bracteas florales, vista lateral.

Estrada & Rivera, Passiflora soliana, una nueva especie de Costa Rica

Fig. 2. A-G. Passiflora soliana A. Vista lateral de boton floral; B. Bracteas florales; C y D. Vista frontal de flor; E. Sepalos y petalos; F. Detalle de operculo

y limen; G. Corte longitudinal de flor; H-J. Passiflorabrevifila H. Flor; I. Detalle de la tercera serie coronal mas interna adyacente al operculo; J. Bractea

floral ypedicelode flor.

Journal of the Botanical Research Institute of Texas 8(1)

84°0

obtusa en la base, venation pinnada (nervios laterales 6 a 8 por lado, arqueado-ascendentes), nervios terciarios

resaltados y conspicuamente reticulados en el enves, membranaceas a cartaceas, lustrosas, margen entero. In-

florescencias axilares, de una flor solitaria, pedunculo de 2.6-2.8 cm, solitarios, articulados en el apice, en la

union con las bracteas, bracteas de 2-2.4 x 2-2.5 cm, foliaceas, enteras, ovadas a sub-orbiculares, agudas en el

apice, connadas en la mitad proximal, margenes doblados hacia afuera (revolutos), abiertas y desplegado antes

de la antesis. Flores 4-4.3 cm de diametro, sin pedicelo o este muy corto ca. 2 mm, con el tubo floral campanu-

lado, sepalos de 1.7-1.8 x 0.7-0.8 cm, verde claro externamente, blancos en el interior, corniculados; petalos de

1.5-1.8 x 0.4-0.6 cm blancos; filamentos de la corona, en dos series, los externos de 7-9 mm, blancos, los inter-

nos de 2-3 mm, purpura-azulados, estrechamente liguliformes; operculo un estrecho borde adjunto a la coro¬

na, liso, margen de ca. 1 mm, crenulado, erecto; limen una delgada membrana en forma de copa, cerrando el

operculo, margen crenulado; androginoforo de 9-10 mm; ovario elipsoide, glabro, verde claro. Frutos descono-

cidos.

Fenologia .—Flores presentes en febrero.

Distribution y Habitat.—Passiflora soliana es una especie endemica de Costa Rica. Se distribuye en

Estrada & Rivera, Passiflora soliana, una nueva especie de Costa Rica

Cuadro 1. Diferencias morfologicas y ecologicas entre Passiflora soliana y Passiflora brevifila.

Passiflora soliana

Passiflora brevifila

Flores sin pedicelo o este muy corto (max. 2 mm)

Flores de 4-4.3 cm de diametro

Bracteas florales de 2-2.3 x 2-2.5 cm, sin una banda marginal

tomentosa, pero con la superficie papilosa adaxialmente,

connadas en toda su mitad proximal, ovadas a sub-orbiculares,

abiertas antes de la antesis, con los margenes revolutos

Corona de filamentos en 2 series. Filamentos mas externos

de la corona de 7-9 mm, blancos

Distribucion alrededor de 300 m, bosques muy humedos,

vertiente pacifica

Flores pediceladas, pedicelos de 5-7 mm

Flores de 6-6.5 cm de diametro

Bracteas florales de 4-7 x 1.8-3.4 cm, con una estrecha banda

marginal tomentosa adaxialmente, superficie no papilosa,

connadas solo en la base (maximo Vio de su longitud),

lanceoladas a estrechamente ovadas, cerradas y cubriendo el

boton floral hasta la antesis, con los margenes rectos, sin

doblarse

Corona de filamentos usualmente con una tercera serie de

filamentos mas interna de ca. 1 mm, adyacente a la base del

operculo. Filamentos mas externos 9-14 mm, bicoloros, blancos

en la mitad distal y purpura en la mitad proximal

Distribucion de 800-2000 msnm, en bosques nubosos, muy

humedos y pluviales, ambas vertientes

bosques muy humedos del Paclfico Sur, alrededor de los 300 m de elevacion. Se ha recolectado en orillas de rlos

en vegetacion secundaria (Fig. 3).

Discusidn.—Passiflora soliana es una especie afln a Passiflora brevifila (Killip 1960; fig. 2, H-J) pero se

distingue de esta por sus flores mas pequenas, sin pedicelos o estos muy cortos, la coloracion y numero de fila¬

mentos de la corona floral y principalmente por la morfologla y menor tamano de sus bracteas florales (ver

Cuadro 1). Adicionalmente Passiflora soliana crece en bosques muy humedos en elevaciones bajas (300 m) del

Paclfico Sur, mientras que P. brevifila crece en bosques nublados, muy humedos y pluviales en elevaciones

mayores (800-2000 msnm), cerca de la division continental en todas los principales cordilleras, en ambas

vertientes. A nivel infragenerico esta especie nueva puede ubicarse, de acuerdo a la clasihcacion de Feuillet &

MacDougal (2003), en el subgenero Passiflora, principalmente por sus peciolos con glandulas y sus 3 bracteas

florales foliaceas, pero no se ajusta claramente a las supersecciones establecidas dentro de este subgenero, lo

cual requiere un estudio mas detallado y especlhco.

Etimologia. —El eplteto hace homenaje a Marisol Nunez, quien con su tiempo, esfuerzo y dedicacion fue

de gran ayuda para descubrir y recolectar esta nueva especie de passiflora.

Paratipo: COSTA RICA. Puntarenas. Osa: Piedras Blancas, La Florida, finca Bellavista, a orillas de rio Bellavista, 8°46.872'N, 83°12.483'W,

297 m, 7 May 2012, G. Rivera & M. Nunez 5090 (CR, MO, USJ).

AGRADECIMIENTOS

Agradecemos a Matt Hogan por las facilidades y apoyo brindado durante la investigacion realizada en su

propiedad Finca Buenavista. A John MacDougal y Christian Feuillet por la revision y sus valiosos aportes al

manuscrito, a Carlos O. Morales (USJ) por su ayuda en la elaboracion de la diagnosis latina y Pedro Juarez por

las ilustraciones realizadas.

REFERENCIAS

Feuillet, C. & J.M. MacDougal. 2003 [2004]. A new infrageneric classification of Passiflora L. (Passifloraceae). Passiflora

13:34-38.

Hansen, A.K, E.G. Lawrence, B.B. Simpson, S.R. Downie, A.C. Cervi, & R.K. Jansen. 2006. Phylogenetic relationships and chromo¬

some number evolution in Passiflora. Syst. Bot. 31:138-150.

Killip, E.P. 1938. The American species of Passifloraceae. Publ. Field Mus. Nat. Hist., Bot. Ser. 19:1 -613.

Killip, E.P. 1960. Supplemental notes on the American species of Passifloraceae, with descriptions of new species. Contr.

U.S. Natl. Herb. 35(1 ):16.

Krosnick, S.E., AJ. Ford, & J.V. Freudenstein. 2009. Taxonomic revision of Passiflora subgenus Tetrapathea including the

monotypic genera Hollrungia and Tetrapathea (Passifloraceae), and a new species of Passiflora. Syst. Bot. 34:375-385.

Journal of the Botanical Research Institute of Texas 8(1)

Krosnick, S.E, K.E Porter-Utley, J.M. MacDougal, P.M. J0rgensen, & L.A. McDade. 2013. New insights into the evolution of

Passiflora subgenus Decaloba (Passifloraceae): Phylogenetic relationships and morphological synapomorphies. Syst.

Bot. 38:692-713.

MacDougal J.M. & C. Feuillet. 2004. Systematics 27-31. In T. Ulmer & J.M. MacDougal, Passiflora: Passionflowers of the

World.Timber Press.

Rodriguez, A. & A. Estrada. 2007. Passifloraceae. In: B.E. Hammel, M.H. Grayum, C. Herrera and N. Zamora, eds. Manual de

plantas de Costa Rica. Vol. 6. Monogr. Syst. Bot. Missouri Bot. Gard. 111:862-891.

CYPERUS STEWARTII (CYPERACEAE), A NEW SPECIES FROM

COCOS ISLAND, COSTA RICA

Gordon C. Tucker

Dept, of Biological Sciences

Eastern Illinois University

Charleston, Illinois 61920, U.S.A.

gctucker@eiu.edu

ABSTRACT

Cyperus stewartii G.C. Tucker, sp. nov. is described. It is known from two collections from Cocos Island (Isla del Coco), Costa Rica, made

in 1905. This species is most similar to C. thyrsiflorus and C. lentiginosus, species of the southern U.S., Caribbean slope of Mexico and Central

America, and northern South America. It differs from both in its smaller anthers, shorter and narrower floral scales, closer spacing of scales on

the rachilla, and very wide spacing of spikelets on the rachilla.

RESUMEN

Se describe Cyperus stewartii G.C. Tucker, sp. nov. Se conoce de dos muestras de la Isla del Coco, Costa Rica, colectadas en 1905. Esta es-

pecie es muy similar a C. thyrsiflorus y C. lentiginosus, especies del sur de EE.UU., vertiente del Caribe de Mexico y America Central y el norte

de America del Sur. Se diferencia de ambos en sus anteras pequenas, escamas florales mas cortas y mas estrechas, escamas de la raquilla mas

densamente espaciadas, y una separacion muy amplia entre las espiguillas sobre la raquilla.

Keywords: Cocos Island, new species, Cyperus, Costa Rica, Central America

INTRODUCTION

The genus Cyperus L. includes about 690 species, occurring worldwide in warm temperate and tropical regions

(Tucker 1994,2001). The distribution of individual species ranges from cosmopolitan, e.g., C. squarrosus L. and

C. odoratus L., to regional and narrow endemics found on all continents except Antarctica and Europe (Kuken-

thal 1935-36). In the New World, areas of high diversity and endemism include the southeastern United States,

Mexico, the Greater Antilles, and eastern Brazil (Tucker 2007.)

During the course of this study, and my preceding studies of the Mexican and Central American species

of Cyperus (Tucker 1983,1994), I have examined some 30,000 herbarium specimens from, or examined at, the

herbaria listed in the acknowledgments. During my earlier studies of Cyperus in Costa Rica and Panama

(Tucker 1983), I did not encounter any material of this new species. Then, in 2009, a single specimen was noted

in a loan of unidentified Cyperus from US. I later contacted staff at CAS (California Academy of Sciences) who

were able to locate two additional sheets of this species. Fortunately, between these three specimens, a com¬

plete description of the species can be made.

TAXONOMIC TREATMENT

Cyperus stewartii G.C. Tucker, sp. nov. (Figs. 1, 2). Type: COSTA RICA: Isla del Coco: “Cocos Island, in low ground near

Wafer Bay, California Academy of Sciences Expedition to Galapagos Islands,” 5-7 Sep 1905, A. Stewart 268 (holotype: CAS; isotype:

US).

Affinis Cyperus lentiginosus et Cyperus panamensis sed differt in antherarum minor, squamae brevior angustiorique, in apice divertente,

propius in rachillarum, et spicularum laxe subsequentes intervallent.

Plants perennial (?), 15-29 cm tall. Rhizome short, 2-3 mm long. Culms 0.7-2.3 mm in diameter, trigonous,

smooth. Leaves 2-3, 6-20 cm long, 1.3-2.8 mm wide, inversely w-shaped, the margins scabrellate. Inflores¬

cence bracts 4-6,4-20 cm long, 0.5-3 mm wide, inversely w-shaped, the margins ciliate-scabrellate, ascendent

at 30-45°. Rays 4-8, 1-6 cm long; prophylls 4-8 mm long, truncate. Spikes (10-)18-34 mm long, (12-)18-30

J. Bot. Res. Inst. Texas 8(1): 25 - 29.2014

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 1. Cyperusstewartii: inflorescence (from isotype, A. Stewart268, US).

mm wide, loosely ovoid to oblong-ellipsoid; rachis 5-10 mm long (spikelets widely spaced, ca. 6-10 spikelets/

cm). Spikelets (7-) 10-30,11-16 mm long, 1.6-1.8 mm wide, linear-lanceolate, compressed; rachilla deciduous,

ca. 0.3 mm wide and 0.1 mm thick, flexuous, green, the wings ca. 0.3 mm wide, hyaline, successive scales 2

mm apart. Floral scales persistent, spreading, 6-12, 3.2 mm long, 1.5 mm wide, ellipsoid, subacute, entire to

Tucker, Cyperus stewartii, a new species from Costa Rica

Fig. 2. Cyperus stewartii: habit; (from holotype, A Stewart268, CAS).

Journal of the Botanical Research Institute of Texas 8(1)

Table 1. Comparison of Cypernsstewartii with C. panamensis and C. lentiginosus.

Cyperus panamensis

Cyperus stewartii

Cyperus lentiginosus

Distribution

Pacific coast, Sinaloa

to Ecuador

Cocos Island, Costa Rica

Caribbean lowlands,Texas

to Venezuela

Spikelet density (# per cm of rachis)

22-34

6-10

20-28

Separation of scales along rachilla (mm)

4-5.6

2.5-3

(2.2-)2.6-3.4

Scale shape

Involute

Conduplicate

Scale apex

Appressed, cuspidate

to mucronulate

Spreading, entire

Appressed, mucronate

Scale color

light brown to golden

brown, medially green

silvery-white, reddish

toward center, medially

green

off-white to stramineous, red

speckled, medially green

Anther length (mm)

0.4-0.7

0.3-0.35

0.5-0.8(-1.4)

Achene length (mm)

2.0-2.8 X 0.8-0.9

1.7-2x0.5-0.65

1.7-2.0 X 0.6-0.7

mucronulate, laterally 3-4 nerved, silvery-white, reddish toward center, medially green, 3-nerved. Stamens 3;

filaments 2.5 mm long; anthers 0.3-0.35 mm long, oblong to ellipsoid, the connective apex not prolonged.

Styles 0.8-1.0 mm long; stigmas 3, ca. 1.0 mm long. Achenes 1.7-2 mm long, 0.5-0.65 mm wide, trigonous,

oblong-ellipsoid, the apex rounded, apiculate from the dark purple style base, sessile to substipitate, the adax-

ial face convex, the abaxial flat, the surface papillose, light brown.

Additional specimen: Costa Rica: Cocos Island, in low ground near Wafer Bay, California Academy of Sciences Expedition to Galapagos

Islands, 5-7 Sep 1905, A. Stewart 267 (CAS).

This new species is endemic to Cocos Island (Isla del Coco), in the Pacific Ocean about 400 km W of Costa

Rica. It is most similar to C. panamensis (Britton) Standi, and C. lentiginosus Millsp. & Chase, species of Mexico

and Central America, and northern South America (Table 1). It differs from both in its smaller anthers, and

very wide spacing of spikelets on the rachilla; additionally the scales of C. stewartii are strongly spreading. In

general appearance, it is also suggestive of C. tenuis, a widespread lowland species of southern Central Ameri¬

ca, the Caribbean, and northern South America; it has been recorded from Isla del Coco (Tucker 1983); how¬

ever, C. tenuis has longer anthers, and much more densely arranged spikelets, typically 25 per cm of rachis. The

holotype was identified in 2002 as “ Cyperus sphacelatus” However, that pantropical species has annual habit,

larger achenes, deciduous floral scales with a large purple spot on each side, and a flexuous, persistent rachilla.

The narrow range and few collections of Cyperus stewartii indicate need for conservation. The type collec¬

tion was made in September. The three available sheets include both flowering and fruiting individuals.

Etymology .—This new species is named for Alban N. Stewart (1875-1940), botanist of the 1905-1906

CAS expedition to the Galapagos Islands, which also visited Cocos Island.

Cocos Island is located in the eastern Pacific Ocean at 5.526°N 87.066°W. The flora consists of 235 vascu¬

lar plants, of which about 30% are endemic (Trusty et al. 2006). Thus far, only one other endemic of the Cypera-

ceae has been described from Isla del Coco, Kyllinga nudiceps Standi. [ Cyperus nudiceps (C.B. Clarke ex Standi.)

O’Neill] (Tucker 1984). Trusty and colleagues visited Isla del Coco several times. They reported five species of

Cyperus: C. aggregatus, C. hermaphroditus, C. ligularis, C. odoratus, and C. tenuis. I have borrowed and examined

specimens deposited at FTG. All were correctly identified with the exception of C. hermaphroditus, which proved

to be C. tenuis as well. Another collection of C. hermaphroditus (Snodgrass & Heller 946, GH, cited by Trusty et al.

2006), proved to be C. tenuis as well. Other than the specimens collected by Stewart in 1905, this newly described

species seems to have never been collected again.

ACKNOWLEDGMENTS

The work was completed during sabbatical leave. I am grateful to curators (Debra Trock, Bruce Bartholomew)

and library staff at CAS (Rebecca Morrin, Christina Fidler) for locating additional specimens and for providing

copies of Alban Stewart’s held notes from his visit to Cocos Island, staff at Harvard University Herbaria (An-

Tucker, Cyperus stewartii, a new species from Costa Rica

thony Brach, Kanchi Gandhi, and Emily Wood) for searching for specimens, the Missouri Botanical Garden

Library for access to literature, and to Barry Hammel for information on locations of Cyperaceae specimens

from Isla del Coco. I appreciate reviews of an earlier draft by Barry Hammel and Barney Lipscomb. Thanks are

extended to curators at the herbaria from which specimens were borrowed or examined for my ongoing studies

of Neotropical Cyperus: AAU, AC, ARIZ, ASU, B, BA, BD, BKL, BH, BM, BR, C, CAS, CGE, CHAPA, CLEMS,

CONN, CORD, CU, DAO, DAV, DS, DUKE, E, ECON, ENCB, EIU, F, FLAS, FTG, G, GH, ILL, ILLS, IND, JE, K,

LCU, LL, M, MASS, MO, MSC, MT, MTMG, NHA, NY, NYS, P, PENN, PH, PMA, POM, PR, PRC, RSA, S, SD,

SIU, SMU, SP, TCD, TEX, TRT, UC, UCR, UEC, US, UTEP, VT, WIS, WRSL, WVA, YU, and Z.

REFERENCES

Kokenthal, G. 1935-36. Cyperaceae-Scirpoideae-Cypereae. In: A. Engler, ed. Das Pflanzenreich IV. 20 (Heft 101 ):1 —671.

Trusty, J.L., H.C. Kesler, & G. Haug-Delgado. 2006. Vascular flora of Isla del Coco, Costa Rica. Proc. Calif. Acad. Sci. (4th Ser.)

57:247-355.

Tucker, G.C. 1983. The taxonomy of Cyperus (Cyperaceae) in Costa Rica and Panama. Syst. Bot. Monogr. 2:1-85.

Tucker, G.C. 1994. Revision of the Mexican species of Cyperus L (Cyperaceae). Syst. Bot. Monogr. 43:1-213.

Tucker, G.C. 2001. Cyperus (Cyperaceae). In: W.D. Stevens, C.U. Ulloa, A. Pool, O.M. Montiel, AT. Arbalaez, D.M. Cutaia, and

V.C. Hollowell, eds. Flora de Nicaragua. Missouri Botanical Garden, St. Louis, Missouri, USA. 1:740-757.

Tucker, G.C. 2007. Systematics of Cyperus L. section Diffusi Kunth (Cyperaceae) in the Neotropics. In: L.M. Barbosa and

N.A. Dos Santos, Jr., eds. A Botanica no Brasil: pesquisa, ensino e politicas publicas ambientais. Sociedade Botanica

do Brasil, Sao Paulo, Brasil. Pp. 311-314.

Journal of the Botanical Research Institute of Texas 8(1)

BOOK REVIEW

Michael W. Beug, Alan E. Bessette, and Arleen R. Bessette. 2014. Ascomycete Fungi of North America: A

Mushroom Reference Guide. The Corrie Herring Hooks Series, Number 69. (ISBN-13: 978-0-292-

75452-2, cloth, alk. 128 gsm matte paper, paper permanence: ANSI/NISO Z39-48-1992 (R1997)). Uni¬

versity of Texas Press, PO Box 7819, Austin, Texas 78713-7819, U.S.A. (Orders: www.utexaspress.com,

1-800-252-3206). $85.00, 502 pp., 843 color photos, glossary, references, index, 3.9 lbs, 7" x 10" x 1.5".

(continued from p. 16 )

fungi in this group merit special recognition, for example, Claviceps purpurea, that produces purple sclerotia in

rye grass florets (ergot). When sclerotia containing toxic alkaloids are ingested by animals and humans, a hor¬

rible gangrenous condition often results in loss of limbs (ergotism). Furthermore, rye flour contaminated with

ergot used in historical bread-making often produced symptoms referred to as St. Anthony’s Fire with associ¬

ated erratic behaviors that led to the infamous Salem, Massachusetts, witchcraft trials (victims found guilty

were hanged). Another group included here are mostly insect parasites that have important medicinal proper¬

ties, represented by Cordyceps militaris; still another hypocrealean example ( Nectriopsis violacea) occurs as a

teleomorph (sexual stage) parasitizing the fruiting body (aethalium) of Myxomycetes, especially Fuligo septica.

There are many more examples included that exemplify this group of Ascomycetes.

Chapter 6, the Feotiomycetes (pp. 353-414), includes the diverse Helotiales represented by wood rot

fungi, root symbionts, endophytes, terrestrial and aquatic saprotrophs, plant pathogens, mycorrhizae, nema¬

tode-trapping fungi, and fungal parasites. Some species are strikingly colorful and fairly common; Chlorocibo-

ria aeruginascens ssp. aeruginascens has distinct bluish-green cups (apothecia) and mycelium that stains bare

decaying wood a bright greenish color.

Chapter 7, the Eurotiomycetes, include mostly microscopic molds, with perhaps Penicillium species one

of the best-known examples because it is the source of the antibiotic penicillin. There is only one taxon de¬

scribed and illustrated, Onygena corvina, that grows on keratinous animal remains such as cow and sheep

horns.

Chapter 8, the Geoglossaceae or earth tongues, has only seven species included, and members of Geoglos-

sum and Microglossum are prominent examples. The Neolectomycetes, Chapter 9, and the Orbiliomycetes,

Chapter 10, have the fewest taxa, one each, and Dothideomycetes, Chapter 11, has two taxa. Chapter 12, the

Taphrinomycotina, contains the dimorphic (yeast and filamentous) genus Taphrina, the causal agent of peach

leaf curl disease

Who should buy this book? Every affiliated club with the North American Mycological Society (NAMA)

should have a copy of this book for fungal forays and as a current reference available for member’s use. Indi¬

vidual NAMA members that represent more than 80 clubs in Canada, Mexico, and the United States of Amer¬

ica should have this reasonably priced book. Professional mycologists as well as amateur mycophiles should

consider this book a high priority for their personal library. College, university, and public libraries will want

to have this book available for the general public to enjoy the biodiversity of fungi. State, national parks, and

conservation agencies should have this book available in their bookstores for visitors, naturalists, and staffers

that interact with the general public. The content of this book will be of the utmost interest to outdoor profes¬

sionals at local nature centers, forest service and state park welcome centers, and for summer campers and

hikers who may wish to learn more about the wonderful world of fungi beneath their feet. The authors should

be congratulated for producing a book that will be at the top of the must-buy book list of many botanists and

mycologists! —Harold W. Keller, PhD, Research Associate, Botanical Research Institute of Texas, Fort Worth, Texas,

USA.

J.Bot. Res. Inst. Texas 8(1): 30.2014

CALATHEA GORDONII (MARANTACEAE), A NEW ENDEMIC

PANAMANIAN SPECIES

Helen Kennedy

UCR Herbarium, Department of Botany and Plant Science

University of California Riverside

Riverside, California 92521, U.S.A.

ganders@maii.ubc.ca

ABSTRACT

Calathea gordonii H. Kenn., sp. nov., endemic to Panama, is described as new for inclusion in the Flora Mesoamericana. It occurs in both

Bocas del Toro and Colon Provinces. Calathea gordonii is characterized by the single elliptic leaf per shoot; the single inflorescence per shoot,

borne either on a separate, leafless, shoot or on the leafy shoot; the densely appressed tomentose bracts and peduncle; the two membranous,

medial, bracteoles and the white flowers. It differs from Calathea basiflora H. Kenn. by the elliptic vs. obovate leaf, the firm, coriaceous,

densely tomentose vs. thin, herbaceous, pilose bracts and the two medial bracteoles membranous vs. claviculate. It differs from C. verecunda

H. Kenn. by the broader leaf blades (14.3-22.4 vs. 4-8 cm) and the bracts with rounded to obtuse vs. acute to acuminate apices and from C.

rhizanthoides H. Kenn. by the generally longer leaves (23.6-41 vs. 17-30 cm), broader angle of divergence of lateral veins from the midrib

(41°-46° vs. 28°-39°) and medial bracteoles membranous vs. claviculate.

RESUMEN

Calathea gordonii H. Kenn., sp. nov., endemica de Panama, es descrita como nueva para inclusion en Flora Mesoamericana. Ella ocurre en

las Provincias de Bocas del Toro y de Colon. Calathea gordonii se caracteriza por tener solo una hoja eliptica por brote; solo una inflorescencia

por brote, en un brote aparte sin hojas o en el brote con la hoja; las bracteas y el pedunculo densamente tomentosos; las dos bracteolas me¬

diates que son membranaceas y la flor blanca. Se diferencia de C. verecunda H. Kenn. por sus laminas foliares mas anchas (14.3-22.4 vs. 4-8

cm) y las bracteas con el apice rotundo u obtuso vs. agudo o acuminado y de C. rhizanthoides H. Kenn. por sus laminas foliares generalmente

mas largas (23.6-41 vs. 17-30 cm), por el angulo de divergencia de las venas laterales del nervio medio mas ancho (41°-46° vs. 28°-39°) y las

bracteolas mediates membranaceas vs. claviculadas.

In preparation for the Flora Mesoamericana treatment, the species of Marantaceae from Panama has been a

special focus. Additional herbarium studies at the Missouri Botanical Garden and the University of Panama

plus more recent collecting in the area of the concession Minera Panama, Colon Prov., have uncovered addi¬

tional new species. Since the publication of the Woodson and Schery (1945) treatment for Flora of Panama,

listing 23 species, the total has significantly increased. By 1972, Dressier (1972:184) reported a total of 35 spe¬

cies for Panama while four years later, Kennedy (1976:312-313) reported an increase to 49 known species and

suggested a possible further increase to 60 or 70. More recently, Kennedy (2011:201) reported a total of 59 spe¬

cies whereas, currently, 68 species are recognized (a 195 per cent increase from the original Flora of Panama

treatment). Nineteen species are recognized as endemic, including the one described herein plus one as yet

undescribed taxa. Additional collecting, especially in adjacent Colombia, most probably will reduce the num¬

ber of endemics.

TAXONOMIC TREATMENT

Calathea gordonii H. Kenn., sp. nov. (Figs. 1, 2). Type: PANAMA. Colon: Distr. de Donoso, Area de concesion Minera Panama,

Coastal Road, 8.5 km, bosque secundario tardio dominado por palmas, 61 m, 8°56 , 40.94"N, 80°4T38.95"W (UTM 0533622 E,

0988753 N), 16 Nov 2013, R. Flores 3512 (holotype: PMA!; isotypes: MO!, UCR!).

Haec species a Calathea basiflora H. Kenn. foliis ellipticis (vs. obovatis), bracteis crassiusculis coraceis dense tomentosis (vs. tenuibus herba-

ceis pilosis) atque bracteolis medialibus membranaceis lanceolatis (vs. induratis claviculatis) differt.

Plants rhizomatous, perennial, herbs, 45-80 cm; cataphylls narrowly ovate, apiculate, purple, densely velvety

tomentose, the hairs 0.3-0.5 mm, innermost cataphyll 11-25 cm. Leaves all basal, 1 per shoot; leaf sheath not

J. Bot. Res. Inst. Texas 8(1): 31 - 35.2014

Journal of the Botanical Research Institute of Texas 8(1)

>:illlllllllllllllllll 99372

rierhario l nivwsidad de Panama (PMA)

Plantas de Panama

Herbario de la Universidad de Panama

Provincia de Colon

; Marantaceae 3512 RF

! Calathea sp. Qet.

Hierba de c. 1 m. Bracteas chocolates, flores blancas.

Provincia de Colon, Donoso. Area de concesion Minera

Panama, Coastal Road, 8 km+500.

Bosque secundario tardfo dominado por palmas.

Coordenadas: UTM 0533622 E, 0988753 IN.

Elevacion:61 m

Fecha de colecta: 16 noviembre-2013

Colector: Rodolfo Flores.

Fig. 1. Calathea gordoniiH. Kenn. Holotype scan provided by Mireya Correa (PMA), {Flores 3512, PMA).

Journal of the Botanical Research Institute of Texas 8(1)

auriculate, green or tinged purple, paler, nearly cream-colored where covered by cataphylls, densely minutely

appressed velvety tomentose, 6-15 cm; petiole green, densely minutely appressed velvety tomentose, (13-)

20.6-30.5 cm; pulvinus elliptic in cross-section, deep olive-green, densely minutely appressed velvety tomen¬

tose throughout, the hairs 0.1-0.3 mm, 1.5-3.6 cm, articulate, nearly 2x diameter of petiole; leaf blade elliptic,

apex obtuse with acumen, base obtuse to rounded, shortly abruptly attenuate, 23.6-41 x 14.3-22.4 cm,

(length:width ratios [1.24-] 1.65-2.1:1) lateral veins 11-19 per 3 cm, cross veinlets 20-23 per 5 mm (veins mea¬

sured at midpoint of each side of blade), vein angle from midrib 41°-46°; adaxial surface of blade glossy green,

glabrous except acumen tomentose along margins and very tip, midrib concolorous, tomentose along center,

the hairs 0.4-0.5 mm; abaxial surface pale grey-green, minutely velvety, appressed tomentose, the hairs 0.2-

0.3 mm, midrib light yellowish tan, minutely velvety, appressed tomentose. Inflorescence basal, terminal, 1

per shoot, borne on a leafless or a leafy shoot, imbricate, ovoid to ellipsoid, 3-4.7 x (1-) 2-3 cm; peduncle

green, densely matted, velvety tomentose, the hairs 0.3-0.6 mm, 5.5-13.4 cm. Bracts 4-7, spirally arranged,

firm, transverse broadly ovate to broadly ovate in upper bracts, apex rounded with acumen in basal bracts, up¬

per ones obtuse, 2-3.1 x ca. 3 cm, each bract subtending up to 7 flower pairs; abaxial surface of bracts purple to

brownish purple, appressed tomentose, more densely so toward base, the hairs 0.6-1 mm, adaxial surface,

glabrous; bicarinate prophyll membranous, elliptic, apex obtuse, glabrous, 2-2.2 x 1-1.3 cm, 0.6-0.7 cm from

carina to carina; secondary bract membranous, broadly elliptic, apex rounded, glabrous, 2-2.2 x 1.4-1.6 cm;

bracteoles 2 per flower pair, medial, membranous, narrowly obovate-elliptic, glabrous, 1.8-1.9 x 0.45-0.5 cm.

Sepals narrowly elliptic, apex slightly cupped, obtuse to rounded, white, glabrous, 23-31 x 4.5-5 mm. Corolla

white, tube glabrous, 35-43 mm; corolla lobes subequal, elliptic, obtuse, glabrous, ca. 14 mm. Staminodes 3,

white; outer staminode obovate, retuse; callose staminode totally callose, subrectangular, apex rounded with

an acumen; stamen with lateral petaloid appendage, anther cream-white, 1.5 mm; style and stigma cream-col¬

ored; ovary smooth, white, glabrous, 3-3.5 x 2 mm. Capsule unknown.

Additional specimens: PANAMA. Bocas del Toro: Hill just S of Chiriqui Grande, at end of pipeline access road 2 mi N of 2nd large bridge N

(10 mi) of Cont. divide, in forest along ridge and draws, 350-500 m, 8°54'N, 82°10'W, 10 Mar 1986, B. Hammel, G. McPherson & L. Sanders

14745 (MO 3398424) (Fig. 2). Colon: Teck Cominco Petaquilla mining concession, forest on slope near coast, 8°58 , 23"N, 80°45 , 27"W, 9 Dec

2007, G. McPherson & H. van der Werff20036 (MO 6252501); Distr. de Donoso, Zona Minera, Coastal road 17.7 km, bosque secundario tardio,

70 m, (UTM 0533818E 0987741N), 13 Oct 2013, R. Flores & R. Vergara 3419 (PMA); Distr. de Donoso, area de concesion, Minera Panama,

Coastal road, bosque secondario tardio dominado por palmas, 43 m, 8°56 , 0.29"N, 80°41 , 20.01"W (UTM 0534171E 0987505N), 24 Oct 2013,

R. Flores 3440 (PMA).

Distribution and habitat.—Calathea gordonii is endemic to Panama. It is known from the Atlantic coastal forest

of both Bocas del Toro and Colon Provinces, from 40-500 m in wet primary or old secondary forest habitat.

DISCUSSION

Calathea gordonii belongs to Calathea sect. Breviscapus Benth. It is characterized by the single elliptic leaf per

shoot; the single inflorescence per shoot, borne either on a separate, leafless shoot or on the leafy shoot; the

densely appressed tomentose bracts and peduncle; the two membranous, medial, bracteoles and the white

flowers. It differs from C. basiflora by the elliptic vs. obovate leaf, the brm, coriaceous, densely tomentose vs.

thin, herbaceous, pilose bracts and the membranous, lanceolate vs. indurate, claviculate medial bracteoles. In

C. verecunda and C. rhizanthoides inflorescences are also borne on separate leafless shoots. However, C. gordonii

differs from C. verecunda by the broader leaf blades (14.3-22.4 vs. 4-8 cm) and the bracts with rounded to ob¬

tuse vs. acute to acuminate apices and from C. rhizanthoides by the broader angle of divergence of the lateral

veins from the midrib (41°-46° vs. 28°-39°) and medial bracteoles membranous, lanceolate vs. indurate, cla¬

viculate. Calathea cleistantha Standi, also has inflorescences borne on separate shoots but has more numerous

leaves per shoot, 2-4(-9) vs. 1 and corolla lobes dark purple vs. white.

Etymology. —The specibc epithet, gordonii, is in honor of Gordon McPherson, Curator, Missouri Botanical

Garden and collector of this new species, for his many excellent Panamanian collections—including several

other previously undescribed Marantaceae—nearly a hundred of which have been the types of new species in

various families in that diverse country.

Kennedy, Calathea gordonii, a new endemic Panamanian species

ACKNOWLEDGMENTS

The Missouri Botanical Garden provided support for my accommodations while working in the MO herbari¬

um (organized, thanks to Olga Martha Montiel). Fred Ganders provided the travel expenses for the trips to MO

and UCR. I am very grateful to Rodolfo Flores for his color photos of this species in the held and for collecting

additional material, including the types. I also thank Gordon McPherson and Barry Hammel for their collec¬

tions of this species which first brought it to my attention. Thanks to Gerrit Davidse, James Solomon and Teri

Bilsborrow for their help in the MO herbarium and Mireya Correa for her help at PMA and the type scan. I

thank Frank Almeda, Barney Lipscomb, and Roy Gereau for helpful suggestions and corrections. Roy Gereau

provided the Latin diagnosis. I am grateful to Andrew Sanders for providing facilities at the UCR herbarium

and to Teresa Salvado for providing accommodations and transportation in Riverside. Missouri Botanical Gar¬

den provided the paratype specimen scan.

REFERENCES

Dressler, R.L. 1972. Terrestrial plants of Panama. Bull. Biol. Soc. Wash. 2:179-186.

Kennedy, H. 1976. Notes on Central American Marantaceae II. New species from Panama and Costa Rica. Bot. Not.

128:312-322.

Kennedy, H. 2011. Three newdistichous-bracted species of Calathea (Marantaceae) from Panama. Novon 21 (2):201 —211.

Woodson, R.E. Jr. & R.W. Schery. 1945. Marantaceae. In: Flora of Panama. Ann. Missouri Bot. Gard. 32:81-105.

Journal of the Botanical Research Institute of Texas 8(1)

BOOK REVIEW

Serge Payette, ed. 2013. Flore Nordique du Quebec et du Labrador. Vol. 1. (ISBN-13: 978-2-7637-2079-1,

hbk; 978-2-7633-7208-7, pdf). Presses de LUniversite Laval, 2180, chemin Sainte-Foy, l er etage, Quebec

(Quebec) G1V 0A6, CANADA. (Orders: www.pulaval.com, 1-800-363-2864). $89.95 CND (~$83.00

USD), 561 pp., 8W' x lOW.

In 1935 Frere Marie-Victorln’s Flore Laurentienne became the standard work for the flora of the province of

Quebec, and now in its third edition, it is still highly regarded and most useful for a large section of the area. As

the title indicates, Flore Nordique covers the more northern areas, from the 54° parallel to the 63° parallel, that

is, from just above James Bay to Hudson Bay, north to the Hudson Strait, and across the provinces to the Labra¬

dor Sea on the east, including all the adjacent islands.

The editor, Serge Payette, director of the Herbarium Louis-Marie at the Universite Laval, has written very

informative, comprehensive, and interesting accounts of the exploration of these territories from the earliest

expeditions to settlements such as those of Moravian settlers in Labrador and on to the very important work of

twelve botanists in the twentieth century. All in their own way contributed to the work, largely through speci¬

mens deposited at the Laval University herbarium and elsewhere. Mr. Payette’s explanations of the geology and

the geography of the area and its five vegetative zones are essential to the understanding of the area covered in

this book. He defines these zones as closed coniferous forest (balsam firs, pines, and a few deciduous trees);

open coniferous forest in drier areas with 10% of regional forest flora; the smaller forested tundra with black

pine, tamarack, other shrubs, and lichens; the shrubby sub-Arctic tundra; and, northernmost, the Arctic tun¬

dra with its mixture of herbaceous plants and scattered shrubs, willows, and Ericaceae. Shaded maps outline

each of these regions. Only one species is strictly endemic: Elatine ojibwayensis Garneau (Elatinaceae), which

does not however fall within the scope of this volume. Other regional endemics are found across the larger

Canadian Shield. The editor considers the relative paucity of plants in the region and the rarity of some as he

discusses the existing populations of species.

Following a key to all of the plant families represented in the entire Flore Nordique, Mr. Payette and six

other botanists present the 32 families covered in this volume. The coverage includes the history of the taxon,

the vernacular names in French and English, a detailed description of the species, and color photographs of its

particular organs or growth patterns, of herbarium specimens, and often of the living plant in habitat. Ap¬

pended to the discussion of the family is a section of physical maps showing range and distribution of each

species and pinpointing where specimens were taken. A bibliography of the literature consulted accompanies

the text.

Coverage includes many pteridophytes, the gymnosperms, and some monocots such as Orchidaceae and

Iridaceae. There is no indication of which families will be covered in each of the subsequent three volumes nor

projected dates of publication.

The appended material is also very valuable. Thirty-three plates offer an entire lexicon of botany, with

precise illustrations and the French term for each illustration. For example, there are on four pages (two plates)

forty-four illustrations with terms for textures used in botanical literature, from “pubescent” to “alveolate.” A

glossary in French, each entry with an English equivalent, defines botanical terms used throughout the book.

There is also an English-French vocabulary of botanical terms that readers may find useful. Indices include

one by scientific name and one by common names, both French and English. Color charts, one bound in and

one removable, are helpful in that they indicate the range of color terms used for the plants.

Flore Nordique is an important contribution to the botany of North America. The editor and co-contribu¬

tors along with the sponsors should be complimented and congratulated on the achievement of this work.

While it will be of primary use to those studying the local flora, the book will also be of interest to those study¬

ing the ecology of plant growth in severe climatic and geographic conditions around the world .—Joann Karges,

Fexas Christian University (retired), Fort Worth, Fexas, USA.

J. Bot. Res. Inst. Texas 8(1): 36.2014

CALATHEA COFANIORUM AND C. SHISHICOENSIS, NEW ENDEMIC SPECIES

OF MARANTACEAE FROM ECUADOR

Helen Kennedy

UCR Herbarium, Department of Botany and Plant Science

University of California Riverside

Riverside, California 92521, U.S.A.

ganders@mail.ubc.ca

ABSTRACT

Calathea cofaniorum H. Kenn. and C. shishicoensis H. Kenn., both endemic to Prov. Sucumbios, Ecuador, are described as new species.

They are most similar in aspect to the Ecuadorian species C. neillii H. Kenn. & C.fredii H. Kenn. with several basal leaves and a single cauline

leaf or bladeless sheath subtending an inflorescence of bright rose-pink bracts. Calathea cofaniorum differs from C. neillii in the usually 3 vs.

7 minor veins between the major veins; the narrower angle of divergence of lateral veins from the midrib, 19°-31° vs. 40°-50°; and the red-

purple vs. yellow petals and staminodes. It differs from C.fredii in the smooth vs. strongly corrugated leaf surface; the narrower, 2.1-4.8 vs.

5.2-10 cm, and abaxially glabrous vs. pilose leaf blades. Calathea shishicoensis differs from C. neillii, C.fredii and C. cofaniorum by its inflo¬

rescence of spirally arranged bracts and broader leaves (length:width ratios of 1.33-1.75 vs. 4.1-7.45:1).

RESUMEN

Se describe Calathea cofaniorum H. Kenn. and C. shishicoensis H. Kenn., ambas endemicas a la Provincia de Sucumbios, Ecuador, como

especies nuevas para la ciencia. Las dos especies se semejan a C. neillii H. Kenn. y C.fredii H. Kenn. de Ecuador con las cuales tienen los car-

acteres de hojas basales y solo una hoja caulina o una vaina sin lamina foliar que subtiende una inflorescenia con bracteas rosadas brillantes.

Calathea cofaniorum se distingue de C. neillii por sus hojas con 3 vs. 7 venas minores entre las venas majores; por el angulo de divergencia de

las venas laterales del nervio medio mas angosto (19°-31° vs. 40°-50°); los petalos y los staminodios rojo-purpureos vs. amarillos. Se dis¬

tingue de C.fredii por la lamina foliar plana vs. fuertemente plegada; mas angosta, 2.1-4.8 vs. 5.2-10 cm, y finalmente, glabra vs. pilosa en la

haz. Calathea shishicoensis se sobresale de C. neillii, C.fredii y C. cofaniorum por sus inflorescencias con bracteas espiraladas vs. disticas y las

hojas mas anchas (la relacion largo/ancho es 1.33-1.75:1 vs. 4.1-7.45:1).

Since the publication of the treatment of Marantaceae for the Flora of Ecuador (Kennedy et al. 1988) there has

been a substantial increase in held work there. Consequently, a number of new species have been collected in

the last decade, including the two being described herein. Of the total of 96 species in the 1988 flora publica¬

tion, 64 were in the genus Calathea. Of these, 32 species were noted as endemic. As the two species described

herein are known only from the types, in Ecuador, they are considered to be endemic. However, since they are

somewhat near the border with Colombia it is possible they could occur there as well but are so far not docu¬

mented. Currently a total of 69 species of Calathea are recognized for Ecuador including the two new species

being described. As with the related C. neillii H. Kenn. and C.fredii H. Kenn., these new species, with their at¬

tractive bright rose-pink bracts, would be excellent garden subjects, though probably not for the lowlands. The

two species described herein seem most likely to be related to either the “C. lanicalis Group” (Kennedy et al.

1988), which includes the red-bracted C. timothei H. Kenn., or C. section Calathea, which has distichous-

bracted species.

TAXONOMIC TREATMENT

Calathea cofanioruin H. Kenn., sp. nov. (Fig. 1). Type: ECUADOR. Sucumbios: foothills of the Andes near the Colombian bor¬

der, access from Bermejo oil field road to Pozo 2, NW between Lumbaqui and Cascales, Rio Bermejo to Cerro Sur Pax, Cofan com¬

munity of Alto Bermejo, steep slopes N of Vista Camp, upper hill forest transition to mountain ridge, steep slopes and rock cliffs,

1300-1600 m, 00°18T3.8"N, 77°24 , 32"W, 29Jul 2001, R. Aguinda, N. Pitman &R. Foster 1415 (holotype: QCNE; isotypes: F 2231893,

UCR).

J. Bot. Res. Inst. Texas 8(1): 37 - 42.2014

Journal of the Botanical Research Institute of Texas 8(1)

Calathea cofaniorum a C. neillii foliis venis minoribus plerumque 3 (vs. 7) inter majores interpositis, lamina foliari adaxialiter glabra (vs.

minute tomentosa secus venas majores), venarum angulo majore (59-71° vs. 40-50°) atque lobis corollinis necnon staminodiis rubropur-

pureis (vs. luteis), a C.fredii foliis laevibus (vs. valde corrugatis) abaxialiter glabris (vs. pilosis) differt.

Plants rhizomatous, caulescent, herbs, 30-55 cm; stem green, glabrous except sparsely minutely tomentose

adjacent to leaf sheath; cataphylls narrowly ovate, apiculate, innermost cataphyll 5.5-13 cm. Leaves 4-8 basal,

none or 1 cauline, a cauline leaf or bladeless sheath borne atop a 20.5-30.5 cm stem internode; leaf sheath not

auriculate, green, wings glabrous along margin, sparsely minutely hispid abaxially along center and adjacent

portion of wing, leaf sheaths 7.5-15.5 cm, bladeless sheath ca. 3 cm, very base of leaf sheath sericeus, the hairs

pale straw-colored, 1 mm; petiole green, minutely subhispid apically, subglabrous to glabrous basally, the hairs

0.2-0.5 mm, 1.2-16.5 cm; pulvinus round in cross-section, olive-green, appressed tomentose in narrowband

adaxially, the rest glabrous, the hairs 0.5-0.7 mm, articulate, noticeably thicker than petiole, 0.5-0.7 cm; leaf

blade narrowly elliptic, apex acuminate-attenuate, base unequal, obtuse; 12.2-24 x 2.1-4.8 cm (length:width

ratios [4.1-] 4.58-6.57:1), generally 3 minor veins between major veins, ratio of width of narrower side of leaf

to wider 1: 1.30-1.41, vein angle from midrib (measured on inner 1/3 to 1/2 of blade) 19°-31°, vein spacing

0.8-1.7 mm between veins, veinlets 31-40 per 5 mm (measured at midpoint of each side of the blade), adaxial

surface green, glabrous, midrib tomentose, hairs along center of midrib, the hairs tan, 0.7-1 mm, subglabrous

to glabrous in apicalmost 0.5-1 cm; abaxial leaf surface grayish green, glabrous, midrib olive-green, sparsely

tomentose to subglabrous along sides, center portion glabrous, the hairs tan, 0.3-0.5 mm. Inflorescence ter¬

minal, 1 per shoot, subtended by a cauline leaf or bladeless sheath, imbricate, subrectangular, laterally com¬

pressed, 3-5.3 x 2-2.3 cm; peduncle olive-green, apical 1 cm tomentose, subglabrous to glabrous basally, 1.9-9

cm. Bracts 6-12, distichous, transverse elliptic, apex retuse, apical margin straight, not recurved, 1.2-1.7 x

2-2.2 cm, each bract subtending 3 or more flower pairs, abaxial surface of bracts rose-red, glabrous, drying

with veins prominent, adaxial surface rose-red, glabrous; bicarinate prophyll membranous, ovate, apex obtuse,

translucent, glabrous, 1.3-1.4 x 0.6-0.8 cm, ca. 0.5 cm wide, carina to carina; secondary bract membrana¬

ceous, ovate-elliptic, apex obtuse, glabrous, ca. 1 x 0.5 cm; bracteole 1 per flower pair, membraneous, narrowly

elliptic, ca. 0.9 x 0.2-0.3 cm. Flowers opening spontaneously. Sepals membranous, narrowly elliptic to nar¬

rowly obovate, obtuse to 90°, glabrous, 12-13 x 2-2.5 mm. Corolla tube light red-purple, glabrous, 12-14 mm;

corolla lobes subequal, elliptic, apex ca. 90°, red-purple, glabrous except for few minute colorless hairs of ca. 0.1

mm at very tip, 5-5.5 x ca. 2 mm. Staminodes 3, red-purple; callose staminode totally callose, ca. 4 mm; cucul-

late staminode, 3-3.5 mm; anther ca. 2 mm, data unavailable on outer staminode due to condition of specimen.

Ovary glabrous, 1.5-2 mm. Capsule obovoid, apical rim irregular, apex concave, glabrous, crowned by a per¬

sistent calyx.

Distribution and habitat.—Calathea cofaniorum is endemic to Ecuador, known only from the type locality

in the Province of Sucumblos, in the foothills of the Andes near the Colombian border, near the Cofan com¬

munity of Alto Bermejo. It occurs at 1300-1600 m elevation in the upper hill forest transition to monutain

ridge on steep slopes and rock cliffs. The type was collected in flower in July.

Discussion.—Calathea cofaniorum, shares the habit of several basal leaves with an inflorescence of disti¬

chous bright rose-pink bracts borne above an elongate stem internode with both C.fredii and C. neillii. Calathea

cofaniorum differs from C. neillii in the usually 3 vs. 7 minor veins between the major veins, the leaf blade ad¬

axially glabrous vs. minutely tomentose along major veins, the steeper vein angle, 19°-31° vs. 40°-50° diver¬

gence from midrib, and the red-purple vs. yellow corolla lobes and staminodes. It differs from C.fredii in the

smooth vs. strongly corrugated leaf surface; the narrower, 2.1-4.8 vs. 5.2-10 cm, and abaxially glabrous vs.

pilose leaf blades and shorter, 12-13 vs. 15-16 mm, sepals. It would key out in Flora of Ecuador (Kennedy

1988:47) under lead 30A because of the distichous bracts.

Etymology. —The specibc epithet, cofaniorum, is in recognition of the Cofan community on whose land

the plant was collected.

Kennedy, Two new endemic species of Calathea from Ecuador

ECUADOR

Prov. de Sucumbfos

MARANTACEAE

Calathea

Rio Bermejo to Cerro Sur Pax: Cofan community of Alto

Bermejo. Access from Bermejo oil field road to Pozo 2, NW

between Lumbaqui and Cascales. Steep slopes N of Vista Camp.

N? 2231893

FIELD MUSEUM

NATURAL HISTORY

SA- EASTERN LOWLANDS

Wet COL/ ECU A/ N.E.PERU

5 cm

00° 18’13.8N, 77°24’32.0W 1300-1600 m

Foothills of the Andes near the Colombian border. Upper hill

forest transition to Mountain Ridge. Steep slopes & rock cliffs.

how-herb to 30 cm. Bracts red-pink, flowers purple.

Roberto Aguinda 1415 29 Julio 2001

Nigel Pitman, Robin Foster

FUNDACION SOBREVIVENCIA COFAN

Fig. 1. Calathea cofaniorum H. Kenn. Isotype {Aguinda, Pitmann & Foster 1415 F). Scan provided by Field Museum.

Journal of the Botanical Research Institute of Texas 8(1)

Sinangoe Station: Shishicho Ridge, Alto Aguarico drainage,

ill Above (south of) Rio Cofanes, west of Puerto Libre, NW of

Lumbaqui. Access from Rio Sieguyo. Ridgeline trail above camp.

N? 2231896

FIELD MUSEUM

NATURAL HISTORY

^ SA- EASTERN LOWLANDS

Wet COL/ ECU A/ N.E.PERU

5 cm

00°12’01.3N, 77°31’54.3W 140Q-150P m

Foothills of the Andes Short, 10-2Qm tall upper hill-forest on

steep ridgeslopes on acid soils.

L ' ^ ' r , ' l

Roberto Aguinda 1173 13 Agosto 2001

Nigel Pitman, Robin Foster ,

FUNDACION SOBREVIVENCIA COFAN

Fig. 2. Calatheashishicoensis H. Kenn. Isotype {Aguinda, Pitmann & Foster 1173 F). Scan provided by Field Museum.

Kennedy, Two new endemic species of Calathea from Ecuador

Calathea shishicoensis H. Kenn., sp. nov. (Fig. 2). Type: ECUADOR. Sucumbios: foothills of the Andes, access from Rio Sie-

guyo, NW of Lumbaqui, W of Puerto Libre, above (S of) Rio Cofanes, Alto Aguarico drainage, Shishico Ridge, Sinangoe Station,

ridgeline trail above camp, short, 10-20 m tall, upper hill-forest on steep ridge slopes on acid soils, 1400-1500 m, 00°18T3.8"N,

77°24 , 32"W, 13 Aug 2001, R. Aguinda, N. Pitman &R. Foster 1173 (holotype: QCNE; isotypes F 2231896, UCR).

Calathea shishicoensis a C. neillii, C. cofaniorum et C.fredii bracteis spiraliter (vs. distiche) dispositis atque foliis ellipticis (vs. anguste ovato-

ellipticis vel anguste ellipticis) longitudinis cum latitudine proportione 1.33-1.75 (vs. 4.10-7.45): 1 differt.

Plants rhizomatous, caulescent, herbs, 50-70 cm; stem green, minutely sparsely tomentose in apical 3 cm (just

below cataphyll subtending the inflorescence), glabrous basally; cataphylls narrowly ovate, apiculate, sparsely

pilose in upper half, glabrous basally, innermost cataphyll 11-16.5 cm. Leaves 2-3 basal, a bladeless sheath

borne atop a ca. 51.5-52.5 cm stem internode; leaf sheaths not auriculate, green, glabrous, 9.4-14 cm; petiole

green, glabrous, 30-34.5 cm; pulvinus round in cross-section, deep olive-green with a purplish band at junc¬

tion to petiole or tinged purple throughout, appressed tomentose in narrow adaxial band, the rest glabrous,

hairs 0.5 mm, 3.1-4.2 cm; leaf blade coriaceous, elliptic, apex obtuse with acumen, base rounded to subtrun¬

cate, shortly abruptly attenuate, 17-21.6 x 9.8-13.1 cm, (length:width ratios 1.33-1.75:1), generally there are 7

minor veins between major veins, vein angle divergence from midrib (measured at midpoint of blade) 36°-48°,

20-30 veins per 3 cm, veinlets ca. 25 per 3 mm (measured at midpoint of each side of the blade); adaxial leaf

surface deep matte green except shiny along margin, major veins darker green, glabrous except sparsely mi¬

nutely (14x magnibcation) tomentose along major veins and on acumen, the hairs colorless, 0.2-0.3 mm,

midrib yellow-green, densely tomentose throughout midrib and just onto adjacent portion of blade, the hairs

colorless 0.3-0.5 mm; abaxial leaf surface semi-shiny gray-green, glabrous except sparsely minutely tomen¬

tose along major veins, glabrous toward margin and along margin in apical 0.5 cm, midrib green, appressed

tomentose but only on sides in basal portion, the hairs colorless 0.5 mm. Inflorescence terminal, 1 per shoot,

subtended by a bladeless sheath 2.5-3 cm, imbricate, fusiform, 4.8-6.5 x 2-2.2 cm; peduncle green, pink right

at junction to lowest bract, sericeous, the hairs colorless 0.25-0.3 mm, ca. 1-1.1 cm. Bracts 11-16, spirally ar¬

ranged, herbaceous, elliptic, apex rounded, dying back and splitting into 2 or 3 irregular segments, outer

margin and apex thin, membranous, straight, 2-2.2 x 1.2-1.4 cm, each bract subtending 5 or more flower pairs,

abaxial surface of bracts deep rose-pink, minutely pilose toward margin and very tip with tuft of hairs, the rest

glabrous, the hairs 0.2-0.3 mm, hairs more evident in lower bracts; bicarinate prophyll membraneous, ovate-

elliptic, apex obtuse to rounded, margin deeply pigmented, probably colored when live, the rest translucent,

sparsely pilose on carina, the rest glabrous, the hairs colorless 0.3-0.5 mm, 1.6-1.7 x ca. 0.8 cm, 0.5-0.6 cm

wide, carina to carina; secondary bracts absent; bracteoles 2 per flower pair, membraneous, medial, both cari¬

nate, narrowly oblong-elliptic, translucent with very margin deeply pigmented (dark red-brown when dried),

glabrous, 1.3-1.5 x 0.2-0.3 cm. Flowers opening spontaneously, pale pink fide label, Aguinda et al. 1173 (F).

Sepals narrowly oblong-elliptic, obtuse to 90°, margins inrolled appearing acute, 14-16 x 2.5-3 mm; corolla

tube glabrous, 15-17 mm; corolla lobes subequal, elliptic, apex acute, darker, purplish pink at apex, pale below,

glabrous except tuft of colorless hairs at apex, the hairs ca. 0.1 mm, 7-8 x 1.5-2 mm. Staminodes 3; outer sta-

minode ca. 4.5 mm; callose staminode totally callose, 6-6.5 mm; cucullate staminode 3.5-4 mm; anther ca. 1.5

mm. Ovary essentially smooth with a thickened, raised rim, apically, the calyx attached in a slight depression.

Capsule and seeds unknown.

Distribution and habitat.—Calathea shishicoensis is endemic to Ecuador, known only from the type local¬

ity in the in the foothills of the Andes, Sucumbios Province, on Shishico Ridge in the Alto Aguarico drainage.

It occurs from 1400-1500 m in the short, 10-20 m tall, upper hill-forest on steep ridge slopes on acid soils. The

type was collected in flower in August.

Discussion.—Calathea shishicoensis shares a similar habit and also the bright rose-pink bracts with C. neil¬

lii, C. cofaniorum and C.fredii, but differs significantly in having spirally arranged bracts and decidedly broader

leaves (length:width ratios of 1.33-1.75 vs. 4.1-7.45:1). Like C. neillii, it has 7 minor veins between the major

veins and also lacks secondary bracts. It would key out in Flora of Ecuador (Kennedy 1988:47) with C. roseo-

Journal of the Botanical Research Institute of Texas 8(1)

bracteata H. Kenn. under lead 40A because of the spirally arranged rose-pink bracts but is distinguished by the

smooth vs. strongly corrugated leaf blade and fewer, 11-16 vs. 25-50, bracts.

Etymology .—The specific epithet, shishicoensis, is in reference to the Shishico Ridge from where it was

collected.

ACKNOWLEDGMENTS

The Missouri Botanical Garden provided support for my accommodations while working in the MO herbari¬

um (organized, thanks to Olga Martha Montiel). Fred Ganders provided the travel expenses for the trips to F,

MO and UCR. I thank R. Aguinda, N. Pitman, and R. Foster for collecting both C. cofaniorum and C. shishicoen¬

sis. I thank Robin Foster for the use of his color photographs of both C. cofaniorum and C. shishicoensis, his

collections and scans of the Field Museum specimens of it. I thank Christine Niezgoda for providing accom¬

modations and transportation and for help in the F herbarium. I gratefully acknowledge the helpful sugges¬

tions and corrections by John Pipoly III, Barney Tipscomb, and Roy Gereau. I am grateful to Andrew Sanders

for providing facilities at the UCR herbarium and to Teresa Salvado for providing accommodations and trans¬

portation in Riverside. Field Museum provided the type scans.

. ,■ REFERENCES ! "

Kennedy, H., L. Andersson, & M. Hagberg 1988. Marantaceae. In: G. Harling & L Andersson, eds. Flora of Ecuador. Berlings,

Arlov, Sweden. 32:11-191.

GLOSSOLOMA VELUTINUM (GESNERIACEAE), A NEW SPECIES

FROM THE CORDILLERA CENTRAL OF THE COLOMBIAN ANDES

Larri A. Rodas

Centro de Estudios e Investigaciones

en Biodiversidad y Biotecnologfa - CIBUQ

Universidad del Quindfo

Armenia, COLOMBIA

maxwellero81 @g mail.com

John L. Clark

Department of Biological Sciences

Box 870345

University of Alabama

Tuscaloosa, Alabama 35487, U.S.A.

jlc@ua.edu

ABSTRACT

A new species of Glossoloma (Gesneriaceae) is described from the Cordillera Central of the Colombian Andes. The new species, Glossoloma

velutinum J.L. Clark & T.A. Rodas, is locally endemic to cloud forests in the provinces of Quindfo and Tolima. The new species is differenti¬

ated from other Glossoloma by the presence of an orange corolla, scandent habit with elongate shoots to 3 m tall, and uniformly velutinous

indumentum on the stems and leaves.

RESUMEN

Una nueva especie de (Gesneriaceae) es descrita en la cordillera central de los andes de Colombia. La nueva especie, Glossoloma velutinum

J.L. Clark & L.A. Rodas, es endemica de las nefosilvas del Quindfo y el Tolima. La nueva especie es diferente de otras Glossoloma por la presen-

cia de una corola de color naranja, de habito escandente llegando alcanzar longitudes de hasta 3 m. y su indumento velutino en tallos y hojas.

Key Words: Cordillera Central, Episcieae, Gesneriaceae, Glossoloma, Quindfo, taxonomy

INTRODUCTION

The genus Glossoloma Hanst. belongs to the New World subfamily Gesnerioideae and subtribe Columneinae

(Weber et al. 2013). Glossoloma is distinguished from closely related genera by the presence of resupinate (up¬

side-down) flowers. Glossoloma ranges from Mexico south to Bolivia and is most diverse in the northern Andes

of Colombia and Ecuador. The genus was recently monographed by Clark (2009) and included 27 species.

Expeditions to Colombia during the previous two years (Clark 2012) have resulted in many discoveries and the

subsequent publication of new species (Clark & Clavijo 2012; Clavijo & Clark 2012; Clavijo & Clark 2014;

Smith et al. 2013). The publication of Glossoloma velutinum increases the total number of species in the genus to

28 with additional discoveries that will be published in the near future.

TAXONOMIC TREATMENT

Glossoloma velutinum J.L. Clark & L.A. Rodas, sp. nov. (Fig. 1) type: Colombia. Quindio: Municipio de Calarca, car-

retera al Campanario, borde de bosque conservado, 4°28'27N, 75°33'25W, 3160-3450 m, 29 Aug 1993, D. Macias , M.L. Chacon &J.C.

Hincapie 101 (holotype: HUQ).

Differs from all other Glossoloma by the presence of a velutinous indumentum, scandent habit, and elongate shoots to 3 m tall.

Terrestrial or epiphytic scandent subshrub; stems erect, rarely branched, to 3 m tall, to 0.7 cm in diameter,

quadrangular, vestiture velutinous to densely villous, woody when mature, succulent to herbaceous when

young, internodes 2.5-8 cm long. Leaves opposite, equal or subequal in a pair; petioles 1.5-2.5 cm long,

sparsely to densely woolly; blades 4-17 x 2.5-7 cm, elliptic-oblong to obovate, base cuneate to acute, occasion¬

ally oblique, apex acuminate, margin denticulate to serrate, adaxially green and velutinous, abaxially much

lighter green and velutinous, coriaceous when dry, lateral veins 6-8 per side. Flowers resupinate, appearing

fasciculate with 2-5 flowers per axil, posture pendent at anthesis, bracteoles 0.5-2.0 x 3-6 mm, ovate; pedicels

1-3 cm long, usually velutinous, rarely villous; calyx lobes nearly free, conduplicate with each lobe appressed

to adjacent lobe and folded lengthwise with the margin curved inward, erect, 4 subequal, 1-3 x 0.6-1.0 cm,

broadly ovate, base truncate, apex acuminate, margin serrate to laciniate, red to pink, abaxially densely veluti¬

nous, adaxially sparsely velutinous; corolla 2.0-3.0 cm long, tubular, gibbous basally on upper surface, long

J. Bot. Res. Inst. Texas 8(1): 43 - 45.2014

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 1. Glossoloma velutinumJ.L. Clark&L.A. Rodas. A. Lateral view of resupinate flower. B. Front view of flower showing androecium on lower surface

and medial lobe on upper surface. C. Habit. (A-C photographic images of LA Rodas 102).

Rodas and Clark, A new species of Glossoloma from Colombia

axis of corolla oblique relative to calyx; base ca. 8 mm in diameter, middle ampliate, becoming apically ventri-

cose on upper surface, throat slightly constricted, not appearing laterally compressed, ca. 1.5 cm wide (at

mouth), outside densely velutinous, inside sparsely velutinous, interior red spotting present, limb spreading,

subregular, ca. 8 mm wide, orange becoming yellow, lobes equal, ca. 2 x 6 mm, rotund, spreading, entire; nec¬

tary a bilobed gland on ventral surface of ovary, sparsely glabrous; filaments curved after anthesis, free por¬

tion to 1.5 cm long, adnate to base of corolla tube for ca. 3 mm, connate for 2.5 mm, forming an open sheath,

glabrous; anthers 1.5 x 3 mm, dehiscing by longitudinal slits; staminode present; ovary ca. 4.5 x 3 mm, ovoid,

densely velutinous, style 0.8-1.5 cm long, glabrous, stigma stomatomorphic. Fruit a fleshy capsule, pendent

when ripe, 1.4 x 1.1 cm, globose to ovoid when immature, velutinous, loculicidally dehiscent and bivalved

when mature, valves not reflexed, reaching a 45°-60° angle when mature. Seeds numerous, ca. 1.0 x 0.5 mm,

elongate, longitudinally- transversely striate, brown.

Additional specimens studied. COLOMBIA. Quindlo: municipio de Salento, Reserva Natural Alto Quindio, Acaime, 4°37'85"N, 75 o 28'0"W,

2780 m, 12 May 1991, CA. Agudelo, L.F. Hoyos, D. Macias &A.L. Lopez. 1736 (HUQ); municipio de Calarca, vereda planadas, microcuenca

de la quebrada la Sonadora, finca la Merced, 75°37'N, 04°26'W, 3200-3500 m, 20 May 2007, L.A. Rodas 21 (HUQ); municipio de Salento,

Reserva Forestal Navarco, 04°38'N, 75°34'W, 2920 m, 18 Apr 1989, G. Arbeldez S., C. Velez N., N. Carvajal D. &J. Uribe M. 2898 (HUQ); mu¬

nicipio de Salento, vereda Cocora Alto, Area de Conservacion la Montana, 4°37'N, 75°28'W, 3120 m, 20 May 2007, P. Sepulveda, O. Martinez

& T. Gomez, 157 (HUQ); municipio de Salento, Reserva Natural Alto Quindio, Acaime, 4°37'N, 75°28'W, 2780 m, 12 May 1991, CA. Agudelo,

L.F. Hoyos, D. Macias &A.L. Ldpez . 1736 (HUQ); municipio de Cordoba, vereda las auras, finca el Cedral, 75°37'N, 04°26'W, 2950-3000 m,

14 Dec 1993, M.C. Velez, D. Macias & L.F. Hoyos 3750 (HUQ). Tolima: municipio de Cajamarca, vereda la luisa, camino hacia la N, 4 0 28'27"N,

75°33 , 25"W, 3000-3300 m, 5 Jun 2013, L.A. Rodas 102 (FAU, HUQ).

Glossoloma velutinum is similar to Glossoloma ichthyoderma (Hanst.) J.L. Clark because of the erect shoots and

subshrub epiphytic habit. It should be noted that Glossoloma ichthyoderma is more frequently found as a terres¬

trial subshrub in contrast to the epiphytic subshrub habit of G. velutinum. The shoots of G. ichthyoderma are

covered in peltate scales in contrast to the velutinous indumentum and absence of peltate scales in G. velutinum.

Distribution and habitat.—Glossoloma velutinum is known form the Cordillera Central of the Colombian

Andes in the departments Quindio (western slopes) and Tolima (eastern slopes) in cloud forests from 2780 to

3300 m.

Etymology. —The specific epithet reflects the velutinous indumentum that covers the stems and leaves.

Phenology. —Collected in flower during March, April, and December, in fruit during May and August.

ACKNOWLEDGMENTS

This study was supported by funds from the National Science Foundation (DEB-841958 and DEB-0949169 to

JLC). A 2012 research expedition to Colombia was a tremendous success because of logistical support from

Laura Clavijo (UNA). Juan David Tovar Duran, Juan Manuel Duque Orozco, and Diego Alejandro Ortiz Mar¬

tinez provided additional logistical support to the environs of Quindio. We gratefully acknowledge Christian

Feuillet and Laurence E. Skog for helpful reviews of the manuscript. The second author would like to thank the

Facultad de Ciencias Basicas y Tecnologlas from the Universidad del Quindio and especially Ana Lucia Lopez

Gonzalez for facilitating a visit to study herbarium specimens (HUQ).

REFERENCES

Clark, J.L. 2009. Systematics of Glossoloma (Gesneriaceae). Syst. Bot. Monogr. 89:1 -126.

Clark, J.L. 2012. Colombia - an unexplored biodiversity hotspot revisited after 25 years. Gesneriads 62:40-47.

Clark, J.L. & L. Clavijo. 2012. Columnea antennifera, a new species of Gesneriaceae from the Cordillera Central of the

Columbian Andes. J. Bot. Res. Inst.Tex. 6:385-390.

Clavijo, L. & J.L. Clark. 2012. Drymonia atropururea (Gesneriaceae), a new species from northwestern South America. J.

Bot. Res. Inst.Tex. 6:71-74.

Clavijo, L. & J.L. Clark. 2014. Drymonia crispa (Gesneriaceae), a new species from northwestern Colombia. Brittonia

66(1 ):65-69. DOI 10.1007/s 12228-013-9310-4:1. [published online July 4, 2013]

Smith, J.F., M. Amaya-Marquez, O.H. Marin-Gomez, & J.L. Clark. 2013. Four new species of Columnea (Gesneriaceae) with

primary distributions in Colombia. J. Bot. Res. Inst. Tex. 7:667-679.

Weber, A., J.L. Clark, & M. Moller. 2013. A new formal classification of Gesneriaceae. Selbyana 31:68-94.

Journal of the Botanical Research Institute of Texas 8(1)

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J. Bot. Res. Inst. Texas 8(1): 46.2014

SYNOPSIS OF GALVEZIA (PLANTAGINACEAE: ANTIRRHINEAE),

INCLUDING A NEW CRYPTIC SPECIES FROM SOUTHERN PERU

Michael O. Dillon

Victor Quipuscoa Silvestre

Botany Department

The Field Museum

1400 South Lake Shore Drive

Chicago, Illinois 60605, U.S.A.

mdillon@fieldmuseum.org

Departamento Academico de Biologia

Universidad Nacional de San Agustfn

Arequipa, PERU

ABSTRACT

A generic synopsis is provided for Galvezia Dombey ex Juss. (Plantaginaceae: Antirrhineae), a genus of shrubs with red to white, tubular

flowers endemic to coastal Ecuador and Peru and the Galapagos Islands (Ecuador). Galvezia elisensii M.O. Dillon & Quipuscoa, a new

cryptic species, is described from southern Peru, bringing the total number of species to four.

RESUMEN

Se presenta la sinopsis del genero Galvezia Dombey exjuss. (Plantaginaceae: Antirrhineae); se trata de un genero pequeno, compuesto por

arbustos con flores de rojo a bianco, tubulares, endemico del lado occidental de Ecuador y Peru y de las Islas Galapagos (Ecuador). Se de¬

scribe ademas, una nueva especie, Galvezia elisensii M.O. Dillon & Quipuscoa, procedente del Sur de Peru, haciendo un total de cuatro

especies en la actualidad.

INTRODUCTION

Galvezia Dombey exjuss. (Plantaginaceae: Antirrhineae) is a genus of shrubs with tubular flowers distributed

along the coast of Ecuador and northern to southern Peru, and one species confined to the Galapagos Islands

(Elisens 1989,1992; Wiggins & Porter 1971; Brako & Zarucchi 1993; Jorgensen & Leon-Yanez 1999). With the

addition of the new species described here, the genus is comprised of four species. The North American taxa

previously attributed to Galvezia are now placed in Gambelia Nutt., e.g., G. speciosa Nutt. (Sutton 1988; Elisens

& Nelson 1993). The phylogeny of the tribe Antirrhineae, including Galvezia and Gambelia , has been investi¬

gated with molecular techniques (Ghebrehiwet et al. 2000; Medhanie et al. 2000; Oyama & Baum 2004; Var¬

gas et al. 2004); however, a more robust sampling will be necessary before relationships can be resolved with

confidence.

During the strong El Nino year of 1983, the senior author collected a specimen of Galvezia from the Lomas

deAtiquipa in Department of Arequipa in southern Peru. A duplicate of that collection (M. Dillon & D. Dillon

3776) was forwarded to Dr. Wayne Elisens for identification and he indicated that it was potentially a new spe¬

cies, distinct from its northern congeners. He subsequently visited southern Peru and made collections

(Elisens 844,845, 847, all OKL) that were included in his study of genetic divergence in Galvezia (Elisens 1992).

While not included in his 1992 study, the sample of M. Dillon & D. Dillon 3776 was included in his analysis of

Gambelia (Elisens & Nelson 1993, p. 456) and termed “G. sp. nov .”

The overall morphological variation in the continental species is not profound, with differences typically

confined to corolla size, anther filament pubescence, and leaf shape. Attention has been drawn to differences

in habit, but these are not useful in species recognition. Authors of recent floras have adopted a broad species

concept for Galvezia by placing several species into the synonymy of G.fruticosa J.F. Gmel (Brako & Zarucchi

1993; Jorgensen & Leon-Yanez 1999). However, data supporting narrower species circumscriptions have been

convincingly presented (Elisens 1992; Elisens & Nelson 1993), even though they prove difficult to quantify

and qualify (i.e., cryptic species). Cryptic species represent a situation where speciation has already broken the

gene flow between populations, but where evolution has not progressed to a point where easily recognizable

adaptations are visually obvious (Bickford et al. 2007; Schlick-Steiner et al. 2007).

J. Bot. Res. Inst. Texas 8(1): 47 - 55.2014

Journal of the Botanical Research Institute of Texas 8(1)

In his discussion of results from allozyme divergence patterns in Galvezia, Elisens argued that data sup¬

ported the taxonomic distinctness of the Galapagos endemic, G. leucantha Wiggins, and three mainland spe¬

cies: G.fruticosa, G. grandiflora (Benth.) Wettst. (as G. ballii Munz + G. lanceolata Pennell), and a “sp. nov.” based

upon his collections from southern Peru (referenced above). He stated that the pattern of allozyme variation

suggested G.fruticosa, G. leucantha, G. sp. nov., and G. grandiflora (as G. ballii) had undergone a gradual genetic

divergence following their reproductive isolation. He stated that the recognition of G. lanceolata in central

coastal Ecuador by Pennell (1946) and Sutton (1988) was not supported by his electrophoretic and morpho¬

logical data. Rather, the data were most concordant with recognition of only one species in northern Peru and

coastal Ecuador (the name G. grandiflora has priority). Finally, Elisens suggested that, “systematic data support

delimitation of an undescribed species, G. sp. nov. In addition to unique leaf and floral characters and an allo-

patric distribution, G. sp. nov. is differentiated from other species in Galvezia by three marker alleles.”

Elisens (1992) accepted three mainland species, G.fruticosa, G. grandiflora (i.e., G. ballii + G. lanceolata)

and an additional taxon (sp. nov.) described here, based upon southern Peruvian material from the Depart¬

ments of lea and Arequipa. We accept Wettstein’s (1891) transfer of Bentham’s Galvezia limensis (Dombey ex

Chav.) Benth. var. grandiflora Benth. as Galvezia grandiflora (Benth.) Wettst. All specimens cited here have

been examined unless otherwise indicated (n.v.).

TAXONOMIC HISTORY

Pennell (1946) provided a detailed discussion of the taxonomic history of Galvezia, but a summation of the

pertinent literature is merited. Galvezia was originally proposed by Joseph Dombey, the French botanist who

accompanied Ruiz and Pavon during their explorations of Peru and Chile. Dombey left South America in 1784,

before Ruiz & Pavon, and, in clear violation of stated protocols, Dombey began publishing selected new taxa he

had encountered. The generic description for Galvezia appeared in J.F. Gmelin’s edition of Linnaeus’ Systema

Naturae (2: 937) in 1791, where Dombey proposed the genus Galvezia in honor of Don Jose de Galvez y Gal¬

lardo, marques de Sonora and minister of the Council of the Indies (b. 1720-d. 1787). A note with Dombey’s

specimen at Kew suggests that his plant was gathered in 1779 near Lima. The alternate spelling, Galvesia, has

been accepted by some databases, but is here rejected given the origin of the generic name.

Ruiz and Pavon did not accept Galvezia sensu Dombey, and believing it to be a species of Dodartia L., a

Palaearctic genus. They published new name, Dodartia fragilis in 1798 based upon the Dombey publication in

Jussieu (1789). They considered that the generic name Galvezia was open and they described another Galvezia

in Florae Peruvianae et Chilensis Prodromus, 56 in 1794 in the Rutaceae. Chavannes in his Monographic des

Antirrhinees (1833), accepted the generic concept of Galvezia, but with Ruiz and Pavon’s genus evidentially

occupied, he proposed Agassizia (p. 180) and described A. limensis for Dombey’s plant collected near Lima. He

provided an illustration (Plate XI) clearly representing G.fruticosa.

When Bentham treated the Scrophulariaceae for DeCandolle’s Prodromus in 1846, he adopted Dombey’s

name and attributed it to Jussieu. He also established a varietal name, Galvezia limensis S grandiflora, for mate¬

rial gathered further north near the port city of Paita and cited F. Hall 10 (K000528872). Galvezia was accepted

in Wettstein’s treatment of the Scrophulariaceae in Engler and Prantl’s Pflanzenfamilien (1895). Wettstein at¬

tempted to provide a new combination for Bentham’s variety; however, he appears to attribute the original au¬

thorship to “(Kell.)” and mentioned California. These errors led Munz to reject the combination by Wettstein

and provide a new name for Bentham’s variety, Galvezia ballii and citing the type locality as Paita (as Payta),

Peru. He based his superfluous name upon J. Ball s.n. (US1323500, US251553). In 1946, Pennell described G.

lanceolata from the region of Manabl, Ecuador, and provided a key to allow for discrimination of G. fruticosa,

G. lanceolata, and G. ballii.

TAXONOMY

Galvezia Dombey ex Juss., Gen. Pi. 119. 1789. Type: G. fruticosa J.F. Gmel., Syst. Nat. 2:937.1791. Non Galvezia Ruiz & Pav., Fl.

Peruv. Prodr. 56.1794.

Dillon and Quipuscoa, Synopsis of Galvezia

Much-branched shrubs to 2 m tall, erect, arching or pendent; glabrous to pubescent. Leaves opposite, or oc¬

casionally 3 at a node, simple, the blades ovate-lanceolate to elliptic, pinnate-nerved, the margins entire, the

bases cuneate to subcordate, the apices acute. Inflorescences paniculate. Flowers axillary; calyx 5-merous,

ovate to lanceolate, the apices acute; corollas red to white, tubular, bilabiate, the tube elongate, subcylindric,

the base expanded to subgibbose, the upper lip erect, bilobed, the lobes ovate, the lower lip trilobed, the lobes

plane to reflexed; stamens didynamous, 4 fertile, one aborted; anthers bilocular; ovary ovoid to spheric. Fruit

capsules, globose to subglobose, cartaceous, chambers dehiscent by 1-2 pores, seeds oblong-truncate. Chro¬

mosome number: unknown.

Galvezia Dombey ex Juss. is a genus of small shrubs with tubular flowers. Four species are recognized

with the addition of this new species from southern Peru; three conbned to western Ecuador and Peru and one

endemic species to the Galapagos Islands (Ecuador). All continental species possess deep red to crimson corol¬

las, whereas the island species have variously colored corollas (see discussion below).

Wiggins (1968) addressed the difference or loss of floral coloration in some of the island species and at¬

tributed the shift to white corollas to the absence of hummingbirds within the islands; however, it has never

been fully documented that hummingbirds were the pollination vectors in the mainland species. Information

on pollination vectors is now available and possible vectors could be members of the Sphingidae, Lepidoptera

(Cock & Boos 2006).

KEY TO GALVEZIA SPECIES

The following key will allow for identification of Galvezia species (adapted from Pennell 1946)

1. Galapagos Islands, Ecuador_ G. leucanthaWiggins

1. Mainland Ecuador and Peru.

2. Cauline leaves elliptic to elliptic-lanceolate, 15-25 mm long, 3-5 mm wide, pedicles 8-12 mm long, stout, straight to

slightly curved, corollas 8-9 mm long (southern Peru)_ G. elisensii M.O. Dillon & Quipuscoa

2. Cauline leaves ovate-lanceolate to lanceolate, 20-25 mm long, 5-15 mm wide; pedicles 8-20 mm or longer, filiform,

distally coiled or strongly incurved, corollas 12-22 mm long (central Peru to Ecuador).

3. Corolla 12-14 mm long, the lips about Vz length of tube; pedicels as long as, or longer than, the subtending leaves,

leaf-blades about 2 cm long, on petioles ca. 2 mm long; shrubby (central to northern Peru)_ G. fruticosa J.F. Gmel.

3. Corolla 14-22 mm long, the lips about Vi length of tube; pedicels shorter than the subtending leaves, leaf-blades

and bracts lanceolate; suffrutescent to herbaceous throughout (northern Peru to Ecuador)_ G. grandiflora (Benth.) Wettst.

1. Galvezia elisensii M.O. Dillon & Quipuscoa, sp. nov. (Figs. 1-3). Type: Peru. Arequipa. Caraveli: Lomas of Atiquipa,

ca. 10.5 km N of turn-off to Atiquipa, km 584 S of Lima, 150-200 m, 1 Nov 1983, M. Dillon & D. Dillon 3776 (holotype: F; isotypes:

CPUN, GH, HUT, K, MO, NY, OKL, TEX).

Galvezia fruticosa J.F. Gmel. simile, sed foliis brevioribus et angustioribus et pedicellis brevioribus.

Much branched shrubs; branches erect to arching, 0.8-1.2 m long; young stems green, maturing reddish.

Leaves opposite, the blades elliptic to elliptic-lanceolate, (10-)15-25(-30) mm long, (2-)3-5(-15) mm wide,

the base narrowly cuneate, the apex acute, glabrous, semisucculent, occasionally sparsely puberulent; petioles

(0.5-)l-2.5(-3) mm long, ca. 0.3 mm in diameter. Inflorescences paniculate. Flowers axillary; the pedicles

stout, (5-)8-12(-18) mm long, glabrous, straight to slightly curved; calyx lobes ovate to lanceolate, c. 2.5 mm

long, c. 1.5 mm wide, apices obtuse, glabrous; corollas tubular, (7-)8-9(-10) mm long, 2.5-3 mm wide, outside

puberulent, red, slightly ampliate, the limb bilabiate, superior lip 3-4.5 mm, bilobed, erect, purberulent, the

lower lip trilobate, 2.5-3.5 mm long, base of lower lip folded, glandular-puberulent; stamens didynamous,

dorsal pair 5.5-7 mm long; ventral pair, 7-9 mm long, filaments glabrous distally, glandular-papillate basally;

anthers thecae 1-1.4 mm long; fifth stamen sterile, c. 1 mm long; ovary ovate, 1.5-2 mm long, 1-1.5 mm wide,

puberulent. Fruit subglobose, glabrous, 4-6 mm long, 4.5-7 mm in diameter, deep red at maturity; seeds

(40-)140-175 per capsule, sub truncate, black, c. 1 mm long.

Etymology. —This species is dedicated to Dr. Wayne J. Elisens, Professor and Curator of the Robert Bebb

Herbarium on the campus of the University of Oklahoma, Norman. His recognition of this taxon, as reflected

in its allelic profiles, has led to its description here.

Journal of the Botanical Research Institute of Texas 8(1)

Field Museum of Natural History

DEPTO: AREQUIPA PROV: CARAVELI

SCROPHULARIACEAE

Galvezia fruticosa Gtnel.

N? 1940835

FIELD MUSEUM

NATURAL HISTORY

Lomas of Atiquipa, oa. 10.5 km N of turn-off to

Atiquipa. KM 584 S of Lima. ca. 150-200 m.

Erect, glutinous shrub to 0.75 m.; flws, red.

1 Nov 1983

M. O. Dillon & D. Dilion

3776

Plants collected under the sponsorship of the National Geographic Society [Grant No. 2706-831

Distributed by Field Museum of Natural History

Fig. 1. Galvezia elisensii. Photograph of holotype collection, M. Dillon & D. Dillon 3776 (FI 940835).

Dillon and Quipuscoa, Synopsis of Galvezia

Fig. 2. Galvezia elisensii. A. Habit in coastal desert of Department of Arequipa, Peru. B. Flower at anthesis, bar = 5 mm. C. Maturing gynoecium, bar

= 2 mm. D. Seeds, bar = 1 mm.

Distribution & Biogeography .—The type locality of this new species is in the area of the Lomas deAtiquipa

(15°48'S, 74°22'W, Department Arequipa), but its distribution extends northward to Palpa (Department lea).

The presence of endemic taxa in southern Peru is a common pattern. From studies of the distribution of

lomas plants in coastal Peru and Chile, floristic sectors have been recognized, including a northern Peru unit

(7°55'S-12°00'S latitude), a south Peru unit (12°S-18°S), and a north Chile unit (20°S-28°S) (Rundel et al.

1991; Dillon et al. 2009). This pattern is found in several groups, including Nolana (Dillon et al. 2009) and hy¬

potheses of pattern formation remain to be tested. Dillon et al. (2009, 2011) discussed the sectoring of coastal

environments. Tong-term climatic changes have been associated with glacial cycles (13,000-200,000 year

cycles); and there have been at least 20 glacial cycles during the Pleistocene, each of approximately 200,000

years. The formation of glaciers on mountains and poles caused sea levels to fluctuate dramatically. Estimates

of sea level fluctuation range between 400-750 ft (120-230 m) and this lowering would have significantly

changed the position of the seashore 18,000 years ago, in relation to that today. This drop would have exposed

a considerable area of the continental shelf and displaced lomas formations, especially between 5°S to 15°S

latitude. Glacial cycles would also have had a profound influence on the flora of the coastal deserts by provid-

Journal of the Botanical Research Institute of Texas 8(1)

-90 -80 -70

ing geographic isolation at certain times, and at other times, opportunities for merging species, thereby allow¬

ing for gene exchange. Paradoxically, this would have also allowed for fragmenting populations, shifting their

ranges in relation to the near-ocean environments, adapting to changing conditions in situ , or undergoing

range reductions and extinction.

The situation in Galvezia seems to conform to the pattern of other species reflecting sectoring and influ¬

ence of past climate changes. The close relationship between G. fruticosa (northern sector) and G. elisensii

(southern sector) suggests a north to south pattern. The relationship with the northern Peruvian and Ecuador¬

ian populations included in G. grandiflora (i.e., G. lanceolata + G. ballii ) suggests a pattern influenced by the

most recent climatic change during the last glacial cycle (Weng et al. 2006).

Evolutionary Relationships. —Elisens (1992) admitted, that while isozyme data could not resolve evolu¬

tionary relationships among G. fruticosa, G. elisensii (as G. sp. nov.), and G. leucantha, the close affinity of these

species was supported by several shared, advanced, morphological characters, a large number of shared alleles

and separation from G. grandiflora (as G. ballii and G. lanceolata) in all networks.

Additional material examined: PERU. Arequipa: Provincia Caraveli, Sur de Nazca entre Km 518 y Km 590,15°26'S, 74°52'W, 80 m, 13 Nov

2005, M.O. Dillon, V. Quipuscoa, E. Ortiz, M. Corrales & G. Castillo 8792,8796 (HUSA, F); Distrito Bella Union: ca. Km 544 Panamericana Sur,

entre Division Puerto Lomas y Chavina, 15°33 , 06.2"S, 74°45 , 11.4"W, 57 m, 09 Jun 2012, V Quipuscoa S., S. Montesinos R., L. Apaza Ch. & F.

Miami Ch. 5058 (HUSA, F, USM, HAO, HUT). Ica: Provincia lea. A mi W of Pan-American Hwy, 15°26 , 30.5"S, 74°52 , 50.7"W, 280 m, 19 Feb

1994, T. Anderson, J. McAulifee, K. &F. Katterman, C. Diaz, C. Ostolaza, G. Fombardi, & W. Hodgson 7874 (ASU0047094, F2144517). Provincia

Dillon and Quipuscoa, Synopsis of Galvezia

Nazca, Marcona: Sur de Nazca entre Km 518 y Km 590,15°26'S, 74°52'W, 80-310 m, 13 Nov 2005, M.O. Dillon, J. Wen, V. Quipuscoa, E. Ortiz,

M. CorrcA.es, & G. Castillo 8792 (F2276589), 8796 (F2276592); Panamericana Km 424 (ca. 2 km S of Mirador Maria Reiche on the Nazca geo¬

glyphs), 700-750 m, 1 Oct 1997, M. Weigend & H. Forther 97/631 (F2184045). Provincia Pisco, 1 km before Puente Huaytara (Km 73 road

Pisco-Ayachucho), 1450 m, 29 Sep 1997, M. Weigend & H. Forther 97/586 (F2184124).

2. Galvezia fruticosaJ.F. Gmel., Syst. Nat. 2:937. 1791. (Fig. 3). Type: PERU. Lima: 1779 j. Dombey 7 (holotype: MPU020307;

isotypes: BM, K000528873).

Dodartiafragilis Ruiz & Pav., Syst. Veg. Fl. Peruv. Chile 1:97.1798. Type. PERU. Lima: Ruiz Lopez &Pavon s.n. (holotype: MA815219, based

upon Galvezia Dombey ex A.L. Jussieu).

Agassizia limensis Dombey ex Chav., Monog. Antirrh. 180, tab 11,1833. Type. PERU. Lima: J. Dombey 7 (holotype: MPU020307; isotypes:

BM001010978, K).

Galvezia limensis (Dombey ex Chav.) Benth., in DC., Prodr. 10:296.1846.

Galveziafruticosa is distributed along coastal Peru from Lima, north to Piura. It exhibits considerable environ¬

mental latitude with populations ranging to interior habitats along the western versant of the Andes. Overall,

the dimensions of leaves are larger, and the blades wider and ovate to lanceolate with distinct petioles. A few

isolated populations in the north have leaf morphologies approaching those exhibited by G. elisensii to which

it is putatively most closely related (Elisens 1992).

Specimens examined: PERU. Ancash: Provincia Casma, ca. 48 km N of Pativilca on PanAmericana Hwy, 10 m, 13 Oct 1984, M.O. Dillon &

M. Whalen 4007 (F1953250). Provincia Huarmey, Panamericana Norte Km 660, 47 m, 10 Oct 2000, M. Weigend, H. Farther, & N. Dostert

2000/667 (F2229797). Provincia Recuay, Km 46 on road from Pativilca to Recuay, ca. 580 m, 27 Jan 1983, M. Dillon, U. Molau, &R Matekaitis

3073 (F1932856). Cajamarca: Provincia Contumaza, Ascope - Algarrobal, 150 m, 29 Dec 1983, A. Sagdstegui A. & J. Mostacero L. 11332

(F1942599); El Portachuelo, 780 m, 20 Apr 1984, A. Sagdstegui A. 11388 (F1949672); Rupe - Huertas, 1200 m, 17Jun 1994, A. Sagdstegui A.,

S. Leiva G., & R LezamaA. 15362 (F2145086); Contumaza, La Paloma, 950 m, 5 May 1984, 1. Sdnchez Vega 3390 (F1953515); San Miguel,

entre Quinden y Platanar, 650-1100 m, 6 Oct 2001, E. Rodriguez R., E. Alvitez I., E. Eopez M. J. Cabrera C. & J. Chdvez G 2411 (F2230009). La

Libertad: Provincia Trujillo, base Cerro Campana, 150 m, 9 Jun 1985, J. Mostacero L. et al. 696 (F1994334). Lima: Provincia Canta, Canta

Valley, 26 km N of Lima, 350 m, 4 Aug 1957, P.C. Hutchison 1001 (F1569722). Provincia Yauyos, road from Pacaran to Yauyos, Km 35.8 after

Pacaran, 1500 m, 12°46 , 12"S, 77°56 , 41"W, 6 Oct 2002, M. Weigend, M. Ackermann, A. Cano, M. I. Fa Torre 7209 (F2247324).

3. Galvezia grandiflora (Benth.) Wettst., Nat. Pflanzenfam. 4(3b):61. 1891. (Fig. 3). Galvezia limensis var. grandiflora

Benth. In: DC., Prodr. 10:296.1846. Type. PERU. Piura: Payta [Paita], E. Hall 10 (Holotype: K000528872).

Galvezia ballii Munz, Proc. Calif. Acad. Sci. ser. 4,15:379.1926. nom. illeg., superfluous renaming.

Galvezia lanceolata Pennell, Notul. Natl Acad. Nat. Sci. Philadelphia 179:5. 1946. Type: ECUADOR. Manabi: above Aguas Blancas, on

the road from Puerto Lopez to Los Penas, 9 Jul 1942,0. Haught 3386 (holotype, PH, n.v.; isotype: US1708104, image 00122131).

Galvezia grandiflora encompasses the species diversity exhibited over a wide range of habitats extending from

northern Peru (Province Paita, Department Piura) to central Ecuador (Prov. Manabi, 1°32'S, 80°44'W). It is

unclear what factors have maintained genetic differentiation between G. grandiflora, with its southernmost

distributional terminus in northern Piura, and G.fruticosa, which extends north to the Sechura Desert (Elisens

1992).

Munz (1926, p. 379-380) discussed the validity of Wettstein’s combination, but rejected it as confusing,

and to eliminate confusion, he proposed Galvezia ballii Munz with collections by J. Ball s.n. types (US251553,

image 00122129, US13223550, image 00122130). With the acceptance of Bentham’s name as combined by Wet-

tstein, G. ballii Munz becomes a superfluous renaming. The acceptance of Bentham’s variety at the level of

species also encompases the taxa formerly treated under G. lanceolata Pennell.

Specimens examined: ECUADOR. Manabi: between Bahia de Caraquez and San Agustin, 0-150 m, 9 May 1980, G. Harling & E. Andersson

18957 (F1930417). PERU. Piura: Paita, Amotape, 4°53'S, 81°1'W, 18 Feb 1984, C.P. Cowan 4487 (F2038375).

4. Galvezia leucantha Wiggins, Occas. Pap. Calif. Acad. Sci. 65:1. 1968. (Fig. 3). Type: Ecuador. Galapagos Islands:

Isla Isabela, 29Jan 1967, E side of ridge near foot of Tagus Cove Mountain, NE of Caleta Tagus (Tagus Cove), ca. 1.5 to 2 km from the

beach, about 95 m, I.L. Wiggins & D. Porter 247 (holotype: CAS633842, image 002778; isotype: K).

Galvezia leucantha, including its various subspecies, is restricted to the Galapagos Islands (Ecuador) and has

been discussed and mapped (Elisens 1989; McMullen & Elisens 2000; Tye & Jager 2000; Jaramillo-Dlaz et al.

2014).

Journal of the Botanical Research Institute of Texas 8(1)

Wiggins (1968) described G. leucantha subsp. leucantha stating (p. 4) the following, “The most striking

feature differentiating this species from those heretofore placed in Galvezia is the waxy white corolla. All the

species of Galvezia described previously have deep red corollas, none of the shades within their range even ap¬

proaching pink or white .” However, far from being a discriminating character, corolla color is variable. McMul¬

len (pers. comm.) has observed that on Rabida Island Galvezia are described as possessing “... outside of the

corolla is completely reddish purple, while the insides ranges from pink to white.” (McMullen & Elisens 2000).

Further, Tye and Jager (2000) described the Galvezia on Santiago Island with the following corolla description

“... exterior magenta with the tips of the upper lobes white, while the interior is pink and white striped.” Wig¬

gins went on to state that “... the calyx lobes of G. leucantha are considerably more slender, longer in relation to

their width, and long in toto than those in G. fruticosa. Whereas the calyx and pedicles of G. fruticosa are

closely glandular-puberulent, the calyces and pedicles of G. leucantha are wholly glabrous.” Obviously, for the

discrimination of these subspecies, it is best to rely upon comparative morphology or geography of collections

rather than their corolla color.

Galvezia leucantha subsp. leucantha Wiggins

Galvezia leucantha subsp. porphyrantha Tye & H. Jager, Novon 10:165. 2000. Type: Ecuador. Galapagos Islands:

Isla Santiago, Cerro entre Bahia Ladilla y Cabo Nepean, 13 Nov 1998, ca. 40 m, H. Jager y A. Tye 53 (holotype: CAS1084133,

bcode#0002779; isotype: CAS1084015, image 0002780).

Galvezia leucantha subsp. pubescens Wiggins, Occas. Pap. Calif. Acad. Sci. 65:6. 1968. Type: Ecuador. Galapa¬

gos Islands: Isla Rabida (Jervis Island), occasional near the shore and from 500-950 ft, 20 Dec 1905, A. Stewart 3441 (holotype: CAS,

n.v.).

ACKNOWLEDGMENTS

We wish to thank Wayne Elisens for calling our attention to this southern Peruvian endemic with his study of

alloenzymes in Galvezia populations. We are grateful for support from the National Geographic Society and

National Science Foundation who funded fieldwork in coastal Peru and Chile between 1983 and 1999. Fred

Barrie is thanked for providing the Fatin diagnosis and help with nomenclatural questions. Billie F. Turner is

acknowledged for reading an early version of the manuscript. We thank Patricia Jaramillo-Dlaz and Conley K.

McMullen for valuable comments during the review process that greatly improved the manuscript. We thank

Christine Niezgoda, Anna Balia, and Daniel Fe who facilitated the imaging of the type specimen (Fig. 1), Ed-

gardo Ortiz for the photograph (Fig. 2C), and Federico Fuebert for preparation of the map (Fig. 3). We acknowl¬

edge the use of digital resources from JSTOR (http://www.jstor.org) and the Biodiversity Heritage Fibrary

(http://www.biodiversitylibrary.org). We thank students Fuz Apaza and Fabiola Miauri from the Biology De¬

partment at Universidad Nacional San Agustln for help with gathering data in the held and laboratory. We also

acknowledge support from members of the Grupo DIBIOS and Instituto MOD, especially Cristian Tejada

Perez, Deysi Caballero, Carmen Fernandez, Smilsa Montesinos, and Raquel Medina.

REFERENCES

Bentham, G. 1846. Scrophulariaceae, Galvezia. In: DC. Prodr. 10:295-296.

Bickford, D., DJ. Lohman, N.S. Sodhi, P.K.L. Ng, R. Meier, K. Winker, K.K. Ingram, & I. Das. 2007. Cryptic species as a window on

diversity and conservation. Trends Ecol. Evol. 22:148-155.

Brako L. & J. Zarucchi. 1993. Catalogue of the flowering plants and gymnosperms of Peru. Monogr. Syst. Bot. Missouri

Bot. Gard.45:1-1286.

Cock, MJ.W. & J.O. Boos. 2006. Observations on Sphingidae (Lepidoptera) from Talara, North central Peru. Rev. pera.

Entomol. 45:75-78.

Chavannes, EJ. 1833. Agassizia limensis Dombey ex Chav., Monogr. Antirrhin. 180.

Dillon, M.O.,T.Tu, L. Xie, V. Quipuscoa S., & J. Wen. 2009. Biogeographic diversification in /Vo/ana(Solanaceae), a ubiquitous

member of the Atacama and Peruvian Deserts along the western coast of South America. J. Syst. Evol. 47:457-476.

Dillon, M.O., S. Leiva G., M. Zapata C., P. Lezama A., & V. Quipuscoa S. 2011 (2012). Floristic Checklist of the Peruvian Lomas

Formations - Catalogo floristico de las Lomas peruanas. Arnaldoa 18(1 ):7-32.

Dillon and Quipuscoa, Synopsis of Galvezia

Eusens, WJ. 1989. Genetic variation and evolution of the Galapagos shrub snapdragon. Natl. Geogr. Res. 5:98-110.

Elisens, WJ. 1992. Genetic divergence in Galvezia (Scrophulariaceae): Evolutionary and biogeographic relationships

among South American Galapagos species. Amer. J. Bot. 79:198-206.

Eusens, WJ. & A.D. Nelson. 1993. Morphological and isozyme divergence in Gambelia (Scrophulariaceae): species delimi¬

tation and biogeographic relationships. Syst. Bot. 18:454-468.

Ghebrehiwet, M., B. Bremer, & M. Thulin. 2000. Phylogeny of the tribe Antirrhineae (Scrophulariaceae) based on morpho¬

logical and ndhF sequence data. PI. Syst. Evol. 220:223-239.

Jaramillo-Diaz, P, A. GuEzou, A. Mauchamp, & A. Tye. 2014. CDF checklist of Galapagos flowering plants - FCD Lista de

especies de plantas con flores de Galapagos. In: F. Bungartz, H. Herrera, P. Jaramillo, N. Trado, G. Jimenez-Uzcategui,

D. Ruiz, A. Guezou, and F. Ziemmeck, eds. Charles Darwin Foundation Galapagos species checklist - Lista de espe¬

cies de Galapagos de la Fundacion Charles Darwin. Charles Darwin Foundation / Fundacion Charles Darwin, Puerto

Ayora, Galapagos: http://www.darwinfoundation.org/datazone/checklists/vascular-plants/magnoliophyta/ Last

updated 21 Jan 2014.

Jorgensen, P.M. & S. Leon-Yanez. 1999. Catalogue of the vascular plants of Ecuador. Monogr. Syst. Bot. 75:1-1181.

McMullen, C.K. & WJ. Eusens. 2000. Variation in the Galapagos shrub snapdragon: is there another subspecies of Galvezia

ieucantha ? Not. Galapagos 61:21 -22.

Medhanie, G., B. Bremer, &M. Thulin. 2000. Phylogeny of the tribe Antirrhineae (Scrophulariaceae) based on a morphologi¬

cal and ndhF sequence data. PI. Syst. Evol. 220:223-239.

Munz, P.A. 1926. The Antirrhinoideae-Antirrhineae of the New World. Proc. Calif. Acad. Sci., Ser.4,15:323-397.

Oyama, R.K. & D.A. Baum. 2004. Phylogenetic relationships of North American Antirrhinum (Veronicaceae). Amer. J. Bot.

91:918-925.

Pennell, F.W. 1946. Species of Stemodia and Galvezia (Scrophulariaceae) on the coast of Ecuador and northern Peru.

Notul. Natl. Acad. Nat. Sci. Philadelphia 179:1-6.

Ruiz Lopez, H. & J.A. Pavon. 1794. Galvezia. FI. Peruv. Prodr. 56.

Ruiz Lopez, H. & J.A. PavOn. 1798. Dodartia fragilis. Syst. Veg. FI. Peruv. Chil. 97.

Rundel, P.W., M.O. Dillon, B. Palma, H.A. Mooney, S.L.Gulmon,&J.R. Ehleringer. 1991.The phytogeography and ecology of the

coastal Atacama and Peruvian deserts. Aliso 13:1-50.

Sutton, D.A. 1988. A revision of the tribe Antirrhineae. Oxford University Press, London, U.K.

Schlick-Steiner, B.C., B. Seifert, C. Stauffer, E. Christian, R.H. Crozier, & F.M. Steiner. 2007. Without morphology, cryptic species

stay in taxonomic crypsis following discovery. Trends Ecol. Evol. 22:391-392.

Tye, A. & H. Jager. 2000. Galvezia Ieucantha subsp. porphyrantha (Scrophulariaceae), a new shrub snapdragon endemic

to Santiago Island, Galapagos, Ecuador. Novon 10:164-168.

Vargas, P, J.A. Rossello, R. Oyama, & J. Goemes. 2004. Molecular evidence for naturalness of genera in the tribe Antirrhineae

(Scrophulariaceae) and three independent evolutionary lineages from the New World and the Old. PI. Syst. Evol.

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in the western Peruvian Andes. Quatern. Res. 66:87-96.

Wettstein, R. 1891. Scrophulariaceae. Nat. Pflanzenfam. 4(3b):61, 39-107.

Wiggins, I.L. 1968. A new species and subspecies of Galvezia (Scrophulariaceae) from the Galapagos Islands. Occas. Pap.

Calif. Acad. Sci. 65:1, f. a-l.

Wiggins, I.L. and D.M. Porter. 1971. Flora of the Galapagos Islands. Stanford University Press, Stanford, CA, U.S.A.

Journal of the Botanical Research Institute of Texas 8(1)

BOOK NOTICE

Paul E. Minnis, ed. 2014. New Lives for Ancient and Extinct Crops. (ISBN-13: 978-0-8165-3062-5, hardcov¬

er). University of Arizona Press, PO Box 210055, Tucson, Arizona 85721-0055, U.S.A. (Orders: www.

uapress.arizona.com, 1-800-621-2736). $65.00, 288 pp., 25 photos, 22 illustrations, 13 tables, index,

6" x 9".

From the publisher: Over many millennia, farmers across the world have domesticated literally thousands of

species and developed tens of thousands of varieties of these plants. Despite the astonishing agricultural diver¬

sity that existed long ago, the world’s current food base has narrowed to a dangerous level. By studying the long

and dynamic history of farming in the ancient past, archaeology can play a part in helping ensure the stability

of the human food supply by identifying once-important crops and showing where and how such crops were

grown in the past. Thanks to this work, extinct crops might even be redomesticated from their wild progenitors.

New Lives for Ancient and Extinct Crops profiles nine plant species that were important contributors to

human diets and had medicinal uses in antiquity: maygrass, chenopod, marshelder, agave, little barley, chia,

arrowroot, little millet, and bitter vetch. Each chapter is written by a well-known scholar, who illustrates the

global value of the ancient crop record to inform the present. From eastern and western North America, Meso-

america, South America, western Asia, and south-central Asia, the contributors provide examples of the unex¬

pected wealth of information available in the archaeological record about ancient and extinct crops.

PAUL E. MINNIS is a professor of anthropology at the University of Oklahoma and former president of the

Society of Ethnobiology. He is editor of Ethnobotany: A reader and coeditor of Biodiversity and Native America.

J.Bot. Res. Inst. Texas 8(1): 56.2014

A NEW SPECIES OF CREMOSPERMA (GESNERIACEAE)

FROM NORTHEASTERN PERU

Brian R. Keener

Department of Biological

& Environmental Sciences

The University of West Alabama

Livingston, Alabama 35470, U.S.A.

BKeener@uwa.edu

John L. Clark

Department of Biological Sciences

Box 870345

The University of Alabama

Tuscaloosa, Alabama 35487, U.S.A.

ABSTRACT

Recent fieldwork and investigations of herbarium specimens during a preliminary revision of Cremosperma (Gesneriaceae) have resulted in

the discovery of a new species. The new species, Cremosperma inversum B.R. Keener & J.T. Clark, is distinguished from other species in

the genus by a suite of characters including orbicular leaves, stoloniferous habit, and tomentose-wooly indument on the upper leaf surface.

The distribution of trichomes on the adaxial leaf surface is clustered in the center of the bullae and away from the more or less glabrous veins

and glabrous on the abaxial leaf surface away from the more densely pubescent veins. This species is endemic to sandstone outcrops in the

Department of Amazonas in northwestern Peru.

RESUMEN

Expediciones recientes al noroccidente de Peru, y el estudio de las colecciones de herbario como parte de la revision preliminar del genero

Cremosperma (Gesneriaceae), han dado como resultado el descubrimiento de una nueva especie. La nueva especie, Cremosperma inver¬

sum B.R. Keener & J.L. Clark, se distingue de las otras especies del genero por un conjunto de caracteres que incluye: hojas orbiculares,

habito estolonifero e indumento tomentoso-lanoso en el haz de la lamina. Los tricomas en la superficie adaxial de la lamina estan agrupados

entre las venas, las cuales son mas o menos glabras, mientras que, en la superficie abaxial las venas son pubescentes y el resto de la superficie

es mas o menos glabra. Esta especie es endemica de los afloramientos rocosos en el departamento de Amazonas, noroccidente peruano.

Key Words: Cremosperma, Gesneriaceae, Taxonomy, Flora of Peru, Amazonas Department, Sandstone

INTRODUCTION

Cremosperma is a genus of terrestrial or saxicolous herbs ranging from Costa Rica to Peru. The genus is a

strongly supported monophyletic lineage in the New World tribe Beslerieae (Clark et al. 2010; Roalson & Clark

2006; Smith 2000). Taxonomic and phylogenetic information for Cremosperma were recently summarized in

Clark and Skog (2011). Until now, the only other known Cremosperma located south of Ecuador was the en¬

demic Peruvian species Cremosperma peruvianum L.E. Skog (Skog 1982).

Kvist and Skog (1988) provided a traditional monograph of the Ecuadorian species and included 14 taxa,

several of which also occur in Colombia and estimated the total diversity of Cremosperma to be 23 species. The

following three Cremosperma species have been described since Kvist and Skog (1988): Cremosperma micrope-

cten Fern. Alonso from Colombia (Fernandez-Alonso 2006); Cremosperma anisophy Hum J.L. Clark & L.E. Skog

from Ecuador (Clark & Skog 2011); and Cremosperma verticillatum J.L. Clark & B.R. Keener from Ecuador

(Clark & Keener 2011). Recent fieldwork in Ecuador, Panama, and Colombia has resulted in the discovery of

additional Cremosperma species that will be described in the near future. The description of Cremosperma in¬

versum brings the total published diversity of the genus to 27 species and the second species from Peru.

TAXONOMIC TREATMENT

Cremosperma inversum B.R. Keener &J.L. Clark, sp. nov. (Fig. 1). Type: PERU. Amazonas: Bagua district, Cerro Tayu, ca.

1 hour from Chiriaco, 05°15 , 56"S, 78°22 , 07"W, 800 m, 19 Mar 2001, H. van der Werjj, R. Vasquez & B. Gray 16240 (holotype: US;

isotypes: F, MO, NY).

Differs from Cremosperma peruvianum by the presence of orbicular leaves (vs. oblong) and tomentose-wooly indument on upper leaf surface

(vs. sparsely villous indument).

J. Bot. Res. Inst. Texas 8(1): 57 - 60.2014

Journal of the Botanical Research Institute of Texas 8(1)

Terrestrial herb; stems 5-25 cm long, creeping to stoloniferous proximally, rooting at nodes, ascending to

erect distally, rarely branched, slightly angled to sulcate, tomentose with septate, uniseriate trichomes, also

with small amber colored glandular protuberances. Leaves opposite, isomorphic, petiolate; petioles terete,

4-15 mm long, tomentose; blades widely elliptic to orbicular, 1.0-3.8 x 0.9-3.5 cm, base symmetrical to

slightly oblique, rounded to slightly cordate, apex rounded, margin crenate with broad shallow teeth, bullate

when fresh, membranaceous and flat when dry, abaxially dark green, tomentose on veins, with glandular pro¬

tuberances in area between lateral veins, adaxially dark green, densely tomentose in area between veins,

slightly tomentose on veins. Inflorescence a reduced pair-flowered cyme, appearing clustered and pseudo-

umbellate, in upper leaf axils, peduncle 2-3 cm long, (1—)2—5 mature flowers/inflorescence, often with rem¬

nant pedicel scars appearing gland-like; bracts absent. Flowers pedicellate, pedicels 2-5 mm long, pilose; ca¬

lyx 3.5-4.5 mm long, lobes 5, fused for 1/2-3/4 of their length, equal, lobes erect during anthesis, persistent

and spreading in fruit to form a splash cup, apex rounded to obtuse, uniformly green, outside pilose, inside

glabrous; corolla 10-11 mm long, tubular, slightly to strongly curved in lower 1/3, base to mid-region 1.5 mm

in diameter, throat 2.5 mm wide at apex, white with yellowish upper region of throat, outer surface of tube

glabrous proximally, pilose distally, throat and corolla lobes abaxially pilose, inner surface of tube glabrous,

throat and base of corolla lobes pilose, corolla lobes glabrous distally, limb bilaterally symmetrical, lobes re¬

flexed and unequal, oblanceolate to spatulate, lower three lobes ca. 3.1 x 3.0 mm, upper two lobes 2.1 x 2.1 mm,

margins entire to slightly crenulate, slightly undulate; stamens 4, didynamous, included; filaments adnate to

corolla tube, abaxial filaments free for 1.5 m, adaxial filaments free for 0.8 mm, glabrous; anthers broader than

long, ca. 0.5 x 1.0 mm; staminode absent; nectary enclosing the ovary on one side, glabrous, ca. 1.1 mm long;

ovary superior, glabrous, ca. 1.1 x 1.5 mm, style and stigma glabrous. Fruit a dry bivalved capsule that dehisces

laterally as it matures and appears 4-valved, globose, ca. 2.0 mm in diameter; seeds numerous, irregularly el¬

liptic to ovoid, often slightly arched, ca. 0.5 x 0.2 mm, reddish brown, surface shallowly alveolate, cavities usu¬

ally longer than wide.

Cremosperma inversum is differentiated from other congeners by the small diminutive habit (to 10 cm

tall). Other small Cremosperma species include C. muscicola L.P. Kvist & L.E. Skog, C. pusillilum C.V. Morton,

and C. veraguanum Wiehler. The isophyllous leaves in C. inversum differentiate it from the anisophyllous leaves

of C. veraguanum and C. muscicola. The orbicular leaves in C. inversum readily differentiate it from congeners

with oblong leaves. The only other known species of Cremosperma from Peru is C. peruvianum. The leaves in C.

peruvianum are oblong in contrast to the orbicular leaves in C. inversum. These two species are geographically

separated by 600+ km with C. peruvianum in the southern department of Huanuco and C. inversum in the

northern province of Amazonas. The leaves in C. inversum are tomentose-wooly on the adaxial leaf surface

with clustered trichomes in the center of the bullae and away from the more or less glabrous veins and glabrous

on the abaxial leaf surface away from the more densely pubescent veins (Fig. 1A, B).

Distribution and habitat.—Cremosperma inversum is endemic to the Bagua district in the Department of

Amazonas of northwest Peru. All of the currently known collections are from sandstone outcrops in a small

area (less than 10 km) in mature lowland rainforest (400-800 m).

Etymology. —The new species name “ inversum ” is in reference to the seemingly inverse pubescence pat¬

tern on the abaxial and adaxial leaf surfaces. Adaxial surfaces are mostly glabrous along the veins and pubes¬

cence between veins. Abaxial surfaces are mostly pubescent only along the veins and glabrous between the

veins (Fig. 1A, B).

Conservation and IUCN Red List category.—Cremosperma inversum is geographically limited to a small

area in northwestern Peru. According to the IUCN Red Fist criteria (IUCN 2001) the limited geographic range

(B2a, less than 10 km 2 and known to exist at only a single location) qualify Cremosperma verticillatum for being

listed in the category CR (Critically Endangered).

Additional specimens studied: PERU. Amazonas: Bagua district, Quebrado El Almendro, 05°14 , 40"S, 78°2r24"W, 430 m, 9 Mar 1998, H. van

der Werff et al 14567 (US); Imaza, Tayu Mujaji, comunidad de Wawas, 05°15 , 25"S, 78°2r41"W, 800 m, 23 Oct 1997, R. Rojas et al 428 (US);

Tayu Mujaji, 05°15 , 56"S, 78°22 , 07"W, 800 m, 16 Feb 2002, R. Vdsquez27590 (US).

Keener and Clark, A new Peruvian species of Cremosperma

Fig. 1 . Illustration of Cremosperma inversum. A. Abaxial leaf surface showing dense pubescence on veins and glabrous between veins. B. Adaxial leaf

surface showing glabrous veins and tomentose between veins. C. Habit showing stolon. D. Inflorescence. E. Front view of corolla. F. Lateral view of

flower. G. Lateral view of open flower showing androecium. H. Calyx showing truncate scale-like gland. I. Seed. J. Mature fruit with persistent calyx.

(A-J from the holotype, H. van der Werffetal. 16240 (US).

Journal of the Botanical Research Institute of Texas 8(1)

ACKNOWLEDGMENTS

This study was supported by funds from the National Science Foundation (DEB-841958 and DEB-0949169) to

JLC. Additional support for the first author was provided the National Science Foundation Research Opportu¬

nity Award (NSF-ROA 1112630). We thank the herbaria F, K, MO, NY, Q, QAP, QCA, QCNE, SEL, and US for

access to their collections. We thank Laura Clavijo for providing the Spanish translation of the abstract. Lau¬

rence E. Skog and Christian Feuillet are gratefully acknowledged for providing helpful reviews. We also thank

Steve Ginzbarg (UNA) for facilitating loans and Sue Blackshear for preparing the illustration.

REFERENCES

Clark, J.L. & L.E. Skog. 2011. Novae Gesneriaceae Neotropica rum XVI: Cremosperma anisophyllum, a new species of

Gesneriaceae from the Choco region of northern Ecuador and southern Colombia. Brittonia 63:133-138.

Clark, J.L. & B.R. Keener. 2011. Cremosperma verticillatum (Gesneriaceae), a new species from northwestern Ecuador. J.

Bot. Res. Inst.Texas 5:499-504.

Clark, J.L., D.A. Neill, J.A. Gruhn, A. Weber, &T. Katan. 2010. Shuaria (Gesneriaceae), an arborescent new genus from the

Cordillera del Condor and Amazonian Ecuador. Syst. Bot. 35:662-674.

FernAndez-Alonso, J.L. 2006. Novedades taxonomicas y nomenclaturales en Cremosperma y Resia (Gesneriaceae) de Co¬

lombia. Rev. Acad. Colomb. Ci. Exact. Fis. Nat. 30:171-180.

IUCN. 2001. IUCN Red List Categories and Criteria, Version 3.1. Prepared by the IUCN Species Survival Commission.

Gland, Switzerland and Cambridge, UK. International Union for Conservation of Nature and Natural Resources.

Kvist, L.P. & L.E. Skog. 1988. The genus Cremosperma (Gesneriaceae) in Ecuador. Nordic J. Bot. 8:259-269.

Roalson, E.H. & J.L. Clark. 2006. Phylogenetic patterns of diversification in the Beslerieae (Gesneriaceae). In: Plant ge¬

nome: Biodiversity and evolution, Phanerogams 1C. A.K. Sharma & A. Sharma, eds. Science Publishers, Enfield, New

Hampshire, USA. Pp. 251-268.

Skog, L.E. 1982. New Gesneriaceae from Peru and Ecuador. Selbyana 7:94-99.

Smith, J.F. 2000. A phylogenetic analysis of tribes Beslerieae and Napeantheae (Gesneriaceae) and evolution of fruit

types: parsimony and maximum likelihood analyses of ndhF sequences. Syst. Bot. 25:72-81.

THREE NEW SPECIES OF SENEGALIA (FABACEAE) FROM BRAZIL

David S. Seigler*

Department of Plant Biology

University of Illinois

Urbana, Illinois 61801, US.A.

seigler@life. illinois. edu

*corresponding author

John E. Ebinger

Department of Biology

Eastern Illinois University

Charleston, Illinois 61920, US.A.

Petala Gomes Ribeiro and Luciano Paganucci De Queiroz

Botanica

Universidade Estadual de Feira de Santana

Av. Transnordestina, s/n, Novo Horizonte

44036-900, Feira de Santana, Bahia, BRASIL

petalagribeiro@gmail. com; lqueiroz@uefs. br

ABSTRACT

Senegalia irwinii Seigler, Ebinger, & RG. Ribeiro from the states of Bahia, Minas Gerais, and Rio de Janeiro; S. harleyi Seigler, Ebinger, &

RG. Ribeiro from the states of Bahia, Minas Gerais, and Parana; and S. hatschbachii Seigler, Ebinger, & P.G. Ribeiro from the states of Minas

Gerais, Parana, and Sao Paulo in Brazil are described, illustrated and compared to their probable nearest relatives, Senegalia mattogrossensis

(Malme) Seigler & Ebinger, Senegalia martiusiana (Steud.) Seigler & Ebinger, and Senegalia tucumanensis (Griseb.) Seigler & Ebinger, re¬

spectively.

Key Words: Fabaceae, IUCN Red List, Mimosoideae, Senegalia

RESUMEN

Se describen, ilustran y comparan, las especies Senegalia irwinii Seigler, Ebinger, & P.G. Ribeiro de los estados de Bahia, Minas Gerais, y

Rio de Janeiro; S. harleyi Seigler, Ebinger, & P.G. Ribeiro de los estados de Bahia, Minas Gerais, y Parana; y S. hatschbachii Seigler, Ebinger,

& P.G. Ribeiro de los estados de Minas Gerais, Parana, y Sao Paulo de Brasil, con las especies afines, probablemente mas cercanas, Senegalia

mattogrossensis (Malme) Seigler & Ebinger, Senegalia martiusiana (Steud.) Seigler & Ebinger, y Senegalia tucumanensis (Griseb.) Seigler &

Ebinger, respectivamente.

The genus Senegalia has previously been treated as part of Acacia s.l., but recent morphological and genetic

studies have shown that this large genus is polyphyletic. Relationships within the genus Acacia s.l., as well as

the position of the genus within the Mimosoideae have been clarified by data from molecular studies (Maslin

et al. 2003a; Miller & Bayer 2003; Luckow et al. 2003; Miller et al. 2003; Rico-Arce & Bachman 2006; Seigler et

al. 2006a; Bouchenak-Khelladi et al. 2010; Gomez-Acevedo et al. 2010; Murphy et al. 2010; Miller & Seigler

2012; Kyalangalilwa et al. 2013). Based on both morphological and molecular data, Acacia s.l. is now regarded

as comprising at least five genera, Acacia s.s., Acaciclla Britton & Rose (1928), Mariosousa Seigler & Ebinger

(Seigler et al. 2006b), Senegalia Raf. (1838), and Vachellia Wight & Arnott (1834) (see Miller & Seigler 2012 for

overview of the new generic classification).

Members of Senegalia are shrubs, trees, or lianas, unarmed or armed with prickles, but without stipular

spines. The prickles usually are scattered, but less commonly are grouped in twos or threes, usually at or near

the nodes (Vassal 1972). Leaves are bipinnate and the petiole and primary rachis have sessile or stipitate glands

of variable position. Flowers possess a more or less tubular nectary below the usually stipitate ovary. Inflores¬

cences are globose heads (capitula) or spikes, often grouped into complex terminal pseudo-inflorescences or

synflorescences. Pods are dehiscent, separating into two valves at maturity, or less commonly indehiscent or

separating into indehiscent one seeded articles. The seeds are uniseriate.

The genus Senegalia consists of approximately 100 taxa in the Americas (unpublished data), as well as 69

J. Bot. Res. Inst. Texas 8(1): 61 - 69.2014

Journal of the Botanical Research Institute of Texas 8(1)

in Africa, 43 in Asia, and two in Australia (Maslin et al. 2003a,b). Eight species occur in two or more areas. Ap¬

proximately half of the American species occur in Brazil. During the course of our work on the genus Senegalia

Raf. of Brazil, three undescribed species were noted from herbarium materials of Bahia, Minas Gerais, Parana,

Sao Paulo, and Rio de Janeiro. These taxa are clearly distinctive and are herein proposed as new species.

Senegalia irwinii Seigler, Ebinger, & P.G. Ribeiro, sp. nov. (Fig. 1). Type: BRAZIL. Minas Gerais: 40 km E of Belo Hori¬

zonte, near BR-31,1800 m, 16 Jan 1971, H.S. Irwin, R.M. Harley &E. Onishi 30523 (holotype: MO; isotypes: MBM, NY).

Senegalia irwinii Seigler, Ebinger & RG. Ribeiro differs from other Senegalia species by leaf size (100-170 mm long), petiolar gland usually

one or two, columnar (1.1-3.1 mm) apex 0.5-0.8 mm in diameter, pinnae 5 to 11 pairs/leaf (60-115 mm), 10-18 mm between pinna pairs,

leaflets 45 to 85pairs/pinna, midvein subcentral; inflorescence a globose head 14-20 mm across, flowers sessile or subsessile, ovary glabrous.

Climbing shrub or small tree to 5 m tall; bark not seen; twigs dark purplish brown, not to slightly flexuous,

terete to slightly ridged, glabrous to lightly puberulent; short shoots absent; prickles dark purplish brown

throughout, flattened, recurved to rarely straight, woody, 1-4 x 1-4 mm at the base, glabrous, scattered along

the twig, petiole and rachis. Leaves alternate, 100-170 mm long; stipules dark brown, linear, symmetrical,

flattened, straight, herbaceous, 2-5 x 0.2-0.5 mm near the base, glabrous to puberulent, early deciduous; peti¬

ole adaxially grooved, 20-38 mm long, lightly puberulent; petiolar gland usually 1 or 2, one located near the

middle of the petiole, one at or near the first pinna pair, columnar, 1.1-3.1 mm long, apex 0.5-0.8 mm across,

orbicular, depressed, glabrous; rachis adaxially grooved, 80-135 mm long, lightly puberulent, a columnar

gland 0.6-1.2 mm long usually between the upper, and sometimes other pinna pairs, apex 0.4-0.8 mm across,

orbicular, depressed, glabrous; pinnae 5 to 11 pairs/leaf, 60-115 long, 10-18 mm between pinna pairs; para-

phyllidia 0.8-1.4 mm long; petiolule 1.9-4.2 mm long; leaflets 45 to 85 pairs/pinna, opposite, 0.6-1.2 mm be¬

tween leaflets, linear, 6-12 x 1.2-1.8 mm, lightly pubescent with appressed hairs beneath, shiny and glabrous

above, lateral veins not obvious, 1 vein from the base, base oblique, truncate on one side, margins lightly ciliate,

apex acute, midvein subcentral. Inflorescence a densely 20- to 40-flowered globose head 14-20 mm across, in

axillary and terminal pseudo-paniculate clusters, the main axis to 300 mm long; peduncles 5-15 x 0.4-0.6 mm

thick, puberulent; receptacle not enlarged, slightly elongated; involucre rarely a single small bract located me¬

dially on the peduncle, early deciduous; floral bracts spatulate, 0.8-1.2 mm long, puberulent, early deciduous.

Flowers sessile or subsessile, cream; calyx 5-lobed, 2.5-3.2 mm long, puberulent; corolla 5-lobed, 3.4-4.2 mm

long, glabrous or nearly so, lobes one-quarter the length of the corolla; stamens 50 to 70; stamen filaments 8-10

mm long, distinct; anther glands absent; ovary glabrous, sessile to short-stalked. Fruits oblong, 75-130 x 22-

28 mm, straight, flattened, not constricted between the seeds, chartaceous, transversely striated, puberulent,

eglandular, dehiscent along both sutures; stipe 8-12 mm long; apex obtuse, short beaked. Seeds not seen.

Distribution and ecology. —Deciduous to evergreen savannas, disturbed second growth forest and thickets

from 1,100 to 1,800 m in the states of Bahia, Minas Gerais, and Rio de Janeiro in eastern Brazil.

Phenology. —Flowering Jan-Mar.

Local Names and Uses.—None known.

Etymology.—Senegalia irwinii is named after Howard S. Irwin, a well-known authority on Fabaceae, who

led eight expeditions to Brazil and Guiana between 1960 and 1972, and collected extensively in northeastern

Brazil.

IUCN Red List category. —DD, data deficient. A rare endemic restricted to the states of Bahia, Minas Gerais

and Rio de Janeiro, Brazil. As we have seen fewer than 10 collections, the species is possibly threatened (IUCN

2001 ).

Paratypes: BRAZIL: Bahia: 3 km da divisa Minas Gerais, 9 Jul 1964, L. Duarte &A. Castellanos 291 (NY, RB). Minas Gerais: 10 km. by road

N of Gouveia, 1320 m, 11 Apr 1973, W.R. Anderson 8664 (CM, F, NY, UB);JardimBotanico, Belo Horizonte, 15 Jan 1934, M. Barreto 6458 (F);

17 km NE of Diamantina, road to Mendanha, 1250 m, 29 Jan 1969, H.S. Irwin, R. Reis dos Santos, R. Souza & S.F. da Fonseca 22870 (MO, NY,

US); Belo Horizonte, campus daUFMG, 4 May 1995 J.A.Lombardi &L.G. Femponi 760 (NY); 1879,J. M iers241b (BM); Brumadinho, Inhotim,

Borda de mata semi decidua da trilha da Caixa DAgua, 20°08 , 21"S, 44°14T3"W, 870 m, 22 Jan 2008, J.G. Oliveira & F.M. Rodrigues 55

(BHCB). Rio de Janeiro: Municipio de Campos dos Goytacazes, Bom Jesus, Assentamento dos Sem Terra (nucleo 1-N1), mata do Bom Jesus,

mato do Caixaou mata da Santa Casa, 3 Oct 2000, J.M.A. Braga 6340 (RB); Rio das Ostras, Reserva Biologica a Uniao, 50 m, 14 Jun 2001,

J.M.A. Braga 6667 (MBM, NY, RB).

Seigler et al., New species of Senegalia from Brazil

Fig. 1. Senegalia irwinii Seigler, Ebinger, & P.G. Ribeiro; A. leaflet, adaxial surface; B. flower; C. portion of twig and petiole with prickles and petiolar

glands; D. leaf; E. pseudo-inflorescence. A, B, D from Irwin etal. 30523 (NY); C, E from Irwin etal. 22870 (MO).

Journal of the Botanical Research Institute of Texas 8(1)

Morphologically similar to Senegalia mattogrossensis (Malme) Seigler & Ebinger, this new species has a compa¬

rable range in eastern Brazil, but grows at a higher elevation, occurring above 1,100 m. Senegalia mattogrossen¬

sis, in contrast, is found below 1,000 m. Morphologically, these two taxa are distinct with S. mattogrossensis

being densely pubescent on the twig; petiole and rachis with yellow hairs to 0.8 mm long. Senegalia irwinii, in

contrast, lacks the yellow pubescence, the twig, petiole, and rachis are glabrous to lightly puberulent with gray

hairs. This taxon also has leaflets that are consistently 6-12 mm long whereas S. mattogrossensis has leaflets

shorter than 6.5 mm in length

Senegalia harleyi Seigler, Ebinger, & P.G. Ribeiro, sp. nov. (Fig. 2). Type: BRAZIL. Bahia: 12 km SE of Barra do Choga, road

to Itapetinga, 700 m, 30 Mar 1977, R.M. Harley, S.J. Mayo, R.M. Storr, T.S. Santos & R.S. Pinheiro 20161 (holotype: NY; isotypes:

CEPEC, IPA, RB, UEC, US).

Senegalia harleyi Seigler, Ebinger & P.G. Ribeiro differs from other Senegalia species by leaf size (60-130 mm long), petiolar glands one to

three scattered along the petiole, sessile, oval to orbicular (0.7-2.5 mm across), pinnae 11 to 25 pairs/leaf, distance between pinna pairs (3-7

mm), leaflets 38 to 80 pairs/pinna, distance between leaflet pairs (0.2-0.4 mm), midvein subcentral; inflorescence a globose head 12-16 mm

across, flowers sessile, anther glands present, ovary glabrous.

Climbing shrub or small tree to 5 m tall; bark not seen; twigs dark brown to dark purplish brown, slightly

flexuous, terete to slightly ridged, densely pubescent with mostly erect dark brown hairs to 0.6 mm long; short

shoots absent; prickles dark brown below, commonly lighter brown above, flattened, commonly recurved,

woody, 1-3 x 1-3 mm at the base, densely pubescent with straight hairs, common in lines on the twig ridges,

also on petiole and rachis. Leaves alternate, 60-130 mm long; stipules dark brown, linear, symmetrical, flat¬

tened, straight, herbaceous, 2-5 x 0.3-0.8 mm, densely ciliate, early deciduous; petiole shallowly adaxially

grooved, 6-15 mm long, densely pubescent; petiolar glands 1 to 3, scattered along the petiole with one just

below the first pinna pair, sessile, oval to orbicular, 0.7-2.5 mm across, apex flattened or the margins raised to

form a cup, glabrous; rachis adaxially grooved, 55-120 mm long, densely pubescent, an oval to orbicular gland

0.4-1.2 mm across between the upper 1 to 6, and sometimes other pinna pairs, apex cup-shaped, glabrous;

pinnae 11 to 25 pairs/leaf, 23-58 mm long, 3-7 mm between pinna pairs; paraphyllidia 0.4-0.8 mm long, com¬

monly absent; petiolule 0.4-1.1 mm long; leaflets 38 to 80 pairs per/pinna, opposite, 0.2-0.4 mm between

leaflets, linear, 2.5-4.4 x 0.4-0.7 mm, glabrous to scattered pubescent below, lateral veins not obvious, 1 vein

from the base, base oblique, margins ciliate, apex acute, midvein subcentral. Inflorescence a densely 20- to

35-flowered globose head 12-16 mm across, in terminal and axillary pseudo-paniculate clusters, the main axis

to 400 mm long; peduncles 4-9 x 0.4-0.7 mm thick, densely pubescent; receptacle elongated, not enlarged;

involucre a small bract located on the upper half of the peduncle, early deciduous; floral bracts spatulate, 0.5-

0.8 mm long, puberulent, early deciduous. Flowers sessile, white to cream; calyx 5-lobed, 1.4-2.1 mm long,

puberulent; corolla 5-lobed, 2.3-3.5 mm long, puberulent, lobes one-sixth the length of the corolla; stamens 50

to 79; stamen filaments 6.5-8.0 mm long, distinct; anther glands present; ovary glabrous, sessile to subsessile.

Fruits oblong, 60-130 x 13-26 mm, straight, flattened, not constricted between the seeds, chartaceous, lightly

transversely striated, pubescent, eglandular, dehiscent along both sutures; stipe 6-11mm long; apex acumi¬

nate to obtuse, short beaked. Seeds 5-6 x 4-5 mm, ovate to elliptic, flattened, smooth; pleurogram U-shaped.

Distribution. —Humid tropical forest, disturbed second growth forests and thickets from 500 to 2200 m

in the states of Bahia, Minas Gerais, and Parana, Brazil.

Phenology. —Flowering Jan-Mar.

Local Names and Uses. —unha-de-gato (Mucuge- Bahia)

Etymology. —Named after Raymond M. Harley, authority on the flora of the caatinga of northeastern Bra¬

zil and collector of many legumes from that area.

IUCN Red List category. —DD, data deficient. Senegalia harleyi appears to be a relatively common species

in eastern Brazil. As we have seen more than 35 collections of this species from Bahia, Minas Gerais, and

Parana, and it often occurs in disturbed second growth forests, it does not seem probable that Senegalia harleyi

is threatened. However, humid tropical forest is disappearing and additional data concerning the future of

Senegalia harleyi need to be obtained (IUCN 2001).

Seigler et al., New species of Senegalia from Brazil

3 mm

Fig. 2. Senegalia harleyi Se\q\er f Ebinger, & P.G. Ribeiro. A. Leaf habit with twig and petiolar glands; B. flower; C. inflorescence; D. pseudo-inflorescence;

E. leaflet adaxial surface; F. fruit. A from de Queirozetal. 7118 (HUEFS); B, C, D from Belem andMendes369 (NY); E, F from de Queirozetal. 7118 (HUEFS).

Journal of the Botanical Research Institute of Texas 8(1)

Paratypes: BRAZIL: Bahia: Mucuge, s.l., 2 Oct 2005J.G. Carvalho-Sobrinho et al. 736 (HUEFS); Estrada para Encruzilhada, ca. 1 km de Di-

visopolis, 6 Feb 2002, PA. Fiaschi et al. 993 (SPF); Ibicoara, Estrada entre Brejo de Cima e a rodovia Mucuge-Barra da Estiva, caminho para

Cascavel, 5 Feb 2003, E. Franca et al. 4320 (HUEFS); Morrao, encosta, 28 Jan 2003, F. Franca et al. 4005 (HUEFS); Mucage na estrada para

Brejo de Campos gerais (cerrado), 31 Jan 2000, A.M. Giulietti & R.M. Harley 1978 (RB); s J., 28 Jan 2010, Divisopolis, estrada para Pedra Azul,

900 m, 6 Feb 2002, M. Groppo,Jr., A.C. Marcato, P. Fiaschi & J.R. Pirani 1062 (F, HUEFS, MBM); M.L. Guedes et al. 16928 (AFCB); Barra da

Estiva, ca. 10 Km N da cidade, rod. p/Ibicoara, prox. ao Rio Preto, 2 Feb 1974, R.M. Harley 15865 (CEPEC, RB); 2 km SWofMorro do Chapeu

on Utinga road, 1000 m, 3 Mar 1977, R.M. Harley, S.J. Mayo, R.M. Storr, F.S. Santos & R.S. Pinheiro 19317 (CEPEC, NY, RB); 3 km S de Candido

Sales, BR-116, 19 Nov 1984, G. Hatschbach 47351 (BR, CEPEC, HBG, INPA, MBM, MO, MU, NY, US, RB); 21 km de Ibicoara na rod. para

Jussiape, 27 Jan 2000, J.G. Jar dim, W.W. Fhomas, S.C. deSantAna, M.V. Albves, A.C. Araujo & B.M. Forke2562 (CEPEC, NY, RB); rodovia para

Utinga, ramal para a torre Telebahia, 8 Sept 1990, H.C. Lima, S. M. de Faria, & H.S. Brito 3884 (CEPEC, RB); Serra do Gobira, 21 Jan 2005,J.

G.Nascimento, F.S. Nunes, and Milton 331 (HUEFS); Povoado de Agua Fria, 16 Feb 2002, F.S.Nunes 885 (HUEFS); Seabra, 900 m, 13 Feb 1987,

J.R. Pirani, R.M. Harley, B.L. Stannard, I. Cordeiro, C. Kameyama & A.M. Giulietti 2013 A (HUEFS, NY); Elisio Medrado, Serra dajiboia, Fazen¬

da Jequitiba, na Estrada para Monte Cruzeiro, 2 Mar 2001, L.P. Queiroz et al. 6466 (HUEFS); Capao do Correia, 2200 m, 24 Jan 2000, L.P.

Queiroz, L.F.P. Gusmao & B.M. da Silva 5635 (CEPEC, HUEFS); 10 km de Jussiape em diregao a Barra da Estiva, 1088 m, 15 Jun 2002, L.P.

Queiroz, E.R. de Souza, J.G. Jar dim, J.G. Sobrinho & B.M. daSilva 7118 (HUEFS); s.l., 12 Dec. 2004, M .F.S. Stradmann&P. Castilho 1005 (ALCB,

CEPEC). Minas Gerais: 30 km de Curral de Bentro para Aguas Vermelhas, 29 Jan 1965, R.P. Belem &J.M. Mendes 369 (NY, RB); 52 km de

Montes Claros para Pirapora, 30Jan 1965, R.P. Belem&J.M. Mendes 405 (CEPEC, NY); Biri-Biri, 23 Jan 1978, G. Hatschbach 40845 (MO, NY,

UC); Curtidor, Felisberto Caldeira, 16 Feb 1973, G. Hatschbach & Z. Ahumada 31675 (NY, TEX); Bern Querer, Cristalia, 850 m, 10 Feb 1991,

G. Hatschbach & O.S. Ribas 54089 (BR, CEPEC, HBG, MBM, MO, NY, US); Serra do Cabral. Joaquim Felicio, 16 Jan 1996, G. Hatschbach, M.

Hatschbach &J.M. Silva 64078 (NY); Chapada dos Gerais, Fazenda Santa Rita, 14 Jan 1996, G. Hatschbach, M. Hatschbach &J.M. Silva 64293

(NY); 3kmSW ofDiamantinaon road to Gouveia, 1300 m, 13 Jan 1969, H.S. Irwin, R. Reisedos Santos, R. Souza & S.F. da Fonseca 21859 (CM,

MO, NY, UB); 33 km SW of Diamantina near Gouveia, 1150 m, 19 Jan 1969, H.S. Irwin, R. Reise dos Santos, R. Souza & S.F. da Fonseca 22286

(MBM, MO, NY); 38 km NE of Francisco Sa, road to Salinas, 1000 m, 13 Feb 1969, H.S. Irwin, R. Reise dos Santos, R. Souza & S.F. da Fonseca

23243 (NY, UB, UEC); 20 km W of Montes Claros, road to Agua Boa, 1000 m, 24 Feb 1969, H.S. Irwin, R. Reise dos Santos, R. Souza & S.F. da

Fonseca 23829 (NY, UB, US); 15 km E of Diamantina, 1100 m, 20 Mar 1970, H.S. Irwin, S.F. da Fonseca, R. Souza, R. Reise dos Santos & J. Ramos

27981 (NY, UB, US); 2 km N of Sao Joao da Chapada, 1200 m, 25 Mar 1970, H.S. Irwin, S.F. da Fonseca, R. Souza, R. Reise dos Santos & J. Ramos

28314 (GH, NY, UB); Grao-Mogol, 11 Mar 1999, M.L. Kawasaki&A. Rapini 1086 (NY, SP); Coronel Morta, M. M agalhdes 15212 (NY); Km 938,

da BR-4 entre Medina e Limeira, G. Pabst & E. Pereira 8341 (R, RB); Km 938 da BR-04, entre Medina e Limeira, 16 Jan 1965, G. Pabst & E.

Pereira 9452 (ILL, R); 7 km SW de Itamarandiba, BR-120,970 m, 1 Dec 1984, B. Stannard, J.D.P. Oliveira & R.M. Harley 36251 (F, NY, RB, SPF).

Parana: Campina Grande de Sul, Ribeirao do Cedro, BR-2,18 Feb 1962, G. Hatschbach 8940 (UPCB, US); Morro Anhangava, Quatro Barras,

14 Feb 1992, Y.S. Kuniyoshi 5445 (MBM, NY).

Nearly all of the specimens of this taxon examined were originally identified as Acacia martiusiana (Steud.)

Burkart [= Senegalia martiusiana (Steud.) Seigler & Ebinger]. These taxa are similar and probably related, but

are easily separated by the structure of the petiolar gland and sessile vs. pedicellate flowers. The petiolar glands

of S. harleyi are sessile, oval to orbicular, 0.7-2.5 mm across, with the apex flattened or the margins raised to

form a cup; those of S. martiusiana, in contrast, have columnar petiolar glands that are 0.5-1.1 mm long, and an

apex that is 0.4-0.8 mm across. Further, all specimens of S. harleyi have sessile flowers whereas all specimens

of S. martiusiana have pedicellate flowers, the stalks 0.9-1.5 mm long. These two taxa occur in eastern and

southern Brazil, S. martiusiana being found in the states of Rio de Janeiro and Sao Paulo, and S. harleyi in Bahia,

Minas Gerais and Parana.

Senegalia hatschbachii Seigler, Ebinger, & P.G. Ribeiro, sp. nov. (Fig. 3). Type: BRAZIL. Minas Gerais: Manhumirim, 9

Feb 1973, G. Hatschbach&A. Ahumada 31392 (holotype: F; isotypes INPA, MBM, MO, MU, NY).

Senegalia hatschbachii Seigler, Ebinger & P.G. Ribeiro differs from other Senegalia species by leaf size (90-180 mm long), a solitary columnar

petiolar gland (1-2.5 mm), apex 0.2-0.6 mm in diameter, columnar rachis glands between the uppermost 1 to 6 pinna pairs, pinnae 13 to 30

pairs/leaf, 4-9 mm between pairs, leaflets 45 to 65 pairs/pinna, distance between leaflet pairs (0.4-0.7 mm), midvein central to subcentral;

inflorescence a globose head (9-15 mm across), ovary pubescent, stipe to 1.1 mm.

Climbing shrub or small tree to 6 m tall; bark not seen; twigs dark purplish brown to dark purple, slightly

flexuous, terete to slightly ridged, lightly puberulent to glabrous; short shoots absent; prickles light brown,

apex usually dark brown to purple, somewhat flattened, mostly recurved, woody, 1-4 x 1-6 mm at the base,

glabrous, scattered along the twig, petiole and rachis. Leaves alternate, 90-180 mm long; stipules light to usu¬

ally dark brown, linear, symmetrical, flattened, straight, herbaceous, 2-7 x 0.6-1.4 mm near the base, glabrous,

early deciduous; petiole adaxially grooved, 7-15 mm long, puberulent; petiolar gland solitary, located on the

upper half of the petiole, columnar, 1.0-2.5 mm long, glabrous, apex 0.2-0.6 across, depressed, glabrous; ra-

Seigler et al., New species of Senegalia from Brazil

Fig. 3. SenegaliahatschbachiiSe\q\er, Ebinger, & P.G. Ribeiro. A. leaf habit with petiolar nectary and twig; B. fruit; C. leaflet, adaxial surface; D. pseudo¬

inflorescence; E. Flower. A, B from Gibbs and Leitao Filho 4029 (F); C, D, E from de Mello s.n. (S).

Journal of the Botanical Research Institute of Texas 8(1)

chis adaxially grooved, 80-170 mm long, puberulent, a columnar gland 0.3-1.0 mm long, between the upper 1

to 6 pinna pairs, apex 0.2-0.5 mm across, depressed, glabrous; pinnae 13 to 30 pairs/leaf, 25-45 mm long, 4-9

mm between pinna pairs; paraphyllidia absent; petiolule 0.4-1.2 mm long; leaflets 45 to 65 pairs/pinna, oppo¬

site, 0.4-0.7 mm between leaflets, linear, 2.2-4.5 x 0.4-1.0 mm, glabrous to lightly pubescent, lateral veins not

obvious, 1 vein from the base, base oblique and obtuse on one side, margins lightly ciliate, apex obtuse, mid¬

vein central to subcentral. Inflorescence a densely 20- to 35-flowered globose head 9-15 mm across, in axil¬

lary and terminal pseudo-paniculate clusters, the main axis to 300 mm long; peduncles 3-14 x 0.3-0.5 mm

thick, puberulent; receptacle slightly enlarged; involucre a single small bract located on the upper half of the

peduncle, early deciduous; floral bracts spatulate, 0.4-0.8 mm long, puberulent, early deciduous. Flowers ses¬

sile, white to cream; calyx 5-lobed, 0.8-1.7 mm long, puberulent; corolla 5-lobed, 1.9-3.1 mm long, glabrous,

lobes one-quarter the length of the corolla; stamens 50 to 70; stamen filaments 5.5-75 mm long, distinct; an¬

ther glands absent; ovary pubescent, on a stipe to 1.1 mm long. Fruits oblong, 70-140 x 17-27 mm, straight,

flattened, not constricted between the seeds, coriaceous, lightly transversely striated, puberulent, eglandular,

dehiscent along both sutures; stipe 6-12 mm long; apex obtuse. Seeds not seen.

Distribution and ecology. —Gallery forests, disturbed wet second growth forests, and thickets from near

sea level to 1,000 m in the states of Minas Gerais, Parana, and Sao Paulo, Brazil.

Phenology. —Flowering Dec-Feb.

Local Names and Uses. —None known.

Etymology. —Named for Gert Guenther Hatschbach (1923-2013) Brazilian botanist and taxonomist; Her¬

barium Director at the Botanical Museum of Curitiba in Parana.

IUCN Red List category. —DD, data deficient. This species has a limited distribution, being known from

only southeastern Brazil. As we have seen fewer than 10 collections, it seems possible that this species pres¬

ently is threatened. Humid tropical forest is disappearing and additional data concerning the future of Senega-

lia hatschbachii need to be obtained (IUCN 2001).

Paratypes: BRAZIL: Parana: Patrimonio, 9 Mar 1915, P. Dusen s.n. (S); Ponta Grossa, 1904, P. Dusen s.n. (G); Capao Grande, 14 Apr 1909, P.

Dusen 7956 (GH, S, US); Jaguariahyva, 770 m, 1 Apr 1915, P. Dusen 16963 (G, MO, S); Orto Florestal, Maringa, 7 Dec 1965, G. Hatschbach, J.

Lindeman & H. Haas 13247 (F, MBM, NY, UPCB, US); Jaguariahyva, 740 m, 3 Jun 1914, G.Jdnsson 509a (F, G, GH, MO, S); Ponta Grossa, Vila

Velha, 800—920 m, 20 Jan 1965, L.B. Smith & R.M. Klein 14887 (FLOR, MICH, NY, R, US). Sao Paulo: Fazenda Santa Genebra, 23 Nov 1976,

P. Gibbs & H.F. Leitao Filho 4029 (F, SP, UEC); Campinas, 12 Mar 1871 ,J.C. de Mello s.n. (S); Santa Maria da Serra, 13 Dec 1976J.J. Famashiro

4183 (F, MBM, NY, UEC).

The columnar petiolar gland with a thin stalk to 2.5 mm long and small bulbous apex separates this taxon from

most other New World members of Senegalia. Quite similar to S. tucumanensis, S. hatschbachii differs in having

twigs that are dark purplish brown to dark purple (usually light brown throughout in S. tucumanensis) ; petioles

7-15 mm long (17-30 mm long in S. tucumanensis); leaflets 2.2-4.2 x 0.5-1.0 mm (5-8 x 1.1-2 mm in S.

tucumanensis); and lateral veins of the leaflets not obvious (lateral veins obvious in S. tucumanensts).

ACKNOWLEDGMENTS

The authors wish to thank several colleagues or advice concerning questions of nomenclature and general

taxonomic advice, in particular, K.N. Gandhi (GH). We thankjennifer Stratton for technical assistance and the

artist Alexa Musgrove for preparing drawings. The review comments of Joseph T. Miller and an anonymous

reviewer on an earlier draft are greatly appreciated. We wish to acknowledge support by the National Science

Foundation (NSF DEB 04-15803), the American Philosophical Society (1992) and the CNPq for financial sup¬

port to PGR, on her master’s degree in the Postgraduate Program in Botany at the Universidade Estadual de

Feira de Santana, Bahia, Brazil. We also gratefully acknowledge the assistance of the curators of the herbaria

that were visited during this study (ALCB, BHCB, CEN, CEPEC, F, FLOR, G, H, HRB, HST, HUEFS, IBGE,

IN PA, IPA, K, MBM, MBML, MO, NY, RB, R, SP, SPF, TEX, UB, UEC).

Seigler et al., New species of Senegalia from Brazil

REFERENCES

Bouchenak-Khelladi, Y., 0. Maurin, J. Hurter, & M. Van der Bank. 2010. The evolutionary history and biogeography of Mimo-

soideae (Leguminosae): An emphasis on African acacias. Molec. Phylogen. Evol. 57:495-508.

Britton, N.L. & J.N. Rose. 1928. Mimosaceae. North Amer. FI. 23:1 -194.

Gomez-Acevedo, S., L. Rico-Arce, A. Delgado-Salinas, S. MagallOn, & L.E. Eguiarte. 2010. Neotropical mutualism between Aca¬

cia and Pseudomyrmex : Phylogeny and divergence times. Molec. Phylogen. Evol. 56:393-408.

IUCN.2001. IUCN Red List categories and criteria, Version 3.1. Prepared by the IUCN Species Survival Commission. IUNC,

Gland, Switzerland and Cambridge, England.

Kyalangalilwa, B., J.S. Boatwright, B.H. Daru, O. Maurin, & M. Van der Bank 2013 Phylogenetic position and revised classifica¬

tion of Acacia s.s. (Fabaceae: Mimosoideae) in Africa, including new combinations in Vachellia and Senegalia. Bot. J.

Linnean Soc., accepted and online.

Luckow, M., J.T. Miller, DJ. Murphy, &T. Livschultz. 2003. A phylogenetic analysis of the Mimosoideae (Leguminosae) based

on chloroplast DNA sequence data. In: Klitgaard B.B. & A. Bruneau, eds. Advances in legume systematics, Part 10:

Higher level systematics. Royal Botanic Gardens Kew. Pp. 197-220.

Maslin, B.R., J.T. Miller, & D.S. Seigler. 2003a. Overview of the generic status of Acacia (Leguminosae: Mimosoideae).

Australian Syst. Bot. 16:1-18.

Maslin, B.R., A.E. Orchard, & J.G. West. 2003b. Nomenclatural and classification history of Acacia (Leguminosae: Mimosoi¬

deae), and the implications of generic subdivision. Available at: http://www.worldwidewattle.com.

Miller, J.T. & RJ. Bayer. 2003. Molecular phylogenetics of Acacia subgenera Acacia and Aculeiferum (Fabaceae: Mimosoi¬

deae), based on the chloroplast matK coding sequence and flanking frn/Cintron spacer regions. Australian Syst. Bot.

16:27-33.

Miller, J.T. & D.S. Seigler. 2012. Evolutionary and taxonomic relationships of Acacia s.l. (Leguminosae: Mimosoideae).

Australian Syst. Bot. 25:217-224.

Miller, J.T., J.W. Grimes, DJ. Murphy, R.J. Bayer, & P.Y. Ladiges. 2003. A phylogenetic analysis of the Acacieae and Ingeae (Mi¬

mosoideae: Fabaceae) based on trnK, matk, psbA-trnH, and trnL/trnF sequence data. Syst. Bot. 28:558-566.

Murphy, D.J., G.K. Brown, J.T. Miller, & P.Y. Ladiges. 2010. Molecular phylogeny of Acacia Mill. (Mimosoideae: Leguminosae):

Evidence for major clades and informal classification. Taxon 59:7-19.

Rafinesque, C.S. 1838. Sylva telluriana. Philadelphia.

Rico-Arce, A.M.L. & S. Bachman. 2006. A taxonomic revision of Acaciella. Anales Jard. Bot. Madrid. 63:189-244.

Seigler, D.S., J.E. Ebinger, & J.T. Miller. 2006a New combinations in the genus Senegalia (Fabaceae: Mimosoideae) from the

New World. Phytologia 88:38-93.

Seigler, D.S., J.E. Ebinger, & J.T. Miller. 2006b. Mariosousa, a new segregate genus from Acacia s.l. (Fabaceae, Mimosoideae)

from Central and North America. Novon 16:413-420.

Vassal, J. 1972. Apport des recherches ontogeniques et seminologiques a I'etude morphologique, taxonomique et

phylogenique du genre Acacia. Bull. Soc. d'Histoire Nat. Toulouse 108:105-247.

Wight, R. & G.A.W. Arnott. 1834. Prodromus Florae Peninsulae Indiae Orientalis 1:1-519 pp.

Journal of the Botanical Research Institute of Texas 8(1)

BOOK NOTICE

James McWilliams. 2013. The Pecan: A History of America’s Native Nut. (ISBN-13: 978-0-292-74916-0, hbk).

University of Texas Press, PO Box 7819, Austin, Texas 78713-7819, U.S.A. (Orders: www.utexaspress.com,

1-800-252-3206). $20.00,192 pp., 3 b&w photos, index, 5 V 2 " x 8 V 2 ".

From the publisher: What would Thanksgiving be without pecan pie? New Orleans without pecan pralines?

Southern cooks would have to hang up their aprons without America’s native nut, whose popularity has spread

far beyond the tree’s natural home. But as familiar as the pecan is, most people don’t know the fascinating story

of how native pecan trees fed Americans for thousands of years until the nut was “improved” a little more than

a century ago—and why that rapid domestication actually threatens the pecan’s long-term future.

In The Pecan, acclaimed writer and historian James McWilliams explores the history of America’s most

important commercial nut. He describes how essential the pecan was for Native Americans—by some calcula¬

tions, an average pecan harvest had the food value of nearly 150,000 bison. McWilliams explains that, because

of its natural edibility, abundance, and ease of harvesting, the pecan was left in its natural state longer than any

other commercial fruit or nut crop in America. Yet once the process of “improvement” began, it took less than

a century for the pecan to be almost totally domesticated. Today, more than 300 million pounds of pecans are

produced every year in the United States—and as much as half of that total might be exported to China, which

has fallen in love with America’s native nut. McWilliams also warns that, as ubiquitous as the pecan has be¬

come, it is vulnerable to a “perfect storm” of economic threats and ecological disasters that could wipe it out

within a generation. This lively history suggests why the pecan deserves to be recognized as a true American

heirloom.

Preface

Acknowledgments

Introduction: Cracking the Nut

Chapter 1. The Native Americans’ Nut

Chapter 2. “Pekan Nuttrees”: Europeans Encounter the Pecan

Chapter 3. “... the Forest into an Orchard”: Passive Cultivation on the Texas Frontier

Chapter 4. Antoine’s Graft: The Birth of the Improved Pecan, 1822-1900

Chapter 5. “To Make These Little Trees”: The Culture of Pecan Improvement, 1900-1925

Chapter 6. “Pecans for the World”: The Pecan Goes Industrial, 1920-1945

Chapter 7. “In Almost Any Recipe ... Pecans May Be Used”: American Consumers Embrace the Pecan, 1940-

1960

Chapter 8. “China Wants Our Nuts”: The Pecan Goes Global

Epilogue. The Future of Pecans

Notes

Bibliographical Essay

Index

JAMES McWILLIAMS is a historian and writer whose books include Just Food: Where Focovores Get It Wrong

and How We Can Truly Eat Responsibly and A Revolution in Eating: How the Quest for Food Shaped America. His

writing on food, agriculture, and animals has appeared in The New York Times, Harpers, The Atlantic, The Wash¬

ington Post, Slate, Forbes, Travel and Teisure, Eos Angeles Times, International Herald Tribune, The Christian Sci¬

ence Monitor, and The Texas Observer, where he has been a contributing writer since 2002. McWilliams is also a

contributor to freakonomics.com and a winner of the Hiett Prize in the Humanities.

J.Bot. Res. Inst. Texas 8(1): 70.2014

A NEW VARIETY OF PHANERA GLAUCA SUBSP. TENUIFLORA

(FABACEAE: CAESALPINIOIDEAE) FROM INDIA

Rajib Gogoi

Subir Bandyopadhyay

Botanical Survey of India

Arunachal Pradesh Regional Centre

Itanagar 791111, INDIA

rajibdzuko@gmail. com

Botanical Survey of India

Central National Herbarium

Howrah 711103, INDIA

subirbandyopadhyay@yahoo.com

ABSTRACT

Phanera glauca subsp. tenuiflora var. gandhiana, var. nov. (Fabaceae: Caesalpinioideae) is described from Arunachal Pradesh, India.

RESUMEN

Se describe Phanera glauca subsp. tenuiflora var. gandhiana, var. nov. (Fabaceae: Caesalpinioideae) de Arunachal Pradesh, India.

Some collections from Anjaw district, Arunachal Pradesh, identified to be of Bauhinia glauca (Benth.) Wall, ex

Benth. subsp. tenuiflora (Watt ex C.B. Clarke) K. Larsen & S.S. Larsen, were seen at ARUN. The collections

definitely belong to ser. Corymbosae in Bauhinia subgen. Phanera sect. Phanera subsect. Fulvae (Wunderlin et

al. 1987), but the identibcation was not correct. These collections have been described here to be of a new vari¬

ety of Phanera glauca Benth. subsp. tenuiflora (Watt ex C.B. Clarke) A. Schmitz because recent phylogenetic

studies based on DNA sequence data (Lewis & Forest 2005; Sinou et al. 2009) suggest that Bauhinia L. subgen.

Phanera (Wunderlin et al. 1987) should be recognized as a genus distinct from Bauhinia.

Phanera glauca Benth. subsp. tenuiflora (Watt ex C.B. Clarke) A. Schmitz var. gandhiana Gogoi & Bandyop.,

var. nov. (Figs. 1, 2 & 3). Type: INDIA. Arunachal Pradesh: Anjaw district, in between Changwanti and Walong, 800 m, 20

May 2011, R. Gogoi 24374 (holotype: CAT 0000025065; isotypes: ARUN, ASSAM).

Differs from Phanera glauca Benth. subsp. tenuiflora (Watt ex C.B. Clarke) A. Schmitz in having fusiform flower buds and hypanthium

shorter than their respective pedicels. In subsp. tenuiflora the flower buds are ovoid and hypanthium longer than their respective pedicels.

Lianas with tendrils, ca. 8 m in height; hairs ferruginous when dry; tendrils flattened, pubescent. Leaves 4.7-

10 x 3-9 cm, ovate or ovate-orbicular, 7-9-nerved, retuse to tapering or bifid to their length into broadly

obtuse lobes at apex, truncate or shallowly cordate at base, glabrous above, pubescent beneath, particularly on

the nerves, later glabrescent excepting the nerves; petioles 0.9-1.9 cm long, pubescent to glabrescent. Stipules

5-9 x 1 mm, linear-oblong, pubescent outside. Racemes corymbose, axillary or terminal, pubescent. Flower

buds 7-7.5 x 3 mm, fusiform, slightly curved at apex, pubescent. Hypanthium ca. 7 mm long, tubular, faintly

striate in dried specimens, pubescent. Pedicels 3-3.6 cm long, slender, pubescent. Bracts 8-9 mm long, linear-

oblong, pubescent; bracteoles 8-9 mm long, bliform, situated near the middle of pedicel. Flowers ca. 1.8 cm

across. Calyx 2-3-lobed. Petals 1-1.1 x 0.4-0.5 cm, white, narrowly to broadly obovate, obtuse at apex, veined,

glabrous inside, glabrescent outside, particularly in the median zone; claw ca. 3 mm long, glabrescent outside.

Fertile stamens 3; blaments 7-9 mm long, white, glabrous; anthers 2.5-3 mm long, purplish, ellipsoid. Re¬

duced stamens 5, ca. 2.5 mm long, with rudimentary anther at tip, swollen and connate at base; bases bright

yellow. Staminodes 2, in between stamens. Gynophore ca. 2 mm long, sparsely pubescent; ovary ca. 5 mm

long, greenish white, sparsely pubescent on the sutures at base; style 3 mm long, greenish white, glabrous;

stigma ca. 1.5 mm across, green, obliquely peltate. Pod unknown.

Distribution and ecology. —India (Arunachal Pradesh, Anjaw district), common at the place of collection

in tropical evergreen forest at an elevation of 800 m. Anjaw district is a newly created district, having been split

from Lohit district in 2004.

Flowering. —May.

J. Bot. Res. Inst. Texas 8(1): 71 - 75.2014

Journal of the Botanical Research Institute of Texas 8(1)

CAL0000025065

HOLOTYPE

BOTANICAL SURVEY OF INDIA

Arunachal Field Station, ttanagar

( A R U N )

FLORA OF... Atyfflt?....!

co,. no SM3.BA.Da,ea.Q.:.0£

Name feauhm, ?./ ,0

Locality : .

Habitat . ...Ll.-£.'§)S'£D..

Distribution.

Notes.... U3

Vern. name &

Phanera glauca Benth. subsp. tenuiflora (Watt ex C.B. Clarke) A. Schmitz var. gandhiana

Gogoi et Bandyop., var. nov.

Fig. 1. Holotype of Phanera glauca subsp. tenuiflora var. gandhiana.

Gogoi and Bandyopadhyay, A new variety of Phanera glauca subsp. tenuiflora

Fig. 2. Phanera glauca subsp. tenuiflora var. gandhiana in flower (Photo: Rajib Gogoi).

Etymology. —The variety has been named to honor Dr. K.N. Gandhi for his valuable contribution in the

held of plant nomenclature.

Chen et al. (2010) while working on the Chinese Bauhinia treated Bauhinia caterviflora H.Y. Chen, B. hu¬

pehana Craib including var. grandis Craib and B. pernervosa H.Y. Chen as synonyms of ‘ Bauhinia glauca var.

tenuiflora (Watt ex C.B. Clarke) K. Larsen & S.S. Larsen’ which were by that time were considered as subspe¬

cies or varieties of Bauhinia glauca. We are accepting here only two subspecies viz. subsp. glauca and subsp.

tenuiflora under P. glauca but at the same time do not agree with the taxonomic treatment of B. hupehana in¬

cluding var. grandis. Chen et al. (2010) in the key characters of subsp. tenuiflora stated that the hypanthium in

subsp. tenuiflora is 2.5-3 cm in length and longer than their respective pedicels. This is, however, not correct

because we have examined the type of B. hupehana (China, W. Hupeh, May 1907, E.H. Wilson 3373 K 000760713

image!) in which of the length of the hypanthium is 1.3-1.6 cm and they are shorter than the their respective

pedicels whereas in subsp. tenuiflora the hypanthium is longer than the respective pedicels. In the type of B.

hupehana Craib var. grandis (China, Western Szechuan, Tung Valley, near Mt. Wa, 500-1000 m, June 1908 &

October 1908, E.H. Wilson 3372 K 000760712 image!) the length of the hypanthium is 1.5-1.8 cm. The leaves

in both these collections are ca. Vi bifid at apex whereas in subsp. tenuiflora the leaves are V9-V5 bifid at apex.

Thus we feel that B. hupehana including var. grandis from Hupeh, Hunan and Szechuan most probably deserves

to be accepted as a variety of subsp. tenuiflora. Both these type specimens though annotated by Supee S. Larsen

as Bauhinia glauca (Wall ex Benth.) Benth. var. tenuiflora (Watt ex C.B. Clarke) K. & S.S. Larsen in 1978, the

identity was not clear to Supee S. Larsen at that time because in a later publication (Larsen & Larsen 1984) they

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 3. Phaneraglauca subsp. tenuiflora var. gandhiana in flower (Photo: Rajib Gogoi).

Table 1 . Comparative account of some morphological features between P. glauca subsp. glauca, P. glauca subsp. tenuiflora and P. glauca subsp. tenuiflora var. gandhiana

Characters

Phanera glauca subsp. glauca

Phanera glauca subsp. tenuiflora

var. gandhiana

Leaf

'A-Vii-Vs) bifid at apex

V9-V5 bifid at apex

Leaves retuse to tapering or bifid

to Vg their length at apex

Inflorescence

Short dense corymbs

Elongate corymbs

Almost like subsp. tenuiflora

Flower bud

Ovoid; glabrous or with

some pubescence at top

Ovoid; pubescent

Fusiform; pubescent

Hypanthium

1 -1.5 cm in length, shorter

than their respective pedicels,

glabrous or sparsely pubescent

2.5-32 cm in length, longer than

their respective pedicels,

pubescent

ca. 7 mm in length, shorter than

their respective pedicels,

pubescent

stated that B. hupehana is very close to subsp. tenuiflora and further studies on Chinese material are necessary

before reaching a final decision on the taxonomic status of B. hupehana.

A comparative account of some morphological features between P. glauca subsp. glauca, P. glauca subsp.

tenuiflora and P. glauca subsp. tenuiflora var. gandhiana are given in Table 1.

There has been no collection of P. glauca subsp. glauca from the Indian region (Bandyopadhyay 2001) and

so we have preferred to describe this variety to be of subsp. tenuiflora which is found in N.E. India.

Gogoi and Bandyopadhyay, A new variety of Phanera glauca subsp. tenuiflora

ACKNOWLEDGMENTS

We thank Paramjit Singh, Director, Botanical Survey of India, P. Satyanarayana, Scientist ‘D’ & HOO, APRC,

BSI and A. A. Mao, Scientist ‘E’ & HOO, ERC, BSI for the facilities. We also thank R.R Wunderlin and an anony¬

mous reviewer for their suggestions.

REFERENCES

Bandyopadhyay, S. 200E On the distribution range of Bauhinia glauca (Benth.) Benth. (Leguminosae: Caesalpinioideae) in

India. J. Bombay Nat. Hist. Soc. 98:149-150. (Errata in J. Econ.Taxon. Bot. 29:800-801.2005 publ. 2006).

Chen, T., D. X. Zhang, K. Larsen, & S.S. Larsen. 2010. Bauhinia. In: A.R. Branch, ed., Flora of China 10:6-21. Science Press,

Beijing, China and Missouri Botanical Garden Press, St. Louis, U.S.A.

Larsen, K. & S.S. Larsen. 1984. Bauhinia. ln:T. Smitinand and K. Larsen, eds., Flora of Thailand 4(1 ):4— 45. The Forest Her¬

barium, Royal Forest Department, Bangkok,Thailand.

Lewis, G.P. & F. Forest. 2005. Tribe Cercideae. In: G. Lewis, B. Schrire, B. Mackinder, and M. Lock, eds. Legumes of the World.

London: Royal Botanic Gardens, Kew. Pp. 57-67.

Sinou, C., F. Forest, G.P. Lewis, & A. Bruneau. 2009. The genus Bauhinia s.l. (Leguminosae): a phylogeny based on the plastid

trnL-trnF region. Botany 87:947-960.

Wunderlin, R., K. Larsen, & S.S. Larsen. 1987. Reorganization of the Cercideae (Fabaceae: Caesalpinioideae). Biol. Skr.

28:1-40.

Journal of the Botanical Research Institute of Texas 8(1)

BOOK NOTICE

Sula Vanderplank, Benjamin T. Wilder, and Exequiel Ezcurra. 2014. Descubriendo la Biodiversidad Terrestre

en la Region de Cabo Pulmo/Uncovering the Dryland Biodiversity of the Cabo Pulmo Region.

(ISBN-13: 978-1-889878-43-0, pbk). Botanical Research Institute of Texas Press, 1700 University Drive,

Fort Worth, Texas 76107-3400, U.S.A., and Next Generation Sonoran Desert Researchers, nextgensd.

com. (Download: http://ow.ly/yhaFW). Free pdf, 118 pp., color photos, tables, maps, 8 V 2 " x 11".

From the publisher: An international multi-disciplinary team of scientists recently published a report on the

terrestrial biodiversity of the Cabo Pulmo region in Baja California Sur, Mexico. These very lands are the site of

the proposed mega development project Cabo Dorado. The scientists demonstrate that the desert lands adjacent

to Cabo Pulmo, the singular coral reef ecosystem of the Gulf of California, harbor high levels of biodiversity,

much of which is only found in this remarkable coastal setting. Their report, Uncovering the Dryland Biodiversity

of the Cabo Pulmo Region, despite the developers’ assessment to the contrary, shows that the project is situated

in an area of extreme conservation value, the center of which is Punta Arena, an idyllic beach setting proposed

to be completely cleared to make way for 20 , 000 + hotel rooms.

The November 2013 survey, despite only a week in duration, documented 560 plants and animals (392

species of plants, 44 mammals, 29 reptiles, and 95 birds) on the land surrounding Cabo Pulmo, 400 more than

were presented in the “Manifestacion de Impacto Ambiental” (environmental impact statement) for the Cabo

Dorado project. Among the species overlooked in the MIA are 27 plants and animals on the Mexican endan¬

gered species list (NOM-059) and 83 endemic species, those found only in this region and nowhere else in the

world.

With an understanding of the biological richness of the beautiful, diverse, and coveted lands of the Cabo

Pulmo region in hand, it is hoped that conservation strategies that can balance ecological integrity and devel¬

opment pressures can be established.

J.Bot. Res. Inst. Texas 8(1): 76.2014

TAXONOMIC STATUS OF KOBRESIA CURVATA AND

KOBRESIA FRAGILIS (CYPERACEAE)

Bikash Jana and R.C. Srivastava

Botanical Survey of India

Kolkata-700 064, INDIA

bikash.janadp@rediffmail.com

rcs_ bsi@yahoo. co. in

ABSTRACT

Critical examination of the type specimens of K. curvata (isolectotype, CAT) and K.fragilis (isotype, CAT) reveals the two taxa are morpho¬

logically distinct and should be regarded as distinct species. Noltie (1993) merged K. curvata under the K.fragilis based on similarities in

inflorescence branching. Based on micro-morphotaxonomic characters, scanning electron microscopy studies and field survey of Kobresia

in Sikkim, Kobresia curvata and K.fragilis should be considered as distinct species.

RESUMEN

El examen critico de los tipos de K. curvata (isolectotipo, CAL) y K. fragilis (isotipo, CAL) revela que los dos taxa son distintos mofologica-

mente y deben tratarse como especies distintas. Noltie (1993) Incluyo K. curvata en K.fragilis basandose en similitudes de la ramificacion de

la inflorescencia. Basandonos en caracteres micro-morfotaxonomicos, estudios de microscopio electronico de barrido y observaciones de

campo de Kobresia en Sikkim, Kobresia curvata y K.fragilis deben considerarse especies distintas.

Boott (1858:2) described Carex curvata as a new species from Sikkim. Unfortunately, the name was illegitimate

when published (non Knaf 1847) (McNeill 2012; Art. 53.1). Clarke (1908) transferred C. curvata to the genus

Kobresia and thus published K curvata C.B. Clarke as a new name. Perhaps unaware of Clarke’s (1908) publica¬

tion, Kukenthal (1909) also published the same new name K curvata (Boott) Kuk., which is treated as an iso-

nym (McNeill 2012; Art. 6, Note 2).

Within the protologue of Carex curvata, Boott (1858:2) mentioned the specimen “HAB. in Himalaya Ori-

entali alpine ad Sikkim, alt. 12,000-14,000 ped. (graminosis), J.D. Hooker s.n.” Noltie (1993) lectotypihed the

name Carex curvata Boott on the basis of J.D. Hooker’s specimens collected from Tungu (Thangu) & Lachen,

localities of Sikkim. One specimen of J.D. Hooker (bearing same data as the type) deposited in CAL is very

similar to the drawing (t.5) of Boott (1858): it is an isolectotype of Carex curvata Boott.

Clarke (1903) provided the following type information for Kobresia fragilis: “Szechuen: Tongolo in Kiala

(Soulie 731). Herb. Kew.” One of the duplicate specimens (det. by C.B. Clarke as “n. sp.”) bearing the same local¬

ity and collection number is deposited in CAL and is the isotype of Kobresia fragilis. Based on this type, Kuken¬

thal (1904) published Schoenoxiphium caricinum Kuk., a superfluous illegitimate name (McNeill 2012; Art.

52.2 (a)). Subsequently, Clarke (1908) transferred K.fragilis to Schoenoxiphium and made the new combination

S. fragile.

In his study of the type specimens of Kobresia curvata and K.fragilis, Noltie (1993) concluded that the in¬

florescence branches are similar; he mentioned that, except for the curvature of the stem, the type of K curvata

is similar to the type of K.fragilis, and suggested the curvature of the stem may result from grazing and tram¬

pling. Thus, Noltie treated K curvata as a synonym of K.fragilis without mentioning any micro-morphological

characters. Srivastava (1996) treated Kobresia curvata as a separate taxon in Flora of Sikkim. During our held

survey of North & East Sikkim, we found K curvata with its typical curved stem on hilly slopes (3500-4000

m), where grazing is impossible. Subsequently, we (present authors) critically examined specimens of these

species and type specimens both. Our analysis of the micro-morphological characters revealed that similarity

in the external appearance of these species is superficial, and we conclude that the two species are distinct (as

depicted in Table 1, Fig. 3).

J. Bot. Res. Inst. Texas 8(1): 77 - 82.2014

Journal of the Botanical Research Institute of Texas 8(1)

Table 1. Comparison of K. curvata C.B. Clarke and K. fragilis U$. Clarke based on the study of the type material.

Characters

K. curvata

K. fragilis

Culm

Inflorescence

Glume

Prophyll

Racheola

Culm curved; base covered with pale brown

prominent leaf sheaths

Linear-oblong, curved; branches overlapping

Glumes of carpellate spikelet aristate or awned,

margin pale green

Oblong-ovate, yellowish brown without any red

spots; opening from apex to middle

Racheola exserted from prophyll, margin scabrid,

longer than style, binerved; nerve brown

Nut surface (SEM study) Reticulate, areole without raised, central silica body

Culm erect; base covered with grayish yellow

not prominent leaf sheaths

Linear, never curved; branches not overlapping

Glumes of carpellate spikelet short mucronate

to obtuse, margin hyaline

Utriculate, yellowish-brown in color with

prominent red spots; open only at the apex

Racheola included in the prophyll, margin

glabrous, shorter than style, binerved; nerve

green

Reticulate, areole with raised, central silica body

TAXONOMIC TREATMENT

Kobresia fragilis C.B. Clarke, J. Linn. Soc. 36:267. 1903. (Figs. 1, 3). Schoenoxiphium caricinum Kiik., Bull. Herb. Boiss.

4(1):49. 1904, nom. superfl. & illegit.; S. fragile (C.B. Clarke) C.B. Clarke, Bull. Misc. Inform. Kew, Addit. Ser. 8:67. 1908. Type:

THIBET ORIENTALI: Tongolo (Principaute de Kiala), 1893, Soulie 731 (holotype: K; isotype: CAL, ace. no. 512696!)

Perennial herbs. Culms erect, 13-31 x 0.08-0.1 cm (excluding inflorescence), trigonous, slender, smooth.

Leaves basal and sub-basal, 6.6-10 x 0.1 cm, shorter than culm; lamina bliform, involute, keeled, margin sca¬

brid; basal sheaths grayish yellow, not prominent. Inflorescence spicate, linear-oblong, 2.6-3 x 0.3-0.5 cm

(excluding awn of proximal glume), yellowish green, axis triquetrous, lateral spikes 4-7, non-overlapping.

Spikes linear, 2.5-4.5 x 1-1.5 mm, proximal spikelets carpellate, 3-5; distal spikelets staminate, 1-3. Proxi¬

mal glume with clasping base, equaling or exceeding the inflorescence, ca. 4.5 cm long (including awn); awn

up to 3-3.5 cm long, margin scabrid. Carpellate spikelets ovate to ovate-oblong, ca. 3-4 x 1 mm. Glumes of

the carpellate spikelets ovate, aristate (awned), ca. 4 x 1 mm (including awn); awn slightly scabrid; yellowish

brown, margin hyaline, nerve 1, green. Prophylls utriculate, ca. 3 x 1 mm, 2-keeled; keels scabrid; yellowish-

brown in color with prominent red spots, open only at the apex. Racheola ca. 1.5 mm, included in the pro¬

phyll, margin glabrous, shorter than style, binerved; nerves green. Gynoecium ca. 3 mm long; ovary ca. 2

mm long, oblanceolate; style ca. 1 mm long; stigmas 3. Staminate spikelets lanceolate, 2-3 x 0.5 mm, brown

in color; stamens 3. Nuts oblong, trigonous, ca. 2 mm, grayish-yellow; appearing smooth at 60x magnification,

with SEM at lOOOx magnification surface reticulate, areoles with raised, central silica body.

Flowering & Fruiting. —-July-Sept.

Distribution. —BHUTAN: Bumthang (above Gortsam) andMongar (Sengor) districts, Thimphu (Pajoding

above Ragyo, mountain E of Thimphu). INDIA: Sikkim (Deosa, Dzongri, Jamlinghang to Bikbari, Lachen,

Karponang, Kyanglasha, Mon Lapcha, Nathula, Tsomgo, Tungu). TIBET.

Specimens examined: INDIA. Sikkim. North Sikkim: Changu Lake, 3640 m, 8 Jul 1996, G.P. Sinha & D.G. Long et al. 17737 (BSHC);

Yumthang, 3520 m, 13 Jul 1996, G.P. Sinha & D.G. Long et al. 17822 (BSHC). TIBET: Tongolo (Principaute de Kiala), 1893, Soulie 731 (isotype,

CAL, ace. no. 512696!, holotype, K).

Kobresia curvata C.B. Clarke, Kew Bull. Addit. Ser. 8:68. 1908. (Figs. 2,3). Carex curvata Boott, Ill. Gen. Carex 1:2, t.5.

1858, non Knaf, 1847. Kobresia curvata (Boott) Kiik. in A. Engler, Pflanzenr. IV. 20 (Heft 38):48.1909, isonym. Type: INDIA: Sikkim,

12,000-13,000 ft,J.D. Hooker s.n. (lectotype, designated by Noltie 1993): K!, bottom right hand specimen; isolectotype: CAL!).

Perennial herbs. Culms curved, 3-4.5 x 0.08-0.1 cm (excluding inflorescence), proximally trigonous, striate.

Leaves basal, 2.2-6.1 x 0.05-0.1 cm, equaling or sometimes exceeding the culm; lamina bliform, convolute,

curved as culm, margin scabrid; basal sheath pale brown, prominent. Inflorescence spicate, linear or linear-

oblong, 1.2-2.7 x 0.25-0.4 cm (excluding awn of proximal glume), curved, yellowish-green, axis triquetrous;

lateral spikes 3-6, overlapping. Spikes oblong or slightly ovoid, 4-7 x 2-3 mm; proximal spikelets carpellate,

4-7; distal spikelets staminate, 2-4. Proximal glumes leafy, exceeding the inborescence, ca. 7 mm (including

Jana and Srivastava, Status of Kobresia curvata and K. fragilis

Fig. 1. Kobresia fragilis C.B. Clarke. A. Habit. B. Spike. C. Carpellate glume. D. Proximal glume. E. Prophyll. F. Staminate spikelet. G. Gynoecium with

racheola. H. Nut surface under SEM.

awn); awn up to 4 cm long, curved, scabrid. Carpellate spikelets ovate, ca. 2.5-3 x 1.2 mm, yellowish. Glumes

of the carpellate spikelets ovate ca. 2 x 1.5 mm, short-mucronate or obtuse, glabrous, yellowish brown, margin

pale green, nerve 1, green. Prophylls oblong-ovate, ca. 2.5-3 x 1-1.2 mm, 2-keeled; keels scabrid; yellowish

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 2. Kobresia curvata C.B. Clarke. A. Habit. B. Spike. C. Carpellate spikelet. D. Carpellate glume. E. Gynoecium with racheola. F. Staminate spikelet.

G. Nut surface under SEM.

brown, opening from apex to middle. Racheola ca. 3 mm long, exserted from prophyll, linear, as long as or

longer than prophyll, margin scabrid, binerved; nerve brown. Gynoecium 3-3.5 mm long; ovary 2 x 0.75 mm,

ovate- elliptic; style 0.5-1 mm long; stigmas 3. Staminate spikelets oblong, 2.5-3 x 1 mm, yellow in color,

margin hyaline; stamens 3. Nuts elliptic to ovoid, trigonous, ca. 1.5x1 mm, reddish brown, appearing smooth

at 60x magnification, with SEM at lOOOx magnification surface reticulate, areoles without central silica body.

Jana and Srivastava, Status of Kobresia curvata and K. fragilis

Fig. 3. A-E. Kobresiafragilis. A. Inflorescence. B. Carpellate spikelet. C. Prophyll with gynoecium. D. Gynoecium with racheola. E. Nut & nut surface under

SEM. F-J. Kobresia curvata. F. Inflorescence. G. Carpellate spikelet. H. Prophyll with gynoecium. I. Gynoecium with racheola. J. Nut surface under SEM.

Journal of the Botanical Research Institute of Texas 8(1)

Flowering & Fruiting. —-July-Aug.

Distribution .—Eastern Himalaya; INDIA: Sikkim (Changu, Karponang, Katao, Kupup, Kyangosla,

Lachen, Lachung, Nathula, Thangu, Tsomgo), 3600-4000 m.

Specimens examined: INDIA. SIKKIM: Himalaya, Cho-le-la, Jul 1879, G. King s.n. (CAL). East Sikkim: 6 km above from Zuluk, 3230 m,

N27°15.431'E 088°47.418', 27Jul 2012, BikashJana 53134 (CAL); Kupup, 4115 m, N 27°18.908' E 088°50.158', 28Jul 2012, BikashJana53169

(CAL). North Sikkim: Katao, Kala Pahar, 28 Jul 1989, N.R. Mondal 10133 (BSHC); Lachen, 2550 m, 07 Jun 1999, D. M aiy 21327 (BSHC, 2

specimens); Thangu, 17 Aug 1989, R.C.Srivastava 12289 (BSHC); Thangu, 17 Aug 1989, R.C. Srivastava 10206 (BSHC). WEST BENGAL:

Darjeeling district, A.B. Chowdhury 53 (CAL).

ACKNOWLEDGMENTS

We are grateful to Paramjit Singh, Director, Botanical Survey of India; P. Lakshminarasimhan, Central Na¬

tional Herbarium, BSI; and A. Bandopadhyay, Head of the Department of Botany, Burdwan University for the

facilities. We greatly appreciate two anonymous reviewers for review comments improving the paper.

RELERENCES

Boon, F.M.B. 1858. Illustrations of the genus Carex, 1:2, t.5. London, UK.

Clarke, C.B. 1903. [Chinese] Cyperaceae. J. Linn. Soc., Bot. 36:202-309.

Clarke, C.B. 1908. New genera and species of Cyperaceae. Kew. Bull. 8:1-196.

Kokenthal, G. 1904. Cariceae novae vel minus cognitae. Bull. Herb. Boiss. 4(1 ):49-60.

Kokenthal, G. 1909. Cobresia Willd. In: H.G.A. Engler, ed. Das Pflanzenreich. Heft 38. Engelmann-Cramer, Weinheim, Ger¬

many. Pp. 40-48.

McNeill, J. & others, eds. 2012. International code of nomenclature for algae, fungi, and plants (Melbourne Code): ad¬

opted by the Eighteenth International Botanical Congress Melbourne, Australia, July 2011. Regnum Veg. 154.

Noltie, H.J. 1993. Notes relating to the flora of Bhutan: XIX. Kobresia (Cyperaceae) in Edinburgh J. Bot. 50:39-50.

Noltie, HJ. & Z. Shuren. 2010. Kobresia Willd. In: Wu, Z.Y. and P.H. Raven, ed. Flora of China. 23:269-285. Science Press,

Beijing & Missouri Botanical Garden Press, Louis, MO, U.S.A.

Srivastava, R.C. 1996. Cyperaceae. In: P.K. Hazra and D.M. Verma, eds. Flora of Sikkim. Botanical Survey of India, Calcutta.

Pp. 198-237.

FIRST VALID PLACE OF PUBLICATION OF DUCHESNEA INDICA

(ROSACEAE: POTENTILLEAE)

James L. Reveal

L.H. Bailey Hortorium

Department of Plant Biology

Cornell University

Ithaca, New York 14853-4301, US.A.

jlr326@cornell.edu

Barbara Ertter

University and Jepson Herbaria

1001 Valley Life Sciences Building #2465

University of California

Berkeley, California 94720-2465, U.S.A.

ertter @berkeley. edu

ABSTRACT

The combination Duchesnea indica traditionally has been attributed to Focke (1888), who based his name on Andrews’s (1807) Fragaria in-

dica. When the genus name was published by Smith (1811), he proposed a superfluous and illegitimate species name, D.fragiformis, so it was

felt that D. indica should have been established before 1888. A search of the literature using on-line resources found that indeed the combina¬

tion was proposed by Teschemacher in 1835. A formal lectotypification of F. indica is proposed.

Keywords: Fragaria indica, Duchesnea fragiformis, Potentilla indica, nomenclature

RESUMEN

Ta combinacion Duchesnea indica ha sido atribuida tradicionalmente a Focke (1888) que baso su nombre en el de Andrews (1807) Fragaria

indica. Cuando se publico el nombre del genero por Smith (1811), el propuso un nombre de especie superfluo e ilegitimo, D.fragiformis, asi

resulto que D. indica habria sido establecido antes de 1888. Una busqueda bibliografica usando recursos on-line encontro que realmente la

combinacion fue propuesta por Teschemacher en 1835. Se propone una lectotipificacion formal de F. indica.

One of the world’s most widely distributed weedy species in the rose family is the mock- or false-strawberry,

Duchesnea indica. A member of the rose family (Rosaceae Juss. trib. Potentilleae Sweet), this native of south-

central Asia is now found on every continent except (for the moment at least) Antarctica. According to Li et al.

(2003), D. indica is one of two species in the genus, the other being D. chrysantha (Zoll. & Moritzal) Miq. of

eastern Asia.

The species was originally described as Fragaria indica by Andrews (1807: ad tab 479) who knew the spe¬

cies as “an ornamental plant, but is in no other respect estimable” being grown in an English garden. Although

superficially resembling the true strawberries ( Fragaria L.) in its fleshy red fruit and ternate leaves, it differs in

having yellow petals, doubly toothed leaflets, and protruding achenes on an essentially dry, almost tasteless

accessory fruit. The genus Duchesnea was subsequently established for the plant by Smith (1811: 372) who

proposed D.fragiformis Smith (1811: 373) as a superfluous and illegitimate name for the species. Since Focke

(1888: 33), most treatments have adopted D. indica or followed Wolf (1904: 661) in treating the species as Po¬

tentilla indica (Andrews) T. Wolf based on the obvious similarities (other than the strawberry-like fruit) with

P. reptans L. and other members of the sect. Potentilla.

Wolf’s proposed relationship has been confirmed by recent molecular analyses (Erikkson et al. 1998;

Lundberg et al. 2009; Dobes & Paule, 2010), in which Potentilla and Fragaria fall into two separate clades, with

Duchesnea indica nested within Potentilla. At the same time, however, Ertter (2007) continues to find the argu¬

ments against paraphyly per se unconvincing and accordingly opts to retain Duchesnea (as well as Ivesia, FLorke-

lia, and Horkeliella) as distinct genera, even though this results in a paraphyletic Potentilla s.s.

When we prepared our treatment of the genus for the Flora of North America Project in 2006, we simply

followed long-established tradition and credited the name Duchesnea indica to Focke and, basically, gave it no

more thought. After all, this bibliographic reference was widely used and no one questioned it. Upon receipt of

page proofs, however, Reveal decided to take advantage of the proliferation of web-based search tools to see if

the combination D. indica might have been validly published prior to 1888. The discovery of an abundance of

J. Bot. Res. Inst. Texas 8(1): 83 - 84.2014

Journal of the Botanical Research Institute of Texas 8(1)

overlooked names in the horticultural literature (Reveal 2012) suggested that this was probably the most likely

place to find the name effectively published. Therefore, using the resources of Biodiversity Heritage Library

(http://www.biodiversitylibrary.org/) and Google search for books (http://books.google.com/), Reveal easily

discovered that, indeed, D. indica was proposed by Teschemacher (1835) long before 1888; without these re¬

sources, finding that place of publication would have been difficult. Thus:

Duchesnea indica (Andrews) Teschem., Hort. Reg. Gard. Mag. 1:460, 1 Dec 1835. Type: as “Duchesnia,” based on

Fragaria indica Andr., Bot. Repos. 7: ad t. 479, Oct 1807 (lectotype, designated here: [icon] Bot. Repos. 7:t. 479.1807).

In proposing the basionym, Andrews knew the plant from a single individual that flowered in the garden of

Charles Francis Greville (1749-1809), probably at his home near Paddington Green in London where he had

glasshouses for his more exotic species, although it is possible the plant was seen at Warwick Castle where

Greville lived late in his life. Andrews stated that the plant was “native of the north-east parts of Bengal” and

this is most likely where Greville obtained his seed having been gathered by one of his many correspondents

or friends as he was also an avid collector of ancient artworks and especially minerals. Andrews left no known

herbarium but did contribute specimens to the herbarium of Sir James Edward Smith. Unfortunately, no origi¬

nal material of Duchesnea indica has been found at the Linnean Society in London. Nonetheless, it seems ap¬

propriate that the name be lectotypihed to assure application of the name and accordingly we have selected the

one original element we are aware of at this point in time.

ACKNOWLEDGMENTS

We wish to thank Luc Brouillet and the staff at the Flora of the North America Project for their patience as we

dealt with this problem at the last minute. Kanchi N. Gandhi (GH), John McNeill (E), and Gerry Moore (USDA)

are acknowledged for their comments and review of the manuscript.

REFERENCES

Andrews, H.C. 1807. Fragaria indica. Indian strawberry. Bot. Repos. 7: ad t. 479.

Dobes, C. & J. Paule. 2010. A comprehensive chloroplast DNA-based phylogeny of the genus Potentilla (Rosaceae): Impli¬

cations for its geographic origin, phylogeography and generic circumscription. Molec. Phylogen. Evol. 56:156-175.

Erikkson,T., M. J. Donoghue, & M.S. Hibbs. 1998. Phylogenetic analysis of Potentilla using DNA sequences of nuclear ribo-

somal internal transcribed spacers (ITS), and implications for the classification of Rosoideae (Rosaceae). PI. Syst. Evol.

211:155-179.

Ertter, B. 2007. Generic realignments in tribe Potentilleae and revision of Drymocallis (Rosoideae: Rosaceae) in North

America. J. Bot. Res. Inst.Texas 1:31-46.

Focke, W.O. 1888. Rosaceae. In: A. Engler & K.A.E. Prantl, eds. Die naturlichen Pflanzenfamilien nebst ihren Gattungen

und wichtigeren Arten insbesondere den Nutzpflanzen, bearbeitet unter Mitwirkung zahlreicher hervorragender

Fachgelehrten.Teil III, Abteilung 3. Wilhelm Engelmann, Leipzig. Pp. 1 -48.

Li, C.-L., H. Ikeda, & H. Ohba. 2003. Duchesnea, pp. 338-339. In: Z.-Y. Wu, P.H. Raven, and D. Hong, eds. Flora of China:

Pittosporaceae through Connaraceae. Volume 9. Science Press, Beijing, & Missouri Botanical Garden Press, St. Louis.

Lundberg, M., M.Topel, B. Eriksen, J. Nylander, &T. Eriksson. 2009. Allopolyploidy in Fragariinae (Rosaceae): Comparing four

DNA sequence regions, with comments on classification. Molec. Phylogen. Evol. 51:269-280.

Reveal, J.L. 2012. A divulgation of ignored or forgotten binomials. Phytoneuron 2012-28:1 -64.

Smith, J.E. 1811. A description of Duchesnea fragiformis, constituting a new genus of the natural order of Senticosae of

Linnaeus, Rosaceae of Jussieu. Trans. Linn. Soc. London 10:371-374.

Teschemacher, J.E. 1835. On artificial rock work. Hort. Reg. Gard. Mag. 1:456-460.

Wolf, T. 1904. "In derTracht der Potentilla reptans nicht unahnlich." In: Ascherson, P.F.A. & K.O.R.P.P. Graebner. Syn. Mit-

teleur. FI. Wilhelm Engelmann, Leipzig. 6(1 ):661.

ERRATA

FOUR NEW ANNUAL SPECIES OF EUPHORBIA SECTION TITHYMALUS

(EUPHORBIACEAE) FROM NORTH AMERICA

Mark H. Mayfield

Herbarium, Division of Biology

Kansas State University

Manhattan, Kansas, USA. 66506-4901, US.A.

markherb@ksu.edu

The following are provided as corrections/additions to Mayfield (2013).

The Perry (1943) reference information was omitted from the references section and it is provided herein.

In Figure 2, the length of the scale bar is 1 mm.

In the key to species on pages 646 and 647, Euphorbia ouachitana was inadvertantly omitted from the key. It

should have been included in a couplet with E. tetrapora under the first part of couplet 13 on page 647. Below,

couplet 13 is revised and a new couplet 14 is added to distinguish E. tetrapora and E. ouachitana. The entire key

is given again for ease of reference and use.

1. Plants biennial; seeds > 1.8 mm long; primary ray bracts about as wide as long or wider, generally suborbicular to

broadly ovate; plants occurring outside of Texas.

2. Seeds rotundly ovoid, strongly pitted, with distinct, round depressions on a generally flat surface; plants of the east¬

ern United States and southern Ontario, Canada (northeastern Oklahoma north to Wisconsin, east to Pennsylvania,

and south to Florida and Mississippi)_ E. commutata

2. Seeds oblong-ellipsoid, weakly dimpled, with shallow, irregularly shaped depressions bordered by weak reticulating

ridges, surface nowhere flat; plants of the western United States (southern California to northwestern Oregon, also

local in southern Colorado and northern New Mexico_ E. crenulata

1. Plants annual; seeds mostly < 1.7 mm long; primary ray bracts usually longer than wide; plants often occurring in Texas

and elsewhere.

3. Stem leaves generally erect-ascending at maturity, if lax, the blades less than 3 mm wide at the widest point.

4. Raylet leaves at least 1.5 x longer than wide, the apices acute.

5. Seeds rotund; stems mostly strict, erect or virgate_ E. austrotexana var. carrii

5. Seeds oblong; stems laxly ascending.

6. Seeds with troughlike and rounded pits, the surface not pimpled_ E. peplidion

6. Seeds without pits, the surface pimpled_ E. exigua

4. Raylet leaves about as long as wide, or wider than long.

7. Leaves linear to linear-oblanceolate_ E. austrotexana var. austrotexana

7. Leaves spatulate to oblanceolate.

8. Seeds uniformly covered with deep, well-defined rounded pits on both surfaces_ E. longicruris

8. Seeds with 4 (or 5) shallow ventral pits, and 4 rows of indistinct pits on the dorsal surface_ E. tetrapora

3. Stem leaves generally divergent or lax at maturity, and over 4 mm wide at the widest point.

9. Stem leaves with petioles or elongated petiole-like bases.

10. Raylet leaves apically obtuse, subdeltate; plants 8-18 cm tall_ E. nesomii

10. Raylet leaves apically rotund, subreniform to scarcely deltate; plants 15-35 cm tall.

11. Capsules with longitudinal wings along the ridges; seeds 1.3-1.5 mm long, bearing two longitudinal sulcae

on the ventral facet_ E. peplus

11. Capsules without longitudinal wings along the ridges; seeds 1.8-2.0 mm long, smooth to pitted, but not

sulcate on the ventral facet.

12. Seeds smooth (lOx), lacking reticulating ridges_ E. helleri

12. Seeds not smooth (1 Ox), with reticulating ridges_ E. roemeriana

9. Stem leaves sessile, or attenuate to a brief, petiole-like base.

J. Bot. Res. Inst. Texas 8(1): 85 - 86.2014

Journal of the Botanical Research Institute of Texas 8(1)

13. Seeds with a few, well-separated pits in vertical rows.

14. Seeds maturing dull grayish brown, the ventral faces with 4 weakly defined pits; plants of the western Gulf

Coastal Plain in Oklahoma, Louisiana and Texas_ E. tetrapora

14. Seeds maturing lustrous reddish-brown, the ventral faces with 6 strongly defined pits; plants of the Interior

Highlands in Arkansas, southwestern Missouri, and central Tennessee_ E. ouachitana

13. Seeds with numerous crowded, deep pits not clearly in vertical rows; plants only occurring on granite outcrops

within the Piedmont Province of Georgia_ E. georgiana

REFERENCES

Mayfield, M.H. 2013. Four new annual species of Euphorbia section Tithymalus (Euphorbiaceae) from North America. J.

Bot. Res. Inst.Texas 7(2):633-647.

Perry, B.A. 1943. Chromosome numbers and phylogenetic relationships in the Euphorbiaceae. Amer. J. Bot. 30:527-543.

THE GENUS ECHINACEA (ASTERACEAE):

FLORAL, STEM, AND PETIOLE MORPHOLOGY

Harold W. Keller

Botanical Research Institute of Texas

1700 University Drive

Fort Worth, Texas 76107, US.A.

haroldkeller@hotmail.com

ABSTRACT

The genus Echinacea (Asteraceae) has importance economically, medicinally, and ornamentally. Endemic to North America, distribution is

centered in the states of Arkansas, Kansas, Missouri, and Oklahoma. Native Americans of the central Great Plains used Echinacea as a

highly prized medicinal plant panacea. This anatomical study is based on R.T. McGregor’s taxonomic treatment of the genus Echinacea that

included 11 taxa: E. angustifolia var. angustifolia, E. angustifolia var. strigosa, E. atrorubens, E. laevigata, E. pallida, E. paradoxa var. neglecta, E.

paradoxa var. paradoxa, E. purpurea, E. sanguinea, E simulata, and E. tennesseensis. Anatomy of Echinacea tennesseensis was not included be¬

cause live plants were not available. Plants were collected at the height of anthesis from the experimental gardens at the University of Kansas.

Samples were prepared for microtome and free-hand sectioning and staining. Macromorphology and microanatomy are described here, and

photomicrographs illustrate the adaxial epidermal cells of ray ligules. Tissue map line drawings illustrate the pattern and distribution of

stem trichomes, epidermal cells, cortex, vascular bundles, and pith. Measurements were included for stem diameters, epidermis, collen-

chyma, parenchyma, xylem vessels, sclerenchyma fibers, xylem and phloem vascularization, protoxylem points, and location and number

of secretory canals for each Echinacea taxon. Sclerenchyma fibers (sclerotic cells with a black phytomelanin substance) are located in the

pith tissue of all the varieties of E. angustifolia. Tissue maps and photomicrographs illustrate petiole transections and the presence of

brachysclerids (stone cells) in E. paradoxa var. neglecta which were found nowhere else in this study. Plants resulting from crossings and

introgression between E. atrorubens and E. angustifolia had many intermediate characteristics and were called “race intermedia .” This name

has no nomenclatural standing but the plants were found to have unique ray ligule adaxial epidermal cells. These multicelluar structures

consist of an enlarged basal cell with a neck and a catenuliform series of one, two, or three discrete pyramidal cells that have not been de¬

scribed for any member of the Asteraceae or other flowering plant. A key to Echinacea taxa that includes the distinctive micromorphology of

ray ligule adaxial epidermal cells is presented. A discussion of the structure and function of ray ligule microanatomy is included as this re¬

lates to insect pollinators. Questions still remain concerning the constancy of anatomical characters over a broad range of habitats based on

statistically sampled populations.

Key Words: dichotomous key, Echinacea (coneflower), endangered species, ligule adaxial epidermal cells, macro-and microanatomy, scle¬

rotic and stone cells

RESUMEN

El genero Echinacea (Asteraceae) tiene importancia economica, medicinal y ornamentalmente. Endemica a Norteamerica, se centra en los

estados de Arkansas, Kansas, Missouri y Oklahoma. Los americanos nativos de las Grandes Planicies (Llanuras Centrales), usaron Echina¬

cea como una planta muy altamente apreciada como panacea medicinal. Este estudio anatomico se basa en el tratado taxonomico del genero

de R.L. McGregor que incluyo 11 taxa: E. angustifolia, E. angustifolia var. strigosa, E. atrorubens, E. laevigata, E. pallida, E. paradoxa var. ne¬

glecta, E. paradoxa var. paradoxa, E. purpurea, E. sanguinea, E. simulata y E. tenneesseensis. Las plantas se colectaron durante el pico de la flo-

racion en los Jardines Experimentales de la Universidad de Kansas para prepararlas para el microtomo, seccionado a mano y tincion. En este

estudio describimos la macromorfologia y la microanatomia, y las fotomicrografias ilustran las celulas epidermicas adaxiales de las flores

radiales. Los dibujos de las celulas epidermicas ilustran los patrones y distribucion de los tricomas del tallo, celulas epidermicas, cortex,

conjuntos vasculares y medula, con medidas de los tipos de celulas que incluyen diametro de los tallos, epidermis, colenquima, parenquima,

vasos del xilema, fibras de esclerenquima, vascularizacion del xilema y liber, puntos del protoxilema y la ubicacion y numero de los canales

secretores para cada taxon de Echinacea. Las fibras del esclerenquima (con una sustancia fitomelanica negra) se localizan en el tejido medu-

lar de E. angustifolia var. angustifolia, E. angustifolia “raza intermedia” y E. angustifolia var. strigosa. Los mapas de tejido y fotomicrografias

ilustran los cortes transversales de los peciolos y la presencia de braquiscleridas (celulas petreas) en E. paradoxa var. neglecta las cuales no se

encontraron en ninguna otra parte en este estudio. Otro taxon llamado E. angustifolia var. angustifolia “raza intermedia” no tiene posicion

nomenclatural pero tambien se describio y se encontro que tiene celulas epidermicas adaxiales unicas en las flores radiales. Estas celulas

multicelulares consisten de una celula basal agrandada con un cuello y una serie cateluniforme de 1, 2 o 3 celulas piramidales discretas que

no se han descrito para ningun miembro de las Asteraceae o alguna otra planta con flores. Se presenta una clave para los taxa de Echinacea

que incluye la micromorfologia distintiva de las celulas epidermicas adaxiales de las hemiligulas. La estructura y funcion de las celulas epi-

J. Bot. Res. Inst. Texas 8(1): 87 -126.2014

Journal of the Botanical Research Institute of Texas 8(1)

dermicas adaxiales de las hemilfgulas tiene implicaciones ecologicas importantes que se relacionan con los insectos polinizadores. Echina¬

cea tennesseensis no se incluyo en este estudio anatomico pero se discutio como un taxon de especial interes como una especie en peligro de

extincion.

INTRODUCTION

Historical Overview

This study was conducted during a two year period (1960-1962) as part of the partial fulfillment of the require¬

ments for the degree of Master of Arts at the University of Kansas (KU) under the direction of R.L. McGregor

(Keller 1962). The author shortly thereafter served as an officer in the United States Army Medical Service

Corps and was not able to publish his thesis at that time. Several years later McGregor (1968) published his

monograph of Echinacea which represented 15 years of held studies throughout the range of the various taxa.

This included growing transplants representing most taxa in the KU experimental garden and greenhouse for

a period of eight years. Several thousand plants were started from seed both in the garden and greenhouse

representing all possible crosses and backcrosses. Chromosome counts were made from over 2100 held collec¬

tions of buds, root tips from transplants, and root tips from germinating seeds (McGregor 1968).

The Comparative Anatomy section of the paper by McGregor (1968) included a summation of taxonomi-

cally signihcant characters for certain taxa associated with stem, petiole, and adaxial ray ligule epidermal cell

anatomy. A key to identify Echinacea taxa was constructed based on the combination of these characters. De¬

tailed descriptions and illustrations of micromorphology for stems, petioles, and floral anatomy for each Echi¬

nacea taxon were not included.

Keller (1962) included a general key based on a combination of anatomical characters that included the

number, size, and location of stem secretory canals, overall stem size and tissue patterns of xylem, presence or

absence of sclerotic cells in pith tissue, presence or absence of trichomes, and differences in ray ligule adaxial

epidermal cells. Additional keys were also included based solely on the marginal ray floret anatomy and stem

secretory system. The 95 pages of text, 56 line drawings, and 34 photomicrographs of microanatomy in the

thesis are included here in part to document a more complete descriptive anatomy of Echinacea taxa (http://hdl.

handle.net/1808/11669).

In a study of root anatomy of Echinacea angustifolia var. angustifolia, E. pallida, and E. purpurea, Mistrlkova

and Vaverkova (2007) noted that “a description of the microscopic characteristics of a cross-section of the

aerial parts of the Echinacea plants is not available.” The question posed here was to determine differences

in stem, petiole, and ray floret microanatomy that could separate and identify taxa as well as contribute to a

better understanding of aerial plant microanatomy and morphology of the genus Echinacea. The purpose of

this study was to emphasize both aspects and, where feasible, devise a key using a combination of macro- and

microanatomical key characters.

Review of Past Literature

Echinacea Moench, Methodus, 591. 1794, is a member of the Asteraceae, a family that exhibits a wide range of

distribution and habit. It is probably the largest family of flowering plants with an estimated 1535 genera and

23,000 species (Bremer et al. 1994) and numbers given as 1353 genera and 23,000 species in Judd et al. (2008).

This family is comprised mostly of herbs, although sometimes shrubby and more rarely woody habits occur.

Echinacea herein is treated as a herbaceous, perennial prairie forb, however, toward the end of the growing

season a small amount of secondary growth may develop, not unlike many herbaceous plants. Echinacea may

also serve as forage for grazing animals and is often an indicator of undisturbed prairies. It is one of many

showy wild flowers that are such a familiar part of the landscape of Kansas. An ever-increasing demand for

horticultural varieties led to the ‘King’, a pink variety, and ‘White Lustre’ among others, both developed from E.

purpurea (McGregor 1968).

The genus Echinacea is native and endemic to central United States of America, extending into Canada,

but is unknown from Mexico. Most taxa are more or less restricted to the states of Arkansas, Kansas, Nebraska,

Oklahoma, Minnesota, Missouri, North Dakota, South Dakota, and Texas with scattered collections from

Keller, Floral, stem, and petiole morphology of Echinacea

Colorado, Georgia, Illinois, Iowa, Kentucky, Louisiana, Massachusetts, Montana, New Mexico, North Caroli¬

na, South Carolina, Tennessee, and Virginia. Highest population density and species diversity occurs in the

states of Arkansas, Kansas, Missouri and Oklahoma with Missouri heading the list (Richter 2013).

The Economic Importance section of McGregor (1968) discusses the use of E. angustifolia and E. pallida

roots as a source of drug extracts for various medical ailments but was limited to early anglo settlers of the

Great Plains region. Uses by Native Americans were not included in his discussion of Economic Importance;

however, an excellent review is available in Kindscher (1989). Native Americans used the prairie forb genus

Echinacea as a general panacea, but some of its reputed medicinal properties were: the root relieved the pain of

toothache; the juice soothed burns and aided healing; and plants placed in steam baths acted as vaporizers

cooling the body from heat discomfort. It even was thought to be beneficial in the treatment of mumps and

distemper in horses. Even today an alcoholic tincture of Echinacea root is used for the healing of wounds and

cure of sore throat (Stevens 1961).

Probably the best source of indigenous names and anglo folk use is detailed for E. angustifolia by Kind¬

scher (1991). The indigenous tribes that had names and uses included the Apache, Cheyenne, Comanche,

Crow, Dakota, Hidatsa, Kickapoo, Kiowa, Lakota, Mesquakie, Omaha, Pawnee, Ponca, Potawatomi, and Win¬

nebago. As the most widely used medicinal plant of the plains, as noted by Kindscher (1991), the collective uses

based on the aforementioned tribes included: the dried seed head was used to comb or brush hair; as a mush¬

room medicine because its seed head was similar in shape to a mushroom; root tissues for an eyewash, coughs,

sore throat, or to stimulate saliva flow; applications for snake bites (rattlesnakes) and other venomous bites,

stings and poisonings; a remedy for hydrophobia, wounds, tonsillitis, stomach ache, pain in bowels; root teas

brewed for sore mouths or gums, rheumatism, arthritis, and measles. Coneflower roots were mixed with fun¬

gal puffballs (Gasteromycetes, Eycoperdon spp.) spores and skunk oil to treat boils. This list of broad medicinal

uses of Echinacea gives a better idea of the past history of possible potential medical efficacy and future herbal

uses in modern human cultures. More recent papers document the enhanced immune-stimulatory and anti¬

inflammatory activity of Echinacea and the impact of overharvesting, especially in areas in the state of Kansas

(Kindscher 1989; Price and Kindscher 2007; Kindscher et al. 2008; Axentiev et al. 2010; Upton 2010).

Metcalf and Chalk (1950) in their classic two-volume work, listed numerous anatomical features that

characterize the Asteraceae, including secretory canals, lacticiferous canals, glandular and nonglandular

hairs, anomalous secondary thickening, and medullary and cortical bundles. Herbaceous stems among mem¬

bers of the family usually exhibit a ring of collateral vascular bundles, each accompanied in the pericyclic re¬

gion by large strands of fibers, often forming distinct “bundle caps.” Even though Echinacea is cited by Soler-

eder (1908) and Metcalf and Chalk (1950), no detailed investigation of anatomical characteristics, either on a

comparative or taxonomic basis, has been published. Since that time additional data were published that ex¬

tends our knowledge of ray floret anatomy in the Asteraceae.

Papers published that relate to Echinacea ray ligule anatomical microcharacters (Table 1, Baagoe 1977a;

1977b) evaluate three epidermal types, including the helianthoid type, consisting of papillose and nearly iso-

diametric cells that come closest morphologically to Echinacea. The illustration of these papillose adaxial epi¬

dermal cells (see Baagoe 1977a, plate lb) shows a three-dimensional SEM view of Rudbeckia sp. and plate 2 c of

an Aster sp. ligule in optical plane cross section showing a light photomicrograph with median wall thickening,

and plate 2e of a Rudbeckia ligule in cross section showing septa. A survey of the 111 genera and 275 species in

the Asteraceae did not mention Echinacea by name or illustrate any of the taxa (Baagoe 1977a). Another paper

by Baagoe (1977b) evaluates ray ligule microcharacters in the Asteraceae as they relate to taxonomic differ¬

ences that separate taxa and applies them as characters in a key to tribes, sub-tribes, and genera. Echinacea is

not mentioned in that paper nor was the Echinacea monograph of McGregor (1968) cited as a source of ana¬

tomical data.

Description of the adaxial epidermal cells of the Sub-tribe Helianthinae that have papillose cells are des¬

ignated as the helianthoid type. These cells represented by Rudbeckia hirta and R. speciosa (see Baagoe 1977b,

fig. 5a SEM) have the largest cells (length-width ratio) and thicker outer cell walls. In other words these cells

Journal of the Botanical Research Institute of Texas 8(1)

are more elongate vertically with a narrow neck and much wider at the base. None of these cells illustrated or

described are multicellular.

Dome shaped adaxial epidermal cells with papillae were shown by Baagoe (1980) using SEM for certain

members of the Lactuceae. Examples (Table 1) were represented by Calycoseris wrightii (fig. IB) with hooked

papillae pointing toward the distal end of the ligule, Hieracium saxifragum (fig. 2A) with more upright papillae,

and Rafinesquia neomexicana (fig. II) with papillose ligule surfaces distally as well as several other examples

(Baagoe 1980). Unfortunately some of the SEMs show distorted and collapsed adaxial epidermal cells caused

by shrinkage after preservation as herbarium specimens. Critical point drying techniques were not used to

preserve cell shapes.

The anatomy of ray florets in the Asteraceae is surprisingly understudied and examples must come from

other unrelated floral taxa. References and examples are summarized in Table 1. These aforementioned com¬

bined images provide a historical context for the different sizes and shapes of ray floret adaxial epidermal cells

observed here in Echinacea taxa.

An exhaustive review of taxonomic treatments and phylogenetic papers including Echinacea is beyond

the scope of this paper; therefore, the Echinacea taxa discussed by McGregor (1968), the taxonomic treatment

by Urbatsch et al. (2006) and by Flagel et al. (2008) will be followed because the anatomical data in Keller

(1962) is associated with those names.

MATERIALS AND METHODS

Collections of Echinacea used in this study were made at the University of Kansas Experimental Gardens,

Lawrence, Kansas, with some additional held collections. A complete list of the taxa with held notes is in¬

cluded under Collections in the thesis of Keller (1962). Voucher specimens were deposited in the R.L. Mc¬

Gregor Herbarium (KANU). Collections were cited by Keller (1962) as the source of live specimens used for

this anatomical study of Echinacea taxa. All plants studied were collected during anthesis from 19 Jun to 22

Junl961. Only specihc portions of the plants were selected: flower head, stem, and node with attached leaf.

Voucher specimens are listed below for the source populations from which the KU garden collections

were grown or from which collections were made.

E. angustifolia var. angustifolia: Kansas, Comanche Co., 17 mi E of Coldwater, prairie hillside, 18 Junl957, E.

Eathrop 3827.

E. angustifolia var. angustifolia race intermedia: Kansas, Mitchell Co., 7 mi N of Hunter, rocky prairie hill¬

side, 22 Jun 1961, B. Menhusen s.n.

E. angustifolia var. strigosa: Oklahoma, Murray Co., 1 mi N of Sulphur, 29 May 1960, R.E. McGregor 15607.

E. atrorubens: Kansas, Douglas Co., 1 mi W and V 2 mi S of KU Experimental Gardens, 22 Jun 1961, H.W. Keller

s.n.

E. laevigata: North Carolina, Durham Co., near Durham, grown from seed sent by Bloomquist 5 (leg. ign. s.n.)

E. pallida: Kansas, Chautauqua Co., 3 mi E and 3 mi N of Sedan, 19 Aug 1959, R.E. McGregor 15042.

E. paradoxa var. neglecta: Oklahoma, Murray Co., rocky prairie hillside common in area at Platt National

Park, 7 Jun 1959, R.E. McGregor 14323.

E. paradoxa var. paradoxa: Missouri, Barry Co., rocky hillside, 3 mi SE jet. Hwy 112 and F, Roaring River

State Park, 12 Jun 1959, R.E. McGregor 14367.

E. purpurea: Arkansas, Baxter Co., wooded hillside, 2 V 2 mi SE of Mountain Home, 6 Augl959, R.E. McGregor

14961.

E. sanguinea: Texas, Angelina Co., sandy open bank at edge of pine forest, 2.6 mi S of Lufkin on Hwy 89, 12

May 1960, R.E. McGregor 15557.

E. simulata: Missouri, Oregon Co., roadside opening in oak-hickory woods, 2.2 mi N of Greer, Clark National

Forest, 3 Jun 1960, V Harms 321.

Live plant samples were based on not more than five plants and at least five transections made of five stems,

petioles, and ray florets. Due to a small sample size taken at one point in time, a statistical comparison of popu¬

lation samples was not attempted. Consequently, the anatomical keys constructed herein are practical only

within certain limitations with the caveat that recognizing qualitative differences in description of cell types

previously unknown merits special consideration.

Keller, Floral, stem, and petiole morphology of Echinacea

Table 1. References describing flower petal and ray ligule adaxial epidermal cells. SEM=scanning electron micrograph; LP=light photomicrograph; LD = line drawing.

Family: Tribe

Taxon

Cell shape

Source & illustration type

Asteraceae: Cichorieae

Calycoseris wrightii

Hooked papillae

Baagoe (1980, Fig. 1B); SEM

Asteraceae: Cichorieae

Hierocium soxifrogum

Upright papillae

Baagoe (1980, Fig, 2A); SEM

Asteraceae: Cichorieae

Rafinesquia neomexicana

Hooked papillae

Baagoe (1980, Fig, 21); SEM

Asteraceae: Heliantheae

Aster sp.

Papillose, dome-shaped

Baagoe (1977a, Plates 2c, 2e); LP

Asteraceae: Heliantheae

Helianthus annuus

Conical

Whitney et al. (2011); SEM

Asteraceae: Heliantheae

Rudbeckio sp.

Papillose, conical

Baagoe (1977a, Plate 1 b); SEM

Gesneriaceae

Saintpaulia ionantha

Rounded papillae

Endress (1994, Fig. 5.10-2); SEM

Lentibulariaceae

Pinguiculo vulgaris

Nipple-like

Eames & MacDaniels (1947, Fig. 169C); LD

Rosaceae

Amelanchier laevis

Hemispherical

Eames & MacDaniels (1947, Fig. 169A); LD

Rosaceae

Rosa sp.

Dome-shaped, columnar

Esau (1960, Fig. 20.1-A); LP

Verbenaceae

Lantana camara

Long, conical papillae

Endress (1994, Fig. 5.10-3); SEM

Violaceae

Viola x wittrockiana cultivar

Sharply pointed conical,

Weryszko-Chmielewska &Sulborska

some with cuticular striation

(2012, Figs. 1C, 5);LP,SEM

Organs such as stem and leaf were collected in the same morphological position on specimens represent¬

ing each taxon to ensure validity of comparison. This was accomplished by determining a point midway be¬

tween ground level and flower attachment which served as the source of material for the stem anatomy pre¬

sented here. The first recognizable leaf (not to be confused with the reduced upper leaf) borne on the stem

below the capitulum was selected for study purposes. This corresponds in some cases to a position just above

leaves borne in somewhat of a rosette-like fashion, especially in the shorter species.

The leaves have either sheathing bases or, in the narrow-leafed species, are distinctly petiolate. Petiole

anatomy was based on samples taken 0.6 cm from the point of departure on the stem axis for sheathing leaf

bases and halfway between the stem axis and the leaf base for distinctly petiolate leaves. Floral parts such as ray

ligules were sectioned at the approximate midpoint gauged by the overall length of the particular structure.

After selection, the designated materials were placed in vials containing formalin-propiono-alcohol (Johansen

1940, p. 42).

Two methods of preparation were employed, each possessing certain merits. Tissue prepared by the free¬

hand sectioning method gave excellent preservation of detail, especially in the secretory canal system. How¬

ever, producing sections uniform in thickness, less than 10 pm and truly parallel to the plane of cutting re¬

quires practice, skill, and patience. Furthermore, sections should be standardized to a certain thickness to

study structural properties on a comparative basis. The microtome method has drawbacks caused by the previ¬

ous dehydrative treatment before sectioning that tends to collapse and distort the thin-walled epithelial cells

that surround the canal cavity. The graded alcohol series and staining in Coplin jars and eventual embedding

in paraffin was a long, time-consuming process prior to sectioning with the microtome.

Free-hand transections were made by orienting live plant material in elderberry pith and then thinly slic¬

ing sections with a single edge razor blade. Transections of ray ligules were mounted directly into glycerol and

photographed with a compound microscope. Stem transections were stained either with safranin and fast

green or phloroglucinol in 18 percent HCL. After passing the stained sections through a graded alcohol series,

they were permanently mounted in picolyte. Temporary phloroglucinol mounts were made by placing sections

in glycerol. Moreover, the true nature of the cells (size, shape, and wall thickness) was greatly enhanced when

fresh material could be cut, stained, and passed directly into glycerol without overuse of a harsh dehydrating

agent such as alcohol. This holds especially for the collenchyma which is rich in water and tends to undergo a

noticeable shrinkage when subjected to dehydration. Shorter preparation time for free-hand sections facilitat¬

ed more rapid analysis.

Sections prepared by the paraffin method had cortical cells more compact, thinner walled, and intercel¬

lular spaces completely occluded. This method more readily demonstrated the vascularization of the stem axis

because whole, intact sections could be obtained. Tissue prepared for the microtome was handled according to

Journal of the Botanical Research Institute of Texas 8(1)

Johansen (1940, p.130) using tertiary butyl alcohol as the dehydrating agent. Following impregnation and

embedding in paraffin, the material was sectioned with a Spencer Rotary Microtome at blade settings of 10 pm

and 15 pm. Difficulty in sectioning was encountered where extensive sclerihcation occurred throughout the

pith region, exhibiting a tough, woody consistency.

Preliminary staining followed the schedule outlined by Johansen (1940, p. 80-82) with slight modifica¬

tions to fit each taxon. Staining time was altered when safranin stain was used with the counterstain fast green

to get brilliant color differentiation. The presence of a carbohydrate compound, presumably starch, was de¬

tected by applying Lugol’s solution (I 2 KI-potassium iodide) to freshly cut sections. Inulin reported in the litera¬

ture as commonly found in roots, and sometimes in stems, gave a negative test. Phloroglucinol indicated the

extent and relative degree of lignihcation among the taxa.

Macerations of stem material were prepared by slicing the stem longitudinally into small slivers to in¬

crease the disintegrating power and lessen the time required to free the cemented cells. Usually thirty minutes

was sufficient time to freely suspend the parenchymatous and collenchymatous cells, but vascular elements

often were teased apart with a pin probe. The macerating fluid was made according to the formula prescribed

byjeffrey in Johansen (1940, p. 104). The range and average measurements of pericyclic fibers, vessels, collen-

chyma, and parenchyma cells were tabulated from a minimum of 30 individual cells of each.

Thin strips of epidermal cells were peeled from the stem. These were then projected and drawn in surface

view noting the frequency of stomata and trichomes, if any, the cuticle characteristics, epidermal patterns, and

cell dimensions. Length in surface view refers to cell elongation and orientation in a vertical plane and width

to that measurement in a horizontal plane.

Measurements made of the ray ligule adaxial epidermal cells have a vertical orientation so that length

refers to the upward, vertical extension exaggerated by their papillose condition. The basal width is in a hori¬

zontal plane parallel with the surface of the ray florets. Trichomes and hairs are terms that also have been used

for stem and leaf pubescence and also for ray florets. However, usage of ray florets and ray ligule adaxial epider¬

mal cells will be followed here to be consistent with McGregor (1968).

The Bausch & Lomb TRI-SIMPLEX Micro-projector was used to project images of tissue sections upon

white paper conveniently placed on a tabletop and then outlining the image to make tissue maps. This appara¬

tus was equipped with a tri-objective revolving nosepiece that gave magnifications of 2.7x, 5x, and 12x. A spe¬

cial attachment, the 5x Huygenian eyepiece made possible even greater magnifications. Drawings of stem and

petiole transections were made at a magnification of 40x. The different tissue systems are represented thusly:

the xylem with associated fibers is vertically lined; the phloem is blank or white; the bundle caps are black¬

ened; and stippling between vascular bundles indicates lignihcation.

Light photomicrographs were taken with a compound microscope with either a low power (lOx) or high

power (43x) objective lens with a lOx eyepiece at a total magnification of lOOx or 430x. Scale bars were not

used; magnifications were calculated using cell measurements. Photographic images were recorded using

Kodak 35mm Panatomic X black and white negative him. The 35mm negatives were scanned at 6300 dpi using

an Imacon Precision II him scanner. The 16-bit gray-scale images were inverted to positive tif images using

Photoshop CS6 software.

GENERAL ANATOMICAL BODY PLAN OL ECHINACEA

Echinacea is a genus of herbaceous perennials usually occurring in undisturbed prairie or glade ecosystems.

Most taxa usually arise from a taproot (only E. purpurea has a rhizome and hbrous root system) as a single erect

stem, are most often unbranched with hairy to smooth leaves, basal and cauline, alternate, petiolate, and ter¬

minating in a single flower head (capitulum). Ray florets (8 to 21) represented by strap-shaped ray ligules are

sterile, forming a showy head of dark purple to rose or pale pink, yellow, or white, either spreading, drooping,

or rehexed ray florets with either two or three notched tips. The central cone is made up of perfect, fertile, disc

flowers that are inconspicuous because of the surrounding colorful paleae (chaffy bracts) with orange to red¬

dish purple ends that create the showy spiny cone.

Keller, Floral, stem, and petiole morphology of Echinacea

The generic name is derived from the Greek word echinos meaning hedgehog that refers to the prominent

spiny cone of disc florets that eventually matures into a bristly seed head. The morphological characters used

to describe the general habit in monographic treatments (McGregor 1968; Urbatsch et al. 2006; Yatskievych

2006) do not mention anatomical characters and that includes stem diameters.

Stem anatomy of plants includes the spatial arrangement of cell types represented by parenchyma, col-

lenchyma, sclerenchyma, and primary and secondary xylem and phloem (vascular tissue). The presence or

absence, location, size, and number of external cells (trichomes) or internal secretory cells that include cavi¬

ties, canals, resiniferous or mucilaginous cells, crystals (cystoliths, druse, raphides), and laticifers serve as di¬

agnostic characters useful in taxonomic work (Eames & MacDaniels 1947; Esau 1958, 1960). Specific regions

of the stem for example, epidermis, cortex, pericycle, endodermis, vascular bundles, and pith, may show

modification due to thickness of cell walls, size, maturation, and taxon-specific differences. The stem tissue is

typical of herbaceous perennial dicots with a pith region that occupies about 75 percent of the central core of

the plant and collateral vascular bundles, forming a ring nearer the periphery and usually separated by inter¬

fascicular parenchyma or sclerenchyma. A descriptive anatomical evaluation of these morphological features

has not been published for the stem, petiole, and ray ligule adaxial epidermal cells for Echinacea taxa.

Petiole anatomy offers additional sources of comparisons between Echinacea taxa as this relates to the

number of major or minor collateral vascular bundles, petiole shape in transectional view, presence or absence

of lacunae, and presence or absence of secretory cells. In Echinacea the petiole is supplied by three major col¬

lateral vascular bundles, a manifestation of departing foliar traces from the stem. Due to different methods of

fusion, division, or twisting of the leaf traces, the number of vascular bundles traversing the cortex may or may

not be the same as the number that enter the leaf. Minor vascular bundles are associated in different numbers

with the major vascular bundles (Eames & MacDaniels 1947).

The arrangement of vascular bundles in the petiole is usually constant for a given species and often for

families (three for the Asteraceae). In addition petiole shape can be used as a taxonomic character. Transverse¬

ly cut petioles can be recognized by shapes, for example, horseshoe-shaped, V-shaped, and cylindrical-shaped.

Moreover, secretory canals, universally present in the genus, differ in size, number and position in the petiole.

Anatomically the petiole contains the same tissues as the stem: epidermis, collenchyma in varying amounts,

and vascular bundles with associated fibrous sheaths.

The main emphasis of this study was the anatomy of the ray florets that are the showy parts of the com¬

posite flower. Petals are leaf-like in form, but they differ histologically in various ways from the typical leaf.

Generally they show some resemblance in their internal structure to mesophytic leaves, although often lacking

differentiated palisade and spongy parenchyma tissue. They consist of ground parenchyma (often called me-

sophyll), a greatly reduced and branched vascular system, and epidermal layers on the adaxial (upper) and

abaxial (lower) surfaces. The vascular supply (here termed veins and veinlets) usually bifurcates more notice¬

ably at the ray ligule tips. Thick-walled supporting tissue often is found surrounding each veinlet. Further¬

more, the vascular tissue often consists of several large veins and a system of smaller veinlets.

Perhaps the most striking anatomical feature of the ray ligules is the peculiar adaxial epidermal cells that

bulge outward, and are modified into various sizes and shapes. The ray ligule adaxial epidermal surface is usu¬

ally modified into the conical-papillate type as in Erysimum cheiri (Weston & Pyke 1999, light photomicro¬

graphs figure 1B, C and figure 2 A SEM shows an adaxial conical-papillate type epidermis with a striated waxy

cuticle epidermis) whereas, the cells of the abaxial epidermis in figure 2B, SEM shows lenticular cells with

stomata. Echinacea closely parallels then the anatomy of most petals as all of the features mentioned previously

are exemplified in its ray florets.

In some flowering plants both floral epidermal surfaces are papillose, but in Echinacea only the adaxial

ligule epidermal surface exhibits this micromorphology. The inner tangential wall is often slightly convex. The

outer wall, by comparison, is often more or less convex or papillose. However, in Viola and Nasturtium, for ex¬

ample, these cells are modified and bear one or more capitate or cone-shaped papillae (not illustrated in Esau

1958, p. 538). Similarly, in E. angustifolia “race intermedia ” the adaxial epidermis of ray ligules partly consists

Journal of the Botanical Research Institute of Texas 8(1)

of cells enlarged basally appearing more bulbous, then capped by one or more (not more than three observed)

catenulate, pyramidal-shaped cells. In many flowering plants the anticlinal walls appear either straight, wavy,

or may bear internal ridges. The undulation and ridging varies widely in degree of expression in different spe¬

cies. Indeed, the epidermis is less simple than its foliar counterpart.

Due to the weak-walled complex nature of the ray ligule adaxial epidermis, a dovetailed arrangement

seems to permit the greatest mechanical support. The functional importance of these highly modified cells is

open to conjecture, but apparently they form a layer mechanically stronger than one of simpler form. Further¬

more, in some plants epidermal anticlinal walls along the veins and at the base of the petal are usually straight,

even if wavy elsewhere. In some cases variability in wall structure gives an assortment of shapes. In Echinacea ,

however, the size and shape of ray ligule adaxial epidermal cells remained somewhat constant for each taxon

at least in the same capitulum. Stomata were not observed on the ray ligule adaxial epidermal cell surface sug¬

gesting these modifications in cell shape play a different functional role perhaps the attraction of insect pollina¬

tors. The ray ligule abaxial epidermis resembles the typical epidermal tissue having stomata, trichomes, and a

heavy cuticular covering and internally often with dense contents consisting of chromoplasts and small par¬

ticles. Chromoplasts occur in the cell sap with an array of colors as can be seen in the drooping or spreading ray

florets.

RESULTS

Description of Ray Florets, Stem, and Petiole Anatomy in Echinacea Taxa

Microanatomy of the following Echinacea taxa is summarized in Table 2 (ray florets), Table 3 (stems), and Table

4 (petioles). Distribution maps (Figs. 1, 2) were created using ArcMap 10.2 showing counties in the U.S.A.

shaded gray, indicating that the taxon occurs there (McGregor 1968; Kartesz 2013; USDA-PLANTS 2013).

1. Echinacea angustifolia DC. var. angustifolia Prodr. 5:554. 1836. (Figs. 1A; 3A, B, E, I, J; 9B; 10A). narrow¬

leaved PURPLE CONEFLOWER, BLACK SAMPSON ECHINACEA, KANSAS SNAKEROOT.

This taxon includes a complex of taxa referred to by various names assigned by McGregor (1968) and followed

during the course of this study (Keller 1962). Its distribution occurs throughout the high plains and drier prai¬

rie areas, barrens, and rocky to sandy-clay soils of Texas, Oklahoma, Kansas, and north to Canada. The west¬

ernmost extension of this taxon includes New Mexico, Colorado, Wyoming, and Montana (Fig. 1A). The

strictly low habit (20-70 cm tall), mostly unbranched, moderately to densely hairy, tuberculate-hirsute to tu-

berculate-hispid stem with ray florets usually purplish to pink, rarely white, yellow pollen, and diploid chro¬

mosome number of 2n = 22 characterize this taxon (Fig. 3A, B).

Ray ligule adaxial epidermal cell microanatomy has a length-width ratio that results in a dome shape with

some cells slightly pinched in part way near the top (Fig. 3E). Secretory chambers of four to six epithelial cells

are found on the abaxial side of the vascular traces (Table 2).

Stem microanatomy features nonglandular trichomes that thickly cover the stem. Epidermal stem cells

have a wide range in length which tends to give an irregular pattern (Fig. 9B). Sclerotic cells with thickened

walls occurring throughout the pith are easily seen in free hand transections but secretory canals are lacking

(Fig. 31, J). Stem interfascicular regions are sclerihed but show no secondary growth (Fig. 10A). The cortex is

largely made up of collenchyma tissue. The secretory system consists of 26 canals restricted to the cortex.

Comparatively this taxon has one of the smaller stems in the genus at ~2 mm in diameter, including a pith di¬

ameter of ~1 mm (Fig. 10A; Table 3). The E. angustifolia complex of taxa all have sclerenchyma cells and lack

secretory canals in the pith which differ from all other Echinacea taxa.

The petiole is smaller and somewhat V-shaped with sides not steeply inclined (Table 4). Outside the lat¬

eral vascular bundles an abrupt delimitation of fundamental tissue occurs, continuing as photosynthetic foliar

tissue. The canal system is greatly reduced in number; thus only one canal 40 pm was seen beside the medial

vascular bundle. The minimum of three vascular bundles traverse the petiole.

2. Echinacea angustifolia var. angustifolia “race intermedia” (Figs. 1C; 3C, D, E; 10B).

Keller, Floral, stem, and petiole morphology of Echinacea

Table 2. Ray ligule micromorphology based on transections. Position of secretory chambers are relative to vascular traces.

Adaxial epidermal cells

Mean

Ligule

No.

Secretory chamber

height

basal

thickness

vascular

position & diameter

Taxon

(pm)

width (pm)

Overall shape

(pm)

traces

(pm)

£. angustifolia

Dome with slight pinching at top

170

abaxial (38)

var. angustifolia

E. angustifolia

100*

45-54

Tapered conical; pyramidal

286

abaxial (31-40)

var. angustifolia

"race intermedia"

catenulate

E. angustifolia var. strigosa

Conical with rounded corners

265

few to lacking

E. atrorubens

Broadened base with "nipplelike"

apex

233

abaxial (55)

£ laevigata

Dome nearly isodiametric

184

lacking

£. pallida

Tapering upward to dome or conical

235

£. paradoxa var. neglecta

Variable tapering upward

abaxial & adaxial

gradually

to a rounded apex

(42-81)

£. paradoxa var. paradoxa

Dome with parallel sides

312

abaxial & adaxial

(42-70)

£. purpurea

Wide ball-like base tapers to

rounded apex

■■

£. sanguinea

115

Elongate "necked" papilla

267

lacking

£. simulata

105

Sharply pointed conical apex

270

tapering upward from wide base

^multicellular (1-3 cells); 138-308 pm in total height

This name “race intermedia ” was used by Keller (1962) but was not validly published. McGregor (1968) noted

that crosses between E. atrorubens and E. angustifolia var. angustifolia produced populations of plants as a re¬

sult of introgression that had many intermediate characteristics, hence the name intermedia.

The ray ligule adaxial epidermal cells observed were of two kinds (unicellular and multicellular) and de¬

scribed in detail and illustrated using light photomicrographs (Keller 1962). Unicellular cells have a tapered

conical shape (Fig. 3D). These specialized multicellular structures appear in tiers mounted on an enlarged

basal cell (Fig. 3C, D). The multicellular structures show a wider range of size: one pyramidal cell (basal cell 90

pm, terminal cell 48 pm, overall length 138 pm (Fig. 3C); two pyramidal cells (basal cell 70 pm, next cell 39

pm, terminal cell 41 pm, overall length 150 pm (Fig. 3C), and three pyramidal cells (overall length 308 pm),

(Fig. 3C; Table 2). Ray ligule adaxial epidermal multicellular structures consist of an enlarged basal cell with a

neck and a catenuliform series of one, two, or three discrete pyramidal cells (Fig. 3C, D). The multicellular

epidermal cells are far fewer in number and scattered among the predominately papillose cells (Fig. 3C). These

multicellular, adaxial epidermal cells appear unique and have not been described for any member of the

Asteraceae or flowering plant and were not observed in any other Echinacea taxa (Table 2). Ray floret secretory

chambers are present usually on the abaxial side of each veinlet. A ring of five elliptical epithelial cells sur¬

round the chamber lumen.

The stem is heavily covered with trichomes that are shorter, 0.5-1.5 mm, and stouter than in other taxa.

Either two or three septa occur with conspicuous lenticular bumps in the trichome wall. A thick cuticle (9 pm)

covers the outer tangential wall and cutinization is evident in the inner tangential wall. Prominent short, spike-

like projections mark the surface of the cuticle. In surface view the epidermal cells appear rectangular and

oblique walled. Stem diameter is -2.6 mm, including a pith diameter of ~1.3 mm with sclerenchyma cells inter¬

spersed throughout. Elliptical pits densely occur in the walls. In transection dark streaks between adjacent

sclerotic cells are the result of a black substance (probably phytomelanin seen in the roots) that occludes

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 1 . Distribution maps. A. Echinacea angustifolia var. angustifolia. B. E. angustifolia var. strigosa. C. E. angustifolia var.angustifolia "race intermedia."

D. E. atrorubens. E. E. laevigata. F. E. pallida. Maps were recreated from Kartesz (2013).

Keller, Floral, stem, and petiole morphology of Echinacea

Fig. 2. Distribution maps. A. Echinaceaparadoxa var. neglecta. B. E. paradoxa var. paradoxa. C. £ purpurea. D. £ sanguinea. E. £ simulata. F. £ tennes-

seensis. Maps were recreated from Kartesz (2013).

Journal of the Botanical Research Institute of Texas 8(1)

Keller, Floral, stem, and petiole morphology of Echinacea

angular intercellular spaces (Table 3). A secretory canal system is absent in the pith but present in the cortex

opposite the vascular bundle caps and interfascicular region. Five to seven celled epithelial rings surround

these smaller canal cavities. Many vessels are tailed with oblique perforations. The vascular bundles are sur¬

rounded by a relatively large amount of fibrous tissue. Several vascular bundles seem to have undergone fusion

undoubtedly caused by this fibrous tissue (Fig. 10B).

The petiole shows an abaxial convexity which tends to be horizontally flattened on the adaxial surface. A fea¬

ture readily recognized is the large amount of collenchymatous supporting tissue occupying a position between

the vascular bundles and abaxial epidermis. Three major vascular bundles are present in transection (Table 4)

3. Echinacea angustifolia DC.var. strigosa McGregor, Trans. Kans. Acad. Sci. 70:366-370. 1968. (Figs. IB;

3F, G, H; 9C; 10C; 11B) . STRIGOSE CONEFLOWER

McGregor (1968) recognized populations in the Arbuckle Mountains of Oklahoma, extending as far north as

Cowley County, Kansas, and as far south as extreme north central Texas as variety strigosa (Fig. IB). Stems

30-60 cm in height, are frequently branched, flexuous and covered with strigose-hirsute trichomes. Variety

strigosa hybridizes with variety angustifolia and E. atrorubens yielding morphologically intermediate popula¬

tions which in some cases are tetraploids.

The ray ligule adaxial epidermal cells are slightly modified into various shapes. Generally these cells tend

to be conical with round corners. The outer tangential wall is slightly drawn out into a papilla. Note the lattice-

work arrangement of the mesophyll tissue and air spaces between cells (Fig. 3F, G). Trichomes are present on

the lower surface of the ray floret. In Echinacea the ground (mesophyllous) tissue is homogeneous and simple

in structure. The thin-walled mesophyllous cells have a central cavity with radiating interconnected arms.

These cells are loosely arranged into a meshwork of lacunose tissue (Fig. 3G). In transection the mesophyllous

cell cavity is cylindrical and elongated in a horizontal plane (running parallel with the vascular system). These

mesophyllous cells have outgrowths (arms) that become septate at a point of juncture (Table 2).

Nonglandular trichomes thickly cover the stem. In surface view stem epidermal cells consist of relatively

small rectangular cells (Fig. 9C). Secretory canals are only present in the cortex. Sclerenchyma cells occur in

the pith (Fig. 3H). Unsclerihed cells range in size from 38 to 88 pm (65 pm). Stem diameter is -2.6 mm, includ¬

ing a pith diameter of ~1.6 mm (Fig. 10C). The discrete vascular bundles are widely separated by an interfas¬

cicular region (Table 3).

This smaller petiole tends to have the outer margins swinging upward slightly. The “wings” show a leaf-

like anatomy made up largely of spongy photosynthetic tissue. Only three vascular bundles supply the petiole.

Secretory canals are relatively small, 30 pm, and are sometimes absent from lateral vascular bundles (Table 4).

Fig. 3. A,B. Echinacea angustifolia var. angustifolia. C, D. E. angustifolia var. angustifolia "race intermedia" (free hand transections). Kansas: Mitchell

County. Ray ligule showing adaxial epidermal cells of two types (unicellular and multicellular). A. Kansas: Trego County, Cedar Bluff Reservoir, on rocky

ledge of limestone cliffs. This site has shallow soil where roots find permanent moisture several feet below, 17 Jun 2010. Note the whitish ray flowers

reflexed vertically downward surrounding stem. B. Kansas: Riley County, Fort Riley Military Reservation on rocky, upland, tallgrass prairie growing in

shallow, calcareous soil, 30 Jun 2003. Note shorter and downwardly reflexed pinkish ray florets with two or three notched ends and insect lower right

(order Coleoptera, family Cerambycidae, subfamily Lepturinae (flower longhorn beetles), Typocerus octonotatus), crawling over the cone of sharply

pointed paleae. C. Lower magnification showing section of ray ligule with specialized upper epidermis and more typical leaf-like lower epidermis (x53).

Close-up insets upper left showing one pyramidal cell and lower right three pyramidal cells. D. Close-up of single pyramidal cell atop enlarged bulbous

basal cell (x232). E. E. angustifolia var. angustifolia. Kansas: Comanche County. Dome-shaped adaxial ray ligule epidermal cells (x200). F. E. angustifolia

var. strigosa. Oklahoma: Murray County. Note modified nearly conical adaxial epidermal ray ligule cells in contrast to the isodiametric shape of abaxial

cells more typical of leaf epidermis (x62). G. E. angustifolia var. strigosa. Oklahoma: Murray County. Ray ligule adaxial epidermal cells showing conical

shape with rounded corners and lattice-work arrangement of the mesophyll tissue with air spaces between cells (x200). H. E. angustifolia var. strigosa.

Oklahoma: Murray Country. Stem microtome transection showing most of pith region with sclerotic cells. Note the black phytomelanin substance

deposited in the angles (intercellular spaces) between cells (x70). I. E. angustifolia var. angustifolia. Kansas: Comanche County. Stem microtome

transection showing thinner walled pith parenchyma cells lacking phytomelanin and the thicker walled sclerotic cells with discrete phytomelanin black

deposits (xl 77). J. E. angustifolia var. angustifolia. Kansas: Comanche County. Stem oblique free hand transection showing thicker sclerotic pith cells

with variable wall thickness with some triangular intercellular spaces free of black phytomelanin and some with heavy deposits filling spaces (x292).

Photo credits: Quinn Long (A), Craig Freeman (B).

Table 3. Stem micromorphology based on transections.

100

Journal of the Botanical Research Institute of Texas 8(1)

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Keller, Floral, stem, and petiole morphology of Echinacea

101

Table 4. Petiole micromorphology based on transections.

Taxon

Shape outline

Shape description

No.

vascular

bundles

Secretory

canals

Canal

diam.

(pm)

Notable features

£. angustifolia

var. ongusti folio

—. r m i j

> Somewhat v-shaped

3 through¬

out

Few (only 1

observed)

Sides not steeply

inclined

E. ongustifolio

j More flattened

Medial

(38)

Collenchyma

var. angustifolia

"race intermedia"

A ® f

% /

adaxially; abaxially

convex

patches between

vascular bundles

and abaxial

epidermis

E. ongustifolio

var. strigoso

l(fV

0 x 7

) Outer margins

swinging upward to

become "wings"

Small

(30)

"Wings" largely

made up of spongy

mesophyll

E. otrorubens

fL A

V \CT|SV

Rounded to horse

shoe with convex

depression

9 (6 minor)

Paired,

adjacent

to each

vascular

bundle

3 large lacunae

running length of

petiole from axil to

leaf base

E. loevigoto

) V

x n

_ .

% /

Horseshoe with

deep adaxial

convexity

Poorly

developed,

medial

(42)

Leaves with

sheathing bases

account for

lengthy "wings"

E. porodoxo

var. neglecto

\«\

j Crescent/lunate/

bow

7 (4 minor)

Relatively

large,

64-84

Brachysclerids

(stone cells) scat¬

^— 79 J

a e

numbering

tered throughout

fundamental tissue

E. porodoxo

\ 0 __

? Thickest with nearly

5 (2 minor)

Paired,

Chlorenchymatous

var. paradoxa

@ j

% /

® /

flat adaxial surface

and exaggerated

abaxial curvature

numerous

pockets toward

abaxial margin

£. purpurea

/"' 9 /

0 J

> Thickened through

medial sector with

gradual flaring and

ascending at ends

5 (2 minor)

Prominent,

adjacent to

medial

vascular

bundle

Scattered

chlorenchymatous

tissue creates

abaxial bulge

£ sanguinea

i ®

(

\ @ .-

\ & \

/ ®

_- a i

Horseshoe nearer

] stem with ends

' folded together,

unfolding near

leaf base

7 (4 minor)

One on

each side

of abaxial

bundle cap

Smallest

overall size

£. simulata

Lateral margins arch

upward, more or less

lunate in outline

7 (4 minor)

Few, poorly

developed

Fan-shaped medial

vascular bundle

with wide

fibrous cap

102

Journal of the Botanical Research Institute of Texas 8(1)

Keller, Floral, stem, and petiole morphology of Echinacea

103

4. Echinacea atrorubens (Nutt.) Nutt., Trans. Amer. Philos. Soc., n. ser. 7:354. 1840. (Figs. ID; 4A, B, C, D;

9D; 10D; 11D, E; 12A) . TOPEKA PURPLE CONEFLOWER.

Plants 50-100 cm tall, mostly unbranched, glabrous below more strigose above, basal leaves petiolate, with the

leaf blade oblong-lanceolate. This taxon is found in a narrow band from Houston, Texas, to Ardmore, Okla¬

homa, northward to Topeka, Kansas (Fig. ID) (McGregor 1968). It occurs on dry limestone or sandstone out¬

crops and prairies and is distinguished from E. paradoxa var. paradox a and E. paradoxa var. neglecta by its dark

purple to dark red and sharply reflexed ray florets that curve inward to a point where they touch the stem.

Disc florets have the typical Asteraceae floral arrangement. Microtome transections near the base show

the style, five stamen filaments, surrounded by a corolla tube (Fig. 4A). Another disc floret transection nearer

the top shows the mature five bilocular, united anthers with stained pollen grains that surround the two-

parted style. The gamopetalous corolla encloses these reproductive structures (Fig. 4B). Transections were

made in June at the height of floral anthesis.

Ray ligule adaxial epidermal cells separate this taxon from all other Echinacea taxa. The distinctive

squatty shapes feature an adaxial epidermal cell with a broadened base and a short “nipple-like” papilla (Fig.

4C, D). Vascular traces are accompanied by very few abaxial secretory chambers (Table 2).

Stem trichomes are sparsely present and often wanting on portions of the stem. Paradermal sections were

peeled easily unlike many other species. The epidermal pattern consisted of rectangular, nearly isodiametric

cells, with mostly straight end walls (Fig. 9D). Some walls are slanted so these areas may be more irregular.

Cortical tissue consists of an outer collenchymatous zone intergrading into an inner parenchymatous region

that extends from 350 to 400 pm. The phloem zone ranges from 42 to 56 pm in radial extent. The pith has pa¬

renchyma cells 92 pm in length (68 pm) and widths range from 41 to 110 pm (78 pm). The secretory system

consists of canals present in both the pith and cortex (Fig. 11D, E). Moreover, an area of thin-walled accessory

tissue is arranged above canals, surrounding canals, and not infrequently in the same morphological positions

of the canals. Epithelial rings contain four to eight rectangular or diamond shaped cells (Fig. 11 D, E). Tangen¬

tial divisions occur halfway around the canal to give a partial double ring. Usually a single well-defined canal

is located opposite the protoxylem points and other canals originate opposite the interfascicular region. The

stem diameter of ~4 mm includes a pith diameter of ~3 mm (Fig.lOD). The stellar pattern has numerous dis¬

crete vascular bundles with prominent bundle caps separated by narrow interfascicular regions (Table 3).

Sclerihed intervening cells between vascular bundles and secondary xylem is lacking as shown in the tissue

map (Fig. 10D).

Petiole outline is more or less rounded to horseshoe shaped except for a convex depression on the abaxial

side. Each major bundle is associated with two resin canals located one on each side adjacent to the abaxial fi¬

brous cap. The canals that accompany the central vascular bundle are relatively large, 40 pm, with five epithe¬

lial cells. Consequently they are conspicuous and well differentiated from the surrounding cells. The vascular

bundles are almost perfectly rounded and completely ensheathed by fibrous tissue. Three air spaces or lacunae

form passageways throughout the length of this petiole (Fig. 11D; Table 4). Serial sectioning demonstrated air

spaces that begin near the stem axis ultimately terminating at the leaf base (Fig. 11D). Although E. paradoxa

var. paradoxa has air spaces, they appear close to the leaf base but beyond the point where sections were made.

Fig. 4. A, B. Microtome transections of Echinacea atrorubens capitulum (x50). C-L. Free hand transections of ray ligules showing shapes of adaxial

epidermal cells. A. Disc floret nearer the base showing the style, the five-parted stamen filaments, surrounded by corolla tube. B. Disc floret nearer

the top showing the two-parted style surrounded by mature five-bilocular, united anthers with stained pollen grains and gamopetalous corolla. Note

absence of sclerotic tissue. This floral arrangement is typical of the Asteraceae. C. £ atrorubens at low magnification showing nipple-like apex in upper

left and more variable cells in lower right (x50). D. £ atrorubens at high magnification of C showing nipple-like apex and broadened base (x200). E.

£ laevigata showing dome shape, undifferentiated mesophyll, lack of secretory chambers, and uniform isodiametric abaxial cells (x50). F. £ pallida

showing dome-shape (x50) and close up inset. G.E. paradoxa var. neglecta tapering upward to rounded apex with secretory chamber below in mesophyll

tissue (x200) and close up inset. H. £ paradoxa var. paradoxa showing dome shape (x200). I. £ purpurea showing ball-like base tapering to rounded

apex (x200). J. £ sanguinea showing elongate"necked"apex and broader base (x200). K. £ simulata showing sharply pointed apex from a wider base

(x50). L. £ simulata higher magnification detail showing conical cells largest in Echinacea (x200).

104

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 5. A, B. Echinacea laevigata in sunny natural habitats of North Georgia, 28 May 2008. A. Stephens County, Chattahoochee National Forest at Currahee

Mountain. The plants shown here occur in small groups scattered among other forbs growing in open forest areas on rocky soil mostly of amphibolite.

Note the long, narrow purplish to white reflexed ray florets. B. Same site as A, showing close up of glabrous stem and long, narrow, drooping, purplish

ray ligules with bifid tips. C-F. £. paradoxavar.paradoxa natural habitats include open sunny glades at Ha Ha Tonka State Park, Camden County, Missouri.

C. Heimbeaugh Hill Glade. This area historically has been maintained with fire and current management includes the removal of Juniperus virginiana

trees and prescribed burns. This more xerophytic west-southwest facing site is characterized by multiple layers of dolomite dating from the early Ordo¬

vician period and these areas are sometimes referred to as "balds" or "barrens." Note that yellow coneflower predominates as a single taxon carpeting

the entire area in late May and early June. D. Close up of previous site from different direction showing synchronous blooming of the taller Echinacea

plants forming a canopy over the shorter prairie grasses with tree line in background. E. Close up of capitulum with prominent raised cone of paleae and

yellow reflexed ray florets with notched ends and crimson coloration at point of insertion shown later in blooming period. F. £. pallida at glade edge. It

typically grows here near the edge or margins of glades with yellow coneflower. Photo credits: Hugh & Carol Nourse (A, B), Christopher D. Crabtree (C-F).

Keller, Floral, stem, and petiole morphology of Echinacea

105

5. Echinacea laevigata (C.L. Boynton & Beadle) S.F. Blake, J. Wash. Acad. Sci. 19:273. 1929. (Figs. IE; 5A, B;

9E; 10E) . SMOOTH PURPLE CONEFLOWER.

Stems erect, usually unbranched and glabrous, 55-110 cm high, leaves petiolate, ray florets purplish or light

pink to whitish, spreading to reflexed. A distinct taxon found outside the distribution range of most Echinacea

taxa in open woods of Pennsylvania scattered in the Piedmont area south to the mountains of Georgia. Natural

habitats occur in sunny openings in forested areas maintained by wildfires and grazing animals. Thus pre¬

ferred sites are open woods, glades, cedar barrens, roadsides, clear cuts, dry limestone bluffs, and power line

rights-of-way associated with soils rich in magnesium, calcium, dolomite or limestone and rocky substrates of

amphibolite (Fig. 5A, B). It is most closely related to E. purpurea but differs in having a vertical caudex and

taproot, unbranched glabrous and glaucous stems, and leaf blades broadly lanceolate or elliptical mostly gla¬

brous, with ray florets longer and more slender (Fig. 5B). This is the second taxon of Echinacea that was Feder¬

ally listed as an Endangered Species (USFWS 1992).

Ray florets are pale pink to purple and lack trichomes, demonstrating the glabrous character of this tax¬

on. The venation pattern consists of 15 vascular traces in transection supported by abundant thick-walled tis¬

sue. The uniformity in size and dome-shape of the ray ligule adaxial epidermal cells is the most obvious char¬

acter. Secretory chambers are apparently lacking (Table 2).

The smooth and glaucous characters of the stem distinguish this taxon from E. purpurea and other taxa of

Echinacea. In every dimension the epidermal cells are larger than in other species. Moreover, a striking pattern

of straight, oblique, and curved walls demonstrates the irregularity in cell shape (Fig. 9E). Only the outermost

layers of the cortex consist of typical collenchyma. The greater part of the cortex is parenchymatous tissue

with large intercellular spaces (10 pm). The pericyclic fibers give a weak phloroglucinol test. They are mostly

thin and needlelike in structure and are easily broken into fragments. The end walls taper to a sharp point with

lengths that vary from 360 to 1150 pm (794 pm) and widths from 8 to 20 pm (13 pm). The phloem zone is

60-70 pm in radial extent. Macerated tissue had vessels that range in length from 159 to 450 pm (302 pm) and

widths range from 23 to 34 pm (29 pm). Some of the scalariform vessels are tailed with oblique perforation

plates. Little sclerihcation occurs between vascular bundles (Fig. 10E) and none was present in the pith. Mea¬

surement of stem diameter was ~5.2 mm, including the pith (~4.2 mm) with pith cell lengths that range from

60 to 174 pm (121 pm) and widths from 60 to 97 pm (80 pm) (Table 3).

The secretory system was represented by canals in the cortex and pith. Those of the cortex are relatively

abundant and large with usually five to eight epithelial cells originating opposite the vascular bundles and/or

interfascicular region. About half of the vascular bundles have single canals and the others occur in pairs. Ca¬

nals in the pith form double rings of epithelial cells as a result of a second periclinal division with an outer ring

of elliptical cells and inner one of rectangular cells. Pith canals have five to eight epithelial cells that surround

the canal cavity (Table 3).

The petiole in transection appears horseshoe-shaped. Leaves of this taxon tend to have sheathing bases

and this accounted for the lengthy “wings” (Table 4). Fundamental tissue extends to the very end of each wing

which fails to show a graduation into foliar anatomy. A uniseriate layer of cutinized cells underlies the epider¬

mis that is structurally indistinguishable from it. This double-layered tissue forms a definite boundary at the

periphery of the petiole. Five vascular bundles are seen in transection (Table 4).

6. Echinacea pallida (Nutt.) Nutt., Trans. Amer. Philos. Soc. II. 7:354. 1840. (Figs. IF; 4F; 5F; 6E; 9A (h); 10F;

11C). PALE PURPLE CONEFLOWER.

This taxon is an apparent segmental allotetraploid possibly derived from a hybrid between E. simulata and E.

sanguinea with a doubling of the chromosome number to 2n = 44. It has white pollen (Fig. 6E) and the largest

pollen grains in the genus, 24-28.5 pm in diameter (26.1 pm) when compared to the most closely related E.

simulata with pollen grains 22.5-24.5 pm in diameter (24.2 pm) and other Echinacea taxa with much smaller

pollen grains (McGregor 1968). This taxon flowers in late spring and early summer in rocky prairies, open

wooded hillsides, savannas, and glades concentrated in eastern Kansas and Oklahoma, western Arkansas, and

throughout Missouri. Plants are rarely branched, 40-90 cm up to 140 cm high, stem trichomes hirsute below

106

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 6. Echinacea purpurea (A-D), E. pallida (E), and £ simulata growing in natural habitats in Arkansas. A. Saline County, open woodland growing in a

clustered group, 16 Jun 2003. B. Saline County, Ouachita National Forest, open woodland along creek in shale barrens growing with Rudbeckia grandi-

floravar.grandiflora, 3 Jul 2003. C. Clark County,Terre Noire Natural Area, 6 Jul 2004, close up of capitulum showing brightly colored paleae, broad ray

ligules with complex venation pattern, and bifid notched ends. D. Saline County, close up of capitulum showing visit of Zebra Swallowtail Butterfly,

order Lepidoptera, family Papilionidae, genus Protographium, Jun 2001. E. £ pallida, Arkansas, Boone County, open areas on acidic chert. Baker Prairie

Natural Area, 8 Jun 2013. Note bumblebee ( Bombus sp.) on dark purplish paleae with contrasting traces of white pollen indicative of this tetraploid. F.

£ simulata, Arkansas, Boone County, dolomite glades, Jun 1996. Note the yellow pollen adhering to the paleae and also the hirsute stem. Photo credits:

John Pelton (A, B, D, F), Mark Clark (C), Joan Reynolds (E).

Keller, Floral, stem, and petiole morphology of Echinacea

107

and more dense above, ray florets reflexed, sparsely hairy abaxially (McGregor 1968; Urbatsch et al. 2006). Ray

florets vary from almost entirely dark rose red to reddish purple in eastern Kansas and throughout Missouri to

whitish that predominates in eastern Oklahoma and western Arkansas and farther south to Texas and Louisi¬

ana (McGregor 1968). An excellent current source of general macromorphological and medical information is

available for E. pallida, including color photographic images of habit and root microanatomy (Upton 2010).

Some irregularity in shape was apparent in the ray ligule adaxial epidermal cells, intergrading from a

short bullet shape to a more conical or slightly dome-shaped papillose cell (Fig. 4F). Noticeable microscopi¬

cally were trichomes of various lengths on the abaxial surface. A large amount of thick-walled supporting tis¬

sue enveloped the 13 vascular traces (Table 2).

Trichomes are moderately scattered over the stem surface described by McGregor (1968) as “hirsute on

both surfaces.” Three to five septa divide each trichome into four or five cells at irregular intervals (Fig. 9A, h).

Generally the trichome is slender and tapered to a sharp point with lengths that range from 1 to 2 mm. In sur¬

face view the epidermis has angulate and rectangular cells; this is expressed in transection notably in the ir¬

regularity of epidermal cell size and shape (Fig. 9F). Paradermal sections were difficult to make due to the

grooves and thick cuticle. The cuticular layer was 7.0 pm thick, bearing tiny spine-like projections on the sur¬

face. The inner tangential walls were heavily cutinized (Table 3).

The cortical zone including the endodermis and epidermis measures 180 pm at the narrowest point and

430 pm at the broadest. Collenchymatous tissue makes up most of the cortex. In macerated preparations col-

lenchyma cells range in length from 110 to 280 pm (189 pm) and in width from 35 to 46 pm (45 pm). Pericyclic

fibers as seen in macerated preparations range in length from 450 to 1450 pm (1056 pm) and in width from 40

to 18 pm (27 pm) with a lumen diameter of 8-16 pm. Xylem elements consist of vessels, fibers, and xylem pa¬

renchyma. Macerations of vessel elements ranged in length from 215 to 610 pm (378 pm) and in width range

from 40 to 51 pm (48 pm). No annular vessels or tracheids are present. All vessels have simple perforation

plates. The protoxylem consists mainly of spiral vessels, and the metaxylem, with scalariform to definite re¬

ticulate pits, seems to be developmentally more advanced than in E. simulata. Pith diameter is ~3.2 mm with no

sclerenchyma cells present (Table 3). The stem tissue map shows cambial activity that unites the collateral

vascular bundles into a mostly solid ring due to some secondary growth in the interfascicular regions (Fig.

10F). The pith consists wholly of parenchymatous tissue with length of cells ranging from 74 to 135 pm (101

pm) and widths that range from 72 to 110 pm (105 pm) (Table 3).

Secretory canals are present in both cortex and pith. Undeveloped canals have approximately four epithe¬

lial cells and a small canal cavity. These arise through radial (anticlinal) divisions sometimes followed by tan¬

gential divisions that give a rectangular shape to epithelial cells. With the completion of these divisions, the

epithelial ring consists of at least six cells with an enlarged cavity. Canals of the pith appear singly, in pairs, or

in triplets opposite the protoxylem points. Three to seven epithelial cells delimit the pith canals with small

patches of thin-walled cells (11-15) designated as accessory tissue staining green with fast green stain. The

pith canals in this species tend to anastomose, still, however, maintaining their individual identity. Cortical

canals are spherical in shape. Each bundle cap has one canal on each side lying adjacent to the endodermis in

the interfascicular region.

7. Echinacea paradoxa (Norton) Britton var. neglecta McGregor, Trans. Kansas. Acad. Sci. 70:366-370. 1968.

(Figs. 2A; 4G; 10G; 11F-G; 12C-F) . bush’s purple coneflower.

Plant stems 30-80 cm high, usually unbranched, yellowish green, sparsely to densely strigose similar to vari¬

ety paradoxa. This taxon is easily distinguished from var. paradoxa by its rose-colored, purple or white ray flo¬

rets and a geographic distribution confined to the Arbuckle Mountains of southwestern Oklahoma in rocky

prairies and open, wooded hillsides (McGregor 1968).

Ray ligule adaxial epidermal cells have one to three secretory chambers arranged on both sides of the

vascular trace and scattered throughout the mesophyll tissue. These cells have a wide range in size that nearly

corresponds to those of the stem, leaf, and petiole secretory systems. Each secretory chamber has an interior

cavity of approximately 50 pm surrounded by three to seven epithelial cells. This is similar to the canals of the

108

Journal of the Botanical Research Institute of Texas 8(1)

stem and petiole, (Fig. 4G). Ray adaxial ligule epidermal cells have outer tangential walls that result in sides

that are parallel about three-fourths the way up, sloping gradually into a rounded apex (Fig. 4G). These cells are

highly variable and in some cases distinctly papillate. The two varieties, paradoxa and neglecta , have adaxial

ray ligule epidermal cells that are similar in size and shape (Table 2).

Trichomes are sparse or lacking but with distinctive epidermal patterns. In surface view epidermal cells

are uniform in shape but significantly smaller in width. The breadth of cortical tissue extends from 230 to 500

pm. Outer portions of the cortex mostly consists of collenchyma while the inner portion intergrades into par¬

enchymatous tissue. The phloem zone extends from 46 to 56 pm. Stem diameter is ~3.8 mm including a pith

diameter of ~2.9 (Fig.lOG; Table 3). Pith cells range from 137 to 257 pm (189 pm) with widths from 33 to 65 pm

(47 pm) and in longisection are densely pitted.

Stem secretory canals are present in both the pith and cortex. Many canals are tangentially flattened espe¬

cially the larger ones. Canals occur either in ones, twos, or threes opposite protoxylem points (Fig. 11D). Epi¬

thelial rings consist of 5-15 cells rectangular in shape. Accordingly the canal cavity is relatively large from 20

to 36 pm surrounded by 8-14 cells (Fig. 11G). Canals originate opposite both the vascular bundle caps and

interfascicular regions. The newly formed canals are much smaller and confined to the endodermal layer;

while those embedded in the cortical tissue are well developed and relatively large (Table 3).

A lunate to bow-shaped petiole is the result of a greater lateral overall thinness (Fig. 12C). Canals are well

formed (distinct epithelial cells surrounding a large cavity) and larger than can be found elsewhere in the ge¬

nus. Comparatively some of these canals are double the size of even the largest canals of other taxa. The total

number of canals present (14) far exceeds counts made for other taxa. These canals are located typically on the

abaxial side of the vascular bundles and some atypically on the adaxial side. The medial vascular bundles had

pairs of canals on both the abaxial and adaxial sides. Well-formed canals have an extremely large cavity 31-43

pm in diameter surrounded by a well-differentiated ring of epithelial cells (Table 4). In other taxa canals are

surrounded by four to six epithelial cells but here 5 to 15 cells make up the epithelial layer. Furthermore, the

epithelial cells have a distinctive shape closely approaching a rectangular shape; while others found in different

Echinacea taxa usually are more elliptic with curved anticlinal walls.

Apparently no taxon in the genus Echinacea has brachysclerids or stone cells in aerial plant parts except

in the petioles of var. neglecta (Fig. 12D, E, F; Table 4). These sclerids occur as single, conspicuous cells with

highly refractive thickened walls that make them appear as shiny, glistening structures reminiscent of the

clustered stone cells in pear fruit mesocarp (Fig. 12E). More noticeable was the concentric layering of the thick¬

ened walls and the branched ramified pit canals revealed by a preferential stain. In this case the walls were

heavily lignihed (stained with safranin) throughout, including the thickened secondary walls (Fig. 12D). The

sclerids were scattered throughout the fundamental tissue as seen in Fig. 12D, E, F). Similar brachysclerids

also were found in roots of E. angustifola (Axentiev et al. 2010) and E. pallida (Upton 2010).

8. Echinacea paradoxa (Norton) Britton var. paradoxa in N.L. Britton and A. Brown, Ill. Fl. N. U.S. ed. 2,

3:476 1913 (Figs. 2B; 4H; 5C, D, E; 9G; 10H) . YELLOW CONELLOWER.

This taxon is easily recognized by its bright yellow ray florets either drooping or reflexed (Fig. 5E). It differs in

color from all other Echinacea species that have deep purple, pinkish, to white ray florets. General habit of the

plant is 40-90 cm tall, stems usually unbranched, yellowish green, sparsely to densely strigose (McGregor

1968). A relatively small distribution area is restricted to rocky and upland prairies, limestone and dolomite

cedar glades, savannas, bald knobs and also roadsides of west-central and southern Ozarks of Missouri and

north central Arkansas (Yatskievych 2006). Native wild populations of strikingly beautiful yellow coneflowers

sometimes dominate the open held landscape in the Ozark cedar glades, especially in Ha Ha Tonka State Park,

Missouri (Fig. 5C, D; Crabtree 2008; Richter 2013; H.W. Keller, pers. obsv.).

Ray ligule adaxial epidermal cells are wide at the base with straight parallel sides that become convex at

the apex (Fig. 4H). The overall dome shape and size are similar in both var. neglecta and var. paradoxa. One

chromoplast occurs in each cell. A secretory system is well differentiated in size, frequency, and arrangement.

Secretory chambers are associated with vascular traces and occur on both adaxial and abaxial sides; each

Keller, Floral, stem, and petiole morphology of Echinacea

109

chamber is conspicuous mainly due to their large cavities and the three to seven peripheral epithelial cells (Fig.

9G). The ray floret secretory system is similar in size and number in both varieties but exceeds in size and

greater numbers the canals in all other Echinacea taxa, including the closely related E. atroruhens (Table 2).

Only a few trichomes were observed. The stem lacks grooves at this position so the paradermal sections

show more uniformity in cell shape. This stem morphology results in a fairly constant rectangular shape of the

epidermal cells in surface view (Fig. 9G). In transectional view many cells are squarish instead of the typical

tabloid epidermal cell. Anticlinal walls of epidermal cells are thin and straight. The cortex is made up of an

outer collenchymatous zone and an inner parenchymatous tissue with large intercellular spaces. Lengths of

collenchyma cells range from 90 to 228 pm (180 pm) and widths range from 28 to 61 pm (45 pm). Pericyclic

fibers range from 475 to 200 pm (1190 pm) and widths range from 16 to 23 pm (20 pm). The average wall thick¬

ness is 6 pm with a lumen of 6-10 pm. The phloem zone is 50-95 pm in radial extent. Vessels range in length

from 325 to 650 pm (485 pm) and widths range from 23 to 50 pm (33 pm). Some secondary growth was ob¬

served in the stem (Fig. 10H). Comparison of stem tissue maps for varieties neglecta (Fig. 10G) and paradoxa

(Fig. 10H) appear almost identical Vascular bundles are connected by interfascicular activity with some

sclerihcation that results in a more or less solid vascular cylinder without any interfascicular areas (Fig. 10H).

Pith parenchyma cells are 83-215 pm in length (190 pm). No thick-walled sclerotized cells are present in

the pith region. The secretory system is represented by canals in both the pith and cortex. Secretory canals oc¬

cur in the pith next to the protoxylem points and in the cortex opposite the vascular bundle caps and interfas¬

cicular regions in both varieties. Secretory canals occur in the pith next to the protoxylem points and in the

cortex opposite the vascular bundle caps and interfascicular regions in both varieties. Epithelial rings consist

of three to six cells positioned to give an hourglass or star-shaped cavity usually much reduced in size adjacent

to the endodermis. Cortical canals originate opposite vascular bundle caps and interfascicular regions. Epithe¬

lial rings consist of five to eight cells (Table 3).

This taxon has the thickest petiole studied with an exaggerated curvature on the abaxial side that is dis¬

cernible even macroscopically. Canals of the secretory system are comparatively numerous occurring in pairs

instead of singly. Toward the abaxial margin of the petiole abundant chlorenchymatous pockets are inter¬

spersed among the collenchyma tissue. Five vascular bundles are noted in transection (Table 4).

9. Echinacea purpurea (L.) Moench., Methodus 591. 1794. (Figs. 2C; 41, 6A, B, C, D; 9H; 101). (2n = 22). eastern

PURPLE CONEFLOWER.

This taxon has a deep fibrous root system that distinguishes it from all other Echinacea taxa. Distribution is

more widespread but concentrated in the central plains states and then north and southeasterly in rocky open

woods, thickets, and prairies (Fig. 2C; McGregor 1968). It is the most widely distributed taxon and character¬

ized by its tall branching habit of 60-180 cm; awns of paleae as long as body; lower leaves ovate and often

toothed; and dull purple to pinkish ray florets, spreading to recurved, 3-8 cm long 7-14 (-19 mm) wide (Fig.

6A-D). The larger colorful ray florets have encouraged the development of hybrids and cultivars grown com¬

mercially and as ornamentals in gardens.

Externally the gross morphology of the ray florets reflects the greater size in breadth but internally size is

expressed by the increased degree of venation and the 31 vascular traces in transection. Comparatively E. pur¬

purea has the broadest ray florets of all the taxa studied (Fig. 6C). Secretory chambers measure 40 pm in diam¬

eter with each canal surrounded by five epithelial cells.

Ray ligule adaxial epidermal cells have a wide ball-like base that gradually taper to a rounded apex (Fig.

41). This shape contrasts with the shorter, convex shape of E. laevigata adaxial epidermal cells. In addition the

broader ray florets of E. purpurea have a more complex venation pattern that differs from E. laevigata with more

narrow ray florets and venation greatly reduced (Table 2).

Nonglandular trichomes thickly cover the stem. The base of the trichome is swollen and accentuated by

an outgrowth of surrounding epidermal cells. Repeated sectioning was required to get thin paradermal peels.

Heavy pubescence along with grooved stems undoubtedly caused this difficulty. In surface view the margin of

the cuticle is wavy and rough in outline. Striae or rod-like bodies decorating the surface of the epidermis were

110

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 7. Echinacea sanguined in sunny open field habitats. A. Wood County, Texas, 3.2 miles southeast of Winnsboro along State Highway 37 at edge of

field near roadside ditch. Scattered groups of plants growing on sandy soil known geologically as the Queen City Sand, 22 May 1989. B. Nearby site

9.4 miles southeast of Winnsboro along State Highway 37 showing four capitula with whitish ray flowers and hirsute stems. Note the smallest stem

diameter found in Echinacea and spindly habit. C. Same site as B showing close up of a single capitulum in early anthesis with spreading white ray

ligules notched at tips. D. Same site as B showing close up of later stage anthesis. Note the reflexed ray florets and faint purplish colors at point of

insertion. Photo credits: Bob O'Kennon.

ascribed to cuticle depositions. Epidermal cells have some rectangular end walls which are usually oblique and

angulate (Fig. 9H; Table 3).

The cortex contains an outer, several-layered, narrow zone of collenchyma cells and an inner, many-lay¬

ered, loosely arranged broad zone of parenchymatous tissue. Collenchyma cell lengths range from 111 to 157

pm (131 pm) and widths from 21 to 37 pm (30 pm). Pericyclic fibers give a moderate phloroglucinol test as

shown by the bundle caps. Fibers range in length from 663 to 2400 pm (1288 pm) with widths from 8 to 22 pm

(15 pm). The average wall thickness is 5 pm and lumen size is 7 pm. The phloem zone radially extends from 59

to 73 pm. Vessel lengths range from 232 to 680 pm and widths from 20 to 42 pm. Pith diameter is ~3.6 with

parenchymatous cell lengths from 92 to 130 pm with widths from 93 to 110 pm. No sclerids are present (Fig.

101 ).

The secretory system consists of canals in both the pith and cortex. Those of the cortex tend to be tangen¬

tially flattened and are located opposite both vascular bundle caps and interfascicular regions. Epithelial cells

are in direct contact with the endodermis and are not well differentiated from surrounding parenchymatous

tissue. Epithelial rings of five to eight cells surround canals. Cortical canals number more than 48 in compari-

Keller, Floral, stem, and petiole morphology of Echinacea

111

son to the less numerous pith canals (~34) centric or excentric in position. Epithelial cells vary from elliptical

to subspherical shapes with no tangential divisions. The vascular stele is broken up into a dictyostele. Medul¬

lary rays pass out into the cortical tissue (Table 3).

Photosynthetic tissue is scattered among the parenchyma cells that make up the abaxial bulge of the

petiole. Secretory canals are prominent when found adjacent to the medial vascular bundle, becoming progres¬

sively smaller as the lateral vascular bundle size decreases. Five vascular bundles are evident in sectional views;

two more are diminutive but recognizable. The central vascular bundle lacks fibrous tissue on the abaxial side

(Table 4)

10. Echinacea sanguinea Nutt., Trans. Amer. Phil. Soc. n. ser. 7:354. 1840. (Figs. 2D; 4J; 7A-D; 9A-i-j, I; 10J;

12B). SANGUINE PURPLE CONEFLOWER.

This is a distinct taxon often confused in the past with F. pallida. However, it differs in having hemispheric

capitula, slender and more spindly stems, 40-90 cm high and sometimes reaching 122 cm in roadside ditches

of East Texas (Fig. 7A-D; Bob O’Kennon pers. obsv.). It has narrower ray florets, elliptical leaves, yellow pollen,

and is a diploid (n = 22). Distribution is more restricted to acidic, sandy soils, open pine barrens, woodlands,

and prairies of southeastern Oklahoma, southwestern Arkansas, western Louisiana, and east Texas (Mc¬

Gregor 1968). It flowers in May in East Texas and gradually becomes a more rose color in a cline northward

with later anthesis (Fig. 7A-D). Ray florets color and flowering times were genetically fixed from the original

habitats while growing in the KU Experimental Gardens (McGregor 1968). This is the only taxon of Echinacea

that has a more southerly distribution in the central states and thus differs from all other taxa.

Ray ligule adaxial epidermal cells have distinctive elongate, “necked” papillae that differ from all other

taxa (Fig. 4J; Table 2). Trichomes are absent. No secretory chambers were present.

Short, stout trichomes thickly cover the stem. The trichome usually has fewer septa with the base heavily

cutinized (Fig. 9A-i-k, I). Epidermal patterns consist of straight, mostly rectangular cells (Fig. 91).

Collenchyma tissue makes up the greater part of the cortex except for several parenchymatous layers

outside of the endodermis. Lengths in these elongated, tapered cells range from 151-373 pm (180 pm) and

widths range from 40-45 pm (43 pm). Breadth of the cortical region ranges from 190-230 pm. Pericyclic fibers

are not fragile or needle-like but are thicker with blunt or truncate ends with lengths from 300-1350 pm (850

pm), and widths 17-21 pm. The phloem zone in radial section is 22-45 pm (Table 3). Vessel lengths range from

275-950 pm (523 pm), and widths range from 20-35 pm (20 pm) (Fig. 11).

Echinacea sanguinea has the smallest stem diameter (~1.7 mm in width and pith diameter of -0.9 mm)

studied which reflects the spindly habit of this taxon (Fig. 10J). No sclerenchyma fibers are present in the pith

(Table 3). Pith parenchyma cells ranged from 89-176 pm in length (137 pm), and widths from 41-105 pm (72

pm). The walls of the parenchyma cells are densely marked with elliptical and conspicuous primary pit helds.

The secretory system has canals that are interfascicular in origin withl3-15 canals in the cortex. A

brownish substance appears in some of the epithelial cells, making their presence conspicuous. Canals range

from 40-48 pm with either a five or six-celled epithelial ring (Fig. 11 A). The relatively small amount of vascu¬

larization reflects in part the spindly habit of the plant. Furthermore, interfascicular rays are sclerihed, previ¬

ously mentioned as a distinguishing character for other species (Table 3).

Gross morphology of the petiole varies based on the location either near or distant to the stem. Close to

the stem the petiole is heart-shaped in transection with the ends folded together. Nearer the leaf base the peti¬

ole unfolds allowing reorientation of the margins. Secretory canals are relatively small arranged one on each

side of the abaxial bundle cap (Fig. 12B). Four minor vascular bundles were present and the medial vascular

bundle lacks an abaxial fibrous cap (Table 4).

11. Echinacea simulata McGregor, Sida 3:282. 1968. (Figs. 2E; 4K-L; 6F; 9A-a-g, k, I; 10K; 11H). wavy-leaf

PURPLE CONEFLOWER.

This species was cited as F. speciosa McGregor by Keller (1962), but that name was never validly published. In

the past it was included in F. pallida , but the pollen size in F. simulata is smaller and yellow as compared to

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Journal of the Botanical Research Institute of Texas 8(1)

white in E. pallida. Stems 60-120 cm tall are mostly unbranched with sparsely to densely pustular based tri-

chomes. It occupies a distinct geographic area restricted mostly to north central Arkansas, eastern Missouri,

western Illinois, and west-central Kentucky (Fig. 2D). The diploid chromosome number of 2n = 22 contrasts

with E. pallida, a polyploid with a chromosome number of 2n = 44 (McGregor 1968).

This taxon has the largest adaxial epidermal ray ligule cells. The distinctive conical shape of the cell with

a wide base and sides tapering upward that forms a sharp point, distinguishes this taxon from all other Echina¬

cea taxa (Fig. 4K-L). Secretory chambers are present having either four or five epithelial cells. Thickness of the

ray ligule is comprised mostly of the larger adaxial and abaxial epidermal cells. Trichomes are present on the

abaxial surface and vascular traces number 13 in transection (Table 2).

Nonglandular trichomes are moderately scattered on the stem described by McGregor (1968) as “hirsute

or somewhat tuberculate hirsute.” These slender, uniseriate hairs gradually taper toward the apex. The termi¬

nal cell is highly variable and often structurally modified into a rounded or sharp point. Most of the trichomes

have three septa, occasionally five in longer ones (Fig. 9A,-k). Marks sculptured in the trichome wall are len¬

ticular in shape. The ontological sequence of the trichome conforms to the pattern in which only epidermal

cells undergo division. However, the trichome is raised on a supporting base formed from both epidermal and

sub-epidermal cells (Fig. 9A-a-g). In surface view they appear morphologically distinct from the surrounding

epidermal cells and have a highly irregular outline, accounting for the wide range in size (Fig. 9J). This no

doubt is due to the grooved surface of the stem which tends to give more rectangular shaped cells in the

grooves, and more angulate cells on the ridges. In transection the cross diameter of epidermal cells is uniform,

unlike the length, which varies considerably (Table 3).

The cortex has small pockets of thin-walled chlorenchymatous cells that underlie each sub-stomatal

chamber. These fan out short distances around the stem and are interrupted at intervals by collenchyma which

directly abuts the epidermis. Generally three types of collenchyma cells are recognized: angular, lamellar, and

tubular. All three types often intergrade, but, the tubular type with intercellular spaces and angular thicken¬

ings is predominant in E. simulata. General shapes consist of elongate cells having unevenly thickened walls,

with either rectangular, oblique, or tapering ends. A transitional region between cortical collenchyma and

parenchyma occurs within one or two rows outside of the endodermis. Size and shape of collenchyma cells

were studied from longitudinal sections and maceration: lengths range from 124 to 280 pm and widths range

from 27 to 41 pm.

The uniseriate endodermis in transection is recognizable by the elliptic, thin-walled cells lacking pits. A

starch test with a potassium iodide solution gave a positive reaction (a dark blue-black color) confined mostly

to the endodermal layer. Instead of giving a blue ring, groups of two or three cells were filled with starch grains;

then, for some distance, cells were void of starch. The starch granules measure 43 pm in diameter and appear

roundish with a roughened surface.

The pericyclic fibers of the bundle caps in transection extended radially from 83 tol35 pm (Fig. 4). Macer¬

ated tissue produced fibers between 322 andl341 pm (761 pm) in length and width 9.7-31 pm (16 pm) that

varied in shape with some tapered to a sharp point, others blunt, and still others, truncate. Lumen diameter

ranges from 4 to 24 pm (11 pm). All fibers have slit-like pits and give a strong positive phloroglucinol reaction.

The phloem zone measures 51-75 pm in radial extent. No crystals or storage products are present. Stem tran-

sections show that the number of rows of metaxylem vessels ranges from three to seven with interspersed fi¬

bers (Table 3). Macerated preparations show vessel lengths vary from 197 to 644 pm. The end walls are com¬

pletely dissolved, resulting in the simple perforation type in end view. Shapes vary from barrel-shaped with

horizontal end walls while others are oblique and pointed. No annular vessels were observed and tracheids

were absent. Vessel widths range from 24 to 42 pm with spirals either loose to close that range from 364 to 1008

pm and pitted vessels range from scalariform to elliptical.

Stem diameter was ~4 mm, including a pith diameter of ~3.3 mm, with intercellular spaces of 1 pm. Pith

cells are nearly isodiametric, loosely arranged, and densely pitted. Cells in the center are larger, merging to¬

ward the periphery into smaller thicker-walled cells. The cells supporting vascular bundles tend to become

Keller, Floral, stem, and petiole morphology of Echinacea

113

sclerotized, especially surrounding the resin canals. No sclerotic cells are present in the center of the pith. Se¬

cretory canals originate adjacent to and through the division of the endodermis forming within the interfas¬

cicular region, the exception being where surrounding cells of the vascular system undergo positional rear¬

rangement, tending to relocate the canal along the ascending arc of the bundle caps. Canals also are found ei¬

ther singly or in pairs closely associated with the vascular bundle at the periphery of the pith (Fig. 11H). These

appear to be an integral part of the vascular bundle, but they actually form outside of and opposite the protoxy-

lem points, becoming surrounded by cells that are highly lignihed (Table 3).

Pith canals number 36 with diameters from 24 to 50 pm. Such a great range can be attributed to the devel¬

opmental stage of the canal. Smaller canals have reduced cavities, and larger, relatively fewer epithelial cells.

The larger canals have an enlarged cavity and smaller, relatively more epithelial cells that range in size from 17

to 22 pm between anticlinal walls. The number of epithelial cells lining the cavity varies from four to seven.

Epithelial cell shape varies from square to oblong rectangular, sometimes ovoid but always the walls form an

oblique angle. Each epithelial cell has a dense cytoplasmic content with a conspicuous nucleus. When present

in pairs they appear in a juxtaposition sometimes spatially separated by as much as 100 pm. Cortical canals

tend to be crushed in sectional view and are not well differentiated from the surrounding cortical parenchyma.

Stem diameter is relatively large in part because of sclerihcation and continuous growth in the interfas¬

cicular region (Fig. 10K). Stellar configuration seems to follow a circular design with secondary growth in only

a portion of the stem. It is interesting to note that both E. pallida and E. simulata have interfascicular growth to

a degree not seen in other Echinacea taxa (Table 3).

A striking feature of the petiole not found in other species is the fan-shaped medial vascular bundle. It has

a wide fibrous cap gradually sloping adaxially as the phloem and xylem diminish in lateral extent. Seven vas¬

cular bundles can be seen in transection. The canal system is not well developed; in fact, canals associated with

the more lateral vascular bundles are reduced greatly in size. The tips of the petiole arch upward and give a

more or less lunate-shape in outline. Some petioles have loosely arranged parenchymatous tissue orientated

adaxially near the epidermis, however, in E. simulata, this area is compacted and constituent cells have smaller

intercellular spaces (Table 4).

12. Echinacea tennesseensis (Beadle) Small, Man. Southeast FI. 1421, 1509. 1933. (Figs. 2F; 8A-F) . TENNESSEE

PURPLE CONEFLOWER.

This taxon’s microanatomy was not included here because live plants were not available in the KU Experimen¬

tal Gardens. McGregor (1968) described its distribution as an endemic restricted to the “dry, gravelly hills and

barrens near LaVergne, Tennessee” (Rutherford County). Specimens were only available from herbaria so Mc¬

Gregor (1968) confined his observations to the type specimen. He recognized E. tennesseensis as a distinct

species most closely related to E. angustifolia var. angustifolia, noting that it was “a very rare or possibly extinct

species endemic to the Cedar Barrens area of central Tennessee.” However, it differs morphologically in the

smaller stature, 10-50 cm tall, softer pubescence, smaller pollen grains, leader stem, and ray florets ascending

rather than drooping (Fig. 8A-F)

A review article by Walck et al. (2002) notes that E. tennesseensis was one of the first plant species listed on

the Federal Endangered Species list in 1979. It was listed in Tennessee as endangered and protected under the

Rare Plant Protection and Conservation Act of 1985. Five additional populations of this species were found in

Davidson, Rutherford, and Wilson counties (Fig. 8A-F) in the vicinity of Nashville as part of the Central Basin

region of Tennessee. Several population sites near Nashville were destroyed when land was cleared for housing

developments. In these localities it occurs in a general area referred to as the Cedar Glades ( Juniper us virginiana

L.) on outcrops of Ordovician-age Lebanon Limestone. The United States Fish and Wildlife Service removed E.

tennesseensis from the Federal List of Endangered and Threatened Plants delisted in the Federal Register 2 Sept.

2011 (USFWS 2011). The discovery and recovery of additional populations of this species represent the coop¬

erative efforts of scientists, especially Elsie Quarterman, and conservation groups over the last 30 years.

Nonglandular trichomes are present on the stem either sparsely or thickly covering the surface in all Echinacea

114

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 8. Echinacea tennesseensis in sunny, open natural habitats in Wilson County, Tennessee, on a site derived from soils of Ordivician age limestone (Lane

Farm State Natural Area). These areas have shallow soils and more xerophytic conditions referred to as cedar (Juniperus virginiana) glades or barrens

often interspersed with Little Bluestem grass. Photographs taken the evening of 21 Jun 2013 and the morning of 23 Jun 2013 at the peak of anthesis.

A. Landscape panoramic view of glade where Eastern Red Cedar trees occur around the marginal edge and Echinacea dominates in patches backlit by

the evening sun. B. Morning hours showing capitula facing the sun. C. Patchy distribution on gravelly, rocky site. D. Top view of capitulum showing 13

marginal ray florets ascending upward and not reflexed downward with two-notched tips. E. Underside view of capitulum showing the hairy whorl

of phyllaries subtended by hairy fluted stem. F. Side view of entire flower and stem showing ascending ray florets with two and three notched tips.

Photo credits: Todd Crabtree.

Keller, Floral, stem, and petiole morphology of Echinacea

115

Fig. 9. Stem paradermal peels showing line drawing illustrations of trichomes. A. Echinaceasimulata developmental sequence of nonglandular, uniseriate,

multiseptate trichome (a-g). Note that the epidermal and sub-epidermal layers give rise to three to five septate trichomes with sharply pointed ends (h,

£ pallida with four septa; i-j, £ sanguinea with three septa, greatly thickened walls and reduced lumen; k, £ simulata with five septa in longer flexuose

trichome) (x78). B-J. Stem paradermal peels showing line drawing illustrations of epidermal cell shapes and sizes (all images x30). B. £ angustifolia

var. angustifolia showing irregular pattern with four anomocytic type stomata in field of view. C. £ angustifolia var. strigosa showing rectangular cells.

D. £ atrorubens showing mostly rectangular cells with straight end walls in middle portion. E. £ laevigata showing the largest cells in irregular sizes

and shapes with mostly oblique to curved walls. F. £ pallida. Note irregular angulate and rectangular cells. G. £ paradoxa var. paradoxa. Note mostly

rectangular cell shapes. H.£ purpurea. Note oblique and angulate end walls. I.£ sanguinea. Note mostly rectangular cells. LE. simulata. Note irregular

outline from trapezoid to rectangular.

116

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 10. Echinacea stem transections compared as tissue maps at same magnification (x8.8). A. E. angustifolia var. angustifolia. B. £ angustifolia var.

angustifolia "race intermedia." C. £ angustifolia var. strigosa. D. £ atrorubens. E. £ laevigata. F. £ pallida. G. £ paradoxa var. neglecta. H. £ paradoxa

var. paradoxa. I. £ purpurea. J. £ sanguinea. K. £ simulata.

Keller, Floral, stem, and petiole morphology of Echinacea

117

taxa with the exception of E. laevigata where they are lacking. These trichomes are simple but variable in size

with three to five septa and usually sharply pointed tips (Fig. 9A). Lenticular markings always appear in the

trichome wall. All trichomes form on a morphologically distinct area of the epidermis (Fig. 9A, a-g). The

grooved (fluted) stem is extended directly below the capitulum for variable distances in different taxa (Fig. 8E).

This grooved surface area accounts in part for the irregular shapes and sizes of stem epidermal cells in parader-

mal peels (Fig. 9B-J).

Throughout the genus only anomocytic stomata occur which have an irregular outline surrounded by a

limited number of cells which cannot be distinguished from other epidermal cells (Fig. 9B). The subsidiary

cells usually found in other stomata types are absent. This type of stomata opening is also called the ranuncu-

laceous type because it commonly is found in the family Ranunculaceae (Fig. 9B; Metcalfe & Chalk 1950).

Surface views of stem epidermal cells show more irregular patterns of straight, oblique, and curved cell

wall shapes with overall larger dimensions as in E. laevigata (Fig. 9E). In striking contrast, E. angustifolia var.

strigosa had the smaller stem epidermal dimensions and a more regular rectangular to squarish cellular pattern

(Fig. 9C). Echinacea paradoxa var. neglecta had a distinctive pattern of rectangular to more trapezoidal-like

shapes with slanted end walls (Table 3). Stem epidermal cells of all 11 Echinacea taxa studied were illustrated

with line drawings by Keller (1962).

Cortical tissue in the near-surface region of the stem outside of the vascular bundles is modified into

chlorenchymatous tissue with neighboring amounts of parenchymatous cells with much larger intercellular

spaces. This photosynthetic tissue is manifest in the greenness of the stem. The cortical zone is comprised

mostly of collenchyma with thickened cell walls that abut on the smaller intercellular spaces (Fig. 11B). All

Echinacea taxa apparently have collenchyma in this region to strengthen and support the aerial system of the

stem. Stem tissue systems in Echinacea lack crystals of any type as well as lacticiferous canals and mucilagi¬

nous cells. Secretory canals are present adjacent to and originate through the division of the endodermis. All

species have secretory canals located either opposite the vascular bundle caps and/or interfascicular region.

When secretory canals occur in the pith, they originate opposite the protoxylem points (Table 3).

All Echinacea taxa have a distinct, continuous endodermal layer or layers. Carbohydrates appear confined

to the endodermal tissue as granular starch grains with roughened surfaces (Fig. 11C). Macerated stem tissues

consisted of xylem elements either spiral, scalariform, and reticulate vessels, xylem parenchyma, fibers but no

tracheids. Phytomelanin-coated sclerids found in the vascular tissue in the roots of E. angustifolia and E. pallida

were not observed in the stem tissue preparations of the xylem and phloem.

GENERAL DICHOTOMOUS KEY TO ECHINACEA

This key to Echinacea taxa is based on macro- and microcharacters of ray florets, stems, and petioles. Echinacea

tennesseensis is not included here and is only known from cedar barrens near Nashville, Tennessee.

1. Stem pith tissue with sclerotic cells scattered throughout_ E. angustifolia var. angustifolia,

E. angustifolia var. strigosa, and hybrids

1. Stem pith tissue lacking sclerotic cells.

2. Secretory canals present only in cortex; stems slender, less than 2 mm diameter_ E. sanguinea

2. Secretory canals present in pith and cortex; stems stouter, more than 2 mm diameter.

3. Stem diameters >4.5 mm; >42 protoxylem points.

4. Ray ligules 5-6 mm; ~15 vascular traces; stems and leaves glabrous; plants with a taproot; distribution restricted

to southeastern U.S.A._ E. laevigata

4. Ray ligules 7-12(19) mm wide traversed by ~31 vascular traces; stems and leaves bearing trichomes; plants with

fibrous root system and horizontal rhizome; distribution mostly in Missouri and Arkansas with scattered popula¬

tions farther east _ E. purpurea

3. Stem diameters <4.5 mm; <42 protoxylem points.

5. Petioles with stone cells_ E. paradoxa var. neglecta

5. Petioles lacking stone cells

6. Ray florets bright yellow_ E. paradoxa var. paradoxa

6. Ray florets purple, red, pink, or white

7. Petioles with three air spaces; ray ligules adaxial epidermal cells with a broadened base and a short"nipple

like"apex_ E. atrorubens

7. Petioles lacking air spaces; ray ligules adaxial epidermal cells dome shaped to conical.

118

Journal of the Botanical Research Institute of Texas 8(1)

8. Ray ligules adaxial epidermal cells distinctly conical and largest in size (83-125 pm in height); pollen

yellow_E. simulata

8. Ray ligules adaxial epidermal cells dome shaped and smaller in size (58-82 pm in height); pollen white_E. pallida

DISCUSSION

Ray ligules

Baagoe (1977a) assessed and discussed the functional role of ray ligule adaxial epidermal cells in the Asteraceae.

Although Echinacea was not included in her study, the highly specialized adaxial epidermal cells occurring

here and in a broad spectrum of unrelated taxa suggest that this epidermal type is a genetically fixed func¬

tional adaptation rather than ontogenetically developed structures based solely on growth rates and other fac¬

tors.

There are two main morphological properties of ray ligule adaxial epidermal cells: their size and shape.

Functionally, these relate to the absorption of visible or ultraviolet light that plays a role in insect pollination

biology or the surface-to-volume ratio as the increased height increases the larger surface area to various exter¬

nal physical properties or physiological activity. It is likely that the interaction between these two selection

pressures, namely pollination biology and physiological adaptations, play a major role in shaping adaxial ray

ligule epidermal cells (Baagoe 1977a).

This was apparent when Noda et al. (1994) reported that petal conical cells in comparison to flat cells in¬

creased the proportion of incident light that entered epidermal cells, enhancing light absorption by the pig¬

ments (anthocyanins) and thus increasing the intensity of petal color. Experimental evidence came from a

mixta mutant of Antirrhinum majus that had petals composed of flat hexagonal-based cells that changed the

mutant cell morphology of petal color to a slightly paler and less velvety surface. Conical adaxial petal cells had

more sparkle that attracted bumble bee pollinators and a velvety surface that facilitated clinging to the petal

surface.

A more recent review paper by Whitney et al. (2011) noted the variation of conical adaxial epidermal cells

of Geranium procurrens, Helianthus annuus, and Hibiscus trionum. These epidermal cells were considered as

they relate to petal color, petal reflexing, petal scent production, petal wettability, and insect pollinator grip on

the flower surface. They recognized at the outset that conical epidermal cells found on adaxial surfaces of flow¬

ers are a special feature of petals rarely found on leaves or any other plant surfaces. Floral conical cells had the

ability to focus light into epidermal vacuoles that contain anthocyanins, increasing color saturation of the

petal and, with the scattering effect from the mesophyll, was more even than found in flat cells and therefore

resulted in a brighter sheen or velvety surface. A series of observations was made noting, (1) petal reflexing

with conical cells that stand more upright and presented a larger surface or target area to attract pollinators

(bumblebees), (2) flat-celled petals were more wettable than conical-celled petals, (3) conical cells were self¬

cleaning and aided in removal of dirt particles or potential fungal pathogenic spores, and (4) more recent con¬

vincing evidence shows that bumblebees can discriminate by tactile touch alone the difference between coni¬

cal and flat-celled surfaces on petals but have a clear preference for the conical surface (Whitney et al 2011).

Echinacea florets are visited and pollinated by bumblebees ( Bombus spp., H.W. Keller, pers. obs.), butter¬

flies, and longhorn beetles (Figs. 3B; 6D, E). The co-evolution of insect pollinators suggests there is selection

pressure toward specialized ray ligule adaxial epidermal cells in Echinacea as observed in the variation of cell

shapes and sizes. There are no flat adaxial epidermal ray flower cells in any Echinacea taxa. The presence of

curious multicellular ray ligule adaxial epidermal cells found only in E. angustifolia “race intermedia ,” and lack¬

ing in the closely related varieties angustifolia and strigosa, merits special consideration because these ana¬

tomical structures previously were not included in publications. Their presence or absence is complicated by a

limited sample size that may have overlooked their presence in other related taxa. Not knowing the cause or

frequency of such a micromorpholgical character in a colony, hybridized population, or species makes it diffi¬

cult to interpret its diagnostic significance and utility. A survey of the literature indicates that these trichomes

are rarely found on the adaxial surface of petals. Their function may be protective against desiccation or preda¬

tion, produce secretions, or as in this case of Echinacea, their function appears to be unknown. These cells are

Keller, Floral, stem, and petiole morphology of Echinacea

119

Fig. 11. Stem transections of Echinacea taxa. A. £ sanguined showing portion of vascular system. Note the absence of secretory canals and sclerotic

cells in the parenchymatous pith region, discrete collateral vascular bundles composed of primary tissue with fibrous bundle caps, partially scarified

interfascicular regions, one small secretory canal middle right in cortex composed mostly of collenchyma cells, and chlorenchyma cells underlying the

epidermis. This microtome section was stained with safranin and fast green (x50). B. £ angustifolia var. strigosa free hand section showing typical

thick-walled, elliptical collenchyma cells of outer cortical region (compare isodiametric parenchyma cells with intercellular spaces in pith see Fig. A). Note

secretory canal in lower cortex (star) (x25). C. £ pallida showing endodermis cell in longitudinal section. Note numerous starch grains with roughened

surfaces ~4.6 pm in diameter (x652). D-H. Stem transections showing secretory canals. D. £ atrorubens with secretory canal lumen surrounded by

thin-walled, rectangular, eight-celled, epithelial ring embedded in thick-walled sderified tissue next to the protoxylem points of the vascular bundle

(x430). E. £ atrorubens with secretory canal embedded in vascular bundle cap (x430). F. £ paradoxav ar. neglecta showing two secretory canals (stars)

opposite the vascular bundle cap near the cortical region (x430). G. £ paradoxam. neglecta with the largest secretory canal located near the protoxylem

points in the pith with a 12-celled epithelial ring (x430). H. £ simulata with paired secretory canals opposite protoxylem points near the pith (x430).

120

Journal of the Botanical Research Institute of Texas 8(1)

widely scattered and therefore do not cover extensive areas nor do they appear to be secretory cells. Additional

studies are justified to elucidate the occurrence and function of these cells.

Groups of Echinacea taxa may share a basic ray ligule adaxial epidermal shape. For example, the dome-

shape adaxial epidermal cell with a rounded apex was observed in E. laevigata, E. paradoxa var. neglecta, E.

paradoxa var. paradoxa, and more modified into a rounded but somewhat pointed apex in E. angustifolia var.

angustifolia, E. angustifolia “race intermedia,” E. angustifolia var. strigosa, E. pallida, and E. purpurea. These are

more or less transitional shapes that could be considered the same basic epidermal cell type. In contrast the

distinctive shapes of E. atrorubens (apex nipple-like), E. sanguinea (apex necked), and E. simulata (apex sharply

pointed conical) appear different from the dome-shaped epidermal cell type. The question remains, however:

how constant will these shapes remain over a broad range of habitats and given statistically significant sampled

populations?

Echinacea simulata has the largest (length and width) ray ligule adaxial epidermal cells in the genus (Fig.

4K, L). The similarity between E. pallida and E. simulata based on macromorphological characters can be dis¬

tinguished by adaxial ray ligule epidermal cells that appear to have distinctive shapes; the more dome shape in

E. pallida in contrast to the more conical shape in E. simulata. This anatomical character alone distinguishes

the two taxa even though the macromorphological characters have created confusion in the past.

Polyploidy, including tetraploids, with the increase in ploidy or sets of chromosomes, usually results in

more robust, larger plants in overall size (stem height and diameter, leaf, flowers) and a proportional increase

in cell size and volume. This increase in the component parts of the plant is called the “gigas effect.” Compari¬

son of ray floret anatomy of E. pallida (a tetraploid) with E. simulata (a closely related diploid with which it has

been confused in the past) resulted in no obvious differences attributed to ploidy. For example, the ligule ray

adaxial epidermal cells in E. pallida average 72 pm in length and 48 pm in width compared to E. simulata that

average 105 pm in length and 75 pm in width. There are no other consistent micromorphological characters

other than pollen size that suggest any differences in cell size between these two taxa.

Microanatomy of the ray florets of E. purpurea described by Upton (2007) noted (1) nonglandular tri-

chomes similar to those on the leaf, (2) papillose epidermal cells of the ligule, (3) secretory ducts along veins,

and (4) epidermis with wavy anticlinal walls, anomocytic stomata, and a light area indicating a secretory duct

beneath a vein. This ray floret description fails to indicate adaxial and abaxial surfaces of the ray ligule, and

indeed, the papillose surfaces in all the Echinacea taxa studied here were modified into various shapes and the

abaxial surface had more epidermal-like trichomes. The line drawing illustration in 6f shows ray ligule tri-

chomes similar to leaf trichomes (Upton 2007). Marginal ray florets in the Asteraceae usually have three teeth

(notched) at the apex with veins outlining the three teeth (Carlquist 1976).

Internal ray floret anatomical characters, apart from epidermal cells—such as the number of vascular

traces (usually 13), thickness of ray florets (170-312 pm), and number and location (adaxial or abaxial surfac¬

es) of secretory chambers—are more or less constant for all Echinacea taxa except for E. laevigata and E. pur¬

purea. Echinacea purpurea has the broadest ray florets of all the taxa and a more complex venation pattern that

distinguishes E. laevigata and E. purpurea based on ray floret anatomy. Here the difference in the size of the ray

floret width is reflected in the number of vascular traces (15 in E. laevigata and 31 in E. purpurea) and the abun¬

dant presence of secretory chambers in E. purpurea and apparent absence in E. laevigata (Table 3).

Stems

Rank ordering of stem diameters from the smallest to the largest was based on measurements in Table 3: E.

sanguinea, E. angustifolia var. angustifolia, E. angustifolia var. angustifolia “race intermedia,” E. angustifolia var.

strigosa, E. paradoxa var. neglecta, E. paradoxa var. paradoxa, E. atrorubens, E. simulata, E. pallida, E. purpurea,

and E. laevigata that follow a general trend of the higher the stem height the greater the stem width with the

exception of E. sanguinea that has the smallest stem diameter and a more spindly habit (Fig. 10J; Table 3).

Samples were taken at the height of anthesis. Comparison of stem diameters of all Echinacea taxa shows a

well-developed pith region usually occupying about 75 percent of the total stem diameter. This contrasts with

the findings of Upton (2007) who reported in the general habit description of E. purpurea a stem diameter of

Keller, Floral, stem, and petiole morphology of Echinacea

121

2-5 mm in transection with the pith hollow or solid. It is difficult to assess the importance and comparison of

these characters with the present study. All of the stem tissue maps and measurements given here show an

extensive pith region in all Echinacea taxa with no sign of tissue disintegration, which might occur later in the

growing season and account for a hollow stem. Transections taken later in the growing season in July and Au¬

gust could possibly result in a hollow pith region and connected interfascicular regions with cambial activity

and some secondary growth.

Perhaps the most striking micromorphological characters of the stem is the presence of sclerenchyma fi¬

bers (sclerotic cells with phytomelanin) in the pith and the absence of secretory canals in the pith and presence

in the cortex of E. angustifolia var. angustifolia, E. angustifolia “race intermedia ,” and E. angustifolia var. strigosa.

This distinguishes the E. angustifolia complex from all other Echinacea taxa.

The secondary xylem and phloem in roots of E. angustifolia, E. atrorubens, E. pallida, and E. purpurea have

sclerids with associated phytomelanin deposition (see table 3; figs. 5d, e; 6a, b, c, g in Upton 2010). This is

similar to the sclerotic cells with associated black substance in the stem pith tissue of E. angustifolia var. angus¬

tifolia, E. angustifolia var. strigosa and hybrids (Fig. 3H, I, J). Thus, in a review of all Echinacea taxa McKeown

(1999) noted that E. angustifolia var. angustifolia has short plant height, short and broad reflexed ray ligules,

yellow pollen color and is a selection candidate for cold hardiness because of adaptation to northern climates.

It is also a potential candidate in breeding for stem strength when coupled with the sclerihed pith tissue

(McKeown 1999).

Stem diameters and secondary growth patterns based on stem anatomy tissue maps appear most similar

when E. pallida and E. simulata are compared. Comparison of Echinacea root anatomy in E. pallida (Upton

2010, see Table 3) noted the presence of phytomelanin-coated sclerids in the secondary phloem and secondary

xylem in roots of E. angustifolia, E. pallida, and E. atrorubens, and the rhizome of E. purpurea pith and second¬

ary phloem. Another paper on the root anatomy of E. angustifolia (Axentiev et al. 2010, see Table 3) also noted

the presence of phytomelanin in the same locations as in E. pallida with the added description that this black

substance fills the triangular intercellular spaces around the sclerids, causing them to appear star-shaped not

unlike the stem anatomy described here for the E. angustifolia complex (Fig. 3H-J). Interestingly, in my ana¬

tomical study of E. atrorubens stem transections, sclerotic cells were not observed, but this taxon is a good

candidate to look for stem sclerotic cells since it hybridizes with E. angustifolia var. angustifolia.

Stem tissue maps for E. laevigata and E. purpurea show anatomical characteristics that distinguish these

two species from other Echinacea taxa. Numerous secretory canals in the pith (~58) and in the cortex (~48) of

E. laevigata compared to E. purpurea with (~34) in the pith and (~50) in the cortex are in contrast to all other

Echinacea taxa with far fewer numbers. These secretory canals originate only opposite the vascular bundles

and interfascicular region. Vascular tissue has 44 protoxylem points in E. laevigata and 42 protoxylem points

in E. purpurea, far greater numbers than in all other Echinacea taxa (Table 3).

The largest stem diameter of ~5.2 mm with pith diameter of ~4.2 mm occurs in E. laevigata (Fig. 10E)

compared to stem diameter of ~4.8 mm with pith diameter of ~3.6 mm in E. purpurea (Fig. 101). These two taxa

are the tallest and overall largest in the genus Echinacea. Stem tissue maps of these two taxa (Fig. 10E, I) are

similar in vascularization and spatial arrangement of the secretory system in development, position, and size.

Cortical tissue in both taxa is composed primarily of parenchyma. The epidermis differs in the presence

of trichomes in E. purpurea and absence in E. laevigata. The more robust habit of these two taxa would suggest

apriori that activity in interfascicular areas would connect the vascular bundles with some degree of sclerihca-

tion. However, vascular bundles are discrete, surrounded and separated by regions of parenchymatous tissue.

Sclerotic cells are lacking in the pith of both species. It appears that macromorphological differences are in

contrast to internal microanatomical similarities.

Petioles

In Echinacea the petiole is supplied by three major collateral vascular bundles, a manifestation of departing

foliar traces from the stem. In addition petiole shape can be used as a taxonomic character. Transversely cut

petioles can be recognized by shapes, for example, horseshoe-shaped, V-shaped, and cylindrical-shaped.

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Journal of the Botanical Research Institute of Texas 8(1)

Fig. 12. Petiole transections. A. £ atrorubens, microtome section showing three major collateral vascular bundles ensheathed by fibrous tissue (vascular

traces) typical for the Asteraceae. Note three air spaces that form passageways throughout the length of the petiole (x20). B. £ sanguined, microtome

section showing central major vascular bundle lacking fibrous tissue on abaxial side and two small secretory canals one on each side (arrows). Note the

two-layered epidermis with similar cells and undifferentiated ground mesophyll (xl 7). C. £ paradoxam. neglecta, lunate to bow-shape with venation

illustrated by line drawing (xl 2). D. £ paradoxam. neglecta, oblique microtome section showing brachysderids (stone cells) as scattered single cells or

in rows (arrows). Larger secretory canal (star) visible near vascular trace (x40). E. £ paradoxa var. neglecta, oblique microtome section showing single,

isolated stone cells with highly refractive thickened cell walls with reduced central lumen. Note the primary pit fields that radiate like the spokes of

a wheel (arrows) (x70). F. £ paradoxam. neglecta, microtome section stained with red safranin and fast green showing central vascular bundle and

cluster of stone cells at lower left (x14).

Keller, Floral, stem, and petiole morphology of Echinacea

123

Moreover, secretory canals, universally present in the genus, differ in size, number and position in the petiole.

The petiole contains the same tissues as the stems that include epidermis, collenchyma in varying amounts,

and vascular bundles with associated fibrous sheaths. However, significant differences were observed for sev¬

eral taxa, for example, in E. purpurea and E. sanguined the medial vascular bundle lacks a fibrous cap (Fig. 12B;

Table 4).

Another apparently unique structure found in E. paradoxa var. neglecta is the brachysclerid or stone cell.

They appear either isolated, clustered, or in rows (Fig. 12D-E). Structurally the petiole of E. atrorubens has

three lacunae situated around the medial vascular bundle (Fig. 12A) and this differs from other Echinacea taxa.

General Habit

Echinacea has mostly a scapose habit with above ground stem that persists in to the fall along with the flower

head that gradually dies and begins to undergo decay. Stems do not survive the winter in the high plains region

of western Kansas due to freezing temperatures and heavy snows that cause lodging, and eventually aerial

parts deteriorate into ground litter as part of the annual life cycle. The stems are not stout enough to withstand

the elements of nature unlike species of Yucca where stems may survive for several years. In grassy habitats

such as prairies and glades Echinacea populations can dominate the landscape, (Figs. 5C-D; 8A-C) represent¬

ing the tallest elements not unlike the forest canopy of trees (Kilgore et al. 2009; Richter 2013). This standing

cover of stems serves to intercept spores of windblown organisms such as myxomycetes (plasmodial slime

molds).

The round spiny cone head also creates a surface area and landing platform for spores. Indeed, a recent

study that is the first of its kind (Kilgore et al. 2009) found that Echinacea species collected from native prairies

in Kansas and Missouri (E. angustifolia, E. pallida , and E. paradoxa var. paradoxa) have a distinct assemblage of

myxomycete species that were isolated in moist chamber cultures. Reproductive structures of herbaceous

plants had greater mean species richness than stems. This study proposed a new term (herbicolous myxomy¬

cetes) for herbaceous, perennial grassland plants associated with this group of myxomycete species (Kilgore et

al. 2009). This above ground canopy of perennial plants should be explored for potential species diversity and

possible myxomycete species new to science.

CONCLUSIONS AND FUTURE CONSIDERATIONS

This study took advantage of more than 15 years of research on the biology of Echinacea by Professor Ronald L.

McGregor at the University of Kansas. His held experience collecting plants throughout the state of Kansas and

as Coordinator of The Flora of the Great Plains research project (Great Plains Flora Association 1986) gave him

a “trained eye” for habitats and species identification, especially Echinacea taxa involving hybrid swarms, in-

trogressed populations, and pockets of typical taxa in part represented here. All of the plants selected for this

study were identified personally by R.L. McGregor. Freshly collected plants were used for this anatomical

survey not herbarium specimens. This avoided using material that often shows shrinkage and distortion of

cells. Therefore, the names and collections used here carry the accuracy and weight of McGregor’s held experi¬

ence.

All of the plants studied here were collected within a few days at the height of anthesis and for the most

part in the same place. This part of the sampling protocol assures that possible variables were at least partially

controlled. Nevertheless, the limited sampling lacked a statistically signihcant number of plant individuals,

different populations from different natural habitats, and ray florets from different capitula. The sample size

places limitations on the use of anatomical characters in dichotomous keys, and therefore, the combination of

macro- and microcharacters used in couplets here. Furthermore, the presence of anatomical characters, while

potentially important in separating taxa—for example, sclerotic pith cells, brachysclerids in petioles, and dif¬

ferences in shapes of adaxial ray ligule epidermal cells—suggests that special caution should be used in the

assumption that these characters are consistent within and across Echinacea taxa. This is especially true of the

multicellular, catenulate, adaxial ray ligule epidermal cells observed in the hybrid population of E. angustifolia

var. angustifolia “race intermedia ” that are present but scattered in small numbers. These epidermal cells were

124

Journal of the Botanical Research Institute of Texas 8(1)

not observed in either of the closely related E. angustifolia var. angustifolia or E. angustifolia var. strigosa.

It is apparent that E. laevigata and E. purpurea are closely related in anatomical characters that include:

stem diameter, cortical breadth comprised mostly of parenchyma cells, vascularization and spatial distances of

tissue systems, diameter of pith, and position, number and size of secretory canals. In contrast ray florets differ

in the number of veins and secretory chambers, but adaxial epidermal cells have similar shapes and sizes noted

previously

Future studies should concentrate on fewer taxa and more critical sampling to more accurately assess the

value of anatomical characters useful in the identification of Echinacea taxa. Nevertheless, this anatomical

study of Echinacea aerial plant parts contributes to a better understanding of structure and function that relates

to similarities and differences in this medically important herbaceous perennial prairie plant.

ACKNOWLEDGMENTS

My gratitude goes to Professor Ronald L. McGregor (recently deceased) for his guidance and encouragement

throughout the course of this study. His suggestion of the problem, helpful criticisms, and stimulating discus¬

sions enhanced the completion of this Master’s Thesis. Credit goes to Azar Kroshovi for drawing the diagrams

of stem and leaf petioles. Professor W.H. Horr gave advice with regard to sharpening and honing microtome

blades. Dr. J.E. Fox assisted with editing the manuscript. And lastly, my appreciation goes to my colleagues and

family for their helpfulness in countless ways. In addition to these acknowledgments that were recorded dur¬

ing the preparation of my thesis the following individuals have contributed to the preparation of this paper by

giving advice, aiding in finding literature sources, reading and editing multiple manuscript drafts: Brooke Best,

Uno Eliasson, Craig Freeman, Kelly Kindscher, Barney Lipscomb, Will McClatchey, Guy Nesom, Kurt Neubig,

Par Ollis, Relf Price, and Sula Vanderplank (who also prepared the Spanish abstract). Brooke Best merits spe¬

cial recognition for the preparation of the tables, illustrations, plates, editing, and formatting for the final

camera-ready version of this paper. Craig Freeman answered questions and provided additional information

about the Echinacea specimens in the R.L. McGregor Herbarium. Kim Norton Taylor prepared the distribution

maps. Richard Sjolund scanned and digitized the black and white 35mm negatives. Marianne Reed and staff at

the University of Kansas Libraries’ Office of Scholarly Communication & Copyright were responsible for digi¬

tizing my thesis and making it permanently available in KU Scholar Works (http://hdl.handle.net/1808/! 1669).

Bob O’Kennon provided held observations and color images of E. sanguinea. The following provided color

plant images from natural habitats: Kelly Kindscher (E. angustifolia var. angustifolia), Chris Crabtree (E. para-

doxa var. paradoxa and E. pallida), Todd Crabtree (E. tennesseensis), and Alan Cressler ( Echinacea laevigata).

Theo Witsell, Botanist at the Arkansas Natural Heritage Commission, assisted with color images of Echinacea

in Arkansas. All photographs not credited were taken by the author. The author wishes to thank two anony¬

mous reviewers who improved the text content and sentence syntax. This paper is dedicated to Ronald L. Mc¬

Gregor for his many years of research devoted to Echinacea, the Bora of Kansas, and the Bora of the Great Plains

region.

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THE FLORAL STRUCTURE OF THREE WEEDY SPECIES OF SIDA (MALVACEAE)

Junior Cezar Muneratto

Luiz Antonio de Souza

Universidade Estadual de Maringa

Departamento de Biologia

Av. Colombo, 5790

(87020-900) Maringa, PR, BRAZIL

Universidade Estadual de Maringa

Departamento de Biologia

Av. Colombo, 5790

(87020-900) Maringa, PR, BRAZIL

lasouza@uem.br

Odair Jose Garcia de Almeida

Instituto de Biociencias

UN ESP - Univ. Estadual Paulista

Programa de Pos-Graduaqao em Biologia Vegetal

Av. 24 A, 1515, (13506-900) Rio Claro, SP, BRAZIL

odair 1000@hotmail.com

ABSTRACT

Malvaceae sensu lato is monophyletic and is characterized by the presence of nectaries of glandular trichomes, located internally at the base

of the calyx. The genus Sida has been heterogeneous since its origin; however, there has been a reduction in the size of the genus with the

removal of many species to other genera, giving a more natural and definable residual group subdivided into 11 sections. The aims of this

work were to describe and compare the floral structure of three weedy species Sida rhombifolia, S. urens, and S. regnellii, using a traditional

anatomical approach, along with scanning electron microscopy analysis. The comparative study of the three Sida species allows us to sug¬

gest that some structural characters are shared, i.e., the hypodermis in the sepal and ovary wall, trichomatous calyx nectary, perianth with

homogeneous mesophyll, and anther with a middle layer and plasmodial tapetum type, and they may reinforce the natural residual group of

Sida. On the other hand, other floral characters have some diagnostic value for classification at the species level, such as the style structure

being solid or rift in the center.

Key Words: Floral anatomy, calyx nectary, trichomes, weedy species

RESUMO

Malvaceae sensu lato e monofiletica e se caracteriza pela presenga de nectarios que consistem de tricomas glandulares localizados interna-

mente na base do calice. O genero Sida e considerado como heterogeneo na sua concepgao inicial, mas ultimamente ele tern sido reduzido de

tamanho com a remogao de varias especies para outros generos, tornando-o um grupo natural residual subdividido em 11 segoes. O objetivo

deste trabalho foi descrever e comparar a estrutura floral de tres especies invasoras, Sida rhombifolia, S. urens e S. regnellii, utilizando-se de

estudo anatomico tradicional e analise em microscopia eletronica de varredura. O estudo comparative das tres especies de Sida permite

sugerir que alguns caracteres estruturais sao comuns as especies, como presenga de hipoderme na sepala e parede do ovario, nectario cali-

cino tricomatoso, perianto com mesofilo homogeneo, e antera com uma camada media e tapete plasmodial, o que reforga o genero Sida como

um grupo natural residual. Por outro lado, outros caracteres florais tern algum valor diagnostico para separagao das especies, como, por

exemplo, a estrutura do estilete que pode ser solido ou apresentar fenda na regiao central.

INTRODUCTION

When circumscribed broadly to include four families (i.e., Tiliaceae, Sterculiaceae, Bombacaceae, and Malva¬

ceae - sensu stricto), Malvaceae sensu lato is monophyletic (Judd et al. 2002). It is characterized by the pres¬

ence of nectaries of glandular trichomes, located internally at the base of the calyx or, with less frequency, on

the petals or on the androgynophore (Judd & Manchester 1997). This circumscription is also supported by

adnation of the androecium to the corolla and usually the presence of unilocular anthers, although Tiliaceae

and Sterculiaceae often retain numerous stamens with 2-locular anthers (Judd et al. 2002). More recently,

Malvaceae sensu lato has been divided into nine subfamilies (as discussed in Tate et al. 2005): Bombacoideae

(formerly Bombacaceae, in part), Brownlowioideae, Byttnerioideae, Dombeyoideae, Grewioideae, Helicteroi-

deae, Malvoideae (formerly Malvaceae), Sterculioideae (formerly Sterculiaceae, in part), and Tilioideae (for¬

merly Tiliaceae, in part). The Malvoideae are sometimes called the “core” Malvaceae s.str., and some authors

J. Bot. Res. Inst. Texas 8(1): 127 -137.2014

128

Journal of the Botanical Research Institute of Texas 8(1)

continue to retain this family name for this core group alone. Prior to that, Bayer and Kubitzki (2003) divided

the subfamily Malvoideae into four tribes: Gossypieae, Hibisceae, Kydieae, and Malveae. Traditionally, mem¬

bers of the Malveae have been characterized by a combination of several morphological characters: schizocar-

pic fruits (sometimes a capsule), three to over 20 mericarps with an equal number of free styles, antheriferous

apex of the staminal column and the absence of lysigenous cavities (“gossypol glands”) (Fryxell 1988; Bayer &

Kubitzki 2003; Tate et al. 2005).

According to Fryxell (1985), the genus Sida has been heterogeneous since its origin as botanists tended to

put into Sida any member of the Malvaceae that are uniovulate and without an involucel. However, there has

been a reduction in the size of the genus with the removal of many species to other genera, giving a more natu¬

ral and definable residual group. Fryxell (1985) subdivided this residual group into 11 sections.

The aims of this study were to describe and compare the floral structure of three weedy species of Sida ,

namely S. rhombifolia L., S. urens L., and S. regnellii R.E. Fr. Economically, the Malvaceae s.l. are important ei¬

ther ornamentally (for instance, the genera Alcea, Hibiscus, and Malvaviscus) or in the textile industry (e.g.

Gossypium and Urena ), or as weedy plants, such as Sida, a genus that can be very detrimental to the agricul¬

tural economy (Bovini et al. 2001).

MATERIAL AND METHODS

Plant material

Floral buds and flowers of the three weedy species were collected in the city of Maringa, Brazil (state of Parana):

S. rhombifolia at 506 m altitude, 23°24T3.3" latitude and 51°56'21.17" longitude; S. regnellii at 559 m altitude,

23°24'43.0" latitude and 51°56'29.6" longitude; and S. urens at 518 m altitude, 23°24'13.1" latitude and

51°56'20.4" longitude. Voucher materials were identified by Massimo Giuseppe Bovini (RB) and deposited at

HUEM (Herbarium of the Universidade Estadual de Maringa) with these accession numbers: S. urens:19909 ( J.

Muneratto 001); S. rhombifolia: 19910 (J. Muneratto 002); and S. regnellii:19911 (J. Muneratto 003).

Anatomical analysis

The material was fixed in glutaraldehyde (1% in 0.1M phosphate buffer, pH 7.2) and then conserved in 70%

ethanol (Johansen 1940). The fixed material was embedded in historesin (Gerrits 1991), sectioned (cross- and

longitudinally) in a rotation microtome, and stained in toluidine blue (O’Brien et al. 1964). Specific micro¬

chemical tests were done for lipid substances (Sudan IV and Sudan Black), tannins (ferric chloride), starches

(iodine-potassium iodide test), lignins (phloroglucin test) and calcium crystals (sulfuric acid) (Sass 1951;

Rawlins & Takahashi 1952; Ruzin 1999). Images were taken using Leica ICC50 and Olympus BX50 optical

microscopes with digital camera attachments.

Scanning Electronic Microscopy analysis

Micromorphological analysis of the floral buds and flowers was done with material fixed in Karnovsky solu¬

tion (Karnovsky 1965). Samples were processed and then mounted on aluminum stubs, gold coated, examined

using scanning electron microscopy (Shimadzu SS-550 Superscan), and digitally photographed.

RESULTS

Perianth

The flowers (Fig. la-c) in all three of the Sida species investigated are actinomorphic, hermaphrodite, with

yellowish (S. urens and S. rhombifolia - Fig. la,b) and whitish (S. regnellii - Fig. lc) petals, distinct from the green

calyx (Table 1).

The five sepals are green in color, connate in the basal half, and with free triangular lobes. The uniseriate

adaxial epidermis is made of cuboid or tabular cells with outer and inner periclinal walls thicker than the an¬

ticlinal ones (Fig. 2a,a’). The abaxial sepal epidermis is uniseriate with trichomes and stomata in all species and

has cells with sinuous anticlinal walls in S. rhombifolia and S. urens. The stomata found in the sepals of the

species are anisocytic and paracytic in S. regnellii and S. rhombifolia and anisocytic in S. urens (Table 1). The

trichomes are non-glandular and glandular (Fig. ld,f,g-i). Unicellular non-glandular trichomes occur only in

Muneratto etal., Floral structure of Sida species

129

Fig. 1. General morphology of the flower in frontal view of Sida rhombifolia (a), 5. urens (b), and 5. regnellii (c); scanning electron micrographs (SEM)

of the calyx surface showing trichomes in 5. regnellii (d-g), d. Abaxial face, apical region, e. Nectary (trichomes). f-g. Abaxial face. Sida urens (h-i).

h. Adaxial surface, i. Abaxial face. Arrowheads indicate unicellular trichomes, long arrow shows stellate trichomes, and short and interrupted arrows

indicate glandular trichomes with a small and long stalk, respectively. Scale bars: 0.5 cm (a,c), 1.5 cm (b), 50 pm (d,f,g), 100 pm (e,h,i).

Table 1. Flower characters which have diagnostic value for separation of Sida rhombifolia; Sida urens, and Sida regnellii.

Characters

Sida rhombifolia

Sida urens

Sida regnellii

Flowers

Yellowish

Whitish

Sepal epidermis abaxial

Sinuous anticlinal walls

Straight anticlinal walls

Sepal stomata

Anisocytic/paracytic

Anisocytic

Anisocytic/paracytic

Unicellular nonglandular trichomes in sepals

Absent

Present

Absent

Sepal spongy parenchyma

2-3 cell layers

2-6 cell layers

4-5 cell layers

Stalk base/glandular trichomes (nectary)

3-4 layers

1-2 layers

Petal shape

Asymmetric

Obcordate

Asymmetric

Petal fasciculate trichome

Present

Absent

Present

Ovary carpels

9-11

Ventral vascular bundles of the ovary

9-11

Style

Hollow/solid in the base

Solid

Solid with reduced rift

S. urens (Fig. li), and this species has epidermal cells completely surrounding the base of the trichome. Stellate,

multicellular, non-glandular trichomes occur on the sepals of all three species (Fig. lg). Also occurring in

these three species are multicellular glandular trichomes (Fig. lg-i), consisting of a small stalk and a head of

several secretory cells or a long stalk and a unicellular secretory head. In the adaxial surface of the Sida sepals

are found crystalliferous cells (druses) in the hypodermis, which have a U-shaped wall thickness (Fig. 2a,a’).

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Journal of the Botanical Research Institute of Texas 8(1)

The S. rhombifolia has a lesser developed hypodermis in the abaxial surface (Fig. 2a,a’). The mesophyll (Fig. 2a)

has spongy parenchyma with mucilaginous cells and cavities, exhibiting two or three cell layers in S. rhombi¬

folia, four or five in S. regnellii, and two to six in S. wrens.

The nectary (Fig. le; 2b-c) occurs close to the base of the calyx (Fig. le) and is a carpet-like tissue made

of multicellular clavate glandular trichomes and a subtending reduced secretory parenchyma, which is sup¬

plied by xylem and a greater proportion of phloem elements. The glandular trichomes have a uniseriate or

pluriseriate stalk and a unicellular head (Fig. 2c). The trichomes are longer in S. rhombifolia than in the other

species; the stalk base of this S. rhombifolia and S. urens has a thickness of three or four cell layers, while S.

regnellii has a base with one or two layers (Table 1).

The corolla shows adnation to the androecium, and the petal shape varies among the Sida species. Sida

urens has symmetrical obcordate petals (Fig. lb; 2f), while in S. rhombifolia (Fig. la) and S. regnellii the petals

are obcordate but very asymmetrically so. The petal epidermis of S. rhombifolia and S. regnellii (Fig. 2d,e, re¬

spectively) is uniseriate with mucilage idioblasts and glandular and non-glandular trichomes. The mesophyll

(Fig. 2d,e) is composed of a homogeneous parenchyma with mucilage cells and cavities. The petal vasculariza¬

tion (Fig. 2d,e) consists of a central bundle and six to nine minor bundles.

Androecium

The androecium consists of filaments joined to form a tube (filament tube) (Figs. 3b; 5e) that surrounds the

style (Figs. 4a,c; 5b-f). The anthers (Fig. 3a) are dorsihxed, bisporangiate, longicidal, and septate monothecal.

The filament tube is composed of internal and external uniseriate epidermis of cuboid or elongated cells and

capitate glandular trichomes; the inner parenchyma has mucilage cells (Fig. 4a,c), and the vascular system

consists of 10 collateral vascular bundles arranged in pairs.

The wall of the young anther (Fig. 3c) has epidermis, endothecium with thin-walled cells, a middle layer

of elongated cells and a tapetum made of a layer of cells. Later, the cell walls of the tapetum undergo disintegra¬

tion, and the protoplasts of the tapetum cells remain available among the pollen grains during their develop¬

ment (Fig. 3d). In the developing anther, the endothecium cells acquire thickenings in the anticlinal walls and

in the inner periclinal wall, the middle layer disintegrates, and the tapetum protoplast is absorbed (Fig. 3e, f).

The connective consists of papillose epidermis, parenchyma with druse idioblasts and one vascular bundle.

Gynoecium

The structure of the style differs among the three species (Table 1) when seen in cross section. In S. rhombifolia

the style is hollow for the most part (Fig. 4b), being solid only in the base; in S. regnellii the style has a reduced

rift (Fig. 4c); and S. urens, in turn, has an entirely solid style (Fig. 4d). The style is composed of uniseriate epi¬

dermis, parenchyma and strands of transmitting tissue, one for each carpel, with 11 in S. rhombifolia (Fig. 4b)

and five in the other two species (Fig. 4d). The stigma consists of papillate epidermis (Fig. 4e,f).

The ovary presents many carpels and locules (Fig. 5a-e) (five in S. regnellii and S. urens, and nine to 11 in

S. rhombifolia) (Table 1). There is one ovule in each locule, and the placentation is axial (Fig. 5b-f). The style

consists of a single column (solid or rifted, depending of the species, according to the description above) (Fig.

5b-f). Close to the apex the style starts to lose the fused shape and separates into free branches, which corre¬

spond to each stigma lobe, in which the number of stigma lobes and locules is the same. The ovary wall (Fig.

6a-f) has glabrous uniseriate outer epidermis with cuboid cells in S. regnellii and S. urens, and cuboid to cylin-

dric in S. rhombifolia. The ovary mesophyll (Fig. 6a-f) is parenchymatous, spongy and homogeneous with two

or three cell layers in the middle region; the remainder (base and apex) of the ovary consists of five cell layers.

A crystalliferous hypodermis (druse cells) is present in the wall ovary (Fig. 6b), although some cells lack crys¬

tals in S. urens. The inner epidermis undergoes cell division periclinally to the ovary surface resulting in a tis¬

sue with two or three cell layers (Fig. 6b), the precursor of the endocarp.

In the ovary, all species studied here had septa (Fig. 6a,c,e) made up of an epidermis of one or two layers

of cells and bi- to multilayered spongy parenchyma tissue with scattered druse idioblasts. The ovary vascula¬

ture is made of a median dorsal bundle (Fig. 6b,d), one or two lateral bundles, and a ventral or placental bundle.

The number of ventral bundles differs among the species, with nine to 11 bundles in S. rhombifolia (Fig. 6e) and

Muneratto etal.. Floral structure of Sida species

131

Fig. 2. Perianth structure of 5. rhombifolia (a,a',d), 5. regnellii (c,e), and 5. urens (b,f), in cross-section and scanning electron micrograph (SEM). a. Calyx,

a'. Details of the epidermis, b. Nectary in SEM. c. Nectary longitudinal section, d-e. Petals, f. Petal base in SEM. Arrow indicates hypodermis cells with

U-shaped wall thickness. Scale bars: 100 pm (a,c,e,f), 30 pm (a'), 50 pm (b,d).

five in S. regnellii (Fig. 6c) and S. urens, which corresponds to the same number as the carpels and styles, and

the same numbers of mericarps in the developed fruit (Muneratto & Souza 2013). In the parenchyma close to

the ventral bundles (Fig. 6a,c,e), there are mucilaginous and druse cells. The ovules (Fig. 6a-f) are bitegmic,

crassinucellate, with an outer integument made up of two layers of cell, and multiseriate inner integument.

„ „ r ,

The trichomatous calyx nectary of the Sida species analyzed here is one of the features that characterize the

core Malvales (Malvaceae s.l. including Bombacaceae, Malvaceae, Sterculiaceae, and Tiliaceae) (Vogel 2000;

132

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 3. Anther structure and filament tube of 5. regnellii (a,b,f); anther structure of 5. urens (c) and 5. rhombifolia (d,e). a. Mature anthers - SEM. b.

Apex of the filament tube with glandular trichomes - SEM. c. Immature anther in cross-section, d. Anther detail visualizing plasmodial tapetum among

microspores, e. Immature anther in cross-section without middle layer and tapetum. f. Mature anther in cross-section with septum. * indicates the

stigma branches/lobes. White arrow shows septum of the anther. Black arrow indicates the tapetum protoplast among microspores in development,

(en = endothecium; ml = middle layer; ta = tapetum). Scale bars: 200 pm (a), 50 pm (b-f).

Leitao et al. 2005). According to Vogel (2000), the trichomatous calyx nectary may have evolved from hyda-

thodes, which provide moisture to protect floral primordia from desiccation, being active in the bud stage. As

discussed by Bernardello (2007), trichomes are by far the most common epidermal nectaries in Angiosperms,

and the nectariferous trichomes may have taxonomic importance in defining related plants groups, such as

Bombacoideae and Malvoideae.

Muneratto etal., Floral structure of Sida species

133

Fig. 4. Filament tube and stigma structure of 5. rhombifolia (a,b), 5. regnellii (c,e-f), and 5. urens (d). a-d. Cross Sections, e. Longitudinal view. f.

Longitudinal section, a. Apex of the filament tube surrounding the stigma branches/lobes, with detail of the epidermis and vascular bundles of the

filament tube. b. Style in cross sections - middle region - with detail of the transmitting tissue, c. Filament tube surrounding the style - middle region,

d. Style - middle region, e. Stigma in SEM. f. Stigma. Asterisk indicates the strands of transmitting tissue, (fb = stigma lobes; st = style). Scale bars:

100 pm (a-d,f),50 pm (e).

The calyx of the three Sida species may develop different functions, acting as a photosynthetic organ, a

protective organ of floral bud, or an attraction for pollinators. The photosynthetic function is related to the

presence of chlorenchyma in the sepal mesophyll. The protective function of the sepal may be executed by the

differentiation of a crystalliferous hypodermis with U-shaped wall thickening cells. Attraction of pollinators

may be performed by means of nectaries that consist of glandular trichomes located at the base of the sepals.

The broad variety of functions of the calyx is known in some groups of plants, such as Fabaceae, Lamiaceae,

134

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 5. Morphology of the ovary of 5. rhombifolia (a,b), 5. urens (c,d), and 5. regnellii (e,f). a, c, e. Ovary/gynoecium in SEM. b, d, f. Flower in longitudinal

section showing ovary and other floral organs, (an = filament tube; ne = nectary; pe = petal; se = sepal; st = style). Scale bars: 500 pm (a,e) 250 pm

(b,c,d,f).

Malpighiaceae, and Ranunculaceae (Roth 1977; Weberling 1992; Endress 1994). Unfortunately we do not have

supplementary data concerning the time of activity about the trichomatous calyx nectary in the Sida species

nor information concerning the visitors of these flowers, in order to ascertain the role of these nectaries in the

pollination system of these species. However, we can assume a nuptial function (i.e., they are involved in pol¬

lination events, sensu Almeida et al. 2013) of the carpet-like nectaries on the sepals of Sida species, since the

flowers of Malvaceae s.l. (Vogel 2000) do not have an annular nectary (nectariferous disk) to attract the pollina-

Muneratto etal., Floral structure of Sida species

135

Fig. 6. Ovary and ovule structure of 5. urens 5. regnellii (c, d), and 5. rhombifolia (e) in cross-sections (a-e) and longitudinal section (f). (ca =

calyx; co = corolla; oi = outer integument; ov = ovule; ow = ovary wall). Asterisk indicates the vascular bundle, and the black arrow shows the septum.

Scale bars: 50 pm.

tors, and these nectaries can be accessed via the gaps at the base of the corolla lobes in the region of adnation

between petals and the filament tube (Vogel 2000; Bernardello 2007).

Dahlgren (1991) introduced a classification based on Davis (1966) of the different types of anther wall

formations, in which four types were presented: basic, dicotyledonous, monocotyledonous, and reduced. He

reported the basic type of anther wall formation, characterized by two middle layers, for some taxa of Malvales.

In contrast, the Sida species studied here has only one single middle layer in the anther wall, placing it clearly

136

Journal of the Botanical Research Institute of Texas 8(1)

in the dicotyledoneous type. Other species of Malvaceae also have only one middle layer in the anther wall,

such as Sida cordifolia L. (Rao 1954) and Modiolastrum malvifolium (Griseb.) K. Schum. (Galati et al. 2007), in¬

dicating the necessity for new studies concerning the anther development in this group of plants.

In all Sida species investigated here, the tapetum is of the plasmodial type, although fusion between pro¬

toplasts of the microspores was not observed. Embryological studies in Malvaceae (Rao 1954) showed that as

the microspores enlarge, the protoplasts of the tapetum cells become smaller and smaller until they are ab¬

sorbed; in addition, according to Rao (1954), they may disappear or their remnants may persist until the pollen

grains have become 2-nucleate and nearly mature, as shown in Sida carpinifolia Mill.

The style may be hollow or solid, depending on the degree of closure of the fused or free carpels (Fahn

1990). The analyzed styles of the Sida flowers are quite distinct. The solid type is restricted to S. wrens , and hol¬

low type with solid base occurs in S. rhombifolia. Unlike these two species, S. regnellii has a narrow single canal

occupying the center of all styles. The style structure might be a reasonable alternative to the identification of

the Sida species. In fact, there has been little research dealing with the style structure. Nevertheless, Souza et

al. (2001) found similar results in three species of Trichilia P. Browne (Meliaceae); that is, T. elegans A. Juss. has

a solid style, T. catigua A. Juss. a hollow style, while T. pallida Sw. has a narrow canal in its style. This variation

in the style structure among species of the same genus may occur in other genera from the clade Eurosids II.

Our results showed mucilaginous cells and cavities in the flowers of the three species, which usually oc¬

curs in the Malvaceae (Metcalfe & Chalk 1957; Esau 1974; Fahn 1979). However, there are differing opinions

concerning the development of cavities in some taxa, as pointed out by Evert (2006). The cavities present in the

flowers of Malvaceae have been reported as both schizogenous and lysigenous (Fahn 1979 and references

therein). Metcalfe and Chalk (1957) recorded lysigenous secretory glands (cavities) in the leaves of Malvaceae

species. Even though for Evert (2006) the concept of lysigenous cavity development is questionable, here, we

consider it as lysigenous since they are formed by the disintegration of mucilaginous cells and surrounding

parenchymatous cells.

The comparative study of these three Sida species allows us to suggest that some structural characters are

shared, i.e., the hypodermis in the sepal and ovary wall, trichomatous calyx nectary, perianth with homoge¬

neous mesophyll, anther with a middle layer and plasmodial tapetum type. These characters may reinforce the

natural residual group of Sida as proposed by Fryxell (1985). On the other hand, Table 1 shows some floral

characters which may have some diagnostic value for classification at the species level, such as the style struc¬

ture being solid or rift in the center.

ACKNOWLEDGMENTS

We thank CAPES (Coordenagao de Aperfeigoamento de Pessoal de Nivel Superior, Brazil) and CNPq (Con-

selho Nacional de Desenvolvimento Cientlhco e Tecnologico, Brazil) for the support granted to realize this

study. The authors are grateful to Denver Falconer, Steven R. Hill, and an anonymous reviewer for critical re¬

view of early versions of the manuscript. We also would like to thank Massimo Giuseppe Bovini for the identi¬

fication of the voucher materials.

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parales, and nonbetalain Caryophyllales. Springer-Verlag, Berlin, Germany. Pp. 225-311.

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Esau, K. 1974. Anatomia das plantas com sementes. Edgard Blucher, Sao Paulo, Brazil.

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malvifolium (Malvaceae). Ann. Bot. 99:755-763.

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138

Journal of the Botanical Research Institute of Texas 8(1)

BOOK NOTICE

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Veteran botanist and educator Roy L. Lehman describes the plants in four major sections, covering the

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ROY L. LEHMAN is professor of biology at Texas A&M University-Corpus Christi, where he is also director of

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Texas A&M University Press in 2005.

J.Bot. Res. Inst. Texas 8(1): 138.2014

A GASTEROID FUNGUS, PALAEOGASTER MICROMORPHA GEN. & SP. NOV.

(BOLETALES) IN CRETACEOUS MYANMAR AMBER

George 0. Poinar, Jr. Donis da Silva Alfredo

Department of Integrative Biology Graduate Program in Systematics and Evolution

Oregon State University Department of Botany and Zoology Center of Biosciences

Corvallis, Oregon 97331, U.S.A. Universidade Federal do Rio Grande do Norte

poinarg@science.oregonstate.edu BRAZIL

luri Goulart Baseia

Department of Botany and Zoology

Center for Biosciences

Universidade Federal do Rio Grande do Norte

BRAZIL

ABSTRACT

A new genus and species of gasteroid fungus, Palaeogaster micromorpha gen. & sp. nov. is described from Early-Mid Cretaceous amber

from the Republic of Myanmar. The species is represented by some 25 complete or partial fruiting bodies in various developmental stages.

Diagnostic characters for the new taxon are its small size, the globose to pyriform shape of the fruiting bodies, mycelial hyphae with clamp

connections and small globose to subglobose spores. It is assigned to the Order Boletales (Sclerodermatineae) and possesses many features

of the family Sclerodermataceae, which includes the earthballs and hard skinned puffballs. Palaeogaster micromorpha represents the first

fossil member of the Sclerodermatineae and the oldest known gasteroid fungus.

RESUMEN

Se describen un genero y especie nuevos de hongo gasteroide, Palaeogaster micromorpha gen. & sp. nov. del ambar del cretacico tempra-

no-medio de la Republica de Myanmar. La especie esta representada por unos 25 cuerpos fructiferos completos o parciales en varios estados

de desarrollo. Los caracteres diagnosticos del nuevo taxon son su pequena talla, cuerpos fructiferos de forma globosa a piriforme, hifas del

micelio fibuladas y esporas globosos a subglobosas pequenas. Se asigna al Orden Boletales (Sclerodermatineae) y tiene muchas caracteristi-

cas de la familia Sclerodermataceae, que incluye los bejines. Palaeogaster micromorpha representa el primer miembro fosil de las Scleroder¬

matineae y el hogo gasteroide fosil mas antiguo conocido.

INTRODUCTION

Aside from containing a variety of animal and plant fossils, amber from Myanmar includes some interesting

fungal remains, such as the Hymenomycete, Palaeoclavaria burmitis Poinar & Brown (2003) and one of the

earliest known mushrooms, Palaeoagaracites antiquus Poinar and Buckley (2007). The present study describes

a gasteroid fungus preserved in Myanmar (Burmese) amber. Fossil gasteroids, which include puffballs, earth-

balls, earthstars and stinkhorn fungi, are exceedingly rare with previous records limited to Lycoperdites tertia-

rius Poinar (2001), from Tertiary Mexican amber, a Late Cenozoic earthstar (Geasteraceae) from Pueblo,

Mexico (Magallon-Pueble & Cervallos-Ferriz 1993) and a subfossil from Holocene deposits in Alaska (Chaney

& Mason 1936).

MATERIALS AND METHODS

The amber piece contains some 25 complete or partial fruiting bodies in various developmental stages. Some

of the opened fruiting bodies near the edge of the piece were sectioned with a diamond saw and mounted in

immersion oil to observe hyphae, and spores. The amber originated from a mine excavated in 2001, in the

Hukawng Valley, southwest of Maingkhwan in Kachin State (26°20'N, 96°36'E) in Myanmar. This location,

known as the Noije Bum 2001 Summit Site, was assigned to the Early-Mid Cretaceous, Upper Albian, on the

basis of paleontological evidence (Cruickshank & Ko 2003) placing the age at 97 to 110 mya. Nuclear mag-

J. Bot. Res. Inst. Texas 8(1): 139 -143.2014

140

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 1. Group of Palaeogaster micromorphain Myanmar amber. Holotype is the specimen with the large opening in the center of the photo. Scale bar= 3 mm.

netic resonance (NMR) spectra and the presence of araucaroid wood fibers in amber samples from the Noije

Bum 2001 Summit Site indicate an araucarian (possibly Agathis) tree source for the amber (Poinar et al. 2007).

Descriptive terminology and taxonomy is based on Guzman (1970), Guzman and Ovrebo (2000), Gurgel, et al.

(2008), Alfredo et al. (2012) and Nouhra et al. (2012).

DESCRIPTION

Boletales (Sclerodermatineae)

Palaeogaster Poinar, Alfredo, & Baseia, gen. nov. (Figs. 1-8), MycoBank no.: MB 801127. Type Species: Palaeogaster

micromorpha Poinar, Alfredo, & Baseia.

Fruiting bodies small, subglobose to pyriform, spore case filling fruiting body; sterile base absent; peridium brown, hard, thick, splitting

irregularly at terminus or subterminally to form large, roundish aperture; gleba firm, then becoming powdery yellow-orange at maturity;

spores small, clear at maturity, globose to subglobose, smooth to slightly irregular surface; capillitium, hymenium and peridioles absent.

Palaeogaster micromorpha Poinar, Alfredo, & Baseia, sp. nov. (Figs. 1-8), MycoBank no.: MB 801127. Type:

MYANMAR (BURMA): Amber mine in the Hukawng Valley, SW of Maingkhwan in Kachin State (26°20'N, 96 0 36'E), 1999, unknown

amber miner s.n. (holotype: the open, centered specimen in Fig. 1; catalogue number B-F-l deposited in the Poinar amber collection

maintained at Oregon State University, Corvallis, Oregon 97331, U.S.A.).

Fruiting bodies from 5-7 mm in length, 3-4 mm in width; peridium persistent, peridium wall 6-12 pm wide;

surface with areas of bne concentric, often intersecting lines; peridium splitting irregularly at terminus or sub¬

terminus to form large, roundish apertures ranging from 2-3 mm in diameter; apertures rimmed with frag¬

ments of original peridium; mature gleba powdery, yellow-orange; spores clear, globose to subglobose, lacking

Poinar, Jr. et al., Palaeogaster micromorpha, a new genus and species of gasteroid fungus

141

Fig. 2. Lateral view of pyriform fruiting body of Palaeogaster micromor¬

pha in Myanmar amber. Scale bar = 2 mm.

Fig. 3. Cross-section of the peridium of a fruiting body of Palaeogaster micro¬

morpha in Myanmar amber. Scale bar = 35 pm.

Fig. 4. Intersecting lines on the peridial surface of a fruiting body of

Palaeogaster micromorpha in Myanmar amber. Scale bar = 27 pm.

Fig. 5. Group of spores in the gleba of a fruiting body of Palaeogaster micromor¬

pha in Myanmar amber. Scale bar = 27 pm.

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 6. Detail of a spore in the gleba of a fruiting body

of Palaeogaster micromorpha in Myanmar amber. Scale Fig. 7. Mycelial hyphae in a fruiting body of

bar = 8 pm. Palaeogaster micromorpha in Myanmar amber.

Scale bar = 100 pm.

Fig. 8. Mycelial hyphae with clamp connections (arrows) in a fruiting body of Palaeogaster micro¬

morpha in Myanmar amber. Scale bar = 80 pm

Poinar, Jr. et al., Palaeogaster micromorpha, a new genus and species of gasteroid fungus

143

a hilum or pedicel, ranging from 4-11 pm in greatest dimension; mycelial hyphae from fruit bodies 7 -10 pm

in width, unpigmented, occasionally branched, thin-walled, with clamp connections.

Habitat. —Caespitose, probably growing on decaying wood.

Etymology. —The generic epithet is from the Greek “palaios” = ancient and the Greek “gaster” = stomach.

The specific epithet is from the Greek “micros” = small and the Greek “morphe” = form.

DISCUSSION

Palaeogaster is distinguished by its small size, shape of the fruiting bodies, large, roundish terminal to subter¬

minal aperture, yellow- orange gleba, non-sculptured spores and absence of a capillitium, hymenium and

peridioles. The subglobose to pyriform fruiting bodies, single layered peridium, large irregular aperture, ab¬

sence of a sterile base, mycelial hyphae with clamp connections and lack of a capillitium align it with the

Sclerodermatineae. Palaeogaster shares with the extant genus Diplocystis (Sclerodermatineae) the habit of

forming aggregates of small fruiting bodies, each forming a leathery, cup-shaped peridium (Touzan et al.

2007). However, the fruiting bodies of Diplocystis have a powdery umber gleba and the grayish-brown spores

are covered with warty or spiny ornamentation. Small fruiting bodies with a mature yellow-orange gleba and

globose to subglobose spores as occur in Palaeogaster are not found in extant representatives of the Scleroder¬

matineae (Arora 1986; Zeller 1949). Palaeogaster micromorpha represents the first fossil member of the Sclero¬

dermatineae and the oldest known gasteroid fungus.

ACKNOWLEDGMENTS

The senior author thanks Roberta Poinar and Art Boucot for comments on an earlier draft of this manuscript.

Two anonymous reviewers carefully examined and offered constructive feedback for improvement.

REFERENCES

Arora, D. 1986. Mushrooms demystified: A comprehensive guide to the fleshy fungi. 2nd, Edition, Ten Speed Press,

Berkeley, California, U.S.A.

Alfredo, D.S., A.G. Leite, R. Braga-Neto, V.G. Cortez, & I.G. Baseia. 2012. Scleroderma minutisporum , a new earthball from the

Amazon rainforest. Mycosphere 3:294-299.

Chaney, R.W. & H.L. Mason. 1936. A Pleistocene flora from Fairbanks, Alaska. Amer. Mus. Novit. 887:1 -17.

Cruickshank, R.D. & K. Ko. 2003. Geology of an amber locality in the Hukawng Valley, northern Myanmar. J. Asian Earth

Sci. 21:441-455.

Gurgel, F.E., B.D., B. Silva, & I.G. Baseia. 2008. New records of Scleroderma from northeastern Brazil. Mycotaxon 105:399-

405.

Guzman, G. & C.L. Ovrebo. 2000. New observations on sclerodermataceous fungi. Mycologia 92:171 -179.

Guzman, G. 1970. Monografia del genero Scleroderma Pers. emend. Fr. (Fungi, Basidiomycetes). Darwiniana 16:233-407.

Louzan, R., A.W. Wilson, M. Binder, & D.S. Hibbett. 2007. Phylogenetic placement of Diplocystis wrightii in the Scleroderma¬

tineae (Boletales) based on nuclear ribosomal large subunit DNA sequences. Mycoscience 48:66-69.

Magallon-Pueble, S. & R.S. Cervallos-Ferriz. 1993. A fossil earthstar (Geasteraceae; Gasteromycetes) from the Late Cenozoic

of Pueblo, Mexico. Amer. J. Bot. 80:1162-1167.

Nouhra, E.R., M.L. H. Caffot, N. Pastor, & E.M. Crespo. 2012. The species of Scleroderma from Argentina, including a new

species from a Nothofagus forest. Mycologia 104:488-495.

Poinar, G.O., Jr. 2001. Fossil Puffballs (Gasteromycetes: Lycoperdales) in Mexican amber. Historical Biol. 15:219-222.

Poinar, G.O., Jr. & A.E. Brown. 2003. A non-gilled hymenomycete in Cretaceous amber. Mycological Res. 107:763-768.

Poinar G.O., Jr. & R. Buckley. 2007. Evidence of mycoparasitism and hypermycoparasitism in Early Cretaceous amber.

Mycological Res. 111:503-506.

Poinar, G.O., Jr., J.B. Lambert, & Y. Wu. 2007. Araucarian source of fossiliferous Burmese amber: spectroscopic and ana¬

tomical evidence. J. Bot. Res. Inst.Texas 1:449-455.

Zeller, S.M. 1949. Keys to the Orders, Families and Genera of the Gasteromycetes. Mycologica 41:36-58.

144

Journal of the Botanical Research Institute of Texas 8(1)

BOOK NOTICE

Richard Primack. 2014. Essentials of Conservation Biology, Sixth Edition. (ISBN-13: 978-1-60535-289-3,

hbk). Sinauer Associates, Inc., PO Box 407, Sunderland, Massachusetts 01375-0407, U.S.A. (Orders:

www.sinauer.com, 1-413-549-4300). $94.95, 603 pp., 294 illustrations, 8 V 2 " x 11".

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RICHARD B. PRIMACK is a Professor in the Biology Department at Boston University. He received his B.A. at

Harvard University in 1972 and his Ph.D. at Duke University in 1976, and then was a postdoctoral fellow at the

University of Canterbury and Harvard University. He has served as a visiting professor at the University of

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versity and a Guggenheim Fellowship. Dr. Primack was President of the Association for Tropical Biology and

Conservation, and is currently Editor-in-Chief of the journal Biological Conservation. Twenty-eight foreign-

language editions of his conservation biology textbooks (the Essentials and the shorter Primer of Conservation

Biology) have been produced, with local coauthors. He is an author of rain forest books, most recently Eropical

Rain Forests: An Ecological and Biogeographical Comparison (with Richard Corlett). Dr. Primack’s research inter¬

ests include: the biological impacts of climate change; the loss of species in protected areas; tropical forest ecol¬

ogy and conservation; and conservation education. He has recently completed a popular book about changes

in Concord since the time of Henry David Thoreau, titled Walden Warming: Climate Change Comes to Thoreau’s

Woods.

J.Bot. Res. Inst. Texas 8(1): 144.2014

XYLARIA ANTIQUA SP. NOV. (ASCOMYCOTA: XYLARIACEAE)

IN DOMINICAN AMBER

George O. Poinar, Jr.

Department of Integrative Biology

Oregon State University

Corvallis, Oregon 97331 USA

poinarg@science.oregonstate.edu

ABSTRACT

Xylaria antiqua sp. nov. is described from Tertiary Dominican amber. Characters of the stroma, perithecia, ascospores and a white bloom

of conidiogenous cells bearing conidiophores and conidia place the fossil in the Xylariaceae and genus Xylaria. The preservation of X. anti¬

qua is excellent and the morphological characters of the hyphae and spores appear unaltered. This is the first record of a fossil fruiting body

of the family Xylariaceae and shows that the basic characteristics of this group were already established some 20-30 million years ago.

RESUMEN

Se describe Xylaria antiqua sp. nov. del ambar dominicano del Terciario. Caracteres del estroma, peritecios, ascosporas y un grupo bianco

de celulas conidiogenas portadoras de conidioforos y conidios colocan al fosil en las Xylariaceae y genero Xylaria. La preservacion de X.

antiqua es excelente y los caracteres morfologicos de las al hifas y esporas parecen sin alterar. Esta es la primera cita de un cuerpo fructifero

fosil de la familia Xylariaceae y muestra que las caracteristicas basicas de este grupo ya estaban establecidas hace unos 20-30 millones de

anos.

INTRODUCTION

Members of the genus Xylaria are very curious fungi in that their fruiting bodies often resemble decaying plant

remains protruding from the ground. One of the more observed species, X. polymorpha, has dark brown to

black club-shaped fruiting bodies that have been aptly named “Dead-Man’s Fingers.” Other species, such as X.

hypoxylon, have antler-like, often arching fruiting bodies that have earned it the common name of “Candle-

Snuff Fungus.” These fungi are common pathogens, saprobes and endophytes that frequently fruit on dead

wood and other plant substrates. They occur in many parts of the world today and are especially interesting

since both asexual and sexual stages often occur on the same stroma, albeit usually not at the same time; co¬

nidia on mature stromata are remains of the earlier asexual phase.

While examining inclusions in amber from the Dominican Republic, a specimen of Xylaria was discov¬

ered. This fossil belongs to the group of Xylaria that are characterized by flattened thick branches rather than

unbranched clubs. This specimen is described and represents the first record of a fossil fruiting body of the

family Xylariaceae, to my knowledge, and shows that the basic characteristics of this group were already estab¬

lished some 20-30 million years ago.

MATERIALS AND METHODS

The description presented here is based on one complete and well-preserved stroma in Dominican amber. The

specimen was obtained from mines in the Cordillera Septentrional of the Dominican Republic. Dating of Do¬

minican amber is still controversial with the latest proposed age of 20-15 mya based on foraminifera (Iturral-

de-Vinent & MacPhee 1996) and the earliest as 45-30 mya based on coccoliths (Cepek in Schlee 1990). Do¬

minican amber was produced by the leguminous tree, Hymenaea protera Poinar, and a re-construction of the

Dominican amber forest based on amber fossils indicated that the environment was similar to that of a present

day tropical moist forest (Poinar & Poinar 1999). The amber piece was repolished to view the conidia and peri¬

thecia on the branches of the stroma. Observations and drawings were made with a Nikon SMZ-10 stereo¬

scopic microscope.

J. Bot. Res. Inst. Texas 8(1): 145 -149.2014

146

Journal of the Botanical Research Institute of Texas 8(1)

DESCRIPTION

Characters of the stroma, perithecia, ascospores and a white bloom of conidiogenous cells bearing conidio-

phores and conidia places the fossil in the Xylariaceae and genus Xylaria (Rogers 1979,1983,1984a, 1984b).

Phylum: Ascomycota

Order: Xylariales

Family: Xylariaceae

Xylaria Hill ex Schrank

Xylaria antiqua Poinar, sp. nov. (Figs. 1-7), MycoBank no.: MB 808649. Type: Dominican repubtic: amber mines

in the northern mountain ranges (Cordillera Septentrional) of the Dominican Republic, Jul 1994, unknown amber miner s.n. (holo-

type: accession # AF-9-12, deposited in the Poinar amber collection, Oregon State University).

Stroma clavate, flattened, broad, many branched from lower third to near apex, with obtuse to blunt apices;

length of entire specimen (substrate base + stoma), 15 mm; length of perithecium-bearing stroma, 8 mm, great¬

est width stroma, 7 mm; stipe short and broad, length stipe, 2 mm; width stipe, 1.5 mm; most perithecia embed¬

ded in stroma with projecting ostioles; some perithecial elevations exposed on surface of stroma; height of

perithecia, 116-130 pm; asci not observed; ejected ascospores single-celled, smooth walled, light brown, rang¬

ing from bean-shaped to ellipsoidal, 15-18 pm x 7-11 pm; germ slit faint, longitudinal, slightly curved.

Asexual (anamorph) state occurring as cream-colored, conidiogenous palisades over apices of stromal

branches; conidia hyaline, smooth, ovate to elongate elliptical, 3-4 pm x 1-2 pm, borne singly or in pairs on

persistent conidiophores covering stroma.

Etymology. —From the Latin “ antiquus ” = old.

Diagnosis. —The broad, flattened, branched, robust stroma with abrupt apices and roughened surface and

the short, broad stipe, small conidia, ascospores and age appear to separate X. antiqua from extant members of

the genus, despite the fact that only one stroma was available and it is unavailable for dissection. The fossil ex¬

hibits some similarities with Xylaria cornu-damae (Schwein.) Berk, and X. digitata (L.) Grev. (Rogers 1984a)

from the Americas; however, both of these species have rounded or pointed apices rather than blunt tipped

ones as in X. antiqua.

The shape of the ascospore and longitudinal germ slit of X. antiqua is similar to that of X. allantoidea

(Berk.) Fr. (Rogers 1984b; p. 918, Fig. 25), a wide ranging species found throughout the Americas and Africa

andX. grandis Peck (Rogers & Callan 1986; p. 396, Fig. 20) from North America. The germ slit in all of these is

depicted under the light microscope by a faint white, somewhat blurred line extending along the side of the

spore. However, both of these extant species have unbranched stromata. In their key to the Xylaria of North

America (Rogers & Callen 1986), which is mainly based on characters of the ascospores, X. antiqua aligns with

X. mali Fromme; however, that species has rounded fertile apices.

discussion

Xylariaceous fungi are primarily parasites and saprophytes of angiosperms and occur on limbs, standing tree

trunks or logs. Most xylariaceous fungi occur in lowland forests, subtropics and cloud forests (Rogers 1979).

Some species of Xylaria appear to be host-specific (Laessoe & Lodge 1994). These authors discuss two species

of Xylaria (X. meliacarum Laessoe and X. guareae Laessoe & Lodge) that occur only on trees in the family

Meliaceae, including on the leaves of Trichilia species in Puerto Rico. This angiosperm family was represented

in the original Dominican amber forest by at least 3 species of Trichilia and one species of Swietenia (Meliaceae)

(Chambers et al. 2011; Chambers & Poinar 2012a, 2012b). However, a few Xylaria species (X. luxurious (Rehm)

C.G. Lloyd, X. ianthino-velutina (Mont.) Fr.) occur on leaves and pods of legumes (Laessoe & Lodge 1994; Den¬

nis 1956), which suggests that the Dominican amber legume tree, Hymenaeaprotera , also could have served as

host. It is likely that X. antiqua was growing on a dead branch of H. protera and after becoming dislodged, pos¬

sibly by some animal, fell into a pool of resin that had collected on one of the lower branches.

The upright stromata of Xylaria species are considered to raise the perithecia above the substrate for more

Poinar, Jr., Xylaria antiqua sp. nov. (Xylariaceae)

147

Fig. 1. Entire specimen of Xylaria antiqua in Dominican amber. Bar = 2.4 mm.

efficient ejection and dispersal of the ascospores (Rogers 1979). Rogers (1979) considered the small, conidia to

represent relictual spermatia and questioned if they were functional. It is now known that conidia of some Xyl¬

aria species are propagules (for example, see Rogers et al. 2008). On X. antiqua, many of the terminal branches

of the stroma appear to be immature, bearing conidia that are the remains of the earlier asexual state.

148

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 3. Layers of conidiophores with

conidia on stroma of Xylaria antiqua

in Dominican amber. Bar = 120 pm.

Fig. 4. Single layer of conidiophores with

conidia on stroma of Xylaria antiqua in

Dominican amber. Bar = 40 pm.

Fig. 2. Mycelial surface of substrate portion of Xylaria antiqua

in Dominican amber. Bar = 510 pm.

Fig. 5. Ostioles of perithecia protruding from stroma of Xylaria

antiqua in Dominican amber. Bar = 50 pm.

Fig. 6. Perithecium on stroma of Fig. 7. Ellipsoidal ascospore of Xylaria antiqua with

Xylaria antiqua in Dominican amber. faint, longitudinal, germ slit (arrow) in Dominican

Released ascospore is on right. Bar amber. Bar = 11 pm.

= 40 pm.

The preservation of X. antiqua is excellent and the morphological characters of the hyphae and spores ap¬

pear unaltered. The pristine condition of amber fossils is thought to be the result of fixation of the tissues from

chemicals in the original resin. In amber embedded arthropods, there may be some collapsing of the append¬

ages due to dehydration, which is also an important stage in the preservation process. However, it is rare to find

any sign of dehydration or collapsing of the tissues of fungi in amber. Even the pileus of the Early Cretaceous

Myanmar amber agaric, Palaeoagaracites antiquus Poinar & Buckley (2000) that was partly decomposed by a

mycoparasite showed no signs of preservation distortion, nor did the much older spores retained on the

Poinar, Jr., Xylaria antiqua sp. nov. (Xylariaceae)

149

phialides of the Early Cretaceous Myanmar amber Hypocreales, Paleoophiocordyceps coccophagus Sung, Poin¬

ar, & Spatafora (2008).

While this is the first fossil record of a fruiting body of a member of the Xylariaceae, as far as I am aware,

the group as a whole has a more extensive history Bharati et al. (2003) reported xylariaceous, single furrowed

amerospores similar to those found in some Hypoxylonites and Spirotremesporites from Late Cretaceous and

Tertiary sediments of northeastern India.

ACKNOWLEDGMENTS

Thanks are extended to Roberta Poinar, Art Boucot, Jack Rogers, and one anonymous reviewer for editorial

comments and additions that greatly improved the manuscript.

REFERENCES

Bharati, N., B. Subhra, & S. Atreyee. 2003. Fossil Xylariaceae spores from the Cretaceous and Tertiary sediments of North¬

western India. Acta Palaeontol. Sinica 42:56-67.

Chambers, K.L., G.O. Poinar, Jr., & A.E. Brown. 2011. Two fossil flowers of Trichilia (Meliaceae) in Dominican amber. J. Bot.

Res. Inst.Texas 5:463-468.

Chambers, K.L. & G.O. Poinar, Jr. 2012a. A Mid-Tertiary fossil flower of Swietenia (Meliaceae) in Dominican amber. J. Bot.

Res. Inst.Texas 6:123-127.

Chambers, K.L. & G.O. Poinar, Jr. 2012b. Additional fossils in Dominican amber give evidence of anther abortion in Mid-

Tertiary Trichilia (Meliaceae). J. Bot. Res. Inst.Texas 6:561-565.

Dennis, R.W.G. 1956. Some Xylarias of Tropical America. Kew Bull. 11:401 -444.

Iturralde-Vinent, M.A. & R.D.E. MacPhee. 1996. Age and paleogeographic origin of Dominican amber. Science 273:1850-

1852.

Laess0e,T. & DJ. Lodge. 1994. Three host-specific Xylaria species. Mycologia 86:436-446.

Poinar, G.O., Jr. & R. Poinar. 1999. The amber forest. Princeton University Press, Princeton, New Jersey, U.S.A.

Poinar, G.O., Jr. & R. Buckley. 2000. Evidence of mycoparasitism and hypermycoparasitism in Early Cretaceous amber.

Mycol. Res. 111:503-506.

Rogers, J.D. 1979. The Xylariaceae: Systematics, biological and evolutionary aspects. Mycologia 71:1-42.

Rogers, J.D. 1983. Xylaria bulbosa, Xylaria curta, and Xylaria longipes in Continental United States. Mycologia 75:457-467.

Rogers, J.D. Xylaria acuta, Xylaria cornu-damae, and Xylaria mali 'm Continental United States. Mycologia 76:23-33.

Rogers, J.D. 1984b. Xylaria cubensis and its anamorph, Xylocoremium flabelliforne, Xylaria allantoidea, and Xylaria poitei'm

continental United States. Mycologia 76:912-923.

Rogers, J.D. & B.E. Callan. 1986. Xylariapolymorpha and its allies in Continental United States. Mycologia 78:391 -400.

Rogers, J.D., R. Yeomans, & MJ. Adams. 2008. The relationship of Xylaria oxyacanthae to seeds of Crataegus monogyna.

North American Fungi 3:1-5.

Sung, G.-H., G.O. Poinar, Jr., & J.W. Spatafora. 2008. The oldest fossil evidence of animal parasitism by fungi supports a

Cretaceous diversification of fungal-arthropod symbioses. Phylogen. Evol. 49:495-502.

Schlee, D. 1990. Das Bernstein-Kabinett. Stuttgarter Beitragezur Naturkunde (C) 28:1-100.

150

Journal of the Botanical Research Institute of Texas 8(1)

JOURNAL NOTICE

Neal K. Van Alfen, Jan E. Leach, and Steven Lindlow, eds. 2013 (Sep). Annual Review of Phytopathology, Vol¬

ume 51. (ISSN: 0066-4286; ISBN: 978-0-8243-1351-7, hbk). Annual Reviews, Inc., 4139 El Camino Way,

RO. Box 10139, Palo Alto, California 94303, U.S.A. (Orders: www.AnnualReviews.org, science@annual-

reviews.org, 1-800-523-8635,1-650-493-4400). $92.00 indiv., 610 pp., 7%" x 9 3 /s".

ABOUT THIS JOURNAL—The Annual Review of Phytopathology, in publication since 1963, covers the significant developments in the field

of plant pathology, including plant disease diagnosis, pathogens, host-pathogen Interactions, epidemiology and ecology, breeding for resis¬

tance and plant disease management, and includes a special section on the development of concepts.

Contents of Volume 51:

1. Will Decision-Support Systems Be Widely Used for the Management of Plant Diseases?— Dani Shtienberg

2. Chemical Signaling Between Plants and Plant-Pathogenic Bacteria— Vittorio Venturi and Clay Fuqua

3. Biology, Epidemiology, and Control of Heterobasidion Species Worldwide—M atteo Garbelotto and Paolo Gonthier

4. Pine Wood Nematode, Bursaphelenchus xylophilus — Kazuyoshi Futai

5. The Life History of Pseudomonas syringae: Linking Agriculture to Earth System Processes— Cindy E. Morris, Caroline L. Monteil, and

Odile Berge

6. Frichoderma Research in the Genome Era— Prasun K. Mukherjee, Benjamin A. Horwitz, Alfredo Herrera-Estrella, Monika Schmoll, and

Charles M. Kenerley

7. Experimental Measures of Pathogen Competition and Relative Fitness— Jiasui Zhan and Bruce A. McDonald

8. Quiescent and Necrotrophic Lifestyle Choice During Postharvest Disease Development—Dov Prusky, Noam Alkan, Fesfaye Men-

giste, and Robert Fluhr

9. Status and Prospects of Plant Virus Control Through Interference with Vector Transmission—C. Bragard, P. Caciagli, O. LemaireJ.J.

Lopez-Moya, S. MacFarlane, D. Peters, P. Susi, and L. Forrance

10. Diversity and Evolution of Root-Knot Nematodes, Genus Meloidogyne: New Insights from the Genomic Era— Philippe Castagnone-

Sereno, Etienne G.J. Danchin, Laetitia Perfus-Barbeoch, and Pierre Abad

11. Antimicrobial Defenses and Resistance in Forest Trees: Challenges and Perspectives in a Genomic Era— Andriy Kovalchuk, Susanna

Kerio, Abbot O. Oghenekaro, Emad Jaber, Fommaso Raffaello, and Fred O. Asiegbu

12. MAPK Cascades in Plant Disease Resistance Signaling— Xiangzong Meng and Shuqun Zhang

13. The Use and Role of Predictive Systems in Disease Management— David H. Gent, Walter F. Mahaffee, Neil McRoberts, and William F.

Pfender

14. Impacts of Resistance Gene Genetics, Function, and Evolution on a Durable Future— Richard W. Michelmore, Marilena Christopou-

lou, and Katherine S. Caldwell

15. Virus-Based Transient Expression Vectors for Woody Crops: A New Frontier for Vector Design and Use— William O. Dawson and

Svetlana Y. Folimonova

16. Paradigms: Examples from the Bacterium Xylella fastidiosa— Alexander Purcell

17. Advances in Understanding Begomovirus Satellites— Xueping Zhou

18. Engineering Plant Disease Resistance Based on TAL Effectors— Sebastian Schornack, Matthew J. Moscou, Eric R. Ward, and Diana M.

Horvath

19. Nonhost Resistance Against Bacterial Pathogens: Retrospectives and Prospects— Muthappa Senthil-Kumar and KirankumarS. Mysore

20. The Role of Prophage in Plant-Pathogenic Bacteria— Alessandro M. Varani, Claudia Barros Monteiro-Vitorello, Helder I. Nakaya, and

Marie-Anne Van Sluys

21. Considerations of Scale in the Analysis of Spatial Pattern of Plant Disease Epidemics— William W. Furechek and Neil McRoberts

22. Pseudomonas syringae pv. tomato DC3000: A Model Pathogen for Probing Disease Susceptibility and Hormone Signaling in Plants

— Xiu-Fang Xin and Sheng Yang He

23. Diseases in Intercropping Systems— Mark A. Boudreau

24. Manipulation of Host Proteasomes as a Virulence Mechanism of Plant Pathogens— Robert Dudler

25. Centrality of Host Cell Death in Plant-Microbe Interactions— Martin B. Dickman and Robert Fluhr

26. Continuous and Emerging Challenges of Potato virus Y in Potato— Alexander V. Karasev and Stewart M. Gray

21. Communication Between Filamentous Pathogens and Plants at the Biotrophic Interface— Mihwa Yi and Barbara Valent

J.Bot. Res. lnst.Texas8(1): 150.2014

FLORA OF THE HALOPHYTIC GRASSLANDS

IN THE VALLE DE JANOS, CHIHUAHUA, MEXICO

Jose Humberto Vega-Mares and Andres Eduardo Estrada-Castillon*

Facultad de Ciencias Forestales

Universidad Autonoma de Nuevo Leon

Km. 145 carr. Nac. Linares-Cd. Victoria

Apartado postal 41

67700 Linares, Nuevo Leon, MEXICO

navega_mares@hotmail.com

Jose Angel Villarreal-Quintanilla Gustavo Quintana Martinez

Departamento de Botanica

Universidad Autonoma Agraria Antonio Narro

Apartado postal Buena Vista

Saltillo, Coahuila, 25315, MEXICO

Facultad de Zootecnia y Ecologfa

Universidad Autonoma de Chihuahua

Periferico Fco. R. Almada

Kilometro 1, colonia Zootecnia

Chihuahua, Chihuahua, MEXICO

ABSTRACT

A study of the flora and plant endemism of the halophytic communities of the Valle de Janos, Chihuahua, was carried out. Documentation

of the flora and endemic species was conducted by collecting plants throughout the area during a two-year period. The vascular plant diver¬

sity accounts for 57 families, 198 genera, and 328 taxa including infraspecific categories. Asteraceae (40 genera, 55 species), Poaceae (29,60),

Fabaceae (14, 28), and Euphorbiaceae (6,22) are the most representative families in genera and species, respectively. Euphorbia (14 species),

Dalea (8), Bouteloua (8), and Opuntia (7) are the most diversified genera. Fifteen of the species recorded are restricted to the Chihuahuan

Desert; three of them are endemic to the State of Chihuahua, and only one species, Dalea janosensis, is restricted to the Valle de Janos halo¬

phytic communities. All species comprise five biological forms, and according to their origin, 96.5% of the genera and 92.1% of the species

are authouchtonous; the rest of them are exotic.

Keywords: Valle de Janos, Chihuahua, Mexico, endemism, halophytic communities

RESUMEN

Se estudio la flora y endemismo de las comunidades halofilas del Valle de Janos, Chihuahua. Para la documentacion de la flora y endemismo

se realizaron recorridos y colectas de material botanico en toda el area de estudio durante un periodo de dos anos. La diversidad floristica

vascular esta integrada por 57 familias, 198 generos y 328 taxones incluyendo categorias infraespecificas. Asteraceae (40 generos y 55 espe-

cies), Poaceae (29, 60), Fabaceae (14, 28) y Euphorbiaceae (6, 22) son las familias mas representativas respecto a numero de generos y espe-

cies. Euphorbia (14 especies), Dalea (8), Bouteloua (8) y Opuntia (7) son los generos con mayor numero de especies. Quince de las especies

registradas estan restringidas al Desierto Chihuahuense, tres de ellas son endemicas del Estado de Chihuahua, y solo una especie, Dalea

janosensis, esta restringida a las comunidades halofilas del Valle de Janos. Todas las especies comprenden cinco formas biologicas princi-

pales. De acuerdo con su origen, 96.5% de los generos y 92.1% de las especies son autoctonos, el resto son exoticos.

Palabras clave: Valle de Janos, Chihuahua, Mexico, endemismo, comunidades halofilas

INTRODUCTION

Halophytic communities are common within the Chihuahuan Desert region of northern Mexico (Henrickson

& Johnston 1997), which are mainly comprised of the families Poaceae, Chenopodiceae, Frankeniaceae, and

Asteraceae (Rzedowski 1978). These associations constitute an important part of the grasslands, since they are

restricted to specific environmental conditions (Herrera 2012), such as species adapted to soils with high salt

concentrations, basic pH, silty-clay texture, and poor drainage (Rzedowski 1978; Gay & Dwyer 1980). The

edaphic factor is the main determinant in the plant composition in these communities (Miranda & Hernan-

dez-X 1963), which are frequently located at the bottom of closed drainage basins that retain water and inter¬

mountain valleys (Rzedowski 1978).

J. Bot. Res. Inst. Texas 8(1): 151 -163.2014

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Journal of the Botanical Research Institute of Texas 8(1)

The Valle de Janos communities have these characteristics, but also, support extensive areas of arid grass¬

lands and associated shrublands composed mainly of mesquite (Prosopis glandulosa var. torreyana) and ephe¬

dra (Ephedra trifurca) (COTECOCA 1978; Royo & Baez 2001). Valle de Janos is located in the Chihuahuan

Desert ecoregion at the northwestern portion of the State of Chihuahua.

The arid zones in northern Mexico support a high number of endemic plant species (Gonzalez-Medrano

& Chiang 1988; Rzedowski 1991). Its flora and vegetation are of particular interest since they are adapted to

specific environmental conditions (Kliem 2000). However, this large region is among the most unknown from

a floristic viewpoint (Rzedowski 1992). Several studies of the halophytic communities in northern Mexico

have been completed (Johnston 1939), including areas in the State of Chihuahua such as Bablcora (Estrada et

al. 1997), the central part of Chihuahua (Estrada & Villarreal 2010), Canon de Santa Elena (SEMARNAT

1997), Campo Experimental La Campana (Royo & Melgoza 2001), Samalayuca (Enriquez 2003), and Presa la

Boquilla (Gonzalez 2005). Undoubtedly, the most important one is the flora of the Chihuahuan Desert

(Henrickson & Johnston 1997), where almost 150 of the species cited occur in the halophytic communities of

Valle de Janos, highlighting those belonging to the families Asteraceae and Chenopodiacaeae. Valle de Janos is

of great importance for natural resource conservation in Mexico and North America (List et al. 2000; Manza-

no-Fischer et al. 2000), and therefore is a North American Grassland Priority Conservation Area (GPCA) (Karl

& Hoth 2005). The National Commission for the Knowledge and Use of Biodiversity (CONABIO) has also

identified it as a Priority Terrestrial Region (RTP-45) (Arriaga 2000) and an important Area for Bird Conserva¬

tion (AICAs-45) (CIPAMEX-CONABIO 1999). Valle de Janos is also part of the complex of priority areas in the

Chiricahua-Peloncillo-Sierra Madre for the conservation of wildlife (Dinerstein et al. 2000). It is noteworthy

that Valle de Janos has the largest colony of the black-tailed prairie dog (Cynomys ludovicianus Ord.) (Ceballos

et al. 1993) in North America, which is an endangered species (SEMARNAT 2010). Nevertheless, this area has

been impacted by anthropogenic activities that threaten the persistence of the endemic flora in these halo¬

phytic communities and the prairie dog populations. These communities have suffered a rapid decline as an

ecosystem over the past 25 years (Ceballos et al. 2010). The main causes of deterioration include livestock

overgrazing, expansion of mechanized agriculture (Ceballos et al. 1993, 2005), and climate change (Pinedo et

al. 2013).

To date, there have been no studies that characterize the plant diversity or the ecology for the region. The

aim of this study is to contribute to the knowledge of the regional flora and plant endemism in the halophytic

communities in the Valle de Janos.

METHODS

Study area

The study area is about 116,000 ha and is located in the northwestern region of the State of Chihuahua, in the

municipality of Janos, 30°54'23"N, 108°38'55"W and 30°53'51"N, 108°13'58"W (Fig. 1). The annual rainfall is

306.7 mm with almost 50% of it occurring between July and September. The mean temperature is 16°C (Rze¬

dowski 1978). The altitude is 1380-1500 m and the topography varies from flat ridges to low round hills with

gentle slopes of 1-8%. The main climate is BSoK type, corresponding to dry and temperate climates according

to the Kopeen Classification System (modified by Garcia 1973). The hydrology of the area belongs to the North

Closed Basin Region, specifically to Rio Casas Grandes Basin (CONAGUA 2009). According to the Guide for

interpreting soils (INEGI2004), INIFAP-CONABIO (1995) and INEGI (2005), the most outstanding soil in the

Valle de Janos are Luvic Xerosol (with clay accumulation in the subsoil, traces of lime or gypsum, with very low

and burdensome physical phase); Haplic Xerosol (with very low organic matter content and very permeable);

Haplic Feozem (in flat or gently wavy relief, permeable); Haplic Yermozol (thin soil, poor in organic matter

content, high content in calcareous material, permeable); Luvic Yermosol (with high clay content in the B ho¬

rizon, and below this may have calcic or gypsic horizon); Orthic Zolonetz (clayey, with an alkaline, saline

phase, waterproof, and poor in organic matter content); Orthic Solonchak (clay, high concentrations of soluble

salts, poor in organic matter and nutrients content, with a sodium phase); Eutric Regosol (coarse texture, poor

Vega-Mares, et al., Halophytic grasslands

153

Fig. 1. Study area. The Valle de Janos (darker shaded area) is located at the northwestern portion in the State of Chihuahua, and, it is the area where

the largest colony of prairie dogs is located in North America.

in organic matter content, nutrient-rich subsoil, very permeable) and Lithosol (stony and very thin, in steep

slopes).

The representative vegetation in the area is salt-tolerant grassland, extensive arid grassland, and micro-

phyllous desert scrub (COTECOCA 1978; Rzedowski 1978) (Fig. 2). The most common shrub species in the

microphyllous desert scrub are Prosopis glandulosa var. torreyana, Ephedra trifurca, Opuntia imbricata, Mimosa

aculeaticarpa, and Atriplex canescens. Also, there are several dominant herbaceous elements such as Gutierre-

zia sarothrae, G. microcephala, Chenopodium album, Muhlenbergiaporteri, Salsola tragus, and Pleuraphis mutica,

while in the natural, undisturbed by agriculture or grazing grasslands, there are Bouteloua spp., Aristida spp.,

Panicum spp., and Eragrostis spp., plus other herbaceous species such as Machaeranthera spp., Sida sp., Eriogo-

num spp., and Eepidium spp. The halophytic grasslands are dominated by species adapted to poorly drained

and saline soils, including Pleuraphis muitca, Sporobolus airoides, Atriplex canescens, and Portulaca mundula.

The study was carried out during the years 2011 and 2012. Specimen collecting was conducted in differ¬

ent seasons in order to document the phenological stages of species in all plant associations. All plants were

properly georeferenced, recording the community or dominant association where the plants were found. Plant

identification was made by using different monographs for the different genera as well as the Flora of the Chi-

huahuan Desert (Henrickson & Johnston 1997). Herbarium voucher specimens were deposited at ANSM

(Saltillo, Coahuila, Mexico) and CFNL (Finares, Nuevo Feon, Mexico) herbaria.

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Journal of the Botanical Research Institute of Texas 8(1)

Fig. 2. Habitat of Pleuraphis mutica, a dominat herbaceous element in a salt-tolerant grassland escosystem in the Valle de Janos, Chihuahua, Mexico.

Other dominates include Prosopisglanduloso var. torreyana and Yucca elata.

RESULTS AND DISCUSSION

Diversity

The flora of the Valle de Janos is represented by 57 families, 198 genera, and 328 species (including infraspe¬

cific categories) of vascular plants (Table 1, Appendix 1). Dicots include 50 families, 161 genera, and 259 spe¬

cies, while monocots are represented by five families, 35 genera, and 67 species. Gymnosperms and ferns are

represented by only one species each. The families with the highest number of genera and species are

Asteraceae (40, 55), Poaceae (29, 60), Fabaceae (14, 28), and Euphorbiaceae (6, 22), and the ten most diversified

families include over half of the genera (122, 61.6%) and species (229, 69.8%) (Table 2). The most diversified

genera are Euphorbia (14), Dalea (8), Bouteloua (8), Opuntia (7), Aristida (6), and Atriplex (5) (Table 3). Accord¬

ing to the plant diversity in Mexico (304 families, 2,804 genera, and 23,424 species) (Villasenor 2004), the State

of Chihuahua encompasses 37.5% (152), 22.2% (890), and 8.1% (4,035) of the families, genera, and species, re¬

spectively. The flora of the Valle de Janos represents 18.8% of the families, 7.1% of the genera, and 1.4% of the

species existing in Mexico, and represents 37.5%, 22.2%, and 8.1% of the families, genera, and species existing

in the State of Chihuahua. The families with the highest diversity in genera and species in most of Mexico are

Asteraceae, Poaceae, and Fabaceae (Rzedowski 1992; Villasenor 2003, 2004), which in turn, have been used as

predictors of diversity in this country (Villasenor et al. 2007). The four genera of Valle de Janos ( Euphorbia ,

Dalea, Bouteloua, and Opuntia) are also among the 17 most numerous genera of Mexico (Villasenor 2004). Ac¬

cording to Rzedowski (1992), the grasslands and shrublands support almost 6,000 species (20% of the total

flora), and 5.5% of these species are found in the Valle de Janos.

Vega-Mares, et al., Halophytic grasslands

155

Table 1. Division of the vascular flora recorded in the study area.

Groups

Families

Genera

Species

Infraspecific categories

Ferns

Gymnosperms

Angiosperms

Dicotyledons

161

259

Monocotyledons

Total

198

328

Table 2. Families with the highest number of genera and species in the study area.

Family

Number of genera

Number of species

Asteraceae

Poaceae

Fabaceae

Malvaceae

Euphorbiaceae

Cactaceae

Solanaceae

Brassicaceae

Nyctaginaceae

Chenopodiaceae

Table 3. Genera with the highest number of species in the study area.

Genera

Number of species

Euphorbia

Dalea

Bouteloua

Opuntia

Aristida

Atrip lex

Muhlenbergia

Sporobolus

Asclepias

Astragalus

Valle de Janos has lower species diversity than other halophytic community areas found in the State of

Chihuahua, such as the center of the State of Chihuahua (Estrada & Villarreal 2010), La Campana Experimen¬

tal Ranch (Royo & Melgoza 2001), and Laguna de Bablcora (Estrada et al. 1997). This difference is related to the

other plant associations present in these areas and absent in Valle de Janos, such as scrublands and pine-oak

forest. However, if we compare the homogenous halophytic communities, such as the areas where prairie dogs

inhabit the northeastern and northwestern regions of Mexico, we can see the northwestern halophytic com¬

munities support greater plant diversity (Table 4). A fairer comparison of the flora of the Valle de Janos is per¬

haps the halophytic grassland of northeastern Mexico (Estrada et al. 2010). In both grasslands Asteraceae,

Poaceae, and Fabaceae are the dominant plant families, followed by others such as Cactaceae, Brassicaceae, and

Solanaceae. Their diversity is low compared with other ecosystems (Rzedowski 1978); both grasslands are

home to the prairie dog, Cynomys mexicanus (northeast grasslands) and Cynomys ludovicianus (northwest

grasslands). Despite being similar in composition and structure, both grasslands show contrast in plant diver¬

sity: Valle de Janos has 14 families, 24 genera, and 102 species more than the northeast grasslands (53 families,

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Journal of the Botanical Research Institute of Texas 8(1)

Table 4. Comparison of the flora in the Valle de Janos against regional floras of Chihuahua and one area from northeastern Mexico. Numbers in parentheses represent

the number of shared taxa.

Families

Genera

Species

Vegetation

Altitude (masl)

Valle de Janos

198

328

Grassland and scrublands

1380-1500

Central Chihuahua

112(51)

493(181)

1322(221)

Grassland and oak-pine forest

1450-2300

La Campana (Chih.)

74(43)

258(130)

433(124)

Grassland and oak-pine forest

1500-2500

Babicora (Chih.)

67(41)

244(93)

476(70)

Grassland and oak-pine forest

2150-2700

NE Mexico

53(37)

174(92)

284(73)

Grassland and scrublands

1550-1950

174 genera, and 226 species) (Estrada et al. 2010). Both grasslands share Dalea, Bouteloua, Opuntia, Aristida,

and Atriplex as the most diversified genera, and the herbaceous plants have the highest diversity, followed by

shrubs and trees. There is a higher plant endemism in the northeast grasslands (17 species, 6%) than in the

northwest ones (1,0.003%). The influence of the different climate, altitude, topography, soil type, and the pres¬

ence of endoergic basins in the northeast grasslands areas have undoubtedly favored this speciation phenom¬

enon, compared to the relatively homogeneous landscape found in the northwest grasslands.

Endemism, growth forms, and flora origin

From the total flora, only fifteen of the species are endemic to the Chihuahuan Desert, including the State of

Chihuahua and Valle de Janos grasslands (see Appendix I). Three of them are endemic only for the State of

Chihuahua: Matelea chihuahuensis, Opuntia orbiculata, and Dalea janosensis. Dalea janosensis is a new species

recently described for this area (Estrada et al. 2013) that was discovered as part of this work. It is associated

only with halophytic communities comprised of Sporoblus airoides, S. cryptandrus, Atriplex acanthocarpa, A.

wrightii, A. canescens, Prosopis glandulosa var. torreyana, Malvella leprosa, Bouteloua aristidoides, and Ipomoea

costellata. According to the NOM-059-SEMARNAT-2010 (SEMARNAT 2010), only one of the species, Amoreux-

ia palmatifida, is protected (special protection, Pr). This species is found only in rocky and undulating ridges

associated with Artemisia frigida, Sphaeralcea wrightii, Tragia nepetifolia, and Heteropogon contortus.

The total taxa include five growth forms: herbaceous (163 perennials, 106 annuals), shrubs (25 thornless

and 8 thorny), fleshy stems (15), rosetophyllous (2), and trees (9). The most abundant annual species in the

study area are Aristida adscensionis, Bouteloua barbata, B. aristidoides, and Panicum hirticaule. The most com¬

mon perennial ones are Pleuraphis mutica, Sporobolus airoides, Bouteloua eriopoda, B. gracilis, Bothriochloa bar-

binodis, and Aristida divaricata. The most common annual forbs are Atriplex wrightii, Eriogonum abertianum,

Portulaca olearacea, Talinum aurantiacum, and Machaeranthera tanacetifolia. The most common perennial forbs

are Salsola tragus, Machaeranthera pinnatifida, Sida abutifolia, Solanum elaeagnifolium, and Evolvulus alsinoides.

The dominant scrublands are comprised of Prosopis glandulosa var. torreyana, Epherda trifurca, Atriplex canes¬

cens, Mimosa aculeaticarpa, and Ziziphus obtusifolia. Salix bonplandiana, Populus fremontii, and Platanus

racemosa are common in riparian areas. Cylindropuntia imbricata, Opuntia macrorhiza var. macrorhiza, O. mac¬

rocentra, and Echinocereus rigidissimus are the most common cacti. Dasylirion wheeleri and Yucca elata are

scarce in the Valle de Janos, found only in the most arid areas.

Several species such as Atriplex canescens, A. acanthocarpa, A. elegans, A. obovata, A. wrightii, Pleuraphis

mutica, Sporobolus airoides, S. pyramidatus, Eragrostis mexicana, and E. neomexicana are common in areas

where sodium or sulfate salt concentrations are very high. These salty soils are frequent in the north of the

study area, such as Rancho Las Arenillas and Rancho el Penasco.

Where there are few hillocks, the topographic diversity highlights the presence of no-saline adpated spe¬

cies, including shrubs such as Aloysia wrightii and Baccharis pteronioides; grasses such as Bouteloua curtipen-

dula, Heteropogon contortus, Bouteloua eriopoda, Eycurus phleoides, Digitaria californica, and Aristida divaricata;

and forbs such as Artemisia frigida, Sphaeralcea wrightii, Tragia nepetifolia, Commelina dianthifolia, and

Amoreuxia palmatifida.

In the Valle de Janos, native species (302 = 92.1%) are, by far, more diverse than the introduced ones (26 =

Vega-Mares, et al., Halophytic grasslands

157

7.9%). Among the most frequent introduced species are Salsola tragus, Chloris virgata, Chenopodium album,

Schismus barbatus, Cyperus esculentus, Eragrostis cilianensis, Urochloa mutica, and Mollugo verticillata (see “i” in

Appendix I). Similarly, the native genera (191 = 96.4%) surpass those introduced (7 = 3.5%). The most frequent

introduced genera are Brassica, Eruca, Cynodon, Schismus, and Salsola. These elements are very common in

abandoned fields in the area. The most common growth forms are herbs (269 species = 82%), shrubs (33 =

10.1%), and trees (10 = 3%). We recorded 76 weed species (23.2% of the total flora), 56 native and 20 exotic ones

(see “w” in Appendix I). The most frequent weed species in the area are Salsola tragus, Sida abutifolia, Machaer-

anthera pinnatifida, Chloris virgata, Pectis prostrata, Chenopodium album, Amaranthus retroflexus, Mollugo verti¬

cillata, Evolvulus alsinoides, and Eragrostis cilianensis. These species are more frequent in areas with exposure

to livestock management (overgrazing) and agriculture (crops abandoned).

CONCLUSIONS

Valle de Janos supports rich plant diversity in spite of its relatively homogenous topography. Its characteristic

plant associations are determined by the arid climate, the edaphic factors, and black-tailed prairie dog, allow¬

ing it the presence of that particular flora. Halophytic communities of northwestern Mexico have higher plant

diversity but lower plant endemism than counterparts found in northeastern Mexico (Estrada et al 2010).

When plant diversity of Valle de Janos is compared with several surrounding areas in the State of Chihuahua,

it is less diverse, since this study counted only the species inhabiting the plains and excluded the oak and oak-

pine forest. This study detected a large quantity of herbaceous species not recorded previously by general bo¬

tanical studies (COTECOCA 1978). We recommend studies focused in ecology and exotic species replacement

and establishment due to soil use and compaction as well as those focused in overgrazing since this ecosystem

is currently highly transformed by agricultural activities. The most common grasses such as Aristida adscen-

sionis, Bouteloua barbata, B. aristidoides, Panicum hirticaule, and Bothriochloa barbinodis and some herbs such as

Sida abutifolia, Machaeranthera pinnatifida, M. tanacetifolia, Solanum elaeagnifolium, and Evolvulus alsinoides are

disturbance indicator plants and are classified as weeds (CONABIO 2013), coupled with some exotic species

such as Salsola tragus, Mollugo verticillata, and Chloris virgata (Villasenor & Espinosa-Garcia 2004).

The capabilities of the plant species to tolerate drought and salinity are causal factors for the presence of

species and communities in different habitat, and the minimum xylem pressure potential are indicative of

drought tolerance, and the minimum cell osmotic potential are indicative of the tolerance of plant species to

salinity (Branson et al 1988). Several species found in this area such as Atriplex obovata, A. canescens, Ephedra

trifurca, Gutierrezia sarothrae, Krascheninnikovia lanata, Earrea tridentata, Prosopis glandulosa var. torreyana,

and Sporobolus airoides. There are over 2,000 known plant species worldwide that have some tolerance to salin¬

ity (Menzel & Lieth 2003); 27 of them are found in the Valle de Janos, and the most outstanding ones in de¬

creasing order with respect to tolerance of salts are Atriplex canescens, Sporobolus airoides, Chenopodium album,

Prosopis glandulosa var. torreyana, and Salsola tragus. At least 60 species found in the Valle de Janos are in¬

cluded into the USDA-ARS data bases (Yensen 2013) as salt-tolerant plants.

The most common forbs associated are Atriplex wrightii, Eriogonum abertianum, E. abertianum Machaer¬

anthera pinnatifida, M. tanacetifolia, Portulaca oleracea, Salsola tragus, Sida abutifolia, Solanum elaeagnifolium,

Talinum aurantiacum, Zinnia grandiflora, and several species of the genus Euphorbia. It is common to find

patches of Pleuraphis mutica in areas where Haplic Yeromozol soils (high calcium contents) are present; it is a

species that grows in environments with moderately saline soils and soils with calcium carbonate, very resis¬

tant to fire and drought (USFS 2014). Several arboreal and shrub species such as Baccharis salicifolia, B. saro-

throides, Platanus racemosa, Populus fremontii, and Salix bonplandiana are frequently found in flat or gently

rolling relief with Haplic Foezem soils. The Yermozol and Solonchak soils are most common at the north-cen¬

tral part of the study area and support mainly Pleuraphis mutica-Sporobolus airoides-Atriplex spp., as well as S.

pyramidatus. This last species is common in salt meadows in Louisiana (Reid et al 2010). According to Rzew-

doski (1978) and COTECOCA (1978), the most important grasses in Valle de Janos in the late 70s and early 80s

were Bouteloua gracilis, B. eripoda, B. hirsuta, B. curtipendula, and Aristida divaricata. This can give an idea of

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Journal of the Botanical Research Institute of Texas 8(1)

the species replacement, although the extent of it is still unknown. With respect to the main grasses ( Pleura-

phis mutica and Sporobolus airoides) in the halophytic communities, the plant physiognomy is almost the same,

but the species composition is different. The discovery of a new taxon (Dalea janosensis ) could imply a new

conservation scheme in this unique halophytic community in the arid land of the Chihuahuan Desert. Last, we

recommend promoting studies in plant diversity in the surrounding regions to complete the total flora of this

interesting and unique area in Mexico.

APPENDIX 1

List of families, genera and species recorded in the Valle de Janos, Chihuahua, Mexico. H.V.M. = Humberto Vega Mares and collection

number. Biological growth forms (t = tree, s = shrub, h = herbaceous, f = fleshy stems, and r = rosetophyllous). Native (n), introduced (i),

weeds (w). Endemic to the Chihuahuan Desert (*), endemic for the State of Chihuahua (•), and endemic to the Valle de Janos (§).

FERNS

Pteridaceae

Astrolepis cochisensis (Goodd) D.M. Benham & Windham ssp. co-

chisensis (Goodd.) D.M. Benham & Windham, H.V.M. 1796; h, n

GYMNOSPERMS

Ephedraceae

Ephedra trifurca Torr. ex S. Watson, H.V.M. 1900; s, n

MONOCOTYLEDONS

Asparagaceae

Dasylirion wheeled S. Watson ex Rothr., H.V.M. 1921; r, n

Yucca elata (Engelm.) Engelm., H.V.M. 1914; r, n

Amaryllidaceae

Allium kunthii G. Don, H.V.M. 1916; h, n

Commelinaceae

Commelina dianthifolia Delile, H.V.M. 1655; h, n

Cyperaceae

Carexfilifolia Nutt., H.V.M. 1531; h, n

Cyperus dipsaceus Liebm., H.V.M. 1595; h, n

Cyperus esculentus L., H.V.M. 1679; h, i, w

Poaceae

Aristida adscensionis L., H.V.M. 1562: h, n, w

Aristida divaricata Humb. & Bonpl. ex Willd., H.V.M. 1584; h, n

Aristida havardii Vasey, H.V.M. 1583; h

Aristida pansa Woot. & Standi., H.V.M. 1482,1604; h, n

Aristida purpurea Nutt. var. longiseta (Steud.) Vasey, H.V.M. 1700; h, n

Aristida schiedeana Jr\r\\us & Ruprecht, H.V.M. 1784; h, n

Bothriochloa barbinodis (Lag.) Herter, H.V.M. 1489; h, n

Bouteloua aristidoides (Kunth) Griseb., H.V.M. 1526; h, n

Bouteloua barbata Lag., H.V.M. 1458; h, n

Bouteloua curtipendula (Michx.)Torr., H.V.M. 1571; h, n

Bouteloua eriopoda (Torr.) Torr., H.V.M. 1498; h, n

Bouteloua gracilis (Kunth) Lag. ex Griffiths, H.V.M. 1543; h, n

Bouteloua hirsuta Lag., H.V.M. 1462,1703; h, n

Boutelouaparryi (E. Fourn.) Griffiths, H.V.M. 1785; h, n

Bouteloua rofhroc/c/7 Vasey, H.V.M. 1606; h, n

Bromus anomalus Rupr. ex Fourn., H.V.M. 1786; h, n

Cenchrus incertus M.A. Curtis, H.V.M. 1663; h, n, w

Cenchrus spinifex Cav., H.V.M. 1788; h, n, w

Chloris virgata Sw., H.V.M. 1563; h, i, w

Cynodon dactylon (L.) Pers., H.V.M. 1789; h, i, w

Dasyochloapulchella (Kunth) Willd. ex Rydb., H.V.M. 1597; h, n

Digitaria californica (Benth.) Henr., H.V.M. 1540; h, n

Digitaria sanguinalis (L.) Scop., H.V.M. 1733; h, i, w

Echinochloa colona (L.) Link, H.V.M. 1725; h, i, w

Echinochloa crusgalli (L.) P. Beauv., H.V.M. 1724; h, i, w

Elyonurus barbiculmis Hack., H.V.M. 1790; h, n, m

Enneapogon desvauxii P. Beauv., H.V.M. 1664; h, n, m

Eragrostis cilianensis (All.) Link ex Vignolo, H.V.M. 1525; h, i, w

Eragrostis lehmanniana Nees, H.V.M. 1607; h, i, w

Eragrostis mexicana (Hornem.) Link, H.V.M. 1560; h, n, w

Eragrostis neomexicana Vasey ex L.H. Dewey, H.V.M. 1561; h, n

Eriochloa gracilis (E. Fourn.) Hitchc., H.V.M. 1723; h, n

Erioneuron avenaceum (Kunth) Tateoka, H.V.M. 1653; h, n

Heteropogon contortus (L.) P. Beauv. ex Roem. & Schult., H.V.M.

1652; h, n

Leptochloa dubia (Kunth) Nees, H.V.M. 1612; h, n, w

Leptochloa filiformis (Pers.) P. Beauv., H.V.M. 1588; h, n, w

Lycurusphleoides Kunth, H.V.M. 1791; h, n, w

Muhlenbergia arenacea (Buckley) Hitchc, H.V.M. 1651; h

Muhlenbergia arenicola Buckley, H.V.M. 1699; h, n

Muhlenbergia monticula Buckley, H.V.M. 1792; h, n

Muhlenbergia ported Scribn. ex Beal, H.V.M. 1603; h, n

Muhlenbergia rigens (Benth.) Hitchc., H.V.M. 1800: h, n

Panicum hirticaule J. Presl, H.V.M. 1564; h, n

Panicum obtusum Kunth, H.V.M. 1585; h, n

Pleuraphis mutica Buckley, H.V.M. 1463; h, n

Schismus arabicus Nees, H.V.M. 1792; h, i, w

Schismus barbatus (L.) Thell., H.V.M. 1678,1726; h, i, w

Scleropogon brevifolius Phil., H.V.M. 1593; h, n, w

Setaria grisebachii E. Fourn., H.V.M. 1605; h, n, w

Setaria leucopila (Scribn. & Merr.) K. Schum., H.V.M. 1673; h, n, w

Sorghum halepense (L.) Pers., H.V.M. 1539; h, i, w

Sporobolus airoides (Torr.) Torr., H.V.M. 1799; h, n

Sporobolus contractus Hitchc., H.V.M. 1608; h, n

Sporobolus cryptandrus (Torr.) A. Gray, H.V.M. 1794; h, n

Sporobolus flexuosus (Thurb. ex Vasey) Rydb., H.V.M. 1702; h, n

Sporoboluspyramidatus (Lam.) Hitchc., H.V.M. 1623; h, n

Tragus berteronianus Schult., H.V.M. 1602; h, i, w

Tridens muticus (Torr.) Nash, H.V.M. 1714; h, n

Urochloa arizonica (Scribn. & Merr.) Morrone & Zuloaga, H.V.M.

1657; h,n

Urochloa mutica (Forssk.) T.Q. Nguyen, H.V.M. 1930; h, i, w

DICOTYLEDONS

Aizoaceae

Trianthema portulacastrum L., H.V.M. 1919; h, n, w

Amaranthaceae

Amaranthus blitoides S. Watson, H.V.M. 1709; h, n

Amaranthus retroflexus L., H.V.M. 1918; h, n, w

*Froelichia arizonica Thornber ex Standi., H.V.M. 1596; h, n

Gomphrena nitida Rothr., H.V.M. 1593; h, n

Guilleminea densa (Humb. & Bonpl. ex Schult.) Moq., H.V.M. 1544;

h, n, w

Tidestromia lanuginosa (Nutt.) Standi., H.V.M. 1917; h, n

Vega-Mares, et al., Halophytic grasslands

159

*Tidestromia suffruticosa (Torr.) Standi., H.V.M. 1513; h, n

Anacardiaceae

Rhus microphyllo Engelm. ex A., H.V.M. 1600; s, n

Apocynaceae

Apocynum androsaemifolium L., H.V.M. 1920; h, n

Asclepios osperulo (Decne.) Woodson, H.V.M. 1530; h, n

Asclepias lotifolio (Torr.) Raf., H.V.M. 1512; h, n

Asclepios oenotheroides Schltdl. & Cham., H.V.M 1749; h, n

Asclepios subverticilloto (A. Gray) Vail, H.V.M. 1717; h, n

•Moteleo chihuohuensis (A. Gray) Woodson, H.V.M. 1736; h, n

Sorcostemmo cynanchoides Decne. ssp. hortwegii (Vail) R. Holm,

H.V.M. 1915; h, n

Aristolochiaceae

Aristolochio longecoudoto S. Watson, H.V.M. 1568; h, n

*Aristoiochio wrightii Seem., H.V.M. 1701; h, n

Asteraceae

Acourtio nono (A. Gray) Reveal & R. M. King, H.V.M. 1519; h, n

Ambrosia artemisiifolia L., H.V.M. 1559; h, i, w

Ambrosia confertiflora DC., H.V.M. 1627; h, n, w

Aphanostephus ramosissimus DC. var. humilis (Benth.) B.L.Turner &

Birdsong, H.V.M. 1743; h, n, w

Artemisia frigido Willd., H.V.M. 1594; h, n

Artemisia ludoviciana Nutt., H.V.M. 1750; h, n, w

Baccharis pteronioides DC., H.V.M. 1670; s, n

Baccharissalicifolia (Ruiz & Pav.) Pers., H.V.M. 1537; s, n

Baccharis sorothroides A. Gray, H.V.M. 1719; s, n

Bahia absinthifolia Benth., H.V.M. 1613; h, n, w

Bohio autumnalis Ellison, H.V.M. 1738; h, n

Bahia pedata A. Gray, H.V.M. 1913; h, n

Boileyo multirodioto Harv. & A. Gray ex A. Gray, H.V.M. 1570; h, n

Berlandiera lyrata Benth., H.V.M. 1527; h, n

Bidens bigelovii A. Gray, H.V.M. 1694; h, n, w

Brickellio eupatorioides (L.) Shinners var. chlorolepis (Wooton &

Standi.) B.L.Turner, H.V.M. 1741; h, n

Brickellio lemmonii A.Gray var. conduplicoto (B.L. Rob.) B.L. Turner,

H.V.M. 1730; h, n

Choetopoppo bellioides (A. Gray) Shinners, H.V.M. 1912; h, n

Chaetopappa ericoides (Torr.) G.L. Nesom, H.V.M. 1640 y 1739; h, n

Cirsium texanum Buckley, H.V.M. 1485; h, n

Conoclinium greggii (A. Gray) Small, H.V.M. 1574; h, n

*Flourensia cernua DC., H.V.M. 1599; s, n

Grindelia squarrosa (Pursh) Dunal, H.V.M. 1630; h, n

Gutierrezia microcepholo (DC.) A. Gray, H.V.M. 1922; s, n

Gutierrezia sarothrae (Pursh) Britton & Rusby, H.V.M. 191; s, n

Helianthus cilioris DC., H.V.M. 1506; h, n, w

Helianthuspetiolaris Nutt., H.V.M. 1751; h, n

Heliomerislongifolia (B.L. Rob. &Greenm.) Cockerell, H.V.M. 1923; h, n

Heterotheca subaxillaris (Lam.) Britton & Rusby, H.V.M. 1545; h, n

Hymenoxys richordsonii (Hook.) Cockerell, H.V.M. 1910; h, n

Laennecia coulteri (A. Gray) G.L. Nesom, H.V.M 1536; h, n

Leuciva dealbata (A. Gray) Rydb., H.V.M. 1616; h, n, w

Machaeranthera pinnatifida (Hook.) Shinners, H.V.M. 1481; h, n, w

Mochoeronthero tonocetifolio (Kunth) Nees, H.V.M. 1508; h, n

Parthenium incanum Kunth, H.V.M. 1541; s, n

Pectis papposa Harv. & A. Gray, H.V.M. 1677; h, n

Pectis prostrata Cav., H.V.M. 1618; h, n, w

Porophyllum ruderole (Jacq.) Cass., H.V.M. 1695; h, n

Psilactis asteroides A. Gray, H.V.M. 1742; h, n, w

Psilostrophe tagetina (Nutt.) Greene, H.V.M. 1549; h, n

Sanvitalia abertii A. Gray, H.V.M. 1547; h, n

Senecio floccidus Less. var. douglosii (DC.) B.L.Turner &T.M. Barkley,

H.V.M. 1509; h, n

Senecio longilobus Benth., H.V.M. 1517; s, n, w

Stephanomeria pauciflora (Torr.) A. Nelson, H.V.M. 1516; h, n

Thelesperma megapotamicum (Spreng.) Kuntze, H.V.M. 1478; h, n

Thymophyllo oceroso (DC.) Strother, H.V.M. 1636; h, n

*Thymophylla aurea (A. Gray) Greene var. polychaeta (A. Gray)

Strother, H.V.M. 1617; h, n

Tithonia tubiformis (Jacq.) Cass., H.V.M. 1740; h, n, w

Trixiscolifornico Kellogg, H.V.M. 1671; s, n

Verbesina encelioides (Cav.) Benth. & Hook. f. ex A. Gray, H.V.M.

1518; h, n, n

Viguiera cordata (Hook. & Arn.) DArcy, H.V.M. 1737; h, n

Viguiero linearis (Cav.) Sch.Bip. ex Hemsl., H.V.M. 1747; h, n, w

Xanthismagracile (Nutt.) D.R. Morgan & R.L. Hartm., H.V.M. 1936; h, n

Xanthium strumarium L., H.V.M. 1909; h, n

Zinnia grandiflora Nutt., H.V.M. 1484; h, n

Bignoniaceae

Chilopsis linearis (Cav.) Sweet, H.V.M. 1689; t, n

Bixaceae

Amoreuxio polmotifido DC. H.V.M. 1665; h, n

Boraginaceae

Cryptantha cinerea (Greene) Cronquist, H.V.M. 1643; h, n

Cryptontho micrantha (Torr.) I.M. Johnst., H.V.M. 1752; h, n

Nama hispida A. Gray, H.V.M. 1908; h, n

Brassicaceae

Brassica rapa L., H.V.M. 1690; h, i, w

Brossico tournefortii Gouan, H.V.M. 1674; h, i, w

Descurainiapinnata (Walter) Britton, H.V.M. 1907; h, n, w

Dimorphocarpa wislizeni (Engelm.) Rollins, H.V.M. 1669; h, n

Erucasativa (L.) Mill., H.V.M. 1727; h, i, w

Lepidium losiocorpum Nutt., H.V.M. 1460; h, n

Lepidium montanum Nutt., H.V.M. 1460; h, n

Lepidium virginicum L., H.V.M. 1924; h, n, w

Physaria gordonii (A. Gray) O'Kane & Al-Shehbaz, H.V.M. 1935; h, n

Cactaceae

Echinocereus fendleri (Engelm.) Sencke ex J.N. Haage var. fendleri,

H.V.M. 1906; f,n

*Echinocereus rigidissimus (Engelm.) F. Haage., H.V.M. 1753; f, n

*Coryphanthascheeri (Kuntze) L.D. Benson var. robustispina (Schott

ex Engelm.) L.D. Benson, H.V.M. 1925; f, n

Cylindropuntio imbricoto (Haw.) F. M. Kunth, H.V.M. 1926; f, n

Mammillaria heyderi Muehlenpf., H.V.M. 1934; f, n

Opuntia discoto Griffiths, H.V.M. 1904; f, n

Opuntia macrocentra Engelm., H.V.M. 1939; f, n

Opuntia mocrorhizo Engelm.var. mocrorhizo, H.V.M. 1927; f, n

Opuntia mocrorhizo Engelm. var. pottsii (Salm-Dyck) L.D. Benson,

H.V.M. 1938; f,n

•Opuntia orbiculata Salm-Dyck ex Pfeiff., H.V.M. 1903; f, n

Opuntia polyacantha Haw. var. trichophoro (Engelm. &J.M. Bigelow)

J.M.Coult., H.V.M. 1928; f, n

Opuntia spinosior (Engelm.) Tourney, H.V.M. 1933; f, n

Thelocactus bicolor (Galeotti) Britton & Rose var. bicolor, H.V.M.

1902; f, n

Chenopodiaceae

Atriplexacanthocarpa (Torr.) S. Watson, H.V.M. 1619; a

Atriplexconescens (Pursh) Nutt., H.V.M. 1503; s, n

Atriplexelegans (Moq.) D. Dietr. H.V.M. 1469; s, n

Atriplexobovata Moq., H.V.M. 1620; s, n

Atriplex wrightii S. Watson, H.V.M. 1631; h, n

Chenopodium album L., H.V.M. 1502; h, i, w

Kroscheninnikovio lonoto (Pursh) A.D.J. Meeuse & Smith, H.V.M.

1504; s, n

Solsolo tragus L., H.V.M. 1634; h, i, w

160

Journal of the Botanical Research Institute of Texas 8(1)

Convolvulaceae

Convolvulus equitons Benth., H.V.M. 1656; h, n

Dichondra argentea Humb. & Bonpl. ex Willd., H.V.M. 1491; h, n, w

Evolvulus olsinoides (L.) L, H.V.M. 1488; h, n, w

Evolvulus sericeus Sw., H.V.M. 1487; h, n

Ipomoeo costelloto Torr., H.V.M. 1582; h, n

Ipomoea cristulata Hallier f., H.V.M. 1686; h, n

Crassulaceae

Sedum wrightii A. Gray, H.V.M. 1754; h, f

Cucurbitaceae

Apodonthero unduloto A. Gray, H.V.M. 1468; h, n, w

Cucurbita digitoto A. Gray, H.V.M. 1467; h, n

Cucurbita foetidissima Kunth, H.V.M. 1901; h, n, w

Ibervillea tenuisecta (A. Gray) Small, H.V.M. 1644; h, n

Ericaceae

Arbutusxalapensis Kunth, H.V.M. 1755; t, n

Euphorbiaceae

Acalypha neomexicana Mull. Arg., H.V.M. 1713; h, n

Acolypho ostryifolio Riddell, H.V.M. 1587; h, n

Argythamnia humilis (Engelm. & A. Gray) Mull. Arg. var. humilis,

H.V.M. 1745; h,n

Argythomnio neomexicono Mull. Arg., H.V.M. 1661; h, n

Croton pottsii (Klotzsch) Mull. Arg., H.V.M. 1712; h, n

Croton suaveolenslorr., H.V.M. 1633; h, n

Euphorbia albomarginata Torr. & A. Gray, H.V.M. 1459; h, n

Euphorbia capitellata Engelm., H.V.M. 1464; h, n

Euphorbia cinereascens Engelm., H.V.M. 1889; h, n

Euphorbia davidii Subils, H.V.M. 1591; h, n, w

Euphorbia dentata Michx., H.V.M. 1554; h, n, w

Euphorbia dioeca Kunth, H.V.M. 1721; h, n, w

Euphorbia exstipulata Engelm., H.V.M. 1555; h, n

Euphorbia heterophylla L. H.V.M. 1522; h, n

Euphorbia hyssopifolia L., H.V.M. 1580; h, n, w

Euphorbia micromera Engelm., H.V.M. 1720; h, n

Euphorbia setiloba Engelm. ex Torr., H.V.M. 1680; h, n

Euphorbia serpyllifolia Pers., H.V.M. 1579; h, n

Euphorbiastictospora Engelm., H.V.M. 1888; h, n

Euphorbia tomentulosa S. Watson, H.V.M. 1685; h, n

Jatropha macrorhiza Benth., H.V.M. 1465; h, n

Tragia ramosa Torr., H.V.M. 1756; h, n

Fabaceae

Acacia angustissima (Mill.) Kuntze, H.V.M. 1471; s, n

Astragalus crassicarpus Nutt., H.V.M. 1632; h, n

Astragalusmollissimuslorr., H.V.M. 1470; h, n, w

Astragalus nuttalianus DC., H.V.M. 1698; h, n

Astragalus sp., H.V.M. 1722; h, n

Caesalpinia jamesii (Torr. & A. Gray) Fisher, H.V.M. 1472; h, n

Chamaecrista nictitans (L.) Moench, H.V.M. 1494; h, n

Crotalaria pumila Ortega, H.V.M. 1550; h, n, w

Da lea brachystachys A.Gray, H.V.M. 1691; h, n

Dalea candidaW\M. var. oligophylla (Torr.) Shinners, H.V.M. 1635; h, n

Daleaformosa Torr., H.V.M. 1641; s, n

Dalea jamesii (Torr.) Torr. & A. Gray, H.V.M. 1705; h, n

§Daleajanosensis A.E. Estrada, Villarreal &H. Vega, H.V.M. 1675; h, n

Dalea nana Torr. var. nana, H.V.M. 1704; h, n

Dalea pogonathera A. Gray, H.V.M. 1565; h, n

Dalea polygonoides A. Gray, H.V.M. 1575; h, n

Desmodium neomexicanum A. Gray, H.V.M. 1598; h, n

Eysenhardtiaspinosa Engelm., H.V.M. 1887; s, n

Hoffmannseggia glauca (Ortega) Eifert, H.V.M. 1493; h, n, w

Macroptilium gibbosifolium (Ortega) A. Delgado, H.V.M. 1457; h, n, w

Mimosa aculeaticarpa Ortega, H.V.M. 1505; s, n

Mimosa dysocarpa Benth., H.V.M. 1886; s, n

Prosopis glandulosa var. torreyana Jorr., H.V.M. 1495; s, n

Rhynchosia senna Gillies ex Hook. &Arn., H.V.M. 1535; h, n

Rhynchosia senna Gillies ex Hook. & Arn. var. angustifolia (A. Gray)

Grear, H.V.M. 1704; h, n

Senna bauhinioides (A. Gray) H.S. Irwin & Barneby, H.V.M. 1520; h, n

Senna durangensis (Rose) H.S. Irwin & Barneby var. durangensis,

H.V.M. 1490; h, n

Senna lindheimeriana (Scheele) H. S. Irwin & Barneby, H.V.M.

1521; s,n

Fouquieriaceae

Fouquieriasplendens Engelm., H.V.M. 1758; s, n

Geraniaceae

Erodium cicutarium (L.) L'Her. ex Ait., H.V.M. 1885; h, i, w

Hydrophyllaceae

Phacelia crenulata Torr. ex S. Watson, H.V.M. 1984; h, n

Krameriaceae

Krameria lanceolata Torr., H.V.M. 1639; h, n

Lamiaceae

Salvia arizonica A. Gray, H.V.M. 1734; h, n

Salvia pinguifolia (Fernald) Wooton & Standi., H.V.M. 1883; s, n

* Salvia subincisa Benth., H.V.M. 1589; h, n

Tetraclea coulteri A. Gray, H.V.M. 1557 y 1628; h, n

Linaceae

Linum aristatum Engelm., H.V.M. 1466; h, n

Loasaceae

Cevallia sinuata Lag., H.V.M. 1667; h, n

Mentzelia multiflora (Nutt.) A. Gray, H.V.M. 1507; h, n

Malpiguiaceae

Janusia gracilis A.Gray, H.V.M. 1610; s, n

Malvaceae

Abutilon palmeri A. Gray, H.V.M. 1609; s, n

Abutilon parvulum A.Gray, H.V.M. 1746; h, n

Abutilon wrightii A. Gray, H.V.M. 1515; h, n

Anoda cristata (L.) Schltdl., H.V.M. 1552; h, n, w

Anoda thurberi A.Gray, H.V.M. 1744; h, n

Hibiscus denudatus Benth., H.V.M. 1659; s, n

Krascheninnikovia lanata (Pursh) A. Meeuse & A. Smith; H.V.M.

1435; h,n

Malvella leprosa (Ortega) Krapov., H.V.M. 1497; h, n

Rhynchosidaphysocalyx (A. Gray) Fryxell, H.V.M. 1601; h, n

Sida abutiifolia Mill., H.V.M. 1650; h, n, w

Sphaeralcea angustifolia (Cav.) G. Don, H.V.M. 1715; h, n, w

Sphaeralcea coccinea (Nutt.) Rydb., H.V.M. 1759; h, n

Sphaeralcea hastulata A. Gray, H.V.M. 1625; h, i, w

*Sphaeralcea wrightii A. Gray, H.V.M. 1666; h

Martyniaceae

Proboscidea louisianica (Mill.) Thell., H.V.M. 1780; h, n, w

Molluginaceae

Mollugo verticillata L., H.V.M. 1499; h, i, w

Moraceae

Morus nigra L., H.V.M. 1687; t, i, w

Nyctaginaceae

*Acieisanthes chenopodioides (A. Gray) R.A. Levin, H.V.M. 1556; h, n

Allionia incarnata L., H.V.M. 1542; h, n

Allioniaincarnata L. var. villosa (Standi.) B.L.Turner, H.V.M. 1572; h, n

Boerhavia erecta L., H.V.M. 1682: h, n, w

Boerhavia gracillima Heimerl, H.V.M. 1662; h, n

Vega-Mares, et al., Halophytic grasslands

161

Boerhovio intermedia M.E. Jones, H.V.M. 1524; h, n

Boerhavia wrightii A. Gray, H.V.M. 1496 y 1546; h, n

Mirabilis longiflora L., H.V.M. 1629; h, n

Mirabilis multiflora (Torr.) A.Gray, H.V.M. 1626; h, n

Oleaceae

Menodora sea bra A. Gray, H.V.M. 1735; h

Onagraceae

Gaura coccinea Pursh, H.V.M. 1477 y 1528; h

Gaura mollis Kunth, H.V.M. 1716; h

Oenothera dissecta A. Gray, H.V.M. 1693; h

Oenothera primiveris A. Gray, H.V.M. 1781; h

Oenothera sp., H.V.M. 1782; h

Orobanchaceae

Orobanche cooperi (A. Gray) A. Heller, H.V.M. 1882; h

Oxalidaceae

Oxalis stricta L., H.V.M. 1681; h, n

Papaveraceae

Argemonepleiacantha Greene, H.V.M. 1929; h, n

Eschscholzia californica Cham., H.V.M. 1881; h, n

Plantaginaceae

Penstemon fendleri Torr. & A. Gray, H.V.M. 1783; h, n

Plantago patagonica Jacq., H.V.M. 1480; h, n

Platanaceae

Platanus racemosa Nutt. var. wrightii (S. Watson) L. D. Benson,

H.V.M. 1534; t, n

Polemoniaceae

Giliastrum rigidulum (Benth.) Rydb., H.V.M. 1642; h, n

Polygalaceae

Polygala obscura Benth., H.V.M. 1523 y 1529; h, n

Polygonaceae

Eriogonum abertianum Torr., H.V.M. 1624,1647 y 1729; h, n

Eriogonum polycladon Benth., H.V.M. 1567; h, n

^Eriogonum rotundifolium Benth., H.V.M. 1614; h, n

Eriogonum wrightii Torr. ex Benth., H.V.M. 1510; h, n

Persicaria maculosa A. Gray, H.V.M. 1648; h, i, w

Rumexcrispus L., H.V.M. 1795; h, i, w

Rumexhymenosepalus Torr., H.V.M. 1931; h, n

Portulacaceae

Portulaca halimoides L., H.V.M. 1932; h, n

Portulaca oleracea L., H.V.M. 1798; h, n, w

Portulaca pilosa L., H.V. M. 1501; h, n w

Portulaca umbraticola Kunth, H.V.M. 1492; h, n

Talinum aurantiacum Engelm., H.V.M. 1466,1476; h, n

Rhamnaceae

Ziziphus obtusifolia (Hook, ex Torr. & A. Gray) A. Gray, H.V.M. 1461; s, n

Rosaceae

Fallugia paradoxa (D. Don) Endl. ex Torr., H.V.M. 1731; ,s, n

Salicaceae

Populus fremontii S. Watson, H.V.M. 1532; t, n

Salixbonplandiana Kunth, H.V.M. 1533; t, n

Sapindaceae

Sapindus saponaria L. var. drummondii (Hook. & Arn.) L.D. Benson,

H.V.M. 1649; t,n

Solanaceae

Chamaesaracha coronopus (Dunal) A. Gray, H.V.M. 1710; h, n

Chamaesaracha pallida Averett, H.V.M. 1645; h. n

*Datura ceratocaula Ortega, H.V.M. 1668; h, n, w

Datura quercifolia Kunth, H.V.M. 1578; h, n, w

Lycium pallidum Miers, H.V.M. 1615; s, n

Lycium torreyi A. Gray, H.V.M. 1684; s, n

Nicotiana fr/gonophylla Dunal, H.V.M. 1660; s, n, w

Physalis acutifolia (Miers) Sandwith, H.V.M. 1586; h, n, w

Solanum elaeagnifolium Cav., H.V.M. 1479; h, n w

Solanum heterodoxum Dunal, H.V.M. 1538; h, n, w

Solanum jamesii Torr., H.V.M. 1676; h, n

Solanum rostratum Dunal, H.V.M. 1718; h, n, w

Ulmaceae

Celtis reticulata Torr., H.V.M. 1511; t, n

ACKNOWLEDGMENTS

We thank James Henrickson and an anonymous reviewer for careful and helpful reviews that improved the

manuscript.

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Contents of Volume 38:

Introduction— Ashok Gadgil and Diana M. Liverman

Who Should Read This Journal?— Ashok Gadgil and Diana M. Liverman

I. Earth’s Life Support Systems

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2. Regional and Global Emissions of Air Pollutants: Recent Trends and Future Scenarios— Markus Amann, Zbigniew Klimont, and

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5. Life-Cycle Assessment of Electric Power Systems— Eric Masanet, Yuan Chang, Anand R. Gopal, Peter Larsen, William R. Morrow III,

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18. Humans and Nature: How Knowing and Experiencing Nature Affect Well-Being— Roly Russell, Anne D. Guerry, Patricia Balvanera,

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19. Preindustrial Human Impacts on Global and Regional Environment— Christopher E. Doughty

Cumulative Index of Contributing Authors, Volumes 29-38

Cumulative Index of Article Titles, Volumes 29-38

J.Bot. Res. Inst. Texas 8(1): 164.2014

PRIMER REGISTRO DE ASTROCASIA PELTATA (EUPHORBIACEAE)

EN COSTA RICA

Irene Calderon Sanou

Escuela de Biolog fa

Universidad de Costa Rica

11501-2060 San Jose ,, COSTA RICA

irecalsa@gmail. com

RESUMEN

Se documenta una poblacion aislada de Astrocasia peltata Standi, creciendo a orillas de la quebrada Brasil en el Parque Nacional Diria, en

Costa Rica. Este es el primer registro de esta especie fuera de Mexico de donde se creia endemica. Se incluye una descripcion actualizada de

la especie, se presenta una clave para distinguir las dos especies de Astrocasia en Costa Rica y se discuten posibles explicaciones de la distri¬

bucion geografica disyunta que se revela para A. peltata con este nuevo descubrimiento.

Palabras Clave: Astrocasia peltata, Euphorbiaceae, Costa Rica, Parque Nacional Diria, distribucion disyunta

ABSTRACT

The discovery of an isolated population of Astrocasia peltata Standi., growing on the banks of Brasil Creek in Parque Nacional Diria, in

Costa Rica, is documented. This is the first record of the species outside of Mexico where it was believed endemic. An updated morphologi¬

cal description of the species is presented and possible explanations for the disjunct geographical distribution are discussed.

Key Words: Astrocasia peltata, Euphorbiaceae, Costa Rica, Parque Nacional Diria, disjunct distribution

El genero Astrocasia B.L. Rob. & Millsp. esta representado por seis especies neotropicales y tiene una distribu¬

cion disyunta desde Mexico y Cuba hasta el sur de Bolivia y Brasil (Webster 1992; Jimenez & Martinez 2001).

Pertenece a la subfamilia Phyllanthoideae, que igual que Oldfieldioideae incluye los generos de Euphorbiaceae

con dos ovulos por loculo del ovario (Webster 1994). Astrocasia se puede distinguir por la combinacion de las

siguientes caracterlsticas: la ausencia de pubescencia en las plantas, hojas deciduas, flores con largos pedice-

los, una corola bien desarrollada con petalos libres, un androceo peltado, discos femeninos en forma de una

cupula y frutos capsulares (Burger & Huft 1995). En Costa Rica se conocla una sola especie, A. tremula

(Griseb.) G.L. Webster, de la cual solo hay un registro recolectado por Jorge Gomez Laurito en un bosque re¬

sidual a orillas de la quebrada Pavas de Bajo Rodriguez, en San Ramon de Alajuela ( Gomez Laurito 12368, USJ).

En una expedicion realizada en julio de 2012, se recolectaron ejemplares con frutos de una planta arbus-

tiva a orillas de la quebrada Brasil, en el Parque Nacional Diria, en Santa Cruz de Guanacaste, que se identified

como A. peltata Standi, y resulto ser un nuevo registro para Costa Rica y America Central (Grayum et al. 2012).

Esta especie se consideraba endemica de Mexico, donde se ha hallado en las costas de Jalisco, las Islas Tres

Marlas y una poblacion aislada en Sinaloa cerca de Mazatlan (Webster 1992). Una distribucion disyunta se ha

observado en otras especies de Astrocasia y este nuevo hallazgo, junto con los descubrimientos recientes de

especlmenes de A. tremula en Costa Rica, Panama, Colombia y Venezuela, sugieren que la distribucion bi-

modal del genero observada por Webster (1992), con un centro en Mexico-Guatemala y otro en Sudamerica,

puede no reflejar su distribucion, sino la falta de informacion en las regiones ubicadas entre estos dos grandes

centros.

En una segunda expedicion realizada en mayo de 2013, se visito el mismo sitio donde se hablan encon-

trado los primeros ejemplares y se observaron plantas en estado reproductivo, con flores femeninas y masculi-

nas enteras. Webster (1992) y los autores anteriores no conocieron los petalos ni los sepalos en flores pistiladas,

que si se observaron en varias plantas de Costa Rica ( Calderon Sanou 59, USJ, duplicado en CR). A continuacion

J. Bot. Res. Inst. Texas 8(1): 165 -168.2014

166

Journal of the Botanical Research Institute of Texas 8(1)

se ofrece una description que combina la informacion de Webster, quien trabajo con los ejemplares de Mexico,

y las medidas realizadas en los ejemplares de Costa Rica.

Astrocasia peltata Standi., Contr. Dudley Herb. 1:74, pl.l, fig. 4. 1927 (Fig. 1). Tipo: Mexico. Nayarit: islasTres

Marfas, Isla Marfa Madre, Ferris 5571 (holotipo: DS).

Arbusto o arbol pequeno, 2-5 m de altura, caducifolio, monoico. Tallo lenoso, con ramas delgadas y largas;

ramitas glabras. Hojas maduras peltadas, con estlpulas de 3.5-7.5 mm de largo; peclolo (1.5)4-7.4 cm de largo;

lamina cartacea o subcoriacea, orbicular u ovado-orbicular, 2-10 cm de largo y 1.6-8.5 cm de ancho, enves

palido; venacion reticulada. Flores axilares, fasciculadas (2-3) o solitarias. Flores estaminadas: pedicelo 6-10

mm de largo; sepalos oblongo-ellpticos, 1.5-2.5 mm de largo y 1.8-2 mm de ancho; petalos obovados, 3 mm de

largo y 2 mm de ancho; estambres 5; androceo ca. 1.1 mm de ancho; columna estaminal ca. 1.7 mm de alto;

pistilodio ca. 0.8 mm transversalmente. Flores pistiladas: pedicelo (25)30-74 mm de largo, menos de 1 mm de

diametro, sepalos oblongos, 2-2.5 mm de largo y 2 mm de ancho; petalos obovados, 4.5-5 mm de largo y 3 mm

de ancho; gineceo ca. 1.3 mm de largo y 1 mm de ancho; estilos bipartitos, ovario con 3(4) carpelos. Frutos

oblados, ca. 8 mm de largo y 10-11.2 mm de ancho; columela ca. 4 mm de largo; semillas plano-convexas, lisas,

3.9-4.8 mm de largo, 3.1-3.9 mm de ancho.

Astrocasia peltata se diferencia de las otras especies del genero por tener hojas fuertemente peltadas,

aunque se ha observado que otras especies de Astrocasia pueden presentar algunas hojas levemente peltadas

(Webster 1992), entre estasA. tremula, por lo que se recomienda tener cuidado si se va a utilizar esta caracterlstica

para diferenciar estas especies. La clave siguiente permite distinguir las especies de Astrocasia de Costa Rica:

1. Hojas maduras peltadas; lamina foliar orbicular u ovado-orbicular; petalos de flores estaminadas elipticos a oblongo-

lanceolados, 2.5-3.5 mm de largo, 0.8-1.1 mm de ancho; habitat bosque seco_ A. peltata

1. Hojas maduras basifijas; lamina foliar ovada u ovado-eliptica; petalos de flores estaminadas obovados, 3 mm de largo,

2 mm de ancho; habitat bosque muy humedo_ A. tremula

Distribucion y habitat de Astrocasia peltata. —En Mexico, en laderas de bosques deciduos y semideciduos de la

costa de Jalisco, Islas Tres Marlas y Sinaloa (Webster 1992, Fig. 5). En Costa Rica, una pequena poblacion ais-

lada creciendo en bosque seco a orillas de la quebrada Brasil, Parque Nacional Diria.

Material examinado: Astrocasia peltata Standi. COSTA RICA. Guanacaste: Parque Nacional Diria, Distrito El Arado, Santa Cruz, a orillas

de la quebrada Brasil, 10°09 , 40"N, 85°36 , 07"W, 360 m, 21 Jul 2012,1. Calderon 9 (CR, USJ); loc. cit., 4 May 2013,1. Calderon 59 (CR, USJ).

MEXICO. Nayarit: Marfa Madre, Tres Marfas Islands, 21 Oct 1925, R.S. Ferris 5571 (MO-251998, foto en www.tropicos.org).

Astrocasia tremula (Griseb.) Webster. COSTA RICA. Bajo Rodriguez: Alajuela, San Ramon, Coope-Zamora, bosque residual a la

orilla de la Quebrada Pavas, 10°18 , 32"N, 85°32 , 07"W, 300 m, 23 Mar 1993,J. Gomez-Laurito et al. 12368 (USJ).

DISCUSION

La distribucion geograhca disyunta de Astrocasia peltata que revela este nuevo descubrimiento podrla tener

varias explicaciones. Por un lado, tanto en Mexico como en Costa Rica se ha encontrado en laderas rocosas, por

lo que es de esperar que las plantas que crecen en estos ambientes de diflcil acceso se mantengan ocultas a los

ojos de los botanicos y exploradores durante las expediciones. Poblaciones pequenas de A. peltata podrlan

permanecer escondidas en otras partes de America Central, esperando ser descubiertas. Asl, lo que se nos

presenta ahora como una distribucion disyunta serla, entonces, solo una parte del mapa de distribucion de la

especie, aun incompleto debido a la falta de informacion entre estos dos puntos. Por otro lado, es posible que la

destruccion del habitat por causa del ser humano o de fenomenos naturales en el pasado hayan provocado la

desaparicion de poblaciones de A. peltata en el resto de Centroamerica. En este caso la observacion del esce-

nario completo se dihculta, porque nos encontrarlamos frente al problema de las especies que poseen poblacio¬

nes relictuales, como menciona Webster (1992).

La distribucion disyunta observada en A. peltata nos hace recordar otros descubrimientos recientes en la

flora costarricense, como es el caso de Pleodendron costaricense N. Zamora, Hammel & R. Aguilar (Canellace-

ae), una rara especie de arbol encontrada en el bosque lluvioso de bajura del Paclhco Sur de Costa Rica, a 2000

km de distancia de la unica otra especie del mismo genero, P. macranthum en Puerto Rico (Hammel & Zamora

Calderon, Astrocasia peltata en Costa Rica

167

Fig. 1. Astrocasia peltata: A) Quebrada Brasil, en cuyas orillas se halla esta especie B) Pequena poblacion creciendo en laderas rocosas C) Habito D) Hojas

y ramitas E) Rama con flor estaminada y boton floral en fasciculos F) Rama con frutos verdes colgantes.

2005). Otro ejemplo interesante de disyuncion en el continente americano es Chiangiodendron mexicanum T.

Wendt (Flacourtiaceae), que se hallo en Costa Rica cuando solamente se conocla del extremo sur del estado de

Veracruz en Mexico (Zamora et al. 2004). El descubrimiento de A. peltata y de otras especies de plantas disyun-

tas es informacion valiosa para entender el complejo origen que pudieron haber tenido los bosques de Cen-

troamerica.

Astrocasia peltata y A. tremula se han considerado como especies hermanas por compartir el mismo habi¬

tat y por la presencia de A. tremula en las mismas regiones donde se crela endemica A. peltata en Jalisco, Mexico

(Webster 1992). En Costa Rica, los habitats en que se han encontrado diberen considerablemente. Astrocasia

168

Journal of the Botanical Research Institute of Texas 8(1)

peltata crece en un bosque caducifolio, similar a su habitat en Mexico; en cambio, A. tremula se halla en un

bosque muy humedo, diferente a lo que se hubiera esperado. Ademas, Webster (1992) menciona que la ma-

yorla de especies de Astrocasia, incluida A. peltata , parecen estar exclusivamente asociadas a suelos calcareos

segun las observaciones de los recolectores. Sin embargo, hasta ahora no sabemos si los especlmenes de A.

peltata recolectados en el Parque Nacional Diria estaban asociados a este tipo de suelo.

AGRADECIMIENTOS

Agradezco a Carlos O. Morales por su gran apoyo y por su ayuda en la revision del manuscrito y la identih-

cacion de la especie. Igualmente a Barry E. Hammel y a Daniel Santamarla, quienes mediante comunicacion

via electronica (cuando Daniel se hallaba en el Herbario MO) en un solo dla pudieron hacer la determinacion.

Este descubrimiento no hubiera sido posible sin la participacion y el apoyo de Elmer G. Garcia, que me per-

mitio conocer la localidad, en el marco del Trabajo Comunal Universitario-54 de la Universidad de Costa Rica.

Un sincero agradecimiento tambien a un re visor anonimo por sus sugerencias y comentarios.

REFERENCIAS

Burger, W.C. & MJ. Huft. 1995. Family 113. Euphorbiaceae. En: W.C. Burger, ed. Flora Costaricensis 36:1 -168.

Grayum, M.FI., B.E. FIammel, & N. Zamora (eds.). 2012. Leaps and bounds - Euphorbiaceae. The Cutting Edge (A quarterly

newsletter in anticipation of Manual de Plantas de Costa Rica) 19(4),October - http://www.mobot.org/mobot/re-

search/edge/octl 2/octl 2lea.shtml

Hammel, B.E. & N.A. Zamora. 2005. Pleodendron costaricense (Canellaceae), a new species for Costa Rica. Lankesteriana

5(3):211-218.

Jimenez, J. & M. Martinez. 2001. Una especie nueva del genero Astrocasia (Euphorbiaceae) del estado de Guerrero, Mexico.

Acta Bot. Mex. 55:1-5.

Webster, G.L. 1992. Revision of Astrocasia (Euphorbiaceae). Syst. Bot. 17(2):311-323.

Webster, G.L. 1994. Synopsis of the genera and suprageneric taxa of Euphorbiaceae. Ann. Missouri Bot. Gard. 81:33—144.

Zamora, N., B.E. Hammel, & M.H. Grayum. 2004. Novedades. En: B. E. Hammel, M.H. Grayum, C. Herrera, & N. Zamora, eds.

Manual de Plantas de Costa Rica. vol. I. Introduccion. Monogr. Syst. Bot. Missouri Bot. Gard. 97:1-300.

PLEURISANTHES ELAVA (ICACINACEAE): A NEW RECORD FOR BRAZIL

Bruno S. Amorim Rodrigo Duno de Stefano

Laboratorio de Morfo-Taxonomia Vegetal Dept Botanica Centro de Investigacion Cientffica de Yucatan

Universidade Federal de Pernambuco A.C. Colonia Chuburna de Flidalgo

CEP: 50670-901, Recife, Pernambuco, BRAZIL CP 97200, Merida, Yucatan, MEXICO

brunosarim@yahoo.com.br

Marccus Alves

Laboratorio de Morfo-Taxonomia Vegetal Dept. Botanica

Universidade Federal de Pernambuco

CEP: 50670-901, Recife, Pernambuco, BRAZIL

ABSTRACT

Pleurisanthes flava Sandwith is reported here for the first time from the Brazilian lowland Amazon Rainforest based on unreported collec¬

tions from 1936,1968, and 1975. The species can be recognized by the elliptic to ovate leaves with entire margin, the axillary or supra-axil-

lary and racemose inflorescence, sessile or shortly pedicellate, and 5-merous flowers.

Key Words: Biodiversity, lowland Amazon Rainforest, South America

RESUMO

Pleurisanthes flava Sandwith e citada aqui pela primeira vez para a Floresta Amazonica brasileira baseada em coletas dos anos de 1936,1968

e 1975 previamente nao publicados. A especie pode ser reconhecida pelas folhas elipticas a ovadas com margens inteiras, pela inflorescencia

racemosa axilar a supra-axilar com pedicelo curto ou sessil e flores 5-meras.

Palavras chave: America do Sul, Biodiversidade, Floresta Amazonica de terras baixas

Icacinaceae (s.l.) comprises approximately 52 genera and 400 species worldwide. It occurs predominantly in

the Tropics and rapidly decreasing in number toward the subtropics. In the Neotropics, the family is repre¬

sented by 12 genera and 54-57 species (Duno de Stefano 2004). Morphological and molecular studies (Kare-

hed 2001) showed Icacinaceae (s.l.) as of polyphyletic origin and under a new circumscription, it was segre¬

gated in four distinct families [Icacinaceae s.s., Cardiopteridaceae Blume, Stemonuraceae (M. Roem.) Karehed,

and PennantiaceaeJ. Agardh].

Icacinaceae s.s. has a pantropical distribution with 30 genera and 140 species (Duno de Stefano 2004;

Duno de Stefano & Amorim 2012; Karehed 2001). Pleurisanthes Baill. is a small genus of woody vines to climb¬

ing shrubs and restricted to the Neotropics. Seven species of Pleurisanthes are recognized from the rainforests

of Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, and Brazil (Duno de Stefano 2004; Howard &

Duno de Stefano 1999; Duno de Stefano & Amorim 2012). The genus is characterized by the flowers not being

articulated at the distal portion of the pedicels (Duno de Stefano et al. 2002) and the species have smaller and

less attractive flowers and fruits than other species in Icacinaceae s.s. The highest diversity of Pleurisanthes spe¬

cies is found in Brazil which comprises five species distributed in the Amazon and Atlantic Forest (Duno de

Stefano & Amorim 2012). Pleurisanthes has a problematic taxonomy and needs a further review. The genus is

poorly known mainly because of the general lack of collections from the Amazon Forest. We publish here the

new record of Pleurisanthe flava from the Amazon forest of Brazil.

Pleurisanthes flava Sandwith has previously been recorded from Guyana (de Roon 1994; Keloff et al. 2011)

and its occurrence in the Brazilian Amazon was suggested by de Roon (1994). However, no Brazilian collec¬

tions were cited by de Roon (1994). In a recent herbarium survey, four vouchers of P. flava from Brazil were

discovered and verified, and are here reported from Brazil for the first time, distant ca. 700 Km from the south¬

ern record of the species in Guyana, expanding the known distribution of the species.

J. Bot. Res. Inst. Texas 8(1): 169 -173.2014

170

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 1. Distribution map of Pleurisanthesflava Sand with in South America, showing records in Guyana and Brazil (States of Amazonas and Para).

MATERIAL AND METHODS

This study was based on herbarium collections of Pleurisanthes from the Amazon rain forest. Specimens from

16 herbaria were studied (BM, C, F, G, IAN, INPA, K, MIRR, MO, NY, P, PH, RB, SP, UFRR, US, and VEN) (her¬

barium abbreviation follows Thiers 2013). Morphological studies of the herbarium collections were carried out

using a stereomicroscope.

RESULTS AND DISCUSSION

A short description with the main diagnostic characters is provided. A more complete description of Pleurisan¬

thesflava can be found in de Roon (1994) and Sandwith (1931).

Pleurisanthes flava Sandwith, Bull. Misc. Inform. Kew 1931 (10):467-468. 1931. (Figs. 1, 2). Type: Guyana:

Bartica, Essequibo River, Moraballi Creek, near sea level, 11 Nov 1929 (fl), N.Y. Sandwith590 (holotype: K!; isotype: K!, NY!).

Woody vines. Leaves 7.5-19.5 x 4-6 cm, elliptic to ovate, apex apiculate, rarely emarginate, basecuneate to

rounded, margin entire, bicolor (dried samples), adaxial surface with midveinsulcate, secondary veins 8-10

pairs, abaxial surface fulvo-stringose; petiole 1-2 cm long. Inflorescence axillary or supra-axillary, simple or

compound racemes, main axis ca. 5-7.5 cm long; flowers sessile to shortly pedicellate, 0.1-0.2 cm long; calyx

5-lobed, 0.05-0.1 cm long, triangular, apex acute, pubescent; corolla 5-lobed, petals oblong to laceolate, 0.2-

0.3 cm long; stamens 5, ca. 2 mm long; ovary 0.1 diam., ovoid, tomentose; style 0.1 cm long, tomentose; stigma

punctate. Fruits drupaceous, 1-1.2 cm diam., ovoid, apex apiculate, shortly hirsute, reddish-brown, wrinkled

when dried.

Voucher specimens/material examined: BRAZIL. Amazonas: Jutai, Rio Solimoes, rio Bia, afluente do rio Jutai, 04 Nov 1975 (fl), L. Coelho et

al. 313 (INPA); Lajes, 17 Km from Manaus, 18 Feb 1975 (fr), G.T. Prance&J.F. Ramos P23278 (INPA, MO, NY); Sao Paulo de Olivenga, basin

of creek Belem, 26 Oct-Nov 1936 (fl), BA. Krukoff8683 (G, K, P, US). Para: Riojari, Planalto do Monte Dourado, 22 Jan 1968 (fl), E. Oliveira

3939 (IAN). GUYANA. Bartica, Essequibo river, Moraballi Creek, near sea level, 11 Nov 1929 (fl), N.Y. Sandwith 590 (K-3 sheets, NY); Maza-

runi Station, 07 Nov 1943 (fl), D.C.O. 67853 (NY); Pomeroon-Supenaam, Mabura, 22 Sep 1992 (fl), B. Hoffman & L. Roberts 2806 (US); Pota-

ro-Siparuni: Paramakatoi and vicinity, 13 Mar 1989 (fr.), W. Han et al. 5661 (K, US); Kaieteur Falls National Park, 28 Jan 1987 (fr) J.J. Pipoly

& G. Gharbarran 10168 (NY, P, US); Upper Demerara, 19 Nov 1986 (fl) J.J. Pipoly & R. Boyan 8845 (FDG, NY, P, US); ibid, 19 Nov 1986 (fl) J.J.

Pipoly & R. Boyan 8933 (FDG, NY, P, US); Upper Takutu-Upper Essequibo, 24 May 1997 (fr), D. Clarke 4932 (NY, US); ibid,15 May 1997 (fl),

2 cm

Amorim etal., Pleurisanthes flava in Brazil

171

Fig. 2. Pleurisanthes flava. A. Fertile branch (from 7.7. Pipoly&R. Boyan 8845); B. Inflorescence (from D.C.0.67853); C. Flower (from 7.7. Pipoly&R. Boyan

8845); D. Fruit (from 7.7. Pipoly&G. Gharbarran 10168).

172

Journal of the Botanical Research Institute of Texas 8(1)

D. Clarke 4552 (US); ibid., 12 Nov 1998 (fl), D. Clarke et al. 7788 (US); Waraputa Compartment, ca. 25 Km of Mabura, 05 Nov 1991 (fl bud),

M. Polak & P.J.M. Maas 506 (K, NY, U, US).

In Brazil, Pleurisanthesflava occurs in the states of Amazonas and Para and in different areas from the Brazilian

Amazon Forest. The species is recorded in lowland Amazon Forest and usually collected near streams and

river banks. The species could possibly be found in the states of Amapa and Roraima based on the current

knowledge and geographic distribution of the species. Pleurisanthesflava is most similar to P. howardii R. Duno,

Riina & P.E. Berry, which is endemic to Venezuela, but the leaves of P.flava are bicolorous, less coriaceous and

with the abaxial surface fulvo-strigose and the fruits are smaller, ovoid, shortly hirsute, reddish-brown, and

with the apex noticeably apiculate (Duno de Stefano et al. 2002). Pleurisanthesflava differs from P. artocarpi

Baill. and P. emarginata Tiegh. by the entire margin (clearly dentate in the other two species), from P. parviflora

(Ducke) R.A. Howard by the shorter pedicel, broader petals, and tomentose style (de Roon 1994; Sandwith

1931), and from P. simplicifolia by the oblong and membranaceous to chartaceous leaves, with entire margin,

and 8-10 secondary veins, (elliptic or suborbicular, coriaceous to subcoriaceouswith denticulate margin and 6

secondary veins, in P. simplicifolia).

KEY TO SPECIES OF PLEURISANTHES BAILL. FROM THE BRAZILIAN AMAZON RAINFOREST

[BASED ON HOWARD ( 1942 ) AND DE ROON ( 1994 )]

1. Leaves coriaceous to subcoriaceous, margin denticulate to dentate; flowers 4-5-merous.

2. Leaves obovate to obovate-elliptic, broadest at the base_ P. artocarpi

2. Leaves elliptic to widely elliptic, broadest at the middle.

3. Leaves with 7-9 secondary veins, flowers 4-merous_ P. emarginata

3. Leaves with 6 secondary veins, flowers 5-merous_ P. simplicifolia

1. Leaves membranaceous to chartaceous, margin entire; flowers 5-merous.

4. Flowers distincly pedicellate (1 cm long)_ P. parviflora

4. Flowers short pedicellate (0.1-0.2 cm long)_ P.flava

ACKNOWLEDGMENTS

We are indebted to the organizations which funded our held research, including CNPq-INCT, the U.S. Na¬

tional Science Foundation (DEB-0946618), Velux Stiftung, and the Benehcia Foundation; PNADB/Capes; The

Department of Botany of the Smithsonian Institution that provided the Cuatrecasas Fellowship to the first

author; and Fundagao de Amparo a Ciencia e tecnologia do Estado de Pernambuco (FACEPE) for the first au¬

thor’s Ph.D. grant. We also thank the curators of the cited herbaria, Aline Melo who facilitated the access to

collections from Northern Brazil, Laurence Dorr and Deborah Bell from US, Jacquelyn Kallunki and Stella

Sylva from NY, Eve Lucas from K, Germinal Rouhan from P, and Nicolas Fumeaux and Louis Nusbaumer from

G for the support; the anonymous reviewer for constructive criticism and Barney Lipscomb for editorial sug¬

gestions; to Regina Carvalho for the illustrations, and M.Sc. Jefferson Maciel for the distribution map.

REFERENCES

Duno de Stefano, R. 2004. Icacinaceae.ln: N. Smith, S.A. Mori, A. Henderson, D.Wm. Stevenson, and S.V. Heald, eds. Flower¬

ing plants of the Neotropics. Princeton University Press, Princeton, New Jersey, U.S.A.

Duno de Stefano, R. & B. Amorim. 2012. Icacinaceae. In: Lista de especies da Flora do Brasil. Jardim Botanico do Rio de Ja¬

neiro. http://floradobrasil.jbrj.gov.br/2011/FB000135.Accessed October 2012.

Duno de Stefano, R., R. Riina, & P.E. Berry. 2002. Pleurisanthes howardii (Icacinaceae), a new species from the Venezuelan

Guayana. Harvard Pap. Bot. 7:13-15.

Howard, R.A. 1942. Studies of the Icacinaceae IV. Consideration of the New World Genera. Contr. Gray Herb. Harvard

Univ. 142:1-59.

Howard, R.A. & R. Duno de Stefano. 1999. Icacinaceae. In: P.E. Berry, K. Yatskievych, and B.K. Host, eds. Flora of the Venezu¬

elan Guayana. Vol 5. Missouri Botanical Garden Press, Saint Louis, Missouri, U.S.A.

KArehed, J. 2001. Multiple origin of the tropical forest tree family Icacinaceae. Amer. J. Bot. 88:2259-2274.

Kelloff, C.L., N. Sara, S.N. Alexander, V.A. Funk, & H.D. Clarke. 2011. Smithsonian plant collections, Guyana: 1995-2004, H.

David Clarke. Smithsonian Contr. Bot. 97. Smithsonian Institution Scholarly Press, Washington, D.C., U.S.A.

Amorim etal., Pleurisanthes flava in Brazil

173

de Roon, A.C. 1994. Icacinaceae. In: A.R.A. Gorts-van Rijn, ed. flora of the Guianas. Koeltz Scientific Books, Koenigstein,

Germany. Pp. 82-109.

Sandwith, N.Y. 1931. Contributions to the flora of Tropical America: VIII. Bull. Misc. Inform. Kew 1931(10):467-468.

Thiers, B. 2013 [continuously updated]. Index Herbariorum: A global directory of public herbaria and associated staff.

New York Botanical Garden's Virtual Herbarium, online at http://sweetgum.nybg.org/ih/.

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Journal of the Botanical Research Institute of Texas 8(1)

JOURNAL NOTICE

Douglas J. Futuyma, H. Bradley Shaffer, and Daniel Simberloff, eds. 2013 (Dec). Annual Review of Ecology,

Evolution, and Systematics, Volume 44. (ISSN: 1543-592X; ISBN: 978-0-8243-1444-6, hbk). Annual

Reviews, Inc., 4139 El Camino Way, RO. Box 10139, Palo Alto, California 94303, U.S.A. (Orders: www.

AnnualReviews.org, science@annualreviews.org, 1-800-523-8635, 1-650-493-4400). $92.00 indiv.,

658 pp., 7 5 /8" x 9%".

ABOUT THIS JOURNAL—The Annual Review of Ecology, Evolution, and Systematics, in publication since 1970, covers significant develop¬

ments in the fields of Ecology, Evolutionary Biology, and Systematics, as they apply to all life on Earth. Essay reviews cover topics ranging

from phylogeny, speciation, and molecular evolution through behavior and evolutionary physiology to population dynamics, ecosystems

processes, and applications in invasion biology, conservation, and environmental management.

Contents of Volume 44:

Genomics in Ecology, Evolution, and Systematics Theme

1. Introduction to Theme “Genomics in Ecology, Evolution, and Systematics”— H.B. Shaffer & M.D. Purugganan

2. Genotype-by-Environment Interaction and Plasticity: Exploring Genomic Responses of Plants to the Abiotic Environment—D.L.

Des Marais, K.M. Hernandez, & T.E.Juenger

3. Patterns of Selection in Plant Genomes— J. Hough, R.J. Williamson, & S.I. Wright

4. Genomics and the Evolution of Phenotypic Traits—G.A. Wray

5. Geographic Mode of Speciation and Genomic Divergence— J.L. Feder, S.M. Flaxman, S.P. Egan, A.A. Comeault, & P. Nosil

6. High-Throughput Genomic Data in Systematics and Phylogenetics—E. M oriarty Eemmon &A.R. Eemmon

7. Population Genomics of Human Adaptation— J. Lachance & S.A. Tishkoff

Topical Reviews

8. Symbiogenesis: Mechanisms, Evolutionary Consequences, and Systematic Implications—T. Cavalier-Smith

9. Cognitive Ecology of Food Hoarding: The Evolution of Spatial Memory and the Hippocampus—V.V. Pravosudov & T.C. Roth II

10. Genetic Draft, Selective Interference, and Population Genetics of Rapid Adaptation—R.A. Neher

11. Nothing in Genetics Makes Sense Except in Light of Genomic Conflict— W.R. Rice

12. The Evolutionary Genomics of Birds— H. Ellegren

13. Community and Ecosystem Responses to Elevational Gradients: Processes, Mechanisms, and Insights for Global Change—M.K.

Sundqvist, N.J. Sanders, & DA. Wardle

14. Cytonuclear Genomic Interactions and Hybrid Breakdown—R.S. Burton, R.J. Pereira, & F.S. Barreto

15. How Was the Australian Flora Assembled Over the Last 65 Million Years? A Molecular Phylogenetic Perspective—M.D. Crisp &

E.G. Cook

16. Introgression of Crop Alleles into Wild or Weedy Populations—N.C. Ellstrand, P. MeirmansJ. Rong, D. Bartsch, A. Ghosh, T.J. dejong,

P. Haccou, B.-R. Eu, A.A. Snow, C.N. StewartJr.,J.E. Strasburg, PH. van Tienderen, K. Vrieling, & D. Hooftman

17. Plant Facilitation and Phylogenetics—A. Valiente-Banuet & M. Verdu

18. Assisted Gene Flow to Facilitate Local Adaptation to Climate Change— S.N. Aitken & M.C. Whitlock

19. Ecological and Evolutionary Misadventures of Spartina—D.R. Strong & D.R. Ayres

20. Evolutionary Processes of Diversification in a Model Island Archipelago—R.M. Brown, C.D. Siler, C.H. 01iveros,J.A. Esselstyn, A.C.

Diesmos, PA. Hosner, C.W. Linkem, A.J. Barley, J.R. Oaks, M.B. Sanguila, L.J. Welton, D.C. Blackburn, R.G. Moyle, A.T. Peterson, &A.C.

Alcala

21. Perceptual Biases and Mate Choice— M.J. Ryan & M.E. Cummings

22. Thermal Ecology, Environments, Communities, and Global Change: Energy Intake and Expenditure in Endotherms—N. Kronfeld-

Schor & T. Dayan

23. Diversity-Dependence, Ecological Speciation, and the Role of Competition in Macroevolution—D.L. Rabosky

24. Consumer Fronts, Global Change, and Runaway Collapse in Ecosystems— B.R. Silliman, M.W. McCoy, C. Angelini, R.D. Holt,J.N.

Griffin, & J. van de Koppel

25. Implications of Time-Averaged Death Assemblages for Ecology and Conservation Biology—S.M. Kidwell &A. Tomasovych

26. Population Cycles in Forest Lepidoptera Revisited— J.H. Myers&J.S. Cory

27. The Structure, Distribution, and Biomass of the World’s Forests—Y. Pan, R.A. Birdsey, O.L. Phillips, & R.B. Jackson

28. The Epidemiology and Evolution of Symbionts with Mixed-Mode Transmission—D. Ebert

Cumulative Index of Contributing Authors, Volumes 40-44

Cumulative Index of Article Titles, Volumes 40-44

J.Bot. Res. Inst. Texas 8(1): 174.2014

ECOLOGY AND CONSERVATION OF ACACIA AND PROSOPIS (FABACEAE)

WOODLANDS OF THE MOJAVE DESERT, U.S.A.

Scott R. Abella 1 and Kenneth L. Chittick

Department of Environmental and Occupational Health

University of Nevada, Las Vegas

Las Vegas, Nevada 89154-3064, U.S.A.

abellaNRC@gmail.com

Present address: Natural Resource Conservation LLC, Boulder City, Nevada 89005, U.S.A.

ABSTRACT

Woodlands of Acacia greggii, Prosopis glandulosa, and Prosopis pubescens are of conservation-priority in the Mojave Desert because of their

wildlife and watershed values. We measured plant community composition, environmental variables (e.g., slope gradient, soil), and ecolog¬

ical condition (e.g., tree recruitment) in 50, 0.1-ha woodland plots within 449,000-ha Take Mead National Recreation Area in the eastern

Mojave Desert in Arizona-Nevada. We classified community types, analyzed vegetation-environment relationships, developed ecological

species groups (species sharing similar distributions), and evaluated woodland condition. Cluster analysis identified 5 community types at

the finest hierarchical level, which were quite distinct floristically (53% mean Sorensen similarity within communities), and included an A.

gpeggii community occupying dry washes, 2 P. glandulosa communities, a mixed community, and a P. pubescens community inhabiting

drainage outflows of springs. We recorded a total of 201 taxa. Mean species richness varied significantly among communities from 10 in P.

pubescens to 35 species/0.1 ha in mixed communities. Environmental variables such as soil texture and cations were related to community

gradients, distributions of tree species, and frequency of the tree parasite desert mistletoe ( Phoradendron californicum). We classified 73

species into 14 species groups, ranging from groups characteristic of uplands (e.g., Larrea tridentata group) to those most frequent in low¬

lands (e.g., Allenrolfea occidentalis group). Ecological condition of the woodlands was characterized by well-distributed tree density among

size classes (except for P. pubescens communities which were dominated by large trees), dominance by native species (94% of total taxa were

native), mistletoe infection on 66% of plots, and infrequent evidence of perceived threats (e.g., woodcutting).

RESUMEN

Los bosques de Acacia greggii, Prosopis glandulosa, y Prosopis pubescens tiene prioridad de conservation en el desierto de Mojave por sus va-

lores ambientales. Se midieron la composicion de la comunidad vegetal, variables ambientales (ej., gradiente de la ladera, suelo), y condicio-

nes ecologicas (ej., reclutamiento de arboles) en 50 parcelas, de 0.1-ha en las 449,000-ha del Lake Mead National Recreation Area en el este

del desierto de Mojave entre Arizona y Nevada. Se clasificaron los tipos de comunidad tipos, se analizaron las relaciones de la vegetacion con

el ambiente, se desarrollaron grupos ecologicos de especies (especies que comparten distribuciones similares), y se evaluo la condicion del

bosque. El analisis cluster identified 5 tipos de comunidad en el nivel jerarquico mas fino, que fueron bastante distintos floristicamente (53%

de media en el indice de similitud de Sorensen en las comunidades), e incluyo una comunidad de A. greggii que ocupa humedales secos, 2

comunidades de P. glandulosa, una comunidad mixta, y una comunidad de P. pubescens que vive en drenajes de manantiales. se registraron

un total de 201 taxa. La riqueza media de especies vario significativamente entre comunidades de 10 en P pubescens a 35 especies/0.1 ha en

comunidades mixtas. Las variables ambientales tales como textura del suelo y cationes estaban relacionadas con los gradientes de la comu¬

nidad, distribuciones de especies arboreas, y frecuencia de muerdago pargasito de los arboles ( Phoradendron californicum). Se clasificaron 73

especies en 14 grupos de especies, variando desde grupos caracteristicos de las tierras altas (ej., grupo de Larrea tridentata ) a los mas fre-

cuentes en tierras bajas (ej., grupo de Allenrolfea occidentalis). Las condiciones ecologicas de los bosques se caracterizaron por su densidad

de arboles bien distribuida entre las clases de tamano (excepto las comunidades de P. pubescens que estaban dominadas por arboles grandes),

dominancia de las especies nativas (94% del total de taxa fueron nativos), infeccion de muerdago en el 66% de las parcelas, y evidencia infre-

cuente de amenazas percibidas (ej., talas).

INTRODUCTION

Riparian plant communities in the arid American Southwest occupy small portions of landscapes but have

disproportionately large habitat value, productivity, and services to humans (Sada et al. 2001; Patten et al.

2008). The valuable functions that riparian ecosystems provide—such as water to sustain human habitations,

agriculture, and ranching—and their native biota are threatened by past and present intensive human use of

these habitats (Deacon et al. 2007). In the eastern Mojave Desert, for instance, Acacia greggii, Prosopis glandu¬

losa, and P. pubescens riparian woodlands have been destroyed or altered through hydrologic changes and ur-

J. Bot. Res. Inst. Texas 8(1): 175 -195.2014

176

Journal of the Botanical Research Institute of Texas 8(1)

ban development in Clark County containing metropolitan Las Vegas, Nevada (Crampton & Sedinger 2011).

Now covered under a multiple species habitat conservation plan (MSHCP) to forestall U.S. Endangered Species

Act listing of associated species, conservation goals for Acacia and Prosopis woodlands in this region include

restoring and maintaining the land area occupied by the woodlands in 2000 (inception of MSHCP), sustaining

protected communities in a healthy ecological condition (e.g., well-distributed tree size classes, moderate in¬

fection of the tree parasite desert mistletoe [Phoradendron californicum], and dominance by native species), and

maintaining species affiliated with the woodlands (Crampton et al. 2006).

However, significant knowledge gaps in our understanding of the ecology and conservation needs for

these woodlands hinder development of conservation strategies (Crampton et al. 2006). For example, commu¬

nity structure, vegetation-environment relationships, understory composition, and ecological condition of the

woodlands including exotic plant invasion status, tree recruitment, desert mistletoe infection, and disturbanc¬

es such as fire or woodcutting, are poorly understood. Some community classification has been performed in

parts of the California Mojave Desert (Evens 2003; Thomas et al. 2004; Keeler-Wolf et al. 2007), but little vege¬

tation-environment research for these communities in the American Southwest has been conducted and con¬

clusions have varied. Some reports in the literature have included that distribution of Prosopis pubescens com¬

munities was unrelated to gradients in soil pH, soluble salts, or texture along the Rio Grande River in central

New Mexico (Campbell and Dick-Peddie 1964). Along the San Pedro River in the Chihuahuan and Sonoran

Deserts, Prosopis velutina patches occupied sites with low frequency of flooding and highest elevations away

from the active flood channel (Bagstad et al. 2006). In Mojave Desert ephemerally moist washes, Evens (2003)

noted that Acacia greggii occurrences correlated to elevation and amount of topographic protection (concave

sites exhibit high protection).

Distinguishing ecological species groups is another means to understand species distributions and vege¬

tation-environment relationships (Goebel et al. 2001). Species groups consist of co-occurring species that

share similar environmental affinities and are based on classifying species (rather than communities) into

groups usually of 2-10 species displaying similar distributions (Kashian et al. 2003). For example, on a north¬

ern Arizona Pinus ponderosa forest landscape to the east of the Mojave Desert, we previously classified 18 spe¬

cies groups ranging from plants inhabiting xeric, volcanic cinder soils, to those typifying moist, silt loam soils

(Abella & Covington 2006). Species groups are based on a premise that once the groups are developed, pres¬

ence of some species of a group suggests that environmental characteristics of a site are within the realized

niche of the group (Kashian et al. 2003). Ecological species groups have been little developed in southwestern

deserts. Species groups have been valuable on other landscapes for understanding vegetation-environment

relationships and for management applications such as matching species for ecological restoration to environ¬

ments where they are best adapted (Goebel et al. 2001).

Exotic plant invasion, tree recruitment, mistletoe infection, and disturbance are additional features re¬

lated to ecology and condition of Acacia and Prosopis communities (Stromberg 1993). For example, riparian

communities can be highly invadable because of their location along seed dispersal corridors and their re-

source-rich environment favorable for plant growth (Tabacchi & Planty-Tabacchi 2005). Exotic plant abun¬

dance is important to evaluate if dominance by native species is considered a measure of woodland health.

Presence of a range of tree size classes is another feature considered desirable for high-quality habitat condi¬

tions (Crampton & Sedinger 2011). Tree size and age are not always correlated, but size class analyses are useful

for identifying trees that became established more recently than the current largest trees (Miller et al. 2001).

Moreover, tree size distribution is important for several other reasons such as suitability of nesting sites for

avian species and amount of parasitic mistletoe a tree can support as a food resource for wildlife (Crampton &

Sedinger 2011). Mistletoe extracts water and nutrients through a vascular connection to the host tree, with

larger trees generally supporting more mistletoe (Watson 2001). Mistletoe is a key food and nesting resource

for Phainopepla nitens, a conservation-priority bird species covered by the MSHCP, so intermediate amounts of

mistletoe are a good indicator of habitat value at a level sustainable to avoid killing trees (Crampton & Sedinger

2011). Disturbances such as fire or woodcutting also can affect ecological condition of woodlands. The wood-

Abella and Chittick, Acacia and Prosopis woodlands of the Mojave Desert

177

land tree species have some resprouting ability when burned or cut, but these disturbances can reduce their

abundance (Stromberg 1993; Busch 1995; Abella 2010).

To help bll knowledge gaps in the ecology of Acacia and Prosopis woodlands and support development of

conservation strategies, we examined plant community structure, vegetation-environment relationships, and

ecological condition of these communities on a Mojave Desert landscape of Lake Mead National Recreation

Area. Under National Park Service protection, this landscape is viewed as a core conservation area by the

MSHCP, which indicates that maintaining quality woodland habitat on this landscape is a key part of conserv¬

ing these communities in the eastern Mojave Desert (Crampton et al. 2006). Specific study objectives were to:

(1) develop a hierarchical classification and identify diagnostic species for Acacia and Prosopis woodlands; (2)

identify vegetation-environment relationships of communities and distributions of tree species; (3) develop

ecological species groups; (4) examine species richness relationships with community types, environmental

gradients, and exotic species; and (5) assess current woodland condition, including tree recruitment, mistletoe

infection, and evidence of disturbances such as fire or woodcutting.

METHODS

Study Area

We conducted this study in Lake Mead National Recreation Area, a 449,000-ha unit (excluding full-pool areas

of Lakes Mead and Mohave) of the National Park Service, in southeastern Nevada and northwestern Arizona

in the eastern Mojave Desert (35°59'N, 114°5PW; Fig. 1). The centrally located Boulder City, Nevada, weather

station has reported the following averages: 14 cm/yr of precipitation, 4°C January daily low temperature, and

39°C July high temperature (768 m elevation, 1937-2004 records; Western Regional Climate Center, Reno,

Nevada). Consistent with the Mojave Desert’s status as a winter rainfall desert (Keeler-Wolf et al. 2007), 70% of

precipitation falls from September through April. Predominant landforms include low mountain ranges, ba-

jadas (coalesced alluvial fans), relatively flat plains, washes serving as intermittent drainageways, and playas

(dry lakes). Mapped soil types include Aridisol and Entisol orders (Lato 2006). Uplands, which occupy >90%

of the area, are dominated by shrublands of Larrea tridentata and Ambrosia dumosa (Abella et al. 2012a). Com¬

munities containing Acacia or Prosopis are associated with washes, springs, and topographically protected sites

(Fig. 2). Major large herbivores include exotic Equus asinus in some areas and native Ovis canadensis and small¬

er animals such as Lepus californicus. Some unauthorized cattle grazing occurs in the northeastern part of the

study area. Human recreation use is concentrated along access points of Lake Mead’s shoreline and Colorado

River south of Hoover Dam and along major roads (Fig. 1).

DATA COTTECTION

We used an existing map of Acacia greggii, Prosopis glandulosa, and Prosopis pubescens distribution within the

study area (Crampton et al. 2006), combined with our own held reconnaissance, to identify 118 polygons >0.25

ha and containing >2% cover of one or more of these species. This cover criterion excluded sampling sites con¬

taining only an individual tree. We randomly selected 50 of these polygons for sampling, ranging in size from

0.25-89 ha. We generated a random Universal Transverse Mercator (UTM) coordinate using a geographic in¬

formation system (ArcMap, Esri Corporation, Redlands, California) within each polygon (subdivided by tree

species) at which to establish a plot. Plots were 0.1 ha and were 20 m x 50 m (45 of the 50 plots) where the

landform allowed; otherwise 33.3 m x 33.3 m (5 plots). We sampled plots from July-October 2011, during the

leaf-on period for the deciduous Acacia and Prosopis.

We measured the plant community on each plot by visually categorizing areal cover of vascular plant

species using cover classes: trace (assigned 0.2% for analyses), < 1% (assigned 0.5%), 1% intervals to 5%, and

thereafter 5% intervals. The same botanist measured all plots for consistency in cover categorization. Along

with live plants, standing dead annual plants, noted to persist for 1-2 years in the Mojave Desert (Beatley

1966), were included in sampling to more thoroughly characterize the annual plant community. Plants not

identifiable in the held were collected, pressed, and keyed to the finest taxonomic level possible. Four speci-

3920000 3960000 4000000 4040000

178

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 1. Distribution of 50 sample plots displayed by community type and locations mentioned in the text for Acacia and Prosopis woodlands of the

eastern Mojave Desert, USA. The inset at the top right includes mapped polygons of the tree species with randomly located plots within. Coordinates

are Universal Transverse Mercator (m). North American Datum 1983.

Abella and Chittick, Acacia and Prosopis woodlands of the Mojave Desert

179

Fig. 2. Examples of woodland community types of the eastern Mojave Desert, USA: [a) Acacia greggii/msh, (b) Mixed/variable, (c) Prosopis glandulosa/

protected, (d) Prosopisgianduiosa/ gypsum, and (e) Prosopispubescens/ spring. Desert mistletoe is shown in the foreground of (f) parasitizing an Acacia

greggii tree.

mens out of 1385 total plant records across plots were not identifiable to at least family and were deleted from

the data set. Classification of taxa to growth form (forb, shrub, or tree), life span (e.g., annual), and native/exot¬

ic status to North America followed USDA, NRCS (2012).

Live and dead individuals (of all sizes including seedlings) for all tree species were counted on each plot

and their height was determined using a measuring pole. We measured diameter at root collar for the largest

stem for all individuals and diameter at breast height (137 cm) for each tree taller than breast height. To meas¬

ure mistletoe infection on each tree, we used the Hawksworth (1977) 6-class mistletoe rating. Infection was

180

Journal of the Botanical Research Institute of Texas 8(1)

recorded as none (assigned 0), light (<50% of branches infected, assigned 1), or heavy (>50% of branches infect¬

ed, assigned 2) for each third of the tree. The values were summed to result in a 0 (no infection) to 6 (heavy)

rating (Hawksworth 1977).

Data regarding depth to groundwater and ground-water chemistry would be desirable. These data were

not available for the study area (Gary Karst, Hydrologist, Lake Mead National Recreation Area, pers. comm.)

and were difficult or not permissible to obtain through drilling monitoring wells. We were able, however, to

collect a variety of environmental data for each plot including location, topography, disturbance, and soil. We

recorded elevation and location (UTM, using a global positioning system, at the southwestern plot corner),

slope gradient (clinometer), aspect (compass), and landform type (e.g., wash; following Lato [2006]). In addi¬

tion to capturing possible variation unaccounted for by other environmental variables, location can represent

influences such as historical disturbance difficult to detect but potentially influencing site-specific vegetation

patterns. We linearized aspect to range from 0 (southwest) to 2 (northeast; Beers et al. [1966]). We obtained

1971-2000 mean annual precipitation and temperature for each plot location from PRISM (Daly et al. 2008).

We qualitatively noted visual evidence of disturbance, such as fire, off-road vehicle tracks, woodcutting, and

livestock presence (animals or dung).

We collected 3 subsamples of the 0-5 cm mineral soil from each of 3 different interspaces >1 m from the

outermost edge of a tree canopy. To measure bulk density, we collected a sample of approximately 400 cm 3

from the same interspaces. Soil samples were composited by plot. We sieved air-dried analytical samples to

pass a 2-mm sieve and analyzed the fine fraction for texture (hydrometer method) following Tan (2005); pH

and electrical conductivity (1:1 soibwater); available P (Olsen sodium-bicarbonate extraction); CaC0 3 (ma¬

nometer method); total C, N, and S (dry combustion, CNS analyzer); organic C (difference between total and

inorganic C); N0 3 , S0 4 , and Cl (ion chromatography); and the water-soluble concentrations of Na, K, Mg, Ca,

Mn, Fe, Ni, Cu, Zn, Co, B, Mo, Pb, and Cd (1:3 soibwater extracts, inductively coupled plasma mass spectros¬

copy) following Burt (2004). We estimated bulk density by sieving through a 2-mm sieve, oven drying the <2-

mm fraction at 105°C for 24 h, and including volume of coarse fragments >2 mm in the total soil volume. We

used bulk density to convert nutrient concentrations to volumetric contents (Burt 2004). Because concentra¬

tions and contents were strongly correlated (e.g., r = 0.95 for organic C, 0.92 for total N, and 1.00 for total S), we

report concentrations.

DATA ANALYSIS

We conducted multivariate plant community and soil analyses using version 6.07 of PC-ORD software (Mc-

Cune and Mefford 1999). We used hierarchical cluster analysis (Sorensen index and flexible beta [p = -0.25]

linkage method) to classify plots by species composition based on relative cover (cover of species/cover of all

species on a plot). To identify species with the greatest fidelity to each hierarchical plot grouping, we used indi¬

cator species analysis to produce an indicator value ranging from 0 (no fidelity) to 100% (highest fidelity) based

on relative cover and relative frequency among the groups (Dufrene & Legendre 1997). We ordinated species

composition (relative cover) with non-metric multi-dimensional scaling through PC-ORD’s autopilot, slow

and thorough routine. Environmental variables and species displaying the strongest correlations with com¬

munity compositional patterns were displayed as vectors scaled to the strength and direction of correlations.

We ordinated soil composition using principal components analysis, with the cross-products matrix derived

from correlation to account for different measurement scales of soil variables.

We used SAS 9.2 software (SAS Institute 2009) to conduct univariate and bivariate analyses. We used a

Kruskal-Wallis test followed by Tukey’s test on ranks for multiple comparisons to compare species richness

among plant community types at the finest level of the cluster analysis community classification. We used

Pearson correlation to assess the relationship of native and exotic species richness.

To identify biophysical correlates with distribution of community types, tree species, and measures of

ecological condition (exotic species richness and cover, tree recruitment, and mistletoe), we used classification

(for categorical response variables) and regression trees (for continuous response variables) in JMP 9 software

(SAS Institute 2010). Regression trees are nonparametric models that partition data into increasingly homog-

Abella and Chittick, Acacia and Prosopis woodlands of the Mojave Desert

181

enous subsets and provide dichotomous keys to estimate a dependent variable at different levels of explanatory

variables (Breiman et al. 1984). Dependent variables were screened for inclusion in models based on a criterion

of minimizing total sums of squares at different splits. Splitting stopped when adding more explanatory vari¬

ables increased r 2 by <0.05 or when the minimum node size (n = 5 for most analyses) was reached. There is

essentially no limit to the number of independent variables that can be input to each model because a screening

process identifies variables with the strongest explanatory power for inclusion in final models (SAS Institute

2010). We employed JMP’s Je-fold crossvalidation (k = 3 or 5) to compute a cross-validated r 2 . We explored

modelling several tree recruitment (e.g., trees/ha or percent of total trees by height and diameter at root collar

and breast height classes) and mistletoe (e.g., proportion of infected trees, infected trees/ha, total Hawksworth

rating) measures. The final model for recruitment portrayed percent of trees in the 1-9 cm root collar diameter

class, because root collar differentiated trees with large stems that might be short in height yet still old (Miller

et al. 2001) and the model displayed the highest r 2 among recruitment measures. We chose the final mistletoe

model to portray infected trees/ha because Crampton and Sedinger (2011) found that this measure was corre¬

lated to Phainopepla nitens nesting preference and this response variable also yielded the highest model r 2 in

our study.

We constructed ecological species groups by: (1) including only species occupying >3 plots; (2) relativiz-

ing species cover by species sums of squares to emphasize habitat preferences, avoiding groupings based on the

commonness or rarity of species (McCune et al. 2000); and (3) grouping species through cluster analysis (So¬

rensen distance and -0.25 Flexible Beta group linkage) in PC-ORD (McCune & Mefford 1999). We used Pear¬

son correlation to relate average cover of species groups to principal components and environmental variables.

RESULTS

Community Classification

At the coarsest grouping, cluster analysis classified plots into an Acacia greggii- dominated group and those

containing Prosopis spp. (Fig. 3). Finer groupings distinguished Prosopis pubescens, two types of Prosopis glan-

dulosa communities, and a mixed community of A. greggii, Prosopis spp., and other species. Sorensen similar¬

ity among plots within a community at the finest level ranged from 42% (Acacia greggii/ wash) to 66% (Prosopis

glandulosa/ protected) and averaged 53 ± 10% (± SD, n = 5 community types). We named the 5 community

types at the finest hierarchical level according to dominant tree species and either a commonly associated

topographic feature or soil parent material (Table 1).

There were significant indicator species at each level of the community hierarchy. At the finest level, un¬

derstory species such as Hymenoclea salsola were significantly associated with the Acacia greggii/ wash com¬

munity; cacti species and Baccharis salicifolia with Mixed/variable; Isocoma acradenia, Atriplex confertifolia,

and Suaeda moquinii with Prosopis glandulosa/ gypsum; and Allenrolfea occidentalis and Distichlis spicata with

Prosopis pubescens/ spring. Cluster analysis combined with indicator species analysis suggested that the vegeta¬

tion was readily distinguishable into community types at multiple hierarchical levels.

Species Richness

A total of 201 taxa (90% identified to species) were detected on plots. This flora consisted of 61 annual forb

(30%), 47 shrub (23%), 41 perennial forb (20%), 14 annual-perennial forb (7%), 9 cactus (4%), 9 perennial

graminoid (4%), 7 tree (3%), 6 annual grass (3%), 5 annual-biennial forb (2.5%), 1 annual-perennial grass

(0.5%), and 1 perennial fern (0.5%). Species richness varied significantly (Kruskal-Wallis % 2 = 18.4; P = 0.001)

among communities from 10 (Prosopis pubescens/ spring) to 35 species/0.1 ha (Mixed/variable; Fig. 4A). Even

including dead annual plants, richness in all communities was dominated by perennials.

Vegetation and Soil Gradients

The vegetation ordination corroborated cluster analysis with distinct community groupings evident in the or¬

dination (Fig. 5A). Whereas Acacia greggii- dominated plots clearly separated from those of other communities,

plots within this community displayed a large spread consistent with their low similarity (42%) in cluster

analysis. Acacia greggii plots in the lower part of the ordination grouping had the greatest relative cover of Aca-

182

Journal of the Botanical Research Institute of Texas 8(1)

Similarity = 42%, n = 29

Acacia greggii 87

Hymenoclea salsola 83

Bromus rubens 62

Bebbia juncea 53

Eriogonum fasciculatum 52

Similarity = 66%, n = 10

Prosopis glandulosa 71

Acacia Prosopis

gregg i i / glandulosa /

wash protected

Similarity = 36%, n =

Prosopis glandulosa

Similarity = 46%, n = 18

Prosopis glandulosa 99

-o-

Similarity = 41%, n = 8

Opuntia basilaris 73

Isocoma acradenia 54

Mixed /

variable

Similarity = 61%, n = 3

Allenrolfea occidentalis 100

Prosopis pubescens 98

Distichlis spicata 63

Similarity = 44%, n = 4

Cylindropuntia acanthocarpa 90

Ferocactus cylindraceus 63

Opuntia basilaris 62

Baccharis salicifolia 50

Similarity = 54%, n = 4

Isocoma acradenia 70

Atriplex confertifolia 67

Suaeda moquinii 66

Stylocline intertexta 65

Descurainia pinnata 52

Atriplex hymenelytra 50

ui_

Prosopis Prosopis

glandulosa / pubescens /

gypsum spring

Fig. 3. Hierarchical community classification of Acacia and Prosopis woodlands of the eastern Mojave Desert, USA. Internal Sorensen similarity, number

of plots, and indicator species and indicator values significant at P <0.05 and >50 are shown at each division. Community types at the finest level of

the classification are named according to dominant tree species and environmental features.

da, whereas plots in the upper part exhibited greater relative cover of species such as Ambrosia dumosa, Hy¬

menoclea salsola, and Eriogonum fasciculatum. Some environmental variables were correlated with vegetation

gradients. Coarse-textured soil, for example, was associated with A. greggii communities, and total S and sev¬

eral cations correlated with Prosopis glandulosa/ gypsum communities.

In contrast to vegetation that clearly grouped into community types, ordination of soil properties did not

display strong grouping (Fig. 5B). Some plots, associated with gypsum, were correlated with soil electrical

conductivity, S0 4 , and various elements along axis 1. Axis 2 displayed few relationships.

Whereas multivariate variation in soil properties was not strongly linked to plant community gradients,

examining means in individual soil properties suggested several findings (Table 2). Variability across commu¬

nities differed among properties, with some properties (e.g., B) displaying extreme variation in orders of mag¬

nitude. Certain properties, such as total S, were highly variable within communities (e.g., 398% coefficient of

variation for S in Acacia greggii/ wash), yet some of these properties like S were still orders of magnitude greater

in one or more communities. A community could occupy a range of values in soil properties, but there were

some properties exhibiting especially large or small values in particular communities.

Community and Tree Species Distribution

Effectiveness of classification or regression tree models and variables they included differed for portraying

distributions of communities and tree species (Fig. 6). A classification tree selected UTM coordinates and sand

concentration as most important for differentiating communities. High sand concentration was again related

to Acacia greggii/ wash communities, as in the ordination. The UTM coordinates corresponded with distribu¬

tion of soil parent materials, such as gypsum, which occupied the northeastern part of the study area, and top¬

ographic features like Black Canyon (below the Hoover Dam; Fig. 1). A classification tree of Prosopis pubescens

with gypsum). A regression tree for A. greggii canopy cover illustrated that soil of low electrical conductivity

Abella and Chittick, Acacia and Prosopis woodlands of the Mojave Desert

183

Table 1. Characteristics of Acacia and Prosopis woodland community types of Lake Mead National Recreation Area, Mojave Desert, USA.

Acacia greggii/

wash

Mixed/

variable

Prosopis glandulosa/

protected

Prosopis glandulosa/

gypsum

Prosopis pubescens/

spring

Number of plots

Live trees/ha (mean±SD)

230±126

330±88

239±79

508±319

197±124

Live trees/ha (range)

50-600

200-390

120-370

170-910

120-340

Acacia greggii (%)

Prosopis glandulosa (%)

100

Prosopis pubescens (%)

Live trees infected

100

(%, mean±SD)

Live trees infected

19±23

16± 15

19±16

11 ±13

50±25

(%, range)

0-86

0-35

0-48

2-29

21-67

Elevation (m, mean±SD)

581±255

457±93

370±170

424±47

460±9

Elevation (m, range)

Slope gradient

201-1154

381-590

198-685

379-469

455-471

(%, mean±SD)

4±6

10±6

17±19

3±11

6±3

Slope gradient (%, range)

1-31

3-18

2-52

2-4

3-9

Topography

Soil classification

Washes

Variable

Canyons, concave

Washes, depressions

Spring drainages

(great group)

Torriorthents

Haplocalcids

Torriorthents

Haplocalcids

Torriorthents

Haplogypsids

Haplocalcids

Petrogypsids

Haplogypsids

had the greatest canopy cover. On soil with higher conductivity, the greatest A. greggii canopy cover occurred

on soil with low S0 4 and high gravel concentration. Canopy cover of Prosopis glandulosa exhibited a different

pattern: it was greatest on soil rich in N0 3 , or on sites with steep slopes when soil N0 3 was low.

Ecological Condition and Species Groups

Thirteen exotic species were detected, including the annual grasses Bromus rubens (86% of plots), Schismus

spp. (76%), Polypogon monspeliensis (10%) and Bromus tectorum (8%); annual forbs Brassica tournefortii (6%),

Salsola tragus (6%), Malcolmia africana (4%), and Sonchus asper (2%); annual-biennial forbs Erodium cicutarium

(28%), Lactuca serriola (2%), and Sisymbrium altissimum (2%); the perennial forb Marrubium vulgare (2%); and

the tree Tamarix ramosissima (22%). Prosopis pubescens/ spring contained no plots with exotic species, and aver¬

age exotic richness ranged from 2-4 species/0.1 ha in the other communities. Native and exotic species rich¬

ness were positively related (r = 0.47; Fig. 4B). This correlation was larger than the next highest correlations that

exotic richness exhibited (r = 0.35 with precipitation and 0.34 with UTMx).

A regression tree portraying exotic species richness accounted for 40% of variance upon crossvalidation

(Fig. 7A), which was higher than the 17% for exotic cover (not shown). The regression tree indicated that sites

with >16 cm/yr of precipitation and silty soil contained the most exotic species. The least exotic rich were drier

sites, especially those with high soil S.

Mistletoe was present on 33 of 50 plots (66%), and the percentage of trees infected was 50% in the Proso¬

pis pubescens/ spring community and lower (11-19%) in other communities (Table 1). The greatest density of

infected trees portrayed by a regression tree occurred on soil with low CaC0 3 concentration, or, if CaC0 3

concentration was high, on higher elevation sites with high tree canopy cover (Fig. 7B).

Tree size-class distributions revealed that densities were variable within diameter classes among sites

within communities (Fig. 8). This resulted from some sites containing few or no trees in some size classes.

Communities as a whole exhibited good representation of small trees, with generally as many or more trees in

small as large size classes. The exception was the Prosopis pubescens/ spring community, which mostly con¬

tained large trees. Density of trees in the smallest size class (1-9 cm diameter at root collar) was positively

correlated with total tree density (r = 0.73). A regression tree showed that the greatest percentage of small trees

occurred on sites with high precipitation and in areas other than the northeastern part of the study area (Fig.

184

Journal of the Botanical Research Institute of Texas 8(1)

T—

.92

Cl)

Acacia greggii/

wash

Mixed/

variable

Prosopis glandulosa/ Prosopis glandulosa/ Prosopis pubescens/

protected gypsum spring

Community

Fig. 4. (a) Species richness by lifeform for Acada and Prosopis woodlands of the eastern Mojave Desert, USA. Error bars are 1 SD for total mean richness,

and means without shared letters differ at P <0.05. (b) Relationship between native and exotic species richness, with a slope and y intercept for a

regression line shown for descriptive purposes.

7C). Prosopis pubescens/ spring communities, which were dominated by large trees, occurred in the study area’s

northeastern corner.

We classified a total of 73 species into 14 species groups comprised of 2-8 species (Appendix 1). Ampli¬

tude and fidelity to community types varied among species groups, with some groups most frequent in one or

a few communities (e.g., Allenrolfea occidentalis group most frequent in Prosopis pubescens/ spring) and others

more widespread yet still often sparse or absent from one or more communities. Examples of species group

distributions include: the Acacia greggii group of Hymenoclea salsola and annual species like Eriogonum palme-

rianum that occupy washes; Larrea tridentata group of dry-site species inhabiting xeric areas within or on the

edges of the sampled riparian patches; Enceliafarinosa group of Acacia greggii/ wash but also of other communi¬

ties, excepting Prosopis pubescens/ spring; Pluchea sericea group with a distribution difficult to characterize;

Suaeda moquinii principally of gypsum soil, including Prosopis glandulosa/ gypsum and Prosopis pubescens/

spring, although some species of the group also frequented other communities; and the Allenrolfea occidentalis

group primarily of Prosopis pubescens/ spring or Prosopis glandulosa/ gypsum, indicating that this group inhab¬

its extreme soil properties. Exotic species did not group together (only one species group contained more than

one exotic species) and instead occurred in a range of species groups with native species.

Abella and Chittick, Acacia and Prosopis woodlands of the Mojave Desert

185

Fig. 5. Ordination of (a) vegetation and (b) soil composition with plots displayed according to community type for Acacia and Prosopis woodlands of

the eastern Mojave Desert, USA. Vectors are scaled in proportion to their correlation with ordination axes. Vectors are shown with r 2 >0.20 for (a) and

>0.50 for (b). Abbreviations for vectors: ACAGRE = Acaciagreggii, ALLOCC= Allenrolfea occidentalis, HYMSAL = Hymenocleasalsola, PROGLA = Prosopis

glandulosa, PROPUB = Prosopis pubescens, and SUAMOQ = Suaedamoquinii.

186

Journal of the Botanical Research Institute of Texas 8(1)

Table 2. Soil properties (0-5 cm) of Acacia and Prosopis woodland community types of Lake Mead National Recreation Area, Mojave Desert, USA.

AG/wash a

M/variable

PG/protected

PG/gypsum

PP/spring

Physical properties

Mean ± SD

Gravel (% weight)

46±14

37±17

43±17

21 ±21

3±3

Gravel (% volume)

28±10

19±17

24±11

13±15

1±1

Bulk density (g/cm 3 )

0.78±0.23

0.83±0.2

0.70±0.13

0.93±0.1

0.67±0.06

Sand (%)

86±13

72±7

72±16

75± 18

53±22

Silt (%)

8±9

22±7

20±13

14±7

40±19

Clay (%)

6±4

7±2

9±4

11±11

7±4

Chemistry

EC b (mS/cm)

2.5±11.0

2±2.3

3.3±8

9.5±10.9

16.9±14.2

8.0±0.3

8±0.5

7.7±0.3

8±0.6

8.4±0.3

Chemical composition

CaC0 3 (%)

9.1 ± 15.6

13.6±5.1

7.4±7.0

26.1 ±12.0

2.7±0.6

Organic C (%)

0.6±0.8

0.7±0.6

0.7±0.5

1.1 ±0.6

Inorganic C (%)

1.1±1.9

1.6±0.6

0.9±0.8

3.1 ±1.4

0.3±0.1

Total C (%)

1.6±2.1

2.3±1.0

1.5±0.8

3.8±1.1

1.4±0.5

NO s (mg/kg)

0.7±0.6

4.9±8.5

6±7.6

18.6±36.3

5.1 ±7.6

Total N (%)

0.04±0.04

0.08±0.06

0.06±0.04

0.07±0.07

0.10±0.05

Olsen P (pg/g)

6±5

15±15

9±7

19±18

13±11

S0 4 (mg/kg)

400±1924

253±417

533±1593

627±611

2243±1935

Total S (%)

0.20±0.81

0.12±0.13

0.15±0.36

0.07±0.04

3.39±2.92

K (mg/kg)

235±834

181 ±178

142±152

376±259

470±336

Ca (mg/kg)

523±1142

1008±1416

543±864

2498±1506

2870±2218

Mg (mg/kg)

81 ±196

173±210

105±195

314±258

497±384

Na (mg/kg)

449±1586

466±532

302±525

752±761

1230±846

Fe (pg/kg)

114±144

55±45

119±103

57±31

100±41

Mn (pg/kg)

0.03±0.04

0.05±0.02

0.03±0.03

0.12±0.09

0.09±0.02

Cu(pg/kg)

33±25

53±20

82±60

59±23

69±25

Zn (pg/kg)

18±44

15±18

46±77

24±18

69±53

Mo (pg/kg)

3.8±17.8

2.5±3.2

3±7.2

3±3

13.7±12.7

B(pg/kg)

657±2537

629±803

1094±2964

2953±4712

19469±24806

Cl (mg/kg)

251±1314

138±250

148±402

856±1256

1264±1130

Ni(pg/kg)

11 ±8

17±7

10±7

16± 12

12±10

Co (pg/kg)

0.7±1.0

1.3±1.3

1.5±1.6

1.8±2

1.1 ±0.6

Cd (pg/kg)

0.09±0.33

0.06±0.04

0.09±0.15

0.11 ±0.06

0.25±0.24

Pb (pg/kg)

0.4±0.4

0.2±0.1

0.6±0.4

0.2±0.1

0.4±0.2

a From left to right, full names of community types are: Acacia greggii/wash, Mixed/variable, Prosopis glandulosa/protected, Prosopis

glandulosa/gypsum, and Prosopispubescens/ spring

b Electrical conductivity

Nine of the 14 species groups had correlation coefficients > 10.401 with at least one principal component or

environmental variable. Among the strongest correlations included the Acacia greggii group with precipitation

(r = 0.55), sand (r = 0.60), and total N (r = -0.46); Pluchea sericea group with Cu (r = 0.72); and the Allenrolfea

occidentalis group with total S (r = 0.64).

DISCUSSION

Community Classification and Gradients

Plant species composition was distinct for each community type, and distributional overlap among the three

tree species was low at our 0.1-ha plot scale. With an internal similarity of 44%, even the mixed/variable com¬

munity was not merely a collection of sites unable to be classified, but rather was a community of unique spe¬

cies composition. Uniqueness of community composition was illustrated by internal similarities of 42-66%

and segregation of community groupings in ordination. Only 1 (2%) of 50 plots contained all three focal tree

species and 10 (20%) contained two species (all co-occurrences of Acacia greggii and Prosopis glandulosa), sug¬

gesting relatively strong partitioning of species distributions. Based on soil texture and landforms, we surmise

Abella and Chittick, Acacia and Prosopis woodlands of the Mojave Desert

187

Fig. 6. Distribution of community types and tree species as a function of environmental variables for Acacia and Prosopis woodlands of the eastern Mojave

Desert, USA. (a) Classification tree for distribution of community types, abbreviated as: AW = Acacia greggii/msh, MV = Mixed/variable, PP = Prosopis

glandulosa/ protected, PG = Prosopis glandulosa/ gypsum, and PS = Prosopispubescens/ spring. The actual proportion of plots is shown at the top of the

tree. For each division, the estimated probability is shown on the left and the actual proportion on the right, with bold font highlighting the community

with the highest estimated probability, (b) Regression tree with estimated mean (± SD) Acacia canopy cover at terminal nodes, (c) Classification tree

of Prosopis pubescens presence/absence, with estimated probability of presence shown on the left and actual proportion of presence on the right for

each division, (d) Regression tree with estimated mean (± SD) Prosopisgianduiosa canopy cover at terminal nodes.

that A. greggii generally occupied the driest sites (coarsest soil and dry washes), P. glandulosa intermediate

(topographically protected and moister washes), and Prosopis pubescens the wettest (outflow of springs, often

with visible surface water). Comparative ecohydrological research (Smith et al. 1998) may be useful for evalu¬

ating if water balances were consistent with this perceived distribution.

Ordinations suggested that vegetation grouped more strongly than did the suite of 31 measured soil vari¬

ables and that plant communities inhabited a range of environmental properties. However, there were some

environmental correlates for the communities and tree species. Coarse-textured soil was associated with Aca¬

cia greggii, reflecting this species’ affinity for dry washes. These washes have coarse soil because periodic

floods carry away fine soil particles, while depositing coarse material from higher elevations (Schwinning et al.

2011). UTM, expressing location, was also an important variable, suggesting that certain communities had

affinity to particular sections of the study area. Prosopis pubescens/ spring communities, for example, were lo¬

cated in the northeastern part of the study area where hydrological conditions associated with outflow of

springs were apparently favorable for development of this community. These areas were also affiliated with

gypsum soil, likely accounting for relationships of Prosopis pubescens/ spring communities with variables such

as total S. Gypsum, comprised of CaS0 4 -2H 2 0, produces soils high in S and salts (Meyer 1986) and can have

extreme properties compared to the rest of the landscape, as we observed in our study. Prosopis pubescens oc¬

cupies non-gypsum soil in other parts of its range (Busch 1995), and it is unclear if P. pubescens simply is toler-

188

Journal of the Botanical Research Institute of Texas 8(1)

a) Exotic species richness

2.5±1.4, n = 50

Precipitation < 16 cm/yr

Total S > 0.15%

Precipitation > 16 cm/yr

r 2 = 0.18

Total S < 0.15%

Silt < 21%

r 2 = 0.30 I

0.4±0.5, n = 5 2.1±0.9, n = 14 Clay < 8%

Silt >21%

r 2 = 0.42 J

Clay >8% 4.4±2.1, n = 5

Cross-validated r 2 = 0.40

J r 2 = 0.53 |

3.1±9, n = 19 1.6±0.8, n = 7

b) Live trees/ha mistletoe infected

43±46, n = 48

CaC0 3 > 2.6%

Elevation < 388 m

CaC0 3 < 2.6%

r 2 = 0.24

Elevation > 388 m

r 2 = 0.35

5±12, n =12 Tree cover < 13%

80±58, n = 13

Tree cover :> 13%

I r 2 = 0.45 J

22±25, n = 13 66±25, n = 10

Cross-validated r 2 = 0.34

c) Percentage of small trees

24±20, n = 48

Precipitation < 19 cm/yr

Species richness < 30

Precipitation > 19 cm/yr

r 2 = 0.25

Species richness > 30 UTMx .> 747216 m

UTMx < 747216 m

j r 2 = 0.60

11±14, n=23

Cross-validated r 2 = 0.52

r 2 = 0.48

30±9, n=9 25±15, n = 10

60±12, n=6

Fig. 7. Regression trees for ecological condition variables of Acacia and Prosopis woodlands of the eastern Mojave Desert, USA. Cumulative r 2 is shown

for each division and estimated means (± SD) of response variables at terminal nodes. Trees could not be counted due to inaccessibility at 2 plots so

sample size is 48 for (b) and (c).

ant of the extreme properties of gypsum in our study or if environmental conditions favorable for its occur¬

rence were related to gypsum soil properties.

Although potential importance of plant correlations with soil variables should not be dismissed, data on

ground-water depth and chemistry might help to account for additional variance in community and tree species

distribution. Few data on groundwater exist for the study area (Gary Karst, Hydrologist, Lake Mead National

Recreation Area, pers. comm.). An unpublished report using six wells found that depth to groundwater was <1

Abella and Chittick, Acacia and Prosopis woodlands of the Mojave Desert

189

Diameter class (cm)

Fig. 8. Diameter at root collar distribution by tree species for Acacia and Prosopis community types of the eastern Mojave Desert, USA. Error bars are 1

SD. Note the difference in y-axis scales among the graphs.

190

Journal of the Botanical Research Institute of Texas 8(1)

m in summer 1992 to 1994 in the drainage of Sacatone Wash containing Prosopis glandulosa and Prosopis pubes¬

cens in the southern part of the study area (Inglis et al. 1996). In an area of the Bluepoint Spring outflow support¬

ing P. pubescens, we also directly observed that depth to groundwater was approximately <2 m based on “sink¬

holes” where flowing groundwater was visible. Groundwater might be predicted to be deeper in Acacia greggii

communities because they occupied dry washes characterized by more ephemeral, rather than perennial, wa¬

ter fluxes (Schwinning et al. 2011). Relationships of groundwater depth with P. glandulosa are unclear because

some locales of this species were in topographically protected sites where shading might reduce evaporation

and surface water might collect (Schwinning et al. 2011). Observed distributional differences in these commu¬

nities afford opportunities for ecophysiological and hydrological research to improve understanding of habitat

partitioning (Busch & Smith 1995). This is especially important for conservation given concerns about poten¬

tial for groundwater pumping to lower regional water tables coupled with climate change (Deacon et al. 2007).

Whereas Judd et al. (1971) concluded that established P. glandulosa could survive pumping-related lowering of

water to 13 m below the surface before the trees died, Patten et al. (2008) suggest that declines in water depths

of even a meter for near-surface groundwater can dramatically impact tree recruitment and associated species.

In addition to depth, groundwater chemistry might affect plant distribution by influencing composition

of water that roots access (Springer et al. 2008). Near springs in Death Valley National Park in the western

Mojave Desert, Hunt (1966) concluded that groundwater chemistry rather than soil chemistry more strongly

correlated with distribution of Prosopis glandulosa.

Species Groups

Little research has examined ecological species groups in arid environments, but our results are consistent

with some general principles of species groups in temperate regions. For instance, our finding that a species

group was not restricted to one community type concurs with the common observation that most groups in¬

habit multiple communities but are quantitatively most abundant in only a few communities (Kashian et al.

2003). Species groups in temperate regions were more strongly correlated with multivariate environmental

gradients than single-factor gradients (Goebel et al. 2001). We also found few strong correlations of groups

with individual environmental variables, and occurrences instead were likely related to multivariate gradients

in groundwater depth, chemistry, soil moisture under textural and topographic control, flooding frequency,

and soil chemistry such as the presence of gypsum (Hunt 1966; Patten et al. 2008; Springer et al. 2008).

Obligate wetland species (e.g .Juncus and Scirpus spp.) were not well represented in the species groups or

in the flora as a whole at these sites. Instead, the flora was dominated by species characterized as transitional

between wetlands and uplands (Patten et al. 2008). Additionally, species of the Acacia greggii/ wash community

in particular are associated with disturbance. For example, abundance of Hymenoclea salsola and Sphaeralcea

ambigua often increases following fire and other anthropogenic disturbances (Abella 2010), consistent with

their occurrence in natural washes that are periodically disturbed by flooding.

Ecological Condition and Conservation Implications

Based on features of favored habitat described by Crampton and Sedinger (2011) for the conservation-priority

bird Phainopepla nitens, many sites in the study area demonstrate favorable characteristics. Almost all nests of

P. nitens are in mistletoe-infected trees (Crampton & Sedinger 2011), and we found that 66% of our 0.1-ha plots

contained >1 infected tree. Phainopepla nitens nest suitability also is correlated with the number of infected

trees, which we found averaged 62 ± 44 trees/ha on plots where mistletoe was present. Surveying abundance of

P. nitens and other priority wildlife species across this network of woodland sites might help improve under¬

standing of landscape-scale distributional relationships of wildlife species with plant communities.

Communities were dominated by native species, which comprised 94% of the total 201 taxa detected. In

regression analysis, native species richness accounted for 22% of the variability in exotic richness, more than

any other variable and consistent with the often-observed positive relationship between native and exotic rich¬

ness (Tabacchi & Planty-Tabacchi 2005). The least species-rich community ( Prosopis pubescens/ spring) was

least invaded, whereas the most species-rich sites generally were most invaded.

Abella and Chittick, Acacia and Prosopis woodlands of the Mojave Desert

191

Of the 13 total exotic species detected, 3 are of greatest current concern to resource managers. Bromus

rubens is a major concern because it increases fine fuel loads to facilitate spread of fire, which is a recent, novel

disturbance to which native Mojave Desert flora is not considered well adapted (Abella 2010). At our study

sites, however, B. rubens cover was low, exceeding 10% (and never more than 20% cover) at only 6 of its 43 oc¬

cupied plots even including cover of dead stalks. Our study landscape as a whole is at a lower elevation than the

middle elevations where B. rubens abundance is greatest in the Mojave Desert, suggesting that B. rubens even

in riparian areas with supplemental moisture at low elevations does not attain the dominance it does in up¬

lands at higher elevations (Abella et al. 2012b). Although we included dead plants as a measure of cumulative

recent cover, B. rubens cover can vary dramatically between multi-year wet and dry periods (Steers et al. 2011)

such that periodic monitoring of these riparian areas is warranted. The second species of greatest concern is

Brassica tournefordi, which also can provide fuel and compete with native plants and predominates at low ele¬

vations (Barrows et al. 2009). We detected this species at only 3 sites, suggesting it is not presently a major

component of these woodlands. The third species, Tamarix ramosissima, can outcompete native species and

alter soil properties through production of salt-rich litter and exudates (Smith et al. 1998). Although we de¬

tected T. ramosissima at 22% of sites, this species and the native trees typically comprised different patches, as

was also noted by Bagstad et al. (2006) along the San Pedro River in Arizona. Future management of this spe¬

cies might be guided by effectiveness of the biocontrol Diorhabda carinulata (tamarisk leaf beetle), presently

moving south and reaching the northern boundary of the study area (Bateman et al. 2010). Although further

monitoring is warranted, these riparian communities have lower exotic plant abundance in comparison to

many other areas of the Mojave Desert including those that have burned by wildfire (Dudley 2009; Steers et al.

2011 ).

Other observations also suggested that threats to these woodlands were less prevalent in our study than

observed in some other areas (Crampton et al. 2006). Qualitative observations indicated no evidence of fire or

woodcutting at most sites, with only minimal (e.g., some branches) and localized cutting noted on plots near

anthropogenic camping locations. Some observation of probable evidence (by recording browsed plants) of

unauthorized livestock grazing was noted in the northeastern part of the study area, but effects to the wood¬

lands are unclear. Lack of tree recruitment is considered a major problem in other areas (Crampton et al. 2006),

but we observed tree densities well distributed among size classes, except in the Prosopis pubescens/ spring

community, which was dominated by large trees. Further investigation of recruitment potential in this com¬

munity is warranted. If desired, it is feasible to actively facilitate establishment of P. pubescens and the other tree

species through planting nursery grown seedlings (Abella & Newton 2009).

In summary, the data suggest that these woodlands were: readily classified into community types that

might exhibit different conservation needs; correlated with some measured environmental variables, but fur¬

ther investigation into groundwater depth and chemistry could be informative; dominated by native species;

inhabited by suites of annual and perennial plants classifiable into species groups displaying unique distribu¬

tions; typified by well-distributed tree density across size classes at most sites; and characterized by low evi¬

dence of threats such as fire noted in other regions.

APPENDIX 1

Ecological species groups for Acocio and Prosopis woodlands of Lake Mead National Recreation Area, Mojave Desert, USA.

Community type

Species group 3 AGW b MV PGP PGG PPS

Acacia greggii

Frequency (%) b

Acacia greggii —catclaw acacia

Bromus arizonicus —Arizona brome

Camissonia boothii —Booth's evening primrose

Eriogonumpalmerianum —Palmer's buckwheat

Erodium cicutarium —redstem stork's bill*

100 100 70

24 25 10

21 0 0

31 0 0

192

Journal of the Botanical Research Institute of Texas 8(1)

Continued

APPENDIX 1

Community type

Species group 3 AGW b MV PGP PGG PPS

Acocio greggii

Hymenodea salsola — cheesebush

Pectocarya setosa —moth combseed

Salvia columbariae —chia

Eriogonum fasciculatum

Eriogonumfasdculatum — eastern Mojave buckwheat

Encelia virginensis-Vi rgin River brittlebush

Phacelia vallis-mortae —Death Valley phacelia

Porophyllum gradle — slender poreleaf

Sphaeralcea ambigua — desert globemallow

Xylorhiza tortifolia — Mojave woodyaster

Ephedra viridis

Ephedra viridis — mormon tea

Amsinckia tessellata —bristly fiddleneck

Draba cuneifolia —wedgeleaf draba

Nemacladus glanduliferus —glandular threadplant

Viguieraparishii — Parish's goldeneye

Tamarix ramosissima

Tamarixramosissima — saltcedar*

Funastrum cynanchoides — fringed twinevine

Nicotiana obtusifoiia — desert tobacco

Stillingia linearifolia — q ueen's-root

Ephedra torreyana

Ephedra torreyana — Torrey's jointfir

Bromus rubens —red brome*

Chorizanthe brevicornu —brittle spineflower

Cryptantha pterocarya —wingnut cryptantha

Cusaitadentiailata — desert dodder

Guillenia lasiophylla —California mustard

Lepidium lasiocarpum —shaggyfruit pepperweed

Schismus spp.—Mediterranean grass*

Ephedra nevadensis

Ephedranevadensis — Nevada jointfir

Acamptopappus sphaerocephalus — rayless goldenhead

Ambrosia eriocentra — woolly fruit bur ragweed

Amsinckia menziesii —Menzies'fiddleneck

Bromus tectorum —cheatgrass*

Eriogonum thomasii —Thomas' buckwheat

Pleuraphis rigida — big galleta

Ericameria paniculata

Ericameriapaniculata — Mojave rabbitbrush

Cryptantha circumscissa —cushion cryptantha

Echinocereus engelmannii — hedgehog cactus

Echinocereuspolyacanthus — Mojave mound cactus

Eriogonum deflexum —flatcrown buckwheat

Yuccaschidigera — Mojave yucca

Larrea tridentata

Larrea tridentata — creosote bush

Cylindropuntia acanthocarpa — buckhorn cholla

Eriogonum inflatum — desert trumpet

Ferocactuscylindraceus — California barrel cactus

Krameria erecta — littleleaf ratany

Opuntia basilaris — beavertail pricklypear

Encelia farinosa

Encelia farinosa — brittlebush

Bebbia juncea — sweetbush

Frequency (%) b

100

Abella and Chittick, Acacia and Prosopis woodlands of the Mojave Desert

193

Continued

APPENDIX 1

Community type

Species group 3

AGW b

PGP

PGG

PPS

En celia farinosa

Chamaesyce polycarpa — smallseed sandmat

Frequency (%) b

20 25

Krameria grayi — white ratany

Stephanomeria paudflora — brown plume wi relettuce

Ambrosia dumosa

Ambrosia dumosa — burrobush

100

Polypogon monspeliensis — annual rabbitsfoot grass

Prosopis glandulosa

Prosopis glandulosa — honey mesq u ite

100

Baccharis emoryi — Emory's baccharis

Pluchea sericea

Pluchea sericea — ar rowweed

Phragmitesaustralis — common reed

Pleurocoronispluriseta — bush arrowleaf

Suaeda moquinii

Suaeda moquinii — Mojave seablite

100

Atriplexconfertifolia — shadscale saltbush

Chorizanthe rigida — devil's spineflower

Descurainia pinnata — western tansymustard

Isocoma acradenia — alkali goldenbush

100

Plantago ovata — desert Indianwheat

Stylocline intertexta — Morefield's neststraw

Vulpia octoflora — sixweeks fescue

Allenrolfea occidentalis

Allenrolfea occidentalis — iod inebush

100

Distichlis spicata — sa 1 tg r ass

Prosopis pubescens — screwbean mesq u ite

100

a Bold = perennial, not bold = annual or biennial, and * = exotic

b Abbreviations and numbers of plots for community types: AGW = Acocio greggii/wash (n = 29), MV = Mixed/variable (n = 4), PGP =

Prosopis glandulosa/protected (n = 10), PGG = Prosopis glandulosa/gypsum (n = 4), and PPS = Prosopis pubescens/ spring (n = 3). Bold

values signify where a species group is overall most frequent

ACKNOWLEDGMENTS

This study was funded through a cooperative agreement organized by Alice Newton between the National Park

Service, Lake Mead National Recreation Area, and the University of Nevada Las Vegas (UNLV). We thankjos-

lyn Curtis and Sylvia Tran (UNLV) for help with fieldwork; Alice Newton and Dara Scherpenisse (LMNRA) for

help with study design and insight on conservation implications; the UNLV Environmental Soil Analysis

Laboratory (in particular Yuanxin Teng and Brenda Buck) for performing soil analyses; Sharon Altman

(UNLV) for creating figures; and J. Andrew Alexander and Walter Fertig for helpful comments on the manu¬

script.

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Hunt, C.B. 1966. Plant ecology of Death Valley. U.S. Geological Survey Professional Paper 509.

Inglis, R., C. Deuser, & J. Wagner. 1996. The effects of tamarisk removal on diurnal ground water fluctuations. Tech. Rep.

NPS/NRWRD/NRTR-96/93. U.S. National Park Service, Washington, D.C., USA.

Judd, B.L., J.M. Laughlin, H.R. Guenther, & R. Handegarde. 1971. The lethal decline of mesquite on the Casa Grande National

Monument. Great Basin Nat. 31:153-159.

Kashian, D.M., B.V. Barnes, &W.S. Walker. 2003. Ecological species groups of landform-level ecosystems dominated by jack

pine in northern Lower Michigan, USA. PL Ecol. 166:75-91.

Keeler-Wolf, T. 2007. Mojave Desert scrub vegetation. In: M.G. Barbour, T. Keeler-Wolf, and A.A. Schoenherr, eds. Terres¬

trial vegetation of California. University of California Press, Berkeley, USA. Pp. 609-656.

Abella and Chittick, Acacia and Prosopis woodlands of the Mojave Desert

195

Lato, L.J. 2006. Soil survey of Clark County area, Nevada. U.S. Department of Agriculture, Natural Resources Conserva¬

tion Service. U.S. Government Printing Office, Washington, D.C., USA.

McCune, B. & M.J. Mefford. 1999. PC-ORD: Multivariate analysis of ecological data. Vers. 4. User's guide. MjM Software

Design, Gleneden Beach, Oregon, USA.

McCune, B., R. Rosentreter, J.M. Ponzetti, & D.C. Shaw. 2000. Epiphyte habitats in and old conifer forest in western Washing¬

ton, USA. Bryologist 103:417-427.

Meyer, S.E. 1986. The ecology of gypsophile endemism in the eastern Mojave Desert. Ecology 67:1303-1313.

Miller, D., S.R. Archer, S.F. Zitzer, & M.T. Longnecker. 2001. Annual rainfall, topedaphic heterogeneity and growth of an arid

land tree (Prosopis glandulosa). J. Arid Environm. 48:23-33.

Patten, D.T., L. Rouse, & J.C. Stromberg. 2008. Isolated spring wetlands in the Great Basin and Mojave Deserts, USA: Poten¬

tial response of vegetation to groundwater withdrawal. Environm. Managem. 41:398-413.

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conserving springs in the western United States. Tech. Ref. 1737-17. Bureau of Land Management, National Science

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SAS Institute. 2010. Using JMP 9. SAS Institute, Cary, North Carolina, USA.

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distributions of Larrea tridentata and Ambrosia dumosa in the Mojave Desert, CA, USA: Patterns, mechanisms and

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Wetlands 18:687-696.

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classification system: Integrating geomorphic, hydrogeochemical, and ecological criteria. In: L.E. Stevens and VJ.

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mapping database. U.S. Geological Survey, Western Regional Science Center and Southwest Biological Science

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249.

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Journal of the Botanical Research Institute of Texas 8(1)

BOOK NOTICE

Johnnie L. Gentry, George P. Johnson, Brent T. Baker, C. Theo Witsell, and Jennifer D. Ogle, eds. 2014. Atlas of

the Vascular Plants of Arkansas, (no ISBN given, pbk). University of Arkansas, Department of Printing

Services, University Services Building, 2801 S. University Ave., Little Rock, Arkansas 72204, U.S.A. (Or¬

ders: University of Arkansas Herbarium, Attn: Jennifer Ogle, Biomass Research Center 141, University

of Arkansas, Fayetteville, Arkansas 72701, U.S.A., www.uark.edu/~arkflora, herb@uark.edu, 1-479-575-

4372). $43.90 ($7.15 shipping, Arkansas Native Plant Society members receive a 10% discount), 709 pp.,

2892 maps, some color introduction maps, size unknown.

From the publisher: The Atlas includes 2,892 taxa representing 187 families, 936 genera and 2,715 species. Each

species is presented with a county-level distribution map along with family and common names; numerical

codes representing special status for certain taxa (nonnative, invasive, endemic, or species of conservation

concern) are also included. This publication also includes the following chapters, featuring full-color maps and

plates.

This 709-page paperback publication features distribution maps for each of the 2,892 native and natural¬

ized vascular plants in Arkansas, representing 2,715 species, 936 genera, and 187 families, introductory chap¬

ters, featuring full-color maps and plates, provide information on the following subjects:

General introduction

History of botanical exploration in Arkansas

An overview of the geology of Arkansas

Effects of physical factors on the distribution of native flora and vegetation in the natural divisions of

Arkansas

Additional sections include:

Floristic summary

Arkansas endemic taxa

Additional taxa reported for Arkansas

Arkansas vascular plants of conservation concern

J.Bot. Res. lnst.Texas8(1): 196.2014

NOTEWORTHY VASCULAR PLANT COLLECTIONS

LROM THE RED RIVER OF ARKANSAS AND LOUISIANA, U.S.A.

Christopher Reid

Louisiana Department of Wildlife & Fisheries

2000 Quail Drive

Baton Rouge, Louisiana 70898, U.S.A.

creid@wlf.la.gov

M. Jerome Lewis

Department of Biological Sciences

Louisiana State University in Shreveport

One University Place

Shreveport, Louisiana 71115, U.S.A.

Iewism06@lsus.edu

ABSTRACT

A plant collecting excursion by boat on a ca. 50 km stretch of the Red River straddling the Arkansas-Touisiana state line yielded several in¬

teresting botanical discoveries. The second record of Loeflingia squarrosa from Arkansas was documented. Collections of Dalea Janata and

Heliotropium convolvulaceum were made from both states. These collections extend the ranges of these taxa several hundred river-km down¬

stream on the Red River. Our collections of D. Janata and H. convolvulaceum in Touisiana represent the first records of these species for that

state.

RESUMEN

Una excursion de recogida de plantas en bote en un trecho de ca. 50 km en el Red River a caballo en la frontera estatal Arkansas-Luisiana

produjo varios descubrimientos botanicos interesantes. Se documento la segunda cita de Loeflingia squarrosa de Arkansas. Se realizaron

colecciones de Dalea Janata y Heliotropium convolvulaceum en ambos estados. Estas colecciones extendieron los rangos de estos taxa varios

cientos de kilometros rio abajo en el Red River. Nuestras colecciones de D. Janata y H. convolvulaceum en Luisiana representan las primeras

citas de estas especies para el estado.

INTRODUCTION

We carried out a plant collecting excursion by boat on the Red River in southwestern Arkansas and northwest¬

ern Louisiana on 15 and 16 August 2012. The objective was to explore the habitats along the river, particularly

sand bars on the river in its present course and relict sand bars abandoned when the river shifted course. In our

region, plant collections from streams and rivers seem to frequently be from easily accessible points such as

bridges and boat launches (pers. obs.). It was hoped that our approach using a boat would yield some interest¬

ing records that may otherwise not be documented.

METHODS

We launched at the Arkansas Highway 160 bridge (latitude and longitude in decimal degrees: 33.089622,

-93.858549) and proceeded upstream for ca. 20 km, then turned around and explored downstream into Loui¬

siana, terminating at the Louisiana Highway 2 bridge (latitude and longitude in decimal degrees: 32.892751,

-93.820295). A stretch of approximately 50 river-km was covered during our held work (Fig. 1). A 14-foot hat

bottom aluminum boat with a shallow water marine drive “go-devil” motor was used for our study. Capable of

navigating water 45 cm deep and handling the occasional underwater sandbar or mudflat, this set-up seemed

most useful. Water levels were slightly below average for that time of the year.

RESULTS AND DISCUSSION

Our held work on the Red River yielded several noteworthy plant records. The diminutive Loeflingia squarrosa

Nutt, was collected from an abandoned point bar in Arkansas. This species is very rare in Arkansas and Loui¬

siana and is ranked as SI in both states (NatureServe 2013). Previous records of L. squarrosa from Arkansas

and Louisiana are from xeric sandhill woodlands associated with Tertiary formations (Singhurst & Holmes

1999; Reid & Faulkner 2006). Our record, only the second from Arkansas, was from a dry sandy grassland

J. Bot. Res. Inst. Texas 8(1): 197 - 201.2014

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Journal of the Botanical Research Institute of Texas 8(1)

Fig. 1 . Map showing Red River in northwestern Louisiana and southwestern Arkansas, U.S.A. The yellow circles delimit the stretch of the River that was

botanically explored on 15-16 August 2012.

developed on recently-deposited sediment. This grassland was dominated by Sporobolus cryptandrus (Torr.) A.

Gray and Eragrostis secundiflora J. Presl on the highest driest areas. Thickets of Prunus angustifolia Marshall

were conspicuous. Other characteristic species included Heterotheca subaxillaris (Lam.) Britton & Rusby, Cro-

ptilon divaricatum (Nutt.) Raf., and Diodia teres Walter. Lower slopes of this abandoned bar had heavy cover of

the exotic Cynodon dactylon (L.) Pers.

Collections of Dalea lanata Spreng. and Heliotropium convolvulaceum (Nutt.) A. Gray were made from Ar¬

kansas and Louisiana. Collections of these taxa in both states are significant. Dalea lanata is state-rare, ranked

as S2, in Arkansas (NatureServe 2013) and was previously known only from sand bars on the Arkansas River

in the central part of the state (Smith 1988; Gentry et al. 2013). In Arkansas, we collected Dalea lanata from the

same site as Loeflingia squarrosa. It was observed at one additional site on a less stable and sparsely-vegetated

sand bar farther upstream. In Louisiana, one population of D. lanata was documented from a dry sand dune

held which is apparently shaped by seasonal hooding. Since this species had not previously been reported for

Louisiana by MacRoberts (1984,1989), Thomas and Allen (1998), USDA, NRCS (2013), or NatureServe (2013),

our Louisiana record of D. lanata is regarded as the hrst for the state. Dalea lanata is a species of the southwest¬

ern U.S. that occurs on sand dunes and in sandy river valleys, with known occurrences on the Red River in

Oklahoma and Texas (Correll & Johnston 1970; McGregor 1986; Hoagland et al. 2004). Our records of D. la-

Reid and Lewis, Plant exploration along the Red River of Arkansas and Louisiana

199

Fig. 2. Top: Sandbar on Red River in Miller County, Arkansas, at latitude/longitude 33.149814, -93.779734 (decimal degrees). Heliotropium convolvu-

laceum was collected at this site and a small population of Dalea lanata was observed. Image was taken facing downstream (west-northwest) where

river channel is visible in background. Bottom: Open dry sand dune field in Bossier Parish, Louisiana, at latitude/longitude 32.956424, -93.831097

(decimal degrees). This dune-like area was associated with a portion of a point bar apparently where seasonal flood waters pass to the east of the

most stable part of the bar which is wooded. Image was taken oriented roughly southwest and facing the tree line on the most stable part of the bar.

Heliotropium convolvulaceum (a few plants flowering in foreground) and Dalea lanata were present on this site.

nata are approximately 500 river-km downstream from nearest reported populations in Bryan Co., Oklahoma,

and Grayson Co., Texas (Turner et al. 2003; Hoagland et al. 2004).

Heliotropium convolvulaceum has a similar distribution as Dalea lanata and is similar ecologically, prefer¬

ring dry loose sands (Correll & Johnston 1970; Kaul 1986). As with D. lanata, it is rare in Arkansas with a

200

Journal of the Botanical Research Institute of Texas 8(1)

ranking of S2 (NatureServe 2013) and was previously known only from sand bars on the Arkansas River

(Smith 1988; Gentry et al. 2013). It is herein reported in Louisiana for the first time as other floristic references

do not include it in the state (MacRoberts 1984,1989; Thomas & Allen 1998; NatureServe 2013; USDA, NRCS

2013). Our collections of H. convolvulaceum are approximately 300 river-km downstream from previously

documented stations along the Red River in McCurtain Co., Oklahoma, and Lamar Co., Texas (Turner et al.

2003; Hoagland et al. 2004).

Dalea lanata and Heliotropium convolulaceum were sympatric on sparsely vegetated sand bars and dunes

(Fig. 2). Close associates of these species included Cycloloma atriplicifolia (Spreng.) J.M. Coult., Cyperus escul-

entus L., Heterotheca subaxillaris, and Sporobolus cryptandrus.

VOUCHER SPECIMENS

Loeflingia squarrosa (Caryophyllaceae)

Voucher Specimen: ARKANSAS. Lafayette Co.: Slay Bend on Red River, ca. 8 river-km upstream from AR 160 bridge, ca. 8.8 air-km NE of

Doddridge, 33.1424840, -93.8342270, abandoned point bar supporting sandy grassland, with Sporobolus cryptandrus, Eragrostis secundiflo-

ra, Heterotheca subaxillaris, Diodia teres, Croptilon divaricatum, and Prunus angustifolia thickets, common and scattered, 15 Aug 2012, Reid

and Lewis 8278 (ANHC, LSU).

Dalea lanata (Fabaceae)

Voucher Specimens: ARKANSAS. Lafayette Co.: Slay Bend on Red River, ca. 8 river-km upstream from AR 160 bridge, ca. 8.8 air-km NE of

Doddridge, 33.1424840, -93.8342270, abandoned point bar supporting sandy grassland, with Sporobolus cryptandrus, Eragrostis secundiflo-

ra, Heterotheca subaxillaris, Diodia teres, Croptilon divaricatum, and Prunus angustifolia thickets, abundant in small area, 15 Aug 2012, Reid

and Lewis 8275 (ANHC, LSU). Miller Co.: sandbar on Red River S of Haley Lake, ca. 14 river-km upstream from AR 160 bridge, ca. 13 air-km

WNW of Bradley, 33.1494510, -93.7790070, only a few plants seen in small area, no voucher specimen collected. LOUISIANA. Bossier Par¬

ish: Red River ca. 10 river-km upstream from LA 2 bridge, ca. 6 air-km E of Mira and ca. 8.7 air-km NE of Hosston, 32.9532900, -93.8315800,

open sparsely vegetated sandy dune field on east side of river bed, dry loose sand with Heterotheca subaxillaris, one large plant ca. 1 m radius

and a few smaller satellite plants, 16 Aug 2012, Reid and Lewis 8299 (LSU).

Heliotropium convolvulaceum (Boraginaceae)

Voucher Specimens: ARKANSAS. Miller Co.: Sandbar on Red River S of Haley Lake, ca. 14 river-km upstream from AR 160 bridge, ca. 13

air-km WNW of Bradley, 33.1494510, -93.7790070, common on high dry sand bar with Sporobolus cryptandrus, Strophostyles helvola (L.)

Elliott, Heterotheca subaxillaris, and Cycloloma atriplicifolia, 15 Aug 2012, Reid and Lewis 8293 (ANHC, LSU). LOUISIANA. Bossier Parish:

Red River ca. 10 river-km upstream from LA 2 bridge, ca. 6 air-km E of Mira and ca. 8.7 air-km NE of Hosston, 32.9532900, -93.8315800,

open sparsely vegetated sand dune field on east side of river bed, dry loose sand, common and scattered with Heterotheca subaxillaris, Cy¬

cloloma atriplicifolium, and Cyperus esculentus, 16 Aug 2012, Reid and Lewis 8300 (LSU).

ACKNOWLEDGMENTS

Gary Hansen and Amanda Lewis of the Red River Watershed Management Institute, Louisiana State Univer¬

sity at Shreveport, and Michael MacRoberts, curator of LSUS Herbarium and associate of the Red River Water¬

shed Management Institute, provided valuable assistance and support in implementing this survey Hubert

Hervey kindly allowed use of his boat and motor. Nicole Lorenz prepared Figure 1 and we greatly appreciate

her contribution. We appreciate the reviews of our manuscript by Michael MacRoberts, David Rosen, Brett

Serviss, and Theo Witsell.

REFERENCES

Correll, D.S. & M.C. Johnston. 1970. Manual of the vascular plants of Texas. Texas Research Foundation, Renner, Texas,

U.S.A.

Gentry, J.L., G.P. Johnson, B.T. Baker, C.T. Witsell, & J.D. Ogle, eds. 2013. Atlas of the vascular plants of Arkansas. University of

Arkansas Herbarium, Fayetteville, Arkansas, U.S.A.

Hoagland, B.W., A.K. Buthod, I.H. Butler, P.H.C. Crawford, A.H. Udasi, WJ. Elisens, & R.J. Tyrl. 2004. Oklahoma Vascular Plants

Database (http://geo.ou.edu/botanical), Oklahoma Biological Survey, University of Oklahoma, Norman, Oklahoma,

U.S.A.

Reid and Lewis, Plant exploration along the Red River of Arkansas and Louisiana

201

Kaul, R.B. 1986. Boraginaceae. In: The Great Plains Flora Association, eds. Flora of the Great Plains. University Press of

Kansas, Lawrence, Kansas, U.S.A. Pp. 683-701.

MacRoberts, D.T. 1984. The vascular plants of Louisiana. An annotated checklist and bibliography of the vascular plants

reported to grow without cultivation in Louisiana. Bull. Mus. Life Sci. 6:1 -165.

MacRoberts, D.T. 1989. A documented checklist and atlas of the vascular flora of Louisiana. Acanthaceae to Fabaceae.

Bull. Mus. Life Sci. 8:257-536.

McGregor, R.L. 1986. Fabaceae. In: The Great Plains Flora Association, eds. Flora of the Great Plains. University Press of

Kansas, Lawrence, Kansas, U.S.A. Pp. 416-490.

NatureServe. 2013. NatureServe Explorer: An online encyclopedia of life [web application]. Version 7.1 NatureServe,

Arlington, Virginia. Available http://www.natureserve.org/explorer. Accessed 22 April 2013.

Reid, C. & P. Faulkner. 2006. Loeflingia squarrosa (Caryophyllaceae): new to Louisiana. Phytologia 88(1):97-99.

Singhurst, J.R. & W.C. Holmes. 1999. Noteworthy collections: Arkansas. Castanea 64:276-277.

Smith, E.B. 1988. An atlas and annotated checklist of the vascular plants of Arkansas. Published by E.B. Smith.

Thomas, R.D. &C.M. Allen. 1998. Atlas of the vascular flora of Louisiana. Vol. III. Dicotyledons: Fabaceae-Zygophyllaceae.

Louisiana Department of Wildlife and Fisheries, Baton Rouge, Louisiana, U.S.A.

Turner, B.L., H. Nichols, G. Denny, & O. Doron. 2003. Atlas of the vascular plants of Texas. Volume 1. Sida Bot. Misc. 24:1-648.

USDA, NRCS. 2013. The PLANTS Database (http://plants.usda.gov). National Plant Data Team, Greensboro, North Caro¬

lina 27401 -4901 U.S.A. Accessed 22 April 2013.

202

Journal of the Botanical Research Institute of Texas 8(1)

BOOK NOTICE

Paul Dowlearn. 2013. The Lazy Man’s Garden. Maximum Return; Minimum Input. (ISBN-13: 978-1-

491060889, pbk). Published by the author, www.wvlandscape.com. (Orders: available atAmazon.com).

$14.95,156 pp., 6" x 9".

From the publisher: The Lazy Man’s Garden: Maximum return; Minimum input, a nonfiction work of 58,600

words, explains practical approaches to gardening with emphasis on xeriscape and adjusting attitudes to prac¬

tical gardening. Dowlearn writes, “Americans are intensively growing millions of acres of lawn grasses. We are

also attempting to grow mostly non-native exotic hybrids for ornamentals. Much of this traditional culture is

not sustainable. Vegetable gardening is one of the few things that survived our landscaping ethic to give the

homeowner benefits that go beyond aesthetics. Utilizing native plants, creating habitat for wildlife, and seek¬

ing out old tried and true heirloom plants are current trends that promise a more relaxed, eco-friendly style.”

The author, PAUL DOWLEARN, is co-owner of Wichita Valley Landscape (Wichita Falls, TX). He does land¬

scape design and installation through his nursery and has focused on xeriscape and low maintenance land¬

scapes for the last twenty years. Dowlearn has authored numerous articles on native and well adapted plants,

plus organic gardening methods for newspapers and magazines, as well as speaking to many organizations

throughout Texas and Oklahoma. He has hosted local radio and TV call-in shows and teaches landscaping

courses at Vernon College. He is past president of the Red River Chapter of the Native plant Society of Texas,

member of the Ladybird Johnson Wildflower Research Center, Riverbend Nature Center, Texas Organic Farm¬

ers & Gardeners Association and several other non-profit organizations.

J.Bot. Res. Inst. Texas 8(1): 202.2014

VEGETATION AND VASCULAR FLORA OF TALLGRASS PRAIRIE AND

WETLANDS, BLACK SQUIRREL CREEK DRAINAGE, SOUTH-CENTRAL

COLORADO: PERSPECTIVES FROM THE 1940s AND 2011

Sylvia Kelso

Department of Biology, Carter Herbarium

Colorado College

Colorado Springs, Colorado 80903, U.S.A.

tkelso@coloradocollege.edu

Leah Fugere

Environmental Program

Colorado College

Colorado Springs, Colorado 80903, U.S.A.

Miroslav Kummel

Sebastian Tsocanos

Environmental Program

Colorado College

Colorado Springs, Colorado 80903, U.S.A.

mkummel@coloradocollege.edu

Environmental Program

Colorado College

Colorado Springs, Colorado 80903, U.S.A.

ABSTRACT

We examined a tallgrass prairie-wetland complex of the Black Squirrel Creek drainage in south-central Colorado to compare the current

grassland composition to its documentation by Robert Tivingston in the early 1940s. Tivingston considered these grasslands as probable

Pleistocene relicts analogous to Midwestern tallgrass prairie with respect to dominant grasses and forbs. Using Livingston’s methodology,

we assessed an area near his original plots to determine whether the dominant grass species had changed in their contributions to cover or

frequency. We found an almost identical suite of species to those documented in the 1940s, with modest differences in frequency and rela¬

tive contribution to cover by the key grasses. We also characterized wetland habitats occurring within the grassland matrix, documented

the vascular flora of mesic and hydric habitats, and analyzed the extent to which they contain species of conservation concern, Midwest

prairie elements, or montane species typically occurring regionally at higher elevations. The tallgrass communities here differ from others

in Colorado and the Midwest in having a lower abundance of Andropogon gerardii, and being dominated by Sporobolus heterolepis along with

xeric species like Bouteloua gracilis and Calamovilfa Xongifolia, and montane species like M uhlenbergia montana. Although the structure of

the extant vegetation remains similar to what existed in the 1940s and continues to be supported by ample groundwater, these grasslands

are now reduced in extent. The vegetation mosaic of tallgrass prairie and wetlands holds a rich flora with numerous elements of phytogeo¬

graphic and conservation interest.

RESUMEN

Examinamos el complejo de praderas de hierbas altas y humedales del desague de Black Squirrel Creek, en el sur-centro de Colorado, para

comparar la composicion actual de las praderas con la que Robert Livingston documento a principios de los anos cuarenta. Livingston con-

sidero que estas eran analogas a las praderas de hierbas altas del Medio Oeste, con respecto a los pastos y formas dominantes, probables re-

lictos de la vegetacion del Pleistoceno. Usando la metodologia de Livingston, evaluamos un area cercana a las parcelas originales para deter-

minar si las especies dominantes de pastos han cambiado en su contribucion a la cobertura o en su frecuencia. Encontramos un conjunto de

especies casi identico al de las documentadas en los cuarenta, con diferencias pequenas en frecuencia y contribucion a la cobertura por

parte de pastos clave. Tambien caracterizamos los humedales presentes en el marco de la pradera, documentamos las plantas vasculares de

habitats mesicos e hidricos, y analizamos en que medida estos contienen especies de interes para la conservacion, elementos de las praderas

del Medio Oeste o especies montanas comunes en regiones de mayor altitud. Las comunidades de pastos altos en este lugar difieren de otras

en Colorado y en el Medio Oeste por la menor presencia de Andropogon gerardii y la presencia dominante de Sporobolus heterolepis, junto con

especies aridas o montanas, como Bouteloua gracilis, Calamovilfa longifolia y Muhlenbergia montana. Aunque la estructura de la vegetacion

existente sigue siendo similar a la de los anos cuarenta y aun cuenta con el soporte de abundantes aguas subterraneas, la extension de estas

praderas se ha reducido. El mosaico de vegetacion de praderas de hierbas altas y humedales conserva una rica flora con numerosos elemen¬

tos de interes para la geobotanica y la conservacion.

INTRODUCTION

Over the past century, Colorado prairies changed and diminished as urban, suburban, and exurban develop¬

ment expanded while bre suppression, invasion of exotic species, and overgrazing altered grassland compo-

J. Bot. Res. Inst. Texas 8(1): 203 - 225.2014

204

Journal of the Botanical Research Institute of Texas 8(1)

nents. Prairie vegetation overall, and tallgrass communities in particular, now elicits special conservation fo¬

cus amid widespread concerns about accelerating loss, fragmentation, and degradation throughout the Mid¬

west and West (Nicholson & Hulett 1969; Samson & Knopf 1996; Bachand 2001; Colorado Natural Heritage

Program 2005; Rondeau et al. 2011). In the early 1940s, ecologist Robert Livingston undertook a detailed study

of tallgrass prairie in south-central Colorado north of Colorado Springs (Fig. 1). He profiled this vegetation in

his graduate theses and related publication (1941,1947,1949,1952) as unique remnant vegetation with strong

floristic similarities to the Midwest prairies. This work provided a portrait of regionally anomalous vegetation

as it existed nearly seventy years ago.

In this study, we reanalyzed the grassland vegetation in the Black Squirrel Creek drainage (Fig. 2) that

was the focus of Livingston’s work. We compared its current composition to the earlier descriptions and added

additional documentation of associated wetland communities and the vascular flora. Although these addi¬

tional components were not a focus in the original Livingston studies, contemporary conservation interest in

these elements suggested their importance as part of the regional ecological profile. The objectives of our study

were to:

1) Document the flora of the mesic and hydric communities and assess the extent to which this flora contains

elements from the Midwest prairie or other regional components such as montane species typically oc¬

curring in the foothills, and highlight species of concern.

2) Document the types of wetland habitats occurring in the grassland matrix and their signature flora, plant

associations and hydrogeomorphic profiles.

3) Compare the current dominant species and composition of the vegetation to the 1940’s profile considered

representative of Midwestern tallgrass prairie. In particular, we examined whether the relative rank of

the dominant grass species had changed with respect to contribution to cover and frequency.

Regional vegetation contexts: shortgrass, mixed grass and tallgrass prairie

The eastern plains of Colorado encompass a wide range of grassland communities across diverse topography

and edaphic substrates (Ramaley 1919; Shantz 1923; Weaver & Fitzpatrick 1934). These grasslands represent

the western edge of the Great Plains, where collective vegetation types and many individual biotic components

have been diminished from their historic presence (Rondeau et al. 2011). Shortgrass prairie (also known as

shortgrass steppe senus Lauenroth et al. 2008) with its signature species Bouteloua gracilis (Willd. ex Kunth)

Lag. ex Griffiths (nomenclature herein follows the U.S. Dept, of Agriculture National Resources Conservation

Service Plants Database 2013 (www.plants.usda.gov); see Appendix 1 for full citations and exceptions) is the

dominant regional vegetation (Weaver 1954; Neeley et al. 2006; Lauenroth et al. 2008). Shortgrass prairie

south of Denver typically lacks extensive amounts of Buchloe dactyloides, a codominant elsewhere in short¬

grass prairie, although it occurs sporadically in the shortgrass matrix here. In the piedmont east of the Colo¬

rado Front Range foothills, a mixed grass prairie prevails, where grama grass is present in conjunction with a

high representation of species in Elymus, Hesperostipa, Muhlenbergia, and Poa along with Schizachyrium sco-

parium (Michx.) Nash), Sporobolus cryptandrus (Torrey) A. Gray, and Koeleria macrantha (Ledeb.) Schult. This

vegetation is extensive north and east of Colorado Springs, where the topographic watershed known as the

Palmer Divide separates the drainages of the South Platte River to the north and the Arkansas River to the

south.

Limited occurrences of tallgrass prairie vegetation (Vestal 1914, 1917, 1919; Moir 1972; Bock & Bock

1998) exist just east of the Front Range in areas where edaphic conditions enhance soil moisture (Branson et

al. 1965). The Colorado Natural Heritage Program (2012) tracks four tallgrass communities of conservation

concern: xeric tallgrass prairie dominated by Andropogon gerardii Vitma and Sporobolus heterolepis (A. Gray) A.

Gray), or A. gerardii and Schizachyrium scoparium (Michx.) Nash), and mesic tallgrass prairie dominated by A.

gerardii and Calamovilfa longifolia (Hook.) Scribn.), or A. gerardii and Sorghastrum nutans (L.) Nash). These as¬

sociations are part of mixed vegetation that comprises the collective Western Great Plains Foothill and Pied¬

mont Grasslands (Colorado Natural Heritage Program 2005).

Kelso et al., Flora of tallgrass prairie and wetlands. Black Squirrel Creek drainage

205

Fig. 1. Generalized area of study and Black Squirrel Creek drainage. Main study site is noted in red shading. The Palmer Divide is the region between El

Paso and Douglas Counties, and represents the hydrological divide between the Platte and Arkansas Rivers.

At the species level, Colorado tallgrass prairies parallel those found in the Midwest where signature taxa

include Andropogon gerardii, Hesperostipa spartea (Trin.) Barkworth), Panicum virgatum L., Sorghastrum nutans,

and Sporobolus heterolepis, as well as the sand prairie tallgrass species Calamovilfa longifolia (e.g., Weaver 1954;

Freeman 1998). The relative amounts of these species typically differ among central and western states (Weav¬

er 1954; Weaver & Albertson 1956) according to precipitation and temperature regimes and soil types; in

206

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 2. Core study site along upper Black Squirrel Creek drainage, east of Colorado Springs. Transect locations are indicated by numbered ellipses; sizes

are exaggerated for clarity.

Colorado, it has long been recognized that these communities similarly vary along a north-south gradient

(Robbins 1910; Vestal 1914,1917,1919).

South of the Palmer Divide, isolated examples of tallgrass prairie occur within a ponderosa pine forest-

grassland matrix known as the Black Forest (Shaddle 1939; Fig. 1). Studies by Vestal (1917), Shaddle (1939),

and Williams and Holch (1946) noted the unusual vegetation and flora here. This region and surrounding

Kelso et al., Flora of tallgrass prairie and wetlands. Black Squirrel Creek drainage

207

grasslands became the focus of studies in which Livingston (1941,1947,1949,1952) highlighted the similarity

of the grassland vegetation and flora to Midwestern “true prairie”, and suggested it represented fragmented

relict communities from the Pleistocene (Livingston 1952; Weaver and Albertson 1956). Although relatively

recent studies of tallgrass vegetation in eastern Colorado have been conducted in the Boulder area ca. 150 km

north of the Black Forest (e.g., Moir 1972; Baker & Galatowitch 1985; Bock & Bock 1998; Neid et al. 2009), no

assessment of the Black Squirrel Creek vegetation has been done since those of Livingston, although surveys

by the Colorado College Herbarium and the Colorado Natural Heritage Program (Doyle et al. 2001a; 2001b)

have shown the area to be rich in rare species and ecological communities.

Study Site

The Black Squirrel Creek system, a complex anastomosed network of drainages into the main creek channel,

begins near the summit of the Palmer Divide (Figs. 2, 3) in the Black Forest and extends southeast, ultimately

draining into the Arkansas River east of the city of Pueblo. In the upper quarter of the drainage, the creek typi¬

cally has perennial flow, but this becomes intermittent aboveground to the south. Our primary study area was

located on the upper Black Squirrel drainage between the municipalities of Falcon and Peyton, with an eleva¬

tion range of ca. 2000 to 2200 m (6500 to 7100 ft). Quantitative data were taken on the core area covering ca.

770 ha (1900 acres) on a ca. 3100 ha (7700 acre) ranch that encompasses the main stem of Black Squirrel Creek

with perennial flowing water, as well as with subsidiary drainages with intermittent flow and standing water.

We took additional floristic and qualitative information south of the core area for a distance of ca. 10 km in

order to encompass the area utilized in the 1940’s studies. The topography consists of gently rolling uplands of

mixed grass prairie on Quaternary deposits of aeolian sand (Morgan & Barkmann 2012) separated by lowland

drainages and swales with small discontinuous wetlands, seeps, springs, and seasonal ponds supported by

groundwater. Our study focused on the mesic and hydric flora and vegetation occurring in these drainages, the

associated wetlands, and streambeds of Black Squirrel Creek and its tributaries rather than the xeric and

mixed grass vegetation of the uplands that is widely represented on the plains.

Climate

Longterm annual precipitation since 1956 for this region averages ca. 38 cm (17 in; Western Regional Climate

Center, data for Eastonville, CO) with ca. 75% of this occurring from spring rains in April to May and a July-

August pulse from thunderstorms. Interannual variation in precipitation can be extreme in Colorado, with

severe droughts occurring in the 1950s, 1970s, early 1980s, and early 2000s (Henz et al. 2004). Local rainfall

tracks the topographic gradient, where higher elevations near the top of the drainage receive more rainfall in

the summer and more winter storm events; these are often highly localized and precipitation events can vary

over short distances. Longterm temperature records (Western Regional Climate Center; data for Colorado

Springs 1948-2005) indicate an average daily high of 26.6° C (79.9° F) - to an average low of 12.7° C (54.8° F)

in the growing season months of June to August, while January temperatures range from an average high of

5.9° C (42.6° F) to a low of 16.6° F (-8.5 C°). These climatic parameters do not differ significantly from those

reported by Livingston (1947).

Geology

Livingston (1952) and Branson et al. (1965) noted the relationship between soils of perennial high moisture

and the persistence of tallgrass prairie vegetation in Colorado; these azonal conditions are promoted by soil

composition and water table dynamics. In our study area, two aquifers play a significant role in local hydrology

and soil moisture: the alluvial Black Squirrel Creek aquifer and the underlying andesitic sandstone Dawson

Formation bedrock aquifer (Bittenger 1976; Robson 1988; Topper 2008; Morgan & Barkmann 2012). Subsur¬

face topography slopes steeply south, shaped by the ancestral Black Squirrel Creek now covered with glacial

alluvium that forms the Black Squirrel Creek aquifer of Pleistocene gravels and coarse sand. This aquifer

ranges in depth from 0-215 m; close to the headwaters of Black Squirrel Creek it is relatively shallow but it

becomes deeper to the southeast. The alluvial aquifer is a significant source of well water for domestic, agricul¬

tural, and municipal uses (Topper 2008) and provides localized high water table occurrences. However, in our

208

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 3. Main stem of Black Squirrel Creek. Upland areas contain mixed grass prairie vegetation and tallgrass vegetation occurs along the side banks.

study area the alluvial layer is relatively thin or absent, and much of the subsurface water here that supplies the

seeps and springs is likely to be primarily a result of the Dawson Formation bedrock aquifer (R. Topper, pers.

comm.), a late Cretaceous-Tertiary sandstone widely exposed along the Palmer Divide and close to the surface

in the upper Black Squirrel Creek drainage. Becausethe aeolian and alluvial surface deposits of the area are

highly permeable, runoff is low, and both aquifers receive recharge from precipitation (Topper 2008). The in¬

terface of the alluvial and bedrock aquifers is regionally complex with no detailed mapping of localized hydro-

logical regimes yet available, and it is likely that both the alluvial and bedrock aquifers supply groundwater

across the entirety of the drainage.

MATERIALS AND METHODS

The study area was surveyed intensively from May to August 2011 throughout the growing season, although

floristic collections occurred regularly since 2000 and sporadically since the 1970s. We conducted qualitative

surveys of topography and floristic composition of significant plant communities on each section of the main

and subsidiary drainages every two weeks during 2011. We documented the vascular flora as completely as

possible, combining new collection with existing recent ones at the Carter Herbarium of Colorado College

(COCO); primary voucher specimens for all taxa are at COCO, with duplicates at COLO and CS. Vegetation

communities and significant species and wetland features along the main drainage and its key tributaries were

mapped using geospatial coordinates.

Our initial surveys along the drainage of Upper Black Squirrel Creek provided qualitative assessments of

the current extent and condition of grassland communities in sites as close as possible to those in the original

Livingston studies, for which only generalized location information was available. Due to landscape modifica¬

tion in housing developments and roads or inability to obtain permission for access, the most intact communi¬

ties in which we were able to take quantitative data were located ca. 10 km north of the original Livingston

Kelso et al., Flora of tallgrass prairie and wetlands. Black Squirrel Creek drainage

209

plains sites, and a similar distance south of his forest-grassland interface plots (Livingston 1947; 1952) within

the same hydrologic system in comparable topography We were able to access the general area where his

original sites were located for floristic information, although we did not take quantitative data due to distur¬

bance from intensive grazing and anthropogenic surface alterations.

Livingston utilized nine 100m line transects in seven locations: four sites, each with a single transect,

were in the Black Forest within a ponderosa pine-savannah community at 2200-2300 m (7200-7500 ft.), and

three locations (one with 3 transects) were along the Black Squirrel Creek drainage in grasslands at ca. 2000 m

(6500 ft). The original transects each encompassed ten quadrats of 1 x 0.5 m; in comparison, we used four 105

m transects, each with 20 similarly sized quadrats spaced 5 m apart. This modification allowed us to assess the

small-scale heterogeneity characteristic of the region. All 2011 transects were located at approximately 2100 m

(7000 ft) in elevation.

Transects 1 and 2 were located along the north-facing bank of the main stem of Black Squirrel Creek

(Figs. 2, 3) in vegetation that met our criteria for sufficient length with the presence of tallgrass indicator spe¬

cies Sporobolus heterolepis, Andropogon gerardii , Hesperostipa spartea , or Sorghastrum nutans. These transects

were parallel, ca. 10 m apart, and offset so that they overlapped by one half their length. We located two addi¬

tional transects (Fig. 2) along subsidiary drainages in vegetation with similar criteria: Transect 3 was in a side

drainage ca. 1 km north of transects 1 and 2, and Transect 4 was located in a tributary drainage, ca. 3 km north

of transect 3. To match the protocol, timing, and data format of Livingston, we surveyed the vegetation in late

August following his methodology for assessing basal and relative cover and frequency of dominant taxa.

Basal Cover and Relative Cover

In each quadrat, we estimated the total basal cover as a percentage of the total area, as well as the percent cover

of bare ground and litter combined, then averaged these over each transect of 20 quadrats. For each quadrat we

estimated the total cover of all plants, the relative cover of combined graminoids (Poaceae, Cyperaceae, Junca-

ceae and Juncaginaceae) as a percentage of the total vegetative cover, and the contribution of each identifiable

species to the total graminoid cover.

Frequency

In each quadrat we recorded the presence of all graminoid species and calculated a frequency metric and rank

of the dominant taxa by summing the quadrat data for each transect individually. We also calculated the mean

frequency and rank frequency of each species across all transects.

RESULTS

Vascular Flora

The flora reported here (Appendix 1) represented only the hydric and mesic habitats and does not encompass

species restricted to the more xeric uplands. Some components of this upland vegetation extended into the

drainages, particularly in the open gravels of stream banks and terraces, and are included on the species list,

noted as a xeric component. We defined notable elements (Table 1) in four categories. Rare taxa were those

tracked as being of conservation concern by the Colorado Natural Heritage Program. Regionally uncommon

taxa were those with few regional herbarium records COCO and so categorized in prior studies by Kelso

(2012), Culver and Lemly (2013) or Weber and Wittmann (2012). Foothills/Montane elements were topo¬

graphic disjuncts that typically occur in higher elevation locations of the Pikes Peak or Front Range foothills as

noted by Weber and Wittmann (2012) and herbarium records at COCO. Midwestern elements were taxa as¬

sociated with the characteristic Midwest Prairie flora as explicitly noted by Shantz (1928), Weaver and Fitzpat¬

rick (1934), Weaver (1954), or Livingston (1952).

In the current study, we documented almost 300 taxa representing 62 families as currently recognized in

the U.S.D.A. Plants Database (www.plants.usda.gov). The highest species richness was in the Asteraceae (50

species), Poaceae (43 species), Cyperaceae (18 species), and Juncaceae (12 species). Additional families with

high species richness included the Fabaceae, Polygonaceae, Rosaceae, and Scrophulariaceae (s. lat.). The flora

included relatively few noxious weeds; Ly thrum salicaria, which existed sporadically in a side drainage of Black

210

Journal of the Botanical Research Institute of Texas 8(1)

Table 1. Notable plant taxa in the upper Black Squirrel Creek drainage. See Appendix 1 for full nomenclature. Midwest Prairie affiliated species are those so noted

by Shantz (1928), Weaver and Fitzpatrick (1934), Weaver (1954) and Livingston (1952). Foothills/Montane distributions in Colorado by Weber and Wittman

(2012), and Kelso (2012). Locally uncommon species are from records in COCO and prior fieldwork by Kelso but not tracked statewide by Colorado Natural Heritage

Program (CNHP); Rare species are those tracked by the Colorado Natural Heritage Program (2012) as being of conservation concern. CNHP state rarity ratings are as

follows: SI=Critically imperiled due to extreme rarity, or factors making it vulnerable to extirpation; 5 or fewer occurrences or less than 1000 remaining individu¬

als; S2=lmperiled, 6-20 occurrences or between 1000-3000 remaining individuals; S3= Vulnerable, 21-100 occurrences or between 3000-10,000 remaining

individuals! S4: Apparently secure, uncommon but widespread, with possible longterm concern (www.cnhp.colostate.edu).

Family

Species

Foothills/Montane Midwest Prairie Locally Uncommon Rare

Anacardiaceae

Toxicodendron rydbergii

Apiaceae

Cicuto douglosii

Asdepiadaceae

Asclepias hallii

X:S3

Asteraceae

Artemisia ludoviciana

Cosmos parviflorus

Erigeron lonchophyllus

Helenium autumnale

Helianthus pauciflorus

Liatris ligulistylis

X:S1/S2

Oligoneuron albidum

X:S2/S3

Oligoneuron rigidum

Packera pseudaurea

Rudbeckia hirta

Solidago missouriensis

Solidago nana

Symphotrichum ericoides

Symphotrichum laeve

Tripleurospermum perforatum

Brassicaceae

Arabis holboelii var. retrofracta

Draba nemorosa

Cactaceae

Pediocactus simpsonii

Campanulaceae

Campanula rotundifolia

Lobelia siphilitica

Caryophyllaceae

Stellaria longifolia

Clusiaceae

Hypericum scouleri

Cyperaceae

Carexaurea

Carexcrawei

X:S1

Carex disperm a

Carex echinata

Carex simulata

Eleocharis quinqueflora

Equisetaceae

Equisetum arvense

Fabaceae

Astragalus canadensis

Dalea Candida

Dalea purpurea

Glycyrrhiza lepidota

Gentianaceae

Gentianopsis amarella

(.strictiflora type)

Gentianopsis virgata

(not yet

CNHP

rated)

Hippuridaceae

Hippuris vulgaris

Iridaceae

Hypoxis hirsuta

X:S1

Juncaceae

Juncus brachycephalus

X:S1

Juncus brevicaudatus

X:S1

Juncaginaceae

Triglochin palustris

Lamiaceae

Lycopus americanus

Monarda fistulosa

Scutellaria galericulata

Lentibulariaceae

Utricularia minor

X:S2

Liliaceae

Lilium philadelphicum

X:S3/S4

Malvaceae

Sidalcea neomexicana

Onagraceae

Gayophytum diffusum

Oenothera flava

Kelso et al., Flora of tallgrass prairie and wetlands. Black Squirrel Creek drainage

211

Table 1. Continued

Family

Species

Foothills/Montane

Midwest Prairie

Locally Uncommon Rare

Orchidaceae

Platanthera aquilonis

Spironthes romonzoffiono

Poaceae

Agropyron cristatum

Agrostis scobro

Alopecurus aequalis

Andropogon gerordii

Bouteloua curtipendula

Colomogrostis stricta

Calamovilfa longifolia

Elymus canadensis

Glyceria elata

Glyceria striata

Hesperostipa spartea

Koeleria macrantha

Muhlenbergia montana

Pan icum virgatum

Poa fendleriana

Poa nemoralis ssp. interior

Poa pratensis

Schizachyrium scoparium

Sorghastrum nutans

Sporobolus heterolepis

Polygonaceae

Polygonum amphibium

Primulaceae

Dodecatheon pulchellum

Lysimachia ciliata

Ranunculaceae

Anemone canadensis

Anemone cylindrica

Rosaceae

Agrimonia striata

Geum aleppicum

Potentilla arguta

Rosa arkansana

Rubiaceae

Galium boreale

Galium trifidum

Salicaceae

Populus angustifolia

Salixirrorata

Selaginaceae

Selaginella densa

Scrophulariaceae

Nuttallanthus canadensis

Pedicularis canadensis

Penstemon glaber

Penstemon gracilis

Veronica serpyHi folia

Sparganiaceae

Sparganium angustifolium

Valerianceae

Valeriana edulis

Violaceae

Viola sororia

Squirrel Creek,

is the only A list species. Although a detailed floristic list was not an objective of the Livingston

studies, we found almost all taxa he noted as still present, with only a few exceptions (Appendix 1).

Over one fifth of the flora was regionally associated with foothills/montane habitats, and at least a compa¬

rable proportion was characteristic of the Midwest prairies. By contemporary phytogeographic perspectives

and greater documentation of the Great Plains flora, the Midwest association was almost certainly an underes¬

timate, but to simplify comparison, we used for reference only those species explicitly listed in early studies as

characteristic of the Midwest. Eight of the “Midwest” species were also locally characteristic of the foothills/

montane zone. Eighteen plant species occurring in the Black Squirrel Creek drainage were locally uncommon,

and nine were tracked by the Colorado Natural Heritage Program for being of conservation concern.

212

Journal of the Botanical Research Institute of Texas 8(1)

Wetland habitat Classifications

We identified eight general wetland habitat types characterized by distinct hydrogeomorphic characteristics

and floristic profiles (Table 2). Our classification follows the Colorado Natural Heritage Program (Carsey et al.

2003) and includes general categories of Riverine Wetlands sourced by ongoing streamflow, Slope Wetlands

supported by groundwater on gentle to moderate slopes, and Depressional Wetlands supported by ground

water hlling a depression on a permanent or intermittent basis. Each habitat type occurs in multiple instances

throughout the Black Squirrel drainage. Vegetation associations listed for each hydrogeomorphic class follow

those used by the Colorado Natural Heritage Program (Carsey et al. 2003; Culver & Lemley 2013) classifica¬

tions as closely as possible.

Riverine Wetlands

Stream Channel Tall Willow Shrubland. —This community occurred in a limited extent on the northwestern

edge of the main Black Squirrel Creek drainage, covering about a kilometer in length; shrub cover diminished

further downstream, but reoccured in patches along the drainage in wide stream meanders with shallow sub¬

surface water. The primary vegetation community was sandbar willow-mesic graminoid shrubland dominated

by Salix exigua with occasional occurrences of other tree and shrub species of willow (e.g., S. irrorata, S. liguli-

folia, and S. amygdaloides); the forb component was limited but included patchy occurrences of Agrimonia

striata, Cirsium canadensis, Glycyrrhiza lepidota, Helianthus nuttallii, Monardafistulosa, and Rudbeckia hirta.

Stream Channel Herbaceous Vegetation. —Open gravels of the main channel and occasional side drainages

supported a linear strip of obligate wetland forbs, sedges, and rushes where stream flow formed riffles around

gravel banks and sandbars. The gravel stream channels were notable for their abundance and diversity of

rushes, including the rare Juncus brachycephalus, as well as Gentianopsis virgata. Both of these species are Mid¬

west prairie elements known in Colorado only from this region. Although dominated by non-woody vegeta¬

tion, the stream channels also supported occasional occurrences of young saplings of Populus deltoides or spe¬

cies of Salix.

Slope Wetlands

Moist Shelves. —These heterogeneous surfaces were located primarily in the main drainage above the creek

channel depression and along some subsidiary drainages. Surfaces were flat to gently sloping, with moisture

accumulating from springs and runoff above. Moister areas held a greater abundance of facultative or obligate

wetland species interspersed with xeric elements. Vegetative cover was primarily composed of graminoids and

mixed forbs with occasional shrub patches. Sporobolus heterolepis was particularly widespread here, and the

shelves supported extensive occurrences of a Sporobolus heterolepis dominated community with occasional

instances of Andropogon gerardii, along with patches of Andropogon gerardii-Sorghastrum nutans associations.

The Sporobolus-dominated communities ranged in width from 5 m to almost 40 m; depending on the topogra¬

phy, lengths could be short patches of 10 m to longer extents over 50 m. Drier areas included heterogeneous

mixed grass vegetation with Calamovilfa longifolia, Koeleria macrantha, Muhlenbergia montana, Poa pratensis,

and Schizachyrium scoparium.

Moist Banks. —These encompassed a significant portion of the drainage system and held some of the high¬

est diversity of forbs. One of the most significant species occurring here was the locally abundant, state-rare

Liatris ligulistylis. The moist banks typically occurred on side drainages with a U-shape profile and received

consistent subsurface moisture from seeps and springs; their surfaces were steeper than moist shelf habitats,

and they usually included seeps that oozed perennial moisture. Plant associations included mixed mesic tail-

grass communities with components of Calamovilfa longifolia, Schizachyrium scoparium, Sorghastrum nutans,

Sporobolus heterolepis, and Stipa spartea. Andropogon gerardii clumps were common, but did not form a domi¬

nant component of the vegetative cover. Like moist shelf communities, bank communities sometimes occurred

as lengthy strips to 50 or more meters, or as shorter patches interspersed with depressional wetlands.

Depressional wetlands

Nebraska Sedge Bogs and Meadows. —These associated habitats were both dominated by Carex nebrascensis and

Kelso et al., Flora of tallgrass prairie and wetlands. Black Squirrel Creek drainage

213

Table 2. Common floristic elements in wetland communities of the Black Squirrel Creek drainage. For full nomenclature see Appendix 1.

Graminoids

Forbs

Shrubs

RIVERINE WETLANDS

Stream Channel Tall

Agrimonio strioto

Solixexiguo

Willow Shrubland

Monarda fistuloso

Rudbeckia hirta

Helianthus nuttollii

Salixirrorata

Solixligulifolio

'Stream Channel

Juncus alpinoarticulatus

Epilobium ciliotum

Herbaceous Wetlands

Juncus brachycephalus

Juncus dudleyi

Juncus interior

Juncus saximontanus

Juncus torreyi

Gentionello strictiflora

Gentianopsis virgoto

Onosmodium bejoriense

Veronica onogollis-oquotico

SLOPE WETLANDS

Moist Shelves

Andropogon gerordii

Antennorio microphyllo

Rosa orkonsono

Calamagrostis stricto

Elymus lanceolatus

Juncus arcticus

Poo pratensis

Schizachyrium scoparium

Helianthus nuttallii

Monordo fistuloso

Pediculoris canadensis

Ratibida columnifera

Rudbeckia hirta

Astragalus canadensis

Erigeron bellidiostrum

Fragaria vesca

Geum oleppicum

Glycyrrhiza lepidota

Symphoricarpos occidentalis

'Moist Banks

Andropogon gerordii

Poo protensis

Sorghastrum nutons

Stipo sporteo

Liotris ligulistylis

Potentillo orguto

Prunella vulgaris

Rudbeckia hirta

Thermopsis montono

Viola sororia

Collomio linearis

Geum aleppicum

Helenium outumnole

Helianthus nuttallii

DEPRESSIONAL WETLANDS

Sedge Meadows & Bogs

Corex nebrascensis

Glycerio eloto

Glycerio striata

Juncus nodosus

Juncus saximontanus

Juncus torreyi

Poa leptocoma

Epilobium leptophyllum

Lobelia siphilitico

Mentha arvensis

Polygonum pensylvanicum

Polygonum punctatum

Scutellaria golericuloto

Stellararia longifolia

Open Seeps

Co rex oureo

Corex crowei

Corexxerontico

Eleochoris quinquefloro

Juncus alpinoarticulatus

Triglochin maritima

Triglochin polustris

Hypoxis hirsuto

Spiranthes romanzoffiana

Dodecatheon pulchellum

Gentianopsis virgata

Pornossio polustris

Platanthera aquilonis

Sisyrinchium montanum

Fens

Carexsimulata

Eleochoris ociculoris

Scirpus pungens

Helenium autumnale

Ponds

Scirpus microcorpus

Alismo triviole

Hippuris vulgaris

Polygonum omphibium

Potamogeton natans

Ranunculus trichophyllus

Sagittaria cuneata

Sogittorio lotifolio

Sparganium angustifolium

Utriculorio minor

214

Journal of the Botanical Research Institute of Texas 8(1)

typically occurred adjacent to streams and marshy areas with a high water table overlain by a layer of sediment

and organic material. In the bogs, Glyceria was often present, along with a limited number of forbs such as

members of the Polygonaceae and Helenium autumnale. Tributary channels above the water flow of drainage

bottoms supported the more abundant sedge meadow community, which also occurred along shallow chan¬

nels with no visible surface water. Sedge meadows were drier and more floristically diverse in hydrophytic

graminoids, with Nebraska sedge occurring along with Juncus arcticus, other sedge species such as Carex di-

sperma , as well as bulrush species in Scirpus and Schoenoplectus. Forbs included hydrophytes such as Lobelia

siphilitica and Scutellaria galericulata, both regionally uncommon but locally abundant here, as well as the

widespread Mentha arvensis and representatives of the Polygonaceae.

Open Seeps .—Open seeps underlain by clay lenses occurred frequently throughout the drainage system.

Groundwater emerged through the soil to create a shallow layer of standing water 1-2 cm deep over saturated

clay-rich mud with little to no vegetative cover. Seeps ranged from ca. 1 m 2 to 100 m 2 in area and were located

above stream level along shallow bank margins. The surfaces were dotted with low hummocks, vegetation-

covered mounds from 10 to 50 cm in height and width. These habitats encompassed an unusual flora com¬

posed of species more typical of higher elevations (e.g., Dodecatheon pulchellum, Eleocharis quinqueflora, and

Parnassia palustris) along with a number of state-rare species, all Midwest prairie elements, such as Carex cra-

wei, Gentianopsis virgata, and Hypoxis hirsuta.

Fens .—This habitat type was a significant wetland community because only a few are known east of the

Front Range, although diverse types occur commonly in higher elevations. Fens are characterized by a deep,

subsurface peat layer (Culver & Lemly 2013) and abundant Carex simulata. A C. simulata matrix is character¬

istic of higher elevation fens and indicative of peatland development (Culver and Lemly 2013). The largest

Black Squirrel Creek fen covered ca. 400 m 2 in a subsidiary drainage north of the main channel below a large

open seep. It was characterized by Carex simulata mats with occasional occurrences of Eleocharis acicularis,

Carex nebrascensis, and Schoenoplectus pungens. The few forbs present included Helenium autumnale and Par¬

nassia palustris. Soils were highly saturated and visibly quaked when stepped upon; the underlying peat layer

was over a meter thick. Smaller apparent fens where the vegetation was dominated by C. simulata occurred

sporadically in subsidiary wet drainages.

Ponds .—A number of small ponds occurred throughout the drainages. These are generally less than 9.3

m 2 (ca. 100 ft. 2 ) in area, with a depth of 0.3 m (1 ft) to over 1.5 m (4 ft), depending on precipitation. The ponds

supported abundant amphibians, aquatic insects, crustaceans, and other larger vertebrates such as minnows,

along with diverse floating and emergent plant species. Pond associations included an emergent Typha marsh

community, and a floating aquatic community with the carnivorous species Utricularia minor in shallow

ponds, along with more common aquatics such as Sagittaria, Alisma, Sparganium, and Potamogeton.

Vegetation transects

Vegetative and Graminoid Cover. —Transects were situated primarily on moist banks and moist shelves, al¬

though they also encompassed Nebraska sedge meadows and bogs as well as open seeps (Fig. 2). They varied

in their vegetative cover and the relative cover of graminoids, forbs, or litter/bare ground (Table 3; Fig. 4). On

average, total basal vegetative cover was slightly over 50%, and graminoids constituted almost 80% relative

cover. The dominant grass contributors to cover (in order of prominence: Sporobolus heterolepis, Muhlenbergia

montana, Schizachyrium scoparium, Sorghastrum nutans, and Calamovilfa longifolia; Table 4) parallelled key

components documented by Livingston (1952). Common but lesser contributors in both studies included An-

dropogon gerardii, Hesperostipa spartea, and Panicum virgatum.

Variation among the transect quadrats reflected the characteristic local heterogeneity of patchy clumps of

vegetation interspersed with open soil. Transect 1 was the most hydric, with abundant subsurface water and a

small seep dominated by Juncus arcticus. It had the highest overall cover, primarily Sorghastrum nutans, Spo¬

robolus heterolepis, and Schizachyrium scoparium, with lesser components of Muhlenbergia montana, Bouteloua

gracilis, Calamovilfa longifolia, Panicum virgatum, and Koeleria macrantha.

Although adjacent to transect 1, transect 2 showed somewhat different community structure where the

Kelso et al., Flora of tallgrass prairie and wetlands. Black Squirrel Creek drainage

215

Table 3. Comparison of total vegetative cover and relative cover of dominant graminoid species, 2011 and Livingston studies of the 1940s. T1-T4 represent 2011

transects. L1-L7 represent data from Livingston (1947, 1952; 1952 transect numbers modified from those used in 1947).

T 4

Mean All Transects

Total % Vegetative

Cover

63.8

58.8

46.5

43.8

53.2

Relative % Cover

69.5

77.7

86.5

85.7

79.9

All Graminoids

Dominant Grasses

Sorghastrum nutans

Sorghastrum

Sporobolus

Muhlenbergia

Sporobolus

(% Relative Cover)

(12)

nutans (25.3)

heterolepis (27.1)

montana (36.8)

heterolepis (15.1)

Sporobolus heterolepis

Schizachyrium

Calamovilfa

Sporobolus

Muhlenbergia

(9)

scoparium (20.2)

longifolia (20)

heterolepis (18.7)

montana (13.2)

Schyzachyrium

Muhlenbergia

Andropogon

Schizachyrium

scoparium (7.5)

montana (8.9)

gerardii (12)

scoparium (17.5)

scoparium (11)

Bouteloua sp. (4.5)

Calamovilfa

Bouteloua gracilis

Hesperostipa

Sorghastrum

longifolia (7.4)

(8.9)

spartea (3.9)

nutans (10.7)

Muhlenbergia

Sporobolus

Muhlenbergia

Sorghastrum

Calamovilfa

montana (4.2)

heterolepis (7.2)

montana (4.9)

nutans (3)

longifolia (7.8)

Calamovilfa longifolia

Bouteloua gracilis

Schizachyrium

(3.2)

(1.3)

scoparium (0.9)

Panicum virgatum

Hesperostipa

(2.8)

spartea (1.2)

Koeleria macrantha

(0.7)

Nasella viridula (1 )

Forest

Mean

Dominant Grasses

Poa pratensis (31.4)

Bouteloua gracilis

Sporobolus

(% Relative Cover)

(14.2)

heterolepis (33.9)

heterolepis (73.4)

heterolepis (36.7)

Sporobolus heterolepis

Poa compressa

Muhlenbergia

Poa pratensis (4.6)

Poa pratensis (9)

(29.1)

(10.5)

montana (9.3)

Muhlenbergia

Sporobolus

Bouteloua gracilis

montana (4.9)

heterolepis (1 0.1)

(6.8)

(5.3)

Muhlenbergia

montana (3.5)

Plains

L7U/M/L

Mean

Dominant Grasses

Andropogon gerardii

Sporobolus

Schizachyrium

Sporobolus

(% Relative Cover)

(28.2)

heterolepis (30.7)

scoparium

(47.2/28.7/55.8)

heterolepis (35)

Bouteloua

Schizachyrium

Bouteloua gracilis

Schizachyrium

gracilis (19.8)

scoparium (25.3)

(31.0/4.9/15)

scoparium (31.4)

Sporobolus heterolepis

Elym us

Calamovilfa

Bouteloua

(12.5)

trachycaulus (8.0)

longifolia

(10.6/0/0.8)

gracilis (15)

Calamovilfa

Andropogon

Sorghastrum

Calamovilfa

longifolia (8.1)

gerardii (5.2)

nutans

longifolia (7.6)

(0.3/30.7/2.1)

Andropogon

gerardii (6.7)

microtopography of the terraces contributed to variable soil moisture and texture. This transect was also

dominated by Sorghastrum nutans, contributing ca. 25% of the vegetative cover, with Schizachyrium scoparium

contributing an additional 20%. Sporobolus heterolepis contributed considerably lesser cover here (7%). In gen¬

eral, this transect showed a greater presence of more xeric elements such as Muhlenbergia montana and Cal-

amovilfa longifolia than transect 1 above it, and somewhat less overall cover.

Transect 3, in a side drainage adjacent to the main channel, had comparatively lower cover than transects

1 and 2, less than 50% overall; 90% of this cover was composed of grasses dominated by Sporobolus heterolepis,

Calamovilfa longifolia and Andropogon gerardii. Together these species accounted for ca. 59% relative cover. This

was the only transect in which A. gerardii appeared to a notable extent, contributing 12% relative cover.

216

Journal of the Botanical Research Institute ofTexas 8(1)

100 %

80%

60%

40%

20 %

0 %

Ground Cover

H % Graminoid Cover

H% Forb Cover

□ % Litter Cover

■ % Bare Ground

Fig. 4. Percent ground cover of graminoids, forbs, litter and bare ground for 2011 transects. T1 -4 refers to transect numbers as described in the text.

Table 4. Comparison of the highest frequency grasses in the Livingston (1947; 1952) study and 2011.T1—T4 are 2011 transects; LI, L4, L5, and L6 are Livingston data

from 1947, republished in 1952. No frequency data were given for transects L2, L3, and L7. Livingston stations L1-L4 were located in the Black Forest ("forest stations"

sensu Livingston) and stations L5-L6 were"plains"transects. 2011 transects were located midway in distance and elevation between the forest and plains transects.

Species

Mean %F in

2011 (rank)

Mean %F-

forest 1952

(rank)

Mean %F -

plains 1952

(rank)

Andropogon gerordii

9(8)

10(6)

50(4)

Bouteloua gracilis

15(6)

100

65(2)

Calamovilfa longifolia

35(4)

60(3)

Hesperostipa spartea

11(7)

25 (5)

Koeleria macrantha

4(9)

30(4)

20(6)

Muhlenbergia montana

43(1)

50(3)

20 (6)

Panicum virgatum

3(10)

20(6)

Poa pratensis

9(8)

60(2)

Schizachyrium scoparium

38(2)

5(7)

93(1)

Sorghastrum nutans

36(3)

23(5)

Sporobolus heterolepis

34(5)

100

85(1)

100

60(3)

In transect 4, located in a large subsidiary drainage with a perennial secondary stream, vegetative cover

was also less than 50%, and composed primarily of Muhlenbergia montana, Sporobolus heterolepis, and

Schizcichyrium scoparium; together these species comprised 73% of the graminoid cover. Hesperostipa spartea

and Sorghastrum nutans were more frequent here than in the other transects, although they contributed rela¬

tively little to cover. Both species are regionally uncommon to rare (but not considered rare statewide), and

when present, they typically occur in disparate clumps. This characteristic pattern was apparent in transect 4,

Kelso et al., Flora of tallgrass prairie and wetlands. Black Squirrel Creek drainage

217

and underscored the decoupled metrics of a species contribution to cover from overall frequency of occurrence

for individual species in these communities.

Grass Species Frequency Rankings

Across all transects, the most common species (Table 3) were Muhlenbergia montana (frequency of ca. 42% of

all quadrats), Schizachyrium scoparium (37% frequency), Sorghastrum nutans (36% frequency), Calamovilfa

longifolia (35% frequency), Sporobolus heterolepis (34% frequency), and Juncus arcticus (32% frequency). Grass

taxa of second tier frequency included Hesperostipa spartea (16% frequency), Bouteloua gracilis (15% frequen¬

cy), Andropogon gerardii (9% frequency), and Poa pratensis (9% frequency). The highest frequency non-grami-

noid taxa (data not shown) included a high representation of Midwest prairie elements, notably Artemisia

ludoviciana, Dalea purpurea, Equisetum arvense, Glycyrrhiza lepidota, Helenium autumnale, Oligoneuron rigidum,

Rosa arkansana, Symphotrichum ericoides, and Symphotrichum laeve. These species were common components

of the regional mixed grass prairie; only Helenium autumnale is an elevational disjunct more common in the

Foothills/Montane zone than on the plains.

DISCUSSION

Comparison of Grassland Vegetation Structure: 1940s and 2011

Both studies document considerable variation within and among transects and the overall means cover wide

ranges at both time frames (Table 3). We found considerably higher basal vegetative cover (typically over 50%)

than what Livingston reported (typically less than 20%). In 2011, graminoids constituted ca. 80% of this cover,

while on the Livingston forest-grassland plots, 85% of this cover was constituted by graminoids, and on the

plains plots, 55%. The dominant grasses were similar, with Sporobolus heterolepis being the top contributor to

cover in both studies. In 2011, Muhlenbergia montana, Schizachyrium scoparium, Sorghastrum nutans, and Cal¬

amovilfa longifolia were also important components; jointly these provided 60% relative cover. In the Livings¬

ton study, in addition to S. heterolepsis (55% relative cover), Poa pratensis, Bouteloua gracilis, and M. montana

contributed most highly on the forest transects while S. scoparium, B. gracilis, C. longifolia, and A. gerardii

played key secondary roles on the plains (95% relative cover).

Comparison of the frequency of species occurrence (Table 4) provides a similar picture of a consistent

suite of species common to both time frames, albeit with different rankings for individual frequencies. Muhlen¬

bergia montana, Schizachyrium scoparium, Sorghastrum nutans, Calamovilfa longifolia, and Sporobolus heterole¬

pis were the most commonly occurring species in 2011, each present in over 30% of the quadrats. On the Liv¬

ingston forest plots, S. heterolepis, Poa pratensis, M. montana, and Koeleria macrantha were most frequently en¬

countered, while on the plains, S. scoparium, B. gracilis, S. heterolepis, C. longifolia, and A. gerardii each occurred

in at least 50% of the quadrats.

The 2011 structure of the tallgrass vegetation of Black Squirrel Creek remained strongly comparable to

what Livingston described in the 1940s with respect to the most frequent grass species and those that contrib¬

uted the most cover. Key species in the 1940s (in particular, Sporobolus heterolepis, Muhlenbergia montana,

Schizachyrium scoparium, and Sorghastrum nutans) remained important now. Andropogon gerardii seems to

have diminished in both frequency and contribution to cover since the 1940s, although even then it was not a

dominant component of the vegetation. However, because the Livingston transects could not be precisely relo¬

cated, our transect positions did not match his sample sites, and at least some of these differences may be arti¬

facts of different transect positions. Other differences may result from limited sampling in a landscape mosaic

where topography, edaphic factors, and at least to some extent, current and past grazing practices, create

patchy vegetation or bare ground. Because our plots had light grazing and no mowing (in comparison to the

Livingston plots that were subjected to grazing and seasonal mowing), this may have allowed for greater cover

to develop in some areas. Alternatively, these differences may reflect real changes in species abundance. Pos¬

sible considerations include recovery lag from the 1930s Dust Bowl decade that might have exerted a lingering

effect during the 1940s, or the converse, when several substantial drought episodes in the intervening decades

since the Livingston work may have influenced the response and recovery of individual species.

218

Journal of the Botanical Research Institute of Texas 8(1)

Although Livingston’s characterization of Midwestern grasslands as “true prairie” may be arguable, his

analogy of the Black Squirrel Creek vegetation to the Midwestern grasslands was then, and remains now, ap¬

propriate with respect to the major grasses present and their relative contributions to cover, as well as to the

forb and shrub elements in the flora and vegetation structure. However, this vegetation is not identical to

iconic tallgrass associations of the central prairies and identified elsewhere in Colorado, where Andropogon

gerardii is the signature species with the highest cover and frequency (Moir 1972; Bock & Bock 1998; Neid et

al. 2009). In the Black Squirrel Creek associations, while A. gerardii never contributed highly to cover, it oc¬

curred relatively frequently. The most prominent grass was Sporobolus heterolepis, occurring with foothills/

montane species such as Muhlenbergia montana and xeric or mixed grass prairie species such as Calamovilfa

longifolia and Bouteloua gracilis. The Black Squirrel Creek vegetation may be better described as Sporobolus

heterolepis-Muhlenbergia montana grasslands with subsidiary components of Calamovilfa longifolia, Schizachy-

rium scoparium, or Sorghastrum nutans rather than the more classical model of tallgrass communities defined

by the dominance of A. gerardii. No comparable associations dominated by Sporobolus heterolepis are currently

listed by NatureServe (2013), or documented in Colorado, although Weaver (1954) described prairie dropseed

communities as a distinctive, albeit minor, component of xeric upland prairie on the midwestern plains.

The Wetland-Grassland Mosaic

We concur with the conclusion reached by Livingston (1952) specifically for the Black Squirrel Creek/Black

Forest area, and Branson et al. (1965) more broadly for the Colorado mountain front, that adequately high soil

moisture is the key factor responsible for the occurrence of mesic tallgrass prairie. As transitional zones be¬

tween aquatic and terrestrial habitats, wetlands play vital roles in linking ecological and hydrological systems

(Culver and Lemly 2013) and are widely recognized for having high biological significance due to their variety

of biodiversity and community types; this is particularly true in the arid Arkansas River drainage of Pueblo and

El Paso Counties (Doyle et al. 2001b) where few wetlands presently occur. In the Black Squirrel Creek drain¬

age, the wetlands were a critical part of the vegetation mosaic of hydric and mesic communities and entwined

biota. As noted in other prairie systems (Semlitsch and Bodie 1998; Leibowitz 2003), even small and superfi¬

cially discontinuous imbedded wetlands support metapopulation, corridor, seasonal or annual habitat dy¬

namics for flora and fauna by connecting subsurface geology and hydrology. In the Black Squirrel Creek drain¬

age, this connectivity sustains the anomalous vegetation and flora and is certainly part of their longevity.

A conclusion of botanical stasis along Black Squirrel Creek is not appropriate, however. Change has oc¬

curred, perhaps with respect to change in frequency of certain species, but certainly visibly with respect to

extent of these grasslands, which now exist as remnants reduced in both number and size since the 1940s, and

even by then probably reduced from their former extent prior to extensive ranching (Livingston 1952). Where

widespread mesic grasslands once prevailed, housing developments and infrastructure or altered grasslands

with adventive, grazing- tolerant, or xeric species sometimes now dominate. In spite of these alterations and

diminished extent of communities flagged as noteworthy 70 years ago, the Black Squirrel Creek drainage re¬

mains a remarkable center of biotic diversity increasingly significant for its numerous rare or uncommon spe¬

cies, and its elevational and longitudinal disjuncts. Its unique grasslands show clear affiliation to their geo¬

graphically and temporally distant cousins that are not structurally identical but nonetheless strongly conspe-

cihc in their floristic profiles.

Although it remains speculative that these communities are in situ Pleistocene relicts, they have endured

substantial climatic vicissitudes. In the past century, interannual droughts have occurred regularly, and severe

droughts within the context of these comparative studies show little apparent major impact in areas directly

supported by ground water. Where this vegetation remains, the Black Squirrel Creek drainage testifies to the

capacity of hydrogeomorphic systems to sustain relative stasis in prairie plant communities and their constitu¬

ent flora through fluctuating temperature and moisture regimes. Whether this system can be self-sustaining

through accelerating anthropogenic and climatic pressures remains an open question and concern.

Kelso et al., Flora of tallgrass prairie and wetlands. Black Squirrel Creek drainage

219

APPENDIX 1

ANNOTATED CHECKLIST OF THE VASCULAR FLORA

UPPER BLACK SQUIRREL CREEK DRAINAGE, EL PASO CO., COLORADO

Species list for vascular flora occurring in mesic and hydric communities of the upper Black Squirrel Creek

drainage from 7000-6500 feet in elevation. The list includes some upland species occurring sporadically in the

drainage system and gravel stream channels, but which are more typically found in the surrounding xeric

mixed grass and shortgrass matrix in the surrounding uplands. Nomenclature (including family designations)

and common names follow the National Resource Conservation Service database (www. Plants.USDA.gov:

accessed 8/2013) except as noted. Nomenclature used in Colorado (e.g., Weber & Wittmann 2012) is given in

curly braces as {name} prior to the common name. Alternative Angiosperm Phylogeny Group familial designa¬

tions (APG; www.mobot.org/MOBOT/Research/APweb) are included for each family. Voucher specimens for

all taxa are at COCO. Noteworthy species are coded as follows:

FM Species generally occurring in Foothills/Montane zone 2000-3200 m (ca. 7000-10,500 ft). Distributions

follow Weber and Wittman 2012; Culver and Lemly 2013; Kelso, 2012); occurrence on the prairie is re¬

stricted.

MWP Species prominent in Midwest prairie as explicitly noted by Shantz (1923), Weaver and Fitzpatrick

(1934: Tables 15; 16), Weaver (1954), and Livingston (1952). Earlier nomenclature used in these publica¬

tions cross-referenced with NRCS Plants Database synonyms.

R Rare species listed as of conservation concern and tracked by the Colorado Natural Heritage Program (2012):

State Conservation rankings are as follows: SI: Critically Imperiled, S2: Imperiled, S3: Vulnerable

U Locally or regionally uncommon or local endemic (Kelso 2012; Weber & Wittmann 2012)

X Xeric element of uplands extending into mesic vegetation

L indicates a species noted in the Livingston studies; square brackets indicates a species observed but not recol¬

lected in our study. Unless otherwise noted, these are locally common components of the xeric plains

flora. Livingston species not observed are so indicated.

Species with no coding are locally widespread components of regional vegetation.

Amaranthaceae

Froehlichia gracilis (Hook.) Moq. (slender snakecotton) X

Alismataceae

Sagittaria cuneata Sheldon (arumleaf arrowhead)

Sagittaria latifolia Willd. (broadleaf arrowhead)

Alisma triviale Pursh (northern water plaintain)

Anacardiaceae

Toxicodendron rydbergii (Small) Greene (poison ivy) FM, MWP

Apiaceae

Cicuta douglasii (D.C.) J.M. Coulter & Rose (western water hemlock)

MWP

Berulaerecta (Hudson) Coville (cutleaf water parsnip)

Apocynaceae

Apocynum cannabinum L. (Indianhemp)

Asdepiadaceae (APG Apocynaceae)

Asclepias hallii A. Gray (Hall's milkweed) R (S3), FM

Asclepiasspeciosa Torr. (showy milkweed)

Asteraceae

Achillea millefolium L. (common yarrow) L

Agoseris glauca (Pursh) Raf. (pale agoseris)

[Ambrosia artemisiifolia L. (annual ragweed) L]

Ambrosiapsilostachya DC. (Cuman ragweed) X

Antennaria microphylla Rydb. (littleleaf pussytoes) X; L as A.

parviflora

[Artemisia campestris L. ssp. caudata (Michx.) Hall & Clements (field

sage wort) L]

Artemisia frigida Willd. (prairie sagewort) X, L

Artemisia ludoviciana Nutt, (white sagebrush) MWP, L as

A.gnaphalodes

Bahia dissecta (A. Gray) Britton (ragleaf bahia) X

Bidens tenuisecta A. Gray (slimlobe beggarticks)

Carduus nutans L. (nodding plumeless thistle) Noxious weed of

limited occurrence on the study site; few individuals.

Cirsium arvense (L.) Scop. [Breea arvense} (Canada thistle) Noxious

weed of limited occurrence on the study site

Cirsium flodmanii (Rydb.) Arthur (Flodman's thistle)

Conyza canadensis (L.) Cronquist (Canadian horseweed) L as

Leptilon canadensis

Cosmos parviflorus (Jacq.) (southwestern cosmos) U Southwestern

species known primarily as regional endemic to the Black Forest

region; locally common along Black Squirrel Creek on gravelly

stream channels.

Erigeron bellidiastrum Nutt, (western daisy fleabane)

Erigeron compositus Pursh (cutleaf daisy) FM

Erigeron divergens Torr. & A. Gray (spreading fleabane)

Erigeron flagellaris A. Gray (trailing fleabane)

Erigeron glabellus Nutt, (streamside fleabane)

Erigeron lonchophyllus Hook. [Trimorpha lonchophylla} FM

Erigeron subtrinervis Rydb. ex Porter & Britton (threenerve fleabane)

Erigeron vetensis Rydb. (early bluetop fleabane) FM

Grindelia squarrosa Dunal (curlycup gumweed)

Helenium autumnale L. (common sneezeweed) FM, MWP, L

Helianthus annuus L. (common sunflower)

Helianthus nuttallii Torr. & A. Gray (Nuttall's sunflower)

Helianthus petiolaris Nutt, (prairie sunflower) L

220

Journal of the Botanical Research Institute of Texas 8(1)

Helianthuspumilus Nutt, (little sunflower)

Helionthus pouciflorus Nutt. ssp. (Rydb.) O. Spring. & E. Schilling

{Helianthus rigidus} (stiff sunflower) MWP, L

Heterotheca canescens (D.C.) Shinners (hoary false goldenaster) L

as Chrysopsis villosa

Lactuca tatarica (L.) Meyer (blue lettuce)

Liatris ligulistylis (A. Nelson) K. Schum. (Rocky Mountain blazingstar)

R(S1/S2),MWP, LCommon in seeps and tallgrass communities

on the study site.

Liatris punctata Hook, (dotted blazingstar) MWP, L

Lygodesmiajuncea (Pursh) D. Don ex Hook, (rush skeletonplant)

Oligoneuron album (Nutt.) G.L. Nesom { Unamia alba} (prairie gold¬

en rod) R (S2/S3), MWP, L

Oligoneuron rigidum (L.) Small (stiff goldenrod) MWP, L as Solidago

rigidum

[Packera neomexicana (A. Gray) W.A. Weber & A. Love var. mutabilis

(Greene) W.A. Weber & A. Love (New Mexico groundsel) L as

Senecio mutabilis )

Packera pseudaurea (Rydb.) W.A. Weber & A. Love (falsegold

groundsel) FM

Packera tridenticulata (Rydb.) W.A. Weber & A. Love (threetooth

ragwort)

Pseudognaphalium canescens (D.C.) W.A. Weber (Wright's cudweed)

Ratibida columnifera (Nutt.) Woot. & Standi, (upright prairie

co nef lower)

Rudbeckia hirta L. (black eyed Susan) FM, MWP, L

[Salsola tragus L. (prickly Russian thistle) L as Salsola pestifer]

Schkuhria multiflora Hook. & Arn. (manyflower false threadleaf) X

Senecio spartioides Torr. & A. Gray (broomleaf ragwort)

Solidago gigantea Aiton (giant goldenrod)

Solidago missouriensis Nutt. (Missouri goldenrod) MWP, L

Solidago nana Nutt, (baby goldenrod) FM

Solidago nemoralis Aiton (gray goldenrod) L

Solidago velutina D.C. (three-nerve goldenrod)

Symphotrichum ericoides (L.) A. Love & D. Love (white heath aster)

MWP, L as Aster multiflorus

Symphotrichum laeve (L.) A. Love & D. Love (smooth aster) FM,

MWP, L as Aster geyeri

Symphotrichum lanceolatum (Willd.) G.L. Nesom (white panicle

aster)

[Thelesperma megapotamicum (Spreng.) Kuntze (Hopi tea green-

thread) L as Thelesperma gracile]

Tragopogon dubius Scop, (yellow salsify)

Tripleurospermum perforatum (Merat) M. Lainz (scentless false

marigold) FM

Tetraneuris acaulis (Pursh) Greene (stemless four-nerve daisy)

Boraginaceae

Cryptantha cinerea (Greene) Cronquist { Oreocarya suffruticosa}

(James'cryptantha) X [Lappula occidentalis (S. Watson) Greene

(flatspine stickweed) L]

Mertensia lanceolata (Pursh) D.C. (prairie bluebells)

Onosmodium bejariense DC var. occidentale (Mack.) B.L. Turner

{Onosmodium molle ssp. occidentale} (softhair marbleseed)

Plagiobothrysscouleri I.M. Johnst. (Scouler's popcorn flower)

Brassicaceae

Arabis holboelii Hornem. var. retrofracta Rydb. {Boechera retrofracta}

(second rockcress) FM

Barbarea orthoceras Ledeb. (American yellowrocket)

Draba nemorosa L. (woodland draba) FM

Sisymbrium loeselii L. (small tumbleweed mustard)

Cactaceae

Pediocactus simpsonii (Engelm.) Britton & Rose (mountain ball

cactus) FM

Opuntia polyacantha Haworth (plains prickly pear) L

Campanulaceae

Campanula rotundifolia L. (bluebell bellflower) FM

Lobeliasiphilitica L. (great blue lobelia) U, MWP, L Common on the

study site; regionally uncommon species

Caprifoliaceae

Symphoricarpos occidentalis Hook, (western snowberry)

Caryophyllaceae

Arenaria hookeri Nutt. {Eremogone hookeri} (Hooker's sandwort)

Stellaria longifolia Muhl. ex. Willd. (longleaf starwort) FM

Paronychia jamesii Torr. & A. Gray (James' nail wort) X

[Silene scouleri Hook, (simple campion) L] Species common in

the foothills/montane zone but not currently known from

this location

Chenopodiaceae (APG Amaranthaceae)

Chenopodium graveololens Willd. {Teloxis graveolens} (fetid goose-

foot)

Chenopodium leptophyllum (Moq.) Nutt, (narrowleaf goosefoot), L

Cycloloma atriplicifolium (Spreng.) J.M. Coulter (winged pigweed)

Suaeda calceoliformis (Hook.) Moq. {Suaeda depressa} (Pursh

seepweed)

Clusiaceae

Hypericum scouleri Hook. {Hypericaceae: Hypericum formosum}

(Scouler's St. Johnswort) FM

Commelinaceae

Tradescantia occidentalis (Britton) Smythe (prairie spiderwort)

Crassulaceae

Sedum lanceolatum Torr. (spearleaf stonecrop)

Cyperaceae

Carexaurea Nutt, (golden sedge) FM, U, L

Carexbrevior (Dewey) Mack, (shortbeak sedge) L

Carexcrawei (Dewey) (Crawe's sedge) R (SI), FM

Carexdisperma Dewey (softleaf sedge) FM

Carexdouglasii Boot (Douglas'sedge)

Carexechinata Murray {Carex angustior}[star sedge) FM, U

{Carex filifolia Nutt, (threadleaf sedge) L]

{Carexheliophila Mack. = C. inops L.H. Bailey ssp. heliophila (Mack.)

Crins (sunsedge) L]

{*Carex oreocharis T. Holm (grassyslope sedge) L- not observed

this study]

Carexpellita Muhl. {Carexlanuginosa} wooly sedge

Carexnebrascensis Dewey (Nebraska sedge) L

{*Carex hallii Olney [C. parryana Dewey ssp. hallii [specimen coll.

R.B. Livingston 1430: @COCO;] (deer sedge)]

{*Carexpraegracilis\N. Boott) L-not observed this study]

Carexsimulata Mack, (analogue sedge)

Carexxerantica L.H. Bailey (whitescale sedge)

Eleocharis acicularis (L.) Roemer & Schultes (needle spikerush)

Eleocharis obtusata (Willd.) Schult. (blunt spikerush)

{Eleocharispalustris (L.) Roem. & Schult. (common spikerush) L]

Eleocharis quinqueflora (Hartmann) O. Schwartz (fewflower spik¬

erush) FM

Cyperus schweinitzii Torr. (Schweinitz' flatsedge) {Mariscus sch-

weinitzii}

Schoenoplectus acutus (Muhl.) A. Love & D. Love {Scirpus acutus}

(hardstem bulrush)

Schoenoplectuspungens (Vahl) Palla (common threesquare)

Schoenoplectus tabernaemontani (C.C. Gmelin) Palla (softstem

bulrush) {Scirpus lacustris}

Scirpus microcarpus Presl. & C. Presl (panicled bulrush)

Kelso et al., Flora of tallgrass prairie and wetlands. Black Squirrel Creek drainage

221

Equisetaceae

Equisetum orvense L. (field horsetail) MWP, L

Equisetum laevigata A. Brown (smooth horsetail) MWP

Euphorbiaceae

Chamaesyce glyptosperma (Engelm.) Small (ribseed sandmat) Las

Euphorbia glyptosperma

Euphorbia brachycera Engelm. (horned spurge) X

Fabaceae

Amorpha fruticosa L. var. angustifolia Pursh (false indigo bush)

Astragalus canadensis L. (Canadian milkvetch) U, MWP

Dalea Candida Michx. exWilld. (white prairieclover) MWP

Dalea purpurea\/ent. (purple prairieclover) MWP, Las Petalostemon

purpureus

Gleditsia triacanthos L. (honeylocust) A single tree occurring on

edge of study site near old ranch buildings.

Glycyrrhiza lepidota Pursh (American licorice) MWP, L

Lathyrus polymorphus Nutt, (manystem pea)

Lupinus pusillus Pursh (rusty lupine)

[Melilotus officinalis (L.) Lam. (sweetclover) L as M. alba]

Oxytropis multiceps Nutt. (Nuttall's oxytrope)

Robinia neomexicana A. Gray (New Mexico locust) Occasional trees

occurring on edges of study site near old ranch buildings.

[Thermopsis montana Nutt, (mountain goldenbanner) L]

[Trifolium pratense L. (red clover) L]

[Trifolium repens L. (white clover) L]

[Vicia americana Muhl. ex Willd. (American vetch) L]

Gentianaceae

[Gentiana affinis Griseb. (pleated gentian) L]

Gentianella amarella (L.) Borner ssp. acuta (Michx.) Gillette [Gen-

tianella strictiflora} (autumn dwarf gentian) FM, L As noted

by Weber and Wittmann (2012), the densely white flowered

form with a stiffly erect inflorescence is very distinctive in this

region in the montane zone and in higher elevations on the

plains; it is easily recognized as separate from the amarella/

acuta form. The form occurring in the Black Squirrel Creek

region is the"strictiflora"form, rather than the purple flowered,

smaller"acuta"form.

Gentianopsis virgata (Raf.) Holub [Gentianopsis procera ssp. crinita; G.

crinita } [lesser fringed gentian] R (CHNP-Not rated), MWP This

species, recently confirmed by Flora of North America experts

in the genus, is only known to occur in the upper Black Squirrel

Creek drainage. Although not yet listed by Colorado Natural

Heritage Program, its state rarity and disjunct connection to

the Midwest prairie flora is notable.

Geraniaceae

Geranium atropurpureum A. Heller. {G. caespitosum ssp. atropurpu-

reum}(western purple crane's bill) FM

Grossulariaceae

Ribesaureum Pursh (golden currant)

Haloragaceae

Myriophyllum sibiricum Kom. (shortspike watermilfoil)

Hippuridaceae (APG Plantaginaceae)

Hippuris vulgaris L. (common mare's tail) FM

Hypoxidaceae

Hypoxis hirsuta (L.) Coville (common goldenstar) R (SI), MWP

Iridaceae

Iris missouriensis Nutt, (wild iris)

Sisyrinchium montanum Greene (strict blue eyed grass) FM, L as

S. angustifolium

Juncaceae

Juncus alpinoarticulatus Chaix (northern green rush)

Juncus arcticus Willd. ssp. littoralis (Willd.) Hulten [J. arcticus ssp.

ater] (mountain rush), Las J. balticus

Juncus brachycephalus (Engelm.)Buchenar (smallhead rush) R(S1),

MWP, L

*Juncus brevicaudatus (Engelm.) Fernald: narrowpanicle rush R

(SI); MWP A specimen of this species under the name J.

brachycephalus {Penland4935; COCO; OSH) was collected by in

the Black Squirrel Creek drainage) and later verified by N. Har-

riman and F. Herrmann (Herrmann, 1975) as the very similar J.

brevicaudatus. We have tentatively identified one of ou r col lec¬

tions as this species. It grows intermixed with J. brachycephalus.

Juncus bufonius L (toad rush)

Juncus dudleyi\N\eg. (Dudley's rush)

Juncus interior\N\eg. (inland rush)

JuncuslongistylisTorr. (longstyle rush) L

Juncus marginatus Rostk. (grassleaf rush)

Juncus nodosus L. (knotted rush)

Juncus saximontanus A. Nelson (Rocky Mountain rush)

Juncus torreyi Coville (Torrey's rush) L

Juncaginaceae

Triglochin maritima L. (seaside arrowgrass) L

Triglochin palustris L. (marsh arrowgrass) FM

Lamiaceae

Lycopus americanus Muhl. ex Bartram (American water horehound)

MWP

Mentha arvensis L. (wild mint)

Monarda fistulosa L. (wild bergamot) MWP

Prunella vulgaris L. (common selfheal) L

Scutellaria galericulata L. (marsh skullcap) U

Stachys palustris L. (marsh hedgenettle)

Lemnaceae

Lemna minor L. (common duckweed)

Lentibulariaceae

Utricularia minor L. R(S2), FM

Liliaceae s.l. (APG Amaryllidaceae)

Allium cernuum Roth (nodding onion) [Alliaceae] L

Calochortusgunnisonnii S. Watson (Gunnison's sego lily)

Liliumphiladelphicum L. (wood lily) R(S1), FM, MWP

Lythraceae

Lythrumsalicaria L. (purple loosestrife) This A list noxious weed has

invaded a subsidiarydrainageof BlackSquirrel Creek, and is the

only significant weed issue. It currently is not a monoculture,

and many of the rare and unusual species are intermixed with

it, making chemical controls problematic.

Malvaceae

Sidaicea neomexicana A. Gray (saltspring checkerbloom) U, FM

Najadaceae (APG Hydrocharitaceae)

Najas guadalupensis (Spreng.)Magnus (spring water nymph)

Nyctaginaceae

Abronia fragrans Nutt, ex Hook, (snowball sand verbena) X

Mirabilis linearis (Pursh)Heimerl [Oxybaphus lanceolatus; Oxybaphus

linearis}{ narrowleaf four o'clock)

Oleaceae

Forestierapubescens Nutt. [Forestiera neomexicana} (stretchberry)

Onagraceae

Calylophusserrulatus (Nutt.) P. H. Raven (yellow sundrops)

Kelso et al., Flora of tallgrass prairie and wetlands. Black Squirrel Creek drainage

223

Potentilla norvegico L. (Norwegian cinquefoil)

Potentillaplattensis Nutt. (Platte River cinquefoil)

Prunus pumila L. var. besseyi (L.H. Bailey) Gleason { Prunus besseyi;

Cerosuspumiloj (western sandcherry)

Prunus virginiana L. { Padus virginiana} (chokecherry) FM

Roso orkonsono Porter (prairie rose) MWP

Rubiaceae

Galium boreale L. { Galium septentrionale} (northern bedstraw) FM

Galium trifidum L. (threepetal bedstraw) MF

Salicaceae

Populus x acuminata Rydb. (lanceleaf cottonwood)

Populus angustifolia James (narrowleaf cottonwood) FM

Populus deltoides Bartram ex Marsh, (plains cottonwood)

Salixamygdaloides Andersson (peachleaf willow)

Salixexigua Nutt, (narrowleaf willow)

Salixirrorata Andersson (deweystem willow) FM

Salixligulifolia (C.R. Ball) C.R. Ball (strapleaf willow)

Santalaceae

Comandra umbellata (L.) Nutt, (bastard toadflax) X, L as C. pallida

Saxifragaceae

Pamassiapalustris L. var. parviflora (DC) Boivin {Parnassiaparviflora }

(smallflower grass of Parnassus) FM

Selaginellaceae

Selaginella densa Rydb. (lesser spikemoss) FM

Scrophulariaceae (s. lat.; APG see also Callitrichaceae,

Orobanchaceae, Phrymaceae)

Mimulusglabratus Kunth (roundleaf monkeyflower)

Nuttallanthus canadensis (L.) D.A. Sutton {Linaria canadensis }

(Canada toadflax) U

Orthocarpus luteus Nutt, (yellow owlclover) L

Pedicularis canadensis L. (Canadian lousewort) FM

[.Penstemon angustifolius Nutt. ex. Pursh (broadbeard beardtongue)

Penstemonauriberbis Pennell (Colorado beardtongue) XThis species

is extremely common south in the Arkansas River drainage but

occurs only sporadically around Colorado Springs.

Penstemon glaber Pursh (sawsepal pentstemon) FM

Penstemon gracilis Nutt, (lilac penstemon )U, FM, MWP Species

uncommon in the foothills and pine savannah regions of the

Black Forest area; found once on our study site.

Penstemon virgatus ssp. asa-grayi Crosswhite (one-sided pen¬

stemon)

Veronica anagallis-aquatica L. (water speedwell) { Veronica catenata}

Veronica peregrina L. (neckweed) L

Veronica serpyllifolia L. ssp. humifusa (Dicks.) Syme (thymeleaf

speedwell) FM

Solanaceae

Physalis virginiana Mill. (Virginia groundcherry)

Sparganiaceae (APG Typhaceae)

Sparganium angustifolium Michx. (narrowleaf bur-reed) FM

Typhaceae

Typha latifolia L. (broadleaf cattail)

Valerianaceae (APG Caprifoliaceae)

Valeriana edulis Nutt. exTorr. & A. Gray (tobacco root) FM

ACKNOWLEDGMENTS

We are deeply grateful to the Lee and Le Compte families for allowing us access to develop this research on

their extensive ranches. In particular, we thank W. Tracy Lee for his exceptional stewardship of 4-Way Ranch

over multiple decades that has preserved this remarkable area. The original Livingston theses were retrieved

from the Duke University archives through the efforts of N. Margolin. Renee Rondeau, D. Culver, andj. Lemly

provided much assistance in delineating and assessing the vegetation communities and informing us about

wetland and grassland vegetation types. Ralph Topper was invaluable in providing information about local

hydrology. Field support was provided by A. Freierman, B. Drummond, and G. Maentz, and GIS support by M.

Gottfried. Funding for the project was provided by the Jack and Martha Carter Fund of the Carter Herbarium

at Colorado College.

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BOOK NOTICE

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69375-3, pbk). Cambridge University Press, 32 Avenue of the Americas, New York, New York 10013-

2473, U.S.A. (Orders: www.cambridge.org, 1-845-353-7500). $39.99, 304 pp., 102 tables, 21 exercises,

index, 6" x 9".

From the publisher: Latin is one of two acceptable languages for describing new plants, and taxonomists must

be able to translate earlier texts in Latin. Providing a simple explanation of Latin grammar along with an in-

depth vocabulary, this is an indispensable guide for systematic botanists worldwide. All relevant parts of

speech are discussed, with accompanying examples as well as worked exercises for translating diagnoses and

descriptions to and from Latin. Guidelines for forming specific epithets are also included. The authors cross-

reference their grammar to Steam’s Botanical Latin and to articles in the International Code of Nomenclature for

Algae, Fungi and Plants. The comprehensive vocabulary is enhanced with terms from recent glossaries for non¬

flowering plants - lichens, mosses, algae, fungi and ferns - making this an ideal resource for anyone looking to

hone their understanding of Latin grammar and to translate botanical texts from the past 300 years.

EMMA SHORT worked for 5 years at the Royal Botanic Gardens, Kew, UK, on the Index Kewensis database and

with R.K. Brummitt on Authors of Plant Names. She checked Latin for Australian Systematic Botany for 18 years,

worked as a freelance translator, and has taught courses in Botanical Latin.

ALEX GEORGE is an Adjunct Associate Professor at the School of Biological Sciences and Biotechnology, Mur¬

doch University, Perth. He was a botanist with the Western Australian Herbarium for 21 years, then Editor of

the Flora of Australia for 12 years. He studied Latin to Tertiary level and has used it in describing some 400 new

taxa, translating for others and editing.

J.Bot. Res. Inst. Texas 8(1): 226.2014

A FLORISTIC INVENTORY OF DAGNY JOHNSON KEY LARGO HAMMOCK

BOTANICAL STATE PARK AND IMMEDIATELY ADJACENT LANDS

(MONROE COUNTY), FLORIDA, U.S.A.

George J. Wilder

Naples Botanical Garden

4820 Bayshore Drive

Naples, Florida 34112-7336, U.S.A.

gwilder@naplesgarden.org

Janice A. Duquesnel

Florida Park Service

77200 Overseas Fiighway

Islamorada, Florida 33036, U.S.A.

Janice.Duquesnei@dep.state.fi.us

Susan V. Sprunt

P. 0. Box 66

Tavernier, Florida 33070, U.S.A.

Spruntsv@gmail.com

Susan F. Kolterman

30 Poinciana Drive

Key Largo, Florida 33037, U.S.A.

sforrest6@bellsouth.net

ABSTRACT

Individuals of ninety-nine families, 302 genera, 417 species, and 421 infrageneric taxa of vascular plants grow wild in Dagny Johnson Key

Targo Hammock Botanical State Park and on immediately adjacent lands (Key Targo, Monroe Co., Florida). Three hundred (71.3%) of the

421 infrageneric taxa are native to Florida. Herein, eight main kinds of habitats are recognized within the study area, and individual taxa

inhabit one or more of these habitats. Fifty-nine presently reported species are listed as Endangered (38 species) or Threatened in Florida (21

species). For South Florida, one species listed as Extirpated and eight species listed as Critically Imperiled were documented during this

study.

Key Words: Dagny Johnson Park, Key Targo, floristic inventory, vascular plants

RESUMEN

Individuos de noventa y nueva familias, 302 generos, 417 especies, y 421 taxa infragenericos de plantas vasculares crecen nativos en Dagny

Johnson Key Targo Hammock Botanical State Park y en lugares inmediatamente adyacentes (Key Targo, Monroe Co., Florida). Trescientos

(71.3%) de los 421 taxa infragenericos son nativos de Florida. Por ello, se reconocen ocho tipos principales de habitats en el area de estudio,

y los taxa individuales habitan uno o mas de estos habitats. Cincuenta y nueve de las especies citadas estan listadas como En Peligro (38

especies) o Amenazadas en Florida (21 especies). Se documentan para South Florida en este estudio, una especie listada como Extinguida y

ocho especies listadas como Criticamente Amenazadas.

INTRODUCTION

This is the fourth of a series of papers focused on the flora of south Florida (Wilder & McCombs 2006; Wilder

& Roche 2009; Wilder & Barry 2012). Presented, herein, are the results of a floristic inventory of the native and

the exotic taxa of vascular plants growing wild in Dagny Johnson Key Largo Hammock Botanical State Park

and on immediately adjacent lands. We call the studied locations, collectively, “the study area.” These locations

are situated within northern Key Largo (Monroe Co., Florida). Key Largo ranks as the largest and among the

northernmost of the Florida Keys (Weiner, 1977-1986).

Dagny Johnson Key Largo Hammock Botanical State Park (hereafter, called the Park) was established in

1982 and additional lands were added later on. The Park is managed by the State of Florida Department of En¬

vironmental Protection Division of Recreation and Parks (DEP). It exhibits diverse habitats, including the

largest remnant of West Indian hardwood hammock (a kind of rockland hammock) remaining within the

continental United States (SFDEPDRP 2004).

The Park consists of discontinuous properties that, collectively, define an approximately linear contour

oriented from southwest to northeast (Fig. 1, areas depicted in green and orange). Privately owned lands and

J. Bot. Res. Inst. Texas 8(1): 227 - 251.2014

228

Journal of the Botanical Research Institute of Texas 8(1)

associated roads occupy spaces between the Park properties. The Park is centered at 25°14'21"N and

80°19T2"W, it is ca. 7.3 miles long (including spaces between properties), it exhibits a maximum width of ca.

O. 75 miles, and it occupies 2454 acres (SFDEPDRP 2004). The maximum altitude is ca. 13 feet above sea level.

Most of the Park is situated within an eastern sector of northern Key Largo (Fig. 1, area colored green).

This portion of the Park is bounded to the east by the Straits of Florida (a body of water contiguous eastward

with the Atlantic Ocean). To the west, this portion is demarcated, jointly, by three entities that are contiguous

with, and that extend parallel to, one another (Fig. 1, the three entities are represented, collectively, by a single

red line). Listed in an order extending from west to east, these entities include: (1) County Route 905 (CR 905),

(2) ruderal land situated east of CR 905, and (3) a right-of-way that includes power lines. A discontinuous lin¬

ear clearing extends beneath the power lines. The right-of-way is owned by Monroe County, but it is assigned

as a utility easement to the Florida Keys Electric Cooperative (FKEC).

A northwestern sector of the Park is unique, because it occurs by the western side of CR 905 and because

its western boundary abuts Card Sound (a body of water that connects, via Florida Bay, to the Gulf of Mexico;

Fig. 1, area colored orange).

Those portions of the study area that lie immediately adjacent to, but that do not belong to, the Park in¬

clude the linear clearing, aforementioned, certain ruderal lands, and oceanic habitat situated at the eastern

terminus of Valois Blvd. (see below, for additional comment about the latter two portions).

Numerous botanists—independently or in small groups—have collected, and deposited in herbaria,

voucher specimens from northern Key Largo. Examples of previous workers include: T.R. Alexander, F. Alm-

eda, D. Austin, G.N. Avery, K.A. Bradley, C.R. Broome, G.R. Cooley, D.S. Correll, F.C. Craighead,J. De Boer,J.A.

Duquesnel, G.D. Gann, O. Lakela, R.W. Long, J.E. Poppleton, R.P. Sauleda, A.G. Shuey, W.L. Stern, R.F. Thorne,

P. B. Tomlinson, and D.B. Ward (Gann et al. 2002; IRC 2013; Wunderlin & Hansen 2013).

Taylor Alexander (1953, 1968 [1969]) wrote two papers about the vascular flora of the Park. The first pa¬

per was a floristic study of land that was initially pine rockland (land that was also studied presently). The

second paper reported Acacia choriophylla (Fabaceae) as a species new to Florida.

Arthur Weiner (1977-1986) authored a voluminous monograph about the ecology of rockland ham¬

mocks of the Florida Keys. Therein, he presented, in part, the results of 26 surveys of different sectors of rock¬

land hammock in Key Largo, that were undertaken by Weiner and Karen Anchor during/between May, 1977

and Feb. 1986. Weiner (1977-1986) included a separate plant list among the results of each survey.

After the Park was established, DEP staff members compiled lists of the plant species found there (SFDEP¬

DRP 2004). The most recent list, not yet formally approved, was assembled while current research was still

underway (SFDEPDRP, Unit Management Plan in preparation); that list reports certain species that were first

observed by us in the study area. In addition, the Institute of Regional Conservation (IRC 2013) maintains a

separate list of species attributed to the Park.

The present study was undertaken for four reasons. (1) Weiner’s (1977-1986) plant lists and DEP’s lists

specified no voucher specimens, and the IRC list specified a limited number of them; thus, each list was impos¬

sible, or was possible only in part, to verify. (2) The Park list did not stipulate, and the IRC list stipulates incon¬

sistently, which workers had identified particular species. (3) For many of the herbarium specimens collected

by previous workers, the specimen labels did not specify—or they specified imprecisely—where in northern

Key Largo (or in Key Largo, overall) collections were made; thus, it was not possible to definitely link such

specimens to lands currently under investigation. (4) We aspired to document species previously unreported

within the study area.

Voucher specimens or photographs are cited herein of all infrageneric taxa (species and varieties) report¬

ed presently. Indicated, also, are taxa collected by previous workers in northern Key Largo that were not ob¬

served during the present study. Reported, too, are the habitat(s) noted for each infrageneric taxon during

current fieldwork.

Climate, Geology and Soils

Key Largo manifests long, hot, humid summers (with frequent cooling by sea breezes) and warm winters (with

Wilder et al., Flora of Dagny Johnson Key Largo Hammock Botanical State Park

229

Fig. 1. Map depicting northern Key Largo and surrounding bodies of water. Green and orange signify the eastern portion and the northwestern portion,

respectively, of Dagny Johnson Key Largo Hammock Botanical State Park. The red line represents, collectively, CR 905, ruderal land situated east of CR

905, and the right-of-way that includes power lines. This map is from State of Florida Department of Environmental Protection Division of Recreation

and Parks (2004) and was modified by Nathan Mitchell.

230

Journal of the Botanical Research Institute of Texas 8(1)

occasional cooling by intrusions of air from the north). Average monthly temperatures vary from 75° F (Janu¬

ary) to 90° F (July, August). Rainfall occurs throughout the year. Average precipitation is highest in September

and is lowest from/including December through April (Hurt et al. 1995; The Weather Channel 2014).

Key Largo Limestone—a “raised coral reef” of Pleistocene origin—is the uppermost bedrock within Key

Largo (Randazzo & Halley 1997). The limestone is eroded, and here and there large solution holes extend

downward from its surface. The holes are commonly wider than deep, they may exceed five feet deep, and they

may contain disproportionate numbers of individuals of particular plant species ( Acrostichum aureum, Annona

glabra). The holes develop naturally because of rainfall, and they sometimes hold standing water. Also, humans

have excavated the limestone in places, producing basins and canals (SFDEPDRP 2004; George Wilder, per¬

sonal observations).

Dispatch Slough, a body of shallow saltwater, extends through the northeastern sector of the Park and

connects Card Sound with the Straits of Florida (SFDEPDRP 2004).

Six soil types occur within the study area: (1) Pennekamp gravelly muck, 0-2% slopes, extremely stony;

(2) Matecumbe muck, occasionally flooded; (3) Rock outcrop, Tavernier complex, tidal; (4) Islamorada muck,

tidal; (5) Keylargo muck, tidal; and (6) Udorthents-Urban land complex. Soil type nos. 1 and 2 are associated

with upland and subtropical hardwood hammock; type nos. 3-5 occur within mangrove swamps, exhibit el¬

evations of less than two feet, and are subject to daily flooding by tides; and type no. 6 occurs within “con¬

structed upland areas adjacent to areas of water” (Hurt et al. 1995).

Habitats

Recognized, herein, are eight main kinds of habitats within the study area: rockland hammock, mangrove,

rock barren, coastal berm, submerged habitat, ruderal land, the linear clearing situated beneath power lines,

and non-ruderal disturbed land (Appendix 1).

Rockland hammock .—This is subtropical, upland forest. Its name signifies the underlying limestone,

which either is exposed (especially in solution holes) or which lies very near the surface (FNAI2010). In ham¬

mock situated within the study area, dicotyledons comprise the majority, both of individuals and of species, of

trees and shrubs; however, one or more palm species also abound(s) in places. The hammock varies from being

young second-growth forest to being older growth. A thick, soft layer of leaf litter characterizes the higher-

quality portions of hammock.

Before the Park was established, a 15-acre tract - now situated within the northeastern sector of the Park

- was pine rockland habitat; however, during the early 1900s the pines were logged and living pines are now

absent (SFDEPDRP 2004). Subsequent to logging, renewed growth transformed this tract. Today most of it

appears, essentially, as rockland hammock, and that majority is here classified as such.

Mangrove .—This is variously dense forest. It comprises fringing vegetation along the coast and also oc¬

curs sporadically a little inshore of the coast (where it may intergrade with rock barren). Too, it occupies Dis¬

patch Slough. Four species of trees dominate: Avicennia germinans (Black Mangrove), Conocarpus erectus (But¬

tonwood), Laguncularia racemosa (White Mangrove), and Rhizophora mangle (Red Mangrove).

Rock barren .—Grouped together here, under this heading are three previously recognized habitat types:

keys tidal rock barren, keys cactus barren, and portions of terrain elsewhere called a “rocky area of scrub man¬

groves” (by the SFDEPDRP 2004). The first two habitat types were described by the Florida Natural Areas In¬

ventory (FNAI 2010).

The “portions of terrain,” aforementioned, occur somewhat inshore of fringing mangrove habitat, they

are insolated, they are relatively free of mangrove species, and they abound with Bads maridma and chenopo-

diaceous species. Aside from having rocky substrate, they resemble Bads marsh (a habitat which manifests

marl soils; Craighead 1971). The three habitat types are grouped together here: (A) for purposes of simplicity,

(B) because all are considerably insolated, (C) because certain species abound in all three types, and (D) be¬

cause some of the types may abut one another (i.e., Keys cactus barren [which has slightly higher elevation]

and the portions of terrain, aforementioned [which have slightly lower elevation]).

Rock barren is widespread at low elevations of the Park. Aside from occurring inshore of the coast, it lines

Wilder et al., Flora of Dagny Johnson Key Largo Hammock Botanical State Park

231

both sides of Dispatch Slough. The substrate is intact- or broken-up limestone. The least elevated parts of rock

barren are commonly flooded, whereas flooding is infrequent in higher parts. Rock barren located by the west¬

ern side of Dispatch Slough abuts rockland hammock to the west and mangrove habitat to the east.

The Florida Natural Areas Inventory (FNAI 2010) ranks the Park as an “exemplary site” for Keys tidal

rock barren and, also, for coastal berm.

Coastal berm. —The Florida Natural Areas Inventory (FNAI 2010) defines coastal berm as “... a short for¬

est or shrub thicket found on long narrow storm-deposited ridges of loose sediment ...” Within the Park a

geographically limited, forested ridge of berm exists just landward of the fringing mangrove habitat. At high

elevations (of ca. a few feet) leaf litter covers the surface, but massive quantities of trash surmount less elevated

berm sectors (see below).

Submerged habitat. —Submerged, aquatic plant species were noted growing in ocean water (at the east¬

ern terminus of Valois Blvd. [a road traversing a space between Park properties]), in Dispatch Slough, and

within an excavated basin of limestone containing shallow water. The first location actually occurs within a

peripheral-most portion of John Pennekamp Coral Reef State Park.

Ruderal land. —We define this in a strict sense as disturbed land bordering or on roads. Ruderal land was

investigated (A) along the eastern side of CR. 905, (B) along roads penetrating the spaces between Park proper¬

ties, and (C) along, and on, roads located within the Park.

The linear clearing situated beneath power lines. —This clearing extends for nearly the entire length of

the Park; however, it is interrupted mainly at two places where the power lines extend especially close to CR

905. At those places the power lines overlie land confluent with, and outwardly indistinguishable from, ru¬

deral land bordering CR 905. At those places land beneath the power lines is classified as ruderal land, rather

than as part of the linear clearing. Most portions of the clearing are bordered on both sides by rockland ham¬

mock.

The linear clearing is mowed once yearly (personal communication from Sara Hamilton of FKEC, of 15

Aug 2013), so that herbaceous vegetation predominates. Depauperate woody plants occur, but the periodic

mowings prevent them from becoming dominant. Because of their restricted development, it was considered

misleading to list for the clearing—and we do not do so here—certain species of the depauperate woody plants

(Appendix 1). Also, the clearing is mowed less frequently than are the ruderal lands associated with the Park,

so that it often manifests more robust vegetation than do the ruderal lands.

Non-ruderal disturbed land. —This habitat includes variously extensive clearings within the Park. Con¬

current with the present study, DEP acted to restore some clearings (by bulldozing and, sometimes, by apply¬

ing mulch). Those efforts, apparently, promoted the arrival of certain species within the Park that were previ¬

ously absent (Ipomoea hederifolia, Trianthemaportulacastrum).

Flotsam-derived Trash

Massive trash accumulations clearly deposited during storms and/or high tides occur a little inshore from the

Straits of Florida. These accumulations appear within portions of fringing mangrove habitat, of coastal berm

at low elevation, and of rock barren. Trash includes glass bottles, plastic bottles, additional plastic objects (e.g.,

styrofoam slabs), buoys, ropes, metal cans, buckets, fishing gear, shoes, boards, poles, etc.

Historical Sketch

Northern Key Largo has a long history of human occupation and of human-induced disturbances. Native

American Indians inhabited “Key Largo Hammock” between 1600 BC and 1200 AD, and they continued their

activities there until the mid-18th century. Included, were members of the Calusa tribe, of Timucuan culture,

and settlers from the Bahamas. Prehistoric Indian sites remain within the Park. They tend to be located near

the water and mainly include shell middens (SFDEPDRP 2004).

The Spanish first explored the northern Florida Keys in 1516. Thereafter, and continuing throughout the

16th, 17th, and 18th centuries, certain species of hardwoods were logged aggressively within rockland ham¬

mock situated there. Examples included Colubrina elliptica (Soldierwood), Eugenia rhombea (Red Stopper),

Guaiacum sanctum (Lignumvitae), Hypelate trifoliata (White Ironwood), and Swietenia mahagoni (Mahogany).

232

Journal of the Botanical Research Institute of Texas 8(1)

Logs were extracted, partly, for shipbuilding, home construction, and cabinetry (Rothing 2009; Weiner 1977-

1986). Later on in northern Key Largo, Pinus elliottii (Slash Pine) and Quercus virginiana (Live Oak) were also

logged (see below; Alexander 1953; SFDEPDRP 2004). Today, within the Park one may still observe in rock-

land hammock the old trunks of previously cut trees.

The first European settlers arrived during the mid-18th century.

During the 1800s, individuals established homesteads and farmed in northern Key Largo (SFDEPDRP

2004; Wilkinson 2013b). For example, Samuel Lowe purchased and homesteaded 900 acres of land. A farmer,

he likely grew pineapples, a crop which most historians believe was then the crop of choice in the northern

Florida Keys. If Lowe raised pineapples, he probably practiced slash-and-burn agriculture and cleared consid¬

erable land (Wilkinson 2013b). We emphasize Lowe, because his land would have been located partly or en¬

tirely within the current study area (a conclusion that we base on Wilkinson’s [2013b] statement that Lowe’s

property encompassed land that was much later developed into a missile base [see below]).

During the 1900s, significant attempts were made to create housing developments within areas now in¬

corporated within the Park. Two examples are provided. (1) The Port Bougainville site, the largest such area,

encompasses ca. 100 acres within a southern sector of the Park. During the late 1970s and early 1980s this site

was subjected to land clearing, road building, and the construction of lakes, a marina, various buildings, an

entrance archway, a tunnel, etc. The site was subsequently restored somewhat, but it still exhibits considerable

cleared land, roads, and additional artifacts. (2) The Carysfort Yacht Club site includes over 40 acres of land

within a northern sector of the Park. That site was cleared to house the Carysfort Yacht Club, but was later

utilized as a campground. A marina was also built. Subsequent restoration attempts entailed removal of the

campground and marina, partial reforestation with native vegetation, and remodeling of the landscape

(SFDEPDRP 2004).

In June 1965, nearly three years after the Cuban missile crisis, the U.S. government made operational the

Key Largo HM-40 Nike Hercules missile base in northern Key Largo. That facility, which housed 120 individu¬

als, was closed in June 1979. It had two main parts: the Integrated Fire Control area (radar/administrative area)

and the Launcher area (which housed missiles in hardened storage bunkers). Today the two parts are separated

by CR 905 and are situated within the Park and within the adjacent Crocodile Lake National Wildlife Refuge,

respectively (Wilkinson 2013a, 2013b). Within the Park, the defunct Integrated Fire Control area occupies ca.

13 acres and manifests deteriorating buildings, pavement, and degraded habitat (SFDEPDRP 2004; George

Wilder, personal observations). Hammock vegetation overgrows parts of the area.

The Park contains portions of two highways: Old CR 905 and Old Card Sound Rd. Below, we apply the

names Old CR 905 and Old Card Sound Rd., solely, to those portions rather than to the entire highways. Old

CR 905 and Old Card Sound Rd. are now closed to ordinary traffic. During the late 1960s, Old CR 905 (which

demarcated the southern boundary of the Integrated Fire Control area) was replaced by a portion of CR 905

oriented west of Old CR 905 (Fig. 1, upper portion of red line). Also, Old Card Sound Rd. once extended east¬

ward to the Straits of Florida, where it culminated in “Dynamite Docks” (a pier utilized for unloading and

transporting explosives). Initially, Old CR 905 and Old Card Sound Rd. traversed Dispatch Slough at two sepa¬

rate locations; however, in 2000, roadway was removed at each location, and the locations are now submerged

(SFDEPDRP 2004; Wilkinson 2013a, 2013b).

Plant-collecting combined with logging during the 20th century have also taken a toll (Roger Hammer,

personal communications of 6 Jan and 10 Jan 2014). Weiner (1977-1988), referring to “upper keys high ham¬

mocks”, including rockland hammock in Key Largo, commented as follows. There are “... very few remaining

populations of lignum vitae, white ironwood, manchineel [ Hippomane mancinella] , soldierwood, red stopper,

satinleaf [ Chrysophyllum oliviforme] and other tropical hardwoods. Orchids and bromeliads ... are now repre¬

sented by vestigial populations. The dollar orchid [ Prosthechea boothiana] has all but disappeared. ... Popula¬

tions of bromeliads are found in only the remotest and least accessible areas surveyed. All of these air plants are

suffering from over-collection and habitat alteration.” (Names between brackets were added by us.)

Wilder et al., Flora of Dagny Johnson Key Largo Hammock Botanical State Park

233

METHODS

The study area was visited 109 times, beginning on 19 March 2012 and ending on 20 Feb 2014. Multiple visits

were made during each month of the year. George Wilder vouchered all but six infrageneric taxa with dried

herbarium specimens; because of their rarity, Susan Kolterman vouchered those six taxa solely with numbered

photographs ( Opuntia corallicola, Phoradendron rubrum, Prosthechea boothiana, Tillandsia fasciculata, Tilland-

sia utriculata, and Vanilla barbellata). All specimens but one and all photographs were deposited in the Her¬

barium of Southwestern Florida (SWF), housed at the Naples Botanical Garden (Appendix 1). The exceptional

specimen (of Sida sp., a species possibly new to Florida) was donated to the USF Herbarium (USF). Examined,

also, were herbarium specimens that previous workers had collected, or had apparently collected, in northern

Key Largo (at Virtual Herbarium 2013, at Wunderlin and Hansen 2013, at FTG, and at SWF). We inventoried

solely wild individuals, i.e., plants which - except in three cases - were deemed to have originated naturally

within the study area. Exceptional, were Acacia choriophylla, Opuntia corallicola, and Phoradendron rubrum;

those species, extirpated from Key Largo years ago, were introduced into the Park by previous workers. We call

the introduced plants wild because they grew in natural habitats, because they were not being maintained ar¬

tificially, because some were robust and many years old, and because the three species were formerly native to

Key Largo. Species are characterized as native, exotic, or endemic to Florida, according to Wunderlin and

Hansen (2011, 2013). Mostly, present nomenclature follows Wunderlin and Hansen (2013); however, Appen¬

dix 1 (footnote 1) specifies nomenclatural differences between that work and the present paper.

RESULTS AND DISCUSSION

Taxonomic analysis of present data

The study area exhibited 99 families, 302 genera, 417 species, and 421 infrageneric taxa (species and varieties)

of vascular plants (Appendix 1). Between parentheses, the numbers of families, genera, and infrageneric taxa

are indicated, respectively, for each of the following major groups: pteridophytes (6, 10, 12), angiosperms (93,

292, 409), monocotyledons (14, 62, 102), and dicotyledons sensu lato (79, 230, 307). For infrageneric taxa of

each of these major groups, their percentage of all 421 infrageneric taxa is listed: pteridophytes, 2.9%; angio¬

sperms, 97.1%; monocotyledons, 24.2%; and dicotyledons sensu lato, 73.1%.

The five largest families of monocotyledons, as gauged by the numbers of infrageneric taxa present, are

Poaceae (56), Cyperaceae (15), Arecaceae (8), Bromeliaceae (8), and Orchidaceae (5) (for each family the num¬

ber of infrageneric taxa is listed between parentheses). The families Poaceae and Cyperaceae, collectively, ex¬

hibited 16.9% of all 421 infrageneric taxa listed (i.e., 71 infrageneric taxa).

The eleven largest families of dicotyledons sensu lato are Fabaceae (43), Asteraceae (29), Euphorbiaceae

(21), Rubiaceae (16), Convolvulaceae (14), Malvaceae (9), Verbenaceae (8), Boraginaceae (7), Cactaceae (7),

Scrophulariaceae (7), and Solanaceae (7). The families Fabaceae and Asteraceae, collectively, exhibited 17.1% of

all 421 infrageneric taxa listed (i.e., 72 taxa).

No gymnosperms were observed, and pteridophytes were noted solely within the Park. Most fern species

were rare or uncommon; however, numerous individuals were observed of Acrostichum aureum (localized

primarily within solution holes) and of Pleopeltis polypodioides (localized within and bordering other portions

of hammock).

Infrageneric taxa situated within the Park vs. on immediately adjacent land

Three hundred and eighty three infrageneric taxa (91.0% of all 421 infrageneric taxa reported) occurred solely

within the Park or both within, and external to, the Park. The remaining 38 taxa grew solely outside of the

Park. Of all 143 infrageneric taxa reported presently for ruderal land, 63 taxa grew on ruderal land that was

located either solely within the Park or that occurred both within, and external to, the Park; the remaining 80

taxa inhabited ruderal land located solely outside of the Park.

Infrageneric taxa and habitats

Habitats are listed for all 421 infrageneric taxa reported here (Appendix 1).

234

Journal of the Botanical Research Institute of Texas 8(1)

Within the entire study area, non-ruderal disturbed land exhibited the highest percentage of infrageneric

taxa. Intermediate percentages of taxa grew in each of rockland hammock and ruderal land. Lowest percent¬

ages occurred within each of the linear clearing situated beneath power lines, berm, rock barren, mangrove

habitat, and underwater habitat.

Supporting data are presented. Each number, below, refers solely to the infrageneric taxa that we noted

inside of a habitat, not to taxa whose sole association with the habitat was occurence at the habitat boundary.

For each habitat indicated, listed between parentheses are the number of infrageneric taxa observed therein

and the percentage which that number represents of all 421 infrageneric taxa reported here: non-ruderal dis¬

turbed land (231; 54.9%); rockland hammock (143,34.0%); ruderal land (143, 34.0%); the linear clearing situ¬

ated beneath power lines (87, 20.7%); berm (56,13.3%); rock barren (52,12.4%); mangrove habitat (34, 8.1%);

and underwater habitat (3, 0.71%).

Also, noted were (A) 50 infrageneric taxa situated at the boundaries between rockland hammock and

other habitats, that were not located inside the hammock itself, and (B) 27 infrageneric taxa situated at the

boundaries between mangrove habitat and other habitats (excluding rockland hammock), that were not lo¬

cated inside the mangrove habitat itself (Appendix 1).

As was stated above, most of that locale which in previous years was pine rockland is classified presently

as rockland hammock. (Hereafter, we designate as Locality A that portion which is now hammock). Accord¬

ingly, we have—in the previous two paragraphs—subsumed within the tallies of taxa of rockland hammock

the counts of all taxa situated within Locality A.

Sixty-seven infrageneric taxa were noted as being situated within, or in several cases bordering, Locality

A (Appendix 1). Although, Locality A is currently interpreted as rockland hammock, that locality differs from

other hammock within the study area, because certain species characteristic of pine rockland persist there.

Within the study area, those species are either confined to Locality A ( Quercus virginiana, Serenoa repens) or

they are represented there disproportionately ( Byrsonima lucida, Myrica cerifera).

Approximately, 60 years ago Taylor Alexander, together with a plant ecology class, investigated Locality

A. They delineated an east-west transect through Locality A, along which they recorded 25 species (Alexander

1953). Noted, presently within Locality A were 22 of those 25 species. Two more of the species were observed

elsewhere in the Park ( Guettarda scabra, Simarouba glauca); however, Alexander (1953) listed one species, Psy-

chotria sulzneri, that was not observed presently within the study area. Coordinate with present findings, Al¬

exander (1953) reported that Locality A “... showed little difference ...” from other areas of typical Key Largo

hammock.

Native and endemic taxa inventoried during the present study

Within the study area 300 (71.3%) of the 421 infrageneric taxa recorded were native to Florida (this calculation

does not include Chenopodium sp., which was insufficiently mature to identify). Between parentheses, the

number and percentage of native infrageneric taxa within each major group of vascular plants are listed, re¬

spectively: pteridophytes (10, 83.3%), angiosperms (290, 70.9%), monocotyledons (66, 64.7%), and dicotyle¬

dons sensu lato (224, 73.0%).

Five taxa were endemic to Florida: Agave decipiens, Argythamnia blodgettii, Chamaesyce conferta, Harrisia

fragrans, and Opuntia corallicola.

Exotic species inventoried during the present study

One hundred and twenty-one infrageneric taxa observed within the study area are exotic within Florida (not

counting Chenopodium sp.; Appendix 1).

The Florida Exotic Pest Plant Council (FLEPPC 2013) has recognized two categories of plant species ex¬

otic within Florida, that pose especial threats to the ecology of the State, overall, i.e., Category I and Category

II (those categories indicate decreasing degree of threat; FLEPPC 2013). Noted presently were 16 Category I

species ( Albizia lebbeck, Asparagus aethiopicus, Casuarina equisetifolia, Colubrina asiatica, Ficus microcarpa,

Jasminum dichotomum, Lantana camara, Manilkara zapota, Melinis repens, Nephrolepis brownii, Neyraudia reyn-

Wilder et al., Flora of Dagny Johnson Key Largo Hammock Botanical State Park

235

audiana, Panicum repens, Pennisetum purpureum, Scaevola taccada, Schinus terebinthifolia, and Thespesia popul-

nea) and 19 Category II species ( Cocos nucifera, Dactyloctenium aegyptium, Eulophia graminea, Flacourtia indica,

Hyparrhenia rufa, Leucaena leucocephala, Macroptilium lathyroides, Panicum maximum, Pennisetum setaceum,

Pteris vittata, Richardia grandiflora, Ricinus communis, Ruellia blechum, Sansevieria hyacinthoides, Sphagneticola

trilobata, Stachytarpheta cayennensis, Terminalia catappa, Tradescantia spathacea, and Tribulus cistoides).

Possibly, there were four additional Category II species: Chamaedorea seifrizii, Phoenix reclinata, Pittospo-

rum pentandrum, and Washingtonia robusta. We did not identify those species, but we did observe sterile plants

or seedlings that are identified here as Chamaedorea sp., Phoenix sp., Pittosporum sp., and Washingtonia sp. The

16 Category I species and the 19 Category II species comprised 20.8% and 23.5% of all 77 Category I species

and 81 Category II species recognized for Florida, respectively.

Within the study area Leucaena leucocephala, Manilkara zapota, and Thespesia populnea ranked among

the most abundant of exotic species listed by FLEPPC (2013). Eulophia graminea, a Category II species native to

Asia and to subtropical islands in the Pacific Ocean, was first observed in Florida in ca. 2006 (Pemberton et al.

2008). Since its discovery in south Florida, E. graminea has spread invasively there, and during the present

investigation it has become considerably more abundant within the study area.

Staff members of DEP and of FKEC, as well as individuals unafhliated with those entities, routinely de¬

stroy exotic plants within the study area. They have, apparently, extirpated Bucida buceras x Bucida spinosa,

Peltophorum pterocarpum (two taxa that were previously listed for the Park [SFDEPDRP 2004]), Hylocereus

undatus, and Melia azedarach.

Native species listed as rare in Florida

The study area manifests an extraordinary number of rare species (Table 1). Fifty-nine presently reported spe¬

cies are State-listed as Endangered (38 species) or Threatened in Florida (21 species; Weaver & Anderson

2010). Within the study area, all but one of these species are confined to the Park; Argythamnia blodgettii, the

sole exception, comprises a small, localized population located in/by the linear clearing situated beneath the

power lines. For South Florida, one species listed as Extirpated and eight species listed as Critically Imperiled

(Gann et al. 2002) were documented during this study.

Herein, two listed species are newly reported for the Park: Celosia nitida and Vallesia antillana.

Two species of Cactaceae are State-listed: Acanthocereus tetragonus and Harrisiafragrans. Excluding cacti,

38 of the State-listed species grow as shrubs (including Phoradendron rubrum) or trees (including Thrinax spp.);

however, other species develop as erect herbs ( Acrostichum aureum, Argythamnia blodgettii, Prosthechea boothi-

ana, Tillandsia spp., Sclerialithosperma, Voyria parasitica) or as herbaceous- or woody vines ( Dalbergiabrownei,

Jacquemontia spp., Microgramma heterophylla, Passiflora multiflora, Rhynchosia swartzii, Vanilla barbellata).

Four of the State-listed species are considered in greater detail, below.

Acacia choriophylla.—Taylor Alexander reported a single tree of A. choriophylla, apparently of natural

origin, in northern Key Largo. The discovery area was “... forested by species of West Indian affinity” ... and

was “... typical of the ecotone between salt water mangrove swamps and high hammock...” (Alexander 1968

[1969]). His report was the first and only indication that A. choriophylla is native to the United States; however,

Gann et al. (2002) concluded that A. choriophylla was likely extirpated in northern Key Largo during the 1970s

and early 1980s. Many years ago a Park biologist planted the individuals of A. choriophylla reported presently.

Hippomane mancinella.—Ten years ago SFDEPDRP (2004) reported fewer than twelve individuals of

this species in the Park. Observed, during the present study were only one mature individual and several seed¬

lings of that species that grew beneath it. The plants were situated within berm habitat. Hippomane mancinella

is poisonous to humans, a circumstance that has led to dramatic human-induced destruction of this species.

Thus, within Florida H. mancinella is now “... only sparsely scattered throughout the Keys as well as near Fla¬

mingo in Everglades National Park” (Nelson 1994).

236

Journal of the Botanical Research Institute of Texas 8(1)

Table 1 . List of the species of rare plants of Dagny Johnson Key Largo Hammock Botanical State Park and immediately adjacent lands. Rankings of rarity are for

Florida (Weaver & Anderson 2010) and for south Florida (Gann et al. 2002). Crit. Imp. = critically imperiled; End. = endangered; Ext. = extirpated; Hist.

= historical; Threat. = threatened.

Taxon

Weaver & Anderson (2010)

Gann etal. (2002)

Acacia choriophylla

End.

Ext.

Acanthocereus tetragonus

Threat.

Acrostichum aureum

Threat.

Argusia gnaphalodes

End.

Argythamnia blodgettii

End.

Bourreria succulenta

End.

Byrsonima lucida

Threat.

Calyptranthes pallens

Threat.

Calyptranthes zuzygium

End.

Canella winterana

End.

Celosia nitida

End.

Chrysophyllum oliviforme

Threat.

Colubrina elliptica

End.

Crossopetalum ilicifolium

Threat.

Crossopetalum rhacoma

Threat.

Dalbergia brownei

End.

Dodonaea elaeagnoides

End.

Drypetes diversifolia

End.

Drypetes lateriflora

Threat.

Erithalis fruticosa

Threat.

Ernodea cokeri

End.

Crit. Imp.

Eugenia confusa

End.

Eugenia rhombea

End.

Crit. Imp.

Exostema caribaeum

End.

Gossypium hirsutum

End.

Guaiacum sanctum

End.

Crit. Imp.

Harrisia fragrans

End.

Hippomane mancinella

End.

Hypelate trifoliata

End.

Jacquemontia havanensis

End.

Crit. Imp.

Jacquemontia pentanthos

End.

Jacquinia keyensis

Threat.

Manilkara jaimiqui

Threat.

Maytenus phyllanthoides

Threat.

Microgramma heterophylla

End.

Opuntia corallicola

End.

Crit. Imp.

Opuntia stricta

Threat.

Pass i flora multi flora

End.

Phoradendron rubrum

End.

Crit. Imp.

Pithecellobium keyense

Threat.

Prosthechea boothiana

End.

Psychotria ligustrifolia

End.

Reynosia septentrionalis

Threat.

Rhynchosia swartzii

End.

Crit. Imp.

Schaefferia frutescens

End.

Scleria lithosperma

End.

Smilax havanensis

Threat.

Solanum donianum

Threat.

Swietenia mahagoni

Threat.

Thrinax morrisii

Threat.

Thrinax radiata

End.

Tillandsia balbisiana

Threat.

Tillandsia fasciculata

End.

Tillandsia flexuosa

Threat.

Tillandsia utriculata

End.

Trema lamarckiana

End.

Vallesia antillana

End.

Crit. Imp.

Vanilla barbellata

End.

Voyria parasitica

End.

Wilder et al., Flora of Dagny Johnson Key Largo Hammock Botanical State Park

237

Phoradendron rubrum. —Plants of P. rubrum parasitize stems of Swietenia mahagoni. During March,

1998, Josef Nemec discovered a previously unknown population (here called Population A) of P. rubrum

within the Park; it was the last remaining original population known to occur there. Janice Duquesnel col¬

lected seeds from that population and planted them onto mahogany trees situated within the Park and else¬

where. The seeds yielded numerous offspring that persist today. Population A itself died-out in 2004, con¬

comitant to death of the host trees.

Gann et al. (2002) concluded that wild plants of P. rubrum were known with certainty from South Florida,

solely, from the Park; however, in 2002, IRC staff members discovered P. rubrum on Sands Key (Miami-Dade

Co., FL; SFDEPDRP 2004). Subsequently, in May 2013, Larry Manfredi discovered P. rubrum growing at an¬

other site in Key Largo.

Opuntia corallicola. —John K. Small (1930) reported O. corallicola from Key Largo, but thereafter the

species was extirpated there. Subsequently, staff members of Fairchild Tropical Botanic Garden (particularly,

Christopher Kernan), of the Florida Park Service (including Janice Duquesnel), and of other institutions col¬

laborated to introduce plants of O. corallicola into the Park. To the present day, Janice Duquesnel—with the

assistance of other individuals—monitors O. corallicola and Phoradendron rubrum within the Park.

Gann et al. (2002) reviewed the history of discovery of O. corallicola in south Florida, and they indicated

efforts to introduce that species in other Florida Keys.

Taxa that are rare or localized within the study area

One hundred and twenty-nine infrageneric taxa (30.6% of all infrageneric taxa reported presently) are judged

to be rare within the study area (Appendix 1). We deem a taxon to be rare there (A) if no more than five indi¬

viduals, thereof (or five clumps of individuals, in the case of herbaceous species), were observed, or (B) if, re¬

gardless of the number of individuals observed, the species occupied an area not larger than a housing lot.

Twelve native species are listed here as rare, that were also characterized above as Endangered or Threat¬

ened in Florida (Weaver & Anderson 2010) or as Critically Imperiled in south Florida (Gann et al. 2002):

Acacia choriophylla, Argythamnia blodgettii, Celosia nitida (1 to several), Crossopetalum ilicifolium, Eugenia rhom-

bea, Hippomane mancinella, Jacquemontia havanensis, Jacquemontia pentanthos, Solanum donianum (1), Trema

lamarckiana (1), Vallesia antillana (2), and Vanilla barbellata (the numbers between parentheses indicate the

number of individuals noted presently for certain species). Another species, Sporobolus pyramidatus, is not

classified as Endangered, Threatened, or Critically Imperiled, but is rare in the study area and was also consid¬

ered rare in Florida, overall (Wunderlin & Hansen 2011).

In contrast to the aforementioned species, various native/non-native species that are rare in the study area

vary from occasional to frequent in Florida, overall (e.g., Amaranthus spinosus, Ambrosia artemisiifolia,

Chamaecristanictitans, Cyperusflavescens, Echinochloa colona, Eclipta prostrata, Eleocharisflavescens, Erechtites

hieraciifolius, Erythrina herbacea, Gaura angustifolia, Eaunaea intybacea, Eeptochloa fusca subsp. fascicular is,

Monanthochloe littoralis, Muhlenbergia capillaris, Oldenlandia corymbosa, Nephrolepis exaltata, Pilea microphyl-

la, Solanum americanum, Sonchus asper, and Toxicodendron radicans (Wunderlin & Hansen 2011).

Some native species that are not rare within the study area exhibit conspicuously restricted distributions,

therein. (1-4) Eugenia confusa, Schoepfia chrysophylloides, Thrinax morrisii, and Thrinax radiata are limited to

rockland hammock situated within the northern portion of the Park (excluding planted individuals of T. ra¬

diata that grow on a plot of disturbed land located elsewhere within the study area); (5) Tillandsia setacea is

confined to the northwestern sector of the Park, where plants grow at/near the boundary between mangrove

habitat and rockland hammock; (6) Argusia gnaphalodes grows wild within disturbed land on a spoil island

bordering the Straits of Florida (albeit, apparently planted individuals occupy disturbed land situated else¬

where); (7) Harrisia fragrans grows primarily at/near the boundary between rockland hammock and either

mangrove habitat or rock barren; and (8) Ernodea cokeri is confined almost entirely within/along that portion

of rockland hammock that was formerly pine rockland (Locality A).

238

Journal of the Botanical Research Institute of Texas 8(1)

Comparisons with previous studies

Examined at FTG and USF, collectively, were herbarium specimens of 26 species and one hybrid that were not

currently observed within the study area, but that previous workers had collected, or had apparently collected,

in northern Key Fargo (imprecise language on some herbarium-specimen labels made it uncertain whether the

corresponding specimens were from northern Key Fargo; Appendix 1). According to the labels, overall, of the 27

taxa, specimens of solely five species and of the hybrid were collected within the Park: Ayenia euphrasiifolia, Cy-

perus croceus, Daturametel, Pectis xfloridana, Thelypteris kunthii, and Vitex trifolia. We could not ascertain wheth¬

er any other of the 27 taxa were collected from the study area. Fifteen (55.5%) of the 27 taxa are native to Florida.

SFDEPDRP (2004) listed 347 infrageneric taxa of vascular plants for the Park. Documented, presently

were all but 34 (9.8%), but possibly up to 37, of those taxa (this imprecision reflects the uncertain identifica¬

tions of some taxa). Of those 34 taxa, nine (26.4%) were native to Florida; remaining taxa were listed as non¬

native or cultivated. The Institute for Regional Conservation (IRC 2013) maintains a database that summarizes

previous floristic work undertaken within the Park and which reports ca. 458 species of vascular plants for the

Park. Documented, during current research were 339 to 345 of those species (six species of four genera were

questionable, because we observed solely sterile adults or seedlings of those genera).

The nature of the data assembled here differs in certain ways from that provided by the latter two sourc¬

es—a circumstance which helps to explain the differences in the numbers of taxa reported presently and by

those sources. A. Inventoried, presently were taxa located both within the Park and on immediately adjacent

land, whereas, both other sources listed taxa solely for the Park. B. Taxa that grew solely under cultivation are

not reported presently, whereas, both other sources reported such taxa. C. Our inventory is conclusive. In

contrast, the IRC database lists certain species as present, but describes other species as being reported (24), as

assumed to be present (10), as doubtfully present (5), as recorded as present in error (12), as possibly extirpated

(6), and as presumed extirpated (13). (Each number between parentheses is the number of species that IRC

assigns to the associated descriptor, that were not presently observed and/or that could not presently be identi¬

fied with certainty [the six species, aforementioned]).

The aforementioned differences in the numbers of reported taxa also reflect the circumstance that certain

taxa, apparently, died-out in, or were extirpated from, the study area before current research began. For ex¬

ample, we repeatedly visited the original collection locality of Ayenia euphrasiifolia (i.e., the Integrated Fire

Control area), but we never observed that species. We speculate that it was shaded-out by growing woody

vegetation. At least one species first recorded for the Park during current research, Turbina corymbosa, was in¬

advertently extirpated there during restoration efforts by DEP.

APPENDIX l 1

Table of species, varieties, a hybrid, and higher-level taxa documented during the present study and by previous workers. Non-bold font

signifies infrageneric taxa (species and varieties) documented within the study area during the present investigation. Infrageneric taxa

that previous workers documented, or apparently documented in northern Key Largo, overall, but that were not observed during the

present study are listed with bold font. For certain species, listed after a species name is/are a relevant synonym (between brackets),

an indication of whether the species is rare within the study area, and/or the designation of the species by the Florida Exotic Plant Pest

Council (FLEPPC 2013). Presented after a species name is the five-digit Wilder and McCombs collection number 2 of a voucher specimen

or of a voucher photograph of that species. Habitat data are provided within the eight vertical columns at the right of this table. For

infrageneric taxa documented solely by previous investigators, data are provided as follows after the Latin name of a taxon: relevant

synonym, if any (between brackets); collector(s); collection number; year of collection; and acronym of the herbarium where the speci¬

men is on deposit. 3 After the name of each family and suprafamilial taxon, between parentheses are included two or four numbers; the

two numbers not in italics—if present—signify, respectively, the numbers reported presently of genera and infrageneric taxa within that

family or suprafamilial taxon; the two numbers in italic—if present—signify, respectively, the sums of numbers reported presently and

by previous workers, of such genera and infrageneric taxa. * = alien to Florida; ? = a taxon that was not identified with sufficient precision

to ascertain whether it is native to, or exotic within, Florida; n = endemic to Florida; Berm = berm habitat; Dist = non-ruderal disturbed

land; FLEPPC I and FLEPPC II = taxa recognized as Category I or Category II species by the Florida Exotic Plant Pest Council (2013); Man =

mangrove habitat; Pow = linear clearing beneath power lines; RH = rockland hammock; Rock = rock barren; Rud = ruderal land; Subm

= underwater habitat; X = occurrence of a taxon within a habitat, away from a habitat boundary (except for X bp and X pi ). For ruderal land,

X signifies occurrence on ruderal land within the Park, whereas, X bp signifies occurrence on ruderal land outside of the Park; for rockland

hammock, X pi signifies rockland hammock which was pine rockland prior to the onset of current research. The following symbols signify

occurrences at habitat boundaries: X d = boundary with non-ruderal disturbed land; X man = boundary with mangrove habitat; X pi ' man =

Wilder et al., Flora of Dagny Johnson Key Largo Hammock Botanical State Park

239

boundary between mangrove habitat and rockland hammock that was formerly pine rockland; X pi r = boundary between ruderal land

and rockland hammockthat was formerly pine rockland; X pi rb = boundary between rock barren and rockland hammock that was formerly

pine rockland; X po = boundary with linear clearing beneath power lines; X r = boundary with ruderal land;X rb = boundary with rock barren.

1 We follow the nomenclature of Wunderlin and Hansen (2013), with two exceptions. (1) We recognize families which they submerged

within, or divided into, other families. Between parentheses, after the name of each family that we recognize is listed the corre¬

sponding family(ies) of Wunderlin and Hansen (2011): Asclepiadaceae (Apocynaceae), Capparaceae (Brassicaceae), Chenopodiaceae

(Amaranthaceae), Euphorbiaceae (Euphorbiaceaesensusfncfo, Phyllanthaceae, Putranjivaceae), Scrophulariaceae (Orobanchaceae,

Plantaginaceae, Scrophulariaceae sensu stricto ), and Sterculiaceae (Malvaceae). (2) We recognize varieties of Conocarpus erectus L.,

Digitaria ciliaris (Retz.) Koeler, Eragrostis ciliaris (L.) R. Br., Paspalum setaceum Michx, and Schizachyrium sanguineum (Retz.) Alston as

did Long and Lakela (1976), Wipff (2003a), Peterson (2003), Allen and Hall (2003), and Wippf (2003b), respectively. Wunderlin and

Hansen (2011) did not do so.

2 Ms. Martha McCombs contributed importantly to SWF; hence, on the label of each herbarium sheet from SWF George Wilder's name

and Martha McCombs' name precede the collection number of each specimen, a circumstance not duplicated in this appendix.

3 Data for previous collections were compiled from Virtual Herbarium (2013), from Wunderlin and Hansen (2013), and during visits to

FTG and USF.

RH Man Rock Rud Pow Dist Subm Berm

X pi

PTERIDOPHYTES (10,12; 77, 73)

DENNSTAEDTIACEAE (1,1)

Pteridium aquilinum var. caudatum

(L.) Sadeb.; 34212

NEPHROLEPIDACEAE (1, 2)

Nephrolepis exaltata (L.) Schott; Rare; 34468

*Nephrolepis brownii (Desv.) Hovenkamp & Miyam.;

remnant of cultivation; Rare; 33548

POLYPODIACEAE (4, 4)

Campyloneurum phyllitidis (L.) C. Presl; Rare; 34416

Microgramma heterophylla (L.) Wherry; 34527

Phlebodium aureum (L.) J. Sm.; 34366

Pleopeltis polypodioides (L.) E.G.

Andrews & Windham; 33622

PSILOTACEAE (1,1)

Psilotum nudum (L.) P. Beauv.; 34498

PTERIDACEAE (2, 3)

Acrostichum aureum L.; 34255

Acrostichum danaeifolium Langsd. & Fisch.; Rare; 34467

*Pteris vittata L.; FLEPPC II; 33816

THELYPTERIDACEAE (7, 7)

Thelypteriskunthii (Desv.) C.V. Morton; G.A. Gann, J.A. Duquesnel 1154; 2003 (FTG)

VITTARIACEAE (1,1)

Vittaria lineata (L.) Sm.; 34470 X, X pi

X,X man ,X pi

X,X man ,X pi ,X rb

X,X pi

XPi-m

MONOCOTYLEDONS (62,102; 64,108)

AGAVACEAE (2, 2] 2,3)

n Agave decipiens Baker; Rare; 33875 X

*Agavesisalana Perrine; R.W. Long, F. Almeda, J. De Boer, C.R. Broome 1860; 1966 (USF)

Yucca aloifolia L.; 33876 X,X pi

AMARYLLIDACEAE (1,1)

Hymenocallis latifolia (Mill.) M. Roem.; 33853 X d

ARECACEAE (7, 8)

*Chamaedorea sp. (sterile); Rare; 34244 X

*Cocos nucifera L. (seedlings); FLEPPC II; 34957 X r

*Phoenix sp. (sterile); Rare; 35409 X

Saba!palmetto (Walter) Lodd. ex Schult. & Schult. X, X pi

f.; 33546

Serenoa repens (W. Bartram) Small; 34214 X pi

Thrinaxmorrisii H. Wendl.; 33511 X, X pi

Thrinaxradiata Lodd. ex Schult. & Schult. f.; 34820 X

*Washingtonia sp. (seedlings); Rare; 35237

ASPARAGACEAE (1, 1)

*Asparagus aethiopicus L.; Rare; FLEPPC I; 33736 X

X X

240

Journal of the Botanical Research Institute of Texas 8(1)

RH Man Rock Rud Pow Dist Subm Berm

X,X man ,XP i

X,XP i ,X rb

XPi

x,x man

X,XP i ,X r

XP',X rb

x,x r

XPi

ASPHODELACEAE (7, 7)

*Aloevera L.; R.W. Long, F. Almeda, J. DeBoer, C.R. Broome 1859; 1966 (USF)

BROMELIACEAE (1, 8)

Tillandsia bolbisiono Schult. & Schult. f.; 34838

Tillandsia fasciculata Sw.; 35497 (photograph)

Tillandsia flexuosa Sw.; 34410

Tillandsiapaucifolia Baker; 34411

Tillandsia recurvata (L.) L.; 34504

Tillandsia setacea Sw.; 34800

Tillandsia usneoides (L.) L.; 33735

Tillandsia utriculata L.; 35498 (photograph)

COMMELINACEAE (1,1)

*Tradescantia spathacea Sw. [Rhoeo discolor

(L'Her.) Hance]; FLEPPC II; 33718

CYMODOCEACEAE (1,1)

Halodule wrightii Asch.; 34167

cyperaceae (5,15; 5, 16)

Cladium jamaicense Crantz; 33455

Cyperus compressus L.; Rare; 33822

Cyperuscroceus Vahl; G.A. Gann, J.A. Duquesnel 1146; 2003 (FTG)

*Cyperus esculentus L.; 34280

Cyperus flavescens L.; Rare; 33986

Cyperus ligularis L.; 33657

Cyperus odoratus L.; Rare; 33987

Cyperus ovatus Baldwin [Cyperus retrorsus

Chapm.]; 34801

Cyperusplanifolius Rich.; 33951 X r

Cyperus polystachyos Rottb.; 33348

* Cyperus rotund us L.; 34105

Eleocharis flavescens (Poir.) Urb.; Rare; 34098

Eleocharis geniculata (L.) Roem. & Schult.; Rare; 35500

Fimbristylis cymosa R.Br.; 33349

Fimbristylis spadicea (L.) Vahl [Fimbristylis castanea

(Michx.) Vahl]; 33588

Scleria lithosperma (L.) Sw.; 34903

HYDROCHARITACEAE (1, 1)

Thalassia testudinum Banks & Sol. ex J. Konig; 34168

ORCHIDACEAE (5, 5)

Encyclia tampensis (Lindl.) Small; 33826

*Eulophia graminea Lindl.; 33658

*Oeceoclades maculata (Lindl.) Lindl.; 33827

Prosthechea boothiana (Lindl.) W.E. Higgins;

35496 (photograph)

Vanilla barbellata Rchb. f.; Rare; 35499 (photograph)

poaceae (34,56; 35,59)

Andropogon glomeratus (Walter) Britton et al.; 33925

Andropogon virginicus L.; 34101

Aristida purpurascens Poir.; 34213

*Bothriochloa ischaemum (L.) Keng; 33926

*Bothriochloapertusa (L.) A. Camus; 33449

Cenchrus echinatus L.; 33952

Cenchrus spinifex Cav. [Cenchrus incertus M.A.

Curtis]; 33817

(hloriselata Desv.; G.N. Avery 1936; 1978 (USF)

*Cynodon dactylon (L.) Pers.; 33768

*Dactyloctenium aegyptium (L.) Willd. ex Asch.

&Schweinf.; FLEPPC II; 33344

Digitaria ciliaris (Retz.) Koeler var. ciliaris; 33345

Digitariainsularis (L.) Fedde [Trichachneinsularis (L.) Nees]; O. Lakela & L. Pardue 31593; 1968

Distichlis spicata (L.) Greene; 34731 X

XPi

X,XP j

x r

X b P

(USF)

X b P

X, X b P X

Wilder et al., Flora of Dagny Johnson Key Largo Hammock Botanical State Park

241

RH Man Rock Rud Pow Dist Subm Berm

*Echinochloa colona (L.) Link; Rare; 34558

*Echinochloo crus-galli (L.) P. Beauv.; Rare; 34559

*Eleusineindica (L.) Gaertn.; 33346

*Eragrostis amabilis (L.) Wight & Arn. ex Hook. & Arn.;

Rare; 34868

*Eragrostis ciliaris (L.) R.Br. var. ciliaris; 34579

*Eragrostis ciliaris (L.) R.Br. var. laxa Kuntze; Rare; 34163

Eragrostis elliottii S. Watson; 34082

*Eremochloa ophiuroides (Munro) Hack.; 34103

Eustachys petraea (Sw.) Desv.; 33655 X r

*Heteropogon contortus (L.) P. Beauv. ex Roem.

&Schult.; 33734

*Hyparrhenia rufa (Nees) Stapf; FLEPPC II; 34473 X r

Lasiacis divaricata (L.) Hitchc.; 33517 X

Leptochloafusca (L.) Kunth subsp. fascicularis (Lam.)

N. Snow; Rare; 35160

Leptochloa dubia (Kunth) Nees; 33451

*Me!inis repens (Willd.) Zizka [Rhynchelytrum

repens (Willd.) C.E. Hubb.]; FLEPPC I; 33821

Monanthochloe littoralis Englem.; Rare; 35399

Muhlenbergia capillaris (Lam.) Trin.; Rare; 34164

*Neyraudia reynaudiana (Kunth) Keng ex X d

Hitchc.; FLEPPC I; 34368

Oplismenus hirtellus (L.) P. Beauv.; 35232 X

Panicum dichotomiflorum Michx. var. bartowense

(Scribn. & Merr.) Fernald; Rare; 34557

'*Panicum maximum Jacq.; FLEPPC II; 34278 X r

*Panicum repens L.; Rare; FLEPPC I; 34104

Panicum virgatum L.; Rare; 34369 X

Paspalum blodgettii Chapm.;33818

Paspalum caespitosum Flugge; Rare; 34477

*Paspalum fimbriatum Kunth; O. Lakela & F. Almeda 30463; 1966 (USF)

*Paspalum notatum Flugge var. notatum; 33820

Paspalum setaceum Michx. var. longipedunculatum

(Leconte) Alph. Wood; 34419

Paspalum setaceum Michx. var. stramineum

(Nash) DJ. Banks; 33452

Paspalum vaginatum Sw.; 33769

*Pennisetum purpureum Schumach.; Rare;

FLEPPC I; 34398

*Pennisetum setaceum (Forssk.) Chiov.; Rare;

FLEPPC II; 34537

*RottboeHia cochinchinensis (Lour.) Clayton; 33552

Schizachyrium sanguineum (Retz.) Alston

var. sanguineum; 33642

Setaria macrosperma (Scribn. & Merr.) X

K. Schum.; 34736

Setariaparviflora (Poir.) Kerguelen

[Setaria geniculata P. Beauv.]; 33454

*Sorghum halepense (L.) Pers.; Rare; 34985

Spartina spartinae (Trin.) Merr. ex Hitchc.; 33972 X r X

Sporobolus dom ingen sis (Trin.) Kunth; 33832

^Sporobolus jacquemontii Kunth; 33927

Sporoboluspyramidatus (Lam.) Hitchc.; 33928

Sporobolus virginicus (L.) Kunth; 33553 X X

Stenotaphrum secundatum (Walter) Kuntze; Rare; 33547

Urochloa adspersa (Trin.) R.D. Webster; 34799

* Urochloa distachya (L.) T.Q. Nguyen

[Urochloa subquadripara (Trin.) R.D. Webster]; 33656

*Zoysiapacifica (Goudswaard) M. Hotta & Kuroki; 34400 X r

X,X b P

X b P X

X,X b P X X

X b P

X b P X X

X b P

X b P X X

X b P X

X b P X X

X X

X,X b P X X

X b P X

X X

X b P X X

X b P

X X

X b P

X b P X X

X X

X b P X X

X b P X

X X

X b P X

x,x rb

242

Journal of the Botanical Research Institute of Texas 8(1)

RH Man Rock Rud Pow Dist Subm Berm

RUPPIACEAE (1,1)

Ruppia maritima L.; 33512

RUSCACEAE (1,1)

*Sansevieria hyocinthoides (L.) Druce; X,X po ,X rb

FLEPPC II; 33738

SMILACACEAE (1,1)

Smilax havanensis Jacq.; 34904 X p ',X r

DICOTYLEDONS SENSU LATO (230, 307; 245,327)

ACANTHACEAE (2, 3,’ 3, 4)

*Asystasiagangetica (L.) T. Anderson; K.A. Bradley 1367; 1998 (FTG)

Dicliptera sexangularis (L.) Juss.; 34450

*Ruellia blechum L. [Blechum pyramidatum

(Lam.) Urb.]; Rare; FLEPPC II; 34539

*Ruellia ciliatiflora Hook.; Rare; 34106

AIZOACEAE (2, 2)

X r

x bp

Sesuvium portulacastrum (L.) L.; 33482

Trianthema portulacastrum L.; Rare; 35193

AMARANTHACEAE (4, 6,’ 5, 7)

Alternanthera flavescens Kunth; 33516

X,X r

*Amaranthus dubius Mart. exThell.; Rare; 34401

*Amaranthus spinosus L.; Rare; 34562

x bp

*Amaranthus viridis L.; 34600

X X

Blutaparon v'erm/culare (L.) Mears; 33483

Celosia nitida Vahl; Rare; 35362

Iresinediffusa Humb. & Bonpl. ex Willd.; R.W. Long 2981

; 1969 (USF)

ANACARDIACEAE (3, 3)

Metopium toxiferum (L.) Krug & Urb.; 33286

X pi , X r

x,x

: r x

*Schinus terebinthifolia Raddi; FLEPPC 1; 33484

X, x pi

Toxicodendron radicans (L.) Kuntze; Rare; 33772

x r

ANNONACEAE (1,1)

Annona glabra L.; 34420

x,x pi

APIACEAE (1, 1)

*Cyclospermum leptophyllum (Pers.) Sprague

x,x bp

ex Britton & P. Wilson; 34454

apocynaceae (5,5; 7, 7)

Harissamacrocarpa (Eckl.) A. DC.; R.W. Long, F. Almeda, J. De Boer, C.R. Broome 1769;

1966 (USF)

*Catharanthus roseus (L.) G. Don; Rare; 33554

Echites umbellatus Jacq.; 33645

x r

*Nerium oleander L.; F. Almeda, J. De Boer, C.R. Broome,

R.W. Long 1764;

1966 (USF)

Pentalinon luteum (L.) B. F. Hansen & Wunderlin; 33385

x pi ,x r

x r

Rhabdadenia biflora (Jacq.) Mull. Arg.; 33623

Vallesia antillana Woodson; Rare; 34541

ASCLEPIADACEAE (2, 3,’ 3, 4)

*Cryptostegiamadagascariensis Bojer ex Decne.; F. Almeda, J. De Boer, C.R. Broome,

R.W. Long 1761; 1966 (USF)

Cynanchum angustifolium Pers.; 33386

x,x r

Cynanchum scoparium; Rare; 35111

Sarcostemma clausum (Jacq.) Roem. & Schulte-

x r

Rare; 33555

ASTERACEAE (23, 29; 24,33)

Ambrosia artemisiifolia L.; Rare; 33773

Baccharis angustifolia Michx.; 34171

x r

Baccharis glomeruliflora Pers.; Rare; 34289

Baccharis halimifolia L.; 34109

x r

Bidens alba (L.) DC.; 33828

X,X bp X

Borrichia arborescens (L.) DC.; 33387

x r

Borrichia frutescens (L.) DC.; 33350

*Calyptocarpus vialis Less.; 34650

x bp

Chromolaena odorata (L.) R.M. King & H. Rob.; 34421

Conyza canadensis (L.) Cronquist; 34907

*Cyanthillium cinereum (L.) H. Rob. [Vernonia

cinerea (L.) Less.]; Rare; 34823

x rb

Wilder et al., Flora of Dagny Johnson Key Largo Hammock Botanical State Park

243

RH Man Rock Rud Pow Dist Subm Berm

Eclipto prostroto (L.) L.; Rare; 33829 X

Emilia fosbergii Nicolson; Rare; 33905

Erechtites hieraciifolius (L.) Raf. ex DC.; Rare; 34824 X bp

Eupatorium capillifolium (Lam.) Small ex

Porter & Britton; 34293

Eupatoriumserotinum Michx.; R.W. Long, F. Almeda, J. De Boer, C.R. Broome 1781; 1966 (USF)

Flaveria linearis Lag.; 33667 X

Flaveria trinervia (Spreng.) C. Mohr; 33669 X

*Helianthusannuus L.; D.S. Correll 48116; 1977 (FTG)

*Launaea intybacea (Jacq.) Beauverd; Rare; 35410

Melanthera nivea (L.) Small; 33668

Mikania scandens (L.) Willd.; Rare; 35307 X r

*Parthenium hysterophorus L.; 33351 X bp

Pectis glaucescens (Cass.) DJ. Keil; 33830

Pectisxfloridana D.J. Keil; G.A. Gann, K.A. Bradley 151; 1995 (FTG)

Pluchea carolinensis (Jacq.) G. Don; 33877 X bp

Plucheaodorata (L.) Cass.; R.W. Long, F. Almeda, J. De Boer, C.R. Broome 1841; 1966 (USF)

*Sonchus asper (L.) Hill; Rare; 34565 X bp

*Sonchus oleraceus L.; 33352 X bp

*Sphagneticola trilobata (L.) Pruski [Wedelia trilobata X

(L.) Hitchc.]; FLEPPC II; 33721

Symphyotrichum bahamense (Britton) G.L. Nesom X

[Symphyotrichum subulatum var. elongatum

(Bosserdet ex A.G. Jones & Lowry) S.D. Sundberg.];

34108

Symphyotrichum tenuifolium (L.) G.L. Nesom

[Symphyotrichum bracei (Britton ex Small)

G.L. Nesom]; 34107

*Tridaxprocumbens L.; 33879

AVICENNIACEAE (1, 1)

Avicennia germinans (L.) L.; 34183 X

BATACEAE (1, 1)

Batis maritima L.; 33545

BIGNONIACEAE (1, 1)

*Tecoma stans (L.) Juss. ex Kunth; Rare; 33556 X d

BORAGINACEAE (5, 7)

Argusia gnaphalodes (L.) Heine; 34651

Bourreria succulenta Jacq. [Bourreria ovata Miers]; X

34070

*Cordia sebestena L.; 34944 X rb

Heliotropium angiospermum Murray; 34456 X r

Heliotropium curassavicum L.; 33847

*Heliotropium procumbens Mill.; Rare; 34528

Toumefortia volubilis L.; 33337 X r

BRASSICACEAE (2, 2)

*Brassicajuncea (I.) Czern.; Rare; 34637

Lepidium virginicum L.; 33627

BURSERACEAE (1, 1)

Bursera simaruba (L.) Sarg.; 33543 X,X r

CACTACEAE (5, 7)

Acanthocereus tetragonus (L.) Hummelinck; 34494 X,X man ,X r

n Harrisia fragrans Small ex Britton & Rose; 34493 X,X man ,X rb

*Hylocereus undatus (Haw.) Britton & Rose; Rare; 35090 X r

*Opuntia cochenillifera (L.) Mill.; Rare; 34249

n Opuntia corallicola (Small) Werderm.; 35494 X

(photograph)

Opuntia stricta (Haw.) Haw.; 34435 X,X rb

*Se!enicereus pteranthus (Link & Otto) Britton & Rose; X d

Rare; 34825

CANELLACEAE (1, 1)

x r

x,x bp

x bp

yman

244

Journal of the Botanical Research Institute of Texas 8(1)

Man

Rock

Rud

Pow

Dist

Subm Berm

Canella winterana (L.) Gaertn.; 34250

X,X man ,XP°

CAPPARACEAE (1,2)

Capparis flexuosa (L.) L.; 34422

X ^man ^rb

Capparis jomoicensis Jacq.

[Capparis cynophallophora L.]; 33806

CARICACEAE (1, 1)

X,X man ,XP°

Carica papaya L.; 33629

CASUARINACEAE (1, 1)

*Casuarina equisetifolia L.; Rare; FLEPPC 1; 33671

CELASTRACEAE (4, 5)

X,XP°

Crossopetalum ilicifolium (Poir.) Kuntze; Rare; 34372

XPi

Crossopetalum rhacoma Crantz; 34583

XXP^X^X^

Hippocratea volubilis L.; 33628

X,X man ,XP i

Maytenusphyllanthoides Benth.; 34543

Schaefferia frutescens Jacq.; 34194

CELTIDACEAE (1, 2)

Trema lamarckiana (Schult.) Blume; Rare; 34334

X, XP°, x r

Trema micrantha (L.) Blume; 33509

x r

X b P

CHENOPODIACEAE (5, 5)

Atriplexpentandra (Jacq.) Standi. [Atriplexcristata

Humb. & Bonpl. ex Willd.]; 33490

tChenopodium sp. (Nonfruiting specimen); Rare; 35170

x rb

Salicornia bigelovii Torr.; 33357

Sarcocornia ambigua (Michx.) M.A. Alonso & M.B.

Crespo [Sarcocorniaperennis (Mill.)

AJ. Scott]; 33356

Suaeda linearis (Elliott) Moq.; 33491

CHRYSOBALANACEAE (1,1)

Chrysobalanus icaco L.; Rare; 34999

COMBRETACEAE (3, 4,’ 4, 5)

XPi

Conocarpus erectus L.; 33492

X,XP j

yman

Conocarpus erectus L. var. sericeus DC.; Rare; 34373

Laguncularia racemosa (L.) C.F. Gaertn.; 34185

Terminaliabuceras (L.) C. Wright[Bucidabuceras L.]; F.C. Craighead s.n.; 1961 (USF)

*Terminalia catappa L.; Rare; FLEPPC II; 33742

CONVOLVULACEAE (6, 14)

Dichondra sp. (sterile); Rare; 34620

X d

X b P

Evolvulus alsinoides (L.) L.; 34735

X b P

Ipomoea alba L.; 33558

Ipomoea hederifolia L.; Rare; 35412

Ipomoea imperati (Vahl) Griseb.; Rare; 33673

Ipomoea indica (Burm.) Merr.; 34374

X, X r

X b P

Ipomoea pes-caprae (L.) R.Br.; 33646

Ipomoea sagittata Poir.; Rare; 33390

XPi

x r

*lpomoea triloba L.; 33674

Ipomoea violacea L.; 33743

x r

X b P

Jacquemontia havanensis (Jacq.) Urb.; Rare; 33391

Jacquemontia pentanthos (Jacq.) G. Don; Rare; 34322

x r

*Merremia dissecta (Jacq.) Hallierf.; Rare; 33861

x r

Turbina corymbosa (L.) Raf.; Rare; 35309

CRASSULACEAE (1,1)

*Kalanchoe delagoensis Eckl. & Zeyh.; 33863

euphorbiaceae sensu lato (10,21)

nArgythamnia blodgettii (Torr. ex Chapm.)

Chapm.; Rare; 34926

Caperonia castaneifolia (L.) A. St.-Hil.; Rare; 33956

X b P

Chamaesyce blodgettii (Engelm. ex Hitchc.)

X,X b P

Small; 33881

n Chamaesyce conferta Small; 34883

X,X b P

Chamaesyce hirta (L.) Millsp.; 33882

X,X b P

Wilder et al., Flora of Dagny Johnson Key Largo Hammock Botanical State Park

245

RH Man Rock Rud Pow Dist Subm Berm

X,XP i ,XP°,X r

X, XP°, x r

X,X b P

X b P

X,X b P

X b P

Chamaesyce hypericifolia (L.) Millsp.; 33907 X,X bp

^Chamaesyce mendezii (Boiss.) Millsp.; 34405 X bp

Chamaesyce mesembrianthemifolia (Jacq.) X d

Dugand; 34072

Chamaesyce ophthalmica (Pers.) D.G. Burch; 33358

Chamaesyce prostrata (Aiton) Small; 33395

Drypetes diversifolia Krug & Urb.; 33542

Drypetes lateriflora (Sw.) Krug & Urb.; 34487

*Euphorbia graminea Jacq.; 33885

* Euphorbia trigona Haw.; Rare; 34412

Gymnanthes lucida Sw.; 34681

Hippomane mancinella L.; Rare; 34692

*Phyllanthus amarus Schumach. &Thonn.; 33397

*Phyllanthus tenellus Roxb.; 33632

Poinsettia cyathophora (Murray) Bartl.; 33497

Poinsettia heterophylla (L.) Klotzsch & Garcke

ex Klotzsch; Rare; 34587

*Ricinus communis L.; Rare; FLEPPC II; 33633

fabaceae (31,43; 32,44)

Acacia choriophylla Benth.; Rare; 34628

Acacia farnesiana (L.) Willd.; 33729

Acacia pinetorum FJ. Herm.; Rare; 34529

*Albizia lebbeck (L.) Benth.; Rare; FLEPPC I; 34458

*Alysicarpus vaginalis (L.) DC.; 34910 X bp

Caesalpinia bonduc (L.) Roxb.; 33779

*Cajanus cajan (L.) Huth; Rare; 34659

Canavalia rosea (Sw.) DC.; 33780

Centrosemavirginianum (L.) Benth.; F. Almeda, J. De Boer, C.R. Broome, R.W. Long 1742; 1966 (USF)

Chamaecrista nictitans (L.) Moench var. aspera

(Muhl. ex Elliott) H.S. Irwin & Barneby; Rare; 34199

*Clitoria ternatea L.; Rare; 33560

*Crotalaria incana L.; Rare; 34721

Crotalaria pumila Ortega; 33293 X

*Crotalaria spectabilis Roth; Rare; 33908

Dalbergia brownei (Jacq.) Schinz; 34295

Dalbergia ecastaphyllum (L.) Taub.; 33398

*Delonix regia (Bojer ex Hook.) Raf.; Rare; 34912

*Desmanthus leptophyllus Kunth; 33539

X r

X d

X r

X, X r

X,X man

XP°,X r

X r

Desmanthus virgatus (L.) Willd.; 33701

*Desmodium incanum DC.; 34888

*Desmodium tortuosum (Sw.) DC.; 33746

*Desmodium triflorum (L.) DC.; 34739

Erythrina herbacea L.; Rare; 35383 X

Galactia striata (Jacq.) Urb.; 34299 X r

*lndigofera spicata Forssk.; 34520

*lndigofera tinctoria L.; Rare; 34325

*Leucaena leucocephala (Lam.) de Wit;

FLEPPC II; 34187

Lysiloma latisiliquum (L.) Benth.; 33959 X,X p ',X r

*Macroptilium lathyroides (L.) Urb.; 34117

*Meiiiotus albus Medik.; 34544

Neptuniapubescens Benth.; 33537

*Parkinsonia aculeata L.; Rare; 34238

Piscidia piscipula (L.) Sarg.; 33886 X

Pithecellobium keyense Britton ex Britton & Rose; 34459 X,X p ',X r

Pithecellobium unguis-cati (L.) Benth.; 34691 X,X r

Rhynchosia minima (L.) DC.; 33363

Rhynchosia swartzii (Vail) Urb.; 34376 X,X d ,X po ,X r

Senna ligustrina (L.) H.S. Irwin & Barneby; 34803

*Senna obtusifolia (L.) H.S. Irwin & Barneby; Rare; 33912

X b P

X,X b P

X r

X b P

X,X b P

X b P

246

Journal of the Botanical Research Institute of Texas 8(1)

RH Man Rock Rud Pow Dist Subm Berm

Sesbania vesicorio (Jacq.) Elliott; Rare; 35183

Sophora tomentosa L.; 33535 X p ',X r

*Stylosanthes homoto (L.) Taub.; 34407

*Tamarindus indica L.; Rare; 34950 X

Vigno luteolo (Jacq.) Benth.; 33784

FAGACEAE(1, 1)

Quercus virginiana Mill.; 34200 X pi

GENTIANACEAE (3, 3)

Eustoma exaltatum (L.) Salisb. ex G. Don; 33887

Sabatiastellaris Pursh; 33677 X pi

Voyria parasitica (Schltdl. & Cham.) Ruyters & Maas X

[Leiphaimosparasitica Schltdl. & Cham.]; 34026

GOODENIACEAE (1, 2)

*Scaevola taccada (Gaertn.) Roxb. var. sericea

(Vahl) H. St. John; FLEPPC I; 34607

*Scaevola taccada (Gaertn.) Roxb. var. taccada;

Rare; FLEPPC I; 33869

HYDROPHYLLACEAE (1,1)

*Nama jamaicensis L.; Rare; 34517

LAMIACEAE (2, 2) 3, 3)

Callicarpa americana L.; 33902

*Hyptis mutabilis (Rich.) Briq.; Rare; 34666

*Vitextrifolia L.; G.A. Gann, J.A. Duquesnel 1159; 2003 (FTG)

LAURACEAE (1,1)

Ocotea coriacea (Sw.) Britton; 33915 X r

MALPIGHIACEAE (1, 1)

Byrsonima lucida (Mill.) DC.; 34812 X pi , X r

MALVACEAE (7, 9)

Abutilon permolle (Willd.) Sweet; Rare; 34773

Gossypium hirsutum L.; 34379 X rb

Herissantia crispa (L.) Brizicky; 34829

* Hibiscus rosa-sinensis L. var. rosa-sinensis; Rare; 34326 X r

Malvastrum corchorifolium (Desr.) Britton ex

Small; 34423

Sida antillensis Urb.; 34591

Si da ci Haris L.; 34891

*Sida sp. (apparently new to Florida); Rare; 33747

*Thespesia populnea (L.) Sol. ex Correa; X

FLEPPC I; 33401

MELIACEAE (1, 1;2,2)

*Meliaazedarach L.; R.P. Sauleda, D.K. Sauleda 4742; 1981 (FTG)

Swietenia mahagoni (L.) Jacq.; 33617 X, X pi

MORACEAE (1, 3)

Ficus aurea Nutt.; 34188 X,X p ',X r

Ficus citrifolia Mill.; 34382 X r

*Ficus microcarpa L. f.; Rare; FLEPPC I; 34462 X

MYRICACEAE (1, 1)

Myrica cerifera L.; 34202 X pi

MYRSINACEAE (2, 2)

Ardisia escallonioides Schiede & Deppe ex X,X pi

Schltdl. & Cham.; 33618

Myrsine cubana A. DC. [Rapanea punctata X, X pi

(Lam.) Lundell]; 33712

MYRTACEAE (2, 6)

Calyptranthes pallens Griseb.; 33580 X,X p ',X po ,X r

Calyptranthes zuzygium (L.) Sw.; 34056 X,X pi

Eugenia axillaris (Sw.) Willd.; 34439 X,X p ',X r

Eugenia confusa DC.; 34208 X,X man

Eugenia foetida Pers.; 33891 X,X pi

Eugenia rhombea Krug & Urb. ex Urb.; Rare; 34531 X

X r ,X rb X

X r

X r X

X d

x bp

X X

x bp

X,X bp X X

XXX

X bp X

x bp

x,x rb

yman

Wilder et al., Flora of Dagny Johnson Key Largo Hammock Botanical State Park

247

RH Man Rock Rud Pow Dist Subm Berm

NYCTAGINACEAE (4, 4)

Boerhovio diffusa L.; Rare; 34029

*Bougainvillea glabra Choisy; Rare; 34440

Guapira discolor (Spreng.) Little; 34032

Pisonia aculeata L.; 34953

OLEACEAE (1, 1)

*Jasminum dichotomum\fah\; Rare; FLEPPC I; 33785

ONAGRACEAE (1,1)

Gaura angustifolia Michx.; Rare; 33786

OXALIDACEAE (1, 1)

Oxalis corniculata L.; 33637

PAPAVERACEAE (1, 1)

Argemone mexicana L.; 33750

PASSIFLORACEAE (1, 2)

Passiflora multiflora L.; 34514

Passiflora suberosa L.; 33529

PHYTOLACCACEAE (1,1; 2,2)

Phytolaccaamericana L.; R.W. Long, F. Almeda, J. De B

Rivina humilis L.; 33814

PITTOSPORACEAE (1,1)

*Pittosporum sp. (sterile); Rare; 33880

PLANTAGINACEAE (1,1)

*Plantago major L.; Rare; 34549

PLUMBAGINACEAE (1,1)

Limonium carolinianum (Walter) Britton; 33870

POLYGONACEAE (1,2)

Coccoloba diversifolia Jacq.; 33916

Coccoloba uvifera (L.) L.; 33527

PORTULACACEAE (2, 2)

Portulaca oleracea L.; 33680

*TaHnum fruticosum (L.) Juss.; Rare; 34071

RHAMNACEAE (4, 5,’ 4, 6)

*Colubrina asiatica (L.) Brongn.; FLEPPC I; 33754

Colubrinacubensis (Jacq.) Brongn. var. floridana M.C.

Colubrina elliptica (Sw.) Brizicky & W.L. Stern; 34175

Gouania lupuloides (L.) Urb.; 33502

Krugiodendron ferreum (Vahl) Urb.; 33713

Reynosiaseptentrionalis Urb.; 34834

RHIZOPHORACEAE (1,1)

Rhizophora mangle L.; 33815

RUBIACEAE (12, 16)

Chiococca alba (L.) Hitchc.; 33917

Erithalis fruticosa L.; 34489

Ernodea cokeri Britton ex Coker; 34394

Exostema caribaeum (Jacq.) Schult.; 34445

Guettarda elliptica Sw.; 33892

Guettarda scabra (L.) Vent.; 33757

Hamelia patens Jacq.; 33730

Morinda royoc L.; 33461

*Oldenlandia corymbosa L.; Rare; 34409

Psychotria ligustrifolia (Northr.) Millsp. [Psychotria

bahamensis Millsp.]; 34816

Psychotria nervosa Sw.; 33919

Randia aculeata L.; 33410

*Richardia grandiflora (Cham. & Schltdl.) Schult.

& Schult. f.; Rare; 34918

Spermacoce remota Lam. [Spermacoce assurgens

Ruiz & Pav.]; 33893

Spermacoce tetraquetra A. Rich.; 33564

*Spermacoce verticillata L.; 33897

X,X b P

X r

X,X d ,XP i ,X r X d ,X r X,X r

X,X r

X r

X b P

X,XP°,X r

X X,X b P

r, C.R. Broome 1843; 1966 (USF)

x r ,x man X

x d

X b P

X X

X, XP j , x r

X man ( XPi,X r x r

X,X b P

X X X b P

Johnst.; F.C. Craighead s.n.; 1961 (USF)

X r

XP°,X r

X,X man ,XP°,X r x d

X,X man ,XP i ,X r ,X rb

X I XX

X,X d ,XP i ,X r

x pi , x r

X pi X P'" r

XP°,X r

X,XP°,X r

x r

X,XP i ,X r

X b P

X,XP i ,XP°,X r

x,x r

X,X man ,XP i

x,x r

X b P

248

Journal of the Botanical Research Institute of Texas 8(1)

RH Man Rock Rud Pow Dist Subm Berm

RUTACEAE (3, 3)

Amyris elemifero L.; 33966

* Citrus sp. (sterile); Rare; 34428

Zanthoxylum fagara (L.) Sarg.; 33524

SALICACEAE (1, 1)

*Flacourtia indica (Burm. f.) Merr.; Rare;

FLEPPC II; 34461

SAMOLACEAE (1,1)

Samolus ebracteatus Kunth; Rare; 33369

SAPINDACEAE (5, 6) 5, 7)

Cordiospermum corindum L.; 33797

X,XP°,X r

X r

X,X man ,XP°,X r

X d ,X r

X r

X, X r

X,XP°,X r

X,XP°,X r ,X rb

X,X r

X,XP j

X,XP°

x,x r

ypi (in opening) y

x r

Cardiospermummicrocarpum Kunth.; F.C. Craighead s.n.; 1962 (USF)

Dodonaea elaeagnoides Rudolphi ex Ledeb. X pi ,X r

&Alderstam; 33799

Dodonaea viscosa Jacq.; Rare; 33507

Exotheapaniculata (Juss.) Radik.; 34465

Hypelate trifoliata Sw.; 34598

Sapindussaponaria L.; 33731

SAPOTACEAE (3, 6)

Chrysophyllum oliviforme L.; 33523

Manilkara jaimiqui (C. Wright ex Griseb.)

Dubard; 33698

* Manilkara zapota (L.) P. Royen; FLEPPC I; 34432

Sideroxylon celastrinum (Kunth) T.D. Penn.; 33412

Sideroxylon foetidissimum Jacq. [Mastichodendron

foetidissimum (Jacq.) HJ. Lam.; 34364

Sideroxylon salicifolium (L.) Lam.; 33800

SCHOEPFIACEAE (1, 1)

Schoepfia chrysophylloides (A. Rich.) Planch.; 34267

scrophulariaceae sensu lato (7,7)

Agalinis maritima (Raf.) Raf.; 34067

Bacopa monnieri (L.) Pennell; 34068

Buchnera americana L.; Rare; 34047

Capraria biflora L.; 33371

Mecardoniaprocumbens (Mill.) Small; Rare; 34777

*Russelia equisetiformis Schltdl. & Cham.; 33899

Scoparia dulcis L.; Rare; 35251

SIMAROUBACEAE (1,1)

Simarouba glauca DC.; 33900 X

SOLANACEAE (3, 7) 6, 10)

Capsicumannuum L. var. glabriusculum (Dunal) Heiser & Pickersgill; J.E. Poppleton & A.G. Shuey s.n.; 1974 (USF)

*Daturametel L.; G.A. Gann, J.A. Duquesnel 1152; 2003 (FTG)

Lycium carolinianum Walter; 33874 X,X r X

*Nicotianatabacum L.; F. Almeda, J. De Boer, C.R. Broome, R.W. Long 1758; 1966 (USF)

Physalis pubescens L.; Rare; 34978

Solanum americanum Mill.; Rare; 33762

Solanum bahamense L.; Rare; 34727

Solanum donianum Walp.; Rare; 33801

Solanum erianthum D. Don; 34835

*Solanum lycopersicum L. [Lycopersicon esculentum

Mill.]; Rare; 33732

STERCULIACEAE (2, 2) 3, 3)

Ayeniaeuphrasiifolia Griseb.; G.A. Gann, J.A. Duquesnel 1140; 2003 (FTG)

*Melochia corchorifolia L.; Rare; 35181

Waltheria indica L.; 34850

STRYCHNACEAE(1,1)

Spigelia anthelmia L.; 34189

SURIANACEAE (1,1)

Suriana maritima L.; 33802

TETRACHONDRACEAE (1,1)

X b P

X,X b P

X b P

X,X b P X

x d

Wilder et al., Flora of Dagny Johnson Key Largo Hammock Botanical State Park

249

Man

Rock

Rud

Pow

Dist

Subm Berm

Polypremum procumbens L.; 33489

THEOPHRASTACEAE (1, 1)

Jacquinia keyensis Mez; 35034

TILIACEAE (1, 1)

Corchorus siliquosus L.; 34524

TURNERACEAE(1, 1)

XPi

X,X b P

*Tumera ulmifolia L.; 33689

URTICACEAE (2, 3)

Parietaria floridana Nutt.; Rare; 34629

X b P

Parietariapraetermissa Hinton; Rare; 33766

Pilea microphylla (L.) Liebm.; Rare; 35182

VERBENACEAE (6, 8)

Citharexylum spinosum L.; 33804

x r

X r

*Duranta erecta L.; Rare; 33693

x r

*Lantana camara L.; FLEPPC 1; 33415

x r

Lantana involucrata L.; 33372

XP°,X r

Phyla nodiflora (L.) Greene; 33903

X b P

Priva lappulacea (L.) Pers.; 33767

X b P

*Stachytarpheta cayennensis (Rich.) Vahl;

Rare; FLEPPC II; 34550

Stachytarphetajamaicensis (L.) Vahl; 34610

VISCACEAE (1, 1)

Phoradendron rubrum (L.) Griseb.; 35495

X, XP j - r

(photograph)

VITACEAE (3, 3)

Cissus verticillata (L.) Nicolson & C.E. Jarvis; Rare; 34980

Parthenocissus quinquefolia (L.) Planch.; 33510

x,x r

Vitis rotundifolia Michx.; 33904

X b P

XIMENIACEAE (1,1)

Ximenia americana L.; 34243

ZYGOPHYLLACEAE (3, 3)

X,XP j ,X r

Guaiacum sanctum L.; 33921

Kallstroemia maxima (L.) Hook. & Arn.; Rare; 34045

*Tribulus cistoides L.; Rare; FLEPPC II; 33665

X, XP°

X b P

ACKNOWLEDGMENTS

We express deep appreciation to Brian Holley and the Naples Botanical Garden for providing laboratory space

for the present study and for housing the SWF Herbarium. We also thank the following individuals and insti¬

tutions for their assistance: Michael Duever; Jim Duquesnel; Allison Duran; Hanina Epstein; Trudy Ferraro,

Pat Wells and the State of Florida Department of Environmental Protection Division of Recreation and Parks;

Peter Frezza, Jerome Lorenz and Audubon, Florida; Susan Gallagher; George Gann and the Institute for Re¬

gional Conservation; Sara Hamilton; Roger Hammer; Bruce Hansen and the USF Herbarium; Patricia Howell;

Brett Jestrow and the FTG Herbarium; Bibijabar; Duane Kolterman; Cyril Marks; Randy Mears; Nathan Mitch¬

ell; Andee Naccarato; George Newman; Karen Relish; Jean Roche; Eileen Watkins; Rebecca Wilder; Jerry

Wilkinson; Emily Wilson; and Joseph Wipff. Too, we express deep appreciation to Loran Anderson and Alan

Franck for their careful reviews of the manuscript of this paper.

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BOOK NOTICE

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Americas, New York, New York 10013-2473, U.S.A. (Orders: www.cambridge.org, 1-845-353-7500).

$99.00,378 pp., 88 b&w illus., 40 color illus., 9 tables, taxonomic and subject indices, 7 V 2 " x 10".

From the publisher: Tracing the evolution of one of the most ancient major branches of flowering plants, this is

a wide-ranging survey of state-of-the-art research on the early clades of the monocot phylogenetic tree. It ex¬

plores a series of broad but linked themes, providing for the first time a detailed and coherent view of the taxa

of the early monocot lineages, how they diversified and their importance in monocots as a whole. Featuring

contributions from leaders in the held, the chapters trace the evolution of the monocots from largely aquatic

ancestors. Topics covered include the rapidly advancing held of monocot fossils, aquatic adaptations in pollen

and anther structure and pollination strategies and floral developmental morphology. The book also presents

a new plastid sequence analysis of early monocots and a review of monocot phylogeny as a whole, placing in an

evolutionary context a plant group of major ecological, economic and horticultural importance.

■ The hrst detailed modern account of the early evolution of the monocots, a plant group of major ecologi¬

cal, economic and horticultural importance which includes grasses, cereals, palms, orchids and yams.

■ Leading scientists in the held provide up-to-date reviews, showing readers how iconic plants like orchids,

lilies and palms originated from largely aquatic ancestors.

■ Includes the hrst survey of early monocot fossils and the new insights they yield on the evolution of Low¬

ering plants, providing an accessible account of the latest advances in this rapidly developing held.

PAUL WILKIN is Lilioid and Alismatid Monocots and Ferns Team Leader in the Herbarium, Library, Art and

Archives Directorate of the Royal Botanic Gardens, Kew. His main research foci are systematics of Dioscoreales

(yams and their allies) and Dracaenoids (dragon trees and mother-in-law’s tongues), lilioid monocots widely

used in human diet and horticulture, with taxa of high conservation and ecological importance. He is principal

investigator of the eMonocot biodiversity informatics project.

SIMON J. MAYO is an Honorary Research Associate at the Royal Botanic Gardens, Kew. Since 1977 he has

worked on the systematics and phylogeny of the Araceae, the largest plant family of the early divergent clades

in monocots. He has been active in postgraduate teaching in Brazilian universities since 1988, focusing on

monocot families and especially the Araceae.

Contributors: W.J.D. lies, S.Y. Smith, S.W. Graham, D.D. Sokoloff, M.. Remizowa, P.J. Rudall, C.A. Furness, F.A.

Jones, B. Sobkowiak, C.D.L. Orme, R. Govaerts, V. Savolainen, D.H. Les, N.P. Tippery, N.Tanaka, K. Uehara, J.

Murata, S.J. Mayo,J. Bogner, N. Cusimano, D. Barabe, R.W. Scotland, M.G. Sajo, R. Mello-Silva, J.I. Davis, J.R.

McNeal, C.F. Barrett, M.W. Chase, J.I. Cohen, M.R. Duvall, T.J. Givnish, G. Petersen, J.C. Pires, O. Seberg,

D.W.M. Stevenson, J. Leebens-Mack

J.Bot. Res. Inst. Texas 8(1): 252.2014

A QUANTITATIVE STUDY OF THE VEGETATION SURROUNDING

POPULATIONS OF ZIGADENUS DENSUS (MELIANTHIACEAE)

AT FORT POLK IN WEST CENTRAL LOUISIANA, U.S.A.

Jacob Delahoussaye, Charles Allen, Stacy Huskins, and Ariel Dauzart

Colorado State University-Fort Polk

1645 23rd Street

Fort Polk, Louisiana 71459, U.S.A.

charles.m.allen l.ctr@mail.mil

ABSTRACT

Osceola’s plume ( Zigadenus densus (Desr.) Fern) is a rare species (S2) in Touisiana with reports from Natchitoches, St. Tammany, Vernon,

and Washington parishes. Quantitative data from 40 populations (clumps) are reported, and the plant community associated with Z. densus

is described. In west central Touisiana, Z. densus is found along the natural levee of baygall streams and is associated with the tree species:

Magnolia virginiana, Pinus palustris, and Nyssa biflora; the shrub species Morelia caroliniensis, Rhododendron oblongifolium, and Acer rubrum;

the woody vine species Rubus argutus, Gelsemium sempervirens, and Smilax laurifolia; and the herbaceous species Dichanthelium dichotomum

var. tenue, Eupatorium rotundifolium, and Osmunda regalis.

RESUMEN

Zigadenus densus (Desr.) Fern es una especie rara (S2) en Tuisiana con citas en la parroquias de Natchitoches, St. Tammany, Vernon, y Wash¬

ington. Se citan datos cuantitativos de 40 poblaciones (matas), y se describe la comunidad vegetal asociada con Z. densus. En el centra oeste

de Tuisiana, Z. densus se encuentra a lo largo de las riveras naturales de torrentes y esta asociado con tres especies: Magnolia virginiana, Pinus

palustris, y Nyssa biflora; las especies arbustivas Morelia caroliniensis, Rhododendron oblongifolium, y Acer rubrum; Fas especies trepadoras

Rubus argutus, Gelsemium sempervirens, y Smilax laurifolia; y las especies herbaceas Dichanthelium dichotomum var. tenue, Eupatorium rotun¬

difolium, y Osmunda regalis.

INTRODUCTION

Osceola’s Plume ( Zigadenus densus (Desr.) Fern) is a rhizomatous herbaceous perennial, 4 to 20 dm in height

with a distinct blue-green foliage color, flowering from March to July. Other common names include black

snakeroot, death camasa, and crow poison. It is reported from Alabama, Delaware, Florida, Georgia, Louisi¬

ana, Mississippi, North Carolina, South Carolina, Tennessee, Texas, and Virginia (USDA NRCS 2013; Nature

Serve 2013). In Flora of North America North of Mexico (Schwartz 2003), it is also reported from Kentucky, New

Jersey, New York, and West Virginia. Globally, Osceola’s plume is ranked G5, but it can be rare in certain parts

of its range; it is ranked SI in VA, S2 in LA, S3S4 in GA, and S4 in NC (Nature Serve 2013). In Louisiana, this S2

species is found in Natchitoches, St. Tammany, Vernon, and Washington parishes (Louisiana Natural Heritage

Program 2011; MacRoberts et al. 2002; Thomas & Allen 1993). In Texas, Z. densus is reported from Anderson,

Henderson, and Tyler counties (Diggs et al. 2006).

This species is reported from hillside bogs and longleaf flatwoods savannahs in Louisiana (Louisiana

Natural Heritage Program 2011) and from wet areas, damp pinelands, and bogs in east Texas (Diggs et al.

2006). The habitats reported for this species in the Flora of North America are pine bogs and flatlands

(Schwartz 2003). In the Carolinas, it is reported from savannahs and pocosins (Radford et al. 1968) and God¬

frey and Wooten (1979) report it from pine savannahs, flatwoods, and bogs.

The objectives of this study were to document the species associated with Z. densus and to describe its

habitat in west central Louisiana. Several populations of Z. densus are known from Fort Polk in Vernon Parish,

west central Louisiana.

J. Bot. Res. Inst. Texas 8(1): 253 - 259.2014

254

Journal of the Botanical Research Institute of Texas 8(1)

METHODS

In May to mid-June 2013, 40 populations (clumps) of Z. densus were surveyed on Fort Polk; most populations

(clumps) had previously been found during vegetation surveys. Most (23) of the 40 sampling sites were located

on Ruston fine sandy loam, others were found on Osier loamy fine sand (5/40), Malbis fine sandy loam (4/40),

Eastwood silt loam (4/40), Beds loamy fine sand (2/40), and Guyton-Iuka complex (2/40) (Soil Survey Division

2003). The area around each population was examined and the 5 nearest individuals were identified and re¬

corded for each of 5 categories including herbaceous, woody vine, shrub (woody non-vines shorter than 1.83

m), shrub/sapling (woody non-vines taller than 1.83 m and dbh less than 5 in), and tree (woody non-vines

taller than 1.83 m and dbh larger than 5 in). The total number of individuals recorded for each plant category

was 200 (40 populations and 5 individuals per population). The relative abundance was calculated by dividing

the number of times a species was recorded by 200, the total number of observations. This was done for all 5

categories and converted to a percent (Allen et al. 2006).

During the same survey, we also recorded the presence of all herbaceous species within 1 m of the center

of the Z. densus population. We didn’t record the sizes of the clumps, but we approximate the average size of the

clumps was about one square meter, with each clump having on average ten to fifteen stems. All woody vine

and all shrub species within 4 m of the center were recorded. All tree or shrub/sapling species within 15 m of

the center were recorded. The relative frequency was calculated by dividing the number of occurrences for a

species by the total number of occurrences recorded for the category. This value was converted to a percent.

Importance value (total = 200) for herbaceous, woody vines, and shrubs was calculated by adding the

relative frequency and the relative abundance (Megyeri & Allen 2011). Importance value (total = 300) for trees/

shrub saplings was calculated by adding the relative abundance for each species in the shrub/sapling category

and tree category to the relative frequency of each species in the tree/shrub/sapling category.

RESULTS AND DISCUSSION

A total of 168 species were recorded including 81 herbaceous, 12 woody vines, and 75 woody non-vines (39

shrubs and 36 trees; note overlap in categories). The species are listed by decreasing importance value. The

most important herbaceous species were Dichanthelium tenue (27.74), Eupatorium rotundifolium (14.74), and

Osmunda regalis (13.55) (Table 1). The woody vine species with the highest importance value were Rubus ar-

gutus (52.38), Gelsemium sempervirens (32.62), and Smilax laurifolia (32.42) (Table 2). The most important

shrub species were Morelia caroliniensis (22.12), Rhododendron oblongifolium (20.29), and Acer rubrum (19.44)

(Table 3). The shrub/sapling and tree species with the highest importance value were Magnolia virginiana

(54.39) and Pinus palustris (53.53) (Table 4).

The Zigadenus densus habitat in west central Louisiana is best described as the natural levee and seepage

areas upslope of baygall streams with the vegetation being typical of such a stream. We found the tree canopy

vegetation associated with Z. densus to be Magnolia virginiana, Pinus palustris, Nyssa biflora, and Acer rubrum

var. drummondii and the shrub canopy to be Morelia caroliniensis, Rhododendron oblongifolium, and Acer ru¬

brum. These woody species are similar to the species reported for baygalls in the area (Allen et al. 2004), in east

Texas (Diggs et al. 2006), and in central Louisiana (MacRoberts et al. 2004).

Allen et al. (2013) and MacRoberts et al. (2004) report two ( Magnolia virginiana and Nyssa biflora) of the

top three tree/shrub species in importance value to be the same as in our study. Our study shares 10 of the top

20 species of shrub/saplings with Allen et al. (2013)’study of yellow root. Allen et al. (2013) shares 8 of the 12

woody vine species with our study, but the 3 most important woody vines in association with Z. densus are the

3 least important woody vines in association with yellow root. Allen et al. (2013) has 6 of the top 15 herbaceous

species in common with our study of Z. densus in their yellow root study.

Our data are the first quantitative report on the vegetation surrounding Zigadenus densus. The vegetation

around the other populations of Zigadenus densus throughout its range should be sampled for comparison with

our data so as to get a better idea of the variation, if any, of its habitat.

Delahoussaye et al., Vegetation studies of Zigadenus densus

255

Table 1. Relative abundance, relative frequency, and importance value for herbaceous species recorded from AO Zigadenus densus sites at Fort Polk in west central

Louisiana.

Species

Relative

Abundance

Relative

Frequency

Importance

Value

Dichanthelium dichotomum (L.) Gould var. tenue (Muhl.) Gould & C.A. Clark

19.50

8.24

27.74

Eupatorium rotundifolium L.

6.50

8.24

14.74

Osmunda regolis L.

9.00

4.55

13.55

Scleria spp.

7.50

3.69

11.19

Mitchello repens L.

5.00

3.41

8.41

Dichanthelium scabriusculum (Ell.) Gould & C.A. Clark

5.00

2.84

7.84

Osmunda cinnamomea L.

3.50

4.26

7.76

Symphyotrichum lateriflorum (L.) A.& D. Love

2.50

4.26

6.76

Woodwardia virginica (L.) Sm.

4.00

2.27

6.27

Viola x primulifolia L. (pro sp.)

1.00

4.26

5.26

Solidago rugosa P. Mill.

3.00

1.70

4.70

Solidago patula Muhl. ex Willd.

1.50

3.13

4.63

Doellingeria sericocarpoides Small

2.50

1.99

4.49

Coreopsis tripteris L.

2.00

2.27

4.27

Dichanthelium acuminatum (Sw.) Gould & C.A. Clark var. acuminatum

2.50

1.14

3.64

Woodwardia areolata (L.) T. Moore

2.50

1.14

3.64

Sarracenia alata Wood.

1.50

1.99

3.49

Rhexia alifanus Walt.

2.50

0.85

3.35

Centella erecta (L. f.) Fern.

1.50

1.70

3.20

Oxypolis rigidior (L.) Raf.

0.50

2.56

3.06

Chasmanthium laxum (L.) Yates

1.00

1.99

2.99

Pteridium aquilinum (L.) Kuhn

1.50

1.42

2.92

Carex lonchocarpa Willd.

2.00

0.57

2.57

Helianthus angustifolius L.

1.00

1.42

2.42

Schizachyrium scoparium (Michx.) Nash

1.00

1.42

2.42

Dichanthelium dichotomum (L.) Gould var. dichotomum

0.50

1.70

2.20

Scleria pauciflora Muhl. ex Willd.

1.00

1.14

2.14

Carexglaucescens Ell.

0.50

1.42

1.92

Eupatorium leucolepis (DC.) Torr. & A. Gray

0.50

1.42

1.92

Melanthium virginicum L.

1.00

0.85

1.85

Panicum virgatum L.

1.00

0.85

1.85

Eupatorium fistulosum Barratt

0.50

1.14

1.64

Rudbeckia scabrifolia L.E. Brown

1.00

0.57

1.57

Lobeliapuberula Michx. var. puberula

0.00

1.42

Mitreola sessilifolia (J.F. Gmel.) G. Don

0.00

1.14

Elephantopus nudatus A. Gray

0.50

0.57

1.07

Eupatorium perfoliatum L.

0.50

0.57

1.07

Gentiana saponaria L.

0.50

0.57

1.07

Ptilimnium costatum (Ell.) Raf.

0.50

0.57

1.07

Scutellaria integrifolia L.

0.00

0.85

Xyris spp.

0.00

0.85

Ambrosia psilostachya DC.

0.50

0.28

0.78

Dichanthelium laxiflorum (Lam.) Gould

0.50

0.28

0.78

Paspalum spp.

0.50

0.28

0.78

Scleria oligantha Michx.

0.50

0.28

0.78

Ambrosia artemisiifolia L.

0.00

0.57

Athyrium filix-femina (L.) Roth

0.00

0.57

Ctenium aromaticum (Walt.) Wood

0.00

0.57

Dichanthelium sphaerocarpon (Ell.) Gould

0.00

0.57

Oldenlandia uniflora L.

0.00

0.57

Rhexia petiolata Walt.

0.00

0.57

Rhynchospora spp.

0.00

0.57

Aristida purpurascens Poir. var. purpurascens

0.00

0.28

Asclepias rubra L.

0.00

0.28

Calopogon tuberosus (L.) B.S.P.

0.00

0.28

Centrosema virginianum (L.) Benth.

0.00

0.28

Chamaecrista fasciculata (Michx.) Greene

0.00

0.28

Chaptalia tomentosa Vent.

0.00

0.28

256

Journal of the Botanical Research Institute of Texas 8(1)

Table 1.continued

Species

Relative

Abundance

Relative

Frequency

Importance

Value

Coreopsis glodioto Walt.

0.00

0.28

Dichanthelium aciculare (Desv. ex Poir.) Gould & C.A. Clark

0.00

0.28

Dichanthelium commutatum (J.A. Schultes) Gould

0.00

0.28

Dichanthelium polyanthes Schult

0.00

0.28

Eryngium integrifolium Walt.

0.00

0.28

Euphorbia corollata L.

0.00

0.28

Galactia volubilis (L.) Britt.

0.00

0.28

Liatris spicata (L.) Willd.

0.00

0.28

Linum medium (Planch.) Britt, var. texanum (Planch.) Fern.

0.00

0.28

Ludwigia glandulosa Walt.

0.00

0.28

Ludwigia hirtella Raf.

0.00

0.28

Lycopodiella caroliniana (L.) Pichi Sermolli

0.00

0.28

Lycopus rubellus Moench

0.00

0.28

Muhlenbergia capillaris (Lam.) Trin.

0.00

0.28

Oligoneuron nitidum (Torr. & A. Gray) Small

0.00

0.28

Plantanthera spp.

0.00

0.28

Pluchea rosea Godfrey

0.00

0.28

Rhexia mariana L.

0.00

0.28

Rhynchosia latifolia Nutt, ex Torr. & A. Gray

0.00

0.28

Solidago odora Ait.

0.00

0.28

Tephrosia onobrychoides Nutt.

0.00

0.28

Vemonia texana (A. Gray) Small

0.00

0.28

Total

100.00

200.00

Table 2. Relative abundance, relative frequency, and importance value for woody vine species recorded from AOZigadenus densus sites at Fort Polk in west central

Louisiana.

Species

Relative

Abundance

Relative

Frequency

Importance

Value

Rubus argutus Link

33.67

18.71

52.38

Gelsemium sempervirens (L.) St. Hil.

20.92

11.70

32.62

Smilax laurifolia L.

14.29

18.13

32.42

Toxicodendron radicans (L.) Kuntze

8.67

11.11

19.78

Bignonia capreolata L.

7.65

11.11

18.76

Smilax glauca Walt.

6.63

9.36

15.99

Smilax rotundifolia L.

3.06

8.19

11.25

Vitis rotundifolia Michx.

1.53

5.26

6.79

Smilax smallii Morong

3.06

2.92

5.98

Smilax waited Pursh

0.00

2.34

Parthenocissus quinquefolia (L.) Planch.

0.51

0.58

1.09

Berchemiascandens (Hill) K. Koch

0.00

0.58

Total

100.00

200.00

Delahoussaye et al., Vegetation studies of Zigadenus densus

257

Table 3. Relative abundance, relative frequency, and importance value for shrub/sapling (< 1.83 m) species recorded from AO Zigadenus densus sites at Fort Polk

in west central Louisiana.

Species

Relative

Abundance

Relative

Frequency

Importance

Value

Coreopsis glodioto Walt.

0.00

0.28

Morelia caroliniensis (P. Mill.) Small

16.00

6.12

22.12

Rhododendron oblongifolium (Small) Millais

16.00

4.29

20.29

Acerrubrum L. var. drummondii (Hook. & Arn. ex Nutt.) Sarg.

12.50

6.94

19.44

Photiniapyrifolia (Lam.) Robertson & Phipps

10.00

5.51

15.51

Perseapalustris (Raf.) Sarg.

8.00

5.71

13.71

Hypericum crux-andreae (L.) Crantz

4.00

4.08

8.08

Alnus serrulata (Ait.) Willd.

3.50

4.49

7.99

Magnolia virginiana L.

2.00

5.71

7.71

Ilexcoriacea (Pursh) Chapman

3.00

4.69

7.69

Toxicodendron vernix{ L.) Kuntze

1.50

5.31

6.81

Morelia cerifera (L.) Small

3.00

3.06

6.06

Nyssa biflora Walt.

3.00

2.86

5.86

Vaccinium elliottii Chapman

2.00

3.67

5.67

Vaccinium fuscatum Ait.

0.50

4.69

5.19

Viburnum nudum L. var. nudum

0.50

4.29

4.79

Hypericum hypericoides (L.) Crantz ssp. hypericoides

2.50

2.04

4.54

Itea virginica L.

2.50

2.04

4.54

Callicarpa Americana L.

0.50

3.47

3.97

Liquidambar styraciflua L.

1.50

2.04

3.54

Pinus taeda L.

1.50

1.84

3.34

Lyonia lucida (Lam.) K. Koch

1.00

1.63

2.63

Rhododendron canescens (Michx.) Sweet

1.00

1.63

2.63

Viburnum nudum L. var. cassinoides (L.) Torr. & A. Gray

0.50

1.84

2.34

Rhus copallinum L.

1.00

1.22

2.22

Ilex opaca Ait.

0.50

1.43

1.93

Hypericum galioides Lam.

0.50

1.22

1.72

Quercus laurifolia Michx.

0.50

1.22

1.72

Quercus falcata Michx.

0.50

1.02

1.52

Chionanthus virginicus L.

0.00

1.22

Symplocos tinctoria (L.) L'Her.

0.00

1.02

Ilex vomitoria Ait.

0.00

0.82

Sassafras albidum (Nutt.) Nees

0.00

0.82

Hypericum frondosum Michx.

0.50

0.20

0.70

Quercus alba L.

0.00

0.61

Pinus palustris P. Mill.

0.00

0.41

Quercus hemisphaerica Bartr. ex Willd. var. hemisphaerica

0.00

0.20

Quercus nigra L.

0.00

0.20

Vaccinium arboreum Marsh.

0.00

0.20

Vaccinium virgatum Ait.

0.00

0.20

Total

100.00

200.00

258

Journal of the Botanical Research Institute of Texas 8(1)

Table 4. Relative abundance, relative frequency, and importance value for tree/shrub/sapling (> 1.83 m) species recorded from 40 Zigadenus densus sites at Fort

Polk in west central Louisiana.

Species

<5"dbh

Relative

Abundance

>5"dbh

Relative

Abundance

Relative

Frequency

Importance

Value

Magnolia virginiana L

23.50

20.00

10.89

54.39

Pinus palustris P. Mill.

7.00

36.50

10.03

53.53

Nyssa biflora Walt.

7.50

25.00

10.32

42.82

Acer rubrum L. var. drummondii (Hook. & Arn. ex Nutt.) Sarg

19.50

5.00

10.32

34.82

Perseapalustris (Raf.) Sarg.

17.50

2.50

8.02

28.02

Ilexcoriacea (Pursh) Chapman

8.50

0.00

5.44

13.94

Pinus taeda L.

1.50

6.50

4.58

12.58

Liquidambar styraciflua L.

2.50

3.00

4.58

10.08

Toxicodendron vernix{ L.) Kuntze

4.00

0.00

4.87

8.87

Ilex opaca Ait.

0.00

1.00

4.01

5.01

Alnus serrulata (Ait.) Willd.

1.00

0.00

3.72

4.72

Viburnum nudum L. var. nudum

1.50

0.00

2.29

3.79

Morelia caroliniensis (P. Mill.) Small

0.50

0.00

2.29

2.79

Callicarpa americana L.

1.00

0.00

1.72

2.72

Quercus laurifolia Michx.

0.00

2.58

Symplocos tinctoria (L.) L'Her.

1.50

0.00

0.57

2.07

Morelia cerifera (L.) Small

0.50

0.00

1.43

1.93

Vaccinium fuscatum Ait.

0.50

0.00

1.43

1.93

Ilex vomitoria Ait.

0.50

0.00

1.15

1.65

Quercus falcate Michx.

0.00

0.50

1.15

1.65

Quercus alba L.

0.00

1.43

Vaccinium elliottii Chapman

0.50

0.00

0.86

1.36

Lyonia lucida (Lam.) K. Koch

0.00

1.15

Photiniapyrifolia (Lam.) Robertson & Phipps

0.50

0.00

0.57

1.07

Viburnum nudum L. var. cassinoides (L.) Torr. & A. Gray

0.50

0.00

0.57

1.07

Rhododendron canescens (Michx.) Sweet

0.00

0.86

Castaneapumila (L.) P. Mill, var .pumila

0.00

0.57

Chionanthus virginicus L.

0.00

0.29

Itea virginica L.

0.00

0.29

Pinus echinata P. Mill.

0.00

0.29

Prunus serotina Ehrh.

0.00

0.29

Quercus hemisphaerica Bartr. ex Willd. var. hemisphaerica

0.00

0.29

Quercus incana Bartr.

0.00

0.29

Quercus marilandica Muenchh.

0.00

0.29

Quercus nigra L.

0.00

0.29

Vaccinium arboreum Marsh.

0.00

0.29

Total

100.00

300.00

ACKNOWLEDGMENTS

We thank Kristen Mayo for assistance in data collection and Michael MacRoberts and an anonymous reviewer

for the suggested corrections in the manuscript.

REFERENCES

Allen, C.M., J. Pate, S. Thames, S.Trichell, & L. Ezell. 2004. Changes in baygall vegetation from 1986 to 2001 at Fort Polk in

west central Louisiana. Sida 21(1 ):419—427.

Allen, C.M., S. Thames, H. Anderson III, B. Newton III, & R. Hampton. 2006. Green fringed orchid (Platanthera lacera, Orchida-

ceae) in Southern Louisiana. Sida 22(1 ):805—809.

Allen, C.M., R. Erwin, J. McMillian, & J. McMillian. 201 3 A quantitative study of the vegetation surrounding a Xanthorhiza

simplicissima (Ranunculaceae) population at Fort Polk in West Central Louisiana. J. Bot. Res. Inst.Texas 7(1 ):519-528.

Diggs, G.M., B.L. Lipscomb, M.D. Reed, & RJ. O'Kennon. 2006. Illustrated flora of East Texas. Sida, Bot. Misc. 26:1-1594.

Godfrey, R.K. & J.W. Wooten. 1979. Aquatic and wetland plants of the Southeastern United States. Univ. Georgia Press,

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Athens, Georgia, U.S.A.

Louisiana Natural Heritage Program. 2011. Louisiana Dept. Wildlife & Fisheries, Natural Heritage Program, Baton Rouge,

Lousiana, U.S.A. Available www.wlf.louisiana.gov/wildlife/rare-plant-species. (Accessed 23 September 2013).

MacRoberts, B.R., M.H. MacRoberts, & L.S. Jackson. 2002. Floristics and management of pitcher plant bogs in northern

Natchitoches and Winn parishes, Louisiana. Proc. Louisiana Acad. Sci. 64:14-21.

MacRoberts, B.R., M.H. MacRoberts, & L.S. Jackson. 2004. Floristics of Baygalls in Central Louisiana. Phytologia 86:1-22.

Megyeri, K. & C.M. Allen. 2011. Additional observations of the associated plant species surrounding populations of Cypri-

pedium kentuckiense (Orchidaceae) in West Central Louisiana. J. Bot. Res. Inst. Texas 5(2):843-847.

NatureServe. 2013. NatureServe Explorer: An online encyclopedia of life [web application]. Version 7.1. NatureServe,

Arlington, Virginia, U.S.A. Available www.natureserve.org/explorer. (Accessed: 23 September 2013).

Radford, A.E., H.E. Ahles, &C.R. Bell. 1968. Manual of the vascular flora of the Carolinas. University of North Carolina Press,

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

Schwartz, F.C. 2003. Zigadenus. In: Flora of North America Editorial Committee, eds. Flora of North America north of

Mexico. Vol 26, Oxford University Press, New York, U.S.A., and Oxford, U.K.

Soil Survey Division. 2003. Soil survey of Vernon Parish, Louisiana. United States Department of Agriculture, Natural Re¬

sources Conservation Service, Washington, D.C., U.S.A.

Thomas, R.D. & C.M. Allen. 1993. Atlas of the vascular flora of Louisiana, Vol. 1: Ferns and ferns allies, conifers, and mono¬

cotyledons. Louisiana Department of Wildlife and Fisheries, Baton Rouge, Louisiana, U.S.A.

USDA, NRCS. 2013. The PLANTS Database. National Plant Data Team, Greensboro, North Carolina, U.S.A. Available at

plants.usda.gov. (Accessed: 23 September 2013).

260

Journal of the Botanical Research Institute of Texas 8(1)

BOOK NOTICE

Peter Wyse Jackson. 2014. Ireland’s Generous Nature: The Past and Present Uses of Wild Plants in Ireland.

(ISBN-13: 978-0915279784, hbk). Missouri Botanical Garden Press, PO. Box 299, St. Louis, Missouri

63166-0299, U.S.A. (Orders: www.mbgpress.org). $60.00 (leather bound $75.00), 768 pp., illustrated.

From the publisher: Dr. Peter Wyse Jackson’s Ireland’s Generous Nature is the brst ever comprehensive account of

the historical and present-day uses of wild plant species in Ireland. It records a wealth of traditional knowledge

about Irish plant use, knowledge that has been disappearing fast. More than 1500 wild plants are detailed in a

systematic list, which gives both their Irish and English names. Many historical references have been included

from a wide range of Irish literature. This lively and scholarly book shows how plants have been used in virtu¬

ally every aspect of human life in Ireland: food, clothes, medicine, construction, drinks, veterinary medicine,

human health and beauty, and even death. The book is richly illustrated with photographs, as well as botanical

paintings by Irish artist Lydia Shackleton (1828-1914). Its blend of scientific and historic facts with myths,

superstition and tales offers an unrivalled account of the rich heritage of Irish plants.

Set offshore at the far fringe of Europe, its historic subsistence rural culture so long remote from indus¬

trial development or bourgeois mores, Ireland should offer an encyclopedic heritage of the use of wild plants as

food and medicine, for everyday utility or symbolic ritual. For early centuries, indeed, the importance of plant

knowledge was tempered as much by a continuing pagan respect for nature as by the herb gardens of the many

monasteries. ... More than half of the 925 native plants of this island have been useful to its people, but no¬

where, until now, have their stories been gathered together so systematically, both for Irish readers and the

wider cultural and scientific audience.

J.Bot. Res. Inst. Texas 8(1): 260.2014

A QUANTITATIVE STUDY OF THE VEGETATION SURROUNDING

POPULATIONS OF UVULARIA SESSILIFOLIA (COLCHICACEAE) AT FORT POLK

IN WEST CENTRAL LOUISIANA, U.S.A.

Ariel Dauzart, Charles Allen, Stacy Huskins, and Jacob Delahoussaye

Colorado State University, Fort Polk

1645 23rd Street

Fort Polk, Louisiana 71459, U.S.A.

ABSTRACT

Sessile leaf bellwort ( Uvularia sessilifolia) is a rare species (S2) in Touisiana with reports from Caddo, Claiborne, Grant, Tincoln, Morehouse,

Ouachita, Union, and Vernon parishes. Quantitative data from 4 populations (56 samples) in Vernon Parish are reported, and the plant com¬

munity associated with U. sessilifolia is described. In west central Touisiana, U. sessilifolia is found along baygall streams and is associated

with the tree species: Ilexvomitoria, Hamamelis virginiana, and Liquidambar styraciflua; the shrub species Vaccinium elliottii,Acer rubrum var.

drummondii, and Pinus taeda; the woody vine species Smilax smallii, Vitis rotundifolia, and Smilax glauca; and the herbaceous species Wood-

wardia areolata, Chasmanthium laxum, and M itchella repens.

RESUMEN

Uvularia sessilifolia es una especie rara (S2) en Luisiana con citas de la parroquias de Caddo, Claiborne, Grant, Lincoln, Morehouse, Ouachi¬

ta, Union, y Vernon. Se citan datos cuantitativos de 4 poblaciones (56 muestras) en la parroquia de Vernon, y se describe la comunidad veg¬

etal asociada con U. sessilifolia. En el cenro oeste de Louisiana, U. sessilifolia se encuentra a lo largo de torrentes y esta asociada con las espe-

cies: Ilexvomitoria, Hamamelis virginiana, y Liquidambar styraciflua; las especies arbustivas Vaccinium elliottii, Acer rubrum var. drummondii,

y Pinus taeda; las especies trepadoras Smilax smallii, Vitis rotundifolia, Smilax glauca; y las especies herbaceas Woodwardia areolata, Chasman¬

thium laxum, y Mitchella repens.

INTRODUCTION

Sessile leaf bellwort (Uvularia sessilifolia L.) is a native rhizomatous herbaceous perennial in the Colchicaceae.

It is reported from the following states in the US: AL, AR, CT, DC, DE, FL, GA, IA, IL, IN, KY, LA, MA, MD, ME,

MI, MN, MO, MS, NC, ND, NH, NJ, NY, OH, OK, PA, RI, SC, SD, TN, TX, VA, VT, WI, and WV; and from the

following provinces in Canada: MB, NB, NS, ON, and QC (USDA NRCS 2013). NatureServe (2013) has a simi¬

lar distribution but also reports this species for Kansas but not for Texas, and in the Flora of North America

treatment for this species, it is not reported for South Dakota (Utech & Kawano 2003). Sessile leaf bellwort is

listed globally as G5 and SI in KS, ND, and OK; S2 in LA; S3? in IL; S3 in IA; S4 in DE, NC, and WV; and S5 in

KY, NJ, NY, and VA (NatureServe 2013). In Canada, it is listed as S2 in MB, S4 in ON, S4S5 in NS, and S5 in NB

and QC. In Louisiana, this S2 species is reported from Caddo, Claiborne, Grant, Lincoln, Morehouse, Ouachi¬

ta, Union, and Vernon parishes (Louisiana Natural Heritage Program 2011; Thomas & Allen 1993). In east

Texas, it is reported from Cass, Jasper, and Newton counties (USDA NRC 2013; Diggs et al. 2006). In Missis¬

sippi and especially in Arkansas, this species is widespread with reports from several counties (USDA NRCS

2013).

This species is reported from forested seeps and bayhead swamps in Louisiana (Louisiana Natural Heri¬

tage Program 2011) and from deep ravines and mesic forests in east Texas (Diggs et al. 2006). The habitats re¬

ported for this species in the Flora of North America are moist hardwood coves, alluvial bottomlands, thickets,

and xeric woods northwards (Utech & Kawano 2003). In the Carolinas, it is reported from alluvial woods and

coves (Radford et al. 1968).

The objectives of this study were to document and quantitatively characterize the dominance of the vas¬

cular plant species associated with sessile leaf bellwort and to describe its habitat in west central Louisiana.

Twenty-eight different clumps of sessile leaf bellwort are reported from Fort Polk in Vernon Parish. Some of

J. Bot. Res. Inst. Texas 8(1): 261 - 266.2014

262

Journal of the Botanical Research Institute of Texas 8(1)

these are small with only a few or only one stem(s). The four largest populations were selected for this study

Three of the populations were on Guyton-Iuka complex and one was partially on Briley loamy fine sand and

partially on the Guyton-Iuka complex (Soil Survey Division 2003).

METHODS

The four largest Uvularia sessilifolia populations included two along Bird’s Creek and one each along Ouiska

Chitto Creek and East Fork Sixmile Creek. At each population, a macroplot that encompassed the entire U.

sessilifolia was created with the width of all macroplots being 8 m and the length variable depending on the

population size. Each macroplot was subdivided into the maximum number of samples (0.5 m x 8 m). A ran¬

dom number generator was used to select which of the samples would be used. At each location, 50 % (V 2 ) of

the samples were selected for sampling and resulted in a total of 56 samples in the four macroplots.

The plant categories sampled included herbaceous plants, woody vines, shrubs and saplings (woody non¬

vine species shorter than 6 ft = 1.83 m), and trees and shrubs (woody non-vine species taller than 6 ft = 1.83m).

During the sampling period, the number of stems in a sample for each species were counted and recorded. For

herbaceous plants, woody vines, and shrubs and saplings, cover was determined by measuring the area occu¬

pied by the plant(s) in the sample. The cover percent was calculated by multiplying the area times the density

and then dividing by the area of the sample (40,000 cm 2 ). The cover was converted to a percent by multiplying

by 100. For the trees and shrubs taller than 1.83 m, the dbh (diameter breast high) was measured at the stan¬

dard 1.37 m height using a diameter tape and recorded to the nearest 0.1 cm.

All data were entered into a Microsoft Excel spreadsheet for storage and calculation of variables. The

mean diversity (richness-species per sample) and mean density (stems per sample) were calculated for all

plants and for each of the four plant categories. Mean cover percent was calculated for herbaceous plants,

woody vines, and shrubs/saplings, and mean dbh (cm per sample) was calculated for trees/shrubs.

The frequency was calculated for each species in a sample group by dividing the number of samples of

occurrence by the total number of samples (56). It was converted to a percent by dividing by 100. The mean

density was calculated for each species in a sample group by totaling the densities from all samples and divid¬

ing by 56. Mean cover percent for each herbaceous, woody vine, and shrub/sapling species was calculated by

totaling the cover percent from all 56 samples and dividing by 56. The mean dbh was calculated for the tree/

shrub species by totaling the dbh from all samples and dividing by 56.

The relative values for each of these variables (frequency, mean density, mean dbh, and mean cover per¬

cent) were calculated by dividing the value for a species by the total for all species within the plant category.

Each relative value was converted to a percent by multiplying by 100. The relative frequency, relative density,

and relative cover percent were totaled to produce the importance value for each herbaceous, woody vine, and

shrub/sapling species. The relative frequency, relative density, and relative dbh were totaled to produce the

importance value for each tree/shrub species.

RESULTS

A total of 90 (31 herbaceous, 14 woody vines, and 45 trees/shrubs/saplings) species were observed in all 56

samples with a mean number of species per sample of 21.79 (Table 1). The mean diversity per sample ranged

from 1.46 species per sample for trees/shrubs to 9.88 species for shrubs/saplings. The mean number of stems

per sample (density) for all plants was 151.32 stems and ranged from 1.91 stems per sample for trees/shrubs to

85.23 for herbaceous plants. The mean cover percent for all plants was 68.43 percent and ranged from 4.76

percent for woody vines to 43.40 percent for herbaceous plants. The dbh averaged 9.17 cm per sample.

The frequency, mean density, relative mean cover percent, and importance value for each herbaceous spe¬

cies (listed in descending importance value) are in Table 2. Since the macroplots for sampling were centered on

Uvularia sessilifolia, it is not surprising that it has the highest importance value. The three species with the next

highest importance value are Woodwardia areolata, Chasmanthium laxum, and Mitchella repens. The frequency,

mean density, mean cover percent, and importance value for each woody vine species (listed in descending

importance value) are in Table 3. The three species with the highest importance value are Smilax smallii , Vitis

Dauzart et al., Vegetation study of Uvularia sessilifolia

263

Table 1 . Communityvariables for 56 sample plots around four Uvularia sessilifolia macroplots on Fort Polk in West Central Louisiana.

All

Plants

Herbaceous

Plants

DIVERSITY

Shrubs/

Saplings

Trees/

Shrubs

Woody

Vines

Mean

21.79

6.63

9.88

1.46

4.73

Std Dev.

4.75

2.11

2.43

1.44

1.84

Range

11-29

1-10

4-15

0-6

0-9

Total Number

DENSITY

All

Herbaceous

Shrubs/

Trees/

Woody

Plants

Saplings

Shrubs

Vines

Mean

151.32

85.23

1.91

45.18

19.00

Std Dev.

79.48

58.38

2.12

21.59

17.17

Range

33-361

9-265

0-9

10-107

0-108

COVER PERCENT

DBH CM

All

Herbaceous

Shrubs/

Trees/

Woody

Plants

Saplings

Shrubs

Vines

Mean

68.43

42.40

21.27

4.76

9.17

Std Dev.

30.77

29.58

15.96

5.28

11.23

Minimum

14.88-154.31

1.37-131.7

2.20-73.32

0-24.05

0-44.4

Table 2. Frequency, mean density, mean cover percent, and importance value for herbaceous species in 56 sample plots around four Uvularia sessilifolia macroplots

on Fort Polk in West Central Louisiana.

Species

frequency

mean

density

mean

cover %

importance

value

Uvulorio sessilifolia

82.14

31.86

5.39

62.49

Woodwardia areolata

44.64

11.30

12.76

50.10

Chasmanthium laxum

82.14

12.04

9.23

48.29

Mitchella repens

80.36

14.77

1.63

33.30

Osmunda cinnamomea

37.50

1.13

8.02

25.89

Dichanthelium commutatum

60.71

2.86

0.58

13.88

Scleria spp.

46.43

3.05

1.25

13.55

Carexdebilis

32.14

2.48

0.98

10.08

Osmunda regalis

21.43

0.46

1.22

6.65

Solidago caesia

23.21

1.63

0.36

6.25

Viola x primulifolia

25.00

0.89

0.08

5.01

Dichanthelium dichotomum

19.64

0.59

0.13

3.95

Chasmanthium latifolium

16.07

0.54

0.32

3.80

Aster lateriflorus

14.29

0.23

0.08

2.63

Dichanthelium boscii

12.50

0.27

0.07

2.37

Athyrium filix-femina

8.93

0.21

0.14

1.92

Tipularia discolor

10.71

0.21

0.01

1.89

Arisaema triphyllum

10.71

0.14

0.00

1.79

Elephantopus tomentosus

5.36

0.07

0.02

0.94

Solidago arguta

5.36

0.09

0.01

0.93

Pteridium aquilinum

3.57

0.13

0.07

0.85

Dioscorea villosa

3.57

0.09

0.02

0.70

Dichanthelium spp.

3.57

0.05

0.00

0.61

Carex intumescens

1.79

0.02

0.33

Lilium michauxii

1.79

0.02

0.01

0.31

Diodia teres

1.79

0.04

0.00

0.31

Dichanthelium laxiflorum

1.79

0.02

0.00

0.30

Pleopeltis polypodioides

1.79

0.02

0.00

0.29

Bidens aristosa

1.79

0.02

0.00

0.29

Botrychium biternatum

1.79

0.02

0.00

0.29

Total

662.50

85.23

42.40

300.00

264

Journal of the Botanical Research Institute of Texas 8(1)

Table 3. Frequency, mean density, mean cover percent, and importance value for woody vine species in 56 sample plots around four Uvulariasessilifolia macroplots

on Fort Polk in West Central Louisiana.

Species

frequency

mean

density

mean

cover %

importance

value

Smilax smallii

55.36

3.43

0.87

47.94

Vitis rotundifolia

51.79

1.36

1.34

46.20

Smilax glauca

80.36

2.88

0.50

42.56

Smilax laurifolia

32.14

4.52

0.04

31.41

Gelsemium sempervirens

44.64

0.96

0.73

29.82

Smilax pum ila

44.64

1.95

0.41

28.21

Parthenocissus quinquefolia

37.50

1.02

0.36

20.82

Smilax rotundifolia

33.93

0.64

0.15

13.65

Toxicodendron radicans

30.36

0.63

0.16

13.12

Bignonia capreolata

26.79

0.95

0.10

12.77

Rubus argutus

17.86

0.29

0.05

6.39

Smilax spp.

7.14

0.29

0.03

3.63

Berchemia scandens

8.93

0.09

0.01

2.59

Smilax bona-nox

1.79

0.02

0.89

Total

473.21

19.00

4.76

300.00

rotundifolia, and Smilax glauca. The frequency, mean density, mean cover percent, and importance value for

each shrub/sapling species (listed in descending importance value) are in Table 4. The three shrub/sapling spe¬

cies with the highest importance value are Vaccinium elliottii, Acer rubrum var. drummondii, and Pinus taeda.

The frequency, mean density, mean dbh, and importance value for each tree/shrub species (listed in descend¬

ing importance value) are in Table 5. The three tree/shrub species with the highest importance value are Ilex

vomitoria, Hamamelis virginiana, and Liquidambar styraciflua.

DISCUSSION

The habitat for Uvularia sessilifolia in west central Louisiana is along the edge of baygalls as indicated by the

association with W oodwardia aerolata, Chasmanthium laxum, and Acer rubrum var drummondii but in the

slightly higher and dryer sites within the baygall as indicated by the association with Mitchella repens, Smilax

smallii, Ilex vomitoria, and Hamamelis virginiana. Allen et al. (2013) reports Chasmanthium laxum and Mitchella

repens as two of the top three herbaceous species in baygalls associated with Xanthorhiza simplicissima Marsh.

These authors also list Vitis rotundifolia and Smilax glauca as two of the top three woody vines in their study.

Vaccinium elliottii, Liquidambar styraciflua, and Acer rubrum var. drummondii were in the top five species of

trees/shrubs/saplings in their study.

We found the tree canopy/subcanopy vegetation associated with Uvularia sessilifolia to be Ilex vomitoria,

Hamamelis virginiana, Liquidambar styraciflua, Magnolia virginiana L., Acer rubrum var. drummondii, Quercus

alba L., Persea palustris (Raf.) Sarg., and Fagus grandifolia Ehrh. Magnolia virginiana and Persea palustris are

very typical baygall plants (Allen et al. 2004; Diggs et al. 2006; MacRoberts et al. 2004), and Ilex vomitoria,

Hamamelis virginiana, Quercus alba, and Fagus grandifolia are indicators of higher dryer sites. The shrub/sap-

ling layer (Table 4) also had a mixture of typical baygall plants ( Vaccinium elliottii, Acer rubrum var. drummon¬

dii, Rhododendron canescens (Michx.) Sweet, and Persea palustris) plus species of higher dryer sites ( Pinus taeda,

Ilex vomitoria, Carpinus caroliniana Walt., Hamamelis virginiana, and Symplocos tinctoria (L.) L’Her.).

The herbaceous associates of Uvularia sessilifolia in our study that are baygall species were Woodwardia

areolata, Chasmanthium laxum, Osmunda cinnamomea L., Carex debilis Michx., Osmunda regalis L., Viola x

primulifolia L., and Dichanthelium dichotomum L. And the associate species that are typical of higher dryer sites

like natural levees are Mitchella repens, Dichanthelium commutatum (J.A. Schultes) Gould, Scleria spp., and

Solidago caesia L. Two woody vine species, Vitis rotundifola and Smilax laurifolia L., were associated with

Dauzart et al., Vegetation study of Uvularia sessilifolia

265

Table 4. Frequency, mean density, mean cover percent, and importance value for shrub/sapling species in 56 sample plots around four Uvularia sessilifolia macroplots

on Fort Polk in West Central Louisiana.

species

frequency

mean

density

mean

cover %

importance

value

Voccinium elliottii

60.71

1.84

5.68

36.91

Acer rubrum var. drummondii

87.50

8.80

1.56

35.70

Pin us toedo

53.57

6.91

0.20

21.68

Ilex vomitoria

60.71

2.18

1.39

17.51

Carpinus Carolinian a

37.50

4.45

0.63

16.62

Hamamelis virginiana

48.21

2.16

1.37

16.12

Symplocos tinctoria

39.29

1.61

1.65

15.29

Rhododendron canescens

51.79

2.39

0.39

12.37

Persea palustris

50.00

1.07

0.79

11.16

Ilex opaca

57.14

1.66

0.33

11.01

Quercus hemisphaerica

42.86

1.13

0.61

9.69

Nyssa biflora

44.64

2.02

0.11

9.52

Quercus alba

37.50

1.20

0.43

8.48

Fagus grandifolia

23.21

0.45

1.08

8.43

Vaccinium virgatum

16.07

0.66

1.05

8.02

Halesia diptera

25.00

0.43

0.87

7.55

Quercus laurifolia

35.71

1.07

0.20

6.91

Hypericum hypericoides

23.21

1.41

0.17

6.25

Morelia caroliniensis

10.71

0.50

0.47

4.41

Prunus serotina

26.79

0.43

0.09

4.09

Lyonia lucida

7.14

0.16

0.60

3.89

Crataegus marshallii

10.71

0.20

0.36

3.23

Viburnum dentatum

17.86

0.46

0.03

2.97

Magnolia virginiana

12.50

0.23

0.20

2.71

Ilex coriacea

16.07

0.29

0.04

2.46

Callicarpa americana

14.29

0.21

0.06

2.20

Corn us florida

12.50

0.20

0.06

2.00

Ostrya virginiana

5.36

0.23

0.19

1.96

Itea virginica

5.36

0.07

0.13

1.33

Liquidambar styraciflua

7.14

0.07

0.09

1.32

Magnolia grandiflora

3.57

0.07

0.13

1.13

Styrax americanus

7.14

0.11

0.03

1.10

Vaccinium fuscatum

3.57

0.09

0.05

0.82

Styrax grandifolius

5.36

0.09

0.01

0.80

Carya texana

5.36

0.05

0.02

0.73

Vaccinium arboreum

1.79

0.02

0.10

0.69

Carya alba

3.57

0.04

0.01

0.48

Sassafras albidum

3.57

0.04

0.00

0.45

Ilex longipes

1.79

0.04

0.03

0.41

Chionanthus virginicus

1.79

0.05

0.02

0.39

Ligustrum sinense

1.79

0.04

0.00

0.26

Viburnum acerifolium

1.79

0.02

0.00

0.24

Triadica sebifera

1.79

0.02

0.00

0.23

Euonymus americana

1.79

0.02

0.00

0.23

Viburnum nudum

1.79

0.02

0.00

0.22

Total

987.50

45.18

21.27

300.00

sessilifolia and are typical baygall species. The other woody vine associates, Smilax smallii, Smilax glauca, Gel-

semium sempervirens (L.) St. Hil., and Smilax pumila Walt., are more typically found in higher dryer sites.

Our data are the first quantitative description on the vegetation surrounding Uvularia sessilifolia. The

vegetation around the other populations of U. sessilifolia throughout its range should be sampled for compari¬

son with our data so as to get a better idea of the variation, if any, of its habitat.

266

Journal of the Botanical Research Institute of Texas 8(1)

Table 5. Frequency, mean density, mean dbh, and importance value for tree/shrub species in 56 sample plots around four Uvulariasessilifolia macroplots on Fort

Polk in West Central Louisiana.

species

frequency

mean

density

mean

dbh (cm)

importance

value

Ilex vomitoria

19.64

0.36

0.84

41.26

Hamamelis virginiana

17.86

0.32

0.74

37.06

Liquidambar styraciflua

10.71

0.13

1.54

30.67

Magnolia virginiana

5.36

0.13

1.35

24.90

Acer rubrum var. drummondii

10.71

0.13

0.50

19.33

Quercus alba

3.57

0.04

0.99

15.10

Persea palustris

7.14

0.09

0.51

15.08

Fagus grandifolia

8.93

0.09

0.27

13.75

Carpinus Carolinian a

7.14

0.07

0.39

12.84

Nyssa biflora

5.36

0.05

0.58

12.73

Quercus laurifolia

7.14

0.09

0.24

12.22

Prunus serotina

7.14

0.07

0.15

10.29

Ilex opaca

7.14

0.07

0.13

10.04

Pin us taeda

3.57

0.04

0.42

8.85

Magnolia grandiflora

3.57

0.04

0.14

5.87

Symplocos tinctoria

3.57

0.04

0.07

5.03

Vaccinium elliottii

3.57

0.04

0.05

4.89

Quercus hemisphaerica

3.57

0.04

4.80

Carya texana

1.79

0.02

0.07

2.93

Ostrya virginiana

1.79

0.02

0.04

2.64

Rhododendron canescens

1.79

0.02

0.04

2.58

Ilex coriacea

1.79

0.02

0.03

2.45

Com us florid a

1.79

0.02

2.35

Morelia caroliniensis

1.79

0.02

2.35

Total

146.43

1.91

9.17

300.00

ACKNOWLEDGMENTS

We thank Kristen Mayo, Brian Early, and Jeff McMillian for assistance in data collection. Appreciation is ex¬

tended to the reviewers, David Rosen and Michael MacRoberts.

REFERENCES

Allen, C.M., J. Pate, S. Thames, S.Trichell, & L. Ezell. 2004. Changes in baygall vegetation from 1986 to 2001 at Fort Polk in

west central Louisiana. Sida 21(1 ):419—427.

Allen, C.M., R. Erwin, J. McMillian, & J. McMillian. 2013 A quantitative study of the vegetation surrounding a Xanthorhiza

simplicissima (Ranunculaceae) population at Fort Polk in West Central Louisiana. J. Bot. Res. Inst.Texas 7(1 ):519-528.

Diggs, G.M., B.L. Lipscomb, M.D. Reed, & RJ. O'Kennon. 2006. Illustrated flora of East Texas. Sida, Bot. Misc. 26:1-1594.

Louisiana Natural Heritage Program. 2011. Louisiana Department of Wildlife & Fisheries, Natural Heritage Program, Baton

Rouge, Louisiana, U.S.A. Available www.wlf.louisiana.gov/wildlife/rare-plant-species. (Accessed 23 September 2013).

MacRoberts, B.R., M.H. MacRoberts, & L.S. Jackson. 2004. Floristics of baygalls in central Louisiana. Phytologia 86:1-22.

NatureServe. 2013. NatureServe Explorer: An online encyclopedia of life [web application]. Version 7.1. NatureServe,

Arlington, Virginia, U.S.A. Available www.natureserve.org/explorer. (Accessed 23 September 2013).

Radford, A.E., H.E. Ahles, &C.R. Bell. 1968. Manual of the vascular flora of the Carolinas. University of North Carolina Press,

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

Soil Survey Division. 2003. Soil survey of Vernon Parish, Louisiana. United States Department of Agriculture, Natural Re¬

sources Conservation Service, Washington D.C., U.S.A.

Thomas, R.D. & C.M. Allen. 1993. Atlas of the vascular flora of Louisiana, Vol. 1: Ferns and ferns allies, conifers, and mono¬

cotyledons. Louisiana Department of Wildlife and Fisheries, Baton Rouge, Louisiana, U.S.A.

USDA, NRCS. 2013. The PLANTS Database (plants.usda.gov, 23 September 2013). National Plant Data Team, Greensboro,

North Carolina, U.S.A.

Utech, F.H. & S. Kawano. 2003. Uvularia. In: Flora of North America Editorial Committee, eds. Flora of North America north

of Mexico. Oxford University Press, New York, U.S.A., and Oxford, U.K. 26:147.

BLYXA AUBERTII (HYDROCHARITACEAE) NEW TO MISSISSIPPI, U.S.A.

Daniel M. McNair

Department of Biological Sciences

University of Southern Mississippi

Hattiesburg, Mississippi 39406, U.S.A.

danielmcnair@gmail.com

Mac H. Alford

Department of Biological Sciences

University of Southern Mississippi

Hattiesburg, Mississippi 39406, U.S.A.

mac.alford@usm.edu

ABSTRACT

Blyxa aubertii Rich. (Hydrocharitaceae) is reported as new to the flora of Mississippi. New and historical records, along with negative survey

results from nearby states, are mapped at the county level. Voucher specimens are cited, and photographs of B. aubertii are provided.

RESUMEN

Se amplia el area de distribucion de Blyxa aubertii Rich. (Hydrocharitaceae) al estado de Mississippi. Se realiza el mapa de distribucion de la

especie por condado basandose en nuevos registros, datos historicos y en resultados obtenidos del inventario de presencia/ausencia de espe-

cies no detectadas en los censos efectuados en estados aledanos. Se listan los ejemplares revisados en este estudio y se presentan fotografias

de B. aubertii.

Blyxa aubertii Rich., a native of tropical and subtropical regions in Asia, Australia, and Africa (Wang et al.

2010), was first reported for North America, in Louisiana, over 40 years ago (Thieret et al. 1969). No new occur¬

rences outside Louisiana have been reported since that time or listed in relevant treatments or databases (God¬

frey & Wooten 1979; Cook & Luond 1983; Thomas & Allen 1993; Haynes 2000; McCook & Kartesz 2000;

Haynes & Holm-Nielsen 2001; Diggs et al. 2006; Weakley 2012; Kartesz 2013; USDA, NRCS 2013). Thus, our

collections of Blyxa aubertii represent a new record for Mississippi, augmenting several other recent records for

the state (Majure 2007, 2008; Majure & Bryson 2008; Nesom 2010; Whitson 2010; Pruski 2011; Alford 2012;

Urbatsch 2013). Since we found B. aubertii in four counties in south Mississippi and it was already known to

occur in an equal number of parishes in south Louisiana, we predicted it might have also expanded its range to

include parts of Texas and Alabama. While this prediction may still prove true, especially with more extensive

surveys, we did not find B. aubertii or records of its occurrence in these states (Fig. 1).

Voucher specimens: U.S.A. MISSISSIPPI. Forrest Co.: Paul B. Johnson State Park, SE end of Geiger Lake, 31.137253°, -89.235334°, 9 Nov

2013, McNair 1652 (USMS). Lamar Co.: Big Bay Lake, 31.201119°, -89.567456°, 8 Nov 2012, Alford 4378 (BRIT, LSU, USMS). Stone Co.: Flint

Creek Water Park, NE edge of lake, near the end of Day Use Road, 30.895945°, -89.124452°, 9 Nov 2013, McNair 1651 (USMS). Wayne Co.:

Maynor Creek Water Park, N of Reservoir Road, 31.65752°, -88.719192°, 8 Nov 2013, McNair 1650 (MISS, MMNS, USMS).

We observed Blyxa aubertii only in anthropogenically disturbed habitats, specifically, in artificial lakes, sub¬

merged and rooted in nutrient poor substrates; we did not observe it in any naturally formed bodies of water

such as oxbow lakes or rivers. Furthermore, the four lakes in which B. aubertii was found were all constructed

between 1943 and 1974. The oldest of these, Geiger Lake, was finished in 1943 and surveyed for vascular plants

in 1965-1966 (Carter & Jones 1968), so it is likely that B. aubertii established in Mississippi sometime after

1966.

Like a number of other aquatic monocots, Blyxa aubertii has septate leaves and roots, and in vegetative

form it might be mistaken for its closest relative in the U.S., Vallisneria americana. The leaves of B. aubertii have

somewhat noticeable midribs (Fig. 2 A, B) and acuminate apices, while the leaves of V americana lack promi¬

nent midribs and have rounded apices. Also, B. aubertii has bisexual flowers (Fig. 2 C) and ridged seeds (Fig. 2

D), while V americana has unisexual flowers and smooth seeds. In general, individuals of Blyxa aubertii are

smaller with shorter and narrower leaves than Vallisneria. Blyxa aubertii may also be confused with Eriocaulon

aquaticum and Lachnocaulon anceps. Common associates include Nymphaea odorata and Juncus repens. We

J. Bot. Res. Inst. Texas 8(1): 267 - 270.2014

268

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 1. Distribution of Blyxa aubertii in the United States.

observed that many of the plants were in bloom from August to December, the same phenology observed for B.

aubertii in Japanese rice paddies (Jiang & Kadono 2001).

While Blyxa aubertii apparently continues to naturalize and increase its range in the U.S., we do not con¬

sider it a serious threat to native plants, at least not to those found in intact habitats, since B. aubertii seems

confined to disturbed areas of artificial lakes. Still, we believe more investigation will be needed to elucidate

this issue, especially if the species is found inhabiting natural bodies of water.

ACKNOWLEDGMENTS

Support for this study was provided by a grant to the University of Southern Mississippi from the National Sci¬

ence Foundation (DBI 1203684). We would like to thank Liliana Hernandez for translating the Spanish ab¬

stract. We would also like to thank Robert and Linda Donnell for a tour of Big Bay Lake. The review comments

of Robert R. Haynes and Charles T. Bryson are greatly appreciated.

REFERENCES

Alford, M.H. 2012. Noteworthy collections: Mississippi and Louisiana. Castanea 77:181 -185.

Carter, J.W., Jr. & S.B. Jones, Jr. 1968. The vascular flora of Johnson State Park, Mississippi. Castanea 33:194-205.

Cook, C.D.K. & R. Loond. 1983. A revision of the genus Blyxa (Hydrocharitaceae). Aquatic Bot. 15:1-52.

Diggs, G.M., B.L. Lipscomb, M.D. Reed, & RJ. O'Kennon. 2006. Illustrated flora of East Texas. Sida, Bot. Misc. 26:1-1594.

Godfrey, R.K. & J.W. Wooten. 1979. Aquatic and wetland plants of southeastern United States: Monocotyledons. Univer¬

sity of Georgia Press, Athens, Georgia, U.S.A.

Haynes, R.R. 2000. Hydrocharitaceae. In: Flora of North America Editorial Committee, eds. Flora of North America north

of Mexico. Oxford University Press, New York, U.S.A., and Oxford, U.K. 22:26-38.

McNair and Alford, Blyxa aubertii new to Mississippi

269

Fig. 2. Blyxa aubertii. A. Detail of leaf. B. Rosette of leaves and roots. C. Flower. D. Seeds. (A-C, McNair 1650, D, Alford4378).

Haynes, R.R. & L.B. Holm-Nielsen. 2001. The genera of Hydrocharitaceae in the southeastern United States. Harvard Pap.

Bot. 5:201-275.

Jiang, M. & Y. Kadono. 2001. Seasonal growth and reproductive ecology of two threatened aquatic macrophytes, Blyxa

aubertii and B. echinosperma (Hydrocharitaceae), in irrigation ponds of south-western Japan. Ecol. Res. (Washington,

DC) 16(2):249—256.

Kartesz, J.T. 2013. North American plant atlas, (www.bonap.org, accessed 15 Dec 2013). The Biota of North America

Program (BONAP). Chapel Hill, North Carolina, U.S.A.

Majure, L.C. 2007. Noteworthy collections: Mississippi. Castanea 72:121 -122.

Majure, L.C. 2008. New records of Geranium molle L. and Erodium cicutarium (L.) L'Her. ex Ait. (Geraniaceae), from Missis¬

sippi and other important collections from the state. S. E. Naturalist (Steuben) 7:367-370.

Majure, L.C. & C.T. Bryson. 2008. Carexbreviculmis R. Br. (Cyperaceae), new to the Flora of North America. J. Bot. Res. Inst.

Texas 2:1381-1387.

270

Journal of the Botanical Research Institute of Texas 8(1)

McCook, L.M. & J. Kartesz. 2000. A preliminary checklist of the plants of Mississippi, (www.herbarium.olemiss.edu/check-

list.html, accessed 15 Dec 2013). University of Mississippi, Oxford, Mississippi, U.S.A.

Nesom, G.L. 2010. Pyracantha (Rosaceae) naturalized in Texas and the southeastern United States. Phytoneuron 2:1-6.

Pruski, J.F. 2011. Hypochaeris microcephala var. albiflora (Hypochaeris albiflora: Asteraceae), new for the vascular flora of

Mississippi and its distribution in North America. J. Bot. Res. Inst. Texas 5:345-348.

Thieret, J.W., R.R. Haynes, & D.H. Dike. 1969. Blyxa aubertii (Hydrocharitaceae) in Louisiana: New to North America. Sida

3:343-344.

Thomas, R.D. &C.M. Allen. 1993. Atlas of the vascular flora of Louisiana, Vol. 1: Ferns and Ferns allies, conifers, and mono¬

cotyledons. Louisiana Department of Wildlife and Fisheries, Baton Rouge, Louisiana, U.S.A.

Urbatsch, L.E. 2013. Plants new and noteworthy for Louisiana and Mississippi. Phytoneuron 2013-14:1-7.

USDA, NRCS. 2013. The PLANTS Database (http://plants.usda.gov, accessed 15 Dec 2013). National Plant Data Team,

Greensboro, North Carolina, U.S.A.

Wang, Q.F, Y.H. Guo, R.R. Haynes, & C.B. Hellquist. 2010. Hydrocharitaceae. In: Flora of China. Science Press and Missouri

Botanical Garden Press, Beijing, China, and St. Louis, U.S.A. 23:91-102.

Weakley, A.S. 2012. Flora of the southern and mid-Atlantic states. University of North Carolina Herbarium (NCU), Chapel

Hill, North Carolina, U.S.A. Working Draft of 30 Nov 2012.

Whitson, M. 2010. Noteworthy collections: Mississippi. Castanea 75:136-137.

FLORISTIC STUDIES IN NORTH CENTRAL NEW MEXICO, U.S.A.

THE SANGRE DE CRISTO MOUNTAINS

Jill Larson

1 072 Hightower Road

Wheatland, Wyoming 82201, U.S.A.

thousandthjill@hotmail.com

B.E. Nelson

Rocky Mountain Herbarium

Department of Botany, Dept. 3165

University of Wyoming

1000E. University Ave.

Laramie, Wyoming 82071, U.S.A.

bnelsonn@uwyo.edu

Brian Reif

214 South Church Street

Silverton, Oregon 97381, U.S.A.

brianreif_2000@yahoo.com

Ronald L. Hartman

Rocky Mountain Herbarium

Department of Botany, Dept. 3165

University of Wyoming

1000E. University Ave.

Laramie, Wyoming 82071, U.S.A.

rhartman@uwyo.edu

ABSTRACT

This represents the second of two papers covering the floristic diversity of North Central New Mexico. It reports on results from the Sangre

de Cristo Mountains, as well as adjacent lands administered by the State of New Mexico, the Bureau of hand Management, the Picuris and

Taos Indian Reservations, and some other private lands. The first paper covered the Jemez and Tusas ranges on the west side of the Rio

Grande. For the sake of continuity, the two papers are treated as self-contained companion works. The goal is to enumerate results of the

most intensive floristic inventory ever conducted in New Mexico. Here we report on 15,298 numbered collections of vascular plants from an

area covering over 1.3 million acres (526,000 ha) (the sum of the entire area covering more than 3.7 million acres (1.5 million ha) is 35,857

new collections). A total of 1226 unique taxa, including 144 infraspecies and 8 hybrids, are documented from 98 families. Of these, 129 are

exotics (12 are designated as noxious in New Mexico), 18 are species of conservation concern, 23 represent first reports or their confirmation

for New Mexico, and finally 12 are endemic to New Mexico. Based on verified material from the University of New Mexico herbarium, 121

additional unique taxa are included in the Annotated Checklist; thus the grand total is 1347.

RESUMEN

Este es el segundo de los articulos que cubren la diversidad floristica del centro norte de Nuevo Mexico. Se senalan los resultados de las

montanas Sangre de Cristo, asi como tierras adyacentes administradas por el estado de Nuevo Mexico, Ta Oficina de Gestion del Territorio,

las Reservas Indias de Picuris y Taos, y algunos otros territories privados. El primer articulo cubrio las cordilleras de Jemez y Tusas en el lado

oeste de Rio Grande. Por el bien de la continuidad, los dos articulos se tratan como companeros. Su objetivo es enumerar resultados del in-

ventario floristico mas exhaustive* llevado a cabo en Nuevo Mexico. Se citan aqui 15,298 colecciones numeradas de plantas vasculares de un

area de mas de 1.3 millones de acres (526,000 ha) (la suma total del area cubre mas de 3.7 millones de acres (1.5 millon de ha) con 35,857

nuevas colecciones). Se documentan un total de 1226 taxa unicos, incluyendo 144 taxones infraespecificos y 8 hibridos, de 98 familias. De

ellos, 129 son exoticos (12 se designan como nocivos en Nuevo Mexico), 18 son especies con rango de conservacion, 23 representan nuevas

citas o su confirmacion para Nuevo Mexico, y finalmente 12 son endemicas de Nuevo Mexico. Basados en material verificado del herbario

de la Universidad de Nuevo Mexico, se incluyen 121 taxa unicos adicionales en el Catalogo anotado; llegando a un total de 1347.

INTRODUCTION

We report on botanical inventories in the eastern portions of the Carson National Forest (CNF) and the Santa

Fe National Forest (SFNF) by Jill Larson (2008) and Brian Reif (2006), respectively. Included are surround¬

ing public lands administered by the State of New Mexico as well as the Bureau of Land Management, the

Picuris and Taos Pueblo Indian Reservations, and some other private lands (Fig. 1). As these segments of the

two forests are defined as the portions occurring east of the Rio Grande, it is restricted to the Sangre de Cristo

Mountains.

This is the second paper on federal and adjoining lands in north central New Mexico. The first focused on

the Jemez and Tusas Mountains and included the Valles Caldera National Preserve and Bandelier National

J. Bot. Res. Inst. Texas 8(1): 271 - 303.2014

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Journal of the Botanical Research Institute of Texas 8(1)

Fig. 1. Location of entire study area in north central New Mexico. This paper covers floristic research conducted to the east of the Rio Grande. Lands to the

west of the river (in more muted colors) are treated in the companion paper (Reif et al. 2009). The location of selected towns and villages are indicated

and federal agency and tribal lands are delineated (see legend, upper left corner of plate).

Larson et al., Floristic studies of the Sangre de Cristo Mountains

273

Monument (Reif et al. 2009). For the sake of continuity, the two papers are treated as self-contained companion

works.

The geographic area covered in this paper is over 1.3 million acres (526,000 ha) (the area inventoried as a

whole in the two thesis projects was over 3.7 million acres (1.5 million ha). Included are western Colfax, west¬

ern San Miguel, western Mora, southeastern Rio Arriba, northeastern Santa Fe, and eastern Taos counties. The

area ranged from just below 5500 feet (1686 m) in Anton Chico, (the extreme southeastern portion of the area)

to 13,161ft (4000 m) on Wheeler Peak, the highest point in New Mexico.

These botanical inventories are part of the larger effort by the Rocky Mountain Herbarium (RM) to map

in relatively fine detail the geographic distribution of species based on vouchered specimens and to produce a

flora of the greater Rocky Mountain region (Hartman 1992; Hartman et al 2009; Hartman & Nelson 2008). To

this end, 61 major floristics inventories (49 as master’s degree projects) have been conducted during the past 35

years in Arizona, Colorado, Idaho, Kansas, Montana, Nebraska, New Mexico, Oregon, South Dakota, Utah,

Washington, and Wyoming. Over 680,000 new collections have been obtained by graduate students, staff, and

research associates of RM. These specimens form the core of the RM Plant Specimen Data Base (830,000

specimen records, 90,000 specimen images, and 4000 held photos) (Hartman et al. 2009).

Topography.—A majority of the area is within the Southern Rocky Mountain Region (Fenneman 1931).

To the north, in Colorado, this region is divided by the headwaters of the Rio Grande, thus forming the treeless

San Luis Valley. Here the mountains to the west are the San Juans, those to the east, the Sangre de Cristos. The

eastern division extends to the south from Colorado into New Mexico as a continuous range to the east of the

Rio Grande River. The southern terminous of the Sangre de Cristo Mountains, near Santa Fe, dissolves into a

land of mesas and plains. The surhcial geology is largely metamorphic and volcanic, but areas covered by sedi¬

mentary formations are also present. On the west side of the Rio Grande, the San Juans extend southeast from

Colorado and are known locally as the Tusas Mountains separated to the south from the Jemez Mountains and

the Sierra Nacimiento by the Rio Chama.

The Taos Plateau is a broad basin and the southern extension of the San Luis Valley of Colorado in the

northern portion of the valley of the Rio Grande. It is peppered with volcanic cones some of which rise 3000 ft

(914 m) or more above the surrounding desert grass-shrublands. To the south, the Rio Grande Rift is the por¬

tion constricted by the mountains and formed by the Embudo Fault (Muehlberger 1978; Muehlberger &

Muehlberger 1982).

Climate.—The climate of north central New Mexico is arid in the lowlands, becoming moister as one

ascends in elevation. Precipitation is significantly affected by the North American monsoon. The monsoon

results in a seasonal pattern of precipitation with a primary maximum during July, August, and September.

Strong diurnal cloud cover and precipitation accompany this season. This corresponds with the peak convec¬

tive heating during the day leading to frequent afternoon thunderstorms (Sheppard et al. 1999). The moun¬

tains exert strong orographic effects on precipitation in direct relationship to elevation, while temperature is

inversely related. New Mexico receives 30 to 40 percent of its annual precipitation during the summer months.

Winter tends to be drier, with a secondary maximum of precipitation occurring November through

March (Sheppard et al. 1999). Winter precipitation is hydrologically important as winter snowpack recharges

surface and ground water (Redmond 2003).

On the west side of the Sangres, from north to south, the annual precipitation is about 13 in (33 cm) at

Cerro (elevation 7592 ft; 2314 m), 12 in (30 cm) at Taos (6952 ft; 2219 m), 10 in (25 cm) at Espanola (5589 ft;

1703 m), 14 in (36 cm) at Santa Fe (6989 ft; 2130 m), and 16 in (41 cm) at Pecos (6923 ft; 2110 m). On the east

side of the project area, from north to south, the annual precipitation is about 21 in (53 cm) at Red River (8650

ft; 2636 m), 15 in (38 cm) at Eagle Nest (8095 ft; 2467 m), 24 in (61 cm) at Gascon (8045 ft; 2452 m), and about

16 in (41 cm) at Las Vegas (6430 ft; 1960 m) (Western Regional Climate Center 2008). There is no long-term

climate data for higher elevations in the mountains.

Drought during a portion of this study had an impact on collecting. Dry conditions were particularly se¬

vere in 2002 and 2003 and again in 2006 until the monsoons developed in late July and early August.

274

Journal of the Botanical Research Institute of Texas 8(1)

Geology and geomorphology.—The current landscape of the Southern Rocky Mountain Region is

mainly a product of the Laramide Orogeny (late Cretaceous into the Tertiary (Eocene epoch), 70 to 55 million

years ago (mya) that produced the Sangre de Cristos and (to the west of the Rio Grande) the Nacimientos. This

uplifting of the Precambrian core caused a warping of the overlying sedimentary strata (anticline). Subsequent

volcanism, erosion, infill, and subsidence are recurring themes throughout the region. Furthermore, many

older rock types of igneous origins have been uplifted and exposed as a result of the complex geological his¬

tory. Now, many summits of the Sangre de Cristo consist of Precambrian rocks (to 1.78 billion years old).

The Taos Plateau is the youngest geologic area. It is a broad, relatively flat surface formed by basaltic lava

flows. These flows issued from hundreds of volcanic vents during the Pliocene (Lipman 1978; Chronic 1987).

The larger shield volcanoes are still present on the Plateau and include Ute Mountain, to the east of the Rio

Grande and San Antonio Mountain, to the west. Rocks from some vents date back 2.2-1.8 mya.

The Rio Grande Rift proper is a major break in the earth’s crust where large slivers (grabens) between two

faults subsided around 30 mya. The rift extends from Colorado south into northern Mexico and is estimated to

have been nearly 5 miles in depth (Chronic 1987). Prior to the Pliocene volcanism, the Rift filled with erosive

deposits from the surrounding mountains. Around Taos it is still actively subsiding relative to the Sangre de

Cristo Mountains (Muehlberger & Muehlberger 1982).

The Rio Grande became a through-flowing river during the Pliocene, having overcome containment in a

succession of closed basins within the Rift (Chronic 1987). On the Taos Plateau, it is now confined to a narrow,

deeply cut canyon called the Rio Grande Gorge. The gorge is up to 800 ft in depth. The east side of the gorge is

marked by alluvium from the Sangre de Cristo Mountains (NMBGMR 2003).

The geology of the Sangre de Cristo Mountains is complex. All of the higher landforms were glaciated

repeatedly during the Pleistocene epoch (Cronic 1987). The glaciers shaped more than 60 cirques in the south¬

ern Sangre de Cristo Mountains, although many are now below treeline (Miller 1963).

The northern portion is an intricate patchwork of volcanic and metamorphic rocks known as the Taos

Range. This range includes the highest and most rugged peaks in the Sangre de Cristo Mountains in New

Mexico. Wheeler Peak, the highest point in New Mexico at 13161 ft, is here located. The range is composed of

exposed Precambrian granitic rocks with remnants of intrusive silicic rocks and basaltic andesites of Tertiary

age (NMBGMR 2003). Faulting is common and adds to the geologic complexity (Shilling 1956). Volcanic activ¬

ity that shaped the Taos Range is evidenced by the ancient Questa caldera and the Fatir volcanic held, which

were active during the Oligocene (25 mya; Meyer 1990). A molybdenum (moly) mine now occupies the Questa

caldera. Along the east-west trending fault just northeast of Taos, the geology changes to sedimentary forma¬

tions. This surface is an artifact of Pennsylvanian “skin,” leaving the older Precambrian core covered. These

sedimentary strata cover a vast area to the south.

The Picuris Mountains are Precambrian quartzite and schist that together with Cerro Azul, on the west

side of the Rio Grande, form a constriction. At this point the Rio Grande Rift takes a decided shift to the west

(NMBGMR 2003). Here a major fault has brought the Precambrian rocks upward in line with the Pennsylva¬

nian strata in the mountains to the east (Bauer & Raiser 1995). hike the Tusas Mountains to the west of the Rio

Grande, the Picuris Mountains experienced volcanism during the middle-Tertiary resulting in thick deposits

of Picuris Tuff (Miller 1963).

At the south end, the Sangres contains thick sedimentary deposits from the Pennsylvanian Period (310-

280 mya). These deposits may be up to 2700 ft in thickness north of Pecos and extend more than 20 miles (38

km) to the north (Sutherland & Montgomery 1975). This forms a tongue of sedimentary strata flanked by

precambrian rock (Chronic 1987). Approximately 26 miles to the north of Pecos in the Truchas Mountains is

Truchas Peak at an elevation of 13102 ft (3993 m).

METHODS

Field work on the Carson and Santa Fe National Forests was conducted during the summers of 2002 through

2006, whereas work on the portion to the west of the Rio Grande was begun a year earlier (Valles Caldera Na-

Larson et al., Floristic studies of the Sangre de Cristo Mountains

275

tional Preserve). Also on the west side of the river, Brian Jacobs conducted an inventory of Bandelier National

Monument from 1986 through 1988 (herbarium now housed at University of New Mexico). These studies

combined represent the most extensive and exhaustive floristic surveys ever conducted in New Mexico.

Botanists have been roaming the country sides throughout the world for centuries, documenting the

riches of floristic diversity. In keeping with the tradition of held botanists, collecting sites were selected and

searched based on the researcher’s judgment. Thus, the “meander search” strategy was employed (Goff et al.

1982; Hartman 1992; Ristau 1998; Hartman & Nelson 2008). As sites were selected subjectively, the result was

the exploration of a much greater diversity of plant communities, soil types, geologic substrates, and topogra¬

phy leading to the documentation of a substantially greater diversity of taxa.

A total of 845 waypoints (each a geographic coordinate determined using a GPS unit) are represented in

this paper (for the two thesis projects as a whole, the total was 1542; Fig. 2). Each waypoint represents a gen¬

eral location for collecting plant specimens, usually within one-half mile (either as a radius from a point or a

trail segment defined by two successive points; notes on community types were recorded). Details concerning

collections as followed in all RM studies are found in Hartman (1992) and Hartman & Nelson (2008).

This paper is based on 15,298 collections from the Sangre de Cristo Mountains and vicinity (total number

of collections obtained for the two thesis projects and the Bandelier National Monument was 35,857 collec¬

tions). This document does not include the northeastern portion of the Sangres in New Mexico. Here, the

contiguous Vermejo Park Ranch was surveyed by Legler (2010) who obtained 7503 specimens (912 mi 2 ;

236,206 ha). A portion of the adjoining Cimarron Range, Philmont National Scout Ranch, was inventoried in

1968 where 1200 collections were taken (210 mi 2 ; 54,389 ha) (Hartman et al. 2009). A complete set of vouchers

from CNF and SFNF as well as the adjoining areas are housed at RM. All authors have made major contribu¬

tions to the collection, identification, and verification of specimens, as well as the writing of this paper.

RESULTS AND DISCUSSION

The following sections will emphasize the results of research in the Sangre de Cristo Mountains and vicinity

with some discussions on various topics. Past discussion referred to the Carson and Santa Fe National Forests

to the west of the Rio Grande (Reif et al. 2009), thus this companion paper completes coverage of these forests.

VEGETATION TYPES

New Mexico’s vegetation has been divided and described in a number of studies at various spatial scales. The

most relevant to this inventory is Dick-Peddie (1993, see Table 12.2). Vegetation types reported here are based

on the application and condensation of the above-mentioned classification as confirmed or modified by our

held observations. We report 16 vegetation types with five broad physiognomic and zonal categories. These

may foster an understanding of the amplitude, common associates, and environmental requirements of the

taxa documented by these floristic inventories.

Alpine

Alpine fellfield and meadow .—Alpine vegetation occurs in the Sangre de Cristo Mountains above upper

treeline. This varies considerably, but generally occurs between 11,100 and 12,000 ft (3380-3650 m) in eleva¬

tion. The lower boundary is in contact with Krummholz or dwarfed conifers. Included in this broad category

are fellfield cushion plant communities, talus slopes, moist to wet meadows, islands of dwarf shrub, alpine

lakes, and small stream drainages. As expected, the species composition of New Mexico’s alpine vegetation

bears greatest affinity to the main Rocky Mountain Cordillera, implicating a north-south migration as the

source of many of its species (Billings 1988; Pase 1994). Consequently, alpine vegetation is regarded as a unit.

It is also acknowledged, however, that site conditions greatly affect species composition, and a significant and

unrelated component of the Bora may be derived from lower elevation. Here is found the most southern alpine

area, Lake Peak just northeast of Santa Fe, in the Rocky Mountain Cordilleran.

At least 31 taxa were found exclusive to this vegetation type. These include Artemisia pattersonii, A. scopu-

lorum, Carex rupestris, Castilleja haydenii, Cymopterus alpinus, Delphinium alpestre, Elymus scribneri, Eritrichi-

um nanum, Paronychia pulvinata, Primula angustifolia, Synthyris alpina, Tonestus pygmaeus, and Trifolium na-

276

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 2. Map of north central New Mexico counties included in the entire study along with 1542 plant collecting waypoints. Triangles are collecting waypoints

associated with the Carson NF portion of the project, diamonds with the Santa Fe NF, and dots with the Valles Caldera NP. The current paper covers floristic

research east of the Rio Grande (shown as the thinner black line), while the companion paper (Reif et al. 2009) covers lands west of the river (see Fig. 1).

num. These taxa demonstrate varying distribution patterns regionally. Some are common throughout the

Rocky Mountains, while others are restricted to the southern region. This applies to Castilleja haydenii and one

of the species of conservation concern, Delphinium alpestre, both known only from north central New Mexico

and adjacent Colorado.

Forests and Woodlands

Bristlecone pine woodland .—These woodlands, fairly limited in distribution, consist of widely spaced, short-

conical individuals of Pinus aristata occurring above 10,000 ft (3050 m) on dry, rocky, exposed slopes and

ridges in surrounding subalpine spruce-fir forests (Peet 1988; Pase & Brown 1994a). Principal associates of

this “scree forest” are Ribes montigenum and Saxifraga bronchialis (Alexander & Ronco 1987). Bristlecone pine

woodland also occurs on deep soils bordering meadows and in association with Festuca thurberi (Alexander &

Ronco 1987). Understory species associated with both subalpine forest and alpine include Castelleja miniata,

Eremogone fendleri, Festuca idahoensis, Helianthella parryi, Juniperus communis, Fuzula spicata, Pedicularis

racemosa var. alba, Phleum alpinum, Frifolium attenuatum.

Larson et al., Floristic studies of the Sangre de Cristo Mountains

277

Spruce-fir forest —These forests occur in subalpine habitats above 9500 ft (2890 m) or on cooler and

more moist slopes at somewhat lower elevations. Picea engelmannii and Abies arizonica are codominants, al¬

though Piceapungens may occur as serai on moist sites (Moir 1993; Pase & Brown 1994a). In the upper 1000 ft

(304 m) Picea engelmannii typically is the sole tree. Scattered stands of Populus tremuloides may be present in

disturbed areas. Conifers of lower elevations, such as Abies concolor, Pinus flexilis, and Pseudotsuga menziesii

may be present in low numbers. Associated shrubs and subshrubs include the abundant and often “impenetra¬

ble” Juniperus communis, as well as Lonicera involucrata, Dasiphorafruticosa, Ribes wolfii, Rubus parviflorus, Sa-

lix scouleriana, Sambucus racemosa, and Vaccinium myrtillus. Associated graminoid species are not well repre¬

sented but may include Bromus ciliatus, Deschampsia cespitosa, and Trisetum montanum. Frequently encoun¬

tered forbs are Actaea rubra, Castilleja sulphurea, Cymopterus lemmonii, Erigeron eximius, E. coulteri, Goody era

oblongifolia, Eigusticum porteri, Packera sanguisorboides, Pedicularis racemosa, Penstemon whippleanus, Pyrola

minor, Solidago simplex, and Viola canadensis.

Mixed conifer forest —Mixed conifer forest occurs throughout the upper montane, roughly between

8000 and 10,000 ft (2440-3050 m). Pseudotsuga menziesii and Abies concolor are the most widespread and im¬

portant conifers. Piceapungens is restricted to locations subject to cold-air drainage including lower slopes and

drainages while Pinus ponderosa and Pinus strobiformis are found in xeric sites at lower elevations (DeVelice et

al. 1986 Moir 1993). The mixed conifer forest is a mix of conifers thus reflecting the heterogeneous landscape.

Common understory shrubs and subshrubs are Acer glabrum, Arctostaphylos uva-ursi, Berberis repens,

Holodiscus dumosa, Jamesia americana, Juniperus communis, J. scopulorum, Paxistima myrsinites, and Physocar-

pus monogynus. Forbs are represented by Allium cernuum, Aquilegia elegantula, Artemisiafranserioides, Castilleja

miniata, Cymopterus lemmonii, Erigeron subtrinervis, Fragaria vesca, Helianthella parryi, Hymenoxys hoopesii,

Lathyrus leucanthus, Lithospermum multiflorum, Packerafendleri, Potentilla gracilis, Solidago simplex, Thalictrum

fendleri, and Thermopsis rhombifolia, while frequent graminoids include Agrostis scabra, Bromus porteri, B. rich-

ardsonii, Carex geophila, C. occidentalis, C. siccata, Elymus trachycaulus, Muhlenbergia montana, and Oryzopsis

asperifolia.

Fire has created a natural mosaic within this vegetation type, with burns ranging from patchy, low inten¬

sity to stand replacing fires (Moir 1993; USDA, Forest Service 1997).

Ponderosa pine forest —In the Sangre de Cristo Mountains, warmer and dryer conditions from 6700-

9000 ft (2040-2740 m) support this forest type, often forming homogenous stands. At its upper limits, Pinus

ponderosa and mixed conifer forests intergrade, while at its lower limit it merges into pinon-juniper woodland.

Where trees are large and scattered the forests may be open and park-like with predominately grasses and

forbs. This is especially true along the western side as well as in the southeastern portions of the range. Distur¬

bance patterns in ponderosa pine forest include frequent, low intensity understory fires (DeVelice et al. 1986;

Moir 1993; USDA, Forest Service 1997).

Representative subshrubs and shrubs include Berberis repens, Ceanothus fendleri, Cercocarpus montanus,

Fallugia paradoxa, Juniperus communis, Quercus gambelii, Rhus trilobata, Ribes cereum, R. inerme, R. leptanthus,

Rosa acicularis, R. woodsii, and Symphoricarpos rotundifolius. Forbs are represented by Achillea millefolium, An-

tennaria parvifolia, Cymopterus lemmonii, Erigeron flagellaris, Heterotheca villosa, Lithospermum multiflorum,

Lupinus argenteus, Penstemon barbatus, and Vicia americana, while common graminoids include Blepharoneu-

ron tricholepis, Carex inops ssp. heliophila, Elymus elymoides var. brevifolius, Festuca arizonica, Koeleria macran-

tha, Muhlenbergia montana, and Poafendleriana.

Pinon-juniperwoodland. — Pinus edulis and Juniperus monosperma are dominant in the foothills largely on

the western and southeastern side of the Sangre de Cristo Mountains where it forms a discontinuous transi¬

tional belt (6000-8500 ft; 1830-2590 m). It is characteristic of escarpments such as Glorieta Mesa where

coarse, rocky soil and enhanced infiltration produces moister site conditions (Dick-Pettie 1993). Pinon-juniper

woodlands represent the lowest elevation forest type. Pinon forms closed woodlands at the upper elevational

ranges, whereas juniper occurs in savanna-like communities at the lower elevational range and interface with

grasslands. Juniperus scopulorum commonly occurs with pinon at higher elevations in mesic settings and has

278

Journal of the Botanical Research Institute of Texas 8(1)

recently encroached into ponderosa pine understory due to absence of fire disturbance (USDA, Forest Service

1997). Understory conditions are dynamic and vary with canopy cover, soil conditions, and land use and bre

history (West 1999).

Shrub cover is variable and includes Quercus gambelii and Q. undulata, but also Artemisia tridentata, Cer-

cocarpus montanus, Gutierrezia sarothrae, Fallugia paradoxa, Ribes cereum, and Rhus trilobata. Frequently en¬

countered forbs are Castilleja integra, Chaetopappa ericoides, Chamaesyce fendleri, Cryptantha cinerea, Eriogo-

num jamesii, Gaillardia pinnatifida, Hymenoxys richardsonii, Lappula occidentalis, Oenothera suffrutescens,

Sphaeralcea coccinea, Tetraneuris argentea, and several succulents including Cylindropuntia imbricata, Opuntia

phaeacantha, O. polyacantha, and Yucca spp. Common graminoids are Achnatherum hymenoides, Aristida pur¬

purea, Bouteloua curtipendula, B. gracilis, Carex geophila, Elymus elymoides var. brevifolius, Hesperostipa comata,

Hilariajamesii, Koeleria cristata, Muhlenbergia torreyi, Piptatherum micranthum, and Poafendleriana.

Historic impacts in woodlands are extensive and include fuel harvest and grazing (Moir 1993). Bark bee¬

tle ( Ips confusus) outbreaks associated with drought stress have resulted in high levels of pinon mortality (San¬

tos & Whitham 2010).

Shrublands

Montane Shrubland .—These shrublands are found throughout the Sangre de Cristo Mountains. They are dis¬

tributed in a patchy manner in ponderosa pine forests and pinon-juniper woodlands. Areas of montane shru¬

bland are often too limited in extent for mapping because they are produced by site-specific factors: distur¬

bance, substrate, and patterns of moisture. Comparably drier and more rocky sites are typical of montane

shrubland, although some associates such as Prunus virginiana and Ptelea trifoliata are found in areas of in¬

creased moisture such as small catchments (Dick-Peddie 1993).

Included in this classification of montane shrubland are thick stands of Quercus gambelii. This species

occurs on soils that are poorly developed and xeric (Brown 1994), and have also been regarded as serai associa¬

tion indicative of past disturbance. Also included here is the “climax” shrubland of Dick-Peddie (1993) or

Cercocarpus montanus in association with Amelanchier alnifolia, Philadelphus microphyllus, Quercus undulata,

and Rhus trilobata. Other shrubs include Ceanothusfendleri, Holodiscus dumosusjamesia americanajuniperus

communis, Physocarpus monogynus, Ribes cereum, R. inerme, Rubus parviflorus, Sambucus racemosa, and Sym-

phoricarpos rotundifolius. Grass and forb species are those of surrounding montane forests and woodlands.

Desert shrubland .—There are two subtypes that fit this descriptor. The first is found along the Pecos

River and in a large area of the Taos Plateau, but less so on the east side of the Rio Grande. This subtype is char¬

acterized by the absence of Artemisia tridentata and the presence of number of other shrubs. Included are Atri-

plex canescens, Chrysothamnus greenei, Ericameria nauseosa, and Gutierrezia sarothrae. Grass cover is often

sparse but may include Achnatherum hymenoides, Aristida purpurea, Bouteloua gracilis, Elymus smithii, and

Muhlenbergia torreyi. Forbs are also scanty, yet represented by Castilleja integra, Chaetopappa ericoides, Eriogo-

num jamesii, Hedeoma drummondii, Lappula occidentalis, and Ehelesperma megapotamicum.

The second subtype is dominated by Artemisia tridentata var. tridentata. It most often occurs in glades

within pinon-juniper woodlands. Atriplex canescens, Cylindropuntia imbricata, Ericameria nauseosa, and Rhus

trilobata are common woody associates. Grasses include Agropyron cristatum, Bromus tectorum, Elymus elymoi¬

des var. brevifolius, Hilaria jamesii, and Muhlenbergia richardsonis. Common forbs are Chaetopappa ericoides,

Echinocereus coccineus, Lappula occidentalis, Opuntia polyacantha, and Plantago patagonica.

Grasslands

Montane meadow and grassland .—This vegetation type occurs from about 8500 ft (2590 m) to the highest

summits. Transitions between forest and grassland vegetation are often abrupt at the upper elevations where

grasslands may represent a climax condition, or are typically gradual at lower elevations where trees can en¬

croach under heavy grazing or have been excluded by past fire (Peedie 1993). Common forbs include Achillea

millifolium, Agoseris aurantiaca, Allium cernuum, Campanula rotundifolia, Castilleja miniata, Frasera speciosa,

Hymenoxys hoopesii, Linum lewisii, Mirabilis melanotricha, and Silene scouleri. Graminoids vary with moisture

and, to a lesser degree, elevation. Moist sites frequently include Carex microptera, C. nova, Deschampsia cespi-

Larson et al., Floristic studies of the Sangre de Cristo Mountains

279

tosa, Phleum pratense, and Poa pratensis. Drier meadows are characterized by Bromus porteri, Festuca arizonica,

Koeleria macrantha, and Blepharoneuron tricholepis.

Plains-desert grassland. —This vegetation type is ecotonal to pinon-juniper woodlands, juniper wood¬

lands, or juniper savannas. Desert grassland occurs primarily along the Rio Grande Rift often up slope to the

pinon-juniper woodland. It has a significant shrub and forb cover. Characteristic grasses are Achnatherum hy-

menoides, Aristida purpurea, Bouteloua gracilis, B. curtipendula, Elymus smithii, Hesperostipa comata, and Hilaria

jamesii, while frequently encountered forbs are Antennaria microphylla, A. rosea, Castilleja Integra, Cryptantha

spp., Glandulariabipinnatifida, Grindelia squarrosa, Oenothera coronopifolia, O. suffrutescens, Opuntia polyacan-

tha, O. phaecantha, Ratibida tagetes, Sphaeralcea coccinea, Teucrium lacinata, and Zinnia grandiflora.

Wetlands

Montane Riparian. —Margins of perennial and intermittent streams support unique species assemblages.

Montane riparian vegetation is found in moist areas within spruce-fir and mixed conifer forests. As with mon¬

tane meadows, the species composition follows an elevational gradient (Dick-Peddie 1993). A rich diversity of

herbaceous and woody vegetation is present. Obligate and facultative riparian species of trees and shrubs can

be arranged along a descending gradient: Picea pungens, Salix amydaloides, S. bebbiana, S. irrota, Alnus incana,

Acer glabrum, Cornus sericea, Populus angustifoliia, and Acer negundo. Additional facultative riparian trees and

shrubs include Populus tremuloides and Prunus virginiana. Among the rich diversity of forbs are Aconitum co-

lumbianum, Caltha leptosepala, Cardamine cordifolia, Dodecatheon pulchellum, Equisetum arvense, Geum macro-

phyllum, Heracleum maximum, Hypericum scouleri, Mertensia franciscana, Mimulus guttatus, Oxypolis fendleri,

Pedicularis groenlandica, Saxifraga odontoloma, Sedum rhodanthum, Veronica americana, and species of Epilobi-

um, Potamogeton, Ranunculus, and Salix. Graminoids are represented by Alopecurus aequalis, Deschampsia

cespitosa, Glyceria grandis, G. striata, Juncus arcticus, Eorreyochloa pallida, and species of Agrostis, Carex, and

Eleocharis.

Floodplain-arroyo riparian. —This type of vegetation occurs at lower elevations on floodplains along the

Rio Grande and the lowest elevations of the Sangres. Many species that thrive here are well adapted to distur¬

bance and dry conditions with periodic flooding. The most common dominant is Populus deltoides, with un¬

derstory shrubs Baccharis salicina, Ericameria nauseosa, Fallugia paradoxa, Forestiera pubescens, Rhus trilobata,

and Salix exigua. The exotics Elaeagnus angustifolia and Eamarix chinensis have proliferated and may persist in

a subclimax state (Dick-Peddie 1993; Minckley & Brown 1994). Arroyo riparian is common to desert shru-

bland and may grade into montane riparian.

Marsh-lacustrine. —The marsh-lacustrine riparian habitat is found around ponds and springs,in other¬

wise arid habitats such as pinon-juniper woodland or desert grassland where the water table remains suffi¬

ciently high or in various montane vegetation types. Along the shoreline, Limosella aquatica, Potentilla anseri-

na, P. norvegia, Ranunculus cymbalaria, Rorippa spp., and Rumex crispus may be encountered. Emergents in¬

clude Scirpus microcarpus, Sparganium emersum, Eypha latifolia and species of Carex, Eleocharis, Juncus, and

Schoenoplectus. Floating and submersed taxa include Callitrichepalustris, Potamogeton spp., Eemna minor, Elo-

dea canadensis, and Ranunculus aquatilis.

Disturbed

Aspen serai forest. — Populus tremuloides, an important serai species and post fire increaser, is widely distrib¬

uted in the Rocky Mountains (Peet 1988). Pure stands of this shade-intolerant species arise through root

sprouting following disturbances (Pase & Brown 1994b). Scattered individuals are also found in late-succes-

sion or near climax stages in forest types and lower subalpine spruce-fir forests (Moir 1993). However, like

many successional communities, aspen forms a distinct assemblage that may persist for long periods of time.

Aspen are found in the various coniferous forest types throughout the Sangre de Cristo Mountains.

Forbs include Campanula rotundifolia, Castilleja miniata, Chamerion angustifolia, Geranium richardsonii,

Ligusticum porteri, Pseudostellaria jamesiana, and Ehalictrum fendleri, while frequently encountered grasses

include Bromus carinatus, B. richardsonii, Festuca arizonica, Poafendleriana, and P. pratensis.

Bum Areas. —Natural and anthropogenic fires have been frequent in most of the vegetation types men-

280

Journal of the Botanical Research Institute of Texas 8(1)

tioned above. This is true for most of the Sangre de Cristo Mountains. In many areas, the vegetation is in vari¬

ous states of succession. While fire is important ecologically, natural succession is often compromised by exot¬

ics. Several major burns have occurred in the Sangres.

Roadside-agricultural. —Here, native vegetation may be largely replaced by exotic and agricultural spe¬

cies. Noxious weeds collected along roads include Aegilops cylindrica, Cirsium vulgare, Convolvulus arvensis,

Lepidium latifolium, Linaria dalmaticum, and Ulmus pumila. Weedy or agricultural plants include Avena sativa ,

Helianthus annuus, Salsola tragus, Sisymbrium altissimum, Tragopogon dubius, and species of Ambrosia, Bromus,

Chamaesyce, Chenopodium, Elymus, Lappula, Lepidium, Medicago, Melilotus, Plantago, Polygonum, and Trifolium.

Roadsides often act as corridors for exotics and thus warrant monitoring.

NOXIOUS WEEDS

Invasive plant species that are particularly damaging or prolific are regulated as noxious weeds (USDA, NRCS

2013). A specific goal of our floristic inventories was to document noxious weeds for the purpose of assisting in

monitoring and control efforts. Of the 35 taxa listed in New Mexico, 13 were encountered at 91 sites. They are

Acroptilon repens, Aegilops cylindrica, Carduus nutans, Centaurea stoebe ssp. micranthos, Cirsium arvense, C. vul¬

gare, Convolvulus arvensis, Elaeagnus angustifolia, Lepidium latifolium, Linaria dalmatica, L. vulgaris, Tamarix

chinensis, and Ulmus pumila. The first of these noxious weeds is based on a specimen at University of New

Mexico. Noxious weeds and other exotics are indicated in the Annotated Checklist by a • or an *, respectively.

TAXA OF CONSERVATION CONCERN

Another primary goal of the inventories was to document the occurrence of rare and endemic taxa. A total of

18 species of conservation concern were documented from 83 sites in the Sangre de Cristo Mountains and vi¬

cinity. These species of conservation concern are listed by Natural Heritage New Mexico (2012) and the New

Mexico Rare Plant Technical Committee (2012) as such. The list is arranged alphabetically and each is followed

by family name, county of occurrence, and collector and associated voucher number(s). They are indicated by

a ♦ in the Annotated Checklist.

Astragalus cyaneus (Fabaceae) Taos: Hartman 80544b

Astragalus iodopetalus (Fabaceae) Taos: Larson 4946,5711,7140

Astragaluspuniceus var. gertrudis (Fabaceae) Taos: Hartman 80541;

Larson 5293

Caiochortus gunnisonii var.perpu/cher (Liliacaeae) Mora, San Miguel:

Reif3026, 7232, 7299, 7614

Cornus canadensis (Cornaceae) Taos: Larson 2799,8035

Cypripediumparviflorum (Orchidaceae) San Miguel: Reif6219

Delphinium alpestre (Ranunculaceae) Taos: Larson 2945, 3004,

3393,8991

Delphinium sapellonis (Ranunculaceae) Mora, San Miguel, Taos:

Nelson 65919, 66309, 69383; Reif 3011, 3773, 4038, 7323, 7641,

8682,8779,8934, 10286

Hackelia hirsuta (Boraginaceae) Colfax, Mora, San Miguel, Taos:

Larson 2303,3093,8186, 10626; Nelson 69141,69442; Reif 3157

Herrickia horrida (Asteraceae) Colfax: Larson 10028

SUMMARY OF TAXA

The first number represents results based on our fieldwork. Parenthetical numbers following most of the for¬

mer are those verified from UNM. The two adjacent numbers below “Exotic taxa” represent the percent of ex¬

otics when compared to total unique taxa.

A total of 1226 unique taxa, including 144 infraspecies and 8 hybrids, are documented from 98 families.

Of these, 129 are exotics (12 are designated as noxious in New Mexico), 18 are species of conservation concern,

23 represent first reports or their confirmation for New Mexico, and finally 12 are endemic to New Mexico.

Based on verified material from the University of New Mexico herbarium, 121 additional unique taxa are in¬

cluded in the Annotated Checklist; thus the grand total is 1347.

Iliamna grandiflora (Malvaceae) Taos: Larson 5404,5470,5848

Parnassia fimbriata (Parnassiaceae) Taos: Larson 3972, 4070, 9775,

9877

Podistera eastwoodiae (Apiaceae) Taos: Larson 1579, 1784a, 1842,

2937, 2982, 3015, 4022, 7989, 8080, 9009, 9966, 10310; Reif

8102, 10214

Salixarizonica (Salicaceae) Mora: Reif 10294

Saxifraga cernua (Saxifragaceae) Santa Fe: Reif 10294

Selaginella weatherbiana (Selaginellaceae) Rio Arriba, San Miguel,

Taos: Hartman 76708, Larson 2509,10291, Reif3111,3876,6107,

7441,8089

Synthyris alpina (Plantaginaceae)Taos: Hartman 81340, Larson 720,

1589, 1769,2991,8153

Trifolium brandegeei (Fabaceae) Taos: Hartman 81284, Larson 1582,

3978,4061,4214, 7212b, 7664, 7687,8127

Larson et al., Floristic studies of the Sangre de Cristo Mountains

281

List by taxonomic category

List by special category

Families

98(103)

Exotic taxa

129(148)

Genera

475 (514)

Percent exotic taxa

10.5(11)

Species

1147(1263)

NM Noxious weeds

12(13)

Hybrids

Taxa conservation concern

Infraspecies

144(149)

NM Endemic taxa

12(13)

State records

Unique taxa

1226(1347)

List of unique taxa by major plant group

Fern Allies 11(21)

Ferns 18(18)

Gymnosperms 13

Angiosperms 1184(1295)

Total 1226(1347)

CONCLUSIONS

This paper represents the second of two contributions that cover the floristic diversity of north central New

Mexico (for the portion west of the Rio Grande, see Reif et al. 2009). The area here covered encompasses the

Sangre de Cristo Range, as well as adjacent lands administered by the State of New Mexico, the Bureau of Land

Management, the Picuris and Taos Indian Reservations, and some other private lands. We report on results of

15,298 numbered collections of vascular plants (total for the two publications covering more than 3.7 million

acres is 35,857 numbered new collections). A total of 1226 (1347) unique taxa, 144 (149) including infraspecies

as well as 8 hybrids, are documented from 98 (103) families. With the addition of 121 taxa (totals within veri¬

fied from RM and UNM), the total for unique taxa rise to 1347. Of these, 129 (148) are exotic taxa of which 12

(13) are designated as noxious in New Mexico, 18 are species of conservation concern, 23 represent first re¬

ports or their confirmation for New Mexico, and 12 (13) are endemic to the state.

THE ANNOTATED CHECKLIST

The checklist is divided into major plant groups (ferns and fern allies, gymnosperms, and angiosperms) each

with alphabetical listing by family and species. Nomenclature follows Allred (2012). In some cases (71) it fol¬

lows that of an antiquated checklist compiled by the staff of the RM. In order to provide an easy “cross walk”

between the companion floristic treatments, that name is maintained between the two and the name used in

Allred 2012 is placed in square brackets [ ] below the alternate name. The original sources used in identifica¬

tion were relevant state and regional treatments and monographs with comparison to authenticated materal,

where possible, in the RM.

Following is a guide to format and abbreviations associated with individual taxa in the checklist. Except

for records based on specimens at RM, the citation of the vouchers are omitted. This is justified as collection

data are available online (Hartman et al. 2009; Symbiota 2013). In the case of Botrychium, all specimens were

collected by Ben Legler on his own.

Taxon Authority (project on the Carson NF or the Santa Fe NF) [3, 9 or 6,-]; COUNTY; elevational range

in ft; GEOLOGIC AREA; habitat type.

[Taxon Authority, name accepted by Allred]

One other attribute includes specimens seen only at the University of New Mexico, [UNM-R. Sivinski

3910]

282

Journal of the Botanical Research Institute of Texas 8(1)

County abbreviations: Geologic area:

A SAn Miguel G

C Colfax L

M Mora P

R Rio Arriba S

S Santa Fe

T Taos

Habitat type:

af Aspen serai forest

am Alpine fellfield and meadow

br Burns

bw Bristlecone pine woodland

ds Desert shrubland

fr Floodplain-arroyo riparian

me Mixed conifer forest

ml Marsh-lacustrine wetland

Symbols by category preceding Taxon:

* Exotic species to New Mexico

• Noxious weed in New Mexico

♦ Species of conservation concern

+ Endemic to New Mexico

! State record for New Mexico

x Hybrid

Rio Grande Rift

Great PLains

Taos Plateau

Sangre de Cristo Mountains

mm Montane meadow and grassland

mr Montane riparian

ms Montane shrubland

pg Plains-desert grassland

pj Pinon-juniper woodland

pp Ponderosa pine forest

ra Roadside-agricultural

sf Spruce-fir forest

FERNS AND FERN ALLIES

Aspleniaceae

Asplenium resiliens Kunze [1,-] A; 5500'; S; fr.

Asplenium trichomones L. [1,-] A; 8900-9500'; S; me.

Dennstaedtiaceae

Pteridium oquilinum (L.) Kuhn var. pubescens Underw. [15,4] A, C,

M, S, T; 7720-10500'; S; af, me, mm, mr, pp, sf.

Dryopteridaceae

Athyrium filix-femina (L.) Roth ex Mert. var. californicum Butters [4,3]

A, S,T; 8300-10500'; S; mr.

Cystopteris frogilis (L.) Bernh. [12,18] A, M, R, S, T; 7740-12700'; S;

am, bw, me, mm, mr, sf.

Cystopteris reevesiono Lellinger [24,24] A, C, M, S, T; 7620-12000';

S; bw, me, mr, sf.

Dryopteris filix-mos (L.) Schott [7,3] A, C, M, T; 7650-10500'; S;

me, mr, pp.

Gymnocarpium dryopteris (L.) Newman [-,1] T; 8450-9600'; S; mr.

Woodsio neomexicana Windham [2,2] R, S, T; 6840-11200'; P, S;

bw, fr, pp.

Woodsio oregano D.C. Eaton var. cathcortiano (B.L. Rob.) Morton

[1,4] A, C,T; 7400-12450'; P, S; am, ds, me.

Woodsio plummerae Lemmon [-,1 ] T; 7400-7550'; P; pj.

Equisetaceae

Equisetum arvense L. [15,24] A, C, M, R, S, T; 5781 -9700'; G, P, S; fr,

ml, mm, mr, ra, sf.

x Equisetum xferrissii Clute [1,2] S, T; 7760-9400'; S; mm, mr.

Equisetum hyemale L. var. affine (Engelm.) A.A. Eaton [8,9] A, C, M,

S,T; 7350-10500'; S; me, mm, mr.

Equisetum laevigatum A. Braun [7,18] A, C, M, R, S,T; 5781-9700';

G, L, P, S; fr, me, ml, mm, mr, ra, sf.

x Equisetum xnelsonii (A.A. Eaton) J.H. Schaffn. [-,1 ] T; 5781'; G; fr.

Lycopodiaceae

Huperzia lucidula (Michx.) Trevis. [UNM-C. Dixon A-289] S; S.

Lycopodium annotinum L. [UNM-N. Osborn 1079] T; S.

Ophioglossaceae

Botrychium echo W.H. Wagner [UNM-B. Legler 11545] T; S.

Botrychium hesperium (Maxon & R.T. Clausen) Wagner & Lellinger

[RM, UNM-B. Legler 11553] T; S.

Botrychium lanceolatum (Gmel.) Angstr. [RM-B. Legler 11567] C; S.

Botrychium lineare W.H. Wagner [RM, UNM- B. Legler 11556] T; S.

Botrychium "neolunaria" in ed. [RM, UNM-B. Legler 11584A] S; S.

Botrychium minganenseM ict. [RM, UNM-B. Legler 11609] T; S.

Botrychium pinnatum H.St. John [RM, UNM-B. Legler 11582] S; S.

Botrychium fun ux Steve ns void & Farrar [RM, UNM-B. Legler 11555]

T; S.

Pteridaceae

Argyrochosma fendleri (Kunze) Windham [1,5] R, S, T; 5800-7550';

G, P, S; ds, fr.

Cheilanthes eatonii Baker [4,2] A, C,T; 5560-8450'; S; fr, pj, pp.

Cheilanthes feei T. Moore [2,-] A; 5650-7150'; S; mr, pj.

Cheilanthes fendleri Hook. [2,1] A, S,T; 7600-8640'; S; me, ms, pp.

Cryptogramma acrostichoides R. Br. [2,6] M, R, T; 9600-12000'; S;

am, bw, me, mm, sf.

Notholaenastondleyi Maxon [2,-] A; 5560-5840'; S; pj.

Pellaea atropurpurea (L.) Link [1,-] A; 5500'; S; pj.

Selaginellaceae

Selaginella densa Rydb. var. densa [1,16] A, T; 7050-12700'; P, S;

am, ds, me, mm.

Larson et al., Floristic studies of the Sangre de Cristo Mountains

Selaginella mutica D.C. Eaton ex Underw. var. limitanea Weatherby

hi ]T, 7600-8450'; S; pp.

Selaginella mutica D.C. Eaton ex Underw. var. mutica [1,-] S; 6540';

S; f r.

Selaginella peruviana (Milde) Hieronymus [1,-] A; 5630'; S; fr.

Selaginella underwoodii Hieronymus [2,-] A, M; 7900-9760'; S; me, sf.

♦ Selaginella weatherbiana R. Tryon [6,2] A, R, S, T; 7750-13024';

S; am, me.

GYMNOSPERMS

Cupressaceae

Juniperus communis L. var. depressa Pursh [34,43] A, C, M, R, S, T;

7620-12430'; P, S; af, bw, me, mm, ms, mr, pp, sf.

Juniperus monosperma (Engelm.) Sarg. [15,18] A, R, S,T; 5560-8720';

G, L, P, S; ds, fr, mr, ms, pj.

Juniperus scopulorum Sarg. [14,44] A, C, M, R, S,T; 6050-10500'; G,

P, S; ds, fr, me, ml, mm, ms, mr, pj, pp, ra.

Pinaceae

Abies arizonica Merriam [15,29] A, C, M, R, S, T; 8500-12400'; S;

me, mm, mr, sf.

Abies concolor (Gord. & Glend.) Hildebr. [32,34] A, C, M, R, S, T;

7600-10500'; P, S; af, me, mm, mr, pj, pp.

Picea engelmannii Parry ex Engelm. var. engelmannii [20,24] A, C,

M, R, S,T; 7720-12300'; S; am, af, bw, me, mm, mr, sf.

Piceapungens Engelm. [22,23] A, C, M, R, S, T; 6840-12000'; S; me,

mm, ms, mr, sf.

Pinus aristata Engelm. [8,21 ] A, C, M, S, T; 9375-13000'; S; am, bw,

me, mm, mr, sf.

Pinus edulis Engelm. [28,29] A, C, M, R, S,T; 5560-10000'; G, P, S; br,

ds, fr, me, mm, ms, pj, pp, ra.

Pinus flexilis E. James [13,10] A, M, S, T; 7740-11209'; P, S; me,

mr, ms, sf.

Pinusponderosa Douglas ex P. Lawson & C. Lawson var. scopulorum

Engelm. [28,29] A, C, M, R, S, T; 5500-9920'; P, S; af, br, fr, me,

mm, ms, mr, pj, pp, ra.

[Pinus scopulorum (Engelm.) Lemmon]

Pinus strobiformis Engelm. [8,5] A, C, M, S, T; 7720-10150'; S; me,

mm, mr.

Pseudotsuga menziesii (Mirb.) Franco var. glauca (Beissn.) Franco

[32,38] A, C, M, R, S, T; 6600-11150'; P, S; af, br, me, mm, ms,

mr, pj, pp.

ANGIOSPERMS

Aceraceae [as Sapindaceae]

Acerglabrum Torr. var. glabrum [29,41 ] A, C, M, R, S,T; 7320-11300';

S; br, me, mm, ms, mr, ra, sf.

[AcerglabrumTorLvaLneomexicanum (Greene) Kearn.&Peeb.]

Acernegundo L. var. interius (Britton) Sarg. [5,6] A, S,T; 5781 -8000';

L, P, S; fr, mr, pj, ra.

Adoxaceae

Adoxa moschatellina L. [UNM-R. Sivinski 3910] S; S.

Sambucus caerulea Raf. var. neomexicana (Woot.) Rehder [UNM-C.

Dixon A-285] S; S.

Sambucus racemosa L. var. melanocarpa (A. Gray) McMinn [3,-] M,

S; 9840-12000'; S; mm, mr.

Sambucus racemosa L. var. microbotrys (Rydb.) Kearn. & Peeb.

[18,21 ] A, C, M, R, S, T; 7840-11500'; S; br, me, mm, mr, sf.

Agavaceae [includes Nolinaceae]

Nolina greenei S. Watson [3,-] A;5700-5800'; L; pj.

Yucca angustissima Engelm. exTrelease var. angustissima [3,-] A, R;

5730-6620'; G, L, S; ds, pj.

283

Yucca baccata Torr. var. baccata [9,6] A, R, S, T; 6036-9450'; G, P,

S; ds, fr, me, pj.

Yucca baileyi Wooton & Standi, var. baileyi [3,-] A, S; 6200-7460';

L, S; pg, pj.

+ Yucca intermedia McKelvey [-,4] R, T; 6050-8150'; G, P, S; br, fr,

Pj, PP-

[Y. baileyi Wooton & Standi, var. intermedia (McKelvey) Reveal; see

Sivinski 2008]

Yucca neomexicana\Nootor\ & Standi. [2,-] A, S; 5700-6540'; G, L; pj.

Alismaceae

Sagittaria cuneata E. Sheld. [UNM-E. Castetter 4792] T; S.

Alliaceae [traditionally in Liliaceae]

Allium cernuum Roth [37,35] A, C, M, R, S, T; 7000-11700'; P, S; af,

br, bw, fr, me, mm, ms, mr, pj, pp, ra, sf.

Allium geyeri S. Watson var. geyeri [10,7] A, M, S, T; 8100-12700'; S;

am, bw, me, mm, ms, mr, pp, sf.

Allium geyeri S. Watson var. tenerum M.E. Jones [1,7] M, T; 9250-

11100'; S; mm, mr.

Allium macropetalum Rydb. [1,-] A; 6200'; L; pg, pj.

Amaranthaceae

Amaranthus albus L. [1,1 ] A, T; 5610-7450'; L, S; fr, ra.

* Amaranthus blitoides S. Watson [UNM-H. Mackay 6T-55] T; S.

Amaranthus powellii S. Watson [3,4] A, C, M, S, T; 7300-9400'; S;

fr, mm, mr, ra.

* Amaranthus retroflexus L. [3,-] A, S; 6840-8000'; S; mm, mr.

Guilleminea densa (Humb. & Bonpl. ex Willd.) Moq. var. aggregata

Uline & Bray [2,-] A;5630-5800'; L; fr, pj.

Anacardiaceae

Rhus glabra L. [1,-] A; 7900'; S; mr, ms.

Rhus trilobata Nutt. var. trilobata [10,18] A, C, M, R, S,T; 5600-8450';

G, L, P, S; ds, fr, ms, mr, pj, ra.

Toxicodendron rydbergii (Small ex Rydb.) Greene [6,2] A, R, S, T;

5800-8300'; G, L, P, S; fr, me, ml, mm, mr.

Apiaceae

Angelica grayi (J.M. Coult. & Rose) J.M. Coult. & Rose [5,5] A, M, R,

S, T; 9800-13024'; S; am, mm, sf.

* Carum carvi L. [-,1 ] T; 8350-9400'; S; me.

Cicuta maculata L. var. angustifolia Hook. [2,1] A, T; 5800-8000';

L, P, S; fr, mr.

Conioselinum scopulorum (A. Gray) J.M. Coult. & Rose [20,14] A, C,

M, R, S, T; 7000-11500'; S; br, me, mr, sf.

* Conium maculatum L. [1,-] A; 5800'; L; fr.

Cymopterus alpinus A. Gray [2,2] A, T; 10000-12584'; S; am.

Cymopterus bakeri (J.M. Coult. & Rose) M.E. Jones [2,8] M, S, T;

11500-13000'; S; am.

Cymopterus constancei R.L. Hartm. [-,6] T; 6600-7500'; P, S; ds, fr, pj.

Cymopterusglomeratus (Nutt.) DC. var. fendleri (A. Gray) R.L. Hartm.

[1,-] A; 6200'; L; pg, pj.

Cymopterus longilobus (Rydb.) W.A. Weber [-,3] R,T; 11600-13024';

S; am, sf.

[Cymopterus hendersonii (J.M. Coult. & Rose) Cronquist, misap¬

plied]

Cymopterus lemmonii (J.M. Coult. & Rose) Dorn [43,69] A, C, M,

R, S,T; 7550-12960'; S; af, am, bw, br, me, mm, ms, mr, pp, sf.

! Cymopterus spellenbergii R.L. Hartm. & J.E. Larson [-,6] R, T;

6200-8763'; G, P, S; fr, pj, pp.

Harbouria trachypleura (S. Watson) J.M. Coult. & Rose [4,-] A, S;

7000-8000'; S; pj, pp.

Heracleum maximum W. Bartram [29,24] A, C, M, R, S, T; 7720-

11 650'; S; af, me, ml, mm, mr, ra, sf.

Ligusticum ported J.M. Coult. & Rose [18,17] A, M, S,T; 8350-11940';

S; af, me, mm, mr, sf.

284

Journal of the Botanical Research Institute of Texas 8(1)

Osmorhiza depouperoto Phil. [21,33] A, C, M, R, S, T; 7620-11800';

S; af, me, mm, mr, sf.

Oxypolis fendleri (A. Gray) A. Heller [35,38] A, C, M, R, S, T; 7620-

1 2000'; S; af, me, mm, mr, sf.

* Pastinaca sativa L. [-,1 ] T; 7240'; S; ra.

* Podistero eostwoodioe (J.M. Coult. & Rose) Mathias & Const.

[2,12] M, R, S, T; 10500-13024'; S; am, mm, mr, sf.

Soniculo morilondico L. [1,-] S; 8600-8840'; S; me.

Apocynaceae

Apocynum ondrosoemifolium L. [9,6] A, C, S, T; 7720-9700'; S; af,

me, mm, mr.

Apocynum cannabinum L. [3,4] R, S, T; 5781-7600'; G, P, S; fr, ml,

mr, ra.

x Apocynum xfloribundum Greene [4,1] A, S, T; 7720-8400'; S;

mm, mr.

[Apocynum medium Greene var. floribundum (Greene) Woodson]

Araliaceae

Aralia racemosa L. ssp. bicrenata (Wooton & Standi.) S.L. Welsh & N.D.

Atwood [5,2] A, S,T; 7640-10500'; S; me, mm, mr.

Asdepiadaceae

Asclepias asperula (Decne.) Woodson var. asperula [4,1] A, C, S;

6540-8600'; G, S; br, fr, pj.

Asclepios engelmonniono Woodson [1,-] A; 5570-5600'; L; fr.

Asclepias involucrata Engelm. exTorr. [3,-] A; 5700-6200'; L; pg, pj, ra.

Asclepias latifolia (Torr.) Raf. [2,-] A; 5650-5840'; L; pg, pj.

Asclepias macrostislon. [2,-] A; 5650-5800'; L; pg, pj.

Asclepias oenotheroides Chamisso & Schlectendal [1,-] A; 5700-

5800'; L; pg, pj.

Asclepiaspumila (A. Gray) Vail [UNM-Nellessen 70] A; S.

Asclepias speciosa Torr. [1,2] A, T; 5800-10200'; L, S; fr, mr, ra.

Asclepias subverticillata (A. Gray) Vail [7,2] A, M, S, T; 5500-7760';

L, S; fr, mm, ra.

Asclepias tuberosa L. ssp. interior Woodson [2,-] A; 7720-7900';

S; mm, ms.

Asclepias viridiflora Raf. [2,-] A; 5700-7000'; L, S; pg, pj, pp.

Funastrum crispum (Benth.) Schlecht. [1,-] A; 5700-5800'; L; pg, pj.

Mateleaproducta (Torr.) Woodson [2,-] A; 5700-5840'; L; pg, pj.

Asparagaceae [traditionally in Liliaeeae]

* Asparagus officinalis L. [2,3] R, S,T; 6050-7860'; P, S; br, ml, mm,

ra.

Asteraceae

Achillea millefolium L. [55,55] A, C, M, R, S,T; 6540-13024'; G, S; am,

af, br, me, mm, mr, pp, ra, sf.

*• Acroptilon repens (L.) DC. [UNM-C.R. Hutchins 6455] T; S.

Ageratina herbacea (A. Gray) R.M. King & H. Rob. [1,-] A; 7900'; S;

me, mm.

Agoseris aurantiaca (Hook.) Greene var. aurantiaca [3,10] M, R, S,T;

8300-13024'; S; am me, sf.

Agoseris aurantiaca (Hook.) Greene var. purpurea (A. Gray) Cronquist

[17,16] A, M, S,T; 8200-12200'; S; am, af, bw, br, me, mm, mr, sf.

Agoseris glauca (Pursh) Raf. var. glauca [-,8] M, R, T; 7050-12450';

S; am, br, mm, mr, sf.

Agoseris parviflora (Nutt.) Greene [-,4] T; 9500-12960'; S; am,

mm, mr.

Amauriopsis dissecta (A. Gray) Rydb. [19,11] A, C, M, R, S, T; 6840-

10200'; S; br, fr, me, mr, pp, ra.

Ambrosia artemisiifolia L. [2,-] A, S; 5610-7000'; L, S; fr, mm, ra.

Ambrosia confertifolia DC. [UNM-R. Fleetwood s.n., 3 Sept 1949] A; S.

Ambrosia psilostachya DC. [8,-] A, M, S; 5500-8575'; L, S; fr, mm,

mr, ra.

Ambrosia tomentosa Nutt. [3,3] A, C, M,T; 7200-9700'; S; ml, mm, ra.

Ambrosia trifida L. var. trifida [-,1 ] T; 7240'; S; ra.

Anaphalis margaritacea (L.) Benth. & Hook. [4,4] A, M, R, T; 7750-

11900'; S; me, mm, mr, sf.

Antennaria marginata Greene [6,15] A, C, M, R, S,T; 7620-10500';

S; af, br, me, mm, mr, pp, sf.

Antennaria media Greene [4,7] C, M, S,T; 7400-12850'; S; am, mm, sf.

Antennaria microphylla Rydb. [1,36] C, M, R, S, T; 7050-13024'; S;

am, af, br, me, mm, mr, pj, pp, ra, sf.

Antennariaparvifolia Nutt. [5,18] C, M, R, S,T; 6600-11115'; G, P, S;

fr, me, mm, mr, pj, pp.

Antennaria rosea Greene [2,26] A, M, S, T; 7740-12960'; S; am, br,

me, mm, mr, sf.

[Antennaria rosea subspecies]

Antennaria rosulata Rydb. [-,1 ] T; 9850-10000'; S; mm.

* Arctium minus (Hill) Bernh. [-,3] M, T; 7200-9400'; S; pj, ra.

Arnicacordifolia Hook. [-,23] R,T; 8300-11500'; S; br, me, mm, mr, sf.

Arnica latifolia Bong. [UNM-R. Jackson 2206] T; S.

! Artemisia borealis Pall. ssp. borealis [-,1 ] T; 11500-12850'; S; am.

Artemisia campestris L. var. pacifica (Nutt.) M. Peck [-,2] C, T;

8500-9500'; S; pp, ra.

Artemisia carruthii A.W. Wood ex Carruth [8,22] A, M, R, S, T;

6980-10500'; P, S; br, mm, mr, ms, pj, ra.

Artemisia dracunculus L. [2,2] A, C,T; 7000-8700'; S; br, mm, mr.

Artemisia franserioides Greene [12,8] A, C, M, R,T; 7840-11500'; S;

af, me, mm, mr, sf.

Artemisia frigida Willd. [2,5] A, C, M,T; 6750-9750'; S; mm, pj, ra.

Artemisia ludoviciana Nutt. var. ludoviciana [8,2] A, M, R, S, T;

8200-12200'; S; me, mm, ms, pp.

Artemisia ludoviciana Nutt. var. mexicana (Willd. ex Spreng.) A.

Gray [2,7] A, M, R,T; 7600-10300'; S; af, br, me, mm, pj, pp, ra.

Artemisiapattersonii A. Gray [-,1 ] T; 11500-12050'; S; am.

Artemisia scopulorum A. Gray [-,8] R, T; 11300-13161'; S; am.

Artemisia tridentata Nutt. var. tridentata [-,16] R, T; 6050-10100';

G, P, S; br, fr, pj, ra.

Artemisia tridentata Nutt. var. wyomingensis (Beetle & A. Young) S.L.

Welsh [-,4] R,T; 5800-7650'; G, S; ds, fr, pj.

Baccharis pteronioides DC. [1,-] A; 5800'; L; fr.

Baccharis salicina Torr. & A. Gray [-,2] R, T; 5781 -6540'; G; ds, fr, ra.

Baccharis wrightii A. Gray [1,-] S; 7320-7370'; S; pj.

Berlandiera lyrata Benth. [6,-] A; 5650-6200'; L; pg, pj.

Bidens cernua L. [-,1 ] C; 8194'; S; ml.

Bidens pilosa L. [1,-] A; 5500'; L; fr.

Bidens tenuisecta A. Gray [3,1] A, M; 7700-9320'; S; mm, ra.

Brickellia brachyphylla A. Gray [1,-] A; 7000'; S; pp.

Brickellia eupatorioides (L.) Shinners var. chlorolepis (Wooton &

Standi.) B.L. Turner [2,1] M,T; 7100-8800'; S; mm, pp, ra.

Brickellia grandiflora (Hook.) Nutt. [10,13] A, C, M, R, S, T; 7650-

12200'; S; am, me, mm, ms, mr, pp, ra, sf.

Brickellia rusbyi A. Gray [UNM-J. McGrath 737] A; S.

Brickelliastrum fendleri (A. Gray) King & H.E. Rob. [10,5] A, R, S, T;

6840-10100'; P, S; fr, me, mm, mr, pp.

*• Carduus nutans L. [5,6] A, C, M, T; 7000-9200'; S; me, ml, mm,

pj, ra.

*• Centaurea stoebe L. ssp. micranthos (S.G. Gmelin ex Gugler)

Hayek [1,-] A; 5500'; L; fr, pj.

Chaetopappa ericoides (Torr.) G.L. Nesom [15,17] A, R, S, T; 5600-

8200'; G, L, P, S; br, ds, pg, pj, ra.

Chrysothamnus depressus Nutt. [-,1] R; 8150'; S; br.

Chrysothamnus greenei (A. Gray) Greene [-,4] T; 7400-8555'; P; ds, fr.

* Cichorium intybus L. [2,1 ] M, S, T; 6840-7700'; S; mm, ra.

*• Cirsium arvense (L.) Scop. [-,12] C,T; 7350-10500'; S; br, ml, me,

mm, mr, ra.

[Cirsium arvense varieties]

Cirsium eatonii (A. Gray) B.L. Rob. var. eriocephalum (A. Gray) Keil

[3,5] M, S, T; 10990-12850'; S; am, mm.

Larson et al., Floristic studies of the Sangre de Cristo Mountains

Cirsium neomexiconum A. Gray [2,6] R, S, T; 6540-8900'; G, S; br,

fr, me, pj, pp, ra.

Cirsium ochrocentrum A. Gray var. ochrocentrum [2,-] A, S; 5760-

7000'; L, S; mm, pg, pj.

Cirsium porryi (A. Gray) Petrak [27,31] A, C, M, S, T; 7400-11500';

S; me, mm, mr, ra, sf.

Cirsium scariosum Nutt. var. coloradense (Rydb.) Keil [-,1] T; 8175';

S; ml, mm.

Cirsium undulatum (Nutt.) Spreng [12,7] A, C, M, S,T; 5570-9700';

L, S; br, fr, me, mm, ms, pg, pj, pp, ra, sf.

*• Cirsium vulgore (Savi) Ten. [4,3] A, C, M, S, T; 6840-9320'; S; fr,

ml, mm, ra.

* Conyza canadensis (L.) Cronquist [7,4] A, C, S,T; 5500-8575'; L,

P, S; br, fr, ml, mm, ra.

Coreopsis lanceolata L. [1,-] A; 7720'; S; pp.

Coreopsis tinctoria Nutt. [1,-] A; 7200-7240'; S; ra.

Cosmosparviflorus (Jacq.) Pers. [2,1 ] A, M, S; 7380-9320'; S; mm, ra.

Crepis runcinata (E. James) Torr. & A. Gray var. runcinata [UNM-J.

Williams 7]T;S.

!* Crepis tectorum L. [-,1 ] C; 8194'; S; ml.

Cyclachaenaxanthifolia (Nutt.) Fresen. [1,1] A, T; 7350-8000'; S; ra.

Dieteria bigelovii (A. Gray) Morgan & R.L. Hartm. var. bigelovii [2,8]

A, C, M, R, T; 6120-9700'; S; br, fr, mm, ms, pj, ra.

Dieteria canescens (Pursh) Nutt. var. ambigua (B.L. Turner) Morgan

& R.L. Hartm. [1,-] A; 5800'; L; fr.

Dieteria canescens (Pursh) Nutt. var. aristata (Eastw.) Morgan & R.L.

Hartm. [-,2] R,T; 5781-7750'; G, S; ra.

Dieteria canescens (Pursh) Nutt. var. glabra (A. Gray) Morgan & R.L.

Hartm. [-,1] C; 7650-8600'; S; br.

* Dyssodiapapposa (Vent.) C.L. Hitchc. [1,2] M, T; 7200-7700'; S;

mm, pj, ra.

Engelmannia peristenia (Raf.) Goodman & Lawson [2,-] A; 5570-

5800'; L; fr.

Ericameria nauseosa (Pall, ex Pursh) G.L. Nesom & G.l. Baird var.

bigelovii (A. Gray) G.L. Nesom & G.L Baird [UNM-E. Wooton

s.n., 24 Aug 1910] S; S.

Ericameria nauseosa (Pall, ex Pursh) G.L. Nesom & G.l. Baird var.

graveolens (Nutt.) Reveal & Schuyler [-,1 ] T; 7100'; S; ra.

Ericameria nauseosa (Pall, ex Pursh) G.L. Nesom & G.l. Baird var .holo-

leuca (A. Gray) G.L. Nesom & G.l. Baird [-,1 ] T; 7700-8555'; P; pj.

Ericameria nauseosa (Pall, ex Pursh) G.L. Nesom & G.L Baird var.

latisquamea (A. Gray) G.L. Nesom & G.l. Baird [UNM-H. Bobisud

37] S; S.

Ericameria nauseosa (Pall, ex Pursh) G.L. Nesom & G.l. Baird var.

oreophila (A. Nelson) G.L. Nesom & G.l. Baird [-,6] C, T; 7240-

9400'; S; br, fr, ml, mm.

Ericameria parryi (A. Gray) G.L. Nesom & G.l. Baird var. affinis (A.

Nelson) G.L. Nesom & G.l. Baird [UNM-A. Cully CU-1 ] T; G.

Erigeron canus A. Gray [1,-] S; 7300-7400'; S; pj.

! Erigeron compositus Pursh [-,1 ] T; 11600-12450'; S; am.

Erigeron concinnus (Hook. & Arn.) Torr. & A. Gray var. concinnus [-,1 ]

T; 6950'; P; pj.

Erigeron coulteri Porter [12,26] A, M, R, S, T; 8200-12000'; S; mm,

mr, sf.

Erigeron divergens Torr. & A. Gray [21,32] A, C, R, S, T; 5730-9400';

G, L, P, S; br, me, mm, mr, ms, pg, pj, pp, ra.

Erigeron elatior (A. Gray) Greene [1,-] M; 9920-9960'; S; af, mm.

Erigeron eximius Greene [15,16] A, C, M, R, S, T; 7840-11800'; S;

am, af, me, mm, mr, sf.

Erigeron flagellaris A. Gray [29,53] A, C, M, R, S, T; 7050-11129'; G,

P, S; af, bw, br, ds, fr, me, mm, ms, mr, pj, pp, ra, sf.

Erigeron formosissimus Greene var. formosissimus [5,10] A, M, R,T;

7850-12000'; S; bw, me, mm, mr, sf.

285

Erigeron formosissimus Greene var. viscidus (Rydb.) Cronquist

[11,23] A, C, M, R, S,T; 7000-11750'; S; me, mm, ms, mr, pp, ra, sf.

Erigeron glabellus Nutt. [-,1] T; 7050'; S; mr.

Erigeron glacialis (Nutt.) A. Nelson var. glacialis [4,6] A, M, R, T;

9700-12000'; S; mm, mr, sf.

Erigeron grandiflorus Hook. [1,9] A, R,T; 9800-13024'; S; am, mm, sf.

Erigeron leiomerus A. Gray [UNM-T. Lowrey 2082] C; S.

Erigeron melanocephalus (A. Nelson) A. Nelson [2,9] M, R, S, T;

9800-13024'; S; am, mm, sf.

! Erigeron nivalis Nutt. [-,1 ] T; 9600-10900'; S; sf.

Erigeron pinnatisectus (A. Gray) A. Nelson [-,5] R,T; 11600-13024';

S; am.

Erigeron pulcherrimus A. Heller [-,4] C, R, T; 5781-8050'; G, S; ds,

fr, mr, pp.

Erigeron speciosus (Lindl.) DC. [6,1] A, M, R, S; 7700-11300'; S; me,

mm, ms, sf.

+ Erigeron subglaber Cronquist [2,-] A, M; 11310-11750'; S; mm.

Erigeron subtrinervis Rydb. ex Porter & Britton [13,35] A, C, M, S, T;

7200-12960'; P, S; am, af, bw, br, ds, fr, me, mm, ms, mr, pj, pp, ra.

Erigeron tracyi Greene [10,11 ] A, R, S,T; 5700-10500'; G, L, P, S; br,

fr, mm, pg, pj, pp, ra.

Erigeron vetensis Rydb. [-,22] C, R, T; 7200-12960'; S; am, bw, me,

mm, mr, pj, pp, sf.

Erigeron vreelandii Greene [6,-] A, S; 7720-9860'; S; af, me, ms, pp.

Gaillardia aristata Pursh [-,3] C, T; 8500-11000'; S; mr, ra.

GaillardiapinnatifidaJorr. [4,1] A, T; 5570-6500'; L, P; pg, pj.

Gaillardia pulchella Foug. [3,2] A, T; 5500-10500'; L, S; fr, pj, ra.

Gnaphalium exilifolium A. Nelson [-,1 ] C; 8194'; S; ml.

Grindeliasquarrosa (Pursh) Dunal [11,10] A, C, M, R, S,T; 5500-9320';

L, P, S; fr, me, ml, mm, ms, pj, pp, ra.

Gutierrezia sarothrae (Pursh) Britton & Rusby [5,7] A, C, M, R, S, T;

5560-8555'; L, P, S; br, mm, ms, pg, pj, pp, ra.

Helenium autumnale L. var. montanum (Nutt.) Fernald [UNM-R.

Wallace 92EM023-F2] M; S.

Helianthella parryi A. Gray [8,30] A, C, M, R, T; 7650-12700'; S; am,

af, bw, me, mm, ms, mr, pp, ra, sf.

Helianthella quinquenervis (Hook.) A. Gray [3,4] M, S,T; 8450-12050';

S; af, me, mm, mr, sf.

Helianthus annuus L. [3,6] A, M, T; 5500-10500'; L, S; fr, mm, pj, ra.

Helianthus nuttallii Torr. & A. Gray [UNM-R. Wallace 92RW002-F3]

T; G.

Helianthuspauciflorus Nutt. var. subrhomboideus (Rydb.) Cronquist

[2,2] A, M, R; 7000-9320'; S; br, pp, ra.

Helianthuspetiolaris Nutt. var. petiolaris [1,-] A; 7000'; S; ms, pp.

Heliomeris multiflora Nutt. var. multiflora [15,11] A, M, R, S, T;

7000-11200'; S; af, br, me, mm, pp, ra, sf.

Heliomeris multifloraNutt. var. nevadensis (A. Nelson) W.F. Yates [2,1 ]

R, S; 7000-8400'; S; br, mm, pp.

Heliopsis helianthoides (L.) Sweet var. scabra (Dunal) Fernald [7,2]

A, C, M,T; 7720-9400'; S; af, me, pj, ra.

♦ Herrickia horrida Wooton & Standi. [-,1 ] C; 7800-8400'; S; me.

Heterotheca villosa (Pursh) Shinners var. minor (Hook.) Semple

[14,21 ] A, C, M, R, S, T; 5800-10500'; G, P, S; br, ds, fr, me, mm,

mr, ms, pj, pp, ra.

Heterotheca villosa (Pursh) Shinners var. nana (A. Gray) Semple

[11,15] A, C, M, R, S, T; 6200-9750'; G, P, S; ds, fr, me, mm, mr,

pj, PP, ra.

Heterotheca villosa (Pursh) Shinners var. villosa [10,12] A, C, M, R, S,

T; 5500-9880'; G, L, S; br, ds, me, mm, pj, ra.

Hieracium fendleri Sch. Bip. [4,8] A, C, M, R, S, T; 7850-11209'; S;

me, mm, mr, pp, sf.

Hieracium pringlei A. Gray [1,-] A; 8000'; S; me, mm.

Hieracium triste Willd. ex Spreng. [2,9] M, R, S, T; 8450-12060'; S;

bw, me, mm, mr, sf.

286

Journal of the Botanical Research Institute of Texas 8(1)

Hymenopoppus filifolius Hook. var. cinereus (Rydb.) I.M. Johnst. [8,12]

A, R, S, T; 5781 -7950'; G, P, S; ds, pj.

Hymenopappus filifolius Hook. var. pauciflorus (I.M. Johnst.) B.L.

Turner [-,1] R; 6380'; G; fr.

Hymenopappus flavescens A. Gray var. canotomentosus A. Gray [2,-]

A; 5570-5800'; L; fr, pj.

Hymenopappus flavescens A. Gray var. flavescens A. Gray [1,-] A;

6200'; L; pg, pj.

Hymenopappus newberryi (A. Gray) I.M. Johnst. [20,9] A, C, M, T;

7720-10700'; S; af, me, mm, mr, ra.

Hymenopappus tenuifolius Pursh [6,-] A; 5700-7500'; L, S; ds, pg, pj.

Hymenoxys brandegeei (Porter ex A. Gray) Parker [5,11] M, R, S, T;

9800-13161'; S; am, mm, mr, sf.

Hymenoxys hoopesii (A. Gray) Bierner [20,21] A, C, M, S, T; 7850-

11800'; S; me, mm, mr, sf.

Hymenoxys richardsonii (Hook.) Cockerell var. floribunda (A. Gray)

Parker [12,20] A, R, S, T; 5800-10500'; G, P, S; ds, fr, me, mm,

mr, pj, pp, ra.

Krigia biflora (Walter) S.F. Blake [1,-] A; 8400-8850'; S; me, mm.

! Lactuca biennis (Moench) Fernald [3,-] A, S; 7760-8600'; S; mm,

mr.

Lactuca canadensis L. [1,2] S, T; 7400-10500'; S; mm, mr.

Lactuca graminifolia Michx. var. arizonica McVaugh [2,-] A;

7720-8000'; S; me, mm.

Lactuca pulchella (Pursh) DC. [-,2] T; 7550-9800'; S; ml, mm.

* Lactuca serriola L. [5,16] A, C, M, S, T; 5500-10100'; L, S; br, fr,

mm, mr, pg, pj, ra.

Laennecia schiedeana (Less.) G.L. Nesom [1,-] M; 8040-8700'; S;

me, ra.

Leibnitzia lyrata (Sch.Bip.) G.L. Nesom [UNM-R. Sivinski 5763] A; S.

* Leucanthemum vulgare Lam. [4,9] A, C, R, T; 7350-10500'; S; fr,

ml, mm, mr, ra. [2,1 ] A, R; 7000-8200'; S; ms, pp.

Liatrispunctata Hook. var. punctata [2,1 ] A, R; 7000-8200'; S; ms, pp.

Lygodesmia juncea (Pursh) D. Don ex Hook. [2,-] A; 7200-7500';

S; ds, pj.

Machaeranthera tanacetifolia (Kunth) Nees [1,-] A; 561 O'; L; fr.

* Matricaria discoidea DC. [-,1 ] T; 7200'; S; mr.

Melampodium leucanthum Torr. & A. Gray [8,-] A; 5630-6750'; L,

S; fr, pg, pj.

Oreochrysum parryi (A. Gray) Rydb. [20,24] A, C, M, R, S, T; 7700-

12450'; S; am, af, me, mm, mr, ms, pp, ra, sf.

Packera dimorphophylla (Greene) W.A. Weber & A. Love var. dimor-

phophylla [-,4] T; 9700-11500'; S; mr.

Packera fendleri (A. Gray) W.A. Weber & A. Love [33,35] A, C, M, R,

S, T; 5781 -11500'; S; af, bw, br, me, mm, mr, ms, pj, pp, ra, sf.

Packera hartiana (A. Heller) W.A. Weber & A. Love [UNM-H. Mackay

6T-207] T; S.

Packera multilobata (Torr. & A. Gray ex A. Gray) Weber & A. Love [1,4]

C, M, R,T; 7250-10500'; S; br, mm, pj, pp.

Packera neomexicana (A. Gray) W.A. Weber & A. Love var. mutabilis

(Greene) W.A. Weber & A. Love [3,12] A, C, R, S,T; 7150-10500';

G, S; me, mr, pj, pp.

Packera pseudaurea (Rydb.) W.A. Weber & A. Love var. flavula

(Greene) D.Trock&T.M. Barkley [-, 1]T; 10500-11000'; S; mm.

+ Packera sanguisorboides (Rydb.) W.A. Weber & A. Love [23,23]

A, C, M, R, S, T; 7840-11940'; S; af, me, mm, mr, sf.

Packera streptanthifolia (Greene) W.A. Weber & A. Love [1,16] R, S,

T; 6600-12450'; P, S; am, af, br, me, mm, mr, sf.

Packera thurberi (A. Gray) B.L. Turner [UNM-H. Dixon V-24] T; S.

Pericome caudata A. Gray [-,1 ] T; 7400-7493'; R; fr.

Petradoria pumila (Nutt.) Greene var. pumila [-,8] R,T; 7200-8763';

P, S; br, pj.

Picradeniopsis oppositifolia (Nutt.) Rydb. [UNM-R. Sivinski 4542] C; S.

Plectocephalus americanus (Nutt.) D. Don [1,-] A; 5650-5740'; L;

P9, Pj-

Pseudognaphalium macounii (Greene) Kartesz [5,-] A, M, S; 8200-

10850'; S; me, mm, ms.

Pseudognaphalium stramineum (Kunth) W.A. Weber [2,-] A, S;

7000-8000'; S; me, mm, mr.

PsHostrophetagetina (Nutt.) Greene [6,-] A; 5570-5840'; L; fr, pg, pj.

Pyrrhopappus pauciflorus (D. Don) DC. [2,-] A, R; 5500-6620'; G,

L; ds, fr.

Pyrrocoma crocea (A. Gray) Greene var. crocea [4,4] A, M, R, T;

8700-11300'; S; mm

Ratibida columnifera (Nutt.) Wooton & Standi. [11,6] A, C, M, T;

5570-10300'; L, S; mm, pg, pj, pp, ra.

[Ratidiba columnifera formas]

Ratibida tagetes (E. James) Barnhart [3,-] A; 5570-7240'; L, S; fr, pj.

Rudbeckia hirta L. var. pulcherrima Farw. [22,12] A, C, M, S, T;

6500-11500'; P, S; fr, mm, mr, ra.

Rudbeckia laciniata L. var. ampla (A. Nelson) Cronquist [32,10] A,

C, M, S, T; 7700-10850'; G, S; me, mm, mr.

! Rudbeckia laciniata L. var. laciniata [2,-] A, S; 7000-7750'; S; mm,

mr.

Sanvitalia abertii A. Gray [UNM-R. Sivinski 2574] S; S.

Schkuhria multiflora Hook. & Arn. [UNM-R. Sivinski 1820] T; S.

* Scorzonera laciniata L. [5,8] A, R, S, T; 5800-8200'; G, L, P, S; br,

fr, ml, pg, pj, ra.

Senecio amplectens A. Gray var. amplectens [4,10] M, R, S, T;

9600-13024'; S; am, bw, mm, mr, sf.

Senecio amplectens A. Gray var. holmii (Greene) Harrington [2,3] M,

R, S, T; 10990-13024'; S; am, sf.

Senecio atratus Greene [2,18] M, R, S, T; 7850-11850'; S; bw, me,

mm, mr, ra, sf.

Senecio bigelovii A. Gray var. hallii A. Gray [15,14] A, M, R, S, T;

7750-12000'; S; me, mm, mr, sf.

Senecio crassulus A. Gray [2,4] M, R, S, T; 8400-13024'; S; am, mm,

PP, sf.

Senecioeremophilus Richardson var. kingii (Rydb.) Greenm. [13,15]

A, C, M, R, S, T; 7400-11500'; S; me, mm, mr, sf.

Senecio flaccidus Less. var. flaccidus [8,3] A, R, T; 5610-8300'; G, L,

P, S; fr, pg, pj, ra.

Senecio fremontii Torr. & A. Gray var. blitoides (Greene) Cronquist

[3,2] M, R, S, T; 11500-13024'; S; am, mm.

Senecio spartioides Torr. & A. Gray [-,1 ] T; 7240'; S; ra.

Senecio taraxacoides (A. Gray) Greene [-,7] T; 11300-13161'; S; am.

Senecio triangularis Hook. [6,25] M, R, S, T; 7850-12000'; S; mm,

mr, sf.

Senecio wootonii Greene [3,8] A, C, R, S,T; 7910-11209'; S; me, mr, sf.

Solidago altissima L. ssp. gilvocanescens (Rydb.) Semple [1,4] A,T;

6980-10093'; P, S; br, me, mm, pj, pp.

Solidago gigantea Aiton [2,-] A; 7750-8000'; S; mr, ra.

Solidago missouriensis Nutt. var. fasciculata Holz. [1,1] A, T;

7520-11200'; S; pp, sf.

Solidago missouriensis Nutt. var. missouriensis [1,4] A, C, T; 7650-

10300'; S; br, me, mm, sf.

Solidago mollis Bartl. [2,1] A, S,T; 7000-8260'; S; pp, ra.

Solidago nemoralis Aiton var. decemflora (DC.) Fernald [5,4] A, R, S,

T; 7950-8880'; S; br, me, mm, ms, pp.

Solidago simplex Kunth var. simplex [28,34] A, C, M, R, S, T; 7840-

13024'; S; am, af, bw, me, mm, mr, pp, ra, sf.

Solidago speciosa Nutt. var. pallida Porter [1,-] A; 7750'; S; mm.

Solidago velutina DC. ssp. sparsiflora (A. Gray) Semple [4,1 ] A, C, M,

S; 7440-9760'; S; me, ra.

Solidago wrightii A. Gray var. adenophora S.F. Blake [9,-] A, M, S;

7840-9760'; S; me, mm, mr, pp, ra.

*Sonchus asper (L.) Hill [2,1 ] A, R,T; 5800-7660'; G, L, S; ds, fr, ml.

Larson et al., Floristic studies of the Sangre de Cristo Mountains

Stephanomeria pauciflora (Torr.) A. Nelson [3,-] A; 5700-5840';

L; P9, Pj-

Symphyotrichum oscendens (Lindl.) G.L. Nesom [2,3] M, T; 8080-

9500'; S; me, mr, ra.

Symphyotrichum eatonii (A. Gray) G.L. Nesom [UNM-R. Wallace

92RW002-F5] T; S.

Symphyotrichum falcatum (Lindl.) G.L. Nesom var. commutatum

(Torr. & A. Gray) G.L. Nesom [-,1 ] R; 8150'; S; br.

Symphyotrichum foliaceum (Lindl. ex DC.) G.L. Nesom var. canbyi (A.

Gray) G.L. Nesom [-,3] M,T; 7400-10700'; S; mm, mr.

Symphyotrichum foliaceum (Lindl. ex DC.) G.L. Nesom var. parryi (D.C.

Eaton) G.L. Nesom [-,2] T; 7850-10500'; S; mr.

Symphyotrichum laeve (L.) A. Love & D. Love var. geyeri (A. Gray)

G.L. Nesom [12,2] A, M, R, S; 7700-9320'; S; af, me, mm, mr,

ms, pp, ra.

Symphyotrichum lanceolatum (Willd.) Nesom var. hesperium (A. Gray)

G.L. Nesom [3,-] A, S; 7400-8260'; S; mm, ra.

Symphyotrichum ported (A. Gray) G.L. Nesom [2,-] A; 7000'; S;

mm, pp.

Taraxacum ceratophorum (Ledeb.) DC. [-,1 ]T; 10500-11000'; S; mm.

* Taraxacum erythrospermum Andrz. ex Besser [1,9] C, R, S, T;

7700-10100'; S; br, mm, ms, mr, pj, pp.

* Taraxacum officinale Weber ex F. H. Wigg. [15,50] A, C, M, R, S,

T; 7050-12000'; G, S; br, fr, me, ml, mm, ms, mr, pj, pp, ra, sf.

Tetradymia canescens DC. [-,3] C,T; 7200-8500'; S; br, pj.

Tetraneuris acaulis (Pursh) Greene var. acaulis [1,2] S,T; 7250-7950';

S; pj, PP-

Tetraneuris acaulis (Pursh) Greene var. arizonica (Greene) Parker [4,-]

A; 5700-6200'; L, S; pg, pj.

Tetraneuris acaulis (Pursh) Greene var. caespitosa A. Nelson [1,4] A,

T; 7380-12600'; S; am, mm, pj.

Tetraneuris argentea (A. Gray) Greene [11,26] A, R, S,T; 6050-8700';

G, P, S; br, fr, pj, pp.

Tetraneuris scaposa (DC.) Greene var. scaposa [UNM-F. Broeke

Co-75] A; L.

Thelesperma fHi folium (Hook.) A. Gray var. intermedium (Rydb.)

Shinners [3,-] S; 7000-7350'; S; mm, pj, ra.

Thelesperma megapotamicum (Spreng.) Kuntze [16,2] A, R, S, T;

5570-7950'; G, L, S; ds, fr, ml, mm, pg, pj, pp.

Tonestus pygmaeus (Torr. & A. Gray) A. Nelson [2,9] R, S, T; 11990-

13024'; S; am, mm.

Townsendia eximia A. Gray [21,18] A, M, R, S,T; 7000-10800'; S; br,

me, mm, ms, pj, pp, ra.

Townsendia exscapa (Richardson) Porter [2,8] A, C, S,T; 5625-8555';

L, P, S; ds, fr, pg, pj, pp.

Townsendia fendleri A. Gray [1,-] S; 6540'; G; pj.

Townsendia leptotes (A. Gray) Osterh. [-,3] T; 7250-8763'; P, S; pj, pp.

* Tragopogon dubius Scop. [27,37] A, C, M, R, S, T; 5750-10500';

G, L, P, S; af, br, ds, me, ml, mm, mr, pj, pp, ra, sf.

* Tragopogon porrifolius L. [2,-] A; 7840-8325'; S; mm, ra.

* Tragopogon pratensis L. [ 11,4] A, M, R, S, T; 7000-10660'; S; me,

mm, mr, ra.

!* Tripleurospermum inodorum (L.) Schultz-Bip. [-,1 ] T; 8175'; S; ml.

Verbesina encelioides (Cav.) Benth. & Hook. f. ex A. Gray [3,1 ] A, S, T;

5500-7450'; L, S; fr, mm, ra.

[Verbesina encelioides var. exauriculata B.L. Rob. & Greenm.]

Xanthisma gracile (Nutt.) Morgan & R.L. Hartm. [1, -] S; 7000'; S;

mm, mr.

Xanthisma grindelioides (Nutt.) Morgan & R.L. Hartm. [1,-] S; 6540';

S; pj-

Xanthisma spinulosum (Pursh) Morgan & R.L. Hartm. var. glaber-

rimum (Rydb.) Morgan & R.L. Hartm. [-,1] G; 6540'; R; fr.

Xanthisma spinulosum (Pursh) Morgan & R.L. Hartm. var .spinulosum

[16,11 ] A, R, S, T; 5610-7600'; G, L, P, S; ds, fr, pg, pj, ra.

287

* Xanthium spinosum L [1,-] A; 561 O'; L; fr.

Xanthium strumarium L. var. canadense (Mill.) Torr. & A. Gray [1,-]

A; 5500'; L; fr.

Zinnia grandiflora Nutt. [7,-] A; 5610-6200'; L, S; pg, pj.

Berberidaceae

Berberis fendleri A. Gray [14,7] A, M, S,T; 7000-9400'; P, S; me, mm,

mr, pj, pp.

Berberis fremontiiTorr. [3,-] A; 5500-6150'; L, S; fr, pj.

Berberis repens Lindl. [7,27] C, R, S, T; 7350-10100'; S; af, br, me,

mm, mr, pj, pp.

* Berberis vulgaris L. [UNM-J. Carter 935] T; G.

Betulaceae

Alnus incana (L.) Moench var. occidentalis (Dippel) C.L. Hitchc.

[34,32] A, C, M, R, S, T; 6040-10500'; S; mr, ra.

[Alnus incana ssp. tenuifolia (Nutt.) Breitung]

Alnus oblongifoliaTon. [1,-] S; 7000'; S; mr.

Betula occidentalis Hook. [-,6] R,T; 6500-9400'; P, S; me, mr.

Boraginaceae

Cryptantha cinerea (Greene) Cronquist var. cinerea [9,3] A, S, T;

5700-7976'; L, S; fr, mm, mr, pg, pj, ra.

Cryptantha crassisepala (Torr. & A. Gray) Greene var. elachantha I.M.

Johnst. [-,2] R,T; 5781-6380'; G; fr, ra.

Cryptantha fulvocanescens (S. Watson) Payson var. fulvocanescens

[-,4] R; 6036-6540'; G, P; ds, fr, pj.

Cryptantha minima Rydb. [5,-] A; 5700-7500'; L, S; ds, pg, pj.

* Cynoglossum officinale L. [1,19] C, M, T; 6055-10500'; S; fr, me,

ml, mm, mr, ra.

Eritrichum nanum (Vill.) Schrad. ex Gaudin var. elongatum (Rydb.)

Cronquist [-,4] T; 11500-13009'; S; am.

Hackelia besseyi (Rydb.) J.L. Gentry [2,-] A, S; 7800-9200'; S; pp, me.

Hackelia floribunda (Lehm.) I.M. Johnst. [3,9] A, C, M,T; 7650-10500';

S; me, ml, mm, mr, ra.

+♦ Hackelia hirsuta [Wooton & Standi.) I.M. Johnst. [1,6] A, C, M,T;

7650-11000'; S; me, mm, ra.

Lappula occidentalis (S. Watson) Greene var. cupulata (A. Gray) L.C.

Higgins [3,1] A, S,T; 6380-7420'; S; mm, pj.

Lappula occidentalis (S. Watson) Greene var. occidentalis [10,40] A,

C, R, S,T; 5781 -9750'; G, P, S; br, ds, fr, me, ml, mm, mr, pj, pp, ra.

* Lappula squarrosa (Retz.) Dumort. [-,1 ] T; 8175'; S; ml.

Lithospermum incisum Lehm. [7,12] A, C, R, S,T; 5750-8830'; G, L,

P, S; ds, fr, me, pg, pj, pp, ra.

Lithospermum macromeria J. Cohen [3,-] A; 7720-8600'; S; mm,

PP, ra.

Lithospermum multiflorum Torr. ex A. Gray [13,14] A, C, S, T;

7000-11500'; S; br, me, mm, mr, pj, pp, ra.

Mertensia alpina (Torr.) G. Don [UNM-H. Mackay 5T-318] T; S.

Mertensia ciliata (E. James ex Torr.) G. Don [-,7] T; 9300-13161'; S;

am, bw, mr, sf.

Mertensia franciscana A. Heller [31,40] A, M, R, S, T; 7350-12850';

S; bw, me, mm, mr, pj, sf.

Mertensia lanceolata (Pursh) DC. [2,18] A, C, R, S, T; 7350-12850';

S; am, br, me, mm, mr, ra.

[Mertensia lanceolata varieties]

* Symphytum officinale L. [UNM-R. Sivinski 3125] S; S.

Brassicaceae

* Alyssum alyssoides (L.) L. [-,1 ] R; 8900-9100'; S; ra.

* Alyssum desertorum Stapf [-,1 ] T; 7200'; S; ra.

* Alyssum simplex Rudolphi [-,15] C, R,T; 5800-10000'; G, P, S; br,

ds, fr, ml, mm, mr, pj, pp, ra.

Arabis pycnocarpa M. Hopkins var. pycnocarpa [-,1] T; 7840-8500';

S; mm.

[Arabis hirsuta (L.) Scop. var. pycnocarpa (M. Hopkins) Rollins]

Barbarea orthoceras Ledeb. [1,1] A, T; 6053-7800'; P, S; mr, ra.

288

Journal of the Botanical Research Institute of Texas 8(1)

* Borboreo vulgaris R. Br. [2,3] A, C, R, S, T; 5800-8000'; L, S; fr, mr.

x Boechera xdivaricarpa (A. Nelson) A. Love & D. Love [-,11 ] C, R,

T; 7150-10500'; G, S; br, me, mm, mr, pj, pp, ra

[A hybrid between Boechera stricta and another taxon]

Boechera fendleri (S. Watson) W.A. Weber [-,19] C, R, T; 6600-10500';

G, P, S; br, me, mm, mr, pj, pp.

Boechera lignifera (A. Nelson) W.A. Weber [-,2] T; 7380-8350'; S; pj.

[error: B. gracilenta (Greene) Windham & Al-Shehbaz]

Boechera pallidifolia (Rollins) W.A. Weber [-,6] T; 7100-8350'; S;

Pj, PP-

Boechera spatifolia (Rydb.) Windham & Al-Shehbaz [2,-] A, S;

7150-841 O'; S; mm, me.

Boechera stricta (Graham) Al-Shehbaz [4,22] A, C, M, S, T; 8400-

12850'; S; am, me, ml, mm, ms, mr, ra, sf.

* Camelina microcarpa Andrz. ex DC. [4,9] C, R, S, T; 7100-8550';

S; ds, me, ml, mm, me, mr, pj, pp, ra.

* Capsella bursa-pastoris (L.) Medik. [9,14] A, M, R, S, T; 7550-

10600'; S; br, me, mm, mr, pp, ra.

Cardamine cordifolia A. Gray var. cordifolia [ 12,21 ] A, C, M, R, S, T;

7900-12000'; S; me, mm, mr, sf.

* Chorispora tenella (Pall.) DC. [-,1 ] R; 6540'; G; ra.

* Conringia orientalis (L.) Dumort. [UNM-E. Castetter s.n., 6 Jul

1935] C; S.

Descurainia californica (A. Gray) O.E. Schulz [-,4] C,T; 7800-10500';

S; br, mm, mr.

Descurainia incana (Bernh. ex Fisch. & C.A. Mey.) Dorn var. incisa

(Engelm.) Kartez & Gandhi [4,9] A, C, M, S,T; 7215-10500'; S;

br, me, mm, mr, pj, pp.

[Descurainia longepedicellata (Fourn.) O. E. Schult]

Descurainia incana (Bernh. ex Fisch. & C.A. Mey.) Dorn var. macro-

sperma (O.E. Schulz) Dorn [3,-] A, M; 8700-10700'; S; mm, mr.

[Descurainia incana]

Descurainia incana (Bernh. ex Fisch. & C.A. Mey.) Dorn var. viscosa

(Rydb.) Dorn [13,9] A, M, R, S, T; 6200-10700'; P, S; ds, fr, me,

mm, pj, ra.

[Not included in taxon count; Descurainia longepedicellata

(Fourn.) O. E. Schult]

Descurainia obtusa (Greene) O.E. Schultz ssp. obtusa [ 1 ,-] S; 6540';

G; mr.

Descurainia pinnata (Walter) Britton var. filipes (A. Gray) M. Peck

[-,1 ] R; 8100'; S; pj.

[Not included in taxon count; Descurainia longepedicellata

(Fourn.) O. E. Schult]

Descurainia pinnata (Walter) Britton var. osmiarum (Cockerell) Shin-

ners [2,1] A; T; 6200-7500'; L, S; ds, pg, pj.

[Not included in taxon count; Descurainia longepedicellata

(Fourn.) O. E. Schult]

* Descurainia sophia (L.) Webb ex Prantl [5,16] A, C, R, S, T;

5781 -9100'; G, L, P, S; br, ds, fr, me, ml, mm, mr, pj, pp, ra.

Draba aureaM ahl ex Hornem. [-,14] R,T; 8350-13024'; S; am, me,

mr, sf.

Draba cana Rydb. [-,1 ] T; 11500-12700'; S; am.

Draba cuneifolia Nutt, ex Torr. & A. Gray var. cuneifolia [-,1] R;

7650'; S; pj.

Draba helleriana Greene var. blumeri C.L. Hitchc. [-,1 ] T; 8500-10500';

S; me.

Draba helleriana Greene var. helleriana [26,34] A, M, R, S, T;

7750-13009'; S; am, bw, me, mm, mr, pj, sf.

Draba helleriana Greene var. patens (Heller) O. E. Schulz [ 6 ,-] A, M;

9000-10660'; S; me, mm, ms.

! Draba nemorosa L. var. nemorosa [-,1 ] T; 7350-7450'; P; ds.

Draba reptans (Lam.) Fernald [-,2] R,T; 7050-7740'; G, S; fr, pj.

Draba spectabilis Greene [1,1 ] S, T; 10150-11900'; S; mr.

Draba streptocarpa A. Gray [2,16] A, C, T; 8300-13009'; S; am me,

mm, mr, sf.

Erysimum capitatum (Douglas ex Hook.) Greene var. capitatum [2,5]

A, M, S, T; 6400-12584'; S; am, bw, me, mm, pj, sf.

Erysimum capitatum (Douglas ex Hook.) Greene var. elatum (Nutt.)

Torr. [31,35] A, M, R, S,T; 6100-13000'; G, P, S; am, br, ds, fr, me,

ml, mm, mr, ms, pj, pp, ra.

[Erysimum capitatum (Douglas ex Hook.) Greene var. purshii

(Durand) Rollins]

Erysimum inconspicuum (S. Watson) MacMill. [2,1] A, M, T; 8680-

12183'; S; am, me, mr.

Hesperidanthus linearifolius (A. Gray) Rydb. [6,9] A, R, S, T; 5500-

8540'; L, P, S; br, ds, fr, pg, pj, pp, ra.

Lepidium alyssoides A. Gray var. alyssoides [-,2] T; 7100-8175'; S;

ml, ra.

* Lepidium campestre (L.) R. Br. [-,1 ] T; 7100-7800'; S; ra.

Lepidium densiflorum Schrad. var. densiflorum [4,3] A, C, S, T;

5750-8000'; L, P, S; ds, ml, pj, ra.

! Lepidium densiflorum Schrad. var. macrocarpum G.A. Mulligan

[-,2] T; 6100-6500'; P; pj.

Lepidium densiflorum Schrad. var. ramosum (A. Nelson) Thell. [1,-]

S; 6540'; G; fr.

Lepidium lasiocarpum Nutt. var. wrightii (A. Gray) C.L. Hitchc. [-,1]

T; 5781'; G; fr, ra.

*• Lepidium latifolium L. [-,3] T; 5781 -8550'; G, P, S; fr, ra.

* Lepidium perfoliatum L. [-,1 ] T; 6053'; P; ra.

Lepidium ramosissimum A. Nelson var. bourgeauanum (Thell.) Rollins

[1,1] S,T; 7000-9750'; S; mm, mr, ra, sf.

Lepidium virginicum L. var. medium (Greene) C.L. Hitchc. [-,3] R, T;

6053-9400'; P, S; fr, pp, ra.

[Lepidium virginicum var. menziesii (DC.) Thell.]

Lepidium virginicum var. pubescens (Greene) Thell. [3,4] A, S, T;

6500-9400'; P, S; me, mr, pp, sf.

[Lepidium virginicum L. var. menziesii (DC.) Thell.]

* Nasturtium officinale R. Br. [-,1 ] T; 6600-6800'; P; fr.

Noccaea fendleri (A. Gray) Holub ssp. glauca (A. Nelson) Al-Shebaz

& M. Koch [8,27] A, M, R, S,T; 7250-11950'; S; bw, br, me, mm,

mr, pj, pp, sf.

Pennellia longifolia (Benth.) Rollins [3,-] A; 8000-8575'; S; me, ms.

Pennellia micranthra (A. Gray) Nieuwl. [1,4] A, C, R,T; 5840-10500';

L, S; me, pg, pj, pp.

Physaria calcicola (Rollins) O'Kane & Al-Shehbaz [2,-] S; 7100-7350';

S; pj-

Physaria fendleri (A. Gray) O'Kane & Al-Shehbaz [1,-] A; 5700-5800';

L; P9, Pj-

Physaria floribunda Rydb. var. floribunda [-,6] T; 7250-8550'; S; br, pj.

Physaria montana (A. Gray) Greene [2,9] R, S,T; 7050-8000'; G, P,

S; ds, fr, pj.

Physaria rectipes (Wooton & Standi.) O'Kane & Al-Shehbaz [1,10] R,

S,T; 6050-8500'; G, P, S; ds, pj.

Physaria valida (Greene) O'Kane & Al-Shehbaz [1,-] S; 7000-7200'; S;

Rorippapalustris (L.) Besser var. fernaldiana (Butters & Abbe) Stuckey

[-,1 ] T; 7240'; S; ra.

[Rorippa palustris var. palustris]

Rorippa sinuata (Nutt.) Hitchc. [UNM-R. Ivey s.n., 1 Sept 1992]T; S.

Rorippa sphaerocarpa (A. Gray) Britton [3,-] A, M; 8375-9600'; S;

me, mm.

* Rorippa sylvestris (L.) Besser [1,-] A; 5500'; L; fr.

* Sisymbrium altissimum L. [8,17] A, C, R, S, T; 5781 -9600'; G, P,

S; br, ds, fr, ml, mm, mr, pj, pp, ra.

* Sisymbrium loeselii L. [-,2] T; 7100-7240'; S; ra.

Streptanthella longirostris (S. Watson) Rydb. [-,1 ] T; 5781'; G; ds, fr.

Streptanthus cordatus Nutt. var. cordatus [-,2] T; 7600-8900'; S; pj, pp.

+ Thelypodiopsis vaseyi (S. Watson ex B.L. Rob.) Rollins [7,3] A, M,

T; 7900-10660'; S; af, me, mm, ra.

Thelypodium wrightii A. Gray ssp. wrightii [-,1 ] T; 7600-8300'; S; pj.

Larson et al., Floristic studies of the Sangre de Cristo Mountains

* Thlaspi arvense L. [-,4] R, T; 5781 -9500'; G, P, S; fr, me, mr, ra.

Turritis globro L. [9,2] A, C, S; 8230-9320'; S; mm, mr, ra.

Cactaceae

Coryphontho viviporo (Nutt.) Britton & Rose [-,3] T; 7600-8450';

S; pj, PP-

Cylindropuntia imbricata (Haw.) Knuth var. imbricata [9,9] A, R, S,T;

5585-8300'; G, L, P, S; ds, fr, pg, pj.

Echinocereus coccineus Engelm. [5,9] A, C, R, S,T; 5625-8900'; G, L,

P, S; ds, mm, me, pg, pj, pp.

Echinocereus triglochidiatus Engelm. [2,10] A, R,T; 5781-8450'; G,

L, P, S; ds, fr, pj, pp, ra.

Echinocereus viridiflorus Engelm. [-,15] C, R, T; 6200-8277'; G, P,

S; ds, fr, pj, pp.

Opuntia engelmannii Salm-Dyck ex Engelm. var. engelmannii [1,-]

A; 5700-5800'; L; pj.

Opunita macrorhiza Engelm. [UNM-E. Castetter 1250] T; S.

Opuntia phaeacantha Engelm. [9,9] A, C, R, S, T; 5730-8300'; G, L,

P, S; ds, mm, pg, pj, pp.

Opuntia polyacantha Haw. var. polyacantha [5,28] A, C, R, S, T;

5730-8900'; G, L, P, S; ds, fr, me, mm, pg, pj, pp, ra.

Pediocactus simpsonii (Engelm.) Britton & Rose [UNM-W. Sedlacek

3769] T;S.

Campanulaceae

Campanula parryi A. Gray var. parryi [2,11 ] A, C, R, T; 7400-10400';

S; me, mm, mr, ra.

Campanula rotundifolia L. [49,51 ] A, C, M, R, S, T; 7000-13024'; S;

am, bw, br, me, mm, ms, mr, pj, pp, ra, sf.

Campanula uniflora L. [UNM-H. Mackay 9T-1 ] T; S.

Lobelia cardinalis L. [UNM-F. Bartlette s.n., Aug 1905] A; S.

Cannabaceae

Cannabis sativa L. var. sativa [1,-] M; 7700'; S; mr.

Celtis occidentalis L. [1,-] A; 5500'; L; fr, pj.

Celtis reticulataJon. [1,-] A; 5700-5800'; L; fr, pj.

Humulus lupulus L. var. neomexicanus A. Nelson & Cockerell [4,1 ] A,

S,T; 7580-8400'; S; mm, mr.

Caprifoliaceae (includes Valerianaceae)

Linnaea borealis L. var. longiflora Torr. [2,4] M, S, T; 8410-10800';

S; me, mr, sf.

Lonicera involucrata (Richardson) Banks ex Spreng. [18,20] A, M, R,

S, T; 7840-11950'; S; me, mm, mr, sf.

Symphoricarpos rotundifolius A. Gray [33,24] A, M, R, S, T; 7250-

10500'; S; br, me, mm, mr, pj, pp.

Valeriana acutiloba Rydb. var. acutiloba [2,9] M, R, T; 7550-11850';

S; br, me, mm, pp, sf.

Valeriana arizonica A. Gray [2,-] A, S; 7740-8180'; S; mr.

Valeriana edulis Nutt, ex Torr. & A. Gray [17,16] A, C, M, S, T; 7050-

12850'; S; bw, br, me, mm, mr, ra, sf.

Caryophyllaceae

Arenaria lanuginosa (Michx.) Rohrb. var. saxosa (A. Gray) Zarucchi,

R.L. Hartm., & Rabeler [15,23] A, C, M, S,T; 7840-12000'; S; af,

bw, me, mm, mr, ms, ra, sf.

[Spergulastrum lanuginosum Michx. ssp. saxosum (A. Gray) W.A.

Weber]

Cerastium arvense L. var. strictum (Gaudin) Koch [-, 20] C, T;

7620-13009'; S; am, mm, mr, sf.

Cerastium brachypodum (Engelm. ex A. Gray) B.L. Rob. [3,-] A, M;

8600-9340'; S; me, mm, mr.

* Cerastium fontanum Baumg. ssp. vulgare (Hartm.) Greuter &

Burdet [17,2] A, M, S,T; 7000-10880'; S; af, me, mm, mr, ra.

* Dianthus armeria L. [-,1 ] R; 7600-7750'; S; pj.

Eremogoneeastwoodiae (Rydb.) Ikonn. var. adenophora (Kearney &

Peebles) R.L. Hartm. & Rabeler [-,3] R; 6050-6500'; G, P; fr, pj.

289

Eremogone eastwoodiae (Rydb.) Ikonn. var. eastwoodiae [2,1] R, S;

6540-7050'; G; ds, fr, pj.

Eremogone fendleri (A. Gray) Ikonn. [22,27] A, C, M, R, S, T; 7000-

13024'; P, S; am, bw, fr, me, mm, mr, pp, sf.

Minuartiaobtusiloba (Rydb.) House [7,20] A, M, R, S,T; 9800-13024';

S; am, mm.

Minuartia rubella (Wahlenb.) Hiern [-,4]T; 10500-12700'; S; am, mm.

Moehringia lateriflora (L.) Fenzl [-,1] T; 9475'; S; mr.

Moehringia macrophylla (Hook.) Fenzl [1,7] S, T; 7620-10500'; S;

br, me, mr.

Paronychia jamesii Torr. & A. Gray [2,-] A; 5730-7500'; L, S; pg, pj.

Paronychia pulvinata A. Gray [-,3] T; 10200-12500'; S; am.

Pseudostellaria jamesiana (Torr.) W.A. Weber & R.L. Hartm. [-,14] R,

T; 7550-11800'; S; br, me, mm, mr, pp, sf.

Sagina saginoides (L.) H. Karst. [-,1 ] T; 10700'; S; mr.

* Saponaria officinalis L. [2,-] A, S; 7400-8000'; S; mm, mr.

Silene acaulis (L.) Jacq. var. subacaulescens (F.N. Wms.) Fern. & St

John [1,7] S,T; 9800-12850'; S; am.

Silene antirrhina L. [-,1 ] T; 7600-8450'; S; pp.

Silene drummondii Hook. var. drummondii [8,16] A, M, R, S, T;

7840-12000'; S; me, mm, mr, pp, sf.

! Silene drummondii Hook. var. striata (Rydb.) Bocq. [-,4] C, M, T;

8300-10986'; S; me, mm, mr, sf.

! Silene hitchguirei Bocq. [-,1 ] T; 11500-12850'; S; am.

Silene latifolia Poiret ssp. alba (Miller) Greuter & Burdet [6,-] A;

7800-9750'; S; me, mm,

mr, ra.

* Silene noctiflora L. [1,-] M; 7700'; S; mm, ra.

Silenescouleri Hook. var. pringlei (S. Watson) C.L. Hitchc. & Maguire

ex Kartesz &

Gandhi [12,9] A, C, M, S,T; 8160-11500'; S; me, mm, mr, sf.

Stellarialongifolia Muhl. exWilld. [1,4] M,T; 7050-11333'; S; mm, mr.

Stellaria longipes Goldie var. longipes [4,16] A, M, S,T; 7850-12700';

S; am, me, mm, mr, pp, sf.

Stellaria umbellata Turcz. ex Karel. & Kir. [1,7] S, T; 9700-12700'; S;

am, me, mm, mr, sf.

* Vaccaria hispanica (Mill.) Rauschert [UNM-F. Bartlette s.n., Jul

1904] A; S.

Celastraceae

Paxistima myrsinites (Pursh) Raf. [14,31 ] A, C, M, R, S,T; 7350-12000';

P, S; br, me, mm, mr, pp, ra, sf.

Chenopodiaceae

Atriplex canescens (Pursh) Nutt. var. canescens [4,8] A, R, S, T;

5800-7550'; G, P, S; ds, fr, pj, ra.

* Bassia hyssopifolia (Pall.) Kuntze [-,2] T; 8550-9675'; G, P, S; br,

me, mr.

Chenopodium atrovirens Rydb. [4,8] A, M, T; 7750-11500'; S; mm,

mr, ra, sf.

Chenopodium berlandieri Moq. var. zschackei (Murr) Murr ex Asch.

[8,2] A, C, M, S; 7400-9200'; S; me, ml, mm, ra.

Chenopodium fremontii S. Watson [5,6] A, M, T; 7350-10500'; S; br,

fr, me, mm, pj, ra.

* Chenopodium glaucum L. var. glaucum [-,1 ] C; 8194'; S; ml.

Chenopodium glaucum L. var. salinum (Standi.) B. Boivin [UNM-E.

Castetter 3951 ]C;S.

Chenopodium incanum (S. Watson) A. Heller var. incanum [5,1] A,

T; 5610-7240'; L, S; fr, ds, mm, pg, pj, ra.

Chenopodium leptophyllum (Moq.) Nutt, ex S. Watson [UNM-K.

Goodrow 558] A; S.

* Chenopodium overi Aellen [7,4] A, M, S, T; 8160-9800'; S; me,

mm, ra.

[Chenopodium capitatum (L.) Ambrosi var. parvicapitatum S.L.

Welsh]

290

Journal of the Botanical Research Institute of Texas 8(1)

Chenopodium pallescens Standi. [1,-] A; 7000'; S; pp.

Chenopodium protericolo Rydb. [-,2] C,T; 7650-9700'; S; pj, mm.

[Chenopodium desiccatum A. Nelson var. leptophylloides (Murr)

Wahl]

Chenopodium watsonii A. Nelson [1,-] A; 5600-5650'; L; pg.

* Dysphonio botrys (L.) Mosyakin &Clemants [UNM-E. Kelley 247]

S; S.

Dysphonio groveolens (Willd.) Mosyakin & Clemants [4,6] A, C, R, S,

T; 7400-9400'; S; br, mm, mr, pp, ra.

* Kochio scoporio (L.) Schrad. [3,2] A, C, S, T; 5730-8750'; R, S, T;

mr, pg, pj, ra.

Kroscheninnikovio lonoto (Pursh) Meeuse & Smit [3,3] A, T; 5730-

8555'; L, P, S; ds, fr, pg, pj.

Monolepis nuttolliono (Schult.) Greene [-,3] T; 9200-11800'; S;

am, mr, sf.

* Solsolo collino Pall. [UNM-E. Kelley 304] S; S.

* Solsolo trogus L. [-,3] C, T; 7240-9400'; S; ml, pj, ra.

Cleomaceae [Capparaceae]

Cleomeserruloto Pursh [UNM-H. Dixon V-97]T; S.

[Peritomo serruloto (Pursh) A. DC.]

Polanisio dodecondro (L.) DC. var. trochyspermo (Torr. & A. Gray) H.H.

litis [2,-] A; 5500-561 O'; L; fr, pj.

Commelinaceae

Commelina dionthifolio Delile [5,1 ] A, C, S; 7000-8260'; S; ms, pp.

Commelino erecto L. var. ongustifolio (Michx.) Fernald. [2,-] A;

5500-5840'; L; ds, fr, pj.

Trodescontio occidentolis (Britton) Smyth var. occidentolis [4,-] A;

5500-6200'; L; fr, pg, pj.

Convallariaceae [traditionally in Liliaceae]

Moionthemum rocemosum (L.) Link var. omplexicoule (Nutt.) Dorn

[14,15] A, C, M, S,T; 7840-10850'; S; me, mm, mr, sf.

Moionthemum stellatum (L.) Link [8,13] A, C, M, S, T; 7620-11650';

S; me, mm, mr, sf.

Polygonotum biflorum (Walter) Elliott [1,-] M; 9760-10600'; S; mr.

Convolvulaceae

*• Convolvulus orvensis L. [10,5] A, M, R, S, T; 5800-9320'; G, L, S;

fr, ml, mm, pg, pj, ra.

Convolvulus equitons Benth. [1,-] A; 5650-5740'; L; pg, pj.

Evolvulus nuttollionus Roemer & Schult. [2,-] A; 5700-5840'; L;

pg, pj, ra.

Evolvulus sericeus Swartz var. sericeus [2,-] A; 5700-5840'; L; pg, pj, ra.

Ipomoeo cristuloto H. Hall [1,-] S; 6840-6880'; S; pj.

Ipomoeo leptophyllolorr. [1,-] A; 5570-5600'; L; fr.

Cornaceae

* Cornus canadensis L. [-,2] T; 9700-11500'; S; me, mm, mr, sf.

Comussericea L. var. sericea [11,12] A, C, S,T; 7400-9400'; S; me, mr.

Crassulaceae

Sedum cockerellii Britton [14,3] A, C, M, S, T; 7750-9400'; S; me,

mr, ms.

Sedum integrifolium (Raf.) A. Nelson ssp. integrifolium [10,19] A, C,

M, R, S,T; 7840-13009'; S; am, me, mm, mr, sf.

Sedum lanceolatum Torr. ssp. lanceolatum [-,18] R,T; 7600-12400';

S; am, bw, me, mm, mr, ra, sf.

Sedum rhodanthum A. Gray [3,4] M, R, T; 9900-12050'; S; am,

mm, mr, sf.

Sedum wrightii A. Gray [UNM-N.D. Atwood 21328] S; S.

Cucurbitaceae

Cucurbita foetidissima Kunth [4,-] A; 5570-7240'; L, S; fr, pj, ra.

Echinocystis lobata (Michx.) Torr. & A. Gray [UNM-R. Jackson 2341 ]

T; G.

Cyperaceae

Carexalbonigra Mack. [1,5] M,T; 10500-12850'; S; am, mm.

Corex oquotilis Wahlenb. var. aquatilis [-,10] M, T; 9700-11200'; S;

mm, mr, sf.

Corexoureo Nutt. [1,5] A, T; 8300-10500'; S; mm, mr.

Corex bello L.H. Bailey [5,19] A, M, R, S,T; 8300-11650; S; bw, mm,

mr, sf.

Corexbrevior (Dewey) Mack. ex. Lunell [1,-] A; 7200-7240'; S; ml.

Corex canescens L. var. canescens [2,3] M, S, T; 8410-12000'; S;

mm, mr, sf.

Corexcapillaris L. [-,1 ] T; 9700-11500'; S; mr.

Corexcholciolepis T. Holm [-,8] T; 10990-13009'; S; am, mm.

! Carexdeweyana Schwein. var. deweyana [1,3] S,T; 7620-9300';

S; mr, pp.

Corex dispermo Dewey [1,3] M, R,T; 8400-10180'; S; mr.

Carexdouglasii Boott [-,5] R,T; 7050-8500'; G, P, S; ds, fr, mm, mr, pp.

Carexduriuscula C.A. Mey. [-,4] C,T; 7200-9700'; S; mm, mr, pj, pp.

Carex ebenea Rydb. [6,12] A, M, R, S, T; 9375-13024'; S; am, bw,

mm, mr, sf.

Carexelynoides Holm [-,8] T; 11500-13161'; S; am.

Carexemoryi Dewey [-,3] R, T; 5781-10000'; G, S; fr, ra.

Carex geophila Mack. [-,30] C, R, T; 6100-11200'; G, P, S; bw, ds, fr,

me, ms, mr, pj, pp.

! Carex gynocrates Wormsk. ex Drejer [-,1 ] T; 9700-11500'; S; mr.

Carexillota L.H. Bailey [-,2] R,T; 11750-12960'; S; mr, sf.

Carex inops L.H. Bailey ssp. heliophila (Mack.) Crins. [2,17] A, C, R, S,

T; 7050-9600'; G, S; af, br, me, mm, ms, mr, pj, pp, ra.

Carex interior L.H. Bailey [1,1] M,T; 9900-12050'; S; mm, mr.

! Carexlenticularis Michx. var. lipocarpa (Holm) L.A. Standi. [2,-] S;

7580-841 O'; S; mr.

Carex micropodo C.A. Meyer [UNM-C. Keller 2218] T; S.

[Carex pyrenaica Wahlenb.]

Carexmicroptera Mack. [19,31 ] A, C, M, R, S,T; 7580-12960'; S; am,

bw, me, mm, mr, ms, sf.

Carex nebroscensis Dewey [2,15] C, M, S, T; 6500-11200'; G, P, S;

ml, mm, mr.

Carex nova L.H. Bailey var. novo [-,16] M, T; 9200-12960'; S; am,

me, mm, mr, sf.

Corex occidentolis L.H. Bailey [14,28] A, C, M, R, S,T; 5750-10000';

G, L, P, S; bw, br, ds, fr, me, mm, mr, ms, pp, ra, sf.

Carexoreocharis Holm [-,1] C; 10000-10600'; S; mm, sf.

Carexpellita Muhl. ex Willd. [2,3] A, C, T; 6600-8400'; P, S; fr, mr.

Corexpetosoto Dewey [-,2] T; 9375-12000'; S; bw, mr.

Corexphoeocepholo Piper [-,1 ] T; 12000-12625'; S; am.

Carex pityophila Mack. [-,9] C,T; 7775-12000; S; am, me, pp sf.

[Carexgeophila Mack.]

Corexproegrocilis Boott [-,4] R, T; 6500-10700'; P, S; fr, ml, mr.

! Corexroseo Schkuhr ex Willd. [2,-] A; 7800-8325'; S; me, mr.

Corexrossii Boott [-,17] R, T; 6036-11000'; P, S; br, me, mr, pp, sf.

Carexrupestris Bellardi ex All. var. drummondiana

(Dewey) L.H. Bailey [-,4] T; 10500-13000'; S; am.

Carexsiccata Dewey [2,16] A, C, M, R, T; 8300-12960'; S; am, me,

mm, mr, sf.

[Corex foeneo Willd. var. foeneo]

Corex stevenii (T. Holm) Kalela [3,8] M, S,T; 7620-11500'; S; mm, mr.

Carex stipata Muhl. ex Willd. var. stipata [6,2] A, T; 7850-10500';

S; mm, mr.

Carexsubfusca Boott [2,-] S; 8260-8940'; S; mr.

Corex tohoensis Smiley [UNM-R. Gierisch 3146] R; S.

Corexutriculoto Boott [2,12] A, C,T; 7050-11209'; S; ml, mm, mr, sf.

Carex vallicola Dewey [-,1 ] T; 7550'; S; br.

Carex vulpinoidea Michx. [1,-] A; 7740-7880'; S; mr.

Corex wootonii Mack. [2,-] A; 8900-9500'; S; me, mr.

Cyperus esculentus L. var. leptostachyus Boeck. [1,-] A; 5500'; L; fr.

Larson et al., Floristic studies of the Sangre de Cristo Mountains

Cyperus fendlerianus Boeck. [17,2] A, C, M, S; 5800-9080'; L, S; me,

mm, mr, pj, pp, ra.

Cyperus retroflexus Buckley var. pumilus (Britton) R. Carter & S.D.

Jones [1 7 -] A; 5500'; L; fr.

Cyperus schweinitzii Torr. [1,-] A; 5630'; L; fr.

Eleochoris bello (Piper) Svenson [1,-] A; 7200-7240'; S; ml.

Eleocharis engelmannii Steud. [1,-] A; 7200-7240'; S; ml.

Eleochoris erythropodo Steud. [UNM- E. Castetter 3390] C; S.

Eleochoris polustris (L.) Roem. &Schult. [1,12] C, M, R,T; 5781-10000';

G, P, S; fr, ml, mm, mr, ra.

Eleochoris quinquefloro (F.X. Hartm.) O. Schwarz [-,5] C, T; 8500-

10500'; S; mm, mr.

Eleochoris rostelloto (Torr.) Torr. [-,1 ] R; 6450'; G; fr.

Eriophorum ongustifolium Honck. ssp. ongustifolium [-,3] T;

10700-12050'; S; mr.

Kobresio myosuroides (Villars) Fiori & Paoli [-,2] T; 11990-12050';

S; am.

Schoenopiectus ocutus (Muhl. ex Bigelow) A. Love & D. Love var.

occidentolis (S. Watson) S.G. Sm. [-,2] T; 7550-9300'; S; ml, mr.

Schoenopiectus omericonus (Pers.) Volkart ex Schinz & R. Keller [-,2]

T; 6500-7660'; P, S; fr, ml.

Schoenopiectuspungens (Vahl) Palla var. longispicotus (Britton) S.G.

Sm. [2,1] M, R; 6380-7700'; G, S; fr, ml, mr.

Elaeagnaceae

*• Eloeognus ongustifolio L. [5,3] A, R, S, T; 5500-7400'; G, L, P, S;

fr, ml, ra.

Shepherdio canadensis (L.) Nutt. [16,14] A, C, M, R, S,T; 7620-10200';

S; me, mm, mr, pp.

Ericaceae

Arctostaphylospungens Kunth [UNM-K. Weissenborn 37] S; S.

Arctostaphylos uva-ursi (L.) Spreng. [18,13] A, C, M, R, S, T; 7720-

11209'; P, S; me, mm, mr, pp, sf.

Chimaphila umbel lata (L.) W.P.C. Barton var. occidentolis (Rydb.) S.F.

Blake [5,1 ] A, S,T; 8385-10440'; S; me, sf.

[Chimaphila umbelloto (L.) Nutt. var. acuta (Rydb.) S.F. Blake]

Goultherio humifuso (Graham) Rydb.[UNM-H. Mackey 6T-169] T, S.

Moneses unifloro (L.) A. Gray var. unifloro [1,16] M, R, S, T; 7850-

11500'; S; mm, sf.

Monotropo hypopithys L. [2,1] C, M, S; 8880-9040'; S; me.

Orthilio secundo (L.) House [11,20] A, M, R, S, T; 7850-12115'; S;

me, mm, sf.

Pterosporo ondromedeo Nutt. [9,3] A, C, S, T; 7580-10500'; S; me.

Pyrolo osorifolio Michx. var. osorifolio [1,3] C, S, T; 8960-10440';

S; me, sf.

Pyrolo chlorontho Sw. [1,6] S, T; 8350-11150'; S; me, mr, sf.

Pyrolo elliptico Nutt. [1,-] S; 8200-8320'; S; mr.

Pyrolo minor L. [4,4] A, M, S, T; 8410-11500'; S; me, mm, mr, sf.

Pyrolo picta Sm. [1,-] A; 8400-8900'; S; me.

Vaccinium myrtillus L. var. oreophilum (Rydb.) Dorn [11,5] A, M, R,

S,T; 8450-13024'; S; me, sf.

Vaccinium scoparium Leiberg ex Coville [-,15] C, T; 9400-12850';

S; am, me, mr, sf.

Euphorbiaceae

Chamaesyce fendleri (Torr. & A. Gray) Small var. chaetocalyx (Boiss.)

Shinners [2,-] S; 6540-7100'; G, S; pj, ra.

Chamaesyce fendleri (Torr. & A. Gray) Small var. fendleri [7,17] A, R,

S, T; 5700-7950'; G, L, P, S; ds, fr, pg, pj, ra.

Chamaesyceglyptosperma (Engelm.) Small [2,1 ] A, S,T; 5760-7100';

L, S; pg, pj, ra.

Chamaesyce serpyllifolio (Pers.) Small [4,7] A, C, M, R, S, T; 5610-

9400'; L, S; br, fr, mr, pj, pp, ra.

Chamaesycestrictospora (Engelm.) Small [UNM-E. Castetter 7078]

T, G.

291

Croton texensis (Klotzsch) Mull.Arg. [5,-] A, S; 5500-6880'; L, S; fr,

pg, pj, ra.

Euphorbia brochycero Engelm. [UNM-R. Sivinski 2221 ] A; S.

Euphorbia davidii Subils [2,-] S; 6840-7420'; S; ra.

Tragia nepetifolia Cav. [2,-] A; 5800-6750'; L, S; pg, pj.

Trogio ramosa Torr. [1,-] A; 5700-5800'; L; fr, pj.

Fabaceae

Amorpho canescens Pursh [3,2] A, C; 7000-8600'; S; ms, pp.

Astragalus ogrestis Douglas ex G. Don [-,1 ] C; 8400-8500'; S; pp.

Astragalus ollochrous A. Gray var. ployonus (M.E. Jones) Isley [-,2] T;

6600-8555'; P; ds, fr, pj.

Astragalus alpinus L. [-,6] T; 8700-11000'; S; mr, sf.

Astragalus crassicarpus Nutt. var. cavus Barneby [1,-] A; 6120-6180';

S; pj-

+♦ Astragalus cyaneus A. Gray [-,1 ] T; 7215'; S; pj.

Astragalus drummondii Douglas ex Hook. [-,8] R,T; 6600-9400'; P,

S; fr, me, pj, pp, ra.

Astragalus flexuosus (Hook.) Douglas ex G. Don var. flexuosus [1,2]

A,T; 7500-9500'; S; mm, mr, sf.

Astragalus grocilis Nutt. [1,-] A; 6200'; L; pg, pj.

Astragalus hallii A. Gray var. hallii [-,3] C, M, T; 8175-9320'; S; ml,

mr, ra.

Astragalus humistratus A.Gray var. humistratus [-,2] C,T; 7650-9750';

S; mm.

♦ Astragalus iodopetolus (Rydb.) Barneby [-,3] T; 7100-7660'; S;

Pj-

Astragalus kentrophyto A. Gray var. tegetorius (S. Watson) Dorn

[UNM-E. Castetter 10533] T; S.

Astragalus laxmannii Jacq. var. robustior (Hook.) S.L.Welsh & Barneby

[-,1]T; 9150'; S; mm.

Astragalus lentiginosus Douglas ex Hook. var. olbiflorus (A. Gray)

Schoener [-,4] R,T; 6036-6700'; P; fr, pj, ra.

Astragalus lonchocorpus Torr. [3,10] R, S, T; 5800-7660'; G, P, S;

ds, fr, ml, pj.

Astragalus lotiflorus Hook. [-,1 ] T; 5781'; G; fr, ra.

Astragalus missouriensis Nutt. var. missouriensis [2,6] A, R, S, T;

5750-8100'; G, L, P, S; ds, fr, pg, pj, pp, ra.

Astragalus multiflorus (Pursh) A. Gray [-,7] R, T; 6380-7950'; G, S;

fr, pj, ra.

[Astragalus tenellus Pursh]

Astragalus praelongus E. Sheld. var. praelongus [1,1] A, R; 6350-

6500'; P, S; pj.

+♦ Astragalus puniceus Osterh. var. gertrudis (Greene) Barneby [-,2]

T; 7215-8000'; S; pj.

Astragalus puniceus Osterh. var. puniceus [-,3] R, T; 6976-7215';

G, P, S; ds, pj.

Astragalus racemosus Pursh var. racemosus [UNM-K. Epperson s.n.,

16 May 2004] T; S.

Astragalus robbinsii (Oakes) A. Gray var. minor (Hook.) Barneby [-,1 ]

T; 9900-10500'; S; sf.

Astragalus scopulorum Porter [1,2] A, R, T; 7150-12200'; S; bw,

me, mr.

Colliondro humilis Benth. var. humilis [3,-] A; 5800-8200'; L, S; ms,

P9, Pj, PP-

Doleo condido Michx. ex Willd. var. oligophyllo (Torr.) Shinners [3,-]

A; 5570-7000'; L, S; pg, pj, pp.

Doleo formosa Torr. [2,1 ] A, R; 5800-6350'; G, L, S; ds, pj.

Dalea jamesii (Torr.) Torr. & A. Gray [2,1 ] A, R; 5750-6500'; L, P; pg, pj.

Doleo purpureoVe nt. var. purpurea [3,-] A; 7000-7520'; S; ms, pp.

Doleo tenuifolio (A. Gray) Shinners [2,-] A; 5700-5800'; L; pg, pj.

Desmanthus cooleyi (Eaton) Trel. [UNM-R. Sivinski 2414] S; S

Desmanthus illinoensis (Michx.) MacMill. ex B.L. Rob. & Fernald

[UNM-C.R. Hutchins 9946] S; S.

Glycyrrhiza lepidota Nutt, ex Pursh [1,1 ] S, T; 5781 -6540'; G; fr, ra.

292

Journal of the Botanical Research Institute of Texas 8(1)

Hedysorum boreole Nutt. var. boreole [UNM-C.R. Hutchins 5859] C; G.

Hoffmonnseggio dreponocorpo A. Gray [1,-] A; 5760'; L; pg, pj.

Lathyrus arizonicus Britton [UNM-C.R. Hutchins 6340] S; S.

Lothyrus eucosmus Butters & H. St.John [1,3] S, T; 7200-7500'; S;

fr, mr, ra.

Lothyrus gram/nfo/Zus T.G. White [UNM-D. Atwood 21434] A; S.

* Lothyrus lotifolius L. [UNM-C.R. Hutchins 8305] S; S.

Lothyrus leuconthus Rydb. [4,17] A, R, S,T; 7620-11500'; S; bw, me,

mm, mr, pj, pp, ra, sf.

Lupinus orgenteus Pursh var. orgenteus [-,14] R,T; 6850-10500'; G,

P, S; br, me, mr, pj, ra.

Lupinus orgenteus Pursh var. orgophyllus (A. Gray) S. Watson [9,10]

A, R, S, T; 6380-8150'; G, S; br, fr, ml, mm, mr, pj, ra.

[Lupinus coudotus Kellogg var. orgophyllus (A. Gray) S.L. Welsh]

Lupinus orgenteus Pursh var. fulvomoculotus (Payson) Barneby [1,1 ]

S,T; 8260-10500'; S; me, mm.

Lupinus orgenteus Pursh var. polmeri (S. Watson) Barneby [UNM-A.

Foster 78] T; S.

Lupinus orgenteus Pursh var. rubricoulis (Greene) S.L. Welsh [-,1] T;

8350-9400'; S; me.

Lupinus brevicoulis S. Watson [-,1 ] T; 7598'; S; pj.

Lupinus kingii S. Watson [1,1] A, T; 7200-8763'; P, S; pj, pp.

* Medicogo lupulino L. [24,22] A, C, M, R, S, T; 5800-10660'; L, P,

S; fr, me, ml, mm, mr, pj, pp, ra.

* Medicogosotivo L. [14,12] A, M, R, S,T; 5781 -10500'; G, L, S; ds,

fr, ml, mm, mr, pp, ra.

* Melilotus olbus Medik. [8,8] A, C, M, R, S, T; 6560-10500'; G, S;

ds, fr, me, mm, mr, pj, ra.

* Melilotus officinalis (L.) Pall. [19,34] A, C, M, R, S,T; 5781 -10500';

G, L, P, S; br, ds, fr, me, ml, mm, ms, mr, pg, pj, pp, ra.

Oxytropis deflexa (Pall.) DC. var. sericea Torr. & A. Gray [-,6] C, T;

8175-11000'; S; ml, mm, mr, sf.

Oxytropis lambertii Pursh var. bigelovii A. Gray [2,6] A, C, R, T;

6560-11200'; G, S; ds, me, mm, ms, pj, pp, ra.

Oxytropis sericea Nutt. var. sericea [2,7] C, M, S, T; 6600-10093'; P,

S; ds, fr, me, pj, pp.

Oxytropis splendens Douglas ex Hook. [1,2] M, T; 8700-10080';

S; mm.

Pomaria jamesii (Torr. & A. Gray) Walp. [1,-] A; 5500'; L; fr.

Prosopis glandulosa Torr. var. torreyana (L. Benson) M.C. Johnston

[1,-] A; 5800'; L; fr, pj.

Psoralidium lanceolatum (Pursh) Rydb. [-,1 ] T; 5781'; G; fr, ra.

Psoralidium tenuiflorum (Pursh) Rydb. [6,-] A; 5570-7240'; L, S;

mm, pg, pj, pp.

Robinia neomexicana A. Gray var. neomexicana [2,-] A, M; 7700-

8000'; S; mm, ms.

Senna bauhinioides (A. Gray) Irwin & Barneby [2,-] A; 5760-5840';

L; P9, Pj-

Thermopsis rhombifolia (Nutt, ex Pursh) Nutt, ex Richardson var.

divaricarpa (A. Nelson) Isely [1,-] M; 10600-10800'; S; mm.

Thermopsis rhombifolia (Nutt, ex Pursh) Nutt, ex Richardson var.

montana (Nutt.) Isley [15,23] A, C, M, R, S,T; 7200-10440'; S;

me, mm, mr, pj, pp, ra, sf.

Trifolium attenuatum Greene [4,21] M, S, T; 9300-12960'; S; am,

bw, me, mm, mr.

* Trifolium brandegeei S. Watson [-,9] T; 9300-12700'; S; am, mr,

sf.

Trifolium gymnocarpon Nutt. [-,1] T; 6600-6800'; P; ds, fr.

* Trifolium hybridum L. [ 1,3] A, C, T; 8050-9700'; S; mr.

Trifolium longipes Nutt. var. reflexum A. Nelson [-,4] T; 9200-11209';

S; mm, mr, sf.

Trifolium longipes Nutt. var. rusbyi (Greene) H.D. Harr. [UNM-W.

Moir 18] T; S.

Trifolium nanum Torr. [-,3] T; 10500-13000'; S; am.

Trifolium parryi A. Gray [-,1 ] T; 12400-12700'; S; am.

* Trifolium pratense L. [35,13] A, C, M, S,T; 6540-10700'; G, S; me,

ml, mm, mr, ra.

* Trifolium repens L. [25,33] A, C, M, R, S, T; 5781 -11300'; G, S; fr,

me, ml, mm, mr, pp, ra.

Trifolium wormskjoldii Lehm. var. arizonicum (Greene) Barneby [-,4]

T; 7550-9700'; S; me, ml, mr.

Trifolium wormskjoldii Lehm. var. wormskjoldii [-,3] T; 7050-11000';

S; mm, mr.

Viciaamericana Muhl. exWilld. [21,26] A, C, M, R, S,T; 7200-11700';

S; br, bw, me, mm, mr, ra, sf.

Vicio ludoviciana Nutt, ex Torr. & A. Gray var. ludoviciana [1,-] A;

6200'; L; pg, pj.

Viciapulchella Kunth [9,-] A, M; 7720-9760'; S; af, me, mm, ms, pp, ra.

* Vicio villosa Roth ssp. varia (Host) Corb. [UNM-C.R. Hutchins

8347] T; G.

Fagaceae

Quercus gambelii Nutt. [43,55] A, C, M, R, S,T; 6200-10500'; G, P, S;

af, br, ds, fr, me, mm, ms, mr, pj, pp, ra.

Quercus grisea Liebm. [11,3] A, R,T; 5640-8000'; L, S; pj, pp, ra.

Quercus turbinella Greene [1,-] A; 5730'; L; pg, pj.

x Quercus xunduloto Torr. [6,4] A, C, S, T; 5725-8900'; L, S; br, ms,

Pj, PP-

Fumariaceae

Corydalis aurea Willd. var. aurea [4,14] A, C, M, S, T; 7550-10600';

S; br, me, mm, mr, ra, sf.

Corydalis aurea Willd. var. occidentalis Engelm. ex A. Gray [-,7] R, T;

6600-10100'; P, S; br, ds, me, pp.

Gentianaceae

Frasera speciosa Douglas ex Griseb. [13,10] A, C, M, S, T; 7900-

12000'; S; af, bw, me, mm, sf.

Gentiana algida Pall. [1,3] M, R, T; 11990-13024'; S; am.

! Gentiana aquatica L. [-,2] T; 9600-10900'; S; mm, mr.

Gentiana bigelovii A. Gray [3,-] A, M; 9160-10000'; S; af, me, mm.

[Gentiana offinis Griseb.]

Gentiana parryi Engelm. [10,8] A, M, R, T; 9320-12000'; S; af, am,

br, mm, sf.

Gentianellaamarella (L.) Borner var. acuta (Michx.) Herder [2,16] A,

M, R,T; 7840-12000'; S; am, me, mm, mr, sf.

Gentianella amarella (L.) Borner var. heterosepala (Engelm.) Dorn

[1,-] A; 8400-8850'; S; mm, mr.

Gentianopsis thermalis (Kuntze) H.H. litis [3,1] M,T; 10200-12500';

S; mm, mr, sf.

Swertiaperennis L. [6,3] A, M, S, T; 9320-11333'; S; mm, mr.

Geraniaceae

* Erodium cicutarium (L.) L'Her. ex Aiton [12,10] A, C, M, R, S, T;

5800-9320'; L, S; br, fr, ml, mm, pj, pp, ra.

Geranium caespitosum E. James [44,24] A, C, M, R, S,T; 6540-10500';

G, S; af, fr, me, ml, mm, mr, ms, pj, pp, ra.

Geranium richordsonii Fisch. & Trautv. [45,55] A, C, M, R, S, T;

7000-11920'; S; af, me, mm, mr, pp, sf.

Grossulariaceae

Ribes aureum Pursh var. aureum [1,2] A,T; 5781-8230'; G, S; fr, mr, ra.

Ribes cereum Douglas [19,30] A, C, M, R, S,T; 6500-10500'; G, P, S;

ds, fr, me, ml, mm, ms, mr, pj, pp, ra, sf.

Ribes inerme Rydb. var. inerme [14,7] A, C, R, S, T; 7600-11290'; S;

mm, mr, ra.

Ribes leptonthum A. Gray [3,4] M, S, T; 6540-11940'; G, S; am, mr.

Ribes montigenum McClat. [13,20] A, C, M, R, S, T; 7580-12500'; S;

am, me, mm, mr, sf.

Ribes wolfii Rothrock [16,17] A, C, M, R, S, T; 8300-12300'; S; me,

mm, mr, sf.

Larson et al., Floristic studies of the Sangre de Cristo Mountains

Haloragaceae

Myriophyllum sibiricum Komarov [-,3] M,T; 9900-10800'; S; mr.

Heliotropaceae

Heliotropium curassavicum L. var. obovatum DC. [-,1 ] C; 8500'; S; ml.

Hydrangeaceae

Fendlera rupicolo A. Gray var. folcoto (Thornber) Rehder [-,1] T;

5781'; G; fr, ra.

Fendlera rupicola A. Gray var. rupicola [-,1 ] T; 7250-8000'; S; pj.

Fendlera rupicola A. Gray var. wrightii A. Gray [-,2] R, T; 6200-6700';

S; pj-

Jamesia americana Torr. & A. Gray var. americana [36,12] A, C, M, S,

T; 7350-11650'; P, S; af, me, mm, mr, ms, sf.

Philadelphus microphyllus A. Gray var. microphyllus [-,1] T; 8300-

9300'; S; mr.

Hydrocharitaceae

Elodea canadensis Michx. [1,1 ] A, T; 7800-8175'; S; ml, mr.

Hydrophyllaceae

Flydrophyllum fendleri (A. Gray) A. Heller var. fendleri [21,11] A, M,

R, S,T; 7620-12850'; S; af, bw, me, mm, mr, sf.

Nama dichotomum (Ruiz & Pavon) Choisy [-,1 ] R; 8200'; S; pp.

Phacelia alba Rydb. [1,1] A, T; 7900-9750'; S; me.

Phacelia bakeri (Brand) J.F. Macbr. [-,2] T; 11120-12960'; S; am, mr.

Phacelia heterophylla Pursh var. heterophylla [25,14] A, M, R, S, T;

7600-11650'; S; af, me, mm, mr, ra, sf.

Phacelia integrifolia Torr. var. integrifolia [1,3] R, S, T; 6100-7950';

G, S; fr, pj, ra.

Phacelia sericea (Graham) A. Gray [UNM-H. Bobisud 126] T; S.

Hypericaceae

Hypericum scouleri Hook. [2,1 ] T; 7720-9675'; S; mr.

Iridaceae

Iris missouriensis Nutt. [19,18] A, C, M, R, S, T; 7050-11300'; S; af,

me, ml, mm, mr, ra, sf.

Sisyrinchium demissum Greene [1,2] C, R, T; 6500-7660'; G, S; ml.

Sisyrinchium montanum Greene var. montanum [2,1] A, T; 8400-

11000'; S; mm, mr.

Juncaceae

Juncus arcticus\N\\\d. var. balticus (Willd.)Trautv. [10,32] A, C, M, R,

S, T; 5781 -12000'; G, P, S; fr, me, ml, mm, mr, ra, sf.

Juncus arcticus Willd. var. mexicanus (Willd. ex Roem. & Schult.)

Balslev [3,-] S; 6840-8760'; S; mr.

Juncus bufonius L. [-,1] C; 8960-9040'; S; ra.

Juncus castaneus Sm. [UNM-J. McGrath 423] C; S.

Juncus confusus Cov. [-,1 ] T; 8100'; S; mr.

Juncus drummondii E. Mey. [6,10] A, M, R, S, T; 9800-12960'; S;

am, mr, sf.

Juncus duof/ey/Wiegand [4,1] A, M,T; 7700-10500'; S; me, mm, mr.

Juncus ensifolius Wikstr. var. montanus (Engelm.) C.L. Hitchc. [17,19]

A, C, M, R, S, T; 6840-12000'; S; me, mm, mr, sf.

Juncus hallii Engelm. [1,-] M; 11940'; S; am.

Juncus interior Wiegand [4,3] A, C, M,T; 5750-8625'; G, L, S; fr, ml, ra.

Juncus longistylis Torr. [2,8] C, M, R, T; 6380-11209'; G, S; fr, ml,

mm, mr, ra, sf.

Juncus mertensianus Bong. [2,-] M; 10800-12000'; S; ml, mm.

Juncus torreyi Coville [UNM-R. Sivinski et al. 2434] A; S.

Luzulaparviflora (Ehrh.) Desv. [17,28] A, M, R, S,T; 7620-12960'; S;

am, me, ml, mr, sf.

Luzula spicata (L.) DC. [3,13] M, R, S, T; 9800-13024'; S; am, bw,

mm, mr.

Juncaginaceae

Triglochin palustris L. [UNM-R. Worthington 32634] T; S.

293

Lamiaceae

Dracocephalum parviflorum Nutt. [21,11 ] A, C, M, R, S,T; 7200-9400';

S; af, br, me, ml, mm, mr, ra.

Hedeoma drummondii Benth. [1,4] A, T; 6980-7550'; P, S; ds, fr,

pj, PP, ra.

* Marrubium vulgare L. [7,4] A, C, M, S, T; 5800-7840'; L, P, S; ds,

ml, pj, ra.

Mentha arvensis L. [4,6] A, C, M,T; 7240-9700'; S; mm, mr, ra.

Monarda fistulosa L. var. menthifolia (Graham) Fernald [7,8] A, S, T;

7400-11115'; S; me, mm, mr, pp.

Monardapectinata Nutt. [2,-] A; 7000-7500'; S; pj, pp.

* Nepeta cataria L. [UNM-R. Sivinski & B. Simpson 2391 ] S; S.

Prunella vulgaris L. var. lanceolata (W.P.C. Barton) Fernald [21,15] A,

C, M, S,T; 7550-11209'; S; me, ml, mm, mr, ra, sf.

Salvia reflexa Hornem. [1,1] M,T; 7350-7700'; S; fr, mr, ra.

Satureja vulgaris (L.) Fritsch [5,1] A, S, T; 7600-8900'; S; me, mm,

mr, ra.

Stachys pilosa Nutt. var. pilosa [-,2] C, T; 8500-9800'; S; mm, mr.

Teucrium laciniatum Torr. [5,-] A; 5650-6200'; L; pg, pj, ra.

Lemnaceae

Lemna minor L. [1,1] A, T; 7550-7880'; S; ml, mr.

Liliaceae (see also Alliaceae, Asparagaceae, Convallariaceae,

Melanthiaceae)

CalochortusgunnisoniiS. Watson var. gunnisonii [-,5]T; 8350-11000';

P, S; me, mm, mr.

+♦ Calochortus gunnisonii S. Watson var. perpulcher Cockerell [4,-]

A, M; 9840-10700'; S; mm, mr, sf.

Calochortus nuttallii Torr. & A. Gray [-,1 ] R; 6050-6350'; G; ds.

Lilium philadelphicum L. [2,-] A; 8325-8900'; S; me, mr.

Lloydia serotina (L.) Rchb. var. serotina [-,3] T; 11200-12500'; S;

am, mr.

Streptopus amplexifolius (L.) DC. [9,14] A, M, R, S, T; 7840-11960';

S; me, mm, mr, sf.

Linaceae

Linum australe A. Heller var. australe [3,3] A, S, T; 7000-9400'; S;

mr, pj, ra.

Linum lewisii Pursh var. lewisii [9,7] A, C, M, T; 5730-10660'; L, S; fr,

me, mm, mr, pg, pj, ra.

Linum pratense (Norton) Small [2,-] A; 5650-5800'; L; pg, pj.

Linumpuberulum (Engelm.) A. Heller [3,-] A; 5650-6750'; L, S; pg, pj.

Loasaceae

Mentzelia laciniata (Rydb.) J. Dari. [1,1] A, T; 5650-7100'; L, S; pg,

mr, ra.

Mentzelia multiflora (Nutt.) A. Gray var. multiflora [9,5] A, R, S, T;

5500-8300'; G, L, P, S; ds, fr, pj, ra.

Mentzelia nuda (Pursh) Torr. & A. Gray var. stricta (Osterh.) Harrington

[-,2] M; 9160-9320'; S; ra.

Mentzelia rusbyi\Nooton [1,-] M; 7700'; S; ms, mr.

Malvaceae

* lliamna grandiflora (Rydb.) Wiggins [-,3] T; 7550-10100'; S; br.

* Malva neglecta Wallr. [4,1 ] A, M,T; 7200-9360'; S; mr, ra.

* Malva parviflora L. [UNM-W. Adair s.n., 21 Sep 1907] T; G.

Sidalcea Candida A. Gray var. Candida [26,17] A, C, M, S, T; 7400-

10800'; S; mm, mr.

Sidalcea neomexicana A. Gray var. neomexicana [-,2] T; 7050-9400';

S; ml, mr.

Sphaeralcea angustifolia (Cav.) G. Don [2,-] A; 5650-7440'; L, S; ra.

Sphaeralcea coccinea (Nutt.) Rydb. var. coccinea [11,7] A, S, T;

5600-8100'; G, L, S; br, ds, fr, pg, pj, ra.

Sphaeralcea fendleri A. Gray var. fendleri [3,3] A, C, M, S, T; 7000-

7800'; S; mr, pj, ra.

294

Journal of the Botanical Research Institute of Texas 8(1)

Sphoerolceo incono Torr. ex A. Gray var. cuneoto Kearney [5,-] A, S;

5610-7900'; L, S; fr, mr, ra.

Sphaeralcea incana Torr. ex A. Gray var. incana [1,-] S; 6840-6880';

S; fr, ra.

Melanthiaceae [traditionally in Liliaceae]

Veratrum califomicum T. Durand var. californicum [9,15] A, C, M, R,

S, T; 8300-12100'; S; me, mm, mr, sf.

Zigodenus elegons Pursh [27,29] A, C, M, S, T; 7050-13161'; S; af,

am, bw, me, mm, mr, sf.

[Anticlea elegans (Pursh) Rydb.]

Myrsinaceae

Lysimachia ciliata L. [UNM-J. McGrath 730] A; S.

Nyctaginaceae

Mirobilisolbido (Walter) Heimerl [-,3] M,T; 8300-10300'; S; me, ra.

Mirabilis linearis (Pursh) Heimerl var. decipiens (Standi.) S.L. Welsh

[1,6] R, S,T; 7350-8320'; S; br, mr, pj, ra.

Mirabilis linearis (Pursh) Heimerl var. linearis [9,2] A, R, S, T; 5600-

8575'; G, L, P, S; ds, fr, mr, pj, pp, ra.

Mirabilis melanotricha (Standi.) Spellenb. [14,-] A, M, S; 7600-9760';

S; af, me, mm, mr, ra.

Mirabilis multiflora (Torr.) A. Gray var. multiflora [2,1] A, T; 5650-

8000'; L, S; fr, pg, pj.

Mirabilis oxybaphoides (A. Gray) A. Gray [UNM-E. Castetter 4034]

T; G.

Oleaceae

Forestierapubescens Nutt. [-,3] R, T; 5781 -6540'; G, P; ds, fr, ra.

Menodora scabra A. Gray [4,3] A, R, S,T; 5570-7950'; L, S; fr, pg, pj.

!* Syringa vulgaris L. [2,-] M, S; 7320-7700'; S; mr.

Onagraceae

Chamerion angustifolium (L.) Holub var. canescens (A.W. Wood) N.

Holmgren & P. Holmgren [19,21] A, C, M, R, S,T; 7750-13024';

S; af, am, br, me, mm, mr, sf.

[Chamerion angustifolium ssp. circumvagum (Mosquin) Hoch ]

Epilobium anagallidifolium Lam. [2,7] A, S, T; 9400-11900'; S; me,

mm, mr.

[Epilobium alpinum L.]

Epilobium ciliatum Raf. var. ciliatum [20,11] A, C, M, S, T; 6840-

11209'; S; ml, mm, mr, ra, sf.

Epilobium ciliatum Raf. var. glandulosum (Lehm.) Dorn [1,5] C, S,T;

7800-11500'; S; me, mr.

Epilobium halleanum Haussk. [7,7] A, M, R, S, T; 7840-11940'; S;

ml, mr, sf.

Epilobium hornemannii Rchb. var. hornemannii [2,6] M, T; 8790-

12000'; S; ml, mr, sf.

Epilobium lactiflorum Haussk. [1,-] S; 8900'; S; me, mr.

Epilobium saximontanum Haussk. [4,7] A, M, S, T; 8350-10986'; S;

af, me, mm, mr, sf.

Gayophytum diffusum Torr. & A. Gray var. strictipes (Hook.) Dorn [-,2]

C, T; 9090-10600'; P, S; me, mm.

Gayophytum ramosissimum Torr. & A. Gray [-,1 ] T; 9200-9750'; S; me.

Oenothera albicaulis Pursh [4,1] A, S, T; 5570-7460'; L, P, S; ds, fr,

P9, Pj-

* Oenothera biennis L. [5,2] A, C, T; 8000-8900'; S; mr, ra.

Oenothera cespitosa Nutt. var. macroglottis (Rydb.) Cronquist [-,13]

M, R, T; 6036-9320'; G, P S; ds, fr, mr, pj, ra.

Oenothera cespitosa Nutt. var. marginata (Nutt, ex Hook. & Arn.)

Munz [2,1 ] R, S, T; 6540-8550'; G, S; fr, pj.

Oenothera coronopifolia Torr. & A. Gray [5,7] A, C, S, T; 7200-9750';

P, S; ds, fr, ml, mm, mr, pj, pp, ra.

Oenothera curtifolia W.L. Wagner & Hoch [-,1 ] T; 7240'; S; ra.

Oenothera elata Kunth var. hirsutissima (A. Gray ex S. Watson) Cron¬

quist [11,2] A, S,T; 7000-8840'; S; pp, me, mr, ra.

Oenothera flava (A. Nelson) Garrett [-,1 ] T; 9950-11000'; S; mr.

Oenothera hartwegii (Benth.) ssp. fendleri (A. Gray) W.L. Wagner &

Hoch. [5,-] A; 5800-7500'; L, S; pg, pj, ra.

Oenothera laciniata Hill [3,-] A; 5700-8840'; L, S; fr, mr, ra

Oenothera pallida Lindl. var. latifolia Rydb. [-,1 ] T; 7350-7450'; S; fr, ra

Oenothera pallida Lindl. var. runcinata (Engelm.) Cronquist [1,-]

A; 5840'; L; pg, pj.

Oenothera serruiata Nutt. [1,-] A; 6200'; L; pg, pj.

Oenothera suffrutescens (Seringe) W.L. Wagner & Hoch [17,6] A, R,

S,T; 5650-7660'; G, L, S; fr, pg, pj, ml, ra.

Oenothera villosa Thunb. var. strigosa (Rydb.) Dorn [5,-] A, S;

7200-8400'; S; mr, pp, ra

Orchidaceae

Calypso bulbosa (L.) Oakes var. americana (R. Br.) Luer [-,4] C, T;

8555-10600'; S; me.

Coeloglossum viride (L.) Hartm. var. virescens (Muhl. ex Willd.) Luer

[1,-] S; 9860-10320'; S; ml, mr.

Corallorhiza maculata (Raf.) Raf. var. maculata [-,8] M, T; 7850-

10700'; S; me, sf.

Corallorhiza maculata (Raf.) Raf. var. occidentalis (Lindl.) Ames [12,9]

A, M, S, T; 7900-10500'; S; af, me, mm, mr, sf.

Corallorhiza striata Lindl. var. striata [2,-] A, S; 7640-9190'; S; mr.

Corallorhiza striata Lindl. var. vreelandii (Rydb.) L.O. Williams [-,2]

T; 7620-8900'; S; mr.

Corallorhiza trifida Chatelain [UNM-R. Jackson 2203] T; S.

Corallorhiza wisteriana Conrad [-,3] R,T; 8000-8800'; S; me, mr, pp.

♦ Cypripedium parviflorum Salisb. var. pubescens (Willd.) O.W.

Knight [1,-] A; 8630-9190'; S; mr.

* Epipactis helleborine L. [1,-] S; 7400-7760'; S; mr.

Goodyera oblongifolia Raf. [11,17] A, C, M, R, S, T; 7800-11500'; S;

af, me, mr, pp, sf.

Goodyera repens (L.) R. Br. ex W.T. Aiton [2,2] A, C, S, T; 8450-9600';

S; me, sf.

Listera cordata (L.) R. Br. var. nephrophylla (Rydb.) Hulten [2,7] M, R,

S, T; 9150-11940'; S; mr, sf.

Malaxis soulei L.O. Williams [1,-] A; 8400-8850'; S; me.

Platanthera aquilonis Sheviak [-,3] T; 9250-11000'; S; me, mr, sf.

Platanthera huronensis (Nutt.) Lindl. [1,13] C, M, S,T; 8300-12000';

S; me, mr, sf.

Platanthera purpurascens (Rydb.) Sheviak &W.F. Jenn. [15,10] A, M,

S,T; 7620-12500'; S; me, ml, mr, sf.

Platanthera sparsiflora (S. Watson) Schltr. [2,-] A; 8325-10180'; S;

me, mr.

Spiranthes romanzoffiana Cham. [UNM-H. Mackay 6T-171 ] T; S.

Orobanchaceae (previously Scrophulariaceae)

Castilleja haydenii {A. Gray) Cockerell [1,9] M,T; 11500-12850'; S; am.

Castilleja integra A. Gray [19,23] A, C, R, S, T; 5630-10200'; G, L, P,

S; br, ds, fr, me, ml, mm, mr, pg, pj, pp, ra.

Castilleja linariifolia Benth. [8,3] A, M, S, T; 7100-9160'; P, S; me,

mr, ms, ra.

Castilleja lineata Greene [-,3] T; 7350-10500'; S; mm, mr, ra.

Castilleja miniata Douglas ex Hook. var. miniata [24,45] A, C, M, R,

S,T; 7850-12430'; S; af, bw, me, mm, mr, pp, sf.

Castilleja exilis A. Nelson [1,-] R; 6500-6620'; G; ml.

[Castilleja minor (A. Gray) A. Gray var. exilis (A. Nelson) J.M. Egger]

Castilleja occidentalis Torr. [2,14] M, R, S, T; 9800-13024'; S; am,

bw, mm, mr, sf.

Castilleja rhexifolia Rydb. [UNM-M. Olsen 204] S; S.

Castilleja sulphurea Rydb. [10,9] A, M, T; 8325-12000'; S; me, ml,

mm, mr.

Conopholis alpina Liebm. var. mexicana (A. Gray ex S. Watson)

Haynes [5,-] A, M, S; 5700-9350'; L, S; fr, mr, pj, pp.

Cordylanthus wrightiiA. Gray ssp. wrightii [-,1] R; 7600-7750'; S; ra.

Larson et al., Floristic studies of the Sangre de Cristo Mountains

Orobanche fasciculata Nutt. [1,5] C, M, T; 6600-9000'; P, S; ds, fr,

me, pj, pp, ra.

Orobanche ludoviciana Nutt. var. multiflora (Nutt.) Beck [2,1 ] A, T;

5610-7450'; L,S;fr,pj, ra.

Orthocarpus luteus Nutt. [3,10] A, C, M, R, T; 8350-11000'; P, S; af,

me, mm, mr, ms, pp.

Orthocarpuspurpureoalbus A. Gray ex S. Watson [-,1 ] T; 7200-7500';

S; pj-

Pedicularis bracteosa Benth. var. paysoniana (Pennell) Cronquist

[-,1 ]T; 10150-11900'; S; mr.

Pedicularis canadensis L. var. fluviatilis (A. Heller) Macbr. [7,11 ] A, C,

M, S, T; 7620-10200'; S; af, me, mm, mr.

Pedicularis centranthera A. Gray [-,14] R,T; 6232-8900'; G, S; fr, pj, pp.

Pedicularis groenlandica Retz. [9,16] A, M, S,T; 9300-12960'; S; am,

ml, mm, mr, sf.

Pedicularis parryi A. Gray [6,7] A, M, S, T; 11310-12850'; S; am,

mm, mr, sf.

Pedicularis procera A. Gray [22,14] A, M, S, T; 7840-12115'; S; af,

me, mm, mr, ra, sf.

Pedicularis racemosa Douglas ex Benth. var. alba (Pennell) Cronquist

[10,20] A, M, R, S, T; 9300-12000'; S; am, bw, me, mr, sf.

Pedicularis racemosa Douglas ex Benth. var. racemosa [-,1] T;

10150-11900'; S;mr.

Oxalidaceae

Oxalis corniculata L. var. wrightii (A. Gray) B.L. Turner [4,-] A, S;

7000-8300'; S; mr, ra.

Oxalis metcalfei Knuth [6,2] A, C, M, T; 7840-10500'; S; mm, mr.

Papaveraceae

Argemone hispida A. Gray [-,2] T; 7400-8100'; S; br, pj.

Parnassiaceae (previously Saxifragaceae)

♦ Parnassia fimbriata Konig [-,4] T; 8450-11500'; S; mr.

Parnassia palustris L. var. montanensis (Fernald & Rydb. ex Rydb.)

C.L. Hitchc. [UNM-J. McGrath 420] C; S.

Plantaginaceae (includes Callitrichaceae, Scrophulariaceae, in

part)

Callitriche heterophylla Pursh var. heterophylla [UNM-R. Sivinski

5611 ] C; S.

Callitriche palustris L. [UNM-J. McGrath 936] R; S.

*• Linaria dalmatica (L.) Mill. var. dalmatica [1,-] S; 7380-7420'; S;

pj, ra.

*•+ Linaria vulgaris Mill. [-,3] T; 7050-10900'; S; mm, mr.

Mimulus glabratus Kunth var. jamesii (Torr. & A. Gray ex Benth.) A.

Gray [-,2] T; 8350-9400'; S; me, mr.

Mimulus guttatus DC. [22,24] A, M, S, T; 7850-11800'; S; ml, mr, sf.

Mimulus rubellus A. Gray [-,1 ] T; 7600-8450'; S; fr, pp.

Mimulus tilingii Regel [4,-] A, M, S; 8200-12000'; S; am, ml, mr.

Penstemon barbatus (Cav.) Roth var. torreyi (Benth.) A. Gray [33,25]

A, C, M, R, S, T; 5500-10500'; G, L, P, S; af, br, fr, me, mm, mr,

ms, pj, pp, ra.

* Penstemon cobaea Nutt. [UNM-C.R.. Hutchins 11490] T; G.

Penstemon crandalliiA. Nelson var. glabrescens (Pennell) G.T. Nisbet

& R.C. Jacks. [-,17] R,T; 7350-9400'; G, S; br, fr, me, pj, pp, ra.

+ Penstemon crandallii A. Nelson var. taosensis (Keck) G.T. Nisbet

& R.C. Jacks. [-,3] T; 7440-8900'; P, S; pj, pp.

Penstemon fendleri Torr. & A. Gray [1,-] A; 6200'; L; pg, pj.

! Penstemon glaber Pursh var. alpinus (Torr.) A. Gray [-,1] T;

10000-12200'; S; bw, mm.

Penstemon griffinii A. Nelson [2,-] S; 7300-8000'; S; pj.

+ Penstemon inflatus Crosswhite [1,6] S,T; 7550-10500'; S; br, me,

mr, pj, pp.

Penstemon jamesii Benth. [10,-] A, S; 5600-7460'; L, S; pg, pj, ra.

Penstemon linarioides A. Gray ssp. coloradoensis (A. Nelson) D.D.

Keck [1,-] S; 8300'; S; mr.

295

Penstemon palmeri A. Gray [UNM-J. McGrath 602] T; G.

Penstemon rydbergii A. Nelson var. rydbergii [-,2] T; 9850-10000';

S; mm.

Penstemon secundiflorus Benth. [3,6] A, R, S, T; 6380-9400'; G, P,

S; fr, me, mr, pj.

Penstemon strictus Benth. [-,7] T; 8350-10000'; S; me, mm, mr.

Penstemon unilateral Rydb. [-,2] C,T; 7350-7450'; S; br, fr, ra

[Penstemon virgatus ssp. asa-grayi Crosswhite]

Penstemon virgatus A. Gray [3,3] A, R, S, T; 6540-8175'; G, S; fr,

pj, PP, ra.

Penstemon whippleanusA. Gray [9,24] A, M, R, S,T; 8650-13024'; S;

am, bw, me, mm, mr, sf.

Plantago argyraea Morris [-,1] R; 8200'; S; pp.

* Plantago lanceolata L. [8,5] A, C, M, R, S, T; 5500-9100'; L, P, S;

fr, mm, mr, ra.

* Plantago major L. [22,10] A, C, M, R, S, T; 5500-10700'; L, S; af,

fr, me, ml, mm, mr, ra.

Plantago patagonica Jacq. [6,4] A, R, S, T; 5700-8175'; G, L, P, S;

ds, fr, ml, pg, pj, ra.

Plantago tweedyi A. Gray [2,-] M, S; 11600-11850'; S; am.

* Synthyris alpina A. Gray [-,6] T; 11500-13161'; S; am.

Synthyris plantaginea (E. James) Benth. [8,10] A, C, M, S, T; 7000-

13009'; S; am, br, me, mm, mr, pp.

Veronica americana Schwein. ex Benth. [9,7] A, S, T; 7850-11209';

S; ml, mr, sf.

* Veronica anagallis-aquatica L. [2,2] S,T; 5781 -7050'; G, S; fr, ml,

mr.

Veronicaperegrina L. var. xalapensis (Kunth.) St. John & F.W. Warren

[5,2] A, S, T; 5781-10440'; G, S; fr, mr, ra.

* Veronica serpyllifolia L. var. humifusa (Dickson) Vahl [7,8] A, M,

S,T; 7775-12000'; S; af, me, mr, sf.

Veronica wormskjoldii Roem. & Schult. [3,16] A, M, S,T; 9150-12960';

S; am, mm, mr, sf.

Poaceae

Achnatherum hymenoides (Roem. & Schult.) Barkworth [5,26] A, R,

S, T; 5781 -8900'; G, L, P, S; br, ds, fr, pg, pj, pp, ra.

Achnatherum lettermanii (Vasey) Barkworth [-,11 ] R,T; 7600-10887';

S; me, mm, mr, ra, sf.

Achnatherum lobatum (Swallen) Barkworth [-,1 ]T; 7600-8300'; S; pj.

! Achnatherum nelsonii (Seribn.) Barkworth var. nelsonii [-,5] A,T;

7550-10150'; S; br, me, mm, mr.

Achnatherum perplexum Hoge & Barkworth [1,-] A; 7720; S; sf.

Achnatherum robustum (Vasey) Barkworth [15,11] A, C, M, S, T;

7000-10300'; P, S; fr, me, ml, mm, mr, pj, pp, ra.

Achnatherum scribneri (Vasey) Barkworth [2,1] A, T; 7400-8000';

P, S; ds, me, ra.

*• Aegilops cylindrica Host [3,3] A, S,T; 5781-7230'; G, L, P, S; fr, pj,

ra.

* Agropyron cristatum (L.) Gaertn. var. cristatum [1,8] A, R, T;

5781 -9400'; G, S; br, ds, fr, ml, mr, pj, ra.

[Agropyron cristatum]

* Agropyron cristatum (L.) Gaertn. var. desertorum (Fisch. ex Link)

Dorn [3,6] A, R, S, T; 6540-8760'; G, P, S; ds, fr, ms, pj, ra.

[Agropyron cristatum]

Agrostis exarata Jr\r\. var. minor Hook. [9,2] A, M, S,T; 7000-11000';

S; mm, mr, sf.

* Agrostis gigantea Roth [19,7] A, C, M, S, T; 6840-10300'; S; af,

me, ml, mm, mr, ra.

Agrostis idahoensis Nash [UNM-H. Mackay 6T-136] T; S.

Agrostisscabra Willd. [28,37] A, C, M, R, S,T; 7000-13024'; S; br, bw,

me, mm, mr, ms, pp, sf.

* Agrostis stolonifera L. [5,5] A, C, M, R, S, T; 6500-10700'; G, S; me,

ml, mm, mr, ra.

Agrostis variabilis Rydb. [1,-] A; 11240-11340'; S; mm, ra.

296

Journal of the Botanical Research Institute of Texas 8(1)

Alopecurus oequolis Sobol, var. oequolis [2,2] A, T; 7050-11209';

S; ml, mr, sf.

* Alopecuruspratensis L. [-,1 ] T; 7050'; S; mr.

Andropogon gerordii Vitman ssp. hallii (Hack.) Wipff [1,-] A; 7000';

S; ms, pp.

Aristida adscensionis L. [1,-] A; 5630'; L; fr.

Aristida divaricata Humb. & Bonpl. exWilld. [UNM-K. Weissenborn

14] A; S.

Aristida havardii Vasey [1,-] A; 5760'; L; pg.

Aristida purpurea Nutt. var. fendleriana (Steud.) Vasey [2,10] A, R, T;

6120-8350'; P, S; ds, fr, mr, pj, pp.

Aristida purpurea Nutt. var. longiseta (Steud.) Vasey [11,12] A, R, S,

T; 5650-7598'; G, L, P, S; ds, fr, pg, pj, ra.

Aristida purpurea Nutt. var. nealleyi (Vasey) Allred [2,-] A; 5600-

5800'; L; pj.

Aristida purpurea Nutt. var. wrightii (Nash) Allred [1,-] A; 5800';

L; pj, ra.

* Arrhenatherum elatius (L.) P. Beauv. ex J. Presl & C. Presl [UNM-H.

Mackay 5T-119]T;S.

* Avena fatua L. [1,-] A; 5800'; L; fr, ra.

* Avena sativa L. [1 ,-] A; 5800'; L; fr, ra.

Beckmanniasyzigachne (Steud.) Fernald ssp. baicalensis (N.l. Kusne-

zow) T. Koyana & Kawano [UNM-N. & P. Holmgren 7245] T; S.

Blepharoneuron tricholepis (Torr.) Nash [18,20] A, C, M, R, S, T;

7750-12700'; S; af, am, br, me, mm, ms, mr, pj, pp, sf.

* Bothriochloaischaemum (L.) Keng var. ischaemum [ 1 ,-] A; 5500';

L; fr, pj-

Bothriochloa laguroides (DC.) Herter ssp. torreyana (Steud.) Allred

& Gould [4,-] A; 5610-5760'; L; fr, pg, pj.

Boutelouacurtipendula (Michx.)Torr. var. caespitosa Gould & Kapadia

[3,2] A, S,T; 5700-7550'; L, P, S; fr, pj, ra.

Bouteloua curtipendula (Michx.) Torr. var. curtipendula [7,7] A, R, S,

T; 5500-8763'; L, P, S; br, ds, fr, ms, pg, pj, pp, ra.

Bouteloua dactyloides (Nutt.) J.T. Columbus [-,1 ] T; 7240'; S; ra.

[Buchloe dactyloides (Nutt.) Engelm.]

Bouteloua gracilis (Willd. ex Kunth) Lag.ex Griffiths [13,18] A, C, M, R,

S,T; 5800-10115'; L, P, S; br, ds, fr, me, mm, mr, ms, pg, pj, pp, ra.

Bouteloua hirsuta Lag. [4,-] A; 5700-7000'; L, S; pg, pj, pp.

Boutelouasimplexlag. [-,2]T; 7200-8500'; S; mr, pj.

Bromus carinatus Hook. & Arn. [10,3] A, M, S,T; 6200-11500'; L, S;

me, mm, mr, ra.

* Bromus catharticus Vahl [3,1 ] A,T; 5500-8450'; L, S; fr, mr, ra.

Bromus ciliatus L. [33,10] A, M, R, S, T; 7380-12050'; S; af, br, me,

mm, mr, pp, sf.

* Bromus inermis Leyss. [29,39] A, C, M, R, S, T; 5800-11500'; G, L,

P, S; af, br, fr, me, ml, mm, mr, ms, pj, pp, ra, sf.

* Bromus japonicus Thunb. ex Murray [11,9] A, C, R, S, T; 5700-

9750'; G, L, P, S; br, fr, me, ml, mm, pj, pp, ra.

Bromus lanatipes (Shear) Rydb. [2,8] A, M,T; 5650-10700'; L, P, S;

fr, me, mm, mr, ms, pj, pp, ra.

Bromus ported (J.M. Coult.) Nash [5,14] A, M, S, T; 8160-10700'; S;

bw, me, mm, mr.

[Bromus anomalus Rupr. ex E. Fourn.]

Bromus richardsonii Link [-,26] C, M,T; 7400-12960'; S; af, me, mm,

ms, mr, ra, sf.

[Bromus ciliatus L.]

* Bromus tectorum L. [9,41 ] A, C, M, R, S, T; 5800-9750'; G, L, P, S;

br, ds, fr, me, ml, mm, ms, mr, pj, pp, ra, sf.

Calamagrostiscanadensis (Michx.) P. Beauv. var. canadensis [10,18]

A, C, M, R, S, T; 7850-11940'; S; am, me, mm, mr, sf.

Calamagrostis purpurascens R. Br. var. purpurascens [-, 6 ] T; 11800-

12850'; S; am, sf.

Calamagrostis stricta (Timm) Koeler ssp. inexpansa (A. Gray) C.W.

Greene [-,1] T; 8700-9700'; S; mm.

Catabrosa aquatica (L.) P. Beauv. var. aquatica [UNM-J. McGrath

434] C; S.

Cenchrus longispinus (Hack.) Fernald [2,-] A; 5500-5610'; L;fr, ra.

Chloris verticillata Nutt. [2,-] A; 5500-5800'; L; fr, ra.

* Chloris virgata Sw. [1,-] A; 8900-9500'; S; me.

Cinna latifolia (Trev. ex Goepp.) Griseb. [4,2] A, C, S,T; 7850-10500';

S; mr.

* Dactylis glomerata L. [30,37] A, C, M, R, S, T; 5800-10900'; G, L,

P, S; fr, me, ml, mm, mr, pj, pp, ra, sf.

Danthonia intermedia Vasey [5,12] A, M, R, T; 9270-12548'; S; am,

mm, mr, sf.

Danthoniaparryi Scribn. [14,19] A, C, M, R, S,T; 7650-12000'; S; am,

br, bw, me, mm, pp, ra.

Danthonia spicata (L.) P. Beauv. ex Roem. & Schult. [9,3] A, C, R, S,

T; 7720-9600'; S; me, mr, pp, ra.

Deschampsia cespitosa (L.) P. Beauv. var. cespitosa [15,41] A, M, R,

S,T; 7050-13024'; S; am, bw, me, ml, mm, mr, sf.

Dichanthelium oligosanthes (Schult.) Gould var. scribnerianum

(Nash) Gould [1,-] A; 7720'; S; pp.

Distichlis spicata (L.) Greene var. stricta (Torr.) Scribn. [-,2] C, T;

5781-8500'; G, S;fr,ml, ra.

* Echinochloa crus-galli (L.) P. Beauv. [2,-] A; 5500-561 O'; L; fr, ra.

* Echinochloa muricata (P. Beauv.) Fernald var. microstachya

Wiegand [1,-] S; 7380-7420'; S; fr, ra.

Elymus baked (E.E. Nelson) A. Love [4,5] A, M, R, T; 10350-13024';

S; am, mm, mr, ra, sf.

Elymus canadensis L. var. canadensis [5,3] A, C, M, S,T; 5500-9320';

L, S; fr, mr, pj, pp, ra.

* Elymus elongatus (Host) Runem. var. elongatus [ 1 ,-] M; 8040-

8700'; S; me, ra.

Elymus elymoides (Raf.) Swezey var. brevifolius (J.G. Sm.) Dorn [39,68]

A, C, M, R, S, T; 5730-10850'; G, L, P, S; br, ds, fr, me, ml, mm,

mr, pg, pj, pp, ra, sf.

[Elymus longifolius (Smith) Gould]

Elymus glaucus Buckley var. glaucus [7,8] A, C, M, S,T; 7840-10500';

S; af, me, mm, mr.

[Elymus hispidus ssp. barbulatus (Schur) Melderis]

* Elymus hispidus (Opiz) Melderis var. hispidus [7,7] A, C, M, R, S,

T; 7500-10580'; S; br, me, mr, ra.

* Elymus hispidus (Opiz) Melderis var. ruthenicus (Griseb.) Dorn

[1,4] M, R,T; 7240-8880'; S; me, pp, ra.

* Elymus junceus Fisch. [4,-] A, S; 7000-7460'; S; pj, ra.

[Psathyrostachys juncea (Fisch.) Nevski]

x Elymus xpseudorepens (Scribn. & J.G. Sm.) Barkworth & Dewey

[6,-] A, M, S; 7900-12100'; S; me, mm, ra.

* Elymus repens (L.) Gould [2,-] A; 7900-8000'; S; me, mr, ra.

Elymus scribneri (Vasey) M.E. Jones [3,9] M, S,T; 10990-12900'; S; am.

Elymus smithii (Rydb.) Gould [11,13] A, C, M, R, S,T; 5570-9675'; G,

L, P, S; br, ds, fr, me, mm, mr, ms, pj, pp, ra

Elymus trachycaulus (Link) Gould exShinners ssp. subsecundus (Link)

A. Love & D. Love [-,4] M, T; 8000-10700'; S; af, me, mm, ms.

Elymus trachycaulus (Link) Gould ex Shinners ssp. trachycaulus

[24,18] A, C, M, S,T; 6500-11700'; P, S; fr, me, mm, mr, pj, pp, ra.

Elymus trachycaulus (Link) Gould ex Shinners ssp. violaceus (Hor-

nem.) A. Love & D. Love [5,15] A, M, T; 8300-12850'; S; am,

me, mm, mr, ra, sf.

Eragrostis curtipedicellata Buckley [1,-] A; 5560-5840'; L; pj.

* Eragrostis curvula (Schrad.) Nees var. curvula [4,-] A; 5570-7840';

L, S; fr, pj, ra.

Eragrostis intermedia Hitchc. [2,-] A; 5700-5840'; L; fr, pj.

Erioneuron pilosum (Buckley) Nash [2,-] A; 6120-6750'; S; fr, pj.

Festuca arizonica Vasey [5,10] A, C, M, T; 7650-10700'; S; af, br,

me, mm, pp, ra.

Larson et al., Floristic studies of the Sangre de Cristo Mountains

* Festuca arundinacea Schreb. [5,6] A, C, M, R, S, T; 5781 -9100';

G, P, S; fr, mm, mr, ra.

Festuca brachyphylla Schult. ex Schult. & Schult. f. ssp. coloradensis

Fred. [-,13] C,T; 7850-12960'; S; am, mr, pp.

Festuca calligera (Piper) Rydb. [-,1 ] T; 10360-10440'; S; ra.

Festuca earlei Rydb. [-,5] R, T; 9800-12050'; S; mm, mr, sf.

Festuca idahoensis Elmer [3,10] A, T; 7215-11000'; S; bw, mm, mr,

pj, PP, ra.

Festuca minutiflora Rydb. [4,-] A, M, S; 11240-12430'; S; am, mm.

* Festucapratensis Huds. [5,2] A, S, T; 5800-9360'; L, S; fr, mr, ra.

Festuca rubra L. ssp. rubra [-,3] M, T; 9900-10800'; S; sf.

Festuca saximontana Rydb. var. saximontana [1,11] A, M, R, T;

8160-13024'; S; am, me, mm, sf.

Festuca sororia Piper [1,2] A,T; 7850-12115'; S; me, mr, sf.

Festuca thurberi Vasey [9,22] A, M, T; 8450-12960'; S; am, bw, me,

mm, mr, sf.

* Festuca trachyphylla (Hack.) Krajina [-,2] C,T; 7650-9700'; S; me,

mm.

Glyceria elata (Nash ex Rydb.) M.E. Jones [-,1 ] C; 8500-8750'; S; mr.

Glyceria grandis S. Watson [1,2] M, T; 8300-9300'; S; mr.

Glyceria striata (Lam.) Hitchc. [18,16] A, C, M, S,T; 7810-10500'; S;

af, me, ml, mr, sf.

Flelictotrichon hooked (Scribn.) Henrard [-,1 ] T; 9250-9500'; S; mm.

Hesperostipacomata (Trin. & Rupr.) Barkworth var. comata [3,12] R,

S, T; 5781 -9400'; G, P, S; br, ds, fr, me, mm, mr, pj, pp, ra.

Flesperostipa comata (Trin & Rupr.) Barkworth var. intermedia (Scribn.

& Tweedy) Dorn [-,2] T; 7850-9750'; S; mm, pp.

Flesperostipa neomexicana (Thurb. ex J.M. Coult.) Barkworth [4,-] A;

5700-6750'; L, S; fr, pg, pj.

Flierochloeodorata (L.) P. Beauv. [-,1] C; 8000-8104'; S; mm.

Flilaria jamesii (Torr.) Benth. [12,7] A, R, S, T; 5600-8000'; G, L, P,

S; br, ds, fr, pg, pj.

[Pleuraphis jamesii Torr.]

Flopia obtusa (Kuntz) Zuloaga & Morrone [2,-] A; 5570-5840'; L;

fr, pg.

Flordeum brachyantherum Nevski [-,2] T; 9250-10500'; S; mr.

Flordeum jubatum L. ssp. intermedium Bowden [2,7] A, C, R, T;

6380-10500'; G, S; fr, me, ml, mr, pp, ra.

Flordeum jubatum L. ssp. jubatum [2,7] A, C, M, S,T; 5800-10500';

L, S; br, fr, me, ml, mr, ra.

* Flordeum murinum L. ssp. glaucum (Steud.) Tzvelev [1,2] A, T;

5781 -7200'; G, L, S; fr, ml, mr, ra.

Flordeum pus ilium Nutt. [1,-] A; 6200'; L; pg, ra.

Koeleria macrantha (Ledeb.) Schult. [35,57] A, C, M, R, S, T; 5700-

12960'; G, L, P, S; af, br, fr, me, mm, mr, pj, pp, ra, sf.

* Lolium perenne L. var. aristatum Willd. [3,1 ] A, C, S; 7650-10800';

S; br, mr, ml, ra.

* Lolium perenne L. var. perenne [2,2] A, T; 8350-11340'; S; me,

mm, mr, ra.

Lycurus setosus (Nutt.) C. Reeder [6,-] A, S; 5700-7000'; L, S; ds,

P9, Pj, PP-

Melica ported Scribn. var. ported [17,7] A, C, M, S, T; 7760-12115';

S; me, mr, sf.

Muhlenbergia arenicola Buckley [2,-] A; 5650-5840'; L; pg, pj.

Muhlenbergia asperifolia (Nees & Meyen ex Trin.) Parodi [2,-] A, R;

5610-6620'; G,L;fr.

Muhlenbergia filiformis (Thurb. ex S. Watson) Rydb. [UNM-A. Fleck

s.n., 19 Sep 1964]T;S.

Muhlenbergia minutissima (Steud.) Swal len [-,1 ] T; 8160-9400'; S; ra.

Muhlenbergia montana (Nutt.) Hitchc. [11,13] A, C, M, R, S, T;

7560-11500'; P, S; af, bw, me, mm, mr, ms, pj, pp, ra.

Muhlenbergiapaucifiora Buckley [2,-] A, S; 5650-8320'; L, S; ds, pj.

Muhlenbergia racemosa (Michx.) Britton, Sterns, & Poggenb. [1,-]

A; 7750'; S; mr, ra.

297

Muhlenbergia repens (J. Presl) Hitchc. [UNM-R. Sivinski 2627] A; S.

Muhlenbergia richardsonis (Trin.) Rydb. [-,1 ] T; 9200-9500'; S; mm.

Muhlenbergia torreyi (Kunth) Hitchc. ex Bush [4,1 ] A,T; 5700-7500';

L, S; ds, pg, pj.

Muhlenbergia wrightii Vasey ex J.M. Coult. [6,4] A, C, M, T; 7200-

9360'; P, S; ds, fr, mr, ms, pj, ra.

Munroa squarrosa (Nutt.) Torr. [1,4] A, T; 5610-7550'; L, P, S; ds,

fr, pj, ra.

Nassella viridula (Trin.) Barkworth [-,2] T; 6500-7450'; P, S; fr, ra.

Oryzopsis asperifolia Michx. [4,13] A, C, M, R, S,T; 7550-10200'; S;

br, me, mr, pp, sf.

Panicum bulbosum Kunth [5,1] A, C, M; 5650-8200'; L, S; pg, mr,

ms, pp.

Panicum capillare L. var. brevifolium (Rydb.) Shear [1,1] S, T;

6840-7500'; S; fr, pj, ra.

Phalaris arundinacea L. var. arundinacea [1,3] A, T; 5781 -8400'; G,

S; fr, ml, mr, ra.

Phleum alpinum L. var. alpinum [10,29] A, M, R, S,T; 8450-13024';

S; am, bw, me, mm, mr, sf.

* Phleum pratense L. var. pratense [34,27] A, C, M, S, T; 7000-

11750'; S; af, me, ml, mm, mr, pp, ra.

Phragmites australis (Cav.) Trin. ex Steud. ssp. berlandieri (E. Fourn.)

Saltonstall & Hauber [1,-] A; 7900'; S; ml.

Piptatherum micranthum (Trin. & Rupr.) Barkworth [7,20] A, R, S,T;

5650-9500'; G, L, P, S; br, ds, fr, me, pj, pp.

Piptochaetium pringlei (Beal) Parodi [3,-] A; 8000-8500'; S; mr,

ms, pp.

Poa alpina L. [1,2] S,T; 9800-12625'; S; am, sf.

* Poa annua L. [8,1 ] A, M, S, T; 8600-11340'; S; mr, ra.

Poa arctica R. Br. var. aperta (Scribn. & Merr.) Soreng [-,2] R, T;

11120-13024'; S; am, mr.

Poa arctica R. Br. var. grayana (Vasey) A. Love, D. Love, & B.M. Kapoor

[-,3] T; 10500-12700'; S; am, sf.

Poa bigelovii Vasey & Scribn. [1,-] A; 6200'; L; pg.

* Poa compressa L. [17,11 ] A, C, M, R, S, T; 7240-10700'; S; af, me,

ml, mm, mr, ms, pp, ra.

Poa fendleriana (Steud.) Vasey [2,51 ] A, C, R, S, T; 6100-12850'; G,

P, S; am, br, bw, ds, fr, me, mm, mr, pj, pp, ra, sf.

[Poa fendleriana subspecies]

Poa glauca Vahl var. glauca [-,3] T; 10603-12050'; S; am, sf.

Poa glauca Vahl var. rupicola (Nash ex Rydb.) B. Boivin [2,10] S, T;

11120-12960'; S; am, mr.

Poa interiorRydb. [-,13] M,T; 8500-12850'; S; am, bw, me, mm, mr, sf.

Poa leptocoma Trin. [3,9] M, R, T; 9400-11940'; S; me, mr, sf.

Poa occidentalis Vasey [3,1 ] A, M, T; 9340-11300'; S; af, me, mm, sf.

Poapalustris L. [1,6] A, C, T; 8000-9300'; S; me, ml, mr.

* Poa pratensis L. [32,65] A, C, M, R, S,T; 5781 -12200'; G, P, S; am,

af, bw, br, fr, me, ml, mm, ms, mr, pj, pp, ra, sf.

[Poa pratensis subspecies]

Poa reflexa Vasey & Scribn. ex Vasey [-,3] T; 9700-12000'; S; sf.

Poasecunda J. Presl [-,2] T; 8500-10000'; S; me, mr.

[Poa secunda subspecies]

* Polypogon monspeliensis (L.) Desf. [2,-] A, R; 5500-6620'; G, L;

fr, ml.

Puccinellia nuttalliana (Schult.) Hitchc. [-,1] T; 8175'; S; ml.

Schedonnardus paniculatus (Nutt.) Trel. [3,-] A, S; 5730-7240'; L,

S; pg, pj, ra.

Schizachnepurpurascens (Torr.) Swallen [2,1] A, T; 8700-9700'; S;

mm, mr.

Schizachyrium scoparium (Michx.) Nash var. scoparium [3,10] A, C,

R,T; 5650-8500'; L, P, S; br, fr, me, ms, pj, pp, ra.

* Secale cereale L. [1,1 ] R, S; 6232-8540'; G, S; br, fr, pj.

Setaria leucopila (Scribn. & Merr.) K. Schum. [1,-] A; 5500'; L; fr.

* Setaria viridis (L.) P. Beauv. [-,1 ] M; 9160-9320'; S; ra.

298

Sorghastrum nutans (L.) Nash [UNM-D. Kennemore 2222] A; L.

* Sorghum halepense (L.) Pers. [1,-] A; 5500'; L; fr.

Sphenopholis obtusata (Michx.) Scribn. var obtusata [1,-] R;

6500-6620'; G; ml.

Sporobolus airoides (Torr.)Torr. [-,2] T; 7100-7240'; S; mr, ra

Sporobolus cryptandrus (Torr.) A. Gray [5,7] A, C, R, S,T; 5570-9400';

L, S; br, ds, fr, pj, pp, ra.

Torreyochloa pallida (Torr.) G.L. Church var. pauciflora (J. Presl) J.l.

Davis [1,-] S; 8940'; S; mr.

Trisetum montanum Vasey [21,13] A, C, M, S, T; 7840-11800'; S; af,

me, mm, mr, sf.

Trisetum spicatum (L.) K. Richt. [9,29] A, M, R, S, T; 8450-13024'; S;

am, me, mm, mr, sf.

Vulpia octoflora (Walter) Rydb. [2,6] A, R, S, T; 6050-7598'; G, L, P,

S; fr, pg, pj, ra.

[Vulpia octoflora varieties]

Polemoniaceae

Aliciellapinnatifida (Nutt, ex A. Gray) J.M. Porter [1,-] S; 8200-8320';

S; mr.

Collomia linearis Nutt. [-,4] T; 8700-11500'; S; me, mm.

Eriastrum diffusum (A. Gray) H. Mason [-,2] R, T; 6036-7450'; P;

ds, fr, pj.

Gilia ophthalmoides Brand [-,4] R,T; 7050-8450'; G, S; fr, pj, pp.

Giliasinuata Douglas ex Benth. [-,1] T; 8400-8900'; S; me.

Ipomopsisaggregata (Pursh) V.E. Grant ssp. Candida (Rydb.) V.E. Grant

& A.D. Grant [-,4] C, T; 8500-10300'; S; mm, ra.

Ipomopsis aggregata (Pursh) V.E. Grant ssp. collina (Greene) Wilken

& Allard [2,2] M,T; 7700-11200'; S; me, mm, sf.

Ipomopsis aggregata (Pursh) V.E. Grant ssp. formosissima (Greene)

Wherry [37,26] A, C, M, R, S, T; 5800-11209'; L, P, S; af, br, fr,

me, mm, mr, ms, pj, pp, ra, sf.

Ipomopsis laxiflora (J.M. Coult.) V.E. Grant [4,1] A, R; 5610-6500';

L, P; ds, fr, pg, pj.

Ipomopsis longiflora (Torr.) V.E. Grant ssp. neomexicana Wilken [1,-]

A; 5500'; L; pj.

Ipomopsis multiflora (Nutt.) V.E. Grant [-,2] R,T; 7350-9700'; S; pj, ra.

Linanthus pungens (Torr.) J.M. Porter & L.A. Johnson [-,2] T; 6900-

7450'; P, S; ds, me, pp.

Microsteris gracilis (Hook.) Greene [-,3] R, T; 7050-8100'; G, S; fr, pj.

Phloxcondensata (A. Gray) E. Nelson [3,-] S; 12160-12500'; S; am.

Phlox longifolia Nutt. ssp. longifolia [1,10] R, S, T; 6976-8100'; G,

P, S; br, fr, pj, ra.

Phloxnana Nutt. [15,-] A, S; 5700-8000'; L, S; pg, pj, pp, ra.

Phloxpulvinata (Wherry) Cronquist [-,3]T; 11500-13161'; S; am.

Polemonium brandegei (A. Gray) Greene [UNM-E. Castetter & H.

Dittmer 9827] T; S.

Polemonium foliosissimum A. Gray [10,3] A, M, T; 8000-10800'; S;

ml, mm, mr.

! Polemonium occidentale Greene var. occidentale [-,1 ] T; 9475';

S; mm, mr.

Polemonium pulcherrimum Hook. var. delicatum (Rydb.) Cronquist

[1,12] A, T; 9800-11960'; S; bw, me, mr, sf.

Polemonium viscosum Nutt. [1,4] S,T; 11500-13000'; S; am.

Polygalaceae

Polygala alba Nutt. var. alba [2,-] A; 5700-5800'; L; pg, pj.

Polygonaceae

Bistorta bistortoides (Pursh) Small [7,16] A, M, R, S,T; 9300-13024';

S; am, mm, mr, sf.

Bistorta vivipara (L.) S.F. Gray [2,9] A, M, T; 8300-13009'; S; am,

ml, mr, sf.

Eriogonum alatum Torr. var. alatum [12,6] A, M, R, S,T; 5800-9640';

G, L, P, S; br, fr, me, mm, ms, pj, pp, ra.

Eriogonum annuum Nutt. [1,-] A; 5570-5600'; L; fr.

Journal of the Botanical Research Institute of Texas 8(1)

Eriogonum jamesii Benth. var . jamesii [16,25] A, C, M, R, S, T;

5600-11200'; L, P, S; br, ds, fr, mm, mr, pj, pp, ra.

Eriogonum lachnogynum Torr. ex Benth. var. lachnogynum [ 1 ,-] A;

5730'; L; ds, pj.

Eriogonum microthecum Nutt. var. simpsonii (Benth.) Reveal [-,3] R,

T; 7200-7500'; S; fr, pj, ra.

Eriogonum racemosum Nutt. [3,13] A, R, S, T; 7000-9400'; P, S; br,

me, mr, pj, pp, ra.

Eriogonum tenellumTorr. [5,-] A; 5570-6200'; L; fr, pg, pj.

* Fallopiabaldschuanica (Regel) Holub[UNM-C.R. Hutchins 5887]

R; G.

* Fallopia convolvulus L. [UNM-R. Sivinski 2427] A; S.

Oxyria digyna (L.) Hill [-,3] T; 11150-12700'; S; am, bw, sf.

* Persicaria lapathifolia (L.) A. Gray [2,2] A, C, T; 5500-8194'; L, S;

fr, ml, ra.

* Persicaria maculosa A. Gray [2,-] S; 6840-7000'; S; fr, ra.

* Polygonum aviculare L. [8,8] A, M, R, S, T; 6540-10887'; G, S; ds,

fr, ml, mm, mr, ra.

Polygonum douglasii Greene [2,8] A, M, R, T; 7200-12050'; S; af,

me, mm, mr, ra, sf.

Polygonum ramosissimum Michx. var. ramosissimum [-,2] T;

8700-9700'; S; me.

Polygonum sawatchense SmaW [-,3] R,T; 7200-8550'; P, S; pj, pp, ra.

* Rumex acetosella L. [21,10] A, M, S, T; 7000-11700'; S; af, me,

mm, mr, ra.

Rumexaltissimus A.W. Wood [1,-] A; 5700-5800'; L; fr, ra.

* Rumexcrispus L. ssp. crispus [23,5] A, C, M, S, T; 6540-9680'; G,

S; me, ml, mr, ra.

Rumexdensiflorus Osterh. [-,3] C,T; 8500-10986'; S; me, mr.

Rumex fueginus Phil. [-,1 ] C; 8194'; S; ml.

[Rumex maritimus L.]

Rumexmexicanus Meisn. [UNM-E. Castetter 3879] T; G.

Rumex occidentalis S. Watson [-,9] M, R,T; 7240-11500'; S; ml, mr, ra.

* Rumexpatientia L. [3,-] A, S; 5800-7580'; L, S; fr, mr, ra.

* Rumexpulcher L. [-,1 ] T; 7350-7450'; S; ra.

Rumex triangulivalvis (Danser) Rech. f. [2,2] A, T; 7200-10660'; S;

ml, mr, ra.

Portulacaceae

Claytonia megarhiza (A. Gray) Parry ex S. Watson [-,2] T; 11500-

13000'; S; am.

Lewisia nevadensis (A. Gray) B.L. Rob. [UNM-R. Sivinski 3921 ] S; S.

Montia chamissoi (Ledeb. ex Spreng.) Greene [-,2] T; 9150-10500';

S; mr.

Phemeranthus brevicaulis (S. Watson) Kiger [UNM-R. Sivinski 4556]

A; S.

Phemeranthusparviflorus (Nutt) Kiger [1,-] A; 7900'; S; ml, ms.

* Portulaca oleracea L. [2,2] A, S,T; 5700-9400'; L, S; fr, ml, pj, pp,

ra.

[Portulaca oleracea subspecies]

Potamogetonaceae (includes Zannichelliaceae)

Potamogeton diversifolius Raf. [1,-] A; 7200-7240'; S; ml.

Potamogeton foliosus Raf. var. foliosus [UNM-J. McGrath 444] C; S.

Potamogeton gramineus L. [UNM-J. McGrath 442] C; S.

Potamogeton natans L. [-,1 ] T; 9375-10625'; S; mr.

Potamogeton nodosus Poir. [1,-] A; 7200-7240'; S; ml.

Potamogeton pusillus L. var. pusillus [UNM-R. Sivinski & B. Simpson

2301] A; S.

Zannichelliapalustris L. [UNM-J. McGrath 437] C; S.

Primulaceae

Androsace chamaejasme Wulfen. var. arctica Kunth [-, 6 ] T; 10500-

13000'; S; am, mm.

[Androsace chamaejasme Wulfen ssp. lehmanniana (Spreng.)

Hulten]

Larson et al., Floristic studies of the Sangre de Cristo Mountains

Androsace occidentalis Pursh [-,1] R; 7150'; S; pj, ra.

Androsoce septentrionolis L. [18,62] A, C, M, R, S,T; 6200-12960'; G,

P, S; am, br, ds, me, mm, ms, mr, pj, pp, sf.

Dodecotheon pulchellum (Raf.) Merr. var. pulchellum [13,13] A, M,

R, S, T; 7740-11500'; S; me, ml, mm, mr, sf.

[Primula pauciflora (Greene) Mast & Reveal]

Primula angustifolia Torr. [-,4] T; 11500-13009'; S; am.

Primulaparryi A. Gray [2,4] M, S, T; 9800-12430'; S; am, mr, sf.

Primula rusbyi Greene [UNM-E. Castetter 3301 -A] A; S.

Ranunculaceae

Aconitum columbianum Nutt. var. columbianum [31,27] A, C, M, S,

T; 7400-12960'; S; mm, mr, sf.

Actaea rubra (Aiton) Willd. var. arguta (Nutt.) Lawson [26,15] A,C,

M, R, S, T; 7775-11115'; S; af, me, mm, mr.

[Actaea rubra ssp. arguta (Nutt.) Hulten]

Anemone canadensis L. [-,3] T; 8350-9400'; S; me, mm, mr.

Anemone cylindrica A. Gray [1,-] A; 7720'; S; pp.

Anemone multifida Poir. [-,4] T; 9250-10500'; S; me, mm, mr.

Aquilegia coerulea E. James var. coerulea [12,18] A, M, S,T; 7720-

12960'; S; af, am, bw, me, ml, mr, sf.

Aquilegia elegantula Greene [8,20] A, C, R, S, T; 7550-11500'; S;

br, me, mr, pp, sf.

Caltha leptosepala DC. [9,15] A, M, S, T; 9300-12960'; S; am, ml,

mm, mr.

Clematis Columbiana (Nutt.) Torr. & A. Gray var. Columbiana [24,23]

A, C, M, R, S, T; 7250-10600'; P, S; am, br, ds, me, mr, pp.

Clematis hirsutissima Pursh var. scottii (Porter) E.O. Erickson [-,1] T;

7350-7500'; P; ds.

Clematis ligusticifolia Nutt. [3,4] A, S,T; 5570-8400'; L, P, S; fr, mr, ra.

♦ Delphinium alpestre Rydb. [-,4] T; 11120-12960'; S; am.

Delphinium barbeyi (Huth) Huth [3,10] M ,S, T; 8450-12000'; S;

me, mr, sf.

Delphinium nuttallianum Pritz var. nuttallianum [-,3] R, T; 7600-

8900'; S; pj, pp.

Delphinium ramosum Rydb. [-,4] C,T; 7840-11500'; S; me, mr, sf.

Delphiunium robustum Rydb. [UNM-M. Schiebout 8846] M; S.

+♦ Delphinium sapellonis Cockerell [9,3] A, M, T; 7840-10800'; S;

af, me, mm, mr, sf.

Delphinium wootonii Rydb. [3,-] A, S; 5700-7200'; L, S; pg, pj, ra.

Pulsatilla patens (L.) Mill. ssp. multifida (Pritz.) Zamels [-,5] C, T;

7350-10600'; P, S; ds, mm, pp.

Ranunculus abortivus L. [UNM-R. Fletcher 6252] T; S.

Ranunculus aquatilis L. var. diffusus With. [-,5] C, T; 8000-9500'; S;

ml, mr, pp.

Ranunculus cardiophyllus Hook. [UNM-J. Williams 53] T; S.

Ranunculus cymbalaria Pursh [-,2] C, T; 8175-8194'; S; ml.

Ranunculus gmeliniii DC. [UNM-E. Castetter 4465] C; S.

Ranunculus inamoenus Greene var. inamoenus [4,9] A, R, S, T;

8400-11500'; S; bw, me, mm, mr, sf.

Ranunculus macauleyik. Gray [2,8] S,T; 10500-13000'; S; am, mr, sf.

Ranunculus macounii Britton [6,-] A, M, S; 7740-8940'; S; me, mr.

Ranunculus pensylvanicus L. f. [1,-] S; 8250-8540'; S; mr, ra.

Ranunculus ranunculinus (Nutt.) Rydb. [2,1] A, C; 7900-8575'; S;

me, pp, ra.

!* Ranunculus repens L. [1,-] A; 8000'; S; mr, ra.

Ranunculus scleratus L. var. multifidus Nutt. [UNM-E. Castetter

4473] C; S.

* Ranunculus testiculatus Crantz [-,1 ] T; 7350-7450'; P; ds.

Ranunculus uncinatus D. Don ex G. Don [-,3] T; 7775-8150'; S; ml, mr.

[Ranunculus uncinatus var. earlei (Greene) L.D. Benson]

Thalictrum alpinum L. [-,2] T; 11500-13009'; S; am.

Thalictrum fendleri Engelm. ex A. Gray [25,20] A, C, M, R, S, T;

7400-12050'; S; br, bw, me, mm, mr, ra.

Thalictrum revolutum DC. [UNM-J. McGrath 729] A; S.

299

Trautvetteria caroliniensis (Walter) Vail [5,4] A, M, R, S, T; 8450-

11900'; S; ml, mr, sf.

Rhamnaceae

Ceanothus fendleri A. Gray [13,6] A, C, M, S, T; 7650-10500'; S; br,

me, ms, pp.

Rosaceae

Agrimonia gryposepala\Na\\r. [2,-] A; 7900-8230'; S; me, mr.

Agrimonia striata Michx. [11,2] A, C, S, T; 7580-10500'; S; mr.

* Alchemilla vulgaris L. [UNM-R. Fletcher 8429] T; S.

Amelanchier alnifolia (Nutt.) Nutt, ex Roem. var. alnifolia [12,7] A,

M, R, S, T; 7620-10500'; S; me, mr, sf.

Amelanchier utahensis Koehne [-,2] T; 7250-9600'; S; br, pj.

Cercocarpus montanus Raf. var. montanus [25,40] A, C, M, R, S, T;

5650-11500'; G, L, P, S; br, ds, fr, me, mm, mr, ms, pj, pp, ra.

Crataegus macracantha Lodd. ex Loud. var. occidentalis (Britton)

Eggleston [2,-] A, S; 7760-8400'; S; mr, ms.

Dasiphora fruticosa (L.) Rydb. [23,29] A, C, M, R, S, T; 8310-13024';

S; af, am, fr, me, ml, mm, ms, mr, ra, sf.

FallugiaparadoxaiD. Don) Endl. ex Torr. [8,18] A, R, S,T; 5570-8900';

G, L, P, S; br, ds, fr, me, pj, pp, ra.

Fragaria vesca L. [16,18] A, C, M, R, S, T; 7350-11900'; S; am, br,

me, mr, pp, sf.

Fragaria virginiana Mill.[7,33] A, C, R, S,T; 8000-11650'; S; bw, br,

me, mm, mr, sf.

Geum aleppicum Jacq. [13,4] A, M, S, T; 7760-10800'; S; af, me,

mm, mr.

Geum macrophyllum Willd. var. perincisum (Rydb.) Raup [13,13] A,

M, R, S, T; 7050-11209'; S; me, mr, sf.

Geum rivale L. [-,2] T; 8450-10500'; S; mr.

Geum rossii (R. Br.) Ser. var. turbinatum (Rydb.) C.L. Hitchc. [5,17] A,

M, R,T; 9800-13161'; S; am, mm, mr.

Geum triflorum Pursh var. ciliatum (Pursh) Fass. [-,2] T; 9850-10500';

S; mm, mr.

Flolodiscus dumosus (Nutt, ex Hook.) A. Heller [14,17] A, C, M, R, S,

T; 7560-11200'; P, S; af, me, mr, ms, pp, ra, sf.

* Maluspumila Mill. [6,2] A, S, T; 6840-8450'; S; mr, ra.

Physocarpus monogynus (Torr.) J.M. Coult. [21,3] A, M, S, T; 7720-

10200'; S; af, me, mr, ra.

Potentilla ambigens Greene [1,-] S; 7600'; S; mr.

Potentilla anserina L. [1,11] A, C, M, R, T; 5500-11000'; G, L, P, S;

fr, ml, mm, mr, ra.

Potentilla concinna Richardson var. bicrenata (Rydb.) S.L.Welsh &

B.C. Johnst. [-,1] T; 7700-8555'; P; me.

[Potentilla bicrenata Rydb.]

Potentilla concinna Richardson var. concinna [2,8] A, C, M, R, T;

8380-13000'; S; am, me, mm, pp, sf.

Potentilla crinita A. Gray var. crinita [2,-] S; 7380-8550'; S; mr, ra.

Potentilla diversifolia Lehm. var. diversifolia [1,13] M, R, T; 7850-

12960'; S; am, af, ml, mr, sf.

! Potentilla fissa Nutt. [1,-] A; 10000-10212'; S; me.

Potentilla gracilis Douglas ex Hook. var. glabrata (Lehm.) C.L. Hitchc.

[1,1] S,T; 9950-12430'; S; am, mr.

Potentilla gracilis Douglas ex Hook. var. pulcherrima (Lehm.) Fernald

[25,24] A, C, M, R, S,T; 7840-12584'; S; af, am, bw, br, me, mm,

mr, ra, sf.

Potentilla hippiana Lehm. var. hippiana [29,28] A, C, M, R, S, T;

7000-11800'; S; af, am, br, me, mm, mr, ms, pp, ra, sf.

x Potentilla hippiana Lehm. xP. gracilis Douglas ex Hook. var.

pulcherrima (Lehm.) Fernald [1,1] A, T; 8160-11300'; S; mr, sf.

Potentilla norvegica L. ssp. monspeliensis (L.) Asch. & Graebn. [3,6]

A, C, T; 5781 -10440'; G, S; fr, me, ml, mr, ra.

[Potentilla norvegica]

Potentilla pensylvanica L. var. pensylvanica [8,13] A, C, M, T; 6900-

10700'; P, S; br, ds, me, mm, mr, ra.

300

Journal of the Botanical Research Institute of Texas 8(1)

Potentilloplattensis Nutt. [UNM-J. Williams 31 ] T; S.

* Potentilla recta L. [UNM-J. McGrath 728] A; S.

Potentilla subviscosa Greene [-,1] C; 8400-8500'; S; pp.

Potentilla thurberi A. Gray var. atrorubens (Rydb.) Kearney & Peebles

[4,-] A; 7720-8575'; S; mr, pp, ra.

Potentilla thurberi A. Gray var. thurberi [1,-] S; 7760'; S; mr.

* Prunus americana Marshall [-,2] R,T; 6540-7300'; G, S; ds, fr, ra.

Prunus virginiana L. var. melanocarpa (A. Nelson) Sarg. [29,14] A, M,

S,T; 5800-9675'; L, P, S; br, fr, me, mm, mr, ra, sf.

Rosa acicularis Lindl. var. sayana Erlanson [23,19] A, C, M, S, T;

5800-11500'; G, S; af, br, me, mr, pp, ra, sf.

Rosa arkansana Porter var. arkansana [15,4] A, M, S,T; 7000-10200';

S; af, me, ml, mm, mr, ra.

Rosa nutkana C. Presl [5,12] A, M, R, S, T; 7775-12000'; S; me, mr,

PP, sf.

[Rosa nutkana ssp. melina (Greene) W.H. Lewis & Ertter]

Rosa woodsii Lindl. var. ultramontana (S. Watson) Jeps. [4,9] A, M,

T; 5800-10000'; L, S; br, fr, me, mm, mr.

Rubus idaeus L. var. aculeatissimus Regel & Tiling [37,20] A, C, M, R,

S,T; 7600-11940'; S; af, am, me, mr, ra, sf.

[Rubus idaeus var. strigosus (Michx.) Maxim.]

Rubus deliciosuslorr. var. neomexicanus (A. Gray) Kearney [UNM-M.

Schiebout 3706] A; S.

Rubus parviflorus Nutt. var. parviflorus [28,18] A, C, M, S, T; 7600-

1 0880'; S; af, me, mr, sf.

* Sanguisorba minor (Scopoli) ssp. muricata (Spach) Nordborg

[UNM -C.R. Hutchins 9683] A; S.

Sibbaldia procumbens L. [4,11] A, M, R, S, T; 9600-13024'; S; am,

mm, mr.

Sorbus dumosa Greene [1,-] M; 9760-10600'; S; mr.

Sorbusscopulina Greene var. scopulina [-,5] R,T; 7900-11500'; S; mr.

Rubiaceae

* Galium aparine L. var. echinospermum (Wallr.) Farw. [-,5] T;

7600-10500'; S; me, mr, ra.

[Gallium aparine]

Galium boreale L. [23,37] A, C, M, S, T; 7440-11150'; S; af, br, me,

ml, mm, mr, pj, ra, sf.

Galium fendleri A. Gray [7,-] A, S; 7900-8900'; S; me.

Galium mexicanum Kunth var. asperrimum (A. Gray) Higgins & S.L.

Welsh [23,2] A, C, M, S, T; 7600-10500'; S; af, me, mm, mr, sf.

Galium trifidum L. var .subbiflorum Wiegand [1,1 ] S,T; 8410-11050';

S; mr, sf.

Galium triflorum Michx. [13,6] A, M, S, T; 7620-11115'; S; me, mm,

mr, sf.

Houstonia acerosa (A. Gray) A. Gray ex Benth. & Hook. var. polypre-

moides (A. Gray) Terrell [1,-] A; 5700-5800'; L; pg, pj.

Houstonia rubra Cav. [3,-] A; 5650-5840'; L; pg, pj.

Stenaria nigricans (Lam.) Terrell var. nigricans [UNM-C.R. Hutchins

9948] S; S.

[Hedyotis nigricans (Lam.) Fosberg var. nigricans]

Rutaceae

Ptelea trifoliata L. [5,7] S,T; 5800-8000'; G, P, S; ds, fr, mr, pj, ra.

[Ptelea trifolata infrataxa]

Salicaceae

x Populus xacuminata Rydb. [2,2] R, S,T; 6380-7950'; G, S; fr, mr.

Populus angustifolia E. James [13,26] A, C, M, R, S, T; 6380-9400';

G, P, S; fr, mm, mr, ra.

Populus deltoides W. Bartram ex Marshall var. wislizenii (S. Watson)

Dorn [4,6] A, R, S, T; 5610-7100'; G, L, P, S; fr, pj, ra.

Populus tremuloides Michx. [41,44] A, C, M, R, S, T; 7000-12300'; S;

af, br, me, mm, mr, pj, pp, sf.

Salixarctica Pall. var. petraea (Andersson) Bebb [-,7] R, T; 11990—

13161'; S; am.

♦ Salixarizonica Dorn [1,-] M; 10500'; S; mr.

Salixbebbiana Sarg. [14,7] A, M, R, S,T; 7750-10500'; S; me, mr, ra, sf.

Salix brachycarpa Nutt. var. brachycarpa [1,6] M, T; 9800-11960';

S; am, mr.

Salix drummondiana Barratt ex Hook. [11,7] A, M, R, S, T; 8000-

10850'; S; mr, sf.

Salixeriocephala Michx. var. ligulifolia (C.R. Ball) Dorn [8,4] A, M, R,

S,T; 6450-8990'; G, S; fr, me, ms, mr, pp.

Salixexigua Nutt. var. exigua [11,22] A, C, M, R, S,T; 5781 -9300'; G,

L, P, S; fr, ml, mr, pj, pp, ra.

♦ Salix fragilis L. [1,-] A; 5500'; L; fr.

Salix glauca L. var. villosa (D. Don ex Hook.) Andersson [-,1] T;

11529-11793'; S; am.

Salixgooddingii C.R. Ball [1,1] A, R; 5800-6540'; G, L; ds, fr, ra.

Salix irrorata Andersson [11,7] A, C, M, R, S, T; 6540-9100'; G, S;

fr, mr, ms, pj.

Salix lasiandra Benth. var. caudata (Nutt.) Sudw. [-,2] R, T; 6380-

7600'; G, S; fr, mr.

[Salixlucida Muhl. ssp. caudata (Nutt.) Murray]

Salix lasiandra Benth. var. lasiandra [5,1] A, S, T; 7850-10500'; S;

me, mr.

[Salixlucida Muhl. ssp. lasiandra (Benth.) Murray]

Salixmonticola Bebb [5,3] A, M, T; 7200-10180'; S; ml, mm, mr.

Salix planifolia Pursh var. planifolia [4,3] M ,S, T; 9200-12000'; S;

am, me, ml, mm, mr.

Salix reticulata L. var. nana Andersson [2,2] S, T; 11500-12430';

S; am.

[Salix reticulata ssp. nivalis (Hook.) A. Love, D. Love, & B.M.

Kapoor]

Salixscouleriana Barratt ex Hook. [8,8] A, M, S,T; 8000-10700'; S;

br, me, mr, sf.

Salix wolfii Bebb var. wolfii [UNM-D. Atwood 21490] A; S.

Santalaceae (Viscaceae)

Arceuthobium divaricatum Engelm. [1,4] R, S,T; 7000-8300'; S; pj.

Arceuthobium douglasii Engelm. [1,4] S,T; 7600-10093'; P, S; me, pj.

Arceuthobium vaginatum (Willd.) J. Presl var. cryptopodium (Engelm.)

Cronquist [-,7] C, R,T; 7600-8900'; S; br, pp.

Comandra umbellata (L.) Nutt. var. pallida (DC.) M.E. Jones [-,7] C,

T; 7550-9675; S; me, pj, pp, ra.

Phoradendron juniperinum Engelm. ex A. Gray var. juniperinum [3,9]

R, S,T; 6500-9400'; G, S; pj, pp.

Saururaceae

Anemopsis californica Hook. & Arn. [UNM- R.C. Sivinski 2471 ] A; S.

Saxifragaceae

Boykiniajamesii (Torr.) Engler [1,-] A; 10100-10200'; S; me.

[Telesonixjamesii (Torr.) Raf.]

Heuchera parvifolia Nutt, ex Torr. & A. Gray [10,33] A, M, R, S, T;

7050-13024'; G, P, S; am, br, ds, me, mm, mr, pj, pp, sf.

+ Heuchera wootonii Rydb. [UNM-E. Castetter & H. Dittmer 9825]

T; S.

Mitella stauropetala Piper var. stenopetala (Piper) Rosend. [-,1] T;

10000-10850'; S; mr.

Saxifraga bronchialis L. var. austromontana (Wiegand) Piper ex

G.N. Jones [20,23] A, C, M, R, S, T; 7750-13024'; S; am, me,

mm, mr, ra, sf.

♦ Saxifraga cernua L. [1,-] S; 11900-11940'; S; am.

Saxifraga chrysantha A. Gray [-,1 ] T; 12144-13009'; S; am.

Saxifraga debilis Engelm. ex A. Gray [UNM-H. Mackay 9T-3] T; S.

Saxifraga flagellaris Willd. ex Sternb. var. crandallii (Gand.) Dorn [-,3]

T; 11500-13009'; S; am.

Saxifraga hirculus L. var. hirculus [UNM-T. Lowrey 2099] C; S.

Saxifraga odontoloma Piper [6,18] A, M, R, S, T; 8410-11960'; S;

me, mm, mr, sf.

Larson et al., Floristic studies of the Sangre de Cristo Mountains

301

[Micranthes odontolomo (Piper) A.A. Heller]

Soxifrogo rhomboideo Greene [2,8] A, C, S, T; 9900-13009'; S; am,

me, ml, mr.

[Micranthes rhomboideo (Greene) Small]

Scrophulariaceae (see also Orobanchaceae, Plantaginaceae)

Scrophulorio lanceolata Pursh [-,1 ] T; 7850-10500'; S; me, mr.

+ Scrophulorio montana Wooton [2,1] A, M, T; 8450-10660'; S;

mm.

* Verbascum thapsus L. [19,1 5] A, C, M, R, S, T; 5570-10500'; L, S;

af, br, fr, me, ml, mr, pj, pp, ra.

Solanaceae

Chamaesaracha coronopus (Dunal) A. Gray [5,-] A; 5700-6200'; L,

S; fr, pg, pj.

Chamaesaracha coniodes (Moric.) Britton [3,-] A; 5570-5800'; L;

fr, pg, pj.

Datura wrightii Regel [1,-] A; 5500'; L; fr.

Lycium pallidum Miers [1,-] A; 5800-5840'; L; pj.

Nicotiono trigonophyllo Dunal [1,-] A; 5600-5800'; L; pj.

Physolis hederifolio A. Gray var. comata (Rydb.) Waterf. [1,-] A;

5700-5800'; L; fr, ra.

Physolis hederifolio A. Gray var. fendleri (A. Gray) Cronquist [3,3] A,

R, S, T; 6232-8000'; G, S; fr, pg, pj, ra.

Physolis longifolio Nutt. var. longifolio [1,-] A; 6980'; S; pp.

Physolissubuloto Rydb. var. neomexicono (Rydb.) Waterf. ex Kartesz

& Gandhi [3,1] A, S,T; 6840-7840'; S; fr, mr, ra.

[Physolis foetens Poir. var. neomexicono (Rydb.) Waterf. ex Kartesz

& Gandhi]

Solanum elaeagnifolium Cav. [11,-] A; 5570-7000'; L, S; fr, mr, pg,

pj, ra.

Solanum jamesii Torr. [UNM-Bamberg 60] A; S.

Solanum nigrum L. [UNM-H. Mackay 6T-58] T; S.

Solanum rostratum Dunal [2,-] A; 5500-561 O'; L; fr.

Sparganiaceae

Sporgonium emersum Rehmann [-,1] T; 9375-9675'; S; ml, mr.

Tamaricaceae

*• Tamarixchinensis Lour. [2,8] A, R,T; 5500-8175'; G, L, P, S; fr, ml,

ra.

Typhaceae

Typho lotifolio L. [1,4] R,T; 6380-7660'; G, S; fr, ml, mr, ra.

Ulmaceae

*• Ulmus pumilo L. [13,11 ] A, R, S, T; 5800-8700'; G, L, P, S; fr, mr,

pj, PP, ra.

Urticaceae

* Urtico dioico L. var. procero (Muhl. ex Willd.) Wedd. [18,22] A, C,

M, R, S, T; 7400-11500'; S; me, ml, mm, mr, ra, sf.

[Urtico dioico ssp. gracilis (Aiton) Selander]

Verbenaceae

Glondulorio bipinnotifido (Nutt.) Nutt. var. bipinnotifido [16,-] A, S;

5600-7500'; L, S; mr, pg, pj.

Phyla cuneifolio (Torr.) Greene [1,-] A; 5500'; L; fr.

Verbena brocteoto Lag. & Rodr. [8,4] A, C, S,T; 5500-9070'; L, S; br,

fr, me, mr, pj, pp, ra.

Verbena macdougalii A. Heller [23,8] A, C, M, S,T; 6840-10500'; S;

af, fr, me, ml, mr, pj, pp, ra.

Violaceae

Viola odunco J.E. Sm. [1,11 ] R, S, T; 8000-12850'; S; am, me, mr, sf.

Viola canadensis L. [19,29] A, C, M, R, S, T; 7320-12000'; S; af, br,

me, mr, ra, sf.

Viola nephrophyllo Greene [1,2] A, R,T; 7740-8700'; S; mr.

Vitaceae

Porthenocissus vitoceo (Knerr) Hitchc. [7,5] A, R, S, T; 5610-7493';

G, L, P, S; fr, mr, pj, pp, ra.

Vitis orizonico Engelm. [3,1 ] A, T; 5610-6540'; G, L; fr, ra.

Zygophyllaceae

* Tribulus terrestris L. [3,-] A, S; 5610-7000'; L, S; fr, mr, ra.

ERRATUM

In the prior article (Reif et al. 2009), we incorrectly reported 3 collections of Asclepias macrosperma. These

specimens are A. macrotis. A corrected checklist entry for this taxon appears below:

Asclepios mocrotisJorr. [1,-,4,3] D,L,R; 5300-6900'; G, P, U; ds, fr, pj.

ACKNOWLEDGMENTS

Staff of Bureau of Land Management (Taos District), Carson National Forest, Santa Fe National Forest, and

Valles Caldera National Preserve are thanked for funding of the projects, the use of facilities, and assistance in

numerous other ways. Key individuals include Chirre Keckler, Randy McKee, Linus Meyer, and Bob Par-

menter. Taos Pubelo and Picuris Pueblo, and Robert Espinosa andjoe I. Quanchello, of their respective tribes,

were instrumental in granting access to reservation lands.

The following individuals are thanked for identification or verification of specimens: Jennifer Ackerheld,

Kelly Allred, Duane Atwood, Mary Barkworth, Karen Clary, Robert D. Dorn, Bill Jennings, Bernadette Kuhn,

Ben Legler, Tim Lowrey, Stewart Markow, and Bob Sivinski.

We acknowledge the valuable contribution by the graduate committees that included Greg Brown, Dave

McDonald, and Steve Prager. Alex Buerkle, Larry Schmidt, and Jeff Lockwood also provided assistance, sup¬

port, and inspiration.

Tim Lowrey, Director of the University of New Mexico Herbarium, Jane Mygatt and later Phil Tonne, both

Senior Collection Managers, provided access to the collections and data base.

Finally, we thank Tim Hogan and an anonymous reviewer for their thoughtful comments.

302

Journal of the Botanical Research Institute of Texas 8(1)

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BOOK NOTICE

James Solomon, Tatyana Shulkina, and George Schatz, eds. 2014. Red List of the Endemic Plants of the Cauca¬

sus: Armenia, Azerbaijan, Georgia, Iran, Russia, and Turkey. (ISBN-13: 978-0915279821, hbk).

Monographs in Systematic Botany from the Missouri Botanical Garden, Volume 125 (ISSN: 0161-1542). Mis¬

souri Botanical Garden Press, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. (Orders: www.mbg-

press.org). Price not given, 768 pp., illustrated.

From the publisher: Red List of the Endemic Plants of the Caucasus: Armenia, Azerbaijan, Georgia, Iran, Russia, and

Turkey provides the brst floristic and conservation analysis of the plants of the Caucasus region, with assess¬

ments for over 60% of the endemic taxa, including top priorities for conservation action. This book was made

possible by an unprecedented collaboration between botanists from all six regions covered within.

J.Bot. Res. Inst. Texas 8(1): 304.2014

LOW GENETIC DIVERSITY AND POOR DISPERSAL, BUT NOT CONSERVATION

STATUS RANK, ARE LINKED TO CLIMATE CHANGE VULNERABILITY

Cynthia M. Morton

Section of Botany

Carnegie Museum of Natural History

4400 Forbes Avenue

Pittsburgh, Pennsylvania 15213, U.S.A.

Matthew D. Schlesinger

Chief Zoologist

New York Natural Heritage Program

625 Broadway, 5th Floor

Albany, New York 12233, U.S.A.

ABSTRACT

Climate change vulnerability assessments for Pennsylvania were completed for 35 plant species using the Climate Change Vulnerability

Index (CCVI v2.0) developed by NatureServe. The CCVI allows the user to examine the exposure and sensitivity of a species to a series of

risk factors associated with climate change. This study, as well as studies from West Virginia (Byers & Norris 2011) and New York

(Schlesinger et al. 2011), indicates that among the top five risk factors, based upon both floral and fauna assessments, appeared to be related

to a lack of dispersal and movement mechanisms along with low genetic diversity. All of the above studies found that conservation status

rankings and vulnerability to climate change were not directly related to one another. In light of these findings conservation protocols need

to be reexamined to direct resources where they will be most effective in furthering the conservation of plant species.

RESUMEN

Se completaron evaluaciones de vulnerabilidad por el cambio climatico en Pennsylvania para 35 especies vegetales usando el Climate

Change Vulnerability Index (CCVI v2.0) desarrollado por NatureServe. El CCVI permite al usuario examinar la exposicion y sensibilidad

de una especie a una serie de factores de riesgo asociados con el cambio climatico. Este estudio, asi como estudios de West Virginia (Byers &

Norris 2011) y Nueva York (Schlesinger et al. 2011), indica que entre los cinco mayores factores de riesgo, basados en evaluaciones de flora y

fauna, parecieron estar relacionados con una falta de mecanismos de dispersion y movimiento junto con una diversidad genetica baja. Todos

los estudios encontraron que las graduaciones de estatus de conservacion y vulnerabilidad al cambio climatico no estaban relacionadas di-

rectamente una con otra. A la luz de estos hallazgos los protocolos de conservacion necesitan ser reexaminados para dirigir los recursos alii

done son mas efectivos el fomento de la conservacion de especies vegetales.

INTRODUCTION

Global climate change (e.g., increasing temperatures, increased carbon dioxide levels, and altered patterns of

precipitation) may alter the distribution of plant species and natural plant communities and may decrease

habitat value for wildlife over broad segments of North America. A number of studies have shown that as cli¬

mates warm, many species will suffer a decline in population and reduced range sizes while others will experi¬

ence an increase in populations and range sizes. The relative vulnerability of species or habitats to climate

change can be used to set goals, determine management priorities, and direct resources where they will be

most effective in furthering the conservation of plant species biodiversity (Glick et al. 2011).

Climate change is only one of many stresses, or pressures from the external environment that plant spe¬

cies and their habitats are currently experiencing. The management strategies traditionally used to address

conventional threats to biodiversity will likely be similar to those needed for threats induced by climate

change. The list of plant taxa which are currently deemed most at risk will change as climate change alters spe¬

cies distribution and population viability, and re-emphasized strategies may become more important as habi¬

tats change.

One planning tool that is increasingly employed for conservation and management decisions is the vul¬

nerability assessment. These assessments typically are models in which the inputs are characteristics of the

species or ecosystems and the output is a rating of relative vulnerability. This type of risk assessment has his¬

torically been used in wildlife management and conservation programs but only recently has been available for

addressing the threat of climate change (Boyce 1992; Ruggiero et al. 1994; Faith & Walker 1996). Vulnerabil-

J. Bot. Res. Inst. Texas 8(1): 305 - 318.2014

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Journal of the Botanical Research Institute of Texas 8(1)

ity assessment can be especially useful to highlight new conservation targets and can be a useful way for states

to address climate change in their state-wide conservation plans or to coordinate broad-scale policy efforts that

span multiple agencies or political boundaries.

Several states such as New York (Schlesinger et al. 2011), Pennsylvania (Furedi et al. 2011), West Virginia,

Nevada, Illinois, and Florida (https://connect.natureserve.org /science/climate-change/ccvi) have implement¬

ed NatureServe’s Climate Change Vulnerability Index (CCVI; Young et al. 2010). One of the chief strengths of

the CCVI is that it is designed to be used in conjunction with NatureServe’s conservation status ranks (S-ranks;

Master et al. 2009), which are an existing global standard for assessing conservation status based on rarity,

trends, and threats. Another important strength lies in its explicit incorporation of scientific uncertainty into

the assessment: assessors are free to pick a range of values for each factor, and this uncertainty is quantified in

a Confidence score. Thus the CCVI considers how susceptible a species or a system is to climate change, while

directly acknowledging inherent uncertainties in future conditions and species responses. Species in the

northeast US are predicted to be exposed to increased temperatures and decreased moisture availability (Cli-

mAid www.nyserda.ny.gov/climaid). However, each individual species is expected to vary in its sensitivity to

these direct climate change impacts. Thus inherent species characteristics, such as dispersal ability, depen¬

dence upon or restriction to specific habitats, interspecific interactions, and genetic variation, will factor into

the species’ vulnerability to climate change.

NatureServe and various member programs, including the Pennsylvania Natural Heritage Program have

assigned conservation status ranks to each species. These ranks provide an estimate of extinction risk for the

plant species. The conservation status ranks, documented at the statewide geographic scale, are based on a one

to five scale, ranging from critically imperiled (SI) to demonstrably secure (S5). The CCVI was designed to be

used in conjunction with S-ranks; integrating climate change vulnerability assessments into existing lists of

at-risk species can be considered a more holistic approach to conservation concerns. This study objective was

to examine 35 plants of SI and S2 ranking to see if there was a relationship between conservation status ranks

and the CCVI index and then use a regression analysis to find the consistent risk factors across all taxonomic

groups examined.

METHODS

Development of a priority assessment list

Rhoads and Klein (1993) reported 3318 taxa of vascular plants for Pennsylvania, which included 2076 native

and 1242 introduced species. It was therefore necessary to develop a more refined list of priority species for the

climate change vulnerability assessment. Previous reports conducted by Byers and Norris (2011), Furedi et al.

(2011), and Schlesinger et al. (2011) used existing lists of species of conservation concern. Understanding the

need for future monitoring of imperiled plant species in danger of extirpation due to climate change, we se¬

lected plant species with a NatureServe conservation status ranking of SI (Critically Imperiled) and S2 (Imper¬

iled) that occur or have been known to occur near one central site in the state. We did not consider habitat

preferences, life forms, tolerance of disturbance, or species distribution patterns within the state of Pennsylva¬

nia. Following the criteria of imperiled conservation status and proximity to one central site, our set of plant

taxa could be efficiently monitored in the future for range expansion, range contraction, extinction or mainte¬

nance, possibly due to climate change. The funding for this project limited our scope to 35 species.

Plant taxa of SI and S2 ranking that have been known to occur within a 10-mile radius of Bedford, Penn¬

sylvania, based upon herbarium label data of specimens from the Carnegie Museum of Natural History, were

chosen for assessment. The city of Bedford is located in Bedford County in the south-central portion of the state

within the Appalachian Mountain section of the Ridge and Valley Physiographic Province. A series of ridges,

namely, Buffalo, Evitts, Tussey, and Polish Mountains and Warrior Ridge, run the length of this 10-mile radius

from southwest to northeast. The Raystown Branch of the Juniata River flows through the circle west to east

and its tributaries generally run in a northeasterly and southeasterly direction. Elevations in the 10-mile radius

range from 2000 to 2500 feet along the ridges to 900 to 1200 feet along the valley floors. At a larger scale, these

Morton and Schlesinger, Low genetic diversity and climate change vulnerability

307

physical features of ridgelines and stream valleys are prominent and extensive features that provide continu¬

ous habitat over many miles. Variations in aspect, slope, and elevation of the ridge and valley province combine

to create different habitats and microenvironments (Wagner 1998).

This 10-mile radius was also selected so that efficient monitoring program within a small geographic area

could be implemented at a future date. Changes in distribution of species and plant communities potentially

due to climate change may be most evident in populations of species occurring at various elevations and in a

variety of habitats from ridgetop to steep slope and to wetland and floodplain areas along streams. For many

imperiled species with relatively few populations overall, or occurring at the edge of their range, climate

change may lead either to their extirpation or expansion in Pennsylvania.

Thirty-five plant species met the above criteria of conservation status and location. The list includes 17

perennial herbs, 1 biennial, and 2 annual herbs, 2 perennial vines, 5 shrubs, 1 subshrub, and 7 graminoids (full

list of taxa at https://connect.natureserve.org sites/default/hles/documents/Pennsylvania-Plant-CCVI-2012.

pdf). Seven species are typically found in wetlands and eight species are typically deemed calciphiles with the

remaining taxa occurring in mixed habitats. Two are known to be parasites and at least five are known to de¬

pend upon a mycorrhizal relationship. The NatureServe Climate Change Vulnerability Index (CCVI) Release

2.0 was applied to each of the 35 plant species.

Examination of species vulnerability to climate change

We scored each of the criteria (described below) using information from peer-reviewed published papers and

reputable websites. Some criteria were more easily scored than others simply because of available information

and previous research. Accurate information on effective pollinators is nonexistent, or untested, for many of

the plant species, and dispersal mechanisms are more often hypothesized than experimentally proven. Fortu¬

nately the index is designed so that the accumulated knowledge of the plant species or genera allows for choos¬

ing a range of values.

Vulnerability to climate change was assessed by considering the two main components of vulnerability as

defined by Williams et al. (2008): the exposure of a species to climate change within a defined area combined

with the sensitivity of a species to climate change. Vulnerability assessment involves describing the severity

and scope of the exposure that species experience, and combining this with species’ sensitivity and capacity to

adapt to climate change. NatureServe’s newly developed Climate Change Vulnerability Index (Young et al.

2010) provides a means of dividing species into groupings of relative risk to climate change and of identifying

key factors causing species to be vulnerable. Used with standard conservation status assessments such as the

NatureServe G- and S-rank system, the Index can help land managers evaluate the likely effectiveness of alter¬

native strategies to promote adaptation of species to climate change as well as select key species to monitor. It

is designed to complement, and not duplicate, information contained in the NatureServe conservation status

ranks (Master et al. 2009), and may be used to update conservation status ranks to include the additional

stressor of climate change. Using regionally specific climate models, the index examines how the changed

climate will impact a species using factors known to be associated with vulnerability to climate change, includ¬

ing species-specific factors as well as external stressors imposed by human actions. Downscaled climate data

representing an ensemble of 16 global circulation models were downloaded from Climate Wizard (Girvetz et

al. 2009) and displayed in a GIS format. Climate data were available on a 4-km grid for historical data, and a

12-km grid for predicted future data. The overlap of changing climate with each species’ range was used to

calculate direct exposure.

The factors considered in evaluating species response might be divided into general categories including

direct exposure, indirect exposure, sensitivity, documented response, and modeled response. Detailed infor¬

mation including the scientific references used to develop each factor and the limitations of the methodology

are given in Young et al. (2010).

Brief definitions of the factors are given below and scored NatureServe’s Climate Change Vulnerability

Index Table (https://connect.natureserve.org/science/climate-change/ccvi).

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Journal of the Botanical Research Institute of Texas 8(1)

A—Direct Exposure

Temperature change is the predicted change in annual temperature by 2050, calculated over the range of the

species in Pennsylvania (ClimateWizard).

Moisture change is the predicted net change in moisture based on the Hamon AET:PET moisture Metric

by 2050, calculated over the range of the species in Pennsylvania (Kartesz 2011; WCRP, Maurer et al. 2007;

ClimateWizard) (Figs. 1 & 2).

B—Indirect Exposure

Bl: Exposure to Sea Level Rise

Weiss et al (2011) predict that only a very small portion of Pennsylvania will be subject to a sea level rise

of 0.5 to 1 meter; accordingly, less than 10% of the range of plant species eligible for this study could be

subject to sea level rise.

B2: Distribution relative to Barriers

Given the topography and geographical context of the state, most plant species in Pennsylvania will not

be subject to natural or anthropogenic barriers such as high mountain ranges, large expanses of water, or

intensive agricultural or urban development.

B3: Predicted impact of land-use changes resulting from human responses to climate change

Forestland in Pennsylvania totals 58% of land cover and is the dominant land class at 166 million acres.

This proportion remained stable from 1989 to 2004; the state’s forest loss (primarily due to residential and

industrial development) was offset by conversion of agricultural land to forest through natural succession

(Pennsylvania’s Forest 2004). Nowak & Walton (2005) predicted that if growth trends of the 1990s con¬

tinued through 2050, urban development could subsume an additional 15 million acres. Even if this loss

cannot be offset by agricultural land conversion, forestland should still be a primary land class. Forest

fragmentation and smaller patch sizes are prevalent in the southeast and west, but, in the north-central

region, forest patches are large and contain more interior forest habitat (Pennsylvania’s Forest 2004).

Twenty-nine percent of the forest land is owned by the state and federal US Forest Service. Govern¬

ment agencies are likely, as part of their management plans, to manage the forests for mitigation-related

carbon storage and carbon sequestration. The majority of forest-land, 71%, is privately owned (Pennsylva¬

nia’s Forest 2004).

C—Sensitivity

C-l: Dispersal—The ability of a species to shift locations in response to climate change (Vittoz & Engler

2007). For seed dispersal distances we used a typology based on dispersal modes and plant traits.

C-2: Predicted sensitivity to temperature and moisture changes: Species requiring specific precipitation

and temperature regimes may be less likely to find similar areas as climates.

C-2-a: Predicted sensitivity to changes in temperatures.

C-2-b: Predicted sensitivity to changes in precipitation, hydrology, or moisture regime.

C-2-c: Dependence on a specific disturbance regime likely to be impacted by climate change. Species

dependent on habitats such as prairies, or are maintained by regular disturbances such as fire or

flooding are vulnerable to climate change.

C-2-d: Dependence on ice, ice-edge, or snow-cover habitats. This factor is of minor significance de¬

pending on a species’ range in PA.

C-3: Physical Habitat Specificity. Species requiring specific soils (limestone outcrops) or physical features

such as caves, cliffs or sand dunes may become vulnerable to climate change.

C-4: Reliance on Interspecific Interactions. Species with tight relationships with other species may be

threatened by climate change.

C-4-a: Dependence on other species to generate habitat

C-4-c: Pollinator versatility

C-4-d: Dependence on other species for propagule dispersal

C-4-e: Forms part of an interspecific interaction not covered above

Morton and Schlesinger, Low genetic diversity and climate change vulnerability

309

Predicted Change in Moisture Availability in 2040-2069

in Pennsylvania

Predicted Change in Moisture

Availability 2040-2069

□ Predicted Future Change in Moisture based on median of

scenario of the Hamon AET:PET Moisutre Metric from Climate

■0.028 to -0.050 (Value 2) Wizard Analysis (negative values indicate net drying)

Fig. 1. Predicted future change in Pennsylvania moisture availability. Based on median of 16 global climate models and a middle-of-the-road emissions

scenario of the Hamon AETrPET moisture metric from Climate Wizard Analysis. Negative values indicate net drying.]

C-5: Genetic Factors—A species’ ability to evolve adaptations to environmental conditions brought about

by climate change is largely dependent on its existing genetic variation.

We used the internal transcribed spacer (ITS) gene because it contained the most data at this generic

level. The ITS region of nuclear ribosomal DNA (nrDNA) has proven to be a valuable resource for plant

systematics as a useful source of characters for phylogenetic studies in many angiosperm families (Bald¬

win et al. 1995). We scored the number of parsimony informative characters, a common measure of ge¬

netic variation. If the number of parsimony informative characters was under 150 then we coded the fac¬

tor as increasing vulnerability to climate change; between 151-250 was coded as somewhat increasing

vulnerability; between 251-350 we coded as neutral and over 351 was coded as somewhat decreasing

vulnerability.

C-6: Phenological response to changing seasonal temperature and precipitation regimes. Recent research

suggests that some phylogenetic groups are declining due to lack of response to changing annual tem¬

perature dynamics (e.g., earlier spring, longer growing season), including some temperate zone plants

that are not moving their flowering times.

D—Documented or Modeled Response to Climate Change

D-l: Documented responses to recent climate change: The results of published research may be available

that document changes within species that can be definitively linked to climate change.

D-2: Modeled future change in range or population size: The change in area of the predicted future range

relative to the current range is a useful indicator of vulnerability to climate change.

D-3: Overlap of modeled future range with current range: The results of future distribution models can be

compared to current range maps to address potential overlap.

D-4: Occurrences of protected areas in modeled future distribution: The results of future distribution

310

Journal of the Botanical Research Institute of Texas 8(1)

Predicted Change in Temperature at 2050

in Pennsylvania

Predicted Change in Temperature in 2050

^ > 5.1 to 5.5 deg F.

4.5 to 5.0 deg F.

Predicted Future Change based on median of 16

Global Climate Models and a middle of the road

emissions scenario from Climate Wizard Analysis

Fig. 2. Predicted future temperature change in Pennsylvania. Based on median of 16 global climate models and a middle-of-the-road emissions scenario

from Climate Wizard Analysis.]

models can be compared to present protected areas to see if future ranges may fall entirely outside of pro¬

tected areas and therefore compromise their long-term viability.

Compile and analyze results: Climate Change Vulnerability Index results were compiled and analyzed in order

to (a) highlight those species most (and least) vulnerable to climate change, (b) identify and rank causative

factors, and (c) identify geographic areas or habitat types at high risk.

Regression analysis

To determine the factors most important in assessing vulnerability, we followed the methodology used by

Schlesinger et al. (2011). We built classification trees using the Random Forests (Breiman 2001; Liaw & Wiener

2002) package in R (R Development Core Team 2011), a technique from the held of machine learning. The

Random Forests routine is to build thousands of classification and regression trees using bootstrap samples of

the data set and predictors. We limited our predictor variables to the exposure and sensitivity variables influ¬

encing vulnerability (i.e., omitting documented and modeled responses) and imputed (estimated) values re¬

corded as “Unknown,” as the routine does not accept missing data.

RESULTS AND DISCUSSION

Documented responses to climate change are incorporated into NatureServes Climate Change Vulnerability

Index Table. The output is one of five categories of vulnerability and one indicating lack of evidence. Defini¬

tions, and the abbreviations that are used throughout this document, follow Young et al. (2010) and are pre¬

sented in Table 1.

Plant species assessed and factors affecting vulnerability

The 35 plants included in this assessment ranged from highly vulnerable to not vulnerable to climate change.

Morton and Schlesinger, Low genetic diversity and climate change vulnerability

311

Table 1. Five Vulnerability Index scores and eight individual Risk Factor scores.

Vulnerability Index

EV (Extremely Vulnerable)

Abundance and/or range extent within geographical area assessed extremely likely to

substantially decrease or disappear by 2050; HV Highly Vulnerable Abundance and/or

range extent within geographical area assessed likely to to decrease significantly by 2050.

MV (Moderately Vulnerable)

Abundance and/or range extent within geographical area assessed likely to decrease by 2050.

PS (Not Vulnerable/Presumed Stable)

Available evidence does not suggest that abundance and/or range extent within

geographical area assessed will change (increase/decrease) substantially by 2050. Actual

range boundaries may change.

IL (Not Vulnerable/Increase Likely)

Available evidence suggests that abundance and/or range extent within geographical area

assessed is likely to increase by 2050.

Risk Factor:

Greatly Increase Vulnerability

Inc

Increase Vulnerability

Somewhat Increase Vulnerability

Neutral

Somewhat Decrease Vulnerability

Decrease Vulnerability

N/A

Not Applicable

Unknown

Vulnerability to climate change was due to a combination of multiple risk factors. Influential risk factors ap¬

pear to be limited dispersal capabilities, decreased genetic variation, and dependence on a specific hydrological

or moisture regime. Plants assessed as stable were often habitat generalists and less dependent upon a wetland

habitat, were able to disperse longer distances, and were genetically more diverse. For example, Astragalus ca¬

nadensis, a plant somewhat dependent upon marshy ground or moist prairie, rarely disperses more than 10

meters when its exploding fruit ejects its seeds (Gleason & Cronquist 1991 ). ITS phylogenetic inference re¬

vealed a low number of parsimony information characters and thus low genetic variation. Its vulnerability in¬

dex score was Moderately Vulnerable. On the other hand, Amelanchier sanguinea, a shrub found in an assort¬

ment of habitats throughout its range, produces sweet and juicy fruits highly palatable to birds (Gleason &

Cronquist 1991 and PNHP Factsheet). ITS phylogenetic inference revealed a higher number of parsimony in¬

formation characters and thus higher genetic variation. Its vulnerability index score was Not Vulnerable.

Fourteen of the 35 (40%) species assessed were determined to be vulnerable (HV or MV) to climate

change. None of the species were rated as “Extremely Vulnerable” and only two as “Highly Vulnerable.” Twen¬

ty species (57%) were rated as “Presumed Stable” and only one as “Increase Likely.” Both of the “Highly Vulner¬

able” species are poor dispersers.

All 35 species assessed were ranked SI, S1S2, or S2. The vulnerability statuses were distributed through¬

out these conservation status ranks. Fifteen of the 35 species examined were SI ranked (imperiled species), 8

of these taxa were presumed stable. SI species did not appear to be more vulnerable to climate change than did

the S2 species (Fig. 3).

A review of global conservations ranks found only four taxa were at some risk to global extinction. These

were ranked G2 (globally imperiled - at high risk of extinction due to very restricted range, few populations,

etc.) or G3 (vulnerable - at moderate risk of extinction due to restricted range, relatively few populations, etc.).

Their vulnerability status was Presumed Stable. The remaining 31 species were ranked G4 (apparently secure)

or G5 (secure), and the vulnerability statuses included Increase Likely, Presumed Stable, Moderately and

Highly Vulnerable (Fig. 4).

An important result of this assessment is that we cannot predict the climate change vulnerability of a spe¬

cies based on its current Conservation Status Rank (G=global or S=state rank). Rare species may not always be

vulnerable to climate change and common species are not necessarily resilient. Each species will behave and

respond according to its unique life history characteristics, habitat requirements, and distribution. The impli-

312

Journal of the Botanical Research Institute of Texas 8(1)

Conservation status rank for Pennsylvania

Fig. 3. Percent of species within state conservation status ranks in each vulnerability category (SI n=15, S1S2 n=3 and S2 n=17). PS= Presumed

Stable; IL= Increase Likely; HV= Highly Vulnerable; MV= Moderately Vulnerable. Adapted from Byers and Norris (2011) and Schlesinger etal. (2011).

NatureServe's Global Conservation status rank

■ PS

■ IL

■ HV

■ MV

Fig. 4. Percent of species within global conservation status ranks in each vulnerability category (G2 n=2; G3 n=2; G4 n=25; and G5 n=6). PS= Presumed

Stable; IL= Increase Likely; HV= Highly Vulnerable; MV= Moderately Vulnerable. Adapted from Byers and Norris (2011) and Schlesinger etal. (2011).

Morton and Schlesinger, Low genetic diversity and climate change vulnerability

313

100%

~ 90%

80%

5 70%

8 60%

t 50%

40%

S 30%

8 20 %

* 10%

Center of range East/west edge Northern edge Southern edge

of range of range of range

Relation of species' range to Pennsylvania

■ PS

■ IL

■ HV

■ MV

Fig. 5. Percent of species in each vulnerability category categorized according to the position of their range relative to Pennsylvania (Center n=4; East/

West n=1 8; North n=11; and Southern n=2). PS= Presumed Stable; IL= Increase Likely; HV= Highly Vulnerable; MV= Moderately Vulnerable. Adapted

from Byers and Norris (2011) and Schlesinger et al. (2011).

cations of this are important to rare plant conservation and management strategies, since climate change may

necessitate the reassessment of conservation status. We need to examine and re-align our ranking process to

best conserve species and habitats with the resources available.

Both species at the southern edge of their range were assessed as Moderately Vulnerable, and therefore

possibly disappearing from the state (Fig. 5), whereas the Center of the range, East/West edge of the range and

Northern edge of the range contained 50% or greater of the Presumed Stable (PS) taxa and were assessed as not

highly vulnerable to climate change.

Dispersal scores ranged from somewhat decrease vulnerability to greatly increase vulnerability (Fig. 6)

and did have an influence. Twelve of the 25 species were scored as GI, Inc, and SI indicating dispersal limita¬

tions and were assessed as Highly Vulnerable (HV) or Moderately Vulnerable (MV). The taxa with overall

neutral and somewhat decreased vulnerable scores consisted of mostly Presumed Stable (PS) taxa (7 of the 10

species) indicating no dispersal or movement influence. These results also agree with the first Pennsylvania

study (Furedi et al. 2011), West Virginia (Byers & Norris 2011) and New York (Schlesinger et al. 2011) climate

change vulnerability assessment reports. These reports indicated that the top risk factors, based upon both

floral and fauna assessments, appeared to be related to dispersal and movement mechanisms. Plants that lack

the specialized structures for dispersal by wind, or lack attractive coloration for animal dispersal, have limited

potential for long-distance dispersal.

Measured genetic variation scores were an influence (Fig. 7). Nine of the seventeen species in the Inc and

SI categories indicated low genetic diversity, assessed as Highly Vulnerable (HV) or Moderately Vulnerable

(MV). Taxa with the overall neutral and somewhat decreased vulnerable scores consisted of mostly Presumed

Stable (PS) taxa (11 out of 16 species) and were assessed as not vulnerable to climate change. This study used

the number of parsimony informative characters of the ITS gene as an indicator of genetic diversity. ITS (inter¬

nal transcribed spacer) region was selected because it is typically used at the species level. Although additional

assessment needs to be done using this technique, it was in agreement with other factors used in this study and

314

Journal of the Botanical Research Institute ofTexas 8(1)

100%

90%

$ 80%

e 70%

'—.

J§ 60%

| 50%

•J 40%

C 30%

& 20%

10 %

Dispersal and movement factor score

■ HV

■ MV

■ PS

■ IL

Fig. 6. Percent of species in each vulnerability category categorized according to the dispersal and movement factor relative to Pennsylvania (GI=Greatly

Increase Vulnerability n=4; lnc=lncrease Vulnerability, n=16; SI=Somewhat Increase Vulnerability, n=5; N=Neutral, n= 3; and SD=Somewhat Decrease

Vulnerability, n=7). PS= Presumed Stable; IL= Increase Likely; HV= Highly Vulnerable; MV= Moderately Vulnerable. Adapted from Byers and Norris

(2011) and Schlesinger et al. (2011).

Measured genetic variation factor score

Fig. 7. Percent of species in each vulnerability category categorized according to the measured genetic variation factor relative to Pennsylvania

(lnc=lncrease, n=10; SI=Somewhat Increase Vulnerability, n=7; N=Neutral, n= 9; SD=Somewhat Decease Vulnerability, n=7; and U=Unknown,

n=2). PS= Presumed Stable; IL= Increase Likely; HV= Highly Vulnerable; MV= Moderately Vulnerable. Adapted from Byers and Norris (2011) and

Schlesinger etal. (2011).

Morton and Schlesinger, Low genetic diversity and climate change vulnerability

315

with the results from West Virginia and New York. The previous Pennsylvania study did not contain enough

genetic data to be significant. As stated, these results also agree with the West Virginia (Byers & Norris 2011)

and the New York (Schlesinger et al. 2011) state climate change vulnerability assessment reports. Both of these

reports indicated that genetic factors predisposing species to potential climate change effects were easily the

most important factor. Species with reduced genetic variation are less likely to be able to respond to environ¬

mental change (e.g., Aitken et al. 2008). In addition, plants with poor dispersal strategies will eventually be

genetically bottlenecked and therefore have low genetic diversity (http://www.nature.com/scitable/dehnition/

population-bottleneck-300).

Regression scores

The most important factors, as indicated by the R 2 values, driving a species’ vulnerability status were as follows:

1. Dispersal and movement;

2. Predicted sensitivity to changes in precipitation, hydrology, or moisture regime;

3. Predicted sensitivity to changes in temperature;

4. Physiological hydrological niche (a dependence on a narrow precipitation/hydrologic regime), and

5. Genetic factors.

These top five factors are probably representative of the most consistent risk factors across all plant taxonomic

groups in the state.

The random forest regression analysis supports the visual assessments of the graphs. Dispersal limita¬

tions, the ability of a species to shift locations in response to climate change, were one of the most important

factors in our assessment. The next most important factors were temperature and moisture. Those species re¬

quiring moist microhabitats will experience stress if these habitats dry up. Finally genetic variation, which

affects a species’ ability to adapt to environmental conditions, was among the top five factors (Fig. 8).

Comparisons to other states

Three states in the Northeast—Pennsylvania, New York and West Virginia—have recently completed CCVI

analyses, but only Pennsylvania and West Virginia included plants. Our study contained 2 out of 35 taxa as

highly vulnerable whereas the first Pennsylvania analysis contained 24 out of 40 taxa as extremely (EV) to

highly vulnerable (HV) and West Virginia contained 7 out of 33 plant taxa as (EV or HV). The proportion of

species assessed as vulnerable depends greatly on the species selected for analysis, as conducting the analysis

on all species is not possible given constraints of funding, time and available information. Other studies aimed

to select species they thought might prove vulnerable to climate change based on habitat; however, we selected

taxa based on their current Conservation Status State Rank (imperiled or endangered species).

Our results agreed with those of West Virginia while there was no overlap between our study and the first

Pennsylvania analysis. Our study and the West Virginia study assessed 2 species in common. The final index

values either matched exactly (Pycnanthemeum torreyi ) or were off by one step (Paxistima canbyi, Presumed

Stable versus Moderately Vulnerable). These differences in index values might result from true differences in

vulnerability among states or differences in interpretation of data; a full analysis of these differences is beyond

the scope of this study.

Management and monitoring recommendation

A complete discussion of management, monitoring and restoration of habitat connectivity is past the capacity

of this paper. However, we can provide some recommendations that are applicable here based on our results.

The fact that the species assessed as Highly Vulnerable in our analyses are associated with the identifica¬

tion of barriers to dispersal as an important component of our vulnerability and regression scores. Maintaining

and restoring habitat connectivity is crucial for many ecological processes, including dispersal, gene flow, and

movement in response to climate change (Mawdsley et al. 2009; Heller & Zavaleta 2009; Byers & Norris 2011;

McRae et al. 2012). This is especially true for vulnerable species restricted to certain habitats. Another valuable

outcome of this procedure is it allows biologists to ascertain which life history traits of a particular species in-

316

Journal of the Botanical Research Institute of Texas 8(1)

Dispersal ability

Exposure to greater change in moisture

Exposure to >5.1°F increase

Exposure to 4.5°-5.0°F increase

Exposure to lower change in moisture

Physiological hydrological niche

Genetic variation

Reliance on pollinators

Position within range

Reliance on physical habitat

G-rank

Reliance on species interactions

Reliance on other species for habitat

Sensitivity to climate-change mitigation

S-rank

Reliance on other species for dispersal

Historical hydrological niche

Phenological response

Relative importance

Fig. 8. Variable importance (decrease in node impurity according to the Gini index) from the random forest analysis of Climate Change Vulnerability

Index scores of 35 Pennsylvania plants.

dicate propensity of that species to be vulnerable to climate change and further highlights other factors that

might pose more immediate threats to certain imperiled species (such as dispersal limitations and low genetic

diversity). Species with good dispersal mechanisms can redistribute themselves, but the key to successful

movement and migration is the presence of contiguous habitats that species are able to colonize or move

across. Protecting large blocks of unfragmented habitats and using linkages and corridors to enhance connec¬

tivity will facilitate this colonization or movement, but this is only one solution. Developing methods to iden¬

tify barriers whose removal would significantly improve connectivity, such as least-cost and simulation model¬

ing, can be cost effective and broaden alternatives available to connectivity conservation. This network of in¬

tact habitats should represent a full range of ecosystems to sustain biodiversity and genetic diversity. Key eco¬

logical processes such as pollination, seed dispersal, nutrient cycling, and natural disturbance cycles will be

maintained under this environment.

Long-term monitoring using multiple taxa and habitats will help 1) test hypotheses about vulnerability;

2) detect changes in species; 3) test hypotheses about consistent risk factors; 4) identify barriers to improve

connectivity; and 5) help examine alternative methods for least-cost simulation modeling to maintain connec¬

tivity. Currently, these data are not readily available and it is imperative for governmental and non-governmen¬

tal organization to have these data to make the most informed conservation and management decisions. The

success in combating these environment changes will only be achievable through an unprecedented level of

Morton and Schlesinger, Low genetic diversity and climate change vulnerability

317

collaboration and cooperation between wildlife managers, other organizations, scientists, and the public.

Building science-driven strategies that maximize the use of scarce resources will be necessary so legislative

support and policy changes can be implemented. In order to facilitate long-term monitoring of multiple taxa

we have used herbarium data to designate a 10-mile radius in Bedford County including most of the plants we

analyzed in this study. Additional sites could be established using the same techniques (i.e. a small number of

species within a defined radius), so a manageable monitoring program could be funded in the future allowing

for improved collaboration and cooperation between organizations which reduce the cost of research yet ob¬

tain a wealth of information so good conservation decisions are made.

Proposed taxa for more detailed monitoring programs

Broad-scale, long-term monitoring of taxa will help test hypotheses about vulnerability, which are essentially

what the CCVI provides. Monitoring can detect unanticipated changes in populations, can identify particular

stressors, and can reveal range shifts and changes in phenology. Long-term monitoring has already revealed

shifts that have been vital in demonstrating responses to climate change (Parmesan et al. 1999; Hitch & Leberg

2007; Zuckerberg et al. 2010). There is a pressing need to establish a solid baseline of data that will allow us to

detect these changes in Pennsylvania and make the most informed conservation and management decisions.

As new factors affect wildlife and habitats, such as changes in phenology and the effects on pollinators and the

increase in invasive species, managers will need to monitor these changes and incorporate them into new ac¬

tion strategies.

ACKNOWLEDGMENTS

Our thanks go to the Wild Resource Conservation Fund of Pennsylvania Department of Conservation and

Natural Resources for partial funding of this research. We would also like to thank J. Corser, K. Perkins, E.

White, and T. Howard for their contributions to this document. Troy Weldy and an anonymous reviewer pro¬

vided useful and helpful review comments.

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Change Biol. 15:1866-1883.

THE VASCULAR FLORA OF FORT SUMTER AND FORT MOULTRIE,

SOUTH CAROLINA, U.S.A.

Richard Stalter, Brent A. Berger 1 Eric E. Lamont John Nelson

Department of Biological Sciences The New York Botanical Garden Department of Biological Sciences

St. John's University 2900 Southern Blvd. University of South Carolina

Jamaica, New York 11439, U.S.A. Bronx, New York 10458, U.S.A. Columbia, South Carolina 29208, U.S.A.

korr. author: brent.a.berger@gmail.com

ABSTRACT

The vascular flora of Fort Moultrie and Fort Sumter, Charleston County, South Carolina, were sampled in 1990, one growing season after

Hurricane Hugo, and during the 2008-10 growing seasons. The flora at Fort Moultrie consisted of 303 taxa, while 77 taxa were identified at

Fort Sumter. The Asteraceae and Poaceae were the largest families in the flora at both sites. Cyperus (9 taxa at Fort Moultrie, 3 taxa at Fort

Sumter) was the largest genus in the flora. The annotated checklist includes the locality and habitat in which each species occurs, the years

the taxa were observed, frequency of occurrence, and pertinent synonyms.

RESUMEN

Se muestreo la flora vascular de Fort Moultrie y Fort Sumter, Charleston County, South Carolina, en 1990, una estacion despues del huracan

Hugo, y durante las estaciones de 2008-10. Fa flora de Fort Moultrie consistio en 303 taxa, mientras que se identificaron 77 taxa en Fort

Sumter. Fas Asteraceae y Poaceae fueron las mayores familias de la flora en ambos lugares. Cyperus (9 taxa en Fort Moultrie, 3 taxa en Fort

Sumter) fue el genero mas grande de la flora. El catalogo anotado incluye la localidad y el habitat de cada especie, los anos en que los taxa se

observaron, la frecuencia de ocurrencia, y los sinonimos pertinentes.

INTRODUCTION

Fort Sumter National Monument in Charleston County, South Carolina, is comprised of Fort Sumter National

Monument (32.75N 79.87W), an island in Charleston Harbor, and Fort Moultrie (32.76N 79.86W) on the

southern rim of Sullivan’s Island north of the harbor (Fig. 1). Two vegetation studies of the sites have occurred

in the last twenty-five years. Stalter and Lamont (1993) sampled the vascular plant species of Fort Moultrie and

Fort Sumter at six-week intervals during the growing season after Hurricane Hugo from March-October 1990.

The combined flora of both sites (ca. 27 ha at Fort Moultrie and ca. 1 ha at Fort Sumter) in 1990 consisted of 63

families, 161 genera and 287 species; 69 species occurred at Fort Sumter, while 218 species occurred at Fort

Moultrie. Subsequently, Schmidt (2004) produced an unvouchered, unpubished list of 265 vascular plant spe¬

cies at Fort Moultrie. One-hundred fifty-four of the taxa (58%) reported by Schmidt were also found by Stalter

and Lamont (1993), while 52 taxa (19%) were not.

METHODS

Four plant communities were documented at Fort Moultrie and Fort Sumter (Stalter & Lamont 1993). Three

occur at both sites (salt marsh, dune and ruderal), while one is exclusive to Fort Moultrie (maritime woodland).

The maritime woodland community was not included in the present study as it was destroyed in Hurricane

Hugo. Remnants of the flora in this community appeared in the earlier paper by Stalter and Lamont (1993).

The vascular plant species at Fort Moultrie and Fort Sumter were sampled at six-week intervals from

April-October 2008-2010. The data was analyzed with previously published data from March-October 1990

(Stalter & Lamont 1993), as well as unpublished data from Schmidt (2004), to better characterize the complete

flora at the two sites. The most intensive collecting was during the 1990 growing season, one year after Hurri¬

cane Hugo. Objectives for each trip included the collection of voucher material and accumulation of informa¬

tion on habitat preference and abundance for ach species. One complete set of vouchers was deposited at the A.

C. Moore Herbarium at the University of South Carolina (USCH). Duplicate vouchers were retained in James

J. Bot. Res. Inst. Texas 8(1): 319 - 331.2014

320

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 1 . General map of South Carolina and the Charleston Harbor Area. The locations of Fort Moultrie (1) and Fort Sumter (2) are designated by gray stars.

Montgomery’s (ferns) private herbarium while and Gordon Tucker’s (Cyperaceae) material is housed at East¬

ern Illinois University Herbarium (EIU). Accession numbers for taxa collected in the 2008-2010 study and

taxa reported exclusively by Schmidt (2004) but not by us are included in the appendix. Because many of

Schmidt’s exclusive taxa from 2004 are not accessioned, we designate these taxa as “sight record.”

The species checklist at both forts contains an inventory of the vascular plant species that reproduce

spontaneously and persist for one growing season, including native species, naturalized and adventive weeds,

and escapes from cultivation. Plant species collected or identified exclusively by Schmidt (2004) are annotated

as such in the checklist. All non-native species are designated using an asterisk (*). Native and non-native sta¬

tus follows that of Gleason and Cronquist (1991) and Wunderlin (1998). The checklist is divided into four cat¬

egories: Polypodiophyta, Pinophyta, Magnoliophyta: Magnoliopsida, and Magnoliophyta: Liliopsida. Radford

et al. (1968) was used as the initial guide for identification, but the names of all taxa were verified using the

working draft from 30 November 2012 of the Flora of the Southern and Mid-Atlantic States (Weakley 2012). If

taxonomy or synonymy was in question, the Integrated Taxonomic Information System (ITIS) and the Inter¬

national Plant Names Index (IPNI) were used for verification. Annotated names reflect currently accepted no¬

menclature from these sources.

RESULTS

Four plant communities as described by Stalter and Lamont (1993) occur at Fort Moultrie: 1) a large salt marsh

community dominated by Spartina alterniflora Loisel; 2) a dune community dominated by graminoids and

forbs; 3) a ruderal community consisting of lawns, fields, roadsides and thickets; and 4) a maritime woodland.

All but the maritime woodland occur at Fort Sumter where a large ruderal, salt marsh and small dune com¬

munity are present. The ruderal community at both forts supports the greatest number of taxa.

Salt Marsh Community .—The salt marsh community is best developed at Fort Moultrie near the dock.

Common species there and at Fort Sumter include Spartina alterniflora, S. patens (Ait.) Muhl., Salicornia de-

pressa Stanl., Sesuvium portulacastrum (F.) F., Borrichia frutescens (F.) DC. and Sporobolus virginicus (F.)

Kunth. The distribution of these taxa along an elevation (tidal) and salinity gradient has been well documented

Stalter et al., Flora of Fort Sumter and Fort Moultrie

321

previously (Stalter 1973; Baden et al. 1973; Stalter and Lamont, 1993). Spartina alterniflora, the most flood toler¬

ant species, occupies the greatest area of salt marsh at both sites.

Dune Community .— Uniola paniculata L. is the dominant species of the primary dune system at Fort

Moultrie. At both Fort Moultrie and Fort Sumter, the state rare Oenothera drummondii Hook, thrives and is es¬

pecially conspicuous when in flower. Additional members of the dune community are Spartina patens (Ait.)

Muhl., Cakile edentula (Bigelow) Hook., Salsola kali L., Yucca aloifolia L., Strophostyles helvola (L.) Ell., Croton

punctatus Jacq., Euphorbia polygonifolia L., Xanthium strumarium L. and Cenchrus tribuloides L. The dune com¬

munity is poorly developed at Fort Sumter, occupying a small area of land outside the western portion of the

fort. At Fort Moultrie an additional species, Ipomoea imperati (Vahl) Griseb., thrives on and in front of the pri¬

mary dune.

Ruderal Community .—The ruderal community was the largest assemblage at Fort Sumter, composed

primarily by lawn inside and outside of the park. The lawn was trimmed several times a month from March-

November, making it difficult for annuals to become established. Common species included Stenotaphrum se-

cundatum (Walter) Kuntze, Hydrocotyle umbellata L., Medicago arabica (L.) Huds., Sisyrinchium rosulatum

Bickn., Sonchus asper (L.) Hill., Oenothera humifusa Nutt, and O. laciniata Hill. Arenaria serpyllifolia L. grows in

brick crevices within both forts along with Centella asiatica (L.) Urb., Polycarpon tetraphyllum (L.) L., Pteris vit-

tata L. and Sagina decumbens (Ell.) Torrey & A. Gray. The weedy border of the visitor center at Fort Moultrie

harbors Vicia sativa ssp. nigra (L.) Ehrh., V hirsuta (L.) Gray, Medicago arabica (L.) Huds., M. lupulina L., Hiera-

cium gronovii L., and Sonchus asper (L.) Hill. A small population of Salvia lyrata L. thrives on the southern

border of Fort Moultrie’s Visitor Center. Additional taxa around Fort Moultrie include Sporobolus indicus (L.)

Br., Lantana camara L., Bromus catharticus Vahl, Oenothera speciosa Nutt., O.fruticosa L., and O. drummondii

Hook. Oenothera speciosa Nutt, was abundant in the unmowed fields at Fort Moultrie in 1990, but was rare in

the often-mowed fields at the fort in 2008-2010. A Florida species, Asparagus densiflorus (Kunth) Jessop., was

observed in a thicket bordering the road on the southeastern side of the fort in 2009 and had spread to addi¬

tional sections of the park by 2011. The species has been previously reported in Beaufort County, South Caro¬

lina (Damrel 2009, pers. comm.). Ranunculus muricatus L. was common at Fort Moultrie in the northern por¬

tion of the lawn, but not observed in 1990. One additional southern taxa observed by us for the first time at Fort

Moultrie was Tillandsia recurvata (L.) L.

Maritime Woodland .—The maritime woodland community occurring exclusively at Fort Moultrie is

comprised of Quercus virginiana Mill., Sideroxylon tenax L., Celtis laevigata Willd., Prunus caroliniana (Mill.)

Ait. and Juglans cinerea L. Additional woody species are Prunus angustifolia Marshall, Zanthoxylum clava-hercu-

lus L., Campsis radicans (L.) Seem, Ilex vomitoria Ait., Ligustrum sinense Lour., and Wisteria sinensis (Sims) DC.

Herbaceous plants along the woodland edges include Canna x generalis L.H. Bailey, Verbena brasiliensis Veil.,

Paspalumjloridanum Michx., Coreopsis basalis (A. Dietr.) S.F. Blake, Chaerophyllum tainturieri Hook., and Sor¬

ghum halepense (L.) Pers.

The vascular flora at Fort Moultrie consists of 303 taxa in 72 families representing 198 genera (see Table

1). The Asteraceae (51 taxa) and Poaceae (40 taxa) were the largest families in the flora comprising 30% of the

taxa sampled. The third largest family was the Cyperaceae with 16 taxa. Cyperus, with 9 taxa, was the largest

genus in the flora. An annotated list of taxa at Fort Moultrie is provided below, while a summary of the flora is

listed in Table 1.

Seventy-five taxa at Moultrie observed by us in 1990 were not reobserved in 2008-2010, including 16

monocots and 59 dicots. Eleven new monocots and 10 new dicots were collected at Fort Moultrie for the first

time. Of the 303 taxa observed in our twenty-year study, 96 taxa (32%) either colonized or were extirpated at

the site.

The vascular flora at Fort Sumter consisted of 77 taxa in 24 families representing 63 genera (see Table 2).

The Poaceae (17 taxa) and Asteraceae (15 taxa) were the largest families comprising 42% of the flora. Cyperus,

with 3 taxa, was the largest genus in the flora. No other genus had more than two taxa. Taxa at Fort Sumter are

listed below in the annotated list, while a summary is provided in Table 2. We observed six new taxa at Fort

322

Journal of the Botanical Research Institute of Texas 8(1)

Table 1. A summary of the vascular flora of Fort Moultrie, South Carolina.

Ferns

Gymnosperms

Dicots

Monocots

Total

Family

Genus

145

198

Native species

159

223 (73.6%)

Non-native species

80 (26.4%)

Total species

222

303

Table 2. A summary of the vascular flora of Fort Sumter, South Carolina.

Ferns

Gymnosperms

Dicots

Monocots

Total

Family

Genus

Native species

51 (65.4%)

Non-native species

27 (34.6%)

Total species

Sumter during the 2008-2010 collecting season, including three new species of Cyperus. During the twenty-

year study, 17 taxa (27%) were identified as colonists or were extirpated. Non-native taxa composed 34.6% of

the flora occurring on land within Fort Sumter. The combined vascular flora at Fort Moultrie and Fort Sumter

one year after Hurrcane Hugo (1990) consisted of 63 families, 161 genera, and 287 species. Fort Moultrie, the

larger of the two sites, supported 218 vascular plant species including one spore plant, three gymnosperms,

185 dicots and 44 monocots. Sixteen fewer taxa were collected at Fort Moultrie during the three-year collecting

period from 2008-2010. Sixty-nine species occurred at Fort Sumter in 1990, while 66 species were observed

during the collecting period from 2008-2010.

DISCUSSION

Four more monocots were found at Fort Moultrie during the 1990 collecting period than in the three-year col¬

lecting period from 2008-2010. The higher number of monocots, immediately after Hurricane Hugo, was most

likely due to the lack of mowing, allowing more successional forbs and grasses to thrive.

Two tropical/subtropical species have been identified at Fort Moultrie —Tillandsia recurvata and Aspara¬

gus densiflorus. Tillandsia recurvata was observed at the Visitor Center growing on live oak ( Quercus virginiana

Mill.) near the NPS dock. This tree was planted by the NPS and probably originated as landscape material from

Florida although no documentation is available. Alan Weakley and Dixie Damrel (pers. comm. 2009) have

reported T. recurvata as introduced in coastal South Carolina in Beaufort, Jasper, Georgetown, and Charleston

counties, brought to South Carolina on nursery stock shipped from Florida, and Carter et al. (2009) report

from it similarly in Georgia.

Asparagus densiflorus has also been reported in Beaufort Co., South Carolina, by Daniel C. Payne (Dam¬

rel, pers. comm.). Material was collected by Payne at 100 Laurens St., Beaufort, South Carolina, on 6 January

2005, the Shell Convience Store, Lady’s Island Shopping Center, Hwy 21, Lady’s Island, South Carolina, on 6

January 2006, and as an epiphyte at the entrance to Hampton Plantation Bluffton, South Carolina, 23 Decem¬

ber 2006. Although Wunderlin (1998) reported that A. densiflorus is common in central Florida and rare north¬

ward, the species can survive the cold temperatures of coastal South Carolina winters, as demonstrated in

2008-2009 when temperatures dropped lower than -6°C.

In summary, the number of taxa at both Fort Moultrie and Fort Sumter decreased from 1990 to 2008-

2010. Fort Moultrie lost 75 of its original taxa while gaining 21 new colonists. Eleven taxa present at Fort Sum¬

ter in 1990 were not observed in the present study; 6 new taxa were observed including three species of Cype¬

rus. Schmidt (2004) reported several taxa at Fort Moultrie that were not observed by us during the 1990 and

Stalter et al., Flora of Fort Sumter and Fort Moultrie

323

2008-2010 collecting periods. Since taxa invade and become extirpated at a site from year to year the differ¬

ences in taxa numbers was expected. Disturbance from Hurrricane Hugo, during September 1989, may have

created new open habitats for taxa to colonize, and the absence of mowing the large held between the dunes

and Fort Moultire may account for the higher number of taxa identified during the 1990 collecting season.

ANNOTATED CHECKTIST OF SPECIES

The vascular plant taxa have been arranged according to the following categories: vascular cryptograms, gym-

nosperms, dicots, and monocots. Within each category, families and lower taxa are arranged alphabetically.

Asterisks (*) are used to designate non-native taxa. Nomenclature at lower taxonomic levels follows the work¬

ing draft from 30 November 2012 of the Flora of the Southern and Mid-Atlantic States (Weakley 2012). When

taxonomy or synonymy was questioned, ITIS and IPNI were used for verification.

FORT MOULTRIE

POLYPODIOPHYTA

Polypodiaceae

Pleopeltis polypodioides ssp. michauxiana (Weath.) E.G. Andrews

& Windham [=Polypodium polypodioides (L.) Watt var. mich-

auxianum Weatherby]. Ruderal—masonry walls: rare. Stalter,

29 Mar 1990.

Pteridaceae

*Pteris vittata L. Ruderal—masonry walls: infreq. Stalter 19479,

2008/10.

PINOPHYTA

Cupressaceae

Juniperus virginiono var. silicicolo (Small) E Murray. [=J. silicicolo

(Small) Bailey]. Dunes—coastal foredunes and swales: infreq.

Stalter, 29 Mar 1990. Stalter 19529, 2008/10.

Taxodium distichum (L.) Rich. Maritime Woodland—ephemeral

ponds: rare. Stalter, 27 Aug 1990. Stalter 19771,2008/10.

Pinaceae

Pinus toedo L. Maritime Woodland: infreq. Stalter, 29 Mar 1990.

Stalter 19530, 2008/10.

MAGNOLIOPHYTA MAGNOLIOPSIDA (DICOTS)

Aizoaceae

Sesuviumportulacastrum (L.) L. Salt Marsh: freq. Stalter, 27 Aug 1990.

freq. Stalter 19772, 2008/10.

Amaranthaceae (including Chenopodiaceae)

*Altemanthera philoxeroides (Mart.) Griseb. Ruderal—roadside

ditches and moist waste areas: rare. Stalter 19803,2008/10.

Amaranthus cannabinus (L.) Sauer. Salt Marsh: infreq. Stalter, 27

Aug 1990.

Atriplex mucronata Raf. [=A. arenaria Nutt.]. Dunes: infreq. Stalter,

26 Oct 1990. Stalter 19805, 2008/10.

Atriplex prostrata Boucher ex DC. [=A hastata L.]. Salt Marsh and

Dunes: freq. Stalter, 27 Aug 1990. Stalter 19804/20254,2008/10.

*Chenopodium album L. Ruderal—roadsides and waste places: freq.

Stalter 19806, 2008/10.

*Dysphaniaambrosioides (L.) Mosyakin &Clemants. [ =Chenopodium

ambrosioides L.]. Ruderal—swales, fields and disturbed sands:

freq. Stalter, 27 Aug 1990. Stalter 19573, 2008/10.

Iresine rhizomatosa Standi. Salt Marsh—edges of tidal marshes:

infreq. Stalter, 26 Oct 1990.

Salicornia virginica L. Salt Marsh: freq. Stalter 19534, 2008/10.

Salicornia maritima Wolff & Jefferies (synonymy incomplete). Salt

Marsh: infreq. Stalter 19807, 2008/10.

Salsola kali L. Dunes—upper beach: freq. Stalter, 26 Oct 1990.

Stalter 19937, 2008/10.

Suaeda linearis (Ell.) Moq. Salt Marsh: freq. Stalter, 26 Oct 1990.

Anacardiaceae

Rhus copallinum L. Maritime Woodland—open, dry, waste places

and swales: freq. Stalter, 27 Aug 1990. Stalter 19532, 2008/10.

Toxicodendron radicans var. radicans (L.) Kuntze [=Rhus radicans L.].

Maritime Woodland—shrublands, disturbed sands; freq. 1990,

2008/10. No specimens collected.

Apiaceae

Centellaasiatica (L.) Urb. [=C. erecta (L. f.) Fernald.]. Ruderal—moist

fields and other open moist, sandy places: freq. Stalter, 26

Oct 1990.

Chaerophyllum tainturieri Hook. Ruderal—fields and waste places:

infreq. to locally freq. Stalter, 29 Mar 1990.

*Daucus carota L. Ruderal—roadsides and waste places: infreq to

locally freq. Stalter, 27 Aug 1990.

Ptilimnium capillaceum (Michx.) Raf. Ruderal—apparently salt

tolerant: infreq to locally freq. Stalter, 29 Mar 1990. Stalter

19938, 2008/10.

Aquifoliaceae

Ilex opaca Ait. Maritime Woodland: infreq. Stalter, 27 Aug 1990.

Stalter 19574, 2008/10.

Ilex vomitoria Ait. Ruderal and Maritime Woodland—swales, thick¬

ets, and fields: freq. Stalter, 29 Mar 1990. Stalter 19575,2008/10.

Araliaceae

Hydrocotyle bonariensis Lam. Dunes—moist, open swales: infreq.

Stalter, 29 Mar 1990. Stalter 20280, 2008/10.

Hydrocotyle umbellata L. Dunes—moist, open swales: infreq. Stalter,

29 Mar 1990.

Hydrocotyle verticillataThunb. Sight record. Schmidt 2004.

Asteraceae

* Achillea millefolium L. Ruderal: freq. Stalter, 14 Jun 1990.

Ambrosia artemisiifolia L. Ruderal: freq. Stalter, 29 Mar 1990. Stalter

19480,008/10.

* Artemisia vulgaris L. Ruderal—disturbed site adjacent to the Visi¬

tor's Center: rare. Stalter, 14 Jun 1990.

Baccharis halimifolia L. Salt Marsh: freq. Stalter, 29 Mar 1990. Stalter

19481,2008/10.

Bidens bipinnata L. Ruderal: infreq. Stalter 20055,2008/10.

Borrichia frutescens (L.) DC. Salt Marsh: freq. Stalter, 27 Aug 1990,

Stalter 20056, 2008/10.

Carphephorus sp. Sight record. Schmidt 2004.

Cirsium nutallii DC. Ruderal—fields: infreq. Stalter, 27 Aug 1990.

324

Conoclinium coelestinum (L.) DC. [=Eupatorium coelestinum L.].

Ruderal—fields: infreq. Stalter, 26 Oct 1990.

*Conyzo bonariensis (L.) Cronq. [ =Erigeron bonariensis L.]. Ru¬

deral—disturbed fields: freq. Stalter, 14 Jun 1990. Stalter

20057, 2008/10.

Conyzo canadensis (L.) Cronq. var. pusilla (Nutt.) Cronq. [=Erigeron

canadensis L. var. pusillus (Nutt.) Boivin, non Ahles]. Dunes: freq.

Stalter, 26 Oct 1990. Stalter 19484, 2008/10.

*Coreopsis basalis (A. Dietr.) Blake. Ruderal—disturbed fields: rare.

Stalter, 14 Jun 1990.

Elephantopus nudatus A. Gray. Maritime Woodland: rare. Stalter,

27 Aug 1990.

Elephantopus tomentosus L Sight record. Schmidt 2004.

Erechtites hieracifolius (L.) Raf. ex DC. Ruderal—disturbed fields and

waste places: infreq. Stalter, 14 Jun 1990. Stalter 20058,2008/10.

Erigeron philadelphicuslvar. philadelphicus. Ruderal—fields: infreq.

Stalter, 29 Mar 1990; 26 Oct 1990.

Erigeron quercifolius Lam. Ruderal—fields, roadsides, and waste

places: freq. Stalter, 29 Mar 1990. Stalter 19486, 2008/10.

Erigeronstrigosus Muhl. ex Willd. Ruderal—fields and waste places:

freq. Stalter, 29 Mar 1990; 14 Jun 1990.

Eupatorium capillifolium (Lam.) Small. Ruderal—fields and swales:

freq. Stalter, 26 Oct 1990. Stalter 20059, 2008/10.

Euthamia graminifolia (L.) Nutt. [=Solidago microcephala (Greene)

Bush]. Ruderal—fields and swales: freq. Stalter, 26 Oct 1990.

Stalter 20062, 2008/10.

Gaillardiapulchella Foug. Ruderal—sandy waste places and swales:

freq. Stalter, 14 Jun 1990; 26 Oct 1990. Stalter 20063,2008/10.

Gamochaeta argyrinea G.L. Nesom. GA Acc#237534. Schmidt 2004.

*Gamochaeta calviceps (Fernald) Cabrera [=G. falcata (Lam.) Ca¬

brera]. Sight record, Schmidt 2004.

Gamochaeta pensylvanica (Willd.) Cabrera. GA Acc# 237530, GA

Acc#237475. Schmidt 2004.

Gamochaeta purpurea (L.) Cabrera. [=Gnaphalium purpureum L. var.

purpureum ]. Ruderal—fields lawns, roadsides, and waste places:

freq. Stalter 19487,1990. Stalter 20061,2008/10.

Heterothecasubaxillaris (Lam.) Britton & Rusby. Ruderal—disturbed

sands and waste places: freq. Stalter, 26 Oct 1990. Stalter

19488, 2008/10.

Hieracium gronovii L. Ruderal—lawns and fields: infreq. Stalter

19489, 2008/10.

*Hypochaeris chillensis (Kunth) Britton. Ruderal—sandy waste

places: rare. Stalter, 29 Mar 1990.

*Hypochaeris glabra L. Ruderal—lawns, fields, and waste places:

infreq. Stalter, 29 Mar 1990. Stalter 19490, 2008/10.

Iva frutescens L. Salt Marsh: freq. Stalter, 29 Mar 1990. Stalter 19491,

2008/10.

Lactuca canadensis L. Dunes—along seashore: infreq. Stalter, 14

Jun 1990. Stalter 19492, 2008/10.

Pluchea camphorata (L.) DC. Ruderal—moist soil and ephemeral

ponds: freq. Stalter 20281 2008/10.

Pluchea carolinensis (Jacq.) G. Don. Schmidt 2004. According to

Wunderlin (1998) this taxon occupies the southern portion of

the Florida penninsula.

Pluchea odorata var. odorata (L.) Cass. Ruderal—roadside ditch and

moist (fresh to brackish) places: infreq to locally freq. Stalter,

27 Aug 1990. Stalter 20253, 2008/10.

Pseudognaphalium obtusifolium (L.) Hillard & Burtt. Ruderal—fields

and swales: freq. Stalter, 29 Mar 1990.

Pyrrhopappus carolinianus (Walter) DC. Ruderal—fields and waste

places: infreq. Stalter, 29 Mar 1990. Stalter 19493, 2008/10.

*Senecio vulgaris L. Ruderal—lawns and waste places: infreq. Stalter,

14 Jun 1990. Stalter 19494, 2008/10.

Solidago odora Ait. Ruderal—thickets: infreq. Stalter, 26 Oct 1990.

Journal of the Botanical Research Institute of Texas 8(1)

Solidago rugosa var. a spera (Ait.) Fernald. Ruderal—fields: infreq.

Stalter, 26 Oct 1990. Stalter 20062, 2008/10.

Solidago mexicana L. Dunes—upper beach and swales: freq. Stalter,

26 Oct 1990. Stalter 20064, 2008/10.

Solidago stricta Ait. Ruderal—roadside ditch: rare. Stalter, 26 Oct

1990.

*Sonchusasper{ L.) Hill. Ruderal—disturbed sands and waste places:

infreq. Stalter, 29 Mar 1990.

*Sonchus oleraceus L. Ruderal—disturbed sands: rare. Stalter, 14

Jun 1990. Stalter 19495, 2008/10.

Symphyotrichum dumosum (L.) Nesom var. dumosum [=Aster du-

mosus L.]. Ruderal—fields: infreq. Stalter, 26 Oct 1990.

Symphyotrichum pilosum (Gray) Nesom var. pringlei [=Aster pilosus

var. demotus Blake]. Ruderal—moist roadsides: infreq. Stalter,

26 Oct 1990.

Symphyotrichum subulatum (Michx.) Nesom [=Aster subulatus var.

subulatus Michx.]. Ruderal—roadside ditches: infreq. to locally

freq. Stalter, 26 Oct 1990.

Symphyotrichum tenuifolium (L.) Nesom [=Aster tenuifolius L.].

Salt Marsh: infreq. to locally freq. Stalter, 26 Oct 1990. Stalter

20065, 2008/10.

* Taraxacum erythrospermum Andrz. ex Besser. [=Taraxacum laev-

igatum (Willd.) DC.]. Ruderal—lawns: rare. Stalter, 29 Mar 1990.

*Taraxacum officinale Weber ex Wigg. Ruderal—lawns and dis¬

turbed sites: freq. Stalter 29 Mar 1990. Stalter 19496, 2008/10.

Xanthium strumarium L. [=X. echinatum Murray]. Dunes: infreq.

Stalter 20066, 2008/10.

*Youngia japonica (L.) DC. [=Crepis japonica (L.) Benth.]. Ruderal—

disturbed sites: rare. Stalter, 14 Jun 1990.

Bignoniaceae

Bignoniacapreolata L. [ =Anisostichus capreolata (L.) Bureau]. Rude¬

ral—thickets: rare. Stalter, 14 Jun 1990.

Campsis radicans (L.) Seem, ex Bureau. Ruderal—edge of field on

trees: rare. Stalter, 14 Jun 1990. Stalter 19535, 2008/10.

Boraginaceae

*Buglossoides arvensis (L.) I.M. Johnst ssp. arvensis. Ruderal—road¬

sides and sandy fields: infreq. Stalter, 14 Jun 1990.

Brassicaceae

Cakile edentula (Bigelow) Hook. Dunes—upper beach: freq. Stalter,

29 Mar 1990. Stalter 19596, 2008/10.

*Cardamine hirsuta L. Ruderal—disturbed sands and roadsides:

infreq. Stalter, 29 Mar 1990. Stalter 19537, 2008/10.

Caradamine pensylvanica Muhl. ex Willd. Ruderal—moist fields and

lawns: infreq. Stalter, 29 Mar 1990.

Descurainia pinnata (Walter) Britton. Ruderal—sandy fields and

waste places: freq. Stalter, 29 Mar 1990. Stalter 19575,2008/10.

*Lepidium didymum L. Ruderal—fields, roadsides, and disturbed

habitats: freq. Stalter, 29 Mar 1990.

Lepidium virginicum L. ssp. virginicum. Ruderal—disturbed sands,

fields, roadsides, and waste places: freq. Stalter, 29 Mar 1990.

Stalter 19577, 2008/10.

Cactaceae

Opuntiaficus-indica (L.) P. Mill. Dunes—swales: rare. Stalter 19538,

2008/10.

Opuntia humifusa (Raf.) Raf. Dunes—swales and sandy waste

places: infreq, but locally abundant. Stalter, 27 Aug 1990.

Stalter 19539, 2008/10.

Opuntia pusilla (Haw.) Nutt. Dunes—swales: rare. Stalter, 27 Aug

1990. Stalter 19540, 2008/10.

Campanulaceae

Triodanisperfoliata (L.) Nieuwl. Ruderal—disturbed sands and waste

places: infreq. Stalter, 29 Mar 1990. Stalter 19576, 2008/10.

Stalter et al., Flora of Fort Sumter and Fort Moultrie

325

Cannabaceae

Celtis loevigoto Willd. Maritime Woodland—salt marsh border

at Visitor's center: infreq. Stalter, 14 Jun 1990. Stalter 19497,

2008/10.

Caprifoliaceae

*Lonicera japonica Thunb. Ruderal—disturbed sands and fields:

freq. Stalter, 29 Mar 1990. Stalter 19498, 2008/10.

Caryophyllaceae

*Arenorio serpyllifolio L. Ruderal—disturbed sands and pavement

cracks: rare. Stalter, 29 Mar 1990. Stalter 19499, 2008/10.

*Cerostium fontonum Baumg. var. vulgore (Hartman) Greuter &

Burdet. Sight record. Schmidt 2004.

*Cerastium g/omerafumThuill. Ruderal—lawns and disturbed sites:

freq. Stalter, 29 Mar 1990.

*Polycorpon tetrophyllum (L.) L. ssp. tetrophyllum. Ruderal—lawns

and disturbed sites: rare. Stalter, 29 Mar 1990.

*Stellorio medio (L.) Vill. Ruderal—lawns, fields, and waste places:

freq. Stalter, 29 Mar 1990. Stalter 19500, 2008/10.

Cistaceae

Lecheo mucronoto Raf. GA Acc# 237706. Schmidt 2004.

Convolvulaceae

Dichondra corolinensis Michx. Ruderal—lawn: rare, only one popu¬

lation seen adjacent to casemate wall. Stalter, 14 Jun 1990.

Stalter 19939, 2008/10.

Ipomoea cornea ssp. fistulosa (Mart, ex Choisy) D. Austin. Schmidt

2004.

Ipomoea imperati (Vahl) Griseb. [=/. stolonifera (Cirillo) J.F. Gmelin].

Dunes—upper beach: infreq. Stalter, 27 Aug 1990. Stalter

20282, 2008/10.

Ipomoea locunoso L. Dunes and Ruderal—waste places: freq. Stalter,

26 Oct 1990. Stalter 19940, 2008/10.

Ipomoea pandurata (L.) G. Mey. Ruderal—fields: infreq. Stalter,

26 Oct 1990.

Ipomoea sagittata Poir. Salt Marsh—upper fringe: freq. Stalter, 14

Jun 1990. Stalter 19941,2008/10.

Cucurbitaceae

*Citrullus lanatus (Thunb.) Matsum. & Nakai ssp. lanatus. Dunes—

upper beach: rare. Stalter, 26 Oct 1990.

Melothria pendula L. Ruderal—swales and fields: infreq. Stalter, 27

Aug 1990. Stalter 19942, 2008/10.

Ebenaceae

Diospyros virginiana L. Ruderal—successonal fields: rare. Stalter,

14 Jun 1990.

Elaeagnaceae

*Elaeagnus pungens Thunb. Ruderal—roadside: rare. Stalter, 29 Mar

1990. Stalter 19541,2008/10.

Euphorbiaceae

Acolypho grocilens A. Gray. Ruderal—disturbed sands: infreq. to

locally freq.. Stalter 19943, 2008/10.

Croton glondulosus L. var. septentrionalis Mull. Arg. Ruderal—swales

and disturbed sands: freq. Stalter, 14 Jun 1990; 26 Oct 1990.

Stalter 20254, 2008/10.

Croton willdenowii Webster. Schmidt 2004.

Croton punctatus Jacq. Dunes: infreq. Stalter, 26 Oct 1990. Stalter

19542, 2008/10.

Euphorbia heterophyllo L. Ruderal—disturbed site near fort: rare.

Stalter, 26 Oct 1990. Stalter 20255, 2008/10.

Euphorbia moculoto L. [=Chomaesyce moculoto (L.) Small]. Ru¬

deral—lawns, roadsides, disturbed places and waste places:

freq. Stalter, 14 Jun 1990. Stalter 19543, 2008/10.

Euphorbia nutans Lag.&Segura [=Chamaesyce nutans (Lag.) Small].

Schmidt 2004.

Euphorbia polygonifolio L. [=Chamaesyce polygonifolio (L.) Small].

Dunes—upper beaches, foredunes and swales: freq. Stalter, 14

Jun 1990. Stalter 19544, 2008/10.

Fabaceae

*Aeschynomeneindica L. Ruderal—wet roadside ditch: rare. Stalter,

27 Aug 1990.

Centrosema virginianum (L.) Benth. Ruderal—fields and disturbed

sites: freq. Stalter, 27 Aug 1990.

Chamaecrista fasciculata (Michx.) Greene var. fasciculata [=Cassia

fosciculoto Michx.]. Ruderal—roadsides and disturbed sands:

freq. Stalter, 29 Mar 1990. Stalter 20256,2008/10.

Chamaecrista nictitons (L.) Moench var. nictitons [=Cossio nictitons

L.]. Ruderal—roadsides and disturbed sands: freq. Stalter, 29

Mar 1990.

Clitoriamariana L. var. mariana. Ruderal—roadside: rare. Stalter, 27

Aug 1990. Stalter 20257, 2008/10.

Crotalaria spectabilis Roth. Ruderal—fields and disturbed sites:

infreq. Stalter, 27 Aug 1990.

Desmodium incanum DC. Sight record. Schmidt 2004.

Galactia volubilis (L.) Britton var. volubilis. Ruderal—fields and

swales: freq. Stalter, 27 Aug 1990. Stalter 20258, 2008/10.

*Medicago arabica (L.) Huds. Ruderal—fields and disturbed soil:

freq. Stalter 19501,2008/10.

*Medicago lupulina L. Ruderal—disturbed sands and roadsides:

infreq. Stalter, 29 Mar 1990. Stalter 19502, 2008/10.

*Medicagopolymorpha L. Ruderal—lawns, fields, and waste places:

infreq. Stalter 19545, 2008/10.

*Melilotus olbus Medik. Ruderal—roadsides and waste places: freq.

Stalter, 29 Mar 1990. Stalter 19503, 2008/10.

*Melilotus officinalis (L.) Pallas. Ruderal—roadsides: infreq. Stalter,

29 Mar 1990.

Senna obtusifolia (L.) Irwin & Barneby. GA Acc#237795. Schmidt

2004.

Sesbania punicea (Cav.) Benth. Dunes: freq. Stalter, 27 Aug 1990.

Stalter 20259, 2008/10.

Sesbania vesicaria (Jacq.) Ell. Ruderal—side of path to beach: rare.

Stalter 20260, 2008/10.

Strophostyles helvola (L.) Ell. Dunes—swales and disturbed sands:

infreq. Stalter, 26 Aug 1990. Stalter 19504, 2008/10.

*Trifolium dubium Sibth. Ruderal—lawns and roadsides: freq.

Stalter, 14 Jun 1990.

*Trifolium repens L. Ruderal—lawns: infreq. Stalter, 14 Jun 1990.

Stalter 19505, 2008/10.

*Vicia hirsuto (L.) Gray. Ruderal—fields: rare. Stalter, 29 Mar 1990.

Vicia sativa L. ssp. nigra (L.) Ehrh. Ruderal—swales and disturbed

sands: infreq. Stalter, 29 Mar 1990; visitor center: freq. Stalter

19506, 2008/10.

*Wisterio sinensis (Sims) DC. Ruderal—border of dune and moist

roadside ditch: rare. Stalter, 14 Jun 1990. Stalter 19579,2008/10.

Fagaceae

Quercus chapmanii Sarg. GA Acc#237568. Schmidt 2004.

Quercus geminata Small. GA Acc#237817. Schmidt 2004.

Quercus lourifolio Michx. Maritime Woodland—thickets: infreq.

Stalter, 29 Mar 1990, Stalter 19546, 2008/10.

Quercus nigra L. Maritime Woodland: rare Stalter, 27 Aug 1990.

Quercus virginiana Mill. Maritime Woodland—fields and scattered

stations: infreq but widespread. Stalter, 29 Mar 1990. Stalter

19547, 2008/10.

Gentianaceae

Sabatiastellaris Pursh. Salt Marsh: rare. Stalter, 14 Jun 1990; Dunes:

infreq. Stalter, 14 Jun 1990. Stalter 19808, 2008/10.

326

Geraniaceae

Geranium carolinianum L. Ruderal—fields and disturbed sands: freq.

Stalter, 29 Mar 1990. Stalter 19548, 2008/10.

Hypericaceae

Hypericum gentianoides (L.) Britton, Sterns & Poggenb. Ruderal—

swales and disturbed sands: locally freq. Stalter, 14 Jun 1990.

Hypericum hypericoides (L.) Crantz. Ruderal—swales: rare. Stalter,

27 Aug 1990. Stalter 19809, 2008/10.

Juglandaceae

*Carya illinoinensis (Wangenh.) Koch. Maritime Woodland—near

fort, persistent after cultivation: rare. Stalter, 26 Oct 1990.

Juglanscinerea L. Maritime Woodland: rare. Stalter 19507,2008/10.

Lamiaceae

Callicarpa americana L. Ruderal—fields and swales: infreq. Stalter,

26 Oct 1990. Stalter 19578, 2008/10.

*Lamium amplexicaule L. var. amplexicaule. Ruderal—lawns, fields,

waste places: freq. Stalter, 29 Mar 1990. Stalter 19508,2008/10.

Salvia lyratal. Ruderal—only along hedgerow near visitor's center:

rare. Stalter, 14 Jun 1990. Stalter 19509, 2008/10.

*Stachys floridana Shuttlw. ex Benth. Previously identified by

Schmidt (2004) as Salvia floridana. Ruderal—field borders: freq.

Stalter 19810, 2008/10.

Stachys hyssopifolia Michx. Ruderal—field: rare. Stalter, 14 Jun 1990.

Teucrium canadense L. Maritime Woodland—edge of thickets:

infreq. Stalter, 26 Oct 1990. Stalter 20269, 2008/10.

*Vitex agnus-castus L. Ruderal—persistant after cultivation near

fort: rare. Stalter, 14 Jun 1990.

Lauraceae

Persea borbonia (L.) Spreng. Maritime Woodland—old stable

dunes at maritime forests: rare. Stalter, 29 Mar 1990. Stalter

19510, 2008/10.

Malvaceae

Hibiscus moscheutos L. Dunes—ephemeral ponds: infreq. Stalter

19811,2008/10.

Kosteletzkya pentacarpos (L.) Ledeb. Dunes—ephemoral ponds:

infreq. Stalter 19812, 2008/10.

Sida rhombifolia L. var. rhombifolia. Ruderal—field: rare. Stalter, 26

Oct 1990. Stalter 19813, 2008/10.

Meliaceae

*Melia azedarach L. Ruderal—fields and roadsides; persistent after

cultivation: infreq. Stalter, 29 Mar 1990. Stalter 19549,2008/10.

Molluginaceae

*Mollugo verticillata L. Ruderal—roadsides and waste places: freq.

Stalter, 14 Jun 1990. Stalter 19774, 2008/10.

Moraceae

*Morus alba L. Ruderal and Maritime Woodland—fields: rare. Stalter,

29 Mar 1990. Stalter 19550, 2008/10.

Morus rubra L. Maritime Woodland—thickets: rare. Stalter, 14 Jun

1990. Stalter 19551,2008/10.

Myricaceae

Morel la cerifera (L.) Small. Ruderal and Maritime Woodland—swales

and edges of fields: infreq. Stalter, 29 Mar 1990. Stalter 19580,

2008/10.

Nyctaginaceae

Boerhavia erecta L. Ruderal—sands: infreq. Stalter 19944,2008/10.

Oleaceae

*Ligustrum japonicum Thunb. Ruderal and Maritime Woodland—

escaped from cultivation near old battery: rare. Stalter, 29 Mar

1990, Stalter 19581/19814, 2008/10.

Journal of the Botanical Research Institute of Texas 8(1)

*Ligustrumsinense Lour. Ruderal and Maritime Woodland—escaped

from cultivation near old battery: infreq. Stalter, 29 Mar 1990.

Stalter 19552, 2008/10.

Onagraceae

Ludwigia maritima R.M. Harper. Ruderal—moist roadside ditch:

infreq. Stalter, 29 Mar 1990. Stalter 19815, 2008/10.

Oenothera biennis L. Dunes: rare. Stalter, 27 Aug 1990. Stalter

19816/19945, 2008/10.

Oenothera drummondii Hook. ssp. drummondii. Dunes: freq. but

rare in the Carolinas (Radford et al. 1968). Stalter, 29 Mar 1990.

Stalter 19817, 2008/10.

Oenothera fruticosa L. Sight record. Schmidt 2004.

Oenothera humifusa Nutt. Dunes: infreq. Stalter, 27 Aug 1990.

Oenothera laciniata Hill. Ruderal—disturbed sands and fields: freq.

Stalter, 29 Mar 1990. Stalter 19818, 2008/10.

*Oenothera speciosa Nutt. Ruderal—fields: infreq. to locally freq.

Stalter, 29 Mar 1990. Stalter 19775, 2008/10.

Oxalidaceae

Oxalis a rticu lata Savigny [=0. rubra A. St.-Hil.]. Ruderal—fields: freq.

Stalter, 29 Mar 1990. Stalter 19512, 2008/10.

Oxalis dillenii Jacq. Ruderal—disturbed sands and waste places:

infreq. Stalter, 29 Mar 1990. Stalter 19511,2008/10.

Phytolaccaceae

Phytolacca americana L. Ruderal—swales and disturbed fields:

infreq. Stalter, 29 Mar 1990. Stalter 19513, 2008/10.

Pittosporaceae

*Pittosporum tobira (Thunb.) Ait. Ruderal—persistent after cultiva¬

tion. Stalter 19581,2008/10.

Plantaginaceae

Nuttallanthus canadensis (L.) D.A. Sutton [=Linaria canadensis (L.)

Dum. Cors.]. Ruderal—swales, disturbed sands, and waste

places: freq. Stalter, 29 Mar 1990. Stalter 19553, 2008/10.

*Plantago aristata Michx. Ruderal—sandy waste places, roadsides,

fields, and lawns: freq. Stalter, 29 Mar 1990; 14 Jun 1990.

*Plantago lanceolata L. Ruderal—disturbed fields and lawns: freq.

Stalter, 29 Mar 1990. Stalter 19554, 2008/10.

Plantago majorl. Ruderal—disturbed fields and lawns: rare. Stalter,

14 Jun 1990. Stalter 19555, 2008/10.

Plantago virginica L. Ruderal—lawns: rare. Stalter, 29 Mar 1990.

Stalter 19556, 2008/10.

* Veronica arvensis L. Ruderal—lawns and waste places: freq. Stalter,

29 Mar 1990. Stalter 19557, 2008/10.

Veronica peregrina L. Ruderal—moist fields: infreq. Stalter, 29 Mar

1990. Stalter 19558, 2008/10.

Polemoniaceae

*Phlox drummondii Hook. Ruderal and Dunes—swales and

disturbed sands: infreq. Stalter, 14 Jun 1990. Stalter 19777,

2008/10.

Polygonaceae

*Persicaria maculosa S.F. Gray [=Polygonum persicaria L.]. Ruderal—

disturbed moist fields: infreq. Stalter, 14 Jun 1990.

Persicaria punctata (Ell.) Small [=Polygonum punctatum Ell.]. Ruder¬

al—fields: infreq. Stalter, 14 Jun 1990. Stalter 19776, 2008/10.

Persicaria virginiana (L.) Gaertn. [=Polygonum virginianum L.]. Ru¬

deral—moist roadside ditch: rare. Stalter, 27 Aug 1990.

Rumex conglomeratus Murray. Ruderal—seen only at one moist

field: rare. Stalter, 27 Aug 1990.

*Rumex crispus L. ssp. crispus. Ruderal—waste places and fields:

infreq. Stalter, 29 Mar 1990. Stalter 19582, 2008/10.

Rumex hastatulus Baldwin. Ruderal—disturbed sands and fields:

infreq. Stalter, 29 Mar 1990. Stalter 19583, 2008/10.

Stalter et al., Flora of Fort Sumter and Fort Moultrie

327

Rumexverticillotus L. Ruderal—moist roadside ditch: rare. Stalter,

27 Aug 1990.

Portulacaceae

*Portuiaca oleracea L. Ruderal—disturbed sands, seen only at Visi¬

tor's Center: rare. Stalter, 14 Jun 1990. Stalter 19584, 2008/10.

*Portuiaca pilosa L. Ruderal—dry sandy fields: rare. Stalter, 14 Jun

1990. Stalter 19585, 2008/10.

Rananculaceae

Ranunculus muricatus L. Ruderal—fields: freq. Stalter 19559,

2008/10.

Rosaceae

Prunusangustifolia Marshall. Dunes: rare. Stalter, 14 Jun 1990. Stalter

19515, 2008/10.

Prunus caroliniana (Mill.) Ait. Maritime Woodland—edge of swales:

infreq. Stalter, 29 Mar 1990. Stalter 19516, 2008/10.

Prunus serotina Ehrh. var. serotina. Maritime Woodland—swales

and fields: infreq. Stalter, 29 Mar 1990. Stalter 19517, 2008/10.

Rubus pensilvanicus Poir. [=/?. argutus Link]. Ruderal and Dunes—

swales and disturbed sands: infreq. Stalter, 29 Mar 1990.

Rubus trivialis Michx. Ruderal and Dunes—swales and fields: freq.

Stalter, 29 Mar 1990. Stalter 19819, 2008/10.

Rubiaceae

Diodia teres Walter. Ruderal—swales and disturbed sands: freq.

Stalter, 14 Jun 1990; 26 Oct 1990. Stalter 19820, 2008/10.

Diodia virginiana L. Ruderal—moist roadside ditch and moist field:

infreq. Stalter, 14 Jun 1990. Stalter 19821,2008/10.

Galium tinctorium (L.) Scopoli var. tinctorium. Ruderal—moist field:

rare. Stalter, 27 Aug 1990.

Oldenlandia uniflora L. Ruderal—fields: rare. Stalter, 14 Jun 1990.

*Richardia scabra L. Ruderal—waste places: rare. Stalter, 27 Aug

1990.

Rutaceae

Zanthoxylum clava-herculis L. Dunes—swales: infreq. Stalter, 29 Mar

1990. Stalter 19513, 2008/10.

Salicaceae

*Populus alba L. Ruderal—waste places: infreq. Stalter, 29 Mar 1990.

Stalter 19514, 2008/10.

Populus heterophylla L. Ruderal—moist field: rare. Stalter, 29 Mar

1990.

Sapotaceae

Sideroxylon lycioides L. [Bumelia lycioides (L.) Pers.]. Maritime

Woodland—swales: rare. Stalter, 29 Mar 1990. Stalter 19560,

2008/10.

Sideroxylon tenax L. [=Bumelia tenax (L.) Willd.]. Maritime Woo¬

dland—thickets and woodland: infreq. Stalter, 27 Aug 1990.

Stalter 19561,2008/10.

Solanaceae

Physalis waited Nutt [=P. viscosa ssp. maritima (M. A. Curtis) Waterf.].

Dunes—swales and dunes: infreq. Stalter, 14 Jun 1990. Stalter

19823, 2008/10.

Solanum americanum Mill. [=5. nigrum var. virginicum L.]. Ruderal—

disturbed sands near margin of salt marsh: rare. Stalter, 29 Mar

1990. Stalter 19822, 2008/10.

Solanum carolinense L. Ruderal—swales and disturbed sands: infreq.

Stalter, 14 Jun 1990.

Solanum rostratum Dunal. Ruderal—disturbed fields: infreq. Stalter,

27 Aug 1990.

Tamaricaceae

*Tamarix gallica L. Salt Marsh—seen only at border between salt

marsh and uplands: rare. Stalter, 29 Mar 1990. Stalter 20283,

2008/10.

Tetrachondraceae (previously in Buddlejaceae)

Polypremum procumbens L. Ruderal—dry sandy soil: freq. Stalter,

14 Jun 1990. Stalter 19518, 2008/10.

Verbenaceae

*Lantana camara L. Ruderal—escaped from cultivation at field near

fort: rare. Stalter, 29 Mar 1990. Stalter 19519, 2008/10.

Phyla nodiflora (L.) Greene. Ruderal—moist disturbed sands: infreq.

to locally freq. Stalter, 14 Jun 1990. Stalter 19946, 2008/10.

*\ /erbena bonariensis L. Ruderal—edge of fields: freq. Stalter, 14 Jun

1990. Stalter 19947, 2008/10.

*\ /erbena brasiliensis Veil. Ruderal—fields: infreq. Stalter, 14 Jun

1990. Stalter 19948, 2008/10.

Vitaceae

Ampelopsis arborea (L.) Koehne. Ruderal—swales: infreq. Stalter, 29

Mar 1990. Stalter 19562, 2008/10.

Parthenocissus quinquefolia (L.) Planch. Maritime Woodland—

swales, thicket: infreq. Stalter, 29 Mar 1990. Stalter 19563,

2008/10.

Vitis aestivalis Michx. Maritime Woodland: infreq. Stalter, 29 Mar

1990. Stalter 19564, 2008/10.

LILIOPSIDA (MONOCOTS)

Agavaceae

*Yucca aloifolia L. Dunes—swales: infreq. Stalter, 29 Mar 1990.

Stalter 19825, 2008/10.

Yucca filamentosa L. Dunes—protected dunes and dry sands: infreq.

Stalter, 29 Mar 1990. Stalter 19826, 2008/10.

Amaryllidaceae

*Nothoscordum bivalve (L.) Britton. Ruderal—fields: rare. Stalter,

29 Mar 1990.

Zephyranthes atamasco (L.) Herb. Ruderal—lawn in front of fort:

infreq. Stalter, 14 Jun 1990. Stalter 19520, 2008/10.

Arecaceae

Saba!palmetto (Walter) Lodd. ex Schult.& Schult. Maritime Wood¬

land—swales: infreq. Stalter, 29 Mar 1990. Stalter 19586,

2008/10.

Asparagaceae

* Asparagus aethiopicus L. [=A. densiflorus (Kunth) Jessop]. Ruderal—

roadsides: rare. Stalter 19587, 2008/10.

* Asparagus officinalis L. Ruderal—roadsides and waste places:

infreq. Stalter, 14 Jun 1990; 26 Oct 1990. Stalter 19588,2008/10.

Bromeliaceae

Tillandsia recurvata (L.) L. Maritime Woodland—epiphyte on Q.

virginiana: rare. Stalter 19521, 2008/10.

Tillandsia usneoides (L.) L. Maritime Woodland—epiphyte on Q.

virginiana: infreq. Stalter, 29 Mar 1990.

Cannaceae

*Canna x generalis L.H. Bailey (pro. sp.) [=C. glauca x indica].

Ruderal—persistent after cultivation near fort; entrance to

path to beach: locally abundant. Stalter, 14 Jun 1990. Stalter

19827, 2008/10.

Commelinaceae

*Commeiina communis L. Ruderal—roadside ditch: locally abun¬

dant. Stalter 19949, 2008/10.

Tradescantiaohiensis Raf. Ruderal—swales and fields: infreq. Stalter,

29 Mar 1990. Stalter 19589, 2008/10.

Cyperaceae

Bolboschoenus robustus (Pursh) Sojak. [=Scirpus robustus Pursh]. Salt

Marsh: rare. Stalter, 14 Jun 1990.

328

Journal of the Botanical Research Institute of Texas 8(1)

Corexlongii Mack. Ruderal—field edges: rare. Stalter, 14 Jun 1990.

Stalter 19950, 2008/10.

Cyperus croceusM ahl. [=C. globulosus AublJ. Ruderal—sandy fields:

rare. Stalter, 26 Oct 1990.

Cyperus echinatus (L.) Wood. [=Cyperus ovuloris (Michx.) Torr.].

Ruderal—fields: rare. 26 Oct 1990.

Cyperus esculentus L. var. leptostochyus Boeck. Salt Marsh—swales:

infreq to locally freq. Stalter, 26 Oct 1990. Stalter 19829,

2008/10.

Cyperus filicinus Vahl. Ruderal—moist soil: infreq. Stalter 19830,

2008/10.

Cyperus odorotus L. Ruderal—moist soil: infreq. Stalter 19831,

2008/10.

Cyperus pseudovegetus Steud. Ruderal—ditch bordering road:

infreq. Stalter 19834, 2008/10.

Cyperus retrorsus Chapm. Ruderal—moist soil: infreq. Stalter, 26 Oct

1990. Stalter 19832, 2008/10.

Cyperus strigosus L. Ruderal—moist soil: infreq. Stalter, 27 Aug 1990.

Stalter 19833, 2008/10.

Cyperus surinamensis Rottb. Sight record. Schmidt 2004.

Eleocharis elliptica Kunth. Ruderal—moist depressions: infreq.

Stalter 19951,2008/10.

Eleocharis flovescens (Poir.) Urban. Ruderal—moist edge of roadside

ditch: rare. Stalter, 29 Mar 1990; 14 Jun 1990.

Fimbristylis castanea (Michx.) Vahl. Dunes—interdunal depressions;

infreq. Stalter 19835, 2008/10.

Fimbristylis puberula (Michx.) Vahl. var. puberula [=F. spadicea (L.)

Vahl.]. Salt Marsh—transition zone between salt marsh and

uplands: infreq. Stalter, 26 Oct 1990. Stalter 19836, 2008/10.

Kyllinga brevifolia Rottb. [=Cyperus brevifolius (Rottb.) Endl. ex

HasskJ. Schmidt 2004.

Juncaceae

Juncus roemerianus Scheele. Salt Marsh—brackish upper of tidal

marshes: infreq. to locally freq. Stalter, 14 Jun 1990. Stalter

19522, 2008/10.

Iridaceae

Sisyrinchium rosulatum Bickn. Ruderal—lawns: freq. Stalter 19523,

2008/10.

Liliaceae

*Allium vineale L. Ruderal—lawns and fields. Stalter, 29 Mar 1990.

Stalter 19524, 2008/10.

*Ornithogalum umbellatum L. Ruderal—lawns and waste places:

infreq. Stalter, 29 Mar 1990. Stalter 19525, 2008/10.

Poaceae

Andropogon glomeratus (Walter) Britton, Sterns & Poggenb. Ru¬

deral—moist sands and old dunes: infreq. Stalter, 26 Oct 1990.

Stalter 19837, 2008/10.

Andropogon gyrans Ashe [=Andropogon elliotii Chapm.]. Ruderal—

swales: infreq. 26 Oct 1990. Schmidt 2004.

Andropogon ternarius Michx. var. ternarius. Ruderal—fields and old

dunes: infreq. Stalter, 26 Oct 1990.

Andropogon virginicus L. var. virginicus. Ruderal—swales, fields and

roadsides: freq. Stalter, 26 Oct 1990. Stalter 19838, 2008/10.

*Arundo donax L. Ruderal—border of visitor center parking lot: rare.

Stalter 19839, 2008/10.

Brizaminor L. Ruderal—disturbed sands: infreq. Stalter, 29 Mar 1990.

Stalter 19565, 2008/10.

*Bromus catharticus Vahl. var. catharticus [=B. wildenowii Kunth.].

Ruderal—roadsides: rare. Stalter, 29 Mar 1990.

Cenchrus longispinus (Hack.) Fernald. Ruderal—sandy fields and

waste places: infreq. Stalter, 26 Oct 1990. Stalter 19778,

2008/10.

Cenchrus tribuloides L. Dunes—swales: freq. Stalter, 26 Oct 1990.

Stalter 19779, 2008/10.

Chasmonthium laxum (L.) Yates [=Uniola laxa (L.) Britton, Sterns

& Poggenb.]. Ruderal—moist, sandy fields: infreq. Stalter, 29

Mar 1990.

*Cynodon dactylon (L.) Pers. Ruderal—lawns, fields, roadsides

and disturbed sites: freq. Stalter, 14 Jun 1990. Stalter 19566,

2008/10.

Dichanthelium acuminatum (Sw.) Gould & Clark var. acuminatum

[=Panicum lanuginosum Ell.]. Ruderal—fields: freq. Stalter,

14 Jun 1990.

Dichanthelium commutatum (Schult.) Gould var. commutatum.

[=Panicum commutatum Schult.]. Sight record. Schmidt 2004.

Dichantheliumscobriusculum (Ell.) Gould &C.A. Clark.Schmidt 2004.

*Digitorio sanguinalis (L.) Scop. Ruderal—roadsides, lawns and

disturbed sites: freq. Stalter, 26 Otober 1990. Stalter 19780,

2008/10.

Digitaria violascens Link. Ruderal—sandy roadside: rare. Stalter,

26 Oct 1990.

Distichlis spicata (L.) Greene. Salt Marsh: freq. Stalter, 26 Oct 1990.

Stalter 19781,2008/10.

*Eleusineindica (L.) Gaertn. Ruderal—roadsides and disturbed sites:

infreq. Stalter, 29 Mar 1990; 14 Jun 1990. Stalter 19782,2008/10.

Elymus virginicus L. var. halophilus (Bickn.) Wieg. Salt Marsh—upland

borders of brackish water and salt marshes: infreq. Stalter, 29

Mar 1990; 14 Jun 1990. Stalter 19840, 2008/10.

Eustachys petraea (Sw.) Desv. [=Chloris petraea Sw.]. Ruderal—

swales: rare. Stalter, 26 Oct 1990.

*Loliumperenne L. var. aristatum. Ruderal—fields and lawns: infreq.

Stalter, 14 Jun 1990. Stalter 19783, 2008/10.

Muhlenbergia capillaris (Lam.) Trin. Dunes: infreq. Stalter 19841,

2008/10.

Muhlenbergia expansa (Poir.) Trinius [ =M . capillaris var. t richopodes

(Ell.) Vasey]. Sight record. Schmidt 2004.

Panicum amarum Ell. Dunes: freq. Stalter, 26 Oct 1990. Stalter

19842, 2008/10.

Panicum dichotomiflorum Michx. Ruderal—moist roadside depres¬

sions and borders of marshes: infreq. Stalter, 26 Oct 1990.

*Paspalum dilatatum Poir. ssp. dilatatum. Ruderal—roadsides and

waste places: freq. Stalter, 14 Jun 1990; 26 Oct 1990. Stalter

19845, 2008/10.

Pospolum floridonum Michx. Ruderal—fields: infreq. Stalter 19843,

2008/10.

Pospolum notatum Flugge. Ruderal—lawns,fields, and waste places:

freq. Stalter, 26 Oct 1990. Stalter 19844, 2008/10.

*Poa annua L. Ruderal—lawns, roadsides and disturbed sands: freq.

Stalter, 29 Mar 1990. Stalter 19526, 2008/10.

Setorio corrugoto (Ell.) Schult. Ruderal—fields: freq. Stalter, 27

Aug 1990.

Sorghum halepense (L.) Pers. Ruderal—disturbed sites, ditches: freq.

Stalter, 26 Oct 1990. Stalter 19846, 2008/10.

Spartina alterniflora Loisel. Salt Marsh: freq. Stalter, 26 Oct 1990.

Stalter 19848, 2008/10.

Spartina patens (Ait.) Muhl. Dunes: infreq. Stalter, 29 Mar 1990; 26

Oct 1990. Stalter 19847, 2008/10.

Sphenopholis nitida (Biehler) Scribn. Sight record. Schmidt 2004.

*Sporobolus indicus (L.) R. Brown. [=5. poiretii (Roem. & Schult.)

Hitchc.]. Ruderal—lawns and fields: freq. Stalter, 27 Aug 1990.

Stalter 19955, 2008/10.

Sporobolus virginicus (L.) Kunth. Salt Marsh: infreq. to locally freq.

Stalter, 14 Jun 1990. Stalter 19849, 2008/10.

Stenotaphrum secundatum (Walter) Kuntze. Ruderal—moist field:

infreq. Stalter, 26 Oct 1990.

Triplasis purpurea (Walter) Chapm. var. purpurea. Dunes—interdu-

Stalter et al., Flora of Fort Sumter and Fort Moultrie

329

nal swales and dry sandflats: freq. Stalter, 26 Oct 1990. Stalter

19850, 2008/10.

Uniolapaniculata L. Dunes: freq. Stalter, 26 Oct 1990. Stalter 19851,

2008/10.

Vulpia octoflora (Walter) Rydb. var. octoflora [=Festuca octoflora

Walter]. Ruderal—lawns and disturbed sands: infreq. Stalter,

14 Jun 1990. Stalter 19956, 2008/10.

Smilacaceae

Smilox ouriculoto Walter. Dunes: rare. Stalter, 26 Oct 1990. Stalter

19527, 2008/10.

Smilaxbona-noxL Maritime Woodland—successional shrublands

and swales: freq. Stalter, 26 Oct 1990. Stalter 19528, 2008/10.

Smilax laurifolia L. Maritime Woodland—thickets: rare. Stalter, 29

Mar 1990.

Typhaceae

Typha latifolia L. Ruderal—wet roadside ditch: rare. Stalter, 26

Oct 1990; ephemeral wet depressions: infreq. Stalter 19784,

2008/10.

FORT SUMTER

POLYPODIOPHYTA

Polypodiaceae

Pleopeltis polypodioides ssp. michauxiana (Weath.) E.G. Andrews

& Windham [=Polypodium polypodioides (L.) Watt var. mich-

auxianum Weatherby]. Ruderal—masonry walls: rare. Stalter,

14 Jun 1990.

Pteridaceae

*Pteris vittoto L. Ruderal—masonry walls: infreq. Stalter 19567,

2008/10.

MAGNOLIOPHYTA MAGNOLIOPSIDA (DICOTS)

Aizoaceae

Sesuviumportulacastrum (L.) L. Salt Marsh: freq. Stalter, 14 Jun 1990.

Stalter 19568, 2008/10.

Amaranthaceae

Atriplexmucronata Raf. [ =A . arenaria Nutt.]. Dunes—upper beach

and dunes: infreq. Stalter, 14 Jun 1990. Stalter 19786,2008/10.

*Dysphania ambrosioides (L.) Mosyakin &Clemants. [ =Chenopodium

ambrosioides L.]. Dunes—upper beach: freq. Stalter, 14 Jun

1990. Stalter 19801,2008/10.

Solicornio virginico L. Salt Marsh: freq. Stalter 19590, 2008/10.

Salsola kali L. Dunes—upper beach: freq. Stalter, 14 Jun 1990.

Stalter 19785, 2008/10.

Suaeda linearis (Ell.) Moq. Salt Marsh: freq. Stalter, 14 Jun 1990.

Asteraceae

*Achillea millefolium L. Ruderal—disturbed sand outside fort: freq.

Stalter, 29 Mar 1990.

Baccharis halimifolia L. Salt Marsh—border: rare. Stalter, 14 Jun

1990; 26 Oct. Stalter 19591,2008/10.

Borrichia frutescens (L.) DC. Salt Marsh: freq. Stalter, 14 Jun 1990.

Stalter 19592, 2008/10.

*Conyza bonariensis (L.) Cronq. [=Erigeron bonariensis L.]. Ruderal—

disturbed soil outside fort: freq. Stalter, 14 Jun 1990. Stalter

19800, 2008/10.

Conyza canadensis (L.) Cronq. var. pusilla (Nutt.) Cronq. [=Erigeron

canadensis L. var. pusillus (Nutt.) Boivin, non Ahles]. Ruderal—

disturbed soil ourside fort: freq. Stalter, 14 Jun 1990. Stalter

20157, 2008/10.

Erigeron quercifolius Lam. Ruderal—lawn: freq. Stalter, 29 Mar

1990. Stalter 20158, 2008/10.

Euthamia graminifolia (L.) Nutt. [=Solidago microcephala (Greene)

Bush]. Dunes: rare. Stalter, 26 Oct 1990. Stalter 19853,2008/10.

Gamochaeta pensylvanica (Willd.) Cabrera [=Gnaphalium pur-

pureum var. spathulatum (Lam.) Ahles]. Ruderal—lawn: rare.

Stalter, 14 Jun 1990.

Gamochaeta purpurea (L.) Cabrera. [=Gnaphalium purpureum L. var.

purpureum ]. Ruderal—lawn: freq. Stalter, 29 Mar 1990; 14 Jun

1990. Stalter 19854, 2008/10.

Heterothecasubaxillaris (Lam.) Britton & Rusby. Dunes: freq. Stalter,

26 Oct 1990. Stalter 19855, 2008/10.

Iva imbricata Walter. Dunes: infreq. Stalter, 26 Oct 1990. Stalter

20159, 2008/10.

Solidago mexicana L. [=5. sempervirens ssp. mexicana (L.) Semple].

Dunes—upper beach and dune: freq. Stalter, 26 Oct 1990.

Stalter 20160, 2008/10.

*Sonchus asper[ L.) Hill. Ruderal—disturbed soil and lawn: infreq.

Stalter, 29 Mar 1990.

*Sonchus oleraceus L. Ruderal—disturbed soil: rare. Stalter, 29 Mar

1990. Stalter 19594, 2008/10.

*Taraxacum officinale Weber ex Wigg. Ruderal—lawns and dis¬

turbed sites: freq. Stalter, 29 Mar 1990. Stalter 19595,2008/10.

Brassicaceae

Cakileedentula (Bigelow) Hook. Dunes—upper beach: freq. Stalter,

29 Mar 1990. Stalter 19857, 2008/10.

Descurainiapinnata (Walter) Britton. Ruderal—disturbed soil: freq.

Stalter, 29 Mar 1990. Stalter 19799, 2008/10.

Lepidium virginicum L. ssp. virginicum. Ruderal—disturbed soil: freq.

Stalter, 14 Jun 1990. Stalter 19798, 2008/10.

Caryophyllaceae

*Polycarpon tetraphyllum (L.) L. Ruderal—lawns and disturbed sites:

rare. Stalter, 14 Jun 1990.

Saginadecumbens (EII.)Torrey & A. Gray. Ruderal—pavement cracks

and waste places: infreq. Stalter, 29 Mar 1990.

*Stellaria media (L.) Vill. Ruderal—lawn and disturbed soil: freq.

Stalter, 29 Mar 1990. Stalter 19599, 2008/10.

Convolvulaceae

Dichondra carolinensis Michx. Ruderal—lawn: rare, only one popu¬

lation seen adjacent to casemate wall. Stalter, 26 Oct 1990.

Stalter 19858, 2008/10.

Ipomoea lacunosa L. Dunes: freq. Stalter, 14 Jun 1990. Stalter

19860, 2008/10.

Euphorbiaceae

Croton glandulosus var. septentrionalis (L.) Mull. Arg. Ruderal—dis¬

turbed soil: freq. Stalter, 26 Oct 1990. Stalter 20284, 2008/10.

Croton punctatus Jacq. Dunes: infreq. Stalter, 26 Oct 1990. Stalter

19859, 2008/10.

Euphorbia maculata L. [=Chamaesyce maculata (L.) Small]. Ru¬

deral—lawns, and disturbed sites: freq. Stalter, 14 Jun 1990;

26 Oct. Stalter 202161, 2008/10.

Euphorbia polygonifolia L. [=Chamaesyce polygonifolia (L.) Small].

Dunes—upper beach: freq. Stalter, 14 Jun 1990. Stalter 20162,

2008/10.

Fabaceae

*Medicago arabica (L.) Huds. Ruderal—disturbed soil: freq. Stalter

19596, 2008/10.

*Medicago polymorpha L. Ruderal—lawns: infreq. Stalter 19597,

2008/10.

330

*Melilotus olbus Medik. Ruderal—disburbed soil: infreq. Stalter, 14

Jun 1990. Stalter 19598, 2008/10.

*Melilotus officinalis (L.) Pallas. Dunes: infreq. Stalter, 29 Mar 1990;

14 Jun 1990.

Strophostyles helvola (L.) Ell. Dunes: infreq. Stalter, 26 Oct 1990.

Vicia sativa L. ssp. nigra (L.) Ehrh. Ruderal—disturbed soil: infreq.

Stalter, 29 Mar 1990. Stalter 19797, 2008/10.

Geraniaceae

Geranium carolinianum L. Ruderal—lawn and disturbed soil: infreq.

Stalter, 29 Mar 1990. Stalter 21163, 2008/10.

Lamiaceae

*Lamium amplexicaule L. var. amplexicaule. Ruderal—lawns and

disturbed soil: freq. Stalter, 29 Mar 1990. Stalter 19600,2008/10.

Malvaceae

Modiola caroliniana (L.) G. Don. Ruderal—seen only at border

between lawn and fort: rare. Stalter, 29 Mar 1990. Stalter

20164, 2008/10.

Onagraceae

Oenothera drummondii Hook. ssp. drummondii. Dunes: freq. but

rare in the Carolinas (Radford et al. 1968). Stalter, 29 Mar 1990.

Stalter 20165, 2008/10.

Oenothera laciniata Hill. Dunes: freq. Stalter, 29 Mar 1990. Stalter

20168, 2008/10.

Oxalidaceae

Oxalis dillenii Jacq. Ruderal—disturbed soil: infreq. Stalter, 29 Mar

1990. Stalter 20167, 2008/10.

Phytolaccaeae

Phytolacca americana L. Ruderal—disturbed soil and dune: infreq.

Stalter, 14 Jun 1990. Stalter 19601,2008/10.

Plantaginaceae

Nuttallanthus canadensis (L.) D.A. Sutton [=Linaria canadensis (L.)

Dum. Cors.]. Ruderal—disturbed soil and dune: freq. Stalter, 29

Mar 1990. Stalter 20285, 2008/10.

*Plantago lanceolata L. Ruderal—disturbed soil and lawn: freq.

Stalter, 29 Mar 1990. Stalter 20286, 2008/10.

*Veronicaarvensis L. Ruderal—lawns and disturbed soil: freq. Stalter,

29 Mar 1990. Stalter 19602, 2008/10.

Veronicaperegrina L. Ruderal—disturbed soil: infreq. Stalter, 29 Mar

1990. Stalter 19603, 2008/10.

Polygonaceae

*Rumexcrispus L. ssp. crispus. Ruderal—disturbed soil: rare. Stalter,

29 Mar 1990. Stalter 19795, 2008/10.

Portulacaceae

*Portu!aca pilosa L. Ruderal—lawns and disturbed soil: infreq.

Stalter, 14 Jun 1990. Stalter 20169, 2008/10.

Rubiaceae

Diodia feres Walter. Ruderal—disturbed sands: freq. Stalter, 14 Jun

1990. Stalter 20287, 2008/10.

Journal of the Botanical Research Institute of Texas 8(1)

LILIOPSIDA (MONOCOTS)

Agavaceae

*Yucca aloifolia L. Dunes: infreq. Stalter, 29 Mar 1990. Stalter 19566,

2008/10.

Cyperaceae

Cyperus compressus L. Dunes: rare. Stalter 20270, 2008/10.

Cyperus croceus Vahl. [=C. globulosus AublJ. Dunes: rare. Stalter

20271,2008/10.

Cyperus gray/Torr. Dunes: rare. Stalter 20272, 2008/10.

Iridaceae

Sisyrinchium rosulatum Bickn. Ruderal—lawn: infreq. Stalter 19570,

2008/10.

Poaceae

*Bromus catharticus Vahl. var. cartharticus [=B. wildenowii Kunth.].

Ruderal—disturbed soil: rare. Stalter, 29 Mar 1990.

Cenchrus longispinus (Hack.) Fernald. Dunes: infreq. Stalter, 14 Jun

1990. Stalter 20273, 2008/10.

Cenchrus tribuloides L. Dunes: freq. Stalter, 14 Jun 1990. Stalter

20274, 2008/10.

*Cynodon dactylon (L.) Pers. Ruderal—lawn: freq. Stalter, 29 Mar

1990; 14 Jun 1990. Stalter 19791,2008/10.

*Digitaria sanguinalis (L.) Scop. Ruderal—lawn and disturbed soil:

freq. Stalter, 14 Jun 1990. Stalter 19792, 2008/10.

*Eleusine indica (L.) Gaertn. Ruderal—lawn and disturbed soil:

infreq. Stalter, 29 Mar 1990. Stalter 20275, 2008/10.

Eustachyspetraea (Sw.) Desv. [=ChlorispetraeaSw.\. Ruderal—lawn

and disturbed soil: rare. Stalter, 29 Mar 1990.

*Loliumperenne L. var. aristatum. Ruderal—lawn: infreq. Stalter, 14

Jun 1990. Stalter 19793, 2008/10.

Panicum amarum Ell. Dunes: freq. Stalter, 26 Oct 1990. Stalter

19794, 2008/10.

*Paspalum dilatatum Poir. ssp. dilatatum. Ruderal—disturbed soil

outside fort: freq. Stalter, 14 Jun 1990. Stalter 20277, 2008/10.

Paspalum notatum Flugge. Ruderal—disturbed soil outside fort:

freq. Stalter, 14 Jun 1990. Stalter 20278, 2008/10.

*Poa annua L. Ruderal—lawn: freq. Stalter, 29 Mar 1990. Stalter

19572, 2008/10.

Spartina alterniflora Loisel. Salt Marsh: freq. Stalter, 26 Oct 1990.

Stalter 19789, 2008/10.

Spartina patens (Ait.) Muhl. Dunes: freq. Stalter, 14 Jun 1990. Stalter

19790, 2008/10.

*Sporobo!us junceus (P. Beauv.) Kunth. [=5. poiretii (Roem. & Schult.)

Hitchc.]. Ruderal—lawns and disturbed soil: freq. Stalter, 26 Oct

1990. Stalter 20279, 2008/10.

Triplasispurpurea (Walter) Chapm. var. purpurea. Dunes: freq. Stalter,

26 Oct 1990. Stalter 19788, 2008/10.

Uniola paniculata L. Dunes: freq. Stalter, 26 Oct 1990. Stalter 19787,

2008/10.

Vulpia octoflora (Walter) Rydb. var. octoflora [=Festuca octoflora

Walter]. Ruderal—lawn and disturbed soil: infreq. Stalter, 14

Jun 1990. Stalter 20276, 2008/10.

ACKNOWLEDGMENTS

The authors thank Bill Dorance, Fort Sumter/Fort Moultrie National Monument for hospitality during our

visits and assistance with boat transportation to and from Fort Sumter and many other details; Gordon Tucker

and Jim Montgomery for determinations of the Cyperaceae and Pteridophytes; undergraduate research stu¬

dents at St. John’s University for assistance in processing voucher material. This paper was improved thanks to

the helpful comments and editorial suggestions of three anonymous reviewers and suggestions from Barney

Lipscomb.

Stalter et al., Flora of Fort Sumter and Fort Moultrie

331

REFERENCES

Baden, J., W.T. Batson, & R. Stalter. 1975. Factors affecting the distribution of vegetation of abandoned rice fields, George¬

town County, South Carolina. Castanea 40:171-184.

Carter, R., W.W. Baker, & M.W. Morris. 2009. Contributions to the flora of Georgia, U.S.A. Vulpia 8:1 -54.

Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 16+ vols. New York and

Oxford.

Gleason, FI.A. & A. Cronquist. 1991. Manual of vascular plants of northeastern United States and adjacent Canada, 2nd ed,

The New York Botanical Garden, Bronx, NY.

Kartesz, J.T. 1994. A synonymized checklist of the vascular flora of United States, Canada, and Greenland. 2nd ed. Volume

1-Checklist.Timber Press, Inc. Portland, OR.

Radford, A.E., H.E. Ahles, & C.R. Bell. 1968. Manual of the vascular flora of the Carolinas, 2nd ed. The University of North

Carolina Press, Chapel Hill.

Schmidt, J.P. 2004. Vascular plant survey of Fort Frederica National Monument, Fort Sumter/Fort Moultrie National Monu¬

ment and Charles Pinckney National Historic Site. Institute of Ecology, University of Georgia, Athens, Georgia 30606.

Stalter, R. 1973. Factors affecting vegetation distribution in the Cooper River Estuary, South Carolina. Castanea 38:18-24.

Stalter, R. & E.E. Lamont. 1993. The vascular flora of Fort Sumter and Fort Moultrie, South Carolina, one year after Hur¬

ricane Hugo. Castanea 58:141 -152.

Stephens, P.F. (2001 onwards). Angiosperm Phylogeny Website. http://www.mobot.org/MOBOT/research/APweb/. Ver¬

sion 12, July 2012 [and more or less continuously updated since]. Accessed 4 November 2013.

Windham, M.D. 1993. New taxa and nomenclatural changes in the North American fern flora. Contr. Univ. Michigan Herb.

19:31-61.

Wunderlin, R.P. 1998. Guide to the vascular plants of Florida. University Press of Florida, Gainesville. P. 806.

332

Journal of the Botanical Research Institute of Texas 8(1)

BOOK NOTICE

Dennis Wm. Stevenson, Roy Osborne, and Alberto Sidney Taylor Blake, eds. 2012. Proceedings of Cycad 2008:

The 8th International Congress on Cycad Biology, 13-15 January 2008, Panama City, Panama.

(ISBN-13: 978-0893275150, hbk). Memoirs of the New York Botanical Garden, Volume 106. The New York

Botanical Garden Press, 2900 Southern Boulevard, Bronx, New York 10458-5126, U.S.A. (Orders: http://

nybgpress.org, 1-718-817-8721, nybgpress@nybg.org). $95.00, 554 pp., b&w and color figures, scientific

name index, subject index, 7" x 10".

From the publisher: Cycads have been around since the Late Permian. During this time they have retained

seemingly unchanged features, yet they have evolved unique characteristics that science continues to uncover.

As the discoveries keep coming, medicine and agriculture find new applications for what cycads have to offer,

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The International Conference on Cycad Biology, held every three years since its inception in 1987, is a

prominent meeting centering on all aspects of the biology of this unusual, important, and interesting group of

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Having survived for 250 million years, cycads now find themselves endangered as a result of human ac¬

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J.Bot. Res. Inst. Texas 8(1): 332.2014

EXPANDED DISTRIBUTION OF GRATIOLA QUARTERMANIAE

(PLANTAGINACEAE) IN TEXAS, U.S.A.

Kimberly Norton Taylor and Robert J. O'Kennon

Botanical Research Institute of Texas

1700 University Drive

Fort Worth, Texas 76107, U.S.A.

ktaylor@brit.org, okennon@brit.org

ABSTRACT

Forty-nine new populations of Gratiola quartermaniae are reported from seven new counties in north central Texas. All populations were

found on seasonally wet Walnut Timestone glades and are the first to be found in the Texas Grand Prairie and Timestone Cut Plain. These

collections indicate that the species and its calcareous glade habitat are much more abundant in Texas than previously thought.

RESUMEN

Se citan cuarenta y nueve poblaciones nuevas de Gratiola quartermaniae de siete condados nuevos en el centro norte de Texas. Todas las

poblaciones se encontraron en los claros estacionalmente humedos de Walnut Timestone y son las primeras que se encuentran en la Grand

Prairie de Texas y Timestone Cut Plain. Estas colecciones indican que la especie y su habitat calcareo son mucho mas abundantes en Texas

de lo que se creia previamente.

Gratiola quartermaniae D. Estes (Plantaginaceae) was first described from Eastern North America by Estes and

Small in 2007. The species typically grows in thin, seasonally saturated soil over exposed limestone or dolo¬

mite bedrock (Estes & Small 2007; Taylor & Estes 2012). This habitat is typically found associated with lime¬

stone glades, barrens, prairies, and alvars. Estes and Small (2007) note that the species is most common in the

limestone cedar glades of central Tennessee and northern Alabama, though it also occurs in primarily calcare¬

ous habitats in northeastern Illinois, central Texas, and southeastern Ontario. Similar disjunction patterns are

seen in several other calciphilous species which grow in association with G. quartermaniae including Clinopo-

dium arkansanum (Nutt.) House, Grindelia lanceolata Nutt, Heliotropium tenellum (Nutt.) Torr., Isoetes butleri

Engelm., Juncusfilipendulus Buckl., and Minuartiapatula (Michx.) Mattf.

Gratiola quartermaniae is known from five collections in Texas (Fig. 1) including one each in Bell (Wolff

2317 , SMU) and Llano (W hitehouse 18477 , SMU, UC, US) counties, and three in Williamson County ( Bodin s.n.,

PH, MIN; Turner & Turner 122 , BRIT, MO, TENN, TEX; Turner & Turner 119, BRIT, GH, MO, TENN, TEX).

While the label data do not indicate the exact location of each site, the Williamson and Bell county specimens

appear to occur in calcareous habitats, which is typical for the taxon. The Llano County site appears to be an

exception, apparently occurring on granite.

In 2007, O’Kennon discovered a population of what he later determined to be Gratiola quartermaniae

from a calcareous Walnut Limestone glade seep in Wise County, Texas ( O’Kennon 20515B, BRIT). This repre¬

sents the first documented population for G. quartermaniae in north central Texas, a disjunction of approxi¬

mately 230 km to the north of the populations in central Texas. Despite the large disjunction, the occurrence

of G. quartermaniae in the Grand Prairie is not surprising. The predominately Cretaceous limestone substrate

that comprises the Edwards Plateau extends northward through the Limestone Cut Plain (Lampasas) and

Grand Prairie (Griffith et al. 2004). With these large extensions of similar, and in some cases identical, sub¬

strate it is not surprising that we find calciphilous species with distributions extending farther to the north

than previously thought (Swadek & Burgess 2012; Taylor et. al 2012; Taylor & O’Kennon 2012).

In the spring of 2012, the authors conducted extensive surveys of limestone prairies, barrens, and glades

in the Grand Prairie and northern Limestone Cut Plain of north central Texas. Areas where limestone outcrops

form “glades” reminiscent of the cedar glades found in the Central Basin of Tennessee were searched for G.

J. Bot. Res. Inst. Texas 8(1): 333 - 337.2014

334

Journal of the Botanical Research Institute of Texas 8(1)

Legend

(*) New Gratiola quartermaniae collections

/C Previous Gratiola quartermaniae collections

Walnut Limestone

□ Cross Timbers Level 3 Ecoregion

Level 4 Ecoregions

| Balcones Canyonlands

y\ Eastern Cross Timbers

| Edwards Plateau Woodland

| Grand Prairie

■ Limestone Cut Plain

Fig. 1. Documented collection sites of Gratiola quartermaniae in Texas. Forty-nine previously unknown sites were documented in 2012 from Walnut

Limestone glade seeps on the western edge of the Grand Prairie and the northern Limestone Cut Plain (McGowen et al. 1987; McGowen et al. 1991;

Griffith etal. 2004).

quartermaniae. Satellite imagery and geologic maps were used to identify other possible locations for explora¬

tion. During this search, 49 new locales were identified for G. quartermaniae. These collections represent 7 new

county records, including Bosque, Denton, Hood, Johnson, Parker, Tarrant, and Wise counties (Fig. 1). At least

one voucher specimen from each county was collected and deposited at BRIT. Specific collection information

for each specimen is available online at atrium.brit.org.

All sites in north central Texas are underlain by Walnut Limestone substrate. This Cretaceous, fossilifer-

ous, erosion resistant limestone forms glades when exposed. In the Grand Prairie these glades form at lower

topographic positions as the overlying substrate erodes (Fig. 2). In the Limestone Cut Plain, Walnut Limestone

glades are found encircling the Edwards or Comanche Peak Limestone mesas that characterize the region (Hill

1901). Seepage from the interbedded limestone and marl layers upslope will often form shallow pools or

streams over exposed Walnut bedrock at the base of the slopes (Swadek & Burgess 2012).

Gratiola quartermaniae was found in very shallow soil directly over limestone bedrock on the edges and

on small soil islands in limestone bedrock creeks and seeps, or in deeper, mucky limestone soil in disturbed

sites including highly grazed cattle ponds (Fig. 3). Gratiola quartermaniae is most abundant in areas with little

competing vegetation or in areas with high levels of disturbance.

The majority of the sites were found on relatively undisturbed, seasonally saturated seeps and streambeds

with little to no soil accumulation and large amounts of exposed limestone bedrock (Fig. 3). The remaining

sites were found in deeper soils over Walnut Limestone but were highly disturbed. Most deeper soil sites were

Taylor and O'Kennon, Distribution of Gratiola quartermaniae in Texas

335

Walnut Limestone

Grand Prairie

Walnut Limestone v glade formation

\ /

Lampasas Cut Plain

Fig. 2. Diagrammatic section of Grand Prairie (top) and Limestone (Lampasas) Cut Plain (bottom), showing position of Walnut Limestone outcrops across

the landscape. In the Grand Prairie Walnut Limestone glades form when Walnut Limestone bedrock is exposed. In the Lampasas Cut Plain glades are

found near the base of the many mesas throughout the region. Diagrams modified from Figures 3 and 5 by Hill (1901).

heavily grazed by livestock and were highly trampled. We believe this high level of disturbance eliminates

competitors allowing G. quartermaniae to persist in the deeper soils. Most populations were small and isolated

with fewer than 50 plants, though a few sites had well over 500 plants. Associated species include Eleocharis

occulta S.G. Sm., Hypoxis hirsuta (L.) Coville, Spiranthes magnicamporum Sheviak, Isoetes butleri Engelm., Noth-

oscordum bivalve (L.) Britton, Allium canadense L. var .fraseri Ownbey, Juncus filipendulus Buckl., Fuirena sim¬

plex Vahl var. simplex, and Leucospora multifida (Michx.) Nutt.

Plants flower from late March through May, and set seed from April through early June. The plant dries

and disappears completely by early summer. This phenology closely matches the hydrological regime of the

limestone glades where the species is found. Saturated conditions in the spring give rise to near drought-like

conditions in the summer and fall as temperatures rise and rainfall decreases. Gratiola quartermaniae appears

to be specifically adapted to complete its life cycle within this narrow hydrological window.

Gratiola quartermaniae is much more widespread in Texas than previously thought. This oversight can be

attributed to the short lifecycle of the species and the lack of thorough botanical exploration in the limestone

prairies of north central Texas. Until recently, the limestone glade habitat, which is prolific on the western edge

of the Grand Prairie, remained relatively unexplored botanically. This is evident by the recent discovery or range

expansion of several species characteristic of glades in the region, including Isoetes butleri (Taylor et. al 2012),

Phemeranthus calcaricus (Engelm.) Kiger (Swadek 2012), and Dalea reverchonii (S. Wats.) Shinners (Taylor &

O’Kennon 2012). Additional exploration of the Grand Prairie and Limestone Cut Plain regions of Texas would

likely reveal the presence of additional glade taxa and more locations for G. quartermaniae. In particular, the

Walnut Limestone formation is quite extensive in the Limestone Cut Plain and warrants further exploration.

Representative Voucher Specimens.—U.S.A. TEXAS. Bosque Co.: Co Rd 2650 ca. 0.7 mi SE of Co Rd 2660, ca. 3.5 air mi NW of Walnut

Springs. 32.087772N, -97.795782W, elev 297 m, abundant along ephemeral glade stream, 27 Apr 2012, Norton & O’Kennon 1673 (BRIT).

Denton Co.: N County Line Rd ca. 0.25 mi S of Hwy 380, ca. 70 m E, Walnut Limestone seep glade, 33.242092N, -97.38987W, elev 226 m, 21

Apr 2012, O’Kennon 24963 (BRIT). Hood Co.: Running Deer Ct ca. 0.8 mi N Cleburne Hwy, Walnut Limestone glade, 32.433357N,

-97.621662W, elev 229 m, 27 Apr 2012, Norton & O’Kennon 1676 (BRIT). Johnson Co.: Running Deer Ct at Cleburne Hwy, Walnut Limestone

glade, 32.427675N, -97.615694W, elev 240 m, 27 Apr 2012, Norton & O’Kennon 1675 (BRIT). Parker Co.: Old Agnes Rd ca. 0.1 mi N of Louis

Scherer Rd on E side of rd, 32.846977N, -97.778998W, elev 355 m, growing in thin soil over Walnut Limestone bedrock, seasonally wet,

abundant, 10 Apr 2012, Norton & O’Kennon 1525 (BRIT). Tarrant Co.: White Settlement, Verna Trail N ca. 0.2 mi N of Stubbs Trail,

32.77301N, -97.5015W, elev 216 m, growing in ephemeral swale over Walnut Limestone, 13 Apr 2012, Norton & O’Kennon 1548 (BRIT). Wise

Co.: Hwy 114 ca. 0.6 mi W of hwy 81 at Rhome. N side of rd, 33.060325N, -97.488865W, elev 247 m, seasonally wet seepage area below pond

damn, thin soil over Walnut Limestone bedrock, growing with Isoetes butleri, Fuirena simplex, and Leucospora multifida, 13 May 2007,

O’Kennon 20515B (BRIT); 8 May 2012, Norton & O’Kennon 1689 (BRIT).

336

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 3. Ephemeral limestone seep with exposed Walnut Limestone bedrock, Denton Co. {O'Kennon 24971, BRIT) (top); Gratiola quartermaniae growing

in shallow soil overtop exposed bedrock. Wise Co. {Norton & O'Kennon 1539, BRIT) (left); close up of 6. quartermaniae, Parker Co. {Norton & O'Kennon

1511, BRIT) (right).

Taylor and O'Kennon, Distribution of Gratiola quartermaniae in Texas

337

ACKNOWLEDGMENTS

We thank Austin Sewell from the Lyndon B. Johnson National Grasslands for his assistance in locating sites

and Dwayne Estes for his assistance in verifying species identification. We thank Dwayne Estes, Allan Nelson,

and Rebecca Swadek for their helpful reviews of the manuscript. We also thank all the landowners who grant¬

ed us permission to collect on their property.

REFERENCES

Estes, D. & R.L. Small. 2007. Two new species of Gratiola (Plantaginaceae) from eastern North America and an updated

circumscription for Gratiola neglecta. J. Bot. Res. Inst. Texas 1:149-170.

Griffith, G.E., S.A. Bryce, J.M. Omernik, J.A. Comstock, A.C. Rogers, B. Harrison, S.L. Hatch, & D. Bezanson. 2004. Ecoregions of

Texas, U.S. Environmental Protection Agency, Corvallis, Oregon. 1:2,500,000.

Hill, R.T. 1901. Geography and geology of the Black and Grand Prairies, Texas. U.S. Geological Survey. Annual Report 21.

United States Geological Survey, Washington, D.C., U.S.A.

McGowen, J.H., C.V. Proctor Jr., W.T. Haenggi, D.F. Reaser, & V.E. Barnes. 1987. Geologic atlas of Texas, Dallas Sheet (Gayle

Scott Memorial Edition). In: V.E. Barnes, ed. Geologic atlas of Texas. Bureau of Economic Geology, University of Texas

at Austin, Austin. 1:250,000.

McGowen, J.H., T.F. Hentz, D.E. Owen, M.K. Pieper, C.A. Shelby, & V.E. Barnes. 1991. Geologic atlas of Texas, Sherman Sheet

(Walter Scott Adkins Memorial Edition). In: Geologic atlas of Texas, V.E. Barnes, ed. Bureau of Economic Geology,

University of Texas at Austin, Austin. 1:250,000.

Swadek, R.K. &T.L. Burgess. 2012. The vascular flora of the north central Texas Walnut Formation. J. Bot. Res. Inst. Texas

6:725-752.

Swadek, R.K. 2012. Phemeranthus calcaricus (Montiaceae) new to Texas. J. Bot. Res. Inst. Texas 6:303-307.

Taylor, K.N. & D. Estes. 2012. The floristic and community ecology of seasonally wet limestone glade seeps of Tennessee

and Kentucky. J. Bot. Res. Inst.Texas 6:711-742.

Taylor, K.N. & RJ. O'Kennon. 2013. Ecology and distribution of the north central Texas endemic Dalea reverchonii

(Fabaceae). J. Bot. Res. Inst. Texas 7:603-610.

Taylor, K.N., RJ. O'Kennon, &T.F. Rehman. 2012. Expanded distribution of Isoetes butleri (Isoetaceae) in Texas. J. Bot. Res.

Inst. Texas 6:753-757.

338

Journal of the Botanical Research Institute ofTexas 8(1)

ANNOUNCEMENT

LANDON E. McKINNEY

(1949-2014)

Source of information: www.schoedinger.com

Landon E. McKinney, age 65, passed away on Thursday, June 5th, 2014 at the Wade Park Veteran’s Medical

Center in Cleveland. He was born May 17, 1949 in Nashville, Tennessee, to the late Lawrence Vern and Con¬

stance Joy McKinney.

A proud Vietnam Veteran of the United States Navy, Mr. McKinney was a Marine Navy Corpsman. He is

survived by his loving wife of 31 years, Lela McKinney; children Adam McKinney, Amanda (Troy) Chitwood,

and Eric (Amy) McKinney; grandchildren Landon, Lucas, Erica, Katelyn, and Whitney; many extended family

members, special friends, and colleagues.

Landon was a friend and colleague and published violet (Viola) articles in Sida and Journal of Botanical

Research Institute of Texas. Landon published his monograph of the acaulescent blue violets in Sida, Botanical

Miscellany (see reference below).— Barney Lipscomb

McKinney, L.E. 1992. A Taxonomic Revision of the Acaulescent Blue Violets (Viola) of North America. Sida, Bot. Misc. 66

pp., 28 b/w figs., photographs, maps.

We plan to have a more complete tribute to Landon in a forthcoming issue.

J. Bot. Res. Inst. Texas 8(1): 338.2014

THE VASCULAR FLORA OF GALVESTON ISLAND STATE PARK,

GALVESTON COUNTY, TEXAS, U.S.A.

David J. Rosen and Shiron K. Lawrence

Department of Biology

Lee College

Baytown, Texas 77522-0818, U.S.A.

drosen@lee.edu

Natural Resources, State Parks

Texas Parks and Wildlife Department

14200 Garrett Road

Houston, Texas 77044, U.S.A.

andrew.sipocz@tpwd. texas.gov

Andrew Sipocz

ABSTRACT

Galveston Island State Park is located near the center of Galveston Island in Galveston County, Texas, U.S.A. The 789.1 ha park lies within

the Western Gulf Coastal Plain ecoregion. A floristic survey was conducted from July 2011 through November 2013, and vouchered speci¬

mens at TEX collected in the early 70s by R.J. Fleetwood verified, with the goal of assembling an annotated checklist of vascular plants. This

resulted in a checklist of 317 species of vascular plants representing 68 families and 221 genera. The largest families were Poaceae (65 spp.),

Asteraceae (36 spp.), Fabaceae (25 spp.), Cyperaceae (24 spp.), and Amaranthaceae (11 spp.). Non-native species account for 16.4% of the

total flora. Seven species in six different families are of conservation interest in that they are endemic to the ecoregion. Fists of species char¬

acteristic of readily recognizable habitat types are provided.

RESUMEN

El Galveston Island State Park esta localizado cerca del centro de la isla Galveston en el condado de Galveston, Texas, U.S.A. Es parque de

789.1 ha esta en la ecoregion de Western Gulf Coastal Plain. Se realizo un estudio floristico de julio de 2011 a noviembre de 2013, y se veri-

ficaron especimenes testigo en TEX collectados a principios de los 70 por R.J. Fleetwood, con el objetivo de realizar un catalogo anotado de

plantas vasculares. Asi se obtuvo una catalogo de 317 especies de plantas vasculares que representan 68 familias y 221 generos. Las familias

mas numerosas fueron Poaceae (65 spp.), Asteraceae (36 spp.), Fabaceae (25 spp.), Cyperaceae (24 spp.), y Amaranthaceae (11 spp.). Las es¬

pecies no nativas fueron el 16.4% de la flora total. Siete especies de seis diferentes familias son de interes para la conservacion ya que son

endemicas de la ecoregion. Lists of species characteristic of readily recognizable habitat types are provided.

INTRODUCTION

Galveston Island State Park is located near the center of Galveston Island, one of a series of barrier islands and

bay/lagoon systems separating most of the Texas mainland from the Gulf of Mexico (Fig. 1). The Texas coast

comprises most of the area designated by Griffith et al. (2004) as the Western Gulf Coastal Plain ecoregion.

The 789.1 ha (1,950 acre) park straddles the Island from the Gulf to West Galveston Bay and is roughly square

in overall dimensions (Fig. 2). Galveston Island’s formation is recent, beginning as a submerged offshore bar no

more than 4,500 to 5,000 years ago that accumulated until about 1900 when it reached its maximum width

(Garner 1997). The bayside region consists of salt marsh fringing a series of peninsulas and intervening la¬

goons perpendicular to the Island’s long axis (Fig. 2). A series of dune ridges and swales parallel the Island’s

long axis from its center to the beach (Fig. 2). These formed as the Island accreted seaward and became too

wide and high for significant wash-over events. The ridges and swales support prairie and freshwater wetlands

respectively. All of these features are well represented in the park.

Galveston Island has been narrowing through erosion and apparent sea level rise since approximately

1900. This is due to several factors including the construction of jetties at the mouth of Galveston Bay which

block the longshore drift and sand supply to the island’s beaches, subsidence of the Island from subsurface

fluid withdrawal, and eustatic sea level rise (Raven et al. 2009). Since the park’s establishment in 1972, its

beach has moved inland approximately 74.1 m (243 feet; Sipocz 2010). The active dune system has been com¬

pletely displaced since the park’s establishment and now overlies what had previously been wetlands or devel¬

oped facilities (Sipocz 2010).

J. Bot. Res. Inst. Texas 8(1): 339 - 352.2014

340

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 1. General location of Galveston Island State Park, Galveston County, Texas.

Climate

Galveston Island occurs within a humid, warm-temperate to marginally subtropical region receiving an aver¬

age of 129 cm (51 in) of rainfall annually (Britton & Morton 1989; National Weather Service 2013). The average

temperature for Galveston Island is 21.8°C (71.2°F), with August being the warmest month (29°C, 85°F) and

January the coldest (12.9°C, 55.2°F; National Weather Service 2013). The cooling effect of sea-breezes from the

Gulf of Mexico moderates summer temperatures (Crenwelge et al. 1988). The predominant wind direction is

southeast, but varies by season. The light southerly winds predominate in spring through early fall, while cold

fronts with strong northerly winds that push water out of the bay punctuate the winter. Tropical storms and

hurricanes are frequent on the Texas coast, striking with a frequency of 0.67 storms per year (Hollingsworth

1998).

Soils and Vegetation

Four different soil series occur at Galveston Island State Park, and all are derived from the inland deposition of

Rosen etal., Flora of Galveston Island State Park, Texas

341

Fig. 2. True-color aerial photograph of Galveston Island State Park (outlined in green), Galveston County, Texas.

beach sands (Fig. 3; Crenwelge et al. 1988). The Karankawa Mucky Loam Series is a bayside salt marsh soil

formed on over-wash-deposited sands with a high amount of partially decayed organic matter resulting from

plant growth coupled with anaerobic soil conditions (Crenwelge et al. 1988).

Prominent dune ridges in the interior of the park consist of Galveston Loamy Fine Sand and Galveston

Fine Sand Series soils (Fig. 3). These are wind accumulated, mildly alkaline, somewhat excessively drained

soils that are very rapidly permeable with fine sand in the upper 183 cm (72 in) to 356 cm (140 in) and support

prairie vegetation (Crenwelge et al. 1988). Upland prairie on Texas’ barrier islands is regionally referred to as

“strand prairie” (Hollingsworth 1998). We are uncertain as to the origin of this seemingly colloquial name, but

it is clear that Hollingsworth (1998) used it to refer to Schizachyrium littorale (Nash) E.P. Bicknell- Paspalum

monostachyum dominated grasslands of Texas’ barrier islands and Coastal Sand Plain (Diamond et al. 1987).

This community is considered an ally of coastal prairie marked by tolerance of occasional tidal over-wash, and

probably more importantly, the salt spray typical of the spring through fall months which produces measur-

342

Journal of the Botanical Research Institute of Texas 8(1)

] Kilometers

Galveston Island State Park

SSURGO SOIL NAME

[GaBj GALVESTON FINE SAND

[GsJ GALVESTON LOAMY FINE SAND

■a KARANKAWA MUCKY LOAM

|Ms] MUSTANG FINE SAND

Mt MUSTANG-NASS COMPLEX

Ns NASS VERY FINE SANDY LOAM

WATER

Fig. 3. Map of major soil series of Galveston Island State Park, Galveston County, Texas.

able soil salinity throughout the park and accumulates on plant surfaces during dry periods. The Galveston

Loamy Fine Sand Series is uniquely underlain by a shell layer and portions of it support live oak (Quercus vir-

giniana) woodlands known colloquially as “live oak mottes.” Just east of the park, a live oak motte on this soil

type contains an extensive Native American burial ground which was described by early Spanish explorers

indicating that the presence of trees on the Island predates European settlement (Ricklis et al. 1994). The

Galveston Fine Sand Series also supports strand prairie but is treeless, not underlain by shell fragments, and

lies closer to the beach.

The majority of the terrestrial park lands consist of the Mustang Fine Sand Series. This strongly alkaline,

somewhat poorly drained, very slowly permeable soil occurs on flats or slightly convex or concave surfaces and

although it includes dune ridges and swales, they are poorly developed (Crenwelge et al. 1988). These soils are

a mixture of wind and washover deposits of slightly coarser sands. The water table is close to and occasionally

at the surface of these soils; normally 25.4 cm (10 in) to 76.2 cm (30 in) in depth (Crenwelge et al. 1988). Vegeta-

Rosen etal., Flora of Galveston Island State Park, Texas

343

tion composition within the Mustang Soil region changes considerably with surface slope and configuration.

The water table elevation follows surface topography and so lies at different elevations throughout the Park

(Lambert 1998). Topographical changes control depth to the water table and affect soil alkalinity, drainage, and

permeability. Therefore, even very slight topographical and slope changes result in markedly different plant

communities. Concave surfaces contain what Texas’ Natural Heritage Program (1993) termed a “Gulf Cord-

grass Series” grassland dominated by Spartina spartinae and S. patens , while convex surfaces support strand

prairie.

Nass Very Fine Sandy Loam Series soils are neutral to moderately alkaline soils in non-tidal swales and

occur as inclusions in each of the three prairie soil types. The water table extends to or above the soil surface in

most years and a permanent water table lies within 15.2 cm (6 in) to 70 cm (24 in) of the surface (Crenwelge et

al. 1988). They are the remainder of lagoons formed as the Island accreted seaward and offshore bars accumu¬

lated sand, rising to form new dune ridges parallel to the beach. The older swales have been broken by over¬

wash deposition and wind erosion into strings of circular, freshwater ponds up to 1.8 m (6 ft) in depth. These

are often surmounted by crescent-shaped dunes along leeward sides that may contain small live oak mottes.

Resource Management

Resource management at the park strives to conserve and restore its pre-European settlement landscape, and

terrestrial and aquatic plant and wildlife communities. Prior to becoming a State Park in 1972 the site had been

intensively grazed by fenced cattle, greatly reducing the dominance of tall grasses such as little bluestem

(.Schizachyrium scoparium) that are typical of strand prairie (Keith 2005). In addition, much of the park had

been invaded by non-native plants including Chinese tallow-tree ( Triadica sebifera) and Japanese honeysuckle

(Lonicera japonica). Mowing, mechanical chipping, herbicide, and prescribed fire have been used to almost

eliminate these species and slowly increase those described as dominants in strand prairie.

Prescribed and wildfires have been documented in the park since 1976 (Creacy 2007). Between 1976 and

2006, when a more rigorous prescribed fire program was implemented, 11 fires burned a total of 985.8 ha

(2,436 acres), mostly within the 356.1 ha (880 acres) containing strand prairie and gulf cordgrass plant com¬

munities though the results were poorly documented. Since 2006 prescribed fires applied to the prairies have

been better documented including the use of permanent vegetation plots for effects monitoring. The entire

prairie area of the park has been burned at least once during the 2007 to present time period.

Present resource management includes the continued use of prescribed fire to burn the prairies on an ap¬

proximate 3 to 7 year rotation as well as spot treatment of non-native, invasive plant species including Guinea

grass ( Megathyrsus maximus), Chinese tallow-tree, Vasey grass ( Paspalum urvillei), black locust ( Robinia pseu¬

doacacia), cabbage palm ( Sabal palmetto), and Mexican fan palm ( Washingtonia robusta). Mechanical treat¬

ments are no longer used as it is thought to spread invasive species into the prairies, and they have been made

unnecessary by the more rigorous use of prescribed fires.

Other management activities include the propagation and planting of species that are uncommon or ab¬

sent from the park’s prairies, but are common to strand prairie on an unfenced reference site on the adjacent

barrier island just south of Galveston. Indeed, a focus on the restoration of the park’s prairies revealed the need

for an intensive baseline inventory of vascular plant species that occurred there. RaymondJ. Fleetwood, a U.S.

Fish & Wildlife Service biologist who worked on the Texas coast beginning in the 1960s, performed what was

probably the first and only effort to survey the vascular plants at Galveston Island State Park, compiling a list

comprising 108 species distributed in 96 genera and 39 families (Fleetwood 1973). Fleetwood also vouchered

a modest exsiccatae of 63 specimens at the University of Texas at Austin Plant Resources Center Herbarium

(TEX). Interest in restoring the park’s prairie flora and continuing the work that Fleetwood began four decades

ago has culminated in the study reported here.

METHODS

Twenty collecting trips were made to the park from July 2011 through November 2013, and in all months ex¬

cept December-February. A complete set of voucher specimens is housed at TEX. Plant identifications were

344

Journal of the Botanical Research Institute of Texas 8(1)

Table 1. Taxonomic summary of vascular plants of Galveston Island State Park, Galveston County, Texas.

Families

Genera

Native

Species

Non-native

Total

Monocots

117

Eudicots

154

171

200

Totals

221

265

317

primarily made using the appropriate volumes of Flora of North America (1993+), Correll and Johnston (1970),

and Gould (1975). When practical, infraspecific names were determined. Classification and author names fol¬

low Tropicos (2013). Nativity and any special conservation status of each species were determined by review of

Flora of North America (1993+) and Correll and Johnston (1970). We follow Nesom (2000) in defining non-na¬

tive species as those originating from a different continent or less commonly in this flora cultivated from out¬

side the geographic extent of the Western Gulf Coastal Plain ecoregion (Griffith et al. 2004).

RESULTS & DISCUSSION

A combination of held work from July 2011 through November 2013 and review of Fleetwood’s specimens

housed at TEX yielded 317 species of vascular plants representing 68 families and 221 genera (Table 1). The

families containing the largest number of species (native and non-native combined) are Poaceae (65 spp.),

Asteraceae (36 spp.), Fabaceae (25 spp.), Cyperaceae (24 spp.), and Amaranthaceae (11 spp.). Large genera in¬

clude Cyperus with nine native and one non-native species, and Juncus with eight native species. Non-native

species account for 16.4% of the total flora. Poaceae included the most non-native species (16). Two families,

Arecaceae and Tamaricaceae, are represented by only non-native species. The two Arecaceae species are not

known to be invasive in Texas (Nesom 2009), but have begun rapidly reproducing on Galveston Island, likely

because of warmer winters (Britton & Morton 1989; National Weather Service 2013), and are a new threat to

the prairie habitats. Seven species in six different families are of conservation interest in that they are endemic

to the ecoregion.

Plant Communities

No effort was made to quantitatively describe plant communities that occur in the park. However, due to the

influence of soil types, wind, wave, tidal action, and sometimes sharp elevation gradients across the landscape,

several habitat types are readily identified in the held simply because the resulting species composition is so

consistent (Fig. 4, Table 2). We believe we provide nearly complete lists of characteristic species for some habi¬

tats (beach, dunes, salt marsh), or at least a list of dominant species (prairie, woodlands). The sea-grass beds in

the park are dominated by a two species, Halodule wrightii and Ruppia maritima. Although a decline and even¬

tual disappearance of sea-grass by the early 1980s was reported for West Galveston Bay (Pulich & White 1991),

sea-grass beds are now frequent along the lagoon and bay-shores in Galveston Island State Park (Fig. 4F). To

what degree this is the result of past efforts to reintroduce plants, natural recruitment, or both is not known.

The park’s prairies have the highest species richness (153 spp.) and include five of the seven endemic species

we collected (Fig. 4D, Table 2, Appendix). The park’s woodlands occur as small stands of trees along natural

berms and ridges (e.g., Fig. 4G).

Endemics and Species of Conservation Interest

Seven endemic species were documented during held work. Digitaria arenicola, a rhizomatous perennial re¬

stricted to prairies in the park, is endemic to deep coastal sands of Texas and is mapped by Wipff and Hatch

(1994) as seeing its northern-most records from Galveston County. Digitaria texana is another sandy-prairie

species known only from the Texas’ coast and Rio Grande plains (Correll & Johnston 1970). Gomphrena neal-

leyi seems to be of restricted distribution, known from sandy or clayey soils in coastal Texas and the Rio Grande

plains and adjacent southwest Texas and Mexico (Correll & Johnston 1970). Herhertia lahue, Fradescantia

Rosen etal., Flora of Galveston Island State Park, Texas

345

Fig. 4. Representative photos of general habitat types at Galveston Island State Park, Galveston County, Texas. A. Beach. B. Dunes (seaward aspect). C.

Dunes (leeward aspect). D. Strand prairie. E. Salt marsh. F. Seagrass bed. G. Woodland. H. Wetland.

346

Journal of the Botanical Research Institute of Texas 8(1)

Table 2. Characteristic native species (listed in alphabetical order) of select habitats of Galveston Island State Park, Galveston County, Texas.

Beach

Amaronthus greggii

Rayjacksonia phyllocephala var. phyllocephala

Cakile constricta

Sesuvium portulacastrum

Cokile geniculoto

Sporobolus virginicus

Eustoma exaltatum

Fimbristylis costoneo

Tidestromia lanuginosa subsp. lanuginosa

Dunes (both seaward & leeward)

Ipomoeo imperoti

Rubus trivialis

Ipomoea pes-coproe subsp. brasiliensis

Spartina patens

Ponicum omorum

Heterotheca subaxillaris

Uniola paniculata

Baptisia bracteata var. leucophaea

Aphanostephus skirrhobosis

Oenothera drummondii

Helianthus praecox

Croton punctatus

Vigna luteola

Salt marsh

Agalinis maritima

Salicornia bigelovii

Botis moritimo

Sarcocornia utahensis

Cuscuta indecora var. indecora

Monanthochloe littoralis

Iva angustifolia

Spartina alterniflora

Iva frutescens

Spartina spartinae

Limonium carolinianum

Spergularia salina

Lycium carolinianum

Strophostyles helvola

Strand Prairie

Agalinis fasciculata

Monarda punctata

Ambrosia psilostachya

Muhlenbergia capillaries

Baptisia bracteata var. leucophaea

Oenothera drummondii

Croton capitatus var. lindheimeri

Panicum virgatum

Croton glandulosus var. lindheimeri

Paspalum monostachyum

Dichanthelium scoparium

Paspalum plicatulum var. plicatulum

Digitaria cognata subsp. cognata

Paspalum setaceum var. stramineum

Eupatorium serotinum

Physalis cinerascens var. spathulifolia

Euthamia leptocephala

Schizachyrium scoparium var. scoparium

Fimbristylis caroliniana

Setaria pumila

Fimbristylis castanea

Spartina patens

Heterotheca subaxillaris

Spartina spartinae

Mimosa strigillosa

Strophostyles leiosperma

Woodlands

Baccharis halimifolia

Paspalum monostachyum

Campsis radicans

Quercus nigra

Celtis laevigata

Quercus virginiana

Erythrina herbacea

Sideroxylon lanuginosum subsp. oblongifolium

Ilex vomitoria

Smilax bona-nox

Indigofera suffruticosa

Vitis mustangensis

Melothria pendula

Zanthoxylum clava-herculis

subacaulis, and Zephyranthes traubii are also prairie species endemic to the either primarily the Western Gulf

Coastal Plain (in the case of H. lahue) or Texas (Correll & Johnston 1970). In the 1970s, RaymondJ. Fleetwood

discovered a population of “corkwood” he identified as Leitneriafloridana in forested wetlands in nearby Bra¬

zoria County. Recognizing the plants were regionally unique and concerned with the conservation of the

population, in 1972, he introduced plants to several suitable sites in the park’s prairies (Fleetwood 1973; David

Riskind, personal communication). Since then, the donor-site (Bird Pond) has been permanently protected

through acquisition by the U.S. Fish & Wildlife Service, and the plants there have been recently described as a

new species, Leitneria pilosa subsp. pilosa, known only from forested wetlands and prairies of the upper Texas

Gulf Coast (Schrader & Graves 2011).

A native plant known to have been extirpated from the park has been successfully re-introduced. Sea-oats

Rosen etal., Flora of Galveston Island State Park, Texas

347

(Uniola paniculata) from native populations on nearby Follet’s Island were established in dunes in the park in

2010. Future efforts to restore the park’s strand prairies will continue and include the use of fire as well as local

cultivation and reintroduction of species absent, but expected in this community type. Interestingly, Schizach-

yrium littorale does not occur in the park as mapped by Diamond et al. (1987), but rather is replaced by S. sco-

parium var. scoparium. The need for protection and floristic inventory of a remnant strand prairie on nearby

Follet’s Island has also come to light during this study, and steps are being taken to bring this about.

Plant introductions during restoration should first be carefully evaluated. Probably in an effort to improve

habitat for migratory songbirds, in about 1990, the non-native black locust ( Robinia pseudoacacia ) was pur¬

posefully planted in the park’s woodlands or in stands to create new-woodlands. This species has since become

invasive and required control.

APPENDIX

ANNOTATED CHECKLIST OF VASCULAR PLANTS AT GALVESTON ISLAND STATE PARK

Families are arranged alphabetically, beginning with monocots and followed by eudicots. Genera, species, and

infraspecific names are arranged alphabetically under families. Some species names are preceded by special

symbols to indicate nativity and conservation interest as follows: (1) non-native species are indicated by an

asterisk (*) based on review of Correll and Johnston (1970); (2) endemic species are indicated by a superscript

dagger (t) based on review of Flora of North America (1993+) and Correll and Johnston (1970). Following each

name is an abbreviation from Palmer et al. (1995) representing one of the following subjective estimates of the

relative abundance of that species in the particular habitat(s) where it was collected: r = rare (very difficult to

find and limited to one or very few locations or uncommon habitats); i = infrequent (difficult to find with few

individuals or colonies but found in several locations); o = occasional (widely scattered but not difficult to find);

f = frequent (easily seen or found in one or more common habitats but not dominant in any common habitat);

and a = abundant (dominant or co-dominant in one or more common habitats; terms in quotes are those of

Fleetwood). Following the relative abundance, the habitat(s) where that species is typically found is indicated

by the following general categories (terms in quotes are those of Fleetwood): beach = from the wave swash zone

to the base of the dunes; dunes (both seaward and leeward) = vegetation stabilized wind deposited mounds

and ridges that parallel the beach; prairie = grasslands throughout the park; woodlands = thickets of woody

species; wetlands = all non-tidal freshwater wetlands including seasonally hooded ponds and swales; salt

marsh = wetlands with rare to daily tidal hooding dominated by halophytic vascular plants; sea-grass beds =

stands of rooted aquatic vascular plants that occur in shallow waters of the bay and lagoons; disturbed = dirt

roads, roadside ditches, fence-lines, campgrounds, and vicinity of man-made structures. Collections are the

first authors with the exceptions of two without number (s.n.) by the third author and those of Fleetwood (RJF)

or William R. Carr (WRC).

MONOCOTS

Alismataceae

Sagittaria longiloba Engelm. ex J.G. Sm., r, wetland, 5592

Amaryllidaceae

Allium canadense L. var. mobilense (Regel) Ownbey, i, prairie, 5997

Nothoscordum bivalve (L.) Britton, o, disturbed, 5959

+ Zephyranthes traubii (W. Hayw.) Moldenke, o, prairie, 6109

Arecaceae

* Saba! palmetto (Walter) Lodd. ex Schult. & Schult. f., o, prairie, 6113

*Washingtonia robusta H. Wendt, o, disturbed, woodland 6137

Asparagaceae

Yucca flaccida Haw., r, prairie, 6140

Commelinaceae

Commelina erecta L. var. angustifolia (Michx.) Fernald, r, prairie, 5488

Tradescantia occidentalis (Britton) Smyth, o, disturbed, 5976

Tradescantia ohiensis Raf., o, prairie, 5553

^Tradescantia subacaulis Bush, i, prairie, 5938

Cymodoceaceae

Halodule wrightii Asch., a, seagrass beds, 6119

Cyperaceae

Carexlongii Mack, o, wetland, 5982

Cyperus acuminatus Torr. & Hook., r, prairie, 6148

Cyperus croceus Vahl, r, prairie, 5760

* Cyperus esculentus L., r, prairie, 5489

Cyperus odoratus L. var. odoratus, r, wetland, 5910

Cyperuspolystachyos Rottb., o, prairie, wetland, 5576

Cyperuspseudovegetus Steud., o, wetland, 5759

Cyperus retrorsus Chapm., o, prairie, 5254

Cyperus strigosus L., "disturbed", RJF 10,912

Cyperus surinamensis Rottb., o, disturbed, 6094

Cyperus virens Michx. var. virens, f, wetland, 5756

Eleocharis albida Torr., o, prairie, wetland, 5993

348

Eleocharis ambigens Fernald, r, wetland, 6133

Eleochoris montevidensis Kunth, o, prairie, wetland, 5599

Eleocharispalustris (L.) Roem. & Schult., o, wetland, 5594

Fimbristylis caroliniana (Lam.) Fernald, o, prairie, 5494

Fimbristylis castanea (Michx.) Vahl, f, beach, prairie, 5300

*isoiepis carinata Hook. & Arn. exTorr., r, prairie, 5558

*isoiepis cernua (Vahl) Roem. & Schult., o, prairie, 5602

*Kyllinga brevifolia Rottb., o, disturbed, 6066

Rhynchospora colorata (L.) H. Pfeiff., o, prairie, 6075

Schoenoplectus americanus (Pers.) Volkart ex Schinz & R. Keller, o,

wetland, 5590

Schoenoplectus californicus (C.A. Mey.) Sojak, a, wetland, 6013

Schoenoplectuspungens (Vahl) Palla, f, prairie, wetland, 5995

Iridaceae

+ Herbertia lahue (Molina) Goldblatt, I, prairie, 5989

Sisyrinchium biforme E.P. Bicknell, r, prairie, 5581

^Sisyrinchium exile E.P. Bicknell, r, prairie, 5578

Sisyrinchium langloisii Greene, o, prairie, 5941

Juncaceae

Juncus acuminatus Michx., o, prairie, 5981

Juncus brachycarpus Engelm., o, prairie, 6016

Juncus dichotomus Elliott, o, prairie, 5992

Juncus marginatus Rostk., o, prairie, 5567

Juncus megacephalus M.A. Curtis, o, wetland, 5755

Juncus roemerianus Scheele, f, wetland, 5572

Juncus validus Coville var. fascinatus M.C. Johnst., o, wetland, 5754

Juncus validus Coville var. validus, o, wetland, 5762

Lemnaceae

Lemna minuta Kunth, a, wetland, 6014

Poaceae

Agrostis hyemalis (Walter) Britton, Sterns & Poggenb., o, prairie, 5988

Andropogon glomeratus (Walter) Britton, Sterns & Poggenb., r,

wetland, 5916

Andropogon virginicus L., o, prairie, 5498

Aristida purpurascens Poir., i, prairie, s.n.

*Arundo donax L., f, wetland, 6139

Axonopus fissifolius (Raddi) Kuhlm., "abundant, grassland", RJF

10,507

*Bothriochloa ischaemum (L.) Keng, o, disturbed, 6136

*Briza minor L., f, disturbed, 5974

Cenchrus spinifex Cav., i, disturbed, 6070

*Cynodon dactylon (L.) Pers., o, disturbed, 5969

*Dactyioctenium aegyptium (L.) Willd., "occasional, disturbed",

RJF 10,542

Dichanthelium acuminatum (Sw.) Gould & C.A. Clark, o, prairie, 6015

Dichanthelium laxiflorum (Lam.) Gould, o, prairie, 5568

Dichanthelium scoparium (Lam.) Gould, o, prairie, 5274

Dichanthelium sphaerocarpon (Elliott) Gould "occasional, dunes,

wetlands", RJF 10,813

f Digitaria arenicola (Swallen) Beetle, i, prairie, 6086

*Digitaria ciliaris (Retz.) Koeler var. ciliaris, o, prairie, 5285

Digitaria cognata (Schult.) Pilg. subsp. cognata, o, prairie, 5487

f Digitaria texana Hitchc., r, prairie, 6151

Echinochloa waited (Pursh) A. Heller, o, wetland, 5593

Elymus virginicus L., i, prairie, 6040

Eragrostis elliottii S. Watson, o, prairie, 5497

Eragrostis secundiflora J. Presl, r, prairie, 5919

Eragrostis silveana Swallen, r, prairie, 6152

Eustachys petraea (Sw.) Desv., o, prairie, 5996

Hordeum pusillum Nutt., f, disturbed, 5967

Leptochloa fusca (L.) Kunth subsp. uninervia (J. Presl) N.W. Snow,

r, wetland, 5266

Journal of the Botanical Research Institute of Texas 8(1)

Leptochloa nealleyiM asey, f, wetland, 5289

Limnodea arkansana (Nutt.) L.H. Dewey, i, woodland, 6003

*Lolium arundinaceum (Schreb.) Darbysh., "infrequent, grassland",

RJF 10,513

*Lolium perenne L., o, disturbed, 6029

* Megathyrsus maximus (Jacq.) B.K. Simon & S.W.L. Jacobs, r, prairie,

5485

Monanthochloe littoralis Engelm., a, salt marsh, 5957

Muhlenbergia capillaris (Lam.) Trin., o, prairie, 5491

Panicum amarum Elliott, f, dunes, 6145

Panicum dichotomiflorum Michx., f, wetland, 6150

*Panicum repens L., o, beach, 6091

Panicum virgatum L., o, prairie, 5298

*Parapholis incurva (L.) C.E. Hubb., o, saltmarsh, 5964

*Paspalidium geminatum (Forssk.) Stapf var. geminatum, f, wet¬

land,6131

Paspalum monostachyum Vasey, o, prairie, woodlands, 5495

Paspalum plicatulum Michx. var. plicatulum, "common", prairie,

RJF 10,541

Paspalum setaceum Michx. var. stramineum (Nash) DJ. Banks, o,

prairie, 5492

*Paspalum urvillei Steud., o, prairie, 6078

Paspalum vaginatum Sw., a, wetland, 5265

Phalaris angusta Nees ex Trin., i, prairie, 5944

Phalaris caroliniana Walter, o, prairie, 5561

Phragmites australis (Cav.) Trin. ex Steud., f, wetland, 6146

*Poa annua L., o, prairie, disturbed, 5937

*Polypogon monspeliensis (L.) Desf., f, saltmarsh, 5963

Sacciolepis striata (L.) Nash, o, wetland, 5915

Schizachyrium scoparium (Michx.) Nash var. scoparium, f, prairie,

5490

Setaria magna Griseb., f, wetland, 5301

Setariaparviflora (Poir.) Kerguelen, o, prairie, 5598

Setariapumila (Poir.) Roem. & Schult., f, prairie, 5261

*Sorghum halepense (L.) Pers., o, disturbed, 6116

Spartina alterniflora Loisel., a, saltmarsh, 6123

Spartinapatens (Aiton) Muhl., a, dunes, prairie, 5257

Spartinaspartinae (Trin.) Merr. ex Hitchc.,f, prairie, saltmarsh, 5589

Sphenopholis obtusata (Michx.) Scribn., o, prairie, 5559

Sporobolus virginicus (L.) Kunth, o, wetland, beach, 5911

Tridens strictus (Nutt.) Nash, i, prairie, s.n.

Tripsacum dactyloides (L.) L., r, prairie, 6041

Uniolapaniculata L., r, dunes, 6144

Vulpia octoflora (Walter) Rydb. var. octoflora, o, prairie, 5575

Ruppiaceae

Ruppia maritima L., o, seagrass beds, wetland, 5971

Smilacaceae

Smilax bona-nox L., o, woodland, 6127

Typhaceae

Typha latifolia L., r, wetland, 5299

EUDICOTS

Adoxaceae

Sambucus nigra L. subsp. canadensis (L.) Bolli, i, prairie, 6042

Aizoaceae

Sesuvium maritimum (Walter) Britton, Sterns & Poggenb., f, wet¬

land, 5268

Sesuvium portulacastrum (L.) L., f, beach, 5934

Amaranthaceae

*Aiternantheraphiloxeroides (Mart.) Griseb., f, wetland, 5927

Amaranthus greggii S. Watson, f, beach, 6087

Rosen etal., Flora of Galveston Island State Park, Texas

349

*?Amaranthus spinosus L., "disturbed", RJF 10,530

*Chenopodium album L., r, prairie, 5311

*Dysphania ambrosioides (L.) Mosyakin & Clemants, f, prairie, 5409

+ Gomphrena nealleyi J.M. Coult. & Fisher, o, prairie, 6110

Gomphrena serrata L, i, prairie, 6024

Salicornia bigelovii Torr., a, salt marsh, 6114

Sarcocornia utahensis (Tidestr.) AJ. Scott, a, salt marsh, 6064

Suaeda linearis (Elliott) Moq., r, wetland, 5303

Tidestromia lanuginosa (Nutt.) Standi, subsp. lanuginosa, o, beach,

6103

Apiaceae

Ammoselinum butleri (Engelm. ex S. Watson) J.M. Coult. & Rose,

o, prairie, 5951

Chaerophyllum tainturieri Hook. var. tainturieri, o, disturbed,

prairie, 5940

Cyclospermum leptophyllum (Pers.) Sprague, o, prairie, 5603

Limnosciadiumpinnatum (DC.) Mathias & Constance, o, prairie, 5586

Limnosciadium pumilum (Engelm. &A. Gray) Mathias & Constance,

f, prairie, 5962

Apocynaceae

Cynanchum angustifolium Pers., o, wetland, 5295

Aquifoliaceae

llexvomitoria Aiton, i, woodland, 6028

Araliaceae

Hydrocotyle bonariensis Lam., o, wetland, 5752

Asteraceae

Ambrosiapsilostachya DC., o, prairie, 6134

Ambrosia trifida L. var. texana Scheele, o, woodland, 6134

Aphanostephus skirrhobasis (DC.) Trel. ex Coville & Branner, o,

dunes, 5929

Baccharis halimifolia L., f, woodland, 6128

Borrichia frutescens (L.) DC., o, wetland, 5307

Cirsium horridulum Michx. var. horridulum, o, prairie, 5998

Conoclinium coelestinum (L.) DC., r, wetland, 5292

Conyza canadensis (L.) Cronquist, o, prairie, 5408

Coreopsis basalis (A. Dietr.) S.F. Blake, o, prairie, 6022

Coreopsis tinctoria Nutt., r, prairie, 5749

Ecliptaprostrata (L.) L., r, wetland, 5286

Erigeron procumbens (Houst. ex Mill.) G.L. Nesom, i, prairie, 5746

Eupatorium capillifolium (Lam.) Small ex Porter & Britton, o, prairie,

6130

Eupatorium serotinum Michx., f, prairie, 5407

Euthamia gymnospermoides Greene, "common, grassland", WRC

10,187

Euthamia leptocephala (Torr. & A. Gray) Greene ex Porter & Britton,

a, prairie, 5499

Gaillardiapulchella Foug., o, prairie, 5277

Helenium amarum (Raf.) H. Rock, r, prairie, 5417

Helianthus petiolaris Nutt, subsp. petiolaris, o, prairie, 5255

Helianthus praecox Engelm. & A. Gray, f, dunes, 6076

Heterotheca subaxillaris (Lam.) Britton & Rusby, o, dunes, prairie,

5753

*Hypochaeris microcephala (Sch. Bip.) Cabrera var. albiflora (Kuntze)

Cabrera, i, disturbed, 6020

Iva angustifolia Nutt, ex DC., o, saltmarsh, 6124

Iva frutescens L., f, saltmarsh, 5269

Krigia wrightii (A. Gray) K.L. Chambers ex KJ. Kim, r, disturbed, 5958

Mikania scandens (L.) Willd., o, wetland, 5291

Pluchea odorata (L.) Cassini var. odorata, o, wetland, 5259

Pseudognaphalium obtusifolium (L.) Hilliard & B.L. Burtt, o, prairie,

5555

Pyrrhopappus carolinianus (Walter) DC., r, prairie, 5748

Rayjacksoniaphyllocephala (DC.) R.L. Hartm. & M.A. Lane var. phyl-

locephala, a, beach, 5932

Rudbeckia hirta L., o, prairie, 5552

Solidago sempervirens L., o, prairie, 5420

Soliva sessilis Ruiz & Pav., f, disturbed, 6025

*Sonchus asper (L.) Hill, i, prairie, 6023

*Sonchus oleraceus L., i, disturbed, 5961

Symphyotrichum subulatum (Michx.) G.L. Nesom, o, prairie, 5914

Bataceae

Batis maritima L., a, saltmarsh, 5305

Bignoniaceae

Campsis radicans (L.) Bureau, o, woodland, 6072

Boraginaceae

Heliotropium curassavicum L. var. curassavicum, f, wetland, 5288

Nama jamaicensis L., "disturbed", RJF 10,905

Brassicaceae

Cakile constricta Rodman, f, beach, 6031

Cakile geniculata (B.L. Rob.) Millsp., f, beach, 6032

Lepidium virginicum L. var. virginicum, o, prairie, 5587

Cactaceae

Opuntia humifusa (Raf.) Raf. var. humifusa, o, prairie, woodland, 6096

Campanulaceae

Triodanis biflora (Ruiz & Pav.) Greene, o, prairie, 5556

Cannabaceae

Celtis laevigata Willd., o, woodland, 6007

Caprifoliaceae

*Lonicerajaponica Thunb, o, woodland, 5985

Valerianella woodsiana (Torr. & A. Gray) Walp., RJF 10,908

Caryophyllaceae

Cerastium glomeratum Thuill., o, prairie, 5936

*Polycarpon tetraphyllum (L.) L., "disturbed", RJF 10,922

*Siiene gallica L., "disturbed", RJF 10,888

Spergularia salina J. Presl & C. Presl, o, saltmarsh, 5965

Stellaria media (L.) Vill., o, prairie, 5953

Celastraceae

Lepuropetalon spathulatum Muhl. ex Elliott, o, prairie, 5573

Cistaceae

Helianthemum rosmarinifolium Pursh, o, prairie, 6125

Lechea mucronata Raf., r, prairie, 5912

Convolvulaceae

Calystegia sepium (L.) R. Br. subsp. limnophila (Greene) Brummitt,

o, disturbed, 6034

Cuscuta indecora Choisy var. indecora, o, saltmarsh (on Iva frute¬

scens), 6118

Dichondra carolinensis Michx., o, prairie, 5973

Ipomoea cordatotriloba Dennst. var. cordatotriloba, i, prairie, 6112

Ipomoea imperati (Vahl) Griseb., f, dunes, 6033

Ipomoea pes-caprae (L.) R. Br. subsp. brasiliensis (L.) Ooststr., f,

dunes, 6090

Ipomoea sagittata Poir., f, wetland, 5302

Cucurbitaceae

Ibervillea lindheimeri (A. Gray) Greene, "occasional, disturbed",

RJF 10,539

Melothria pendula L, o, woodland, 6147

Euphorbiaceae

*Chamaesyce maculata (L.) Small, o, prairie, 5284

*Chamaesyce nutans (Lag.) Small, o, disturbed, 6143

350

Croton capitatus Michx. var. lindheimeri (Engelm. & A. Gray) Mull.

Arg., o, prairie, 5922

Croton glandulosus L. var. lindheimeri Mull. Arg., f, prairie, 5256

Croton punctatus Jacq., f, dunes, 6077

*Triadica sebifera (L.) Small, o, woodland, 6126

Fabaceae

Acocio fornesiono (L.) Willd., o, prairie, 5983

*Aeschynomene indica L., r, wetland, 5414

Astragalus leptocarpus Torr. & A. Gray, i, prairie, 5949

Baptisia bracteata Muhl. ex Elliott var. leucophaea (Nutt.) Kartesz &

Gandhi, f, dunes, prairie, 5580

Centrosema virginianum (L.) Benth., r, prairie, 5418

Chamaecrista fasciculata (Michx.) Greene, o, prairie, 5406

Erythrina herbacea L., f, woodland, 5984

*Glottidium vesicarium (Jacq.) R.M. Harper, r, prairie, 5496

Indigofera miniata Ortega, i, prairie, 6035

Indigoferasuffruticosa Mill., o, woodland, 6129

*Leucaena leucocephala (Lam.) de Wit, o, woodland, 6121

*Medicagopolymorpha L., o, disturbed, 5977

*Meli lotus indicus (L.) All., r, prairie, 5994

Mimosa strigillosa Torr. & A. Gray, o, prairie, 5278

Pediomelum rhombifolium (Torr. & A. Gray) Rydb., "infrequent,

disturbed", RJF 10,535

Rhynchosia americana (Mill.) Metz, o, prairie, 6071

*Robiniapseudoacacia L., i, woodland, 6074

Sesbania drummondii (Rydb.) Cory, o, wetland, 5294

Strophostyles helvola (L.) Elliott, f, prairie, slatmarsh, 6117

Strophostyles leiosperma (Torr. & A. Gray) Piper, f, prairie, 5262

Tephrosia onobrychoides Nutt., r, prairie, 5744

Trifolium carolinianum Michx., i, disturbed, 5991

Vida ludoviciana Nutt, ex Torr. & A. Gray, o, disturbed, 5972

Vida minutiflora D. Dietr., o, prairie, 5948

Vigna luteola (Jacq.) Benth., f, wetland, dunes, 5283

Fagaceae

Quercus nigra L, i, woodland, 6011

Quercus virginiana Mill., f, woodland, 6008

Gentianaceae

*Centaurium pulchellum (Sw.) Druce, r, disturbed, 5966

Eustoma exaltatum (L.) Salisb. ex G. Don, r, beach, prairie, 5419

Sabatia campestris Nutt., r, prairie, 5579

Geraniaceae

Geranium carolinianum L., f, prairie, 5952

Geranium texanum (Trel.) A. Heller, o, prairie, 5577

Hypericaceae

Hypericum drummondii (Grev. & Hook.) Torr. & A. Gray, o, prairie, 5921

Lamiaceae

Monardapunctata L., o, prairie, 5583

Scutellariaparvula Michx., o, woodland, 6073

Teucrium canadense L., f, prairie, 5272

Linaceae

Linum medium (Planch.) Britton var. texanum (Planch.) Fernald, o,

prairie, 5750

Malvaceae

Callirhoe involucrata (Torr. & A. Gray) A. Gray var. lineariloba (Torr. &

A. Gray) A. Gray,"disturbed", RJF 10,926

Hibiscus laevis All., i, wetland, 6095

Kosteletskya virginica C. Presl, f, wetland, 5263

Sida ciliaris L., o, prairie, 6079

Sida rhombifolia L., o, disturbed, 6005

Journal of the Botanical Research Institute of Texas 8(1)

Melastomataceae

Rhexia mariana L. var. mariana, o, prairie, 5745

Myricaceae

Morelia cerifera (L.) Small, i, prairie, woodlands, 6012

Onagraceae

Gaura filiformis Small, f, prairie, 5271

Gauraparviflora Douglas ex Lehm., i, prairie, 6027

Ludwigia glandulosa Walter, o, wetland, 6081

*Ludwigia grandiflora (Michx.) Greuter & Burdet subsp. hexapetala

(Hook. & Arn.) G.L. Nesom & Kartesz, o, wetland, 6082

Ludwigia linearis Walter, o, wetland, 5758

Ludwigia repens J.R. Forst., o, wetland, 6065

Oenothera drummondii Hook., f, dunes, prairie, 5751

Oenothera laciniata Hill, o, prairie, 5970

Oenotheraspeciosa Nutt., f, prairie, 5956

Orobanchaceae

Agalinis fasciculata (Elliott) Raf., f, prairie, 5276

Agalinis heterophylla (Nutt.) Small, i, prairie, 6141

Agalinis maritima (Raf.) Raf., o, salt marsh, 6069

Buchnera americana L., r, prairie, 6111

Oxalidaceae

*Oxalis corniculata L., o, prairie, 5943

Oxalis violacea L., r, disturbed, 5918

Phytolaccaceae

Phytolacca americana L., o, prairie, 5282

Plantaginaceae

Bacopa monnieri (L.) Wettst., f, wetland, 5267

Callitrichepeploides Nutt., r, wetland, 5597

Callitriche terrestris Raf., o, wetland, 5596

Nuttallanthus texanus (Scheele) D.A. Sutton, o, prairie, 5584

Plantago hookeriana Fisch. & C.A. Mey.,"disturbed", RJF 10,904

Plantago virginica L., f, disturbed, 5968

Plumbaginaceae

Limonium carolinianum (Walter) Britton, o, saltmarsh, 5913

Polygalaceae

Polygala verticillata L., o, prairie, 5588

Polygonaceae

Persicaria hydropiperoides (Michx.) Small, "common, wetland",

RJF 10,527

Persicaria punctata (Elliott) Small, f, wetland, 5591

Rumexcrispus L., o, wetland, 5978

Rumexhastatulus Baldwin, o, prairie, 5582

Rumexverticillatus L., i, wetland, 5945

Polypremaceae

Polypremum procumbens L., r, prairie, 5920

Portulacaceae

*Portulaca oleracea L., o, disturbed, 6068

Portulacapilosa L., o, disturbed, 6108

Primulaceae

Anagallis arvensis L., o, prairie, 5605

Anagallis minima (L.) E.H.L. Krause, r, prairie, 5557

Samolus ebracteatus Kunth, o, prairie, 6000

Ranunculaceae

Ranunculus muricatus L., o, disturbed, 5980

Ranunculuspusillus Poir., f, wetland, 5595

Rosaceae

Rubus trivialis Michx., f, dunes, 5924

Rosen etal., Flora of Galveston Island State Park, Texas

351

Rubiaceae

Diodio teres Walter, o, prairie, 5923

Diodia virginiana L., o, wetland, 5296

Galium aparine L., o, disturbed, prairie, 5955

Galium tinctorium L., f, wetland, 5597

Galium uniflorum Michx., "common, wetlands", RJF 10,510

Oldenlandia uniflora L., r, prairie, 5747

Rutaceae

Zanthoxylum clava-herculis L., f, woodland, 5987

Salicaceae

Salixnigra Marshall, i, wetland, 6083

Sapotaceae

Sideroxylon lanuginosum Michx. subsp. oblongifolium (Nutt.) T.D.

Penn., o, woodland, 6122

Simaroubaceae

+ Leitneriapilosa J.A. Schrad. & W.R. Graves subsp. pilosa, i, wetland,

6138

Solanaceae

Lycium carolinianum Walter, a, salt marsh, 6149

Physalis angulata L. var. angulata, r, wetland, 5297

Physalis cinerascens (Dunal) Hitchc. var. spathulifolia (Torr.) J.R. Sul¬

livan, o, prairie, 6085

Physalispubescens L. var. pubescens, o, wetland, 5306

Solanum ptychanthum Dunal, r, prairie, 5304

Tamaricaceae

*Tamarixramosissima Ledeb., f, wetland, 5933

Urticaceae

Urtica chamaedryoides Pursh, "wetlands", RJF 10,818

Verbenaceae

*Lantana camara L., o, prairie, 5281

Phyla lanceolata (Michx.) Greene, f, wetland, 5975

Phyla nodiflora (L.) Greene, "common", RJF 10,532

Verbena halei Small, o, prairie, 5942

*Verbena brasiliensis Veil., o, disturbed, 6010

Vitaceae

Ampelopsis arborea (L.) Koehne, r, prairie, 5412

Cissus incisa Des Moul., o, wetland, 5413

Vitis mustangensis Buckley, o, woodland, 5986

ACKNOWLEDGMENTS

We are grateful to David Riskind for enthusiasm and support during this project, and for providing accounts

and records detailing the efforts of R.J. Fleetwood, a conservationist to whom we are also grateful, and who

seemingly (from what we’ve learned about him) recognized the importance of floristics and plant conservation

as fundamental to wildlife conservation and management. We are also grateful for the support of the park staff,

especially Sean Rogers, for always managing to make an all-terrain vehicle available which made the going

much easier. Special thanks to Jennifer Estes for preparing Figures 1-3. We are also thankful for the helpful

reviews by Bill Carr and Eric Keith. Lastly, the first two authors thank Tom Wendt for facilitating their visit to

TEX to review Fleetwood’s specimens.

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Texas Natural Heritage Program. 1993. Plant communities of Texas (Series Level). Unpublished document. Texas Parks and

Wildlife Department, Austin,Texas, U.S.A.

Tropicos.org. Missouri Botanical Garden, http://www.tropicos.org. Accessed January 2014.

Wipff, J.K. & S.H. Hatch. 1994. A systematic study of Digitaria - sect. Pennatae (Paniceae) in the New World. Syst. Bot.

19:613-627.

AN ANNOTATED FLORA OF REED PLATEAU AND ADJACENT AREAS,

BREWSTER COUNTY, TEXAS, U.S.A.

Wendy Weckesser

7 02 Miguel Angel Court

Del Rio, Texas 78840, U.S.A.

wendy week 7 3@gmail.com

Martin Terry

Sul Ross State University

Box C-64

Alpine, Texas 79832, U.S.A.

ABSTRACT

Reed Plateau is a geographic feature in south Brewster County, Texas (U.S.A.) that exhibits a floral array characteristic of the northern Chi-

huahuan Desert region. Held as private property by many landowners, Reed Plateau and areas adjacent to it have never been the focus of a

botanical study. A survey of Reed Plateau and adjacent areas was conducted from August 2004 through November 2007, with a total of 1065

specimens collected. The flora consists of 262 taxa, including 1 subspecies and 15 varieties, in 188 genera and 63 families. The best-repre¬

sented families are the Asteraceae (33 species), Poaceae (23 species), Fabaceae (18 species), Cactaceae (17 species), and Euphorbiaceae (13

species). One federally listed threatened species, Echinomastus mariposensis, was identified within the study area. The only known U.S.

populations of Genistidium dumosum occur on Reed Plateau. The occurrence of Stemodia coahuilensis represents a new county record, and a

new Hibiscus hybrid was described. Two Big Bend and three Trans-Pecos endemics were documented. Four non-native species were col¬

lected, three of which are considered noxious or invasive. The vegetation associations found in the Reed Plateau study area strongly reflect

the predominantly limestone substrate of the Terlingua-Solitario structural block. Diverse geographic factors within the relatively small

study area support floral diversity patterns which are compared to studies from nearby Big Bend National Park and Big Bend Ranch State

Park, as well as the Southwestern United States and the Chihuahuan Desert region.

RESUMEN

Reed Plateau es un elemento geografico en el sur del condado Brewster, Texas, que exhibe una coleccion floral caracteristica del norte del

Desierto Chihuahuense. Compuesto de terrenos privados de muchos propietarios, Reed Plateau y sus alrededores nunca han sido el sujeto

de un estudio botanico. Una investigation fue conducida entre agosto 2004 y noviembre 2007, con 1065 especimenes recogidos. Ta flora

consiste en 262 especies, con una subespecie y 18 variedades, de 188 generos y 63 familias. Fas familias mejor representadas fueron las

Asteraceae (33 especies), Poaceae (23 especies), Fabaceae (18 especies), Cactaceae (17 especies) y Euphorbiaceae (13 especies). Una especie

amenazada en la lista federal, Echinomastus mariposensis, fue identificada dentro de la zona del estudio. Fas unicas poblaciones de Genis¬

tidium dumosum conocidas en los E.U. ocurren en Reed Plateau. Ta ocurrencia de Stemodia coahuilensis representa una especie nueva para el

condado de Brewster, y un hibrido nuevo de Hibiscus fue descrito. Dos especies endemicas de Big Bend y tres del Trans-Pecos fueron docu-

mentadas. Cuatro especies no nativas fueron recogidas, tres de ellas consideradas nocivas o invasivas. Fas asociaciones vegetales encontra-

das en Reed Plateau reflejan fuertemente el sustrato predominantemente calizo del bloque estructural Terlingua-Solitario. Factores geogra-

ficos diversos dentro del area relativamente pequena del estudio apoyan patrones de diversidad floral, los cuales se comparan con los

patrones de diversidad encontrados en estudios que se han hecho en sitios cercanos, como el parque nacional de Big Bend y el parque estatal

de Big Bend Ranch, y ademas con estudios del Suroeste de los E.U. y la region del Desierto Chihuahuense.

INTRODUCTION

Reed Plateau is a geographic feature located near Terlingua in southern Brewster County, Texas. Two protected

areas flank Reed Plateau, Big Bend National Park (BBNP) to the east and Big Bend Ranch State Park (BBRSP) to

the west (Fig. 1). While much floristic work has been done in the protected areas (Butterwick & Lamb 1976;

Butterwick & Strong 1976a, 1976b, 1976c; Powell 1985; Worthington 1995; Louie 1996; Bartel 2002; Henklein

2003), Reed Plateau is held as private property in its entirety and has never been the focus of a botanical survey.

The compilation of an annotated flora for Reed Plateau and adjacent areas (RP) was the primary objective of

this study. In conjunction with the flora, we have described the vegetation associations in the study area. The

species list generated by the current effort has been compared with the floras resulting from similar studies in

the nearby protected areas (Hardy 1997; Fenstermacher 2008), as well as a Chihuahuan Desert (CD) flora

(Henrickson & Johnston 2004) and a Southwest United States (SW) flora (McLaughlin 1986). The annotated

RP list can be used as baseline information for future studies, as the level of human activity on and near RP

changes.

J. Bot. Res. Inst. Texas 8(1): 353 - 379.2014

354

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 1. Map of Trans-Pecos Texas with locations of Reed Plateau study area (RP), Solitario Dome (SD), and Dead Horse Mountains (DH) noted (courtesy

of Sul Ross State University Department of Biology, with modifications.)

Site description

Physical features of the Terlingua-Solitario structural block (Erdlac 1990) define the study area: the Terlingua

uplift, the Terlingua monocline and the Long Draw drainage. The portion of the Terlingua uplift that lies to the

south of Highway 170 with a general east-west orientation is identified on United States Geologic Survey

(USGS) topographic and geologic maps (USGS 1971a, 1971b; Barnes 1979) as Reed Plateau. North of Highway

170, the uplift continues uninterrupted in a more northwesterly direction to the point where Long Draw, from

its origins in the vicinity of Black Mesa, cuts through the cliffs. The Terlingua uplift is seen as steep to near¬

perpendicular cliff faces cut by narrow drainages and canyons. The plateau rises 70-100 m (230-340 ft) in el¬

evation above the surrounding desert floor, with a maximum elevation of 1012 m (3380 ft). Long Draw, a broad

creek bed that is normally dry but sporadically flooded, follows the base of the cliff formed by the uplift until it

cuts through the cliff once again to head in a southerly direction. The Terlingua monocline falls away to the

south and west, with wide slopes and cliff faces that are less steep than those of the north side. The southern

slopes and cliffs are cut by wide drainages and canyons or abruptly drop off with steep limestone outcrops.

The study area is approximately 3,237 ha (8,000 ac; 31 km 2 ; 12.5 sq mi). Long Draw forms the northern

and eastern limits. The Terlingua monocline loosely delineates the southern and western boundaries of the

study area; investigational forays into the lower elevations extended only to where there seemed to be no fur¬

ther variation in vegetation.

The east-west section of RP is 7.2 km (4.5 mi) from the eastern intersection of Long Draw and the Terlin-

gua uplift to the Terlingua Sinkhole. Approximately 3.2 km (2 mi) to the northwest beyond the Terlingua

sinkhole, Long Draw again cuts through the plateau, forming the northern boundary of RP. The study area is

approximately 1.6 km (1 mi) wide, measuring from Long Draw to the elevations of the Terlingua monocline

where no further variations in vegetation were observed.

RP experiences hot summers and cool winters. Temperatures range from 43°C (110°F) in midsummer to

below freezing in winter. Average rainfall in the area varies from 16 to 28 cm (6 to 11 in), depending on eleva¬

tion. Most precipitation occurs in late summer, as scattered afternoon downpours. There is a strong relation

between the occurrence of high summer temperatures and the onset of the rainy season in late summer

Weckesser and Terry, Flora of Reed Plateau, Texas

355

(Schmidt 1986). Winter precipitation occurs infrequently, with intermittent snowfall more common in the

higher elevations (pers. obs.; Arbingast et al. 1973; Morafka 1977; Schmidt 1979, 1986). During the course of

the study, precipitation was above average for late summer 2004 and midsummer 2005 (unpublished data,

BBNP and BBRSP).

The geology of RP is largely Cretaceous limestone. Across the Terlingua uplift, Lower to Upper Creta¬

ceous formations are present (Erdlac 1990). The uplift is comprised of Buda limestone, Del Rio clay, and Santa

Elena limestone (Barnes 1979). This stratigraphy is also found in BBNP (Maxwell et al. 1967) and in the Soli-

tario of BBRSP (McCormick et al. 1996). Alluvium, colluvium, and surface caliche of Old Quaternary deposits

are found in the drainages of and at the base of the Terlingua monocline (Barnes 1979). At the base of the Santa

Elena limestone cliffs, the surface strata in Long Draw are identified as Young Quaternary deposits (Barnes

1979). An outcrop of the Pen Formation is located within the study area in Long Draw drainage near the Rain¬

bow Mine, just west of the Terlingua Ghost Town.

Current classification of soils from the U.S. Department of Agriculture’s Natural Resources Conservation

Service (NRCS) shows that the most common soil type on the uplands of Reed Plateau is the Blackgap-Rock

outcrop complex. In Long Draw, the soils are identified as the Riverwash-Pantera complex. The Geefour silty

clays complex is associated with the Del Rio and the Pen Formations, and is also found at the eastern end of the

study area and in scattered pockets north of Hwy 170 (Web Soil Survey 2013).

Vegetation Associations

The vegetation associations of the Chihuahuan Desert region (CD) described by Henrickson and Johnston

(1986) provide the basis for the description of the vegetation associations of RP. Communities are broadly

categorized as desert scrub, woodland, grassland, and chaparral, with multiple subcategories and variations

that reflect the complexity and diversity of CD vegetation patterns found on RP.

Cultural History

Substantial evidence of human habitation in the Terlingua area dates to the late Archaic Period, from approxi¬

mately 1000 BC to AD 900 (Hudson 1976; Ing et al. 1996). In the Prehistoric Period (900 AD-1530 AD), evi¬

dence of permanent habitation and agriculture appears, especially in the area of the confluence of the Rio

Grande and the Rio Conchos (Ing et al. 1996). In the Historic Period (1530 AD to present), the sedentary peo¬

ples of the rivers and deserts came in contact with Spanish explorers; the Mescalero Apache, in their seasonal

migration, moved through the area; and, later, Comanche groups supplanted the Apaches. The Comanche

moved continually through the area until their complete subjugation by United States forces in the late 1880s

(Ragsdale 1976).

The first geologic survey was conducted in the area in the 1880s. From that time until the 1920s ranching

was the dominant way of life in the region. With little water and, reportedly, not much grass, Terlingua was the

last area to be impacted by ranching.

An awareness of cinnabar in the area was longstanding, and in 1884 the first mercury mining claim was

put into operation (Ragsdale 1976). The Colquitt Tigner Mine, still identified by name on the Amarilla Moun¬

tain topographic map (USGS 1971a), was in a drainage immediately to the west of the northern extent of the

study area. The Waldrop Mine is within the study area of this project. The furnaces to process the cinnabar

were bred with cottonwood and mesquite, gathered from the Rio Grande corridor, and probably Long Draw, to

the point of being completely depleted by the 1930s (Ragsdale 1976).

The mercury was carted overland to Marfa. Two-track roads from Villa de la Mina, within the study area,

and Lone Star Mine, to the west of RP, still exist that eventually reach Marfa (Ragsdale 1976). These old roads

provided access to some of the more remote sections of the study area.

When the demand for mercury declined in the 1940s, the area population plummeted, and Terlingua be¬

came a ghost town (Ragsdale 1976). One activity that continued through the 1950s and into the 1970s was can-

delilla wax processing (B. Pittman, pers. comm.). Euphorbia antisyphilitica was harvested and processed for its

wax, used in cosmetics. At least three processing sites were within RP. Another extractive activity from the 1970s

that continues today is the removal of cacti and other succulents for commercial purposes (Harrington 1980).

356

Journal of the Botanical Research Institute of Texas 8(1)

In the late 1970s, the population of the area slowly began to increase. BBNP had been established in the

1940s, and it was a draw for outdoor adventurers. River-tour companies started providing guided trips on the

Rio Grande. Current land division dates to that time, with home construction the predominant use of the land.

The pace of development had been slow until approximately 10 years ago, when the rate of land sales and new

home construction increased, compared to the previous twenty years.

MATERIALS AND METHODS

Herbaria search

Herbaria at Angelo State University (SAT), the Botanical Research Institute of Texas (BRIT), Texas A&M Uni¬

versity (TAES and TAMU), and University of Texas Austin (TEX-LL) were contacted via email with requests

for electronic searches. A manual search was conducted at the A. Michael Powell Herbarium at Sul Ross State

University (SRSC). In addition, the Flora of Texas Consortium website was used for an online search of elec¬

tronic databases (Texasflora.org 2003).

Field collections

Permission for access to the privately held lands of RP was documented with written statements from land-

owners. Variation in geology and terrain guided held efforts, in an attempt to End as much diversity within the

study area as possible. USGS maps of the Amarilla Mountain, Texas (1971a), and Terlingua, Texas (1971b),

quadrangles were references for elevation and topography. Access to portions of the study area was possible by

vehicle along old two-track mining roads, but much of the study area is accessible only on foot. A modified

meander-search method, based on Goff et al. (1982), was employed to collect enough specimens to reflect the

diversity of plant life on RP. Specimens were collected from August 2004 through November 2007, with 127

days in the held. Collection effort, over 500 hours, was greatest in spring and early fall, especially after rains.

Principal sources for identification and nomenclature were Correll and Johnston (1970), Powell (1994,

1998), and Powell and Weedin (2004), as well as online sources (ITIS 2007; Tropicos.org 2014; USDA-PLANTS

2014). Access to Powell’s key (in prep.) to Trans-Pecos non-woody plants was especially helpful. Billie L.

Turner and A. Michael Powell verified specimen identifications. Taxonomy follows Evert and Eichhorn (2013)

at the phyla level. Additional sources for identification and nomenclature were Jones et al. (2003), Turner et al.

(2003), and Yarborough and Powell (2002). Warnock (1970, 1974, 1977) proved useful for visual identifica¬

tions. Current nomenclature and authors were found on two online sources (Tropicos.org 2014; ehoras.org

2014). Author abbreviations follow Brummitt and Powell (1992). A representative collection of each taxon is

housed at SRSC, with selected duplicates sent to BRIT and TEX-LL.

RESULTS AND DISCUSSION

During the three-year study period, 1065 specimens were collected. The flora consists of 262 taxa, including 1

subspecies and 15 varieties, in 188 genera and 63 families. Phylum Anthophyta, with 254 species, forms 97%

of the floral array. The families best represented in the flora are the Asteraceae (33 species), Poaceae (23 spp.),

Fabaceae (18 spp.), Cactaceae (17 spp.), and Euphorbiaceae (13 spp.). These five families comprise 40% of the

flora. Other well-represented families include the Boraginaceae, Solanaceae, Malvaceae, Nyctaginaceae,

Verbenaceae, Brassicaceae, Pteridaceae, and Onagraceae, which together constitute 19% of the flora. Thirty-

eight species are the sole representatives of their families on RP. Five species endemic to the Big Bend area or

the Trans-Pecos region ( Kallstroemia perennans , Lycium puberulum var. berberioides , Lycium texanum, Chamae-

syce perennans , Thelypodium texanum) were documented. Four species with conservation status ( Genistidium

dumosum, Oenothera boquillensis, Lycium texanum, Echinomastus mariposensis) and four non-native species

(Cynodon dactylon, Salsola tragus, Tamarix chinensis, Pennisetum ciliare) were identified. Further distributional

data for RP, along with taxonomic distributions of the Dead Horse Mountains (DH) and Solitario Dome (SD)

floras, are presented in Table 1.

Herbaria search

The amount of material from the study area previously disseminated to herbaria in Texas seems to be sparse,

Weckesser and Terry, Flora of Reed Plateau, Texas

357

Table 1 . Taxonomic composition of three Big Bend floras [RP = Reed Plateau study area; DH = Dead Horse Mountains (Fenstermacher 2008); SD = Solitario Dome

(Hardy 1997)].

Taxa

Families Species

Families

Species

Families Species

Lycopodiophyta

Monilophyta

Gnetophyta

Coniferophyta

Monocotyledones

107

Dicotyledones

225

525

434

Totals

262

662

532

which may be expected since RP has never been the sole focus of a floristic survey. In addition, limited avail¬

ability of herbaria information in electronic format meant that online searches would be far from exhaustive.

The electronic search of the TAMU database resulted in a list of over 6,000 records from Brewster County, with

no feasible means of narrowing the search parameters. No further examination of that database was per¬

formed. No results from search requests were returned from BRIT or TAES since their collections were not

completely accessioned electronically or not available via the Internet. The probability of locating other collec¬

tions from RP and the Terlingua area in herbaria such as BRIT and TAES will increase when their collections

are fully accessible in an electronic format. No information was available from the herbarium at Angelo State

University (SAT).

The search of the University of Texas at Austin (TEX-LL) database resulted in a list of over 500 collections

from the Terlingua area, including RP. A manual search of SRSC resulted in a number of collections from RP,

some of which were duplicated in the TEX-LL list. Eleven collections of species not encountered during the

current study are stored at SRSC and TEX-LL. These are included in the annotated species list for RP.

Taxonomic breakdown

A comparison of the RP flora with the DH (Fenstermacher 2008) and the SD (Hardy 1997) floras (Table 1)

shows expected similarities in species diversity and taxonomic composition, with minor variations that can be

explained by the differences in scope of study and geographic variability. The SD study area encompassed igne¬

ous as well as limestone substrates and canyon environments more developed than those seen in RP. The DH,

with the inclusion of the Rio Grande corridor, presents a much greater range of elevation and habitat variation

than either RP or SD. In addition to the riparian habitat included in DH, both DH and SD contain springs and

seeps. There is no permanent aquatic habitat in RP.

The Gnetophyta are represented in the three study areas by the genus Ephedra (Ephedraceae). While sev¬

eral species were found on DH and SD, only E. aspera was found on RP. The relative homogeneity of the habitats

on RP may account for Ephedra being represented by a single species. Of the Coniferophyta, species from two

families were documented on the DH and the SD, but no conifers were documented from RP in this study.

While all species of the Monilophyta of RP are also found on both DH and SD, there are additional species

from this group in DH and SD that do not occur in RP. The taxonomic breakdown of the RP flora shows a lower

percentage of ferns and fern allies, as well as a lower proportion of the monocots, with a concomitant slightly

higher proportional representation of the eudicots, when compared to the other Big Bend floras (Table 1). Ad¬

ditional representation among the ferns and monocots on SD and DH, with their springs and riparian habitats,

is especially notable at the family level. The monocot families Amaryllidaceae, Orchidaceae, Commelinaceae,

Juncaceae, and Typhaceae are represented on both SD and DH. The absence of these families from RP may ex¬

plain the seemingly anomalous dominance of the eudicots on RP. One possible explanation for this is the lack

of mesic habitats on RP. There are species from these families present in the SD and DH that require moister

habitats than are found on RP. The ferns and fern allies show this same trend. Another contributing factor to

the lower percentage of monocots could be the under-representation of the family Poaceae in the RP flora.

358

Journal of the Botanical Research Institute of Texas 8(1)

Table 2. Taxonomic breakdown of families and species of local and regional [SW = Southwest U.S. (McLaughlin 1986); CD = Chihuahuan Desert region (Henrickson

pers. comm.); DH = Dead Horse Mountains (Fenstermacher 2008); SD = Solitario Dome (Hardy 1997); RP = Reed Plateau study area].

Taxonomic Group

Families

Ferns and fern allies

3.2

3.3

2.3

7.3

7.0

Gymnosperms

1.6

3.3

2.3

2.7

2.4

Monocots

6.3

11.0

11.6

14.0

7.9

Eudicots

88.9

82.4

83.7

114

76.0

105

82.7

Species

Ferns and fern allies

2.7

3.5

3.8

2.9

120

2.2

Gymnosperms

0.4

1.1

0.8

1.1

0.7

Monocots

11.1

107

16.2

13.9

515

1S9

723

13.2

Eudicots

225

85.9

525

79.3

734

81.6

2588

80.0

4579

83.9

In a regional context (Table 2), trends emerge in the broad scope of their relationships to each other and to

floras of the SW (McLaughlin 1986) and the CD (Henrickson & Johnston 2004). The taxonomic distribution

of the DH, easternmost of the three Big Bend study areas, shows influences from the Great Plains and the Chi¬

huahuan Desert. The SD flora aligns more closely with the SW flora. RP, closer geographically to the SD than

to the DH, but in between the two, presents influences from both regions. The gymnosperms are poorly repre¬

sented on RP when compared to regional as well as local floras. The representation of gymnosperms for DH is

the same as that for the CD, and the SD shows similar proportions to those of the SW.

In contrast to the lack of gymnosperms on RP, ferns and fern allies are represented in the RP flora at a

level commensurate with their representation in the CD. Both DH and SD show greater percentages of ferns

and fern allies in their floras, more than either regional flora as well.

The percentage of monocots in the CD flora is higher than that of the SW flora. The greater number of

graminoid species, as a reflection of the influence of the grasslands to the east of the CD, has been proposed as

the point of departure (Fenstermacher 2008). As with the local comparisons of monocots and eudicots, RP

does not support the level of diversity of monocot families seen in the SW or CD, given its relatively small area,

the limited range of habitat variability, and the lack of permanent water.

Life forms

The most common life forms found on RP are, in order of decreasing abundance, perennial herbs, shrubs, sub¬

shrubs, and annual herbs. Vines, trees, and semi-succulents make up less than 4% each of the flora. The life

form categories used in the Big Bend studies and the SW are not identical, but regrouping some of the data al¬

lows for an indirect comparison (Weckesser 2008). No summary of life forms from the CD is available. Peren¬

nial herbs are the dominant life forms for the three Big Bend study areas, as well as in the SW flora. Annual

herbs constitute the next most dominant form on DH, SD, and the SW. In contrast, the shrubs are more abun¬

dant than annual herbs in RP. With its dry, rocky ridges and gullies, RP supports more shrub species through¬

out the year than annual herbs in season.

Six species in RP are categorized as trees. Two species (Acacia roemeriana and Prosopis glandulosa) that

can grow to tree form in appropriate conditions are most commonly found as shrubs on RP. Quercus vaseyana

is usually found as a major shrub component of the chaparral association, but on RP it occurs, unexpectedly, as

a glade of trees in a steep-sided, narrow canyon. All of the tree species encountered on RP are also found on DH

and SD.

Floral diversity

The most abundant families, i.e., those represented by the greatest proportion of species in each flora, are the

Asteraceae, Poaceae, Fabaceae, Cactaceae, and Euphorbiaceae. The Asteraceae are most numerous in each

study area. But on DH, the Asteraceae and Poaceae are of nearly equal abundance, reflecting the influence of the

proximity to the Great Plains (Fenstermacher 2008). The Fabaceae comprise the third most abundant family

Weckesser and Terry, Flora of Reed Plateau, Texas

359

in both DH and RP floras. The Tamaulipan thornscrub vegetation zone, to the east and south of the Big Bend

region, apparently influences the make-up of the flora of the DH in this regard (Fenstermacher 2008), and this

influence may extend to RP. The Poaceae constitute the second most abundant family in each of the three

floras, but on RP the family makes up a smaller portion of the flora than seen in DH or SD. This may be a result

of the greater range of elevation in the DH and a wider range of habitat on SD, or it may be due to under-repre¬

sentation of grasses in the RP collections.

In the context of the Big Bend region, the family representations in the floras of RP and SD seem to have

more in common with each other than with DH, and this fits with the proximity of RP to SD and their common

geologic past. The variations in habitat within these two study areas are not as extreme as those in the DH, with

range in elevation, perhaps, being the most significant physical difference.

In contrast, fifty-one species from the RP flora were not documented on SD, which is to say that nearly

20% of the RP flora is distinct from the SD flora. Thirty species, over 10% of the RP flora, were not found in the

DH. Nine species from RP were not found on either SD or DH. Four of these species are endemic to the immedi¬

ate vicinity of RP. Three of these species are associated with gypsiferous soils. Of these species unique to RP, 11

not on SD are known or suspected gypsophiles. Eight known or suspected gypsophiles are documented on RP

that are not found on DH.

When looking at the taxonomic make-up of the three Big Bend floras in the context of the CD and SW

floras (Table 2) it is clear that the DH, at the species level, shows a stronger affinity for the CD flora than for the

SW flora. The SD shows a slightly greater affinity for the floral assemblage of the SW than for that of the CD. RP

also shows a slightly greater affinity for the SW flora, even as its dominant shrub component reflects the influ¬

ence of the Tamaulipan thornscrub vegetation zone.

Noteworthy Collections, Rare Taxa, Non-natives

Many species of concern have been identified in the protected areas of BBNP and BBRSP. For example, Echino-

cereus chisoensis var. chisoensis is endemic to one locale within BBNP and is federally listed as a threatened

species (Louie 1996). Echinomastus mariposensis, found in both parks, is listed as threatened, according to both

state and federal criteria. Quercus hinckleyi is known from only two areas in the United States: Shafter, Texas,

and the Solitario Dome in BBRSP (Hardy 1997; Powell 1998). In contrast, plant life on the greater expanses of

private land is less well understood, due to limited access and scarce funding for research on private lands.

However, as a result of change in land ownership, work on private land to the west of Terlingua has uncovered

a species not previously described (Turner & Nesom 2003). This brings to light the importance of work by

trained botanists on private property. For example, communication among local botanists about the presence

of Stemodia coahuilensis on RP enabled recognition of the species when it was subsequently recognized in

BBNP. In other regions of the country, similar work is being done to compare the floras of adjacent protected

and private lands (Chester 2003). Expanding held effort on private lands with owners increasingly aware of

conservation issues will only increase our understanding of the botanical resources of the Chihuahuan Desert.

Stemodia coahuilensis is a new record for Brewster County. It had been considered a strictly Mexican spe¬

cies, but work in recent years has resulted in collections in Presidio and Jeff Davis counties ( Worthington 25254

SRSC, Turner 24-492B SRSC). The existence of a fourth Texas S. coahuilensis population has been strongly sug¬

gested by a photograph taken by Roy Morey in the Ernst Tinaja area on the east side of BBNP (Fig. 2).

A new naturally occurring hybrid between Hibiscus coulteri and H. denudatus has been described and

named H. x sabei (Weckesser 2011). It has been documented in four sites in the Big Bend area. In habit it re¬

sembles H. coulteri (Fig. 3), but all specimens consistently differ from both parent species in Bower color and

several vegetative characters (Fig. 4).

The endemic Bora of gypseous deposits in the Chihuahuan Desert is one of the largest but least studied

restricted floras in North America (Moore & Jansen 2007). The Pen formation, described as gypsiferous (Max¬

well et al. 1967), is exposed within the study area along a stretch of the Long Draw drainage. It supports popu¬

lations of several gypsophilous taxa: Anulocaulis leiosolenus var. lasianthus, A. eriosolenus, Acleisanthes parvifo-

lius, Tiquilia gossypina, and T. hispidissima. Xylorhiza wrightii, common on the clay flats of the Pen Formation

360

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 2. Stemodia coahuilensis, near Ernst Tinaja, Big Bend National Park, Texas (photograph by Roy Morey).

and on other known gypsiferous sites in the Big Bend area, is a suspected gypsophile (M. Powell, pers. comm.).

The ubiquitous nature of gypsum in clay soils of RP is demonstrated by the distribution of gypsophiles on soils

not otherwise identified as gypsiferous. Known gypsophiles were found in the Long Draw drainage, on the Del

Rio clay, and on other clayey soils. These include Mentzelia mexicana, Psathyrotopsis scaposa, and Haploesthes

greggii var. texana.

Over 50% of the species included in the proceedings of the Texas Plant Conservation Conference (Clary

et al. 2002) are found in Trans-Pecos Texas. Several species from the current work are endemic to the Trans-

Pecos or to the Big Bend Region and are of special conservation concern.

Genistidium dumosum (Fig. 5) is a monotypic genus, endemic to the Chihuahuan Desert (Correll & John¬

ston 1970). Genistidium dumosum has been documented on only three sites in the U.S., all located on RP (Poole

1992), and five populations have been recorded in Mexico (Clary et al. 2002). It is ranked G1S1 by NatureServe

(2013) and it is under review for threatened or endangered status by the USFWS (2009). Attempts to relocate

the RP populations have had mixed results. Three previously documented populations were not relocated. A

population reported to have approximately 100 plants (M. Powell, pers. comm.) was not located. However, in

the search for that population, a population of 12 plants was located.

Kallstroemiaperennans is an endemic of the Trans-Pecos, its range limited to the western Edwards Plateau

near Langtry, Texas, southwest Brewster County, and adjacent areas of Presidio County (Fig. 6). It is ranked

G1S1 by NatureServe (2013) due to its limited distribution.

Chamaesyceperennans (Fig. 7) is a Big Bend endemic, restricted to the Terlingua-Lajitas area and adjacent

Chihuahua, Mexico. It is ranked G3S3 (NatureServe 2013) due to its limited distribution.

Lycium puberulum var. berberidoides (Fig. 8) and Lycium texanum (Fig. 9) are endemic to Trans-Pecos

Texas. Lycium texanum is considered a species of concern and is ranked G2S2 (NatureServe 2013).

Weckesser and Terry, Flora of Reed Plateau, Texas

361

Fig. 3. Hibiscus x sabei in flower. Highway 170 at Pepper's Hill. Reed Plateau study area, Brewster County, Texas.

Thelypodium texanum (Fig. 10) is endemic to the Big Bend region. Given its restricted distribution, it is

considered vulnerable and ranked G3S3 by NatureServe (2013).

Oenothera boquillensis (Fig. 11) is ranked as G3S2 (NatureServe 2013). Its range is limited to Brewster and

Presidio counties in Texas and the Mexican states of Chihuahua, Coahuila and Nuevo Leon. Oenothera boquil¬

lensis was also found in the Dead Horse Mountains of BBNP (Fenstermacher 2008), and though Hardy (1997)

did not find it in the SD, she noted that it was previously observed or collected there.

Echinomastus mariposensis (Fig. 12) was listed as a threatened species under the federal Endangered Spe¬

cies Act in 1979 and was listed in Texas in 1983 (NatureServe 2013). Its range is limited to Cretaceous lime¬

stone in the Big Bend area and in Coahuila, Mexico; it is common and locally abundant in the Terlingua area.

The non-native species of RP were initially encountered on or near roadways, homes, or construction

sites. Since the study period, the spread of invasive exotics such as Salsola tragus and Pennisetum ciliare is in¬

creasing dramatically, with highway and canyons serving as vectors from roadways and human habitation into

desert terrain.

Salsola tragus is common along the dirt road at the east end of RP that leads onto the plateau. It is abun¬

dant along the shoulders of Highway 170 and at the construction site near Villa de la Mina. Salsola tragus is

spreading along the drainages and dirt roads throughout RP and the drainages that cut across the highway.

Pennisetum ciliare was observed on the south slopes of the east end of RP and in Coultriris Canyon, a large

canyon that drains to the south of RP. It is common and abundant in the channels and gullies of Long Draw,

having spread from Hwy 170 after construction in 2003. Dense stands of P. ciliare have spread to the north past

362

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 4. Hibiscus x sabei buds and leaves. Highway 170 at Pepper's Hill, Reed Plateau study area, Brewster County, Texas. Note length of bracts in relation

to length of sepals and color of petals in bud (photograph by Betty Alex).

Weckesser and Terry, Flora of Reed Plateau, Texas

363

Fig. 5. Genistidium dumosum, in recently discovered population on Reed Plateau study area, Brewster County, Texas.

Villa de la Mina, following the 2-track road and spreading into the drainages that cross the road. This hardy

non-native grass, difficult to manage once established, has a high degree of reproductive vigor, a wide range of

adaptability, and few pests and predators (NatureServe 2013).

Tamarix chinensis is common in the Big Bend region. Tamarix chinensis is established but not yet a domi¬

nant species in Long Draw. It has also appeared in the right-of-way of Highway 170 since the road construction

in 2003.

Sorghum halepense is established in thick stands where roads cross low, wide gullies or drainages in the

Terlingua area. Though not collected, it was found in a construction site on RP.

Cynodon dactylon is a competitive, invasive weed (NatureServe 2013). Once established, it is difficult to

remove. It does not spread, though, beyond a water supply, so it remains restricted to irrigated, landscaped

home sites, or over septic systems.

Vegetation Associations

The topographic variation of RP—hills, ridges, drainages, arroyos, canyons, flats, gradual and steep slopes, and

cliffs—sets the stage for vegetation associations typical of the Chihuahuan Desert. In Henrickson and John¬

ston’s scheme (1986) topography plays a significant role in defining some of the associations (e.g., sandy arroyo,

canyon, or dune associations), and Butterwick and Lamb (1976) and Butterwick and Strong (1976a, 1976b,

1976c) based their vegetation associations on type of terrain in BBRSP. Fenstermacher (2008), also referring to

Henrickson andjohnston (1986), added elevation to the criteria used to describe the vegetation associations of

the DH. But in these works the mosaic patterns of the assemblages of plants is stressed; trends and patterns

364

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 6. Kallstroemiaperennans, endemic to Trans-Pecos Texas. Reed Plateau study area, Brewster County, Texas.

exist and associations form with predictability, especially in topographically distinct areas, but there are not

always clear demarcations between adjacent associations.

The vegetation of RP closely follows the association descriptions outlined by Henrickson and Johnston

(1986), with some variations and modifications. The Mixed Desert Scrub association is widespread, intergrad¬

ing with Larrea Scrub, Lechuguilla Scrub, Gypsophilous Scrub, Sandy Arroyo Scrub, and Canyon Scrub asso¬

ciations.

The Yucca Woodland or Dasylirion Scrub association of Henrickson and Johnston (1986) refers to exten¬

sive areas dominated, at least visually, by Yucca spp. or Dasylirion spp. with a mixed grass-shrub understory.

No extensive area of RP is truly dominated by Yucca spp. or Dasylirion spp., but on some of the slopes and

higher ridges of RP there are stands of numerous Y. torreyi (Fig. 13) or D. leiophyllum (Fig. 14).

With changes in surface features and elevation, the grasses, especially Bouteloua ramosa, can dominate a

slope or a ridge top, forming a Grama Grassland association (Fig. 15). Bouteloua ramosa is by far the most plen¬

tiful grass on RP. The presence of this association demonstrates that a Grama Grassland association need not

be restricted to coarse, sandy soils of volcanic origin, as stated by Hendrickson and Johnston (1986).

Chaparral and Oak Woodland associations are not to be expected in low, desert terrain, yet elements of

these higher-elevation associations are combined here in one intriguing site, the previously mentioned canyon

that drains to the north into Long Draw (Fig. 16). Amidst the expected vegetation of the Canyon Scrub asso¬

ciation (Henrickson & Johnston 1986) there is one anomalous addition: a stand of Quercus vaseyana trees,

with a dense Q. vaseyana shrub understory. A species normally associated with shrub-dominated Montane

Chaparral, Q. vaseyana reaches the height of 3-4 m (10-12 ft) in this canyon. The presence of Q. vaseyana as

Weckesser and Terry, Flora of Reed Plateau, Texas

365

Fig. 7. Chamaesyceperennans, endemic to Big Bend region, Texas. Reed Plateau study area, Brewster County, Texas (photograph by Betty Alex).

woodland trees as well as chaparral shrubs speaks to the unique conditions in that canyon, especially deep and

narrow with sheer rock walls, unlike other canyons on RP. While it could be a relictual population, this seems

unlikely given its proximity to Villa de la Mina and the wood-bred mercury processing furnace in operation

during the brst half of the last century (Ragsdale 1976). The stand may represent new growth in the last 70

years.

A similar situation was encountered in SD (Hardy 1997), where elements of higher elevation and mesic

associations were encountered at lower elevations in protected canyons. In several small canyons, Quercus

grisea, also identibed with the Chaparral association, approaches the tree form expected in the Oak Woodland

association, along with Juniperus pinchotii of the Pinyon-Juniper association.

An especially diverse blend of associations occurs along portions of Long Draw, where the main course of

the dry stream bed butts against the base of the cliffs of the Terlingua monocline. The monocline is a generally

north-facing escarpment, irregularly cut by numerous canyons, drainages, and pour-offs. The water channeled

over the cliffs and the north exposure creates a community comprised of elements of the Sandy Arroyo and

Canyon Scrub associations (Henrickson & Johnston 1986). At the base of the cliffs, fairly steep scree slopes of

Santa Elena limestone breakdown, strewn with boulders, have become overgrown with large shrubs and sev¬

eral tree species. There is a dense understory of smaller shrubs, subshrubs, and perennial herbs in the resulting

soils, high in organic matter, and a number of annuals present after summer rains. Several species normally

associated with the more mesic habitats in canyons or at higher elevations are intermixed with species ex¬

pected in the dry arroyos of Mixed Desert Scrub and Sandy Arroyo associations. This particular assemblage is

another example of the overlap and intergradation of the vegetation associations of the CD.

366

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 8. Lycium puberulum var. berberidoides, endemic to Trans-Pecos Texas. Reed Plateau study area, Brewster County, Texas.

An additional association not described by Henrickson and Johnston merits consideration. In several lo¬

cations in the Big Bend area, including RP, stands of Fouquieria splendens (ocotillo) are so dense as to create the

appearance of a forest. In the F. splendens associations of RP, the understory varies, from sparse vegetation of

widely scattered shrubs to dense vegetation with shrubs and grasses from the desert scrub associations, Farrea

tridentata often appearing as a co-dominant. A recent vegetation survey in BIBE has generated descriptions of

vegetation associations, including a series of associations with F. splendens as a nominal component (Lea 2014).

The Ocotillo-Creosotebush-Lechuguilla Desert Scrub association describes the most common association on

RP, with a significant variation. As with the Grama Grassland association of Henrickson and Johnston (1986),

the occurrence of the Ocotillo-Creosotebush-Lechuguilla Desert Scrub association on RP demonstrates that

this association is not restricted to non-calcareous substrates, as described for BIBE. Further description and

delineation of the Desert Scrub vegetation association continuum on RP is warranted.

ANNOTATED CHECKTIST OF THE SPECIES OF REED PLATEAU

AND ADJACENT AREAS

The annotated species list for RP is arranged phylogenetically following Evert and Eichhorn (2013), with Phy¬

lum Anthophyta divided into Monocotyledones and Eudicotyledones. Family, genus, species, and lower rank¬

ings are listed alphabetically. All collections of the first author cited in the list, designated WW, are housed at

SRSC. Taxa previously collected on RP, as documented by voucher specimens in SRSC and TEX-LL, are in¬

cluded in the list with collector and herbarium information. Taxa observed but not collected during the cur¬

rent study are included in the list and noted as such.

Weckesser and Terry, Flora of Reed Plateau, Texas

367

Fig. 9. Lycium texanum, endemic to Trans-Pecos Texas. Reed Plateau study area, Brewster County, Texas.

The information for each species includes scientific name and authorship, common name, nativity, spe¬

cial status, abundance, habitat, and the author’s collection number of a representative specimen. Where appli¬

cable, notations for status for invasive, threatened, or endangered species are also included. Significant addi¬

tional information is included at the end of the description. Native species are noted with N; non-native species

with I (Texaslnvasives.org 2008). Species endemic to Trans-Pecos Texas (E-TP) or to the Big Bend Region (E-B)

are noted (Correll & Johnston 1970; Clary et al. 2002; TAM-BWG 2007). Species ranked for conservation pur¬

poses, with state or federal protection status, are also noted (USDA-PLANTS 2014; NatureServe 2013).

Habitat terms are general descriptions of common habitat for a given species. Habitat descriptors used in

the species list are: alluvium - sand and/or gravel with organic debris in arroyos, drainages or canyon bottoms;

arroyo - dry wash or stream bed, seasonally flooded; drainage - cut in side of slope, with varying width and

steepness; canyon - vertical-sided cut through limestone; clay flat - flat to gentle slope of clay substrate; cliffs

- sheer-faced limestone, as seen in the Terlingua uplift or in canyons; disturbed site - current home site or

historically disturbed areas; ridges - top of slopes, knife-edge summits, or saddles between higher hills; road¬

side - along dirt roads or on shoulders of paved highway; rock outcrop - exposed rock surface or blocks; slope

- relatively smooth terrain tilted with varying aspect and degrees of steepness; ubiquitous - occurs in most or

all habitats throughout study area.

A scheme to describe abundance was derived from Palmer et al. (1995). Determination of abundance was

based on held observations during the course of the study. The terms used here are: rare (R); uncommon (U);

occasional (O); common (C); and abundant (A).

368

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 10. Thelypodium texanum (white flowers), endemic to the Big Bend Region, Texas, with Namahavardii (pink flowers), along Highway 170. Reed

Plateau study area, Brewster County, Texas.

LYCOPODIOPHYTA

Selaginellaceae

Selaginella lepidophylla (Hook. &Grev.) Spring, Resurrection fern, N,

A, slopes, canyons, ridges, open rocky areas, WW184

MONILOPHYTA

Pteridaceae

Argyrochosmo microphyllo (Mett. ex Kuhn) Windham, Littleleaf

cloakfern, N, U, sheltered rock outcrops, WW79A

Astrolepis cochisensis (Goodd.) D.M. Benham & Windham, Cochise

scaly cloakfern, N, U, sheltered rock outcrops, WW78

Astrolepis integerrimo (Hook.) D.M. Benham & Windham, Wholeleaf

cloakfern, N, U, sheltered rock outcrops, WW77

Cheilanthes alabamensis (Buckley) Kunze, Alabama lipfern, N,

sinkhole, Barbee & Powell 4, SRSC

Cheilanthes horridula Maxon, Rough lipfern, N, U, sheltered rock

outcrops, WW 675

Weckesser and Terry, Flora of Reed Plateau, Texas

369

Fig. 11. Oenothera boquillensis, found in Long Draw at base of cliffs, Reed Plateau study area, Brewster County, Texas.

Cheilanthesvillosa Davenp. ex Maxon, Villous lipfern, N, U, sheltered

rock outcrops, WW679

GNETOPHYTA

Ephedraceae

Ephedra aspera Engelm. ex S. Watson, Rough jointfir, N, C, ubiq¬

uitous, WW369

ANTHOPHYTA, Monocotyledones

Agavaceae

Agave lechuguilla Torr., Lechuguilla, N, A, ubiquitous, WW1004

Bromeliaceae

Hechtia texensis S. Watson, Texas false agave, N, C, limestone out¬

crops, slopes, WW433

Cyperaceae

Eleocharis montevidensis Kunth, Sand spikerush, N, R, drainage,

one population was found after significant rain at the edge of

a water-filled earthen tank, WW 1008

Liliaceae

Allium kunthii G. Don, Kunth onion, N, U, north-facing gravelly

slopes, WW34

Dasylirion leiophyllum Engelm. ex Trel., Sotol, N, O, slopes, drain¬

ages, WW804A

Yucca torreyi Shafer, Torrey yucca, N, O, slopes, drainages, WW276

Poaceae

Aristidapurpurea var. longiseta (Steud.) Vasey, Red three-awn, N, O,

gravelly slopes, WW750

Aristida purpurea var. purpurea Nutt., Purple three-awn, N, O, slopes,

ridges, WW76I

Aristida purpurea var. nealleyi (Vasey) Allred, Blue three-awn, N, O,

slopes, ridges, WW606

Bothriochloa barbinodis (Lag.) Herter, Cane bluestem, N, D.5. Correll

& M.C. Johnston 24466 TEX-LL

Bothriochloa laguroides (DC.) Herter var. torreyana (Steud.) M. Marchi

& Longhi-Wagner, Silver bluestem, N, U, drainage, disturbed

site, WW575

Bouteloua barbata Lag., Six-week grama, N, U, drainage, disturbed

site, WW571

Bouteloua curtipendula (Michx.)Torr., Side-oats grama, N, O, gravelly

drainage, clay, WW731

Bouteloua ramosa Scribn. ex Vasey, Chino grama, N, A, ubiquitous,

WW601

Bouteloua trifida Thurb., Red grama, N, O, gravelly slopes, WW605

Cathestecum erectum Vasey & Hack., False grama, N, slopes, al¬

luvium, Hughes 337 SRSC

370

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 1 2 . Echinomastus mariposensis in flower with Bahia absinthifoliavar. dealbata, near Villa de la Mina. Reed Plateau study area, Brewster County, Texas.

Cynodon dactylon (L.) Pers., Bermuda grass, I, U, disturbed sites,

WW568

Dasyochloa pulchella (Kunth) Willd. ex Rydberg, Fluff grass, N, A,

ubiquitous, WW565A

Digitaria californica (Benth.) Henrard, California cottontop, N, clay

flats with gravel, Powell5362 SRSC

Digitorio cognoto (Schult.) Pilg., Fall witchgrass, N, O, gravelly

drainage, WW825

Enneopogon desvouxii P. Beauv., Feather pappusgrass, N, O, slopes,

drainages, WW820

Heteropogon contortus (L.) P. Beauv. ex Roem. & Schult.,Tanglehead,

N, C, ridges, slopes, drainages, WW789

Muhlenbergio ported Scribn. ex Beal, Bush muhly, N, U, drainages,

canyon slopes, WW 1046

PanicumhalliiVasey, Hall panicgrass, N, U, gravelly slopes, WW 1003

Pappophorum vaginatum Buckley, Whiplash pappusgrass, N, O,

disturbed clay flats, drainages, WW573

Pennisetum ciliore (L.) Link, Buffelgrass, I, C, roadside, disturbed sites,

drainages, spreading into canyons and drainages since 2003

road construction, WW574

Pleurophis mutica Buckley, Tobosa grass, N, O, disturbed slopes,

limestone and clay, WW 1009

Poo bigelovii Vasey & Scribn., Bigelow bluegrass, N, O, canyons,

drainages, WW 163

Setorio mocrostochyo Kunth, Streambed bristlegrass, N, O, canyon,

drainages, clay flats, WW1015

Sporoboluspyromidotus (Lam.) Hitch., Madagascar dropseed, N, O,

disturbed sites, clay flats, Weckesser 572

Tridens muticus (Torr.) Nash, Slim tridens, N, O, canyon bottom,

drainages, Weckesser 947A

ANTHOPHYTA, Eudicotyledones

Acanthaceae

Corlowrightio orizonico A. Gray, Arizona carlowrightia, N, U, steep

slopes, WW415

Ruellio porryi A. Gray, Parry wild petunia, N, C, slopes, drainages,

canyon, WW366

Stenandrium borbotum Torr. & A. Gray, Shaggy stenandrium, N, O,

restricted to rock outcrops, WW237

Amaranthaceae

Iresine leptoclada (Hook, f.) Henrickson & S.D. Sundb., Texas shrub,

N, base of cliffs, DS.Correll30709 TEX-LL

Tidestromia carnosa (Steyerm.) I.M. Johnst., Fleshy tidestromia, N,

U, gravelly slopes, WW755

Tidestromia lanuginosa var. lanuginosa (Nutt.) Standi., Wooly

tidestromia, N, C, gravelly slopes, WW59

Tidestromia suffruticosa (Torr.) Standi., Shrubby honeysweet, N, C,

gravelly slopes, alluvium, 1/1/1/1/85

Anacardiaceae

Rhusmicrophylla Engelm., Littleleaf sumac, N, O, canyons, drainages,

alluvium, WW471

Weckesser and Terry, Flora of Reed Plateau, Texas

371

Fig. 13. Yucca torreyi, Reed Plateau study area, Brewster County, Texas.

Rhus virens Lindh. ex A. Gray, Evergreen sumac, N, U, steep sided

canyons, WW553

Apocynaceae

Amsonialongiflora Torr.,Tubular slimpod, N, U, clay flats, drainages,

canyons, WW 199

Mandevilla macrosiphon (Torr.) Pichon, Rocktrumpet, N, C, slopes,

drainages, alluvium, WW585

Aristolochiaceae

Aristoiochia coryi I.M. Johnst., Dutchman's pipe, N, R, drainage, one

population found at the edge of a water filled earthen tank

after significant rain, WW903

Asdepiadaceae

Asclepias asperula (Decne.) Woodson, Antelope-horns milkweed,

N, U, slopes, drainages, WW375

Asclepias oenotheroides, Schltdl. & Cham., Zizotes milkweed, N, U,

gravelly ridges, WW805

Funastrum crispum (Benth.) Schltr., Wavyleaf twinevine, N, R, gravelly

slope, WW486

Funastrum torreyi (A. Gray) Schltr., Soft twinevine, N, R, canyons,

WW907

Matelea parvifolia (Torr.) Woodson, Spearleaf milkvine, N, R, rock

crevices, WW395

Asteraceae

Artemisia ludoviciana Nutt., White sagebrush, N, U, alluvium, WW

1049

Bahia absinthifolia var. dealbata (A. Gray) A. Gray, Dealbata bahia,

N, C, slopes, drainages, ridges, WW213

Chaetopappa bellioides (A. Gray) Shinners, Manyflower leastdaisy,

N, O, slopes, drainages, WW465

Cirsium turned Warnock, Turner thistle, N, U, restricted to cliff faces

of canyons, WW 930

Conoclinium dissectum A. Gray, Palm-leaf mistflower, N, R, canyon,

WW816

Dyssodia acerosa DC., Prickleleaf dogweed, N, C, slopes, ridges,

WW831

Dyssodiapentachaeta (DC.) B.L. Rob., Prickly dogweed, N, A, slopes,

ridges, alluvium, drainages, WW60

Erigeron bigeloviiA. Gray, Bigelow fleabane, N, slopes, ridges, along

cliffs, Worthington 968 TEX-LL

Erigeron tracyi Greene, Running fleabane, N, C, slopes, ridges,

drainages, WW664

Gymnosperma glutinosum (Spreng.) Less., Tatalencho, N, C, ubiq¬

uitous, WW 76

Flaploesthes greggii var. texana (J.M. Coult.) I.M. Johnst., False

broomweed, N, O, clay flats, alluvium, slopes, drainages, 1/1/1/1/58

Flelenium microcephalum var. ooclinium (A. Gray) Bierner, Sneeze-

weed, N, R, drainage, one population found at the edge of a

water filled earthen tank after significant rain, WW905

372

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 14. Dasylirion leiophyllum, Reed Plateau study area, Brewster County, TX.

Heterotheca fulcrata (Greene) Shinners, Rocky goldaster, N, U, north¬

facing canyon, 1/1/1/1/929

Jefea brevifolia (A. Gray) Strother, Shorthorn zexmenia, N, O, canyon

sediment, WW657

Melampodium leucanthum Torr. & A. Gray, Blackfoot daisy, N, O,

drainages, canyons, slopes, WW536

Parthenium confertum A. Gray, Lyreleaf parthenium, N, O, alluvium,

gravelly slopes, WW9

Parthenium incanum Kunth, Mariola, N, C, ubiquitous, WW592

Perityle parryi A. Gray, Heartleaf rockdaisy, N, U, restricted to cliff

face, WW 1053

Perityle vaseyi J.M. Coult., Margined perityle, N, U, restricted to

gypseous clay, WW847

Porophyllum scoparium A. Gray, Shrubby Poreleaf, N, O, gravelly

slope, canyon, 1/1/1/1/393

Psathyrotopsis scaposa (A. Gray) H. Rob., Naked brittlestem, N, U,

restricted to gypseous clay, WW 193

Psilostrophe tagetina (Nutt.) Greene, Wooly paperflower, N, A,

ubiquitous, WW 141

Stephanomeria pauciflora (Torr.) A. Nelson, Brownplume wirelettuce,

N, U, drainage, alluvium, 1/1/1/1/77

Thelesperma longipes A. Gray, Longstalk greenthread, N, O, slopes,

drainages, 1/1/1/1/90

Thelesperma megapotamicum (Spreng.) Kuntze var. megapota-

micum, Rayless greenthread, N, C, slopes, drainages, canyon

bottom, WW890

Trixis californica Kellogg, American trixis, N, O, drainages, canyons,

WW 73

Viguiera dentata (Cav.) Spreng., Sunflower goldeneye, N, U, al¬

luvium, WW 1050

Viguiera stenoloba S.F. Blake, Skeletonleaf goldeneye, N, A, ubiq¬

uitous, WW 22

Xanthismaspinulosum var. chihuahuanum (B.L.Turner & R.L. Hartm.)

D.R. Morgan & R.L. Hartm., Lacy tansyaster, N, O, disturbed sites,

clay flats, 1/1/1/1/272

Xanthium strumarium L., Cocklebur, N, R, alluvium, 1/1/1/1/95

Xylorhiza wrightii (A. Gray) Greene, Gyp daisy, N, U, restricted to

gypseous clays; 1/1/1/1/377

Xylothamia triantha (S.F. Blake) G.L. Nesom, Trans-Pecos desert

goldenrod, N, R, clay flats with limestone gravel, WW578

Zinnia acerosa (DC.) A. Gray, Spinyleaf zinnia, N, C, slopes, ridges,

drainages, canyons, WW456

Berberidaceae

Mahonia trifoliolata Mode., Agerita, N, R, canyons, deep drainages,

WW283

Bignoniaceae

Chilopsis linearis (Cav.) Sweet, Desert willow, N, U, alluvium, WW 1035

Tecomastans (L.) Juss. ex Kunth,Trumpetflower, N, O, slopes, drain¬

ages, canyons, WW 19

Weckesser and Terry, Flora of Reed Plateau, Texas

373

Fig. 15. South-facing slope dominated by Boutelouaramosa. Reed Plateau study area, Brewster County, Texas.

Boraginaceae

Cryptantha coryi I.M. Johnst., Cory cryptantha, N, R, north-facing

slopes, gravel and clay, WW206

Cryptanthamexicana (Brandegee) I.M. Johnst., Mexican cryptantha,

N, O, ubiquitous, WW385

Heliotropiumpowelliorum B.L.Turner, Powell heliotrope, N, C, slopes,

drainages, arroyos, WW513

Lappula redowskii (Hornem.) Greene, Flatspine stickseed, N, U,

alluvium, WW 164

Omphalodes aliena A. Gray ex Hemsl., Mexican navelwort, N, U,

rocky slopes, canyons, WW 114

Tiquilia canescens (A. DC.) A.T. Richardson, Woody crinklemat, N, A,

ubiquitous, WW582

Tiquilia gossypina (Wooton & Stand.) A.T. Richardson, Texas crinkl¬

emat, N, U, clay and gravel, slopes, ridges, WW569

Tiquilia greggii (Torr. & A. Gray) A.T. Richardson, Plumed tiquilia, N,

C, slopes, arroyos, WW493

Tiquilia hispidissima (Torr. & A. Gray) A.T. Richardson, Rough cold-

enia, N, O, slopes, ridges, clay flats, drainages, WW734

Tiquilia mexicana (S. Watson) A.T. Richardson, Mexican crinklemat,

N, C, ubiquitous, WW406

Brassicaceae

Descurainiapinnata (Walter) Britton,Tansymustard, N, R, alluvium,

WW151

Draba cuneifolia Nutt, ex Torr. & A. Gray, Whitlow-wort, N, R, al¬

luvium, 1/1/1/1/7 75

Nerisyrenia camporum (A. Gray) Greene, Bicolor mustard, N, A,

ubiquitous, WW 175

Physaria fendleri (A. Gray) O'Kane & Al-Shehbaz, Fendler bladderpod,

N, C, slopes, drainages, alluvium, WW 109

Streptanthus carinatus C. Wright ex A. Gray, Lyreleaf twistflower, N,

U, north-facing rocky slopes, alluvium, WW 166

Thelypodium texanum (Cory) Rollins, Texas thelypody, N, C, E-BB,

slopes, canyon bottoms, alluvium, clay flats, WW 120

Cactaceae

Ariocarpus fissuratus (Englem.) K. Schum., Living rock cactus, N, C,

slopes, drainages, WW80

Coryphantha echinus (Engelm.) Britton & Rose, Sea-urchin cactus, N,

O, slopes, ridges, WWs.n., photographic record only

Coryphantha sneedii var. albicolumnaria (Hester) A.D. Zimmerman,

Silverlace cactus, N, U, G2G3S2S3, slopes, ridges north end of

RP, WWs.n., photographic record only

Echinocactus horizonthalonius Lem., Eagle claw cactus, N, C, slopes,

ridges, arroyos, Weckessers.n., photographic record only

Echinocereus dasyacanthus Engelm., Rainbow cactus, N, C, slopes,

ridges, arroyos, WW475

Echincereus enneacanthus Engelm., Strawberry cactus, N, O, slopes,

drainages, WW997

374

Journal of the Botanical Research Institute of Texas 8(1)

Fig. 1 6 . Tree form of Quercus vaseyana in steep-sided limestone canyon. Reed Plateau study area, Brewster County, Texas.

Echinocereus stramineus (Engelm.) F. Seitz, Strawberry pitaya, N,

A, slopes, ridges, drainages, arroyos, WWs.n., photographic

record only

Echinomostus mariposensis Hester, Mariposa cactus, N, U, G2S2, LT/T,

slopes, ridges, drainages, WW996

Epithelontho bokei L.D. Benson, Boke's button cactus, N, U, gravel-

covered flat to gently sloping rock outcrops, WW564A

Ferococtus homotacanthus (Muehlenpf.) Britton & Rose, Giant fish¬

hook cactus, N, U, slopes, ridges, arroyos, canyons, WW s.n.,

photographic record only

Glondulicoctus uncinatus var. wrightii (Engelm.) Backeb., Eagleclaw

cactus, N, U, slopes, ridges, WW s.n., photographic record only

Mammillaria lasiacantha Engelm., Golfball cactus, N, U, slopes,

ridges, WW565

Mammillaria pottsii Scheer exSa\m-Dyck, Rattail mammillaria, N,C,

slopes, ridges, clay flats, drainages, WW563

Opuntia aggeria Ralston & Hilsenb., Dog cholla, N, C, ubiquitous,

WW476

Opuntia camanchica Engelm. & J.M. Bigelow, Comanche pricklypear,

N, A, drainages, arroyos, slopes, ridges, WW923

Opuntia engelmannii Salm-Dyckex Engelm., Engelmann pricklypear,

N, O, slopes, drainages, arroyos, WW 1058

Opuntia leptocaulis DC., Christmas cactus, N, O, drainages, slopes,

arroyos, WW848A

Opuntia rufida Engelm., Blind pricklypear, N, C, drainages, slopes,

arroyos, canyons, WW995

Celastraceae

Mortonia scabrella A. Gray, Tickbush, N, O, clay flats, drainages,

WW497

Chenopodiaceae

Atriplexcanescens (Pursh.) Nutt., Four-wing saltbush, N,0, alluvium,

drainages, WW643

Salsola tragus L., Tumbleweed, I, C, roadsides, arroyos, drainages,

spreading into canyons and drainages following road construc¬

tion in 2003, WW848

Convolvulaceae

Bonamia repens (I.M. Johnst.) D.F. Austin & Staples, Creeping lady's

nightcap, N, O, rock outcrops, rocky slopes, WW804

Convolvulus equitans Benth.,Texas bindweed, N, O, disturbed sites,

clay and gravel, WW950

Evolvulus alsinoides (L.) L., Ojo de vibora, N, A, slopes, drainages,

canyons, WW988

Ipomoeacostellatalorr., Crestrib morning glory, N, R, sediments of

canyon bottom, WW648

Cucurbitaceae

Ibervillea tenuisecta (A. Gray) Small, Slimlobe globeberry, N, O, ar¬

royos, alluvium, drainages, WW609

Ebenaceae

Diospyros texana Scheele, Texas persimmon, N, O, deep drainages,

arroyos, canyons, WW669

Weckesser and Terry, Flora of Reed Plateau, Texas

375

Euphorbiaceae

Bernardia obovato I.M. Johnst., Myrtlecroton, N, O, ridges, gravelly

slopes, Weckesser 688

Chomoesyce olbomorginoto (Torr. & A. Gray) Small, Whitemargin

sandmat, N, gravelly slope, disturbed site, Bennack 108B SRSC

Chomoesyce perennons Shinners, Perennial sandmat, N, C, E-BB,

ubiquitous, WW738

Chomoesyce serrulo[Er\ge\rr\.)\Nootor\ & Standi., Sawtooth sandmat,

N, gravelly slope, disturbed site, Bennock 108A SRSC

Chomoesyce stictosporo (Engelm.) Small, Slimseed spurge, N, U,

arroyo, WW 632

Croton fruticulosus Engelm. ex Torr., Bush croton, N, U, canyons,

steep drainages, WW660

Croton pottsii (Klotzsch.) Mull. Arg., Leatherweed Croton, N, A,

ubiquitous, WW964

Argythomnio neomexicono Mull. Arg., New Mexico wild mercury, N,

O, drainages, canyons, WW904

Euphorbia antisyphiliticaZucc., Candelilla, N, A, ubiquitous, WW 966

Euphorbia exstipulata Engelm., Squareseed spurge, N, base of cliff,

arroyo, Bennack 112 SRSC

Jatropha dioica var. graminea McVaugh, Leatherstem, N, A, ubiq¬

uitous, WW985

Phyllanthus polygonoidesNutt. ex Spreng., Knotweed leafflower, N,

C, drainages, arroyos, canyons, WW 104

Tragia amblyondonta (Mull. Arg.) Pax & K. Hoffm., Dogtooth nose-

burn, N, U, shaded arroyo, WW 1048

Tragia ramosa Torr., Branched noseburn, N, 0, drainages, arroyos,

canyons, WW49

Fabaceae

Acacia greggii A. Gray, Catclaw acacia, N, C, slopes, drainages,

ridges, WW458

Acacia neovernicosa Isely, Varnished acacia, N, C, slopes, drainages,

ridges clay flats, WW43

Acacia roemeriana Scheele, Roemer acacia, N,0, ubiquitous, WW891

Acacia schottii Torr., Schott acacia, N, U, slopes, WW 1010

Calliandra iselyi B.L.Turner, False mesquite, N,0, ubiquitous, WW 584

Daleaformosa Torr., Feather dalea, N, 0, ubiquitous, WW309

Dalea neomexicana (A. Gray) Cory, New Mexico dalea, N, 0, slopes,

ridges, WW398

Daleapogonathera A. Gray, Bearded dalea, N, U, canyon, WW328

Dalea wrightii A. Gray, Wright dalea, N, C, ubiquitous, WW 15

Desmanthus glandulosus (B.L. Turner) Luckow, Bundleflower, N,

slopes, drainages, canyon, WW663

Genistidium dumosum I.M. Johnst., Johnston bushpea, N, R, G1S1,

rocky slopes, only known populations in U.S.A., WW946

Mimosa emoryana Benth., Emory mimosa, N, 0, drainages, canyons,

WW436

Mimosa texana (Gray) Small, Catclaw mimosa, N, 0, slopes, canyons,

disturbed sites, WW429

Pomaria melanosticta S. Schauer, Parry holdback, N, 0, slopes,

drainages, canyons, WW5

Prosopisglandulosalon., Mesquite, N, C, ubiquitous, WW856

Senna lindheimeriono (Scheele) H.S. Irwin & Barneby, Lindheimer

senna, N, C, ubiquitous, WW524

Sennapilosior (B.L. Rob. ex J.F. Macbr.) H.S. Irwin & Barneby,Trans-

Pecos senna, N, 0, drainages, arroyos, slopes, WW590

Vida ludoviciono Nutt. exTorr.& A. Gray, Deer pea vetch, N, U, ar¬

royos, WW 162

Fagaceae

Quercus voseyono Buckley, Sandpaper oak, N, R, restricted to one

narrow, steep-sided canyon, WW517

Fouquieriaceae

Fouquieria splendens Engelm., Ocotillo, N, C, slopes, ridges, drain¬

ages, WW405

Gentianaceae

Zeltnera arizonica (A. Gray) A. Heller, Arizona centaury, N, R, shaded

slope, one population found after significant rain, WW357

Hydrophyllaceae

Nama havardii A. Gray, Havard nama, N, C, ubiquitous, WW220

Nama hispida A. Gray, Rough nama, N, U, alluvium, canyon bot¬

tom, WW179

Phocelio congesto Hook., Bluecurls, N, U, shaded alluvium, canyons,

WW165

Krameriaceae

Krameria erecta \N\\\d. ex Schult., Range ratany, N, C, slopes, ridges,

arroyos, canyons, WW403

Krameria grayi Rose & Painter, White ratany, N, C, slopes, ridges,

arroyos, canyons, WW69

Lamiaceae

Hedeoma drummondii Benth., Drummond pennyroyal, N, R, allu¬

vium, found once on gravel bar in dry wash, WW865

Hedeoma nana (Torr.) Briq., Dwarf false pennyroyal, N, U, slopes,

drainages, ridges, canyons, WW 129

Linaceae

Linum berlandieri Hook., Berlandier flax, N, C, ubiquitous, WW 110

Linum rupestre (A. Gray) Engelm. ex A. Gray, Rock flax, N, U, can¬

yons, WW714

Loasaceae

Cevallia sinuata Lag., Stinging cevallia, N, 0, slopes, drainages,

disturbed areas, WW914

Eucnidebortonoides Zucc., Yellow rocknettle, N, R, restricted to rock

walls of steep drainages and canyons, WW447

Mentzelia mexicana H.J.Thomps. &Zavort., Mexican blazingstar, N,

0, slopes, drainages, ridges, arroyos, WW839

Mentzeliapochyrhizo I.M. Johnst., Coahuila blazingstar, N, R, slope,

WW 1044

Loganiaceae

Buddlejo morrubiifolio Benth., Butterfly bush, N, C, arroyos, drain¬

ages, canyons, WW670

Malpighiaceae

Janusia gracilis A. Gray, Helicopter bush, N, U, canyons, steep

drainages, WW958

Malvaceae

Abutilon crispum (L.) Medik., Netvein mallow, N, slopes, drainages,

Powell 3185 SRSC

Abuliton fruticosum Guill. & Perr., Pelotazo, N, C, slopes, drainages,

canyons, WW490

Abutilonmalacum S.Watson, Yellow indian mallow, N, U,disturbed

site, WW1041

Abutilon wrightii A. Gray, Wright abutilon, N, U, cliff face in steep

canyon, WW327

Hibiscus coulteri Harv. ex A. Gray, Desert rosemallow, N, O, slopes,

ridges, WW6!

Hibiscus x sabei Weckesser, Ken's rosemallow, N, R, roadside, WW

645

Hibiscus denudatus Benth., Paleface rosemallow, N, O, slopes, flats,

drainages, WW97

Sida abutifolia Mill., Spreading sida, N, O, slopes, drainages, rock

outcrops, WW 12

SidalongipesA. Gray, Stockflowerfanpetals, N, O, slopes, drainages,

clay flats, WW216A

376

Nyctaginaceae

Acleisanthes angustifolia (Torr.) R.A. Levin, Narrowleaf moonpod, N,

O, slopes, drainages, ridges, clay flats, WW72 1

Acleisanthes longiflora A. Gray, Angel trumpets, N, O, drainages,

canyons, WW770

Acleisanthesparvifolia (Torr.) R.A. Levin, Small leaf moonpod, N, U,

restricted to gypseous clay, WW36

Allionia incarnata L.,Trailing four-o'clock, N, A, ubiquitous, WW639

Anulocaulis eriosolenus (A. Gray) Standi., Big Bend ringstem, N, R,

restricted to gypseous clay, WW 1045

Anulocaulis leiosolenus var. lasianthus I.M. Johnst., Ringstem, N, U,

restricted to gypseous clay flats, WW 1042

Cyphomeris gypsophiloides (M. Martens & Galeotti) Standi., Birdfruit,

N, R, cliff base, WW1055

Mirabilis texensis (J.M. Coult.) B.L. Turner, Texas mirabilis, N, R, cliff

base, WW 1056

Oleaceae

Forestiera angustifolia Jorr., Desert olive, N, C, ubiquitous, WW318

Menodora longiflora Engelm. ex A. Gray, Twinpod, N, U, arroyo,

WW 102

Menodorascabra A. Gray, Rough menodora, N,0, ubiquitous, WW35

Onagraceae

Oenothera boquillensis (P.H. Raven & D.P. Greg.) W.L. Wagner & Hoch,

Boquillas gaura, N, R, G2S2, arroyo margin, WW 1054

Oenothera brachycarpa A. Gray, Shortpod evening primrose, N,

slopes, drainages, clay flats, disturbed areas, WW723

Oenothera hartwegii Benth., Hartweg sundrops, N, U, arroyos,

WW1032

Oenothera kunthiana (Spach) Munz, Kunth sundrops, N, shaded

slope, drainage, Hughes s.n., SRSC

Oenothera rosea L'Her. ex Aiton, Rose sundrops, N, U, drainages,

slopes, disturbed areas, WW256

Oenothera triloba Nutt., Stemless evening primrose, N, U, slopes,

clay, WW226

Orobanchaceae

Castilleja rigida Eastw., Broadbract paintbrush, N, O, slopes, drain¬

ages, canyons, WW778

Orobanche multicaulis Brandegee, Spiked broomrape, N, R, alluvium,

drainage, middle of a gravelly wash, WW462

Phytolaccaceae

Rivinahumilis L., Pigeonberry, N, R, one plant at cliff base, WW 1047

Plantaginaceae

Penstemon baccharifolius Hook., Baccharis-leaf penstemon, N, R,

restricted to crevices in canyon walls and bottoms, WW899

Plantago helleri Small, Heller plantain, N, U, shaded alluvium,

slopes, WW299

Stemodia coahuilensis (Henrickson) B.L. Turner, Coahuila twintip,

N, R, canyon bottom, alluvium, new Brewster County record,

WW690

Polemoniaceae

Gilia rigidula subsp. acerosa A. Gray, Bluegilia, N,0, ridgesand slopes

of Santa Elena limestone, WW 136

Giliapurpusii subsp. stewartii (I.M. Johst.) J.M. Porter, Stewart gilia,

N, C, ubiquitous, 1/1/1/1/87

Polygalaceae

Hebecarpa barbeyana (Chodat) J.R. Abbott, Narrowleaf milkwort,

N, C, arroyos, drainages, canyons, WW26

Hebecarpa macradenia (A. Gray) Abbott, Glandleaf milkwort, N, C,

rock outcrops, WW491

Rhinotropis lindheimeri var. parvifolia (Wheelock) R.R. Abbott,

Shrubby milkwort, N, R, gravelly clay flats, WW881

Journal of the Botanical Research Institute of Texas 8(1)

Polygalascoparioides Chodat, Broom milkwort, N, C, arroyos, drain¬

ages, canyons, slopes, WW724

Portulacaceae

Portulaca pilosa L., Kiss me quick, N, O, ridges, slopes, flats with

gravel surface, WW961

Ranunculaceae

Clematis drummondiiJorr.8iA. Gray, Virgin's bower, N, C, drainages,

arroyos, canyons, WW94

Resedaceae

Oligomeris linifolia (Vahl) Macbride, Desert spike, N, C, drainages,

arroyos, canyons, WW911

Rhamnaceae

Condalia ericoides (A. Gray) M.C. Johnst., Javelinabush, N, O, ubiq¬

uitous; WW473

Condalia warnockii M.C. Johnst., Warnock condalia, N, Powell3633

SRSC

Ziziphus obtusifolia (Hook, ex Torr. & A. Gray) A. Gray, Lotebush, N,

O, ubiquitous, WW894

Rosaceae

Fallugia paradoxa (D. Don) Endl. ex Torr., Apache plume, N, O, ar¬

royos, drainages, canyons, WW 105A

Amelanchier denticulata (Kunth) K. Koch, Serviceberry, N, R, drain¬

age, northern-most extent of Long Draw, WW279

Purshia ericifolia (Torr. ex A. Gray) Henrickson, Heath cliffrose, N, R,

cliff faces of canyons and steep drainages, WW 949

Rubiaceae

Galium proliferum A. Gray, Limestone bedstraw, N, U, slopes, drain¬

ages, WW 263

Hedyotis nigricans (Lam.) Fosberg, Prairie Bluet, N, O, drainages,

slopes, canyons, especially in rock crevices, WW43

Rutaceae

Thamnosma texanum (A. Gray) Torr., Dutchman's britches, N, C,

ubiquitous, WW 140

Sapindaceae

Ungnadia speciosa Endl., Mexican buckeye, N, R, arroyos and

canyons, WW319

Scrophulariaceae

Leucophyllum minus A. Gray, Big Bend silverleaf, N, C, slopes, drain¬

ages, arroyos, WW960

Maurandella antirrhiniflora Humb. & Bonpl. ex Willd., Snapdragon

vine, N, O, shaded arroyo, base of cliff, WW530

Solanaceae

Chamaesaracha pallida Averett, False nightshade, N, O, slopes,

ridges, WW238

Chamaesaracha coniodes Britton, Hairy false nightshade, N, O,

slopes, ridges, clay flats, WW413

Lycium berlandieri Dunal, Berlandier wolfberry, N, O, ridges, clay

flats, WW753

Lycium puberulum var. berberidoides (Correll) F. Chiang, Downy

wolfberry, N, O, E-TP, slopes, clay flats, WW392

Lycium texanum Correll, Texas wolfberry, N, R, G2S2, E-TP, slope,

WW 1029

Nicotiana obtusifolia M. Martens & Galeotti, Desert tobacco, N, O,

ubiquitous, WW74

Physalis hederifolia A. Gray, Heartleaf groundcherry, N, O, drainages,

canyons, disturbed sites, WW210

Physalis lobata (Torr.) Raf., Purple groundcherry, N, O, clay flats,

arroyos, disturbed areas, WW62

Solanum elaeagnifolium Cav., Silverleaf nightshade, N, O, clay flats,

disturbed sites, WW381

Weckesser and Terry, Flora of Reed Plateau, Texas

377

Sterculiaceae

Ayenio microphyllo A. Gray, Dense ayenia, N, C, ubiquitous, WW539

Tamaricaceae

Tamarixchinensis Lour., Saltcedar, I, U, arroyos, roadsides, WW906

Ulmaceae

Celtis iguonoeo (Jacq.) Sarg., Spiny hackberry, N, U, canyon, drain¬

ages, WW526

Urticaceae

Parietaria pensylvanica Muh\. exWilld., Pennsylvania pellitory, N, R,

shaded base of cliff, WW160

Verbenaceae

Aloysia gratissima (Gillies & Hook.) Tronc., Beebrush, N, O, arroyos,

drainages, WW620

Aloysia wrightii A. Heller, Oreganillo, N, U, rocky slopes, drainages,

WW703

Bouchea linifolia A. Gray exTorr., Groovestem bouchea, N, O, drain¬

ages, canyons, WW51

Glandularia bipinnatifida (Nutt.) Nutt. var. ciliata (Benth.) Turner,

Dakota mock vervain, N, R, drainage, northern-most extent of

Long Draw, WW280

Glandularia quadrangulata (A. Heller) Umber, Beaked verbena, N,

R, drainage, WW 146

Lantana achyranthifolia Desf., Veinyleaf lantana, N, O, drainages,

arroyos, canyons, WW656

Tetraclea coulteri A. Gray, Stinkweed, N, O, gravelly slopes, WW389

Vitaceae

Cissusincisa Des Moul., Ivy treebine, N, R, arroyo, WW641

Zygophyllaceae

Guaiacum angustifolium Engelm., Guayacan, N, C, ubiquitous,

WW 868

Kallstroemiaperennans B.L.Turner,Turner's desert poppy, N, R, E-TP

clay flats or Del Rio clay outcrops, WW33

Larrea tridentata (Sesse & Moc. ex DC.) Coville, Creosotebush, N, A,

ubiquitous, WW855

ACKNOWLEDGMENTS

By allowing access to their property, the landowners on and near Reed Plateau demonstrated their interest in

understanding and protecting the community in which they live. This study never would have taken place

without their involvement. Many friends, including one particular dog, accompanied us in the held. The first

author wishes to thank J.C. Zech for steadfast support and assistance during the course of the study. Special

thanks go to A.M. Powell for his untiring efforts in maintaining the herbarium at Sul Ross State University and

assistance with plant identification. Dr. Powell’s accumulated botanical knowledge of the Trans-Pecos region is

an invaluable resource that he shared without reservation. Billie L. Turner verified the plant collections and

offered unwavering encouragement for further studies. Without the gracious hospitality of Dr. and Mrs. M.

Terry much of WW’s time on campus would have been impossible. Much of the authors’ discussions took place

over cups of hot tea in the kitchen. NRCS personnel in the Alpine, Texas office provided an introduction to the

Soil Survey website. Jackie Poole offered insight and discussion on Genistidium dumosum. Sul Ross State Uni¬

versity’s Mary Moll Jennings Memorial and Read Memorial Scholarships and the Coors CLASE Scholarship

partially funded this work.

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380

Journal of the Botanical Research Institute of Texas 8(1)

The 2014 Applications for the Delzie Demaree Travel Award

Applications for the 2014 Delzie Demaree Travel Award should include a letter from the applicant telling how

symposium attendance will benefit his/her graduate work and letter of recommendation sent by the major

professor. Please send letters of application to: Dr. Donna M.E. Ware, P.O. Box 8795, Herbarium, Biology De¬

partment, The College of William and Mary, Williamsburg, Virginia 23185-8795, U.S.A. 1-757-221-2799;

Email: ddmware@wm.edu. Applications maybe sent to: Barney Lipscomb, 1700 University Drive, Fort Worth,

Texas 76107-3400, U.S.A. 1-817-332-7432; Email: barney@brit.org. 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 be¬

fore applying. The Systematics Symposium dates for 2014 are 10-11 October 2014 (dates tentative and subject

to change).

The Delzie Demaree Travel Award was established in 1988 honoring Delzie Demaree who attended 35

out of a possible 36 symposia before he died in 1987. Delzie Demaree was a frontier botanist, explorer, discov¬

erer, and teacher. His teaching career as a botanist began in Arkansas at Hendrix College in 1922. He also

taught botany at the University of Arkansas, Navajo Indian School, Yale School of Forestry, Arkansas A&M,

and Arkansas State University at Jonesboro where he retired as professor emeritus in 1953. One of the things he

enjoyed most as a botanist was assisting students with their held botany research.

J.Bot. Res. Inst. Texas 8(1): 380.2014

ADDENDUM TO THE VASCULAR FLORA OF NASH PRAIRIE, TEXAS, U.S.A.

David J. Rosen

Department of Biology

Lee College

Baytown, Texas 77522-0818, U.S.A.

drosen@lee.edu

ABSTRACT

Nash Prairie is a 120 ha remnant of Texas upper coastal prairie with a previously reported native vascular plant flora of 289 species. Miscel¬

laneous collections made over the past seven years are reported here, increasing the known native flora to 301 species.

RESUMEN

Nash Prairie es un resto de 120 ha de la pradera costera superior de Texas con una flora vascular nativa previa de 289 especies. Se citan aqui

diversas colecciones realizadas en los ultimos siete anos, que incrementan la flora nativa conocida a 301 especies.

Nash Prairie, a 120 ha (296.5 acre) remnant of Texas upper coastal prairie in Brazoria County, remains, in my

estimation, the largest and best example of this rare plant community throughout its historic range (Rosen

2007). Since publishing a checklist of vascular plants for this site that included 289 native species, two impor¬

tant events have occurred: 1) the tract (along with Mowotony Prairie; Rosen 2010) has been purchased by the

Nature Conservancy of Texas, ensuring its conservation in perpetuity, and 2) additional collections made from

2007 through 2013 have revealed the following native species not previously reported from the site.

APOCYNACEAE

Asclepias viridiflora Raf.

Voucher specimen: infrequent and widespread in uplands of the prairie, 9 Aug 2011, D.J. Rosen 5403 with W.R. Carr (TEX).

CLUSIACEAE

Hypericum drummondii (Grev. & Hook.) Torr. & A. Gray

Voucher specimen: rare in uplands near the north hayfield road, 27 Oct 2007, D.J. Rosen 4644 (TEX).

LAMIACEAE

Monarda citriodora Cerv. ex Lag.

Voucher specimen: rare in uplands near the north hayfield road, 20 May 2010, D.J. Rosen 5013 (TEX).

Salvia azurea Michx. ex Vahl var. grandiflora Benth. Throughout the upper coastal prairie, I’ve only observed

this species in high quality remnants.

Voucher specimen: rare and seemingly represented by a few individuals restricted to a single pimple mound in the southwest quadrant of

the prairie, 24Jul 2010, D.J. Rosen 5036 (TEX).

ONAGRACEAE

Oenothera lindheimeri (Engelm. & A. Gray) W.L. Wagner & Hoch. A coastal prairie endemic based on my

observations and the distribution indicated by Correll and Johnston (1970).

Voucher specimen: rare and represented by a few individuals in a single location at the south-central boundary of the prairie, 9 Aug 2011,

D.J. Rosen 5404 with W.R. Carr (TEX).

PHYTOLACCACEAE

Phytolacca americana L.

Voucher specimen: locally frequent in a pond recently disturbed by removal of Chinese tallow tree, 18 Oct 2013, D.J. Rosen 6157 (TEX).

J. Bot. Res. Inst. Texas 8(1): 381 - 382.2014

382

Journal of the Botanical Research Institute of Texas 8(1)

POACEAE

Andropogon virginicus L. var. virginicus

Voucher specimen: occasional in low-lying places in the southeast quadrant of the prairie, 18 Oct 2013, D.J. Rosen 6156 (MO, TEX).

Aristida longespica Poir. var. longespica

Voucher specimen: rare on pimple mounds near the center of the prairie, 6 Oct 2012, D.J. Rosen 5902 (TEX).

Aristida oligantha Michx.

Voucher specimen: rare along the north hayfield road, 27 Oct, 2007, D.J. Rosen 4643 (TEX).

Panicum hallii Vasey subsp. filipes (Scribn.) Freckmann & Lelong

Voucher specimen: rare along the north hayfield road,18 Oct 2013, D.J. Rosen 6158 (TEX).

Schedonnardus paniculatus (Nutt.) Branner & Coville. A widespread monotypic prairie and plains species

of North America (Snow 2003).

Voucher specimen: rare along the south hayfield road, 1 Jul 2007, D.J. Rosen 4290 (BRIT, TEX).

PORTULACACEAE

Portulaca umbraticola Kunth subsp. lanceolata J.F. Matthews & Ketron. This southwestern species is de¬

scribed as occurring in disturbed sites and granitic and sandstone outcrops (Matthews 2003); and, in Texas, it

is restricted to prairies, mesquite thickets, and saltmarshes (Correll & Johnston 1970).

Voucher specimen: occasional on the north hayfield road, 18 Oct 2013, D.J. Rosen 6159 (TEX).

The addition of the 12 species reported here increases the known native vascular flora of the Nash Prairie to

301 species. Five new native species increases the already rich grass flora to 64. New families (Phytolaccaceae

and Portulacaceae) and genera ( Monarda, Phytolacca, Schedonnardus, and Portulaca) also increase the numbers

previously reported to 65 and 201 respectively. This report is also offered to emphasize that a floristician’s work

is never done. As suggested by Prater et al. (2004a), our understanding of biodiversity and the soundness of our

conservation decisions are advanced by continued plant collecting. Even in well explored states and ecore-

gions, continued plant collecting should be encouraged and supported (Prather et al. 2004b).

ACKNOWLEDGMENTS

I am grateful to Bill Carr, Barney Lipscomb, and an anonymous reviewer for helpful comments.

REFERENCES

Correll, D.S. & M.C. Johnston. 1970. Manual of the vascular plants of Texas. Texas Research Foundation, Renner, Texas,

U.S.A.

Matthews, J.F. 2003. Portulaca. In: Flora of North America Editorial Committee, eds. Flora of North America north of

Mexico. Oxford University Press, New York, U.S.A., and Oxford, U.K. 4:496-501.

Prather, L.A., O. Alvarez-Fuentes, M.FI. Mayfield, & CJ. Ferguson. 2004a. The decline of plant collecting in the United States:

A threat to the infrastructure of biodiversity studies. Syst. Bot. 29:15-28.

Prather, L.A., O. Alvarez-Fuentes, M.FI. Mayfield, & CJ. Ferguson. 2004b. Implications of the decline in plant collecting for

systematic and floristic research. Syst. Bot. 29:216-220.

Rosen, DJ. 2007. The vascular flora of Nash Prairie: A coastal prairie remnant in Brazoria County, Texas. J. Bot. Res. Inst.

Texas 1:679-692.

Rosen, DJ. 2010. The vascular plants of Mowotony Prairie: A small remnant coastal grassland in Brazoria County, Texas.

J. Bot. Res. Inst.Texas 4:489-495.

Snow, N.W. 2003. Schedonnardus. In: Flora of North America Editorial Committee, eds. Flora of North America north of

Mexico. Oxford University Press, New York, U.S.A., and Oxford, U.K. 25:228-230.

New Titles from BRIT Press

Manual of Montana Vascular Plants

August 2013

Second printing with corrections and errata, 28 May 2014

Montana is the fourth largest state in the United States. It includes portions of the

Northern Great Plains and the Rocky Mountains. The vegetation of Montana

is diverse, due primarily to the size of the state and its great topographic relief

which provide strong variation in environmental factors. Montana has a rela¬

tively large flora for a northern continental region due to being at the intersec¬

tion of the Cordilleran, Great Plains and Boreal floristic provinces. This book is a

comprehensive held guide to the more than 2,500 species of Montana’s vascular

plants. It contains descriptions as well as habitat and distribution information

based on specimens housed at the states two major herbaria. Portraits or illus¬

trations of diagnostic structures are provided for nearly one-third of the species.

Lesica, P, with contributions by M. Lavin and P.F. Stickney. Illustrations by Debbie McNiel, Rich Adams,

Claire Emery. 2012 (2nd printing with corrections/errata). Manual of Montana Vascular Plants. (ISBN-13:

978-1-889878-39-3, pbk.). Botanical Research Institute of Texas Press, 1700 University Dr., Fort Worth, Texas

76107-3400, U.S.A. (Orders: shop.brit.org, orders@brit.org, 817-332-4441 ext. 264, fax 817-332-4112). $55.00,

6.5"x9.5" (pbk), 780 pp., 2000 + maps + 128 plates. $6.00 shipping ($3.00 each additional copy), outside the

U.S.A. contact orders@brit.org, Texas residents add 8.25% ($5.03) to subtotal including postage for each book.

The Ferns and Lycophytes of Texas

February 2014

Texas has a surprising number of native ferns and lycophytes—127 in all, the

most of any state in the continental U.S.A. This is particularly unexpected

given that most people associate ferns and related plants with humid, even

tropical, conditions, just the opposite of much of Texas. This book explains

why and looks at the fascinating world of Texas ferns, ranging from the swamp

forests of far East Texas, to the hidden canyons of the Edwards Plateau, and

even to the high mountain “sky islands” of such places as Big Bend National

Park. Each species has an illustration page with a color photo, a line drawing,

and detailed maps. Be ready to be surprised by this special group of Texas

plants.

George M. Diggs, Jr. and Barney L. Lipscomb. The Ferns and Lycophytes of

Texas. (ISBN-13: 978-1-889878-37-9, flexbound). Botanical Research Institute

of Texas Press, 1700 University Dr., Fort Worth, Texas 76107-3400, U.S.A. (Orders: shop.brit.org, orders@brit.

org, 817-332-4441 ext. 264, fax 817-332-4112). $29.95, 392 pp., color photos, distribution maps, 7"xl0". $4.50

shipping ($3.00 each additional copy), outside the U.S.A. contact orders@brit.org, Texas residents add 8.25%

($2.43) to subtotal including postage for each book.

For more information on these titles and others, please visit us at shop.brit.org.

Press

THE FERNS AND

LYCOPHYTES

Barney L. Lipscomb OF TEXAS