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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Journal of the Botanical Research Institute of Texas is

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

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International Standard Serial No. (ISSN 1934-5259)

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

(summer/fall) as one volume

by the Botanical Research Institute of Texas.

VOLUME 8 NUMBER 2 25 NOV 2014

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Table of Contents

SYSTEMATICS

Recent findings on the gypsum flora of the rim of the Guadalupe Mountains, New Mexico, U.S.A.:

A new species of Nerisyrenia (Brassicaceae), a new state record, and an updated checklist

Patrick J. Alexander, Norman A. Douglas, Helga Ochoterena, Hilda Flores-Olvera,

and Michael J. Moore 383

Lomatium swingerae: a new species of Lomatium (Umbelliferae) from the Joseph Plains,

Idaho, U.S.A.

Richard P. McNeill 395

New geographical and morphological data for Sideroxylon reclinatum subspecies

austrofloridense (Sapotaceae), a taxon endemic to southeastern peninsular Florida, U.S.A.

Paul T. Corogin and Walter S. Judd 403

A morphologically based taxonomic reevaluation of the genus Stipulicida (Caryophyllaceae),

with comments on rank

Derick B. Poindexter, Kateland E. Bennett, and Alan S. Weakley 419

A new hedge-nettle (Stachys: Lamiaceae) from South Carolina, U.S.A.

John B. Nelson and Douglas A. Rayner 431

Agave x madrensis (Asparagaceae), a putative hybrid from the Sierra Madre Oriental, Mexico

Jose A. Villarreal Quintanilla, Abraham Ramirez GAmez, Eduardo Estrada Castillon,

Dino U. GonzAlez Uribe, and Diana Jasso de Rodriguez 441

Dos nuevas especies de Myrcia (Myrtaceae), del Paclhco sur de Costa Rica

Daniel Santamaria Aguilar, Armando Estrada Ch. y Reinaldo Aguilar 449

Taxonomic studies in the Miconieae (Melastomataceae). XII. Revision of Miconia

sect. Miconiastrum, with emphasis on the Miconia bicolor complex

Walter S. Judd, James Dan Skean, Jr., Eldis R. Becquer, and Lucas C. Majure 457

Calathea rubribracteata, a new endemic species of Marantaceae from Colombia

Helen Kennedy 493

Five new Myrtaceae from southeastern Brazil

Marcos Sobral, Marcelo da Costa Souza, Ludovic Jean Charles Kollmann,

Osny Tadeu Aguiar, Andre Luis Casarin Rochelle, and Kelly Antunes 497

Browneopsis puyensis (Leguminosae: Caesalpinioideae: Detarieae), a new species

from Amazonian Ecuador

David A. Neill and Mercedes Asanza 511

A new species of Garnotia (Poaceae) from Kerala, India

C.N. Sunil, V.V. Naveen Kumar, and M.G. Sanilkumar 517

Tripogon malabarica (Poaceae: Chloridoideae: Chlorideae: Tripogoninae),

a new species from Kerala, India

Thoiba Kottekkattu and A.K. Pradeep 523

Wandersong (Rubiaceae), a new genus from the Greater Antilles

David W. Taylor 529

DEVELOPMENT AND STRUCTURE

Variability of vegetative flush colors in Zamia (Cycadales)

Claudia Calonje, Lindy Knowles, Michael Calonje, and Chad Husby 531

Limitations to natural production of Lophophora williamsii (Cactaceae) III. Effects of

repeated harvesting at two-year intervals for six years in a South Texas (U.S.A.) population

Martin Terry, Keeper Trout, Bennie Williams, Teodoso Herrera, and Norma Fowler 541

PALEOBOTANY

Distigouania irregularis (Rhamnaceae) gen. et sp. nov. in Mid-Tertiary amber from the

Dominican Republic

Kenton L. Chambers and George O. Poinar, Jr. 551

Ticodendron palaios sp. nov. (Ticodendraceae), a Mid-Tertiary fossil flower in Dominican amber

Kenton L. Chambers and George O. Poinar, Jr. 559

FLORISTICS, ECOLOGY, AND CONSERVATION

Vegetation patterns in the Mediterranean-desert ecotone of Baja California, Mexico

Sula Elizabeth Vanderplank, Jose Delgadillo, Exequiel Ezcurra, and Lucinda A. McDade 565

Estructura de un zacatal de toboso (Hilaria mutica: Poaceae) asociado a sustrato

Igneo en el noreste de Coahuila, Mexico

Juan A. Encina-Domi'nguez, Jesus Valdes-Reyna y Jose A. Villarreal-Quintanilla 583

Jarilla chocola (Caricaceae), un nuevo genero y especie para la flora de El Salvador

Frank Sullyvan Cardoza Ruiz, Jose Ledis Linares y Ana Eugenia Aguilar Grijalva 595

Redescubrimiento y nuevos registros de Cuscatlania vulcanicola (Nyctaginaceae)

para El Salvador

Jose Ledis Linares, Frank Sullyvan Cardoza Ruiz y Patricia HernAndez-Ledesma 603

Vascular plant flora of the alpine zone in the Southern Rocky Mountains, U.S.A.

James F. Fowler, B.E. Nelson, and Ronald L. Hartman 611

The first naturalized occurrence of the Cannaceae family in the Arkansas (U.S.A.) flora,

with additional new and noteworthy angiosperm records for the state

Brett E. Serviss, James H. Peck, and Tiffany A. Graves 637

American mistletoe (Phoradendron leucarpum ssp. leucarpum, Viscaceae) occurrence in host

trees within the city of Cairo, Alexander County, Illinois, and its incidence in Illinois, U.S.A.

Ralph L. Thompson 641

Tetragonia tetragonioides (Aizoaceae) discovered in Louisiana, U.S.A.

Sairah M. Javed and Lowell E. Urbatsch 661

Amsonia ludoviciana (Apocynaceae) new to the flora of Texas, U.S.A.

Chris Doffitt, Charles Allen, Patricia Lewis, and David Lewis 663

The vascular flora and plant communities of Candy Abshier Wildlife Management Area,

Chambers County, Texas, U.S.A.

Jason R. Singhurst, Amos Cooper, David J. Rosen, and Walter C. Holmes 665

In Memoriam

Landon E. McKinney (1949-2014)

Ronald L. Jones and Ralph L. Thompson

Book Reviews, Notices, and Announcements 394,402,418,448,456,492, 522, 528, 558, 582,

602, 610, 640, 676

Reviewers for Volume 8 (2014)—679

Index to Volume 8 (2014)—680

Titles of Articles with Authors—680

Authors—682

Botanical Names and Subjects—683

New Names and Combinations—685

INDEX to new names and new combinations in J. Bot. Res. Inst Texas 8(2), 2014

Agave x madrensis Villarreal, A. Ramirez, & A.E. Estrada, hybrid nov.—443

Browneopsis puyensis D.A. Neill & Asanza, sp. nov.—511

Calathea rubribracteata H. Kenn., sp. nov.—493

Calyptranthes solitaria Sobral, Aguiar, & Antunes, sp. nov.—498

Calyptranthes ubatubana Sobral & Rochelle, sp. nov.—500

Distigouania K.L. Chambers & Poinar, gen. nov.—552

Distigouania irregularis K.L. Chambers & Poinar, sp. nov.—552

Eugenia dipetala Sobral & Kollmann, sp. nov.—502

Garnotia variyamensis Sunil, Naveen Kumar, & Sanilkumar, sp. nov.—517

Lomatium swingerae R.P. McNeill, sp. nov.—395

Miconia barbata (Borhidi) Judd, Becquer, & Majure, comb. nov.—478

Miconia bicolor var. patenti-setosa (Borhidi) Judd, Becquer, & Majure, comb. nov.—474

Miconia cajalbanensisjudd, Becquer, & Majure, nom. nov.—483

Miconia cristalensis (Borhidi) Judd, Becquer, & Majure, comb. nov.—485

Miconia guajaibonensis Judd, Becquer, & Majure, nom. nov.—474

Miconia impressa (Urb.) Judd, Becquer, & Majure, comb. nov.—476

Miconia karsticola Judd, Becquer, Skean, & Majure, nom. nov.—463

Miconia maestrensis Judd, Becquer, & Majure, nom. nov.—487

Miconia sect. Miconiastrum (Bonpl. ex Naudin) Judd, Becquer, & Majure, comb. nov.—461

Myrcia cacuminis Kollmann & Sobral, sp. nov.—504

Myrcia paulii-jonesii Aguilar, D. Santam., & A. Estrada, sp. nov.—449

Myrcia riverae A. Estrada, D. Santam., & Aguilar, sp. nov.—452

Nerisyrenia hypercorax P.J. Alexander & M.J. Moore, sp. nov.—384

Plinia ambivalens M. Souza & Sobral, sp. nov.—507

Stachys carolinianaJ.B. Nelson & D.A. Rayner, sp. nov.—431

Stipulicida lacerata (C.W. James) D.B. Poind., K.E. Bennett, & Weakley, comb, etstat. nov.—426

Ticodendron palaios K.L. Chambers & Poinar, sp. nov.—560

Tripogon malabarica Thoiba & Pradeep, sp. nov.—523

Wandersong D.W. Taylor, gen. nov.—530

Wandersong exserta (DC.) D.W. Taylor, comb. nov.—530

Wandersong seminervis (Urb. & Ekman) D.W. Taylor, comb. nov.—530

677

RECENT FINDINGS ON THE GYPSUM FLORA OF THE RIM OF THE

GUADALUPE MOUNTAINS, NEW MEXICO, U.S.A.: A NEW

SPECIES OF NERISYRENIA (BRASSICACEAE), A NEW

STATE RECORD, AND AN UPDATED CHECKLIST

Patrick J. Alexander Norman A. Douglas Helga Ochoterena

Department of Biology, MSC 3AF

New Mexico State University

Las Cruces, New Mexico 88003, U.S.A.

paalexan@polyploid.net

Department of Biology

Oberlin College

Oberlin, Ohio 44074, U.S.A.

ndouglas@oberlin.edu

Departamento de Botanica

Instituto de Biolog fa

Universidad Nacional Autonoma de Mexico

Apartado Postal 70-367

Mexico, D.F. 04510, MEXICO

helga@ib.unam.mx

Hilda Flores-Olvera

Michael J. Moore

Departamento de Botanica

Instituto de Biolog fa

Universidad Nacional Autonoma de Mexico

Apartado Postal 70-367, Mexico, D.F. 04510, MEXICO

mahilda@ib. unam. mx

Department of Biology

Oberlin College

Oberlin, Ohio 44074, U.S.A.

mmoore@oberlin.edu

ABSTRACT

Exposures of Yeso Formation gypsum along the western escarpment (The Rim) of the Guadalupe Mountains in southeastern New Mexico

were first explored botanically in 1996, which revealed the existence of two gypsophilic taxa, Anulocaulis leiosolenus var. howardii and Men -

tzelia humilis var. guadalupensis , both of which are only known from that area. Fieldwork by the authors has revealed another gypsophile

restricted to The Rim, Nerisyrenia hypercorax, which is here described. The new species is similar to N. gypsophila and N. mexicana , from

which it differs in having shorter, crispate fruits and smaller floral parts. We also report a new state record of Paronychia wilkinsonii in New

Mexico and make additional observations regarding the gypsum flora of The Rim.

RESUMEN

Las exposiciones de yeso de la “Formacion Yeso” en la escarpa occidental de las Montanas de Guadalupe, conocida como “The Rim,” en el

sureste de Nuevo Mexico fueron inicialmente exploradas botanicamente en 1996, cuando se descubrio la existencia de dos taxones gipsofi-

los, Anulocaulis leiosolenus var. howardii y Mentzelia humilis var. guadalupensis , conocidos solamente de esa area. El trabajo de campo real-

izado recientemente por los autores, permitio descubrir otra planta gipsofila endemica restringida a dicha formacion y localidad, Nerisyre¬

nia hypercorax, que se describe aqui. La nueva especie es similar a N. gypsophila y N . mexicana pero difiere de ellas en los frutos mas cortos,

crispados y las partes florales mas pequenas. Tambien registramos Paronychia wilkinsonii por primera vez para Nuevo Mexico y proporcio-

namos observaciones adicionales sobre la flora de la region.

INTRODUCTION

Gypsum exposures are distributed in an island-like fashion throughout the Chihuahuan Desert region and

host a diverse array of over 200 gypsophilic (i.e. occurring only on gypsum) plant species in over 35 families

(Powell & Turner 1977; Moore & Jansen 2007). In the US portion of the Chihuahuan Desert, gypsum deposits

are common in much of central and southern New Mexico and adjacent west Texas (Weber & Kottlowski

1959; Anderson & Dean 1995) and host a number of gypsophilic taxa that vary in distribution across the re¬

gion. The dominant gypsophilic taxa in New Mexico and Texas typically include Sporobolus nealleyi (Poaceae),

Tiquilia hispidissima (Boraginaceae), Dicranocarpus parviflorus (Asteraceae), Sartwellia flaveriae (Asteraceae),

Oenothera hartwegii subsp. filifolia (Onagraceae), Nerisyrenia linearifolia (Brassicaceae), and Acleisanthes

lanceolata (Nyctaginaceae). In addition, a number of narrowly distributed gypsophiles are found in New

Mexico and west Texas. In northern New Mexico, for example, exposures of Todilto Formation gypsum are

J. Bot. Res. Inst. Texas 8(2): 383 - 393.2014

384

Journal of the Botanical Research Institute of Texas 8(2)

home to the narrow endemics Abronia bigelovii Heimerl (Nyctaginaceae), Mentzdia todiltoensis N.D. Atwood &

S.L. Welsh (Loasaceae), Townsendia gypsophila Lowrey & P.J. Knight (Asteraceae), and Phacelia sivinskii N.D.

Atwood, P.J. Knight & Lowrey (Boraginaceae).

In southern New Mexico and adjacent west Texas a number of locally endemic gypsophiles can be found

near the Guadalupe Mountains (New Mexico Rare Plant Technical Council 1999-2012). Although the moun¬

tain range itself is composed primarily of Permian limestone, gypsum is present on both the western and

eastern sides of the Guadalupe Mountains as part of the Castile, Seven Rivers, and Yeso formations, and as

Quaternary lacustrine deposits that are ultimately derived from the Yeso and Castile formations (Boyd 1958;

King 1948; Scholle 2003). Four narrowly endemic gypsophiles are known only from the eastern side of the

Guadalupe Mountains. Two of these species are found only on the Castile Formation [Astragalus gypsodes

Barneby (Fabaceae) and Linum allredii Sivinski & M.O. Howard (Linaceae)], whereas the other two are found

on both the Castile and Seven Rivers formations [Amsonia tharpii Woodson (Apocynaceae) and Eriogonum

gypsophilum Wooton & Standi. (Polygonaceae)]. The narrowly endemic Lepidospartum burgessii B.L. Turner

(Asteraceae) is known only from the southwestern side of the Guadalupe Mountains, on the east side of Crow

Flats and Salt Basin in Quaternary lacustrine gypsum deposits. The western escarpment of the Guadalupe

Mountains, known as The Rim, is composed mostly of San Andres Formation limestone, but gypsum of the

Yeso Formation outcrops frequently near the base of The Rim, occasionally forming extensive exposures such

as those near the mouth of Pup Canyon (Figs. 1, 2). These gypsum outcrops were unexplored by botanists

prior to 1996-97, when the Pup Canyon gypsum was first systematically collected by M. Howard, R. Spellenberg,

and T. Wootten (Spellenberg & Wootten 1999). These explorations led to the discovery and description of two

gypsophiles, Anulocaulis leiosolenus var. howardii (Nyctaginaceae) and Mentzelia humilis var. guadalupensis

(Loasaceae).

The gypsum at the base of The Rim is remote, and much of it is difficult to access; consequently, the flora

of this area remains relatively poorly known. In August and September 2013 the authors visited the gypsum

exposures along The Rim of the Guadalupe Mountains in support of an ongoing project to understand the

evolutionary history of the Chihuahuan Desert gypsophilic flora. This fieldwork revealed the existence of a

new gypsophilic taxon that appears to be restricted to The Rim, Nerisyrenia hypercorax, which is here de¬

scribed. We also report new floristic observations for this region, including a state record of the rare Paronychia

wilkinsonii S. Watson and an updated checklist of plants currently known from the gypsum of The Rim

(Appendix 1).

A NEW SPECIES OF NERISYRENIA

Nerisyrenia Greene is a small genus, including 7 (Rollins 1993), 8 (Al-Shehbaz 2012), 9 (Bacon 1978), or 11

(Turner 1993) species. The most recent monograph of the genus is that of Bacon (1978). Nomenclature of Neri¬

syrenia and characteristics of previously-published species below are based on this work and the subsequent

paper by Turner (1993). Nerisyrenia is restricted to the Chihuahuan Desert region and has a remarkable affin¬

ity for gypsum. All the species are gypsophiles except the widespread and variable Nerisyrenia camporum (A.

Gray) Greene. These gypsophilic taxa are largely allopatric with respect to one another, and collectively they

occupy gypsum exposures from central New Mexico to northern San Luis Potosl (Bacon 1978). In the United

States, only two taxa were previously known: N. camporum and N. linearifolia. The discovery of N. hypercorax

adds yet another gypsophilic species to the genus and to the flora of New Mexico. The visit to Pup Canyon by

the authors on 20 Aug 2013 coincided with the filming of Episode 5 of the educational video series Plants Are

Cool Too!, which can be viewed at http://www.youtube.com/watch?v=al6mBFTkrks.

Nerisyrenia hypercorax P.J. Alexander & M.J. Moore, sp. nov. (Figs. 3,4) Type: u.S.A. New Mexico. Otero Co.: W-facing

gypsum slopes of The Rim of the Guadalupe Mountains, just N of Pup Canyon, 32.37803°N, 105.07308°W, 1339 m; scattered in

diverse gypsophilic community on relatively barren gypsum, with no single dominant species; subshrubs to 25 cm tall, petals

white, not senescing lavender, leaves somewhat succulent, never linear, 20 Aug 2013 (fl, fr), M.J. Moore et al 2272 (holotype: NMC;

isotypes: MEXU, OC, TEX/LL, UNM).

Alexander et al., A new species of Nerisyrenia from New Mexico

385

Fig. 1 . Known distribution of N. hypercorax ("+"), closest known population of N. linearifolia ("x"), and approximate distribution ofYeso Formation

gypsum outcrops along the Guadalupe Mountains rim (shown in white with dark outline).

Similar to N. gypsophila J.D. Bacon and N. mexicana (J.D. Bacon) B.L. Turner but differing in having shorter (usually 5-12 mm, vs. > 15 mm),

crispate, incurved fruits and smaller floral parts; crispate fruits are apparently unique in the genus.

Suffrutescent perennials or subshrubs, not rhizomatous; individuals more or less hemispherical, 1-2.5

(-3.5) dm tall. Moderately pubescent throughout, trichomes mostly appressed, sessile or short-stalked (stalks

to 0.05 mm), predominately dendritically 5-branched (rays to 0.25 mm), some trichomes 3-rayed to dendritically

386

Journal of the Botanical Research Institute of Texas 8(2)

• \v *..*»'

Fig. 2. Habitat of N. hypercorax : A. at the type locality; B. at the site of Alexander 1330.

7-branched. Stems branched throughout, 3-6 from a branched, woody caudex, older woody stems to 1 cm

in diameter, herbaceous stems to 1 mm diameter. Cauline leaves succulent, strongly overlapping, 3-6 times

longer than wide, the larger (18-)22-40(-48) mm long, (4—)6—12(—14) mm wide, oblanceolate to spatulate,

attenuate at the base and obtuse to acute at the apex, margins entire to weakly sinuate or, rarely, obscurely

sinuate-dentate. Flowers with sepals (3.0-)4.0-6.0 mm long, 1.0-1.5 mm wide, broadly lanceolate in outline.

Petals white, not fading purple on senescence, (rarely fading very pale lavender), 7-9 mm long, 3.5-4.5 mm

wide, obovate to spatulate in outline, blade margins entire, dilate and denticulate at base. Stamens weakly

tetradynamous, 2.5-5.5 mm long, anthers medihxed, straight and 1.5-1.8 mm long at anthesis, curling with

Alexander et al., A new species of Nerisyrenia from New Mexico

387

Fig. 3. Line drawing of N. hypercorax, based on P.J. Alexander 1324 and M.J. Moore et al. 2272. A. Plant habit. B. Mature fruit and seeds. C. Apex of a

cauline leaf and enlarged view of leaf trichome forms. D. Flowers and immature fruit. Illustration by Avery Liell-Kok.

388

Journal of the Botanical Research Institute of Texas 8(2)

Fig. 4. Nerisyrenia hypercorax. A. Flower at the type locality. B. Fruit at the site of P.J. Alexander 1324. C. Plant habit at the type locality.

age, filaments 2.0-4.5 mm long. Ovary tomentose, 2-4 mm long, 0.8-1.5 mm wide, style glabrous, 1.3-2.2 mm

long, stigma deltoid-sagittate, 0.5-0.8 mm long, somewhat decurrent on the style. Infructescences compact,

(1.5-)2.0-4.5(-6.5) cm long, with 5-15(-20) fruits, middle internodes 1-5 mm long, pedicels straight, ascending

to, infrequently divaricate, (3 —)4—9(— 13) mm long. Siliques obcompressed (angustiseptate), oblong, rounded-

Alexander et al., A new species of Nerisyrenia from New Mexico

389

truncate at apex, not tapered to style, incurved, crispate (tortuose), 5—12(—16) mm long, 2.2-3.0 mm wide, in¬

ner surfaces of the valves glabrous; replum 0.8-1 mm wide. Ovules 20-40 per silique, seeds broadly elliptic to

broadly ovate, 0.8-1 mm long, 0.6-0.7 mm wide, somewhat flattened, about half as thick as wide, yellow-orange.

Etymology. —The specific epithet refers to the distribution of the species, above Crow Flats.

Phenology. —Flowering specimens have been collected from late July to early September, while specimens

with mature fruits have been collected in August and early September.

Geography and habitat.—Nerisyrenia hypercorax occurs on gypsum of the Yeso Formation on the west

side of the Guadalupe Mountains, at the mouth of Pup Canyon and north-northwest for ca. 20 km, between

1300 and 1600 m. Nerisyrenia hypercorax has been found at every site within this band that has been visited by

botanists.

Distinction from other Nerisyrenia. —In the treatment by Bacon (1978), N. hypercorax keys to Nerisyrenia

gypsophila J.D. Bacon, a gypsophile found throughout east central Chihuahua. However, it is perhaps morpho¬

logically closer to Nerisyrenia mexicana (J.D. Bacon) B.L. Turner, a gypsophile of southeastern Coahuila,

southern Nuevo Teon, and adjacent San Tuis Potosl and Tamaulipas, which has similarly short pedicels and

styles. Nevertheless, N. hypercorax differs from these two species by its shorter fruits and smaller floral parts.

For example, it is distinguished from both species by its shorter siliques (5-125-12 mm vs. 17-30 for N. gyp¬

sophila and 15-27 for N. mexicana), shorter anthers (1.5-1.8 mm vs. 2.4-3.4 and 2.5-3.5, respectively), smaller

petals (7-9 mm long, 3.5-4.5 mm wide, vs. 9.5-11.5 x 4.5-8.5 in N. gypsophila, and 8.5-13 x 5.0-8.5 in N.

mexicana), usually shorter filaments (2.5-4.5 mm vs. 4-6 in both N. gypsophila and N. mexicana), and usually

shorter infructescences (2-4.5 cm vs. 4-30 and 6-19, respectively). From N. gypsophila it is further distin¬

guished by its shorter sepals (4-6 mm vs. 6.5-8.5), usually shorter pedicels (5-9 mm vs. 8-12), shorter styles

(1.3-2.2 mm vs. 2.7-4.0), and incurved rather than straight siliques. The crispate fruits of N. hypercorax, pres¬

ent in all individuals observed, are apparently unique in the genus and most readily distinguish this species

from other Nerisyrenia.

Previously, only two species of Nerisyrenia were known from New Mexico: the gypsophilic N. linearifolia

and the morphologically and edaphically variable N. camporum. Nerisyrenia linearifolia is found on gypsum

throughout the southern two-thirds of the state and in adjacent western Texas, while N. camporum is the most

widespread species in the genus, common along the Rio Grande Valley and the southern third of the New

Mexico as well as in southwestern Texas and the Mexican states of Chihuahua, Coahuila, Durango, Nuevo

Teon, Tamaulipas, and Zacatecas. While N. hypercorax shares the broader leaves of N. camporum, it differs from

this species in a number of respects, principally in its more suffruticose habit (vs. herbaceous to weakly suf-

frutescent perennial), entire leaf margins (vs. typically dentate leaves), relatively compact inflorescences that

only slightly exceed the leaves (vs. elongate inflorescences that greatly exceed the leaves), and its petals, which

do not turn pale lavender or purple upon senescence as in N. camporum. Indeed, N. hypercorax shares all of

these character states with its gypsum-loving cousin N. linearifolia, although the broad, shorter leaves, gener¬

ally smaller flower parts, and shorter, crispate fruits of the former serve to easily distinguish it from the latter.

A key to Nerisyrenia in New Mexico is provided below.

KEY TO THE NERISYRENIA SPECIES OF NEW MEXICO

1 . Leaves linear, all less than 5 mm wide_ N. linearifolia

1. Leaves oblanceolate, spatulate, or obovate, the larger more than 5 mm wide

2. Fruits crispate, less than 15 mm long; infructescences less than 7 cm long; petals less than 5 mm wide, remaining

white_ N. hypercorax

2. Fruits not crispate, more than 15 mm long; infructescences more than 7 cm long; petals more than 5 mm wide, fading

lavender_ N. camporum

Although N. camporum and N. linearifolia can be found growing together in New Mexico, so far as is known

neither species co-occurs with N. hypercorax. The authors have found N. hypercorax at five sites but did not

observe any other Nerisyrenia at these sites. There are only two previous Nerisyrenia specimens from this band

of gypsum in regional herbaria, both N. hypercorax.

390

Journal of the Botanical Research Institute of Texas 8(2)

Paratypes. — U.S.A. New Mexico. Chaves Co.: Lincoln National Forest, W slope of the Guadalupe Mountains ca. 1.5 mi N of Otero county

line, 32.53752°N 105.14758°W, elev. 1579 m, limestone interbedded with gypsum, 23 Jul 2012 (fl), K.D. Heil 34399 (SJNM). Otero Co.: W

base of the Guadalupe Mountains E of Pinon Creek, 1.9 mi S of the Chaves County line, 1.7 miNE of Tanner Ranch, 32.4936°N 105.1399°W

(WGS84), elev. 1540 m, along small incised ravines in a SW-facing, reddish gypsum outcrop, 2 Sep 2013 (fl, fr), P.J. Alexander 1324 (DUKE,

MEXU, MO, NMC, OC); W base of the Guadalupe Mountains E of Pinon Creek, 2.9 mi S of the Chaves County line, 1.2 mi ENE of Tanner

Ranch, 32.4786°N 105.1369°W (WGS84), elev. 1550 m, along a narrow arroyo deeply incised in gypsum, 1 Sep 2013 (fl, fr), P.J. Alexander

1330 (DUKE, MEXU, MO, NMC, OC); 49 km NNE of Dell City, Texas, W base of Guadalupe Mts., N of mouth of Pup Canyon, 32.37917°N

105.06650°W, elev. 1460 m, gypseous slopes on upper bajada, W-facing, 5 Aug 1997 (fr), R.W. Spellenberg 12442 (NMC); west base of the

Guadalupe Mountains, 2.0 mi E of Johnson Tanks and 2.6 mi WNW of the mouth of Pup Canyon, 32.38921°N 105.09743°W (WGS84), elev.

1330 m, small outcrop of gypseous clay near the mouth of a small limestone canyon, 28 Aug 2014 (fl, fr), P.J. Alexander & M.O. Howard 1463

(MO, NMC, OC); west base of the Guadalupe Mountains, 2.2 mi E of Johnson Tanks and 2.6 mi NW of the mouth of Pup Canyon,

32.39478°N 105.09323°W (WGS84), elev. 1360 m, gypseous clay cutbanks in a small limestone canyon, immediately below large expanses

of open gypsum, 28 Aug 2014 (fl, fr), P.J. Alexander & M.O. Howard 1464 (NMC, OC, UNM).

FLORA OF THE YESO FORMATION GYPSUM ALONG THE RIM

Spellenberg and Wootten (1999) compiled a list of 59 vascular plant taxa occurring on Yeso Formation gyp¬

sum at The Rim from their fieldwork on the north side of Pup Canyon and a site northeast of Tanner Ranch.

Fieldwork by the authors and additional visits to the area by K. Heil and associates at San Juan College and R.

Worthington of the University of Texas at El Paso have increased our knowledge of the gypsum flora of The

Rim from Pup Canyon and to the north-northwest for ca. 20 km. This expanded list includes 74 vascular plant

taxa in 27 plant families and is provided in Appendix 1. The endemics Mentzdia humilis var. guadalupensis and

Nerisyrenia hypercorax are found throughout this band of gypsum, while Anulocaulis leiosolenus var. howardii

has been found only on gypsum in the immediate vicinity of Pup Canyon.

There are several additional small outcrops of Yeso Formation gypsum southeast of Pup Canyon on the

east side of Big Dog Canyon that have not previously been botanically explored. The first author visited one of

these sites in September 2013 to determine if N. hypercorax is present. None of the narrow endemics of The Rim

were found at this site. Instead of N. hypercorax and M. humilis var. guadalupensis, the more widespread N.

linearifolia and M. humilis var. humilis were found at Big Dog Canyon, while no Anulocaulis was seen. A list of 37

vascular plant taxa in 20 families observed at the site is provided in Appendix 2.

PARONYCHIA WILKINSONII , A NEW STATE RECORD

On limestone adjacent to Yeso Formation gypsum, the first author found Paronychia wilkinsonii, here first re¬

ported from New Mexico: U.S.A. New Mexico. Otero Co.: west base of the Guadalupe Mountains east of Pinon

Creek, 2.9 miles south of the Chaves County line, 0.9 miles east-northeast of Tanner Ranch, 32.4786°N

105.1421°W (WGS84), elev. 1490 m, gentle west slope, limestone cobble, 2 Sep 2013 (fl, fr), P.J. Alexander 1329

(NMC, OC, RM). This is a rare species previously known only from outcrops of novaculite (a form of chert

comprised primarily of microcrystalline quartz) in the Marathon Basin of western Texas and a few scattered

sites in the Mexican states of Chihuahua and Coahuila (Poole et al. 2007). This population in New Mexico is

ca. 300 km disjunct from the nearest known populations near Marathon.

DISCUSSION

The presence of at least three locally endemic gypsophiles at Pup Canyon and neighboring gypsum sites along

The Rim implies that little gene flow has occurred between gypsum at The Rim and other gypsum exposures,

allowing allopatric speciation, and it further raises the possibility that conditions supporting the long-term

persistence of gypsophiles may have characterized The Rim for all or much of the Pleistocene. During full-

glacial periods of the Pleistocene, New Mexico experienced a significantly cooler and wetter climate, resulting

in the replacement of “typical” Chihuahuan Desert vegetation with grassland and savanna that is similar to

that seen in central New Mexico today (Van Devender 1990, Elias & Van Devender 1992). The southerly loca¬

tion of The Rim in New Mexico, its relatively low elevation, and its west-facing aspect may have resulted in a

warmer, drier microclimate during full-glacial periods, helping to maintain the semi-arid character of the site

Alexander et al., A new species of Nerisyrenia from New Mexico

391

over time. Even today the gypsum exposures along The Rim are relatively thinly vegetated compared to most

other gypsum deposits in the Chihuahuan Desert (Fig. 2), which likely results in part from this warmer, drier

microclimate. Moreover, the gypsum substrate itself may have helped promote community stability for gypso-

philic taxa during the wetter periods of the Pleistocene by reducing or preventing the growth of non-gypso-

philic plant taxa, which often have great difficulty establishing and persisting on gypsum (Damschen et al.

2012; Escudero et al. 2014; Moore et al. 2014). Ongoing phylogenetic and phylogeographic studies of the Chi¬

huahuan Desert gypsum flora in the lab of Michael Moore at Oberlin College will help to test this hypothesis

and will shed light on the evolutionary relationships of the unique gypsum flora at Pup Canyon.

APPENDIX 1

The following list includes all species known to occur on gypsum of The Rim from Pup Canyon northward. All species listed by Spellen-

berg and Wootten (1999) are included. The list is supplemented by observations made by the authors and specimens collected by R.

Worthington, K. Heil, S. O'Kane, D. Schleser, and L. Urban. Species are followed by all known herbarium specimens and deposition of

specimens is indicated by herbarium codes following Index Herbariorum (Thiers 1997-2013). If no specimens are listed, the species has

been observed in the field, either by Spellenberg and Wootten (1999) or by the present authors, but no voucher has been collected. Unfor¬

tunately, the remoteness of the area and logistic constraints has prevented complete collection of the gypsum flora of The Rim. We have

opted to provide as complete an account as possible, despite lack of vouchers for some species. Nomenclature below the rank of family

follows Allred and Ivey (2012), families follow APG III (2009), and most herbarium specimen data are available online (SEINet 2009-2013).

Amaranthaceae: Atriplex canescens (Pursh) Nutt.; Tidestromia

suffruticosa (Torr.) Standi, var. suffruticosa: Spellenberg & Woot¬

ten 12481 (NMC), Moore etal. 2278 (OC, NMC, TEX/LL, MEXU).

Anacardiaceae: Rhus microphylla Engelm.

Apocynaceae: Amsonia longifloro Torr. var. solpignotho (Woodson)

McLaughlin: Spellenberg etol. 12434 (NMC, NY, UNM), Spellen¬

berg etal. 12441 (NMC, NY), Spellenberg & Wootten 72474 (NMC,

NY), Spellenberg & Wootten 12501 (NMC, NY); Asdepias macrotis

Torr.: Spellenberg & Wootten 12507 (NMC, NY), Heil34405 (SJNM).

Asparagaceae: Dasylirion leiophyllum Engelm. ex Trel.; Nolina

texana S. Watson: Heil 34400 (SJNM); Yucca elata (Engelm.)

Engelm.; Yucca treculeana Carriere.

Asteraceae: Artemisia ludoviciana Nutt.; Brickellia laciniata A. Gray;

Gaillardia multiceps Greene; Gutierrezia microcephala (DC.) A.

Gray; Haploesthes greggii A. Gray var. texana (J.M. Coult.) I.M.

Johnst.: Spellenberg et al. 12437 (NMC), Heil 34397 (SJNM),

Moore etal. 2274 (OC, NMC,TEX/LL, MEXU); Parthenium inca-

num Kunth: Heil34394 (SJNM); Porophyllumscoparium A. Gray:

Spellenberg etal. 12436 (NMC, UC), Worthington 30289 (UNM,

UTEP); Sartwellia flaveriae A. Gray: Worthington 30282 (UNM,

UTEP); Sidneya tenuifolia (A. Gray) E.E. Schill. & Panero; Thele-

sperma megapotamicum (Spreng.) Kuntze: Moore et al. 2279

(OC, NMC,TEX/LL, MEXU); Thymophylla acerosa (DC.) Strother:

Spellenberg & Wootten 12484 (NMC); Thymophyllapentachaeta

(DC.) Small var. belendinium (DC.) Strother: Spellenberg &

Wootten 12475 (NMC, NY), Spellenberg & Wootten 12483 (BRIT);

Xanthisma spinulosum (Pursh) D.R. Morgan & R.L. Hartm. var.

chihuahuanum (B.L. Turner & R.L. Hartm.) D.R. Morgan & R.L.

Hartm.: Spellenberg & Wootten 12505 (NMC).

Boraginaceae: Nama carnosum C.L. Hitchc.: Spellenberg & Woot¬

ten 12503 (NMC), Alexander 1325 (NMC, OC); Tiquilia greggii

(Torr. & A. Gray) A.T. Richardson: Moore etal. 2281 (OC, NMC,

TEX/LL, MEXU); Tiquilia hispidissima (Torr.) A.T. Richardson:

Spellenberg & Wootten 12502 (NMC), Moore etal. 2275 (MEXU,

NMC, OC, TEX/LL).

Brassicaceae: Nerisyrenia hypercorax PJ. Alexander & MJ. Moore:

Spellenberg et al. 12442 (NMC), Heil 34399 (SJNM), Moore et al.

2272 (MEXU, NMC, OC, TEX/LL), Alexander 1324 (DUKE, MEXU,

MO, NMC, OC), Alexander 1330 (DUKE, MEXU, MO, NMC, OC).

Cactaceae: Coryphantha tuberculosa (Engelm.) A. Berger: Spellen¬

berg & Wootten 12482 (NMC); Cylindropuntia imbricata (Haw.)

F.M. Knuth; Echinocactus horizonthalonius Lemaire; Echino-

cereus dasyacanthus Engelm.: Spellenberg & Wootten 12486

(NMC); Opuntia macrocentra Engelm.: Spellenberg & Wootten

12487 (NMC), Spellenberg & Wootten 12488 (NMC); Opuntia

phaeacantha Engelm.

Ephedraceae: Ephedra aspera S. Watson: Spellenberg & Wootten

12473 (NMC).

Euphorbiaceae: Chamaesycefendleri (Torr. & A. Gray) Small: Spellen¬

berg & Wootten 12458 (NMC), Spellenberg & Wootten 12471 (NY);

Chamaesyceserrula (Engelm.) Wooton & Standi.: Alexander 1332

(NMC); Croton dioicus Cav.

Fabaceae: Daleaformosa Torr.; Dermatophyllum guadalupense (B.L.

Turner & A.M. Powell) B.L. Turner: Urban 5072 (NMC); Vachellia

vernicosa (Britton & Rose) Seigler & Ebinger.

Fouquieriaceae: Fouquieria splendens Engelm.

Krameriaceae: Krameria erecta Schult.: Spellenberg & Wootten

12478 (NMC).

Lamiaceae: Hedeoma nana (Torr.) Briq.: Spellenberg & Wootten

12479 (NMC), Heil34396 (SJNM).

Linaceae: Linum vernale Wooton: Spellenberg & Wootten 12480

(NMC).

Loasaceae: Cevalliasinuata Lag.: Worthington 30285 (UNM, UTEP);

Mentzelia humilis (A. Gray) J. Dari. var. guadalupensis Spellenb.:

Spellenberg & Wootten 12440 (NMC), Spellenberg & Wootten

12455 (NMC, NY, TEX, UNM), Spellenberg & Wootten 12500

(NMC, RM), Worthington 30287 (UCR, UTEP), Heil & O'Kane 33495

(SJNM), Heil & Schleser 34725 (SJNM), Moore et al. 2273 (MEXU,

NMC, OC, TEX/LL).

Malvaceae: Sphaeralcea coccinea (Nutt.) Rydb.: Spellenberg &

Wootten 12476 (NMC).

Nyctaginaceae: Acleisanthes lanceolata (Wooton) R.A. Levin:

Spellenberg & Wootten 12452 (NMC), Spellenberg & Wootten

12499 (NMC), Heil & Schleser 34726 (SJNM), Moore et al. 2277

(MEXU, NMC, OC, TEX/LL); Allionia incarnata L. var. incarnata;

Anulocaulis leiosolenus (Torr.) Standi, var. howardii Spellenb. &

Wootten: Wootten & Howards.n. (NMC), Spellenberg etal. 12433

(NMC, NY, UNM), Spellenberg et al. 12435 (NMC), Spellenberg

et al. 12438 (NMC), Worthington 30288 (UCR, UTEP), Heil &

O'Kane 33493 (SJNM), Moore etal. 2270 (MEXU, NMC, OC,TEX/

LL); Cyphomeris gypsophiloides (M. Martens & Galeotti) Standi.:

Heil & Schleser 34722 (SJNM); Mirabilis linearis (Pursh) Heimerl:

Spellenberg & Wootten 12506 (NMC).

Oleaceae: Menodora scabra A. Gray: Heil34403 (SJNM).

392

Journal of the Botanical Research Institute of Texas 8(2)

Onagraceae: Oenothera hartwegii Benth. subsp. filifolia (Eastw.) W.L.

Wagner & Hoch: Spellenberg & Wootten 12472 (NMC), Heil34401

(SJNM), Moore etal. 2276 (OC, NMC, US); Oenothera suffrutescens

(Seringe) W.L. Wagner & Hoch.

Poaceae: Achnatherum curvifolium (Swallen) Barkworth: Spellen-

berg & Wootten 12477 (NMC, NY); Aristida purpurea Nutt. var.

nealleyi (Vasey) Allred: Spellenberg & Wootten 12453 (NMC);

Aristida pansa Wooton & Standi, var. pansa: Spellenberg &

Wootten 12456 (NMC); Bothriochloa laguroides (DC.) Herter

subsp. torreyana (Steud.) Allred & Gould: Spellenberg & Wootten

12459 (NMC); Bouteloua wamockii Gould & Kapadia: Spellenberg

et al. 12443 (NMC), Heil 34392 (SJNM); Dasyochloa pulchella

(Kunth) Willd. ex Rydb.; Digitaria cognata (Schult.) Pilg. subsp.

pubiflora Wipff & Hatch: Spellenberg & Wootten 12457 (NMC); En-

neapogon desvauxii P. Beauv.; Setaria leucopila (Seribn. & Merr.)

K. Schum.; Sporobolus cryptandrus (Torr.) A. Gray: Spellenberg

& Wootten 12504 (NMC); Sporobolus nealleyi Vasey: Moore et

al. 2271 (MEXU, NMC, OC, TEX/LL); Tridens muticus (Torr.) Nash

var. muticus: Spellenberg & Wootten 12454 (NMC), Heil&Schleser

34720 (SJNM).

Polygonaceae: Eriogonum havardii S. Watson: Spellenberg et al.

12439 (NMC), Heil & O'Kane 33494 (SJNM, UNM), Heil 34404

(SJNM).

Pteridaceae: Astrolepis cochisensis (Goodd.) D.M. Benham &

Windham subsp. chihuahuensis D.M. Benham: Spellenberg etal.

12444 {NMC), Alexander 1333 (NMC); CheilanthesfeeiT. Moore:

Alexander 1331 (WICH).

Rosaceae: Fallugiaparadoxa (D. Don) Endl.

Rubiaceae: Hedyotis nigricans (Lam.) Fosberg var. nigricans: Spellen¬

berg etal. 12445 (NMC), Moore etal. 2282 (MEXU, NMC, OC,TEX/

LL); Hedyotis cf. nigricans (Lam.) Fosberg: Moore etal. 2284 (OC,

NMC, TEX/LL, MEXU).

Solanaceae: Chamaesaracha pallida Averett: Heil 34402 (SJNM),

Alexander 1326 (NMC); Nicotiana trigonophylla Dunal.

Verbenaceae: Aloysia wrightii A. Heller.

APPENDIX 2

The following list includes all species observed by the first author

and nomenclature are as described for Appendix 1.

Amaranthaceae: Atriplex canescens.

Asparagaceae: Dasylirion leiophyllum; Yucca elata.

Asteraceae: Bahia absinthifolia Benth.; Gutierrezia microcephala;

Haploesthes greggii var. texana: Alexander 1318 (NMC, OC);

Parthenium incanum; Porophyllum scoparium; Sartwellia

flaveriae; Sidneya tenuifolia; Thelesperma megapotamicum;

Thymophylla acerosa (DC.) Strother.

Boraginaceae: Tiquilia hispidissima: Alexander 73 77 (NMC, OC).

Brassicaceae: Nerisyrenia linearifolia (S. Watson) Greene: Alexander

1319 (NMC, OC).

Cactaceae: Echinocereus dasyacanthus; Opuntia phaeacantha.

Ephedraceae: Ephedra aspera.

Euphorbiaceae: Chamaesyce fendleri; Croton dioicus.

Fabaceae: Dalea wrightii A. Gray.

Fouquieriaceae: Fouquieria splendens.

gypsum on the east side of Big Dog Canyon. Specimen vouchers

Lamiaceae: Salvia lycioides A. Gray: Alexander 1315 (NMC).

Loasaceae: Cevalliasinuata;Mentzelia humilis (A. Gray) J. Dari. var.

humilis: Alexander 1321 (NMC, OC).

Nyctaginaceae: Acleisanthes lanceolata: Alexander 1316 (NMC, OC);

Allionia incarnata var. incarnata.

Oleaceae: Menodora scabra.

Onagraceae: Oenothera suffrutescens.

Poaceae: Bouteloua curtipendula Torr.; Enneapogon desvauxii;

Muhlenbergia ported Scribn. ex Beal; Sporobolus nealleyi: Alex¬

ander 1320 (NMC, OC); Tridens muticus var. muticus.

Pteridaceae: Astrolepis cochisensis subsp. chihuahuensis.

Rosaceae: Fallugia paradoxa.

Rubiaceae: Hedyotis nigricans var. nigricans.

Verbenaceae: Aloysia wrightii.

ACKNOWLEDGMENTS

We would like to thank the following individuals for assistance with this study: Michael Howard of the BLM

Field Office, Las Cruces, helped arrange site access; George Rauch and Jonna Lou Schafer, ranchers on Crow

Flats, allowed access to Pup Canyon across private land; Rich Spellenberg, Wynn Anderson, Chris Martine,

and Krissa Skogen assisted with held work; Ken Heil and Gregory Penn provided information on specimens at

SJNM. This work was supported by National Science Foundation grant DEB-1054539 and by the National

Geographic Society. The careful reviews of Ihsan Al-Shehbaz and an anonymous reviewer are greatly appreci¬

ated.

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BOOK REVIEW

Thomas N. Taylor, Michael Krings, Edith L. Taylor. 2015. Fossil Fungi. (ISBN 978-0-12-387731-4, hbk.). Aca¬

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$150.00,398 pp., illustrated, 10.75" x 8.5".

The primary focus of this book is to record the history of fungi through time and highlight fossil associations

between fungi and other organisms. The work is arranged in 13 chapters with Chapter 1 defining the held of

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discusses the formation of fossil fungi and procedures used in their investigation. Chapter 3 examines the age

of fungi while Chapters 4-9 cover the various taxonomic groups: Chytridiomycota, Blastocladiomycota, Zygo¬

mycetes, Glomeromycota, Ascomycota, and Basidiomycota. Chapter 10 deals with fossil lichens, Chapter 11

examines fungal spores, Chapter 12 investigates interactions between fungi and other organism, and Chapter

13 explores bacteria and fungus-like organisms, including the enigmatic Peronosporomycetes, puzzling or¬

ganisms that may represent the first eukaryotes on Earth. Also included are a glossary, references, and index.

Many discoveries of the earliest known representatives of fungal groups have been made from the mar¬

velously preserved fossils of the Early Devonian Rhynie chert beds. The Rhynie chert site, dated at some 410

million years BP, represented a subtropical wetland ecosystem consisting of an alluvial plain with small bodies

of water, including hot springs. Both aquatic and terrestrial life forms were preserved, and one of the many

fungal organisms recovered at this site was Paleopyrenomycites devonicus, now considered the earliest known

ascomycete (Fig. 1). A section in the book covering fossil parasitic fungi shows that these organisms have been

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ogy, and the pleasant writing style and beautiful color photos make it a pleasure to read. —George Poinar, Jr.,

Department of Integrative Biology, Oregon State University, Corvallis, Oregon 97331, USA.

Fig. 1 .The spectacular preservation of Early Devonian Rhynie chert fossils is exemplified by these ascomycetous perithecia of Paleopyrenomycites devonicus

developing in the vascular plant Asteroxylon (photo courtesy of T.N. Taylor).

J. Bot. Res. Inst. Texas 8(2): 394.2014

LOMATIUM SWINGERAE: A NEW SPECIES OF LOMATIUM (UMBELLIFERAE)

FROM THE JOSEPH PLAINS, IDAHO, U.S.A.

Richard P. McNeill

1507 S Gonzales Street

Las Vegas, New Mexico 87701, U.S.A.

juniper. botany@gmaii. com

ABSTRACT

Lomatium swingerae R.R McNeill is a new species described from one population on the Joseph Plains, between the Salmon River and

Snake River, Idaho, U.S.A. It co-occurs with L. cous and is morphologically most similar to L. bicolor var. leptocarpum. It is easily distin¬

guished from L. cous based on morphology of the ultimate segments, involucral bractlets and mericarps, and from L. bicolor var. leptocarpum

based on habitat, overall size of the plant, structure of the primary and secondary umbels, the ultimate segments and phenology.

RESUMEN

Lomatium swingerae R.P. McNeill es una nueva especie que se describe de una poblacion de Jose Llanos, entre el rio Salmon y el rio Snake,

Idaho, EE.UU. Se da junto conL. cous y es morfologicamente muy similar a L. bicolor v ar. leptocarpum. Se distingue facilmente deL. couspor

la morfologia de los segmentos finales, bracteas involucrales y mericarpos, y de L. bicolor v ar. leptocarpum por su habitat, el tamano total de

la planta, la estructura de las umbelas primarias y secundarias, los segmentos finales y fenologia.

Keywords: New species, Lomatium, Lomatium swingerae, Umbelliferae, Apiaceae, Salmon River, Snake River, Idaho, Endemic, Joseph Plains

Distinctive landforms and geomorphic processes in conjunction with large-scale patterns of climate change

have resulted in unusual patterns of plant distribution in North Idaho. Consequently, it is home to many rare

and endemic species, and many disjunct populations of species that are normally distributed either west of the

Cascade Mountains or east of the Rocky Mountains (Brunsfeld et al. 2001; McNeill 2012). In 2008,1 discovered

an anomalous taxon of Lomatium on the Joseph Plains, the most northern point of land separating the Salmon

River and the Snake River (Fig. 1). It was not possible to key this taxon reliably in the Flora of the Pacific North¬

west or the Intermountain Flora (Cronquist et al. 1994; Hitchcock et al. 1961). I revisited the site four times

between 15 May 2008-3July 2014 attempting to collect specimen with mature fruit and to locate other popula¬

tions. Mature fruits were finally collected on 3 July 2014 and no other populations have been found. The taxon

was locally common, with approximately several thousand individuals occurring at the site, 86 individuals

were collected for use in this description. Morphologically this species is most similar to Lomatium bicolor JM.

Coult. & Rose var. leptocarpum (Torr. & A. Gray) Schlessman and co-occurs with L. cous J.M. Coult. & Rose.

Lomatium swingerae R.P. McNeill, sp. nov. (Figs. 2—3). Type: U.S.A. IDAHO. Idaho Co.: on ridge top along Divide Creek

Road, on Joseph Plains between the Salmon River and the Snake River, ca. 8 km from junction with Flynn Creek Rd. and ca. 35 km

W of Grangeville, elev. 1285 m, S slope of 0-15 %, 45.851830557-116.57732222°, dry open meadow in Pinus ponderosa forest, shal¬

low, dry soils, very rocky, with mosses filling most of the rock interspaces, soil and rock derived from middle Miocene basalt flows,

27 May 2014, Rick McNeill s.n. (holotype: ID; isotypes: to be distributed).

Lomatium swingerae differs from congeners by geographical and/or ecological separation, phenology, leaf morphology, umbel size/structure,

involucel bractlet shape, pedicel length, mericarp size/structure and root morphology.

Plants acaulescent, perennial, 6.5-32.0(-42.0) cm tall, glabrous to papillate. Roots tuberous thickened, mo-

niliform, or globose, with a simple or rarely branched, subterranean caudex, 1.9-6.6 cm below ground level.

Leaves basal, glabrous, 1-3 leaves per stem; petioles sheathing the stem, extending 0-80.8 mm beyond

sheathing, old leaf bases ± present; blades 19.5-127.7 mm long, 18.8-229.2 mm wide, tripinnate, generally

withered by the time the fruits are mature; ultimate segments 14-28 per 1 cm 2 , acerose, with mucronate api¬

ces, 1.2-14.5 mm long, 0.1-0.9 mm wide, erect from either side of the rachis, pseudofasciculate and appearing

J. Bot. Res. Inst. Texas 8(2): 395 - 401.2014

396

Journal of the Botanical Research Institute of Texas 8(2)

bsmatium swingerae

Idaho

Salmon River

• Lomatium swingerae

Snake River

Vfiz&mJTta

0 2 4 Kilometers

Fig. 1. Joseph Plains and the location of Lomatium swingerae.

McNeill, A new species of Lomatium from Idaho

397

Fig. 2. Lomatium swingerae: habit (from holotype, Rick McNeil s.n., ID), note subterranean caudex and pseudofasciculation of the leaflets with secund

appearance.

398

Journal of the Botanical Research Institute of Texas 8(2)

Fig. 3. Lomatiumswingerae{a \\scale is mm ora 15 cm ruler): A. Root morphology and simple/branched caudices. B. Involucel bractletsand immature fruits.

C. Ultimate segments of leaf: notice acerose form, pseudofasciculation and secund appearance. D. Mericarps: dorsal and ventral view, and E. Habitat.

McNeill, A new species of Lomatium from Idaho

399

secund. Scapes 1-3, 3.3-28.0(-38.5) cm long, ascending, corrugated, glabrous to papillate. Inflorescence

compound umbel, glabrous to papillate; involucre absent; rays (1—)2—10, 10.9-60.4 mm long, ascending to

erect. Flowers sepals 5, connate, inconspicuous; petals 5, yellow, obcordate; ovary superior, two carpellate,

two styles, two yellow, exserted anthers, filaments yellow; pedicels l-4.3(-6.2) mm long; involucel bractlets

spreading to slightly reflexed at maturity, 0.3-2.l(-3.5) mm long, to 0.2 mm wide, narrowly lanceolate to nar¬

rowly oblanceolate, withering by the time the of mature fruits; mericarps ensiform or gladiate, deplanate or

revolute, 10.0-16.5(-18.3) mm long, (0.87-)1.9-3.3(4.0) mm wide, lateral wings ± present, up to 0.5 mm wide,

less than or equal to A as wide as the body, dorsal ribs 3(4); vittae interval 1 between each rib, commissural

2-4(-6), maybe obscure, or incomplete; stylopodia yellowish tan, smooth to paleaceous; carpophore yellow¬

ish tan, 9.3-15.0 mm long.

Distribution and Phenology. —Occurring in a dry, open, rocky meadow. Flowering March through June

and fruitingjune through July.

Etymology. —The specific epithet is for Lindsay Ann Swinger, botanist and friend.

Lomatium swingerae is only superficially similar to L. cous, both tending to be small for Lomatiums; L.

swingerae typically under 3.2 dm and L. cous typically under 3.5 dm. They can be distinguished from each

other by the morphology of the mericarps, involucral bractlets, and the size and shape of the ultimate seg¬

ments. The wings on the mericarps of L. cous are as wide or nearly as wide as the body, while in L. swingerae the

lateral wings may or may not be present, if present, are up to l h as wide as the body. Lomatium cous has broad,

distinctive involucel bractlets that are oblanceolate, obovate or elliptic, while the bractlets of L. swingerae are

narrowly lanceolate to narrowly oblanceolate. Lomatium cous has relatively broad (up to 3mm) and leaf-like

ultimate segments, while on L. swingerae the ultimate segments are narrow (0.1-0.9 mm) and acerose. Loma¬

tium swingerae can be distinguished from Lomatium bicolor var. leptocarpum by the arrangement and shape of

the ultimate segments, the arrangement of the primary and secondary umbels, the size of the plant and habitat.

Lomatium bicolor var. leptocarpum has ascending ultimate segments that are long (10-50 mm) and broad

(0.5-2 mm), while the ultimate segments of L. swingerae are erect from both sides of the leaf rachis, pseudofas-

ciculate, appearing secund, shorter (1.2-14.5 mm long) and narrower (0.1-0.9 mm wide). The primary umbel

of Lomatium bicolor var. leptocarpum is appressed, while L. swingerae is erect to ascending. Lomatium bicolor

var. leptocarpum has very crowded mericarps due their size and the short pedicels, while L. swingerae generally

has smaller mericarps and longer pedicels that result in less crowding. Lomatium bicolor var. leptocarpum is

generally 1-5 dm tall, while L. swingerae is generally no taller than 3.2 dm rarely, 4.2 dm, and finally L. bicolor

var. leptocarpum occurs in wet meadows, and ephemeral pools, usually with high clay content soils, while L.

swingerae is found in a dry, rocky meadow (Schlessman 1984).

Lomatium swingerae has only been found in one location, on the Joseph Plains, which is a high plateau

separating the Salmon River and Snake River approximately 35 km west of Grangeville, ID. The Salmon River

and Snake River have created canyons on three sides of the Joseph Plains that are approximately 980 m lower

than the plains, and up to 16 km wide. Lomatium swingerae occurs in an open rocky meadow with very little

soil development. The rocky interspaces are filled with mosses, soil, and roots. This soil and rock are derived

from the middle Miocene flood-basalt flows, occurring between 16-11.6 million years ago (Bond et al. 1978).

Basalt-derived soils tend to be influenced more by climate than the mineral composition of the parent material,

so the uniqueness of the habitat is likely due to the landscape position on top of an exposed, dry ridge rather

than the result of any special properties of the basalt (Kruckeburg 2002). Lomatium swingerae flowers from

March to June and the fruit mature in late June through July, with timing largely dependent on seasonal varia¬

tion. It is also distinctive from the other Lomatiums on the Joseph Plains in that it flowers and fruits before the

five other species: Lomatium ambiguum J.M. Coult. & Rose, L. cous, L. dissectum (Nutt, ex Torr. & A. Gray)

Mathias & Constance, L. bicolor var. leptocarpum and L. triternatum J.M. Coult. & Rose ssp. platycarpum

(Torr.) Cronquist. On 27 May 2014, Lomatium swingerae had green fruits, while all the other Lomatiums were in

flower, and on 3 July 2014, both L. swingerae and L. cous had mature fruits, but L. bicolor var. leptocarpum, L.

triternatum ssp. platycarpum and L. dissectum had immature fruits. Only Lomatium cous and L. triternatum ssp.

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Journal of the Botanical Research Institute of Texas 8(2)

platycarpum have been found within 2 km of the population of L. swingerae. Lomatium cous and L. swingerae

grow next to each other in the rocky areas of the meadow. In the nearby soils that are less rocky, deeper or wet¬

ter, only L. cous is found. Lomatium triternatum ssp. platycarpum is found on the edge of the meadow, in less

rocky, deeper soils that are partially shaded by the trees. Lomatium swingerae occurs in association with the

following species: Allium tolmiei Baker exJ.M. Coulter var. platyphyllum (Tidest.) Ownbey, Balsamorhiza incana

Nutt., Castilleja chromosa A. Nelson, Crepis atribarba A. Heller ssp. atribarba , Delphinium depauperatum Nutt.,

Erigeron bloomeriA. Gray, Erigeron englemannii A. Nels., L. cous J.M. Coult. & Rose, Microsteris gracilis (Hook.)

Greene var. gracilis, Orobanche uniflora L. var. purpurea (Heller) Achey, and Poa bulbosa L.

Paratypes: U.S.A. IDAHO. Idaho Co.: plateau between the Salmon and Snake Rivers, elev. 1259 m, 45°51.392'N, 116°34.493'W, S + N slope

1-40%, meadow in Pinus ponderosa woodland, rocky soil, 21 May 2008, Rick McNeill s.n. (ID); on ridge top along Divide Creek Road, on Jo¬

seph Plains between the Salmon River and the Snake River, ca. 8 km from junction with Flynn Creek Rd, and ca. 35 km W of Grangeville,

elev. 1285 m, S slope: 0-15 %, 45.851830557-116.57732222°,dry open meadow in Pinus ponderosa forest, shallow, dry soils, very rocky, with

mosses filling most of the rock interspaces, soil and rock derived from middle Miocene basalt flows, 3 Jul 2014, Rick McNeill s.n. (ID).

Eomatium swingerae is endemic to northern Idaho, known from only one population and should be considered

for listing as a sensitive species. The limited geographic range and single known population make it extremely

vulnerable to extirpation.

KEY TO LOMATIUM TAXA OF THE JOSEPH PTAINS, IDAHO

1. Involucel bractlets absent_ L. ambiguum

1. Involucel bractlets present.

2. Involucel bractlets broadly oblanceolate to broadly obovate or elliptic, wing of the mericarp equal or nearly as wide

as the body, root tuberous thickened or globose_ L. cous

2. Not with the above combination of characters.

3. Flowers purple, plants 5-15 (20) dm tall, large woody taproot_ L. dissectum

3. Flowers yellow, if plants over 5 dm tall, then slender taproot, or cormose thickened, not large and woody.

4. Wing of the mericarp nearly as wide as the body, ultimate segments few, less than 10 per cm 2 _ L. triternatum

ssp. platycarpum

4. Wing of the mericarp narrow, less than Vs of the body, ultimate segments many, more than 10 per cm 2 .

5. Plant growing in wet meadows or ephemeral pools, ultimate segments ascending, 1-5 cm long, 0.5-2 mm

wide, rays of primary umbels appressed, mericarps crowded, root short and cormose thickened to more

elongate and slender_ L. bicolor var. leptocarpum

5. Plants growing in dry rocky meadows, ultimate segments erect, pseudofasciculate, appearing secund, 0.1-1.5cm

long, 0.1-0.9 mm wide, rays of primary umbels erect or ascending, mericarps not crowded, root globose,

moniliform or tuberous thickened_ L. swingerae

ACKNOWTEDGMENTS

I would like to thank Todd Ott for his help with collections, Cort Anderson for providing accommodations

while I was collecting and Don Mansfield, and Stephen R. Downie for their thoughtful and timely reviews. The

following herbaria contributed specimen, use of equipment and space to sort collections: the Stillinger Her¬

barium (ID) University of Idaho, the Rocky Mountain Herbarium (RM) University of Wyoming, and the Mar¬

ion Ownbey Herbarium (WS) Washington State University. Finally, I would like to thank Barney Lipscomb for

his help and patience.

REFERENCES

Bond, J.G., J.D. Kauffman, D.A. Miller, & R. Venkatakrishnan. 1978. Geologic map of Idaho. Idaho Bureau of Mines and Geol¬

ogy, with contributions from U.S. Geological Survey, scale 1:500,000.

Brunsfeld, S.J., J. Sullivan, D.E. Soltis, & P.S. Soltis. 2001. Comparative phylogeography of northwestern North America: a

synthesis. In: J. Silverton, and J. Antonovics, eds. Integrating ecological and evolutionary processes in a spatial con¬

text. Blackwell Science, Oxford, UK. Pp. 319-339.

Cronquist, A., N.H. Holmgren, & P.K. Holmgren. 1994. Intermountain flora: vascular plants of the intermountain west, U.S.A.,

Vol. 3: part A subclass Rosidae (except Fabales). New York Botanical Garden Press, Bronx, NY, U.S.A.

Hitchcock, C.L., A. Cronquist, M. Ownbey, & J.W. Thompson. 1961. Vascular plants of the Pacific Northwest, part 3: Saxifraga-

ceae to Ericaceae. University of Washington Press, Seattle, U.S.A.

McNeill, A new species of Lomatium from Idaho

401

Kruckeburg, A.R.2002.Geology and plant life: the effects of landforms and rock types on plants. University of Washington

Press, Seattle.

McNeill, R.P. 2012. Lomatium brunsfeldianum: a new species of Lomatium (Umbelliferae) from Northern Idaho. J. Bot.

Res. Inst.Texas 6(1 ):29-36.

Schlessman, M.A. 1984. Systematics of tuberous Lomatiums (Umbelliferae). Syst. Bot. Monogr. 4:1-55.

402

Journal of the Botanical Research Institute of Texas 8(2)

BOOK REVIEW

Jacques Cayouette. 2014. A la Decouverte du Nord: Deux siecles et demi d’exploration de la flore nordique

du Quebec et du Labrador. (ISBN-13: 978-2-89544-471-8, hbk.). Editions MultiMondes, 930 me Pou-

liot, Quebec, Quebec G1V 3N9, CANADA (Orders: www.multim.com, 1-800-840-3029). $45.00, 363

pp., color photos, index, 8" x 10". French language text throughout.

The two and a half centuries covered in this book included many excursions into land that had scarcely been

explored, the prodigious activity of botanists and others interested in the rich and unique life of a forbidding

landscape. The Moravians, Europeans of one religion and several nationalities, came from their settlement in

Greenland to establish a colony, Nain, in Labrador in 1771. With their respect for all creation, they collected

plants which eventually found their way into the hands of private collectors in England, Germany, Sweden,

and elsewhere and later into the herbariums of universities and museums. Their activity continued into the

1870s, with botanists such as Johannes Lundberg and Samuel Weiz continuing the work of their predecessors.

From England, France, Denmark, Sweden, and the United States botanists, geologists, and zoologists came to

collect, many taking specimens for deposit in their national and university herbariums. There was much to be

found, not only in vascular plants but probably even more importantly in the bryophytes, lichens, and algae.

Species of Carex and other Cyperaceae were of special interest because of the many forms these plants took in

various habitats and different soil types.

From 1842 the Company of Hudson Bay also took a large role in exploration as did later the Canadian

Geological Commission with the Geological and Natural History Survey of Canada. Both of these governmen¬

tal units contributed much to the knowledge of the plants of northern Quebec and Labrador. However, it was

work in the early and middle twentieth century that really led to a complete understanding of the floral heritage

of these regions of Canada, knowledge which resulted in the first volume of the comprehensive Flore Nordique

in 2013. Notable individuals among the botanists active in this period include the indomitable priest Marie-

Victorin (1885-1944), Jacques Rousseau (1905-1970), Nicholas Polunin (1909-1997), Ilmari Hustich (1911—

1982), and Marcel Raymond (1915-1972).

This work is not only a detailed, scholarly history of the development of knowledge of the alpine-boreal

plant communities, but in fact is a kind of festschrift to all those individuals who have contributed so much to

the science. There are pictures of some of their mounted herbarium specimens and of their publications, scien¬

tific and popular, and color photographs of plants in situ that they wrote about. The numerous color photos of

beautiful plants and geographic regions add much to the pleasure of reading about those who discovered and

described them. The bibliography lists the publications, some extensive, of these persons as they made known

to the world their discoveries. Of interest to many botanists may be particular entries in the Index, for example

56 taxa of Carex, 14 of Eriophorum, 11 of Draba. This volume is an important addition to the botanical history

of North America. —Joann Karges, Texas Christian University Library (retired), Fort Worth, Texas, USA.

J. Bot. Res. Inst. Texas 8(2): 402.2014

NEW GEOGRAPHICAL AND MORPHOLOGICAL DATA LOR

SIDEROXYLON RECLINATUM SUBSPECIES

AUSTROFLORIDENSE (SAPOTACEAE), A TAXON ENDEMIC

TO SOUTHEASTERN PENINSULAR FLORIDA, U.S.A.

Paul T. Corogin and Walter S. Judd

Department of Biology

220 Bartram Hall

and

Florida Museum of Natural History

University of Florida

Gainesville, Florida 32611, U.S.A.

pcorogin@ufl.edu; lyonia@ufl.edu

ABSTRACT

Sideroxylon redinatum Michx. subspecies austrofloriden.se (Whetstone) Kartesz & Gandhi is re-diagnosed in light of new morphological

data, compared with its presumed closest relatives, and the extent of its distribution is clarified. Sideroxylon redinatum subsp. austro-

floridense is endemic to subtropical rockland habitats of extreme southeastern peninsular Florida, U.S.A. These habitats face threats from

exotic species invasions, fire suppression, and sea-level rise due to climate change. Sideroxylon rufohirtum Herring & Judd, sometimes con¬

sidered a subspecies of S. redinatum Michx., is here recognized as a separate species.

RESUMEN

Se vuelve a diagnosticar Sideroxylon redinatum Michx. subspecie austrofloridense (Whetstone) Kartesz & Gandhi a la luz de nuevos datos

morfologicos, comparados con los de sus presuntos parientes mas proximos, y se clarifica la extension de su distribucion. Sideroxylon

redinatum subsp. austrofloridense es endemico de habitats rocosos subtropicales del extremo sureste peninsular de Florida, U.S.A. Estos

habitats se enfrentan a amenazas de invasiones de especies exoticas, supresion por fuego, y elevacion del nivel del mar por cambio climatico.

Sideroxylon rufohirtum Herring & Judd, a veces considerado como subespecie de S. redinatum Michx., se reconoce aqui como una especie

diferente.

INTRODUCTION

Sideroxylon L. (Sapotaceae) is a pantropical genus comprising ca. 80 species, occurring most abundantly in

tropical America, but also in Macaronesia, Africa, Madagascar, and the Mascarene Islands, with few species in

southeast Asia (Govaerts et al. 2001; Smedmark & Anderberg 2007). Prior to Pennington’s monographs of the

Sapotaceae (1990, 1991), the Neotropical species of Sideroxylon, centered in the Caribbean and Central Amer¬

ica and extending into the temperate southeastern U.S., were split among the genera Bumelia Sw., Dipholis

A.DC., and Mastichodendron (Engl.) H.J. Lam. The species endemic to the continental U.S. (and extreme north¬

ern Mexico) were assigned to Bumelia, along with many tropical taxa (Cronquist 1945, 1946, 1949). They are

trees and shrubs of mainly dry habitats, usually armed with thorns, with leaves alternate, often becoming

fascicled on short shoots (brachyblasts), inflorescences fasciculate, axillary or in axils of fallen leaves, flowers

with a single whorl of quincuncial sepals, corolla lobes with a median segment and two lateral segments, pet-

aloid staminodes alternating with epipetalous stamens, and the seed having its hilum scar in a basal position,

with endosperm scanty or lacking (Cronquist 1945). Bumelia was revised by Gray (1886), Small (1900), Clark

(1942), and Cronquist (1945, 1949), all of whom differed concerning the number of U.S. species. Pennington

(1990) transferred all of Bumelia, Dipholis, and Mastichodendron to a broadly circumscribed Sideroxylon, and

for species limits he generally followed Cronquist, recognizing eight U.S. species. More recently, phylogenetic

analyses using both morphological and molecular data have shown that Sideroxylon sensu Pennington is

monophyletic including Argania from Morocco (Anderberg & Swenson 2003; Swenson & Anderberg 2005;

J. Bot. Res. Inst. Texas 8(2): 403 - 417.2014

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Journal of the Botanical Research Institute of Texas 8(2)

Smedmark et al. 2006), but excluding two taxa from the Arabian peninsula and Socotra (Smedmark & Ander-

berg 2007; Gautier et al. 2013; Stride et al. in press). Although relationships among the major subclades of Sider-

oxylon are unclear, molecular phylogenies reveal two well-supported Neotropical subclades, one correspond¬

ing to Dipholis/Mastichodendron, and the other to Bumelia, including the species endemic to North America

(Smedmark & Anderberg 2007; Stride et al. in press).

Currently eleven species of Sideroxylon are recognized in the continental United States (Govaerts et al.

2001; Allison 2006; Wunderlin & Hansen 2008). Of these eleven species, eight are endemic to an area encom¬

passing much of the southeastern U.S. and a bit of extreme northern Mexico, most being concentrated on the

coastal plain from Texas through the Carolinas, with Florida as their center of diversity (Govaerts et al. 2001;

Allison 2006; Wunderlin & Hansen 2008). These eight are: Sideroxylon alachuense L.C. Anderson, S. lanugino-

sum Michx., S. lycioides L., S. macrocarpum (Nutt.) J.R. Allison, S. reclinatum Michx., S. rufohirtum Herring &

Judd, S. tenax L., and S. thornei (Cronquist) T.D. Penn. The above-listed taxa, which likely form a monophy-

letic group (Smedmark & Anderberg 2007; Corogin and Judd, unpublished DNA sequence data), are distin¬

guished morphologically from their tropical congeners by having a combination of fascicled leaves, conspicu¬

ously reticulate tertiary venation, short styles, small fruits, and a bipartite hilum scar (Pennington 1990).

Common names include “buckthorn,” “bully,” and “bumelia.”

Sideroxylon reclinatum is a mostly low-growing thorny shrub (occasionally a small tree) of calcareous or

sandy, wet to mesic wooded habitats of Florida and the outer coastal plain from southern Georgia to Louisiana.

Cronquist (1945) recognized two varieties, Bumelia reclinata (Michx.) Vent. var. reclinata, and B. reclinata var.

rufotomentosa Cronquist, the latter having been previously recognized as a species (B. rufotomentosa) by Small

(1900). Whetstone (1985) named a third variety, B. reclinata var. austrofloridensis Whetstone (“Everglades

buckthorn”), based on morphologically distinct collections from subtropical rocklands in the vicinity of Long

Pine Key in the Florida Everglades. Varieties rufotomentosa and austrofloridensis were transferred to Sideroxy¬

lon by Kartesz and Gandhi (1990), and recognized as S. reclinatum subsp. rufotomentosum (Small) Kartesz &

Gandhi, and S. reclinatum subsp. austrofloridense (Whetstone) Kartesz & Gandhi, respectively. Based on con¬

sistent differences in morphology, habit, and habitat, Herring and Judd (1995) recognized S. reclinatum subsp.

rufotomentosum as a distinct species, Sideroxylon rufohirtum Herring & Judd, (see Godfrey 1988), and this

designation is here recognized. S. reclinatum, S. rufohirtum, and a third species, S. macrocarpum (Nutt.) J.R. Al¬

lison, are here referred to as the Sideroxylon reclinatum complex, as they share many similarities in morphology

and habit and are traditionally presumed closely related. We discuss how S. reclinatum subsp. austrofloridense

is distinguished from S. reclinatum subsp. reclinatum, and also from the other two species in the complex.

Our close examination of specimens including more recent collections, along with held observations of

both S. reclinatum subsp. reclinatum and S. reclinatum subsp. austrofloridense, reveal that the two subspecies are

most reliably distinguished by differences in the micromorphology of the abaxial leaf epidermis, and that the

extent of distribution of subsp. austrofloridense, while limited to extreme southern peninsular Florida, is some¬

what larger than was previously supposed.

The focus of this paper is the southern Florida subtropical rockland populations of Sideroxylon reclinatum

that form subspecies austrofloridense. Our aim is to present new morphological and geographical data on these

populations, to revise the circumscription and range of the subspecies in light of this new data, to compare the

subspecies with its presumed closest relatives, and to discuss its importance and conservation status as part of

the endemic flora of southern Florida rockland habitats. A comprehensive taxonomic treatment of S. reclina¬

tum, including both subspecies, will be presented in an upcoming revision of the clade of Sideroxylon endemic

to the continental U.S. and extreme northern Mexico.

MATERIALS AND METHODS

The morphological findings of this study are based on close examination of numerous specimens, both fresh

and dried. Dried flowers were rehydrated by boiling in water, and dissected with the aid of a Wild M5A dissect¬

ing microscope (Aarau, Switzerland). Dimensions of trichomes were measured with the aid of a Carl Zeiss

Corogin and Judd, Sideroxylon redinatum ssp. austrofloridense

405

4311036 compound microscope. Scanning electron microscopy (SEM) was employed for examination of leaf

surface micromorphology. Leaf material from dried herbarium specimens of taxa being compared was mount¬

ed on carbon adhesive tabs on aluminum specimen mounts. Samples were rendered conductive by coating

with a gold-palladium alloy in argon vacuum for 90 seconds using a Denton Desk V sputter coater (Denton

Vacuum, Moorestown, NJ, USA). Samples were examined with a Hitachi S-4000 beld-emission scanning elec¬

tron microscope (Hitachi High Technologies America, Inc. Schaumburg, IL), and digital micrographs were

acquired with PCI Quartz software. SEM work was conducted at the Electron Microscopy Core of the Interdis¬

ciplinary Center for Biotechnology Research (ICBR) at the University of Florida. From Sideroxylon reclinatum

subsp. reclinatum and S. reclinatum subsp. austrofloridense, twelve individuals of each taxon were sampled;

from S. rufohirtum and S. macrocarpum, five individuals of each taxon were sampled. Conclusions regarding

the geographical range of Sideroxylon reclinatum subsp. austrofloridense are based on extensive held observa¬

tions and data from herbarium specimen labels.

RESULTS AND DISCUSSION

Morphology .—The species of the Sideroxylon reclinatum complex share some morphological similarities such

as low habit, small leaves with prominent vein reticulum, and abaxial leaf pubescence that is usually sparse, if

present, and often deciduous. They are Sideroxylon reclinatum, S. macrocarpum, and S. rufohirtum. Sideroxylon

reclinatum ranges throughout Florida, southwestern Georgia, and westward along the coastal plain to Louisi¬

ana, growing in mesic to wet, sandy to calcareous habitats. This species is not typically highly clonal, is gener¬

ally low-growing but can reach heights of ca. 3 m as many-trunked shrubs or occasionally small trees, and has

small fruits (10 mm diameter or less). It is usually a glabrous plant, any leaf or young stem pubescence quickly

deciduous, except for the south Florida subspecies austrofloridense, in which pubescence can be persistent. S.

macrocarpum is a low-growing, highly clonal shrub narrowly endemic to a few counties in southeast Georgia.

It is similar in habit and habitat to S. rufohirtum, a putative close relative endemic to a few counties in north and

west peninsular Florida (Allison 2006). These two species prefer dry sandy habitats, are both highly clonal and

low-growing, rarely exceeding 1 m in height, and they have large fruits (> 10 mm diameter), young stems pu¬

bescent through the first season, and abaxial leaf pubescence that is sparse and often persistent.

Scanning electron microscope (SEM) images of the abaxial leaf cuticular surfaces of these species (Figs. 1,

2) reveal that the surface features of these taxa are quite distinct from one another. Previous SEM study has

shown that taxa of Sideroxylon can be distinguished by such characters (Anderson 1996). Stomata in all cases

are recessed inside chambers with generally elliptical openings. In both subspecies of Sideroxylon reclinatum,

each chamber opening is surrounded by a series of irregularly concentric cuticular ridges, often also with

ridges radiating out perpendicularly from the opening. The features of Sideroxylon macrocarpum and S. rufohir¬

tum are rather different from those of S. reclinatum (Figs. 1, 2). The stomatal chambers of these species are

somewhat larger than those of S. reclinatum. In Sideroxylon macrocarpum, each elliptic-shaped stomatal cham¬

ber opening sits atop a small dome raised above the epidermal surface, and surrounding the dome is a border

of one or more concentric ridges; at least one of these ridges tends to form a striking, neatly defined oval border

around the dome. The epidermal surface is strongly patterned, with impressed grooves defining cell outlines.

The stomatal structures of Sideroxylon rufohirtum are distinctive as well, with stomatal chamber openings

centered on conspicuously donut-shaped, broad, flat to slightly cupped, elevated platforms bordered by con¬

centric cuticular ridging, and the cuticular surface between stomata is relatively smooth but with epidermal

cell outlines visible and defined by impressed grooves.

At the macromorphological level, the two subspecies of Sideroxylon reclinatum are not always easily dis¬

tinguishable (Table 1). Subspecies reclinatum is typically a glabrous plant. Abaxial leaf pubescence, if present at

all, tends to be grey to white in color, the trichomes straight or weakly curled, and it is never dense or persistent;

trichomes quickly slough off as the leaf matures, persisting only along the abaxial midvein if at all. Sepals and

pedicels are glabrous, and the ovary is glabrous or with only a few scattered trichomes at anthesis (Fig. 3). The

population of subspecies austrofloridense at Long Pine Key in the Everglades is morphologically quite distinct,

406

Journal of the Botanical Research Institute of Texas 8(2)

Fig. 1. SEM images of abaxial epidermal leaf surface of four taxa of Sideroxylon, showing differences in stomatal structures and cuticular surfaces. A: 5.

redinatum subsp. austrofloridense (Corogin 1055, FLAS); B: 5. redinatum subsp. redinatum (Corogin 1060, FLAS); C: 5. macrocarpum (Allison 9404, USCH);

D:5. rufohirtum (Corogin 1000, FLAS). (Magnification 500x).

Table 1. Comparison of key morphological differences between Sideroxylon redinatum subsp. austrofloridense and 5. redinatum subsp. redinatum.

subsp. austrofloridense

subsp. redinatum

Pubescence, abaxial

surface of leaf

Ovary at anthesis

Sepals at anthesis

Stomatal structures

Cuticular surface, leaf

abaxial (at high

magnification)

Sparse to dense, finely wooly, matted, sometimes

obscuring leaf surface. Persistent across all of

lamina, or quickly deciduous, leaving surface

glabrous. Most trichomes strongly curled,

ascending, light brown, fading to gray.

Sparsely to densely pubescent.

Glabrous to densely pubescent.

Cuticular ornamentations around chamber

openings are of low to moderate relief.

Epidermal cell outlines always clearly visible

and marked by an impressed groove. Surface

elaborately ornamented with reticulate

pattern in strong relief.

Sparse to absent. Quickly deciduous, persistent

only along midvein if at all. Trichomes fine,

straight to weakly curled, appressed or

ascending, light brown to white.

Glabrous or with only a few scattered trichomes.

Cuticular ornamentations around chamber

Openings are of moderate to strong relief

Epidermal cell outlines not always clearly visible,

and not marked by an impressed groove.

Surface generally smooth and irregularly

undulating.

readily distinguishable at a glance from subspecies redinatum by the persistent matted wooly brown pubes¬

cence on the abaxial leaf surfaces and often also on the sepals and pedicels, and the ovary is densely to sparsely

pubescent at anthesis (Fig. 3). Whetstone (1985) diagnosed subspecies austrofloridense based on specimens

having these features. But our examination of many south Florida specimens, and recent observations in the

held, indicate that plants from locations outside of Long Pine Key (and even a few at Long Pine Key) have a

Corogin and Judd, Sideroxylon redinatum ssp. austrofloridense

407

Fig. 2. SEM close-up views of stomatal structures of four taxa of Sideroxylon. At magnification 3500x; A: 5. redinatum subsp. austrofloridense (Corogin

1055, FNPS); B: 5. redinatum subsp. redinatum {Godfrey81581, FTG). At magnification 3000x; C: 5. macrocarpum {Allison 9404, USCH); D: 5. rufohirtum

{Ward 1206, FLAS). Note the relative sizes—structures of 5. macrocarpum and 5. rufohirtum are somewhat larger than those of 5. redinatum subsp.

austrofloridense and 5. redinatum subsp. redinatum.

sparser and less persistent leaf indumentum, appearing nearly glabrous, many of them most closely resem¬

bling subspecies redinatum. Whether pubescent or glabrous, however, all of the plants within the range of

subspecies austrofloridense exhibit characters at the micromorphological level that distinguish them from

plants outside this range. It turns out that the two subspecies are most reliably distinguishable by differences

in epidermal and cuticular patterns of the abaxial leaf surface (Table 1). These features are visible with good

lighting using a high-quality dissecting microscope (such as the Wild M5A) at lOOOx, but scanning electron

microscopy reveals these features in dramatic clarity. In subspecies austrofloridense (Fig. 4), the cuticular

features are of moderate to strong relief, and the surface between stomata is elaborately ornamented with a

reticulating pattern of grooves and ridges. Epidermal cell outlines are always prominently visible and marked

by an impressed groove or by a ridge with a groove within it (See arrows, Fig. 4). In subspecies redinatum , by

comparison (Fig. 5), the cuticular ornamentations are of low to moderate relief, and the surface between sto¬

mata is relatively smooth and irregularly undulating. Epidermal cell outlines, if visible, are marked by raised

ridges that lack an impressed groove along their middle (See arrows, Fig. 5). Thus at the micromorphological

level it is easy to distinguish the two subspecies.

KEY TO SPECIES OF THE SIDEROXYLON RECLINATUM COMPLEX

1. Plants highly stoloniferous, often forming clonal patches, aerial shoots rarely more than 1 m tall; fruit 10-14 mm long,

plant of dry sandy habitats.

2. Pubescence on young twigs dense, rusty dark red-brown, often persisting well past the first season; abaxial pubescence

408

Journal of the Botanical Research Institute of Texas 8(2)

Fig. 3. (A-B): Ovary, A: Sideroxylon reclinatum subsp. reclinatum ; B: 5. reclinatum subsp. austrofloridense. (C-D): Flowering twig; C: 5. redinatum subsp.

redinatum; D: 5. redinatum subsp. austrofloridense. E-H: 5. redinatum subsp. austrofloridense, Long Pine Key, Everglades National Park, E: Shrub in

marl prairie, ca. 1.5 m tall (pine rockland in background); (F) Flowering branch; (G) Fruiting branch; (H) Older branch, ca. 3 cm diameter, with leaves

and flowers on brachyblasts.

on young leaves rusty red-brown, sparse to moderately dense, giving surface a distinctly reddish hue; staminodes

nearly as long as median corolla lobes; ovary densely pubescent; seed variegated; stomatal chamber openings

located on raised, round, donut- shaped platforms_ S. rufohirtum

2. Pubescence on young twigs moderate to dense, blonde to light brown, rarely persisting past the first season; abaxi-

al pubescence on young leaves blonde to light brown, sparse; staminodes only Vi to % the length of median corolla

lobes; ovary glabrous or with a few trichomes medially; seed uniformly brown; stomatal chamber openings atop

raised elliptical domes neatly bordered by one or more concentric ridges_ S. macrocarpum

1. Plants not highly stoloniferous, growing as erect shrubs with one to several stems, or occasionally small trees, often

taller than 1 m; fruit 7-10 mm long, plant of mesicto wet habitats_ S. reclinatum

Delimitation of the subspecies of Sideroxylon reclinatum. —It can be seen that Sideroxylon reclinatum subsp.

austrofloridense is easily and reliably diagnosable from S. reclinatum subsp. reclinatum by a set of distinct char¬

acter state differences at the micromorphological level. At the macromorphological level, the distinction is less

reliable, as glabrescent specimens of subspecies austrofloridense can be confused with the typically glabrous

subspecies reclinatum. The two taxa are also separated eco-geographically. The more widespread subspecies

reclinatum ranges in coastal plain areas from south Georgia west to Louisiana, and throughout Florida as far

south as Broward County in the east, and Collier and Monroe Counties in the west, and its habitats feature

sandy soils often with limestone near the surface. Subspecies austrofloridense, on the other hand, is a narrow

endemic, its range restricted to subtropical rockland and marl prairie habitats in a well-defined area of extreme

southeast peninsular Florida, and its habitats feature exposed limestone, or limestone thinly overlain with

clayey marl soils. All specimens we examined that were collected within the range of subspecies austro¬

floridense, whether glabrous or pubescent, exhibit the unique micromorphological cuticular surface characters

described above (Figs. 4, 5). The only place where plants of both micromorphological types occur together is

Corogin and Judd, Sideroxylon redinatum ssp. austrofloridense

409

Fig. 4. SEM images of abaxial leaf surface of four different individuals of Sideroxylon redinatum subsp. austrofloridense. A: Corogin 1055 (FNPS), Long Pine

Key, Everglades National Park; B: Possley67 (FTG), urban Miami-Dade County, FL, Larry & Penny Thompson Park; C: Bradley 1547 (FTG), Collier County, FL,

near Monument Lake, Big Cypress National Preserve; D: V.l. Sullivan s.n. (FTG), Long Pine Key, Everglades National Park. Note that epidermal cell outlines

are defined in all cases by an impressed groove or a ridge with an impressed groove within it (see arrows). (Magnification 500x).

where the ranges of the two subspecies come into contact, at the western fringes of the range of subspecies

austrofloridense , in Big Cypress National Preserve (Fig. 6). The two subspecies are thus clearly defined, and es¬

sentially allopatric. Because S. reclinatum subsp. austrofloridense is a narrow endemic restricted to a particular

habitat and geographical area, and because it is allopatric with respect to subspecies reclinatum and diagnos-

able from it, we consider Sideroxylon. reclinatum subsp. austrofloridense to be worthy of taxonomic recognition.

Since the diagnosing characters are micromorphological and somewhat cryptic, and pubescence characters

intergrade, we consider recognition at the subspecific rank to be appropriate. Thus the current classification of

these plants is maintained, but the distribution of subspecies austrofloridense is broadened, and it is best diag¬

nosed by the newly discovered micromorphological features.

KEY TO THE SUBSPECIES OF SIDEROXYLON RECLINATUM

1. Young twigs and leaves glabrous or sparsely pubescent with quickly deciduous, fine, blonde to light brown trichomes;

mature leaves glabrous or with a few blonde to light brown trichomes persisting along midvein; sepals, pedicels and

ovary glabrous or with very few pale trichomes; abaxial leaf epidermal cell outlines obscure, or if visible, defined by raised

ridges that lack an impressed groove along their middle. _S. reclinatum subsp. reclinatum

1. Young twigs and leaves densely to sparsely pubescent with brown, finely wooly trichomes; mature leaves glabrous to

densely pubescent across abaxial lamina with finely wooly, matted, brown to grey indumentum; sepals, pedicels, and

ovary glabrous to densely pubescent with brown trichomes; abaxial epidermal cell outlines always clearly visible, strik¬

ingly defined by impressed grooves, or raised ridges with impressed grooves along their middle _ S. reclinatum

subsp. austrofloridense

410

Journal of the Botanical Research Institute of Texas 8(2)

Fig. 5. SEM images of abaxial leaf surface of four different individuals of Sideroxylon reclinatum subsp. reclinatum. A: Corogin 1060 (FLAS), Gilchrist

County, FL; B: Godfrey81312 (FLAS), Taylor County, FL; C: Correll42244 (FTG), Monroe County, FL, at Pine Crest in Big Cypress National Preserve; D: Godfrey

81581 (FTG), Columbia County, FL. Note that epidermal cell outlines, if visible at all, are defined by raised ridges without the presence of an impressed

groove (see arrows). (Magnification 500x).

TAXONOMIC TREATMENT

Sideroxylon reclinatum Michx. subsp. austrofloridense (Whetstone) Kartesz & Gandhi, Phytologia 68:425.

1990. Bumelia reclinata (Michx.) Vent. var. austrofloridensis Whetstone, Ann. Missouri Bot. Gard. 72:545.1985. Type: U.S.A. Flor¬

ida. Dade Co.: Everglades National Park, 7 July 1984, R.D. Whetstone 14459 (holotype: JSU).

Shrub, single to many-stemmed, occasionally a small tree, ca. 0.5-3.5 m tall, trunk diameter ca. 1-7 cm. Indu¬

mentum of unicellular, light brown to grey-white, occasionally red-brown, T-shaped trichomes, symmetric or

asymmetric, terete and hollow to flattened in cross-section, having a short stalk and a longer upper portion of

varying length, the upper portion curled and ascending. Vegetative buds densely wooly-pubescent with short,

curled, ascending, red-brown T-shaped trichomes, the stalk 0.025-0.06 mm, the upper portion 0.2-0.6 mm.

Young twigs (long shoots) terete, sparsely lanulose to glabrate, quickly becoming glabrous, smooth, red-

brown, prominently dotted with lenticels; thorns often present at nodes, especially on young stems, 3-20 mm

long, some becoming short spur shoots or lengthening to become side branches. Mature twigs glabrous, the

smooth outer surface becoming longitudinally fissured and fading to very light grey by the second season, be¬

coming rough and gnarled in subsequent seasons, transverse fissures appearing along with deepening longitu¬

dinal fissures, giving bark of older stems a somewhat rectangularly-plated reticulate appearance, the leaves

clustered on stumpy brachyblasts (short shoots), internode length 7-33 mm. Leaves with petiole 4-5 mm long

on young long shoots, 2-4 mm on short shoots, the petioles glabrous or sometimes with a wooly indumentum,

light brown and moderately dense to sparse on young petioles, becoming grey-white and sparse on older peti¬

oles; blade 29-50 mm long, 16-21 mm wide, rhombic to elliptic to obovate on young long shoots, 8-52 mm

Corogin and Judd, Sideroxylon redinatum ssp. austrofloridense

411

Fig. 6. Distribution of Sideroxylon redinatum subsp. austrofloridense (circled dots) and 5. redinatum subsp. redinatum (squares). The range of 5. redinatum

subsp. austrofloridense is centered on the Miami rock ridge, with outliers historically as far south as Key Largo and Flamingo, and more recently as far

west and north as Big Cypress National Preserve. Map shows vouchered occurrences only.

long, 5-19 mm wide, narrowly obovate to elliptic (on short shoots), coriaceous, the apex rounded to emargin-

ate, the base cuneate to acute, the margin entire, blades of mature leaves often markedly involute; venation

pinnate and conspicuously finely reticulate, brochidodromous, with secondary veins slightly decurrent,

branching off midvein in opposite to alternate arrangement, more or less in 6-10 irregularly spaced pairs, with

an intersecondary vein occasionally occurring between secondaries, the tertiary and quaternary veins irregu¬

lar-reticulate, the quinternary veins irregular-reticulate to freely ramifying, areolation moderately developed,

the marginal ultimate venation looped; adaxial surface dark green, semi-glossy, sparsely pubescent when

young with fine trichomes, becoming glabrous, the veins impressed (fresh leaf) to conspicuously raised (dried

leaf), giving the surface a finely textured appearance; abaxial surface glabrous or often sparsely to densely

covered with a variably persistent finely wooly indumentum, on young leaves light brown, sparse to moder¬

ately dense and matted, often at least partially obscuring the leaf surface, on older leaves fading to grey-white

and becoming less dense to glabrate, the trichomes T-shaped, with stalk 0.02-0.08 mm long, upper portion

0.3-1.5 mm long, the upper portions mostly curled and ascending, abaxial epidermal cell boundaries evident

under high magnification and marked by impressed grooves, the veins flush to slightly raised (fresh leaf) to

raised (dried leaf). Inflorescence a fascicle borne in a leaf axil, these often clustered on short shoots in axils of

fallen leaves, each cluster having 3-30 pedicellate flowers, the pedicels 2.5-11.8 mm long, glabrous to densely

wooly-strigose at anthesis, often becoming glabrous in fruit. Calyx of 5 (-6) quincuncial, free sepals, the outer

two sepals 1.8-2.0 mm long, 1.5-1.7 mm wide, ovate to suborbicular, the apex rounded, the margin entire,

412

Journal of the Botanical Research Institute of Texas 8(2)

opaque, the inner three sepals 1.8-2.2 mm long, 1.7-2.4 mm wide, orbicular, the apex rounded, the margin

entire to erose with broad zone of thin translucent tissue, all sepals glabrous to moderately densely pubescent

with light brown to red-brown T-shaped trichomes, the pubescence, if present, wooly on the outer, sericeous

on the inner. Corolla white, cyathiform, sympetalous, the tube 1.5-1.8 mm long, enclosed within the calyx, the

lobes 5 (-6), exserted and spreading, glabrous, each with a median lobe and two lateral segments, the median

lobe 1.5-2.0 mm long, 1.5-2.0 mm wide, orbicular, clawed, cupped around a stamen, the lateral segments

1.8-2.2 mm long (from junction with tube), 1.5-1.8 mm long (from junction with median lobe), 0.7-1.0 mm

wide, lanceolate, margins erose, translucent. Stamens 5 (-6), epipetalous, opposite the corolla lobes, exserted;

filament 1.4-1.8 mm long, 0.4 mm wide, fixed at top of corolla tube, dilated proximally and narrowing toward

anther attachment; anther 1.0-1.1 mm long, sagittate, ventrihxed at the point of the sinus, extrorse, opening by

two longitudinal slits. Staminodes petaloid, alternating with stamens, each staminode 2.2 mm long, 1.5 mm

wide, deltoid to ovate, margin erose, translucent. Ovary superior, 5-(6-) loculate, locules uniovulate with

basal-axile placentation, ovary ca. 1.0-1.3 mm long, 1.0 mm wide, globose to ovoid, tapering abruptly to 1 mm-

long style, the ovary at anthesis glabrous to densely pubescent with a sericeous indumentum of straight, ap-

pressed, white to slightly tawny T-shaped trichomes, the trichomes longer on distal part of ovary. Fruit a berry,

one (rarely 2) - seeded, 6.3-11.0 mm long, 6-10 mm wide, ellipsoid to slightly obovoid to subglobose, black,

glossy, glabrous, borne on pedicels 2.5-11.8 mm long. Seeds 4.2-70 mm long, 3.6-6.0 mm wide, subglobose to

ellipsoid, the testa hard, smooth, glossy, dark to light brown, the color solid to boldly mottled; the hilum scar

basal, usually bipartite, the larger part basal, oval to half-moon shaped, 0.8-2.2 mm long, 1.2-2.8 mm wide,

with a smaller, laterally elongate part abaxial to the other part, 0.8-2.8 mm long, 0.2-1.2 mm wide, the two

parts well-separated by a bridge of testa, the scar being the exposed portion of a hollow cavity under the testa

extending a small distance around the basal end of the seed, the cavity filled, especially on the abaxial side,

with a yellowish fatty substance.

Distribution and habitat.—Sideroxylon reclinatum subsp. austrofloridense is found only in Miami-Dade,

Monroe, and Collier Counties, Florida, where it is restricted to pine rocklands, marl prairies, and edges of

tropical rockland hardwood hammocks (Fig. 6). The range of this taxon is centered in Miami-Dade County on

the Miami rock ridge, which extends from Long Pine Key in the Everglades northward through urban Miami

to the Miami River. It is locally abundant at Long Pine Key, and is found at Pine Island, and around the edges of

Royal Palm Hammock and Paradise Key, in Everglades National Park. It has been collected in several extant

rockland fragments in the urbanized areas of the Miami rock ridge as far north as the Miami City Cemetery

(1944, A. Korsakoff s.n.). Historically, it was collected as far south as Key Largo (1948, C.R. Jackson s.n .) and

Flamingo (1924, J.K. Small 11537) in Monroe County. More recently it has been collected farther west and north

at locations in Big Cypress National Preserve: Monument Lake in Collier County (2014, Sadie 630; 1998, Brad¬

ley 1547), and the Loop Road area (1964, Godfrey 63520) and Lostman’s Pines area (2003, W oodmansee 1121) in

Monroe County (Hodges & Bradley 2006; Gann et al. 2014). There are unvouchered reports of additional oc¬

currences on the Miami rock ridge and in Big Cypress National Preserve (Gann et al. 2014). All locations where

Sideroxylon reclinatum subsp. austrofloridense has been collected have limestone bedrock either exposed at the

surface or covered with only a thin layer of marl-based soil. These soils become more sandy toward the north¬

western limits of the range, where S. reclinatum subsp. reclinatum is also found.

Phenology. —Flowering April-May, fruit ripening in June-July.

Additional specimens examined: USA. Florida. Collier Co.: Big Cypress National Preserve, Monument Lake Campground, 22 Mar 1998,

Bradley 1547 (FTG); Big Cypress National Preserve, Monument Lake Campground, marl prairie E of rd. to campground and N of radio

tower service rd., UTM 17N 488611E, 2860886N, 29 May 2014, Sadie 630 (FNPS). Miami-Dade Co.: Everglades National Park, Long Pine

Key, 2.5 mi W of campground T5,7S, R37E, SWA Sec 32,11 May 1999, Anderson 18620 (FSU); Everglades National Park, Long Pine Key near

Mosier Hammock, 11 May 1999, Anderson 18621,18622 (FSU); Everglades National Park, bordering Pine Island Rd., 0.3 mi S of main road,

T58S, R37E, SWA Sec 12,11 May 1999, Anderson 18623 (FSU); Frog Pond, just W of C-l 11 canal, 3.5 miles N of Rt 27, ca. 4 mi W of Florida

City, 11 May 1999, Anderson 18625, (FSU); Lucille Hammock, 0.9 mi W of 217th Ave., ca. 3 mi W of Florida City, T57S, R38E, SWA Sec. 29,

11 May 1999 , Anderson 18629 (FSU); Pine Ridge Sanctuary, 21100.SW 300 St., ca. 2 air mi NW of Homestead, 11 May 1999, Anderson 18631

(FSU); Everglades National Park, Long Pine Key, May 1908, E.A. Bessey 75 (A); Everglades, Paradise Key, 5 May 1908, E.A. Bessey 2 (G); Frog

Pond, 25°26.837'N, 080°33.757'W, 6 Aug 1997, Bradley & Woodmansee 422 (FTG); NW of Royal Palm Park, 3 May 1938, Buswell s.n. (FTG);

Corogin and Judd, Sideroxylon redinatum ssp. austrofloridense

413

Everglades National Park, Long Pine Key, N and W of campground, 26 Jul 2011, 25.40341°N, 080.65797°W, Corogin 1045 (FNPS); ibid.,

25.40335°N, 080.65838°W, Corogin 1046 (FNPS); ibid., 25.40244°N, 080.66216°W, Corogin 1047 (FNPS); ibid., 25.40138°N, 080.66822°W,

Corogin 1048 (FNPS); ibid., 25.40172°N, 080.66997°W, Corogin 1049 (FNPS); ibid., 27 Jul 2011, 25.40463°N, 080.67649°W, Corogin 1050

(FNPS); ibid., 25.40342°N, 080.67348°W, Corogin 1051 (FNPS); ibid., 28 Jul 2011, 25.40359°N, 080.66031°W, Corogin 1052 (FNPS); ibid.,

25.40442°N, 080.65965°W, Corogin 1053 (FNPS); ibid., 25.40061°N, 080.66123°W, Corogin 1054 (FNPS); ibid., 25,40061°N, 080.66103°W,

Corogin 1055 (FNPS); ibid., 25.40096°N, 080.66393°W, Corogin 1056 (FNPS); ibid., 11 May 2013, 25°24 , 5.6"N, 080°39 , 35.9"W, Corogin 1273

(FNPS); ibid., 25°24 , 6.2"N, 080°39 , 38"W, Corogin 1274 (FNPS); ibid., 25°24 , 7.4"N, 080°39 , 42.9"W, Corogin 1275 (FNPS); ibid., 25°24 , 6.2"N,

080°40 , 11.9"W, Corogin 1276 (FNPS); ibid., 26 Jul 2013, 25°24 , 6.2"N, 080°40 , 11.9"W, Corogin 1286 (FNPS); Everglades National Park, Long

Pine Key, 9 Jul 1961, Craighead s.n. (USF); Everglades National Park, Long Pine Key near Redd Hammock, 22 Dec 1958, Craighead s.n.

(FLAS); West Avocado Drive, 15 Feb 1955, F.C.C. 352 (FTG); near Naranja, 21 Jan 1937, Fennell 301 (A); Everglades National Park, Royal

Palm Hammock, 6 May 1928, Fisher 54 (US); Everglades National Park, Long Pine Key, 18 Apr 1964, Godfrey 63460 (FSU); Everglades Na¬

tional Park, 3.2 mi N of Rock Reef Pass, 17 Apr 1964, Godfrey 63394 (FSU); Everglades National Park, Long Pine Key, 4 May 1979, Godfrey

77025 (FSU); Everglades, N end Paradise Key, 29 Mar 1909, R.M. Harper 110 (G); Everglades National Park, pinelands auto trail,1.5 mi W of

Long Pine Key rd., 21 Jun 1975, Hill & Harvey 3215 (FTG); Miami City Cemetery, 30 Mar 1944, Korsakoff s.n. (FLAS); Larry & Penny Thomp¬

son Park, 25.59887°N, 080.40135°W, 26 May 2004, Possley 67 (FTG); Quail Roost Pineland, 25.57630°N, 080.42754°W, 26 Apr 2011, Possley

109 (FTG); Sunny Palms Pineland, 25.42835°N, 080.52053°W, 26 May 2011, Possley 110 (FTG); Everglades National Park, Paradise Key, 3

May 1920, Rehder 898 (A); Everglades National Park, marl prairie near edge of Royal Palm Road, N of Royal Palm Hammock, UTM 17N

538844E, 2808076N, 28 May 2014, Sadie 627, 628 (FNPS); Everglades National Park, marl prairie adjacent to wet pine rocklands at western

edge of Pine Island, just N of Main Park Rd., UTM 17N 541041E, 2808752N, 28 May 2014, Sadie 629 (FNPS); Everglades National Park,

Royal Palm Hammock/Paradise Key and vicinity, 21 Sep 1917, Safford &Mosier 22 (US); Everglades, S of Long Key, 18 Jan 1909, J.K. Small &

J.J. Carter s.n. (FTG); Hammocks, Homestead to Big Hammock Prairie, 15 Feb 1911, J.K. Small & J.J. Carter s.n. (FTG); Everglades, Camp

Jackson to Camp Longview, 21 Feb 1911, J.K. Small & J.J. Carter s.n. (FTG); Everglades National Park, Long Pine Key, 6 May 1904, J.K. Small

& P. Wilson s.n. (US); Everglades National Park, Long Pine Key, 5 Apr 1968, Sullivan s.n. (FTG); Everglades National Park, Long Pine Key near

picnic ground, 8 Apr 1971, Fomlinson & Avery s.n. (FTG); Everglades National Park, 4.6 mi S of park entrance on main rd. to Flamingo, 23

Apr 1958, Fraverse 592 (G); Everglades, Long Pine Key, 5 Jul 1950, Woodbury s.n. (FTG). Monroe Co.: Near Fla. Rt 94, 3.8 mi W of Dade-

Monroe County line, 19 Apr 1964, Godfrey 63520 (FSU); Key Largo, 27 Feb 1948 Jackson s.n. (FTG); Hammock back of Flamingo, Cape Sable

region, 21 Jul 1924, J.K. Small et al. s.n.(G); Big Cypress National Preserve, Lostmans Pines area, ca. 11.13 km S of Loop Rd. (State Rt 94), 17.9

km W of the Miami-Dade County line, just N of border with Everglades National Park, 25.66063° N, 081.04996° W, 26 Mar 2003, Woodma-

nsee 1121 (FNPS).

Endemism in South Florida rockland habitats. —The southern tip of peninsular Florida, including the Florida

Keys, is one of the state’s hotspots of rare and endemic plant taxa (Ward 1979; Chaplin et al. 2000). Unusual for

the continental United States, the flora of this region is dominated by tropical species of West Indian origin.

The uniqueness of Sideroxylon reclinatum subsp. austrofloridense is enhanced by the fact that it is narrowly en¬

demic to the pyrogenic pine rockland habitat of this area. Featuring an overstory of south Florida slash pine

(Pinus elliottii Engelm. var. densa Tittle & Dorman), these rocklands, dotted with small tropical hardwood

hammocks and surrounded by marl prairies and marshes, thrive on exposed karstic limestone bedrock only

partially covered with the thinnest of soils, at an elevation of only 1-5 m above sea level (Snyder et al. 1990).

The understory features a diverse mix of tropical and temperate shrubs, palms, and herbs, with a large number

of endemic taxa present in the species-rich herbaceous layer (Toope & Avery 1979; Avery & Toope 1980; Ol¬

msted et al. 1983; Snyder et al. 1990). South Florida pine rocklands share functional, structural, and floristic

similarities with tropical rocklands of the Bahamas, Cuba, Hispaniola, and Central America, which are also

pyrogenic communities on exposed limestone with a pine overstory, highly diverse understories, and high

endemism (Judd 1987; O’Brien et al. 2008; Myers & Rodriguez-Trejo 2009). Although pine rocklands exist in

the lower Florida keys, Sideroxylon reclinatum subsp. austrofloridense has never been collected there (Hodges &

Bradley 2006). Its known current range is limited to extant rocklands on the mainland, although an early col¬

lection from northern Key Fargo exists (1948, C.R. Jackson s.n., Fig. 6).

Around thirty plant taxa endemic to south Florida are found in pine rocklands; around a dozen of these

are restricted to pine rockland (Avery & Toope 1980; Snyder et al. 1990; Bradley & Gann 1999; FNAI 2010;

Gann et al. 2014). Many of these endemics are listed as endangered or threatened at the state or federal level,

and several are candidates for listing (Gann et al. 2014). Endemic associates of Sideroxylon reclinatum subsp.

austrofloridense at Tong Pine Key include Chamaesyce deltoidea (Engelm. ex Chapm.) Small subsp. pinetorum

(Small) A. Herndon, C. porteriana Small, Euphorbia pinetorum (Small) G.T. Webster, Jacquemontia curtisii Peter

ex Small, Eantana depressa Small var. depressa (Sanders 1987), Melanthera parvifolia Small, Phyllanthus pen-

414

Journal of the Botanical Research Institute of Texas 8(2)

taphyllus C. Wright ex Griseb. var. floridanus G.L. Webster, Schizcichyrium rhizomatum (Swallen) Gould, and

Tragia saxicola Small.

The shrub layer of pine rocklands features more than ninety taxa, mostly of West Indian affinity (Olmsted

et al. 1983). Woody associates of Sideroxylon reclinatum subsp. austrofloridense at Long Pine Key include two

species of Sideroxylon, S. celastrinum (Kunth) T.D.Penn. and S. salicifolium (L.) Lam; both are tropical, ranging

from southern Florida into the Caribbean, Mexico, and Central America. Sideroxylon reclinatum subsp. reclina¬

tum has not been collected in Everglades National Park, but it has been collected in Monroe County at Big Cy¬

press National Preserve (Gann et al. 2014), where the ranges of the two subspecies come into contact.

Other woody understory associates include Ardisia escallonioides Schiede & Deppe ex Schltdl. & Cham.,

Bourreria cassinifolia (A.Rich.) Griseb., Byrsonima lucida (Mill.) DC., Chiococca alba (L.) A.S. Hitchc., Chryso-

balanus icaco L., Chrysophyllum oliviforme L., Coccothrinax argentata (Jacq.) L.H. Bailey, Conocarpus erectus L.,

Croton linearis Jacq., Dodonaea viscosa Jacq., Forestiera segregata (Jacq.) Krug & Urb., Guapira discolor (Spreng.)

Little, Guettarda elliptica Sw., G. scabra (L.) Vent., Ilex cassine L., Metopium toxiferum (L.) Krug & Urb., Morinda

royoc L., Mosiera longipes (O. Berg) Small, Myrica cerifera L., Myrsine cubana A. DC., Randia aculeata L., Rhus

copallinum L., Tetrazygia bicolor (Mill.) Cogn., and Vachellia farnesiana (L.)Wright & Arn. var. pinetorum

(F.J.Herm.) Seigler & Ebinger.

Conservation status.—Sideroxylon reclinatum subsp. austrofloridense is a candidate for federal listing under

the Endangered Species Act, under the common name “Everglades Bully” (USFWS 2013a). Although the main

population of S. reclinatum subsp. austrofloridense is locally abundant and located on protected land, the U.S.

Fish and Wildlife Service considers that a change in listing status is warranted but precluded by higher priori¬

ties (USFWS 2013b; Sadie, pers. comm.). The Institute for Regional Conservation in Miami considers Sideroxy¬

lon reclinatum subsp. austrofloridense to be imperiled in south Florida (Gann et al. 2002, 2014). The pine rock-

land plant community is ranked critically imperiled both globally and in Florida by the Florida Natural Areas

Inventory (FNAI2010). The pine rocklands of mainland south Florida once extended from Long Pine Key, the

principal upland of Everglades National Park, northeastward along the Miami rock ridge as far as the Miami

River, but urban and agricultural development has extirpated the native plant cover from all but Long Pine Key

and a handful of small patches on private land or preserved as county parks (Olmsted et al. 1983; Bradley &

Martin 2012). The chief immediate threats to Sideroxylon reclinatum subsp. austrofloridense and its habitat are

fire suppression and exotic species invasion. Recommended conservation actions currently consist of habitat

protection and maintenance by prescribed fire, control of invasive non-native species, efforts to acquire, moni¬

tor and protect remaining rockland fragments, and continued monitoring and study to watch for adverse ef¬

fects that may arise from human activities such as water management policies and the Everglades restoration

program (Gann et al. 2002; FNAI 2010; USFWS 2013b).

Long-term sea level rise due to climate change also threatens the rockland flora. Discovery of 8600-year-

old submerged remains of pine trees 60 km west of Key West suggests that since the last glacial maximum (ca.

18,000 years before present), south Florida pinelands have been in retreat as rising seas have inundated a once

more extensive range; during the past century that retreat has continued measurably in the Florida keys (Ross

et al. 1994, 2009). As sea level rises, outright inundation is preceded by saltwater intrusion, which is currently

producing changes to the species composition of vulnerable south Florida coastal habitats (Saha et al. 2011).

Sea level is predicted to rise 1-2 m by 2100 (URS Corporation et al. 2007; Saha et al. 2011), and elevation at Long

Pine Key is 2 m or less. Sideroxylon reclinatum subsp. austrofloridense and its rare and endemic associates

clearly face an uncertain future.

APPENDIX

Additional specimens examined for SEM study, and for the south Florida distribution map (Fig. 6). Additional specimens of these taxa

from their entire range were also examined and will be cited in an upcoming taxonomic revision.

Sideroxylon reclinatum Michx subsp. reclinatum.—USA. 2 mi E of Collier-Seminole State Park, by US 41,1 May 1979, Godfrey

Florida. Broward Co.: Cypress Creek Hammock near Margate, 28 77002 (FSU); Cross State Turnpike, Miles City, W of FL 29, 27 May

Apr 1977, Correll 48485 (FTG); Fern Forest Nature Center, Coconut 1966, Lakela & DeBoer 29857 (USF); Deep Lake, 3 mi N of Jerome, 1

Creek,T59S, R42E, Secs. %, 8 May 2001, Howell389 (USF). Collier Co.: mi E of FL 29,21 Aug 1965, Lakela & Laker29124 (USF); Fakahatchee

Corogin and Judd, Sideroxylon redinatum ssp. austrofloridense

Strand State Preserve, 0.3 mi N of Gate 1, W of James Scenic Drive,

22 May 1999, Pires FS0356 (USF); Fakahatchee, margin of hammock,

6 Mar 1965, Lakela & Long 28181 (USF); off rd. to Bird Sanctuary,

Corkscrew, 7 Mar 1965, Lokelo & Long 28215 (USF); Kissimmee Billy,

N of Alligator Alley, 28 Apr 1976, Correll 47094 (FTG). Flagler Co.:

N side of Fla. 305,5.7 mi E of Volusia Co. line, S32, T13S, R29E, 7 Jul

1965, Ward4725 (FLAS). Gadsden Co.: Banks of Apalachicola River

at Aspalaga, 7 Jul 1955, Godfrey53598 (G); by County Rd. 269,0.5 mi

from railroad crossing at River Junction, 1 Jul 1991, Godfrey 84198

(TENN); Chattahoochee, bluff along Apalachicola River overlooking

ClydeT. Hopkins Municipal Park on River Landing Rd., 8 May 1995,

MacDonald 8652 (TENN). Gilchrist Co.: Banks of Suwannee River,

near public boat ramp W of Otter Springs, 5 Jun 2012, Corogin 1060

(FLAS); NW of Trenton, ca. 1 mi W of dead-end of County Rd. 344,

at Suwannee River, ca. 0.5 mi W of Hart Springs Park, 6 Sep 2003,

Judd8047 (FTG). Hendry Co.: Big Cypress Reservation, 8 May 1959,

Sturtevant 146 (US). Lee Co.: Fort Myers, 26 Apr 1934, Buswell s.n.

(FTG). Monroe Co.: Big Cypress National Preserve, near Pine Crest,

along Loop Rd. 94, 28 Apr 1974, Correll 42244 (FTG). Palm Beach

Co.: Yamato Scrub Natural Area, ca. 0.25 mi E of Congress Ave., N

of Clint Moore Rd., 26°24.58'N, 80°05.944'W, 12 May 1998, Bradley

& Woodmansee 977 (FTG). Taylor Co.: Near Steinhatchee River, by

US Rt 98/19, NW of Cross City, 19 May 1984, Godfrey 81312 (FLAS).

415

Sideroxylon macrocarpum (Nutt.) J.R. Allison.— USA.

Georgia. Jeff Davis Co: powerline right of way immediately W

of Appling Co. line and 0.6 mi from Perry Miller Rd., 31 0 52.85'N,

82°26.75'W, 26 May 1999, Nelson 20576 (GA). Laurens Co.: ca. 19.2

air mi SE of Dublin, ca. 9.2 air mi ENE of Cedar Grove, W side of GA

Hwy 19, ca. 0.3 road mi S of Buck Creek, 21 Aug 1996, Allison 9404

(FSU). Pierce Co.: ca. 2 mi NE of Blackshear, N of Shiloh Church,

29 Sep 1999, Allison 12228 (GA). Tattnall Co.: Gordonia Altamaha

State Park, along rd. S of golf course, 32°05 , 08"N, 082°08 , 03.8"W,

24 Sep 2012, Corogin 1179 (FLAS). Wheeler Co.: ca. 11.3 linear mi

NNE of Alamo, S side Oconee River, % mi E of GA Hwy 46, 26 Jul

1996, Allison 9356 (GA).

Sideroxylon rufohirtum Herring & Judd.— USA. Florida.

Alachua Co.: Near Kanapaha Prairie southwest of Gainesville,

29.572°N, 082.426°W, 19 Jun 2009, CoroginlOOO (FLAS). Dixie Co.:

Lower Suwannee National Wildlife Refuge, Yellow Jacket Loop Rd.,

7 May 2004, Barichivich s.n. (FLAS). Lake Co.: Roadside on Hwy 19

about 200 yds S of jet with old 441, Tavares, 20 Sep 1971, Wofford

71-558-A (TENN). Putnam Co.: 75 yds WNW of intersection of trails

#10 and #12, 2000 ft E of Camp Area water tower, Univ. of Florida

Conservation Reserve, Welaka,T12S, R26E, 27 Mar 1959, Ward 1206

(FLAS). Suwannee Co.: T5S, R14E, SI 9, ca. 5 mi NW of Ichetucknee

Springs, ca. 1.5 mi NNE of Beachville, Oct 1981, Simons s.n. (FLAS).

ACKNOWLEDGMENTS

We wish to thank the following herbaria for loans of specimens: A, AUA, FNPS, FSU, FTG, GA, GH, LSU, MISS,

OKL, TENN, US, USCH, USF, and UNA. We thank the following people without whose help this study could

not have succeeded: Kent Perkins of the University of Florida Herbarium (FLAS) for his generous assistance

with processing specimen loans; Brett Jestrow of Fairchild Tropical Garden (FTG) and Jennifer Stafford of the

National Park Service herbarium (FNPS) for facilitating our examination of south Florida herbarium speci¬

mens; Jimi Sadie, Everglades National Park botanist, for assisting with held work; Karen Kelley, electron mi¬

croscopy manager, and Kim Backer-Kelley, lab technician, of the ICBR Electron Microscopy Core lab at the

University of Florida, for their kindness and expertise; and we thank Dr. Ulf Swenson and an anonymous re¬

viewer for their helpful feedback on this manuscript.

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BOOK REVIEW

Judy Barrett. 2013. Yes, You Can Grow Roses. (ISBN-13: 978-1-62349-027-0, flexbound). Texas A&M Uni¬

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JUDY BARRETT is the author of W hat Can I Do with My Herbs? How to Grow, Use, and Enjoy These Versatile

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J.Bot. Res. Inst. Texas 8(2): 418.2014

A MORPHOLOGICALLY BASED TAXONOMIC REEVALUATION

OF THE GENUS STIPULICIDA (CARYOPHYLLACEAE),

WITH COMMENTS ON RANK

Derick B. Poindexter 1 , Kateland E. Bennett, and Alan S. Weakley

UNC Herbarium (NCU)

North Carolina Botanical Garden

University of North Carolina

Chapel Hill, North Carolina 27599, USA.

and

1 Botanical Research Institute of Texas

1700 University Drive

Fort Worth, Texas 76107, US.A.

poindexter@unc.edu

ABSTRACT

Stipulicida is a narrowly endemic, monotypic genus as currently circumscribed. Previous studies have attempted to diagnose additional taxa

within the genus, but only one species comprised of two varieties is currently accepted. These two varieties, Stipulicida setacea var. setacea

and S. setacea var. lacerata, demonstrate significant phenotypic divergence and idiosyncratic biogeographical patterns. We utilized multi¬

variate analyses and nonparametric analyses of variance to ascertain morphological distinctions between these two taxa and assess bio¬

logically appropriate taxonomic rank. Based on our morphometric data and biogeography, we elevate S. setacea var. lacerata to species and

provide a revised key to the species in the genus.

RESUMEN

Stipulicida es un pequeno genero endemico, monotipico tal como se circunscribe normalmente. Estudios previos habian intentado diag-

nosticar taxa adicionales en este genero, pero solo se acepta normalmente una especie con dos variedades. Estas dos variedades, Stipulicida

setacea var. setacea y S. setacea var. lacerata, muestran una divergencia fenotipica y patrones biogeograficos significativos. Utilizamos un

analisis multivariante y un analisis no-parametrico de varianza para establecer diferencias morfologicas entre estos dos taxa y valorar el

rango taxonomico apropiado biologicamente. Basados en nuestros datos morfometricos y biogeografia, elevamos S. setacea var. lacerata a

especie y proporcionamos una clave revisada de especies del genero.

INTRODUCTION

The monotypic genus Stipulicida Michx. (Caryophyllaceae; Paronychioideae; Polycarpeae) is endemic to the

southeastern United States and Cuba (Swanson & Rabeler 2005; Weakley 2012; Kartesz 2013). It consists of

minimalist plants with slender stems, ephemeral basal rosettes, short awl-shaped opposite leaves with incised

stipules, and diffuse, divaricately-branched inflorescences. The flowers are the most conspicuous morphologi¬

cal feature, consisting of 5 dimorphic sepals (often referred to as “inner” and “outer”), 5 white petals, with the

terminal clusters subtended by bracts similar to the leaves. The genus was first described by Andre Michaux

(1803) from collections made “in sabulosis aridis Carolinae ” (= sandy, dry soils of the Carolinas) (Fig. 1). Stipu¬

licida as a genus has remained unscathed by current phylogenetic and systematic studies of the Caryophylla¬

ceae, perhaps only because of its omission from them. In the most recent molecular phylogenetic studies of the

Caryophyllaceae (e.g., Fior et al. 2006; Harbaugh et al. 2010; Greenberg & Donoghue 2011), Stipulicida has

been one of the few genera omitted. Consequently, exact subfamilial and tribal placement has not been con¬

firmed and current circumscription is based solely on morphology. At the time of its description, the genus

was represented by a single species, Stipulicida setacea Michx.

Nash (1895) added another putative species from central peninsular Florida, Stipulicida filiformis Nash,

which allegedly differed from S. setacea based on a much more slender habit, fewer, sessile flowers, and shorter

bracts. Small (1903, 1913, 1933) maintained this taxon as distinct within his influential manuals and added

J. Bot. Res. Inst. Texas 8(2): 419 - 430.2014

420

Journal of the Botanical Research Institute of Texas 8(2)

Fig. 1. One of two presumed isotypes of Stipulicida setacea ( Michauxs.n ., P, image #P00156962).

Poindexter et al.. Reevaluation of Stipulicida (CaryophyIlaceae)

421

petal shape (constricted at the middle vs. spathulate) as an additional discriminating feature. Judd (1983)

sought to address the taxonomic status of S.filiformis by conducting extensive fieldwork and observing varia¬

tion in situ, in addition to morphometric analyses from herbarium specimens. He focused on two quantitative

features, stem diameter and number of flowers/inflorescence. Judd (1983) concluded that the plants intergrade

broadly with respect to these characters, while also stating that bract size and petal shape failed to provide

separation into two taxa, and consequently recognized S. filiformis as a mere morphological extreme or eco¬

type of S. setacea, though commenting that its apparent limitation to the Central Florida Ridge was noteworthy

due to the number of taxa that are distributed similarly as endemics of xerophytic sands of this region. Ward

(2001) felt this pattern (in combination with intergrading morphological distinctions) was sufficient to war¬

rant maintaining it at varietal rank, Stipulicida setacea var. filiformis (Nash) D.B. Ward, in order to preserve

some taxonomic distinction.

James (1957) described the only additional member of this genus to date by recognizing a new variety,

Stipulicida setacea var. lacerata C.W. James, restricted to Florida and the Isle of Pines (now the Isla de la Juven-

tud), Cuba. This variety was considered distinguishable from the typical variant by the presence of lacerate

sepals and outer sepals that were mucronate, as well as “essentially allopatric” distributions. James’ (1957) ra¬

tionale for infraspecific rank was explicit:

“The collection of F.W. Hunnewell (no. 8955, Highlands Co., Fla.), from the eastern limits of the range of var. lacerata was the only one seen

which consisted of plants of both varieties. The specimen of var. lacerata in this collection is somewhat atypical. This, in addition to the fact

that there is not a clear-cut discontinuity in the shape of the sepals, indicates that there may be some gene flow between these taxa.”

During the course of his study, Judd (1983) also noticed the distinctive nature of lacerate-sepaled plants and

felt they were adequately assigned to varietal status due to presence of intermediate plants from Highlands and

Volusia Counties.

At present, the general consensus (as shown by recent floras covering all or parts of the distribution of the

genus: Wunderlin & Hansen 2011; Swanson & Rabeler 2005; Weakley 2012) has been to follow the taxonomic

schema established by Judd (1983) by recognizing a single species within the genus, with two relatively well-

marked varieties, Stipulicida setacea v ar. setacea (incl. S.filiformis) andS. setacea var. lacerata. The distribution

of the genus as a whole is narrow, restricted to the Atlantic and Gulf Coastal Plain (Fig. 2). Of these two variet¬

ies, var. setacea is the most widespread, extending from southeastern Virginia to Florida and west to eastern

Louisiana. The second variety, var. lacerata , is much more narrowly distributed in only peninsular Florida, and

disjunct in Cuba (Isla de la Juventud) (Wunderlin & Hansen 2005; Weakley 2012).

Previous classification attempts have lacked a comprehensive, systematic approach in documenting key

morphological differences between these two entities and have failed to account for what we believe are sig¬

nificant characters, in addition to the seemingly conspicuous sepal morphology. Consequently, we chose to re¬

evaluate the distinctiveness of the two currently recognized varieties using a broader morphometric analysis.

METHODS

We examined selected specimens from across the entire distribution of the species (Fig. 2). Physical material

was examined from GH, NCU, and USF. The southernmost locality (Isla de la Juventud, Cuba) was examined

digitally ( Britton 14200, NY, image #NY01511627). We also studied the original type description and illustra¬

tion of Stipulicida setacea from Michaux (1803), as well as the putative isotypes from the Michaux Herbarium

at the Museum National d’Histoire Naturelle ( Michaux s.n., P, image #P00156961 and #P00156962). In addi¬

tion, we examined digital images of types of the other named taxa, including S.filiformis (Nash 14, GH, image

#GH00038011, MICH, image #MICH1111011, MIN, image #MIN1002897, NY, image #NY00353092, P, image

#P04925565 and #P04925568, US, image #US00103374 and #US00931431) and Stipulicida setacea var. lacerata

(Tracy 6828, GH, image #GH00038012, MSC, image #MSC0092943, NY, image #NY00353093, US, image

#US00103375). We selected 41 representative specimens of Stipulicida setacea (without discrimination be¬

tween var. setacea and var. filiformis) and 26 specimens of var. lacerata for morphometric analysis to assess

422

Journal of the Botanical Research Institute of Texas 8(2)

Fig. 2. Distribution ofStipulicidasetacea specimens examined for this study. Records that have not been physically examined but are from reliable sources

(Wunderlin & Hansen 2008; Kartesz 2013) are included. State-level occurrence is indicated by shading.

previously described features (see Nash 1895; James 1957; Judd 1983) and to seek additional diagnostic fea¬

tures. Specimens measured are denoted within the Appendix (below).

We examined nine continuous characters (Table 1). All characters were selected at random, with the ex¬

ception of the length of the longest mucro (LOM), which necessitated a comprehensive scan of each specimen.

In addition, we made general observations regarding the degree of laceration and fimbriate nature of speci¬

mens, as well as overall gestalt. All measurements were made with a TDI International, Inc. Micro Ruler.

To determine specimen groups and elucidate the most diagnostic variables, we used principal compo¬

nents analysis (PCA). To avoid weighting characters, we examined pairwise correlation coefficients (CC) that

Poindexter et al.. Reevaluation of Stipulicida (CaryophyIlaceae)

Table 1 . Morphological characters measured from herbarium specimens with

abbreviations in parentheses. Those characters used in multivariate analyses

are denoted by an asterisk.

1 Length of outer sepal (LOS)

2 Length of inner sepal (LIS)*

3 Width of outer sepal (WOS)*

4 Width of inner sepal (WIS)

5 Stem width below terminal flower cluster (SWF)*

6 Stem width below lowest branching node (SWN)*

7 Length of longest mucro of the outer sepal (LOM)*

8 Difference between inner and outer sepal length (DSL)

9 Difference between inner and outer sepal width (DSW)

ters in this analysis: LIS, LOM, SWF, SWN, and WOS.

We performed a Bartlett test for homogeneity of group variances on each variable. Variables that demon¬

strated variances that were significantly heterogeneous were log-transformed before further analysis. We test¬

ed each variable for deviations from normality using the Goodness-of-Fit test. Nearly all variables deviated

significantly from a normal distribution. We chose to maintain a conservative approach using the nonpara-

metric Kruskal-Wallis test and employed the Steel-Dwass method (i.e., a nonparametric version of the Tukey

Honestly Significantly Different test) to further detect the amount of differences between groups. Principal

components and nonparametric analyses were conducted using JMP® version 11 software (SAS Institute,

Cary, NC) on an Apple ® MacBook Pro™ computer.

RESULTS

Principal components analysis yielded strong resolution with Stipulicida setacea var. lacerata and var. setacea

clearly separating within the ordination space (Fig. 3). Principal component 1 was responsible for 57.3% of the

observed variation and principal component 2 was responsible for 16.7%, for a total of 74.0% of the variance

explained. An examination of specimens likely to be assigned to Stipulicida setacea var. fdiformis based on their

“wiry” habit and geographic distribution revealed that they were embedded within the var. setacea data cluster

(data not shown). Likewise, the Kruskal-Wallis test supported two distinct groups. Both taxa differed from

each other with respect to means of nearly all measured characters except LOS and LSW (Table 2).

DISCUSSION

The genus Stipulicida has recently been treated as monospecific and is currently represented by one species,

Stipulicida setacea, and two varieties (Swanson & Rabeler 2005; Weakley 2012): S. setacea var. setacea and S.

setacea var. lacerata. The paucity of distinctive morphological characters found within the genus have perhaps

led to a general acceptance of this current taxonomic treatment and a lack of interest in further study. This re¬

search sought to reevaluate the distinctiveness of these two entities through a more rigorous morphometric

approach that implemented multivariate analyses, in an attempt to address appropriate taxonomic ranks. We

selected several characters that had previously been emphasized by other authors, but also generated new

characters (e.g., outer and inner sepal length differences) that have aided in the further discrimination these

taxa. Despite considerable overlap, many diagnostic trends are apparent. Though both taxa have dimorphic

sepals, those of var. lacerata are distinctly lacerate/fimbriate and usually unequal in length (Fig. 4A-B). Con¬

versely, the sepals of var. setacea are mostly entire margined and often even in length (Fig. 4C-D). Stipulicida

setacea var. lacerata exhibits longer and wider inner sepals on average, and outer sepals that are generally wider

and with a more pronounced mucronate apex (which is usually obsolete in var. setacea). Variety lacerata also

appears generally stouter than var. setacea in all aspects as evidenced by significantly greater average stem

widths (Table 2).

A combination or suite of characters rather clearly defines these two taxa within ordination space (Fig. 3).

423

may indicate genetically linked characters and ex¬

cluded those that exceed a standard threshold of >

0.7. The only exception to this concerned the charac¬

ters LOM and WOS, which exhibited a strong correla¬

tion, but we believe that this correlation may be artifi¬

cial, as it is very unlikely that these two traits would

be due to allometric scaling. Thus, we have decided to

retain them both within our analyses. For PCA, we

conducted analyses on a correlation matrix, which

consisted of data that is standardized so that each

variable had a mean of 0 and standard deviation of 1.

We used only the following five continuous charac-

424

Journal of the Botanical Research Institute of Texas 8(2)

Table 2. Means ± 1 SD and ranges for morphological characters measured.

Character abbreviations correspond to those in Table 1. All measurements are

in millimeters. N equals the sample size. Within a row, means with different

superscripts differ significantly (Kruskal-Wallis, P < 0.05).

Character

S.setacea

(N=41)

S. lacerata

(N=26)

1. LOS

1.4 ± 0.2 a

1.5 ± 0.29 a

(0.9 -1.8)

(1.0-2.0)

2. LIS

1.5 ± 0.21 a

1.8 ± 0.32 b

(1.0-1.9)

(1.1 -2.2)

3.WOS

0.4 ± 0.1 a

0.8 ± 0.1 b

(0.2 - 0.7)

(0.6-1.1)

4. WIS

0.6 ± 0.1 a

1.0 ± 0.2 b

(0.4 - 0.9)

(0.7 -1.4)

5. SWF

0.3 ± 0.1 a

0.4 ± 0.1 b

(0.2 - 0.6)

(0.3 - 0.5)

6. SWN

0.5 ± 0.1 a

0.8 ± 0.1 b

(0.3 - 0.9)

7.LOM

0.0 ± 0.04 a

0.3 ± 0.1 b

(0.0- 0.1)

(0.2 - 0.5)

8. DSL

0.1 ±0.1 a

0.3 ± 0.2 b

(-0.2 - 0.5)

9. DSW

0.2 ± 0.1 a

(0.1 - 0.5)

(0.0 - 0.4)

As a secondary measure, we selected specimens that

were either originally determined as Stipulicida seta¬

cea var. filiformis (or S. filiformis) or were within the

originally described distribution and ecological pref¬

erence of this taxon. We mapped these specimens

within our PCA and found them to be centrally nest¬

ed within S. setacea var. setacea (data not shown),

which further supports Judd’s (1983) conclusion that

S. filiformis is not distinct from var. setacea. These

data, in conjunction with known biogeographic pat¬

terns, suggests that Stipulicida setacea var. lacerata

and S. setacea var. setacea are sufficiently distinct to be

recognized as discrete entities. However, previous

authors (James 1957; Judd 1983) have indicated that a

handful of specimens in areas of sympatry are inter¬

mediate. The one specimen (Highlands County, FL,

Hunnewell 8955, GH) cited byjames (1957) as a mixed

collection was included in our analyses, focusing on

the portion he described as “atypical” Stipulicida

lacerata and it was nested within the var. setacea data

cluster. Though we did not examine two of the addi¬

tional specimens mentioned as intermediate by Judd

(1983), we did examine one from Volusia County, FL

(D.S. & H.B. Correll 51934, USF). This specimen ap¬

peared to be a mixed collection with one branch that

was unassociated with a parent plant, but mounted on top of another such that it was not easily detectable as

separate. We measured both this stem, which based on our PCA was nested within var. lacerata, and another

full plant on the sheet separately, which was nested within var. setacea. Since we have not examined other puta¬

tive intermediates reported, we cannot attest to their identity.

We also recognize that some within taxon variation is apparent and could lead to perceived intermediacy.

For instance, some trends were observed that may not be readily apparent, e.g., northernmost populations of

var. setacea exhibit acute to slight retuse outer sepal apices, with progressively more retuse to strongly obcor-

date apices as one progresses further south. This pattern is also seen with northern populations possessing

inner and outer calyces that are narrow and subequal in length, while those of southern populations are wider

and often unequal, as well as shorter in comparison. Even the amount of lacerations/fimbriae of var. lacerata

can vary somewhat and an examination of an entire specimen is necessary to evaluate the plasticity in this

qualitative character state. James (1957) also remarked that the specimens from Cuba do not have as strongly

lacerate sepals as those from Florida, though we found this to be within the range of variation of this taxon.

How does this analysis translate to taxonomic decision-making? We find that (despite variation within

each) two morphologically distinctive entities (“lacerata” and u setacea s.s.”) are consistently separable and

identifiable using a suite of morphological characters. The several morphological characters are correlated

with one another, and do not randomly assort. An ordination confirms the more subjective assessment that

two morphologically distinct entities can be recognized. No true intermediates have been seen; and a few “pu¬

tative intermediates” cited by previous workers prove not to be, based on careful analysis. Neither are there

situations in which the correlation of the seemingly independent characters breaks down. When mapped,

these two morphologically distinct entities have distinctive and different distributions, with a substantial area

of overlap in peninsular and northeastern Florida. In areas of distributional overlap, intermediates are not

seen, although at least one studied collection is mixed. Each of the distribution patterns shown by the two enti-

Poindexter et al.. Reevaluation of Stipulicida (CaryophyIlaceae)

425

Principal Component 1

Fig. 3. Principal components analysis (PCA) of seven morphological characters (LOC, SWN, SWF, OSL, ISL, OSW, ISW) measured from Stipulicida lacerata

and 5. setacea.

ties is a distribution pattern represented by a large number of other angiosperm (and even animal) species: the

“se. Virginia to s. Florida, west to e. Louisiana” distribution of “setacea s.s.” and the “Florida peninsula” distri¬

bution (with optional disjunction into the West Indies) of “ lacerata ”—both commonly repeated distributions

across many organism groups and therefore likely to show “biogeographic signal” (see Sorrie & Weakley 2001,

2006).

If two taxonomic entities are warranted, what should their taxonomic rank be? Based on the analysis

presented above, we believe that these two entities represent independent evolutionary entities (by biological,

evolutionary, and/or phylogenetic criteria) at this time—species per most modern species concepts. A fre¬

quently used, though usually not explicitly stated, assumption suggests that the magnitude of morphological

difference is well-correlated with taxonomic rank—a paradigm that might be characterized as “if you can tell

two entities apart with readily observable, macroscopic characters, they are species, but if the characters re¬

quire magnification or are ‘fussy’, they are infrataxa (subspecies or varieties)”. Ward (2012), in a paper propos¬

ing the reduction of dozens of Southeastern United States species of vascular plants species to varietal status,

makes some of these points more explicitly, decrying the proliferation of “microspecies”. The following state¬

ment regarding Conradina etonia may be regarded as an example: “Details of indumentum, of leaf venation, of

size and pilosity, and of stamen pubescence, while wholly persuasive of the taxon’s genetic separateness, do not

rise to the level of difference to be found among related species” (Ward 2012). But modern biological and phy¬

logenetic species concepts are not based on the magnitude of morphological variation but on the degree to

which two entities are demonstrably separate biological/evolutionary entities—not determinable by the size of

their characters but by inductive reasoning (as in the previous paragraph). The consistency and evolutionary

meaningfulness of the characters is what tells the tale, not their size; small plants with small characters are not

less significant evolutionary entities.

426

Journal of the Botanical Research Institute of Texas 8(2)

Gill (2014) has also recently, and significantly, raised the issue of “burden of proof” in modern taxonomic

decision-making. When previous taxonomic studies have proposed a range of conclusions (splitting to lump¬

ing), what is the null hypothesis and on whom does the burden of proof lie; to disprove previous lumping, or to

disprove previous splitting? Gill (2014) argues that in bird studies, allopatric “populations” have almost always

been shown, after detailed genetic study to warrant taxonomic recognition at species rank, and that the burden

of proof should now be (based on this preponderant pattern) on those who favor “lumping”. Additionally, this

assumption is more conservationally helpful and parsimonious, in providing a more accurate hypothesis of

significant evolutionary entities requiring conservation attention. In much less well-studied groups (such as

vascular plants of the southeastern United States), patterns of previous studies are less clear, but generally also

show that recent rigorous and/or molecular-based studies show a greater general reliability of older morpho¬

logical studies (reflected in the treatments of Small, 1903, 1913, and 1913) than the casual lumping seen in

many late 20th century and early 21st century floras (e.g., Radford et al. 1968; Wunderlin & Hansen 2011). As

demonstrated by Weakley (2005), eastern North American floras from the 1960s to 1990s instituted a trend of

lumping, often without additional studies of the plants themselves either in the held or in herbaria. This has led

to a current misimpression that many “splits” (recognized, for instance by J.K. Small in the Southeast and Fer-

nald in the Northeast) have been disproven, when most have received little or no additional study since the first

half of the 20th century.

For Stipulicida, previous taxonomic studies have concluded that there are one, two, or three entities, and

at various ranks. We concede that we have not presented the ultimate and final analysis, based on an ideal

and complete (as of methodologies available in 2014) synthesis of molecular data, morphological character

analysis, and population biology studies. Such a study would require substantial effort, and with thousands

of such taxonomic problems remaining in the southeastern United States flora and few investigators and even

less funding, it is not our highest priority. We offer this analysis and set of taxonomic conclusions (resulting

in the recognition of two taxa, not one or three) as a current, yet disprovable hypothesis of the taxonomy of

Stipulicida.

We conclude that the best evidence available supports the recognition of two specifically distinct entities

within Stipulicida, and therefore here elevate var. lacerata to the rank of species and provide a revised identifica¬

tion key to members of the genus.

Stipulicida lacerata (C.W. James) D.B. Poind., K.E. Bennett, & Weakley, comb, etstat. nov. Stipulicida setacea var.

lacerata C.W. James; Rhodora 59:98. 1957. Type: UNITED STATES. Florida. Pinellas Co.: Dunedin, 14 Apr 1900, Tracy 6828 (holo-

type: GH!, image; isotypes: MSC!, image, NY!, image, US!, image).

KEY TO SPECIES OF STIPULICIDA

1. Sepal margins lacerate-fimbriate; outer sepals often much shorter than inner sepals (usually by 0.1-0.5 mm); tips of the

outer sepals with longest mucro (0.1 -)0.2-0.4(-0.5) mm; [of FL and Cuba (Isla de la Juventud)]_S. lacerata

1. Sepal margins entire or frayed (not lacerate-fimbriate); outer sepals subequal to slightly shorter or longer than inner

sepals (usually by 0.0-0.2 mm); tips of the outer sepals acute to obtuse or retuse-obcordate; with longest mucro or

apicule 0.0-0.1 mm; [of se. VA south to s. FL, west to e. LA]_S. setacea

Additional studies of the interesting, and seemingly isolated, genus Stipulicida are warranted. Population ge¬

netic and molecular studies could elucidate additional patterns of variation, and more definitively clarify the

status of the alleged “filiformis” entity and the widely disjunct population of S. lacerata in the Isla de la Juventud

(Cuba), as well as providing additional confirmation of our proposed species-level distinction of S. setacea and

S. lacerata. Additionally, the failure to include Stipulicida in recent molecular studies of the family leaves its

phylogenetic affinities and exact taxonomic placement within the Caryophyllaceae uncertain and conjectural.

Poindexter et al.. Reevaluation of Stipulicida (CaryophyIlaceae)

427

Fig. 4. Mature capsules, calyces and calyx lobes (inner lobes = left, outer lobes = right) of A-B) Stipulicida lacerata (Sarasota County, Florida, Perkins

s.n., NCU) and C-D) 5. setacea (Bamberg County, South Carolina, Ahles25982, NCU). Scale bar = 2 mm.

428

Journal of the Botanical Research Institute of Texas 8(2)

APPENDIX

List of representative specimens. Those utilized for morphometric analysis are denoted by an asterisk.

Stipulicida lacerata (C.W. James) D.B. Poind., K.E. Bennett, &

Weakley—U.S.A. FLORIDA. Charlotte Co.: NE of Pirate Harbor,

10 May 1990, Orzell 13438 (USF); Prairie/Shell Creek, 11 May 2009,

Fronck 1250 (USF*). Collier Co.: Immokalee city limits, 7 Apr 1968,

Lokelo 31361 (USF*); vicinity of Lake Trafford, 31 Jul 1968, Lokelo

31560 (USF*). De Soto Co.: W side of Peace River, 31 Jul 2008, Franck

771 (USF*). Flagler Co.: NE of US 1, 1 May 2006, Slaughter 14843

(USF*). Glades Co.: Fisheating Creek Wildlife Management Area,

30 May 2010, Franck2184 (USF*). Hardee Co.: AMAX property, 26

Mar 1979, Arcuri 856 (USF); Goose Pond Rd., 9 Apr 1981, Robinson

129 (USF*). Highlands Co.: S of Old Venus, 1 May 1958, Cooley 6137

(USF*); Archibold Biological Station, 24 Dec 1960, Craighead s.n.

(USF). Hillsborough Co.: Tarpon Springs, 23 Mar 1923, Churchill

s.n. (GH*); Little Manatee River, 21 Feb 1974, Shueys.n. (USF); Little

Manatee River State Park, 11 Apr 1999, Myers 342 (USF*). Lee Co.:

Myers, 1900, Hitchcock 5 (GH*); Fort Meyers, May 1928, Small s.n.

(NCU*); Koreshan State Park, US 41, 20 Apr 1964, Lakela 26983

(USF); Bonita Springs, 13 Feb 1965, Foster s.n. (USF); Koreshan State

Park, 1 mi S of Estero, 1 Mar 1969, McCart 10,630 (NCU*). Manatee

Co.: Wingate Creek State Park, 14 Mar 1991, Weber WC0046 (USF);

Lake Manatee State Recreation Area, 14 Apr 1992, Weber LM0039

(USF*). Martin Co.: Jonathan Dickinson State Park, 9 Feb 1969,

McCart 10,522 (NCU*); Hobe Mountain, Sep 1987, Woodbury 87

(USF*). Okeechobee Co.: Kissimmee Prairie, 27 Apr 1923, Small

10898 (GH*). Osceola Co.: Kempfer Ranch, 9 Mar 1976, Huck 1728

(NCU*). Pasco Co.: SE of Lacoochee,29Jun 1992, Orzelll9761 (USF);

Starkey Wilderness Park, 20 Mar 2004, Ferguson 659 (USF). Pinellas

Co.: NE of Clearwater, 3 Apr 1970, Lelong 5344 (NCU*); Highway

19, 3 Apr 1970, Rogers 2877-B (NCU*); Dunedin Hammock, 1 May

1977, Genelle 2728 (USF*); Magnolia Creek, 24 Mar 1988, Fleming

3981 (USF*). Sarasota Co.: Sarasota, 3 Jan 1943, Perkins s.n. (NCU*);

Oscar Scherer State Park, 27 Aug 1994, Cole 0S0257 (USF); Deep

Prairie Creek Preserve, 23 Aug 2007, Franck 332 (USF). St.Johns Co.:

Hastings, 11 Jun 1975, Godfrey 74329 (NCU*). Volusia Co.: Route

415,26 May 1981, Cornell51934-B (USF*);Tomoka State Park, 15 Mar

2006, Kunzer 1454 (USF*).

Stipulicida setacea Michx.—U.S.A. ALABAMA. Baldwin Co.: Point

Clear, 20 Mar 1883, Mohr s.n. (NCU); Orange Beach, 4 Jul 2012,

Spaulding 13613 (NCU). Henry Co.: NE of Abbeville, 8 May 1989,

Orzell 9538 (NCU). Mobile Co.: Little Dauphin Island, 15 Jul 1965,

Bray 442 (NCU); S of Audubon Bird Sanctuary, 15 Aug 1966, Bray

D995 (NCU); Dauphin Island, 15 May 1972, Lelong 6502 (NCU*).—

FLORIDA. Broward Co.: Deerfield Beach, 17 Jun 1978, Cornell49877

(NCU*); Franklin Co .: near Appalachicola, s.d., Curtiss exsiccata no.

336 (NCU*); S of US Route 98, 22 May 1971, Godfrey 70233 (NCU*).

Gadsden Co.: near Lake Talquin, 29 Apr 1977, Anderson 4430

(NCU*). Highlands Co.: Sebring, 7 Mar 1924, Hunnewell8955 (GH*);

S of Childs, 25 Apr 1960, Ray 9788 (NCU*); S of Sebring, 25 Apr 1960,

Ray 9731 (NCU*). Hillsborough Co.: W of Plant City, 8 Mar 1963,

Wall30 (NCU). Indian River Co .: near Sebastian, 3 Apr 1921, Small

9864 (NCU*). Lake Co.: Howey-in-the-Hills, 29 April 1960, Cooley

7352 (NCU*). Martin Co.: Hobe Sound, 22 May 1960, AtwaterM-214

(NCU*). Palm Beach Co.: W of Lantana, 5 Aug 1952, Howard 12959

(NCU*); West Palm Beach, 14 Feb 1969, Cassen 461 (NCU*). Seminole

Co.: S of Wagner, 3 May 1960, Cooley 7612 (NCU*). St. Lucie Co.: E

of US 1,28 Jul 1962, Lakela 25219 (NCU). Volusia Co.: Route 415,26

May 1981, Cornell 51934-A (USF*). Wakulla Co.: Mashes Island, 24

Apr 1955, Godfrey53207 (NCU*).— GEORGIA. Ben Hill Co.: NNE of

Fitzgerald, 4 May 1968, Faircloth 5189 (NCU*). Candler Co.: 32°16'N,

81°57'W, 3 May 1965, Cook 167 (NCU*); E of the Canoochee River,

13 May 1975, Walker 134 (NCU). Echols Co.: Mayday, 15 May 1969,

Faircloth 5831 (NCU*). Emanuel Co.: N of the Ohoopee River, 18

May 1976, Boufford 18438 (NCU*). Evans Co.: NW of Claxton, 9

May 1977, Oliver 96 (NCU). Hancock Co.: NE of Sparta, 3 Jun 1972,

Williams s.n. (NCU*). Laurens Co.: E of Dublin, 26 Jul 1967, Logue

2113 (NCU). Long Co.: near Beard's Bluff, 10 Apr 1963, Bozeman 2543

(NCU). Marion Co.: S of Geneva, 17 May 1974, Faircloth 7770 (NCU).

McDuffie Co.: S ofThomson, 29 Jul 1950, Duncan 11558 (NCU); N of

Thomson, 18 May 1952, Duncan 13568 (NCU). McIntosh Co.: NW of

Cox, 4 Jun 1962, Bozeman 710 (NCU). Richmond Co.: near August

Airport, 17 Jun 1950, Radford 5358 (NCU*); E of Briar Creek, 16 May

1958, Duke 614 (NCU). Tattnall Co.: NW of Reidsville, 11 Jun 1961,

Ahles 54183 (NCU*). Taylor Co.: W of Butler, 16 May 1974, Faircloth

7559 (NCU). Warren Co.: NE of Sparta, 3 Jun 1972, Williams s.n.

(NCU).— LOUISIANA. St. Tammany Co.: E of Slidell, 17 May 1983,

Thomas 83658 (NCU*). — MISSISSIPPI. Forrest Co.: Hattiesburg, 29

Jun 1971, Rogers 6585 (NCU*).— NORTH CAROLINA. Anson Co.: SE

of Lilesville, 30 Jun 1956, Radford 13452 (NCU). Bladen Co.: Sandy

places, 10 Jun 1898, Ashe s.n. (NCU); W of White Oak, 22 Jun 1957,

Ahles 29167 (NCU); S of Cumberland County line, 12 May 1971,

Leonard4784 (NCU). Brunswick Co.: Boiling Springs Lakes Preserve,

6 Jun 2006, Morris 052605-04 (NCU). Carteret Co.: E of Bogue, 21

Jun 1947, Wood 7014 (NCU); N of NC 24,15 May 1976, Snyder 814

(NCU); Patsy Pond, 22 May 1976, Wilson 1782 (NCU). Craven Co.: W

ofVanceboro, 19 Jul 1958, Radford37647 (NCU). Cumberland Co.:

S of Fayetteville on NC 53,11 May 1941, Radford 1063 (NCU); N of

Cedar Creek, 2 Aug 1954, Munson s.n. (NCU); SE of Spring Lake, 4

May 1957, Ahles 24290 (NCU); WSW of Cumberland, 27 Jun 1957,

Ahles 29729 (NCU). Duplin Co.: NE of Magnolia, 27 Apr 1957, Ahles

24047 (NCU*); N of Scotts Store, 15 Jun 1957, Ahles 28407 (NCU*).

Gates Co.: White Oak Pocosin, 30 May 1958, Duke 805 (NCU*);

near Chowan River, 10 Jun 1989, Musselman s.n. (NCU*). Harnett

Co.: SE of Spout Springs, 7 May 1946, Downs 13446 (NCU*); N of

Lillington, 11 May 1946, Radford 3047 (NCU*). Hoke Co.: SW of

Bowmore, 12 May 1957, Ahles 25089 (NCU); Ashley Heights, 26

Jun 1957, Ahles 29353 (NCU). Lee Co.: N of Moore County line,

24 May 1940, Radford 137 (NCU); SE of St. Andrews Church, 7 Jun

1958, Stewart 469 (NCU). Lenoir Co.: W of Deep Run, 22 Jun 1957,

Radford25701 (NCU). Moore Co.: Weymouth Woods State Park, 13

Jun 1965, Ahles 59622 (NCU); Weymouth Woods-Sandhills Nature

Preserve, 12 May 1973, Carter 556 (NCU); N of Southern Pines, 11

Jun 1978, Reed 106667 (NCU). New Hanover Co.: Wilmington,

May 1867, Canby s.n. (NCU*); NW of Carolina Beach, 12 Jun 1958,

Bell 12691 (NCU); UNC-Wilmington, 2 Jun 1975, Sieren 1330 (NCU).

Pender Co.: SE of Montague, 26 Jul 1953, Radford 7400 (NCU); NE

of Hampstead, 13 Jun 1957, Ahles 28003 (NCU). Richmond Co.: S

of Moore County Line, 4 Jun 1958, Duke 976 (NCU). Robeson Co.:

S of Red Springs, 21 Jun 1957, Ahles 28922 (NCU). Sampson Co.:

SE of Highsmith, 5 May 1957, Ahles 24643 (NCU); W of Clement, 28

Jun 1957, Ahles 30058 (NCU). Scotland Co.: W of Laurinburg, 8 May

1957, Ahles 24817 (NCU); S of Hoke - Scotland County line, 20 Jun

1957, Ahles 28580 (NCU); NW of US 15-501,18 May 1973, Sharp 1592

(NCU); Green Pond, 2 Nov 1984, Berg 1198 (NCU); Scotland Road

Annual Burn Site, 2 Jun 2007, McCormick s.n. (NCU). Wayne Co.: W

of Seven Springs, 21 Jun 1957, Radford25480 (NCU). Wilson Co.: NW

of Sims, 28 Jul 1958, Radford 38003 (NCU).— SOUTH CAROLINA.

Darlington Co.: Cannon's, 8 Jul 1909, Coker s.n. (NCU); Hartsville,

15 May 1910, Coker s.n. (NCU); W of Black Creek, 12 Jul 1920, Norton

s.n. (NCU); W of Hartsville, 22 Apr 1921, Norton s.n. (NCU); Hartsvlle,

10 May 1932, Smith s.n. (NCU); near Black Creek along Highway 34,

23 May 1940, Smith 1664 (NCU); Kalmia Gardens., 27 Sep 1940, Smith

Poindexter et al.. Reevaluation of Stipulicida (CaryophyIlaceae)

429

1668 (NCU); Witherspoon Island, 3 Jun 1941, Smith 612 (NCU); E side

of Lynches River above Clyde, 18 Jul 1941, Smith 777 (NCU); Coker

College Arboretum, 26 Apr 1944, Matthews s.n. (NCU). Dillon Co.:

NNE of Oak Grove, 18 Apr 1957, Ahles 23281 (NCU); NW of Dillon,

12 Jun 1957, Ahles 27802 (NCU); Edgefield Co.: S of Trenton, 4 Jul

1957, Radford26445 (NCU). Fairfield Co.: W of Lugoff, 24 Aug 1961,

Radford 44256 (NCU). Florence Co.: NE of Lynches River, 30 Jun

1958, Bell 13449 (NCU). Horry Co.: Myrtle Beach, 23 Jun 1931, Coker

s.n. (NCU). Jasper Co.: Turtle Island, 14 Jun 1997, McMillan 2492

(NCU). Kershaw Co.: N of Bethune, 3 Aug 1958, Duke 1846 (NCU).

Lancaster Co.: SW of Heath Springs, 6 Jun 1957 , Ahles 27403 (NCU).

Lee Co.: NE of Bishopville, 6 Jun 1957, Radford24295 (NCU); NE of

Lucknow, 26 Jul 1957, Radford 17340 (NCU). Lexington Co.: SE of

Lexington, 27 May 1957, Radford23315 (NCU). Marlboro Co.: Little

Pee Dee River, 10 Jun 1956, Radford 12493 (NCU). Orangeburg Co.:

W of Orangeburg, 18 May 1957, Ahles 25242 (NCU). Richland Co.: E

of Ft. Jackson entrance, 13 May 1958, Duke 529 (NCU). Saluda Co.: S

of Ridge Spring, 26 May 1957, Radford23128 (NCU). Sumter Co.: N

of Pinewood, 5 Jun 1957, Radford24011 (NCU). Williamsburg Co.:

N ofMouzon, 12 Jun 1957, Radford24825 (NCU).— VIRGINIA. Isle

of Wight Co.: S of Lees Mill, 29 May 1966, Svenson 13932 (NCU*).

ACKNOWLEDGMENTS

We would like to thank the curators at GH, MICH, MIN, MSC, NY, P, US, and USF for providing specimen

loans or digital images, and Carol Ann McCormick and Shanna Oberreiter (NCU) for assistance with

databasing and curation of these collections. We are grateful to the Museum National d’Histoire Naturelle

(MNHN) - Paris Herbarium (P) for giving us permission to publish the digital image of an isotype of Stipulicida

setacea from the Michaux Herbarium. We extend our appreciation to the UNC Chapel Hill Biology Department

and UNC Herbarium (NCU) for providing resources to conduct this study. The quality of this manuscript was

improved by the reviews of Richard Rabeler and Daniel Spaulding. This project was partially funded by the

Charles T. Mohr Internship Fund of the UNC Herbarium / North Carolina Botanical Garden.

REFERENCES

Fior, S., P. Ola Karis, G. Casazza, L. Minuto, & F. Sala. 2006. Molecular phylogeny of the Caryophyllaceae (Caryophyllales)

inferred from chloroplast matK and nuclear rDNA ITS sequences. Amer. J. Bot. 93:399-411.

Gill, F.B. 2014. Species taxonomy of birds: Which null hypothesis? The Auk 131:150-161.

Greenberg, A.K. & MJ. Donoghue. 2011. Molecular systematics and character evolution in Caryophyllaceae. Taxon

60:1637-1652.

Harbaugh, D.T., M. Nepokroeff, R.K. Rabeler, J. McNeill, E.A. Zimmer, &W.L. Wagner. 2010. A new lineage-based tribal classifica¬

tion of the family Caryophyllaceae. Int. J. PI. Sci. 171:185-198.

James, C.W. 1957. A new variety of Stipulicida setacea. Rhodora 59:98.

Judd, W.S. 1983. The taxonomic status of Stipulicida filiformis (Caryophyllaceae). Sida 10:33-36.

Kartesz, J.T. 2013. Taxonomic Data Center, (http://www.bonap.net/tdc). The Biota of North America Program (BONAP).

Chapel Hill, North Carolina, U.S.A. [maps generated from Kartesz, J.T. 2013. Floristic synthesis of North America, Ver¬

sion 1.0. Biota of North America Program (BONAP). (in press)]

Michaux, A. 1803. Flora Boreali-Americana. Vol. 1. Levrault, Paris, France.

Nash, G.V. 1895. Notes on some Florida plants. Bull.Torrey Bot. Club 22:141

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.

Small, J.K. 1903. Flora of the southeastern United States, being descriptions of the seed-plants, ferns and fern-allies

growing naturally in North Carolina, South Carolina, Georgia, Florida, Tennessee, Alabama, Mississippi, Arkansas,

Louisiana, and in Oklahoma and Texas east of the one hundredth meridian. Published by the author, New York, New

York, U.S.A.

Small, J.K. 1913. Flora of the southeastern United States, being descriptions of the seed-plants, ferns and fern-allies

growing naturally in North Carolina, South Carolina, Georgia, Florida, Tennessee, Alabama, Mississippi, Arkansas,

Louisiana, and in Oklahoma and Texas east of the one hundredth meridian, second edition. Published by the author,

New York, New York, U.S.A.

Small, J.K. 1933. Manual of the southeastern flora, being descriptions of the seed plants growing naturally in Florida,

Alabama, Mississippi, eastern Louisiana, Tennessee, North Carolina, South Carolina, and Georgia. University of North

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

Sorrie, B.A. & A.S. Weakley. 2001. Coastal Plain vascular plant endemics: Phytogeographic patterns. Castanea 66:50-82.

Sorrie, B.A. & A.S. Weakley. 2006. Conservation of the endangered Pinus palustris ecosystem based on Coastal Plain cen¬

tres of plant endemism. Appl. Veg. Sci. 9:59-66.

430

Journal of the Botanical Research Institute of Texas 8(2)

Swanson, A. & R.K. Rabeler. 2005. Stipulicida. In: Flora of North America Editorial Committee, eds. Flora of North America

north of Mexico. Vol. 5. Magnoliophyta: Caryophyllidae, part 2. Oxford University Press, New York, U.S.A. Pp. 27-28.

Ward, D.B. 2001. New combinations in the Florida flora. Novon 11:360-365.

Ward, D.B. 2012. New combinations in the Florida flora III. Phytologia 94:459-485.

Weakley, A.S. 2005. Change overtime in our understanding of the flora of the southeastern United States: Implications

for plant systematics, bioinformatics, and conservation. Ph.D. dissertation, Duke University, Nicholas School of the

Environment and Earth Sciences, Durham, North Carolina, U.S.A.

Weakley, A.S. 2012. Flora of the southern and mid-Atlantic states, working draft of 30 November 2012. University of

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

Wunderlin, R.P. & B.F. Hansen. 2008. Atlas of Florida vascular plants, www.plantatlas.usf.edu [S.M. Landry and K.N. Camp¬

bell (application development), Florida Center for Community Design and Research.] Institute for Systematic Botany,

University of South Florida, Tampa, Florida, U.S.A. Accessed 28 Feb 2014.

Wunderlin, R.P. & B.F. Hansen. 2011. Guide to the vascular plants of Florida, third edition. University Press of Florida,

Gainesville, Florida, U.S.A.

A NEW HEDGE-NETTLE ( STACHYS: LAMIACEAE)

FROM SOUTH CAROLINA, U.S.A.

John B. Nelson

A. C. Moore Herbarium

Department of Biological Sciences

University of South Carolina

Columbia, South Carolina 29208, U.S.A.

Douglas A. Rayner

Department of Biology

Wofford College

Spartanburg, South Carolina 29345, U.S.A.

ABSTRACT

A distinctive and geographically restricted species of Stachys, S . caroliniana (Tamiaceae), or “hedge-nettle,” is described from South Caro¬

lina, as a member of the flora of the Coastal Plain of the southeastern U.S.A.

RESUMEN

Se describe del Carolina del Sur una especie distinta restringida geograficamente de Stachys, S . caroliniana (Lamiaceae), o “hedge-nettle,”

como miembro de la flora de la Llanura Costera de sureste de Estados Unidos.

Stachys is one of the larger genera of the subfamily Lamioideae within the Lamiaceae, encompassing nearly 300

species, and with a nearly cosmopolitan distribution, absent from Australia and the western Pacific (Epling

1934; Nelson 1981; Mulligan & Munro 1989). Its centers of species diversity include warm-temperate portions

of western Asia and the Mediterranean, southern Africa, and portions of North and South America (ilgim et al.

2008). The genus in North America, north of Mexico, contains about 45 species, with species particularly con¬

centrated west of the Rocky Mountains (especially in the Pacific states) and in the east along the Atlantic sea¬

board states and Appalachian region. Southeastern species occur in a wide variety of habitats, most often in

mesic sites or wetlands, and at a wide variety of regional elevations. An additional species, known only from

the Santee River delta of South Carolina, deserves recognition.

Stachys caroliniana J.B. Nelson & D.A. Rayner, sp. nov. (Figs. 1-7). Type: UNITED STATES. South Carolina. George¬

town Co.: grassy margin of freshwater pond, Cat Island, Tom Yawkey Wildlife Center, 7 Jul 2005, J.B. Nelson 25466 (holotype:

USCH; isotypes: BRIT, CLEMS, FSU, GH, NBYC, NCU, NY, US).

Stachys caroliniana differs from S. pilosa, S. arenicola, and S. aspera in having only short, dense stem pubescence on both stems sides and

angles, the hairs of uniform size; in having shorter calyx lobes; and in having nearly white corollas.

Perennial herbs, with fibrous roots from pale, shallow rhizomes; stems strictly erect, to 0.5 m tall, scarcely

branched except occasionally at lowest node of inflorescence (unless injured), the angles copiously short-hairy

with stiff, short, reflexed or somewhat spreading hairs, the sides similarly pubescent, and also with rare,

nearly sessile glandular hairs which become progressively more abundant upward; nodes slightly bearded;

leaves nearly sessile, or the most basal, or largest midstem leaves with short petioles 2-5 mm; midstem leaves

generally the largest, the blades l-2(-2.5) cm broad, 2.5-6(-8) cm long; rounded to truncate at base, the lowest

blades prominently obtuse-rounded apically, then progressively more acute distally along the stem; blades

lanceolate, lance-elliptic; leaves spreading, upper surface abundantly pubescent or scaberulous with appressed

non-glandular hairs, the hairs up to 1.5 mm, sharply pointed, uniformly distributed across the surface, the

upper surface dark green, tending toward a dark bluish-green, especially in shade plants, or with an ashy-green

appearance; lower surface pale green, copiously pubescent with erect, soft, sharp-pointed hairs to 1 mm, thus

soft and “felty,” the foliage when crushed without any particular fragrance; inflorescence a series of inter¬

rupted verticils, commonly elongated with age; bracts rapidly reduced upward from lowest fertile node (or in

very robust flowering plants, more gradually reduced), the lowest bracts lanceolate, progressively trullate to

ovate, pubescent with eglandular and glandular hairs; axillary cymules each with 3 flowers, the verticils thus

J. Bot. Res. Inst. Texas 8(2): 431 - 440.2014

432

Journal of the Botanical Research Institute of Texas 8(2)

Fig. 1. Flowering stems ofStachyscaroliniana. Photo by Jim Fowler.

Nelson and Rayner, A new species of Stachys from South Carolina

433

Fig. 2. Open flowers at a single node of Stachys caroliniana. Photo by Jim Fowler.

6-flowered; bracteoles narrow, spinulose, glandular and pubescent, 0.5-0.8 mm long; flowers perfect and fer¬

tile; calyx campanulate, densely short-soft pubescent, with abundant delicate, stipitate glands; fruiting calyx

5.5-7 mm long, the tube (to base of lobe sinus) 3.5-4 mm, the lobes 2-3 mm; lobes abundantly hairy with eg-

landular and stipitate glands, especially along veins, deltoid to lanceolate, mostly V 2 the length of the tubes,

each lobe terminated by a pale apiculum; corolla tubular and galeate, from base to galea apex 13 mm long at

full anthesis; corolla tube prominently saccate toward base on lower side, internally glabrous, but with promi¬

nently slanting (oblique) annulus, copiously pubescent with soft, bulbous trichomes; corollas white, scarcely

pinkish, save for scattered spots on lower lip, the lower corolla lip declined up to 90° from tube, to 7 mm long,

the central lobe nearly rotund, 4-5.5(-6) mm broad; filaments pink, with short, capitate glands as well as bul¬

bous trichomes; mericarps 1-1.2 mm broad, 1.6-1.8 mm long, dark brown, minutely pebbled.

Additional collections examined: UNITED STATES. South Carolina: Georgetown Co.: first collection from type locality, 22 Jun 199 OJ.B.

Nelson 9256 (USCH); same locality, 12 Aug 1993 ,J.B. Nelson 14649 with C.N. Horn (USCH). Charleston Co.: Santee Coastal Reserve, along

old rice canal, 10 Aug 1977, DA. Rayner 1011 (USCH).

The first known collection of this plant is from Charleston County, South Carolina, on the south side of the

Santee River within the Santee Coastal Reserve, collected by D.A. Rayner (1011, USCH 20756) in 1977, a uni-

cate, which remained a questionable collection of Stachys aspera Michx. (as Stachys hyssopifolia Michx. var. am-

higua Gray) for some time. The Santee Coastal Reserve is an extensively managed, large holding of land (nearly

24,000 acres) owned and administered by the South Carolina Department of Natural Resources (SCDNR). For

decades, this property has been managed dynamically for wildlife habitat, although its 18th-19th Century

history was devoted in considerable part to rice culture. The original population was located between pine

434

Journal of the Botanical Research Institute of Texas 8(2)

Fig. 3. Flowers in early fruit ofStachys caroliniana. Photo by Jim Fowler.

flatwoods and a freshwater marsh; this population has not been seen since its first collection. The same plant

was collected again in 1990 in Georgetown County, in its only other known locality (the type locality), ap¬

proximately 8 air miles (13 km) to the northeast of the first collection, this on the north side of the Santee River.

This second site is 10.4 air miles (17 km) south of the city of Georgetown on the west-central part of Cat Island,

within the Tom Yawkey Wildlife Center, also managed for wildlife by SCDNR, in Georgetown County (Nelson

2002). The site of the type collection is barely 0.4 miles (about 600 m) east of the Intracoastal Waterway. The

Yawkey Center collection is in habitat similar to that of the Charleston County population, along the south

side of the Santee River. [The watershed of the Santee River effectively drains nearly half of South Carolina: a

common misconception is that the Savannah River is the major river system of South Carolina, when in fact

its watershed is much more important—biologically and culturally—in neighboring Georgia (Edgar 1998)].

This taxon is instantly differentiated from all other taxa in the southeastern U.S.A. by its dense, short,

uniform stem pubescence, its relatively short calyx lobes, and its white corollas. The other species of Stachys

Nelson and Rayner, A new species of Stachys from South Carolina

435

Fig. 4. Midstem leaves and stem of Stachys caroliniana. Photo by Jim Fowler.

known to occur in Charleston and Georgetown counties are the two relatively uncommon natives S. aspera

and S. tenuifolia, as well as the widespread and weedy S.floridana. These three sympatric taxa, as well as others,

are discussed below.

Stachys aspera Michx. is a wide-ranging species, from northern Florida north into Maryland, and includ¬

ing scattered sites in the upper Midwest, and as far south as Missouri. The type (P!) was presumably collected

by Andre Michaux in Charleston County, within which modern locations are still known (e.g., Nelson 10849,

USCH, collected 25 Jul 2001, which incidentally agrees well with the type at P) as well as slightly more distant

counties (e.g., Dorchester, Nelson 25149; Berkeley, Nelson 30626, Porcher 2457b; Colleton, Bourgeois s.n., July

2009, all USCH). Stachys aspera is a potentially tall (stems over 1 m, although usually much shorter) perennial,

the stems commonly sprawling and supported by other vegetation. Its stems are typically glabrous on the sides,

with glabrate to hispid angles. Its leaves are generally lanceolate to narrowly elliptic and although hairy (gla-

brate to hispidulous/hispid), are never soft and felt-like to the touch. Its corollas are pink. The mature calyx

lobes tend to be straight or barely spreading. Although distinctive in South Carolina, S. aspera is clearly related

to S. hispida Pursh from farther north. The separation of these two taxa throughout their range is not straight¬

forward, and there is some discussion concerning their conspecihcity.

Stachys tenuifolia is an even more widespread species, known on the Coastal Plain from the Gulf Coast

(eastern Texas) to New Jersey, most commonly in considerable shade along river bottoms along the lower

Mississippi and its upper tributaries, well into Arkansas, Oklahoma, Missouri, Iowa, Illinois, Indiana, Ohio,

436

Journal of the Botanical Research Institute of Texas 8(2)

Fig. 5. Habitat at type locality of Stachys caroliniana, John Cely pictured. Photo by John Nelson.

and West Virginia and is known additionally from a number of upland, “interior” sites. This is a variable taxon,

most often with glabrous/glabrate stems, these not infrequently slightly retrorse-hispidulous. This taxon al¬

ways features prominently petioled leaves with blades which are lanceolate or elliptical, and usually with

strongly serrated margins, with the plants commonly branching well below the first fertile node. Its corollas

are distinctly pink. Calyx lobes of S. tenuifolia, in fruit, are commonly curled backward. Stachys tenuifolia has

also been regarded as a near relative of S. hispida, the two taxa sometimes intergrading (Cooperrider & Sabo

1969).

Stachys floridana similarly has pink corollas, as well as prominently petioled leaves which are crenate (to

serrate) on the blade margins, and commonly cordate at the base. Plants commonly produce crisp, somewhat

swollen, bulbous, moniliform rhizomes (which are edible). This is the earliest-flowering perennial species

known in the southeastern U.S.A., sometimes blooming in early April on the Coastal Plain of South Carolina,

and has earned considerable notoriety as an aggressive weed. Above-ground short systems commonly senesce

and die by early June, normally disappearing entirely by late summer.

There is no question that the plant from Charleston and Georgetown Counties, South Carolina, is distinc¬

tive from all other commonly known southeastern taxa. The only other possibility of close taxonomic relation¬

ships involves those with Stachys pilosa Nuttall and Stachys arenicola Britton, the ranges of which are consider¬

ably far removed from S. caroliniana.

Stachys pilosa is an extremely variable and softly hairy plant, occurring from Ontario into much of the

American Midwest, which surely represents the broadest distribution of any North American Stachys; it is not

a part of the flora of the southeastern U.S.A. Its stems feature long, spreading hairs on the angles and usually at

Nelson and Rayner, A new species of Stachys from South Carolina

437

Fig. 6. Location of known populations of Stachys carolin iana. G = Georgetown County (type locality), C = Charleston County, Rayner collection (approxi¬

mate). Inset map indicates borders of South Carolina, Georgetown County, Charleston County, and the drainage of the Santee River.

438

Journal of the Botanical Research Institute of Texas 8(2)

Collected from SOUTH side of the road...

PLANTS OF THE ATLANTIC COASTAL PLAIN

SOUTH CAROLINA, U.S.A.

STACHYS

GEORGETOWN COUNTY: Foliage stems somewhat velvety

with abundant soft hairs, stem hairs spreading and retrorse; leaves

dull green, with somewhat ashy look, the lowest blades

oblong/obluse, becoming more acule/Ianeeolate upward; plants

rarely branched; corollas essentially white, some very pale pink;

abundant in shade and in full sun on grassy ground, with

Dichanthelium, Anipetopsis arborea, Parthenodsstts\ both sides of

dirt road on causeway; west-central side of Cal Island,Tom Yawkey

Wildlife Center. Ill air miles SSSE of central Georgetown; elev. 6.

Extremely hot, humid day.

John B. Nelson 2546b ] vl > 2005

with John Cely, Hob Joyner

Collected permission of South Carolina Department of Natural Resources

A. C. Moore Herbarium (USCH)

_University of Smith Carolina

A. C, Moore Herbarium, University ofSouth Carolina

t y\ i

TJcJi. *

John B. Nelson _ / H _ 2014

HOLOTVPE

98304

Qjnounted January 21

Nelson 25466 USCH 98304

Calyx lobe Calyx tube Ratio

2mm

4mm

2.1

3.5

2.1

3.6

.58

Fig. 7. Holotype of Stachys caroliniana [Nelson 25466).

Nelson and Rayner, A new species of Stachys from South Carolina

439

least a “subcanopy” of glandular and eglandular hairs on the sides, commonly forming a two-tiered appear¬

ance. Stachys pilosa has pink corollas, featuring a lance-oblong lower lip commonly notched distally.

Stachys arenicola, similarly with pink corollas, was described by Britton from plants with coarsely sca¬

brous stems and foliage, the stems featuring both coarse and fine eglandular hairs, these predominantly down¬

wardly angled, on the angles as well as the sides. Its southernmost occurrences are apparently in Virginia

(Prince William County, Fleming 7164,13408,13414 [USCH]; Frederick County, Nelson 13174 USCH).

Stachys pilosa, as widespread as it is and as a frequent resident of wet prairies, could conceivably be trans¬

ported by waterfowl along migratory flyways from more northern sites into the Southeast; the same is at least

marginally true for Stachys arenicola. And, the South Carolina sites discussed so far within this article have

historically had heavy visitation by migratory waterfowl. Stachys caroliniana, however, is narrowly restricted in

its range and apparently so differentiated morphologically that its conspecihc status with either S. pilosa or S.

arenicola is most unlikely.

KEY TO STACHYS CAROLINIANA AND ITS GEOGRAPHIC AND MORPHOLOGIC RELATIVES

1. Leaves with well-developed petioles Vs (or longer) the length of the blade; summer or spring-blooming.

2. Plants with slender rhizomes; stem side glabrous (rarely glabrate); stem angles glabrate (rarely hairier); blade bases

rounded; calyx glabrous to hispidulous, without glandular hairs; summer blooming_ S. tenuifolia

2. Plants with swollen, crisp, moniliform tubers; stem angles hispidulous to hirsute with reflexed hairs; blade base

truncate to cordate; calyx abundantly pubescent with glandular and eglandular hairs; spring blooming_ S. floridana

1. Leaves sessile or with petioles less than Vs length of blade; summer blooming.

3. Stem sides glabrous (to glabrescent)_ S. aspera

3. Stem sides abundantly pubescent, tomentulose, hispidulous, or hispid.

4. Corollas pink; calyx lobes V 2 the length of calyx tube; lower leaf blades acute-acuminate; lower lip of corolla ovate

to lance-ovate; VA and north.

5. Leaf blades elliptic; hairs of stem angles and sides spreading, with stipitate glands on sides_ S. pilosa

5. Leaf blades lanceolate; hairs of stem angles and sides retrorse (spreading), without stipitate glands_ S. arenicola

4. Corollas white; calyx lobes < V 2 length of calyx tube; lower leaf blades obtuse; lower lip of corolla rotund-ovate;

endemic to SC_ S. caroliniana

In spite of the large amount of acreage along the Atlantic Seaboard (New York to Florida) which more or less

duplicates landscape features of the Santee River delta in South Carolina, no other collections of this hedge-

nettle are known: indeed, its morphology is distinctive and unique. Despite the relatively high amount of bo¬

tanical interest in and around this area, both historic and recent, only two collections—and populations—are

known. The two populations are apparently from presumably similar habitats, and they are relatively near

each other. They are not weedy plants and not represented from other places by known herbarium collections.

Stachys caroliniana is here proposed as occurring in relict populations and should be considered very rare and

probably in danger of extinction.

Etymology. —Known only from South Carolina, the epithet "caroliniana ” is chosen to reflect its natural

and only known home to date.

ACKNOWLEDGMENTS

We are grateful to the administrators and personnel of the South Carolina Department of Natural Resources

for access to The Santee Coastal Reserve and the Tom Yawkey Wildlife Center, with special appreciation for the

help, respectively, of Tommy Strange and Bob Joyner. Field efforts at the Yawkey Center site were augmented by

the assistance of Devin Kennemore (USCH), Charles Horn (NBYC), and more recently by Bert Pittman, John

Cely, and Jamie Dozier (all SCDNR). Herrick Brown (USCH) provided valuable assistance in designing our

figures. We thank Jim Fowler, of Greenville, SC, for the excellent photographs of Stachys caroliniana. Our

manuscript was improved by the suggestions offered by Gary Fleming (VA Natural Heritage), Derick Poindex¬

ter (NCU), and Michael Vincent (MU). Publication costs for this paper were defrayed by the W.T. Batson En¬

dowment for the A.C. Moore Herbarium at the University of South Carolina.

440

Journal of the Botanical Research Institute of Texas 8(2)

REFERENCES

Cooperrider, T.S. & R.F. Sabo. 1969. Stachys hispida and S. tenuifolia in Ohio. Castanea 34:432-435.

Edgar, W.T. 1998. South Carolina: A history. University of South Carolina Press, Columbia, U.S.A.

Epung, C.C. 1934. Preliminary revision of American Stachys. Repert. Spec. Nov. Regni Veg. 80:1-75.

Ilcim, A., M. £enet, & M.Y. Dadandi. 2008. Stachys marashica (Lamiaceae), a new species from Turkey. Ann. Bot. Fennici

45:151-155.

Mulligan G.A. & D.B. Munro. 1989. Taxonomy of North American species of Stachys (Labiatae) found north of Mexico.

Naturaliste Can. 116:35-51.

Nelson, J.B. 1981. Stachys in southeastern United States. Sida 9:104-123.

Nelson, J.B. 2002. Floristicand Landscape Inventory of Tom Yawkey Wildlife Center, Georgetown County, South Carolina.

Unpublished final report: Tom Yawkey Foundation.

AGAVE x M ADRENSIS (ASPARAGACEAE), A PUTATIVE HYBRID FROM THE

SIERRA MADRE ORIENTAL, MEXICO

Jose A. Villarreal Quintanilla and

Abraham Ramirez Gamez

Universidad Autonoma Agraria Antonio Narro

Departamento de Botanica

Buenavista, Saltillo 25315, Coahuila, MEXICO

javillarrealOO@hotmail.com

ica_ab.ram@hotmail.com

Dino U. Gonzalez Uribe

Universidad Autonoma Agraria Antonio Narro

Departamento de EstadfsticayCalculo

Buenavista, Saltillo 25315, Coahuila, MEXICO

digon_mx@yahoo. com

Eduardo Estrada Castillon

Facultad de Ciencias Forestales

Universidad Autonoma de Nuevo Leon

Apartado postal 41

Linares, 67700, Nuevo Leon, MEXICO

aeduardoestradac@prodigy.net.mx

Diana Jasso de Rodriguez

Universidad Autonoma Agraria Antonio Narro

Departamento de Fitomejoramiento

Buenavista, Saltillo 25315, Coahuila, MEXICO

ABSTRACT

A putative hybrid in Agave is described and illustrated from southern Nuevo Teon state in Mexico. Agave x madrensis is believed to be a

natural, crossbreeding hybrid between Agave gentryi and Agave lechuguilla. A morphometric analysis between the parental species and the

hybrid was created to assess the parentage of the individuals in question. A multivariate analysis (MANOVA) and a discriminant analysis are

used to support the hypothesis of the existence of a natural occurrence of this hybrid. Plant illustrations are included.

Key Words: Agave x madrensis, Asparagaceae, Nuevo Teon, Mexico, hybridization, morphometric analysis

RESUMEN

Se describe un supuesto hibrido en Agave con material proveniente del sur del estado de Nuevo Leon, Mexico. Agave x madrensis se cree es

un cruzamiento natural entr e Agave gentryi y Agave lechuguilla. Se realizo un analisis morfometrico de las especies involucradas para evalu-

ar la posible relacion en el parentesco del hibrido. Un analisis multivariado (MANOVA) y uno discriminante se usan para apoyar la hipotesis

de la existencia de un hibrido natural intermedio entre las especies parentales. Se incluyen ilustraciones de la planta.

Agave is a genus endemic to America. It has approximately 200 species and 247 taxa, of which 75 per cent are

found in Mexico, most of them growing in arid and semiarid regions (Garcia 2002). Natural hybridization is

not rare in Agave (Gentry 1967), as shown below. Hybrids are known in elements of the same subgenus, as in A.

nickelsiae Gosselin and A. lechuguilla Torr., both of subgenus Littaea (Gonzalez et al. 2011), but are more fre¬

quent between species of different subgenera, as in A. marmorata Roezl (subg. Agave) and A. kerchovei Lem.

(subg. Littaea) of central Mexico (Valverde et al. 1996), A. asperrima Jacobi (subg. Agave) and A. nickelsiae Gos-

selin (subg. Littaea) (Gentry 1982; Gonzalez et al. 2011) and A. asperrima Jacobi (subg. Agave) and A. victoria-

reginae T. Moore (subg. Littaea) both from northern Mexico (Verduzco et al. 2009). Most of them were de¬

scribed at the specibc level. Several putative hybrids are also known from the southwestern United States and

include A. x arizonica Gentry & J.H. Weber, A. x ajoensis W.C. Hodgs. and probably A. schottii Engelm. var.

treleasei (Tourney) Kearney & Peebles (Reveal & Hodgson 2001).

Sixteen taxa are reported to occur in Nuevo Leon (Villarreal & Estrada 2008). A recent collection from

the Sierra Madre Oriental of Mexico included a specimen of Agave that is morphologically unlike any known

species. Plants with large leaves and a racemose inflorescence led us to the idea of hybridization between Agave

gentryi and A. lechuguilla, the only two species growing in the area. As both species are frequent in the moun¬

tains of north and central Mexico and the probability of a potential hybrid between them is present, the hybrid

deserves to be named and recognized as a taxonomic element. A statistical analysis of morphometric traits was

J. Bot. Res. Inst. Texas 8(2): 441 - 447.2014

442

Journal of the Botanical Research Institute of Texas 8(2)

Table 1 . Means and standard deviations of the characters used in the morphometric study.

A. lechuguilla

A. gentryi

putative hybrid

Plant diameter (d)

68.600±5.621

248.100±18.935

131.667±10.408

Leaf length (II)

40.900±4.483

84.800±9.601

65.000±15.000

Leaf width (Iw)

2.960±0.212

16.500±1.958

10.333±1.528

Number of leaves (nl)

39.200±5.116

48.500±6.346

49.000±7.937

Spines distance (sd)

1.840±0.196

3.380±0.319

2.467±0.503

Number of spines (ns)

15.400±2.171

30.300±3.622

14.667±2.517

Terminal spine length (tsl)

1.560±0.232

7.270±0.460

3.333±0.577

Valvedrian distance (vd)*

2.390±0.614

12.000±3.197

9.667±1.528

Inflorescence length (il)

1.170± 1.526

2.250±3.631

5.333±4.726

Inflorescence width (iw)

4.700±6.075

42.300±68.131

23.333±20.817

*valverdian distance = distance between the upper spine and the terminal spine

Table 2. Means and standard deviations of the ratios used in the morphometric analysis.

A. lechuguilla

A. gentryi

putative hybrid

Plant diameter/number leaves

4.530±0.695

8.288±1.087

9.078±0.885

Leaf length/width

13.853±1.602

5.211 ±0.889

6.241 ±0.599

Spines number/distance

8.395±1.016

9.097±1.787

5.972±0.293

Terminal spine length/valverdian distance*

0.701 ±0.233

0.669±0.277

0.293±0.346

Inflorescence length/width

1.358±1.785

0.184±0.297

1.431 ±1.344

*valverdian distance = distance between the upper spine and the terminal spine

used to test the hypothesis that a hybrid should be expected to be morphologically intermediate between the

putative parents.

MATERIALS AND METHODS

Ten plants belonging to each parental species were randomly selected in two populations in the area, and three

plants of the putative hybrid. Three leaves were evaluated and the average computed for each individual. The

morphometric measures are: rosette diameter (d), number of leaves (nl), leaf length (11), leaf width (lw), num¬

ber of spines by side (ns), distance between spines (ds), terminal spine length (tsl), and distance between the

last and the terminal spine-valverdian distance (vd). The length (il) and width (iw) of three inflorescences of

parental and putative hybrid were included.

The data were statistically evaluated by a discriminant analysis using a multivariate analysis (MANOVA)

with the Wilk’s test (Everitt & Hothorn 2010), included in the PAST software, version 2.01 (Hammer &

Harper 2006).

RESULTS

Means and standard deviations of the characters used in the morphometric study are shown in the Table 1. Five

character means (leaf length, leaf width, number of leaves, valverdian distance and inflorescence length) are

closer to Agave gentry, while four (plant diameter, spines distance, number of spines and the terminal spine

length) are for A. lechuguilla; and one (inflorescence width) is intermediate between both species. Seven means

of the putative hybrids lay between the means of the parental species, and three (number of leaves, number of

spines and inflorescence length) are slightly exceeded from a parental mean.

The ratios and standard deviation used in the discriminant analysis are shown in the Table 2. The plot

derived from the analysis (Fig. 1) includes three groups clearly separated in the discriminant space. It can be

concluded that the ratios of A. lechuguilla and A.gentryi are different and the putative hybrid is intermediate,

according the Wilk’s test (p < 0.001) with a Wilk’s Fambda = 0.01692. However, direct breeding studies are

Villarreal Q. et al., Agave x madrensis, a putative hybrid

443

Fig. 1. Plot that shows the results of the use of the Discriminant analysis applied to the five morphometric ratios listed in the Table 2.

needed for more conclusive evidence. As the two parental species belong to different subgenera we are consid¬

ering that the plant is a putative intersubgeneric hybrid and is described as a new taxonomic entity.

Agave x madrensis Villarreal, A. Ramirez, & A.E. Estrada, hybrid nov. (Figs. 2-3). Type: Mexico. Nuevo Leon: Mu-

nicipio de Galeana, cerro El Potosf, camino a las antenas. Bosque esparcido de Pinus hartwegii y encinar enano, 2654 m, 21 Aug

2012, A. Ramirez 106 (holotipo: MEXU; isotypes: ANSM, CFNL, ENCB, TEX; to be distributed).

Similar to Agave montium-sancticaroli Garcia-Mend., but with lanceolate leaves 50-80 cm long, inflorescence 7-9 m long with 80-150 florif-

erous branches and distribution in the Sierra Madre Oriental.

Perennial single plants, acaulescent, moderately suckering; rosettes hemispheric, openly spreading, 120-140

cm wide, 80-100 cm high. Leaves 40-55 per plant, lanceolate, concave toward apex, 50-80 cm long, 9-12 cm

wide, ascending, glaucous to green-yellowish when mature, the base slightly enlarged, the apex acuminate, the

margins straight, the terminal spine 3-4 cm long and 6-7 mm wide at the base, brown, the marginal teeth

12-17 pairs, straight to slightly retrorse, 2-3 cm apart, 4-7 mm high, brown. Flowering stalk 7-9 m tall, the

inflorescence a racemose panicle, congested, largely oblong (fusiform), 4-5 m long and 30-40 cm wide, the

peduncles 8-12 cm diameter, bracteates; bracts long triangular, 20-28 long, 5-6 cm wide at the base, charta-

ceous, the margins entire, the apex with a short and weak spine; branches of the inflorescence 15-20 cm long,

spaced 10-15(-20) cm apart, green-purple; bracts like those of the peduncle, largely triangular, 12-18 cm long.

Flowers 18-26 per umbel, 5-6 cm long, narrowly campanulate, succulent, green-yellowish, flushed with

purple; pedicels 4-12 mm long; tepal lobes largely triangular, 18-20 mm long, 3-5 mm wide, succulent, the

outer ones slightly larger than the inner, the margins involute, the apex cucullate; floral tubes 5-6 mm long,

7-10 mm diameter distally, slightly sulcate; filaments 45-55 mm long, inserted above mid-tube (3-4 mm), yel¬

lowish distally flushed with purple; anthers dorsihxed, 20-25 mm long, 2 mm wide, yellow; ovary cylindric,

20-26 mm long, 6-8 mm wide, green, the neck 2-4 mm long, the style 5-6 cm long, the stigma

444

Journal of the Botanical Research Institute of Texas 8(2)

[\g.2. Agave xmadrensis. A. Flowering plant. B. Rosette of leaves. C. Inflorescence close-up. D. Close-up of flowers and pollinators. Photos by A. Ramirez G.

claviform,trilobate. Capsules oblong to slightly ovate, 4-6 cm long and 20-25 mm wide, rostrate, dark brown;

seeds lunulate, flattened, 5-6 mm long, 4-5 mm wide, black.

Distribution and ecology .—The plant grows in southern Nuevo Leon state on steep slopes in limestone

soils, at 2500 to 2700 m. The area is dominated by a dwarf oak community, mainly of Quercus greggii, and

other shrubs such as Arctostaphylos pungens, Arbutus xalapensis, Ceanothus buxifolius, Pinus culminicola, Cerco-

carpus sp., and Agave gentryi, with sparse forest components of Pinus hartwegii and Pseudotsuga menziesii. Ad¬

ditional immature specimens believed to be conspecific were observed, considering a probable wider distribu¬

tion, possibly tracking the occurrence of Agave gentryi in the Sierra Madre Oriental. Four plants were found in

the area, all of them close to a roadside, as might be expected of an association with disturbance. The incidence

of hybrids in disturbed environment was discussed by Grant (1971).

Phenology .—Flowering during the summer, fruiting at the end of summer and beginning of autumn.

Etymology .—The specific epithet refers to the Sierra Madre Oriental where the plant is distributed.

Villarreal Q. et al., Agave x madrensis, a putative hybrid

445

Fig. 3. Agave x madrensis. A. Leaf. B. Leaf margin. C. Bract. D. Cluster of flowers. E. Flower. F. Fruit. G. Seed.

446

Journal of the Botanical Research Institute of Texas 8(2)

Table 3. Comparison of select character states of Agave Xmadrensis, A. gentryi,A. lechuguilla, and A. montium-sancticaroli.

Character

A. x madrensis

A.gentryi

A. lechuguilla

A. montium-sancticaroli

Leaf length

50-80 cm

60-100 cm

25-50 cm

100-120 cm

Leaf shape

lanceolate

triangular

linear-lanceolate

Leaves/plant

40-55

30-45

50-80(100)

Flower stalk

7-9 m

3-5 m

2.S-3.S m

5.5-7 m

Infl. length

3 /s

'A-Vi

V 3 -V 4

y 2 - 5 /6

Infl. shape

fusiform

elliptic

cylindrical

fusiform

Infl. branches

80-150

16-20

80-120

60-80(140)

Branch length

15-20 cm

50-80 cm

4-6 mm

8-13 cm

Flower length

50-60 mm

70-90 mm

30-45 mm

40-50(-60) mm

Distribution

SMO

CDR

LLCG y SSC

SMO = Sierra Madre Oriental, CDR = Chihuahuan Desert Region, LLCG = Llanura Costera del Golfo, SSC = Sierra de San Carlos.

The plants exhibit an inflorescence morphology which is intermediate between the broad panicles of

subg. Agave with many-flowered partial inflorescences (branches), and the narrow-elongate, spike-like inflo¬

rescences of subg. Littaea with few-flowered partial inflorescences (branches), as shown in the Table 3. The

plant seems to represent a natural hybrid between a member of subg. Agave (the sympatric A. gentry B. Ullrich)

and a member of subg. Littaea (A. lechuguilla Torr.).

The plant is morphologically similar to A. montium-sancticaroli Garcla-Mend. from Tamaulipas (Garcia et

al. 2007). Both have a similar racemose paniculate inflorescence, but A. x madrensis differs by the shape and

size of the leaves (see Table 3), lack of small interstitial teeth between the large teeth found in A. montium-

sancticaroli, peduncular bracts 20-26 cm long (vs. 11-16 cm) (Table 1), and occurring in the Sierra Madre

Oriental (vs. between Llanura Costera del Golfo and Sierra de San Carlos), at 2500-2700 m (vs. 150-800 m) in

forest to oak scrubland habitat (vs. tamaulipan thornscrub habitat). Agave montium-sancticaroli is published at

the rank of species, but exhibits intermediate inflorescence morphology and might also represents a natural

hybrid between A. angustifolia Haw. and A. lophantha Schiede ex Kunth. In the protologue, A. montium-sanctic¬

aroli is compared to A. x glomeruliflora (Engelm.) A. Berger, the latter suggested to be a series of hybrids involv¬

ing A. lechuguilla Torr. of subg. Littaea and A. gracilipes Trel., A. havardiana Trel. and A. parryi Engelm. var.

ncomcxicana (Wooton & Standi.) McKechnie of subg. Agave (Gentry 1982). Agave lechuguilla is considered a

“dono-species” by Gentry (1967), hybridizing with several other taxa.

ACKNOWLEDGMENTS

We thank Wendy C. Hodgson and anonymous reviewer for their comments and suggestions to improve the

manuscript. We are grateful to the Consejo Nacional de Ciencia y Tecnologla for the financial support through

the scholarship 369816 and to the Universidad Autonoma Agraria Antonio Narro through research project

36142227.

REFERENCES

Everett, B. &T. Hothorn. 2010. A handbook of statistical analyses using R. 2nd ed. CRC Press, Boca Raton, Florida, U.S.A.

Gentry, H.S. 1967. Putative hybrids in Agave. J. Heredity 58(1 ):32—36.

Gentry, H.S. 1982. Agaves of continental North America. University of Arizona Press, Tucson, Arizona, U.S.A.

Garcia M., A. 2002. Distribution of the genus Agave (Agavaceae) and its endemic species in Mexico. Cactus Succ. J. (Los

Angeles) 74:177-187.

Garcia M., A.J., C. Jacques H., & A. Salazar B. 2007. Una nueva especie de Agave subgenero Littaea (Agavaceae) de Tamau¬

lipas, Mexico. J. Bot. Res. Inst.Texas 1 (1 ):79—84.

Gonzalez E., M.S., M. Gonzalez E., I.L. Lopez E., L. RezEndiz R., J.A. Tena F., & F.L. Retana R. 2011. El complejo Agave victoria-

reginae (Agavaceae). Acta Bot. Mex. 95:65-94.

Grant, V. 1971. Plant speciation. Columbia University Press, New York, New York, U.S.A.

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Hammer, 0. & DAT. Harper. 2006. Paleontological data analysis. Blackwell Publishing, Malden, Massachusetts, U.S.A.

Reveal, J. L. & W.C. Hodgson. 2001. Agave. In: Flora of North America Editorial Committee, eds. 1993 +. Flora of North

America North of Mexico. 16+ vols. New York & Oxford. 26:442.

Valverde, P.L., F. Vite, & J.A. Zavala H. 1996. A morphometric analysis of a putative hybrid between Agave marmorata Roezl

and A. kerchovei Lem.: Agavepeacockii Croucher. Bot. J. Linn. Soc. 122(2):155—161.

Verduzco M., J., C.L. Predo R., & R. Mercado H. 2009. Caracterizacion e identificacion taxonomica del maguey. Memorias

del VII Simposium-Taller Produccion y Aprovechamiento del Nopal en el Noreste de Mexico. Revista Salud Publica y

Nutricion 2:75-90.

Villarreal Q., J.A. & E. Estrada C. 2008. Listados floristicos de Mexico. XXIV. Flora de Nuevo Leon. Instituto de Biologia,

Universidad Nacional Autonoma de Mexico. Mexico D.F.

448

Journal of the Botanical Research Institute of Texas 8(2)

BOOK REVIEW

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ments for the garden. Colored photos of plants and butterflies and tables of plants, bloom times, and height add

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J. Bot. Res. Inst. Texas 8(2): 448.2014

DOS NUEVAS ESPECIES DE MYRCIA (MYRTACEAE),

DEL PACIFICO SUR DE COSTA RICA

Daniel Santamaria Aguilar

Armando Estrada Ch.

Direction Actual: Harvard University Herbaria

22 Divinity Avenue

Cambridge, Massachusetts 02138-2020, U.S.A.

daniel.santamaria366@gmail.com

Museo National de Costa Rica

Apartado Postal 749-1000

San Jose, COSTA RICA

aestrada@museocostarica.go.cr

Reinaldo Aguilar

Los Char cos de Osa, Centro de

Diver si dad de Plantas Regionales

Apartado Postal 76-8203

Peninsula de Osa, Puntarenas, COSTA RICA

RESUMEN

Se describen e ilustran dos nuevas especies de Myrcia: Myrcia paulii-jonesii y Myrcia riverae, endemicas de Costa Rica. Se comentan sus

afinidades, distribucion, fenologia y habitat.

ABSTRACT

Two new species of Myrcia endemic to Costa Rica are described and illustrated: Myrcia paulii-jonesii and Myrcia riverae. Their affinities,

distribution, phenology, and habitat are discussed.

Myrcia DC. es uno de los generos mas diversos y complejos de la familia Myrtaceae, con alrededor de 350 espe¬

cies y una amplia distribucion en America tropical y subtropical (Lucas et al. 2011; Parra-O 2013). Se distingue

de los otros generos de la familia por sus inflorescencias paniculadas, las flores con el caliz abierto en boton,

con cinco sepalos distintos (raro 4), persistentes, ovario con 26 3 (-4) loculos, 2 ovulos por loculo y las semillas

16 2, con el embrion myrcioide (Holst & Kawasaki 2007; Lucas et al. 2011).

El genero fue subdividido por McVaugh (1969) en las secciones Aulomyrcia (O. Berg) Griseb., Armeriela

McVaugh y Myrcia, de las cuales solo la seccion Myrcia ha resultado monohletica (Lucas et al. 2011). Por otra

parte, el genero Gomidesia O. Berg, un grupo numeroso de especies suramericanas con indumento pardo-ro-

jizo o ferruglneo, lobos del caliz truncados y anteras con tecas desplazadas o reflexas (McVaugh 1968; Holst &

Kawasaki 2009; Nic Lughadah et al. 2010), es actualmente incluido dentro de Myrcia respaldado por estudios

moleculares recientes, los cuales sugieren ademas que nueva clasihcacion subgenerica es requerida (Holst &

Kawasaki 2009; Lucas et al. 2011).

En Costa Rica el genero esta representado por ocho especies (sin incluir las dos aqul descritas), aunque se

indica que algunas de estas entidades son tentativas o bien podrlan comprender varias especies distintas, por

tanto muy posiblemente la diversidad de especies en el pals sea mayor (Holst & Kawasaki 2007).

Myrcia paulii-jonesii Aguilar, D. Santam., & A. Estrada, sp. nov. (Figs. 1, 2). Tipo: Costarica. Puntarenas: Canton Osa.

Peninsula de Osa, Bahia Chal-Bajo San Juan, Finca Rafael Granados Altamirano, 08 o 45'20"N, 83 o 30'52"W, 53 m, 01 Ago 2013 (fls),

N. Zamora, R. Aguilar, S. Kim, G. Villalobos, A. Azofeifa, D. Solorzano 6494 (holotipo: CR, isotipos: F, GH, K, MO, NY, SEL, USJ).

Myrcia paulii-jonesii sp. nov. is similar to Myrcia pentagona McVaugh and Myrcia antioquensis Parra-O. It differs from M. pentagona by its

appressed pubescence, not pannose or deciduous in groups in drying (vs. pubescence pannose and deciduous in groups in drying), its

wider leaves (6.1-13.3 cm vs. 5-7.5 cm), the midvein that is slightly raised towards the distal half (vs. sulcate), a short and slightly revolute

leaf base (0.5-1 cm vs. 1-5 cm from the base) and its smooth and unridged hypanthium (vs. a hypanthium with 5 prominent ridges). It differs

from M. antioquensis by its appressed pubescence (vs. tomentose), its narrower leaves (6.1-13.3 cm vs. 7.8-16.5 cm), its smooth, unridged

hypanthium (vs. a hypanthium with 5 prominent ridges), its smooth fruits (vs. fruits with 5 distinct ridges), and by its shorter inflores¬

cences (6-17 cm vs. (8-) 14-32 cm in length).

J. Bot. Res. Inst. Texas 8(2): 449 - 455.2014

450

Journal of the Botanical Research Institute of Texas 8(2)

Fig. 1. k-H.Myrcia paulii-jonesii. A. Haz. B. Enves. C. apice. D. Base. E. Rama con inflorescences. F. Flor y botones florales. G. Rama con infrutescencias.

H. Frutos. (A-D, G, H de /?. Aguilar 12300 ; E, F de N. Zamora etal. 6494, holotipo). Fotos por R. Aguilar.

Arbusto o arbolito 2-10 m de alto, hasta 25 DAP; yemas y ramitas mas jovenes densamente adpreso-pubescen-

tes, pronto y abruptamente glabrescentes (dejando una marca o llnea de cambio), los tricomas ca. 0.1-0.5 mm

de largo, simples, blanquecino-grisaceos a blanquecino-amarillentos, ramitas aplanadas hacia la parte distal,

sulcadas, glandulosas y fistulosas. Laminas foliares (15.0-) 17.6-34.5 x 6.1-13.3 cm, obovadas a mas comun-

mente ellpticas, 2-2.8 veces mas largas que anchas, secando cafe-rojizo a verde-oliva, el haz glabro, excepto la

vena media esparcidamente adpreso pubescente a glabrescente (algunas veces los nervios laterales diminuta-

mente pubescentes), lustroso, el enves diminuto pubescente, cubierto por tricomas blanquecinos o amarillen-

tos, ca. 0.1-0.3 mm de largo; glandulas punctatas a veces visibles en el enves, negruzcas, indistintas en el haz;

vena media en el haz sulcada o plana de la mitad hacia la base de la lamina y ligeramente elevada de la mitad

hacia el apice, en el enves prominentemente elevada; nervios laterales (13-) 17-22 pares por lado, sin incluir los

intermedios, marcadamente impresos en el haz, prominentes en el enves; nervio marginal ligeramente ar-

queado entre los laterales, impreso en el haz y elevado e igual de prominente que los laterales en el enves, 0.3-

0.5 cm del margen; base cuneada, leve y cortamente decurrente y revoluta (0.5-1 cm desde la base); apice

abruptamente acuminado; peclolos (0.3-)1.0-1.4 cm de largo, densamente adpreso pubescente a glabrescente

con la edad, sulcados. Inflorescencias panlculas, axilares o subterminales, 6-17 cm de largo, erectas, solitarias,

ejes y ramitas irregularmente aplanadas a estrechamente 4 anguladas, densamente adpreso pubescentes, trico¬

mas blanquecinos; bracteas y bracteolas posiblemente deciduas, no observadas; pedunculos 2-5.5 cm de largo,

3-3.5 mm de ancho bajo el primer nodo, aplanados. Flores 5-meras; botones 4-4.5 mm de largo, sesiles o cor¬

tamente pedicelados, ca. 0.1-1 mm de largo, hipanto 2-2.5 mm, densamente adpreso-pubescentes hacia la

base, tricomas blanquecino-amarillentos, liso (a veces arrugado longitudinalmente producto del secado), lo-

bos del caliz 1.5-2 mm x 2-2.5 mm, desiguales, imbricados, ± en forma de “D,” el apice redondeado o agudo,

adpreso pubescente en ambas superficies, persistentes y extendidos en el fruto; petalos cuando la flor en boton

ca. 2.5.-4.0 x 2.9-3.6 mm, blancos, esparcidamente pubescentes en ambas superficies, el apice redondeado;

Santamaria et al., Dos nuevas especies de Myrcia

451

Fig. 2. A-F. Myrcia paulii-jonesii. A. Ramitas adpreso pubescentes y luego abruptamente glabrescentes (marca de cambio). B. Yemas y ramitas nuevas

denso adpreso-pubescentes. C. Base revoluta y decurrente. D. Ramita de inflorescencia con botones florales. E. Boton floral. F. Frutos. (A- E de N. Zamora

6494; F de /?. Aguilar 3582). Fotos por A. Estrada.

estambres + 160 por flor, filamentos 3.0-6.0 mm de largo, glabros, blancos, anteras ca. 0.2-0.5 mm, el apice del

conectivo con una glandula conspicua; estilo 4.5-6 mm de largo, pubescente en la mitad proximal, glabro

distalmente. Frutos 0.8-1.2 x 0.4-0.7 cm, generalmente elipsoides, glandulosos, esparcidamente pubescentes,

tricomas mas abundantes hacia el apice y los lobos del caliz, rosados o rojo-rosados; semilla 1, 0.5-0.7 x 0.25

cm cafe oscuro a negruzcas, lisas a un poco rugosas.

Fenologia. —Flores en julio y agosto. Frutos en agosto y septiembre.

Distribution y Habitat.—Myrcia paulii-jonesii es una especie endemica de Costa Rica. Se distribuye en los

bosques muy humedos en el Paclfico Sur costarricense, en la Peninsula de Osa (Bahia Chal y alrededores), en-

tre 10 y 100 m de elevacion (Fig. 4). Se ha recolectado en orillas de caminos y claros naturales, en terrenos con

ligera pendiente. Se ha observado que localmente es abundante y crece junto a elementos florlsticos restringi-

dos a esta region del pals, como: Byttneria osaensis (Malvaceae), Cremastosperma sp. nov. (Annonaceae), Inga

golfodulcensis (Fabaceae), Miconia osaensis (Melastomataceae), Schnella bahiachalensis (Fabaceae), Williamo-

dendron glaucophyllum (Lauraceae).

Discusion.—Myrcia paulii-jonesii es una especie llamativa, que se caracteriza por sus ramitas jovenes

aplanadas, glandulares y densamente cubiertas por tricomas adpreso-blanquecinos; laminas foliares relativa-

mente grandes, blanquecino-grisaceas en el enves (cuando secas), con la base levemente revoluta, nervios

laterales y marginal muy conspicuos, fuertemente impresos en el haz y elevados en el enves y por sus frutos

elipsoides rosados, glandulosos y esparcidamente pubescentes. En estado juvenil esta especie llega a tener

452

Journal of the Botanical Research Institute of Texas 8(2)

laminas foliares mucho mas grandes ca. 34.5-53.5 x 11.4-13 cm, con el apice cirriforme y tricomas ± largos,

blanquecinos y erectos que cubren las ramitas, hojas y peclolos.

Esta especie es similar a Myrcia pentagona McVaugh (de la Amazonia peruana) y a Myrcia antioquensis

Parra-O. (de los bosques subandinos de Colombia), con las cuales comparte las siguientes caracterlsticas: hojas

grandes (> 14 cm), con la vena marginal y las venas laterales muy prominentes en el enves e impresas en el haz,

estas ultimas numerosas (usualmente mas de 20), la vena media ligeramente elevada en la mitad distal (solo

compartido con M. antioquensis) y la base de alguna forma revoluta. Ademas de las inflorescencias grandes y

robustas, con ejes aplanados a cuadrangulares. No obstante se diferencia de M. pentagona por su pubescencia

adpresa, no afelpada ni caediza en grupos al secar (vs. pubescencia afelpada y caediza en grupos al secar), por

sus hojas mas anchas (6.1-13.3 cm vs. 5-7.5 cm), con venas terciarias mas marcadas y evidentes, la vena media

ligeramente elevada hacia la mitad distal (vs. sulcada), la base foliar leve y cortamente revoluta (0.5-1 cm vs.

1-5 cm desde la base) y su hipanto liso, no acostillado (vs. hipanto con 5 costillas prominentes). De M. antio¬

quensis se diferencia por su pubescencia adpresa (vs. tomentosa), sus hojas mas estrechas (6.1-13.3 cm vs.

7.8-16.5 cm), su hipanto liso, no acostillado (vs. hipanto con 5 costillas prominentes), sus frutos lisos (vs. fru-

tos con 5 costillas evidentes) y sus inflorescencias mas cortas (6-17 cm vs. (8-) 14-32 cm de largo).

Otra especie con la que podrla ser comparada por su venacion prominente y la vena media elevada en el

haz, es Myrcia crassimarginata McVaugh de Peru, pero se distingue de esta por sus hojas mas grandes (17.6-

34.5 x 6.1-13.3 cm vs 16-20 x 7-8 cm), corto-pubescentes y lisas en el enves (vs. enves casi glabro y verrucu-

loso), base cuneada (vs. base redondeada), peclolos mas largos (10-14 mm vs 3-6 mm), inflorescencias mas

cortas (6-17 cm vs 15-23 cm de largo) y bracteas y bracteolas deciduas antes de la antesis (vs. persistentes

hasta la antesis).

Entre las especies costarricenses se parece un poco a Myrcia sp. A (Holst & Kawasaki 2007), del Caribe de

Costa Rica y Panama, ya que ambas comparten laminas foliares relativamente grandes y venacion prominente

e impresa; no obstante se diferencia de esta especie por sus peclolos mas largos (1.0-1.4 vs. 0.5-0.8 cm), hojas

con la vena media elevada en la mitad distal y esparcidamente pubescente (vs. sulcada y densamente pubes-

cente), los nervios secundarios menos numerosos (17-22 vs. 20-30) y la base foliar revoluta y decurrente (vs.

base recta). Se diferencia ademas por sus ramitas con pubescencia adpresa y blanquecina, pronto glabrescentes

(vs. tricomas patentes, amarillos a ferruglneos, persistentes).

Etimologia. —Esta especie es nombrada en honor al Sr. Paul Jones, en reconocimiento por apoyar los es-

fuerzos en la conservacion y la investigacion de uno de los lugares mas abundantes en biodiversidad del pla-

neta, la Peninsula de Osa.

Paratipos. COSTA RICA. Puntarenas: Osa, Sierpe. Reserva Forestal Golfo Dulce. Peninsula de Osa. Los Mogos. Bahia Chat Entrada a

Chocuaco, 08 o 46'20"N, 83°22 , 40"W, 100 m, 24 Ago 1994 (fr), R. Aguilar et al 3582 (INB); Reserva Forestal Golfo Dulce. Bahia Chal, a 35 km

de Chacarita, entrada a Chocuaco, entre cruce de Bajo Chocuaco y el bajo quebrada Lagunas, 08 o 43'45"N, 83 0 27'25"W, 10 m, 24 Sep 2009

(fr), R. Aguilar 12300 (GH, MO); misma localidad que R. Aguilar 12300, 24 Mar 2012 (Est), D. Santamaria & R. Aguilar 9355 (CR, GH, MO);

Bajo San Juan a 1500 m al Suroeste del pueblo, rfo Chocuaco arriba, finca del Estado, 08°45 , 33"N, 83°30 , 45"W, 40 m, 24 Ago 2013 (bot. fls),

R. Aguilar 14572 (CR); Los Mogos, 08°43 , 20"N, 83°26 , 30"W, 20 m, 21 Jul 1993 (bot. fls), R. Aguilar 2016 (CR, GH, MO).

Myrcia riverae A. Estrada, D. Santam., & Aguilar, sp. nov. (Fig. 3). Tipo: Costa Rica. Puntarenas: Canton de Golfito. Refu¬

gio Nacional de Vida Silvestre Golfito. Terrenos estatales al E del area, camino a las tones del ICE, 08°38' 44"N, 83 o 09'56"W, 300 m,

03 May 2011 (bot. fls), D. Santamaria, L. Diego Vargas, G. Villalobos 9416 (holotipo: CR; isotipos: F, GH, K, MO, NY, SEL).

This new species is distinguished within Myrcia by the combination of the following distinctive characters: dense velutinous, brownish-red

pubescent covering almost all the plant, large leaf blades (15.5-27.2 x 7-11.6 cm), with very prominent venation on the underside of the leaf,

straight leaf margins and being glandular-punctate on both leaf surfaces (though more visible on the underside), flowers with truncate calyx

lobes, and oblate or globose fruits with persistent calyx lobes that are either erect or connivent.

Arbol o arbolito 8-12 m de alto, hasta 15 cm DAP; yemas y ramitas jovenes densamente velutinas, los tricomas

ca. 0.2-0.8 mm de largo, simples, erectos, cafe-rojizos a dorado-cafe, ramitas cillndricas a ligeramente aplana-

das hacia la parte distal, generalmente sulcadas y hstulosas. Laminas foliares (13.2-)15.5-27.2 x (5.5-)7-11.6

cm, obovadas o mas comunmente amplio-ellpticas, 1.5-2.6 veces mas largas que anchas, secando cafe-rojizas,

el haz glabro (a veces escasa y esparcidamente pubescente), excepto en la vena media, densamente cubierta por

Santamaria et al., Dos nuevas especies de Myrcia

453

Fig. 3. A-F. Myrcia riverae. A. Rama con inflorescencias. B. Botones florales, mostrando los lobos del caliz. C. Ramitas y yemas velutinas. D. Ramita

velutina. E. Fruto con lobos del caliz erectos. F. Frutos. (A, B de D. Santamaria etai. 9416, holotipo). Fotos por L.D. Vargas; (C de G. Herrera 4167, D, E y F

de G. Rivera 4557). Fotos por A. Estrada.

tricomas simples, cafe-rojizos, la pubescencia disminuyendo hacia el apice, opaco, el enves densamente pubes-

cente, principalmente sobre los nervios, tricomas cafe-rojizos; glanduloso-punteadas en ambas superficies,

aunque mas visibles en el enves, glandulas cafe-rojizas a negruzcas; vena media en el haz sulcada hacia la base

y plana (a veces levemente elevada) hacia el apice, en el enves prominentemente elevada; nervios laterales 12-

20 pares por lado, sin incluir los intermedios, impresos o pianos en el haz, elevados en el enves; nervio mar¬

ginal ligeramente arqueado entre los laterales, impreso en el haz y elevado e igual de prominente que los latera¬

les en el enves, 0.1-0.5 cm del margen; base obtusa a cuneada, los margenes rectos; apice cortamente acumi-

nado o agudo; peclolos (0.7-)0.9-1.3(-1.6) cm de largo, densamente pubescente, ligeramente sulcado. Inflores¬

cencias panlculas, axilares o subterminales, (5.2-)6.5-13 cm de largo, erectas, de 1—3(—4) ejes por axila (decre-

cientes), ejes y ramas cillndricas o aplanadas, densamente velutinas, cafe-rojizos a dorado-cafe; bracteas y

bracteolas posiblemente deciduas, no observadas; pedunculos 1.2-4.7 cm de largo, 2-2.5 mm de ancho bajo el

primer nodo, aplanados. Flores 5-meras; botones 4.5-5 mm de largo, sesiles o cortamente pedicelados, 0.1-1.6

mm de largo; hipanto 3.5-4 mm, uniformente denso-velutino, tricomas cafe-rojizos a dorado-cafe, lobos del

caliz ca. 1.1 x 1.5 mm, imbricados, el apice truncado o depreso, densamente velutinos, adaxial y abaxialmente,

persistentes y erectos o conniventes en el fruto; petalos 3.5-4 x 2.5-3 mm, blancos, serlceos en la parte adaxial,

glabros abaxialmente, el apice redondeado; estambres + 130 por flor, hlamentos 3-5 mm de largo, glabros,

blancos, anteras ca. 0.3-0.4 mm de largo, el apice del conectivo con una glandula conspicua; estilo ca. 5.5-6.5

mm de largo, pubescente en la mitad proximal, glabro distalmente. Frutos 0.8-1 x 1-1.3 cm, globosos a obla-

tos, densamente velutinos, amarillo a anaranjados; semillas 16 2, 0.7 x 0.9 cm, lisas, brillantes.

Fenologia .—Flores en abril y mayo. Frutos en julio y agosto.

Distribution y Habitat.—Myrcia riverae es una especie endemica de Costa Rica. Se distribuye en bosques

muy humedos en el Paclhco Sur costarricense, region de Golfo Dulce (Golhto y Peninsula de Osa). Entre los

300 y 754 m de elevacion (Fig. 4). Se ha recolectado en orillas de caminos y hordes de bosques.

Discusidn .—Esta especie se puede reconocer por su densa pubescencia aterciopelada, cafe-rojiza, pre¬

sente en ramitas jovenes, hojas, inflorescencias y frutos; ademas por sus laminas foliares grandes, con venacion

muy prominente en el enves, glanduloso-punteadas en ambas superficies (mas visibles en el enves), flores con

454

Journal of the Botanical Research Institute of Texas 8(2)

84°0

los lobos del caliz truncados y sus frutos oblatos o globosos, con los lobos del caliz persistentes, erectos o con-

niventes. Estas caracterlsticas relacionan a M. riverae con el grupo Gomidesia, aunque muy distintiva de cual-

quier otra especie del grupo. En Mesoamerica Myrcia riverae junto a Myrcia aliena McVaugh, son los unicos

representantes de Gomidesia en la region. Estas especies comparten la pubescencia velutina, cafe rojiza y en los

lobos del caliz cortos y truncados; no obstante se diferencia de M. aliena por sus laminas foliares mas grandes

(15.5-27.2 x 7-11.6 vs. 5-12.5 x 2.3-7 cm), densamente pubescentes (vs. glabrescentes), botones florales mas

grandes (4.5-5 vs. 2-2.5 mm) y frutos densamente velutinos (vs. glabrescentes).

Etimologia .—Esta especie es nombrada en honor a Gerardo Rivera, en reconocimiento por sus in-

cansables esfuerzos de estudio, recolecta y valoracion de la flora costarricense.

Paratipos. COSTA RICA. Puntarenas: Canton de Golfito. Jimenez, Dos Brazos de Rio Tigre. Cerro Rincon, fila noroeste, 08 o 31'35"N,

83°28 , 12"W, 754 m, 28 Ago 1990 (fr), G. Herrera 4167 (CR, INB, MO). Canton de Osa. Piedras Blancas, La Florida, finca Bellavista, 08°

46,872'N-83° 12,483'W, 297 m, 30 Ago 2011 (fr), G. Rivera & M. Nunez 4557 (CR, MO, USJ). Reserva Forestal Golfo Dulce. Peninsula de Osa,

Bahia Chal, Los Mogos, 08°43 , 20"N, 83°26 , 30"W, 20 m, 21 Jun 1993 (bot. fls), R. Aguilar 2016 (CR).

Santamaria et al., Dos nuevas especies de Myrcia

455

AGRADECIMIENTOS

Agradecemos a Chris Davidson, Sharon Christoph, de hnca Playa Rare y la fundacion Blue Moon por su apoyo

al Proyecto de las Plantas Vasculares de la Peninsula de Osa, Costa Rica; a Matt Hogan por las facilidades y

apoyo brindado durante la investigacion realizada en su propiedad Finca Buenavista, ubicada en Piedras Blan¬

cas de Osa. Ademas a Luis Diego Vargas por las fotograflas de Myrcia paulii-jonesii y a Joaquin Sanchez del He-

bario Nacional de Costa Rica (CR) por su colaboracion en la elaboracion del mapa de distribucion de las espe¬

cies. A Ranchi Ghandi por el asesoramiento sobre la forma correcta de escribir el eplteto de Myrcia paulii-jone¬

sii y a Laura Lagomarsino por las traducciones al ingles. Tambien agradecemos a Bruce R. Holst y a Lucia Ra-

wasaki, por sus valiosos aportes y comentarios en la revision del manuscrito.

Daniel Santamaria agradece a los curadores y asistentes del herbario de la Universidad de Harvard por las

facilidades brindadas durante la estancia en dicho herbario; as! como al Missouri Botanical Garden por el

apoyo economico mediante la beca Elizabeth E. Bascom para Botanicas (os) Latinoamericanas (os), as! como a

todo el personal por las facilidades brindadas. Tambien agradecemos a los siguientes herbarios por permitir el

acceso a sus colecciones A, CR, F, GH, INB, LPB, MO, MOL, NY, PMA, USM.

REFERENCIAS

Holst, B.K. & M.L. Kawasaki. 2007. Myrcia DC. Myrtaceae. In: B.E. Hammel, M.H. Grayum, C. Herrera, & N. Zamora, eds.

Manual de plantas de Costa Rica. Vol. 6. Monogr. Syst. Bot. Missouri Bot. Gard. 111:764-768.

Holst, B.K. & M.L Kawasaki. 2009.177. Myrcia DC. Myrtaceae. In: G. Davidse, M. Sousa Sanchez & A.O. Chater, eds. FI. Me-

soamer. Universidad Nacional Autonoma de Mexico, Mexico, D.F. 4(1 ):66—150.

Lucas, E., K. Matsumoto, S. Harris, E. Nic Lughadha, B. Bernardini, & M. Chase. 2011. Phylogenetics, morphology, and evolution

of the large genus Myrcia s.l. (Myrtaceae). Int. J. Pl.Sci. 172(7):915-934.

McVaugh, R. 1968. The genera of American Myrtaceae: An interim report. Taxon 17:354-418.

McVaugh, R. 1969. Myrtaceae. In: B. Maquire & JJ. Wurdack, eds. The botany of the Guayana Highland. Part.VIII. Mem.

New York Bot. Gard. 18(2):55-286.

Nic Lughadha, E., K. Slade, L. Jennings, H. Boudet-Fernandes, & E. Lucas. 2010. Three new species of Myrcia section Gomidesia

(Myrtaceae)-from Espirito Santo, Brazil. Kew Bull. 65:21-28.

Parra-O., C. 2013. Una especie nueva de Myrcia (Myrtaceae) y nuevos registros de la familia para Colombia. Caldasia

35(2):293-298.

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Journal of the Botanical Research Institute of Texas 8(2)

BOOK REVIEW

L.B. (Bert) McCarty and David W. Hate. 2011. More Turfgrass and Related Weeds: Beyond the Color Atlas.

(ISBN-13: 978-0-9798777-5-9, pbk). Clemson University Public Service Publishing, Clemson University,

Clemson, South Carolina 29634-0129, U.S.A. (Orders: https://shopping.clemson.edu, 1-864-656-3311).

$25.00,380 pp., color photos, glossary, index, 7" x 10".

This is a guide for identifying weeds, ostensibly for those who are weed managers. For those interested in iden¬

tification of the variety of grasses, plants, and numerous Bowers in the wild, it gives a clear format of indices to

check.

The introduction provides a way to categorize a plant through its roots, stems, vascular systems, leaves,

flowers, fruits, seeds, etc. Each page names a plant within its family, the species, and also notes its growing

season and life cycle. Every plant has several close-up photos on the page which are illustrative of its habit and

other specific characteristics. The page also includes shape, size, propagation, and habitat along with its distri¬

bution in the U.S. (initiating primarily from Florida) and worldwide.

I found the photos to be very helpful as they provided a clearly recognizable vehicle for identification

when viewing a plant. The other details added credence when further distinguishing was needed. Within the

descriptive narrative there are often extra snippets of interest such as what plants are used for medicinal pur¬

poses or for teas, if edible or poisonous, and whether drought tolerant or invasive.

Since this reading I have identified plants which have heretofore gone unrecognized, and now provide me

with the knowledge to either keep or dispense with as per information gained.— Pat Gerard, Botanical Research

Institute of Texas Volunteer, Fort Worth, Texas, U.S.A.

J.Bot. Res. Inst. Texas 8(2): 456.2014

TAXONOMIC STUDIES IN THE MICONIEAE (MELASTOMATACEAE). XII.

REVISION OF MICONIA SECT. MICONIASTRUM,

WITH EMPHASIS ON THE MICONIA BICOLOR COMPLEX

Walter S. Judd

Department of Biology and

Florida Museum of Natural History

University of Florida

Gainesville, Florida, U.S.A.

wjudd@botany.ufl.edu

Eldis R. Becquer

Jardm Botanico Nacional

Universidad de La Habana,

Carretera "El Rocio" Km 3.5, Calabazar, Boyeros, CP 19230

La Habana, CUBA

erbecquer@fbio.uh.cu, pachyanthus@gmail.com

James Dan Skean, Jr.

Department of Biology

Albion College

Albion, Michigan 49224, U.S.A.

dskean@albion.edu

Lucas C. Majure

Department of Biology and

Florida Museum of Natural History

University of Florida

Gainesville, Florida, U.S.A.

Imajure@ufl.edu

ABSTRACT

A new section, Miconia sect. Miconiastrum, is recognized (and includes species previously placed in Calycogonium, Charianthus, Miconia,

and Tetrazygia). Within this clade, the species of the Miconia bicolor complex are revised. For the nine species of the M. bicolor complex,

descriptions, nomenclatural information (including four new names: Miconia karsticola, M. guajaibonensis, M. cajalbanensis, and M.

maestrensis), specimen citations, and eco-geographical characterizations are presented, along with an identification key to members of the

section. Miconia sect. Miconiastrum represents a monophyletic group within the Caribbean clade of tribe Miconieae (Melastomataceae) and

the group is restricted to southern Florida, the Bahamas, Cuba, and Hispaniola. The clade is characterized by hypanthia strongly constricted

in fruit, more or less reduced calyx teeth, obovate to spathulate petals, stamens dropping before the petals abscise, and leaves with globular-

stellate to stellate hairs or peltate scales on the abaxial surface. Most species possess mite domatia formed from a dense tuft of elongate,

multicellular, barbate, eglandular hairs in the axils of the midvein and major secondary veins, and they have seeds with smooth to slightly

bulging testa cells. Within sect. Miconiastrum a large subclade, i.e., the M. bicolor complex, is diagnosed by 5- or less commonly 6-merous

flowers with the petals abaxially papillose-granulose. All members of this clade, except for M. karsticola, also possess paniculate-cymose

inflorescences of numerous flowers and abaxial leaf surfaces usually densely covered with well-developed stellate hairs or peltate scales.

Within the M. bicolor complex, indumentum characters (as observed on the abaxial leaf surface), form of the calyx tube, size of the calyx

lobes, and presence/absence of mite domatia are taxonomically significant.

RESUMEN

Se reconoce una nueva seccion, Miconia secc. Miconiastrum (que incluye especies previamente conocidas bajo los generos Calycogonium,

Charianthus, Miconia, y Tetrazygia), y dentro de este clado se revisa la taxonomia de los miembros del complejo Miconia bicolor. Para las

nueve especies del complejo M. bicolor, presentamos descripciones, informacion nomenclatural (incluyendo cuatro nombres nuevos: Mico¬

nia karsticola, M. guajaibonensis, M. cajalbanensis, y M. maestraensis), citaciones de especimenes, caracterizaciones ecogeograficas y

una clave de los miembros de la seccion. Miconia sect. Miconiastrum representa un grupo monofiletico dentro del clado caribeno de la tribu

Miconieae (Melastomataceae) y el grupo es restringido al sur de la Florida, Fas Bahamas, Cuba y Fa Espanola. El clado se caracteriza por los

hipantos fuertemente constringidos en el fruto, los dientes del caliz ligera o extremadamente reducidos, los petalos de aovados a espatula-

dos, los estambres caducos antes de la absicion de los petalos, y las hojas con pelos estrellados, globular-estrellados, o escamas estrelladas

peltadas en el enves de la hoja. Fa mayoria de especies poseen acarodomacios formados por un mechon de pelos alargados, multicelulares,

barbados y eglandulares que se presentan en las axilas de la vena primaria con las venas secundarias mayores y tambien tienen semillas con

las celulas de la testa lisas y ligeramente hinchadas. Dentro de la sect. Miconiastrum, un subclado grande, ej. el complejo M. bicolor, se distin¬

gue por sus flores 5- o menos comun 6-meras, con los petalos abaxialmente papiloso-granulosos. Todos los miembros de este clado, excepto

M. karsticola, tambien poseen inflorescencias paniculadas de numerosas flores y el enves de la hoja cubierto por un indumento denso de

pelos estrellados, globular-estrellados o escamas peltadas bien desarrolladas. Dentro del complejo M. bicolor los caracteres del indumento

(del enves de la hoja), la forma del tubo del caliz, el tamano de los lobulos del caliz, y la presencia o ausencia de acarodomacios tienen un

significado taxonomico.

Keywords: Miconia, Tetrazygia, Miconia sect. Miconiastrum, Miconieae, Melastomataceae

J. Bot. Res. Inst. Texas 8(2): 457 - 491.2014

458

Journal of the Botanical Research Institute of Texas 8(2)

During the course of work on the systematics and phytogeny of Miconieae (Melastomataceae), we have become

aware of a number of taxonomic problems, especially in the species of the Greater Antilles. Recent phyloge¬

netic analyses (Becquer et al. 2008; Goldenberg et al. 2008; Martin et al. 2008; Michelangeli et al. 2008) showed

nearly all of the genera of Miconieae, including Miconia Ruiz & Pav., itself, to be highly polyphyletic, a result

that is in keeping with the history of problematic generic delimitations within the tribe (Cogniaux 1891; Glea¬

son 1925, 1932, 1958; MacBride 1941; Wurdack 1972, 1980; Judd 1986, 1989; Judd & Skean 1991). Alain

(1957), Proctor (1972), and Wurdack and Krai (1982) also have noted the problematic separation of Tetrazygia

from Miconia. As a result, major changes in generic circumscriptions are required. It is clear that nearly all cur¬

rently recognized genera, many with numerous Caribbean species, e.g., Calycogonium DC., Clidemia D. Don,

Leandra Raddi, Miconia, Ossaea DC., Pachyanthus A. Rich., and Tetrazygia Rich., are not supported. The most

workable solution to this classihcatory problem (as clearly evident in Figs. 1-3 of Goldenberg et al. 2008; or Fig.

1 of Michelangeli et al. 2008) is to place most species of Miconieae (including those treated in this paper)

within a greatly expanded Miconia, comprising the DNA-supported clade within Miconieae that can be diag¬

nosed by the synapomorphy of berry fruits (Goldenberg et al. 2013; Ionta et al. 2012; Ionta & Judd 2012; Ma-

jure & Judd 2012) and to conserve the large and megadiverse Miconia over the much smaller Tococa (Michel-

angeli, in prep.).

The authors are currently engaged in collaborative research with F. A. Michelangeli, F. Almeda, R. Gold¬

enberg, and G.M. Ionta on the systematics of tribe Miconieae focusing on the systematics and phylogeny of

Caribbean members of the tribe. In the course of this research we discovered, based on careful consideration

of both molecular and morphological evidence, a distinctive clade that is most diverse in Cuba but also occurs

on Hispaniola, Jamaica, the Bahamas, and in southern Florida. The purpose of this paper is to characterize this

group of nine species, i.e., the Miconia bicolor complex, and to place this clade within the more inclusive clade

(comprising ca. 11 species), here characterized morphologically and named Miconia sect. Miconiastrum (Bonpl.

ex Naudin) Judd, Becquer & Majure.

The monophyly of Miconia sect. Miconiastrum is supported both by morphology and DNA-sequence data

(Goldenberg et al. 2008; Michelangeli et al. 2008, who included in their analyses the species M. barbata (Bo-

rhidi) Judd, Becquer & Majure, M. bicolor (Mill.) Triana, M. cajalbanensis Judd, Becquer & Majure (as Tetra¬

zygia coriacea Urb.), M. cristalensis (Borhidi) Judd, Becquer & Majure, M. guajaibonensis Judd, Becquer & Ma¬

jure (as T. lanceolata Urb.). Additionally, Majure et al. (2014) included the species M. angustiflora (Benth.)

Naudin (as T. pollens (Spreng.) Cogn.), M. karsticola Judd, Becquer, Skean & Majure (as Calycogonium saxicola

Britton & P. Wilson), of the M. bicolor complex, and the basally diverging M.fadyenii (Hook.) Judd & Skean (as

T.fadyenii Hook.) and showed high support for the clade (bootstrap = 96). Michelangeli et al. and Majure et al.

(unpubl. data) have also included the two subsequent sister species of the clade, Miconia delicatula A. Rich, and

Clidemia swartzii Griseb., and recovered the clade with high support. Putative synapomorphies of this clade

include the constricted hypanthium (in fruit; Figs. ID, 3E, 6D), more or less reduced calyx teeth (Figs. 1A, B,

3A. D. E. 6A-D), obovate/spathulate petals (Figs. 1,3, 6), and the stamens dropping before the petals (Figs. IB,

3A, 6A-C). The flowers range from 4- to 6-merous. Preliminary DNA-based phylogenetic analyses (Michel-

angeli et al., unpublished) suggest that Miconia delicatula (Fig. 1A) is sister to the remaining members of the

clade, and the characteristic mite domatia, consisting of a tuft of elongate, multicellular, eglandular, barbate

hairs in the axil of the midvein and major secondary veins, likely are synapomorphic for all members of the

section (see Figs. 1C, 6A) except for M. delicatula (although they have been lost in a few species, e.g., M. cris¬

talensis and M. maestrensis Judd, Becquer & Majure; Fig. 6D). All have stellate hairs, although polarities regard¬

ing hair form are unclear. The hairs of M. fadyenii and M. karsticola are quite reduced, while those of M. deli¬

catula are large and globular-stellate. Well-developed, usually dense, and more or less flattened, stellate hairs

(with arms radiating mainly parallel to the epidermal surface, thus termed “stellate” following nomenclature

of Wurdack 1986) or peltate scales (“lepidote” hairs in the terminology of Wurdack 1986) are characteristic of

the remaining species. Miconia fadyenii likely was the next species to diverge (possibly along with Clidemia

swartzii), and this putative clade is sister to the M. bicolor complex (Michelangeli et al., unpublished analyses).

Judd et al., Revision of Miconia sect. Miconiastrum

459

Miconia fadyenii (Fig. IB) is especially distinctive because of its adaptations for hummingbird pollination, and

for this reason has often been placed in Charianthus D. Don (see for example, Proctor 1972), but the close rela¬

tionship of this species to M. bicolor was first noted by Penneys and Judd (2003, 2005). Within sect. Miconias¬

trum the species of the M. bicolor complex, here studied in detail, can be diagnosed by their 5- or 6-merous

flowers with papillose to granulate petals (either abaxially, or both adaxially and abaxially). This morphologi¬

cally cohesive subclade is also strongly supported by molecular data (Becquer et al. 2008; Goldenberg et al.

2008; Michelangeli et al. 2008, and unpublished analyses) and includes M. angustiflora, M. barbata, M. bicolor,

M. cajalbanensis, M. cristalensis, M. guajaibonensis, M. impressa (Urb.) Judd, Becquer & Majure, M. karsticola,

and M. maestrensis.

As noted above, Clidemia swartzii may also belong in Miconia sect. Miconiastrum, but this Jamaican en¬

demic, which has axillary, 4-merous flowers and large leaves with numerous elongate, multicellular, eglandu-

lar hairs along with minute globular-stellate hairs, has none of the putative morphological synapomorphies of

this clade. Its placement in sect. Miconiastrum, therefore, requires confirmation through additional DNA

samples, as well as analyses including additional species. We note that M. fadyenii also is 4-merous, and this

character maybe a synapomorphy of a hypothesized C. swartzii + M. fadyenii clade.

Although not included in the DNA-based analysis of Goldenberg et al. (2008) and Michelangeli et al.

(2008), the rare Cuban endemic, Miconia ancistrophora (C. Wright) Triana, is similar to M. delicatula in its de¬

ciduous habit, flowers with distinctly clawed petals and stamens often forming two groups, and like M. deli¬

catula it may belong in sect. Miconiastrum. This hypothesis is supported by several morphological characters:

its constricted hypanthium, short calyx teeth, obovate petals, and subulate stamens with a small dorsal pore.

We note that its flowers are 4- or 5-merous, the ovaries also 4- or 5-locular, and the seeds somewhat different

from those of other species of sect. Miconiastrum, i.e., they are 1.3-2.7 mm long and the raphe is broad and

deeply sunken (Becquer, unpublished data). Miconia ancistrophora is easily distinguished from all the species

here placed in sect. Miconiastrum by the bulla-based hairs on its adaxial leaf surfaces. However, definitive

placement of this species must await molecular investigation.

Miconia bicolor, the most common and broadly distributed species in the M. bicolor complex, was the first

to be described, as Melastoma bicolor Miller (1768), and this species was transferred to Tetrazygia by Cogniaux

(1891). The Jamaican species, M. angustiflora, was next described, as Diplochita angustiflora Bentham (1846),

although this taxon long has been known under the misapplied name Melastoma pollens Sprengel (1822; trans¬

ferred to Tetrazygia pollens (Spreng.) Cogniaux in 1891). Miconia maestrensis was described next, as Miconias¬

trum lambertianum Bonpl. ex Naudin (1850), and occurs in the Sierra Maestra region, Cuba. This taxon was

long misunderstood and usually has been considered within synonymy (under either M. bicolor or M. angusti¬

flora). The Pinar del Rio, Cuba, endemic, Miconia karsticola was described by Britton and Wilson (1920, as Ca-

lycogonium saxicola) and placed in Calycogonium (instead of Tetrazygia) because of its reduced inflorescences

and small leaves, which convergently suggested a relationship with species such as C. glabratum (Sw.) DC., i.e.,

the glabrate-domatial clade (Judd & Skean 1991) within Calycogonium. The glabrate-domatial clade, however,

has 4-merous flowers and differently formed domatia, i.e., the hairs are usually non-barbate and connate.

Then, in a major paper, Urban (1926) described the narrowly endemic Cuban species M. guajaibonensis (as

Tetrazygia lanceolata), M. impressa (in which we include T. versicolor Urb. and T. minor Urb., also described in

the same publication) and M. cajalbanensis (as T. coriacea). Lastly, Borhidi (1977) described the eastern Cuban

endemics M. cristalensis (as T. cristalensis Borhidi) and M. barbata (as T. barbata Borhidi).

This brief account clearly demonstrates that these species usually have been treated within Tetrazygia;

they are considered within this genus in all Antillean or Florida floras (Small 1913; Urban 1920-21; Barker &

Dardeau 1930; Moscoso 1943; West & Arnold 1956; Alain 1957; Proctor 1972; Long & Lakela 1978; Correll &

Correll 1982; Liogier 2000; Nelson 2011; Wunderlin & Hansen 2011; Michelangeli & Becquer 2012) except for

the phenetically divergent, M. karsticola, which has been placed in Calycogonium (Alain 1957; Michelangeli &

Becquer 2012). Not only have these species been placed in Tetrazygia, many have been confused with the wide¬

spread and variable T. bicolor (as evidenced by frequently misidentihed specimens) and considered to be

closely related (Cogniaux 1891; Urban 1926; Proctor 1972).

460

Journal of the Botanical Research Institute of Texas 8(2)

Within the M. bicolor complex, Miconia karsticola (Fig. 1C) is probably sister to the remaining species

(Michelangeli et al., unpublished analyses). It differs from them in having very reduced inflorescences, smaller

leaves, a shorter habit, the much smaller stellate hairs on the abaxial leaf surface, and flowers with a shorter

hypanthium and more conspicuous calyx teeth (see key). The remaining species are hypothesized to form a

clade, with the putative morphological synapomorphy of leaves with a dense indumentum of stellate hairs (or

peltate scales). Among these species, those with an abaxial leaf indumentum of stellate hairs, e.g., M. angusti-

flora, M. barbata, M. cristalensis, and M. maestrensis (Fig. 6A, B, D), are likely early divergent, while M. bicolor

and M. guajaibonensis (Figs. ID, 3A-D) are presumably derived (and closely related) as suggested by their in¬

dumentum of stellate-peltate scales. The scales of these two species may have evolved from stellate hairs

through the fusion of the individually radiating arms of the hairs.

Miconia sect. Miconiastrum and the M. bicolor complex are restricted to the Caribbean region, occurring

only on Cuba, Jamaica, Hispaniola, the Bahamas, and in southern Florida. Miconia sect. Miconiastrum is most

diverse in Cuba, which has nine species, eight of these in the M. bicolor complex, and all endemic except for M.

bicolor, itself, which is widespread (occurring on Cuba, Hispaniola, the Bahamas, and in southern Florida; Fig.

4). Jamaica is next with two species, i.e., M. angustiflora of the M. bicolor complex, and the early divergent M.

fadyenii, both endemic to the island. Hispaniola, southern Florida, and the Bahamas each contain only a single

species, the widespread M. bicolor. It is of interest that M. bicolor does not occur in Jamaica, which instead has

the phenetically similar M. angustiflora that has sometimes been considered conspecihc. Miconia bicolor also is

not common in the “Oriente” region of Cuba, where M. cristalensis, M. barbata, and M. maestrensis, all of which

also have been confused with M. bicolor, are restricted. However, a few problematic collections (including the

type of Tetrazygia acunae Borhidi) from the provinces of Guantanamo, Holguin and Santiago de Cuba are ten¬

tatively placed within this species (see discussion under M. bicolor). The region of highest species diversity is

the Prov. of Pinar del Rio, Cuba, in which five species of the M. bicolor complex occur: M. bicolor, M. cajalbanen-

sis, M. guajaibonensis, M. impressa, and M. karsticola (Figs. 2, 4, 5). All of these, except for M. bicolor, are en¬

demic to the province. We note that M. cajalbanensis, M. guajaibonensis, and M. karsticola occur in areas of high

endemism (Fig. 5): Pan de Guajaibon, the Cajalbana region, and the Sumidero region, respectively (Borhidi &

Muniz 1986; Borhidi 1991). For information on the vegetation and floristics of the plant communities in which

these species occur the reader should consult Alain (1946), Berazaln (1987), Borhidi (1991), Carabia (1945),

Borhidi & Muniz (1986), Seifriz (1943), Smith (1954), and Vasquez etal. (2006) for Cuba; Ciferri (1936), Garcia

and Mejia (2008), Hager & Zanoni (1993), Lorenzo et al. (1997), Mejia et al. (2011), and Zanoni et al. (1990) for

Hispaniola; Coker (1905), Correll and Correll (1982), Smith et al. (1992), and Nickrent et al. (2008) for the Ba¬

hamas; and Alexander (1967), Myers and Ewel (1990), Olmsted et al. (1983), and Phillips (1940) for Florida.

As stated previously, molecular analyses (Majure et al., unpublished data; Michelangeli et al., unpublished

data) support the hypothesis that Miconia delicatula, an endemic to Pinar del Rio, Cuba, is sister to the remain¬

ing species of sect. Miconiastrum. Miconia karsticola, another Pinar del Rio endemic, is sister to the remaining

species of the Miconia bicolor complex. These patterns suggest that sect. Miconiastrum originated in western

Cuba, where it is still most diverse, and dispersed twice to Jamaica: an early dispersal event leading to M. fady¬

enii (and possibly also Clidemia swartzii, if this species actually belongs in the clade) and a later event resulting

in M. angustiflora, along with other dispersal events to central or eastern Cuba. Some of these dispersal events

may have occurred earlier and resulted in M. cristalensis, M. barbata and probably also M. maestrensis, all of the

“Oriente” region, and some later, involving M. bicolor, which occurs in central Cuba, (especially in Prov. Villa

Clara and Sancti Splritus), and sparingly also in the “Oriente” region (Prov. Guantanamo, Holguin, Santiago de

Cuba). Of course, M. bicolor also has dispersed to Hispaniola, where the plants are distinctive in having consis¬

tently erect, dendritic to globular-stellate hairs or scales on the stems, and also southern Florida and the Baha¬

mas, where the plants have stems covered with more or less appressed stellate-peltate scales.

Chromosome counts are known from only two species, M. bicolor and M. angustiflora, and these have n =

17, which is the most common number in the tribe (Solt & Wurdack 1980).

Judd et al., Revision of Miconia sect. Miconiastrum

461

MEASUREMENTS, TERMINOLOGY, METHODS, AND SPECIES CONCEPTS

Nearly 700 specimens of the Miconia bicolor complex (of sect. Miconiastrum) were examined for this study. The

second author has collected M. barbata, M. bicolor, M. cajalbanensis, M. cristalensis, M. guajaibonensis, M.

karsticola, and M. maestraensis, while the first and third authors have collected M. bicolor and M. angustiflora

(see specimens examined). Herbaria from which specimens were studied are listed in the Acknowledgments.

Abbreviations follow Thiers (2012). Acronyms and citation of collection numbers for Cuban specimens follows

Regalado et al. (2010). One hundred and forty-three different characters were measured or observed for each

species.

All measurements included in the key and descriptions of species were taken directly from dried materi¬

al, with the exception of plant height and flower/fruit color, which were taken from information on specimen

labels, observed in the held, or in photographs, and floral measurements, which were taken from rehydrated

material. Colors of floral structures, placement of petals and stamens, and calyx tube length were all assessed

at anthesis. Terminology and measurement of floral parts follow Judd (2007), Judd & Ionta (2013), and Judd et

al. (2014).

Species delimitations were based on the morphological-phenetic species concept (Judd 2007) and the

diagnostic species concept (Wheeler & Platnick 2000). Finally, the lead researchers in our Planetary Biodiver¬

sity Inventory, N.S.F. grant, i.e., Fabian Michelangeli, Frank Almeda, Renato Goldenberg, and Walter Judd,

have agreed that it is most appropriate, at this time, to treat these species within an expanded Miconia.

TAXONOMIC TREATMENT

Miconia sect. Miconiastrum (Bonpl. ex Naudin) Judd, Becquer, & Majure, comb. nov. Miconiastrum Bonpl. exNau-

din, Ann. Sci. Nat., Bot. ser. 3, 15:341. 1850. Tetrazygia sect. Miconiastrum (Bonpl. ex Naudin) Cogn., in A. de Candolle & C. de

Candolle, Monogr. Phan. 7:724. 1891. Type: Miconia maestrensis Judd, Becquer, & Majure, basionym Miconiastrum lambertianum

Bonpl. ex Naudin.

Naudinia A. Rich, in R. de la Sagra, Hist. Phys. Cuba, Pi. Vase. 10:266.1846. Lectotype, here designated: Naudinia chrysophylla A. Rich. (=

Miconia bicolor (Mill.) Triana).

Evergreen or deciduous shrubs; young stems ± rectangular to terete in cross section, lacking longitudinal

ridges, the indumentum of moderate to dense, ± ferrugineous, dendritic to globular-stellate or stellate hairs,

these sometimes very reduced (in M.fadyenii and M. karsticola), or peltate scales, and rarely also with elongate,

eglandular, multicellular hairs (in M. bicolor var. patenti-setulosa). Leaves ± isophyllous, the blade ovate to ob-

ovate or oblong, coriaceous to chartaceous, the apex attenuate to obtuse, the base attenuate, cuneate, or acute

to rounded or cordate, the margin entire to undulate; secondary veins 2 pairs, basal, the tertiary veins percur-

rent, oriented subperpendicular to midvein, connected by quaternary veins, or separated by composite inter-

tertiary veins, the midvein, major secondary veins flat to moderately impressed, tertiary veins flat to slightly

impressed on adaxial surface; the midvein strongly raised, the major secondary veins moderately to slightly

raised, tertiary and higher order veins slightly raised to flat on abaxial surface; adaxial surface with scattered to

abundant druse crystals just below surface, the indumentum initially of globular-stellate or stellate hairs, or

stellate-peltate scales, but usually quickly glabrescent; abaxial surface moderately to densely covered with fer¬

rugineous to white, or pink, globular-stellate or stellate hairs, these sometimes minute, or with moderate to

dense, ± ferrugineous stellate-peltate scales; mite domatia consisting of a tuft of elongate, ± barbate, eglandular,

multicellular hairs in the junction of midvein and major secondary veins (and occasionally in other vein axils)

in most species (but entirely absent in M. cajalbanensis, M. cristalensis, M. delicatula, and M. maestrensis). Inflo¬

rescences terminal (but also axillary in M. delicatula), paniculate cymes of numerous flowers (but reduced to a

solitary flower or a 3-flowered cyme in M. karsticola), with shoots producing several nodes before becoming

reproductive. Flowers 5- or 6-merous (but 4-merous in M.fadyenii). Hypanthium terete, constricted above the

ovary (at least in fruit), the free portion ± flaring, the outer surface with scattered, globular-stellate or stellate

hairs, or peltate scales, an androecial fringe absent. Calyx lobes valvate or open (except in M. guajaibonensis,

462

Journal of the Botanical Research Institute of Texas 8(2)

where imbricate), separate in bud (but ± fused as a calyptra in M. impressa), the tube short, sometimes tearing

between the calyx lobes, which are triangular to nearly obsolete; calyx teeth short to obsolete, usually shorter

than the calyx lobes (but longer in M. karsticola), ± terete. Petals ± obovate, slightly to clearly asymmetric, usu¬

ally spreading (but often ± erect in M. fadyenii), white (but red in M. fadyenii) at anthesis, the apex rounded,

sometimes slightly asymmetrically notched, the base slightly narrowed to a broad attachment (abruptly nar¬

rowed or clawed in M. delicatula), the adaxial and abaxial surfaces smooth to granulate, the margin entire.

Androecium zygomorphic due to stamens positioned all on one side of flower (most species), in two groups

(often in M. delicatula), or actinomorphic (in M. fadyenii); stamens twice the number of petals, dropping before

the petals, isomorphic, geniculate; staminal filament glabrous, but often papillose, anther thecae subulate,

straight to curved, opening by a small dorso-terminal pore, at anthesis yellow, without a dorso-basal append¬

age, the connective not basally projecting (except in M. fadyenii). Ovary 3-locular (but 2-locular in M. fadyenii),

2/3-4/5-inferior, with conical to cylindric-conical apex, with a short, slightly ridged collar, but lacking crown,

with axile placentation, the ovules numerous, borne on slightly to strongly expanded placenta, style curved

distally, stigma punctate to truncate, minutely papillose. Berries globose to ellipsoid, purple-black at maturity.

Seeds angular-obovoid, with smooth, ± rectangular, non-bulging raphe (but broad and deeply sunken in M.

delicatula), the testa smooth to minutely roughened due to bulging cells; appendage absent.

At this time we can confidently recognize eleven species in this section (although a few specimens from

the “Oriente” region of Cuba are only tentatively placed, and more held work may result in the recognition of

additional species). All eleven species are included in the following key. However, only the species of the Mico-

nia bicolor complex are revised here, and these nine species are numbered in the key. A detailed taxonomic

treatment of Miconia fadyenii (including description, nomenclature, and specimen citations) has been pre¬

sented by Penneys and Judd (2003; see also Fig. IB here). A description (and photograph) of M. delicatula can

be found in Alain (1957; see also Fig. 1A here).

KEY TO THE SPECIES OF THE MICONIA BICOLOR COMPLEX

AND THE RELATED M. FADYENII AND M. DELICATULA

1. Inflorescences terminal and axillary; plants deciduous, blooming when leaves of the previous season have abscised and

new leaves are just flushing out_ Miconia delicatula

1. Inflorescences exclusively terminal; plants evergreen.

2. Petals 4, bright red, abaxially smooth, erect to slightly spreading; stamens 8; ovary 2-locular, with placentas deeply

intruded_ Miconia fadyenii

2. Petals 5 or 6, white, abaxially ± granulate, spreading; stamens 10 or 12; ovary 3-locular, with placentas shallowly

intruded_ Miconia bicolor complex [species 1-9]

3. Leaves 0.8-3.7 cm long; stellate hairs on abaxial leaf surface minute-globular, the hairs 0.02-0.08 mm across; inflo¬

rescences of 1 -3 flowers; low shrub to 1 m; hypanthium 2.8-3.5 mm long; calyx teeth usually ± conspicuous, 0.6-3

mm long_ 1. Miconia karsticola

3. Leaves usually longer than 3 cm; hairs on abaxial leaf surface peltate scales or obviously stellate, the hairs 0.07-0.28

across; inflorescences paniculate cymes of numerous flowers; shrub to tree, to 1.5-10 m tall; hypanthium 4.5-8 mm

long; calyx teeth usually inconspicuous, 0.1 -0.5(-1) mm long.

4. Hairs on abaxial leaf surface stellate-peltate, i.e., the arms ± connate (nearly completely to predominantly so)

and forming a peltate head.

5. Calyx lobes imbricate in young bud, unequal, i.e., the 2 outer lobes larger than the rest, and lobes always de¬

veloped; calyx tube irregularly tearing between lobes; leaves small, the lamina usually less than 8.5 cm long;

petals 4.5-6.3 mm long; Pinar del Rio, at Pan de Guajaibon_ 3. Miconia guajaibonensis

5. Calyx lobes valvate or open in young bud, equal, i.e., all ± the same size, and usually inconspicuous or nearly

obsolete; calyx tube not tearing between lobes (rarely with a few irregular tears); leaves larger, the lamina

usually more than 8 cm long; petals 5-11.5 mm long; plants of Cuba, U.S.A. (Florida), the Bahamas, and

Hispaniola_ 2. Miconia bicolor

4. Hairs on abaxial leaf surface stellate, i.e., the arms not connate or only slightly so, with longer arms proximally

and short to extremely short ones distally.

6. Leaf domatia present (and formed by elongate, barbed, multicellular, eglandular hairs) at junction of the 2

major secondary veins and the midvein.

7. Flower buds with a nearly closed, conical calyptra and the petals thus hidden; calyx tube always regularly

to irregularly tearing between the lobes; calyx lobes 0.15-0.5 mm wide (but appearing broader due to

tearing of the tube); hairs of abaxial leaf surface white to very pale ferrugineous; plants of Pinar del Rio

_4. Miconia impressa

Judd et al., Revision of Miconia sect. Miconiastrum

463

7. Flower buds apically open and the petals exposed; calyx tube usually not tearing at maturity, but occasion¬

ally with a few tears between lobes; calyx lobes 1.5-3.5 mm wide; hairs of abaxial leaf surface ± ferrugine-

ous; plants of Jamaica or the Oriente region, Cuba.

8. Domatia usually well developed, conspicuous, with elongate hairs densely distributed along 1-11 mm

of midvein distal to point where joined by major secondary veins (and sometimes also at junctions with

primary and tertiary veins), and all plants with domatia; plants of the central and northern "Oriente" region,

i.e., Sierra de Nipe, mountains near Bayate, Monte Verde, and the vicinity of Moa and Baracoa, Cuba

_5. Miconia barbata

8. Domatia not well developed, usually inconspicuous, with elongate, ± sparse hairs and distributed along

0.25-3(-4) mm of midvein distal to point where joined by major secondary veins, and some plants

entirely without domatia; plants of Jamaica_ 6. Miconia angustiflora

6. Leaf domatia absent.

9. Flower buds with a nearly closed, conical calyptra and the petals thus hidden; calyx tube always regularly

to irregularly tearing between the lobes; calyx lobes 0.15-0.5 mm wide (but appearing broader due to

tearing of the tube); anthers 4.5-5 mm long; hairs of abaxial leaf surface ± white; plants of Pinar del Rio

_4. Miconia impressa

9. Flower buds apically open and the petals exposed; calyx tube usually not tearing at maturity; calyx lobes

1-4 mm wide; anthers 4.8-8.3 mm long; hairs of abaxial leaf surface variously ferrugineous; plants of Ja¬

maica, Pinar del Rio, or the "Oriente" region of Cuba.

10. Abaxial epidermis of lamina distinctly bullate due to the bulging of individual epidermal cells; testa

roughened, with bulging cells; calyx tube 2-3.3 mm long; plants of Pinar del Rio_ 7. Miconia cajalbanensis

10. Abaxial epidermis of lamina ± smooth; testa smooth to roughened, the cells flat or ± bulging; calyx tube

0.8-2 mm long; plants of Jamaica or the "Oriente" region of Cuba.

11. Hairs of abaxial leaf surface with arms predominantly radiating outward, parallel with the lamina

surface, the hairs sparse to dense, and epidermis often visible; plants of Jamaica_ 6. Miconia angustiflora

11. Hairs of abaxial leaf surface with arms pointing upward to radiating outward, the hairs ± dense, so

epidermis obscured; plants of the Sierra de Cristal and Sierra Maestra, Cuba.

12. Calyx lobes 1.7-3.5 mm long; leaves with tertiary veins raised abaxially; plants of the Sierra de

Cristal_ 8. Miconia cristalensis

12. Calyx lobes 0.3-0.6 mm long; leaves with tertiary veins flat to slightly raised abaxially; plants of

the Sierra Maestra_ 9. Miconia maestrensis

1. Miconia karsticolajudd, Becquer, Skean, & Majure, nom. nov. (Fig. 1C). Calycogonium saxicola Britton & P. Wilson,

Mem. Torrey Bot. Club 16:91.1920, non M. saxicola Brandegee, Zoe 5:215.1905. Type: CUBA, Prov. Pinar del Rio: S of Sumidero, Si¬

erra Caliente, 15, 18 Aug 1915, fl, J.A. Shafer 13770 (holotype: NY!; isotypes: A on-line image #00072013!, CAS on-line image

#0002427!, NY!, US!).

Evergreen shrub up to 1 m. Young stems terete to slightly quadrangular, the indumentum of moderate, ferru¬

gineous, globular-stellate hairs, without elongate, multicellular, non-glandular hairs, internodes 0.5-3 cm

long, lacking longitudinal ridges, nodal line absent. Leaves isophyllous or nearly so; petiole 0.3-2 cm long, the

indumentum of moderate, minute, ferrugineous, globular-stellate hairs; the blade 0.8-3.7 x 0.35-2 cm, ovate to

elliptic or obovate, thinly coriaceous, the apex acute to obtuse or shortly acuminate, the base cuneate or obtuse

to rounded, the margin plane, entire; secondary veins a single pair, conspicuous to inconspicuous, acrodro-

mous, ± basal, the pair joining midvein at base to 1.5 mm above the leaf base, the secondary veins placed 0.5-

3.5 mm in from margin, tertiary veins percurrent, oriented subperpendicular to midvein, 1-4.5 mm apart,

obscure, the higher order veins also obscure, the midvein and secondary veins slightly impressed to flat, ter¬

tiary and higher order veins flat on adaxial surface; the midvein slightly to moderately raised, the secondary

veins slightly raised to flat, the tertiary and higher order veins flat on abaxial surface; adaxial surface appearing

slightly wrinkled after drying, with scattered druse crystals, and drying only slightly darker than the abaxial

surface, the indumentum initially of scattered, ferrugineous, minute, globular-stellate hairs, but very quickly

glabrescent; abaxial surface pale green, the surface smooth, with moderately scattered, ferrugineous, globular-

stellate hairs, 0.02-0.08 mm across (appearing under low magnification as punctations), the veins with similar

hairs, such hairs on lamina and veins persistent; mite domatia in axils of midvein and secondary veins, consist¬

ing of elongate, eglandular, smooth to slightly barbate, ± ferrugineous hairs, 0.1-0.5 mm long (often associated

with a distinct depression in the leaf surface). Inflorescences terminal, a 1-3-flowered cyme, 1.3-3 cm long,

0.8-2 cm across, with no major branch pairs, the peduncle 0.4-1.7 cm long, the ultimate axes (pseudopedicels)

1.5-6 mm long, and flowers (when more than one present) well separated from each other; bracts 1.5-1.8 x

0.15-0.2 mm, narrowly triangular, with acute apex, deciduous; bracteoles 0.4-1 x 0.1-0.15 mm, narrowly

464

Journal of the Botanical Research Institute of Texas 8(2)

Fig. 1. A. Miconia delicatula {Becquer& Abbott8242 1). B. M. fadyenii {Judd5300). C. M. karsticola {Ekman 18182). D. M. bicolor (fruiting plant, from Long

Pine Key, Everglades National Park, Miami-Dade Co., Florida).

triangular, with acute apex, ± deciduous. Flowers 5-merous, zygomorphic (due to positioning of the stamens),

with pedicel 1.4-4 mm long. Hypanthium 2.8-3.5 mm long, terete and slightly flaring, not constricted above

ovary, the free portion 1.5-1.7 mm long, 2.5-3 mm wide at the torus, the outer surface with sparse to moderate,

minute, globular-stellate hairs, the internal surface ± smooth, glabrous. Calyx lobes 5, separate in bud, the tube

1-1.5 mm long, not or sometimes slightly tearing between lobes, with sparse minute, globular-stellate hairs

Judd et al., Revision of Miconia sect. Miconiastrum

465

abaxially, and glabrous adaxially, the lobes 0.2-0.4 x 1.5-3 mm, broadly triangular to nearly obsolete, the apex

obtuse to rounded, with hairs similar to those of the tube but more sparsely distributed, green to red-tinged;

calyx teeth present, 0.6-3 mm long, ± triangular to conic-terete, green to red, with acute apex, and hairs similar

to those of calyx. Petals 5, 7-10 x 5-6.5 mm, asymmetrically obovate, spreading, white, glabrous but abaxial

surface densely papillose-granulate, the apex rounded, sometimes slightly notched, the base slightly narrowed

to a broad attachment, the margin entire. Stamens 10, isomorphic, ± geniculate near filament apex; staminal

filament 4.7-5.3 mm long, glabrous, pale yellow, anther thecae 4.3-4.7 x ca. 1 mm, subulate, straight to slightly

incurved, opening by a small dorso-apical pore, yellow, the connective not prolonged below the thecae, gla¬

brous, but the filament emerging from slight depression in sterile, minutely dorso-lobed anther base. Ovary

3-locular, 1/2-2/3 inferior, globose to shortly ovoid, 2.5-3 mm long, 1.4-2.5 mm in diameter, apically conical

to cylindric-conical, glabrous, the apex with small, ridged collar but lacking crown, with axile placentation,

the ovules numerous, borne on small placenta; style 7-9.5 mm long, distally curved, white, glabrous; stigma

punctate, minutely papillose, 0.1-0.2 mm wide. Berries 4-9 mm long, 4.5-9 mm in diameter, subglobose,

purple-black, with scattered, minute, globular-stellate hairs, the hypanthium constricted, 2.5-3.3 mm wide at

narrowest point and 2.5-3.7 mm wide at torus. Seeds 1-1.6 mm long, angular-obovoid, with ± rectangular

raphe, the testa ± smooth; appendage absent.

Phenology. —This poorly collected species is only known in flower from May, August and November.

Distribution and habitat.—Miconia karsticola is endemic to the vicinity of Sumidero in Pinar del Rio, Cuba

(Fig. 2), occurring in rocky areas and cliffs, at ca. 70 m, on limestone.

Additional specimens examined: CUBA. Prov. Pinar del Rio: Minas de Matahambre, Sumidero, Cueva del Resolladero en el valle de Pica

Pica, 19 May 1984, Alvarez et al. HFC-54676 (HAJB); Minas de Matahambre, Salon de los Gigantes, cerca del resollarero del rio Sumidero, 70

m, 4 Apr 2008, Becquer et al. HFC-85175 (HAJB); Minas de Matahambre, Sumidero, Valle de Pica Pica, alrededores de Cueva Clara, 18 Nov

1978, Bisse et al. HFC-38804 (HAJB); Sumidero, in Sierra Caliente, 24 Nov 1923, Ekman 18182 (S, US); Loma Arenales, S of Sumidero, 20 Aug

1912, Shafer 13797 (US); Paredon calizo de sombra, a la entrada de la Cueva de los Gigantes, Sumidero, 18 May 1991, Urquiola et al. HPR-7569

(HAJB).

Miconia karsticola is easily distinguished from the other members of M. bicolor complex (within Miconia sect.

Miconiastrum ) by the characters presented in the key. It is especially distinctive because of its reduced inflores¬

cences, flowers with short hypanthia, and small leaves, and these characters led to its being treated within the

variable and polyphyletic genus Calycogonium. However, its constricted hypanthium/calyx in fruit, the leaves

with mite domatia formed by clustered elongate, multicellular, eglandular, at least sometimes barbate hairs in

the axils of the midvein and major secondary veins, hve-merous flowers with 3-loculate gynoecia, abaxially

granulose petals and seed features argue for a relationship with Miconia bicolor and its close relatives (in sect.

Miconiastrum). A phylogenetic relationship with M. bicolor and relatives also is supported by DNA sequence

data (Michelangeli et al., unpublished data).

The new name, M. karsticola, is required because the epithet “saxicola” is already occupied in Miconia

(i.e., M. saxicola Brandegee, a Mexican species). The epithet “karsticola” refers to the floristically diverse, karst-

ic habitat of this species and also echoes the original specific epithet. The flora in the vicinity of Sumidero, in

the Sierra de los Organos, is an important center of speciation, possessing ten local endemics (Borhidi & Mu¬

niz 1986; Borhidi 1991). This species is considered vulnerable (VU) by Borhidi and Muniz (1983) and included

in this category in the Red Tist of Cuban Flora (Berazaln et al. 2005) due to its limited geographical range.

2. Miconia bicolor (Mill) Triana, Trans. Finn. Soc. Tondon 28:103. 1872.

Evergreen shrub or tree to 10 m. Young stems terete to slightly quadrangular, the indumentum of moderate to

dense, ± ferrugineous, peltate scales, or globular-stellate to dendritic hairs, these persistent to ± deciduous and

stems then glabrescent, with or without elongate, multicellular, non-glandular hairs, internodes 0.8-8.5 cm

long, lacking longitudinal ridges, nodal line present, faint to well-developed transverse ridge. Leaves isophyl-

lous or nearly so; petiole 1.2-4.7 cm long, the indumentum of dense, peltate scales, but sometimes glabrescent;

the blade 5-21.2 x 1.4-6.6 cm, ovate to elliptic or oblong, coriaceous to thinly coriaceous, the apex acuminate

to attenuate, the base obtuse to rounded or very slightly cordate, the margin plane to slightly revolute, entire to

466

Journal of the Botanical Research Institute of Texas 8(2)

slightly undulate; secondary veins 2 pairs, one pair conspicuous and one pair inconspicuous, acrodromous,

basal or suprabasal, the innermost pair joining midvein at base to 15 mm above the leaf base, the conspicuous

secondary veins placed 1.5-12 mm in from margin, the inconspicuous secondary veins intramarginal to 1.8

mm in from margin, tertiary veins percurrent, oriented subperpendicular to midvein, 2-12 mm apart, con¬

nected by 1-3 quaternary veins, which are ± reticulate, or not connected by quaternary veins, and separated by

composite inter-tertiary vein, the higher order veins reticulate, the midvein and major secondary veins hat to

slightly impressed, tertiary veins hat to slightly impressed, remaining veins hat on adaxial surface; the midvein

strongly raised, the major secondary veins moderately to slightly raised, the minor secondary veins and ter¬

tiary veins slightly raised to hat, and the higher order veins hat on abaxial surface; adaxial surface appearing

slightly wrinkled or punctate after drying, with scattered druse crystals, and drying darker than the abaxial

surface or sometimes with a distinctive yellowish color when dried, the indumentum initially of dense, pale

ferrugineous to translucent, stellate-peltate scales, but quickly glabrescent, although often with a few hairs

proximally on midvein, or occasionally even scattered on the surface; abaxial surface pale green but ferrugine¬

ous to white due to indumentum, the epidermal surface smooth, with dense ferrugineous to white, or translu¬

cent, stellate-peltate scales, 0.13-0.28 mm across, the veins with similar scales, such scales on lamina persis¬

tent to ± deciduous, and persistent to deciduous on the primary to tertiary veins, and lamina surface occasion¬

ally sticky; mite domatia present or absent, usually (when present) at junction of midvein and major secondary

veins, but sometimes also at junction of midvein and minor secondary veins, junction of tertiary veins and

midvein, and/or junction of tertiary veins and major secondary veins, composed of elongate, multicellular, ±

ferrugineous, yellow to reddish, eglandular, barbate hairs, 0.2-1.6 mm long. Inflorescences terminal, panicu¬

late cymes, 3.5-19 cm long, 2-13.5 cm across, with (1-) 2-6 major branch pairs, the peduncle 1-6.7 cm long,

the ultimate axes (pseudopedicels) 1.5-7 mm long, and the numerous flowers well separated from each other;

bracts 0.3-8.5 x 0.15-1 mm, triangular or narrowly triangular to linear, with acute to rounded apex, quickly

deciduous; bracteoles 0.3-0.6 x 0.15-0.3 mm, narrowly triangular, with acute apex, deciduous. Flowers 5- or

Judd et al., Revision of Miconia sect. Miconiastrum

467

less commonly 6-merous, zygomorphic (due to positioning of the stamens), with pedicel 1-3 mm long. Hypan-

thium 3.7-7.5 mm long, terete and moderately to not at all constricted above ovary, the free portion 2-4 mm

long, 2-3.5 mm wide at the torus, the outer surface with sparse to dense stellate-peltate scales (and sometimes

sticky), the internal surface smooth to very slightly longitudinally ridged, glabrous. Calyx lobes 5 or 6, separate

(and valvate or open) in bud, the tube 0.5-2 mm long, not tearing between lobes (but rarely with a few irregular

tears), with moderate to dense stellate-peltate scales abaxially, and sparse to moderate branched to stellate-

peltate scales adaxially, the lobes obsolete or 0.1-1 x 1.5-3 mm, obsolete (with a rim-like calyx) or obscurely to

clearly developed, ± broadly triangular to triangular, the apex rounded to obtuse or acute, with hairs similar to

those of the tube, green, but often red-tinged or red; calyx teeth absent or present, 0-0.3 mm long, round or

ellipsoidal bump (near apex of lobes), green, often red-tinged or red, with rounded to acute apex, and hairs

similar to those of calyx. Petals 5 or 6, 5-11.5 x 27-6.7 mm, asymmetrically obovate, spreading, white, occa¬

sionally red-tinged, glabrous but both surfaces densely papillose-granulate, the apex rounded, ± shallowly

notched, the base slightly narrowed to a broad attachment, the margin entire. Stamens 10 or 12, isomorphic,

± geniculate near filament apex; staminal filament 4-8 mm long, glabrous, pale yellow, occasionally pink-

tinged, anther thecae 4.5-7.3 x 0.8-1.5 mm, subulate, straight to slightly incurved, opening by a small dorso-

apical pore, yellow, the connective not prolonged below the thecae, glabrous, but the filament emerging from

slight depression in sterile, minutely dorso-lobed anther base. Ovary 3-locular, 2/3-4/5 inferior, ellipsoidal to

oblong or ovoid, 3-5 mm long, 1.5-4 mm in diameter, apically cylindric-conical, glabrous, the apex with short,

slightly ridged collar but lacking crown, with axile placentation, the ovules numerous, borne on slightly ex¬

panded placenta that extends only slightly into locule; style 8-18 mm long, distally curved, white to rose, gla¬

brous; stigma punctate, minutely papillose, ca. 0.15 mm wide. Berries 5-14 mm long, 4-14 mm in diameter,

globose to ellipsoid, purple-black, with scattered stellate-peltate scales hairs, the hypanthium constricted,

1.5-2.3 mm wide at narrowest point and 2-3.5 mm wide at torus, but appearing strongly constricted due to

flaring calyx tube. Seeds 1-1.9 mm long, angular-obovoid, with ± rectangular raphe, the testa completely

smooth, or smooth on flattened distal surface, and ± slightly roughened on the sides, due to slightly bulging

cells; appendage absent.

1. Stems and inflorescence axes without elongate, eglandular hairs_var. bicolor

1. Stems and inflorescence axes with elongate, eglandular hairs_var. patenti-setulosa

2a. Miconia bicolor (Mill.) Triana var. bicolor, Trans. Linn. Soc. London 28:103. 1872. (Figs. ID, 3A, B). M elas-

toma bicolor Mill., Gard. Diet., ed. 8, no. 6.1768. Tetrazygia bicolor (Mill.) Cogn. in A. de Candolle & C. de Candolle, Monogr. Phan.

7:724.1891. Type: Loc. Natalis [country unknown], P. Miller s.n. (lectotype, designated here, following the suggestion of W.T. Gillis,

annotation on sheet, 19 Jun 1973, BM, on-line image #001008022!).

M elastoma pallens Spreng., Neue Entdeck. Pflanzenk. 3:62. 1822. Miconia pallens (Spreng.) Triana, Trans. Tinn. Soc. Tondon 28:103.

1872. Tetrazygia pallens (Spreng.) Cogn. in a A. de Candolle & C. de Candolle, Monogr. Phan. 7:724. 1891. Type: Hispaniola (not

seen).

Naudinia argyrophylla A. Rich, in R. de la Sagra, Hist. Phys. Cuba, Pi. Vase. 10:265. 1845. Tetrazygia angustiflora (Benth.) Griseb. var.

argyrophylla (A. Rich.) Griseb., Fl. Brit. W.I.: 254.1860. Miconia angustiflora (Benth.) Griseb. var. argyrophylla (A. Rich.) M. Gomez,

Dice. Bot. Nombres Vulg. Cub. Puerto-Riquenos: 42.1889. Tetrazygia argyrophylla (A. Rich.) Millsp., Publ. Field Columbian Mus.,

Bot. Ser., 1(6):431. 1900. Type: CUBA [Prov. Pinar del Rio]: Vuelta de Abajo, J.M. Valenzuelas s.n. (P?, not seen, and if lost could be

typified by t. 44(1) in de la Sagra, 1850).

Naudinia chrysophylla A. Rich, in R. de la Sagra, Hist. Phys. Cuba, Pi. Vase. 10:266. 1845. Type: CUBA [Prov. Pinar del Rio]: Vuelta de

Abajo, in locis altis montium (P?, not seen).

Tetrazygia acunae Borhidi, Acta Bot. Acad. Sci. Hung. 23:38. 1977 [1978], Type: CUBA. Prov. Oriente [Prov. Santiago de Cuba]: Sierra

Maestra, Pico Turquino, Jul 1936, fr, J. Acuna 19452 (holotype: HAC!), non Miconia acunae Borhidi, Novenyrendsz Novenyfaldr.

Tansz. Eotros Forand Tudomanyegyet. Budapest 4:23.1976.

Stems and inflorescence axes lacking elongate, multicellular, eglandular hairs.

Chromosome number. —2n = 34 (Solt & Wurdack 1980).

Phenology. —Flowering has been recorded in every month of the year.

Distribution and ecology. — Miconia bicolor var. bicolor occurs natively in southern Florida (Miami-Dade

Co.), the Bahamas (Andros, Eleuthera, Grand Bahama, Great Abaco, and New Providence Islands), western Cuba

468

Journal of the Botanical Research Institute of Texas 8(2)

Fig. 3. A, B: Miconia bicolor var. bicolor. A. Inflorescence. B. Flower (plants in cultivation, Florida). C, D: M. guajaibonensis : C. Plant with young inflores¬

cences. D. Flower {Becquer& Abbott 82431). E ,M. impressa {Ekman 17539).

(mainly in Prov. Pinar del Rio, Isla de la Juventud), central Cuba (mainly in Prov. Cienfuegos, Sancti Splritus,

and Villa Clara) and, less commonly, the “Oriente” region (Prov. Guantanamo, Holguin and Santiago de Cuba),

and on Hispaniola (widespread in the Dominican Republic and Haiti), growing in thickets or coppice, moist

to dry broadleaved forests, and open to dense pine forests, pine and/or palm savannas, or disturbed habitats,

on limestone, serpentine, or gravely to sandy soils, from near sea-level to 1200 m (Fig. 4). The species has been

introduced into Hawaii, where it is locally naturalized in mesic to wet forests near Hilo (Wagner et al. 1990).

Judd et al., Revision of Miconia sect. Miconiastrum

469

Additional specimens examined: BAHAMA ISLANDS. Andros: Mangrove Cay, 18 Aug-10 Sep 1906, Brace 4960 (NY, US); Nicholl’s Town,

5-6 Mar 1907, Brace 6733 (NY); N of Love Hill, 29 Jan 1974, D.S. Correll & Godfrey 41267 (FTG); S part of island, W end of airstrip, 24 Sep

1974, D.S. Correll 43527 (FTG, NY); just S of Stafford Creek Bridge, 5 Dec 1976, D.S. Correll & Proctor 47757-B (FTG); 16-18 mi S of Andros

Town on Fish Camp rd, 2 Mar 1966, Dawson 26653 (DUKE, US); Coakley Town to Calabash Bay, 8 Mar 1966, Dawson 26799 (US); Con-

gotown District, E bank of South Bight, 5 mi S of Drigg’s Hill jetty, 19 Mar 1986, Eshbaugh & Wilson 86-75 (FTG); 2 mi N of Fresh Creek, 16

Mar 1966, Gillis 6101 (MSC); along Queen’s Hwy, 100 m S of Stafford Creek bridge, near Forfar Research Station, 27 Feb 1976, Gillis 12859

(MSC); Forfar Field Station, town of Stafford Creek, 24°53 , 51"N, 77°55'56"W, 20 Jul 2002, Goldman2458 (FTG, GH); 2 mi NW of Love Hill,

24 May 1975, Hill 3044 (FTG); near IFS Field Station, near Blanket Sound settlement, 15 Jul 1990, Kjellmark 15 (DUKE); ibid., 0.5 mi S of

main rd, 20 Jun 1991, Kjellmark 58 (DUKE, MO); near San Andros Motel, 13 Jun 1987, Nickrent 2644 (NY); Nicols Town, May 1890,J. I. &A.R.

Northrop 127a (GH); Mangrove Cay, 8 May 1973, Popenoe 194 (FTG); near Lisbon Creek, Mangrove Cay, 16-19 Jan 1910, Small & Carter 8468

(GH, NY, US). Eleuthera: Harbour Island to Lower Bogue, 18 Feb-4 Mar 1907, E.G. Britton 6443 (NY); near turnoff to the Bluff from rd to

Ridley Head, 8 Jan 1974, D.S. Correll 41034 (FTG, NY); just E of The Bluff, 12Jan 1974, D.S. Correll 41207 (FTG); along Queen’s Highway, N of

Lower Bogue, 7 Apr 1978, D.S. &H.B. Correll 49551 (FTG, NY). Grand Bahama: Queens Way, W of Freeport, 18 Mar 1971, Austin & Conroy

4587 (FTG, NY); 7 mi NE of Freeport, 12 Feb 1966, Beckner 824 (FLAS); Eight Mile Rocks, 5-13 Feb 1905, N.L. Britton & Millspaugh 2451

(NY); Gold Rock Creek, near E end of island, 28 Apr 1951, Carr s.n. (FLAS); Gold Rock Creek, near High Rock, 14 Sep 1951, Carr s.n. (FLAS,

GH); near jet of Sandcombe Rd and Grasmere Dr, Freeport, 4 Nov 1973, D.S. Correll 40481 (FTG); intersection of Settlers Way and East Mall,

Freeport, 14 Aug 1974, D.S. Correll & Krai 42816 (FTG, NY); Rand Memorial Park Nature Center Preserve, 1.5 mi from Freeport along Set¬

tlers Way, 31 Mar 1969, Gillis 7758 (FTG). Great Abaco: Marsh Harbor, 9 Dec 1904, Brace 1600 (NY); NW edge of Marsh Harbour, 15 Jun

1981, D.S. Correll & Wasshausen 52032 (FTG, US); N of Coopers Town, 26°53.978'N, 77°33.097'W, 17 May 2000, Freid 00-070 (FTG); Abaco

National Park, 7Jun 2004, Freid 04-023 (FTG, NY); 1 miEofairstripatMarsh Harbour, 28 Mar 1966, Perry 1776 (DUKE); 0.5 mi N ofDundas

Town, 14 Dec 1979, Wunderlin et al. 8265 (FLAS, GH, MO); main rd, 2.9 mi S of Wilson City rd, 16 Dec 1979, Wunderlin et al. 8524 (GH, MO).

New Providence: without definite locality, 2 Feb 1877, Brace 465 (NY); Killarney pine barrens, 12-24 Mar 1907, E.G. Britton 6536 (NY, US);

Blue Hills Rd, 7-8 Apr 1904, N.L. Britton 8 (NY); Farringdon Rd, 24 Aug 1904, N.L. Britton & Brace 232 (NY); near Nassau airport, 19 Apr

1971, Burch 4204 (MICH, MO, NY); without definite locality, 1859, Cooper 82 (GH, NY); Thompson Blvd. near Portago Rd, Nassau, 23 Mar

1977, D.S. & H.B. Correll 48265 (FTG, NY); Garden Hill Estates, Blue Hill area, Nassau, 4 Jun 1976, D.S. Correll 47225 (FTG, NY); near Nas¬

sau, Jan-May 1903, Curtiss 41 (A, GH, HAC, MIN, NY, P on-line image, US); near Harrold and Wilson Ponds, 21 Feb 1946, Degener 18801

(GH, NY); S of Prospect Ridge, Nassau, 15 Feb 1946, Degener 18836 (GH, NY); in pinetis, 22 Feb 1888, Eggers 4180 (MICH, NY, P on-line

image, US); Wend of island, 23 Mar 1963, Gillis 5347 (MSC); SW corner of island, 1 mi SWofLyford Cay, 7Jul 1969, Gillis 8350 (FTG); along

470

Journal of the Botanical Research Institute of Texas 8(2)

Coral Harbour Rd, 4 May 1970, Gillis 9202 (FTG); pinelands, 3 Jul 1948, Ledin 268 (FTG); ibid., 18 Aug 1948, Ledin 315 (FTG); Nassau, 15 Jan

1890, J.I. &A. R. Northrop 127b (NY); S of Windsor Airport, W of Nassau, 2 Apr 1965. Perry 1620 (DUKE); without definite locality, 3-5 Sep

1952, von Reis 231 (MICH, US); N of Winsor Field, 77°28.5'W, 25°3.5'N, 13 Jul 1960, Webster et al. 10457 (DUKE, US); 1 mi S of Nassau, 24

Feb 1905, Wight 128 (GH, NY); Grantstown, 28-29 May 1909, Wilson 8171 (NY). CUBA. Prov. Camaguey: Cespedes, El Quemado, 15 May

1915, Roig et al. H-ROIG 898 (HAC); Queen City to Minas, 21 Nov 1909, Shafer 2925 (F, NY, US). Prov. Ciego de Avila: Sierra de Judas, Cu-

nagua, 1-7 Aug 1985, Cano O. & Herrera 146 SV-34725 (HAC). Prov. Cienfuegos: Cumanayagua, Sierra del Escambray, subida al Pico San

Juan, 7 Nov 1987, Arias et al. HFC-62964 (HAJB); ibid., en el camino entre Los Tornos y El Naranjo, 3 Nov 1987, Arias et al. HFC-62782

(HAJB); ibid., lomas alrededor de Los Tornos, 4 Nov 1987, Arias et al. HFC-62809 (HAJB); ibid., San Bias, 300-400 m, 5 Aug 2004, Becquer et

al. HFC-83168 (HAJB); ibid., Las Vegas entre el cafetal y San Bias, cafetal de Buenos Aires, 29 Oct 1985, Berazain et al. HFC-58009 (HAJB);

ibid., 8 Oct 1986, Gonzdlez L. et al. HFC-60246 (HAJB); Cumanayagua, Sierra del Escambray, loma al sur de Pico San Juan, 700-800 m, Nov

1975, Bisse et al. HFC-28707 (HAJB); Cieneguita, 7Jun 1895, Combs 128 (F, GH, MICH, MO, NY, P on-line image); Cumanayagua, Complejo

San Juan, 2 Nov 1986, Exp. Conjunta P. Herrera et al. SV-35912 (HAC); Las Lagunas, Buenos Aires, 2500 ft, 6 Dec 1928 Jack 6884 (A, NY, P

on-line image, US); Soledad, Cienfuegos, 30 Mar 1936 Jack 8758 (GH, P on-line image). Prov. Habana: al E de playa Baracoa, 23 Oct 1966,

Bisse HFC-189 (HAC); Loma La Pita, entre Campo Florido yjaruco, 31 Nov 1974, Bisse et al. HFC-25316 (HAJB). Prov. Guantanamo: Irmas,

Maisi, Tres Piedras, 700-900 m, 27 Jul 1984, Alvarez et al. HFC-55049 (HAJB); pinar de Monte Cristi, 500-600 m, Jun 1967, Bisse & Rojas

HFC-3614 (HAJB); Monte Cristi, 700 m, May 1968, Bisse & Kohler HFC-9327 (HAJB); Felicidad de Yateras, en la zona de Monte Cristi, 23 Aug

1971, Bisse HFC-20223 (HAJB); orillas del rio Baez, cerca del campamento “Los Naranjos”, 1-3 Aug 1975, Bisse et al. HFC-26997 (HAJB); aser-

rio Nuevo Mundo, al S del aserrio, 300 m, Apr 1975, Bisse et al. HFC-25816 (HAJB); Felilcidad de Yateras, Pinar de Monte Cristi, 700 m, 13

May 1983, Bisse et al. HFC-49401 (HAJB); al S de la Reservacion de Cupeyal, 11 Feb 1970, Borhidi et al. SV-35467 (HAC). Prov. Holguin: Si¬

erra Cristal, valle del rio Lebisa desde Mandinga hacia arriba, Apr 1968, Bisse & Kohler HFC-7412 (HAJB). Prov. Isla de la Juventud (Isla de

Pinos): Sta. Fe, John Jungle, 19-22 Jun 1922, Acuna SV-22673 (HAC); camino de cayo Piedras a Punta del Este, 4 Nov 1981, Alvarez et al.

HFC-45623 (HAJB); camino hacia Puenta del Este, desde Cayo Piedras, 13 Apr 1974, Areces & Berazain HFC-24876 (HAJB); Reserva Ecologi-

ca “Los Indios”, centro-oeste de la isla, al N del Hotel Colony entre el rio Los Indios y el rio Itabo, por el sendero ecologico desde la Estacion

Ecologico, 0-5 m, 31 Mar 2008, Becquer et al. HFC-85059 (HAJB); Nueva Gerona, cerca del Motel Rancho del Tesoro, 22 Dec 1966, Bisse

HFC-723 (HAJB); orillas de la laguna Santa Barbara, 23 Dec 1966, Bisse HFC-881 (HAJB); cerca de las Delicias, 23 Jul 1971, Bisse HFC-19828

(HAJB); camino de cayo Piedras a Punta del Este, 24 Jul 1971, Bisse HFC-19848 (HAJB); ibid., 23 Oct 1976, Bisse et al. HFC-32751 (HAJB); Si¬

erra de la Canada, 50-200 m, 1 May 1970, Bisse et al. HFC-26297 (HAJB); near Nueva Gerona, 5 Apr 1904, Curtiss 414 (F, GH, HAC, MIN, NY,

P on-line image, US); Santa Barbara, 3 Nov 1920, Ekman 12104 (S); vicinity of Nueva Gerona, 6 May 1910, Jennings 57 (NY); arroyo E of Los

Indios, 18 May 1910, Jennings 347 (GH); along rd to San Francisco de las Piedras, 10 Feb 1955, Killip 44741 (GH, HAC, US); rd from Nueva

Gerona to Santa Barbara, 19 Nov 1955, Killip 45110 (US); 2.5 km NW of Santa Barbara on rd to Westport, 26 Jan 1956, Killip 45471 (HAC, US);

near Nueva Gerona, 30 Jun 1900, Palmer & Riley 906 (NY); Sta. Fe, Hacienda San Juan, 12 Aug 1919, Roig & CremataSV-7117 (HAC); without

definite locality, 25 Jun-lOJul 1901, Faylor 138 (GH); ibid., Faylor 138b (F, MO, NY, US); ibid., Faylor 139 (F, GH, MO, NY, US). Prov. Las

Tunas: Cerro de Dumaiiuecos, near Manati, 31 Mar 1939, Bros. Leon & Alain 18914 (GH, NY). Prov. Matanzas: Sabana de Cascajal, 30 Sep

1947, Acuna SV-14784 (HAC); Sabanas de Lacret, casi en el limite con Las Villas, 30 Sep 1954, Bro. Alain 4055 (GH, HAC); Jovellanos, San

Miguel de los Banos, alrededores Loma Jacan, 21 Dec 1976, Berazain et al. HFC-33581 (HAJB); Playa Giron, monte seco, Nov 1967, Bisse &

Rojas HFC-4683 (HAJB); Playa Larga, peninsula de Zapata, montes entre Caletones y Santo Tomas, 19 Feb 1977, Bisse et al. HFC-34499

(HAJB); Aguada de Pasajeros, sabana al N del km 193 de la Autopista Nacional, 11 May 1984, Bisse et al. HFC-54772 (HAJB); Cienaga de Za¬

pata, Batey Casa Nueva, 7 Jun 1978, Oviedo et al. SV-30513 (HAC). Prov. Pinar del Rio: Alturas Pizarrosas del Sur, Zona de Pizarras a lo largo

de la carr. Entre Pinar del Rio y Vinales, km 12-15, 22°50'N, 83°42'W, 140 m, 7 Jul 1993, Acevedo-Rdgz. et al. 5705 (HAC); Vinales, Santo

Tomas, May, Acuna SV-9917 (HAC); San Vicente, Valle del Ancon, 2 Jun 1953, Acuna & Correll SV-18664 (HAC); pinares prox. a Vinales, 30

Nov 1947, Acuna SV-22688 (HAC); Sabanalamar, El Sabalo, Finca Fueyo, 2-5 Apr 1950, Bro. Alain 1335 (HAJB, NY); entre San Vicente y El

Ancon, 17 Mar 1957, Bro. Alain & Proctor 6187 (HAC); Guanae, El Pitirre, en el terraplen desde Hermanos Barcon hasta la carretera de la

Coloma, 19 Oct 1983m Alvarez et al. HFC-50957 (HAJB); Bahia Honda, Sierra de Cajalbana, subida al Tecnologico Invasion de Occidente, 300

m, 15 Jul 1989, Alvarez & Pujadas A. HFC-70164 (HAJB); Sandino, Carabelita, costa entre La Barca y el entronque de Carabelita, Guanaha-

cabibes, 22 Nov 1973 , Areces A. s.n. (HAJB); Herradura, 20 Sep 1904, Baker SV-2145 (F, GH, HAC, HAJB, NY, US); ibid., 10 Apr 1905, Baker &

Diminoch SV-4860 (HAC, HAJB); ibid., 10 Apr 1905, Baker SV-5086 (HAC); Corral Viejo, cerca de la laguna vieja de Santa Teresa, 5-10 m, 6

May 2004, Becquer & Abbott HFC-82286 (FLAS, HAJB); Guane, El Sabalo, pinares de Sabanalamar, al S del pueblo, 5-20 m, 3 Apr 2001,

Becquer & Veloso HFC-79848 (HAJB); Vinales, pinares alrededores de la carretera entre el entronque Ancon y el valle Ancon, 200-300 m, 13

Nov 2002, Becquer & Veloso HFC-81071 (HAJB); Sandino, potreros cerca de La Fe, 6 May 2006, Becquer & Morejon R. HFC-83986 (HAJB);

Sanadino, San Ubaldo, reserva floristica, 8 May 2006, Becquer CMorejon HFC-84011 (HAJB); Vinales, Moncada, Alturas de Pizarras del Sur,

pinares al S de la Escuela de Espeleologia, 11 May 2006, Becquer & Morejon R. HFC-84044 (HAJB); Guanes, cerca de La Fe, Mar 1967, Bisse &

Rojas HFC-1805 (HAJB); Guanes, laguna de Los Indios, Mar 1967, Bisse & Rojas HFC-1833 (HAJB); Guanes, peninsula de Guanahacabibes,

Mar 1967, Bisse & Rojas HFC-3078 (HAJB); La Palma, pinares al N de San Andres, Dec 1967, Bisse & Rojas HFC-4950 (HAJB); San Diego de los

Banos, pinares al N de la Sierra de la Guira, Jul 1968, Bisse et al. HFC-9534 (HAJB); Vinales, pinares al S del pueblo, 16 Nov 1968, Bisse & Lip-

pold HFC-9646 (HAJB); ibid., Bisse & Lippold HFC-9702 (HAJB); Vinales, pinares al S del pueblo, Nov 1986, Bisse & Lippold HFC-10866

(HAJB); Las Ovas, lagunas al NO del pueblo, 1970, Bisse & Lippold HFC-17390 (HAJB); La Guira, las Bermejales, al N de la Sierra de Guira,

200-300 m, Dec 1970, Bisse & Lippold HFC-18380 (HAJB); Vinales, Canada de un arroyo en los pinares al SO del pueblo, Dec 1974, Bisse et al.

HFC-26430 (HAJB); lagunas al SO de Las Ovas, 5 Apr 1976, Bisse et al. HFC-31122 HAJB); Sumidero, cerca de la confluencia de arroyo Cayo

Judd et al., Revision of Miconia sect. Miconiastrum

471

Malo y arroyo Camarones, 12 Dec 1978, Bisse et al. HFC-38545 (HAJB); Mantua, monte al E de la desembocadura al arroyo Camarones, 27

Mar 1982, Bisse et al HFC-43361 (HAJB); Las Ovas, laguna, 4 km al SO del pueblo, 50 m, 31 Mar 1982, Bisse et al HFC-46704 (HAJB); San

Luis, Las Llanadas, 50 m, 19 Dec 1982, Bisse et al HFC-48582 (HAJB); Minas de Matahambre, Sumidero, zona Cejas de Lrancisco, 20 Jan

1984, Bisse et al HFC-51457 (HAJB); Mantua, las Clavellinas, cayo Cienaga del Prado, 16 May 1984, Bisse et al HFC-54448 (HAJB); brazo del

rio Lrio al E de Mantua, 24 Nov 1975, Borhidi & Capote SV-32857 (HAC); Sandino, peninsula de Guanahacabibes, 25 Oct 1975, Borhidi &

Delgado F. HPR-4185 (HAJB); vicinity of Herradura, 26-30 Aug 1910, N.L. Britton et al 6603 (NY, US); San Diego de los Banos, 31 Aug-3 Sep

1910, N.L. Britton et al 6746 (NY, US); vicinity of Pinar del Rio, 5-12 Sep 1910, N.L. Britton et al 7116 (NY); pres de Vinales, 22°37'N, 83°3'W,

130 m, 17Jun 1990, Dechamps et al NH:12468 FW:49941 (MO); Municipio Las Ovas, Reserva Natural de El Punto, 83°33'W, 22°21'N, 10 Mar

1997, Dressier 192 (F); Herradura, 30 Mar 1907, Earle 656 (F, NY, US); ibid., 16 Aug 1907, Earle 730 (NY); Santa Barbara, 5 Dec 1901, Earle s.n.

(NY); Herradura, 12 Apr 1920, Ekman 10774 (S); S of Las Mangas, 25 Jul 1921, Ekman 13066 (S); Herradura, 27 Oct 1923, Ekman 17791 (NY,

S); Consolacion del Sur, Hato de las Vegas, El Tablazo, 26 May 1918, Fortun & San Pedro SV-6887 (HAC); Consolacion del Norte, Jun 1920,

Ganganelli R-ROIG 2076a (HAC); Sandino, San Ubaldo, laguna del Toro, 30 Jan 1985, GonzdlezL. et al HFC-55343 (HAJB); La Fe, 7 Jan 1971,

Grudzinskaya & Imchanitzkaja 768 (HAC); Consolacion del Sur, Autopista Nacional entre Entronque de Herradura y foresa San Diego, 7 Feb

1992, Gutierrez HFC-70380 (NY); Vinales, en el km 13, alrededores de la carretera de Vinales a Pinar del Rio, 9 Feb 1991, GutierrezJ. & Panjet

C. HFC-69550 (HAJB); vicinity of San Diego de los Banos, 5 Jul 1915, Bro. Leon LS-5155 (HAC, NY); SE of Los Palacios, 7 Aug 1917, Bro. Leon

& M. Roca LS-7377 (NY); entre Mangas y Candelaria, 27 Jan 1931, Bro. Leon LS-14766 (HAC, NY); S of Herradura, 7 Apr 1933, Bro. Leon LS-

15936 (HAC, NY); Sabana cerca de Corojal, Laguna de Piedras, Candelaria, Apr 1937, Bro. Ledn 16820 (HAC, NY); San Luis, Sabana de Santa

Marta, 1 May 1940, Bro. Leon et al LS-17821 (HAC); Laguna de Piedras, Mangas, 2 Mar 1939, Bro. Leon LS-18898 (HAC, NY); la Laguna de

Piedras, Mangas, 20 May 1941, Bro. Leon 20078 (GH, HAC, NY); cercanias de La Fe, Feb 1942, Bros. Leon & Marie-Victorin LS-20497 (HAC,

NY); montes cerca de la Jaula, 23 Feb 1977, Lepper L. et al HFC-34367 (HAC, HAJB); Lagune de la Maquina, 4 Mar 1943, F. Marie-Victorin

58352 (GH); near Pinar del Rio, 24 Feb 1900, Palmer & Riley 49 (US); near Coloma, 18 Mar 1900, Palmer & Riley 352 (US); Ancon, vereda de

mina del Gato, a orillas del rio Sajanal, 28 Jun 2007, Panjet C. HFC-85610 (HAC); without definite locality, Jul 1910, Ponce & Ramos H-ROIG

257 (HAC); 4 mi E of Candelaria, 16 Mar 1957, Proctor 16293 (A); Consolacion del Sur, 29 Nov 1915, Roig & Van Hermann SV-6377 H-ROIG

1128 (HAC); San Diego, La Guira, Apr 1966, Samek V. SV-26081 (HAC); without definite locality, 23 Apr 1903, Shajer274 (HAC, NY); Herra¬

dura to railroad, then to Paso Real, 11 Jan 1912, Shafer 11785 (A, F, MO, NY, US); ibid., Shafer 11787 (NY); Los Palacios to San Juan de Zayas,

17Jan 1912, Shajer 11807 (A, F, MO, NY, US); ibid., Shafer 11809 (A, MO, US); San Juan y Martinez, cayo Ratones, 15 Feb 1989, Urquiola et al.

HPR-5091 (HAJB); Ovas, El Punto, 5 Feb 1990, Urquiola et al. HPR-5811 (HAJB); Herradura, 1904, Van Hermann SV-270 (NY, P on-line image,

US); ibid., 1905, Van Hermann SV-560 (NY, P on-line image); ibid., 1905, Van Hermann SV-573 (NY); ibid., 1905, Van Hermann SV-596 (NY);

ibid., 26 Aug 1905, Van Hermann SV-773 (HAC); ibid., 10 Sep 1905, Van Hermann SV-867 (HAC); ibid., 21 Mar 1906, Van Hermann 2780 (A,

GH, HAC, US); ibid., 23 Jan 1905, Van Hermann SV-4478 (HAC); Puerta de Golpe, 27 Jan 1915, Van Hermann SV-5849 (HAC); entre Candela¬

ria yArtemisa, 17 Sep 1904, Wilson 1713 (HAC, HAJB, NY); Vueltabajo, 23 Jul, and Retiro [near Taco Taco], 22Jun, Wright 1222 p.p. (GH, NY,

S); Retiro [near Taco Taco and possibly other localities], June, Wright 1222a (GH, NY, P on-line image, US); Retiro [near Taco Taco], 21 Jun,

and Vueltabajo, 24 Jul, 1860-1864, Wright 2507 (GH, HAC, MO); Los Remales, 1660-1864, Wright s.n. (GH); without definite locality, Wright

s.n. “308”, 1863 (S); ibid., 1865, Wright s.n. (S); ibid., Wright s.n. “980” (HAC); ibid., Wright s.n. (HAC); Sanjulian, granja Sandino, Feb 1966,

Yero M. SV-25996 (HAC); Vinales, cerca del camino, 9 Jun 1995, Yero D. HFC-81190 (HAJB). Prov. Sancti Splritus: Sierra del Escambray,

Topes de Collantes, Mogote mi Retiro, 79°50'W, 22°22'N, 850 m, 2Jul 1993, Acevedo-Rdgz. et al. 5593 (HAC, US); Trinidad, Topes de Collan-

tes, Pico Potrerillo, 750-950 m, 13 Apr 1991, Alvarez & Gutierrez HFC-69444 (HAJB); falda S de las lomas de Banao, Areces et al. HFC-28997

(HAC, HAJB); Banao, camino entre el monument de Cantu y el tope de La Diana, 26 Oct 1986, Arias et al. HFC-59737 (HAJB); Jatibonico, El

Patio, alrededor de la laguna Botijuela, 12 Feb 2002, Becquer & Veloso HFC-79904 (HAJB); Sierra del Escambray, loma al O de la carretera, 5

km al S de Topes de Collantes, 600 m, 7 Nov 1968, Bisse & Lippold HFC-9662 (HAJB); Fomento, camino entre Santa Teresa y Gavilanes, 8 Nov

1979, Bisse et al. HFC-41022 (HAJB); ibid., Bisse et al. HFC-41023 (HAJB); Fomento, valle del arroyo Gavilancito, 300-400 m, 10 Nov 1979,

Bisse et al. HFC-41077 (HAJB); Trinidad Mts, slopes of Pico Potrerillo, 700-900 m, 12Jun 1922, Ekman 13999 (F, S, US); Cabaiguan, El Arrie-

ro, 28 Jul 1918, Fortun SV-6927 (HAC); Trinidad, El Tibisial, alrededores de loma Gavilanes, 29-30 Sep 2002, Gutierrez et al. HFC-80247

(HAJB); Loma de la Jagua, Los Helechales, Banao, 10 Aug 1915, Bros. Leon & Clement LS-5395 (HAC, HAJB, NY); from Trinidad to Tope de

Collantes, Jul 1938, Bro. Leon LS-18430 (GH, HAC, NY); Loma de la Gloria, Banao Mts, 29 Jul 1918, Bro. Leon & M. Roca LS-8003 (HAC, NY);

Lomas de Banao, May 1920, Luua591 (F, HAC); Loma Caballete de Casa, 400-700 m, 8 Nov 1979, Imchanitzkaja & Herrera 448 (HAC). Prov.

Santiago de Cuba: Gran Piedra, carretera de la Piedra a la Mercedita, 1200 m, 25 May 2004, Becquer et al. HFC-82538 (HAJB); Guama, Mo¬

gote Peak [Pico Turquino], 27 Feb 1939, Bucher 116 (HAC); Guama, Sierra del Cobre, subida a la loma El Espejo, 30 Apr 1989, Dietrich H. et

al. HFC-67756 (HAJB). Prov. Villa Clara: Sabanas de Manacas, 6 Jun 1950, Acuna & Correl SV-18663 (HAC); Sabanas de Manacas, orillas de

la Carretera Central, 13 Apr 1954, Bro. Alain 4013 (GH, HAC, US); Manacas, al N del pueblo, 19 Nov 1966, Bisse & Rojas HFC-435 (HAJB);

ibid., al S del pueblo, Feb 1967, Bisse & Rojas HFC-1497 (HAJB); Cartagena, cerca de Santiago, Apr 1967, Bisse & Rojas HFC-2142 (HAJB);

Huffs farm, near Manacas, Jun 1941, Howard 5500 (GH, NY, US); Placetas del Sur, 29 Jul 1916, Bro. Leon LS-6405 (HAC, NY, US); Motembo,

10 Aug 1918, Bro. Leon & M. Roca LS-8213 (NY); Sabana de Motembo, 28 Aug 1922, Bro. Leon LS-11369 (HAC, NY); near Manacas, 11 Jul

1936, Smith & Hodgdon 3097 (F, GH, MICH, MO, NY, P on-line image, S, US). DOMINICAN REPUBLIC. Distrito Nacional: Sierra de Ya-

masa, Sierra Prieta, proximo al nacimiento del arroyo Tosa, 80 m, 12Jun 2005, Clase & Angeles 4060 (FLAS); El Manielito, 23 Jan 1929, Ek¬

man H11313 (S, US); Sierra Prieta, 18.65386°N, 69.9689°W, 120-125 m, 20 Jul 2012,Jestrow 2012-244 (FLAS); Sierra Prieta, Villa Mella,

150-200 m, 24 Oct 1975, Liogier 24117 (JBSD, NY); Sierra Prieta, al NE de Villa Mella, camino a Yamasa, a la orilla del arroyo Mina proximo

a un potrero, 18°19'N, 69°58'W, 70 m, Veloz et al. 103 (FLAS, JBSD); Sierra Prieta, NE de Villa Mella, camino a Yamasa, 18°19'N, 69°58'W, 60

472

Journal of the Botanical Research Institute of Texas 8(2)

m, Veloz et al. 323 (FLAS, JBSD). Prov. Dajabon: Cerro de Chacuey, Partido, Dajabon, 300 m, 18 Aug 1973, Liogier 20004 (F); Cordillera

Central, Municipio de Partido, Cerro de Chacuey, Loma Las Mercedes, 19°26 , 55.1"N, 71°36 , 7.9"W, 400 m, 11 May 2005, Veloz & Garda 3757

(FLAS, FTG, JBSD); Municipio de Partido, Cerro de Chacuey, entrando por Cajuil, 19°26 , 31.9"N, 71°34 , 52.8"W, 385 m, 11 May 2005, Veloz

& Garda 3821 (JBSD). Prov. Hato Mayor: Sabana de la Mar, 8 Jul 1930, Ekman H15599 (S). Prov. La Vega: Cordillera Central, 10 km S de la

carretera La Vega - Santiago, en la carretera hacia Jarabacoa, 19 0 11.5'N, 70°35'W, 490 m, 11 Dec 1995, Garda et al. 6067 (F, JBSD, MO); Cor¬

dillera Central, 7.5 km NE ofjarabacoa on rd to Bayacanes and La Vega, 630 m, 25 May 1992 Judd 6623 (FLAS, JBSD); Loma del Puerto, La

Vega, 400 m, 18 Jun 1969, Liogier 15723 (NY, US). Prov. Maria Trinidad Sanchez: Los Haitises, La Manaclita, 2Jul 1930, Ekman H15528 (A,

S, NY, US). Prov. Monsenor Nouel: Cordillera Central, SE of Bonao, Falcombridge Dominicana mine, at top of Loma Larga, 650 m, 14 May

1992 Judd 6516 (FLAS, JBSD); Loma Peguera, Barrancon, from Bonao to Hato Viejo, 200-300 m, 17 Apr 1969, Liogier 14874 (GH, JBSD, NY,

US); Cordillera Central, Falcombridge Dominican mine in the area of Loma Peguera, Loma Larga, on the top, 650 m, 14 May 1992, Skean

3125 (MICH, MSC); 5 km SE of Banao, at Falconbridge Dominicana Mine, 18°45'N, 70°15'W, 28Jul 1981, Watson 1191 (FLAS, FTG); ibid., 29

Jul 1981, Watson 1264 (FLAS, FTG); SE of Bonao, Loma La Peguera vicinity, Falcombridge Dominicana mine, on Loma Larga, 18°50'N,

70°25'W, 28 Jul 1981, Zanoni et al. 15665 (FLAS, JBSD, NY, US). Prov. Monte Plata: 12.5 km NE de Bayaguana y 3.5 km NW del poblado

Trinidad, en el lugar llamado Sabana Larga, 18°50'N, 69°32'W, 190 m, 14 Sep 1987, Garda & Pimentel 2347 (FLAS, JBSD); Batey Guanuma,

100 mde la finca experimental de laUniv. “CDEP”, 60 m, 31 Jul 1992, F. Jimenez 583 (FLAS, JBSD); Rincon Dajao on Yamasa rd, 10 mi N of

Santo Domingo city, 150 m, 26 Jun 1968, Liogier 11841 (GH, JBSD, NY, P on-line image, US); Los Haitises, proximo al Rio Comate, 1 km NE

del poblado Comatillo (al NE de Bayaguana), 18°48'N, 69°36'W, 100 m, 10 May 1991, Zanoni & F. Jimenez 45222 (FLAS, JBSD). Prov. Puerto

Plata: Puerto Plata, Samanas rd, 10 Feb 1871, Wright et al. 522 (US). Prov. Samana: Peninsula de Samana, Cabo Cabron, 100 m, 12 Nov

1993, Salazar et al. 1260 (FLAS, JBSD). Prov. San Cristobal: 30 km from Santo Domingo City on Duarte Highway, 28 Jun 1972, Liogier 18648

(F,JBSD, NY, US). Prov. Sanchez Ramirez: Campamento Los Pinos, Maimon, 250 m, 31 May 1977, A. &P. Liogier 26656 (JBSD, MO, NY, US).

Prov. Santiago: Cordillera Central, Municipio San Jose de las Matas, seccion Los Ramones, en las carretera hacia Moncion, despues de Ji-

come, 689 m, 20 Aug 2011, Clase et al. 6896 (FLAS, JBSD); Jicome, San Jose de las Matas, 600-700 m, 21 Jun 1933, Valeur935 (F, MO, NY, US).

Prov. Santiago-Rodriguez: between Sabaneta and Santiago de la Cruz, 24 Apr 1970, J. Jimenez &ArizaJulia 5824 (NY); ibid., 25 Apr 1971, J.

Jimenez &Ariza Julia 5924 (FLAS, GH, UCMM, US); Cordillera Central, along rd between La Leonor and El Aguacate, 3 km W of La Leonor,

600 m, 30 May 1992 Judd 6669 (FLAS, JBSD); Cordillera Central, El Cajuil, 10.2 km al SE de Moncion, 19°2TN, 7TT2'W, 600 m, 16 Jul 1985,

Mejia et al. 1408 (FLAS, JBSD, NY). HAITI. Dept, de Nippes: Massif de la Hotte, Corail, near Dutreuil, 200 m, 29 Sep 1928, Ekman H10765

(A, S, US). Dept, de L’Ouest: lie La Gonave, Lotoroue, 650 m, 28Jul 1927, Ekman H8722 (GH, NY, S, US); Gonave Island, near Pte. a Raquette,

1927, Eyerdam 240 (F, GH, NY, US). Dept, du Nord: Massif du Nord, Acul du Nord, Morne Haut du Cap, hill at the Acul Bay, Morne Bois-Pin,

100 m, 16 Dec 1924, Ekman H2888 (S, US). Dept, du Nord Ouest: vicinity of Jean Rabel, S of town, 1-13 Mar 1929, E.C. & G.M. Leonard 13629

(GH, MO, NY, US). UNITED STATES OF AMERICA. Florida. Miami-Dade Co.: F.I.U. Preserve, 25°45.24'N, 80°22.57'W, 24 Apr 2005,

Adonizio2 (FTG); off US 1 between Homestead and Naranja, 9 Jun 1973, Almeda 1937 (USF); Long [Pine] Key, Everglades, May 1908, Bessey

24 (MSC); ibid., Bessey 50 (MSC); Pine Hammock Trail [Long Pine Key], Everglades National Park, 6 Jul 1965, Bourdo s.n. (MSC); Florida

City, 16Jul 1965, Bourdo s.n. (MSC); Goulds Pineland, SW120 Ave at 224 St, 7Jun 1998, Bradley 1808 (FTG); Lucille Hammock Preserve, 0.8

mi W of SW 217 Ave, 0.6 mi S of 344 St, 25.438°N, 80.555°W, 5 Jul 1998, Bradley 1908 (FTG); near Homestead, Feb 1929, Buswell s.n. (FTG);

ibid., 8 Jul 1934, Buswell s.n. (FTG); Long Pine Key, 17 Jul 1937, Buswell s.n. (FTG); Timb’s Hammock, Homestead, 5 Mar 1952, Caldwell 8758

(FLAS, FTG, MIN); 6 mi SW of Homestead along Rt 27,12 Mar 1956, Churchill s.n. (MSC); Everglades National Park, Palma Vista Hammock,

SW of Homestead, 12 Oct 1962, Cooley et al. 9201 (USF); Miami, Timbs Hammock, 26 Feb 1953, Diaz-Piferrer 49 (HAJB); near Naranja, 27

Apr 1937, Fennell 382 (NY); Sykes Hammock, 4 mi NW of Homestead, 27 Jan 1916, Harris C1626 (MIN), ibid., 15 Mar 1917, Harris C17366

(MIN); Murden Hammock, 2 mi W of Goulds, 28 Jan 1916, Harris C1639 (MIN); ibid., 27 Apr 1923, Harris N23226 (MIN); near LongView

School, 15 Mar 1917, Harris C17386 (MIN); Hattie Bauer Hammock, 19 Mar 1917, Harris C17398 (MIN); Royal Palm Hammock, 14 Dec 1922,

Harris C22937 (MIN); Tim’s Hammock, 27 Apr 1923, Harris N23158 (MIN); 0.5 mi S of Naranja, 27 Apr 1923, Harris N23257 (MIN); Royal

Palm State Park, 6 Nov 1927, Hawkins s.n. (FLAS); Everglades National Park, E end of Long Pine Key, 25 Jul 1987, Herndon 1760 (FLAS, FTG,

NY); Everglades National Park, Long Key Campground, 80°39.28'W, 25°24.24'N, 5 Apr 1999, Hess et al. 8560 (FLAS, NY); S of 216 St, 7.4 mi

SW of jet with 194 Ave, W of Goulds, 12 Mar 1975, Hill & Popenoe 2573 (FTG); Long Pine Key, Everglades National Park, 25 Jun 1970, Inman

20 (FTG); Tamiami Trail to Florida Keys, Dec 1929, Jennings s.n. (USF); Subtropical Horticultural Research Station, Chapman Field, Coral

Gables, 21 Jul 1989 Judd 5900 (FLAS); Goulds, near jet of248th Stand 159th Ave, 17 Jul 1995 Judd 7073 (FLAS); Girl Scout Park, Homestead,

13 Aug 1963, Lakela 26183 (USF); Homestead, 20 Jul 1964, Lakela 27323 (FLAS, USF); near North Homestead, 25 Jun 1968, Lakela 31900

(USF); U.S. Coast Guard Station, 25.64°N, 80.30°W, 18 May 2001, Lane et al. s.n. (FTG); Castellow Hammock, 25°3.6'E x 80°5.TE, 8 Jun

1999, Loho & Beck-King 031 (FTG); SE corner of SW 224 St and SW 120 Ave, Homestead, 30 Nov 1991, M ears s.n. (USF); SW 78 Ave & 176 St,

9 Feb 1974, M dinar s.n. (FTG); 1 mi W of Homestead, off Avocado Dr, Fuchs Hammock, 26 Jul 1990, Morris 4087 (FLAS); 0.2 mi W of Tower

Rd, 0.2 mi N of Lucille Dr (SW 360th St), 1.5 mi W of Florida City, Navy Wells Pineland, Royal Palm Ranger Station, 35°26 , 07"N,

80°30 , 23"W, 8 ft, 4 Sep 1999, Orzell & Bridges 18168 (FTG, USF); near Camp Osaissa Bauer, 3 mi N of Homestead, 5 Jul 1976, Popenoe 656

(FTG); Royal Palm State Park, 5 Mar 1944, Rhodes s.n. (FLAS); Everglades National Park, near Long Pine Key campground, 17 Dec 1971,

Poppleton s.n. (USF); Castellow Hammock, Dec 1966, Roberts 59 (FTG); Homestead, 7 Jun 1960, Rossbach 1876 (FLAS); Homestead, 7 Mar

1908, Sargent s.n. (A); SW 117 Ave and Killian Dr, Miami, 17 Jun 1980, Sauleda 3592 (USF); 2 mi N of Rt 27 on SW 217 Ave, 20 Oct 1968,

Schmal et al. s.n. (FTG); Homestead, 7 Jun 1937, Scull s.n. (FLAS); near Homestead Rd, between Cutler and Longview Camp, 9-12 Nov 1903,

Small & Carter 994 (MIN, NY); ibid., Small & Carter 1106 (NY); between Homestead and Camp Jackson, 1 Nov 1906, Small & Carter s.n.

(MIN); Long [Pine] Key, 6-7 May 1903, Small & Wilson 1649 (NY); near the Homestead Trail, near Camp Longview, 13-16 May 1904, Small

Judd et al., Revision of Miconia sect. Miconiastrum

473

& Wilson 1696 (NY); ibid., Small & Wilson 1932 (NY); between Homestead and Camp Jackson, 1 Nov 1906, Small & Carter 2625 (NY); be¬

tween Homestead and Cross Key, 21-22 Nov 1906, Small & Carter 2634 (NY); Long [Pine] Key, Everglades, 18-26 Jan 1909, Small & Carter

3060 (NY); near Goodburn Hammock, 21 Jun 1915, Small & Mosier 6377 (NY); ibid., Small & Mosier 6389 (NY); about Sykes Hammock, 2 Jul

1915, Small et al. 6749 (FLAS); Everglades National Park, Dryopteris Hammock, 9 Aug 1961, Smith 676 (USF); Everglades, 3 Jul 1971,

Sreemadhavan 4980 (USF); N of South Dade Baptist Church, Florida City, corner of 296 St and 172 Ave, 27Jul 1990, TanAV7 (USF); Fire gate

rd #1, Fong Pine Key, Everglades National Park, Tomlinson 21-V11-63C (FTG); Homestead, NE 10th St, 1.5 blocks E of Krome Ave, 28 Apr

1958, Traverse 650 (FTG); SW of Homestead, 14 May 1919, Small & Cuthbert s.n. (FFAS); Homestead, 10 May 1929, Weber s.n. (FFAS); 0.7 mi

W of Eureka Rd-Old Cutler Rd jet, R40E, T56S, 21 Jun 1960, Webster & Williams 10050 (FTG); N of Homestead, 18 Aug 1934, Welch 1525

(NY); Everglades National Park, near jet of Anhinga Rd off Research Center Rd, 7 Jul 1984, Whetstone 14428 (NY); between Homestead and

Florida City, near FF 27,4Jun 1966, Whitney 39 (USF); Redlands, Silver Palm Hammock, 23 Jun 1952, Wilson F-9 (USF); Tamiami Pineland

Complex Addition, 15 mi SW of downtown Miami, S of 12260 SW 130th St, 25.644°N, 80.393°W, 28 Apr 1999, Woodmansee 314 (FFAS,

FTG).

Miconia bicolor var. bicolor is the most common and widely distributed taxon of Miconia sect. Miconiastrum. It

is also the most variable. Plants of the Bahamas and southern Florida are phenetically most similar to those of

western Cuba (i.e., Prov. Pinar del Rio), however, they consistently have mite domatia, while those of Pinar del

Rio are variable: some having domatia and some lacking these structures. Some plants of Pinar del Rio and Isla

de la Juventud have the stems, abaxial surface of their leaves, and hypanthia nearly to somewhat glabrescent

and covered with a sticky secretion (e.g., Acevedo-Rdgz. et al. 5706, Acuna SC-22615, Becquer et al. HFC-85059,

Killip 44741, Roig & Cremata SV-7117, and Yero D. HFC-81190). Another group of plants have stems, abaxial

surface of their leaves, and hypanthia conspicuously lepidote, and the adaxial leaf surface often drying a dis¬

tinctive yellowish color (e.g., Becquer & Abbott HFC-82286, Becquer & More]on R. HFC-83986, Becquer & More-

jon R. HFC-84011, and Bro. Feon et al. FS-17821). Variation within Pinar del Rio may be ecologically correlated,

but more fieldwork (and intensive molecular sampling) is required. Plants of Hispaniola are distinctive in

having stems with hairs more or less globular-stellate, stellate-peltate, to dendritic and erect (see numerous

specimens cited above), while stems with appressed stellate-scales or branched hairs characterize plants of

other regions, except that a few specimens with erect, dendritic hairs also occur in plants of central Cuba (in

the Tope de Collantes, and Pico Potrerillo regions, Prov. Sancti Splritus; e.g., Acevedo-Rdgz. et al. 5593, Ekman

13999, Feon FS-18430). These Cuban collections also are distinctive in usually having obvious calyx lobes with

evident teeth, while most populations of M. bicolor var. bicolor have lobes that are essentially obsolete and lack

calyx teeth, although some plants with well-developed, or at least evident, lobes also have been collected in

Pinar del Rio (e.g., Britton et al. 7116, Farle 656, Shafer 11787). We note also that some collections from Prov.

Sancti Splritus (e.g., Loma los Helechales, Feon & Clement FS-5593 and Loma la Gloria, Feon & Roca FS-8003)

have well-developed calyx lobes, but stems with appressed hairs. The Hispaniolan collections, as is typical of

M. bicolor, usually have calyx lobes that are obsolete or nearly so, but some have more or less evident lobes and

teeth (e.g., Ekman H8722, Jimenez &ArizaJulia 5824, Feonard & Feonard 13629, Valeur 935). Some Hispaniolan

specimens also have stems and young leaves with a sticky secretion (as also seen in some western Cuban

plants). Most plants of central Cuba (Prov. Villa Clara, Lomas de Banao and Jatibonico in Sancti Splritus and

Cieneguitas, Cienfuegos) are distinctive in their often extremely pale to white-lepidote beneath (e.g., Alain

4013, Becquer & Veloso HFC-79904, Combs 128, Howard 5500, Feon FS-11369, and Smith & Hodgdon 3097). On

the other hand, material from Cumanayagua, Cienfuegos, are more or less ferrugineous beneath (and these

plants also have strongly plinerved leaves; see specimens examined). Leaf shape also varies greatly within M.

bicolor var. bicolor, and plants of Hispaniola tend to have leaves more similar to those of M. angustiflora than to

those of populations of M. bicolor var. bicolor in western and central Cuba, the Bahamas, or Florida. The abax¬

ial leaf indumentum of stellate-peltate scales, however, clearly distinguishes these Hispaniolan populations

from M. angustiflora, which has stellate hairs abaxially.

In summary, it is clear that several poorly differentiated geographical races exist within the array of popu¬

lations here considered as M. bicolor var. bicolor. These are not provided formal nomenclatural recognition be¬

cause individual specimens are not always clearly diagnosable as to the entity characteristic of each geographi¬

cal region. These geographically-correlated entities are 1) the populations of southern Florida and the Bahamas,

2) those of Prov. Pinar del Rio and the Isla de la Juventud, Cuba (and within this region there seems to be at least

474

Journal of the Botanical Research Institute of Texas 8(2)

two phenetic groups, perhaps ecologically correlated), 3) those of Prov. Villa Clara, Cuba, 4) those of the Tope

de Collantes and Pico Potrerillo regions, Prov. Sancti Splritus, Cuba, 5) those from Cumanayagua, Cienfuegos,

Cuba, and 6) those of the island of Hispaniola. The Tope de Collantes/Pico Potrerillo and Hispaniolan entities

are the most phenetically divergent. Finally, there are a few collections of M. bicolor from the “Oriente” region,

especially Guantanamo and Santiago de Cuba, and these require more study. Miconia bicolor appears to be

much less common in this region than it is in central and western Cuba, and some of these specimens are diffi¬

cult to distinguish from M. barbata or M. maestrensis (see discussion under these two species). More held work

in these eastern provinces of Cuba is needed, along with population-level molecular study. Some specimens,

e.g., Alvarez et al. HFC-55049, Becquer et al. HFC-82538, Bisse et al. HFC-26997, Bisse & Rojas HFC-3614, Bisse

& Kohler HFC-7412, are tentatively treated within M. bicolor, while others, e.g., Bros. Feon & Alain FS-19256,

Michelangeli et al. 1559, Shafer 2975 and Shafer 3765, although somewhat similar, are considered within M. bar¬

bata (because of the form of their hairs). We note that M. barbata and M. maestrensis have stellate hairs (not the

stellate-peltate scales of M. bicolor) on their adaxial leaf surfaces (see key for other differentiating characters).

It is obvious that Miconia bicolor is most diverse on the island of Cuba, and it is noteworthy that var. paten-

ti-setosa is endemic to Cuba. We hypothesize that the species originated on this island, only later dispersing to

Hispaniola, the Bahamas, and southern Florida.

Miconia bicolor was first reported from the United States by J.K. Small (1905) based on populations col¬

lected in pinelands ( Pinus elliottii Engelm.) of Miami-Dade county. The species is restricted to this county, and

is most characteristic of pinelands on Miami oolitic limestone. These pinelands largely have been destroyed by

agriculture and urban growth (Myers & Ewel 1990) and the species thus is considered threatened by the state

of Florida (Weaver & Anderson 2010). Miconia bicolor is occasionally used as an ornamental (Meerow 1991;

Osorio 1991; Haynes et al 2001;). Beautiful illustrations have been published by de la Sagra (1850), Sargent &

Faxon (1911), Correll & Correll (1982), and Wurdack & Krai (1982).

2b. Miconia bicolor (Mill. ) Triana var. patenti-setosa (Borhidi) Judd, Becquer, & Majure, comb. nov. Tetmzygia

bicolor (Mill.) Cogn. var. patenti-setosa Borhidi, Acta Bot. Acad. Sci. Hung. 23:39. 1977 [1978], Type: CUBA. Prov. Pinar del Rio:

Cerro de Cabras, 10 Oct 1954, fr, Acuna & Torres 19461 (holotype: HAC!; isotype: HAC!, HAJB!).

Stems and inflorescence axes with scattered, elongate, multicellular, eglandular hairs.

Phenology. —The taxon has been collected in fruit only in October.

Distribution.—Miconia bicolor var. patenti-setosa is only known from Cuba, Prov. Pinar del Rio, in the

Cerro de Cabras.

Miconia bicolor var. patenti-setosa is known only from the type; additional collections are needed, and the

taxon is here only provisionally recognized. It is of interest that no other taxon of Miconia sect. Miconiastrum

has elongate, multicellular hairs on the stems; such hairs are only known from mite domatia.

3. Miconia guajaibonensisjudd, Becquer, & Majure, nom. nov. (Fig. 3C, D). Tetrazygia lanceolata Urb., Repert. Spec.

Nov. Regni Veg. 22:223.1926, non M. lanceolata DC., Prodr. 3:190.1828. Type: CUBA. Prov. Pinar del Rio: Pan de Guajaibon, on the

very top of the mountain, 750-800 m, 9 Jan 1921, fl, E.L. Ekman 12767 (lectotype: S!, here designated, because holotype at B de¬

stroyed; isolectotype: NY!, fragment).

Evergreen shrub up to 4.5 m. Young stems terete to slightly quadrangular, the indumentum of dense, multicel¬

lular, ferrugineous, matted to erect, dendritic to globular stellate hairs or appressed stellate-peltate scales, be¬

coming ± glabrate with age, without elongate, multicellular, non-glandular hairs, internodes 0.5-1.8 cm long,

lacking longitudinal ridges, nodal line present, faint. Leaves isophyllous or nearly so; petiole 0.7-2.5 cm long,

the indumentum of dense, ± matted, stellate-peltate scales to stellate-peltate scales, partially deciduous with

age; the blade 3-8.6(-9.3) x 0.7-2.4(-3.3) cm, ovate, usually falcate, chartaceous, the apex acuminate to attenu¬

ate, the base obtuse to rounded, the margin plane to slightly revolute, entire; secondary veins two pairs, one

pair conspicuous and one pair inconspicuous, acrodromous, basal, the innermost pair joining midvein at base

to 1 mm above the leaf base, the conspicuous secondary veins placed 0.8-3.5 mm in from margin, the incon¬

spicuous secondary veins intramarginal, tertiary veins percurrent, oriented subperpendicular to midvein,

Judd et al., Revision of Miconia sect. Miconiastrum

475

0.7-4 mm apart, quaternary veins obscure to visible, ± reticulate, not connecting adjacent tertiary veins or with

1-3 quaternaries connecting adjacent tertiary veins, the higher order veins reticulate; the midvein and major

secondary veins slightly impressed to flat, tertiary and higher order veins flat on adaxial surface, the midvein

moderately to strongly raised, the major secondary veins flat to moderately raised, the minor secondary veins

and tertiary veins flat, and the higher order veins flat on the abaxial surface; adaxial surface appearing smooth

to slightly wrinkled after drying, with scattered to abundant druse crystals, and drying darker than the abaxi¬

al surface, the indumentum initially of dense, pale ferrugineous, stellate-peltate scales, but very quickly gla-

brescent, although a few scales often retained on midvein proximally; abaxial surface pale green but pale fer¬

rugineous due to dense covering of stellate-scales, the epidermal surface ± smooth, with dense, appressed,

ferrugineous or pale ferrugineous, stellate-peltate scales, 0.09-0.17 mm across, the veins with similar scales,

but sometimes more irregular in form, such hairs on lamina and veins ± persistent; mite domatia present or

absent, at junction of midvein and major secondary veins, consisting of dense cluster of multicellular, elongate,

pale ferrugineous to red, barbate, eglandular hairs, 0.5-1 mm long. Inflorescences terminal, paniculate

cymes, 3.5-9 cm long, 2.5-6.5 cm across, with 2-4 major branch pairs, the peduncle 1.7-3.8 cm long, the ulti¬

mate axes (pseudopedicels) 1-5.5 mm long, and numerous flowers well separated from each other; bracts 0.7-1

x 0.2-0.4 mm, but probably also larger, ± triangular or narrowly so, with acute to obtuse apex, early deciduous;

bracteoles 0.4-0.7 x 0.1-0.2 mm, narrowly triangular, with acute apex, ± deciduous. Flowers 5-merous, zygo-

morphic (due to positioning of the stamens), with pedicel 1.3-3 mm long. Hypanthium 4.5-6 mm long, terete

and ± slightly constricted above ovary, the free portion 1.8-3.3 mm long, 2.2-2.8 mm wide at the torus, the

outer surface with moderate to dense, stellate-peltate scales, the internal surface smooth, glabrous. Calyx lobes

5, separate (and imbricate) in bud, the tube 1-1.7 mm long, irregularly tearing between lobes, with moderate to

dense stellate-peltate scales abaxially, and moderate to dense stellate-peltate to globular stellate hairs adaxially,

the lobes unequal, usually the 2 outer lobes larger than the rest, 0.45-1.2 x 1-2.4 mm, triangular to shortly

triangular, the apex rounded to obtuse, with hairs similar to those of the tube, green, often red-tinged; calyx

teeth present, ±0.1 mm long, merely a dorsal, slightly ellipsoidal bump, with rounded apex, and hairs similar

to those of calyx. Petals 5,4.5-6.3 x 2.5-4 mm, obovate, spreading, white, glabrous but both surfaces densely

papillose-granulate, the apex rounded and asymmetrically notched, the base slightly narrowed to a broad at¬

tachment, the margin entire. Stamens 10, isomorphic, ± geniculate near filament apex; staminal filament

4.5-5.5 mm long, glabrous, pale yellow, anther thecae 5-5.8 x 0.6-0.8 mm, subulate, straight to slightly in¬

curved, opening by a small dorso-apical pore, yellow, the connective not prolonged below the thecae, glabrous,

but the filament emerging from slight depression in sterile, minutely dorso-lobed anther base. Ovary 3-locular,

ca. 4/5-inferior, 3.4-3.7 mm long, 1.4-2 in diameter, oblong to ellipsoid, apically conic, unridged, glabrous, the

apex with a short, slightly ridged collar but lacking a crown, with axile placentation, the ovules numerous,

borne on a small placenta not extending or only slightly extending into locule; style 10-13 mm long, distally

curved, white, glabrous; stigma punctate, minutely papillose, 0.15-0.2 mm wide. Berries 5-8 mm long, 4.3-

6.5 mm in diameter (excluding persistent hypanthium and calyx tube), globose to ellipsoid, purple-black, with

scattered stellate-peltate scales, the hypanthium constricted, 1.3-1.7 mm wide at narrowest point and 2-3 mm

wide at torus, but appearing strongly constricted due to flaring calyx tube. Seeds 1.1-1.6 mm long, angular-

obovoid, with ± rectangular raphe, the testa ± smooth; appendage absent.

Phenology. —Recorded in flower injanuary, March, May and November.

Distribution and habitat.—Miconia guajaibonensis is endemic to Pan de Guajaibon, in Pinar del Rio, Cuba

(Fig. 5), from 680-800 m, in thickets over limestone.

Additional specimens examined: CUBA. Prov. Pinar del Rio: Pan de Guajaibon, la Mulata, 16 May 1953, Acuna & Bro Alain SV-18519 (HAC);

en la cumber del Pan de Guajaibon, 750 m, 2 May 1959, Bro. Alain 6778 (HAC); Bahia Honda, Pan de Guajaibon, ladera N, cerca de la cima,

680-700 m, 30 Mar 2004, Becquer HFC-81700 (HAJB); cima del Pan de Guajaibon, 700 m, Jul 2005, Becquer & Abbott HFC-82431 (FLAS,

HAJB); La Palma, cima del Pan de Guajaibon, 720 m, 28 Dec 1970, Bisse HFC-15640 (HAJB); Consolacion del Norte, Pan de Guajaibon, 14

Oct 1976, Bisse et al. HFC-32501 (HAJB); segunda mitad del Pan de Guajaibon, 14 Apr 1980, Herrera et al. SV-29487 (HAC); ibid., Herrera

et al. SV-29488 (HAC); Guajaibon, 4 Nov 1860-1864 [probably 1862], Wright 1222 p.p. (CAS on-line image, GH, MO, P on-line image, S).

476

Journal of the Botanical Research Institute of Texas 8(2)

Few collections of this rare and geographically restricted species are known, and Miconia guajaibonensis is

morphologically quite uniform, although it seems to vary in the development of mite domatia, as in the west¬

ern Cuban populations of Miconia bicolor. These two species likely are closely related, as both are characterized

by an indumentum of stellate-peltate scales on the abaxial leaf surface; see key for their differentiating charac¬

teristics.

The new name, Miconia quajaibonensis, is needed because the epithet “lanceolata” is occupied in Miconia,

i.e., M. lanceolata DC. The new epithet highlights the only known locality at which this rare (and possibly en¬

dangered) species occurs. Currently, six species, including M. guajaibonensis, are known to be endemic to Pan

de Guajaibon (Borhidi & Munoz 1985; Borhidi 1991; Judd et al. 2008; Morejon & Sanchez 2012).

4. Miconia impressa (Urb.) Judd, Becquer, & Majure, comb. nov. (Fig. 3E). Tetrazygia impressa Urb., Repert. Spec. Nov.

Regni Veg. 22:225. 1926. Type: CUBA. Prov. Pinar del Rio: Sierra de los Organos, Grupo del Rosario, Sierra de Pendejeral, on top of

the ridge, 14 Sep 1923, fl, fr, E.L. Ekman 17539 (lectotype: S!, here designated, because specimen at B destroyed; isolectotype: NY!,

fragment).

Tetrazygia versicolor Urb., Repert. Spec. Nov. Regni Veg. 22:224. 1926. Type: CUBA. Prov. Pinar del Rio: Sierra del Sitio Santo Tomas,

steep limestone cliffs, 500 m, 10 Jun 1923, fl, E.E. Ekman 16625 (lectotype: S!, here designated, because specimen at B destroyed;

isolectotype: US, on-line image #00120756!).

Tetrazygia minor Urb., Repert. Spec. Nov. Regni Veg. 22:224. 1926. Tetrazygia lanceolata Urb. subsp. minor (Urb.) Borhidi, Acta Bot.

Acad. Sci. Hung. 23:79.1977 [1978]. Type: CUBA. Prov. Pinar del Rio: San Diego de los Banos, at the foot of Sierra de la Guira, 24 Oct

1923, sterile, E.L. Ekman 17785 (lectotype: S on-line image #05-3527!, here designated, because specimen at B destroyed; isolecto¬

type: NY!, fragment, on-line image #00099743).

Evergreen shrub up to ca. 3 m. Young stems terete to slightly quadrangular, the indumentum of dense, matted,

pale ferrugineous, stellate to dendritic hairs, becoming only sparse with age, without elongate, multicellular,

non-glandular hairs, internodes 0.5-4.2 cm long, lacking longitudinal ridges, nodal line absent or present,

faint. Leaves isophyllous or nearly so; petiole 0.7-3.7 cm long, the indumentum of dense, stellate to dendritic

Judd et al., Revision of Miconia sect. Miconiastrum

477

hairs; the blade 2.1-9.7 x 0.8-3.6 cm, ovate to elliptic, not falcate, chartaceous to thinly coriaceous, the apex

acute to acuminate, the base obtuse to rounded or slightly cordate, the margin plane to slightly revolute, entire;

secondary veins in two pairs, one pair conspicuous and one pair inconspicuous, acrodromous, basal, the in¬

nermost pair joining midvein at base to 1.5 mm above the leaf base, the conspicuous secondary veins placed

1.3-6 mm in from margin, the inconspicuous secondary veins intramarginal to 1 mm in from margin, tertiary

veins percurrent, oriented subperpendicular to midvein, 1-5.5 mm apart, connected by a single quaternary

vein, or quaternary veins reticulate, not connecting tertiary veins, the higher order veins reticulate, the mid¬

vein and major secondary veins moderately to slightly impressed (occasionally flat), tertiary veins slightly im¬

pressed to flat, remaining veins flat on adaxial surface; the midvein strongly raised, the major secondary veins

moderately to slightly raised, the minor secondary veins and tertiary veins slightly raised to flat, and the

higher order veins flat on abaxial surface; adaxial surface appearing slightly wrinkled after drying, with scat¬

tered druse crystals, and drying darker than the abaxial surface, the indumentum initially of dense, white to

very pale ferrugineous, globular stellate to stellate hairs, but very quickly glabrescent, although sometimes a

few hairs remaining on proximal portion of midvein; abaxial surface pale green, the surface smooth to very

slightly bullate, with ± dense, white to very pale ferrugineous, stellate hairs, usually 0.12-0.24 mm across, the

veins with dense, similar hairs, such hairs on lamina and veins persistent to rarely somewhat deciduous; mite

domatia usually present at junction of midvein and major secondary veins, at least on some leaves of each

shoot, but sometimes such domatia absent, when present well to poorly developed, composed of elongate,

multicellular, eglandular, pale yellow to ferrugineous, barbate hairs, 0.2-1 mm long. Inflorescences terminal,

paniculate cymes, 4-8.5 cm long, 2.5-4.5 cm across, with 2-4 major branch pairs, the peduncle 1-5 cm long,

the ultimate axes (pseudopedicels) 1-4 mm long, and the numerous flowers well separated from each other;

bracts quickly deciduous, not seen, probably ± triangular; bracteoles 0.5-1.2 x 0.15-0.2 mm, narrowly triangu¬

lar, with acute apex, deciduous. Flowers 5-merous, zygomorphic (due to positioning of the stamens), with

pedicel 0.5-1.5 mm long. Hypanthium 5-6 mm long, terete and not to only slightly constricted above ovary,

the free portion 25 - 1.1 mm long, 1.5-2.5 mm wide at the torus, the outer surface with ± dense stellate hairs,

the internal surface smooth to very slightly longitudinally ridged, glabrous. Calyx lobes 5, fused in bud as a

conical calyptra, but with a minute, apical opening, the tube 1.4-1.6 mm long, consistently and usually regu¬

larly tearing, with calyptra-like remnants consisting of 2 or 3 lobes, occasionally tearing more regularly be¬

tween lobes, and at anthesis the tube thus appearing only ca. 0.5 mm long, with moderate to dense stellate hairs

abaxially, and sparse to moderate branched to stellate hairs adaxially, the lobes 0.3-0.6 x 0.15-0.5 mm, ± tri¬

angular (but actual calyx lobes not evident at anthesis, appearing as a small apiculum at apex of torn, irregular

to ± triangular pseudo-lobes), the apex acute, with hairs similar to those of the tube, green; calyx teeth present,

0.1-0.2 mm long, represented merely by an ellipsoidal bump, nearly filling each of the reduced calyx lobes,

green, with rounded apex, and hairs similar to those of calyx. Petals 5, 4.5-6 x ca. 3.5-4.2 mm [but possibly

larger, as values based on slightly immature flower and original description], asymmetrically obovate, spread¬

ing to reflexed, white, glabrous but both surfaces densely papillose-granulate, the apex rounded, slightly

notched, the base slightly narrowed to a broad attachment, the margin entire. Stamens 10, isomorphic, ± ge¬

niculate near filament apex; staminal filament 3.5-6 mm long [measurements based on slightly immature

flower and original description], glabrous, white, anther thecae 4.5-6 x 0.8-1 mm, subulate, straight to

slightly incurved, opening by a small dorso-apical pore, yellow, the connective not prolonged below the thecae,

glabrous, but the filament emerging from slight depression in sterile, minutely dorso-lobed anther base. Ovary

3-locular, ca. 3/4 inferior, ± ellipsoid, 2-4.5 mm long, 1.5-2.5 mm in diameter, apically cylindric-conic, gla¬

brous, the apex with small slightly ridged collar but lacking crown, with axile placentation, the ovules numer¬

ous, borne on a small placenta that extends only slightly into locule; style 9-11 mm long, nearly straight to

distally curved, white to pink, glabrous; stigma punctate, minutely papillose, ca. 0.15 mm wide. Berries 4-8.5

mm long, not including the persistent hypanthium and calyx, 3.5-7 mm in diameter, globose, subglobose, or

ellipsoid, purple-black, with scattered stellate hairs, the hypanthium constricted, 1.3-2.5 mm wide at narrow¬

est point and 1.7-3.5 mm wide at torus, but appearing strongly constricted due to expanded calyx tube. Seeds

1-1.7 mm long, angular-obovoid, with ± rectangular raphe, the testa ± smooth; appendage absent.

478

Journal of the Botanical Research Institute of Texas 8(2)

Phenology. —Collected in flower in June, July and September.

Distribution and habitat.—Miconia impressa is endemic to Pinar del Rio, in western Cuba (Fig. 5), occur¬

ring from 500-614 m in thickets over limestone.

Additional specimens examined. CUBA. Prov. Pinar del Rio: Vinales, cresta del mogote al O del sendero Coco Solo-Palmarito, 22°37.409N,

83°43.282W, 377 m, 21 Jul 2013, Barrios et al. (HAJB); Banos San Vicente, rocky limestone hill, 12-16 Sep 1910, N.L. Britton et al. 7464 (NY,

US); sierra Sitio del Infierno, base S, ascenso y cima del mogote de El Americano, cumber mas alta, 614 m, 22 Jul 1991, Sdnchez HFC-69824

(HAJB); mogote de Dos Hermanas, Vinales, 7 Oct 1989, Urquiola et al. HPR-5629 (HAJB, HPPR); cima de la Sierra del Infierno, Vinales, 4Jul

1990, Urquiola et al. HPR-6004 (HAJB, HPPR); without definite locality, Wright s.n. (GH, NY).

Miconia impressa is distinctive, and separabie from other species of the M. bicolor compiex, because of its flower

buds that have a nearly closed (apically), conical calyptra (formed from the calyx tube), so the petals are hidden

in bud (although minute, free calyx lobes are still present at the apex of the calyptra). As the flowers mature, the

calyx tube usually tears completely and irregularly as a calyptra (of one or two pieces, each formed by fused

lobes) or occasionally into more or less regular “lobes,” allowing the petals and stamens to emerge. It is also

distinctive in having leaves with a dense, abaxial indumentum of more or less white, stellate hairs (see key).

The species is somewhat variable in leaf size, the impression of the veins adaxially, and in the presence, and

prominence of mite domatia. Thus, Tetrazygia versicolor and T. minor are here considered within the circum¬

scription of M. impressa.

5. Miconia barbata (Borhidi) Judd, Becquer, & Majure, comb. nov. (Fig. 6A). Tetrazygia barbata Borhidi, Acta. Bot.

Acad. Sci. Hung. 23:38. 1977 [1978], Type: CUBA. Prov. Oriente [Holguin]: Playa de Moa, 25 Jul 1941, Bro. Leon LS-20286, with Bro.

Clemente, &R. Howard (holotype: HAC!; isotypes: NY!, HAC!).

Evergreen shrub or tree to 5.5 m. Young stems terete to slightly quadrangular, the indumentum of dense,

brunnescent to pale ferrugineous, stellate, globular-stellate to dendritic hairs, without elongate, multicellular,

non-glandular hairs, internodes 0.7-8 cm long, lacking longitudinal ridges, nodal line present, faint. Leaves

isophyllous or nearly so; petiole 1.6-3 cm long, the indumentum of brunnescent to ± ferrugineous, globular-

stellate to stellate hairs to stellate-peltate scales with ± free arms; the blade 4.5-19 x 1.6-5.6 cm, ovate to elliptic

or oblong, ± falcate, chartaceous to thinly coriaceous, the apex acuminate to attenuate, the base obtuse to

rounded, the margin plane to slightly revolute, entire to slightly undulate; secondary veins 2 pairs, one pair

conspicuous and one pair inconspicuous, acrodromous, basal, the innermost pair joining midvein at base to 4

mm above the leaf base, the conspicuous secondary veins placed 1.7-11 mm in from margin, the inconspicu¬

ous secondary veins intramarginal to 2.3 mm in from margin, tertiary veins percurrent, oriented subperpen¬

dicular to midvein, 1.5-9 mm apart, connected by 1 or 2 quaternary veins, or such veins ± reticulate and not

connecting the tertiaries, the higher order veins reticulate, the midvein and major secondary veins slightly

impressed, tertiary veins slightly impressed to flat, remaining veins flat on adaxial surface; the midvein strong¬

ly raised, the major secondary veins moderately to slightly raised, the minor secondary veins and tertiary veins

slightly raised to flat, and the higher order veins flat on abaxial surface; adaxial surface appearing slightly

wrinkled after drying, with scattered druse crystals, and drying darker than the abaxial surface, the indumen¬

tum initially of ± dense, brunnescent to pale ferrugineous, globular-stellate hairs and stellate-peltate scales

with ± free arms, but very quickly glabrescent, although sometimes a few hairs persistent on proximal portion

of midvein; abaxial surface pale green but ± brunnescent due to indumentum, the surface smooth to bullate,

with moderate to dense, brunnescent to pale ferrugineous, globular-stellate hairs and stellate-peltate scales

with ± free arms, 0.13-0.24 mm across, the veins with similar hairs, such hairs on lamina persistent to partly

deciduous and those on the veins also persistent to deciduous, especially on midvein and major secondary

veins, but occasionally also on tertiary veins; mite domatia well-developed (i.e., to 11 mm in length), consis¬

tently present, at junction of midvein and major secondary veins, sometimes also at junction of midvein and

tertiary veins, and formed by tuft or ± dense mass of elongate, barbate, pale yellow to ferrugineous, eglandular

hairs, from 0.2-1.7 mm long. Inflorescences terminal, paniculate cymes, 6-16 cm long, 3.5-11 cm across,

with 3-5 major branch pairs, the peduncle 1.4-5 cm long, the ultimate axes (pseudopedicels) 1-4.5 mm long,

and the numerous flowers well separated from each other; bracts 0.5-1 x 0.2-0.3 mm, but probably also larger,

Judd et al., Revision of Miconia sect. Miconiastrum

479

Fig. 6. A. Miconia barbata {Ekman 9484). B. M. angustiflora {Judd5302). C. M. cajalbanensis {Becquer& Abbott 82423). D. M. maestrensis (Ekman 14920).

480

Journal of the Botanical Research Institute of Texas 8(2)

narrowly triangular to triangular, the apex acute, quickly deciduous; bracteoles 0.3-0.5 x 0.1-0.3 mm, nar¬

rowly triangular to triangular, with acute apex, deciduous. Flowers 5- or occasionally 6-merous, zygomorphic

(due to positioning of the stamens), with pedicel 1-2.5 mm long. Hypanthium 4.5-7.5 mm long, terete and

slightly constricted above ovary, the free portion 2.5-4 mm long, 2-2.7 mm wide at the torus, the outer surface

with moderate to dense stellate hairs, the internal surface smooth to very slightly longitudinally ridged, gla¬

brous. Calyx lobes 5 (6), separate in bud, the tube 0.8-2.3 mm long, usually not tearing between lobes, with

moderate to dense stellate hairs abaxially, with moderate to dense branched to stellate hairs adaxially, the lobes

0.3-1.5 x 1.5-3 mm, triangular to broadly triangular, the apex acute to obtuse or rounded, with hairs similar

to those of the tube, green, sometimes red-tinged; calyx teeth present, 0.1-0.15 mm long, merely a small bump

near apex of lobe, green, with rounded apex, and hairs similar to those of calyx. Petals 5 (6), 6-8.5 x 3.5-5.5

mm, asymmetrically obovate, spreading, white (but pink-tinged with age), glabrous but both surfaces densely

papillose-granulate, the apex ± rounded, sometimes obscurely notched, the base slightly narrowed to a broad

attachment, the margin entire. Stamens 10 (12), isomorphic, ± geniculate near filament apex; staminal fila¬

ment 5.3-7 mm long, glabrous, pale yellow, anther thecae 5-7.7 x 0.9-1.2 mm, subulate, straight to slightly

incurved, opening by a small dorso-apical pore, yellow, the connective not prolonged below the thecae, gla¬

brous, but the filament emerging from slight depression in sterile, minutely dorso-lobed anther base. Ovary

3-locular, ca. 3/4 inferior, ellipsoid to ± oblong, apically conical, glabrous, the apex with a small, slightly ridged

collar but lacking a crown, with axile placentation, the ovules numerous, borne on small placenta not or only

slightly extending into locule; style 10-15 mm long, distally curved, white (but pink-tinged with age), gla¬

brous; stigma punctate, minutely papillose, ca. 0.15 mm wide. Berries 6-13 mm long, 4-10 mm in diameter,

ellipsoid to nearly globose, purple-black, with scattered, stellate hairs, the hypanthium constricted, 1.5-2.5

mm wide at narrowest point and 2-3 mm wide at torus, but appearing strongly constricted due to flaring calyx

tube. Seeds 1-2 mm long, angular-obovoid, with ± rectangular raphe, the testa slightly to distinctly roughened

due to bulging cells, except on distal surface where ± smooth, the raphe also smooth; appendage absent.

Phenology .—Recording in flower in March (and likely also April), July, August, October, and December,

and probably flowering year around.

Distribution and habitat.—Miconia barbata is restricted to the central and northern “Oriente” region,

mainly in Prov. Guantanamo and Holguin, Cuba, occurring in the Sierra de Nipe, mountains near Bayate,

Monte Verde, and in the vicinity of Moa and Baracoa (Fig. 7), from near sea-level to 745 m, in pinelands, thick¬

ets, forests along arroyos, and semideciduous forests on limestone.

Additional specimens examined: CUBA. Prov. Guantanamo: prope Bayate, Sabana Miranda, 11 July 1914, Ekman 1946 (NY, S) Bayate, Cayo

del Rey, 21 Feb 1915, Ekman 4694 (S); Charrascal de la Cuaba, Baracoa, 12 Apr 1969, L. Figueiras UO-778 (HAJB); Baracoa, Taco Bay, camino

de Yamaniguey, 12 Apr 1960, L. Figueiras UO-778 (HAC); ibid., 24 Jul 1969, L. Figueiras UO-2166 (HAC, HAJB); Baracoa, Camino Alto La

torre, Nibujon, SE de Taco Bay, 20° 30T.8" N, 74° 394-3.2" W, 150 m, 23 Oct 2009, M ichelangeli et al. 1504 (FLAS); Monte Verde, La Perla, 4

May, 2 Jul,1859-1864, Wright 1222 p.p. (CAS on-line image, MO, P on-line image). Prov. Holguin: Sierra de Nipe, loma al S de la Unidad Sil-

vicola Forestal “Pinal Redondo,” cerca del arroyo La Chivera, 480 m, 25 Apr 2004, Becquer HFC-82254 (FLAS, HAJB); Moa, Rio Quesigua,

donde cruza la carretera de Moa-Baracoa, 5-10 m, 27 Mar 2009, Becquer et al. HFC-85468 (HAJB); Sierra de Nipe, orillas del rio Piloto, cerca

de las cabezadas, Apr 1968, Bisse & Kohler HFC-8529 (HAJB); Cuevas de Purnio, 5 Nov 1971, Bisse et al. HFC-20767 (HAJB); Moa, Los Faral-

lones de Moa, 300 m, 31 Mar 1972, Bisse & Berazain HFC-22229 (HAJB); Mayan Abajo, Sierra de Nipe, orillas de arroyos en el camino a

Woodfred, 600 m, 2 Jun 1979, Bisse et al. HFC-36020 (HAJB); Moa, Monte la Brena alrededores del campamento Los Carboneros camino

hacia el rio Yagrumaje, 300 m, 17 Apr 1981, Bisse et al. HFC-44469 (HAJB); Mayan, Pinares de Mayan, cerca del campamento de pineros en

el valle superior del rio Piloto, 400 m, 31 May 1983, Bisse et al. HFC-50610 (HAJB); Moa, Cerro de la Embocadura del rio Yamaniguey, 24 Mar

1970, Borhidi et al. SV-27946 (HAC); ibid., Borhidi et al. SV-27947 (HAC); Moa, Dec 1939, Mrs. Bucher 144 (HAC, NY); Moa, Bucher SV-11442

(HAC); Sierra de Nipe, El Taller (ad Rio Piloto), 21 Jul 1914, Ekman 2089 (S, US); Sierra de Nipe, ad Rio Piloto, 5 Oct 1919, Ekman 9684 (NY,

S, US); Pinares de Mayan, Sierra de Nipe, 27-31 May 1960, L. Figueiras UO-952 (HAC, HAJB); Moa, camino a mina Mercedita, 16 Mar 2000,

Fritsch et al. SV-41362 (HAC); 5 km S of Woodfred, Sierra de Nipe, Jul 1941, Howard 6149 (GH, MO, NY); Moa, 30 Jul or 3 Aug 1945, Bro. Feon

et al. FS-22506 (GH, HAC, HAJB); Charrasco de la Cueva, Sierra de Nipe, 700 m, 27Jul 1940, Bros. Feon & Alain 19256 (NY); Moa, Charrascal

1.5 km al E de Yamaniguey, carretera Moa-Baracoa, 20°34 , 22.8"N, 74°4446.2"W, 10 m, 26 Oct 2009, M ichelangeli et al. 1539 (FLAS, HAJB,

NY); Mayan, via Cayo Mujeres, 3 km al N y luego 1 km al O por la carretera desde el Hotel Pinares de Mayan, 20 o 31'39.6"N, 75 0 45'34.8"W,

745 m, M ichelangeli et al. 1559 (FLAS, HAJB, NY); crest of Sierra Nipe, 600-700 m, 16-17 Oct 1941, Morton & Acuna 3044 (DUKE, F, FLAS,

FTG, MO, NY, US); Sierra Nipe, near Woodfred, 500-650 m, 4 Dec 1909, Shafer 2975 (NY); near base of Loma Mensura, 680 m, 1-3 Feb 1910,

Shafer 3765 (NY, US).

Judd et al., Revision of Miconia sect. Miconiastrum

481

Miconia barbata shows some geographically correlated variation, with plants of the Sierra de Neiba often hav¬

ing a denser abaxial leaf indumentum than those of other regions; additionally, the plants of the Moa region

have abaxial epidermal cells that are more clearly bullate (and their abaxial epidermis is clearly visible, because

many of the stellate hairs are deciduous on older leaves).

The species often has been confused with Miconia bicolor (see specimen annotations) even though it is

readily distinguished by the indumentum of the abaxial leaf surface, i.e., with stellate hairs or stellate-peltate

scales with largely free arms in M. barbata and with stellate-peltate scales (i.e., the arms usually strongly fused)

in M. bicolor (see also Borhidi, 1978, who described the indumentum of M. barbata as radiate scales). Abaxial

mite domatia are consistently present in M. barbata (although they range from prominent to occasionally more

or less inconspicuous), while they may or may not occur in M. bicolor. The two species are partly geographi¬

cally isolated (Figs. 4, 7) but likely co-occur (more held work is needed in the “Oriente” region). Miconia bar¬

bata is actually much more difficult to distinguish from M. angustiflora, a Jamaican endemic, which has less

well developed mite domatia (and frequently entirely lacks such domatia; see key).

6. Miconia angustiflora (Benth.) Naudin, Ann. Sci. Nat. Bot. ser. 3, 16:246. 1850. (Fig. 6B). Diplochita angustiflora

Benth., Pi. Hartw. 263.1846. Tetrazygia angustiflora (Benth.) Griseb., Goett. Abh. 9:53.1861. Type: JAMAICA: Stony Hill (not seen).

Tetrazygia ovata Cogn., Symb. Antill. 5(3):447.1908. Type: JAMAICA. [St. Catherine Parish]: Holly Mount, prope Ewarton, 867 m, 2600

ft, 11 Aug 1896, fl, W. Harris 6450 (holotype: B, destroyed; isotypes: F!, NY!, 2 sheets, on-line images #00099744 and 00099745!).

Evergreen shrub or tree to 10 (-15) m. Young stems terete to slightly quadrangular, the indumentum of dense,

ferrugineous to pale ferrugineous, stellate to globular-stellate hairs, without elongate, multicellular, non-glan-

dular hairs, internodes 0.5-11.3 cm long, lacking longitudinal ridges, nodal line present, faint. Leaves isophyl-

482

Journal of the Botanical Research Institute of Texas 8(2)

lous or nearly so; petiole 1.3-6.6 cm long, the indumentum of dense to moderate, stellate hairs; the blade 5.3-

20 x 1.8-8.3 cm, ovate to elliptic or obovate, falcate or not, coriaceous, the apex acuminate to attenuate, the

base narrowly cuneate or acute to obtuse or rounded, the margin plane to slightly revolute, entire to slightly

undulate; secondary veins 2 pairs, one pair conspicuous and one pair inconspicuous, acrodromous, basal, the

innermost pair joining midvein at base to 10 mm above the leaf base, the conspicuous secondary veins placed

3-13 mm in from margin, the inconspicuous secondary veins intramarginal to 3 mm in from margin, tertiary

veins percurrent, oriented subperpendicular to midvein, 2-9 mm apart, connected by 1-3 quaternary veins, or

not connected and quaternary veins reticulate, the higher order veins reticulate, the midvein and major sec¬

ondary veins slightly impressed to flat, tertiary veins slightly impressed to more commonly flat, remaining

veins flat on adaxial surface; the midvein strongly raised, the major secondary veins slightly to moderately

raised, the minor secondary veins and tertiary veins slightly raised to flat, and the higher order veins flat on

abaxial surface; adaxial surface appearing smooth and punctate to slightly wrinkled after drying, with scat¬

tered druse crystals, and drying darker than the abaxial surface, the indumentum initially of dense, pale fer-

rugineous, stellate hairs, but very quickly glabrescent, although scattered hairs often retained on lamina, and

numerous hairs often retained on midvein and major secondary veins; abaxial surface pale green, the surface ±

smooth, with dense to moderate (rarely sparse), ferrugineous to pale ferrugineous, stellate hairs (with arms

predominantly radiating outward), 0.13-0.25 mm across, the veins with similar hairs, such hairs on lamina

usually ± persistent, and those of primary, secondary, and even tertiary veins often deciduous (but sometimes

persistent as well), mite domatia present or absent, usually not well developed, at junction of midvein and ma¬

jor secondary veins, with elongate, ± ferrugineous, barbate, eglandular hairs, 0.1-1 mm long. Inflorescences

terminal, paniculate cymes, 5-22 cm long, 4-13 cm across, with 3-7 major branch pairs, the peduncle 1-6.6

cm long, the ultimate axes (pseudopedicels) 1-7 mm long, and the numerous flowers well separated from each

other; bracts 0.7-1.5 x 0.2-0.3 mm, but probably also larger, narrowly triangular, with acute apex, quickly de¬

ciduous; bracteoles 0.4-1 x 0.1-0.25 mm, narrowly triangular, with acute apex, deciduous. Flowers 5- or oc¬

casionally 6-merous, zygomorphic (due to positioning of the stamens), with pedicel 1.5-6 mm long. Hypan-

thium 4.5-6 mm long, terete and slightly constricted above ovary, the free portion 2.5-3.5 mm long, 2-3 mm

wide at the torus, the outer surface with moderate to dense stellate hairs, the internal surface smooth to very

slightly longitudinally ridged, glabrous. Calyx lobes 5 (6), separate in bud, the tube 0.8-1.5 mm long, not tear¬

ing between lobes, with moderate to dense stellate hairs abaxially, with sparse to moderate branched to stellate

hairs adaxially, the lobes 0.2-0.8 x 2-3.5 mm, broadly triangular to nearly obsolete, the apex obtuse to round¬

ed, with hairs similar to those of the tube, green, sometimes red-tinged; calyx teeth present, 0.1-0.2 mm long,

a small bump near apex of each lobe, green, with rounded apex, and hairs similar to those of calyx. Petals 5 (6),

7-11 x 3.5-6.5 mm, asymmetrically obovoid, spreading, white, glabrous but both surfaces densely papillose-

granulate, the apex rounded and ± notched, the base slightly narrowed to a broad attachment, the margin en¬

tire. Stamens 10 (12), isomorphic, ± geniculate near filament apex; staminal filament 5.5-8 mm long, gla¬

brous, pale yellow, anther thecae 5.2-7.9 x 0.8-1.2 mm, subulate, straight to slightly incurved, opening by a

small dorso-apical pore, yellow, the connective not prolonged below the thecae, glabrous, but the filament

emerging from slight depression in sterile, minutely dorso-lobed anther base. Ovary 3-locular, 3/4-4/5 inferi¬

or, ellipsoid to oblong, 4-4.5 mm long, 1.7-3 mm in diameter, apically conical, glabrous, the apex with small,

ridged collar, but lacking crown, with axile placentation, the ovules numerous, borne on small placenta that

only slightly extends into locule; style 12-18.5 mm long, distally curved, white to rose, glabrous; stigma punc¬

tate, minutely papillose, 0.15-0.2 mm wide. Berries 6-10 mm long, 5.5-10 mm in diameter, globose to ellip¬

soid, purple-black, with scattered, stellate hairs, the hypanthium constricted, 1.5-2.7 mm wide at narrowest

point and 2.5-3.3 mm wide at torus, but appearing strongly constricted due to flaring calyx tube. Seeds 1-2

mm long, angular-obovoid, with ± rectangular raphe, the testa smooth to ± roughened due to bulging cells, but

the raphe smooth; appendage absent.

Chromosome number. —2n = 34 (Wurdack & Solt 1980).

Phenology .—Recorded in flower in every month except April, October, and December, and likely bloom¬

ing year around.

Judd et al., Revision of Miconia sect. Miconiastrum

483

Distribution and habitat.—Miconia angustiflora is endemic to Jamaica, occurring nearly throughout the

island (Fig. 8), usually in dry to moist thickets or forests over limestone, from 90-900 m.

Additional specimens examined. JAMAICA. Clarendon Parish: between Lluidas Vale and Croft Hill, 520 m, 16 Jul 1962, Fosberg 42715

(US); Peckham woods, 2500 ft, 6 Jul 1911, Harris 10961 (NY, US); ibid., 27 Sep 1912, Harris 11176 (F, NY, US). Hanover Parish: Campbellton

Hall, 300-400 ft, 22 Dec 1960, Adams 8649 (MO); Green Island and vicinity, Fish River Mountains, 13-15 Mar 1908, N.L. Britton & Hollick

2165 (NY); Hopewell, 500 ft, 26 Jun 1975, Clarkson & Kress 75-123 (DUKE); Askenish, Dolphin Head, 29 Jun 1975, Clarkson et al. 75-142

(DUKE); NE of Dolphin Head, 20 Mar 1908, Harris 10316 (F, NY, US); Dolphin Head, SW of Askenish, ca. 7 mi S of Lucea, 270-300 m, 12 May

1987, Judd 5302 (F, FLAS, GH, IJ, MO); Dolphin Head, E side, 1200 ft, 22 Aug 1952, Proctor 7184 (IJ); lower slopes between Askenish and

Dolphin Head, 900-1250 ft, 28 Jun 2001, Proctor 51899 (IJ); Dolphin Head, on the top, 535 m, 9 Jul 1986, Skean 1841 (IJ, NY); NE slopes of

Dolphin Head, 330 m, 9 Jul 1986, Skean 1854 (IJ, MO, US); Quashiba Mountain, 1 mi W of Georges Plain, 1100-1300 ft, 29 Jul 1954, Webster

& Wilson 5092 (IJ, MICH, S, US). Manchester Parish: Lincoln, 3-7 Sep 1908, N.L. Britton 3108 (NY); Marshall’s Pen, 21-23 Sep 1908, N.L.

Britton 3704 (NY); vicinity of Mandeville, 15-26 Feb 1910, Brown 266 (NY, US); Limestone Hill at Banana Ground, 2900 ft, 14 Jul 1963,

Crosby et al. 656 (DUKE, F, MICH, MO, MSC, NY); near Troy, 2000 ft, 30 Jun 1904, Harris 8744 (F, FLAS, NY, S, US); 0.5 mi NW of Christia¬

na, 3000 ft, 3 Jul 1955, Howard & Proctor 14341 (A, IJ); 2.25 mi NW of Mandeville, Marshall’s Pen, 18°5'N, 77°30'W, 700 m, 1 Jun 1983,

Landrum 4745 (NY); Mandeville, Marshall’s Pen, 750 m, 19 Nov 2000, Parker & Proctor 3447 (IJ); 0.5 mi NW of Christiana, 3000 ft, 22 Oct

1955, Proctor 11040 (IJ, NY, US); Marshall’s Pen, 2.25 mi NW of Mandeville, 2100-2300 ft, 28 Jul 1968, Proctor 28893 (IJ, NY, US); Cross Keys,

2000-3000 ft, Aug 1954, Robertson 1249 (NY). Portland Parish: slopes above Uncommon Hill, above Fruitful Vale, 1000-2250 ft, 17 Aug

1962, Proctor22615 (IJ, MICH, NY). St. Andrew Parish: Red Hills, 1500 ft, 7July 1960, Adams 7525 (DUKE); Mona Hill, vicinity of Kingston,

11-12 Sep 1906, N.L. Britton 366 (NY); Cooper’s Hill, Red Hills, NW of Kingston, 2300 ft, 7 Aug 1963, Crosby & Anderson 1117 (DUKE, F,

MICH, MSC, NY); Stony Hill, Nov 1898, Fawcett & Harris 7462 (A, GH, NY, S, US); Halls Delight, 334 m, 15 Jun 1895, Harris 5512 (F, US);

Constant Spring to Bardowie, 800 ft, 27Jul 1915, Harris 12089 (F, GH, MO, NY, S, US); between Mavis Bank and Gordon Town, between mile

marker 13 and 14, 760 m, 18 May 1987 Judd 5363 (IJ); Hermitage Dam and vicinity, 500 m, 3 Jun 1926, M axon 8787 (GH, S, US); Stony Hill,

19 Jul 1949, Newill s.n. (IJ); ibid., 1500 ft, 14 Jul 1952, Newill s.n. (IJ). St. Ann Parish: 1 mi S of Crescent Park, vicinity of Lydford Post Office,

1750 ft, 20-31 Dec 1953, Howard & Proctor 13548 (A, IJ); Hopewell, 2 Feb 1938, Hunnewell 15328 (GH, NY); near Bamboo, 11 Feb 1948, Hun-

newell 18834 (NY); Mount Diablo region, lower slopes of Hollymount, 1 mi S of Faiths Pen, 500-575 m, 28 May 1987, Judd 5512 (F, FLAS, IJ,

NY); vicinity of Lime Hall, 1000-1250 ft, 25 Jun 1954, Lewis s.n. (IJ); vicinity of Mosely Hall Cave, nearBlackstonedge, 2000 ft, 24 Aug 1952,

Proctor 7206 (HAC, IJ, NY); rd from Jamaica A-l to Hollymount on Mt. Diablo, 1700 ft, 6 Jul 1967, R.E. & S. Weaver 902 (DUKE, GH). St.

Catherine Parish: on A-l 0.7 mi S of St. Ann/St. Catherine boundary, lower slopes of Mt. Diablo, 18°12.392'N, 77°5.675'W, 526-530 mjudd

8309 (FLAS, IJ, NY); 0.6 mi SE of Guys Hill, 1600 ft, 3 May 1952. Proctor 6653 (IJ, US); 3 mi W of Lluidas Vale, 1750 ft, 11 Jun 1965, Proctor

26441 (IJ); near Sligoville, 9 Aug 1947, van der Porten s.n. (IJ); 1 mi E of Top Hill, 550 m, 10 Oct 1969, Wurdack & Solt 2616 (NY, US). St. Eliza¬

beth Parish: Santa Cruz Mountains, 12 Sep 1907, N.L. Britton 1296 (NY); Munro College, 2600 ft, Aug 1944, Snee s.n. (IJ). St. Mary Parish:

Cabarita Island, off Port Maria, 150 ft, 29 Dec 1952, Proctor 7552 (IJ). St. Thomas Parish: Serge Island, 15-19 Sep 1908, N.L. Britton 3655

(NY). Trelawny Parish: from Burnt Hill to Ramgoat Cave, 18°25'N, 77°33'W, 500 m, Acevedo-Rdgz. 9569 (US); Cockpit Country, near Bar¬

becue Bottom, 14 Aug 1963, Crosby & Anderson 1209 (DUKE, F, GH, MICH, MSC, NY); 2-3 mi N by road of Burnt Hill crossroads, 1300 ft,

16 Aug 1965, Hespenheide 1289 (DUKE, FLAS, GH, MICH, MO, MSC, NY, US); 0.6-1 mi N of Spring Garden, 1750-2000 ft, 1 Nov 1975,

Proctor 35429 (IJ, MO).

Miconia angustiflora is variable in the development of mite domatia; some individuals have mite domatia on the

abaxial leaf surface, while others lack them. The domatia, when present, are usually poorly developed. The

species often has been confused with T. bicolor (Goldenberg et al. 2013), although the form of the hairs (i.e.,

stellate hairs vs. stellate-peltate scales) easily differentiate the two taxa. It is actually much more similar to M.

barbata and M. maestrensis, two species of the “Oriente” region of Cuba (see key for differentiating characters),

although we note that the latter species also have been confused with T. bicolor.

The species has almost always been known as Tetrazygia pollens (see, for example, Proctor 1972) although

this name actually is a synonym of T. bicolor (see Michelangeli & Becquer 2012, and nomenclatural discussion

under Miconia bicolor, above). The misapplication of the name Tetrazygia pollens, originally described from

Hispaniolan material, came about as a result of the broad circumscription of Tetrazygia pollens in Cogniaux

(1891), which included plants from Jamaica, Hispaniola, and eastern Cuba; the name T. pollens was then erro¬

neously applied to the Jamaican populations when the circumscription of this species was restricted.

7. Miconia cajalbanensis Judd, Becquer, & Majure, nom. nov. (Fig. 6C). Fetrazygia coriacea Urb., Repert. Spec. Nov.

Regni Veg. 22:225. 1926, non Miconia coriacea DC., Prodr. 3:189. 1828. Type: CUBA [Prov. Pinar del Rio]: without definite locality,

1860-1864, fl, Wright 1222 p.p. (lectotype, here designated, because holotype at B destroyed: S!; isolectotypes: CAS on-line image

#0004793!, HAC!, NY!, S!).

Evergreen shrub or tree to 5 m. Young stems terete to slightly quadrangular, the indumentum of dense,

484

Journal of the Botanical Research Institute of Texas 8(2)

Fig. 8. Distribution of Miconia angustiflora in Jamaica.

ferrugineous, stellate hairs, without elongate, multicellular, non-glandular hairs, internodes 0.6-3.3 cm long,

lacking longitudinal ridges, nodal line absent. Leaves isophyllous or nearly so; petiole 0.7-2.7 cm long, the in¬

dumentum of dense, ferrugineous, stellate hairs; the blade 3.4-10 x 1.3-3.5 cm, ovate to elliptic or oblong, not

falcate, coriaceous, the apex acuminate, the base obtuse to rounded (or very slightly cordate), the margin plane

to revolute, entire; secondary veins 2 pairs, one pair conspicuous and one pair inconspicuous, acrodromous,

basal, the innermost pair joining midvein at base to 1 mm above the leaf base, the conspicuous secondary veins

placed 1.8-5.5 mm in from margin, the inconspicuous secondary veins intramarginal to 1 mm in from margin,

tertiary veins percurrent, oriented subperpendicular to midvein, 1.7-6 mm apart, connected by 1 quaternary

vein, or not connected by quaternary veins, and these reticulate, the higher order veins reticulate, the midvein

and major secondary veins impressed, tertiary veins very slightly impressed to flat, remaining veins flat on

adaxial surface; the midvein strongly raised, the major secondary veins moderately to slightly raised, the minor

secondary veins and tertiary veins flat to very slightly raised, and the higher order veins flat on abaxial surface;

adaxial surface appearing slightly wrinkled after drying, with scattered druse crystals, and drying darker than

the abaxial surface, the indumentum initially of dense, ferrugineous, stellate hairs, but very quickly glabres-

cent, although a few often persisting on lamina, and numerous such hairs often persisting on primary and

secondary veins, especially the proximal portions thereof; abaxial surface pale green but appearing ferrugine¬

ous due to the indumentum, the surface distinctly bullate, with moderate to dense, ferrugineous, stellate hairs,

0.07-0.22 mm across, the veins with similar hairs, such hairs on lamina and veins persistent; mite domatia

absent. Inflorescences terminal, paniculate cymes, 4-15.3 cm long, 4-8 cm across, with 2-5 major branch

pairs, the peduncle 1.8-5 cm long, the ultimate axes (pseudopedicels) 1.7-5.5 mm long, and the numerous

flowers well separated from each other; bracts 0.5-1.5 x 0.15-0.3 mm, and probably also larger, narrowly tri¬

angular, with acute apex, very quickly deciduous; bracteoles 0.3-0.8 x 0.1-0.2 mm, narrowly triangular, with

acute apex, quickly deciduous. Flowers 5- or occasionally 6-merous, zygomorphic (due to positioning of the

stamens), with pedicel 1-3 mm long. Hypanthium 6.5-7.5 mm long, terete and not or only slightly constricted

above ovary, the free portion 3.5-4.5 mm long, 2.5-4 mm wide at the torus, the outer surface with moderate to

dense stellate hairs, the internal surface smooth to very slightly longitudinally ridged, glabrous. Calyx lobes 5

(6), separate in bud, the tube 2-3.3 mm long, not tearing between lobes, with moderate stellate hairs abaxially,

and moderate to dense branched to stellate hairs adaxially, the lobes 0.2-0.7 x 2-4 mm, broadly triangular to

nearly obsolete, the apex rounded, with hairs similar to those of the tube, green, often red- tinged; calyx teeth

present, 0.2-0.5 mm long, a rounded to conical bump near apex of each lobe or merely a raised line along

Judd et al., Revision of Miconia sect. Miconiastrum

485

midrib of lobe, green to red, with rounded to acute apex, and hairs similar to those of calyx. Petals 5 (6), 9-11

x 4.5-6.5 mm, asymmetrically obovate, spreading, white, often pink- tinged, glabrous but both surfaces

densely papillose-granulate, the apex ± rounded, the base slightly narrowed to a broad attachment, the margin

entire. Stamens 10 (12), isomorphic, ± geniculate near filament apex; staminal filament 7-7.5 mm long, gla¬

brous, pale yellow, often pink-tinged, anther thecae 6.5-8.3 x 0.8-1.3 mm, subulate, straight to slightly in¬

curved, opening by a small dorso-apical pore, yellow, often pink-tinged, the connective not prolonged below

the thecae, glabrous, but the filament emerging from slight depression in sterile, minutely dorso-lobed anther

base. Ovary 3-locular, ca. 3/4 inferior, ellipsoid to oblong, 4.3-5 mm long, 1.7-3 mm in diameter, apically

conical or cylindric-conical, glabrous, the apex with a small, ridged collar, but lacking a crown, with axile

placentation, the ovules numerous, borne on a small placenta that only slightly extends into locule; style 15-17

mm long, distally curved, white to pink, glabrous; stigma punctate, minutely papillose, 0.15-0.2 mm wide.

Berries 5-9.5 mm long, 5.5-8 mm in diameter, globose to ellipsoid, purple-black, with scattered stellate hairs,

the hypanthium constricted, 2-2.3 mm wide at narrowest point and 2.5-3 mm wide at torus, but appearing

strongly constricted due to flaring calyx tube. Seeds 1.3-2 mm long, angular-obovoid, with ± rectangular ra¬

phe, the testa distinctly roughened due to bulging cells, except at distal end where cells ± smooth, however,

raphe smooth; appendage absent.

Phenology. —Collected in flower from April to July.

Distribution and habitat.—Miconia cajalbanensis is endemic to the Cajalbana region, Prov. Pinar del Rio,

Cuba (Fig. 5), where it occurs in xeromorphic scrub and pinelands on serpentine soil from 150-470 m.

Additional specimens examined: CUBA. Prov. Pinar del Rio: La Palma, La Cajalbana, 26 Jan 1950, Acuna H-ROIG-9043 (HAC); La Cajal¬

bana, 10 Jun 1950, Acuna et al. SV-16187 (HAC); Cajalbana, La Palma, 1 Oct 1949, Bro. Alain & Acuna 1119 (HAC, NY); Cuabales de Cajal¬

bana, La Palma, 3 Dec 1949, Bro. Alain & Acuna 1199 (HAC); ibid., Acuna & Bro. Alain SV-15699 (HAC); W of Loma Cajalbana, La Palma, 3

Apr 1954, Bro. Alain 3899 (GH, HAC, NY); W of Cajalbana, 7 Dec 1955, Bro. Alain 4495 (GH, HAC, US); Bahia Honda, Sierra de Cajalbana,

subida al Tecnologico Invasion de Occidente, 300 m, 15 Jul 1989, Alvarez de Zayas & Pujadas HFC-70172 (HAJB); ibid., Alvarez de Zayas &

Pujadas HFC-70175 (HAJB); ibid., Alvarez de Zayas & Pujadas HFC-70176 (HAJB p.p.); ibid., Alvarez de Zayas & Pujadas HFC-70177 (HAJB);

Cajalbana, Oct 1975, Armenteros s.n. (HAJB); Mpio. La Palma, Cajalbana, carretera a Mil Cumbres, sendero ecoturistico, 7 May 2004, Becquer

& Abbott HFC-82423 (FLAS, HAJB); La Palma, Loma de Cajalbana, falda O, Apr 1967, Bisse & Rojas HFC-1942 (HAJB); Bahia Honda, parte

oriental de la Loma de Cajalbana, Jul 1968, Bisse HFC-9567 (HAJB); La Palma, pinares de Cajalbana, Oct 1975, Bisse et al. HFC-28608 (HAJB);

La Palma, Cajalbana, 28 Dec 1974, Bisse et al. HFC-29316 (HAJB); Consolacion del Norte, falda S de la loma peluda (Preluda) de Cajalbana, 15

Oct 1976, Bisse et al. HFC-32604 (HAC, HAJB); La Palma, Altiplano de Cajalbana, cerca d el Tecnologico forestal, 350 m, Dec 1980, Bisse et al.

HFC-45507 (HAJB); Loma Cajalbana, pinar de la cumbre, 450-470 m, 5 Jul 1970, Borhidi & Vdsquez 8000 SV-33414 (HAC); ibid., Borhidi &

VdsquezSV-33417 (HAC); La Palma, ladera S de Cajalbana, entre Sagua y El Sitio, 22°45'N, 83°30'W, 150 m, 16 Jul 1996, Cedeno 1234 (NY);

Loma de Cajalbana in “Cuabal”, 10 Mar 1920, Ekman 10481 (S); Cajalbana, 200-300 m, Nov 1975, Garcia & Sdnchez s.n. (HAJB); La Palma,

Arroyo de Morto, Sierra de Cajalbana, ladera S, 22°46'NH, 83°26'W, 14 Sep 1999, Kuba-Exkursion Uni Frankfurt/M 1999 no. 13 (FR n.v., S);

ibid., 17 Sep 1999, Kuba-Exkursion Uni Frankfurt/M 1999 no. 112 (FR n.v., S); Cajalbana, ladera al NE del Tecnologico Forestal, 400 m, 21 Jan

1981, Martinez C. SV-36145 (HAC); Monte La Cajalbana, 4 Feb 1956, Morton 9820 (US); Cajalbana, 1 Nov 1975, Plasencia s.n. (HAJB); pinares

sobre serpentine de Cajalbana, en la ladera S, 5 May 1988, Urquiola et al. HPR-4501 (HAJB); La Cajalbana, Jan 1967, Yero M. 677 (HAC); ibid.,

Jul 1965, Yero M. 1093 (HAC).

Miconia cajalbanensis is phenetically distinctive because of its fairly small stellate hairs and distinctly bullate

abaxial leaf epidermis (see key). Its seeds are also noteworthy, having bulging testa cells, in contrast to most

species of the M. bicolor complex, which have more or less smooth testa cells. Despite these distinctive charac¬

ters, it has frequently been confused with the more widespread and common M. bicolor, a species with an ab¬

axial leaf indumentum of stellate-peltate scales.

The new name M. cajalbanensis, required since the epithet “coriacea” is occupied in Miconia, reflects the

fact that this species is endemic to the vicinity of Cajalbana, Cuba. This is a region of exceptional endemism,

with ca. 40 endemic species and several endemic genera (Borhidi and Muniz 1985; Berazaln 1987; Borhidi

1991). Miconia cajalbanensis is considered to be critically endangered in Cuba (Berazaln 2005).

8. Miconia cristalensis (Borhidi) Judd, Becquer, & Majure, comb. nov. Fetrazygia cristalensis Borhidi, Acta Bot. Acad.

Sci. Hung. 23:37.1977 [1978], Type: CUBA. [Prov. Santiago de Cuba:] entre Los Mulos y Corea, 2100-2200 ft, Sierra de Cristal, 27-28

Aug 1959, M. LopezFigueiras UO-279 (holotype: HAC!; isotypes: HAC!, HAJB!, NY!).

486

Journal of the Botanical Research Institute of Texas 8(2)

Evergreen shrub to small tree up to 4 m. Young stems terete to quadrangular, the indumentum of dense, fer-

rugineous, globular-stellate to dendritic hairs, without elongate, multicellular, non-glandular hairs, internodes

0.9-3.5 cm long, lacking longitudinal ridges, nodal line present, faint to prominent. Leaves isophyllous or

nearly so; petiole 1-2.6 cm long, the indumentum of dense, ferrugineous, globular-stellate hairs; the blade

4.6- 14 x 1.8-4.4 cm, ovate to elliptic, coriaceous, not falcate to slightly falcate, the apex acuminate, the base

obtuse to rounded, the margin plane to revolute, entire to slightly undulate; secondary veins two pairs, one pair

conspicuous and one pair inconspicuous, acrodromous, basal, the innermost pair joining midvein at base to 2

mm above the leaf base, the conspicuous secondary veins placed 2-7.5 mm in from margin, the inconspicuous

secondary veins intramarginal to 1.5 mm in from margin, tertiary veins percurrent, oriented subperpendicu¬

lar to midvein, 1.5-7 mm apart, connected by 1 or 2 quaternary veins, or quaternary veins reticulate, not con¬

necting tertiary veins, the higher order veins reticulate, the midvein and major secondary veins slightly to

moderately impressed, tertiary veins slightly impressed to flat, remaining veins slightly impressed to flat on

adaxial surface; the midvein strongly raised, the major secondary veins strongly to moderately raised, the mi¬

nor secondary veins and tertiary veins slightly raised, and the higher order veins slightly raised to flat on ab-

axial surface; adaxial surface appearing slightly wrinkled after drying, with scattered druse crystals, and dry¬

ing darker than the abaxial surface, the indumentum initially of dense, brunnescent to ferrugineous, globular-

stellate hairs, but quickly glabrescent, although some hairs persisting proximally on midvein and major sec¬

ondary veins; abaxial surface pale green but appearing brunnescent to ferrugineous due to indumentum, the

epidermal surface ± smooth, with dense, brunnescent to ferrugineous, globular-stellate hairs (with arms

pointing upward to radiating outward), 0.16-0.28 mm across, the veins with similar hairs, such hairs on lam¬

ina and veins persistent, mite domatia absent. Inflorescences terminal, paniculate cymes, 3-5 cm long, 2-4

cm across, with 1-3 major branch pairs, the peduncle 1.5-3.3 cm long, the ultimate axes (pseudopedicels) 2-7

mm long, and numerous flowers well separated from each other; bracts not seen, presumably triangular, very

quickly deciduous; bracteoles not seen, presumably ± triangular, deciduous. Flowers 5-merous, zygomorphic

(due to positioning of the stamens), with pedicel 1-2 mm long. Hypanthium 6.5-8 mm long, terete and slight¬

ly to distinctly constricted above ovary, the free portion 2.5-4 mm long, 2.3-3 mm wide at the torus, the outer

surface with moderate to dense globular-stellate hairs, the internal surface very slightly longitudinally ridged,

glabrous. Calyx lobes 5, separate in bud, the tube 0.9-2 mm long at late anthesis (only old flowers seen), not

tearing between the lobes, with moderate to dense globular-stellate hairs adaxially and abaxially, the lobes

1.7- 3.5 x 1-3.5 mm, ± triangular, the apex acute to rounded, with hairs similar to those of the tube, green, often

red-tinged; calyx teeth present, 0.2-1 mm long, merely a ridge along lobe to a ridge with short apical bulge or

projection, green or reddish, with rounded apex, and hairs similar to those of calyx. Petals 5, 6-7.5 x 5-6 mm,

asymmetrically obovate, spreading, white, glabrous but both surfaces densely papillose-granulate, the apex

rounded, the base slightly narrowed to a broad attachment, the margin entire. Stamens 10, not seen. Ovary

3-locular, 2/3-4/5 inferior, apically conical, glabrous, the apex with a small, slightly ridged collar but lacking a

crown, with axile placentation, the ovules numerous, borne on a small placenta not extending or only slightly

extending into locule; style not seen. Berries 6-8 mm long, 5-6 mm in diameter, ellipsoid, purple-black, with

sparse to moderate globular-stellate hairs, the hypanthium strongly constricted, 2-2.5 mm wide at narrowest

point and 3-3.5 mm wide at torus. Seeds 1.7-2 mm long, angular-obovoid, with ± rectangular hilum, the testa

± smooth; appendage absent.

Phenology. —Only collected in flower in August.

Distribution and habitat.—Miconia cristalensis is endemic to the Sierra de Cristal (Prov. Santiago de Cuba),

Cuba (Fig. 7), occurring from 630-800 m in pine forests and thickets on serpentine or gabbro.

Additional specimens examined: CUBA. Prov. Santiago de Cuba: Sierra de Cristal, Mpio. Segundo Frente, ladera SE del Pico Cristal, subida

al firme desde la interseccion del camino del Oro a Batista y La Zanja, 700 m, 26 Apr 2004, Becquer HFC-82262 (FLAS, HAJB); Segundo Fr¬

ente, Sierra Cristal, camino de San Nicolias al Oro, cerca del cruce del rio Miguel, 3 Nov 2005, Becquer et al. HFC-83729 (HAJB); Mayan Ar¬

riba, Sierra de Micara, 800 m, Jun 1967, Bisse & Rojas HFC-4233 (HAJB).

Miconia cristalensis is very poorly collected, but apparently is most closely related to M. macstrensis (see key for

differentiating characters) and M. barbata , from which it differs in the lack of mite domatia and in its longer

Judd et al., Revision of Miconia sect. Miconiastrum

487

calyx lobes. The three species are likely reproductively isolated, as M. maestrensis occurs only in the Sierra

Maestra and M. barbata in the Sierra de Nipe and other areas in northern “Oriente” (Fig. 7); however both M.

maestrensis and M. barbata likely co-occur with the widespread M. bicolor. They are at least broadly sympatric

geographically, and more fieldwork is needed to investigate subtle ecological differences.

9. Miconia maestrensis Judd, Becquer, & Majure, nom. nov. (Fig. 6D). Miconiastrum lambertianum Bonpl. exNaudin,

Ann. Sci. Nat.,Bot.,ser. 3.15:341,1.17.1850, non Miconia lambertiana DC., Prodr. 3:185.1928. Type: CUBA [Prov. Santiago de Cuba]:

St. Jago de Cuba, Nima-nima, Aug 1844, Linden 2081 (lectotype, chosen by Martin & Cremers, J. Bot. Soc. Bot. France 37:40. 2007,

P on-line image #481686!; isolectotype, NY, 2 sheets!, P on-line image #481687!).

Evergreen shrub or tree to 5 m. Young stems terete to slightly quadrangular, the indumentum of dense, fer-

rugineous, globular-stellate to dendritic hairs, without elongate, multicellular, non-glandular hairs, eventually

± glabrescent, internodes 0.7-10.8 cm long, lacking longitudinal ridges, nodal line present, faint. Leaves iso-

phyllous or nearly so; petiole 1.8-3 cm long, the indumentum of dense, ferrugineous, globular-stellate to stel¬

late hairs; the blade 4.7-15.5 x 1.8-5.6 cm, ovate to elliptic, not falcate to slightly so, ± coriaceous, the apex

acuminate to attenuate, the base acute to obtuse or rounded, the margin plane to slightly revolute, entire; sec¬

ondary veins 2 pairs, one pair conspicuous and one pair inconspicuous, acrodromous, basal, the innermost

pair joining midvein at base to 3 mm above the leaf base, the conspicuous secondary veins placed 3-8 mm in

from margin, the inconspicuous secondary veins intramarginal to 1.3 mm in from margin, tertiary veins per-

current, oriented subperpendicular to midvein, 1.7-10 mm apart, connected by 1 or 2 quaternary veins, or not

connected by such veins, and quaternary veins ± reticulate, the higher order veins reticulate, the midvein and

major secondary veins slightly impressed to flat, tertiary veins very slightly impressed to flat, remaining veins

flat on adaxial surface; the midvein strongly raised, the major secondary veins moderately to slightly raised, the

minor secondary veins and tertiary veins flat to very slightly raised, and the higher order veins flat on abaxial

surface; adaxial surface appearing slightly wrinkled after drying, with scattered druse crystals, and drying

darker than the abaxial surface, the indumentum initially of ± dense, stellate hairs, but very quickly glabres¬

cent, although a few hairs often persisting on proximal portion of midvein and major secondary veins; abaxial

surface pale green, but appearing ferrugineous to pale ferrugineous due to dense indumentum, the epidermal

surface ± smooth, with dense, ferrugineous to pale ferrugineous, stellate hairs (with arms pointing upward to

radiating outward), 0.13-0.24 mm across, the veins with similar hairs, such hairs on lamina and veins persis¬

tent; mite domatia absent. Inflorescences terminal, paniculate cymes, 3.5-13 cm long, 2.5-6 cm across, with

1-4 major branch pairs, the peduncle 0.8-4 cm long, the ultimate axes (pseudopedicels) 1.5-5.5 mm long, and

the numerous flowers well separated from each other; bracts not seen, but presumably triangular, very quickly

deciduous; bracteoles ca. 0.4 x ca. 0.15 mm, narrowly triangular, with acute apex, deciduous. Flowers 5-mer-

ous, zygomorphic (due to positioning of the stamens), with pedicel 1-3 mm long. Hypanthium 6-6.5 mm long,

terete and slightly constricted above ovary, the free portion 2.4-2.8 mm long, 2-2.5 mm wide at the torus, the

outer surface with dense stellate hairs, the internal surface smooth to very slightly longitudinally ridged, gla¬

brous. Calyx lobes 5, separate in bud, the tube 0.7-1 mm long, not tearing between lobes, with ± dense stellate

hairs adaxially, and dense to moderate branched to stellate hairs adaxially, the lobes 0.3-0.6 x 2.5-3 mm,

broadly triangular, the apex obtuse to rounded, with hairs similar to those of the tube, green to red-tinged;

calyx teeth present, 0.1 mm long, merely a small bump near apex of lobe, green, with rounded apex, and hairs

similar to those of calyx. Petals 5, 6.3-7 x 3.4-5 mm, asymmetrically obovate, spreading, white, glabrous but

both surfaces densely papillose-granulate, the apex rounded, the base slightly narrowed to a broad attachment,

the margin entire. Stamens 10, isomorphic, ± geniculate near filament apex; staminal filament 5.5-7 mm long,

glabrous, color unknown, anther thecae 4.8-6 x 0.7-1 mm, subulate, straight to slightly incurved, opening by

a small dorso-apical pore, yellow, the connective not prolonged below the thecae, glabrous, but the filament

emerging from slight depression in sterile, minutely dorso-lobed anther base. Ovary 3-locular, 2/3-4/5 inferi¬

or, apically conical, glabrous, the apex with a small, slightly ridged collar but lacking a crown, with axile pla-

centation, the ovules numerous, borne on a small placenta not or only slightly extending locule; style 12-16

mm long, distally curved, white, glabrous; stigma punctate, minutely papillose, 0.15-0.2 mm wide. Berries

488

Journal of the Botanical Research Institute of Texas 8(2)

[nearly mature] 5.5-10 mm long, 4.5-6 mm in diameter, globose to ellipsoid, green [but presumably turning

purple-black], with scattered to moderate, stellate hairs, the hypanthium strongly constricted, 1.5-2.3 mm

wide at narrowest point and 2.3-2.7 mm wide at torus, but appearing wider due to flaring calyx tube. Seeds

1.5-2 mm long, angular-obovoid, with ± rectangular hilum, the testa ± smooth; appendage absent.

Phenology. —Only known to flower in August, but very poorly collected, and surely with a broad flower¬

ing period, as mature fruits have also been collected in August.

Distribution and habitat.—Miconia maestrensis occurs in Cuba, in the Sierra Maestra and associated lower

elevation habitats, in southern “Oriente” (Prov. Granma, and Santiago de Cuba), in moist forests, on limestone,

near rivers or arroyos, at low elevations to 935 m (Fig. 7).

Additional specimens examined: CUBA. Prov. Granma: Sierra Maestra, Mun. Buey Arriba, Barrio Nuevo a Pata la Mesa, cerca 4 km al SE de

El Manguito, 13 May 1988, Alvarez deZayas et al. HFC-64310 (B, HAJB); Buey Arriba, Guama, Barrio Nuevo a Peladero, 24 May 1988, Alvarez

deZayas et al. HFC-65244 (HAJB); Sierra Maestra, Rio Yara, 6Jul 1922, Ekman 14188 (S); Sierra Maestra, on the edge of Arroyo Corrojo, near

Nagua, 20 Aug 1922, Ekman 14920 (GH, S); Peninsula de Cabo Cruz, between Rio Nuevo and the coast, 16 Jan 1923, Ekman 16150 (GH, HAC,

S); Bartolome Maso, Parque Nacional Turquino, sendero Alto del Naranjo-Pico Turquino, entre Alto del Naranjo y La Platica, 20 o 0'53"N,

76°53 , 44.5"W, 935 m, 8 Nov 2013, M ichelangeli et al. 2209 (HAJB, NY). Prov. Santiago de Cuba: St. Jago de Cuba, Nima-nima, 1834-1844,

Linden 2091 (NY, P on-line image).

Miconia maestrensis, although early described, has long been known as either Tetrazygia pollens (see Cogniaux

1891) or T. bicolor (e.g., identifications on specimens such as Ekman 14188 or Ekman 14920). This species is

distinguished from T. bicolor by its indumentum of stellate hairs (not stellate-peltate scales, see key), and is

phenetically most similar to M. cristalensis. It differs from M. cristalensis by the characters indicated in the key

(note especially its much shorter calyx lobes). Miconia maestrensis is easily distinguished from M. barbata by

the lack of mite domatia. It appears to be reproductively isolated from both M. cristalensis and M. barbata, since

they occur in northern “Oriente” (Fig.7). Miconia maestrensis has been very poorly collected, and additional

collections would improve our understanding of its pattern of variation and geographical distribution. The

specimen Alvarez deZayas et al. HFC 64310 is tentatively considered to represent this species; only a photo has

been seen by the first author, and it appears to be more densely pubescent than other material examined. Per¬

haps the variation exhibited among the specimens here considered within M. maestrensis actually represents

that of more than one species, but a broad circumscription is here advocated due to the current paucity of

herbarium material.

An epithet is unavailable in Miconia since Miconiastrum lambertianum cannot be transferred. We have

therefore chosen the new name Miconia maestrensis, highlighting the distinctive geographical range of this

species.

ACKNOWLEDGMENTS

We thank Norris Williams, Keeper of the University of Florida Herbarium (FLAS), Kent Perkins, Collections

Manager, and Gretchen Ionta for assistance in processing specimens. We also thank the curators and staff of

the herbaria (A, F, FTG, GH, JBSD, MICH, MO, NY, S, and US) who generously provided specimens on loan.

Milclades Mejia, Thomas A. Zanoni, Daisy Castillo, Alain H. Liogier, Teodoro Clase, Ricardo Garcia, Francisco

Jimenez, and Brlgido Peguero assisted in several visits of the first and third authors to the herbarium (JBSD) Q f

the Jardin Botanico Nacional, Dominican Republic. Keron C. St. E. Campbell graciously assisted in the visit of

the first author to the herbarium (IJ) of the Natural History Museum of Jamaica, Institute of Jamaica. Bret Jestro

assisted during the visit of the first author to the herbarium (FTG) of the Fairchild Tropical Botanic Garden;

Anita Cholewa assisted during the visit to the University of Minnesota herbarium (MIN) and Alan Prather

during the visit to the Michigan State University herbarium (MSC). We thank the administrators of the Her-

bario Rafael M. Moscoso (UCMM) at the Pontihcia Universidad Catolica Madre y Maestra, Santiago, Republica

Dominicana, the Herbario de la Academia de Ciencias (HAC), La Habana, Cuba, the Herbario del Jardin Bo¬

tanico Nacional (HAJB), La Habana, Cuba, and the herbarium of Instituto Superior Pedagogico de Pinar del Rio

(HPPR) for their assistance. Finally, we thank Bruce F. Hansen for making available digital images of the hold¬

ings of Miconia bicolor at the herbarium (USF) of the University of South Florida. Numerous individuals helped

Judd et al., Revision of Miconia sect. Miconiastrum

489

in field work, most notablyJ. Richard Abbott, Reed Beaman, Gretchen M. Ionta, John Kress, and Tim McDow¬

ell. We are grateful that Richard Abbott has allowed use of his beautiful photos of Miconia bicolor (Fig. 3A, B),

M. cajalbanensis (Fig. 6C), M. delicatula (Fig. 1A), and M. guajaibonensis (Fig. 3C, D), and to Nicolas Garcia for

his photo of fruiting M. bicolor (Fig. ID). One anonymyous reviewer and Lynn Bohs (U) provided helpful com¬

ments on an earlier version of the manuscript. This research was supported, in part, by National Science Foun¬

dation Grant BSR-0818399, and we thank Fabian A. Michelangeli for his administrative and professional as¬

sistance in connection with this grant-supported project.

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492

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BOOK REVIEW

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., color photos

and illustrations, 8.25" x 10.25".

This comprehensive study of the ethnobotany of Ireland provides an in-depth, interesting, and enjoyable dis¬

cussion of the historic and current uses, as well as the cultural significance, of native and introduced plants

that are found throughout the Irish countryside.

At 750 pages, this volume contains information on more than 1000 species of plants, of which more than

600 have known ethnobotanical uses. The first 150 pages of the book provide the reader with historical and

cultural context as well as general information related to the plants. The remainder of the book is devoted to an

alphabetical listing, narrative descriptions, and common uses of flowering plants, grasses and sedges, trees

and shrubs, ferns and fern allies, algae, and miscellaneous plants.

The narrative on each species includes facts, lore, legend, and traditions associated with that plant. There

are 17 categories of ethnobotanical uses, all represented by icons. The icons are displayed along with the nar¬

rative descriptions for quick reference. Many of the descriptions are also accompanied by a small color photo

of the plant. Ethnobotanical uses such as plants used for food, beverages, medicines, potions, cosmetics, tex¬

tiles, construction materials, arts and crafts, dyes, poisons, cultural and religious symbols, currency, and many

other uses are noted as appropriate for each species.

The first appendix is a listing of plant names that are part of local Irish place names. Appendix II is a list¬

ing of plants with the scientific plant name, the Irish common name, and the Gaelic name (with pronunciation

guides!) for each plant. These listings are followed by a section which describes the exhibitions and traditional

folklife locales where plants and their varied uses can be seen and studied. Finally, there is a glossary which is

useful in understanding the botanical and taxonomic terminology that is commonly used in Ireland.

This unique book would be enjoyable and useful to botanists as well as casual readers who have a special

interest in ethnobotany, the natural resources of Ireland, or the social and cultural history of Ireland .—Dan

Caudle, Independent Grassland Consultant, Weatherford, Texas, U.S.A.

J. Bot. Res. Inst. Texas 8(2): 492.2014

CALATHEA RUBRIBRACTEATA, A NEW ENDEMIC SPECIES

OF MARANTACEAE FROM COLOMBIA

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 rubribracteata H. Kenn. endemic to Dept. Valle de Cauca, Colombia, is described as new. It is similar in aspect to the Ecuadorian

species C. neillii H. Kenn., C.jredii H. Kenn., C. cofaniorum H. Kenn. and C. shishicoensis H. Kenn. and to the Colombian species C. trianae

T.B. Sm. & Idrobo and C. timothei H. Kenn. with several basal leaves and a single cauline leaf or bladeless sheath subtending an inflorescence

of bright red or rose-pink bracts. Calathea rubribracteata differs from C. trianae, C. neillii, and C.jredii in the spirally arranged vs. distichous

bracts and relatively wider leaf blades (length:width ratios 3.03-3.58:1 vs. 3.55-7.45), from C. timothei by the fewer bracts (6-11 vs. 40-ca.

100) and the usually 3 vs. 7 minor veins between the major veins and from C. shishicoensis by the relatively narrower leaf blades

(length:width ratios 3.03-3.58:1 vs. 1.33-1.75:1) and yellow vs. pale pink flowers.

RESUMEN

Calathea rubribracteata H. Kenn. endemica del Depto. de Valle de Cauca, Colombia, es descrita como nueva especie. Ella es similar en

aspecto a las especies C. neillii H. Kenn., C.jredii H. Kenn., C. cojaniorum H. Kenn. y C. shishicoensis H. Kenn. de Ecuador y C. trianae L.B.

Sm. & Idrobo y C. timothei H. Kenn. de Colombia que tienen varias hojas basales y solo una hoja caulinar o una vaina sin lamina foliar que

subtiende una inflorescencia con bracteas rojas o rosadas brillantes. Calathea rubribracteata se distingue de C. trianae, C. neillii y C.jredii por

sus inflorescencias con bracteas espiraladas vs. disticas y las hojas relativamente mas anchas (la relacion largo/ancho es 3.03-3.58:1 vs.

3.55-7.45), de C. timothei por las pocas bracteas (6-11 vs. 40-ca. 100) y con la lamina foliar que lleva generalmente 3 vs. 7 venas menores

entre las venas mayores, y de C. shishicoensis por las hojas relativamente mas estrechas (la relacion largo/ancho es 3.03-3.58:1 vs. 1.33-

1.75:1) y las flores amarillas vs. rosado palidas.

Both Colombia and Ecuador have a rich diversity of Marantaceae. Collecting in the montane forests in both

countries have yielded some interesting new, red-bracted species of Calathea. There are currently 69 species of

Calathea recognized for Ecuador (Kennedy 2013, 2014), but so far only 49 described species are recorded for

Colombia. With such diversity and additional collections and herbarium work, the number should soon equal

that for Ecuador as at least ten more species are known but as yet undescribed, including the one herein. As

with C.trianae and C. timothei from Colombia, and C. neilli, C.jredii, C. cofaniorum and C. shishicoensis from

Ecuador, this new species, with its attractive bright red bracts, would make an excellent ornamental species.

TAXONOMIC TREATMENT

Calathea rubribracteata H. Kenn., sp. nov. (Fig. 1). Type: COLOMBIA. Valle del Cauca: Km 18 y Km 20 de la carretera de Cali

a Buenaventura entrando por la finca Zingara, cumbre de la cordillera occidental, selva climax andina, nublada y humeda con

abundante capote, relictus de las selva nublada y bastante intervenida antropogenicamente, 1500-2000 m, 03°28'N, 76°35'W, 28

Feb 1988, 1. Cabrera & H. van der Werjj15829 (holotype: MO 3765450).

Haec species a Calathea neillii H. Kenn., C.jredii H. Kenn. et C. cojaniorum H. Kenn. bracteis spiraliter dispositis (vs. distichis) atque lamina

foliari pro ratione latiore (longitudinis cum latitudine proportione 3.03-3.58 vs. 4.80-7.45), a C. trianae L.B. Sm. & Idrobo inflorescentia

turbinata (vs. rectangulari; longitudinis cum latitudine proportione ca. 1.2 vs. 6.00-7.33), a C. timothei H. Kenn. bracteis paucioribus (6 ad

11 vs. 40 ad ca. 100) atque venis minoribus plerumque 3 (vs. 7) inter majores, a C. shishicoensi H. Kenn. lamina foliari pro ratione angustiore

(longitudinis cum latitudine proportione 3.03-3.58 vs. 1.33-1.75) atque floribus flavis (vs. pallide roseis) differt.

Plants rhizomatous, caulescent, herbs, 35-60 cm high; stem green, internodes between basal leaves sericeous,

uppermost, elongate, portion glabrous; cataphylls appressed tomentose, more densely so at apex, margins and

J. Bot. Res. Inst. Texas 8(2): 493 - 496.2014

494

Journal of the Botanical Research Institute of Texas 8(2)

MISSOURI

BOTANICAL GARDEN

HERBARIUM

N® 3765450

cm ..duplicates at MO

SPECIMEN-DATE _

MARANTACEAE

Depto. VALLE: Km 18 y Km 20 de la carre-

Let a de Cali a Buenaventura entrando

1j tinea Z ingara . Cumbre de la cordill e¬

ra occidental. Preci pi taci on pluvial

pot ano. Re 1ictus de la selva nublada

y bastante intervenida antropoqinica-

mente.

1500-2000 m

PI ant a erect* que crece dentro dy la

selva cluuax andinat nublada > burned*

con abundant# capote. Ilojas coriaceaa y

bractea s rojas muy vistosust clocea

amar ill as. Ss muy bell* p*r* ornamental.

Isidore C*br«r* R. 2$ rebtero IMi

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MISSOURI BOTANICAL tiARPKN HSKRARIUN l«Oi

Fig. 1. Calathea rubribracteata H. Kenn. Holotype. (/. Cabrera &H. van der Werff 15829 MO).

Kennedy, Calathea rubribracteata, a new endemic Colombian species

495

base. Leaves 4-8 basal, none or 1 cauline, a cauline leaf or bladeless sheath borne atop a ca. 36 cm stem inter¬

node; leaf sheath not auriculate, green, wings lighter, tinged reddish, apical 1 cm subglabrous, appressed to-

mentose basally, the hairs more dense and longer toward the base, the hairs 0.25-1 mm, 13-17.5 cm; petiole

green, tomentose in a band adaxially in apical 3-4 cm, glabrous basally, bearing a shallow groove apically,

9-23.5 cm; pulvinus articulate, noticeably thicker than petiole, olive-green or tinged with red, tomentose in a

narrowband adaxially, the hairs 0.2-0.3 mm, the rest glabrous, 1-2.6 cm; leaf blade coriaceous, elliptic, apex

acuminate-attenuate, base obtuse to rounded, 18.2-23.5 x 5.7-7 cm (lenght:width ratios 3.03-3.58:1), gener¬

ally 3 minor veins between major veins, vein angle of divergence from midrib (measured at midpoint of blade)

34°-43°, 13-15 veins per 3 cm, 22-27 veinlets per 5 mm (measured at midpoint of each side of the blade), ad-

axial leaf surface deep green, glabrous except appressed tomentose on blade in apical 2 cm, more densely so

toward apex, midrib yellowish green, minutely tomentose, the hairs faint straw-colored, 0.25 mm; abaxial leaf

surface grey-green, glabrous except tuft of hairs on apicalmost 2-3 mm, midrib yellow-green, essentially gla¬

brous except for a few sparse colorless hairs along sides in basal half, the hairs to 0.2 mm (seen at 30x magnifi¬

cation). Inflorescence terminal, 1 or 2 per shoot, first borne terminally the second in the axil of the subtending

leaf or bladeless sheath, lax, turbinate, ca. 4 x 3.3 cm; peduncle red, glabrous, ca. 1 cm. Bracts 6-11, spirally

arranged, lowermost one elliptic, upper ones broadly elliptic to broadly obovate, apex obtuse, margin and apex

spreading, 2.2-2.5 x 1.6-2 cm, abaxial surface of bracts red, sericeous at very base, sparely pilose along basal

margin, the hairs 0.3 mm, adaxial surface red, glabrous, rachis sericeous; bicarinate prophyll membranous,

obovate, apex obtuse, red apically, sparsely scattered pilose on distal surface of the basal half of the carina, the

hairs 0.7-1 mm, ca. 2.1 x 0.65 cm, ca. 0.5 cm carina to carina; secondary bract membranous, narrowly elliptic,

apex obtuse, glabrous, 1.8-1.95 x 0.5-0.65 cm; bracteoles 2 per flower pair, membraneous, medial, one cari¬

nate, one channeled, narrowly elliptic, acute, glabrous, 1.55-1.7 x 0.2-0.3 cm. Flowers opening spontaneous¬

ly, but only partially open as the petals, though separated, do not reflex. Sepals narrowly oblong-elliptic, ob¬

tuse, yellow apically, cream below, glabrous, 14.5-17 x 2.5-3 mm. Corolla yellow, glabrous, tube 16-18 mm;

corolla lobes subequal, elliptic, obtuse, margins inrolled, 8.5-9 x 3.5-4.5 mm. Staminodes yellow; outer sta-

minode not found in material available; callose staminode totally callose, spathulate, apex rounded to shal¬

lowly retuse, ca. 8 x 5-5.5 mm; cucullate staminode 5-5.5 mm; stamen yellow with lateral petaloid appendage

to 1.5 mm wide, extending half way along the anther, anther 2 mm; style and stigma yellow, stigma blunted.

Ovary ca. 1.5 x 1 mm. Capsule unknown.

Distribution and habitat.—Calathea rubribracteata is endemic to Colombia, known from the type locality

in the Department of Valle de Cauca, in the Cordillera Occidental, and in cultivation in a nearby, private gar¬

den, at ca. Km 22 on the Cali to Buenaventura road. It occurs at 1500-2000 m elevation in a relictual climax

Andean cloud forest. The type was collected in flower in February.

Discussion.—Calathea rubribracteata shares the general habit of several basal leaves with an inflorescence

of bright red or rose-pink bracts borne above an elongate stem internode with C.fredii, C. neillii, C. cofaniorum,

and C. shishicoensis from Ecuador and C. trianae and C. timothei from Colombia. It differs from C. trianae, C.

neillii, C.fredii, and C. cofaniorum in the spirally arranged vs. distichous bracts and relatively wider leave blade

(length:width ratios 3.03-3.58:1 vs. 3.55-7.45), from C. timothei by the fewer bracts (6-11 vs. 40-ca. 100) and

the usually 3 vs. 7 minor veins between the major veins, and from C. shishicoensis by the relatively narrower

leaf blade (length:width ratios 3.03-3.58:1 vs. 1.33-1.75:1) and yellow vs. pale pink flowers. It is interesting to

note that C. timothei lacks the outer staminode and has a blunted stigma (Kennedy 1977). Whether C. rubri¬

bracteata in fact also lacks the outer staminode was not clear from the material at hand, but it appeared to have

a similar blunted stigma and also the non-reflexed petals. Many of these red to rose-pink bracted species are

found in the upper altitudes for Marantaceae distribution, 800-2100 m, in premontane to montane cloud forest

habitats.

Etymology. —The specific epithet, rubribracteata, refers to its bright red bracts.

496

Journal of the Botanical Research Institute of Texas 8(2)

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 covered the travel expenses for the trips to MO

and UCR. I thank Isidore Cabrera R. and Henk van de Werff for collecting C. rubribracteata. I am grateful to

Andrew Sanders for providing facilities at the UCR herbarium, to Teresa Salvado for providing accommoda¬

tions and transportation in Riverside and to James Solomon for the loan material from MO. I thank Carla Black

and Emilio Constantino for sharing their color photos of the plant. I thank Luz Stella Suarez S., Barney

Lipscomb, and anonymous reviewer for helpful suggestions and corrections. Roy Gereau provided the Latin

diagnosis.

REFERENCES

Kennedy, H. 1977. Unusual floral morphology in a high altitude Calathea (Marantaceae). Brenesia 12/13:1-9.

Kennedy, H. 2013. Two new endemic species of Calathea (Marantaceae) from Ecuador. J. Bot. Res. Inst. Texas 7(1 ):161 -167.

Kennedy, H. 2014. Calathea cofaniorum and C. shishicoensis, new endemic species of Marantaceae from Ecuador. J. Bot.

Res. Inst.Texas 8(1 ):37-42.

FIVE NEW MYRTACEAE FROM SOUTHEASTERN BRAZIL

Marcos Sobral

DCNAT-UFSJ

SaoJoao del-Rei, MG, BRAZIL

marcos_sobral@hotmail.com

Marcelo da Costa Souza

Dep. Botanica - UFRRJ

Seropedica, RJ, BRAZIL

souza.m@gmail.com

Ludovic Jean Charles Kollmann

Museu de Biologia Mello Leitao

Santa Teresa, ES, BRAZIL

ludovic@limainfo.com.br

OsnyTadeu Aguiar

Instituto Florestal de Sao Paulo

Sao Paulo, SP, BRAZIL

tadeu@iflorestal.sp.gov.br

Andre Luis Casarin Rochelle

Dep. Botanica - Unicamp

Campinas, SP, BRAZIL

rochelle@unicamp.br

Kelly Antunes

Dep. Botanica - Museu Nacional

Rio de Janeiro, RJ, BRAZIL

klybio@yahoo.com.br

ABSTRACT

We here describe, illustrate, and evaluate the conservation status of five new species from southeastern Brazil: Calyptranthes solitaria, C.

ubatubana, Eugenia dipetala, Myrcia cacuminis, and Plinia ambivalens. Calyptranthes solitaria, collected in the states of Minas Gerais,

Rio de Janeiro, and Sao Paulo, is apparently related to Calyptranthes tricona, from which it is kept apart by the uniflorous inflorescences and

glabrous adult leaves. Calyptranthes ubatubana is apparently restricted to the state of Sao Paulo and is related to Calyptranthes lanceolata,

from which it is distinguished by its pilose inflorescences and flowers. Eugenia dipetala and Myrcia cacuminis are from the state of Espirito

Santo. Eugenia dipetala is distinguished from other Brazilian species by flowers showing two petals; Myrcia cacuminis is related to M. capi-

tata, from which it is distinguished by the petiolate leaves, lanceolate blades, and triflorous inflorescences. Plinia ambivalens, from Espirito

Santo and Bahia, is related to Plinia involucrata, from which it differs by the leaves with larger petioles, venose blades, and ovaries with

multiovulate locules.

Key Words: Mata Atlantica, Calyptranthes, Eugenia, Myrcia, Plinia

RESUMO

Sao descritas, ilustradas e avaliadas quanto a sua conservagao Calyptranthes solitaria, C. ubatubana, Eugenia dipetala, Myrcia cacumi¬

nis e Plinia ambivalens. Calyptranthes solitaria, coletada nos estados de Minas Gerais, Rio de Janeiro e Sao Paulo, e aparentemente proxima

de Calyptranthes tricona, diferindo pelas inflorescencias unifloras e folhas adultas glabras. Calyptranthes ubatubana, aparentemente restrita

ao estado de Sao Paulo, e proxima de C. lanceolata, diferindo pelas inflorescencias e flores pilosas. Eugenia dipetala e Myrcia cacuminis ocor-

rem no estado do Espirito Santo. Eugenia dipetala difere de todas as especies brasileiras pela presenga de duas petalas; Myrcia cacuminis e

proxima de Myrcia capitata, diferindo pelas folhas pecioladas, laminas lanceoladas e inflorescencias trifloras. Plinia ambivalens, dos estados

da Bahia e Espirito Santo, e proxima de Plinia involucrata, diferindo pelas folhas com peciolos mais longos, laminas com venagao evidente e

ovarios com loculos multiovulados.

Key Words: Mata Atlantica, Calyptranthes, Eugenia, Myrcia, Plinia

INTRODUCTION

Myrtaceae comprise 22 native genera and about 990 species in Brazil (Sobral et al. 2014), from which about 680

grow in the coastal Brazilian Atlantic rainforest, which extends about 5,000 km from northeastern to southern

Brazil (Silva & Casteleti 2005), embracing a considerable part of Brazilian biological diversity and being con¬

sidered by Myers et al. (2000) as one of the diversity hotspots worldwide. However, because it has been subject

to such intense anthropogenic pressures such as urban expansion, farming, mining and legal or illegal logging

that reduced its cover to less than 10% of its original extent (Galindo-Leal & Camara 2005). Nevertheless, its

diversity is still poorly surveyed, and intensive collection efforts in this biome frequently bring to light several

undescribed and in some cases extremely local species. The examination of recent collections from southeast-

J. Bot. Res. Inst. Texas 8(2): 497 - 510.2014

498

Journal of the Botanical Research Institute of Texas 8(2)

ern Brazil have revealed to us some species that we consider as new, which are described here. For the evalua¬

tion of their conservation status, we used the conservation criteria proposed by IUCN (2012). In order to get an

estimate of the collection efforts in the municipalities cited in this paper, we compared the available data on

collection numbers (CRIA 2014) and the total area (IBGE 2014) of each municipality; although not especially

informative for conservation issues, this estimate can be interesting, since the scarcity of specimens from a well

surveyed location may suggest a real rareness of one species. For this, we consider as moderately surveyed ar¬

eas those with more than three gatherings/km 2 (an index proposed by Shepherd 2003); since the average of

gatherings in Brazil is smaller than one collection/km 2 (Sobral & Stehmann 2009), the Shepherd index may

serve as a good parameter for the intensity of botanical survey.

NEW SPECIES

Calyptranthes solitaria Sobral, Aguiar, & Antunes, sp. nov. (Figs. 1-2). Type: brazil, sao Paulo: Mun. Salesopolis,

Parque Estadual da Serra do Mar, trilha da Torre, 23 Feb 2005, E.M. Souza , M.T.Z. Toniato & D. Souza 495 (holotype: SPSF; isotype:

HUFSJ).

This species is apparently related to Calyptranthes tricona, differing through its coriaceous blades that easily fracture when bent (versus

chartaceous blades, bending without fracturing), which are mostly glabrous abaxially when adult (vs. visibly pilose), its uniflorous inflores¬

cences with bracts, if present, deciduous early before anthesis (vs. triflorous inflorescences with lanceolate bracts subtending the flowers

and frequently persisting through anthesis) and flowers with one or two petals (vs. petals absent).

Tree 2-14 m high. Twigs terete, dichotomously branched, the young ones velutinous, with simple brown tri-

chomes to 2 mm, the older ones entirely glabrous, the young cortex exfoliating, in some internodes tearing

longitudinally and falling as one entire piece; internodes 25-40 x 1-2.5 mm, the nodes slightly swollen and

with circular scars due to the exfoliation of the cortex, more visible at the proximal side of the node. Leaves

with petioles 4-10 x 1-2 mm, glabrous; blades elliptic, narrowly elliptic or ovate, 40-90 x 16-35 mm, 1.7-2.8

times longer than wide, coriaceous when adult, the young ones markedly discolorous, green and with scat¬

tered simple trichomes to 0.5 mm adaxially, brown and densely covered with an arachnoid indumentum of

curled simple trichomes 0.5-1 mm, along with some scattered straight simple trichomes to 2 mm, mature

leaves glabrous or with very scattered trichomes on the abaxial surface, slightly discolorous, the adaxial side

green or dark brown and with the venation very delicately sulcate, the abaxial surface light, dull green, with

glandular dots about 0.1 mm in diameter, ca. 5/mm 2 , visible only when examined against light; apex acumi¬

nate, 2-8 mm; base cuneate; midvein moderately sulcate adaxially and prominent abaxially; lateral veins 10 to

18 at each side, plane or finely sulcate adaxially, moderately prominent abaxially, departing from the midvein

at 45-60°; marginal vein 1-2 mm, occasionally a second vein visible at 0.2-0.3 mm from the margin, the mar¬

gin itself revolute and with a pale yellow thickening 0.1-0.2 mm wide. Inflorescences uniflorous, axillary, one

or two per axil, the pedicel 20-25 x 1-1.5 mm, with a dense indumentum of simple brown erect or ascending

trichomes to 1 mm; bracts not seen, possibly early deciduous; bracteoles not seen, apparently deciduous before

anthesis due to evident scars at the base of the flowers; flower buds ovate, 6-8 x 4 mm, densely covered with

trichomes as the inflorescences, with an evident apiculum to 1 mm, opening through a calyptra to 1 x 4 mm,

glabrous internally; petals one or two, rounded, to 1 x 1 mm (in bud), glabrous; stamens about 200, to 3 mm (in

bud), the anthers globose, 0.3 x 0.3 mm, eglandular; staminal ring to 2 mm in diameter; calyx tube 1-2 mm

deep, glabrous; style to 4 mm (in bud), the stigma acute, minutely papillose; ovary with two locules and two

ovules per locule. Fruits elliptic, immature, with trichomes as the flowers, 10 x 8 mm; seeds not examined.

Habitat, distribution and phenology.—Calyptranthes solitaria is a rainforest tree from the states of Minas

Gerais, Rio de Janeiro and Sao Paulo, growing at altitudes between 850-1500 m; flowers were collected in Feb¬

ruary and April and fruits in May.

Conservation. —This species was collected in four municipalities of Minas Gerais, Rio de Janeiro and Sao

Paulo, which total area is of 1,640 km 2 (IBGE 2014); these municipalities have been intensively surveyed, with

6,920 collections from Rio Preto, 2,262 from Salesopolis, 17,777 from Itatiaia and 18,585 from Ubatuba (data

from CRIA 2014), with a collection density of 27.7 gatherings/km 2 , which is notably higher than the Brazilian

average (Sobral & Stehmann 2009); the relative scarcity of collections of this species maybe an indicative of its

Sobral et al., New Myrtaceae from Brazil

499

herbArio d.bento pickel

SPSF- 34767

MYRTACEAE

Calyptranthes sp5

Del: O. T. Aguiar, 3/5/2005.

Brasil, Sao Paulo, Municlpio de Salesopolis, Parque Estadual

da Serra do Mar, Estrada Intermediary da PETROBRAS,

Trilha da Torre (SABESP). FlOresta Ombrofila Densa

Montana. (SAD69, 7386493S, 429146W, 1312m), 2/23/2005.

Arvore ca. 14 m, botoes ferrugineos.

Col.: F. M. Souza, 495. M. T. Z. Toniato; D. Souza.

Instituto Florestal — Sao Paulo

Fig. 1. Calyptranthes solitaria. Isotype at HUFSJ.

Journal of the Botanical Research Institute of Texas 8(2)

500

rareness. According to IUCN criteria (IUCN

2012), this species can be considered as vul¬

nerable (VU), btting criteria B1 ab(iii), since

its extent of occurrence is less than 20,000

km 2 (criterion Bl), its habitat is severely frag¬

mented (criterion a) and the habitat quality

and area are prone to decline, since two col¬

lection sites are not included in biological re¬

serves and are under anthropogenic pressures

such as farming and agriculture.

Affinities. —Due to its inflorescence

structure, it is apparently related to Calyp-

tranthes tricona D.Legrand (for description

see Legrand & Klein 1971; Tressens & Ro¬

driguez 1996; or Sobral 2003), a species from

the Atlantic forest growing from Minas Gerais

to Rio Grande do Sul, from which it can be

distinguished by the combination of charac¬

ters given in the diagnosis. The uniflorous

inflorescences of this species are reminis¬

cent of Calyptranthes pauciflora O.Berg (for

description see Berg 1857-1859:543), from

southeastern Brazil, which also show mostly

inflorescences with solitary flowers but sometimes also triflorous. Nevertheless, in spite of the inflorescence

character, this species does not seem related to Calyptranthes solitaria, due to its narrowly elliptic blades and

the inflorescences and flowers with densely appressed trichomes.

Etymology. —The epithet is derived from the Latin word for “lonely” or “solitary,” alluding to the uniflo¬

rous inflorescences.

Paratypes. BRAZIL. Minas Gerais: Mim. Rio Preto, Serra Negra, propriedade do sr. Nene Rock, 24 Apr. 2013, K. Antunes & R.J.V. Alves 455

(CESJ, HUFSJ, R). Rio de Janeiro: Mim. Itatiaia, Parque Nacional de Itatiaia, trilha do hotel Simon para os Tres Picos, 19 May 1999, A. Quinet

226 (BHCB, RB). Sao Paulo: Mum Ubatuba, 19 Apr 2007, R. Bertoncello 122 (BHCB, UEC).

Calyptranthes ubatubana Sobral & Rochelle, sp. nov. (Figs. 3-4). Type: BRAZIL. Sao Paulo: mun. Ubatuba, Picinguaba,

fazenda Capricorniojul 2007, A. Rochelle228 (holotype: UEC, isotypes: HUFSJ, RB).

This species is related to Calyptranthes lanceolata, from which it is distinguished by the densely pilose inflorescences with flowers crowded

at their apices (versus inflorescences and flowers glabrous, the inflorescences with flowers regularly distributed along axis).

Tree 3-7 m. Twigs terete, the young ones with a dense arachnoid indumentum of simple light brown to ochra-

ceous trichomes, 2-2.5 mm long, soon deciduous, then the twigs completely glabrous, grey to dark brown, the

distal internodes 35-50 x 2-3 mm, the one or two most distal internodes with stipule-like, lanceolate, adaxi-

ally folded cataphylls 40-50 x 8-10 mm, these occasionally with scattered asymmetrical dibrachiate trichomes

0.2-0.3 mm. Leaves with petioles 12-17 x 2 mm, glabrous or the young ones with trichomes as the twigs;

blades oblong to oblong-lanceolate, 170-200 x 32-60 mm, 4 to 7 times longer than wide, glabrous or the young

ones with scattered trichomes as the twigs along the midvein at the abaxial surface, concolorous when dry,

with 4 to 5 glandular dots/mm 2 , these smaller than 0.1 mm in diameter, visible abaxially; apex acuminate to 20

mm; base cuneate; midvein adaxially sulcate and prominent abaxially; lateral veins 25 to 35 at each side, visible

and prominent on both sides, more so abaxially, departing from the midvein at 45-50°, with one to three

secondary lateral veins between each pair; marginal veins two, 2-3 mm and 0.2-0.3 mm from the margin.

Inflorescences paniculiform, axillary or terminal, ramified one to three times, pendulous, emerging with

young shoots, the main axis 150-220 x 2-2.5 mm, slightly complanate, with a dense arachnoid indumentum

Fig. 2. Calyptranthes solitaria. Detail of inflorescences, from isotype (scale = 10 mm).

Sobral et al., New Myrtaceae from Brazil

501

Fig. 3. Calyptranthes ubatubana. Unmounted isotype at HUFSJ.

502

Journal of the Botanical Research Institute of Texas 8(2)

of simple light brown or ochraceous trichomes to 2.5

mm, these occasionally deciduous at anthesis, flowers

45 to 60, crowded apically in capituliform structures

formed by three to five extremely reduced axes; basal

bracts lanceolate, glabrous, adaxially folded, to 40 x 6

mm; bracts subtending the first ramifications lanceolate

or ovate-lanceolate, glabrous, 10-12 x 3 mm; bracts sub¬

tending the reduced axes widely ovate, to 5-7 x 7 mm,

slightly acuminate, with light brown, occasionally de¬

ciduous trichomes to 1 mm abaxially, the adaxial sur¬

face glabrous, the ultimate segments with three to five

flowers, each subtended by one bract similar to the basal

bract and with two elliptic to lanceolate bracteoles, 5-6

x 2-3 mm, glabrous adaxially and abaxially with brown

simple trichomes to 1 mm, occasionally enveloping the

flower buds, these elliptic to obovate, 5-6 x 2-3 mm,

uniformly covered with brown trichomes 0.5-1 mm,

with the calyx completely fused and a blunt apiculum to

r , „ , , ^ « * 1 mm; flowers opening through an irregular calyptra to

Fig. 4. Calyptranthes ubatubana. Detail of inflorescence; from isotype r

(scale = 10 mm) 2 mm in diameter; petals not seen in the flowers exam¬

ined, probably abortive; stamens 100 to 120, 6-7 mm,

the anthers globose to oblate, 0.4-0.5 x 0.4-0.6 mm, eglandular, opening through longitudinal slits; staminal

ring to 2 mm in diameter; calyx tube 2-2.5 mm deep; style 10-11 mm, the stigma slightly capitate and papil¬

lose; ovary with two locules and two ovules per locule. Fruits not seen.

Habitat, distribution and phenology.—Calyptranthes ubatubana is a tree from coastal rainforests of the

northern portion of the Brazilian state of Sao Paulo, where it was collected at altitudes 350-400 m. Flowers

were collected in April and July, but they were also observed in March (Rochelle, pers. obs.).

Conservation. —Considering the limited information currently available for this species, known for the

moment from two collections, assigning a conservation status for it is premature, even considering the fact that

it comes from the Brazilian Atlantic coastal forest, one of the most significant hotspots for conservation in the

planet (Myers et al. 2000). Also, the existence of only two collections being known from a location intensively

surveyed (20,324 gatherings in 723 km 2 , with an average of 28 collections/km 2 ; data from CRIA 2014 and IBGE

2014) may be an indicative of its rarity. Nevertheless, as there are no additional environmental informations,

we think it is convenient for the moment scoring it as Data Deficient (DD), according to IUCN criteria (IUCN

2012 ).

Affinities. —This species is related to Calyptranthes lanceolata O. Berg (for description see Legrand & Klein

1971), a species also found in the Brazilian coastal Atlantic forest from the states of Rio de Janeiro to Santa Ca¬

tarina, from which it can be distinguished through the characters cited in the diagnosis.

Etymology. —The epithet is derived from the municipality where the type specimen was collected.

Paratype. BRAZIL. Sao Paulo: Mim. Sao Sebastiao, Parque Estadual da Serra do Mar, 23°45'S, 45°36'W, 20 Apr 2000, A.A. de Oliveira et al.

(plus 9 collectors) 3624 (HUFSJ, UEC).

Eugenia dipetala Sobral & Kollmann, sp. nov. (Figs. 5-6). Type: BRAZIL. EspIrito Santo: Mun. Santa Maria de Jetiba, 13 Mar

2003, L. Kollmann 6035 (holotype: MBML; isotype: BHCB).

This species resembles in the leaf morphology Eugenia gracillima, but the blades are thicker, strongly bullate and markedly narrower in

outline (versus more or less membranous, smooth and elliptic in outline). Additionally, the presence of only two petals distinguishes it from

E. gracillima as well as from other species of Eugenia from southeastern Brazilian.

Shrub to 4 m. Twigs grey, subterete, with scattered white simple trichomes to 0.2 mm; internodes 20-25 mm.

heaves with petioles 1.5-2 x 0.4 mm; blades narrowly ovate, narrowly elliptic or oblong-ovate, 40-60 x 10-16

Sobral et al., New Myrtaceae from Brazil

503

Museu de Biologia Mello Leitao 020078

Santa Teresa - ES - Brasil _ _ MBML

MYRTACEAE

Loc.: Brasil. ES, Mun. de Santa Maria de Jetiba, fcerreno de

Reinaldo Berger.

Col.: L. Kollmann 6035, M.V.S. Berger.

Data: 13/03/2003.

Fig. 5. Eugenia dipetala. Isotype at BHCB.

Journal of the Botanical Research Institute of Texas 8(2)

mm, slightly discolorous when dry, markedly

bullate and glabrous adaxially, with scattered

white trichomes to 0.2 mm abaxially; glandular

dots visible against light, smaller than 0.05 mm

in diameter and about 30/mm 2 ; apex acuminate

to 10-12 mm; base acute or obtuse; midvein sul-

cate adaxially and prominent abaxially; lateral

veins 15 to 20 pairs, leaving the midvein at angles

of 80-90 degrees, sulcate adaxially and promi¬

nent abaxially; intramarginal vein 0.3-0.5 mm

from the revolute margin. Flowers 1 or 2 per ax-

ile; pedicels 16-20 x 0.4-0.5 mm, pilose as the

Fig. 6. Eugenia dipetala. Detail of old flower, from isotype (scale = 10 mm). blades; bracteoles triangular-lanceolate, 0.8-1 x

0.6-0.8, with cilia to 0.1 mm, persisting after an-

thesis; flower buds globose to obovate, to 4 x 3 mm, with white trichomes to 0.1 mm, these occasionally more

dense on the ovary; calyx lobes four, elliptic to rounded, concave, somewhat unequal, 2-3.5 x 1.8-4 mm; petals

two in the flowers examined, rounded, glabrous, white, markedly concave, to 3-5 x 3-5 mm; stamens 70 to 80,

3.5-4.5 mm; anthers elliptic, 1 x 0.5-0.6 mm, eglandular; staminal ring 2-3 mm in diameter; style 4-5 mm,

stigma punctiform or slightly capitate, minutely papillose; ovary locules 2; ovules 4 to 5 per locule. Fruits un¬

known.

Habitat, distribution and phenology. —This species is a shrub from forests of the montane region of central

Esplrito Santo. It is presently known only from the type collection, from the municipality of Santa Maria de

Jetiba. Flowers were collected in March.

Conservation. —The municipality of Santa Maria de Jetiba encompasses an area of 735 km 2 (IBGE 2014),

from from which 2,233 collections have been made (CRIA 2014), with a moderate average of 3 collections/km 2 ;

considering this, the uniqueness of the type specimen may be an indicative of its rareness. Nevertheless, con¬

sidering the absence of additional information, it seems more adequate to score this species as Data Deficient

(DD), according to IUCN criteria (IUCN 2012).

Etymology. —The epithet alludes to the two petals in all presently examined flowers of the species, a curi¬

ous feature evident among all flowers examined in our study, but otherwise unknown for species of the genus,

at least in southeastern Brazil

Affinities. —The leaves of this species somewhat resemble those of Eugenia gracillima Kiaerskou (for de¬

scription see Kiaerskou 1893:120), differing through the characters cited in the diagnosis.

Myrcia cacuminis Kollmann & Sobral, sp. nov. (Figs. 7-8). Type: BRAZIL. EspIrito Santo: Mun. Aguia Branca, 18°56'39"S,

40°47 , 55"W, 6 Sep 2006, L. Magnago, V. Demuner, T. Cruz & E. Bausen 1341 (holotype: MBML; isotype: BHCB).

This species is apparently related to Myrcia capitata, from which it is distinguished by the petiolate, narrowly elliptic leaves with mostly

glabrous adult blades (vs. sessile, ovate, abaxially pilose blades) with cordate bases (vs. cuneate bases) and up to three flowers per inflores¬

cence (vs. five to seven flowers).

Shrub 2-3 m. Twigs terete, 0.8-1.2 mm in diameter at the most distal internode, glabrous or with simple white

trichomes to 0.3 mm. heaves with petioles 4-5 x 0.3-0.7 mm, glabrous or with trichomes to 0.2 mm; blades

ovate to narrowly elliptic, 40-54 x 13-22 mm, 2.4-3 times longer than wide, glabrous or with scattered tri¬

chomes 0.3-0.4 mm along the midvein adaxially, the indumentum sometimes dense in young blades, concol-

orous when dry; glandular dots about 0.05 mm in diameter, 10 to 20/mm 2 , visible on both sides; apex and base

acute; midvein adaxially sulcate and abaxially prominent; lateral veins 12 to 17 at each side, visible and moder¬

ately prominent on both sides, leaving the midvein at angles 60-70 degrees; marginal veins two, 1-1.3 mm and

0.2-0.3 mm from the margin, the margin itself with grey cilia to 0.3 mm at least on young leaves. Inflores¬

cences emerging below the leaves, but coeval with them, one pair in each branch, the axis 20-35 x 0.5-0.6 mm,

with simple white trichomes 0.5-0.6 mm, with 2 or 3 sessile and apically crowded flowers; bracts three to five,

Sobral et al., New Myrtaceae from Brazil

505

Museu de Biologia Mello Leitao

Santa Teresa - ES - Brasil

MYRTACEAE

Det.:

MBML

Brasil, ES, Aguia Branca, Rochedo, area de afloramento rochoso, propr.r

Ailton Corteleti. , Alt.: 300-560m/s.m.

Coord. 18°56'39' 'S e 40°47*55"W

Col.: L. F. S. Magnago 1341, V. Demuner, T. Cruz, Elias Bausen

Data: 6/9/2006

Arbusto 2,5m de alt. Frutos imaturos verdes. Vegetagao rupestre.

Fig. 7. Myrcia cacuminis. Isotype at BHCB.

506

Journal of the Botanical Research Institute of Texas 8(2)

Fig. 8. Myrcia cacuminis. Detail of inflorescence, from isotype (scale = 10 mm)

Sobral et al., New Myrtaceae from Brazil

507

foliaceous, directly subtending the flowers, narrowly ovate to ovate, 9-10 x 4-4.5 mm, with cilia to 0.3 mm;

bracteoles foliaceous, ovate to narrowly ovate, to 5 x 2 mm, ciliate as bracts; flower buds globose or obovate, 4

x 3-4 mm, with white trichomes to 0.3 mm, the hypanthium densely pilose; calyx lobes five, subequal, trian¬

gular, 2.5-3 x 2 mm; petals rounded or oblate, 3-3.2 x 3.5 mm, white, ciliate; stamens about 100 (based on

scars over staminal ring); staminal ring 2-3 mm in diameter; calyx tube 0.5-1 mm deep; style to 9 mm, stigma

punctiform and minutely papillose; ovary bilocular, ovules 2 per locule. Fruits globose or oblate, 5-6 x 6-7

mm, unripe in the specimens examined, seeds 2 to 3, reniform, 4-5 x 4 mm, testa light brown, glossy and eas¬

ily detachable; embryos not completely developed.

Habitat, distribution and phenology.—Myrcia cacuminis is a shrub from rocky outcrops at altitudes from

100 to 900 m; it is presently known for the municipalities of Aguia Branca and Sao Roque do Canaa, in the

center of the southeastern Brazilian state of Esplrito Santo. Flowers and fruits were collected in October.

Conservation .—This species comes from two municipalities from central Esplrito Santo, Aguia Branca,

and Sao Roque do Canaa, with respectively 454 and 342 km 2 (IBGE 2014) and 2,447 and 1,601 collections from

each (CRIA 2014), resulting in an average of about five collections/km 2 , a reasonable amount of gatherings.

Considering this, the small amount of collections may be an indicative of its rareness. Additionally, it fulfills

criteria B1 ab(iii) for Endangered (IUCN 2012), since the presently known range of the species is less than

5,000 km 2 (criterium Bl), its habitat also is severely fragmented (Aguiar et al. 2005) (criterium a) and presents

a continuing decline in extension (criterium b(iii)), since municipalities in central Esplrito Santo have been

intensely deforested. (See Mendes & Padovan (2000) for data regarding the neighbouring municipality of

Santa Teresa.)

Affinities .—Due to its inflorescence structure, it is apparently related to Myrcia capitata O. Berg (for de¬

scription see Berg 1857-1859:154), a species collected in the cerrados of the state of Minas Gerais, from which

it is kept apart by the characters given in the diagnosis.

Etymology .—The epithet is derived from the Latin word for “summit,” and alludes to the rocky outcrops

on mountain tops where the species was collected.

Paratypes. BRAZIL. Esplrito Santo: Mim. Aguia Branca, 18°56 , 39"S, 40°47 , 55"W, 6 Sep 2006, L. Magnago et al. 1358 (BHCB, MBML, RB);

18°58 , 40"S, 40°39 , 56"W, 18 Oct 2006, V. Demuner et al. 2948 (MBML); mun. Sao Roque do Canaa, Sao Jacinto, Militon, 16 Oct 2004, L. Koll-

mann et al. 7069 (BHCB, MBML); idem, Misterioso, Pedra dos Tres Carneiros, 24 Oct 2004, Kollmann et al. 7126 (BHCB, MBML).

Plinia ambivalens M. Souza & Sobral, sp. nov. (Figs. 9-10). Type: BRAZIL. Bahia: Mun. Arataca, RPPN Caminho das Pe-

dras, em trilha de acesso ao Peito de Moga; 15°10 , 25"S, 39°20 , 30"W, 6 Aug 2006, M.M.M. Lopes, P. LabiakJ. Paixao & L. Gomes 1015

(holotype CEPEC; isotypes BHCB, NY).

This species is apparently related to Plinia involucrata, from which it is distinguished by the leaves with petioles to 4 mm (versus up to 3 mm

in P. involucrata ) and blades with midvein and lateral veins visible and moderately raised on both faces (vs. scarcely visible on both faces) and

locules with three or more ovules (vs. biovulate locules).

Tree to 13 m. Twigs with white appressed trichomes to 0.5 mm; internodes 15-25 x 1-2 mm, subterete,

sometimes longitudinally striate. Leaves with petioles 7-13 x 0.9-1.2 mm; blades elliptic, narrowly elliptic or

elliptic-obovate, 65-90 x 30-40 mm, 2-2.3 times longer than wide, slightly discoloured, with white appressed

trichomes to 0.5 mm scattered adaxially, more dense but not concealing the surface abaxially, glandular dots

smaller than 0.1 mm in diameter, sometimes visible on the surfaces, more visible against light, to 10/mm 2 ;

apex acuminate to 5-7 mm; base cuneate or attenuate; midvein plane, biconvex or salient adaxially, prominent

abaxially; lateral veins 15-20 pairs, visible and moderately salient on both faces; marginal vein 1-2 mm from

the revolute margin. Inflorescences axillary, glomeruliform, the axis 1-2 x 1.5 mm, with 2 to 4 sessile flow¬

ers; bracts ovate to widely ovate, basally with 4 to 10 linear colleters 0.3-0.4 x 0.05 mm adaxially, glabrous or

with cilia to 0.1 mm, in 3-4 series, progressively larger distally, the basal ones to 1 x 1.5 mm, the most distal

ones to 2 x 2.5 mm, these occasionally tearing in two halves due to the development of the flowers; bracteoles

apparently in two pairs (the proximal pair more properly a bract, that is, the axis from where the flower arises

must be considered a brachyblast), the proximal pair elliptic-ovate, 2-2.2 x 1.5 mm, with scattered simple

trichomes to 0.1 mm, initially fused at least partly, the distal pair elliptic-rounded, to 3 x 2 mm, glabrous or

508

Journal of the Botanical Research Institute of Texas 8(2)

MYRTACEAE

Arvore ca. 12m alt. Folhas subcoriaceas, discolores. Calice e corola

verde-claros; estames brancos. Foto.

Lopes, M. M. M. 1015 P. Labiak, J. Paixao & L. Gomes 6/8/2005

Det:

Fig. 9. Plinia ambivalens. Isotype at BHCB.

Sobral et al., New Myrtaceae from Brazil

509

with scattered trichomes to 0.1 mm, with evident cilia to

0.1 mm, partially fused at base, persisting after anthesis;

flowers buds obovate, 4-5 x 4 mm, uniformly covered with

white or grey simple trichomes 0.5-0.6 mm, with calyx

lobes fused; calyx lobes three to four, tearing irregularly,

to 1-2 x 2 mm, generally one lobe evidently larger than the

others, calyptriform; petals, in the flowers examined, two

to three, obovate, to 3 x 2 mm, somewhat vestigial; stamens

about 100, 5-6 mm, anthers elliptic, 0.3-0.4 x 0.2, some¬

times with one connectival gland; staminal ring 2-3 mm in

diameter; calyx tube 1 mm deep, glabrous, persisting after

anthesis; style to 6 mm, glabrous, stigma punctiform, pap¬

illose; ovary with two internally glabrous locules and three

to six ovules per locule. Fruits unknown.

Distribution, habitat and phenology. —This species is

known from montane rainforests of central Esplrito Santo

and southern Bahia, where it grows at altitudes 800-1000 m

above sea level. Flowers were collected in July and August.

Conservation.—Plinia ambivalens was collected in lo-

i i , Fig. 10. Plinia ambivalens. Detail of flower bud, from isotype

cahties more than 500 km apart, which makes its possible _ -jq

occurrence area larger than 20,000 km 2 , suggesting it could

be scored, according to IUCN criteria (IUCN 2012) as a

species of least concern (LC).

Affinities. —This species shows a troubling combination of characters that puts it on the border of two

closely related genera, Plinia and Neomitranthes Kausel ex D. Tegrand (for description see Tegrand & Klein

1977). Both genera are nested in the Plinia group as defined by Tucas et al. (2007), characterized by em¬

bryos with plano-convex cotyledons and a mean of nine ovules per locule, among other features. In practical

grounds, Neomitranthes and Plinia can be distinguished by the completely closed calyx that opens through a

calyptra and a high number of ovules per locule in Neomitranthes, while Plinia presents a calyx incompletely

closed, rarely forming a calyptra, and mostly two ovules per locule. The combination of characters present in P.

ambivalens leads to a dilemma, since it has a high number of ovules per locule (typical of Neomitranthes) com¬

bined with an incompletely closed calyx that tears in irregular lobes, occasionally one of them calyptriform

(typical of Plinia). Our decision for assigning the present species to Plinia rather than to Neomitranthes resides in

the arrangement of involucral bracts in the inflorescence, a feature apparently absent in Neomitranthes. Another

quite plausible possibility is that we are dealing with only one variable genus instead of two distinct lineages; if

this comes to be proved as the case, then Plinia would have priority over Neomitranthes, and our choice would

at least avoid one more combination in the overburdened nomenclature of the family. Additionally, we are

not aware of any species of Neomitranthes that should be compared with Plinia ambivalens, while among the

presently known species of Plinia it shows great similarity with the Amazonian Plinia involucrata (O. Berg)

McVaugh (for description see Berg, 1857-1859:375, under Myrciaria involucrata; McVaugh 1958:779 under

Plinia pinnata and McVaugh 1969:228), from which it is distinguished by the characters listed in the diagnosis.

Etymology. —The epithet alludes to the mixture in this species of characters of the genera Neomitranthes

and Plinia as they are presently defined.

Paratypes.— BRAZIL. Esplrito Santo: Mim. Santa Teresa, Sao Antonio, 20 Jul 1999, L. Kollmann, W. Pizziolo & E. Bausen 2704 (BHCB,

MBML, RB); idem, Parque Natural de Sao Lourengo, arvore 13 m, DAP 57 cm, flor branca, meia encosta, 9 Sep 2003, T.A. da Cruz, IN. Subtil

& D.T. da Cruz 1 (BHCB, MBML, RB).

Observations. —The collection Cruz et al. 1 is registered under this number in MBMT and in speciesTink (CRIA

2014); some duplicates, however, have been distributed as Cruz et al. 669 (the data in the labels are identical,

and at least the duplicate at BHCB has the number 669 erased and replaced by number 1).

510

Journal of the Botanical Research Institute of Texas 8(2)

ACKNOWLEDGMENTS

We are grateful for the collectors of the cited specimens and for the curators of cited herbaria for their valuable

help, for Rafaela Forzza, Erika von Sohsten Medeiros and Mariana Bunger for their help in preparing the illus¬

trations and to Barney Lipscomb and two anonymous reviewers for very useful suggestions that improved our

paper.

REFERENCES

Aguiar, A.P., A.G. Chiarello, S.L. Mendes, & E.N. de Matos. 2005. Os corredores central e da Serra do Mar na mata atlantica

brasileira. In: C. Galindo-Leal & I.G. Camara, eds. Mata atlantica, biodiversidade, ameagas e perspectivas. Belo Hori¬

zonte, Fundagao SOS Mata Altantica/Conservagao Internacional. Pp. 119-132.

Berg, 0.1857-1859. Myrtaceae. In: K.F.P. von Martius, org. Flora Brasiliensis 14(1 ):1 —656.

CRIA (Centro de Referenda em Informagao Ambiental). 2014. Specieslink (http://www.splink.org.br/index?lang=pt).

Galindo-Leal, C. & I.G. Camara. 2005. Status do hotspot mata atlantica: uma sintese. In: C. Galindo-Leal & I.G. Camara, eds.

Mata atlantica. Biodiversidade, ameagas e perspectivas. Belo Horizonte, SOS Mata Atlantica, Conservagao Internacio¬

nal. Pp. 3-11.

IBGE (Instituto Brasileiro de Geografia e Estatistica). 2014. IBGEcidades@ (http://www.ibge.gov.br/cidadesat/default.

php).

IUCN (International Union for the Conservation of Nature). 2012. IUCN red list categories and criteria: Version 3.1.2nd ed.

IUCN, Species Survival Commission, Gland, Switzerland, (on-line version at http://www.iucnredlist.org/technical-

documents/categories-and-criteria)

Kiaerskou, H. 1893. Enumeratio myrtacearum brasiliensium quas collegiunt Glaziou, Lund, Mendonqa, Raben, Reinhardt,

Schenck, Warming alique. In: E. Warming, ed. Symbolarum ad floram Brasiliae Centralis cognoscendam 39:1-199.

Landrum, L.R. & M.L. Kawasaki. 1997. The genera of Myrtaceae in Brazil: An illustrated synoptic treatment and identifica¬

tion keys. Brittonia 49:508-536.

Legrand, C.D. & R.M. Klein. 1971. Mirtaceas - Calyptranthes. In: R. Reitz, org. Flora ilustrada Catarinense, Itajai. Pp. 489-552.

Legrand, C.D. & R.M. Klein. 1977. Mirtaceas - Campomanesia, Feijoa, Britoa, Myrrh in ium, Hexachlamys, Siphoneugena, Myr-

cianthes, Neomitranthes, Psidium. In: R. Reitz, org. Flora ilustrada Catarinense, Itajai. Pp. 571-730.

Lucas, E.J., S.A. Harris, F.F. Mazine, S.R. Belsham, E.M. NicLughadha, A. Telford, P.E. Gasson, & M.W. Chase. 2007. Suprageneric

phylogenetics of Myrteae, the generically richest tribe in Myrtaceae (Myrtales). Taxon 56:1105-1128.

McVaugh, R. 1958. Flora of Peru - Myrtaceae. Publ. Field Mus. Nat. Hist., Bot. Ser. 13(4):569—819.

McVaugh, R. 1969. Flora oftheGuayana Highland - Myrtaceae. Mem. New York Bot. Gard. 18:55-286.

Mendes, S.L. & M.P. Padovan. 2000. A Estagao Biologica de Santa Lucia, Santa Teresa, Espirito Santo. Boletim do Museu de

Biologia Mello Leitao (nova serie) 11/12:7-34.

Myers, N., R. Mittelmeier, C. Mittelmeier, G.A.B. Fonseca, & J. Kent. 2000. Biodiversity hotspots for conservation priorities.

Nature 403:853-858.

Silva, J.M.C. & C.H.M. Casteleti. 2005. Estado da biodiversidade da mata atlantica brasileira. In: C. Galindo-Leal & I.G. Ca¬

mara, eds. Mata atlantica. Biodiversidade, ameagas e perspectivas. Belo Horizonte, SOS Mata Atlantica, Conservagao

Internacional. Pp. 43-59.

BROWNEOPSIS PUYENSIS (LEGUMINOSAE: CAESALPINIOIDEAE: DETARIEAE),

A NEW SPECIES FROM AMAZONIAN ECUADOR

David A. Neill and Mercedes Asanza

Universidad Estatal Amazonica

Puyo, Pastaza, ECUADOR

davidneill53@gmail.com

gman2260@gmail.com

ABSTRACT

Browneopsis puyensis D. A. Neill & Asanza (Teguminosae: Caesalpinioideae: Detarieae), a new species known only from a single adult tree

in a riverside urban park in the city of Puyo, the capital of Pastaza province at the western edge of the Amazon basin in Ecuador, is described

and illustrated. The new species is a large canopy tree attaining 30 m, and the ramiflorous inflorescences are borne on large horizontal

limbs. The white, nocturnal flowers are protogynous, a feature not reported previously for Browneopsis or related genera in the tribe Detar¬

ieae.

RESUMEN

Se describe y se ilustra Browneopsis puyensis D.A. Neill & Asanza (Leguminosae: Caesalpinioideae: Detarieae), una especie nueva cono-

cida unicamente de un solo arbol adulto en un parque urbano ribereno en la ciudad de Puyo, la capital de la provincia de Pastaza en la margen

occidental de la llanura amazonica de Ecuador. La especie nueva es un arbol grande de 30 m de altura, y las inflorescencias ramifloras salen

de las ramas principales horizontales. Las flores blancas y nocturnas son protoginas, una caracteristica no reportada previamente para

Browneopsis o para generos afines de la tribu Detarieae.

The genus Browneopsis Huber comprises seven known species from the western Amazon basin in Ecuador,

Peru and western Brazil, the Choco biogeographic region along the Pachic coast of Colombia and Ecuador, in¬

cluding the Magdalena valley of Colombia, and the Darien region of eastern Panama (Klitgaard 1991; Lewis et

al. 2005; Silverstone-Sopkin 2010). Browneopsis is separated from the related genus Brownea Jacq. by several

floral characteristics: Browneopsis lacks bracteoles and some or all of the petals are reduced in size, or entirely

lacking, and the floral parts of Browneopsis are white or cream whereas those of Brownea are red, orange, pink

(yellow in one recently described species, Brownea jaramilloi A.J. Perez & Klitg.; Perez et al. 2012). The floral

features are correlated with differences in pollinators: the red, diurnal flowers of Brownea pollinated by hum¬

mingbirds and the white, nocturnal flowers of Browneopsis pollinated by bats or moths. A molecular phylogeny

using chloroplast trnL intron sequences (Bruneau et al. 2001) suggested that Browneopsis and Brownea are not

monophyletic, but that more comprehensive sampling is needed (Lewis et al. 2005).

Browneopsis ucayalina Huber is locally common in Amazonian Ecuador and Peru, with up to 47 individu¬

als recorded in one-hectare forest inventory plots (data from unpublished appendix to ter Steege et al. 2013),

but most of the other species are evidently quite rare, and several are listed as Critically Endangered (CR) or

Endangered (EN) according to the IUCN Red List categories (IUCN 2001): B sanintiae Silverst. in Colombia

and B. macrofoliolata Klitg. and B. disepala (Little) Klitg. in coastal Ecuador (Silverstone-Sopkin 2010; Neill

2011). We describe herein an additional species of Browneopsis from Amazonian Ecuador that, based on avail¬

able evidence, is also rare and endangered.

Browneopsis puyensis D.A. Neill & Asanza, sp. nov. (Figs.l, 2). Type. ECUADOR: Pastaza province: Pastaza canton, Puyo,

River walk along the Rio Puyo, the “Paseo Tunstico” between the campus of the Universidad Estatal Amazonica and the “malecon”

near the city center, 1°28T0"S 77°59 , 40"W, 950 m, 26 Mar 2013 (fl.), David Neill, Javier Robayo & Mercedes Asanza 17614 (holotype:

ECUAMZ: isotypes: K, MO, NY, QCNE).

J. Bot. Res. Inst. Texas 8(2): 511 - 516.2014

512

Journal of the Botanical Research Institute of Texas 8(2)

30 cm

Fig. 1. Browneopsis puyensis D.A. Neill & Asanza. A. Leaf. B. Inflorescence bud. C. Inflorescence bract. D. Protogynous inflorescence in the female phase.

E. Open flower with sepals. F. Flower after the sepals are shed. G. Sepals, showing range of sizes. H. Inflorescence with pistils after sepals and stamens

have been shed; no fruit set was observed after this stage of flowering. Drawn from the type collection, Neill etal. 17614.

This species differs from other taxa of Browneopsis in the combination of characters including its large stature, a 30-m canopy-level tree; the

number of leaflets, with 7(-8) pairs of leaflets with long-acuminate apices; inflorescence position, with inflorescences ramiflorous, borne

on large horizontal limbs, not cauliflorous, axillary or terminal; and presence of only one vestigial, minute petal, or petals entirely absent.

Tree attaining 30 m, with trunk 66 cm diameter. Bark densely lenticellate, gray. Wood very hard and heavy.

Crown cylindrical with large horizontal limbs. Young branchlets, petioles and adaxial leaf surfaces densely

pubescent with light brown simple hairs, all parts becoming glabrescent with age. Leaves alternate, paripin-

nate, with 7(—8) pairs of leaflets; leaflets opposite to subopposite. Petiole + rachis 20-27 cm. Blades coriaceous,

Neill and Asanza, A new species of Browneopsis from Amazonian Ecuador

513

Fig. 2. Photograph of protogynous inflorescence of Browneopsispuyensis, in the female phase, shortly after anthesis in late afternoon before the first

night of flowering; the styles are exserted and beginning to straighten out, but the staminal filaments remain curled and covered by the cucullate

sepals until the following evening.

glaucous with a waxy coat on adaxial surface, becoming glabrescent with age. Proximal pair of leaflets shorter

than the 6(-7) distal pairs, broadly ovate, 4.5-6 cm x 1.8-3.2 cm, base rounded and mostly asymmetrical, apex

long-acuminate. Distal leaflets elliptic to narrowly lanceolate, 7-12 x 3-4 cm, base rounded and asymmetrical

or nearly symmetrical, apex long-acuminate. Inflorescences ramiflorous, borne on short shoots emerging from

large horizontal limbs, or on medium-diameter branches but not on small distal leafy branches, usually erect

but sometimes descending. Inflorescence bud globose, 6-8 cm in diameter, enclosed by 4-6 inflorescence

bracts. Bracts cucullate, 5-6 cm x 4-5 cm, apex rounded, base truncate, white, with dense light brown pubes¬

cence on abaxial surface; the adaxial surface glabrous or with a few scattered simple brown hairs. Flowers ca.

30-40 per capitulum, sessile, bracteoles absent; hypanthium 12-15 mm x 5-6 mm, glabrous; sepals 3-4, with

2 larger sepals and 1-2 smaller ones, larger sepals 35-40 x 15-18 mm, spathulate, cucullate, rounded at apex,

white with light brown pubescence on lower third of abaxial surface, smaller sepals 20-25 x 3-4 mm, nar¬

rowly spathulate, apex acute, petals 0-1, vestigial, ca. 2-3 x 1-2 mm, subulate; stamens 14-16, basally connate,

514

Journal of the Botanical Research Institute of Texas 8(2)

tube + free filaments 55-65 mm, filaments fused in basal 15-20 mm, tube open on one side, white, glabrous,

anthers versatile, ca. 6 x 1.5 mm; ovary + style 65-70 mm, ovary 15-20 x 3-4 mm, densely villous with light

brown hairs; style and stigma glabrous, stigma subcapitate. Fruit not seen.

Browneopsis puyensis is a large, canopy-level tree attaining 30 m in height, whereas most other species of

Browneopsis are smaller understory or subcanopy trees in humid lowland forests (Klitgaard 1991). An excep¬

tion may be Browneopsis excelsa Pittier in the Darien region of Panama, which has been described as a large

30-m tree (Schery 1951). With its 7(—8) pairs of leaflets, B. puyensis differs from its congeners in numbers of

leaflets: 2-3 pairs in B. excelsa, B. macrofoliolata and B. peruviana (J.F. Macbr.) Klitg., 2-4 pairs in B. cauliflora

(Poepp. & Endl.) Huber, 4-6 pairs in B. sanintiae, and 8-11 pairs in B. disepala. The leaves of B. puyensis are

remarkably similar to those of Brownea grandiceps Jacq. in the size, shape and number of leaflets. The latter

species occurs in Amazonian Ecuador, but at lower elevations and farther east than B. puyensis, and is a small

understory tree with terminal inflorescences and red flowers; in sterile condition, juvenile plants of these two

species in different genera may be quite indistinguishable. The position of the inflorescence in B. puyensis,

ramiflorous and usually erect on large principal branches, is unique among its congeners: B. ucayalensis and B.

cauliflora have cauliflorous inflorescences along the trunk, and the remaining species have terminal, subtermi¬

nal or axillary inforescence on small branches.

Distribution and ecology.—Browneopsis puyensis is known only from a single large adult individual, grow¬

ing in the middle of a brick sidewalk constructed around the base of the tree in recent years, along the “Paseo

Turlstico” on the banks of the Puyo River in the city of Puyo, the capital of Pastaza province, Ecuador, at 950 m

elevation. The site is about 100 m from the entrance to the campus of the Universidad Estatal Amazonica,

within a small and narrow remnant of primary forest trees along the river. Beside the adult tree is a 4 m juvenile

treelet of the same species, and several small saplings less than 1 m tall are in the forest understory near the

adult. The original forest in the area around Puyo has almost entirely been deforested and the land converted

to pastures, sugar cane fields and urban-suburban development, and a search for additional adult trees of this

species was not successful. Mr. Rafael Sancho, the former mayor of Puyo, described to the authors his initiative

to protect the riverine forest during his mayoral tenure, 1978-1982; at that time, a larger area of relatively intact

forest was still extant along the Puyo River, but subsequent municipal administrations did not continue the

conservation efforts and most of the remaining primary forest was lost during the following decades. The pres¬

ent brick walkway, the “Paseo Turlstico Rio Puyo”, was built by the Ministry of Tourism in 2010 and the walk¬

way is a favorite site for residents and visitors to stroll, jog and sometimes to swim in the river; besides Browne¬

opsis puyensis, several other rare and locally endemic tree species occur in the narrow belt along the riverine

park within the urban core of Puyo. The site is 20 km east of the eastern base of the Andes with a wet asea-

sonal climate; annual precipitation in Puyo is about 4000 mm; in the recently completed vegetation map of

Ecuador (Ministerio del Ambiente 2012), the original vegetation of the area is designated “Bosque siempre-

verde piemontano del Norte de la Cordillera de los Andes”. Growth rates of this species are not known, but

judging from the very dense heavy wood, evident in the branch wood, and the slow growth rates observed by

the authors in other genera and species of Detarieae in Amazonia, the 30-m, 66-cm diameter individual of B.

puyensis is probably several centuries old and predates the late 19th-century founding of the city of Puyo by a

wide margin.

Etymology .—The specific epithet refers to the city of Puyo, the capital of Pastaza province, where the only

known adult and juvenile individuals are known. The city s name is derived from the word “puyu” in Kichwa,

the language of the indigenous people of the region, and signifies “mist,” a term which aptly describes the per-

humid climate of the site.

Phenology and protogyny .—The large adult individual of Browneopsis puyensis flowered abundantly for

about a month each year it was observed, during late March to late April in 2013 and again in 2014. No fruits

were set in either year; however, the caretakers of the urban park along the Puyo River informed the authors

that in some past years, fruits matured with viable seeds during August. The presence of a few juvenile indi¬

viduals near the base of the large flowering tree suggests that seed production has indeed occurred. No addi-

Neill and Asanza, A new species of Browneopsis from Amazonian Ecuador

515

tional adult trees are known, and if the species is mostly self-incompatible but with occasional fruit set from

geitonogamous pollination, a low incidence of mature seed production could be expected.

The flowers of Browneopsis puyensis are protogynous at the level of the inflorescence. We observed that

the inflorescence buds open in late afternoon, and during the first night of anthesis, the styles are fully exserted

and straight, but the stamens remain curled up within the “pockets” of the two large cucullate sepals (Fig. 2.).

On subsequent nights, the sepals fall away and the staminal filaments are fully extended. The female phase of

the protogynous inflorescences may sometimes last more than one night before the sepals dehisce and the

staminal filaments unfurl completely. Protogyny in inflorescences of Browneopsis disepala from coastal Ecua¬

dor was previously described by Knudsen and Klitgaard (1998) as part of an extensive study of the pollination

biology and floral scent of that species. At night, we observed the inflorescences with the aid of binoculars,

looking for floral visitors, and detected hawkmoths in the vicinity of the inflorescences but were not able to

confirm that they actually visited the flowers.

IUCN Red List Category.—Browneopsis puyensis is known at present from a single adult individual with

flowers that may be mostly, though perhaps not completely, self-incompatible. Like most taxa in the tribe De-

tarieae, the species is probably slow-growing and confined to primary forest under natural conditions. The

region around the city of Puyo where this single tree was found has been largely deforested in recent decades

and no other fragments of primary forest have been found with this species present. In lieu of a comprehensive

search and a population study, it is logical to place the new species in the highest threat category of the IUCN

Red List system (IUCN 2001): Critically Endangered (CR). Given the easy accessibility of the one known flow¬

ering individual, it will be monitored in future years for possible fruit set, and vegetative propagation ex situ of

this species will be carried out by the Universidad Estatal Amazonica, following the guidelines of the Global

Strategy for Plant Conservation (http://www.cbd.int/gspc/strategy.shtml) adopted by the nations that are par¬

ties to the Convention on Biological Diversity.

ACKNOWLEDGMENTS

We thank Julio Cesar Vargas, Rector of the Universidad Estatal Amazonica (UEA), for his support for our

research and for the development of the UEAs new herbarium, the Herbario Amazonico del Ecuador

(ECUAMZ); Rafael Sancho, former mayor of Puyo, prefect of Pastaza and legislative representative for the prov¬

ince, for his efforts to promote the conservation of the Puyo River forest remnants, and his descriptions of the

development of the city over the past 50 years; Javier Robayo, for his help in obtaining the type specimens; and

Eric Meza Andrade for the line drawing of the new species. We thank Bente Klitgaard (k) and one anonymous

reviewer for helpful comments.

REFERENCES

Bruneau, A., F. Forest, P.S. FIerendeen, B.B. Klitgaard, & G.P. Lewis. 2001. Phylogenetic relationships in the Caesalpinioideae

(Leguminosae) as inferred from chloroplast trnL intron sequences. Syst. Bot. 26(3):487-514.

IUCN (International Union for Conservation of Nature and Natural Resources). 2001. IUCN Red List Categories and Cri¬

teria, Version 3.1. IUCN, Gland, Switzerland, and Cambridge, U.K. http://www.iucn.org.

Klitgaard, B.B. 1991. Ecuadorian Brownea and Browneopsis (Leguminosae-Caesalpinioideae): Taxonomy, palynology,

and morphology. Nordic J. Bot. 11:433-449.

Knudsen, J.T. & B.B. Klitgaard. 1998. Floral scent and pollination in Browneopsis disepala (Leguminosae: Caesalpinioideae)

in western Ecuador. Brittonia 50(2):174-182.

Lewis, G.P., B. Schrire, B. Mackinder, & M. Lock. 2005. Legumes of the World. Royal Botanical Gardens, Kew, Richmond,

Surrey, U.K.

Ministers del Ambiente (Ecuador). 2012. Mapa de ecosistemas del Ecuador Contiental. Ministerio del Ambiente, Quito,

Ecuador.

Neill, D.A. 2011. Fabaceae. In: S. Leon-Yanez, R. Valencia, L. Endara, C. Ulloa Ulloa, & H. Navarrete, eds. Libro rojo de las

plantas endemicas del Ecuador, 2 a edicion. Pontificia Universidad Catolica del Ecuador, Quito, Ecuador. Pp. 325-335.

516

Journal of the Botanical Research Institute of Texas 8(2)

Perez, A.J., B.B. Klitgaard, C. Saslis-Lagoudakis, & R. Valencia. 2012. Brownea jaramilloi (Leguminosae: Caesalpinioideae), a

new, over-looked species endemic to the Ecuadorian Amazon. Kew Bull. 68:157-162.

Schery, R.W. 1951. Flora of Panama: Leguminosae-Caesalpinioideae. Ann. Missouri Bot. Gard. 38(1 ):1 —94.

Silverstone-Sopkin, P.A. 2010. A new species of Browneopsis (Leguminosae, Caesalpinioideae) from the Cauca valley, Co¬

lombia. Novon 20:207-211.

ter Steege, H. (and 119 others). 2013. Hyperdominance in the Amazonian tree flora. Science 342(6156):1243092.

DOI:10.1126/science. 1243092

A NEW SPECIES OF GARNOTIA (POACEAE) FROM KERALA, INDIA

C.N. Sunil, V.V. Naveen Kumar, and M.G. Sanilkumar

Post Graduate and Research Department of Botany

SNM College, Maliankara

Ernakulam District, Kerala, INDIA

dr.cnsunil@yahoo.in

ABSTRACT

Garnotia variyamensis Sunil, Naveen Kumar, & Sanilkumar, a new species of Garnotia from Ernakulam District of Kerala, India, is de¬

scribed and illustrated. The species is distinguishable by the presence of dissimilar spikelet pairs, with one sessile or nearly so with an un-

awned lemma with empty or male flowers, the other pedicelled with a shortly bifid lemma with a geniculate awn in the sinus and bisexual

flowers. The species is also notable by the awned glumes and broad axes of the panicle and its branches. The novelty is probably allied to

Garnotia courtallensis.

Keywords: Poaceae, Garnotieae, Garnotia variyamensis, new species, India, Kerala, Ernakulam District

RESUMEN

Garnotia variyamensis Sunil, Naveen Kumar, & Sanilkumar, es una nueva especie de Garnotia de Ernakulam Distrito de Kerala, India, que

se describe y se ilustra. La especie se distingue por la presencia de pares de espiguillas disimilares, una sesil o casi con un lema mutico con

flores vacias o masculinas, la otra pedicelada con un lema cortamente bifido con una arista geniculada en el seno y flores bisexuales. La

especie es tambien notable por sus glumas muticas y ejes de la panicula y sus ramas anchos. La novedad esta relacionada con Garnotia

courtallensis.

INTRODUCTION

The genus Garnotia Brongn. (Subfamily-Panicoideae, Tribe-Garnotteae)is represented by ca. 39 species dis¬

tributed in Asiatic and Pacibc regions of the world (Mabberley 2008). Gould (1972) while revising Garnotia,

recognized 29 species and six varieties and categorised them under two sections, sect. Garnotia and sect.

Scoparia. In India, Garnotia is represented by about 12 species (Prakash &Jain 1979) and most of them are dis¬

tributed in Peninsular India. In Kerala, this genus has so far been represented by 10 species (Nayar etal. 2006).

The present new taxon from Kerala state is an additional species to the known species of the genus.

During floristic exploration in Ernakulam district of Kerala, India, the authors came across an interesting

Garnotia from a moist rocky area of Variyam in Edamalayar Forest range. A critical examination revealed that

the specimen was distinct from the hitherto known species of the genus. It is described and illustrated here as

a new species.

Garnotia variyamensis Sunil, Naveen Kumar, & Sanilkumar, sp. nov. (Figs. 1,2). Type: India, kerala: Ernakulam

District, Edamalayar Forest Range, Variyam, 10°12.595'N &76°52.44TE, elev. ca. 794 m, 16 Oct 2013, Sunil & Naveen Kumar 6193

(holotype: CAL; isotype: MH).

Differs from all G. courtallensis in its lower glume shorter than the upper glume and lemma with geniculate, twisted awn, but differs by the

large leaves, dissimilar spikelet pairs, awned glumes, and long-awned lemma of pedicelled spikelets.

Annual herbs. Culms 15-105 cm long, erect or geniculate, generally purplish; nodes villous, lower ones root¬

ing. Leaf sheaths upto 8 cm long, slightly keeled, striate, tuberculate hairy, margins ciliate; ligules ca. 0.5 mm

long, membranous; blades 4-15 x 0.8-2 cm, ovate-lanceolate or broadly lanceolate, base rounded to shallowly

cordate, margins scabrid towards apex, apex acuminate, softly pubescent, articulated to sheath, membranous,

lower side generally purplish, upper side greenish, collar hairy. Panicles 9-30 cm long, narrowly contracted;

branches 2-4 per node in distant fascicles, 1.8-4 cm long, appressed to main axis, erect; rachis triquetrous to

flattened, 1-1.3 mm broad, convex on back, scabrid on outer margins. Spikelets paired, dissimilar, one sessile

and the other pedicelled, 3-3.3 x 0.5-0.6 mm, lanceolate, acuminate at apex, 1-flowered; callus shortly bearded

J. Bot. Res. Inst. Texas 8(2): 517 - 521.2014

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Journal of the Botanical Research Institute of Texas 8(2)

Fig. 1 . Gamotia variyamensis. A. Habit; B. Base of Leaf; C. Single pedicelled spikelet; D. Sessile spikelet; E. Lower Glume; F. Upper Glume; G. Lemma of

pedicelled spikelet; H. Lemma of sessile spikelet; I. Palea; J. Stamen; K. Pistil; L. Caryopsis. Based on Sunil & Naveen Kumar 6193 (CAL).

Sunil et al., A new species of Garnotia from India

519

Fig. 2. Garnotia variyamensis. A. Habit; B. Inflorescence node showing racemes; C. Villous nodes; D. Leaf base; E. Lower glume; F. Upper glume; G. Lemma

of pedicelled spikelet; H. Palea. Photos by C.N. Sunil (type collection: Sunil&Naveen Kumar 6193 (CAL)).

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Journal of the Botanical Research Institute of Texas 8(2)

or glabrous; pedicels 2-3 mm long, triquetrous, scabrid on margins. Sessile spikelets empty or male or rarely

with rudimentary pistil, usually awnless; pedicelled spikelets bisexual, awned. Lower glume 2.6-2.9 x 0.5-0.6

mm, lanceolate, pale green, chartaceous, 3-nerved, lateral nerves scabrid, awned; awn upto 2.5 mm long, sca¬

brid. Upper glume 3-3.2 x 0.5-0.6 mm, lanceolate, chartaceous, 3-nerved, lateral nerves scabrid, awned; awn

upto 2 mm long, scabrid. Lemma 2.7-3 x 0.5-0.6 mm, lanceolate, margins in-rolled, acuminate and unawned

or shortly aristate in sessile spikelets and shortly 2-lobed, awned from sinus in pedicelled spikelets, subcoria-

ceous, hyaline, 3-nerved; awns 11-14 mm long, geniculate, scabrid, column 2-2.5 mm long, dark brownish,

twisted. Palea 2-2.2 x 0.4-0.5 mm, lanceolate, acute at apex, 2-nerved 2-keeled, hyaline. Stamens 3; anthers

0.5-0.7 mm long, oblong, cream-yellow.Ovary ca. 0.3 mm long, ellipsoid; style ca. 0.5 mm long, slender; stigma

0.5-0.7 mm long, purple, feathery. Lodicules 2, ca. 0.2 mm long, obovate, truncate. Caryopsis 1-1.2 x ca. 0.5

mm, oblong, pale brownish.

Flowering & Fruiting. —August-No vember.

Etymology .—The species is named after its type locality, Variyam, a tribal settlement within the Western

Ghats.

Habitat and Ecology.—Garnotia variyamensis grows on wet exposed rocky grassland among mosses in

association with Fripogon sivarajanii Sunil, T. wightii Hook.f., Dimeria spp., Exacum sessile L., Impatiens viscosa

Bedd., and Arundina graminifolia (D. Don) Hochr.

Faxonomic note.—Garnotia variyamensis resembles Garnotia courtallensis Thwaites in its lower glume

shorter than the upper glume and lemma with geniculate, twisted awn, but differs by the large leaves, dissimi¬

lar spikelet pairs, awned glumes, and long-awned lemma of pedicelled spikelets. Diagnostic morphological

characters of Garnotia variyamsis and Garnotia courtallensis are provided in Table 1 and in the following key.

KEY TO SECTIONS OF GARNOTIA IN INDIA

1. Leaf blade and leaf sheath junction distinct, articulated; leaf sheath margins glabrous to ciliate but not woolly_Sect. 1.

Garnotia

1. Leaf blade and leaf sheath junction indistinct, not articulated; leaf sheath margins woolly near base_Sect. 2. Scoparia

Sect. 1. Garnotia

1. Lemma with straight awn.

2. Some spikelets awned, others awnless; spikelets upto 3 mm long_G. micrantha var. micrantha

2. All spikelets awned; spikelets 3-5 mm long.

3. Leaf- blades with a row of long hairs immediately behind the ligule_G. fergusonii

3. Leaf- blades without a row of long hairs behind the ligule_G. acutigluma

1. Lemma with geniculate awn.

4. Awn of lemma 16-26 mm long.

5. Plants 5-25 cm long, usually growing with moss on tree trunks and rocks; lemma apex bifid, but the lobes not

setaceous; panicle lax, few flowered, spreading_G. arborum

5. Plants 20-50 cm long, not growing in moss on trees and rocks; lemma apex with narrow setaceous lobes; panicle

contracted, many flowered_G. polypogonoides

4. Awn of lemma 5-14 mm long.

6. Awn of lemma without well- marked twisted column; apex of leaf- blades ending in a filiform tip_G. arundinacea

6. Awn of lemma with well- marked twisted dark column; apex of leaf blades without filiform tips.

7. Lower glume longer than the upper one or subequal.

8. Awns geniculate at or near the base; leaf blades with a dense row of hairs immediately behind the ligule;

collar densely hairy_G. fergusonii

8. Awns geniculate well above the base; leaf- blades without a dense row of hairs behind the ligule; collar

glabrous_G. tenella

7. Lower glume shorter than the upper one.

9. Leaf- blades 1 -2 cm broad; spikelets dissimilar, one sessile and the other pedicelled, sessile spikelets male or

empty, its lemma awnless; glumes awned; awns on lemma 11-14 mm long_G. variyamensis

9. Leaf- blades 0.3-1 cm broad; spikelets similar, in unequally pedicelled pairs, all bisexual, its lemma awned;

glumes awnless; awns on lemma 3-7 mm long_G. courtallensis

SECTION 2. SCOPARIA

1. Spikelets awnless; leaf-blades oblanceolate, flat_G. exaristata

1. Spikelets awned; leaf- blades linear, canaliculated, triangular in section.

2. Awn geniculate at or near the base without well- marked twisted column_G. scoparia

2. Awn geniculate well above the base with a well- marked twisted column.

Sunil et al., A new species of Garnotia from India

521

Table 1 . Diagnostic morphological characters of Garnotia variyamensis and Garnotia courtallensis.

Character

Garnotia variyamensis

Garnotia courtallensis

Culm

15-105 cm high.

10-60 cm high.

Leaf blades

4-15 cm long, 0.8-2 cm broad.

2-9 cm long, 0.2-1 cm broad.

Panicles

9-30 cm long, axis and branches very broad,

triquetrous to flattened, convex on back.

4-15 cm long, axis and branches narrow and slender.

Panicle branches

Erect, more or less appressed to main axis.

Widely spreading or compacted.

Spikelets

Paired, dissimilar, one sessile and the other

pedicelled, sessile spikelets empty or male,

its lemma unawned

In unequally pedicelled pairs, similar, all bisexual

and awned or lower ones of panicle unawned.

Glumes

With up to 2.5 mm long awn.

Acute, unawned.

Lemma

Of pedicelled spikelets shortly 2-lobed at apex

and awned from sinus and that of sessile

spikelet acuminate and unawned.

Entire or sometimes two lobed, acute, and awned from

apex or rarely lower ones of panicle unawned.

Lemma awn

11-14 mm long.

3-7 mm long.

3. Spikelets 5-7.5 mm long; panicle 9-25 cm long, congested; awn of lemma 5-12 mm long_ G. schmidii

3. Spikelets 3-5.5 mm long; panicle 28-60 cm long, not congested; awn of lemma 3.5-5 mm long_ G. elata

Conservation status .—The distribution of Garnotia variyamensis showed that there were less than 245 mature

individuals restricted to an area of 5km 2 . By following IUCN criteria for assessing the status of rare and threat¬

ened plants (IUCN, 2001), Garnotia variyamensis is belonging to critically endangered (CR) category.

ACKNOWLEDGMENTS

The authors are grateful to the University Grants Commission, New Delhi, for financial assistance for the

study. We express our thanks to the Kerala Forest Department for permitting collection of specimens. Sincere

thanks are expressed to the manager, SNM College, Maliankara, for providing the facilities. Help rendered by

K.D. Girija and other members of the Department of Botany, SNM College, Maliankara, are also thankfully

acknowledged. We greatly appreciate careful reviews by Alok Chorghe, Jeffery Saarela, and an anonymous re¬

viewer.

REFERENCES

Bor, N.L. 1960. The grasses of Burma, Ceylon, India and Pakistan (excluding Bambuseae). Pergamon Press, London,

Oxford, New York, Paris.

Gould, F.W. 1972. A systematic treatment of Garnotia (Gramineae). Kew Bull. 27(3):515—562

IUCN. 2001. IUCN Red list categories and criteria (version 3.1). IUCN, Gland, Switzerland and Cambridge, U.K.

Mabberley, D.J. 2008.Mabberley's plant-book: A portable dictionary of plants, their classification and uses. Third Edition.

Cambridge University Press, Cambridge, U.K.

Nayar, T.S., A. RasiyaBeegam, N.Mohanan, & G. Rajkumar. 2006. Flowering plants of Kerala. Tropical Botanical Garden and

Research Institute, Palode,Thiruvananthapuram, India.

Prakash, V. & S.K. Jain. 1979. Poaceae: Garnotieae. In: Fasc. FI. India 3:1-16, Botanical Survey of India, Calcutta, India.

522

Journal of the Botanical Research Institute of Texas 8(2)

BOOK REVIEW

Kenneth D. Heil, Steve L. O’Kane, Jr., Linda Mary Reeves, and Arnold Clifford. 2013. Flora of the Four Corners

Region: Vascular Plants of the San Juan River Drainage. Arizona, Colorado, New Mexico, and

Utah. (ISBN-13: 978-1-930723-84-9, hbk). Missouri Botanical Garden Press, P.O. Box 299, Saint Louis,

Missouri 63166-0299, U.S.A. (Orders: www.mbgpress.info, orders@mbgpress.org, 1-877-271-1930).

$72.00,1098 pp., 8.5" x 11.25".

There is always an element of hubris inherent in any attempt at compiling a flora for a region. Plants, after all, grow to

suit their own needs, and trying to slot such diversity into neat human-friendly delineations requires a powerful

faith. How much more true is this when confronting a landscape as wild and varied as the Four Corners region?

And yet, the collaborative effort that is Flora of the Four Corners Region has kept the faith and created a remark¬

able text for a magnificent but too-often neglected region. As the authors themselves recognized, the Four Corners

region was often a marginal element of other floras that were based on political, rather than eco-regional, boundaries.

The argument in favor of floras based on ecoregions is a cogent one. After all, location may be the one definitive

fact attributable to a specimen of unknown origin. By limiting a flora to one region, the taxa treated can likewise be

limited to only those one is likely to encounter in a particular area. From the user’s perspective, it can be tremen¬

dously frustrating to be comparing a specimen to descriptions of species occurring hundreds of miles distant.

From the perspective of natural history, eco-regional floras allow a more comprehensive treatment of organ¬

isms that share a similar region, if not similar habitats. The likelihood of gene flow decreases sharply as linear dis¬

tance increases. Even though the Four Corners region contains over 2000 m of elevational variation, the region is

still united by aridity and a high (average) elevation. Individual populations have many opportunities to meet and

mingle under these conditions. The eco-regional approach of Flora of the Four Corners allows a systematic account of

this diversity considered in context.

That intensity of focus is the real strength of this work: scholarship backed by extensive field work and her¬

barium studies to provide real heft to the keys and descriptions contained within. The treatments show this attention

to detail. I am not qualified to review every key presented, but I have been spending an inordinate amount of time

focusing on the genus Astragalus in New Mexico, and I am, by now, intimately familiar with the keys provided by

Barneby in his seminal Atlas of North American Astragalus (Memoirs of the New York Botanic Garden, vol. 13,1964).

I have also spent significant time with the Flora of New Mexico by Martin and Hutchins, and Flora Neomexicana by

Allred and Ivey. I will not attempt direct comparisons, since each covers such diverse geographic regions. I can, how¬

ever, say that Flora of the Four Corners Region takes an approach to the genus that offers more than a simple pruning

of Barneby’s more geographically comprehensive work. In other words, I can attest that this work offers a new and

interesting approach to at least one diverse and difficult taxon.

Finally, the Flora includes many notes of ethnobotanical interest. Other works, of course, will offer much

greater depth on this subject, but it is profoundly refreshing to see traditional knowledge treated with respect and

inclusion. The authors of this work rightly recognize that they are the most recent in a long history of people interact¬

ing with the flora of this region.

My only wish for the Flora is that more illustrations had been included. As a researcher working primarily with

herbarium specimens, many of them more than three decades old, simple line drawings illustrating the fine points

of stipules, reproductive structures, and hair morphology would be of tremendous use. I recognize that this volume

was already decades in the making, and the inclusion of monochromatic line drawings would have been a significant

additional complication. Nevertheless, those drawings could be the single most effective means of elucidating an

identification.

Illustrations notwithstanding, however, this Flora of the Four Corners Region is a remarkable and extremely use¬

ful guide to the plants of some of the most rugged and beautiful terrain in the United States. The authors have created

a classic that wifi remain the definitive work for a long time to come.— Brian Witte, PhD, Research Associate, Botanical

Research Institute of Texas, Fort Worth, Texas, U.S.A.

J.Bot. Res. Inst. Texas 8(2): 522.2014

TRIPOGON MALABARICA (POACEAE: CHLORIDOIDEAE: CHLORIDEAE:

TRIPOGONINAE), A NEW SPECIES FROM KERALA, INDIA

Thoiba Kottekkattu and A.K. Pradeep

Department of Botany, University of Calicut

Kerala 673 635, INDIA

thoibakk@gmail.com; akpradeep1@rediffmail.com

ABSTRACT

A new species of Poaceae, Tripogon malabarica Thoiba & Pradeep from Malabar Wildlife Sanctuary, Western Ghats of Kerala, India, is

described and illustrated.

Keywords: Tripogon, Poaceae, new species, Western Ghats, India.

RESUMEN

Se describe e ilustra una nueva especie de Poaceae, Tripogon malabarica Thoiba & Pradeep de los Malabar Wildlife Sanctuary, Western

Ghats de Kerala, India.

INTRODUCTION

The genus Tripogon Roem. & Schult. (Poaceae: Chloridoideae: Chlorideae: Tripogoninae) is represented by 44

species distributed in Africa, Australia, and Temperate and Tropical Asia (Clayton et al. 2006). The genus is

known to have 20 species in India, which includes the recently described four new species (Murugesan &

Balasubramaniam 2008; Newmaster et al. 2008; Kabeer et al. 2009; Chorghe et al. 2013). With the addition of

the present species, the total number of species in India has become 21.

The authors, during the course of revisionary studies on Eragrostidinae s.l. in South India came across an

interesting population of Tripogon growing along steep granitic cliffs in Malabar Wildlife Sanctuary, Kakkay-

am along the Western Ghats of Kerala, in South India. Critical studies revealed it to be an undescribed species

of Tripogon, though they showed some similarities with T. lisboae and T. vellarianus (Table 1).

TAXONOMIC TREATMENT

Tripogon malabarica Thoiba & Pradeep, sp. nov. (Figs. 1-2). Type: INDIA. Kerala. Kozhikode Dt.: Malabar Wildlife Sanc¬

tuary, Kakkayam, 990 melev., 11°33.037'N, 075°55.024'E, 19 Sep 2013, Thoiba Kottekkattu 134436 (holotype: BRIT; isotypes: CALI,

K, MH).

Tripogon malabarica is closely allied to T. lisboae Stapf and T. vellarianus Pradeep but differs in having linear, coriaceous, villous leaf sheaths;

13-14 mm long spikelets with 4-7 florets; lemma with straight or geniculate, scabrid awn and keeled palea with a subulate apex.

Tufted perennial herb. Culms 25-65 cm high; nodes glabrous. Leaf blades 30-70 x 02 - 0.8 cm, linear-lanceo¬

late, margins slightly scabridulous or scabrid, upper surface pubescent with short white papillose hairs, gla¬

brous abaxially. Leaf sheaths linear-lanceolate, coriaceous, villous adaxially, 10-18 cm long, apex pubescent

with a tuft of 2.5-3.5 mm long hairs at both ends; ligules indistinct. Racemes terminal, 15-40 cm long, with

30-55 spikelets; rachis stout, glabrous. Spikelets 13-14 mm long, linear, distant, dorsiventrally flattened,

4-7-flowered; callus bearded. Lower glumes 2.5-4 x 1-1.5 mm, coriaceous, ovate-lanceolate, 1-keeled,

1-nerved, keels slightly scabrid, with acuminate apex. Upper glumes 4.5-6 x 1-1.5 mm, lanceolate, coriaceous,

1-keeled, scabrid, 3-nerved, acute at apex. Lemmas 5 x 1.5-2 mm, ovate-lanceolate, 3-nerved, slightly keeled,

scabrid; 1-awned, 3 mm long, scabrid, sometimes slightly geniculate towards apex. Paleas 3-5 x 1-1.5 mm,

obovate, hyaline, 2-keeled, ciliate, apex subulate, rachilla 1mm long, glabrous. Lodicules 2, c. 0.25-0.5 mm

long, truncate apex coarsely 3-toothed. Stamens 3; anthers 1.5-2 mm long, oblong; filaments 1mm long, slen¬

der, glabrous. Ovary 0.5-0.75 x 0.25-0.5 mm, obovate, styles 2, slender, hyaline, 0.5-0.75 mm long; stigma

0.75 mm long, plumose, creamy white. Grain not seen.

J. Bot. Res. Inst. Texas 8(2): 523 - 527.2014

524

Journal of the Botanical Research Institute of Texas 8(2)

Table 1. Diagnostic morphological differences between Tripogon lisboae, Tripogon vellarianus, and Tripogon malabarica.

Tripogon lisboae

Tripogon vellarianus

Tripogon malabarica

Culms

20-60 cm high

30-90 cm high

25-65 cm high

Ligule

an eciliate membrane

indistinct or absent

indistinct, a fringe of hairs at the ends

of leaf-sheath; hairs 2-3.5 mm long

Leaf-sheath

linear, flat or convolute, rigid

linear, closely clasping, rigid

linear-lanceolate, coriaceous,

and glabrous

villous adaxially

Racemes

7-25 cm long

30-40 cm long

15-40 cm long

Spikelets

3-12-flowered

8-10-flowered

4-7-flowered

Lower glume

asymmetrical, membranous,

symmetrical, membranous,

symmetrical, coriaceous, 1.5-4 mm

1 mm long, scabrid

4-5 mm long, glabrous

long scabrid

Upper glume

elliptic, 3-3.5 mm long,

elliptic-lanceolate, 6-7.5 mm

lanceolate, 2-6 mm long, coriaceous,

subcoriaceous,1 -nerved, slightly

long, membranous, 3-nerved,

3-nerved, scaberulose, acute at apex

scabrid, mucronate at apex

glabrous, acute at apex

Lemma

3-5 mm long, 2-fid, 1-awned,

5-10 mm long (excluding awn),

4-5 mm long (excluding awn), 2-fid,

awns 1-2 mm long, glabrous,

2-fid, 1 - awned, awns 3 mm long,

1 -awned, awns 3 mm long, scabrid,

straight

glabrous, straight

straight or geniculate at maturity

Palea

2-4 mm long, oblong, keels

4-8 mm long, narrowly elliptic,

2-4.5 mm long, oblong-lanceolate,

scaberulose, obtuse at apex

winged, keels puberulous,

acute at apex

keels scaberulose, subulate at apex

Caryopsis

narrowly oblong, terete

not seen

Distribution. —Presently Tripogon malabarica is known only from the Malabar Wildlife Sanctuary, Kak-

kayam in Kozhikode district of Kerala.

Ecology and Phenology. —This species grows from 950-1500 m elevation on steep granitic cliffs (Fig. 2),

road cuts and wet rocky hillsides and found growing in association with Chlorophytum malabaricum Baker,

Pouzolzia auriculata Wight, P. bennetiana Wight, Impatiens scapiflora Heyne ex Roxb., I. gardneriana Wight,

Kleinia grandiflora (Wall, ex DC.) Rani, Themeda triandra Forssk., and Panicum spp.; flowering late July to No¬

vember.

KEY TO SPECIES OF TRIPOGON IN SOUTH INDIA

1. Culms thickened below by the persistent leaf-sheaths; leaves < 2 cm long, equitant, rigid, pungent_ T. pungens C.E.C. Fisch.

1. Culms not thickened below by the persistent leaf sheaths; leaves > 2.5 cm long, not equitant, not rigid or pungent.

2. Lemma cleft at apex into 2 lobes, awned in the cleft; lobes awned or not.

3. Central awn of the lemma flexuous, capillary, and several times as long as lemma_ T. capillatus Jaub. &Spach

3. Central awn of the lemma straight or curved, not more than twice as long as the lemma.

4. Annuals; rachilla internodes 2mm; lemma apex 3-awned_ T. cope Newmaster, V. Balas., Murug. & Ragup.

4. Perennials; rachilla internodes 1mm; lemma apex 1-awned.

5. Upper glumes 5.5-7 mm long; awns of the lemmas 6-8 mm long, straight or curved_ T. wightii Hook.

5. Upper glumes 8-9.5 mm long; awns of the lemma 10.5-12 mm long, always straight_ T. velliangiriensis

Murug. &V. Balas.

6. Perennials with wiry roots forming tufts; leaf blade glabrous or scabrid; lower glumes asymmetrical.

7. Leaves and culms glaucous, leaves 5-20 cm long, involute, filiform; ligule very short but definite, ciliate

_T. jacquemontii Stapf

7. Leaves and culms not glaucous, leaves 30-60 cm long, 4-8 mm wide, usually flat, sometimes rolled; ligule

obsolete_ T. lisboae Stapf

6. Perennials with fibrous roots, forming a close turf; leaf blade glabrous or villous adaxially; lower glumes

symmetrical or asymmetrical.

8. Leaves flat, ligules indistinct; lower glume symmetrical.

9. Ligules indistinct, a fringe of hairs at the ends of leaf sheath; spikelets with 4-7 florets; lemma 4-5 mm

long; keels of palea scaberulose, subulate at apex_ T. malabarica Thoiba & Pradeep

9. Ligules indistinct or absent; spikelets with 8-10 florets; lemma 5-10 mm long; keels of palea puberulous,

acute at apex_ T. vellarianus Pradeep

8. Leaves flat to convolute, ligules membranous, lower glume asymmetrical.

10. Ligules membranous; lemma base glabrous, median awn as long as or shorter than the lemma;

inflorescence 20-45 cm long; spikelets with 5-8 florets_ T. sivarajanii Sunil

1 mm

Thoiba and Pradeep, A new species of Tripogon from Kerala, India

525

Fig. 1. Tripogon malabarica Thoiba & Pradeep. A. Habit. B. Spikelet. C. Lower glume. D. Upper glume. E. Lemma. F. Palea. G. Stamen. H. Pistil. A-H,

drawn from holotype.

526

Journal of the Botanical Research Institute of Texas 8(2)

Fig. 2. Steep granitic cliffs at Malabar Wildlife Sanctuary, Kakkayam, Kozhikode, the natural habitat of Tripogon malabarica Thoiba & Pradeep. Photo

taken by Thoiba Kottekkattu, 21 Aug 2014.

Thoiba and Pradeep, A new species of Tripogon from Kerala, India

527

10. Ligules ciliate, membranous; lemma base bearded, median awn 1.5 times longer than lemma;

inflorescence 15-20 cm long; spikelets with 9-10 florets_ T. tirumalae

Chorghe, Rasingam, Prasanna, & Sankara Rao

2. Lemma cleft at apex into 4 or 6 lobes or with a definite lobes between each lateral awn and central; outer lobes if

present awned or not.

11. Leaves hairy; ligules inconspicuous; keels of the palea scabrid_ T. bromoides Roth

11. Leaves glabrous or hairy; ligules conspicuous, thin, membranous; keels of the palea ciliate or not.

12. Lemma cleft at apex into 6- lobes.

13. Upper glume 7-9 mm long, notched at apex with small awn between; keels of palea ciliate; stamens 1.3-2

mm long_ T. anantaswamianus Sreek., VJ. Nair, & N.C. Nair

13. Upper glume 3 mm long, acute at apex, deeply 2-lobed with arista 0.5 mm long; keels of palea not ciliate;

stamens upto 0.8 mm long_ T. borii Kabeer, VJ. Nair, & G.V.S. Murthy

12. Lemma cleft at the apex into 4- lobes.

14. Culms 10-20 cm tall; lower glumes shallowly lobed on one-side above the middle; central lobes of the

lemmas ovate-acute_ T. narayanae Sreek., VJ. Nair, & N.C. Nair

14. Culms 24-70 cm tall; lower glume deeply lobed on one-side below the middle; central lobes of the lemmas

lanceolate_ T. ravianus Sunil & Pradeep

ACKNOWLEDGMENTS

We would like to thank Kerala Forest Department for permission to do field studies at Malabar Wildlife

Sanctuary; University of Calicut for providing financial support; and two anonymous reviewers for comments

that improved the manuscript.

REFERENCES

Chorghe, A., L. Rasingam, P.V. Prasanna, & M.S. Rao. 2013. Tripogon tirumalae (Poaceae), a new species from the Seshacha-

lam Hills of Andhra Pradesh, India. Phytotaxa 131(1 ):17—22.

Clayton, W.D., M.S. Vorontsova, K.T. Harman, & H. Williamson. 2006 onwards. GrassBase-the online world grass flora. Avail¬

able at http://www.kew.org/data/grass.db.html. Accessed 10 Aug 2013.

Kabeer, K.A.A., VJ. Nair, & G.V.S. Murthy. 2009. Tripogon borii —a grass species new to science from India. Bull. Bot. Surv.

India 50:115-118.

Murugesan, M. & V. Balasubramaniam. 2008. Tripogon velliangiriensis (Poaceae)—a new species from Tamil Nadu, India.

Indian J. Forest. 31:109-111.

Newmaster, S.G., V. Balasubramaniam, M. Murugesan, & S. Ragupathy. 2008. Tripogon cope (Poaceae: Chloridoideae), a new spe¬

cies supported by morphometric analysis and a synopsis of Tripogon in India. Syst. Bot. 33:695-701.

528

Journal of the Botanical Research Institute of Texas 8(2)

BOOK NOTICE

Richard B. Taylor. 2014. Common Woody Plants & Cacti of South Texas. (ISBN-13: 978-0-292-75652-6,

flexbound). University of Texas Press, PO Box 7819, Austin, Texas 78713-7819, U.S.A. (Orders: www.

utexaspress.com, 1-800-252-3206). $22.95, 152 pp., 117 color photos, illus. glossary, bib., index, 4.5" x

7.25".

This is a held guide for the identification of and critical information related to the values and uses of the 50 most com¬

mon trees, shrubs, cacti, yucca, and succulents of the South Texas “brush country”. It was written by a retired Texas

Parks & Wildlife Department biologist. Its primary focus is wildlife, but it includes a great deal of valuable informa¬

tion about resource management and values for a variety of other concerns.

The book’s compact size makes it an ideal reference to take along on hikes, held trips, and explorations. The 50

plants selected for inclusion in the book are the most signihcant of the more than 281 woody species and 32 species

of cacti, yucca, and succulents found in the region. They represent more than 75% of the total tree, shrub, and cacti

component of South Texas.

The book begins with a historical perspective on the social, cultural, and economic factors that have influenced

the plant communities, ranching enterprises, and wildlife habits of South Texas. That is followed by a description of

the history of brush management activities and efforts in recent history. Also included in the introductory section is

general guidance on habitat appraisal and evaluation techniques.

The Quick Key for plant identification is one of the most useful keys I have seen for woody plants. Species are

quickly grouped into three broad categories: (1) thorned woody plants, (2) thornless woody plants, and (3) cacti,

yucca, and succulent plants. From there each broad category is further subdivided and systematically narrowed

down using easily identified characteristics. For example the thorned woody plants are subdivided into two catego¬

ries: (1) those with straight thorns and (2) those with curved thorns. Thornless plants are placed into subcategories

based on growth form (tree-like or shrub-like). As the user progresses through the Quick Key using the recognizable

characteristics to limit the possibilities, even relatively inexperienced botanists can easily narrow the list of potential

species until finally the user has a positive identification, or at least a very small group of potential plants.

Following the quick key are narrative descriptions of each of the 50 plants. Species are listed by scientific name

but also include one or more common names (including Spanish common names). The descriptions are well-written,

easy to understand and very informative. They are accompanied by at least one, and often two or more, color photo¬

graphs showing the plant in its natural habitat. The descriptions and pictures help the user verify the identity of each

plant. The narrative description of each species also includes a value rating of the plant for wildlife, livestock, and

other uses.

In addition to the typical information listed above, this guide also includes at the end of each narrative descrip¬

tion the crude protein content of the leaves and/or fruits of the plant during each season of the year. This is critical

information for wildlife biologists, ranch managers, landowners, and others who depend on these browse plants to

provide the nutritional needs of wildlife and livestock. The information is a valuable reference source to aid in wild¬

life management.

Appendix I is a table showing the habitat values (food, cover, nesting, roosting, water, etc.) that the included

plants provide for the following categories of wildlife: game birds, song birds, insects, small mammals, deer, and

reptiles. Appendix II is a table listing the palatability and browse preference of each plant species for white-tailed

deer. Plants are described as 1st choice, 2nd choice, 3rd choice, etc. Appendix III lists the percentages of crude pro¬

tein, digestible protein, and digestible dry matter for each plant across the seasons. And Appendix IV is a listing of the

common names and scientific names of every plant, vertebrate, and invertebrate mentioned in the book. A two-page

illustrated glossary, a bibliography, and an index round out the book’s end matter.

This held guide is an invaluable resource for wildlife managers, biologists, ranchers, conservationists, amateur

botanists, students, and anyone else interested in the vegetation of the South Texas “brush country .”—Dan Caudle,

Independent Grassland Consultant, Weatherford, Texas, U.S.A.

J.Bot. Res. Inst. Texas 8(2): 528.2014

WANDERSONG (RUBIACEAE), A NEW GENUS FROM THE GREATER ANTILLES

David W. Taylor

Department of Biology

University of Portland

Swindells Hall 108, MSC 163

5000 N. Willamette Blvd.

Portland, Oregon 97203, U.S.A.

taylorda@up.edu

ABSTRACT

A revision of Chione DC. (Rubiaceae) has excluded two species, C. exserta (DC.) Urb. and C. seminervis Urb. & Ekman, from the genus on the

basis of morphology. The designation Colleteria D.W. Taylor was proposed to house these species, but it was invalidly published. The new

genus Wandersong D.W. Taylor is here erected to accommodate these two species. The new combinations Wandersong exserta (DC.) D.W.

Taylor and Wandersong seminervis (Urb. & Ekman) D.W. Taylor are here effected under this new generic name. The new genus name

Wandersong D.W. Taylor honors two Andersons, Dr. GregoryJ. Anderson and Dr. William R. Anderson, for their outstanding and influential

achievements in the field of botany. The name Wandersong is constructed with the surname Anderson shared between the first initials of

these two botanists.

RESUMEN

Una revision taxonomica de Chione DC. (Rubiaceae) ha excluido dos especies, C. exserta (DC.) Urb. y C. seminervis Urb. & Ekman, del ge-

nero debido a su morfologia. Se propuso la designacion Colleteria D.W. Taylor para acomodar estas especies, pero su publicacion fue invali-

da. Aqui se erige el nuevo genero Wandersong D.W. Taylor para acomodar estas dos especies. Aqui tambien se efectuan las nuevas combina-

ciones Wandersong exserta (DC.) D.W. Taylor y Wandersong seminervis (Urb. & Ekman) D.W. Taylor. El nombre del nuevo genero,

Wandersong D.W. Taylor, honra a dos botanicos con el apellido de Anderson, Dr. GregoryJ. Anderson y Dr. William R. Anderson, por sus

logros tan sobresalientes y sus investigaciones sumamente influyentes en el campo profesional de la Botanica. El nombre Wandersong esta

construido con el apellido Anderson compartido entre las primeras iniciales de los nombres de estos dos botanicos.

Keywords: Wandersong D.W. Taylor, Wandersong exserta (DC.) D.W. Taylor, Wandersong seminervis (Urb. & Ekman) D.W. Taylor, Colleteria

D.W. Taylor, Colleteria exserta (DC.) D.W. Taylor, Colleteria seminervis (Urb. & Ekman) D.W. Taylor, Chione exserta (DC.) Urb., Chione semi¬

nervis Urb. & Ekman, Dr. William R. Anderson, Dr. GregoryJ. Anderson.

During a taxonomic revision of the genus Chione DC. (Rubiaceae), two species, C. exserta (DC.) Urb. and C.

seminervis Urb. & Ekman, were found to be morphologically outside the circumscription of the genus, and

were not referable to any existing genus (Taylor 2003a). The designation Colleteria D.W. Taylor was proposed

to house these two species (Taylor 2003b), but inadvertently used a morphological term, which is not permit¬

ted (Article 20.2 of the International Code of Nomenclature for algae, fungi, and plants (Melbourne Code);

International Association for Plant Taxonomy website; http://www.iapt-taxon.org/nomen/main.

php?page=art20; website accessed June 4, 2014). The term “colleteria” is the plural of “colleterium,” a technical

entomological term for “an accessory glandular structure of oviduct which manufactures and secretes viscid

material used to cement eggs together” (Gordh & Headrick 2011). Thus, the designation Colleteria was not

validly published under Art. 32.1. A new genus name, Wandersong D.W. Taylor, is here published to accommo¬

date Chione exserta and Chione seminervis. The new combinations Wandersong exserta (DC.) D.W. Taylor and

Wandersong seminervis (Urb. & Ekman) D.W. Taylor are here effected under this new generic name.

The new genus name Wandersong D.W. Taylor honors two Andersons, Dr. GregoryJ. Anderson and Dr.

William R. Anderson, for their outstanding and influential achievements in the held of botany. The name Wan¬

dersong is constructed with the surname Anderson shared between the first initials of these two botanists. The

author wishes to honor these two botanists specifically because of their extraordinary mentorship, first in their

roles as his undergraduate and Ph.D. advisors, respectively, at the University of Connecticut and the University

J. Bot. Res. Inst. Texas 8(2): 529 - 530.2014

530

Journal of the Botanical Research Institute of Texas 8(2)

of Michigan, and then later as professional colleagues. Both of these Andersons, through their great dedication,

generosity, and insight, have been instrumental in shaping the author’s career as a botanist.

The three nomenclatural changes proposed here are therefore:

1. Wandersong D.W. Taylor, gen. nov. Type: Wandersong exserta (DC.) D.W. Taylor.

A full description and diagnosis of this new genus is found under the designation Colleteria D.W. Taylor

(2003b:203).

The genus Wandersong D.W. Taylor can be diagnosed by a combination of characters, some of which are:

colleters present consistently on adaxial face of stipules and often on adaxial face of calyx, corolla lobes imbri¬

cate in bud, anthers inserted at or below middle of corolla tube, fruit with two pyrenes, the pyrenes adaxially

concave, abaxially convex and ridged, with marginal germination slits extending from apex to middle, each

pyrene with one locule, ovules solitary and pendulous in locules. Chione is distinguished from Wandersong by

many vegetative and reproductive features (see Taylor 2003a and 2003b), including the absence of colleters and

each fruit having a single, sclerenchymatized pyrene enclosing two locules. The fruit type of Wandersong, in¬

cluding pyrene architecture, is similar to that reported for some members of the Psychotrieae (Taylor 1989,

1996; Andersson 2002), but Wandersong is distinct in having apical placentation.

2. Wandersong exserta (DC.) D.W. Taylor, comb. nov. Basionym: Psychotria exserta DC., Prodr. 4:517. 1830. Chione exserta

(DC.) Urb., Symb. antill. 8:675.1921. Colleteria exserta (DC.) D.W. Taylor, Syst. & Geogr. Pi. 73(2):204. 2003, nom. inval. Type: DO¬

MINICAN REPUBTIC: Santo Domingo, Bertero s.n. (holotype: G-DC, not seen, microfiche, photo: MO).

Taylor (2003b) has written a detailed description of this species, under the designation Colleteria exserta (DC.)

D.W. Taylor.

3. Wandersong seminervis (Urb. & Ekman) D.W. Taylor, comb. nov. Basionym: Chione seminervis Urb. & Ekman, Ark.

Bot. 20A(5):59.1926. Colleteria seminervis (Urb. & Ekman) D.W. Taylor, Syst. & Geogr. Pi. 73(2):206.2003, nom. inval Type: HAITI.

Massif de la Selle: Nouvelle Touraine, high ridge towards Morne La Visite, (Petionville, Roberjat), in forest on eruptive, ca. 1700 m,

Ekman H1537 (lectotype, here designated because holotype at B destroyed: S; isotypes: F, G, GH, S, US).

Taylor (2003b) has written a detailed description of this species, under the designation Colleteria seminervis

(Urb. & Ekman) D.W. Taylor.

ACKNOWLEDGMENTS

The author wishes to express his appreciation to Christiane Anderson, Mona Anderson, Rev. Robert Antonelli,

Fred Barrie, Roy Gereau, Joseph Kirkbride, Jr., Barney Lipscomb, Charlotte Taylor, and an anonymous re¬

viewer for their very kind help. The author is extremely grateful to both Gregory J. Anderson and William R.

Anderson for their countless thoughtful contributions to his life.

REFERENCES

Andersson, L. 2002. Relationships and generic circumscriptions in the Psychotria complex (Rubiaceae, Psychotrieae).

Syst. & Geogr. PI. 72:167-202.

Gordh, G. & D. Headrick. 2011. A dictionary of entomology, 2 nd edition, p. 333. CABI, Cambridge, Massachusetts, U.S.A.

International Code of Nomenclature for algae, fungi, and plants (Melbourne Code). 2012. Available at the International Associa¬

tion for Plant Taxonomy website; http://www.iapttaxon.org/nomen/main.php?page=art20. Accessed June 4, 2014.

Taylor, C.M. 1989. Revision of Palicourea (Rubiaceae) in Mexico and Central America. Syst. Bot. Monogr. 26:1-102.

Taylor, C.M. 1996. Overview of the Psychotrieae (Rubiaceae) in the Neotropics. In: E. Robbrecht, C. Puff & E. Smets, eds.

Second international Rubiaceae conference proceedings. Opera Bot. Belg. 7:261-270.

Taylor, D.W. 2003a. A taxonomic revision of the genus Chione (Rubiaceae). Syst. & Geogr. PI. 73(2):171-198.

Taylor, D.W. 2003b. Colleteria (Rubiaceae), a new genus from the Caribbean. Syst. & Geogr. PI. 73(2):199-208.

VARIABILITY OF VEGETATIVE FLUSH COLORS IN ZAMIA (CYCADALES)

Claudia Calonje

Montgomery Botanical Center

11901 Old Cutler Rd.

Coral Gables, Florida, 33156 USA.

Michael Calonje

Montgomery Botanical Center

11901 Old Cutler Rd.

Coral Gables, Florida, 33156 US.A.

and Florida International University

Miami, Florida 33174, US.A.

michaelc@montgomerybotanical.org

Lindy Knowles

Bahamas National Trust

Nassau, THE BAHAMAS

Chad Husby

Montgomery Botanical Center

11901 Old Cutler Rd.

Coral Gables, Florida, 33156 US.A.

ABSTRACT

The color of new vegetative leaf flushes can vary among species of cycads. It is often used as a diagnostic character in taxonomy and is one of

the most appreciated features of the group in ornamental horticulture. However, little is understood about the variability of flush color

within species and wild populations. This paper discusses variability in flush color within the genus Zamia and uses a comprehensive ex situ

conservation collection of three Zamia species (Z. angustifolia, Z. integrifolia, and Z. lucayana ) collected from six islands in The Bahamas

(Abaco, Andros, New Providence, Eleuthera, Grand Bahama, and Tong Island) as a case study to examine this variability between species,

between and within populations, and within progeny of individual mother plants. Flush color was found to be exclusively brown in Z. an¬

gustifolia and green in Z. lucayana. For Zamia integrifolia, flush color was exclusively green in seedlings derived from Abaco, Grand Bahama,

and Eleuthera, but was variable in seedlings derived from Andros in New Providence, where both colors were present. On these two islands,

flush color varied between and within populations, as well as within progeny of individual mother plants.

The observed variability in flush colors suggests that a large number of in situ observations from multiple plants in several populations

would be required to determine if a particular flush color character state is fixed and therefore useful as a diagnostic character. This can only

be done through long term study of in situ populations, as leaf production is typically seasonal and is not necessarily synchronized or uni¬

versal within wild populations. Therefore, we recommend against the use of flush color as a diagnostic character in cycad taxonomy.

RESUMEN

El color de las hojas nuevas de cicadas puede variar entre diferentes especies. A menudo se utiliza como caracter diagnostico en taxonomia

y es una de las caracteristicas mas apreciadas del grupo en horticultura ornamental. Sin embargo, se sabe muy poco sobre la variabilidad en

el color de hojas nuevas dentro de las especies y poblaciones silvestres. En este articulo se discute la variabilidad en el color de hojas nuevas

dentro del genero Zamia y se usa una coleccion extensa de conservacion ex situ de tres especies de Zamia (Z. angustifolia, Z. integrifolia, y Z.

lucayana ) colectadas de seis islas de las Bahamas (Abaco, Andros, New Providence, Eleuthera, Grand Bahama, y Long Island) como un estu-

dio de caso para examinar esta variabilidad entre especies, en y entre poblaciones, y en descendientes de la misma planta madre. El color de

hojas nuevas fue exclusivamente cafe en Z. angustifolia y verde en Z. lucayana. Para Zamia integrifolia, el color de hojas nuevas fue exclusi-

vamente verde en plantulas derivadas de Abaco, Grand Bahama, y Eleuthera, pero variables en plantulas de Andros y New Providence,

donde ambos colores estaban presentes. En estas dos islas, el color de hojas nuevas variaba en y entre poblaciones asi como tambien entre

descendientes de una misma planta madre.

La variabilidad observada sugiere que un numero grande de observaciones in situ de multiples plantas en multiples poblaciones serian

requeridas para determinar si el estado de caracter para el color de hojas nuevas es un caracter fijo que pueda servir como caracter diag¬

nostico. Esto solo se podria hacer a traves de estudios de poblaciones in situ a largo plazo porque la produccion de hoja nueva ocurre por

temporadas y no necesariamente esta sincronizado o es universal en las poblaciones silvestres. Por estas razones recomendamos que el color

de las hojas nuevas no se utilice como caracter diagnostico en la taxonomia de las cicadas.

INTRODUCTION

Cycads are highly appreciated as ornamental plants due to their ease of culture, long life span, and distinctive

appearance. They reproduce by means of pollen and seed cones that are always produced on separate plants

(i.e. they are dioecious), and can be quite distinctive in some species. The seed cones in particular are one of the

J. Bot. Res. Inst. Texas 8(2): 531 - 540.2014

532

Journal of the Botanical Research Institute of Texas 8(2)

most appreciated ornamental characteristics of cycads. They can be quite large in some species, as exemplified

by Eepidozamia peroffskyana Regel (Fig. 1A), that has cones reaching 1 m in length and 40 kg in weight (Nor-

stog & Nicholls 1997:235) and are considered the largest cones in the plant kingdom. In addition to attaining

impressive dimensions in some species, seed cones can often be quite colorful, particularly in the African ge¬

nus Encephalartos as exemplified by the bright orange cones of E.ferox G. Bertol (Fig. IB).

In addition to being appreciated for their highly ornamental strobili, cycads are also highly prized for

their foliage. Cycads have distinctive pinnately compound leaves which are held in a rigid crown. Although

cycad leaves are relatively conservative in their morphology compared to some other plant groups (Norstog &

Nicholls 1997:49), they can be quite variable in terms of texture, leaflet arrangement, and the color of new leaf

flushes. This variation in foliage is best exemplified by the neotropical genus Zamia , which is considered to be

the most morphologically diverse genus in the order Cycadales.

Zamia, consisting of 75 accepted species (http://cycadlist.org), is the most species-rich cycad genus in the

New World and has the broadest geographic distribution, ranging from Southern Georgia to Bolivia. Through¬

out its vast geographic range, the genus occupies a variety of diverse environments ranging from open, desert-

like environments, to dense rainforests in some of the rainiest places on earth. Adaptation to these different

environments has resulted in remarkable variation in leaf form within the genus.

Most species of Zamia may hold multiple leaves per crown, but some species, such as Z. meermanii Calo-

nje, typically hold only 1-3 leaves per crown. Leaflets in Zamia vary in texture from soft and paper-like as in Z.

vazquezii D.W. Stev., Sabato & De Luca (Fig. 2A), to rigid and cardboard-like, as in Z. encephalartoides D.W.

Stev. (Fig. 2B), a desert-dwelling species from Colombia. Some species, such as Z. hamannii A.S. Taylor, J.L.

Haynes & Holzman (Fig. 2C), have broad, deeply-veined leaflets.

Leaflets vary in size from the diminutive leaflets of Z. pygmaea Sims, which may represent the smallest

leaflets in the cycadales (Fig. 2D), to the large paddle-like leaflets of Z. imperialis A.S. Taylor, J.L. Haynes &

Holzman which at up to 75 cm in length and 21 cm in width (Taylor et al. 2008) may be the largest leaflets

found in living cycads (Fig. 2E). Leaflets also vary in arrangement from being widely spaced along the rachis,

such as those of Z. imperialis, to closely spaced and overlapping, such as in Z. chigua Seem. (Fig. 2F).

In addition to the remarkable diversity in leaflet shape, size, texture and arrangement, the genus Zamia

exhibits the greatest diversity in the color of new vegetative flushes. The color of these young developing leaves

is perhaps one of the most ornamentally appreciated features of the genus. New leaves are produced in an array

of different colors, including bright orange (Z. pyrophylla Calonje, D.W. Stev., & A.Lindstr.; Fig. 3A), light green

(Z. nesophila A.S. Taylor, J.L. Haynes & Holzman; Fig. 3B), reddish brown (Z. lacandona Schutzman & Vovides;

Fig. 3C), white (Z. stevensonii A.S. Taylor & Holzman; Fig. 3D), reddish-orange (Z. hamannii; Fig. 3E), and

reddish-brown (Z. purpurea Vovides, J.D. Rees & Vazq. Torres; Fig. 3F).

Flush color appears to be a fixed trait in many Zamia species and is sometimes used as a diagnostic taxo¬

nomic character. For example, Z. hamannii is distinguished from closely related species by its reddish-orange

to rosy-pink flushes (Fig 3E; Taylor et al. 2008), whereas Z. stevensonii (Fig. 3D) is distinguished from its puta¬

tive sister species Z. elegantissima Schutzman, Vovides & R.S. Adams by its white flushes compared to the

bright yellow flushes ofZ. elegantissima (Taylor & Holzman 2012).

Although flush color appears to be fixed in many species, it can also be variable in some species. For ex¬

ample, a wide array of colors has been observed in cultivated individuals of some species such as Z. standleyi

Schutzman andZ. vazquezii (Broome 2002). As the exact provenance and pedigree of ornamentally cultivated

cycads is often unknown, it is unclear whether the variation observed in some cultivated species is the result of

source germplasm derived from separate populations with distinct flush colors, or if this variation can be

found within individual wild populations.

Little has been published regarding the variability of flush color in wild populations of Zamia other than

Schutzmaris (1984) report of separate populations of Z. splendens Schutzman (currently considered synonym

of Z. katzeriana (Regel) E. Rettig, sensu Nicolalde-Morejon et al. 2009) which produce either brilliant red or

light green leaf flushes. However, the extent to which flush color may vary among and within populations of

Zamia or even among the progeny of a single mother plant has not been previously examined.

Calonje et al., Vegetative flush colors in Zamia

533

Fig. 1. A. Mature cone oUepidozamiaperoffskyana held by Dr. Andrew Vovides. B. Seed cone of Encephalartos ferox.

Fig. 2. Leaf variability in Zamia. Leaves of: A)Z. vazquezii, B)Z. encephalartoides, C)Z. hamannii, D)Z. pygmaea, E)Z. imperials, and F)Z. chigua.

534

Journal of the Botanical Research Institute of Texas 8(2)

Fig. 3. Variability of flush color in Zamia. A )Z pyrophylla, B)Z nesophila, C )Z. lacandona, D)Z stevensonii, E)Z imperialis, F)Z purpurea, G)Z amplifolia,

H)Z integrifolia from New Providence, Bahamas displaying variable flush color in two plants derived from same mother plant.

Calonje et al., Vegetative flush colors in Zamia

535

Since flush color is often used as a diagnostic character, understanding flush color variability could have

taxonomic implications. Similarly, understanding the variability in flush colors may inform future selective

breeding of desirable flush colors for ornamental horticulture.

In this paper we explore variability of flush color by examining a taxonomically and geographically com¬

prehensive ex-situ conservation collection of Bahamian Zamia plants cultivated at Montgomery Botanical

Center (Coral Gables, Florida, USA). This collection was developed from 2009-2011 as part of a conservation

genetics project focused on Zamia lucayana (see Calonje et al. 2013).

MATERIALS AND METHODS

Seeds were collected from wild populations to establish ex-situ conservation collections at Montgomery Bo¬

tanical Center (MBC) and The Retreat of Bahamas National Trust (Nassau, New Providence, The Bahamas).

Sampling encompassed the entire known distribution range for the genus in The Bahamas, including multiple

populations on six different islands (Abaco, Andros, Eleuthera, Grand Bahama, New Providence, and Long

Island) and representing three different species according to current species delimitations (Osborne et al.

2012). The three sampled species were: Zamia angustifolia from Eleuthera, Z. lucayana from Long Island, and

Zamia integrifolia, which as currently circumscribed has a wide distribution including Abaco, Andros, Eleu¬

thera, Grand Bahama, and New Providence (Fig. 4). Zamia angustifolia is distinguished by having very narrow

leaflets, whereas Z. lucayana has the broadest leaflets. Zamia integrifolia has leaflets that vary in width on dif¬

ferent islands; populations from Andros, New Providence and Eleuthera have narrower leaflets and popula¬

tions from Grand Bahama and Abaco have broader leaflets (Fig. 5).

Sampling of seeds followed protocols developed at Montgomery Botanical Center that aim to maximize

the genetic diversity of the ex-situ collections by sampling seeds from up to five mother plants per population

(see Walters 2003). Seeds derived from individual mother plants, representing half siblings, are kept separate

and provided their own accession number.

Seeds were germinated and seedlings cultivated at MBC. Flush color was documented on plants with new

leaves during the period of 2012-2013. New vegetative flushes were observed on a total of 1133 seedlings com¬

prising 62 separate accessions derived from a total of 14 wild populations.

The distribution of flush color in the seedlings was analyzed using JMP 10.0.2 statistical software (SAS

Institute, Inc, Cary, North Carolina, USA). Contingency table analysis was used to detect variation in leaf color

distribution across islands and across species. Following significant contingency table homogeneity tests, bi¬

nomial analyses were used for detailed comparisons among species, populations, accessions, and islands.

RESULTS

Flush color by species and island

At the species level, flush color was exclusively brown in Z. angustifolia (from 2 populations in Eleuthera), ex¬

clusively green in Z. lucayana (from 3 populations in Long Island), and variable in Z. integrifolia, where exclu¬

sively green flushes occur on Abaco, Grand Bahama, and Eleuthera, and both colors occur on Andros and New

Providence (Fig. 3). It should be noted that species delimitation in Caribbean Zamia is particularly unclear,

and species determinations are tentative pending ongoing genetic studies of Caribbean Zamia (e.g., Calonje et

al. 2013; Salas-Leiva et al. 2013; Meerow et al. 2012) that seek to clarify the genetic relationships among these

populations and shed some fight on their taxonomy and nomenclature.

Green leaf flushes were observed on seedlings from all six islands, whereas brown flushes were limited to

seedlings from New Providence, Andros, and northern Eleuthera (Figs. 6-8). The three islands where brown

flushes occur are adjacent to each other and were part of a single landmass known as Paleoprovidence in the

Pleistocene when sea levels were lower. Along these three islands, there appears to be a longitudinal gradient

in flush color proportion, as seedlings of Z. angustifolia from northern Eleuthera were entirely brown emergent

(100% brown), those from New Providence mostly brown (85% brown), and those from Andros mostly green

(33% brown). Contingency table analysis showed significant variation among islands (p < 0.0001) and species

(p < 0.0001) (Tables 1, 2).

536

Journal of the Botanical Research Institute of Texas 8(2)

0 25 50 100

Kilometers

Species

A Z. angustifoiia

0 Z. integrifolia

O Z. lucayana

Fig. 4. lamia collection sites in The Bahamas.

Flush Color by Population

Of the three Bahamian species, only Z. integrifolia exhibited variation in flush color, and only on the islands of

Andros and New Providence (Figs. 6-7), where variation was found among populations, within populations,

and within progeny of individual mother plants which were treated as separate accessions (Fig. 7). On Andros,

observed Zamia populations were restricted to North Andros, and only the southernmost sampled populations

yielded offspring with brown-emergent leaves (Fig. 7). In New Providence, only a small number of accessions

yielded offspring with green flushes. Contingency table analysis showed significant variation among popula¬

tions (p < 0.0001) (Table 3 for detailed comparisons).

DISCUSSION

Flush color in Bahamian Zamia species appears to be genetically determined, as variation in flush color was

found within wild populations on Andros and New Providence, and within the progeny of individual mother

plants cultivated under identical conditions in MBC’s ex-situ conservation collection. Plants with different

flush colors from both islands were otherwise consistent in vegetative morphology.

Cycads are diploid, meaning that they have two sets of homologous chromosomes which may carry two

different forms of a particular gene (alleles) at the same location. We have observed only green and brown leaf

flushes on Bahamian Zamia , indicating that there may be at least two different alleles in a single gene for flush

color present in Bahamian Zamia populations. The variability observed suggests that the alleles for particular

flush colors may be fixed in some species, populations, and half-sibling cohorts, but polymorphic in others.

Calonje et al., Vegetative flush colors in Zamia

537

Eleuthera

Andros

New Providence

Grand Bahama

Abaco

Long Island

Fig. 5. Characteristic leaflet shapes of Bahamian Zamia populations.

Table 1. Numbers of brown and green emergent plants on different islands in the Bahamas.

Island

# Brown

emergent

# Green

emergent

Lower 95% CL* for %

Brown emergent

Upper 95% CL* for %

Brown emergent

Abaco

280

0.00%

1.31%

Long Island

192

0.00%

1.90%

Eleuthera

158

0.00%

2.31%

Grand Bahama

138

0.00%

2.64%

Andros Island

224

13.58%

23.03%

New Providence

124

77.45%

89.82%

Eleuthera

176

97.93%

100.00%

* CL = Confidence Limit

Table 2. Numbers of brown and green emergent plants for different Zamia species in the Bahamas.

# Brown

# Green

Lower 95% CL* for %

Upper 95% CL* for %

Species

emergent

Brown emergent

Zamia lucayana

192

0.00%

1.90%

Zamia integrifolia

173

823

15.07%

19.87%

Zamia angustifolia

176

97.93%

100.00%

* CL = Confidence Limit

The differences in allele frequency among Zamia populations are likely the result of a combination of evo¬

lutionary forces including natural selection, genetic drift (particularly in small populations), and gene flow

between populations (migration).

Assuming this trait is controlled by a single gene, it is not known whether a particular allele for flush color

538

Journal of the Botanical Research Institute of Texas 8(2)

_[topic p! fornper

Z. integrifolia

n= 258

ubpopulations= 3

Accessions= 11

Z. integrifolia

n= 115

Subpopulations= 1

Accessions= 5 %

Z. angustifolia

n= 163

lSubpopulations= 2

Accessions= 7

Z. integrifolia

n= 163

Subpopulations= 5

Accessions= 8

New

Providence

Eleuthera

Z. integrifolia

n= 128

| Subpopulations= 1

Accessions= 5

Z. integrifolia

n= 128

Subpopulations= 3

Accessions= 11

Isla'nd

Z. lucayana

n= 178

Subpopulations= 4

Accessions= 15

| Proportion of brown flushes

Proportion of green flushes

Fig. 6. Flush color distribution by species and island. Every island is represented by a single species except Eleuthera, which has separate populations of

Z. angustifolia and Z. integrifolia with distinct flush colors.

may be dominant over the other, or whether this dominance may be consistent among different taxa. As the

ex-situ collection at MBC is now beginning to reach reproductive maturity, the inheritance patterns of flush

color may be determined by conducting controlled crosses between different plants and analyzing the flush

color proportions of the resulting progeny Caribbean species of Zamia are ideal for experimental crossing,

because they can reach reproductive maturity in two to three years, which is relatively fast compared to other

cycad species which may need a decade or longer to reach reproductive maturity

It is not clear whether the brown flush color offers a selective advantage to cycads, but there are several

possible hypotheses suggesting an advantage to brown or reddish flushes. The brown tint is believed to be

produced by anthocyanins which may have fungicidal properties, protect leaves against UV damage, or make

them inconspicuous to herbivores (Queenborough et al. 2013).

CONCLUSIONS

Flush color in Bahamian zamias varies geographically as well as at the species, population, and progeny levels,

with color appearing to be fixed in some populations and variable in others. This variability suggests that to

determine whether flush color is a fixed and therefore a diagnostic character for a given species, one would have

to record flush color data from multiple individuals in each population and sample multiple populations. This

is challenging data to collect because leaf production is seasonal and may not be synchronized among popula¬

tions of a single species or individual plants within a population. Furthermore, a proportion of the population

may not produce any leaves during a particular growing season, so collecting enough data to determine this

would require long term study. Even if a fixed flush color were observed for a large sample of wild plants, at

Calonje et al., Vegetative flush colors in Zamia

539

Fig. 7. Flush color ofZ. integrifolia by accession in northern Andros and New Providence. Each bar represents a sampled accession, the height of the bar

depending on the number of seedlings sampled from each accession, and the colors representing the proportion of flush colors found in the seedlings

sampled from each accession.

Table 3. Numbers of brown and green emergent plants for different Zamia populations in the Bahamas.

Species

Population

# Brown

emergent

#Green

emergent

Lower 95% CL* for

% Brown emergent

Upper 95% CL* for %

Brown emergent

Zomio lucoyono

192

0.00%

1.90%

Zamia integrifolia (Eleuthera)

158

0.00%

2.31%

Zamia integrifolia (Grand Bahama)

138

0.00%

2.64%

Zamia integrifolia (Abaco)

130

0.00%

2.80%

Zamia integrifolia (Abaco)

0.00%

4.02%

Zamia integrifolia (Abaco)

0.00%

5.96%

Zamia integrifolia (Andros Island)

0.00%

6.16%

Zamia integrifolia (Andros Island)

0.00%

10.28%

Zamia integrifolia (Andros Island)

26.02%

73.98%

Zamia integrifolia (Andros Island)

44.13%

71.86%

Zamia integrifolia (New Providence)

74.09%

88.63%

Zamia integrifolia (New Providence)

77.23%

99.15%

Zamia angustifolia

176

97.93%

100.00%

* CL = Confidence Limit

present it would be difficult to use this character state diagnostically in the absence of similar data available for

other species.

Although a large number of leaf flushes were observed in this study, the flush color proportions reported

for a given wild population are based on the progeny of a relatively small number of sampled mother plants,

540

Journal of the Botanical Research Institute of Texas 8(2)

and are not necessarily representative of the populations as a whole. Consequently, we believe that flush color

information should be included in cycad species descriptions but should not be used as a diagnostic character

for taxonomy

Flush color is variable in some species and appears to be genetically determined. Future studies to deter¬

mine the heritability of this characteristic would be useful to inform selective breeding of cycads to emphasize

desirable flush colors for ornamental horticulture.

ACKNOWLEDGMENTS

Field work in The Bahamas was conducted in close collaboration with The Bahamas National Trust, USDA

(Subtropical Horticulture Research Station, Miami), and Fairchild Tropical Botanic Garden, and was support¬

ed by The Mohamed bin Zayed Species Conservation Fund (project number 0925331) and matching funds

from the Montgomery Botanical Center.

Some of the photographs were kindly provided by Greg Holzman (Figs. 3D-3F) and Bart Schutzman (Fig.

1A). David Knowles, Camilla Adair, Russell Adams, Sarah Gilmer, Louis Johnson, Javier Francisco-Ortega,

Alan Meerow, and Claudia Calonje participated in the held work resulting in the ex-situ collections studied

here. Javier Francisco-Ortega, Patrick Griffith, and Andrew Vovides reviewed an earlier version of the

manuscript.

REFERENCES

Broome, T. 2002. A celebration of red and brown emergent cycads. Cycad Newslett. 25:4-7.

Calonje, M., A.W. Meerow, L. Knowles, D. Knowles, M.P. Griffith, K. Nakamura, & J. Francisco-Ortega. 2013. Cycad biodiver¬

sity in the Bahamas Archipelago and conservation genetics of the threatened Zamia lucayana (Zamiaceae). Oryx

47:190-198.

Cycadlist.org. The world list of cycads. Accessed 03 Feb 2014. http://cycadlist.org

Meerow, A.W., J. Francisco-Ortega, M. Calonje, M.P. Griffith, T. Ayala-Silva, D.W. Stevenson, & K. Nakamura. 2012. Zamia (Cyca-

dales: Zamiaceae) on Puerto Rico: Asymmetric genetic differentiation and the hypothesis of multiple introductions.

Amer.J. Bot. 99:1828-1839.

Nicolalde-MorejOn, F., A.P. Vovides, & D.W. Stevenson. 2009. Taxonomic revision of Zamia in Mega-Mexico. Brittonia

61:301-335.

Norstog, K.J. &T.J. Nichols. 1997. The biology of the cycads. Cornell University Press, Ithaca, New York, U.S.A.

Osborne, R., M. Calonje, K.D. Hill, L. Stanberg, & D.W. Stevenson. 2012. The world list of cycads. Mem. New York Bot. Gard.

106:480-510.

Queenborough, S.A., M.R. Metz, R. Valencia, & SJ. Wright. 2013. Demographic consequences of chromatic leaf defence in

tropical tree communities: Do red young leaves increase growth and survival? Ann. Bot. 112:677-684.

Salas-Leiva, D.E., A.W. Meerow, M. Calonje, M.P. Griffith, J. Francisco-Ortega, K. Nakamura, D.W. Stevenson, C.E. Lewis, & S. Namoff.

2013. Phylogeny of the cycads based on multiple single-copy nuclear genes: Congruence of concatenated parsi¬

mony, likelihood and species tree inference methods. Ann. Bot. 112:1263-1278.

Schutzman, B. 1984. A new species of Zamia L. (Zamiaceae, Cycadales) from Chiapas, Mexico. Phytologia 55:299-303.

Taylor, A.S., J.L. Haynes, & G. Holzman. 2008. Taxonomical, nomenclatural and biogeographical revelations in the Zamia

skinneri complex of Central America (Cycadales: Zamiaceae). Bot. J. Linn. Soc. 158:399-429.

Taylor, A.S. & G. Holzman. 2012. A new Zamia species from the Panama Canal Area. Bot. Rev. 78:335-344.

Walters, T. 2003. Off-site collections. In: J. Donaldson, ed. Cycads: Status survey and conservation action plan. IUCN,

Gland, Switzerland and Cambridge, UK. Pp. 48-53.

LIMITATIONS TO NATURAL PRODUCTION OF LOPHOPHORA WILLIAMSII

(CACTACEAE) III. EFFECTS OF REPEATED HARVESTING AT TWO-YEAR

INTERVALS FOR SIX YEARS IN A SOUTH TEXAS (U.S.A.) POPULATION

Martin Terry

Sul Ross State University

Department of Biology

Alpine, Texas 79832, U.S.A.

Bennie Williams

Cactus Conservation Institute

RO. Box561

Alpine, Texas 79831, U.S.A.

Keeper Trout

Cactus Conservation Institute

P.O.Box 561

Alpine, Texas 79831, U.S.A.

Teodoso Herrera

Rio Grande Native American Church

P.O. Box460346

San Antonio, Texas 78246, U.S.A.

Norma Fowler

The University of Texas at Austin

Department of Integrative Biology C0930

205 W 24th St

Austin, Texas 78712, U.S.A.

ABSTRACT

Here we report the 6-year results of a long-term study of the effects of harvesting on a wild population of the cactus Lophophora william-

sii (peyote). Harvesting was performed using the best known technique: removing only the crown from the top of the plant. The two-year

interval between harvests was chosen because it was similar to that observed by persons who harvest peyote for legally protected religious

use by members of the Native American Church. Plants in the study were divided into three treatment groups: (1) control plants that were

never harvested, (2) plants that were harvested only once, at the beginning of the study, and (3) plants that were harvested at the beginning

of the study and every two years thereafter. Over the last two years of the study (2012-2014), the survival rate was significantly lower (77%)

in the plants harvested every two years than in the once-harvested plants (100%) and the unharvested control plants (98%). At the end of the

6th year of the study, average volume of living crown tissue per plant was significantly and substantially lower in the plants harvested every

two years than in the once-harvested plants and the unharvested controls. The average volume of once-harvested plants was 27% lower than

that of the controls, although this latter difference was not statistically significant. The modal number of crowns per plant varied with treat¬

ment and over time; in the plants harvested every two years it underwent a progression from 1 to 2 to 3 to 1 in response to successive har¬

vests. The results of this study indicate that a six-year recovery period, following the harvesting of peyote in natural habitats, is probably not

long enough to ensure long-term sustainability.

Keywords: cactus conservation, peyote harvest, cactus overharvesting, Native American Church, peyote conservation status

RESUMEN

Aqui reportamos los resultados de seis anos de un estudio a largo plazo sobre los efectos de cosechar individuos del cactus Lophophora wil-

liamsii (peyote) en una poblacion silvestre. La cosecha se hizo utilizando la mejor tecnica que se conoce, recogiendo solamente la corona de

la parte superior de la planta. El intervalo de dos anos entre cosechas fue similar al utilizado por personas que cosechan peyote para uso ju-

ridicamente protegido por los miembros de la Native American Church. Las plantas en el estudio fueron divididas en tres grupos de trata-

miento: (1) las plantas de control que nunca fueron cosechadas, (2) las plantas que fueron cosechadas solo una vez, al comienzo del estudio,

y (3) las plantas que fueron cosechadas al comienzo del estudio y posteriormente cada dos anos. En los ultimos dos anos del estudio (2012-

2014), la tasa de supervivencia fue significativamente menor (76,5%) en las plantas cosechadas cada dos anos, que en las plantas cosechadas

una sola vez (100%) y las plantas de control que no fueron cosechadas nunca (97,7%). Al final del sexto ano del estudio, el volumen promedio

del tejido vivo de la corona por planta fue significativamente menor en las plantas cosechadas cada dos anos, que en las plantas cosechadas

una sola vez y las plantas de control que nunca fueron cosechadas. El numero modal de coronas por planta vario con el tratamiento y con el

tiempo; en las plantas cosechadas cada dos anos hubo una progresion desde Ia2a3alen respuesta a las cosechas sucesivas. Los resultados

de este estudio indican que un periodo de recuperacion de seis anos despues de la recoleccion del peyote en el habitat probablemente no es

suficientemente largo como para asegurar la sostenibilidad a largo plazo.

J. Bot. Res. Inst. Texas 8(2): 541 - 550.2014

542

Journal of the Botanical Research Institute of Texas 8(2)

INTRODUCTION

Lophophora williamsii (Lem. ex Salm-Dyck) J.M. Coult. (Cactaceae), commonly known as peyote, is a small

globular cactus of northeastern Mexico and adjacent border areas of Texas. The crowns (apical chlorophyllous

portions of the stem) of these plants are approximately hemispherical in shape and generally protrude a few cm

above ground level. Some plants are caespitose, i.e., they have multiple crowns arising from a single nonchlo-

rophyllous (generally subterranean) stem. What was known of the biology of this species up to the mid-1990s

is summarized by Anderson (1996). Early suggestions that the species might be threatened by overharvesting

came from Morgan (1976), Anderson (1995), and Trout (1997).

The harvested crowns of peyote are collected and sold by licensed distributors to the Native American

Church (NAC) for religious use as protected by U.S. law. The anatomy of harvesting and the process of regen¬

eration of new crowns in response to harvesting are described by Terry and Mauseth (2006). It is now abun¬

dantly clear that the current rate of harvesting of peyote from wild populations is not sustainable (IUCN 2013;

NatureServe 2012; Terry et al. 2011, 2012). In March 2008 we began the first experimental investigation of the

effects of harvesting on peyote plants in situ. In previous papers we reported the effects that were detectable

two years (Terry et al. 2011) and four years (Terry et al. 2012) after the initial harvest. The present report fo¬

cuses on effects detectable six years after the initial harvest, in once-harvested plants and/or in plants har¬

vested every two years, the latter treatment representing the harvest frequency observed by persons who har¬

vest peyote for legally protected religious use by members of the Native American Church.

MATERIATS AND METHODS

The study site in the Tamaulipan thornscrub of South Texas was described previously by Terry et al. (2011).

The study design was described in detail by Terry et al. (2012). In summary, there were three treatment groups:

(1) a group of 50 plants which were unharvested control plants; (2) a group of 25 plants which were harvested

only once, at the beginning of the study; (3) a group of 25 plants which were harvested at the beginning of the

study and reharvested every two years thereafter. All plants were harvested using the best known technique,

removing only the crown from the top of the plant with a sharp knife. All plants in the study were individually

numbered and tagged in situ, along a transect through the population. Data collected on each plant at each

census (0, 2, 4 and 6 years) included number of crowns, number of ribs on each crown, and diameters of the

crowns. All statistical analyses were done with SAS 9.3 (SAS Institute, Cary, NC, USA).

Statistical analyses of survival. —Because a census of plants was conducted every two years (2008, 2010,

2012, 2014), each census interval was two years long. Harvesting was done directly after each census, and

therefore survival rates reflect the effects of treatments of preceding years, but not of the treatment of the cen¬

sus year. Survival rates were calculated for each census interval separately. Only plants present at the begin¬

ning of the given interval (and not dug up by feral hogs during the interval) were used to calculate survival

rates. Four plants were dug up by hogs during the first interval and were therefore dropped from all survival

calculations, and one plant was dug up during the third interval and was therefore dropped from the calcula¬

tion of survival rate during the third interval. Figure 1 is not an exact representation of cumulative survival

rates (which would have been affected by hog-caused mortality), but instead shows the products of the calcu¬

lated interval survival rates for each treatment (e.g., the proportion of plants surviving to census 3 is the prod¬

uct of the survival rate from census 1 to census 2, times the survival rate from census 2 to census 3).

A single survival rate was calculated for harvested plants for the first census interval (2008-2010), be¬

cause both groups of harvested plants were harvested after the first census. The “plants harvested once” were

not harvested again. Plants harvested once and plants harvested multiple times were separated in the analyses

of survival during the second (2010-2012) and the third (2012-2014) census intervals. Fisher’s exact test was

used to compare treatments within each of the three census intervals, because the expected values of some

cells were < 5, making ordinary % 2 tests inappropriate.

Statistical analyses of size. —Our primary measure of plant size was estimated total above-ground volume. It

Terry et al., Natural production of Lophophora williamsii

543

(D 40 -

••<>•

plants harvested every 2 years

— o

plants harvested once

•

plants never harvested

0 -I - 1 - 1 - T

2008 2010 2012 2014

census

Fig. 1. Cumulative proportions of plants remaining alive after excluding plants dug up by hogs from the calculations (see Methods). Plants were harvested

immediately after each census. Plants harvested once were harvested only after the first census (2008). For statistical tests, see Table 1.

was calculated from the diameter of each crown by assuming that each crown was a hemisphere: estimated

crown volume = % n (crown diameter/2) 3 . If a plant had multiple crowns in a given census, the estimated vol¬

umes of all of its crowns were summed to obtain estimated total above-ground volume of that plant. As is

usual with plant size measurements, the distribution of volume was skewed, with a right tail (i.e., a few large

plants, more smaller plants). No single transformation of total volume produced residuals that met the assump¬

tions of analysis of variance for all years. Instead, volumes from censuses in 2008 and 2012 were square-root

transformed before analysis, and volumes from censuses in 2010 and 2014 were log-transformed before analy¬

sis. (The log-transformation over-corrected skewness in the censuses of 2008 and 2012; the square-root trans¬

formation under-corrected it in the censuses in 2010 and 2014). Treatments were then compared with analysis

of variance (ANOVA). If the effect of treatment was significant and there were >2 treatments to be compared,

individual treatments were compared, using the Tukey adjustment for multiple testing. For graphical presenta¬

tion (Fig. 2), the mean, mean plus 1 standard error, and mean minus 1 standard error of each treatment were

separately back-transformed with the appropriate function, using the standard error of each treatment from

the corresponding ANOVA.

Another measure of size is the number of crowns per plant. When the experiment was initiated, each

experimental plant included in the groups to be harvested had exactly one crown. That constraint was not

imposed on the control plants, however, and the few multi-crowned individuals that occurred along the study

transect were not rejected from inclusion in the control group. This was done in order to ensure that the control

group was representative of the population in terms of the full range of crown numbers per individual plant.

The number of crowns/plant in each subsequent census was found to be Poisson distributed, and treatments

each year were therefore compared with a generalized linear model with a Poisson distribution for which the

544

Journal of the Botanical Research Institute of Texas 8(2)

census

plants never harvested

plants harvested once

plants harvested every 2 years

Fig. 2. Plant size, measured as total above-ground volume. Points represent back-transformed means; error bars are means ± 1 standard error, separately

back-transformed.

log is the link function. To supplement this analysis, we also calculated the change in number of crowns of

each surviving plant during each of the last two census intervals, and compared the distributions of size

changes among treatments with Fisher’s exact test.

RESULTS

Survival. —Survival rates of control plants remained high: more than 95% in each two-year census interval

(Table 1, Fig. 1). Survival rates of plants harvested every two years were consistently lower than those of control

plants, and decreased over time; the rate at which these plants survived the third census interval, by which

time they had been harvested three times, was only 77%. There is evidence that the negative effect of harvest¬

ing on survival lasts more than two years: survival rates of once-harvested plants did not return to control

levels until four years had passed after they were harvested (i.e., until the third census interval, 2012-2014).

Size. —The once-harvested plants appeared to continue to be in the process of recovering from the adverse ef¬

fects of being harvested six years before (Table 2, Fig. 2). Six years after harvesting, we no longer were able to

detect a significant difference in size (total above-ground volume) between never-harvested and once-harvested

Terry et al., Natural production of Lophophora williamsii

545

Table 1 . Survival rates during each census interval. Plants were harvested immediately after each census; plants harvested once were harvested only after the first

census. NS, not significant.

2008-2010

2010-2012

2012-2014

control plants

97.9%

95.7%

97.7%

plants harvested once

89.6%

82.6%

100%

plants harvested every two years

89.6%

85%

76.5%

P = 0.013

P = 0.0097

sample size

N = 96

N = 89

N = 80

Table 2. Results of ANOVAs comparing plant volumes among treatments. All harvests took place immediately after the census in the calendar year indicated. Analyses

for census years 2008 and 2010 compare unharvested and harvested plants; the first harvest (2008) occurred after the first census. Analyses for census years 2012

and 2014 compare unharvested plants, plants harvested once (after the 2008 census), and plants harvested every two years (i.e., after the censuses of 2008,2010,

and 2012). Control, never harvested; once, harvested once after the first census; multi, harvested every two years after each census.

census year

transformation used

treatment

multiple comparisons

2008

square root

^95 = 0.59

control v once

control v multi

once v multi

2010

log

F, ,88 = 58.31

< 0.0001

— a

2012

square root

f 2i76 = 28.73

< 0.0001

<0.0001

0.13

2014

log

F 2 , 7 , =45.29

< 0.0001

0.31

<0.0001

a no multiple comparisons needed; treatment P value compares control and harvested plants

plants. This of course does not mean that the once-harvested plants had completely recovered, especially given

that small sample sizes reduced the power of our test. A power analysis of our results indicates that, given the

average control plant size of 48 cm 3 in the 2014 census and the observed distribution of plant sizes in that cen¬

sus, we could have detected a significant difference between the control and the once-harvested plants only if

the once-harvested plants had been < 28 cm 3 in size on average (42% smaller); in fact, they were 35 cm 3 in size

on average (only 27% smaller). In contrast, the plants harvested every two years remained very small. We were

not able to detect a temporal trend in the volume of those plants, perhaps because very small plants simply

died.

As the experiment proceeded, the distribution of crown numbers in control (i.e., never harvested) plants

remained quite constant (Figs. 3 and 4), with the average number of crowns per control plant remaining close

to 1.7 (Table 3). The majority of control plants always had one crown, but note the long right tails of the distri¬

butions in Fig. 3, each of which includes a few control plants with five or more crowns each.

The first harvest (2008) caused a significant increase in the number of crowns per harvested plant, with a

modal value of 2 crowns per plant and a mean of 2.58 (Table 3, Fig. 3). (However, these new crowns were quite

small; recall that harvesting reduced average plant volume substantially). Over the next four years plants that

were not re-harvested continued to have a modal value of 2 crowns per plant, with little change in their distri¬

bution (Figs. 3 and 4).

The second harvest of plants harvested every two years shifted the modal value to 3 crowns per plant

(mean 2.56) in 2012, which was more crowns/plant than the once-harvested plants in the same census had

(mode 2, mean 2.16 crowns/plant) (Table 3; Fig. 3, census 2012). However, 31% (5 of 16) of plants harvested

every two years decreased in crown number during this period (Fig. 4,2010-2012). The statistical test compar¬

ing crown number distributions among treatments in 2012 did not quite reach significance (Table 3: P = 0.09),

nor did the statistical test comparing changes in crown number (P = 0.18).

The third harvest of plants harvested every two years reduced the average number of crowns per plant in

2014, and one crown per plant was once again the most common value in these plants (Fig. 3, census 2014,

percentage of plants percentage of plants percentage of plants

546

Journal of the Botanical Research Institute of Texas 8(2)

crowns/plant

Fig. 3. Distributions of numbers of crowns per plant. For statistical tests, see Table 3.

Terry et al., Natural production of Lophophora williamsii

547

change in number of crowns

i i plants never harvested

ix- w i plants harvested once

plants harvested every two years

change in number of crowns

Fig. 4. Distributions of changes in the number of crowns per plant.

plants harvested every two years). Of plants harvested every two years, 58% (7 of 12) had fewer crowns at the

end of this census interval (2014) than they did at its beginning (2012) (Fig. 4, 2012-2014, plants harvested

every two years). While the differences in crown number among treatments did not reach statistical signifi¬

cance in 2014 (Table 2 :P = 0.31), the statistical comparison of the changes in crown number (Fig. 4,2012-2014)

among treatments was highly significant (P < 0.0001).

DISCUSSION

During the fifth and sixth years of this ongoing experimental study of a peyote population, we saw few chang¬

es in the behavior of never-harvested (control) plants. Once-harvested plants continued to recover from har¬

vesting, but plants harvested every two years continued to experience lowered survival rates and to be quite

548

Journal of the Botanical Research Institute of Texas 8(2)

Table 3. Numbers of crowns per plant.

comparison of treatments

average no. of

crowns/plant

EM)

census year 2010

8.06(1,88)

0.006

control

1.70

harvested

2.58

census year 2012

2.46 (2,77)

0.09

control

1.69

once-harvested

2.16

harvested every 2 yrs

2.56

census year 2014

1.19(2,71)

0.31

control

1.74

once-harvested

2.33

harvested every 2 yrs

2.08

small, but for the first time also showed a reduced capacity to produce new crowns. The survival rate and indi¬

vidual sizes of never-harvested plants indicate that the site continued to be suitable for this species. However,

the loss of plants to digging, apparently by feral hogs, did not cease; it continued to be a concern.

Effects of harvesting.—The recovery of once-harvested plants from their harvest in 2008 appeared to be

nearly complete in terms of survival: their survival from 2012 to 2014 was not different from that of the never-

harvested plants. This delayed but marked increase in the survival rate of the once-harvested group attests to

the resilience of the species when it is allowed a long enough respite from further harvesting. But note that this

recovery was delayed and occurred only in the third two-year period after harvesting; harvesting reduced

survival for at least four years.

By our best measure of size, the estimated above-ground volume of each plant, the once-harvested plants

also continued to recover in terms of size (Fig. 2). However, inspection of Fig. 2 suggests that this recovery was

not yet complete in 2014. Although the difference in size between never-harvested and once-harvested plants

was no longer significant in 2014, this does not mean that the two groups did not differ. A power analysis of our

data indicated that, due to plant-to-plant variability and small sample sizes, only differences of 42% or more

from average control plant size would have been detected as statistically significant. In other words, the rela¬

tively low power of the analysis made a type II error (false negative, i.e., failure to reject the null hypothesis)

very likely as soon as the once-harvested plants were about half the size of the never-harvested plants. If we had

been able to design the study with a total of 500 plants instead of 100 (which would have implied finding a

landowner who was willing to allow 250 peyote plants to be harvested from his property), we would have had

substantially greater statistical power, which might well have resulted in a P value less than 0.05 for the com¬

parison of control and once-harvested plants. The pattern evident in Fig. 2, these statistical considerations, and

the precautionary principle, strongly support the assumption that six years is not long enough to wait after a

population of peyote is harvested before going back and re-harvesting the population, if the goal is a sustain¬

able regimen of harvesting.

In contrast to the ongoing recovery of once-harvested plants, the plants harvested every two years contin¬

ued to die at significantly higher rates during 2012-2104, and the survivors remained extremely small. All evi¬

dence continues to support the conclusion that harvesting this species every two years, even using best-prac¬

tice harvesting methods, is not sustainable.

Can the increase in crown number compensate for the negative effects of harvesting?—If the death of one

crown on a plant is independent of the death of another crown on the same plant, then having more crowns

would increase the plant’s chance of survival. It is not clear what mechanisms would create this scenario, be¬

cause both resource competition and herbivory likely affect all the crowns of a plant at once. However, it is

possible that this scenario was in part the reason that the survival rate of once-harvested plants recovered from

Terry et al., Natural production of Lophophora williamsii

549

harvesting before their volume did. In any case, the multiple crowns of the repeatedly-harvested plants did not

prevent their survival rate from remaining lower than that of control plants.

By 2014 the number of crowns of the repeatedly-harvested plants had declined from its peak in 2012,

while the above-ground volume of these plants remained extremely low. It seems likely that these repeatedly-

harvested plants had exhausted most of their stored resources by 2014 and therefore their ability to regrow

following harvest was reduced. Presumably each harvest used up stored carbohydrates and other resources, as

well as reducing the length of time during which the plant could accumulate new resources via photosynthe¬

sis. Viable areoles on the nonchlorophyllous/subterranean portion of the stem are probably also depleted by

repeated regrowth after harvesting. Such areoles are not replaced, as they are generated early in the life of the

plant at the apical meristem and migrate radially as the plant grows, ultimately becoming incorporated into the

basal (non-chlorophyllous) portion of the stem, below the crown. Once such an areole is used up in producing

regrowth in the form of an axillary branch with its new crown, that areole is permanently extinguished by the

harvesting of that crown.

Management implications. —The continuing decline of the plants harvested every two years clearly and

strongly supports not harvesting wild-grown peyote plants this frequently. As we argued in a previous paper

(Terry et al. 2012), frequent harvests are producing a classic case of the overharvesting of a wild population.

We also conclude that even six years is probably not long enough for plants to recover from harvesting,

even using best-practice harvesting methods (clean knife cut, leaving the non-chlorophyllous stem and roots

intact). It is possible that eight years between harvests would allow recovery; we do not have data to address

this hypothesis as yet. However, for the purpose of considering conservation options, let us make the assump¬

tion that eight years would be sufficient for plants to completely recover from a single harvest of their crowns

by the current best-practice method. We have heard several objections to the proposition that cultivation of

peyote is the single most viable solution to the current problem of reduced availability of peyote. One of the

most frequently heard objections is that “10 years is too long” to wait for peyote planted from seed to reach

maturity in a greenhouse so that it would be available for harvesting for ceremonial use by the Native American

Church. But if that 10-year waiting period for sustainable production of cultivated peyote is now compared to

an 8-year waiting period required for sustainable production of peyote in its natural habitat, perhaps the 10-

year wait for greenhouse cultivation to come on line to allow adequate annual production for the Church’s

ceremonial needs is not so unreasonable after all. A greenhouse-grown supply of peyote would also eliminate

all the uncertainty of relying on collection from the wild and greatly reduce the real possibility that this species

will soon become commercially extinct in the wild, that is, so rare that it is no longer economically feasible to

collect wild plants.

ACKNOWLEDGMENTS

We thank C.W. Hellen Ranches, Ltd. - La Mota Division - Charles W. (Bill) Hellen, Managing Partner, for

providing access to a site where a long-term study such as this can be carried out safely. Essential funding for

the study was generously provided by the Alvin A. and Roberta T. Klein Foundation, Sul Ross State University

(Research Enhancement grant and use of the SRSC herbarium), and all the donors supporting the scientific

work of the Cactus Conservation Institute, Inc. We thank A. Michael Powell and Billie L. Turner for prompt

and careful reviews.

REFERENCES

Anderson, E.F. 1995. The "Peyote Gardens" of South Texas: A conservation crisis? Cact. Succ.J. (U.S.) 67:67-73.

Anderson, E.F. 1996. Peyote: The divine cactus. University of Arizona Press, Tucson, U.S.A.

International Union for the Conservation of Nature (IUCN). 2013. IUCN Red List of threatened species. Lophophora williamsii

assessment: Vulnerable, www.iucnredlist.org/details/151962/0 (Accessed: 6 December 2013.)

Morgan, G.R. 1976. Man, plant, and religion: Peyote trade on the Mustang Plains of Texas. Ph.D. Thesis, University of

Colorado.

550

Journal of the Botanical Research Institute of Texas 8(2)

NatureServe. 2012. NatureServe Explorer: An online encyclopedia of life [web application]. Version 7.1. NatureServe,

Arlington, Virginia. Available at http://www.natureserve.org/explorer (Accessed: July 3, 2012.)

Terry, M. & J.D. Mauseth. 2006. Root-shoot anatomy and post-harvest vegetative clonal development in Lophophora wil-

liamsii (Cactaceae: Cacteae): limplications for conservation. Sida 22:565-592.

Terry, M., K. Trout, B. Williams, T. Herrera, & N. Fowler. 2011. Limitations to natural production of Lophophora williamsii

(Cactaceae) I. Regrowth and survivorship two years post harvest in a South Texas population. J. Bot. Res. Inst. Texas

5:661-675.

Terry, M., K. Trout, B. Williams, T. Herrera, & N. Fowler. 2012. Limitations to natural production of Lophophora williamsii

(Cactaceae) II. Effects of repeated harvesting at two-year intervals in a South Texas population. J. Bot. Res. Inst. Texas

6:567-577.

Trout, K. 1997. Sacred cacti. Narayan Publications, Sedona, Arizona, U.S.A.

DISTIGOUANIA IRREGULARIS (RHAMNACEAE) GEN. ET SR NOV.

IN MID-TERTIARY AMBER FROM THE DOMINICAN REPUBTIC

Kenton L. Chambers

George O. Poinar, Jr.

Department of Botany and Plant Pathology

Oregon State University

Corvallis, Oregon 97331, U.S.A.

chamberk@science.oregonstate.edu

Department of Integrative Biology

Oregon State University

Corvallis, Oregon 97331, U.S.A.

poinarg@science.oregonstate.edu

ABSTRACT

Distigouania irregularis (Rhamnaceae) is described as a new genus and species of fossil angiosperm from deposits of Mid-Tertiary amber

in the Dominican Republic, on the Caribbean island of Hispaniola. The fossil is represented by a single staminate flower at anthesis and

displays an irregular corolla in which 4 petals are rhombic-lanceolate, spreading, and sepal-like, while the 5th is shorter, erect, and slightly

cupped. The stamens are dimorphic; the 4 that are opposite the spreading petals have dehisced anthers on erect filaments that diverge from

the petals, while the 5th stamen adjoins the erect, cupped petal and has a larger, unopened anther. The hypanthial disc is densely pubescent

with tangled trichomes, and on the disc margin, at the base of each sepal, is an enlarged, bilobed, probably glandular appendage. There are

minute papillate trichomes on the disc appendages and on the epidermis of the anthers. These epidermal papillae, as well as the pubescent

disc and bilobed appendages, are similar to features of staminate flowers in various extant American species of Gouania. However, the fossil

is unique within family Rhamnaceae in its irregular corolla and androecium, and it stands out as an evolutionary experiment with zygomor-

phy in this otherwise actinomorphic family.

RESUMEN

Distigouania irregularis (Rhamnaceae) se describe como genero y especie nuevos de angiosperma fosil de los depositos de ambar del Ter-

ciario medio de la Republica Dominicana, en la isla caribena de La Espanola. El fosil esta representado por una flor simple estaminada en

antesis que muestra una corola irregular cuyos cuatro petalos son rombico-lanceolados, patentes, y semejantes a sepalos, mientras que el

quinto es mas corto, erecto, y ligeramente acopado. Los estambres son dimorficos; los 4 que estan opuestos a los petalos desplegados tienen

anteras abiertas en filamentos erectos que divergen de los petalos, mientras que el quinto estambre se acerca al petalo erecto acopado, y tiene

una antera mas grande no abierta. El disco hipantico es densamente pubescente con tricomas enmaranados, y en el margen del disco, en la

base de los sepalos, esta un apendice ensanchado bilobulado, probablemente glandular. Hay tricomas con papilas diminutas en los apendi-

ces del disco y en la epidermis de las anteras Estas papilas epidermicas, asi como el disco pubescente y los apendices bilobulados, son simil-

ares a las caracteristicas de las flores estaminadas de varias especies americanas actuales de Gouania. Sin embargo, el fosil es unico en la fa-

milia Rhamnaceae por su corola irregular y androceo, y se mantiene como un experimento evolutivo con la zigomorfia en esta familia acti-

nomorfica por lo demas.

INTRODUCTION

Amber mines in the mountains of central Hispaniola have yielded numerous fossil flowers from Mid-Tertiary

tropical forests in the Caribbean region. The present flower is the 16th species to be described from these de¬

posits. Others are 1 species of Fabaceae (Poinar 1991), 3 of Arecaceae (Poinar 2002a, 2002b), 2 of Poaceae

(Poinar &Judziewicz 2005; Poinar & Columbus 2012), 1 of Chrysobalanaceae (Poinar et al. 2008a; revised by

Chambers & Poinar 2010), 2 of Lauraceae (Chambers et al. 2011a, 2012), 3 of Meliaceae (Chambers et al. 2011b;

Chambers & Poinar 2012), 1 of Burseraceae (Chambers & Poinar 2013), 1 of Myristicaceae (Poinar & Steeves

2013), and 1 possibly of Moraceae (Poinar et al. 2008b). The moist forests of this period have been described by

Poinar and Poinar (1999), based on numerous amber-embedded insect species together with whatever plant

fossils were then available. The resin that trapped these organisms was produced by Hymenaeaprotera Poinar

(Poinar 1991), a progenitor of the modern-day forest species H. courharil L. Fossils of Rhamnaceae, principally

leaves and fruits from the Late Cretaceous and Early Cenozoic, have been the object of numerous studies (see

references in Jones & Dilcher 1980; Burge & Manchester 2008; Correa et al. 2010) and have contributed to

phylogenetic and historical phytogeographic analyses of the family (Richardson et al. 2000a, 2000b, 2004).

J. Bot. Res. Inst. Texas 8(2): 551 - 557.2014

552

Journal of the Botanical Research Institute of Texas 8(2)

MATERIALS AND METHODS

This and the other fossils referred to above were collected from amber mines in the Cordillera Septentrional,

between Puerto Plata and Santiago, Dominican Republic. Different dates have been ascribed to the marine

sediments in which the amber is found. A younger proposed age of 20-15 Ma is based on foraminifera present

in the deposits (Iturralde-Vinent & MacPhee 1996). An older age of 45-30 Ma is based on marine coccoliths

(Cepek in Schlee 1999). The amber is in turbiditic sandstones of the Upper Eocene to Lower Miocene Mamey

Group (Draper et al. 1994). Examination and photographs were made with a Nikon stereoscopic microscope

SMZ-10-R at 80x and a Nikon Optiphot microscope at 800x.

DESCRIPTION

Distigouania K.L. Chambers & Poinar, gen. nov. (Figs. 1-5) .Type species: Distigouania irregularis K.L. Chambers & Poinar,

sp. nov.

Flower staminate, pedicel glabrous, hypanthium dish-shaped, glabrous (Fig. 5), calyx radially symmetrical,

sepals 5, equal, spreading (Fig. 1), corolla irregular, with 4 spreading, petals, not cupped or hooded, and 1

shorter, cupped, erect petal (Figs. 1, 3, 4), stamens dimorphic, the 4 opposite the spreading petals erect on

slender filaments, with dehisced, bilocular anthers, stamen opposite the cupped petal with the anther larger, ±

appressed to petal, indehiscent or possibly tardily dehiscent, disc flat, densely pubescent with tangled tri-

chomes (Fig. 2), appendages of disc 5, enlarged, bilobed, each opposite the base of a sepal (Fig. 2), epidermis of

disc appendages and anthers minutely papillate, style none. Pistillate flower unknown.

Etymology. —From Latin “disto,” to stand apart, be separate, and Gouania, a genus of Rhamnaceae.

Distigouania irregularis K.L. Chambers & Poinar, sp. nov. Type: HISPANIOLA. Dominican Republic: amber mine in the

northern mountain ranges (Cordillera Septentrional), 1998, unknown amber miner s.n. (holotype: catalogue number Sd-9-178, de¬

posited in the Poinar amber collection maintained at Oregon State University, Corvallis, Oregon 97331, U.S.A.)

Sepals glabrous, ovate, 0.9-1.2 mm long, 0.7-0.8 mm wide, tip acute to acuminate, not thickened, midline

ridged (Figs. 1, 2), corolla glabrous, the 4 spreading petals rhomboid-lanceolate, 0.7-0.9 mm long, 0.4-0.5 mm

wide, acute (Fig. 1), 5th petal obovate, obtuse, ca. 0.4 mm long, 0.35 mm wide, slightly cupped (Figs. 1, 3, 4),

filaments of 4 erect, dehiscent stamens 0.014-0.027 mm long, valves of dehisced anthers 0.023-0.030 mm

long, 0.013-0.018 mm wide (Figs. 2,3), undehisced anther valves of 5th stamen ca. 0.032 mm long, 0.022 mm

wide (Fig. 3), filament of undehisced anther ca. 0.015 mm, disc appendages 0.025-0.028 mm wide, 0.018-

0.022 mm long (Fig. 2), pedicel 1.2 mm long.

Etymology. —From Latin “irregularis,” divisible into 2 equal halves along one plane only.

DISCUSSION

Our original comparison of the fossil was with Gouania, a pantropical genus of ca. 50 species (Medan & Schi-

rarend 2004; Pool 2014). Examination of published illustrations and a limited sample of herbarium specimens

has not shown any examples of irregular corollas or anthoecia, nor do standard descriptions of the genus men¬

tion these floral features. In such a large genus, it is not inconceivable that irregular flowers may have evolved

in some extant species; however, if present, they seem to have escaped notice. With respect to a shallow hypan¬

thium, small petals cupping the anthers, a flat, pubescent disc, bilobed disc appendages, and lack of a style in

staminate flowers, the fossil resembles Gouania taxa such as G. hypoglauca Standi., a Central American species.

A review of Gouania in the Caribbean, Mexico, and Central America was recently published by Pool (2014),

giving a basis for comparing Distigouania with modern species of this region. Treatments of the Rhamnaceae

as a whole (e.g., Sussenguth 1953; Medan & Schirarend 2004) describe the family as having radially symmetri¬

cal flowers. This fossil is so unusual in its floral structure, therefore, that we believe it merits description as a

separate genus.

A feature of the flower not yet remarked on is the strong similarity of the 4 laterally spreading petals to the

sepals. Except for being smaller, these petals are like the sepals in their shape and textural appearance, as fos¬

silized. Their color, in life, may also have been sepal-like. A further similarity is the presence of a partial mid-

Chambers and Poinar, Distigouania irregularis from Mid-Tertiary amber

553

: V

Fig. 1. Distigouania irregularis. Apical view of flower. Upper arrow is on sepal, middle arrow indicates one of the 4 spreading petals, lower arrow shows

erect, cupped petal. Note ridge on sepal midline. Scale bar = 0.7 mm.

Fig. 2. Distigouania irregularis. Enlarged apical view. Long black arrow indicates tangled trichomeson hypanthial disc, 2 short black arrows are on bilobed

disc appendages, white arrows point to 2 dehisced anthers. Scale bar = 0.49 mm.

554

Journal of the Botanical Research Institute of Texas 8(2)

Fig. 3. Distigouania irregularis. Apical view. Long arrows are on dehisced anthers, short arrow, top middle, points to undehisced anther, shortest arrow,

top right, indicates cupped petal subtending this anther. Scale bar = 0.675 mm.

Fig. 4. Distigouania irregularis. Lateral view. Arrow indicates the erect petal, which is cupped and subtends the stamen with undehisced anther. Scale

bar = 0.26 mm.

Chambers and Poinar, Distigouania irregularis from Mid-Tertiary amber

555

Fig. 5. Distigouania irregularis. Basal view. Right arrow is on pedicel, upper left arrow indicates location of 5th, non-spreading petal, lower arrow is on

hypanthium. Scale bar = 0.736 mm.

line ridge, visible in the proximal half of petals in the 2 o’clock and 4 o’clock positions in Figure 1. The 5th

petal, erect and cupped against the undehisced anther (Fig. 1, lower arrow, Fig. 4) is like the limb of a typical

petal of Rhamnaceae in size, shape, and texture. At a developmental- genetic level we speculate that the sepal-

oid appearance of 4 of the second whorl organs could result from the partial loss of expression of B class genes

in this whorl (from the ABC model of flower development, see Bowman et al. 1989; Theissen et al. 2000), as

occurs in a variety of extant taxa (e.g., Kramer et al. 2003). Additionally, loss of radial symmetry and petal

showiness could result from changes in the expression pattern of flower symmetry genes (Preston et al. 2011).

Therefore, if extant members of Rhamnaceae with a similar phenotype were found, it could lead to interesting

comparative studies of the genetics of candidate genes for flower development, such as orthologs of PISTILLA-

TA, APETALA3, CYCHLOIDEA and DICHOTOMA. We might then come closer to explaining the morphology

of this unusual Mid-Tertiary fossil.

If allied to Gouania, as hypothesized, Distigouania is a member of Tribe Gouanieae (Richardson et al.

2000a:333; Medan & Schirarend 2004:330). In a molecular phylogenetic analysis of Rhamnaceae using the

plastid genes rbcE and trnE-F, Richardson et al. (2000b) found 3 well-supported clades; however, these could

not be characterized morphologically, at least with presently available data. They were given the informal

names of the rhamnoid clade, the ampeloziziphoid clade, and the ziziphoid clade. Gouanieae belong to the zizi-

phoid clade as a well-supported, monophyletic group, consisting of Crumenaria, Gouania, Helinus, Pleurantho-

des, and Reissekia. In their discussion of the biogeography of Rhamnaceae, these authors propose a phylogeny

extending back to the Mid-Cretaceous with possible relationships to the Rose Creek flower of Basinger and

Dilcher (1984), which may establish 94-96 Ma as the minimum age of the family. 1 For some genera, at least, a

Assigning the Rose Creek flower to Rhamnaceae is not accepted by some other authorities, however (Calvillo-Canadell & Cevallos-Ferriz 2007:1667).

556

Journal of the Botanical Research Institute of Texas 8(2)

widespread Old and New World distribution may be attributed to their ancestors’ initial presence in Gondwa-

naland prior to its breakup. The occurrence of modern genera of Rhamnaceae in the late Cretaceous has not yet

been firmly established in the fossil record. Flowers of this family, not specifically assignable to an extant ge¬

nus, were discovered in Mexican strata of probable late Campanian age, ca. 74 Ma (Calvillo-Canadell & Ceval-

los-Ferriz 2007). Upper Maastrichtian leaves of Berhamniphyllum (Jones & Dilcher 1980; Correa et al. 2010)

are similar to those of Rhamnidium and Berchemia , while the winged fruit of Archeopaliurus (Correa et al. 2010)

compares well with that of Paliurus. Taking into account both fossil and molecular evidence, Calvillo-Canadell

and Cevallos-Ferriz (op. cit.) propose an Eocene/Oligocene origin for genera such as Gouania, Ziziphus,

Berchemia , Karwinskia, and Paliurus.

The present fossil, from the Oligocene or early Miocene, stands out as an evolutionary experiment with

zygomorphic flowers in this otherwise actinomorphic family. Because floral zygomorphy is frequently associ¬

ated with insect pollination, we suggest that the notably enlarged, bilobed appendages of the hypanthial disc

may have been nectariferous, supplying a reward for insect visitors. The function of the undehisced anther,

which stayed closed while the other 4 anthers opened and released their pollen, remains unexplained. Given

the absence of living relatives having a comparable floral syndrome, we choose not to speculate on whether, in

this species, it also may have had a particular function in insect pollination.

ACKNOWLEDGMENTS

We thank the herbaria of Harvard University, the New York Botanical Garden, and the Missouri Botanical

Garden for lending us specimens for this study. Amy Pool, Richard Abbott, and Barbara Rye provided useful

observations and suggestions. Veronica Di Stilio was our consultant in developmental floral genetics and

helped to prepare the paragraph on that subject in the Discussion section.

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Poinar, G.O., Jr., K.L. Chambers, & A.E. Brown. 2008b. Trochanthera lepidota gen. and sp. nov., a fossil angiosperm inflores¬

cence in Dominican amber. J. Bot. Res. Inst.Texas 2:1 1 67-1 1 73.

Poinar, G.O., Jr. & J.T. Columbus. 2012. Alarista succina gen. et sp. nov. (Poaceae: Bambusoideae) in Dominican amber.

Historical Biol. 1-6.

Poinar, G.O., Jr. & R. Steeves. 2013. Virola dominicana sp. nov. (Myristicaceae) from Dominican amber. Botany 91:530-534.

Pool, A. 2014. Taxonomic revision of Gouania of North America. Ann. Missouri Bot. Gard. 99:490-552.

Preston, J.C., C.C. Martinez, & L.C. Hileman. 2011. Gradual disintegration of the floral symmetry gene network is implicated

in the evolution of a wind-pollination syndrome. Proc. Natl. Acad. Sci. 108:2343-2348.

Richardson, J.E., M.F. Fay, Q.C.B. Cronk, & M.W. Chase. 2000a. A revision of the tribal classification of Rhamnaceae. Kew Bull.

55:311-340.

Richardson, J.E., M.F. Fay, Q.C.B. Cronk, D. Bowman, & M.W. Chase. 2000b. A phylogenetic analysis of Rhamnaceae using rbcL

and trnL-F plastid DNA sequences. Amer. J. Bot. 87:1309-1324.

Richardson, J.E., L.W. Chatrou, J.B. Mols, R.HJ. Erkens, & M.D. Pirie. 2004. Historical biogeography of two cosmopolitan fami¬

lies of flowering plants: Annonaceae and Rhamnaceae. Phil. Trans. Roy. Soc., B, Biol. Sci. 359:1495-1508.

Schlee, D. 1999. Das Bernstein-Kabinett. Stuttgarter Betr. Naturk. Ser. C, 28.

Sussenguth, K. 1953. Rhamnaceae. In: H. Melchior & E. Werdermann, eds. Engler & Prantl's Die Naturlichen Pflanzenfami-

lien, Band 20d, Duncker & Humblot, Berlin, Germany. Pp. 7-173.

Theissen, G., A. Becher, A. Di Rosa, A. Kanno, J.T. Kim, T. Munster, K-U. Winter, & H. Saedler. 2000. A short history of MADS-box

genes in plants. PI. Molec. Biol. 42:115-149.

558

Journal of the Botanical Research Institute of Texas 8(2)

BOOK REVIEW

Roy L. Lehman. 2013. Marine Plants of the Texas Coast. (ISBN-13: 978-1-62349-016-4, flexbound). Texas

A&M University Press, 4354 TAMU, College Station, Texas 77843-4354, U.S.A. (Orders: www.tamu-

press.com, 1-800-826-8911). $32.00, 205 pp., 264 color photos, map, bibliography, index, 6" x 9".

This 205-page book is an important resource for amateur plant enthusiasts, biologists, conservationists, stu¬

dents, and others with a special interest in coastal ecosystems and natural resources. It includes narrative de¬

scriptions, color photos, and specific information on a total of 140 individual species representing 81 genera

from 52 families.

A logical, well organized, straightforward, and user-friendly key provides a useful tool to aid in identifica¬

tion of the plants included in the book. One of the details that I found most helpful and practical to a broad

audience is that all measurements are listed first in English increments and followed by the metric equivalent

in parentheses. The book is organized into four distinct plant groupings: shoreline plants, seagrasses, man¬

groves, and seaweeds.

The section on shoreline plants includes a small number of the most significant/common grasses, sedges,

rushes, and forbs that provide a buffer between the tidal waters and the coastal marshes. Many of these are

halophytes (salt-tolerant species) that survive and thrive in the hostile environment of the Texas coast where

they are subject to high salinity, periodic inundation, wind and wave action, and harsh xerophytic conditions

at times.

The seagrass section contains information about some of the most common seagrasses found in the shal¬

low waters of the coast. In reality, these are not grasses; rather they are submerged aquatic species that have a

similar appearance to terrestrial grasses and sedges.

Mangroves are woody species with specialized structural and seed dispersal properties that occur in the

muddy tidal flats and shallow tidal waters. They perform many important functions including stabilization of the

soil. They also trap debris, filter runoff, and prevent terrestrial organic matter from reaching the coastal waters.

Mangroves and seagrasses serve as nurseries and essential habitat for fish, shellfish, and aquatic invertebrates.

The last half of the book is dedicated entirely to seaweeds (macroalgae) of the Texas coastal waters. This

section includes 86 species and 45 genera and is subdivided into three subcategories based on color: red,

brown, or green. While I have no knowledge and little interest in seaweed, this is apparently one of the most

comprehensive discussions of that subject that has been published in a book intended for general use by a wide

audience.

The appendices include a glossary and a section on plant collection and preparation techniques. I found

the glossary to be very helpful for those, like me, who are not familiar with botanical terminology that is spe¬

cific to aquatic vegetation and ecosystems. The discussion of plant collection and preparation techniques

would be especially useful to those who are collecting and preparing specimens of marine plants. And the

color photographs are an interesting and helpful addition to the information provided for each species.

The sections on seagrasses and mangroves were appropriate and provided adequate information on those

plant categories. I would have preferred the book to include more species of the major shoreline plants, par¬

ticularly the grasses and sedges, and much less coverage of the seaweeds. However, this book would be a valu¬

able tool for anyone with an interest in the vegetation of the Texas coast .—Dan Caudle, Independent Grassland

Consultant, Weatherford, Texas, U.S.A.

J.Bot. Res. Inst. Texas 8(2): 558.2014

TICODENDRON PALAIOS SP. NOV. (TICODENDRACEAE),

A MID-TERTIARY FOSSIL FLOWER IN DOMINICAN AMBER

Kenton L. Chambers

George O. Poinar, Jr.

Department of Botany and Plant Pathology

Oregon State University

Corvallis, Oregon 97331, U.S.A.

chamberk@science.oregonstate.edu

Department of Integrative Biology

Oregon State University

Corvallis, Oregon 97331, U.S.A.

poinarg@science.oregonstate.edu

ABSTRACT

Ticodendron palaios is described as a new species of fossil flower preserved in amber from Mid-Tertiary deposits in the Dominican Repub¬

lic. The genus, newly described in 1989 (Gomez-Taurito & Gomez-P. 1989), is arborescent and limited to regions of wet primary forests in

Central America, ranging from southern Mexico to Panama. It was made a separate family, Ticodendraceae, by Gomez-Taurito and Gomez-

P. (1991) and assigned to the hamamelidalian order Fagales. The fossil species is represented by a single pistillate flower and consists of a

partial inflorescence with 1 short-pedicelled, inferior ovary subtended by 4 large bracts and surmounted by 3 short sepals and 2 papillate

styles. The sepals, ovary, and pedicel are densely hispid-setose. The flower shows an overall resemblance to the original species, T. incogni-

tum, with some important differences, one being that its styles, instead of being extended, are appressed to the ovary. However, this could be

due to the flower’s position at the edge of the resin exudate before it was fossilized.

RESUMEN

Se describe Ticodendron palaios como una nueva especie fosil de flor preservada en ambar en depositos del Terciario medio en la Republi-

ca Dominicana. El genero, descrito en 1989 (Gomez-Taurito & Gomez-P. 1989), es arborescente y limitado a las regiones de bosques pri¬

maries humedos en America Central, que van del sur de Mexico a Panama. Se hizo una nueva familia, Ticodendraceae, por Gomez-Taurito

y Gomez-P. (1991) y se assigno al orden Fagales de las hamamelidas. Ta especie fosil esta representada por una flor simple pistilada y consiste

en una inflorescencia parcial con un ovario infero cortamente pedicelado, subtendido por 4 grandes bracteas y sobremontado por tres cortos

sepalos y dos estilos 2 papilados. Eos sepalos, ovario, y pedicelo son densamente hispido-setosos. Ta flor muestra un parecido general con la

especie original, T. incognitum, con algunas diferencias importantes, una son sus estilos, que en vez de ser patentes, son apresados al ovario.

Sin embargo, esto puede estar debido por la posicion de la flor en el borde del exudado de resina antes de fosilizarse.

INTRODUCTION

Dominican amber is obtained from mines in the mountains of central Hispaniola and contains fossilized plant

materials and animals, especially insects, which make it sought after by collectors. The marine strata contain¬

ing the amber have been placed in the Mid-Tertiary period, but the exact dating is uncertain, due to two differ¬

ent published estimates derived from fossil organisms in the deposits. An age of 20-15 Ma was proposed

(Iturralde-Vinent & MacPhee 1996) based on foraminifera, while studies of coccoliths yielded an estimate of

45-30 Ma (Cepek in Schlee 1999). The strata are composed of turbiditic sandstones of the Upper Eocene to

Lower Miocene Mamey Group (Draper et al. 1994).

The resin giving rise to the amber was produced by a leguminous tree, Hymenaea protera (Poinar 1991), a

relative of the present-day Neotropical species H. courharil L. The suite of amber-embedded organisms is rep¬

resentative of a moist tropical forest, which was described by Poinar and Poinar (1999) based on the known

insect fauna together with whatever plant fossils were then available. The present flower is the 17th angio-

sperm species to be described from Dominican amber. Others are 1 species of Fabaceae (Poinar 1991), 3 of

Arecaceae (Poinar 2002a, 2002b), 2 of Poaceae (Poinar & Judziewicz 2005, Poinar & Columbus 2012), 1 of

Chrysobalanaceae (Poinar et al. 2008a, revised by Chambers & Poinar 2010), 2 of Lauraceae (Chambers et al.

2011a, 2012), 3 of Meliaceae (Chambers etal. 2011b, Chambers & Poinar 2012), 1 of Burseraceae (Chambers &

Poinar 2013), 1 of Myristaceae (Poinar & Steeves 2013), 1 of Rhamnaceae (Chambers & Poinar 2014), and 1

possibly of Moraceae (Poinar et al. 2008b).

J. Bot. Res. Inst. Texas 8(2): 559 - 564.2014

560

Journal of the Botanical Research Institute of Texas 8(2)

Our proposal to place the fossil in Ticodendron came from its similarity to illustrations and descriptions of

T. incognitum published in Kubitzki (1993), and this relationship was later confirmed by examination of her¬

barium specimens. Ticodendron which was formally described only recently (Gomez-Faurito & Gomez-P. op.

cit.) is a dioecious, arborescent, monotypic genus occurring in moist tropical forests of Central America, from

southern Mexico to Panama. It had been collected infrequently in the past and was most often mistaken for

Alnus, due to a similarity in leaf architecture (Hickey & Taylor 1991). The small, unisexual flowers were not

well understood by earlier taxonomists, who were unable to place them in a known genus. Discovery of a fossil

species of Ticodendron , if correctly assigned,, will help in understanding the history of the family and of the

Tertiary Laurasian forests of which it is characteristic (Hammel & Burger 1991). These relationships are dis¬

cussed in a later section.

MATERIALS AND METHODS

The fossil is contained in a rectangular block of amber measuring 8x4x4 mm. It was obtained from mines in

the Cordillera Septentrional, between Puerto Plata and Santiago, Dominican Republic. Examination and pho¬

tographs were made with a Nikon stereoscopic microscope SMA-10-R at 80x and a Nikon Optiphot microscope

at800x.

DESCRIPTION

Ticodendron palaios K.L. Chambers & Poinar, sp. nov. (Figs. 1-2). Type: HISPANIOLA. Dominican Republic: amber mine

in the northern mountain range (Cordillera Septentrional), 2013, unknown amber miner s.n. (holotype: catalogue number Sd-9-192,

deposited in the Poinar amber collection maintained at Oregon State University, Corvallis, Oregon 97331, U.S.A.).

Flower single, pistillate, in a peduncled partial inflorescence (Tobe 1991), peduncle 1.9 mm long, lightly hispid,

arising from a terminal or axillary bud with numerous, spirally arranged bud scales (Fig. 1), bracts subtending

inflorescence 4, separate, lanceolate, acute, up to 1.9 mm long, 0.6 mm wide, glabrous (Figs. 1, 2), axillary

scales, if any, not visible, ovary inferior, oblong-cylindric, 2.8 mm long, 1.5 mm wide, densely hispid-setose

(Figs. 1, 2), pedicelled, pedicel 0.8 mm long, hispid-setose (Fig. 2), sepals 3 (2 visible), 0.7 mm long, 0.4 mm

wide, oblong-lanceolate, obtuse, hispid-setose abaxially (Fig. 2), petals 0, styles 2 (?), densely papillate (Fig. 2),

probably stigmatic throughout, unnaturally appressed to apex of ovary due to position of flower in the block of

amber; staminate inflorescence unknown.

Etymology .—From Greek “palaios,” ancient.

DISCUSSION

The apex of the flower is pressed against the edge of the block of amber (see Fig. 2), and there are an air bubble

and granular artifact on one side (Fig. 1) that make interpretation of the styles and perianth difficult. Two se¬

pals are evident in the other lateral view (Fig. 2), and the base of one folded-over style can be seen between

them, bending sideways across the flower apex (Fig. 2). The styles otherwise can’t be observed or measured. Air

bubbles are present among the bracts (Figs. 1, 2), but these do not interfere with an interpretation of the inflo¬

rescence.

There are a number of significant differences between Ticodendron palaios and the extant species, T. incog¬

nitum. In the latter taxon, the inflorescence does not arise directly from a basal bud. Instead, it is terminal on a

short lateral branch at whose tip is a pair of bracteate leaves (Gomez-Faurito & Gomez-P. 1989, Fig. 6F). Also,

the two lower bracts subtending the flower are connate basally and a third bract is separate and displaced up¬

ward (pers. observ.); all 3 are densely strigillose-pubescent abaxially and glabrous adaxially (pers. observ.). In

the fossil species, the bracts are 4, distinct throughout, and glabrous on both sides. The flower of T. incognitum

is sessile, whereas in T. palaios it is borne on a short pedicel. The pubescence of the ovary in the extant species is

lightly appressed-pilose, but in the fossil, it is densely hispid-setose. The sepals in T. incognitum are abundantly

pilose-hispid adaxially, and these trichomes persist around the base of the styles in the developing fruit (pers.

observ.). No such pubescence is seen in T. palaios. Feaf morphology, as well as the nature of the staminate inflo¬

rescence, is unknown in T. palaios, so that further comparisons with T. incognitum are not possible at present.

Chambers and Poinar, Ticodendron palaios a Mid-Tertiary fossil flower

561

Fig. 1. Ticodendron palaios. A: Terminal portion of one style lying over top of ovary. B and C: 2 of the 4 bracts subtending the flower. D: Peduncle of the

inflorescence. E: Scale of the basal bud. The air bubble and granular material at upper left are artifacts. Scale bar = 1.3 mm.

The anatomy, morphology, and relationships of Ticodendron were the subject of a series of papers pub¬

lished shortly after the genus was first described (Behnke 1991; Carlquist 1991; Feuer 1991; Hammel & Burger

1991; Hickey & Taylor 1991; Tobe 1991). The species had been collected a number of times in Central America,

but it was not recognized as a unique genus until its alder-like leaves were brought together with its incon-

562

Journal of the Botanical Research Institute of Texas 8(2)

Fig. 2. Ticodendron palaios. Reverse lateral view of inflorescence. A: base of style. B: 1 of 2 visible sepals. C: ovary. D: floral pedicel. Scale bar = 1.5 mm.

spicuous male and female inflorescences by Gomez-Laurito and Gomez-P. (1989). The authors writing in 1991,

above, considered several possible taxonomic orders in which to place the genus, including Juglandales, Myri-

cales, Fagales, and Urticales. The totality of its wood and bark anatomy, pollen morphology, phloem sieve-ele¬

ments, leaf architecture, and floral morphology led to its being classified as a monotypic family in order Fagales

(Gomez-Laurito & Gomez-P. 1991).

Chambers and Poinar, Ticodendron palaios a Mid-Tertiary fossil flower

563

Ecological relationships of the modern species, Ticodendron incognitum, were given special attention by

Hammel and Burger (1991), who noted its occurrence in diverse, moist, evergreen cloud forests of middle-ele¬

vations, found on both sides of the continental divide throughout Mesoamerica. According to these authors,

the fossil history of associated taxa indicates that they are members of a once widespread northern (Laurasian)

forest flora of the Tertiary Period (Hammel & Zamora 1990). Studies of the leaf architecture of T. incognitum by

Hickey and Taylor (1991) suggested a relationship with an Oligocene fossil taxon from Colorado, the genus

Fagopsis (Manchester & Crane 1983). This relationship was confirmed in cladistic analyses of 48 principally

vegetative characteristics (Hickey & Taylor, op. cit., Figs. 19-22). Even more convincing is its relationship to

the Middle Eocene fossil fruit Ferrignocarpus bivalvis from the Clarno Formation in Oregon, assigned to Tico-

dendraceae by Manchester (2011). In this paper, Manchester also described identical fruits from the Early Eo¬

cene London Clay flora in England, which had been informally named u Carpolithus sp. 38” by Reid and Chan¬

dler (1933). In his discussion, Manchester (op. cit.) noted that this bicontinental disjunction “implies that the

genus spread across higher latitudes and via the North Atlantic land bridge during the Late Paleocene and/or

Early Eocene, when climatic conditions were suitable for the establishment of thermophilic taxa farther

north.” This analysis parallels the observations by Hammel and Burger (1991) and Hammel and Zamora

(1990), mentioned above, concerning the history of present-day forests occupied by Ticodendron in Central

America. From our studies of Dominican amber fossils, cited earlier, the following genera can be added to this

forest type, as represented in the Caribbean region of the late Oligocene or early Miocene: Roystonea, Frithri-

nax, Palaeoraphe, Alarista, Pharus, Hymenaea, Ficania, Trochanthera, Swietenia, Trichilia, Persea, Treptostemon,

Virola, Protium, Distigouania, and Ticodendron.

ACKNOWLEDGMENTS

We thank Alex E. Brown for supplying us with the fossil and the curator of the Missouri Botanical Garden

herbarium for providing specimens of Ticodendron incognitum. Helpful comments and suggestions from Barry

Hammel and a second reviewer are gratefully acknowledged.

REFERENCES

Behnke, H.-D. 199E Sieve-element characters of Ticodendron. Ann. Missouri Bot. Gard. 78:131-134.

Carlquist, S. 1991. Wood and bark anatomy of Ticodendron. Ann. Missouri Bot. Gard. 78:96-104.

Chambers, K.L. & G.O. Poinar, Jr. 2010. The Dominican amber fossil Lasiambix (Fabaceae: Caesalpinioideae?) is a Licania

(Chrysobalanaceae). J. Bot. Res. Inst.Texas 4:217-218.

Chambers, K.L., G.O. Poinar, Jr., & A.E. Brown. 2011a. A fossil flower of Persea (Lauraceae) in Tertiary Dominican amber. J.

Bot. Res. Inst.Texas 5:457-462.

Chambers, K.L., G.O. Poinar, Jr., & A.E. Brown. 2011 b. Two fossil flowers of Trichilia (Meliaceae) in Dominican amber. J. Bot.

Res. Inst.Texas 5:463-468.

Chambers, K.L. & G.O. Poinar, Jr. 2012. 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., & A.S. Chanderbali. 2012. Treptostemon (Lauraceae), a new genus of fossil flower from Mid-

Tertiary Dominican amber. J. Bot. Res. Inst.Texas 6:551-556.

Chambers, K.L. & G.O. Poinar, Jr. 2013. A fossil flower of the genus Protium (Burseraceae) in Mid-Tertiary amber from the

Dominican Republic. J. Bot. Res. Inst. Texas 7:367-373.

Chambers, K.L. & G.O. Poinar, Jr. 2014. Distigouania irregularis (Rhamnaceae) gen. et sp. nov. in Mid-Tertiary amber from

the Dominican Republic. J. Bot. Res. Inst.Texas 8:551-557.

Draper, G., P. Mann, & J.F. Lewis. 1994. Hispaniola. In: S. Donovan and T.A. Jackson, eds. Caribbean geology: an introduc¬

tion. The University of the West Indies Publishers'Association, Kingston, Jamaica. Pp. 129-150.

Feuer, S. 1991. Pollen morphology and the systematic relationships of Ticodendron incognitum. Ann. Missouri Bot. Gard.

78:143-151.

Gomez-Laurito, J. & L.D. Gomez-P. 1989. Ticodendron: a new tree from Central America. Ann. Missouri Bot. Gard. 76:1148-

1151.

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Gomez-Laurito, J. & L.D. Gomez-P. 1991. Ticodendraceae: a new family of flowering plants. Ann. Missouri Bot. Gard.

78:87-88.

Hammel, B.E. & N.A. Zamora. 1990. Nyssa talamancana (Cornaceae), an addition to the remnant Laurasian Tertiary flora of

southern Central America. Brittonia 42:165-170.

Hammel, B.E. & W.G. Burger. 1991. Neither oak nor alder, but nearly: the history of Ticodendraceae. Ann. Missouri Bot.

Gard. 78:89-95.

Hickey, L.H. & D.W. Taylor. 1991. The leaf architecture of Ticodendron and the application of foliar characters in discerning

its relationships. Ann. Missouri Bot. Gard. 78:105-130.

Iturralde-Vinent, M.A. & R.D.E. Macphee. 1966. Age and paleogeographic origin of Dominican amber. Science 273:

1850-1852.

Kubitzki, K. 1993. Ticodendraceae. In: K. Kubitzki, J.G. Rohwer, and V. Bittrich, eds. The families and genera of flowering

plants. II. Flowering plants—Dicotyledons. Springer-Verlag, Berlin. Pp. 594-596.

Manchester, S.R. 2011. Fruits of Ticodendraceae (Fagales) from the Eocene of Europe and North America. Int. J. PI. Sci.

172:1179-1187.

Manchester, S.R. & P.R. Crane. 1983. Attached leaves, inflorescence, and fruits of Fagopsis, an extinct genus of fagaceous

affinity from the Oligocene Florissant flora of Colorado, U.S.A. Amer. J. Bot. 70:1147-1164.

Poinar, G.O., Jr. 1991. Hymenaea protera sp. n. (Leguminosae: Caesalpinioideae) from Dominican amber has African

affinities. Experientia 47:1075-1082.

Poinar, G.O., Jr. & R. Poinar. 1999. The amber forest. Princeton University Press, Princeton, New Jersey, U.S.A.

Poinar, G.O., Jr. 2002a. Fossil palm flowers in Dominican and Mexican amber. Bot. J. Linn. Soc. 138:57-61.

Poinar, G.O., Jr. 2002b. Fossil palm flowers in Dominican and Baltic amber. Bot. J. Linn. Soc. 139:361 -367.

Poinar, G.O., Jr. & EJ. Judziewicz. 2005. Pharusprimuncinatus (Poaceae: Pharoideae: Phareae) from Dominican amber. Sida

21:2095-2103.

Poinar, G.O. Jr., K.L. Chambers, &A.E. Brown. 2008a. Lasiambixdominicensis gen and sp. nov., a eudicot flower in Dominican

amber showing affinities with Fabaceae subfamily Caesalpinioideae. J. Bot. Res. Inst. Texas 2:463-471.

Poinar, G.O. Jr., K.L. Chambers, & A.E. Brown. 2008b. Trochanthera lepidota gen. and sp. nov., a fossil angiosperm inflores¬

cence in Dominican amber. J. Bot. Res. Inst.Texas 2:1167-1173.

Poinar, G.O. Jr. & J.T. Columbus. 2012. Alarista succina gen et sp. nov. (Poaceae: Bambusoideae) in Dominican amber.

Histor. Biol. 1-6.

Poinar, G.O. Jr. & R. Steeves. 2013. Virola dominicana sp. nov. (Myristicaceae) from Dominican amber. Botany 91:530-534.

Reid, E.M. & M.EJ. Chandler. 1933. The London Clay flora. British Museum (Natural History), London. 561 pp.

Schlee, D. 1999. Das Bernstein-Kabinett. Stuttgarter Beitr. Naturk. Ser. C, 28.

Tobe, H. 1991. Reproductive morphology, anatomy and relationships of Ticodendron. Ann. Missouri Bot. Gard. 78:

135-142.

VEGETATION PATTERNS IN THE MEDITERRANEAN-DESERT ECOTONE

OF BAJA CALIFORNIA, MEXICO

Sula Elizabeth Vanderplank

Botanical Research Institute of Texas

1700 University Dr.

Fort Worth, Texas 76107 US.A.

svanderplank@BRIT.org

Exequiel Ezcurra

University of California, Riverside

Department of Botany and Plant Sciences

900 University Ave.

Riverside, California 92507 U.S.A.

Jose Delgadillo

Universidad Autonoma de Baja California

Facultad de Ciencias

Carretera Transpeninsular Ensenada-Tijuana, No. 3917

Fraccionamiento Playitas

Ensenada, Baja California 22860 MEXICO

Lucinda A. McDade

Rancho Santa Ana Botanic Garden

and Claremont Graduate University

1500 North College Ave.

Claremont, California 91711 U.S.A.

ABSTRACT

The Pacific coast of NW Baja California is a hotbed of plant species richness and endemism. Greater San Quintin (GSQ), Baja California,

Mexico, is located near the southernmost reaches of the California Floristic Province in a vegetation transition zone. Perennial species

distributions were recorded and used to assess the distribution and richness of rare, narrowly endemic and native taxa. A Principal Com¬

ponents Analysis assessed the major elements in the perennial flora, and separated the habitats to indicate the primary factors affecting

plant distributions. Distribution of plant species is related to elevation and soil type. The data analysis identifies three distinct areas of spe¬

cies richness, each on a different substrate. Three major habitats were resolved in the analysis; however, indicator species provide greater

resolution of habitats with similar soils. Range size on a regional scale was not related to local abundance of perennial species. Across the

landscape, native, rare and endemic species richness patterns are very similar, suggesting that native plant richness can be used as a proxy

for identifying areas rich in sensitive species. This method provides useful data to conservation efforts. Coastal habitat is essential for many

species and conservation recommendations for the region are detailed.

RESUMEN

Ta costa pacifica del noroeste de Baja California es un foco de riqueza y endemismo para numerosas especies vegetales. Ta region de San

Quintin, Baja California, Mexico, se localiza cerca de los limites australes de la Provincia Floristica de California, en una zona de vegetacion

de transicion. Se registro la distribucion de especies perennes para evaluar la riqueza y distribucion de taxa nativos raros y de endemismos

restringidos. Un Analisis de Componentes Principales evaluo los elementos primordiales en la flora perenne y separo los habitats para indi¬

car factores primarios que afectan la distribucion de la vegetacion. Ta distribucion de las plantas esta relacionada con la elevacion y el tipo

de suelo. El analisis de datos identifica la riqueza de especies vegetales en tres distintas areas, cada una en un substrato diferente. Se deter-

minaron tres habitats principales en el analisis; sin embargo, las especies indicadoras aportan mayor resolucion de habitats con suelos simi-

lares. El tamano del rango a escala regional no tuvo relacion con la abundancia local de especies perennes. A traves del paisaje, los patrones

de riqueza de especies nativas, raras y endemicas son muy similares, sugiriendo que la riqueza de flora nativa puede ser utilizada como un

agente para identificar areas ricas en especies sensibles.

Key Words: Endemism, mapping, maritime succulent scrub, rarity, species richness, San Quintin, Baja California, vernal pools

INTRODUCTION

Around the mid-latitudinal belt of the globe, the western side of continents is occupied by deserts while the

eastern side is normally covered by forests. This distribution of coastal drylands is driven by the global circu¬

lation of oceans: Equator-bound currents on the eastern side of large oceans are deflected westwards by the

rotational force of the Earth, to form the equatorial currents and the easterly trade winds. As the westbound

surface waters move away from the continents, they pull cold, nutrient-rich waters to the surface that generate

a cool, stable coastal atmosphere, with little evaporation from the sea and very low rainfall other than morn¬

ing fogs. In coasts neighboring these oceanic upwellings, typical coastal fog deserts tend to develop, forming

some of the driest ecosystems on earth. Thus, the large-scale circulation of the ocean is the main reason why

J. Bot. Res. Inst. Texas 8(2): 565 - 581.2014

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Journal of the Botanical Research Institute of Texas 8(2)

coastal deserts are always found on the west side of continents, such as the Succulent Karoo in Africa, Atacama

in Chile, the Atlantic Coastal Desert of Morocco, or the deserts of Baja California. All these fog-driven drylands

are adjacent to winter-rain Mediterranean ecosystems in higher latitudes, and large coastal Mediterranean-

to-desert ecotones driven by fog, more than rains, occupy the transition areas along these coasts (Ezcurra &

Mellink 2013).

The over-arching pattern of global species diversity is of increased species numbers and decreased range

sizes toward the equator (Rapoport’s rule, Stevens 1989). However, regions of very high species diversity and

endemism are also found in areas of Mediterranean climate at intermediate latitudes. The California Floristic

Province (CFP) has been designated as a global biodiversity hotspot, moreover an area of high endemism that

has been heavily impacted by human activity (Myers et al. 2000). The geographic distribution of the California

Floristic Province (CFP) essentially coincides with the presence of winter-spring rainfall and the absence of

summer precipitation (Minnich & Franco-Vizcalno 1998; Caso et al. 2007).

At the southern extreme of the CFP, the desert-to-Mediterranean transition in northwestern Baja Cali¬

fornia has a large proportion (-10%) of endemism, including 172 locally endemic species, many of which are

micro-endemic to areas of less than 50 hectares (O’Brien et al., in press). Fittle is known about the factors con¬

trolling the distributions of these narrowly endemic species in these ecotonal drylands.

As we enter a period of accelerated global environmental change, it is predicted that we will see extinc¬

tions on an unprecedented scale (Thomas et al. 2004). The CFP is no exception and current estimates are that

66% of endemic species will experience up to 80% range reductions in the next century under even conserva¬

tive climate change scenarios (Foarie et al. 2008). However, the impacts of climate change have been difficult

to assess due to the absence of baseline data from which we can establish distribution patterns and assess the

fate of rare and endemic taxa (Jansson 2009).

The southern limit of the CFP has been the subject of debate for decades. Rather than a sharp boundary,

there is a gradual Mediterranean-scrub-to-desert transition that begins near the U.S./Mexico border and ends

near the 30 th parallel near El Rosario according to Shreve (1936). This transitional zone has the greatest floris¬

tic diversity in the state of Baja California and transitional vegetation associations recorded here have the high¬

est endemic community values (i.e., greatest densities of narrowly endemic taxa, Peinado et al. 1995a). In the

present study we focus on the area between parallels 30 and 31 on the Pacific Coast of Baja California, Mexico,

here referred to as ‘Greater San Quintln’ (GSQ; Fig. 1). Although rainfall is low for most of the year, with an an¬

nual average of 105-137 mm (Vanderplank 2011), San Quintln experiences frequent heavy fogs caused by the

cool upwelling of the California current off-shore (Herbert et al. 2001), and the associated humidity provides

significant extra water resources, to which the local vegetation is adapted (Rundel et al. 1972).

The ecotonal position of San Quintln (where the Mediterranean CFP transitions into the Sonoran desert

to the south) often makes species distributions patchy and discontinuous, with the vegetation being very

heterogeneous in this region (Vanderplank 2013). Greater San Quintln lies at the southern end of this tran¬

sitional vegetation and is home to sixty-seven taxa that are endemic or near-endemic to NW Baja California

(Vanderplank 2011). The unusual mix of CFP species with plants from the desert area to the south make GSQ

a floristically rich area with 433 taxa documented in recent years (Vanderplank 2011). Thirty-four percent of

the species that comprise the flora of GSQ are rare and/or locally endemic, making GSQ a priority region for

conservation (Vanderplank 2011).

Greater San Quintln is currently under consideration for declaration as a natural protected area or Area

Natural Protegida (ANP) under Mexican law. Focal conservation organizations have identified four distinct

terrestrial habitats in need of conservation in GSQ: salt marsh, dunes and beaches, rivers and riparian areas,

and the maritime succulent scrub (note that “riparian” areas in this region are most often dry washes with san¬

dy substrates). The habitats have been similarly categorized by the Instituto Nacional de Estadistica Geografia e

Informatica, Mexico (INEGI), although upper salt flats are distinguished from inundated marshes.

Conservation efforts are unavoidably biased by available biodiversity data. The challenge of identifying

core areas for conservation given the rich assemblage of rare and endemic plants in GSQ is heightened by

Vanderplank et al., Vegetation patterns in a Mediterranean-Desert ecotone

567

Fig. 1. Topography and local place names within Greater San Quintin. N 30.18-30.45 and W115.48-116.03.

568

Journal of the Botanical Research Institute of Texas 8(2)

competition for land use and the heterogeneity of the vegetation itself. Despite the very limited global extent

of the maritime succulent scrub (Cartron et al. 2005), the saltmarsh habitat is most frequently the primary

conservation target in this region. The saltmarshes of GSQ provide a suite of ecosystem services and provide

a model system for restorations in other areas along the Pacific coast (West 2000) yet are the most floristically

depauperate habitat, and harbors only one endemic taxon (Bell 2010; Vanderplank 2011).

Fine-scale phytosociological studies in this region have shown diverse shrub associations in northwest¬

ern Baja California, where the climate gradient from north to south correlates with increased floristic diversity

(both for endemic species and diversity of life-forms) as aridity increases (Peinado et al. 1995a, 1995b). This is

the first study conducted at a macroecological scale in this region (lxl km 2 ) and brings new perspective on

the larger factors controlling vegetation distributions in this region.

A goal of this study was to investigate the distribution of the long-lived taxa that dominate the vegetation

and can be identified year-round. This study determines the current status of the terrestrial perennial vegeta¬

tion and provides baseline data from which future changes in the flora can be assessed and monitored. We here

try to assess which factors are dominant in the distribution of vegetation, and in particular, in the distribution

of rare and endemic species.

MATERIALS AND METHODS

Sampling methods. —A one-km grid of cells was created in ESRI ArcMap and georeferenced to overlay the

study area (the top-left NW corner of the grid lies at N 30.738 and W 116.039). Each grid cell became the mac¬

roecological sampling unit for the terrestrial vegetation. TerraSync software was used with a Trimble Juno GPS

unit in the held. Grid cells that contained only agricultural fields or urbanized areas, as well as cells with less

than 10% cover of natural vegetation were excluded from the analysis.

Quantitative data on presence and abundance of perennial species and selected invasive annual species

was gathered within each grid square (the estimation method is described below). All perennial plants that

could be identified year-round were included in the analyses; geophytes and herbaceous perennials that could

not be consistently identified throughout the year were excluded. Non-native annual taxa that were selected

for inclusion were abundant, identifiable year-round, and judged to be potentially invasive. The goal here was

to provide spatial information on the current extent of the most abundant non-native species. This study fol¬

lows the vascular plant checklist for Baja California (Rebman in review) for taxonomy and nativity.

Each square km that had native vegetation was visited to assess a number of traits of the vegetation. A

visual assessment was made to estimate the percent natural vegetation and to determine species present. Trials

showed that this method encountered a significantly higher number of species than alternative methods (e.g.,

quadrats or transects) that subsample smaller areas. Several taxa cannot readily be differentiated from one

another at all times of the year, these were pooled at a taxonomic level higher than species. For example, the

three species of the genus Lycium reported from the San Quintln area (these drought-deciduous shrubs cannot

easily be distinguished without leaves). Admittedly less than ideal, this approach was preferred over omitting

these taxa from the study (see supplemental information table for taxon list).

Abundance per km cell was estimated using the method of Nichols (1930) and the following categories

of abundance: (1) rare: fewer than 3 individuals encountered; (2) scarce: 3-10 individuals; (3) uncommon:

>10 but < 100 individuals (1/10,000 m 2 ); (4) frequent: 100-1,000 individuals (1/1,000 m 2 ); (5) common:

1,000-10,000 individuals (1/100 m 2 ); (6) abundant: 10,000-100,000 individuals (1/10 m 2 ); and (7) dominant:

>100,000 individuals (1/1 m 2 ). Estimates were based on number of plants seen and their distribution patterns.

When estimating numbers of plants that sprout from roots or rhizomes (e.g., salt grasses such as Distichlis

spp.), the numbers of individual sprouts were considered, such that these should not be taken to estimate num¬

bers of genetic individuals.

Analysis. —The resulting data matrix for each cell listed perennial and selected non-native annual taxa scored

for presence/absence. Perennial species richness was calculated for each square. Pearson’s residuals (the stan¬

dardized difference between the observed species richness in each grid square and the mean over all squares,

Vanderplank et al., Vegetation patterns in a Mediterranean-Desert ecotone

569

Everitt 1992) were calculated. Because Pearson residuals are distributed as normal deviates, those squares

that departed significantly (P < 0.01) from the mean richness were identified (Everitt 1992) to show areas of

significantly high or low richness.

Each taxon was classified according to (1) its origin (native/non-native), (2) endemism (endemic/non¬

endemic), (3) regional rarity, and (4) local rarity or abundance. All taxa that are restricted to the Baja California

peninsula (most of which are also endemic, or nearly so, to the CFP within the peninsula) were classified as

endemic. For rarity we followed O’Brien et al. (in press), which is based on previous work by the California

Native Plant Society: taxa on any of three lists of concern (i.e., rare globally, rare in CFP Baja California but

more common elsewhere, and watch-list taxa) were together categorized as “rare” for our analysis (in the few

incidences where multiple taxa were lumped, the category of the most abundant taxon was used). Data on

abundance or rarity within Greater San Quintln were drawn from the vegetation data. Species found in more

than 40 one-km cells were classified as locally abundant, species found in less than 10 one-km cells were clas¬

sified as locally scarce, and all species between the two thresholds were classified as intermediate. We tested

for association between regional and local rarity, using X 2 contingency-table analysis.

ESRI ArcMap was used to map species distributions across the study area. Data from the matrix were

attached to the grid layer using spatial coordinates for each cell. Relevant data layers were obtained from the

Instituto Nacional de Estadlstica y Geografla, Mexico (INEGI 1997, including shore outline, elevation, relief,

soil, and habitat types. Following analysis of the data matrix, squares with extreme species richness or paucity

(Pearson’s residuals, P < 0.01) were mapped. The same was done for species richness for three subsets of taxa:

native, endemic, and rare species.

Multivariate analysis.—To examine the distribution of species across habitat types as distinguished by

The Nature Conservancy of Baja California, a non-standardized principal components analysis (PCA) of the

presence-absence matrix for 163 native perennial taxa was conducted. Because the cross-product of two spe¬

cies vectors with presence-absence is simply the number of sites shared by both species, and, reciprocally, the

cross-product of two presence-absence site vectors is the number of shared species, PCA on presence-absence

data will reflect the number of co-occurrences as a measure of ecological similarity (Ezcurra 1987). The result¬

ing PCA axes were tested for significance using the broken-stick model (Jackson 1993). These scores were then

mapped to extract the dominant habitat type for each km square in the analysis. This map was compared to

the habitat maps from INEGI and habitat maps generated using indicator species. An indicator species from

each of the primary habitats was selected based on observed consistent presence in, and restriction to, specific

habitat.

RESULTS

Vegetation patterns.—The analysis of vegetation tallied a total of 163 taxa, including 140 native perennials,

across the 206 one-km cells (Table 1). This analysis allowed individual taxa to be mapped across the study area

and their abundance displayed for each km square. The abundance distribution of endemic and rare species

did not differ significantly from that of the total native flora within the Greater San Quintln area (X 2 = 1.86;

df = 2, P = 0.39; Table 1), indicating that plants that are rare at a regional scale may be locally abundant within

our study site, and, conversely, that plants that are regionally common may be locally scarce within our site.

Total species richness within each one-km cell varies across the study region (Fig. 2). The spatial distribu¬

tion of significantly (P > 0.01) high and low Pearson’s residuals for the whole flora shows some well-defined

patterns (Fig 3a), especially for high-diversity cells, that tend to occur in three distinct spatial clusters. These

spatial patterns of high richness are repeated in a similar manner for subsets of the flora (total native, Fig. 3b;

endemic, Fig. 3c; rare taxa, Fig. 3d).

Species richness is low adjacent to agriculture and urbanized areas (compare Fig. 3 to Fig. 5b). Richness

in the salt marshes and saline flats is much lower than in other habitats. This is particularly evident north of

the volcanic held and south of the Santa Marla Escarpment in the area surrounding Faguna Mormona (note

bands of pale yellow squares along the coast; Fig. 3a & b). Non-native taxa are especially rich in riparian areas,

570

Journal of the Botanical Research Institute of Texas 8(2)

Table 1. Native species total (140 species) and parsed into two categories of regional rarity within the CFP in Baja California (endemism and rarity), and by their

local abundance category based on their frequency within Greater San Quintin (scarce, intermediate, or abundant).

Local Rarity

Endemic

Rare

Total native

Scarce

Intermediate

Abundant

Totals

140

and their distributions follow all major drainages. The non-native iceplant Mesembryanthemum crystallinum

occurs almost ubiquitously across the study area. It is interesting to note that there was no correlation between

percent natural vegetation in a square and species richness (r = 0.11, df = 196; P = 0.13). This absence of a clear

species-area relationship at the one-km cell scale is most likely due to heterogeneity of the landscape; three

habitat types coexist in a fraction of a one-km cell in whereas in others an entire km cell of degraded salt marsh

that might have only four perennial species.

Areas of high species richness.—Three primary regions of high species richness emerge from the vegetation

study: at the northern and southern extremes of the study area and along the coast adjacent to the volcanic held.

Northern biodiversity hotspot: The northern region of high species richness has two distinct subareas:

Santo Domingo wash (northern boundary of the study area) and a clay mesa (slightly southeast from the

wash). The Santo Domingo wash shows high species richness when all taxa are considered (Fig. 3a); however,

richness here decreases somewhat when non-native taxa are excluded (Fig. 3b), and the area has few rare (Fig.

3d) or endemic (Fig. 3c) species. In contrast, the clay mesa in Colonia Vicente Guerrero, north of the Santa

Maria escarpment, presents a strong signal of high species richness for native, endemic, and rare species (i.e.,

compare Figs. 3a-d). This area is a disturbed patch of vegetation with diverse succulent scrub and some small

remnant-but-degraded vernal pools. Like the Santo Domingo wash this area has significantly more non-native

species than typical of the study area as a whole.

Volcanic coast hotspot: The coastal strip near the volcanic held that includes the tip of the sandy peninsula

at Monte Mazo is particularly diverse. This area includes a complex patchwork of habitats along the coast

which likely contributes to high species richness. The precise cells that show significant richness among the

analyses vary slightly (i.e., compare Figs. 3a-d), but the coastal strip is always rich in native, rare, and endemic

species. There are not significantly high numbers of non-natives in this area (consistent with the relatively

intact nature of the habitats).

Southern dunes hotspot: Near the southern end of the study area, the El Socorro Dunes stand out as the

largest area of high species richness. Because only one habitat type is represented across several cells here, the

observed high richness is not the result of habitat heterogeneity. Although there are some differences in the

pattern of cells that harbor significantly high numbers of native, endemic, and rare taxa, the area is rich in all

of these.

Multivariate analysis.—The Principal Components Analysis showed two axes that, according to the broken

stick test, departed significantly from a random model (Fig. 4). The first axis is positively correlated with topo¬

graphic elevation (r = 0.41, df = 204, P < 0.0001) and displays a salinity-elevation gradient: plants of coastal

habitats are on the left and those of maritime succulent scrub are on the right (positive on axis 1). The second

axis reflects soil type on coastal habitats and separates saltmarshes (positive on axis 2) from freshwater and

sand dune species (negative on axis 2). The corresponding species plot, accordingly, shows three distinct clus¬

ters of species dominating in three habitat types: (a) scrub, (b) saltmarshes, and (c) sandy soil plants regardless

of whether on dunes or in riparian areas (Fig. 4.). When mapped back onto the study area, the three habitats

identified by the PCA bi-plots produced a map that was very consistent with the vegetation chart (INEGI1997

produced with completely different methods (Fig. 5).

Vanderplank et al., Vegetation patterns in a Mediterranean-Desert ecotone

571

Total species richness

• 4-9

O 9-13

O 13-19

• 19-27

0 27-38

£ 38-55

Fig. 2. Total species richness in 206 one-km cells for perennial vegetation of Greater San Quintin. As indicated at lower left, the size of the circle reflects

number of species recorded in each cell.

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Journal of the Botanical Research Institute of Texas 8(2)

Fig. 3. Spatial pattern of 206 cells with significantly higher (red) and lower (yellow) species richness than expected using Pearson's residuals (P < 0.01)

to test departure from random distributions for (a) all species; (b) native species; (c) endemic species; (d) rare species.

Vanderplank et al., Vegetation patterns in a Mediterranean-Desert ecotone

573

Fig. 4. First two axes of PCA biplot for (a) species and (b) one km cells. Axis 1 separates maritime succulent scrub species and sites (red) which occur in

the higher parts of the area, from the more coastal vegetation types. Axis 2 separates saltmarsh species and sites (green) from dune species (orange)

and sandy soil communities (yellow).

574

Journal of the Botanical Research Institute of Texas 8(2)

Fig. 5. (a) Spatial pattern of habitats defined by the first two PCA axes (see Fig. 4). (b) Vegetation map of Greater San Quintin (GSQ) modified from

Geographic Information System layers provided by INEGI (gray indicates areas without vegetation).

DISCUSSION

Patterns in the perennial vegetation

The analysis of perennial vegetation clearly identified overlapping areas of high species richness for the analy¬

sis of the regional endemic flora and for native plants as a whole. This suggests that endemics are distributed

across GSQ in the same pattern as native species, and as a result, species rich areas will have many range-re¬

stricted taxa. On the other hand, local rarity was found to be independent of regional rarity—taxa that are rare

at the regional scale are equally likely to be rare or abundant at the local scale—suggesting that many regional

endemics may be abundant where found, a result of importance for conservation.

The multivariate analysis identified groups of species whose distribution can be readily interpreted as

related to elevation and soil type. This suggests that these two variables are key factors in species distributions.

Many species are in the central cluster with variation in their distributions unexplained by these axes. Some

are generalist species that do not have marked habitat preference and whose occurrence is not highly associ¬

ated to the presence of other species. Additionally, because the sampling regime (i.e., one-km 2 cells) does not

necessarily correspond exactly with habitats (i.e., several habitats may fall within any given cell), many taxa

that occupy different habitats may be co-mingled in the data matrix, in effect limiting the discriminatory

power of the analysis.

More attention should be paid to both the geologic origins and soil types of the region and the extent of

the salt spray zone. It is likely that these factors affect plant distributions throughout GSQ. The vegetation of

the region can be seen to clearly differ among sand, mud, volcanic rock, or clay substrates. As a direct result

of this, carefully selected indicator species can give finer scale resolution of habitats (Fig. 6). Many species are

restricted to entirely coastal habitats (e.g., dunes), highlighting the importance of coastal ecosystems to biodi¬

versity conservation.

Vanderplank et al., Vegetation patterns in a Mediterranean-Desert ecotone

575

Regions of high perennial species richness

Coastal volcanic field: The coastal strip along the volcanic held of San Quintln includes many species that

should be a high priority for conservation in Mexico due to their limited distribution. The unique geology of

the volcanic held is connected to a number of edaphically restricted endemic species (Vanderplank 2011]); this

area should be prioritized in regional conservation efforts. Conservation of the maritime succulent scrub that

occurs on clay soils outside the volcanic held will require more regional analysis beyond the boundaries of

GSQ where this habitat occurs in larger intact areas.

El Socorro Dunes: The El Socorro Dunes represent an unusually large and intact dune system that should

be highly prioritized for protection. It is perhaps the area most in need of conservation. The unusual composi¬

tion and high species diversity found here are not matched elsewhere in Mexico (Rodrlguez-Revelo et al. 2014).

There is a wealth of archeological data from the area in the form of coastal shell middens and shards (Moore

1999) that also provides signihcant justihcation for conservation of this region. In December 2010, the entire

dune system was slated for development and was being divided into narrow strips that connect the main road

to the beach.

Clay mesa of Vicente Guerrero : The clay mesa just north of the Santa Marla Escarpment, in Colonia Vi¬

cente Guerrero, is used as a village commons by the local ejidatarios. The area is often used as a latrine and as

a trash dump. Restoration of the vernal pools here may be feasible but would require protection and monitor¬

ing. Although small in size, the vernal areas here could serve as a re-introduction site for species extirpated

elsewhere in Greater San Quintln. This area may be ideal for a community outreach and education project

encompassing habitat restoration.

CONCLUSIONS

Patterns in the flora are strongly related to elevation or distance from the sea, and soil conditions. On a broad

scale these parameters characterize four broad habitat types, yet on a finer scale, specialized micro-habitats

with range-restricted taxa are also present. Native species richness is the primary indicator for rare and en¬

demic species richness in the flora. Conservation of biodiversity here will conserve the majority of endangered

taxa; however specific measures (e.g., protection of key areas) are necessary to conserve micro-endemics and

the endemics of micro-habitats.

The perennial flora of the area forms a rich heterogeneous matrix, with high levels of endemism and rare

plant species throughout. As a transitional area in a global biodiversity hotspot, the heterogeneity of GSQ may

provide critical refugia for many species under a changing climate regime. Ongoing assessment of habitat

quality and the status of the globally rare and locally endemic taxa in this region are strongly recommended.

The results presented here provide valuable data for planning in this region and may be used to model

faunal distributions for taxa that exhibit species-specific and habitat-specific interactions. This methodology

is recommended for future floristic studies seeking to provide baseline data to conservation.

APPENDIX 1

Table of perennial and selected non-native species from Greater San Quintln. Their status (non-native, rare, endemic) is given as assessed for each taxon in the

study (rarity and endemism follows combined values from B. O'Brien et al. (In Press, taxonomy and nativity follow Rebman (In Review).

Family Taxon Non-native Rare Endemic

Agavaceae

Agave shawii subsp. shawii

Agavaceae

Yucca schidigera

Aizoaceae

Carpobrotus chilensis

Aizoaceae

Mesembryanthemum crystallinum

Aizoaceae

Mesembryanthemum nodiflorum

Anacardiaceae

Malosma laurina

Anacardiaceae

Rhus integri folia

Anacardiaceae

Toxicodendron diversilobum

Apiaceae

Foeniculum vulgare

Apocynaceae

Asclepias subulata

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Journal of the Botanical Research Institute of Texas 8(2)

Fig. 6. Distribution and abundance of the main indicator species (compare to habitats in Fig. 5). Intensity of color reflects number of individuals per

km 2 , (a) Ambrosiachenopodiifolia, an indicator for maritime succulent scrub; (b) Ambrosiamonogyra, indicator for riparian habitat; (c) Jaumeacarnosa,

indicator for saltmarsh habitat; (d) Helianthus niveus, indicator for dunes and other sandy soils (note that some plants also occur in sandy riparian

areas near the coast).

Vanderplank et al., Vegetation patterns in a Mediterranean-Desert ecotone

577

APPENDIX 1

(continued)

Family

Taxon

Non-native

Rare

Endemic

Apocynaceae

Funastrum orenorium

Asteraceae

Ambrosia chamissonis

Asteraceae

Ambrosia chenopodifolia

Asteraceae

Ambrosia confertiflora

Asteraceae

Ambrosia monogyra

Asteraceae

Artemisia californica

Asteraceae

Artemisia dracunculus

Asteraceae

Artemisia dracunculus

Asteraceae

Artemisia tridentata

Asteraceae

Baccharis pilularis

Asteraceae

Baccharis salicifolia

Asteraceae

Baccharis sarothroides

Asteraceae

Bahiopsis laciniata

Asteraceae

Bahiopsis triangularis

Asteraceae

Bebbia juncea

Asteraceae

Brickellia californica

Asteraceae

Centauria melitensis

Asteraceae

Chrysanthemum coronarium

Asteraceae

Leptosyne maritima

Asteraceae

En celia as peri folia

Asteraceae

Encelia californica

Asteraceae

En celia farinosa

Asteraceae

Ericameria palmeri

Asteraceae

Eriophyllum confertiflorum

Asteraceae

Hazardia berberidis

Asteraceae

Helianthus niveus

Asteraceae

Heterotheca grandiflora

Asteraceae

Isocoma menziesii subsp. menziesii

Asteraceae

Isocoma menzeisii subsp. decumbens

Asteraceae

Iva haesiana

Asteraceae

Jaumea carnosa

Asteraceae

Pluchea sericea

Asteraceae

Pseudognaphalium biolettii, P. leucocephalum

Asteraceae

Senecio lyonii

Asteraceae

Stephanomeria exigua, 5. diegensis

Asteraceae

Trixis californica

Bataceae

Batis maritima

Brassicaceae

Brassica nigra

Brassicaceae

Brassica tournfordii

Brassicaceae

Dithyrea maritima

Brassicaceae

Sisymbrium irio

Cactaceae

Bergerocactus emoryi

Cactaceae

Cylindropuntia alcahes

Cactaceae

Cylindropuntia californica subsp. rosarica

Cactaceae

Cylindropuntia cholla

Cactaceae

Cylindropuntia molesta

Cactaceae

Cylindropuntia prolifera

Cactaceae

Echinocereus maritimus

Cactaceae

Ferocactus fordii

Cactaceae

Lophocereus schottii

Cactaceae

Mammillaria brandegeei

Cactaceae

Mammillaria dioica, M. louisae

Cactaceae

Myrtillocactus cochal

Cactaceae

Opuntia littoralis, Opuntia oricola, 0. phaeacantha

Cactaceae

Stenocereus gummosus

Cleomaceae

Peritoma arborea

Caryophyllaceae

Cardionema ramossissima

Chenopodiaceae

Allenrollfea occidentalis

578

Journal of the Botanical Research Institute of Texas 8(2)

APPENDIX 1

(continued)

Family

Taxon

Non-native Rare

Endemic

Chenopodiaceae

Arthrocnemum subterminole

Chenopodiaceae

Atriplex canescens

Chenopodiaceae

Atrip lex juloceo

Chenopodiaceae

Atriplex leucophylla

Chenopodiaceae

Atriplex semiboccoto, A. suberecto

Chenopodiaceae

Atriplex watsonii

Chenopodiaceae

Chenopodium murole

Chenopodiaceae

Salsola tragus

Chenopodiaceae

Salicornia pacifica

Chenopodiaceae

Suaeda spp. (S. esteroa, 5. nigra, 5. taxifolia )

Crassulaceae

Dudleya anthonyi

Crassulaceae

Dudleya attenuata

Crassulaceae

Dudleya cultrata

Cyperaceae

Cyperus esculentus

Cyperaceae

Bolboschoenus maritimus

Ephedraceae

Ephedra califomica

Equisetaceae

Equisetum laevigatum

Ericaceae

Arctostaphylos glauca

Euphorbiaceae

Acalypha californica

Euphorbiaceae

Croton californicus

Euphorbiaceae

Euphorbia misera

Euphorbiaceae

Riccinus communis

Euphorbiaceae

Stillingia linearifolia

Fabaceae

Astragalus anemophilus

Fabaceae

Astragalus fastidius, A. gruinus, A. trichopodus

Fabaceae

Acmispon niveus

Fabaceae

Acmispon distichum

Fabaceae

Acmispon heermanii

Fabaceae

Acmispon watsonii

Fabaceae

Lupinus cf. longifolius

Fabaceae

Prosopis glandulosa

Frankeniaceae

Frankenia palmeri

Frankeniaceae

Frankenia salina

Geraniaceae

Erodium spp.

Grossulariaceae

Ribes tortuosum

Hippocastanaceae

Aesculus parryi

Boraginaceae

Eriodictyon angustifolium

Boraginaceae

Eriodictyon sessilifolium

Boraginaceae

Phacelia ixoides

Juncaceae

Juncus acutus subsp. leopoldii

Juncaceae

Juncus mexicanus

Juncaginaceae

Triglochin maritina

Lamiaceae

Hyptis emoryi

Lamiaceae

Salvia apiana

Lamiaceae

Salvia brandegeei

Loasaceae

Petalonyx linearis

Malvaceae

Malacothamnus fasciculatus

Malvaceae

Malva parviflora

Malvaceae

Sphaeralcea ambigua

Malvaceae

Sphaeralcea fulva, S. axillaris

Ncytaginaceae

Mirabilis laevis var. laevis

Nyctaginaceae

Abronia maritima

Onagraceae

Camissonia cheiranthifolia

Onagraceae

Camissonia crassifolia

Papaveraceae

Argemone munita

Phrymaceae

Mimulus puniceus

Plantaginaceae

Gambelia juncea

Plantaginaceae

Penstemon spectabilis subsp. subinteger

Vanderplank et al., Vegetation patterns in a Mediterranean-Desert ecotone

579

APPENDIX 1

(continued)

Family

Taxon

Non-native Rare Endemic

Platanaceae

Platan us racemosa

Plumbaginaceae

Limonium californicum

Poaceae

Arundo donax

Poaceae

Bromus madritensis

Poaceae

Cynodon dactylon

Poaceae

Distichlis littoralis

Poaceae

Distichlis spicata

Poaceae

Melica imperfecta, M. frutescens

Poaceae

Pennisetum ciliare

Poaceae

Phalaris minor

Poaceae

Schismus barbata

Poaceae

Spartina foliosa

Poaceae

Stipa lepida

Polygonaceae

Eriogonum fasciculatum

Polygonaceae

Harfordia macroptera subsp. galioides

Polygonaceae

Polygonum aviculare, P. agyroceolum

Polygonaceae

Rumexpulcher

Polypodiaceae

Ch eilan thes/Pen togramma/Aspidotis

Polypodiaceae

Pellaea mucronata

Polypodiaceae

Polypodium californicum

Ranunculaceae

Clematis pauciflora

Rhamnaceae

Ceanothus leucodermis

Rhamnaceae

Rhamnus crocea

Rosaceae

Adenostoma fasciculatum

Rosaceae

Heteromeles arbutifolia

Rosaceae

Rosa minutifolia

Salicaceae

Salixexigua var. hindsiana

Salicaceae

Salixlasiolepis

Saururaceae

Anemopsis californica

Simmondsiaceae

Simmondsia chinensis

Solanaceae

Lycium spp. (L brevipes, L californicum, L andersonii, L fremontii)

Solanaceae

Nicotiana glauca

Solanaceae

Solanum americanum

Solanaceae

Solanum hindsiana

Solanaceae

Solanum palmeri

Tamaricaceae

Tamarix ramosissima

Typhaceae

Typha dominguensis

ACKNOWLEDGMENTS

Funding was generously provided by The Jiji Foundation, Rancho Santa Ana Botanic Garden, and Claremont

Graduate University, Claremont University Club, BJ Ledyard, Tony LaFetra, and the California Native Plant

Society We thank the staff of the RSA-POM, BCMEX, and SD herbaria for their assistance with herbarium

specimens. Thanks to Giandiego Campetella, Laco Mucina, Phil Rundel, and an anonymous reviewer for their

helpful edits and suggestions. Many people helped with held work. Vanderplank is especially grateful to Jorge

Ochoa and Sergio Mata for their held assistance and to Naomi Fraga and Richard Felger for their insightful

comments and long-term mentoring. For institutional support Vanderplank thanks UCMEXUS (The Univer¬

sity of California Institute for Mexico and the United States); Rancho Santa Ana Botanic Garden; San Diego

Natural History Museum; Terra Peninsular A.C.; and the Next Generation of Sonoran Desert Researchers.

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582

Journal of the Botanical Research Institute of Texas 8(2)

BOOK REVIEW

Ricky J. Linex. 2014. Range Plants of North Central Texas — A Land User’s Guide to Their Identification,

Value and Management. (ISBN-13: 978-1-4951-2165-4, pbk., spiral-bound). USDA Natural Resources

Conservation Service, 532 Santa Fe Drive, Weatherford, Texas 76086, U.S.A. (Orders: shop.brit.org,

1-817-332-4441). $20.00,345 pp., color photos, illus. glossary, glossary, bib., index, 9.5" x 11.25".

Wildlife Biologist Ricky Linex has created a fine resource for botanists, farmers, land managers, wildlife manag¬

ers, plant enthusiasts, students, and anybody interested in the flora of Texas and the southern plains states.

Though the title specifies its regional focus as North Central Texas, the book should also prove useful to readers

in Oklahoma and even parts of New Mexico.

The concept for the book came from a previous USDA-NRCS publication, Common Rangeland Plants of the

Texas Panhandle by Clint Rollins. Linex took the idea and repeated it for an area that includes not only the Rolling

Plains vegetational area, but also the Cross Timbers, Blackland Prairie, and Post Oak Savannah areas.

The book is laid out in sections, with forbs, grasses, and woody plants being separated out and arranged

taxonomically by first family then genus. Though some of the family names are old (Asclepiadaceae and Cheno-

podiaceae instead of Apocynaceae and Amaranthaceae), the author does include synonyms for a good portion of

the 324 taxa, ostensibly for “those of us who first learned the scientific names more than a few years ago”.

Each taxon page is headed in large print by the most-used common name, with alternative common names

listed out to the side. A small area in the upper left or right corner of each page includes the following items: sci¬

entific name, family name, common name for family, origin, longevity, season of growth, bloom period, and

distribution relative to vegetational areas. In addition, each page includes three narrative sections on identifica¬

tion, value, and management (hence the book’s subtitle), and it is these last two that make this book different

from other regional guides. The value section on Justicia americana (water-willow), for example, states “Live¬

stock and deer graze upon water-willow, and the plants will be utilized extensively when other forage becomes

scarce during hot, dry summers ...” The management section for the same species reads, “Under heavy use by

livestock and native or exotic animals, water-willow will be browsed close to the ground. Proper stocking rates

and rotational grazing will ensure this forb will remain rooted in wet areas where it helps provide stability to

stream banks.”

Each page boasts from 3 to 6 color photographs (typically 4 or 5), many of which were taken by the author

himself. A good number of pages contain a standardized seed photo—several seeds against an orange back¬

ground with a Vie inch ruler for scale. This standardization makes these photos easy to pick out among the photo¬

heavy pages, and I am thankful that the author made this decision.

Lots of extras are sprinkled into the book. Between the forb and grass section one can find the “Checklist

and Value of Cattle Forage Plants of North Central Texas” as well as a nice little essay on grass by John James In¬

galls (1833-1900). Between the grass and woody sections are “Preference Categories of Deer Food Plants” written

by Steve Nelle, “Checklist and Value of Deer Food Plants of North Central Texas,” and a poem about mesquites by

Frank Grimes. All of this is capped off with an illustrated glossary (mostly about leaf characteristics and grass

morphology), standard textual glossary, list of references, common name index, and scientific name index.

The book itself is printed on thick gloss paper (likely at least 80-100 lb text) where the color images and the

large (likely 12 pt) font really stand out nicely. The book is spiral bound and thus lies flat when open, and the

cover is heavy gloss/laminated card stock. Since I’m almost certain the first run will sell out in quick measure, the

only suggestion I have for the author for his second printing is to switch from the spine-over-spiral cover to a

basic spiral/comb cover. (Spine-over spiral gives spiral books a printable spine by using one continuous sheet that

wraps around the wire spiral/comb to create both covers and a spine, with only the back portion actually attached

to the spiral.) Although it makes the book look great on a shelf, it’s a bit clumsy to handle, and I for one plan to be

using my copy so much that it might NEVER sit on the shelf!— Brooke Byerley Best, PhD, Editor & Botanist,

Botanical Research Institute of Texas, Fort Worth, Texas, U.S.A.

J.Bot. Res. Inst. Texas 8(2): 582.2014

ESTRUCTURA DE UN ZACATAL DE TOBOSO (HILARIA MUTICA: POACEAE)

ASOCIADO A SUSTRATO IGNEO EN EL NORESTE DE COAHUILA, MEXICO

Juan A. Encina-Dommguez

Laboratorio de Sistemas de Informacion Geografica

Universidad Autonoma Agraria Antonio Narro

Buenavista, Saltillo 25315

Coahuila, MEXICO

jaencinad@gmail.com

Jesus Valdes-Reyna y

Jose A. Villarreal-Quintanilla

Departamento de Botanica

Universidad Autonoma Agraria Antonio Narro

Buenavista, Saltillo 25315

Coahuila, MEXICO

RESUMEN

Con el objetivo de evaluar la estructura y diversidad de especies en un zacatal de toboso ( Hilaria mutica (Buckley) Benth.) en la Planicie

Costera del Golfo, se establecieron 20 sitios, en parcelas de 100 m 2 . Se cuantificaron las arbustivas, las herbaceas se midieron en tres parcelas

de 2 m 2 por sitio, en ambos estratos se tomo altura y cobertura de las especies. Se calcularon atributos de densidad, cobertura y frecuencia,

ademas del valor de importancia relativo (VIR) por especie. Se calculo la diversidad con el indice de Shannon-Wiener. La flora esta integrada

por 32 familias, 89 generos y 109 especies, las familias mas importantes son: Asteraceae con 16 especies, Fabaceae (12), Euphorbiaceae (10)

y Poaceae (9). Se registraron 17 especies consideradas como malezas ruderales. Se cuantificaron arbustos espinosos e inermes como: Acacia

rigidula con 625 ind/ha, Opuntia lindheimeri 650 ind/ha, y Aloysia gratissima 469 ind/ha, su densidad es mayor en areas disturbadas debido

al pastoreo. El estrato herbaceo esta dominado por Hilaria mutica con dominancia relativa de 64.96% y VIR de 30.65%, la riqueza se incre-

menta en epoca de lluvias, donde las anuales mas abundantes son: Ambrosia confertiflora, Aphanostephus ramosissimus y Ratibida colum-

nifera. El estrato arbustivo tiene una riqueza de 33 especies, indice de diversidad 2.67 nats. Para el estrato herbaceo se registraron 55 especies

y el indice de diversidad es 3.050 nats. El zacatal presenta evidencias de sobrepastoreo y puede ser invadido por malezas ruderales, y a medio

plazo ser sustituido por el matorral espinoso que crece en areas aledanas.

Palabras Clave: Floristica, pastizal, riqueza de especies, vegetacion

ABSTRACT

With the objective to survey and delineate the structure and diversity of species of a toboso grassland ( Hilaria mutica (Buckley) Benth.) in

the Gulf Coastal Plain (Coahuila, Mexico), 20 sampling sites were established in plots of 100 square meters. The survey quantified shrubs,

while the forbs within 3 plots of 2 square meters per site, for both strata, height and species coverage were measured. Density, canopy cover,

frequency and the value of relative importance (VIR) per species were calculated. The diversity was estimated using the Shannon-Wiener in¬

dex. The flora includes 32 families, 89 genera, and 109 species; the families with the highest number of species are: Asteraceae (16), Fabaceae

(12), Euphorbiaceae (10), and Poaceae (9). There were 19 species recorded as weeds. The density of several shrubs such Acacia rigidula 625

ind/ha, Opuntia lindheimeri 650 ind/ha, and Aloysia gratissima 469 ind/ha is related to the degree of disturbance by cattle grazing. The her¬

baceous stratum is dominated by Hilaria mutica with relative dominance of 64.96% and 30.65% VIR. The richness of the herbaceous flora

increases during the rainy season. The most abundant herbs are: Ambrosia confertiflora, Aphanostephus ramosissimus y Ratibida columnifera.

The shrub stratum has a richness of 33 species, with a diversity index of 2.67 nats. For the herbaceous stratum there were 55 species re¬

corded and the diversity index is 3.050 nats. The grassland shows evidence of overgrazing and being invaded by ruderal weeds. In the short

term it can be replaced by thorn scrub from surrounding areas.

Key Words: Grassland, floristic, species richness, vegetation

Los zacatales son comunidades vegetales con dominancia de especies de la familia Poaceae (gramlneas o

zacates) (Rzedowski 1966), se desarrollan en valles donde los suelos son de mediana profundidad, as! como

en laderas poco inclinadas y mesetas, con amplia distribucion en Mexico y Norteamerica (Rzedowski 1975,

2006). En Coahuila se distribuyen en porciones aisladas que varlan en tamano a traves del estado y ocupan un

area aproximada del 8% de la superficie estatal (Villarreal & Valdes 1992-93). Segun el ordenamiento ecologi-

co de Coahuila, los zacatales ocupan 6.18% de la superficie estatal (ICE 2001) e incluye el zacatal natural o

climatico, as! como el gipsohlo (Pinkava 1984) y halohlo (Estrada et al. 2010) determinados por condiciones

edahcas. Es frecuente que se presenten asociados con otro tipo de vegetacion, pero con mayor frecuencia con el

J. Bot. Res. Inst. Texas 8(2): 583 - 594.2014

584

Journal of the Botanical Research Institute of Texas 8(2)

Matorral Xerohlo, por lo que la composicion de especies es variada, sin embargo, domina el estrato herbaceo,

con arbustos y arboles que crecen aislados.

Los zacatales dominados por el zacate toboso ( Hilaria mutica) son frecuentes en los margenes de lagunas

intermitentes y valles de cuencas endorreicas en areas con suelos arcillosos, limosos y profundos con alto con-

tenido de sales, se ubican en altitudes entre 1,300 a 1,600 m, son propios del oeste de Coahuila (Muller 1947),

centro-este del estado de Chihuahua (Shreve 1942; Henrickson & Johnston 1986), en el Valle de Janos, en el

noroeste de Chihuahua (Vega-Mares et al. 2014), en el noreste de Durango (Gentry 1957; Gonzalez et al. 2007)

y en el noroeste de Nuevo Leon (Briones & Villarreal 2001). Como parte de la composicion se presentan arbus¬

tos bajos y esparcidos de chamizo ( Atriplex canescens), nopales ( Opuntia spp.) y mezquite ( Prosopis glandulosa),

entre otras especies. De acuerdo con Rzedowski (2006), los zacatales de toboso no estan restringidos a suelos

alcalino-salinos, sino que tambien prosperan en otros que tienen drenaje algo dehciente y textura hna.

En Mexico, estos zacatales se han utilizado para el pastoreo intensivo de ganado bovino, los cuales son

de excelente valor forrajero cuando se aprovechan aplicando una administracion adecuada de agostaderos, sin

embargo, el mal manejo al que han estado sometidos, no permite obtener el mayor rendimiento (Rzedowski

2006). Los zacatales naturales se encuentran dentro de los ecosistemas que han sido mas perturbados por la

influencia humana (Hannah et al. 1995). Se ha documentado que el sobrepastoreo impide el desarrollo de

especies forrajeras, propiciando la invasion de plantas indeseables como anuales de tipo ruderal y arbustos

espinosos que son dispersados por el ganado, ademas, de que el pisoteo excesivo causa compactacion del suelo,

lo que reduce la cobertura de la vegetacion (Hobbs & Huenneke 1992; Rzedowski 2006) y, de acuerdo con Kah-

men et al. (2002) altera la composicion, diversidad y estructura de la vegetacion.

En Chihuahua y el oeste de Coahuila la superhcie de los zacatales de Hilaria mutica se ha reducido debido

a la apertura de areas agricolas, ya que algunos zacatales se ubican en terrenos fertiles con baja salinidad y tex¬

tura arcillosa, por lo que son utilizados para cultivar alfalfa, algodon y malz, con elevado suministro de agua a

traves de riegos periodicos (obs. pers.).

En el noreste de Coahuila se presenta un zacatal de Hilaria mutica sobre un afloramiento de roca Ignea,

el cual incluye una elevada riqueza de herbaceas. En esta comunidad se asocian arbustos espinosos e inermes

propios del Matorral Tamaulipeco como Acacia rigidula, Opuntia lindheimeri y Prosopis glandulosa var. glandu¬

losa, su abundancia esta relacionada con el grado de disturbio debido al sobrepastoreo. Con el presente estudio

se pretende evaluar la riqueza y diversidad de especies presentes, ademas de describir la estructura del estrato

herbaceo y arbustivo del zacatal.

Area de estudio

El area de estudio se localiza en el municipio de Sabinas al noreste del estado de Coahuila, con extremos de

latitud entre 27°45' y 26°20'N y de longitud entre 101°40' y 100°30'W (Fig. 1). Forma parte de la provincia hsi-

ograhca Grandes Llanuras de Norte America y la subprovincia Llanuras de Coahuila y Nuevo Leon, conforma-

das por extensas planicies, lomerlos bajos y escasas montanas con intervalos altitudinales entre 150 y 1,000 m

(ICE 2001). La altitud media en el area es de 350 m. Se ubica en la region hidrologica RH-24 “Bravo-Conchos”,

la corriente permanente mas importante es el rlo Sabinas que se origina en la Sierra Santa Rosa, el cual fluye a

la presa Venustiano Carranza (Don Martin). En el area dominan las rocas extrusivas del Pleistoceno formadas

por basalto. Los suelos son aluviales, de textura hna y media, de profundidad variable, moderadamente alcali-

nos, con bajo contenido de materia organica, de color cafe o gris (ICE 2001).

El area se ubica en una planicie donde las altitudes oscilan entre 355 a 415 m, sobresale el cerro de Agua

Dulce, su parte mas alta tiene de 500 m de altitud, la llanura corresponde a un afloramiento Igneo que ocupa

un area aproximada de 29,340 ha, provenientes de roca Ignea extrusiva, tipo basalto, del Cenozoico, de la era

Cuaternaria. Los suelos son arcillosos, que se mezclan, con alta proporcion de arcillas expandibles, forman

grietas anchas y profundas cuando estan secos; posee un 40% o mas de gravas u otros fragmentos gruesos

promediado en una profundidad de entre 50 a 100 cm de la superhcie del suelo (FAO 2007). Como unidad co-

dominante, se presenta el suelo vertisol, asociado con calihcadores del tipo calcarico-endoleptico. Son suelos

de textura hna donde la fase rudica es pedregosa.

Encina et al., Estructura de un zacatal de toboso (Hilaria mutica)

585

Fig. 1. Localization del area de estudio y ubicacion de los sitios de muestreo.

586

Journal of the Botanical Research Institute of Texas 8(2)

El clima de acuerdo con el sistema de clasihcacion de Koppen (modihcado por Garcia 1973) es seco muy

calido y calido BSO(h’). La precipitacion promedio es 454 mm al ano y la temperatura media es 22°C, el mes

mas frlo es enero con 3 a 4°C y el mas caliente es julio, con una variacion de 35 a 37°C, la epoca lluviosa ini-

cia en mayo y termina en octubre, se presenta sequla intraestival entre junio y julio, la temporada de lluvias

coincide con los meses mas calientes del ano. Los vientos dominantes provienen del noreste, en el invierno se

presentan vientos frlos procedentes del hemisferio norte (ICE 2001).

La cubierta vegetal del area segun Villarreal & Valdes (1992-93) es de porte arbustivo, la vegetacion mas

abundante es Matorral Tamaulipeco (Muller 1947), el cual se presenta en planicies y lomerlos, entre 240 y

850 m de altitud, ubicados al este de la sierras Santa Rosa, Obayos y Pajaros Azules, donde dominan arbustos

espinosos y algunos inermes, las especies mas frecuentes son: Acacia rigidula, Celtis pallida , Karwinskia hum-

boldtiana, Leucophyllumfrutescens, Opuntia lindheimeri y Prosopis glandulosa var. glandulosa, las herbaceas mas

abundantes son gramlneas de los generos: Aristida, Bouteloua y Tridens. El area estudiada es parte de ranchos

cinegeticos, donde se presenta una densidad alta de venado cola blanca ( Odocoileus virginianus var. texanus)

ademas de que los zacatales son utilizados para realizar el pastoreo intensivo de ganado bovino.

METODOLOGIA

Para la medicion de la vegetacion se utilizo el metodo de parcela (Mueller-Dombois & Ellenberg 1974). Se le-

vantaron 20 sitios de muestreo donde las especies arbustivas se evaluaron en parcelas de 100 m 2 , mientras que

las especies del estrato herbaceo se midieron en tres parcelas de 2 m 2 por sitio de muestreo, en ambos estratos

se tomo la altura y cobertura de las especies. En cada sitio se registro la altitud y coordenadas geograhcas con

un geoposicionador. Se realizaron colectas de material botanico durante el verano y otono, las cuales fueron

identificadas y depositadas en el herbario ANSM (Saltillo, Coahuila, Mexico) (Holmgren & Holmgren 1990).

Los nombres cientlficos validos as! como los autores son de acuerdo a la base de datos del Missouri Botanical

Garden (http://www.tropicos.org). Se calcularon los atributos de la vegetacion como la densidad, frecuencia y

cobertura de copa para cada especie, los valores relativos de estas variables se sumaron para calcular el valor de

importancia relativo (en lo sucesivo VIR) (Mueller-Dombois & Ellenberg 1974). Tambien se calculo la diver-

sidad con el Indice de Shannon-Wiener (H’), donde se utilizo logaritmo natural (Magurran 2004), as! como la

equitatividad (E) de acuerdo con Pielou (1966) y el Indice de riqueza de especies de Margalef (Magurran 2004).

RESULTADOS Y DISCUSION

Como parte de la flora se registraron 109 especies, distribuidas en 32 familias y 89 generos (ver Anexo 1), esta

cifra representa el 3.58% de la flora reportada para Coahuila (Villarreal 2001). De acuerdo con Rzedowski

(1992) los pastizales y matorrales albergan aproximadamente 6,000 especies (20% de la flora total), el 1.81%

de esas especies se presentan en el zacatal de Hilaria mutica. Las familias mas importantes son: Asteraceae

con 16 especies, Fabaceae (12) y Euphorbiaceae (10) (Tabla 1), la misma tendencia se presenta en el pastizal

halohlo del noreste de Mexico estudiado por Estrada et al. (2010) y de forma general con los pastizales calcihlos

de Guanajuato (Rzedowski & Calderon de Rzedowski 1995). Tales familias son de igual forma, las de mayor

riqueza de especies en pastizales halohlos del Valle de Janos, Chihuahua (Vega-Mares et al. 2014) y en un pas¬

tizal del suroeste de Queretaro (Gomez-Sanchez et al. 2011). Los generos con mayor riqueza, con al menos tres

taxones, son: Acacia, Croton, Euphorbia y Verbena. De la flora observada, las monocotiledoneas representan

el 13.34% y las dicotiledoneas el 86.66%. Con relacion a las formas de vida, dominan las hierbas, de estas, 55

especies son perennes y 17 son anuales, 30 especies son arbustos y 7 crasicaules. De las 32 familias registra-

das, 47% (15 familias) estan representadas por una especie. La riqueza de Asteraceae puede atribuirse a sus

adaptaciones evolutivas como fertilidad alta, ehciencia en dispersion y plasticidad qulmica (Villasenor 1993).

El estrato arbustivo tiene una riqueza de 33 especies, posee un Indice de diversidad 2.675 nats, equitatividad

76.53%. Por su parte, el Indice de riqueza es de 4.846. Para el caso del estrato herbaceo, se registro una riqueza

de 55 especies, posee una diversidad de 3.050 nats, una equitatividad 76.12% y un Indice de riqueza de 6.887.

En el caso del Indice de diversidad obtenido para el estrato arbustivo es medio y para el herbaceo es alto de

acuerdo con el criterio de Margalef (1972) que indican que valores superiores a 3.0 nats son considerados altos.

Encina et al., Estructura de un zacatal de toboso (Hilaria mutica)

587

Tabla 1 . Familias con mas riqueza de generos y especies en la flora del zacatal de Hilaria mutica.

Familias

Numerode generos

Numerode especies

Asteraceae

Fabaceae

Euphorbiaceae

Poaceae

Verbenaceae

Malvaceae

Cactaceae

Brassicaceae

Solanaceae

Lamiaceae

De las 109 especies registradas para el zacatal de Hilaria mutica, 105 especies (96%) estan reportadas

para la flora de las planicies del sur de Texas de acuerdo con Hatch et al. (1990), esto, debido a que comparten

condiciones ecologicas, pues ambas areas se ubican en la provincia florlstica de la Planicie Costera del Noreste,

la cual tiene influencia tropical (Rzedowski 2006). Con respecto a la vegetacion del ecotono para el noroeste de

Nuevo Leon reportado por Briones & Villarreal (2001), tiene una elevada similitud ya que comparte 79 espe¬

cies (71%). Gran parte de las especies de la familia Fabaceae son comunes en el norte del estado de Nuevo Leon

(Estrada et al. 2005). Por otro lado, con los pastizales halohlos del noreste de Mexico estudiados por Estrada

et al. (2010), comparte 17 especies (15.6%), la poca similitud es debido a que estos pastizales se localizan en la

provincia del Altiplano Mexicano y que incluyen flora de abnidad xerica. La mayor parte de los generos de la

flora presentan ahnidad geograhca meridional (Rzedowski 1965), por lo que presentan un vinculo neotropi¬

cal, algunas especies son exclusivas de la provincia florlstica Planicie Costera del Noreste como Acacia rigidula

y Leucophyllum frutescens, entre otras. Especies como Hilaria mutica, Jatropha dioica y Larrea tridentata son

propias de la Altiplanicie Mexicana y extienden su distribucion hasta el noreste de Nuevo Leon (Briones &

Villarreal 2001).

En el area se presentan dos especies bajo estatus de conservacion en la NOM-059-SEMARNAT-2010

(SEMARNAT 2010), las cuales son: Amoreuxia wrightii en la categorla de “en peligro de extincion” y de acuerdo

con Calderon de Rzedowski (1994) la localidad tipo de la especie esta en Texas, donde habita desde el suro-

este de tal estado y en el norte y noreste de Mexico, con distribucion disyunta hasta Yucatan y Sudamerica;

en el area es abundante en el estrato herbaceo del zacatal, con densidad de 2,083 ind/ha, su presencia es mas

evidente en la estacion lluviosa. Se localiz6 tambien a Echinocereus poselgeri, especie con distribucion en el Ma-

torral Tamaulipeco, en el noreste de Mexico (Guzman et al. 2003), registrada para la flora de Texas por Hatch

et al. (1990), esta catalogada como “sujeta a proteccion especial” en la mencionada norma. Por otra parte se

colectaron dos plantas endemicas de la region centro de Coahuila, de acuerdo con Villarreal & Encina (2005)

como: Gaillardia coahuilensis, comun en la epoca de lluvias y Mimosa turneri, con pocos individuos que crecen

aislados. Las especies conocidas en el area estudiada a partir de uno o pocos individuos se les puede consid-

erar raras (Rabinowitz et al. 1986), en el zacatal estudiado se encontraron en esta condicion a las siguientes:

Apodanthera undulata, Euphorbia johnstonii, Jatropha cathartica, Malvella sagittifolia, Mimosa unipinnata y Rivina

humilis.

Como parte de la flora se colectaron 17 especies (15.6% de la flora del zacatal estudiado) consideradas

por la CONABIO (2014) como malezas de tipo ruder al, las que se han registrado en varios estados de Mexico,

la mayorla de las especies ruderales tienen estatus migratorio como nativas de Mexico y solo dos Malvastrum

coromandelianum y Solarium elaeagnifolium son consideradas probablemente nativas. En la Tabla 2 se comparan

13 especies que se presentan en otros pastizales y matorrales de Mexico, de estas, comparte cuatro especies

con los pastizales calcihlos citados para Guanajuato por Rzedowski & Calderon de Rzedowski (1995), ocho

son parte de la vegetacion de la region de Ojuelos, Jalisco reportada por Harker et al. (2008), siete son ruderales

con distribucion en los pastizales halohlos del noreste de Mexico (Estrada et al. 2010), mientras que con los

588

Journal of the Botanical Research Institute of Texas 8(2)

Tabla 2. Comparacion de las principales especies ruderales del zacatal de Hilaria mutica con otros pastizales y matorrales de Mexico.

Especie

Este

estudio

Guanajuato.

Rzedowski &

Calderon de

Rzedowski

(1995)

Ojuelos,

Jalisco.

Harkeretal.

(2008)

Noreste de

Mexico.

Estrada et al.

(2010)

Janos,

Chihuahua.

Vega-Mares etal.

(2014)

Ambrosia confertiflora

Aphanostephus ramosissimus

Apodanthera undulata

Desmanthus virgatus

Euphorbia nutans

Glandularia bipinnatifida

Lepidium virginicum

Parthenium hysterophorus

Portulaca pilosa

Sida abutifolia

Solanum elaeagnifolium

Sphaeralcea angustifolia

Sphaeralcea hastulata

pastizales halohlos del Valle de Janos, Chihuahua estudiados por Vega-Mares et al. (2014) comparte nueve

especies. Algunas malezas como Aphanostephus ramosissimus, Sida abutifolia, Solanum elaeagnifolium y Sphaer-

alcea angustifolia, estan presentes de la mayorla de las localidades, esto debido a que son comunes en el centro-

norte de Mexico, mientras que las otras especies tienen menor rango de distribucion o bien se tienen menos

reportes de su presencia en comunidades naturales.

En areas con sobrepastoreo, el zacatal incluye un estrato arbustivo aislado que integra un matorral abi-

erto, las especies dominantes son arbustos espinosos que poseen alturas de 0.98 a 1.50 m (Fig. 2), los que tienen

mayor densidad y VIR son: Acacia rigidula con 625 ind/ha y VIR de 17.53%, Opuntia lindheimeri con 650 ind/ha

y VIR de 12.84%, ademas del arbusto inerme: Aloysia gratissima con 469 y VIR de 10.17% (Tabla 3), se presenta

un estrato subarbustivo con altura de 0.50 a 0.54 m, donde domina Lippia graveolens con 613 ind/ha y Jatropha

dioica con 706 ind/ha. Se registraron colonias aisladas de Echinocereus stramineus con densidad de 50 ind/ha,

ademas de Echinocactus texensis y Sclerocactus scheeri, las que tienen densidad inferior a 10 ind/ha. En areas

con suelos profundos y areas donde se acumula la humedad se presenta Prosopis glandulosa var. glandulosa, su

altura media es 1.42 m, con densidad de 156 ind/ha y VIR de 4.99%, tales especies lenosas son reportadas por

Mata (2010) para asociaciones del matorral xerohlo ubicado en el centro-este de Coahuila, en el matorral espi-

noso reporta para A. rigidula una densidad de 1,675 ind/ha, la cual es mas elevada que en el zacatal estudiado.

Un zacatal similar es reportado por Briones & Villarreal (2001) para el noroeste de Nuevo Leon, donde el es¬

trato herbaceo alcanza hasta 50 cm de altura y esta representado por Hilaria mutica, crecen arbustos esparcidos

de 2.0 m de alto de los generos Acacia, Opuntia y Prosopis, as! como Flourensia cernua y Eycium berlandieri.

En el estrato herbaceo el zacate toboso ( Hilaria mutica) tiene alta dominancia relativa con 64.963% y VIR

de 30.65%, abundante en los sitios con menor intensidad de pastoreo del ganado bovino, el cual forma “macol-

los” con alturas de 20 a 30 cm (Tabla 4 y Fig. 2). La riqueza del estrato herbaceo se incrementa en la temporada

de lluvias y en areas con mayor disturbio por el pastoreo, dos aspectos mencionados por Pausas & Austin

(2001) como influyentes en la riqueza de especies. De forma codominante se presentan tres especies anuales

como: Ratibida columnifera (con VIR de 7.42%), Ambrosia confertiflora (VIR 6.45%) y Aphanostephus ramosissi¬

mus (VIR 5.95%), las dos ultimas consideradas malezas ruderales, tales especies son dispersadas por el ganado,

el que ademas facilita su establecimiento (Hobbs & Huenneke 1992). Algunas especies perennes que registran

mayores valores de densidad y VIR son: Desmanthus virgatus (con VIR de 3.67%), Bouteloua trifida (VIR 3.47%)

y Phyllanthus polygonoides (VIR 3.46%). El zacate Hilaria mutica se presenta en tres de las cinco asociaciones del

matorral xerohlo estudiado por Mata (2010), es el dominante en el estrato herbaceo del matorral de Prosopis

Encina et al., Estructura de un zacatal de toboso (Hilaria mutica)

589

Fig. 2. Arriba: zacatal dominado por Hilaria mutica con arbustos aislados de Opuntia lindheimeri, principal especie en las areas con sobrepastoreo,

Abajo: zacatal de Hilaria mutica con presencia de arbustos de Acacia rigidula y Opuntia lindheimeri, al fondo se observa el cerro de Agua Duke principal

elevacion del area de estudio.

590

Journal of the Botanical Research Institute of Texas 8(2)

Tabla 3. Atributos estructurales de las especies arbustivas del zacatal de Hilaria mutico.

Especie

Altura media (cm)

Dom. Rel. (%)

Dens, (ind/ha)

Dens. Rel. (%)

Free. Rel. (%)

VIR (%)*

Acacia rigidula

150.93

30.727

625

13.550

8.333

17.537

Opuntia lindheimeri

98.34

13.832

650

14.092

10.606

12.843

Aloysia gratissima

115.82

10.505

469

10.163

9.848

10.172

Lippia graveolens

54.21

5.280

613

13.279

7.576

8.712

Jatropha dioica

50.81

2.737

706

15.312

5.303

7.784

Prosopis glandulosa

142.71

5.537

156

3.388

6.061

4.995

Eysenhardtia texana

134.89

6.260

194

4.201

3.788

4.750

Acacia constricta

94.17

3.573

2.033

4.545

3.384

Larrea tridentata

100.78

4.200

138

2.981

2.273

3.151

Cylindropuntia leptocaulis

69.11

1.782

106

2.304

5.303

3.130

Karwinskia humboldtiana

85.81

2.194

2.033

4.545

2.924

Viguiera stenoloba

73.14

2.187

150

3.252

1.515

2.318

Guaiacum angustifolium

74.07

0.503

1.220

3.788

1.837

Echinocereus stramineus

24.60

0.263

1.084

3.788

1.712

Ziziphus obtusifolia

123.75

0.969

0.542

3.030

1.514

Leucophyllum frutescens

130.70

1.590

1.355

1.515

1.487

Croton incanus

88.16

0.826

119

2.575

0.758

1.386

Mimosa turned

93.13

2.035

1.084

0.758

1.292

Castela erecta

130.00

0.718

0.407

2.273

1.132

Yucca treculeana

175.00

0.487

0.407

2.273

1.055

Echinocactus texensis

12.00

0.003

0.136

0.758

0.299

Sclerocactus scheeri

13.00

0.002

0.136

0.758

0.298

Echinocereus poselgeri

34.00

0.001

0.136

0.758

0.298

Otras 10 especies

3.791

200

4.336

9.848

5.992

Total

100.00

4,613

100.00

* Dom. Rel. = Dominancia relativa, Dens. = Densidad, Dens. Rel. = Densidad relativa, Free. Rel. = Frecuencia relativa, VIR (Valor de impor-

tancia relativo) = Dom. rel. + Dens. rel. + Free. rel. / 3.

glandulosa-Castela texana , el cual tiene una densidad de 26,500 ind/ha, dominancia relativa de 34.64% y VIR

de 18.79%, tales valores son inferiores a los registrados en el presente estudio.

La elevada riqueza de especies herbaceas registrada en el zacatal estudiado esta influenciada ademas por

el tipo de suelo, el cual se deriva de roca Ignea, los cuales aportan mas cantidad de nutrientes (Buckman &

Brady 1970; Pausas & Austin 2001) que incrementan el numero de especies, en especial de herbaceas. Por otra

parte, se ajusta al modelo de disturbio intermedio citado por Hobbs & Huenneke (1992), el cual senala que, a

niveles intermedios de disturbio, la riqueza alcanza su mayor valor.

La distribucion del matorral espinoso en algunos sitios donde el estrato herbaceo esta dominado por

zacate toboso ( Hilaria mutica) al parecer corresponde a una condicion intermedia de sucesion ecologica, in-

ducida por el apacentamiento excesivo, ya que arbustivas como Opuntia lindheimeri, Acacia rigidula y Prosopis

glandulosa var. glandulosa son invasoras comunes en zacatales degradados (Golubov et al. 2001; Sanchez et al.

2007), de igual manera, el caracter invasivo tambien se reconoce para Castela erecta en el sur de Texas (Taylor

et al. 1999). Algunas especies de mezquite ( Prosopis spp.) son consideradas invasoras agresivas de agostaderos

(Andrade et al. 2007). En varios estados del norte de Mexico especies como Prosopis glandulosa var. glandulosa,

Acacia berlandieri, A. constricta y A. rigidula son abundantes sobre grandes extensiones, asociados a comuni-

dades de matorral xerohlo. Algunas especies dominan en areas con disturbio y constituyen un problema en

campos de cultivo abandonados, donde su erradicacion es diflcil, destacando en este grupo Acacia jarnesiana y

Mimosa aculeaticarpa (Estrada & Martinez 2003).

La fuerte presion por el pastoreo desordenado de bovinos ha provocado el sobrepastoreo del zacatal estu¬

diado (obs. pers.), lo cual ha propiciado la invasion de arbustos no apetecibles y especies ruderales (Richardson

et al. 2000, Gomez-Sanchez et al. 2011) y ha disminuido la capacidad forrajera de la comunidad al ocasionar

en algunas areas la conversion del zacatal por un matorral espinoso debido a la invasion de especies lenosas

Encina et al., Estructura de un zacatal de toboso (Hilaria mutica)

591

Tabla 4. Atributos estructurales de las especies herbaceas del zacatal de Hilaria mutica.

Especie

Altura media (cm)

Dom. Rel. (%)

Dens, (ind/ha)

Dens. Rel. (%)

Free. Rel. (%)

VIR (%)*

Hilaria mutica

25.82

64.963

112,917

21.330

5.677

30.657

Ratibida columnifera

29.08

8.178

46,875

8.855

5.240

7.424

Ambrosia confertiflora

14.76

5.661

42,500

8.028

5.677

6.455

Aphanostephus ramosissimus

15.53

3.874

41,667

7.871

6.114

5.953

Desmanthus virgatus

9.08

1.052

27,292

5.155

4.803

3.670

Bouteloua trifida

6.99

1.271

39,167

7.399

1.747

3.472

Phyllanthus polygonoides

9.84

0.581

19,583

3.699

6.114

3.465

Euphorbia prostrata

4.77

0.967

14,167

2.676

3.930

2.525

Sida abutifolia

8.29

0.787

11,875

2.243

3.930

2.320

Argythamnia humilis

5.78

1.094

13,333

2.519

3.057

2.223

Melampodium cinereum

11.90

1.157

12,083

2.283

2.620

2.020

Gutierrezia sphaerocephala

20.95

0.385

14,375

2.715

2.620

1.907

Thymophylla pentachaeta

9.22

0.507

11,667

2.204

2.620

1.777

Thymophylla micropoides

3.68

0.605

17,917

3.384

1.310

1.766

Aneilema karwinskiana

19.88

0.815

4,167

0.787

2.620

1.408

Sphaeralcea hastulata

7.40

0.312

8,125

1.535

2.183

1.344

Parthenium hysterophorus

14.22

0.687

5,208

0.984

2.183

1.285

Urochloa fasciculata

8.58

0.487

8,542

1.614

1.310

1.137

Amoreuxia wrightii

14.69

0.250

2,083

0.394

2.620

1.088

Dalea pogonathera

10.40

0.113

3,542

0.669

2.183

0.988

Gutierrezia texana

25.08

0.415

8,750

1.653

0.873

0.980

Gaillardia coahuilensis

19.75

0.911

3,750

0.708

0.873

0.831

Scutellaria potosina

6.50

0.021

625

0.118

0.873

0.338

Otras 32 especies

4.905

59,167

11.177

28.821

14.968

Total

100.00

529,375

100.00

* Dom. Rel. = Dominancia relativa, Dens. = Densidad, Dens. Rel. = Densidad relativa, Free. Rel. = Frecuencia relativa, VIR (Valor de impor-

tancia relativo) = Dom. rel. + Dens. rel. + Free. rel. / 3.

(Halpern et al. 2010; Rice et al. 2012; Amodeo & Zalba 2013). Desde hace casi cuatro decadas, de acuerdo con

Rzedowski (1975), en Mexico se reconocla que la presencia humana en pastizales producla una fuerte influen-

cia en su extension, composicion y dinamica, sin embargo, la magnitud de estos cambios es diflcil de evaluar.

Por lo anterior se preve, que de continuar el libre pastoreo en el zacatal estudiado, las malezas registradas po-

drlan ser mas abundantes e inclusive reemplazar algunas especies nativas, lo anterior de acuerdo con Baruch

et al. (1989) quienes mencionan que las ruderales son mejores competidoras y capaces de desplazar a las espe¬

cies nativas. El zacatal podrla ser ademas reemplazado por arbustos espinosos como Acacia rigidulay Opuntia

lindheimeri, propios de los matorrales adyacentes, estas son especies que el ganado no consume y dominan en

sitios sobrepastoreados, de acuerdo con Nai-Bregaglio et al. (2002) son llamadas especies crecientes. Por otra

parte, las especies preferidas por el ganado bovino como Hilaria mutica que son mas nutritivas (Sternberg et al.

2000) pueden disminuir o desaparecer de este zacatal.

ANEXO 1

Listado floristico del zacatal de Hilaria mutica en el centra de Coahuila.

Las colectas corresponden al primer autor, los numeros de colecta los antecede la inicial E. Despues del nombre cientifico se cita (A)

para las especies anuales, el resto son perennes. Para las especies consideradas malezas de tipo ruderal de acuerdo con el criterio de

CONABIO (2010) se anota (M).

MAGNOLIOPHYTA

DICOTYLEDONEAE (MAGNOLIOPSIDA)

Acanthaceae: Ruellia occidentalis (A. Gray)Tharp. & Barkley, E1560,

2472, 2501, 2593, 2737; R. nudiflora (Engelm. & A. Gray) Urb.,

El 639,1950, 2659.

Apocynaceae: Mandevilla macrosiphon (Torr.) Pichon, E1952,2408,

2430, 2602, 2640.

Asdepiadaceae: Asclepias asperula (Decne.) Woodson, E 2739.

Asteraceae: Ambrosia confertiflora DC., (M) E 2490a; Aphanoste-

phus ramosissimus DC., (A, M), E 2363, 2403, 2523, 2549, 2975;

Chaetopappa bellioides (A. Gray) Shinners, E 2481,2498,2562;

Journal of the Botanical Research Institute of Texas 8(2)

592

Gaillardia coahuilensis B.L.Turner, (A), E 2422,2849; Gutierrezio

sphaerocephala A. Gray, (A), E 2359; G. texana (DC.) Torr. & A.

Gray, (A), E 1836,2522,2915,2925; Melampodium cinereum DC.,

E 2365,2417,2597,3010; Palafoxia texana DC., (A), E 1944,2738;

Parthenium hysterophorus L., (A, M), E 2461, Ratibida columnifera

(Nutt.) Wooton & Standi., E 2374, 2397, 2405, 2424, 2548;

Tetragonotheca texana A. Gray & Engelm., E 2423, 2528, 2846;

Tetraneuris linearifolia (Hook.) Greene, (A), E 2530; Thymophylla

micropoides (DC.) Strother, (A), E 1987,2431,2460,2939,3102;

T. pentachaeta (DC.) Small var. belenidium (DC.) Strother, E1949,

2744,2914,3020; Viguiera dentata (Cav.) Spreng., (M) E 1571; V.

stelonoba S.F. Blake, E 2493,3116.

Bixaceae: Amoreuxia wrightii A. Gray, E 1947,2399,2583.

Boraginaceae: Heliotropium confertifolium (Torr.) Torr. ex A. Gray,

E 1919, 1985, 2479, 2614, 2649; H. torreyi I.M. Johnst., E 2433;

Tiquilia canescens (DC.) A.T. Richardson, E 1974,2923,3072.

Brassicaceae: Lepidium virginicum L., (A, M), E 2821,3084; Physaria

fendleri (A. Gray) O'Kane & Al-Shehbaz, E 2499, 2697, 3087;

Nerisyrenia incana Rollins, E 2541.

Cactaceae: Cylindropuntia leptocaulis (DC.) F.M. Knuth, E 2963;

Echinocactus texensis Hopffer; Echinocereus poselgeri Lem.; £.

stramineus (Engelm.) F. Seitz; Mammillariaheyden Muehlenpf.;

Opuntia lindheimeri Engelm., E 2959; Sclerocactusscheeri (Salm-

Dyck) N.P. Taylor.

Cleomaceae: Polanisia uniglandulosa (Cav.) DC., (A, M), E 1597,

2401,2535,2578.

Celastraceae: Schaefferia cuneifolia A. Gray, E 2610, 2936, 3026,

3110.

Convolvulaceae: Ipomoea costellatalorr., (A), E 2651.

Cucurbitaceae: Apodanthera undulata A. Gray, (M); Ibervillea tenui-

secta (A. Gray) Small, E 1969, 2526, 2604, 2928.

Euphorbiaceae: Argythamnia humilis (Engelm. & A. Gray) Mull. Arg.,

E 2426, 2534, 2898; Croton incanus Kunth, E 1629,1924, 2414;

C. monanthogynus Michx., (A), E 2442; C. poffs//(Klotzsch) Mull.

Arg., E 2585; Euphorbia johnstonii Mayfield, E 2432; £ nutans

Lag., (A, M), E 2570, 2577; £. prostrata Aiton (A), E 2881, 2994;

Jatropha cathartica Teran & Berland.; J. dioica Cerv., E 2946;

Phyllanthuspolygonoides Nutt, ex Spreng., E 2404,2511,3019.

Fabacea e: Acacia constricta Benth., E 2740; A. gregg/7 A. Gray, E 2354,

2445,2542,3078; A. rigidula Benth., E 3098; Calliandra conferta

Benth., E 1933,2495,3101; Cercidium texanum A. Gray, E 1961,

2927,3107; Dalea aurea Nutt, ex Pursh, E 2398; D. pogonathera

A. Gray var.pogonathera, E 1977,2386,2459,2482,2527,2897,

3006, 3086; Desmanthus virgatus (L.) Willd., (M), E 2381, 2586,

2654, 2743; Eysenhardtia texana Scheele, E 1962, 2413, 2438;

Mimosa turned Barneby, E 1918; M. unipinnata D.B. Parfitt &

Pinkava, E 2407; Prosopis glandulosa Torr. var. glandulosa, E

2794, 2957,3031.

Hydrophyllaceae: Nama hispida A. Gray, E 2370,2835.

Lamiaceae: Salvia ballotiflora Benth., E 1630, 1929, 3013, Scutel¬

laria potosina Brandegee, E 2357,5. microphylla Moc. & Sesse

ex Benth., E 2371.

Malvaceae: Abutilon wrightii A. Gray, E 2415; Herissantia crispa (L.)

Brizicky, (M) E 1644, 1931, 2458, 2480, 2933, 2942; Hibiscus

coulteri Harv. ex A. Gray, E 1988,2467,2573,2903; Malvastrum

coromandelianum (L.) Garcke, (M), E 2394; Malvella sagittifolia

(A. Gray) Fryxell, E 2360; Sida abutifolia Mill., (M), E 1937,2454,

2529,2734; Sphaeralcea angustifolia (Cav.) G. Don, (M), E 2882,

2974, 2993; S. hastulata A. Gray, (M), E 2584.

Oleaceae: Forestiera angustifolia Torr.

Phytolaccaceae: Rivina humilis L., E 2490.

Portulacaceae: Portulaca pilosa L., (A, M), E 2636,2918,3081

Rhamnaceae: Karwinskia humboldtiana (Schult.) Zucc., E 1960,

2373, 2543, 2552, 2666, 2937, 3015, 3113; Ziziphus obtusifolia

(Hook, ex Torr. & A. Gray) A. Gray.

Rubiaceae: Hedyotis nigricans (Lam.) Fosberg, E 2532.

Rutaceae: Thamnosma texanum (A. Gray) Torr., E 2469, 2567,

2647, 2943.

Scrophulariaceae: Leucophyllumfrutescens (Berland.) I.M. Johnst.,

El 587,1917, 2491,2571,3001.

Simaroubaceae:Casfe/aerectaTurpin subsp. texana (Torr. & A. Gray)

Cronquist, E 2652, 2926, 2965, 2989, 3039, 3109.

Solanaceae: Chamaesaracha crenata Rydb., E 2380,2694; Quincula

lobata (Torr.) Raf., E 2379, 2569; Solanum elaeagnifolium Cav.,

(M) E 2657, 2813.

Talinaceae: Talinum aurantiacum Engelm., E 2366,2566,3010,3029.

Verbenaceae: Aloysia gratissima (Gillies & Hook.) Tronc., E 1965,

2355, 2589, 2689, 3032; Citharexylum brachyanthum (A. Gray

ex Hemsl.) A. Gray, E 1963,1972,2609,2615,2615,2935; Glan-

dularia bipinnatifida (Nutt.) Nutt., (A, M), E 2671,2831,3095; G.

delticola (Small) Umber, (A), E 2489; Lantana camara L., E 1627,

1973,2565,2804; L. macropoda Torr., E 1564,1626,1966,2392,

2608a, 3002; Lippia graveolens Kunth, E 1625,1928,1964,2412,

2434, 2440, 2492, 2524, 2594; Verbena neomexicana (A. Gray)

Briq., E 2356, 2531,2682.

Zygophyllaceae: Guaiacum angustifolium Engelm., E 2421,2538,

2954, 3041; Larrea tridentata (Sesse & Moc. ex DC.) Coville, E

2687, 2795, 2944.

MONOCOTYLEDONEAE (LILIOPSIDA)

Asparagaceae: Yucca treculeana Carriere

Amaryllidaceae: Cooperia drummondii Herb., E 3008.

Commelinaceae: Aneilema karwinskiana Woodson, E 2579, 2638,

2741; Commelina erecta L. var. angustifolia (Michx.) Fernald,

E 2418.

Poaceae: Aristidapurpurea Nutt. var. purpurea, E1981,2637; Boutel-

oua mf/c/o Thurb., E 1980,2435,2599,2653; Digitaria californica

(Benth.) Henrard, E 1978, 3017; Dinebra panicea (Retz.) Ohwi

subsp. mucronata (Michx.) P.M. Peterson & N. Snow, E 2568,

2581; Hilaria mutica (Buckley) Benth., E 1946,1975,2402,2439,

2533, 2582; Panicum hallii Vasey var. hallii, E 1970, 2419, 2682;

Pappophorum bicolor E. Fourn., E 1955,2909,2948; Tridens texa-

nus (S. Watson) Nash, E 2400, 2420, 2580; Urochloa fasciculata

(Sw.) R.D. Webster, (A, M), E 1959,2395,2453,2596,2990,3089.

AGRADECIMIENTOS

Estamos agradecidos con el herbario ANSM por las facilidades para la consulta e identificacion de especies

registradas en esta investigacion. Se agradece a dos revisores anonimos quienes aportaron valiosos comen-

Encina et al., Estructura de un zacatal de toboso (Hilaria mutica)

593

tarios a la primera version. Efren Mata Rocha apoyo en la toma de datos y captura de la informacion de campo.

Agradecemos a Felipe N. Hernandez Soto la elaboration de la figura 1.

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JARILLA CHOCOLA (CARICACEAE), UN NUEVO GENERO Y ESPECIE

PARA LA FLORA DE EL SALVADOR

Frank Sullyvan Cardoza Ruiz

Consultor independiente

Magister ofScientiae en Monejo y Conservation

de Bosques Tropicalesy Biodiversidad

Centro Tropical Agronomico de

Investigacion y Ensenanza (CATIE)

San Salvador, EL SALVADOR

fscardozan@gmail.com

Jose Ledis Linares

Catedratico de botanica

Centro Universitario Regional

del Litoral Atlantico (CURLA)

Universidad Nacional Autonoma de Honduras (UNAH)

La Ceiba, HONDURAS

Tin aresj_ 98@yahoo. com

Ana Eugenia Aguilar Grijalva

Biologa, Universidad de El Salvador (UES)

Ministerio de Medio Ambiente y Recursos Naturales (MARN)

San Salvador, EL SALVADOR

aeugenia_bio@yahoo.com

RESUMEN

Se reporta la especie Jarilla chocola Standi. (Caricaceae) que constituye el primer registro del genero para El Salvador. Las localidades de

colecta se convierten en las mas orientales para todo el genero, conocido solo de Mexico y Guatemala; representando una importante e

interesante adicion a la flora salvadorena. Se describe el genero tal como ocurre en el pais y se ilustra fotograficamente por primera vez

la especie. Asimismo, se presenta informacion ecologica de las localidades donde se ha encontrado el genero hasta ahora en El Salvador.

Palabras clave: El Salvador, Caricacea e, Jarilla chocola, distribucion, Mexico

ABSTRACT

Jarilla chocola Standi. (Caricaceae) is reported new to the flora of El Salvador. The collection localities represent the easternmost known col¬

lections from Mexico and Guatemala, and represent an important and exciting addition to the Salvadoran flora. Jarilla chocola is described

in detail along with ecological and habitat information and photographs are provided.

Keywords: El Salvador, Caricacea e, Jarilla chocola, distribution, Mexico

INTRODUCCION

El Salvador, ha sido durante mucho tiempo el pals mas pobremente colectado de la region Centroamericana

y probablemente sea el mas desconocido en el aspecto florlstico. Esto ha generado un importante atraso con

respecto a los demas palses vecinos, siendo el mas pequeno (21,040.79 kilometros cuadrados), el mas densa-

mente poblado (6.3 millones de habitantes) y probablemente el mas deforestado del hemisferio despues de

Haiti; siendo esto, la falta de interes para investigadores de otros palses que preberen concentrar sus esfuerzos

en areas relativamente bien conservadas con abundante vegetacion (Linares 2003). En un area tan pequena,

la demanda por recursos se hace cada vez mas prioritaria y la presion sobre los mismos mas crltica, manifes-

tandose en el deterioro de los ecosistemas, perdida de biodiversidad y altos niveles de contaminacion en sus

recursos (MARN 2005).

Los bosques primarios han desaparecido casi totalmente y en la franja costera todavla quedan restos de

la antigua selva tropical, con areas de mangles que en algunos casos el deterioro por actividades agricolas y

deforestacion es evidente. Sin embargo, debido a lo accidentado del relieve y a la continua actividad volcanica,

subsisten en todo el territorio pequenas areas o relictos donde aun sobrevive algun tipo de vegetacion y en don¬

de se desarrollan diferentes etapas de sucesion, encontrandose generalmente en pendientes escarpadas o en

recientes campos de lava y la presion del hombre, sobre ellas, es relativamente pequena (Linares 2003). A pesar

J. Bot. Res. Inst. Texas 8(2): 595 - 601.2014

596

Journal of the Botanical Research Institute of Texas 8(2)

de esta problematica ambiental y el limitado territorio, existe una considerable riqueza en los ecosistemas por

su posicion geograhca, latitudinal, historia geologica y su compleja topografla (MARN 2005).

Genero Jarilla

El genero Jarilla de la familia Caricaceae, fue descrito por primera vez en 1832 por La Llave bajo el nombre de

Mocinna. Desde la publicacion inicial del genero se han descrito diferentes especies, subespecies y variedades,

todas las cuales fueron posteriormente concentradas en un solo taxon Jarilla heterophylla Rusby. Sin embargo,

es un genero de taxonomla complicada; porque algunos autores solo reconoclan una especie variable, mientras

que otros distinguen tres: J. caudataj. chocola, J. heterophylla y un hlbrido natural: J. caudata x J. heterophylla.

Estas especies se desarrollan en los bosques caducifolios de Mexico y Guatemala (Romero 2013). Jarilla, esta

distribuida principalmente en Mexico, con una especie (J. chocola) que se extiende hasta Guatemala (Diaz &

Lomeli 1992).

DESCRIPCION DE LA ESPECIE

Jarilla chocola Standi. Field Mus. Nat. Hist., Bot. Ser. 17(2):200. 1937. (Fig. 1). Tipo: MEXICO: Chihuahua: Guasaremos,

Rio Mayo, 10 Aug 1936, Howard Scott Gentry 2366 (holotype: F).

Ilustraciones. —Diaz Luna y Lomeli Sencion, Acta Bot. Mex. 20:85, t. 3,1992.

Plantas erguidas de 20-50 cm de alto, generalmente sin ramificaciones. Tuberculos fusiformes de cerca de 10

cm, ocasionalmente 20 cm de longitud. Tallo aereo verde de 1-8 mm de diametro, suculento, generalmente

glabro. Hojas con un peclolo de 3-20 cm de longitud y de 0.5-4 mm de diametro, con presencia de latex en la

base del peclolo. Limbo glabro, de 5-17.5 cm de longitud y de 5-18.3 cm en su parte mas ancha, generalmente

mas ancho que largo, de color verde oscuro en el haz y mas claro en el enves, generalmente 3-5 lobulado,

bordes lisos, los lobulos pueden ser insinuados o profundamente hendidos, a veces subdivididos, de apice agu-

do, redondeado o acuminado, con salientes pronunciadas o dientes. Base truncada, concava, obtusa, cordada,

redondeada o bilobulada, a veces con salientes hasta de 2 cm de largo. Borde entero o con salientes aristadas,

hendido con lobulos redondeados o agudos. Apice acuminado o caudado. Con 3-5 nervaduras basales, que

generalmente terminan en los lobulos.

Inflorescencias masculinas con 8-40 flores en un pedunculo de 6.8-20.5 cm de longitud, filiforme o has¬

ta de 3 mm de diametro. Flor masculina con el caliz de 0.4-1.3 mm de largo, los segmentos ellpticos de apice

agudo. Corola de color bianco de 5.1-8.6 mm de largo, tubo de 2.3-4.8 mm de largo, con lobulos de aproxima-

damente la misma longitud, de 1.3-1.7 mm de ancho y apice redondeado. Estambres del grupo superior con

anteras de 1.4-1.8 mm de largo y hlamentos de 1.2-1.4 mm de longitud; el grupo inferior con anteras de 1.8-1.9

mm de largo y hlamentos de 0.3-0.7 mm de longitud. Rudimento del ovario de 3 mm de largo.

Flores femeninas solitarias, sobre un pedunculo filiforme de 8-23 mm de largo. Caliz con segmentos

triangulares de apice agudo, de aproximadamente 1.4 mm de largo, alternos a los segmentos corolinos. Corola

con lobulos largamente ellpticos de apice redondeado, de 8-9 mm de largo y de 2-3 mm de ancho. Ovario de

forma ascoide con base truncada, presenta exteriormente cinco alas longitudinales prolongadas cada una en

un apendice basal, alternandose con una sutura interalar; estigmas densamente papilosos, de forma sigmoide,

de 2.2-2.9 mm de largo por 0.4 mm de diametro (Fig. 2)

Frutos solitarios en un pedunculo de 0.2-3.2 cm de longitud; globosos-alargados, ascoides u ovoides, de

color verde uniforme, de 2.3-7 cm de longitud total y de 1.6-4 cm de diametro en su parte media; su porcion

basal truncada o concava, sin cuello; presenta cinco alas longitudinales de 9 mm de alto que se prolongan en la

base en cinco apendices de 4-9 mm de longitud; su porcion apical redondeada, prolongandose en un apendice

de 3-5 mm de longitud en donde convergen las alas y las suturas intercalares (Fig. 3). Con 40-100 semillas por

fruto. Semillas de 3.2-5 mm de largo con testa de color negro o pardo claro, superhcie casi lisa con granulacio-

nes y surcos semireticulados mas obscuros ligeramente hendidos, porcion anterior aguda, observandose en su

cara ventral un arilo alargado de color claro y sabor agridulce con olor afrutado acido.

Habitat. —Esta asociado a bosque tropical deciduo y subdeciduo, bosque de encino, matorral xerohlo,

pastizal y formas perturbadas de estos tipos de vegetacion, en altitudes que van de 330 a 2,700 msnm. En estos

Cardoza et al., Jarilla chocola en El Salvador

597

Fig. '\. Jarilla chocola Standi. A. Planta completa en su habitat natural; B. Flores; C. Botones; D. Latex. Fotos: Frank Sullyvan Cardoza.

598

Journal of the Botanical Research Institute of Texas 8(2)

ecosistemas es donde se encuentra una diversidad muy apreciable que posee especies aun no registradas para

El Salvador como el caso de Jarilla chocola. Esta especie herbacea perenne es un habitante inconspicuo de estos

tipos de bosques y probablemente debido a su porte pequeno, su floracion eflmera y al hecho de perder toda la

parte aerea durante gran parte del ano, ha pasada desapercibida durante largo tiempo. Ademas, estos tipos de

bosque han sido relativamente poco colectados en el pals y en muchos casos son considerados como de poca

diversidad e importancia. En El Salvador, el genero crece en selvas bajas caducifolias, asociadas a antiguos

campos de lava en grandes acumulaciones de humus y en una pequena franjas de vegetacion de transicion

entre la selva baja y el bosque de robles ( Quercus spp.). Estos bosques tienen una estacion seca muy fuerte con

escasas precipitaciones. La distribucion altitudinal varla desde los 250 hasta los 800 msnm.

Fenologia .—El tallo aereo generalmente emerge a principios de junio; florece desde principios de julio

hasta fines de agosto (Fig. 3); fructihcacion, mediados de julio hasta fines de septiembre (Diaz & Lomeli 1992).

En El Salvador los tallos emergen muy a principio de la temporada lluviosa (principios de mayo) y la floracion

se observa en las primeras semanas de abril y fructihcacion es en junio del siguiente ano.

Material examinado. —EL SALVADOR. Santa Ana: Municipio Texistepeque, Canton Valle Nuevo. Localidad: Cerros Pedregosos de Valle

Nuevo, 4 km al S de Texistepeque, 14°05 , 26.54"N, 89°30 , 25.14"W, 10 Ago 2009J.L. Linaresy C.A. Martinez 13882 (MEXU); Municipio Texis¬

tepeque, Canton Cujucuyo, Caserio Valle Nuevo, Aproximadamente 14 km al nor-este de la ciudad de Santa Ana en carretera CA-12 hacia

Metapan, cerca de entrada principal al Canton Cujucuyo. Coordenadas 14 o 05'25"N y 89°30T5"W, vegetacion bosque tropical semideciduo

en zona perturbada en areas de potreros a una altitud 450 msnm, 2Jun 2013, J.L. Linares, Frank Sullyvan CardozayA. Eugenia Aguilar 15,025

(MEXU); Municipio de Metapan, Aproximadamente 4.5 km al sur-este de la ciudad de Metapan en carretera CA-12 (Alrededores de una

pequena laguna al lado oriente de la carretera a Santa Ana), coordenadas 14 0 17'43.76"N y 89 0 27'47.74"W, vegetacion bosque sub-tropical se¬

mideciduo en zona de pendientes con rocas basalticas creciendo en el sotobosque entre mucha hojarasca a la sombra de Ficus ovalis, altitud

472 msnm, 13 Abr 2014 J.L. Linares, Frank Sullyvan CardozayA. Eugenia Aguilar 16,005 (MEXU).

Etimologia .—El genero fue establecido en 1921 por Rusby, por el nombre comun de “jarrilla” dado en el oc-

cidente de Mexico a la especie Jarilla heterophylla. El eplteto chocola, probablemente aluda a algun nombre

comun dado en Guatemala a esta planta.

Caracteristicas de la especie .—Esta especie se caracteriza por su tallo erguido, ovario y fruto con alas lon-

gitudinales y hojas generalmente palmadas, mas anchas que largas. Las semillas requieren de un perlodo de

reposo para presentar germinacion aceptable; la emergencia ocurre entre 10 y 17 dlas despues de la siembra

(Diaz & Lomeli 1992; Willingham & White 1976). Segun Diaz & Lomeli (1992) Jarilla chocola crece en bosque

tropical deciduo, y zonas perturbadas de estos tipos de vegetacion, a una altitud de 330-1200 msnm. En El

Salvador esta especie crece en selvas bajas caducifolias (bosque tropical deciduo), selvas bajas subcaducifolias

(bosque tropical subdeciduo) entre 200-800 msnm y en ecotonos de bosque de pino-roble y selva baja cadu-

cifolia a una altura cercana a 1000 metros. Algunos nombres comunes son “chocola,” “kapiya” (Chihuahua);

“bonetillo” (Jalisco); “berenjena” (Michoacan); “papayillo” (Nayarit); “chocola,” “kapiah” (Sonora); “talguen-

to” (Guatemala). En El Salvador no se han registrado nombres, ya que ha pasado desapercibida durante mucho

tiempo.

Distribucion geografica .—Se encuentra en Mexico en los estados de Chihuahua, Sonora, Sinaloa, Nayarit,

Jalisco, Michoacan y Chiapas. En Guatemala se le conoce del departamento dejutiapa (Diaz & Lomeli 1992).

En El Salvador se ha colectado en los departamentos de Ahuachapan y Santa Ana, especlhcamente en la parte

norte del macizo del Imposible (departamento Ahuachapan) en selvas bajas junto con especies tlpicas de este

tipo de vegetacion como Bursera simaruba, Spondias purpurea , Spondias mombim, Machaerium spp. y Com-

bretum spp. En el departamento de Santa Ana, se ha colectado en la parte norte en las cercanlas de la Laguna

de San Diego creciendo en acumulacion de humus en rocas volcanicas recientes con las especies Gyrocarpus

jatrophifolius, G. mocinoi, Omphalea oleifera, Poeppigia procera, Bursera simaruba y Jacaratia mexicana. En las

cercanlas de Candelaria de la Frontera, crece tambien en humus acumulado en suelos blancos probablemente

originados de tobas volcanicas del terciario junto con especies caracteristicas de la selva baja caducifolia. Siem-

pre en Santa Ana, se ha encontrado en el Canton Cujucuyo creciendo en vegetacion de bosque tropical semide¬

ciduo; especlhcamente en zonas perturbadas en areas de potreros a una altitud 450 msnm asociada con arboles

dispersos de Guazuma ulmifolia, Luehea Candida, Spondias purpurea, Bursera simaruba, Ficus ovalis, Sapranthus

Cardoza et al., Jarilla chocola en El Salvador

599

Fig. 2. Jarilla chocola Standi. A. Detalle de las hojas; B. Detalle de las flores e inflorescencia. Fotos: Frank Sullyvan Cardoza.

f\G.3. JarillachocolaStan<\\. A. Plantascontuberculoyfruto; B.Tuberculofusiformescon parte aerea;C. Detalledelosfrutos; D. Frutosecoy verdecon

mediciones; E. Sacando tuberculo en su habitat. Fotos: Frank Sullyvan Cardoza.

600

Journal of the Botanical Research Institute of Texas 8(2)

Fig. 4. Distribution geografica y localidades en Mexico, Guatemala y El Salvador deJarillachocola Standi

violaceus, Calycophyllum candidissimum. Suelos rocosos y expuestos de tobas volcanicas con pendientes entre

25-40% (Fig. 4).

CONCLUSIONES

Esta especie es poco frecuente considerandose extinta en muchas zonas del pals debido a la degradation y frag¬

mentation de su habitat. El Estado de acuerdo con los criterios de la Lista Roja de Especies Amenazadas (UICN

2012) y segun las observaciones de los autores, encontrada en areas de condiciones muy especlficas sujetas a

impactos y perturbaciones; se considera que la categorla deberla ser Vulnerable, VU Al(d).

AGRADECIMIENTOS

Nuestra gratitud a los dos revisores anonimos por sus valiosos comentarios y observaciones. Tambien a Barney

Lipscomb por su valiosa revision y apoyo en esta publication que contribuye a la divulgation del conocimiento de

la flora salvadorena. Un sincero agradecimiento tambien a un re visor anonimo por sus sugerencias y comentarios.

REFERENCIAS

Diaz, C. & J. Lomeli. 1992. Revision del Genero Jarilla Rugby (Caricaceae). Acta Bot. Mex. 20:77-99.

Linares, J.L. 2003 [2005]. Listado comentado de los arboles nativos y cultivados en la republica de El Salvador. Ceiba

44(2):105-268.

MARN (Ministerio de Ambiente y Recursos Naturales de El Salvador). 2005. Estado del Conocimiento de la Biodiversidad en El

Salvador. MARN, San Salvador, El Salvador.

Cardoza et al., Jarilla chocola en El Salvador

601

Romero, J.A. 2013. Manejo y conservacion de germoplasma de la familia Caricaceae. Postgrado de Recursos Geneticos y

Productividad. Colegio de Postgraduados. Campus Montecillo.Tesis para Doctor en Ciencias.

UICN (Union Internacional para la Conservacion de la Naturaleza). 2012. Categorias y criterios de la lista roja de la UICN: Ver¬

sion 3.1. Segunda edicion. Gland, Suiza y Cambridge, Reino Unido.

Willingham, B.C. & G.A. White. 1976. Agronomic evaluation of prospective new crop species. V. Jarilla chocola: A protein¬

ase source. Econ. Bot. 30(3):189-192.

602

Journal of the Botanical Research Institute of Texas 8(2)

BOOK NOTICE

Jennifer Ackerfield. Spring 2015. The Flora of Colorado, (pbk). Botanical Research Institute of Texas Press,

1700 University Drive, Fort Worth, Texas 76107 U.S.A. (Orders: shop.brit.org, 1-817-332-4441). $65.00,

approx. 880 pp., color plates, distribution maps, 6.5" x 9.5".

From the Publisher: Colorado has a rich diversity of ecosystems and vegetation communities, ranges from 3,500

ft to over 14,000 ft in elevation, and ultimately supports over 3,300 vascular plant species. This book is a com¬

prehensive guide to the vascular plants in Colorado, and contains descriptions, distribution maps, habitat in¬

formation, flowering times, and elevation ranges for all species. Color photographs for nearly V 3 of the species

are also included. The aim of this guide is to enable students, researchers, amateur and professional botanists,

or anyone interested in the flora of Colorado to successfully identify plants with confidence and satisfaction.

JENNIFER ACKERFIELD has been studying the flora of Colorado for nearly 20 years. During this time, she

has traveled extensively across the state of Colorado documenting its rich floristic diversity. She is currently

the collection’s manager at the Colorado State University herbarium and teaches Plant Identification at CSU.

She has served on the board for the Colorado Native Plant Society, and written several articles on the flora of

Colorado. She enjoys hiking, photographing wildflowers, botanical illustration, and educating students on the

wonderful world of botany.

J.Bot. Res. Inst. Texas 8(2): 602.2014

REDESCUBRIMIENTO Y NUEVOS REGISTROS DE CUSCATLANIA VULCANICOLA

(NYCTAGINACEAE) PARA EL SALVADOR

Jose Ledis Linares

Catedratico de botdnica

Centro Universitario Regional

del Litoral Atlantico (CURLA)

Universidad Nacional Autonoma de Honduras (UNAH)

La Ceiba, HONDURAS

jose.linares@unah.edu.hn

Frank Sullyvan Cardoza Ruiz

Consultor independiente

Magister Scientiae en Manejoy Conservacion

de Bosques Tropicalesy Biodiversidad

Centro Tropical Agronomico

de Investigacion y Ensenanza (CATIE)

San Salvador, EL SALVADOR

fscardozan@gmail.com

Patricia Hernandez-Ledesma

Departamento de Botdnica, Instituto de Biologfa

Universidad Nacional Autonoma de Mexico (UNAM)

Mexico, D.F., MEXICO

phl@ibunam2. i biolog ia. unam. mx

RESUMEN

En el transcurso de varias decadas, el botanico Paul C. Standley realizo diversos tratamientos y describio una gran cantidad de especies de

Nyctaginaceae; que incrementaron el conocimiento de la familia en Norte y Centroamerica, convirtiendose en el principal especialista

hasta el momento. En su expedicion por El Salvador en 1922, colecto en la base del Volcan de San Vicente una peculiar nictaginacea de flores

vistosas, que describio en 1923 como Cuscatlania vulcanicola, unica especie del genero que permanecio practicamente desconocida para los

taxonomos, pues solo se conocian dos ejemplares, el holotipo y un isotipo. Se presentan por primera vez fotografias de plantas vivas de esta

especie, despues de 91 anos de haberse publicado, considerandose extinta por no haberse reportado ni en la zona original de colecta ni en

otros lugares. Sin embargo, ahora se reportan nuevos registros de otra localidad en el Occidente del pais (Sonsonate), lo que representa un

redescubrimiento para la flora salvadorena. Tambien se presenta una descripcion mas detallada de esta especie, haciendo algunos comen-

tarios sobre su morfologia, taxonomia y proporcionando datos de distribucion y habitat.

Palabras clave: Cuscatlania, nictaginacea, redescubrimiento, Sonsonate, Standley, El Salvador

ABSTRACT

Over the course of several decades botanist Paul C. Standley worked on various taxonomic treatments and described a large number of spe¬

cies that increased our knowledge of the Nyctaginaceae family in North and Central America, becoming the leading specialist so far. In his

expedition to El Salvador in 1922, a peculiar Nyctaginaceae with showy flowers was collected at the base of Volcan San Vicente, which he

described in 1923 as Cuscatlania vulcanicola, which is only species of the genus and remained virtually unknown to taxonomists because

only two specimens (the holotype and isotype) are known. Presented for the first time here are photographs of living plants of this species,

after 91 years of being considered extinct since it has not been reported either in the original collection area or elsewhere since the original

publication. But now new records of other localities in the west of the country (Sonsonate) are reported, representing a rediscovery for the

Salvadoran flora. A more detailed description of this species is also presented, with some comments on the morphology, taxonomy, and

distribution and habitat data.

Key Words: Cuscatlania, rediscovery, Nyctaginaceae, Sonsonate, Standley, El Salvador

INTRODUCCION

En las exploraciones y colectas botanicas que realizo Paul C. Standley (1884-1963) en Centroamerica, particu-

larmente en marzo de 1922 en El Salvador; colecto en el Departamento de San Vicente, en la base del Volcan

San Vicente, una interesante nictaginacea de flores llamativas de color rosado, siendo una nueva especie para

la ciencia que describio en 1923 como Cuscatlania vulcanicola.

Standley realizo diversos tratamientos y describio una gran cantidad de especies de Nyctaginaceae (p.

ej. Standley 1909, 1911, 1918, 1931) que incremento el conocimiento de la familia en Norte y Centroamerica,

J. Bot. Res. Inst. Texas 8(2): 603 - 609.2014

604

Journal of the Botanical Research Institute of Texas 8(2)

convirtiendose en el principal especialista hasta el momento. Este hecho fue muy relevante para el estudio de

la familia, pues ademas se incremento la diversidad generica con este nuevo descubrimiento. Hasta enton-

ces, Cuscatlania vulcanicola se conocla solo de la localidad tipo; inclusive, en la obra “Flora Salvadorena” que

Standley realizo junto con Salvador Calderon (Standley & Calderon 1941) y en la cual culminaron sus estudios

en El Salvador, para la especie se menciona la misma colecta de 1922: Standley 21678 , que corresponde al ejem-

plar tipo.

Estas especie permanecio durante todas estas decadas desconocida para los taxonomos, pues solamente

se conoclan dos ejemplares, el holotipo (Fig. 1) depositado en el United States National Herbarium (US) y el

isotipo, depositado en el herbario del Missouri Botanical Garden (MO). En este artlculo, presentamos una

descripcion mas detallada y fotograflas de Cuscatlania vulcanicola por primera vez desde su descripcion en

1923; es decir, 91 anos despues de su publicacion. Durante todo este tiempo, aparentemente nunca habla

sido reportada, hasta que el primer autor (Jose Linares) la colecto en el departamento de Sonsonate. Tambien

hacemos algunos comentarios sobre su morfologla y taxonomla y proporcionamos los datos de distribucion y

habitat.

El redescubrimiento

Durante los recorridos realizados por medio del Proyecto “Mejor Manejo y Conservacion de Cuencas Hi-

drograhcas Crlticas” (MMCCHC), al occidente del pals en los meses de diciembre de 2007 y enero 2008,

Linares y Cardoza colectaron a Cuscatlania vulcanicola aparentemente por primera vez desde que Standley

la colecto hace mas de 85 anos. Es as! como Standley publico en el Journal of the Washington Academy of Sci¬

ence, en 1923, la especie Cuscatlania vulcanicola. El ejemplar tipo se encuentra depositado en United States

National Herbarium (US) con el numero 21,678. A diferencia de Standley, Cuscatlania vulcanicola se colecto en

el departamento de Sonsonate, municipios de Izalco y Caluco, mientras que la localidad tipo es en el Departa¬

mento de San Vicente, municipio de San Vicente en el centro oriente del pals; ahora estos nuevos datos son un

redescubrimiento y representan nuevos registros para El Salvador, ademas de ampliar su area de distribucion

en el pals. A continuacion se presenta la descripcion de la especie y las primeras fotograflas (en su habitat y

cultivadas) despues de 91 anos de su publicacion. Durante uno de los recorridos se encontro una poblacion

en el Area Natural Protegida de San Marcelino, especlhcamente en el bosque de Las Lajas (Canton Las Lajas,

Departamento de Sonsonate), probablemente con mas de 30 plantas (Linares 2009). Este importante hallazgo

de redescubrimiento botanico hace enfasis en la importancia de la localidad como un reservorio de plantas

raras y unicas en el mundo (Fig. 2).

DESCRIPCION DE LA ESPECIE

Cuscatlania vulcanicola Standi., J. Washington Acad. Sci. 13(20):437.1923. Tipo: el Salvador. Departamento de San

Vicente: collected at 500m, in a ravine near the base of the San Vicente Volcano, Mar 1922, Standley 21678 (holotipo: US-imagen;

isotipo: MO).

Hierba postrada, ascendente o decumbente, tallo pardo oscuro, brilloso, ligeramente estriado, glabro, con

ramas puberulentas o vellosas hacia el apice. Hojas distribuidas a lo largo de todo el tallo, opuestas, evidente-

mente de diferente tamano las de cada par, ovadas, oblongo ovadas u oblongo-lanceoladas, glabrescentes,

las hojas jovenes pubescentes, con tricomas glandular-estipitados, septados, hialinos, distribuidos en toda

la lamina y nervadura; apice agudo, acuminado o largamente acuminado, base cuneada, asimetrica, margen

entero; hojas basales 6.0-11.0 cm de largo, 3.0-4.8 cm de ancho, reduciendose de tamano hacia el apice; peclo-

los 1.0-3.0 cm, pubescentes en las hojas jovenes. Inflorescencias cimosas, con flores casmogamas terminales

y cleistogamas axilares, no ramihcadas formando cimas compactas y densas, las inflorescencias de flores

casmogamas con numerosas bracteas foliaceas verde purpureo en el haz y purpureas en el enves.

Flores sostenidas por 4-8 bracteas foliaceas verde purpureas, ovadas, lanceoladas, oblanceoladas a

oblongo-ellpticas, 10-12 mm de largo, 2.5-5.5 mm de ancho, pubescentes, con tricomas glandular-estipitados,

septados, hialinos, margen ciliado, sesiles, formando un involucro de bracteas libres sobre un pedunculo de

ca. de 6 mm de largo, pubescente, sosteniendo 1—2(—3) flores (Fig. 2). Perianto infundibuliforme, magenta,

Linares et al., Cuscatlania vulcanicola en El Salvador

605

ailiiili"

hh-

No 2/^/^ paul c - standley - 6

C/3*

Fig. 1. Imagen tipo de Cuscatlania vulcanicola Standi. Depositada en el US National Herbarium en 1922.

606

Journal of the Botanical Research Institute of Texas 8(2)

Fig. 2. Cuscatlania vulcanicola Standi. Plantas y flores como ocurren en el campo en la localidad del Lago de Coatepeque (Canton Las Lajas, Sonsonate)

Foto: Frank Sullyvan Cardoza.

puberuleto, con rafidios en la superficie, 2.9-3.3 cm de largo, limbo de 12-13 mm de ancho, tubo ensanchan-

dose gradualmente hacia la base de 1 hasta 3 mm de ancho, estambres (3)5-6, exertos, filamentos desiguales,

insertos en la pared del tubo del perianto; estigma capitado, estilo filiforme, ovario oblongo, ca. 1 mm de largo.

Fruto un antocarpo, verde claro a pardo claro, oblongo-elipsoide, ligeramente constrenido en el apice y base,

6.4-7.8 mm de largo, 2.2-3.7 mm de ancho, puberulento, tricomas blanquecinos septados, rafidios en la su¬

perficie, 10-costillado, costillas y espacios intercostales ± del mismo ancho, no mucilaginoso cuando humedo

(Fig. 3).

Habitat .—Selva alta perennifolia y selva mediana subperennifolia. Los dos nuevos registros (localidades)

para El Salvador se encuentran en el Departamento de Sonsonate. Una localidad es el Municipio de Caluco, en

el Area Natural Protegida Plan de Amayo y la otra localidad esta en Canton Las Lajas, en las cercanlas al Lago

Coatepeque dentro del Area Natural Protegida San Marcelino (Fig. 4).

Fenologia .—Floracion de diciembre-marzo y fructihcacion de enero-abril.

Material examinado.— EL SALVADOR. Sonsonate. Municipio de Izalco: A la mitad del camino entre el cementerio El Pacun y Lajamaya,

13°49 , 52.10"N, 89°34 , 00.37"W, 950 msnm, 15 die 2007, Linares 12938 (MEXU). Municipio Caluco: Area Natural Protegida Plan de Mayo,

en el sector conocido como El Sunzalito ± a 100 m al N del centra de Operaciones, 13 0 41'128"N, 89 0 38'38"W, 325 msnm, 24 ene 2008, Linares

13440 (MEXU).

Etimologia .—El genero Cuscatlania fue dedicado por el prollhco botanico estadounidense Paul C. Standley

(1884-1963), maestro e investigador, quien visito y recorrio gran parte del pals por un perlodo de 6 meses

(diciembre de 1921 y mayo de 1922). Antes de la llegada de los espanoles en 1524, el pueblo de Cuscatlan, era la

Linares et al., Cuscatlania vulcanicola en El Salvador

607

provincia nativa mas importante de la actual Republica de El Salvador (Amaroli 1986). El eplteto vulcanicola,

hace referenda al habitat donde crece la especie y especlficamente al lugar donde se encontro y describio que

fue el Volcan de San Vicente, uno de los mas importantes del pals.

Comentarios. —Esta especie se distingue entre toda la familia, como lo menciona Standley (1923) y Cal¬

deron y Standley (1925), por la insercion de los estambres en la pared del tubo del perianto y no por debajo del

ovario formando una estructura a manera de copa, como se presenta en los demas representantes de la familia.

Clasificacion y afinidades filogeneticas. —Tradicionalmente la clasificacion de Nyctaginaceae que prevale-

cio por mas de 15 anos fue la de Bittrich y Kuhn (1993); en este sistema Cuscatlania fue clasihcada junto con

Mirabilis, Nyctaginiay Allionia (Tabla 1) dentro de la tribu Nyctagineae subtribu Nyctagininnae; dicha subtribu

fue caracterizada por incluir hierbas anuales o perennes, con inflorescencias parciales o flores solitarias invo-

lucradas. Involucro con bracteas connatas o libres, el estigma papiloso o hmbriado, el polen pantoporado de

100-200 pm de diametro, con numerosos poros. Recientemente, Douglas y Spellenberg (2010), basados en la

hipotesis hlogenetica de Douglas y Manos (2007) realizaron algunas modihcaciones a la clasificacion tribal

de Nyctaginaceae y aunque en la hipotesis hlogenetica no se incluyo a Cuscatlania, los autores consideran

que por similitud morfologica es probable que el genero se incluya en la tribu en la que tradicionalmente fue

clasihcado; sin embargo en este nuevo sistema de clasificacion, los autores no reconocieron grupos a nivel de

608

Journal of the Botanical Research Institute of Texas 8(2)

Tabla 1. Afinidades morfologicas de los generos de la subtribu Nyctagininnae (Bittrich & Kuhn 1993).

Morfologia

Allionia

Cuscatlania

Mirabilis

Nyctaginia

Presentation de las bracteas involucrales

Libres

Principalmente connatas

Libres

Duracion de las bracteas involucrales

Persistentes

Desarrollo de las bracteas involucrales

No acrescentes

Acrescentes

No acrescentes

Inflorescencias

Glomerulo

Cimosas

Capitadas

Numero de bracteas por involucro

4-8

6-20

Numero de flores por involucro

2-3

1-16

Mas de 3

Simetria de las flores

Zigomorfa

Actinomorfa

Numero de estambres

4-7

3-5

3-6

5-8

sutribus. A simple vista, en su aspecto general, el tipo de inflorescencia, numero de flores por involucro y color

de las flores, Cuscatlania recuerda a diversas especies de Mirabilis (p. ej., M.jalapa, M. gracilis, M. sanguined).

Sin embargo, tomando en cuenta el involucro de bracteas libres no acrescentes, Cuscatlania recuerda al genero

m. onotlpico Nyctaginia.

CONCLUSIONES

En referencia al Estado de Conservacion de esta especie, alto grado de deterioro y fragmentacion de su habitat;

la hacen poco frecuentes, considerandose extintas en otras zonas del pals. De acuerdo con los criterios de la

Lista Roja de Especies Amenazadas (UICN 2012) y segun las observaciones de los autores, estas plantas son

raras y poco comunes, siendo la categorla para esta especie Vulnerable, VU Al (d).

AGRADECIMIENTOS

Nuestra gratitud a George Pilz y un revisor anonimo por sus valiosos comentarios y observaciones al manu-

Linares et al., Cuscatlania vulcanicola en El Salvador

609

scrito y a Barney Lipscomb por su valiosa revison y apoyo en todo el proceso de edicion de esta publicacion que

contribuye al conocimiento de la flora salvadorena.

REFERENCIAS

Amaroli, P. 1986. En la Busqueda de Cuscatlan: Un proyecto etnohistorico y arqueologico. Manuscrito inedito, Direccion

del Patrimonio Cultural, San Salvador.

Bittrich, V. & U. Kohn. 1993. Nyctaginaceae. In: K. Kubitzki, J.G. Rohwer, & V. Bittrich, eds. The families and genera of vas¬

cular plants - Dicotyledons 2. Springer-Verlag, Berlin, Germany.

Douglas, N.A. & P.S. Manos. 2007. Molecular phylogeny of Nyctaginaceae: Taxonomy, biogeography and characters as¬

sociated with a radiation of xerophytic genera in North America. Amer. J. Bot. 94(5):856-872.

Douglas, N. & R. Spellenberg. 2010. A new tribal classification of Nyctaginaceae. Taxon 59(3):905-910.

Linares, J.L. 2009. Flora inventory in southwestern El Salvador. In: O. Komar, ed. Comprehensive inventories of selected

biological resources within targeted watersheds and ecological corridors of southwestern El Salvador. Improved

Management and Conservation of Critical Watersheds Project, USAID, SalvaNatura, and Gobierno de El Salvador.

Standley, P.C. 1923. New species of plants from El Salvador. J. Washington Acad. Sci. 13(20):437.

Standley, P.C. & S. Calderon. 1925. Lista preliminar de las plantas de El Salvador. Tipografia La Union, Dutriz Hermanos,

San Salvador, El Salvador.

Standley, P.C. &S. Calderon. 1941. Flora Salvadorena. 2da. Edicion. Imprenta Nacional, San Salvador, El Salvador.

610

Journal of the Botanical Research Institute of Texas 8(2)

JOURNAL NOTICE

Sabeeha S. Merchant, Winslow R. Briggs, and Donald Ort, eds. 2014 (Apr). Annual Review of Plant Biology,

Volume 65. (ISSN: 1543-5008; ISBN: 978-0-8243-0665-6, hbk). Annual Reviews, Inc., 4139 El Camino

Way, P.O. Box 10139, Palo Alto, California 94303, U.S.A. (Orders: www.AnnualReviews.org, science@

annualreviews.org, 1-800-523-8635,1-650-493-4400). $101.00, 790 pp., 7.5" x 9.25".

ABOUT THIS JOURNAL— The Annual Review of Plant Biology, in publication since 1950, covers the signih-

cant developments in the held of Plant Biology, including Biochemistry and Biosynthesis, Genetics, Genomics

and Molecular Biology, Cell Differentiation, Tissue, Organ and Whole Plant Events, Acclimation and Adapta¬

tion, and Methods and Model Organisms.

Contents of Volume 65:

Introduction—Sabeeha S. Merchant

1. Our Eclectic Adventures in the Slower Eras of Photosynthesis: From New England Down Under to Biosphere 2 and Beyond— Barry

Osmond

2. Sucrose Metabolism: Gateway to Diverse Carbon Use and Sugar Signaling— Yong-Ling Ruan

3. The Cell Biology of Cellulose Synthesis— Heather E McFarlane, Anett Boring, and Staffan Persson

4. Phosphate Nutrition: Improving Low-Phosphate Tolerance in Crops— DamarLizbethLopez-Arredondo, Marco Antonio Leyva-Gonzalez, Sandra Isabel Gonzalez-

Moral es, Jose Lopez-Bucio, and Luis Herrera-Estrella

5. Iron Cofactor Assembly in Plants— Janneke Balk and Theresia A. Schaedler

6. Cyanogenic Glycosides: Synthesis, Physiology, and Phenotypic Plasticity— RoslynM. Gleadow and Birger Lindberg Mailer

7. Engineering Complex Metabolic Pathways in Plants— Gemma Fane, Dieter Blancguaert, Teresa Capell, Dominigue VanDerStraeten, PaulChristou, andChangfuZhu

8. Triterpene Biosynthesis in Plants— RameshaThimmappa, Katrin Geisler, Thomas Louveau, Paul O'Maille, and Anne Osbourn

9. To Gibberellins and Beyond! Surveying the Evolution of (Di)Terpenoid Metabolism— JiachenZi,SibongileMafu, and Reuben J. Peters

10. Regulation and Dynamics of the Light-Harvesting System— Jean-DavidRochaix

11. Gene Expression Regulation in Photomorphogenesis from the Perspective of the Central Dogma— Shu-Hsing Wu

12. Light Regulation of Plant Defense— CarlosL Ballare

13. Heterotrimeric G Protein-Coupled Signaling in Plants— Daisuke Hrano and Alan M. Jones

14. Posttranslationally Modified Small-Peptide Signals in Plants— YoshikatsuMatsubayashi

15. Pentatricopeptide Repeat Proteins in Plants— Alice Barkan and Ian Small

16. Division and Dynamic Morphology of Plastids— Katherine W. Osteryoung and Kevin A. Pyke

17. The Diversity, Biogenesis, and Activities of Endogenous Silencing Small RNAs in Arabidopsis — Nicolas G. Bologna and Olivier Voinnet

18. The Contributions of Transposable Elements to the Structure, Function, and Evolution of Plant Genomes— Jeffrey L. Bennetzen andHao Wang

19. Natural Variations and Genome-Wide Association Studies in Crop Plants— Xuehui Huang and Bin Han

20. Molecular Control of Grass Inflorescence Development— Dabing Zhang and Zheng Yuan

21. Male Sterility and Fertility Restoration in Crops— Letian Chen and Yao-GuangLiu

22. Molecular Control of Cell Specification and Cell Differentiation During Procambial Development— KaoriMiyashima Furuta, Eva Hellmann, and

Yka Helariutta

23. Adventitious Roots and Lateral Roots: Similarities and Differences— Catherine Bellini, Daniel I. Pacurar, and Irene Perrone

24. Nonstructural Carbon in Woody Plants— Michael C. Dietze, Anna Sala, MariahS. Carbone, Claudia I. Czimczik, Joshua A. Mantooth, Andrew D. Richardson, and Rodrigo

Vargas

25. Plant Interactions with Multiple Insect Herbivores: From Community to Genes— JeltjeM. Stam, AnnekeKroes, Yehua Li, Rieta Gols, Joop J.A. van Loon,

Erik H. Poelman, and Marcel Dicke

26. Genetic Engineering and Breeding of Drought-Resistant Crops— Honghong Hu and Lizhong Xiong

TJ. Plant Molecular Pharming for the Treatment of Chronic and Infectious Diseases— EvaStoger, Rainer Fischer, Maurice Moloney, and Julian K-C. Ma

28. Genetically Engineered Crops: From Idea to Product— Jose Rafael Prado, Gerrit Segers, Toni Voelker, Dave Carson, Raymond Dobert, Jonathan Phillips, Kevin Cook,

Camilo Cornejo, Josh Monken, Laura Grapes, Tracey Reynolds, and Susan Martino-Catt

J.Bot. Res. Inst. Texas 8(2): 610.2014

VASCULAR PLANT FLORA OF THE ALPINE ZONE IN THE

SOUTHERN ROCKY MOUNTAINS, U.S.A.

James F. Fowler

B.E. Nelson and Ronald L. Hartman

US. Forest Service

Rocky Mountain Research Station

2500 S. Pine Knoll Drive

Flagstaff, Arizona 86001, U.S.A.

jffowler@fs. fed. us

Rocky Mountain Flerbarium

Department of Botany, 3165

1000E. University Ave.

Laramie, Wyoming 82071, U.S.A.

bnelsonn@uwyo.edu; rhartman@uwyo.edu

ABSTRACT

Field detection of changes in occurrence, distribution, or abundance of alpine plant species is predicated on knowledge of which species

are in specific locations. The alpine zone of the Southern Rocky Mountain Region has been systematically inventoried by the staff and

floristics graduate students from the Rocky Mountain Herbarium over the last 27 years. It is centered on the mountain ranges of Colorado

and extends north to the Medicine Bow Mountains in southeast Wyoming and south into the Sangre de Cristo Range in north central New

Mexico. It also includes the Ta Sal Mountains of Utah and the San Francisco Peaks in northern Arizona. The alpine meadow and treeline

ecotone flora of the Southern Rocky Mountains includes 609 unique taxa of vascular plants comprising 581 species. The richest families

are Asteraceae (104 species), Poaceae (58 species), Cyperaceae (57 species), and Brassicaceae (42 species). The central Colorado subregion

is the most taxon rich (499) with richness tapering off to the north, southeast, and southwest. Non-endemic alpine zone taxa occur more

frequently elsewhere in the Rocky Mountain Cordillera Floristic Region (515) than in the Madrean (373), Circumboreal (226), or North

American Atlantic Floristic Regions (120). Tevels of endemism within the flora of the alpine zone in the Southern Rocky Mountains range

from single mountains (7) to the flora as a whole (59) including 25 taxa endemic to both the Southern Rockies and to its alpine zone. This

checklist is based on vouchered specimens and should be most useful to botanists and land managers determining what taxa are likely to

occur within their area of interest.

RESUMEN

Ta detection de cambios en ocurrencia, distribution, o abundancia de especies de plantas alpinas se predica como conocimiento de que es-

pecies estan en lugares especificos. Ta zona alpina del sur de la Region de las Montanas Rocosas ha sido inventariada sistematicamente por

el personal y estudiantes de floristica del Rocky Mountain Herbarium en los ultimos 27 anos. Esta centrado en las cordilleras de Colorado y

se extiende hacia el norte hasta las montanas Medicine Bow en el sureste de Wyoming y hacia el sur hasta la cadena Sangre de Cristo en el

norte centro de Nuevo Mexico. Tambien incluye las montanas Ta Sal de Utah y los picos San Francisco en el norte de Arizona. Ta pradera

alpina y la flora del ecotono de la linea arbolada del sur de las Rocky Mountains incluye 609 taxa unicos de plantas vasculares que compren-

den 581 especies. Fas familias mas ricas son Asteraceae (104 especies), Poaceae (58 especies), Cyperaceae (57 especies), y Brassicaceae (42

especies). Ta subregion de Colorado central es la mas rica en taxa (499) cuya riqueza disminuye hacia el norte, sureste, y suroeste. Eos taxa

no endemicos de la zona alpina ocurren mas frecuentemente en cualquier lugar de la Region Floristica de la Cordillera Rocky Mountain

(515) que en la Madreana (373), Circumboreal (226), o las Regiones Floristicas Atlanticas Norte Americanas (120). Eos niveles de endemis-

mo en la flora de la zona alpina de las Southern Rocky Mountains varia de montanas simples (7) a la flora como conjunto (59) que incluye 25

taxa endemicos de las Southern Rockies y su zona alpina. El catalogo esta basado en especimenes testigo y puede que sea util a los botanicos

y gestores del territorio para determinar que taxa puede que existan en su area de interes.

INTRODUCTION

Field detection of changes in occurrence, distribution, or abundance of alpine plant species is predicated on

knowledge of which species are in specific locations. Baseline surveys for this purpose are anchored in the elu¬

cidation of regional species pools; therefore, what species could occur at the chosen baseline survey sites. High

quality baseline study design to detect change at local and regional scales also hinges on the synergistic abil¬

ity of plant systematists, ecologists, and land managers to identify and track individual species as systematic,

ecological, and evolutionary entities. Alpine species in their mountaintop habitats have limited opportunity to

migrate upslope or northward and may be more vulnerable to local extinction (Chapin & Korner 1994; Grab-

herr et al. 1994; Theurillat & Guisan 2001). Concerns of biodiversity managers regarding the potential loss of

alpine vascular plant species due to global warming often center on possible inadequate rates of species migra-

J. Bot. Res. Inst. Texas 8(2): 611 - 636.2014

612

Journal of the Botanical Research Institute of Texas 8(2)

tion or microevolutionary adaptation. Recent studies have shown or predicted a variety of species responses to

increased temperatures and changes in precipitation patterns in montane and alpine zones (e.g., Crimmins et

al. 2011; Pauli et al. 2012). In the Southern Rocky Mountains, determination of the regional alpine plant species

pool should allow baseline studies to be more effective at detecting real change.

Completion of 4,000-13,000 km 2 scale floristic inventories containing alpine areas by the staff and flo-

ristic graduate students from the Rocky Mountain Herbarium (RM) over the past 27 years (Kastning 1990;

Vanderhorst 1993; Lyon 1996; Chumley 1998; Moore 1998; Elliott 2000; Taylor 2000; Nunn 2003; Arnett

2002; Holt 2002; Foley 2006; Reif 2006; Flaig 2007; Farson 2008; Fegler 2010; Fukas et al. 2012; Brummer

2014; and Kirkpatrick 2014; also Nelson 1974,1984) presents an opportunity to synthesize species occurrence

and distribution information for the entire Southern Rocky Mountain Region. The ongoing efforts of the Flora

of North America (FNA 1993+) project to standardize species circumscriptions and nomenclature transcends

the more fragmented data contained in state floras such as Weber and Wittmann (2012), Dorn (2001), and

Allred and Ivey (2012). The recently completed Intermountain Flora (Cronquist et al. 1972; Cronquist et al.

1977,1989,1997,1994; Cronquist et al. 1984; Holmgren et al. 2005; Holmgren et al. 2012) illustrates what can

be achieved on a physiographic scale. There are a few alpine specific checklists for small areas within the Rocky

Mountain Cordillera (kittle 1941; Johnson & Billings 1962; Spence & Shaw 1981; Rundel et al. 2008) but not

for the entire Southern Rocky Mountain Region. The purpose of this study is to document the occurrence of

vascular plant species in the alpine zone of the Southern Rockies using voucher specimen information from

the two largest regional herbaria: Rocky Mountain Herbarium and University of Colorado Herbarium (COFO).

Species distribution within the Southern Rocky Mountain region and by habitat within the alpine zone is a

secondary goal. The resulting checklist will serve as a reference for both land managers and researchers when

establishing site specific baseline studies to detect changes in plant distribution and occurrence.

Study Area

The Southern Rocky Mountain Region (Arno & Hammerly 1984, Fig. 1) is the southern portion of the pro¬

posed boundaries for the Flora of the Rocky Mountains Project (Hartman 1992). It is centered on the moun¬

tain ranges of Colorado and extends north to the Medicine Bow Mountains in southeast Wyoming and south

into the Sangre de Cristo Range of north central New Mexico. It also includes the Fa Sal Mountains of Utah

(Hartman 1992) and adds the San Francisco Peaks in northern Arizona due to floristic similarities (kittle 1941;

Schaak 1983). Colorado was divided into four contiguous subregions (Fig. 1) based on state lines (Wyoming

and New Mexico) and by major highway corridors (1-25,1-70, US Hwy. 50, and by US Hwy. 285 south of US

Hwy. 50). Thus Colorado is separated into north, central, southeast, and southwest geographic components.

Our area of interest is the alpine zone comprised of meadow and treeline ecotone areas of the Southern

Rockies as defined by Korner (1998, 2003). Timberline is the local upper elevational limit of closed canopy

forest; treeline is the general upper limit of scattered clumps of upright trees greater than three meters in

height; treeline ecotone is the transition region between timberline and treeline with a mix of upright trees and

herbaceous vegetation; and alpine meadow is the herbaceous plant dominated region above treeline, but often

includes dwarf shrubs, scattered single trees, krummholz, and patchily vegetated talus, rock outcrops, and

peaks. In the Southern Rockies, the alpine zone starts at 3350 m in southeast Wyoming and gradually rises to

begin at 3540 m in northern New Mexico.

Most of the study area is within the Southern Rocky Mountain Province of the Rocky Mountains Physio¬

graphic System (Hunt 1974; Brouillet & Whetstone 1993). The predominantly north-south oriented mountain

ranges from the southeast Wyoming subregion through the north, central, and southeast Colorado subregions

to the northern New Mexico subregion are underlain by weathered granitic formations while the San Juan

Mountains in the southwest Colorado subregion are of volcanic origin. Both the Utah and Arizona subregions

are within the Colorado Plateau Physiographic Province. The Fa Sal Mountains are the result of laccolithic

intrusions and have exposed igneous rocks above timberline (Baars 1983; Blakey & Ranney 2008). The San

Francisco Peaks are the collapsed caldera edge of a stratovolcano (Nations & Stump 1981). Surface topography

within the study area is quite variable and rugged due to uplift, erosion, and glaciation, but it also includes

Fowler et al., Alpine flora in the southern Rocky Mountains

613

Fig. 1. Study area map for the alpine zone vascular plant flora of the Southern Rocky Mountains. Triangles indicate the 245 collections sites each of which

may represent several individual collections at the RM and COLO. The eight sub-regions within the Southern Rockies are by state (AZ, NM, UT, WY) and

within Colorado by four sub-regions using highways that follow major physiographic separations. N = northern Colorado mountain ranges, C = central

Colorado mountain ranges, SW = San Juan Mountains, SE = southeastern Colorado mountain ranges.

extensive alpine meadows on round topped ridges that have escaped glaciation since before the Wisconsin

epoch (Marr 1967).

The climate of the Southern Rockies has two primary periods of precipitation, summer and winter (Arno

& Hammerly 1984). Most summer precipitation originates from southern air masses while winter snows enter

from the North Pacific (Arno & Hammerly 1984). At Niwot Ridge (3740 m elevation) in northern Colorado,

July mean temperature is 8°C (Billings 2000) and summers are cool with frequent thunderstorms, sometimes

with hail and sleet, with a short growing season of 45 days (Marr 1967). Winters are long, cold, and windy with

frequent blizzards and a minimum temperature of about -28°C at Niwot Ridge (Marr 1967); January mean

temperature is -14°C (Billings 2000). Mean annual temperature at Niwot Ridge is -3.8°C; mean annual precipi¬

tation ranges from 66-86 cm; and mean annual windspeed is 25 km/hr (Marr 1967).

METHODS

The alpine areas have been systematically inventoried by the staff and floristics graduate students from the

RM. This study is largely a synthesis of RM collections verified by Hartman or Nelson over the past few de-

614

Journal of the Botanical Research Institute of Texas 8(2)

cades, as well as those housed at the University of Colorado (COLO). The area covered is southeast Wyoming,

Colorado, north central New Mexico, the La Sal Mountains in southeast Utah, and the San Francisco Peaks in

northern Arizona. From the two databases, we chose 245 collection sites. In searching the databases, the term

“alpine” was first sought followed by elevation adjusted by latitude. Sites chosen from the RM database were

known and collected by at least one of the authors and associated graduate students. From the COLO database,

sites were sorted by county and if site elevation was above the known alpine zone and had more than 20 col¬

lections, specimen labels were searched for “alpine” and if present, included on the initial species list. This list

was extensively vetted for specimens that appeared to be out of geographic or elevational range. Taxonomic

and geographic data were pulled from the RM and COLO plant specimen databases. Habitat data was not

always available from the specimen record, in which case, habitat assignment was based on the authors’ knowl¬

edge of the site and the collector. Lists of species from each location were recorded in an expanded spreadsheet.

Systematics and nomenclature follow the published volumes of Flora of North America (FNA 1993+) with

minor exceptions and follow current taxonomic usage at the RM for families not yet revised by FNA. Excep¬

tions to FNA include recognition of Dorn’s treatment for Salix (Dorn 2010), Packera oodes (Rydb.) W. A. Weber

as a segregate of Packera streptanthifolia, Trisetum montanum Vasey as a segregate of Trisetum spicatum, and

Abies arizonica Merriam as a segregate of Abies bifolia. Floristic region classification is based on Thorne’s (1993)

phytogeography treatment for FNA. Levels of endemism were determined by the authors based on distribu¬

tion maps published in FNA, from RM and USDA Plants online databases, and from geographic distributions

published in the Intermountain Flora (Cronquist et al. 1972; Cronquist et al. 1977,1989,1997,1994; Cronquist

et al. 1997; Holmgren et al. 2005; Holmgren et al. 2012) and state floras. The RM database was thoroughly

searched (beyond our sites) for putative Southern Rockies and alpine endemics to verify our designations. Dis¬

junct populations of taxa are separated by at least one state or province. Our use of the term “alpine endemic”

indicates vascular plant taxa that are distributionally restricted to the alpine zone of the Rocky Mountain

Cordillera, unlike Korner (2003) who includes these taxa along with taxa that are distributionally centered in

the alpine zone as “alpine taxa.” Threatened, Endangered, and Sensitive (TES) species designation were taken

from each state’s natural heritage/rare plants database.

RESULTS

The alpine meadow, talus, and treeline ecotone flora of the Southern Rocky Mountains includes 609 unique

taxa (species, varieties, and subspecies) of vascular plants comprising 581 species (Table 1). The richest fami¬

lies are Asteraceae (104 species), Poaceae (58 species), Cyperaceae (57 species), and Brassicaceae (42 species)

(Table 2). The most speciose genera are Car ex (Cyperaceae, 51 species), Draba (Brassicaceae, 23 species), Erig-

eron (Asteraceae, 18 species), and Poa (Poaceae, 16 species).

Within the eight subregions of this flora, distribution and frequency of occurrence varies by taxon. Oc¬

currences in Table 2 show subregional distribution and give some indication of relative abundance within

each taxon. Between taxa frequency-of-occurrence comparisons may not be valid due to collection biases.

Examples range from the widespread and common Geum rossii to the infrequent single mountain endemic,

Seneciofremontii var. inexpectatus.

The central Colorado subregion is the most taxon rich at 499. Richness tapers off to the north, southeast,

and southwest, dropping to 127 and 211 at the north and south ends of the contiguous Southern Rockies in

Wyoming and New Mexico, respectively. The east-west trending Hoosier Ridge in central Colorado, which has

been repeatedly surveyed, has 180 taxa alone. The more insular La Sal Mountains of Utah are similar to the

north (Wyoming) end with 130 taxa, and the yet more distant San Francisco Peaks of Arizona have only 58

vascular plant taxa in our data set.

Species occurrences between alpine meadow/treeline ecotone areas are also variable. Some species such

as Minuartia obtusiloba, Carex elynoides, and Hymenoxys grandiflora occur mostly in alpine meadows whereas

many species occur throughout both areas; for example, Micranthes rhomboidea (Table 2). Trees define the

treeline ecotone area, but very few herbaceous species occur only there and those few are probable outliers

Fowler et al., Alpine flora in the southern Rocky Mountains

615

Table 1. Summary of search results for Southern Rocky Mountain vascular plants in alpine and treeline ecotone habitats. The survey includes collections from 245

sites in the USFS, RM, and COLO herbaria databases. Disjunct populations are separated from larger populations by at least one state or province. Threatened,

Endangered, and Sensitive (TES) species are officially listed under the Endangered Species Act or by US National Forest Regions.

List by Taxonomic Category

List by special category

Families

Endemic to Southern Rocky Mountains

Genera

213

Endemic to Alpine/treeline ecotone habitats

Species

581

Endemic to a single mountain range

Infraspecies

Disjunct population(s)

Unique taxa

609

TES species

Exotic to Southern Rocky Mountains

from lower elevation habitats. Even the conifers Abies bifolia and Picea engelmannii occur as single trees above

the upper treeline ecotone boundary within alpine meadow, although their main abundance is within the

closed forest below timberline. Occurrences shown in Table 2 indicate relative habitat preference within the

alpine zone of each subregion.

The non-endemic Southern Rocky Mountain alpine zone taxa occur more frequently elsewhere in the

Rocky Mountain Cordillera Floristic Region (515) than in the Madrean (373), Circumboreal (226), or North

American Atlantic Floristic Regions (120). Most (21 of 24) of the disjunct taxa in this flora are shared with the

Circumboreal Floristic Region and northern part of the Rocky Mountain Cordillera Floristic Region. For ex¬

ample, the Colorado and Utah (Uintah Mountains) populations of Armeria maritima ssp. sibirica are separated

by over 1500 kilometers from its main distribution in the circumboreal and arctic regions.

The levels of endemism within the flora of the alpine zone in the Southern Rocky Mountains (Table 1)

range from a single peak or mountain range (7 taxa) to the flora as a whole (59 taxa). Notably 25 taxa are en¬

demic to both the Southern Rockies and to its alpine zone. The alpine zone endemics represent 13 families and

24 genera with Poa and Draba each having four alpine endemic taxa. Alpine zone endemics also include two

Endangered Species Act (ESA) Threatened species: Packerafranciscana from the San Francisco Peaks of north¬

ern Arizona and Eutrema penlandii from the Mosquito Range of central Colorado. The latter species is listed as

Eutrema edwardsii in Table 2 since F. penlandii was recently synonymized prior to publication of the Flora of

North America. However, the disjunct Colorado population of F. edwardsii (from northern British Columbia)

is still officially listed as Threatened F. penlandii pending further review and analysis. The remaining 18 Sensi¬

tive species (Table 1) are listed by individual National Forest Regional Offices (Table 2).

DISCUSSION

This study was intended to provide a species checklist for the entire Southern Rocky Mountain alpine zone

along with some indication of subregional distribution. We chose a more conservative approach to species

occurrence that leans toward errors of omission rather than errors of commission that might result from mis-

identihed specimens in a database only approach. Finer scale distribution by major mountain range within

subregions awaits further biogeographical study.

Scott (1995) indicated 609 alpine plant species for the Middle Rockies, however, Hadley (1987) counted

619 alpine species for the Middle and Southern Rocky Mountain regions combined. Our checklist shows 581

alpine zone species from the Southern Rockies (Table 1). These counts are comparable to Korner’s (2003) esti¬

mate of 600 alpine species for larger mountain systems based on a known alpine species richness for the entire

Swiss Alps of approximately 650 and data showing a 150-550 alpine plant species richness range for many

smaller Swiss alpine floras (10-100 km 2 ). Species richness data from smaller North American alpine floras are

also within Korner’s observed range: Ruby Range of Colorado (220 species, Hartman & Rottman 1987), Teton

Range of Wyoming (260 species, Spence & Shaw 1981), Beartooth Plateau of Montana (210 species, Johnson

& Billings 1962), and White Mountains of California (163 species, Rundel et al. 2008). The larger scale esti¬

mates from the Rocky Mountains also support Korner’s conjecture that smaller mountain range components

616

Journal of the Botanical Research Institute of Texas 8(2)

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Fowler et al., Alpine flora in the southern Rocky Mountains

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Fowler et al., Alpine flora in the southern Rocky Mountains

621

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Journal of the Botanical Research Institute of Texas 8(2)

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623

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624

Journal of the Botanical Research Institute of Texas 8(2)

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Journal of the Botanical Research Institute of Texas 8(2)

Lloydia serotina (L.) Rchb. var. serotina 1A6C 18A19C 9A3T5C 11A1C

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Fowler et al., Alpine flora in the southern Rocky Mountains

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630

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Fowler et al., Alpine flora in the southern Rocky Mountains

631

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634

Journal of the Botanical Research Institute of Texas 8(2)

represent approximately half of the species richness of larger mountain systems. Future synthesis of the Mid¬

dle Rockies floristic inventories at the RM may clarify the discrepancy between the Scott (1995) and Hadley

(1987) estimates, but our results seem to fit the broader alpine pattern.

The checklist (Table 2) is based on vouchered specimens and should be most useful to botanists and land

managers determining what taxa are likely to occur within their area of interest. The frequency values within

Table 2 are best interpreted as relative probabilities for within taxon comparisons between subregions or be¬

tween alpine meadow and treeline ectotone areas within the alpine zone of a subregion. Relative probability

of occurrence between taxa should be limited to small discrete areas of interest. The overall lower number of

occurrences within the treeline ecotone probably reflects its relative narrowness compared to more expansive

alpine meadow and talus slopes. We caution against over interpretation of the frequency counts. Some taxa

with a high number of occurrences, such as Achillea millefolia and Cymopterus lemmonii, are common in lower

elevation habitats as well as in the alpine zone. Other high occurrence taxa, such as Geum rossii, are predomi¬

nantly alpine zone species. Similarly, a low number of occurrences could indicate taxa that infrequently range

up to the treeline ecotone, taxa that are narrowly distributed geographically, or prominent tree species that are

less often collected.

The levels of endemism and floristic region distribution (Table 2) provide a convenient synthesis of the

best available knowledge for botanists and biodiversity managers that recommend TES designations based on

geographic distribution. Most of the U.S. Forest Service Sensitive species (Table 2) have some level of ende¬

mism within the Southern Rockies. The 25 taxa endemic to the alpine zone of the Southern Rockies would also

make a key set of biogeographic/population ecology/species migration studies for the effects of future climate

change and possible species loss.

ACKNOWLEDGMENTS

We are indebted to the graduate students from our program in covering alpine areas in their studies. We thank

Tim Hogan, COLO, for making their database available and Josh Irwin for preparing Figure 1. Salary sup¬

port for this manuscript was provided by the Rocky Mountain Herbarium and by the Forest and Woodlands

Ecosystems Program, Rocky Mountain Research Station, U.S. Forest Service. We thank Walter Fertig and an

anonymous reviewer for their contribution to this paper.

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THE FIRST NATURALIZED OCCURRENCE OF THE CANNACEAE FAMILY IN

THE ARKANSAS (U.S.A.) FLORA, WITH ADDITIONAL NEW

AND NOTEWORTHY ANGIOSPERM RECORDS FOR THE STATE

Brett E. Serviss

Department of Biology

Henderson State University

Arkadelphia, Arkansas 71999-0001, U.S.A.

servisb@hsu.edu

James H. Peck

P.O.Box 705

Cedar Key, Florida 32625, U.S.A.

jhpeck@ualr.edu

Tiffany A. Graves

Department of Biology

Henderson State University

Arkadelphia, Arkansas 71999-0001, U.S.A.

TR180094@reddies.hsu.edu

ABSTRACT

Two species of monocotyledonous angiosperms, Canna indica L. and Miscanthus sinensis Anderss., are reported as new for Arkansas. The

Canna indica record is the first documentation of the Cannaceae family in the state’s flora outside of cultivation. Two additional species of an¬

giosperms, Deutzia scabra T. and Miscanthus sacchariflorus (Maxim.) Hack., are documented for only their second occurrences in the state.

RESUMEN

Se dan como nuevas para Arkansas dos especies de monocotiledoneas, Canna indica L. y M iscanthus sinensis Anderss. La cita de Canna indica

es la primera documentada de la familia Cannaceae para la flora del estado fuera de cultivo. Dos especies adicionales de angiospermas,

Deutzia scabra L. y M iscanthus sacchariflorus (Maxim.) Hack., se documentan como segundas occurrencias en el estado.

INTRODUCTION

The Cannaceae, or canna family, is a group of monocotyledonous angiosperms represented by a single genus,

Canna, and about 10 species of perennial, rhizomatous herbs, principally distributed over subtropical and trop¬

ical regions of the Americas. The precise native range of the genus, however, is difficult to determine, as many

species of Canna are, at present, distributed over subtropical and tropical regions worldwide (Kress & Prince

2000; Wu & Kress 2000). Species of Canna are important as ornamentals for their showy, colorful flowers, large

foliage leaves, and ease of cultivation. The U.S. flora boasts two native species of Canna, C.flaccida Salisb. and C.

glauca L., along with the well-naturalized introduction, C. indica L. Other Canna species and hybrid taxa have

been introduced via the horticultural trade, and along with the aforementioned species, are frequently culti¬

vated in the southeastern U.S., including Arkansas (Bailey 1949; Bailey & Bailey 1976; Kress & Prince 2000).

Canna species reproduce by seed and/or via vegetative/clonal reproduction from an extensive rhizome system.

Vegetative reproduction can generate entire colonies of plants, allowing rapid establishment of populations,

persistence long after cessation of cultivation practices, and potential spread and naturalization from areas of

cultivation. Additionally, seeds or rhizome fragments can also be dispersed through natural or anthropogenic

means, potentially resulting in the establishment of remote escaped populations. The origin of the Canna

indica population in Arkansas is not known; however, plants were apparently not persisting from cultivation

and were observed spreading at the site, where both reproductively mature and smaller plants were present.

Though documented outside of cultivation in several southeastern states, and sometimes cultivated in Ar¬

kansas, our discovery of C. indica represents the first occurrence of the Cannaceae family outside of cultivation

in the state (Kress & Prince 2000; Arkansas Vascular Flora Committee 006; Gentry et al. 2013; SDA, NRCS

2014).

J. Bot. Res. Inst. Texas 8(2): 637 - 639.2014

638

Journal of the Botanical Research Institute of Texas 8(2)

ADDITIONS TO THE ARKANSAS FTORA

Canna indica L. (Cannaceae), Indian shot. Canna indica is a perennial, rhizomatous herb that is considered to

be native to the neotropics, although its precise native range is obscure. At present, it is widely cultivated and

naturalized throughout the subtropics and tropics (Kress & Prince 2000; Wu & Kress 2000). Canna indica is

also well-established in the U.S. flora from eastern Texas and Louisiana to Florida and the Carolinas (Kress

& Prince 2000; USDA, NRCS 2014). In addition to ornamental use, C. indica is sometimes grown for its edible

rhizomes which are rich in starch (Bailey 1949; Wu & Kress 2000).

Voucher specimen: ARKANSAS. Pulaski Co.: a few scattered plants with flowers, escaped and “seeding” in other plants—some plants

smaller, Boyle Park vicinity, Cantrell Hill, wooded bluff and ravine above park, deciduous forest, Tittle Rock, 12 Sep 2002, Peck 2002110

(HEND).

Miscanthus sinensis Anderss. (Poaceae), Maiden grass; Chinese silvergrass. Miscanthus sinensis is a caespi-

tose perennial that is native to China, Japan, and Korea (Hitchcock 1950; Barkworth 2003; Chen & Renvoize

2006). It is naturalized over much of the eastern U.S. from Louisiana, Missouri, and Illinois to New York

southward to Florida. It has also been documented from California and Colorado (Barkworth 2003; USDA,

NRCS 2014). Miscanthus sinensis is readily available through the horticultural trade, frequently encountered in

cultivation, and has become invasive in several states (Barkworth 2003; Smith 2008; Quinn et al. 2010; Meyer

2011). The origin of the Garland County plant of M. sinensis is unknown, but no other plants of M. sinensis were

observed in the vicinity, nor was there any evidence of cultivation. The plant was well-established and thriving

when discovered.

Voucher specimen: ARKANSAS. Garland Co.: one plant along roadside, Hwy. 70 E of Hot Springs near bypass, 3 Nov 2006, Peck 06-777

(HEND).

SECOND OCCURRENCES IN THE ARKANSAS FLORA

Deutzia scabra L. (Hydrangeaceae), Roughleaf deutzia. Deutzia scabra is a deciduous, scandent shrub that is

native to China and Japan (Bailey 1949). It is naturalized in the eastern U.S. from Illinois westward to New

York and Vermont, and southward to North Carolina. It has also been documented from Utah and Florida

(USDA, NRCS 2014). Deutzia scabra has only previously been documented in Arkansas from Garland County.

Voucher specimen: ARKANSAS. Clark Co.: several scandent shrubs from less than 1 m to more than 3 m (a few climbing into the lower

canopy) growing along banks of Mill Creek, semi-wooded riparian zone, University Drive and Hwy. 7/67, Arkadelphia, 19 Sep 2013, Serviss

8049 (HEND).

Miscanthus sacchariflorus (Maxim.) Hack. (Poaceae), Amur silvergrass. Miscanthus sacchariflorus is a rhi¬

zomatous perennial that is native to China, Japan, Korea, and Russia (Hitchcock 1950; Barkworth 2003; Chen

& Renvoize 2006). It is naturalized in the north central and northeastern U.S. from western Nebraska and

Missouri to Minnesota, New York, and Connecticut (Barkworth 2003; USDA, NRCS 2014). Miscanthus sac¬

chariflorus has only previously been documented in Arkansas from Prairie County.

Voucher specimen: ARKANSAS. Garland Co.: a few plants along shoreline, south shoreline of Lake Hamilton near Andrew Hulsey State

Fish Hatchery, 20 Oct 2006, Peck 06-726 (HEND).

KEY TO ARKANSAS SPECIES OF MISCANTHUS

Miscanthus sacchariflorus and M. sinensis are morphologically similar, but can be distinguished using the fol¬

lowing key.

1. Spikelets prominently awned; trichomes of callus about as long as spikelet_M. sinensis

1. Spikelets generally lacking awns; trichomes of callus about two times longer than spikelet_M. sacchariflorus

ACKNOWLEDGMENTS

We sincerely thank Brent Baker (Arkansas Natural Heritage Commission) and Barney Lipscomb (Botanical

Research Institute of Texas) for their helpful comments and suggestions regarding this paper. We also thank

Serviss et al., Cannaceae family in the Arkansas flora

639

the Henderson State University Biology Department and the University of Arkansas at Little Rock Biology De¬

partment for supporting this work.

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JOURNAL NOTICE

Neal K. Van Alfen, Jan E. Leach, and Steven Lindlow, eds. 2014 (Aug). Annual Review of Phytopathology,

Volume 52. (ISSN: 0066-4286; ISBN: 978-0-8243-1352-4, hbk). Annual Reviews, Inc., 4139 El Camino

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ABOUT THIS JOURNAL—The Annual Review of Phytopathology, in publication since 1963, covers the signih-

cant developments in the held of plant pathology, including plant disease diagnosis, pathogens, host-pathogen

Interactions, epidemiology and ecology, breeding for resistance and plant disease management, and includes a

special section on the development of concepts.

Contents of Volume 52:

1. How Way Leads on to Way— Isaac Barash

2. Harnessing Population Genomics to Understand How Bacterial Pathogens Emerge, Adapt to Crop Hosts, and Disseminate— Boris A.

Vinatzer, Caroline L Monteil, and Christopher R. Clarke

3. New Insights into Mycoviruses and Exploration for the Biological Control of Crop Fungal Diseases— JiataoXieandDaohongJiang

4. Altering the Cell Wall and Its Impact on Plant Disease: From Forage to Bioenergy— Qiao Zhao and Richard A. Dixon

5. Network Modeling to Understand Plant Immunity— Oliver Windram, Christopher A. Penfold, and Katherine J. Denby

6. The Role of Trees in Agroecology and Sustainable Agriculture in the Tropics— Roger R.B. Leakey

7. Plant-Parasitic Nematode Infections in Rice: Molecular and Cellular Insights— TinaKyndt, Diana Fernandez, andGodelieve Gheysen

8. Mechanisms of Nutrient Acquisition and Utilization During Fungal Infections of Leaves— Jessie Fernandez, Margarita Marroquin-Guzman, and Richard

A. Wilson

9. Governing Principles Can Guide Fungicide-Resistance Management Tactics— Frank van den Bosch, Richard Oliver, Femke van den Berg, and Neil Paveley

10. Virus Infection Cycle Events Coupled to RNA Replication— Pooja Saxena and George P. Lomonossoff

11. Novel Insights into Rice Innate Immunity Against Bacterial and Fungal Pathogens— WendeLiu, JinlingLiu, Lindsay Triplett, JanE. Leach, andGuo-Liang

Wang

12. The Activation and Suppression of Plant Innate Immunity by Parasitic Nematodes— AskaGoverseandGeertSmant

13. Protein Kinases in Plant-Pathogenic Fungi: Conserved Regulators of Infection— DavidTurra, David Segorbe, and Antonio Di Pietro

14. Speciation in Fungal and Oomycete Plant Pathogens— Silvia Restrepo, Javier F.Tabima, Maria EMideros,NiklausJ.Grunwald, and Daniel R.Matute

15. The ABCs and 123s of Bacterial Secretion Systems in Plant Pathogenesis— JeffH. Chang, Darrell Desveaux, and Allison L Creason

16. Induced Systemic Resistance by Beneficial Microbes— CorneMJ. Pieterse, ChristosZamioudis, RoelandL. Berendsen, DavidM. Weller, Saskia C.M. Van Wees, and

Peter A.H.M. Bakker

17. Fifty Years Since Silent Spring— Lynn Epstein

18. Localizing Viruses in Their Insect Vectors— Stephane Blanc, Martin Drucker, andMarilyne Uzest

19. Plant Cell Wall-Degrading Enzymes and Their Secretion in Plant-Pathogenic Fungi— Christian P. Kubicek, Trevor L. Starr, andH. Louise Glass

20. Meta-Analysis and Other Approaches for Synthesizing Structured and Unstructured Data in Plant Pathology— H. Scherm, C.S. Thomas, K.A.

Garrett, andJ.M. Olsen

21. Networks and Plant Disease Management: Concepts and Applications— M.W. ShawandM. Pautasso

22. Small RNAs: A New Paradigm in Plant-Microbe Interactions— Arne Weiberg, Ming Wang, Marschal Bellinger, and Hailing Jin

23. Predisposition in Plant Disease: Exploiting the Nexus in Abiotic and Biotic Stress Perception and Response— RichardM. Bostock, MatthewF

Pye, and Tatiana V. Roubtsova

24. Susceptibility Genes 101: Howto Be a Good Host— Chris C.N. van Schie and Frank L.W. Takken

25. Horizontal GeneTransfer in Eukaryotic Plant Pathogens— Darren Soanes and Thomas A. Richards

J. Bot. Res. Inst. Texas 8(2): 640.2014

AMERICAN MISTLETOE (PHORADENDRON LEUCARPUM SSP. LEUCARPUM,

VISCACEAE) OCCURRENCE IN HOST TREES WITHIN THE CITY OF CAIRO,

ALEXANDER COUNTY, ILLINOIS, AND ITS INCIDENCE IN ILLINOIS, U.S.A.

Ralph L.Thompson

Hancock Biological Station, Murray State University

Murray, Kentucky 42701, U.S.A.

Botanical Research Institute of Texas

1700 University Drive, Fort Worth, Texas 76107, U.S.A.

Berea College Herbarium, Biology Program

Berea College

Berea, Kentucky 40404, U.S.A.

ABSTRACT

A comprehensive survey of American mistletoe [Phoradendron leucarpum (Raf.) Reveal & M.C. Johnston ssp. leucarpum, Viscaceae] occur¬

rence in host trees was conducted within the incorporated city limits of Cairo, Illinois, from 2011-2013. An extensive herbaria search for

Illinois mistletoe specimens was completed in 2012, and a mistletoe reconnaissance was also made in 2013 to verify its present occurrence

within 14 southern Illinois counties. Cairo, a historic river city and the county seat of Alexander County, is the southernmost city in Illinois,

near the confluence of the Ohio and Mississippi Rivers. In Cairo, Phoradendron leucarpum ssp. leucarpum was observed in 547 host trees

among 13 tree species (9 native, 4 introduced exotics). The two predominate host trees documented were Acer saccharinum (304 trees) and

Ulmus americana (111 trees). The mistletoe occurrence value of Cairo is 13.41 host trees per km, the highest incidence of American mistletoe

and greatest number of host species for a city of its size in Illinois. A statewide assessment of 19 southern Illinois counties resulted in 225

mistletoe specimens examined: 52 specimens collected in Cairo, 97 annotated from herbarium searches, and 76 additional specimens col¬

lected during the 14 county mistletoe host tree survey. Twenty-one host tree species were documented among those herbarium specimens.

The top three host tree species by Illinois counties were Ulmus americana (16), Acer saccharinum (8), and Nyssa sylvatica (8). New North

American mistletoe host trees documented were Acer palmatum, Betula utilis ssp .jacquemontii, and Styphnolohiumjaponicum. Duration of

low winter temperatures has been the determining factor in the northern extension of mistletoe in southern Illinois over time.

Keywords: American mistletoe, distribution, host trees, mistletoe occurrence, Phoradendron leucarpum ssp. leucarpum, Viscaceae, Cairo,

Alexander County, Illinois counties

RESUMEN

Un estudio exhaustivo de la ocurrencia de muerdago americano [Phoradendron leucarpum (Raf.) Reveal & MC Johnston ssp. leucarpum,

Viscaceae] en los arboles hospedadores se llevo a cabo dentro de los limites incorporados de la ciudad de Cairo, Illinois, entre 2011-2013. Se

completo una extensa busqueda para especimenes de herbario de muerdago de Illinois en 2012, y tambien se hizo un reconocimiento se-

cundario de muerdago en 2013 para verificar su ocurrencia actual en 14 condados del sur de Illinois. Cairo, una ciudad riberena historica

vieja y el asiento del Condado de Alexander, es la ciudad mas meridional de Illinois, cerca de la confluencia de los rios Ohio y Mississippi.

Phoradendron leucarpum ssp. leucarpum se observo en 547 arboles huespedes entre 13 especies de arboles (9 nativas, 4 exoticas introduci-

das). En Cairo, los dos arboles hospedadores predominante documentados eran Acer saccharinum (304 arboles) y Ulmus americana (111 ar¬

boles). El valor ocurrencia de muerdago de Cairo es 13.41 arboles hospedadores por km, la mayor incidencia de muerdago americano y el

mayor numero de especies huespedes para un area de su tarnano en Illinois. Una evaluacion del estado resulto en 225 especimenes de

muerdago examinados: 52 muestras recogidas en Cairo, 97 anotadas en la busqueda de herbario, y 76 ejemplares adicionales recogidos du¬

rante una encuesta de los arboles huespedes de muerdago en los condados del sur de Illinois. Veintiun especies de arboles de acogida dife-

rentes fueron documentados de esos especimenes de herbario en 19 condados del sur de Illinois. Las tres principals especies de arboles

hospedadores por condados de Illinois eran Ulmus americana (16), Acer saccharinum (8), y Nyssa sylvatica (8).

Palabras Clave: muerdago americano, distribucion, arboles hospederos, presencia de muerdago, Phoradendron leucarpum ssp. leucarpum,

Viscaceae, Cairo, condado de Alexander, condados de Illinois

INTRODUCTION

A survey of the incidence of Phoradendron leucarpum ssp. leucarpum (Viscaceae), herein, American mistletoe

or mistletoe, in host trees within Cairo was conducted from March-December 2011, June 2012, and March-

J. Bot. Res. Inst. Texas 8(2): 641 - 660.2014

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Journal of the Botanical Research Institute of Texas 8(2)

April 2013. This Cairo survey is the first comprehensive American mistletoe survey for any Illinois town, city,

or county

Cairo, a famous historic river city in Alexander County, Illinois, was chosen as an excellent survey site for

American mistletoe. This decision was made following preliminary reconnaissance in March 14, 2010, which

revealed several species of mistletoe-infested host trees within the incorporated city boundary (Fig. 1). A com¬

parable investigation of host trees infested with American mistletoe was recently completed for the historically

significant city of Berea, Kentucky (Thompson et al. 2008).

While documenting mistletoe in Cairo, it also became evident little was known regarding the statewide

distribution of this taxon. To conduct a complete study of American mistletoe for Cairo and the state of Il¬

linois, a second essential objective of this study was to annotate the incidence of mistletoe documented by

Illinois herbarium specimens. Fourteen state, regional, private, and major herbaria were surveyed for Illinois

mistletoe specimens. All available mistletoe herbarium specimens from Illinois were examined to gather data

on distribution in Illinois counties, collectors, dates, and host tree specificity through time. A third objective

was to make a reconnaissance in 14 southern Illinois counties to currently document incidence of mistletoe-

infested host trees.

AMERICAN MISTLETOE BIOLOGY

Reveal and Johnston (1957) established Phoradendron leucarpum (Raf.) Reveal & M.C. Johnston as the nomen-

claturally correct name for American mistletoe. Abbott and Thompson (2011) made new combinations for

three subspecific taxa within P. leucarpum , of which P. leucarpum ssp. leucarpum is the only infraspecific taxon

native to the eastern United States.

American mistletoe is an epiphytic, dioecious, obligate hemiparasite on numerous deciduous host species

in the eastern United States. This evergreen shrub is characterized by aerial shoots with opposite, simple, oval

to ovate, coriaceous leaves, small spikes of tiny staminate and pistillate flowers on separate plants, and viscid,

pearl-white, translucent one-seeded globular berries (Kuijt 1982, 2003; Fig. 2).

The principal dispersal agents of mistletoe fruits and seeds are avian vectors (Kuijt 2003). Birds spread

mistletoe by ingesting the berries, defecating on branches, dislodging fruits, and wiping bills and feet during

feeding, perching, and roosting (Thompson & Poindexter 2005; Thompson et al. 2008). The availability of host

trees and birds are two important variables; moreover, the Mississippi Flyway corridor, which extends from

upper Canada through Illinois to the Gulf Coast into South America, constitutes one of the most important

flight pathways for bird travel and migration in the eastern United States. Nearly 50 percent of the 700-900

bird species in North America spend at least part of their lives in the Mississippi Flyway (Audubon 2013).

Mistletoe has a clumped or aggregated spatial distribution pattern characteristic of ornithophily (Thompson

& Noe, Jr. 2003; Thompson & Poindexter 2005).

In his Monograph of Phoradendron, Kuijt (2003) mapped the distribution range of American mistletoe [as:

Phoradendron serotinum (Raf.) M.C. Johnston ssp. serotinum]. Its geographical range extends from New Jersey

and Maryland southward through all Atlantic and Gulf Coastal States westward to central Texas, northeast

Oklahoma, southeastern Kansas, east to Arkansas and Tennessee, southern Missouri, Illinois, Indiana, and

Ohio, throughout Kentucky to west-central West Virginia and southern Pennsylvania. The geographical range

of Phoradendron leucarpum ssp. leucarpum in the southeastern United States is southern intraneous (Thomp¬

son & Jones 2001) based on the geographical affinity model of Cain (1930). The geographical affinity of Ameri¬

can mistletoe in southern Illinois is southern extraneous with its northernmost distribution limits primarily

within a 160 km radius of the Mississippi Gulf Coastal Plain Province.

Winter climate is the determining factor of the northernmost extension of American mistletoe. Several

botanists have reported the controlling factor on the northern distribution limits of mistletoe as low and pro¬

longed winter freezing temperatures, e.g., Illinois (Schneck 1884a, b), Indiana (Deam 1932), Ohio (Spooner

1983), and Kentucky (Garman 1913; Thompson 2005). Coder (2008) reported climatic limits were the con¬

trolling features of the distributional range of American mistletoe and assessed the northern boundary to be

Thompson, Phoradendron in Cairo, Illinois, and incidence in Illinois

643

Fig. 1. Cairo incorporated city limits (red-lined boundary) in southern Alexander County near confluence of the Ohio and Mississippi Rivers.

Fig. 2. Pistillate clump of American mistletoe (Phoradendron leucarpum ssp. leucarpum) in Callery pear (Pyrus calleryana).

644

Journal of the Botanical Research Institute of Texas 8(2)

determined by the December minimum daily temperature being greater than -4.0°C. She also considered that

the western edge of mistletoe was bounded by areas with a mean greater than 63.5 cm annual precipitation.

Spooner (1983) determined the principal limiting factor of the northern expansion of mistletoe was directly

correlated with the -4.5°C January mean minimum isotherm. He specifically mentioned that one of two excep¬

tions to this isotherm was along the Wabash River Valley in southern Illinois. At this location, microclimates

exist that are not incorporated into the gross climatological data. In southern Illinois, the northern range

of American mistletoe lies within USDA Plant Hardiness Zone 6b (-17.78°C to -20.56°C) temperature range

(USDA, ARS 2013).

cairo: the study site

History

Cairo, the southernmost city in Illinois and the county seat of Alexander County, served as an important

steamboat port for cargo and commerce to New Orleans in the 19th century. The city is situated on a near level

peninsula surrounded entirely by levees just north of Fort Defiance Park at the junction of the Ohio and Mis¬

sissippi Rivers (Fig. 1). The Mississippi River is the largest river in discharge flow at its mouth and the Ohio

River is the third largest river in discharge flow at its mouth in the United States (Kammerer 1990). Fort Defi¬

ance Park, a Union campsite and fort during the American Civil War, represents the southernmost land border

within the Cairo Corporate Boundary and the lowest elevation in Illinois at 84 m above sea level (asl).

The first Europeans to discover the junction of the Ohio and Mississippi Rivers were the French Jesuit

Jacques Marquette and explorer Louis Jolliet on their trip to map the Mississippi River in 1653. French Jesuit

explorer Louis Hennepin made a campsite on the Cairo peninsula in 1660, and the first fort and tannery were

built in 1702 by Pierre Charles Juchereau de St. Denys. By 1703, Native Americans had driven off the French,

destroyed the village, and effectively curtailed European settlement for over 110 years (Weiser 2012).

On November 16, 1803, Meriwether Lewis, William Clark, and the Corps of Discovery Expedition “...

landed on the point at which the Ohio and Mississippi form there [sic] junction...” (now Cairo, Alexander

County, Illinois). The exploration party remained until November 21, 1803, to conduct scientific research

before continuing up the Mississippi on their historic westward journey (Lewis et al. 2005). In 1818, John

Comegys obtained a charter to establish a city on the peninsula by the convergence of the two rivers, but his

plans never materialized due to lack of financial support. Nevertheless, Comegys coined “Cairo” as the name

for the future city from his deduction that the Ohio River delta resembled the delta of the Nile River in Cairo,

Egypt. Thereafter, “Cairo” and the “Little Egypt” area became standard usage throughout southern Illinois

(Bradsby 1883; Lansden 1910; Lantz 1972). Cairo was founded in 1837 by the Cairo City and Canal Company,

incorporated into a city in March 1858, and designated the county seat of Alexander County in 1860 (Bradsby

1883; Lansden 1910; Lantz 1972).

The 19th century history of Cairo has been well recorded including accounts of its pioneer settlements,

major shipping port, building of factories, warehouses, levees, and canals, completion of the Illinois Central

Railroad in 1854, occupation by the Union Army and Navy in April 1861 during the Civil War, decline of

steamboat river commerce, and the severe effects of great floods (Bradsby 1883; Lansden 1910; Lantz 1972;

Beadles 1990). Other past occurrences, such as the turbulent events of the 1967-1973 Civil Rights Movement,

general decay of the city, and demographic changes in the mid-20th century, have also been reported (Lantz

1972; Ewing & Roddy 1996; Weiser 2012). According to the 2010 United States Census, the population of

Cairo was 2381 individuals (Illinois Demographics 2013). Weiser (2012) described Cairo, Illinois, as a “ghost

town,” a historical city that left behind evidence of its previous glory.

Historical Mistletoe Literature Near Cairo

White (1997) presented significant references on the flora and vegetation of the nearby Cache River, and some

reports directly pertained to American mistletoe along the Ohio River near or at the junction with the Missis¬

sippi River in Alexander County, Illinois.

Thompson, Phoradendron in Cairo, Illinois, and incidence in Illinois

645

The Journals of the Lewis and Clark Expedition (1803-1806) originally written by Meriwether Lewis were

edited by G.E. Moulton two centuries later at the University of Nebraska (Lewis et al. 2005). These documents

provide a unique introduction into the past vegetation of this area. Lewis et al. (2005) noted the abundance of

large silver maple (Acer saccharinum L.), American elm ( Ulmus americana L.), eastern cottonwood ( Populus del-

toides W. Bartram ex Marshall), and American sycamore ( Platanus occidentalis L.) floodplain forests, and river

cane [Arundinaria gigantea (Walter) Muhl.] thickets along the Ohio and Mississippi shorelines. On November

21,1803, while moored near the confluence of the Ohio and Mississippi Rivers, Lewis et al. (2005) recorded:

.. .“from this place, I observed a large quantity of Misseltoe [sic] on the trees bordering on the river, on the main shore... in descending the

Ohio, I furst [sic] observed this plant about the mouth of the Muskingum river [joins Ohio River, near Marietta ].. .it [mistletoe] continued

increasing until I arrived at the mouth of the Ohio [junction of the Ohio and Mississippi, Alexander County, Illinois] and still continues in

larger quantities on this [Mississippi] river than on the Ohio, in so much that the trees at this place were perfectly loaded with it.”

On his trip down the Ohio River to the confluence of the Mississippi, Judge James Hall (1838) recorded in his

Notes on the Western States:

“The mistletoe is seen hanging from the branches of the trees throughout the whole course of the Ohio. It becomes more abundant after pass¬

ing Cincinnati, and is seen in the greatest profusion between Louisville and the mouth of the river [at Cairo], This little plant never grows

upon the ground, but with a very poetic taste, takes up its attic residence upon the limbs of the tallest trees. The berry which contains the

seed, is so viscous as to adhere to the feet of birds, which carry it from tree to tree, and thus contribute to the propagation of this ornamental

parasite.”

In a letter to his northern Illinoisan friendj. Danforth, in December 24,1861, Captain George Dodge (1861) at

Camp McClernand wrote on the presence of mistletoe and the living conditions at Cairo:

“We are located about one and a half miles north of this city [Cairo], and almost on the only ground that I have seen which is susceptible

of being drained. Standing here and there, thro’ our camp is a number of stately sycamore trees, upon whose boughs hang pendant tufts of

mistletoe, and which would, in almost any other place look very pretty, but here in this low, flat, muddy sink and hot-bed of diarrheas, ague,

and fevers, although a rare thing to us of the north, [mistletoe] is hardly even looked at, and seldom mentioned.”

Misdetoe was even commemorated during the Civil War, when a small steamboat tug, built in 1861 under the

name Restless at St. Louis, was purchased by the Union Army early in the Civil War for service in the Western

Naval Flotilla. In 1862, the tugboat was renamed, USS Mistletoe, after the “parasitic green shrub” (Mooney

1968). The tugboat then was transferred to the Navy at Cairo, Illinois, where it joined the Mississippi River

Squadron toward the 1863 assault upon Vicksburg. After the Civil War in 1864, USS Mistletoe was decommis¬

sioned and sold at public auction in Mounds City, Illinois. It was renamed Ella Wood in 1866 and remained in

merchant service until 1871 (Mooney 1968).

Cairo Location and Description

The land area inside the Cairo Corporate Boundary encompasses 1750 ha, and consists of Fort Defiance Park

(22 ha), Angelo Towhead (311 ha), agricultural bottomland and floodplain forest (661 ha) southwest of Cairo,

the Future City (40 ha), the Cairo Regional Airport (190 ha), and the incorporated city (526 ha) of Cairo (Soil

Survey Staff 2013; Fig. 1). The study area for this mistletoe survey was restricted to the incorporated Cairo

city limits situated within the Cairo Quadrangle (Fig. 1). The southern boundary lies just south of the Cairo

Sewage Disposal Plant and Fevee Road between latitude 36°59'38.976"N and longitude 89°9T5.768"W (at 95

m asl), and the northernmost border of the Illinois Central Railroad overpass levee (The Cairo Gate) is located

at latitude 37°1T4.998"N and longitude 89°11T7.998"W (96 m asl). The eastern boundary is created by the

concrete floodwall adjacent to the Ohio River (100 m asl), while the western boundary is determined by the top

of the long earthen Fevee Road (105 m asl) of the Mississippi River. The eastern and western levees then adjoin

the northern levee of the 1903 Illinois Central Railroad bridge (Fig. 1).

Cairo is broadly bisected into western and eastern portions by US 5 IN to Washington Avenue then north-

northeast on Sycamore Street towards Mounds, Illinois (Fig. 1). The major Cairo population resides in older

neighborhoods primarily in the western section beyond Washington Street and Sycamore Street. The eastern

half of the city is comprised of many fewer residences, a soybean oilseed processing company, the Bunge Cor-

646

Journal of the Botanical Research Institute of Texas 8(2)

poration of North America, and numerous abandoned, decaying commercial buildings, vacant lots, houses,

and rubble among the dilapidated “Historic Downtown Cairo” district. However, Cairo has some bne ex¬

amples of historic architecture listed in the National Register of Historic Places; e.g., Riverlore Mansion (1865),

Magnolia Manor (1872), the U.S. Custom House Museum (1872), A.B. Safford Memorial Library (1883), at least

nine stone churches dating from 1857 to 1964, and the American sculptor, George Grey Barnard’s The Hewer

(1906) bronze statue (Pilotlight 2005).

Physiography and Geology

Fenneman (1938) mapped extreme southern Illinois within the Southeastern Lowlands of the Mississippi Al¬

luvial Plain of the Gulf Coastal Plain Province. Keys et al. (1995) classified extreme southern Illinois into the

North Mississippi River Alluvial Plain Subsection, Mississippi Alluvial Basin Section of the Eastern Broadleaf

Forest. Omernik (2007) mapped the southern Illinois area of Alexander County into the Mississippi Allu¬

vial Plain. The Cairo Peninsula lies entirely within the Bottomlands Section of the northern extension of the

unglaciated Gulf Coastal Plain Province (Leighton et al. 1993). Specifically, Cairo is situated within the Bot¬

tomland Section of the Gulf Coastal Plain Division of Alexander County (Schwegman et al. 1973; White 1997).

Although the Bottomland Section of the Coastal Plain Division was not glaciated, the overall effects of outwash

and alluvial deposits have largely determined the present physiography, soil development, and vegetation.

Nelson (2008) described the geology of Cairo and environs in detail from the Cairo Quadrangle. The

geology of the Cairo Peninsula belongs to the Cahokia Formation with clayey deposits ranging from 8 to 52

m deep from the Holocene Stage of the Quaternary System. The superficial sediments from 5 to 10 m in depth

consist of mottled medium to dark gray clay, silty clay, and silt containing very fine to fine sands with quartz

and chert among organic matter (Nelson 2008).

Soils

The topography of incorporated Cairo is nearly level (<1.0-2.0% slope) with an elevation gradient of 94 to 97 m

asl from southern to northern boundaries. Much of the soil in Cairo has initially been excavated and redepos¬

ited during construction to level roads, streets, and railways, levees, homes, and business enterprises due to its

location on the great floodplain of the Mississippi and Ohio Rivers. The soil data are derived from Williams et

al. (2007) and Soil Survey Staff (2013). The soils of Cairo (526 ha) are comprised mainly of one artificial earthy

fill soil and seven major alluvial soil series on the broad floodplain bottomlands prior to enclosing the corpo¬

rate city area within levees and flood walls.

Orthents (215 ha/40.9 % of Cairo area) are excavated, earthy fill materials that had been redeposited over

the generally level Cairo floodplain and in construction of levees. Orthents are undulating to hilly 15-200 cm

deep silty loams, moderately well drained, and strongly acidic in reaction (Williams et al. 2007; Soil Survey

Staff 2013). The seven naturally deposited soils are Beaucoup (28 ha/5.4%), Cairo (27 ha/5.2%), Darwin (51

ha/9.6%), Gorham (49 ha/9.2%), Riley (37 ha/7.2%), Tice (89 ha/16.9%), and Ware (30 ha/5.6%). The seven

alluvial silty clay to silty clay loam soils are very poorly drained to poorly drained to rarely moderately well-

drained with very slow to slow permeability, and strongly acidic to moderately acidic to near neutral in reac¬

tion. The depth of these soils varies from 50 to 203 cm. Under natural seasonal bottomland flooding condi¬

tions, these alluvial soils are annually flooded and are deposited on existing sediments by the Mississippi and

Ohio Rivers outside of the Cairo boundary (Parks & Fehrenbacker 1968).

Climate

The climate of Cairo is of the humid subtropical (Koppen Cfa) type (Rosenberg 2013). Precipitation at this

mild mid-latitude tends to be spread relatively evenly throughout the year without a significant dry season.

Summers are hot and humid with temperatures frequently nearing 32.2°C. The low elevation and proximity

to the Mississippi and Ohio Rivers hold in the summer heat and the high humidity creates hot, muggy condi¬

tions. Winters are generally cool with mild periods due to the elevations and proximity to the rivers preventing

strong winter lows and plunging temperatures. Winter climate is the major limiting factor to mistletoe distri¬

bution along the Ohio River in southern Illinois, Indiana, and Ohio, and the Mississippi River in southeastern

Missouri and southern Illinois.

Thompson, Phoradendron in Cairo, Illinois, and incidence in Illinois

647

The Cairo area has the mildest climate in Illinois. Climate data for 1971-2000 are from Cairo 3 N Station

111166 (Midwest Regional Climate Center 2013). Mean annual temperature in Cairo is 14.7°C with the coldest

month injanuary (0.7°C) and the warmest month in July (26.9°C). Mean annual precipitation is 121.5 cm and

a mean snowfall of 24.4 cm. The least rainfall occurs in September (7.7 cm) and the most rainfall in April and

May (each 12.1 cm). The longest growing season is 270 days, based on 0°C, and the median growing season is

228 days. The last freeze approximates April 9 and the first frost occurs around October 22.

Vegetation and Plant Communities

Braun (1950) classified the forest region at the northern extension of the Mississippi Gulf Coastal Plain in

southern Illinois as Southeastern Evergreen Forest of Bottomland Hardwood Forest composition, while

Kuchler (1964) placed vegetation of the region in the Southern Floodplain Forest. Dyer (2006) in a reclassifica¬

tion of Braun’s forest regions, subsequently mapped her Southeastern Evergreen Forest of the Mississippi Gulf

Coastal Plain as the Mississippi Alluvial Plain Mesophytic Forest. Voigt and Mohlenbrock (1964) classified

the lowland plant communities as Bottomland Forest in Alexander, Pulaski, Massac, and Pope Counties of

southern Illinois. They recognized five floodplain types of vegetation-moisture classes in the adjacent Power

Wabash Valley that ranged from heavy-wet (littoral), wet (overflow bottom), wet-moist (overflow terrace), very

moist (aggraded terrace), to moist floodplain. In the Bottomland Section of southern Alexander County, bot¬

tomland forests are mainly mesic floodplain, wet-mesic floodplain, and wet floodplain forest based on natural

physical features (topographic elevation and aspect, soil moisture, reaction, and permeability, frequency and

duration of flooding, geological substrate and glacial history), and species composition of woody flora and ex¬

isting vegetation (White & Madany 1978; Taft & Mankowski 1997). The characteristic forest communities of

the Bottomland Section at Horseshoe Fake Conservation Area in Alexander County, 27 km northwest of Cairo

at 98 m elevation asl, are characterized as swamps, wetland marsh, wet floodplain, and wet-mesic floodplain

forest (Basinger et al. 1997). Hosner and Minckler (1963) described tree species composition in a successional

study of bottomland hardwood forest in southern Illinois, which included three Alexander County sites, as

representative of a mixed soft-hardwoods forest.

Near Cairo and environs, lands disturbed by annual flooding and siltation are composed of wet flood-

plain forest, while seasonally flooded terraces are primarily wet-mesic floodplain forest. Species composition

of wet floodplain forest along the Ohio and Mississippi Rivers is comprised of riparian trees, most of which

also inhabit wet-mesic floodplain forest. Wet floodplain indicator trees are American sycamore, black willow

(Salix nigra Marshall), box elder (Acer negundo F.), eastern cottonwood, river birch ( Betula nigra F.), and sand¬

bar willow ( Salix exigua Nuttall). Characteristic hardwoods of wet-mesic forests include American elm, green

ash ( Fraxinus pennsylvanica Marshall), hackberry ( Celtis occidentalis F.), honey locust ( Gleditsia triacanthos F.),

pin oak ( Quercus palustris Muenchh.), red elm (JJlmus rubra Muhl.), sweetgum ( Liquidambar styraciflua F.),

and sugarberry ( Celtis laevigata Willd.). These taxa are all present inside the levees of Cairo. Woody species

richness and species diversity become higher in bottomlands as the moisture gradient progresses from wet

floodplain to wet-mesic floodplain to mesic floodplain forest.

Nevertheless, the predominate plant community within the incorporated boundaries of Cairo consists

of woody vegetation including natural remnants and cultivated taxa interspersed among the open graminoid

Culturally Derived Community. This major community has been maintained throughout the past to present

by anthropogenic disturbances (e.g., lawns, yards, streets borders and medians, parks, vacant lots, woodlots,

and numerous other ruderal disturbed habitats).

The woody plant species growing and populating the Culturally Derived Community are mainly native

species characteristic of wet floodplain and wet-mesic floodplain forests. Mesic floodplain forest of drier habi¬

tats includes those trees of wet and wet-mesic forest with the addition of black cherry ( Prunus serotina Ehrh.),

black walnut ( Juglans nigra F.), pecan [Carya illinoensis (Wang.) K. Koch], red maple ( Acer rubrum F.), and

sugar maple (A. saccharum Marshall), among other species. Trees differ significantly in species composition

in the open habitats through selective planting of ornamental native and exotic trees. Some natural wet-mesic

floodplain woodlots and groves are established in Cairo through volunteering of natives and naturalization

648

Journal of the Botanical Research Institute of Texas 8(2)

of some exotic trees, e.g., Callery pear ( Pyrus calleryana Decne.), paper mulberry [Broussonetia papyrifera (L.)

L’Her. ex Vent.], and tree-of-heaven [ Ailanthus altissima (P. Mill.) Swingle]. The more abundant mistletoe-in¬

fested trees of floodplain habitats in the Culturally Derived Community tended to be characteristically mature,

taller, open-canopied, and located in insolated areas.

METHODS AND MATERIALS

An intensive American mistletoe held survey was conducted within the incorporated city limits of Cairo

during March-December 2011, June 2012, and March-April 2013. A vehicle odometer was utilized to record

mileage traveled on all paved streets and unpaved gravel roads. Other terrain was traversed through walking

reconnaissance. Cairo street maps from City Hall and enhanced Cairo topographic maps (Google Earth 2013)

were extensively utilized as street, terrain, and GPS guides.

Nikon Monarch 5 ATB™ 8 x 42 binoculars were used to identify each specific host tree with visible

aerial signs of mistletoe infestation or malformation (i.e., sprigs, clumps, clusters, cankerous swellings, leafless

brooms, limb dieback). Each host was recorded by tree species, degree of mistletoe infestation, and precise

Cairo location. Mohlenbrock (2002) was followed for native tree nomenclature, and USDA, NRCS (2013) was

used for selected non-native trees.

The Mistletoe Infestation Index (MIS), a scale of infestation categories (Thompson et al. 2008), was fol¬

lowed and an inclusive index was given for host tree species: light infestation (1-10 clusters), moderate infesta¬

tion (11-30), heavy infestation (31-100), and extensive infestation (101+). Hemmerly (1989) was used to derive

a Mistletoe Occurrence Value (MOV), a spatial distribution pattern and relative abundance value of host trees

per kilometer. This density factor is calculated by the total number of host trees divided by the total number of

kilometers travelled.

Documentation of selected host trees was by collecting representative voucher specimens with a 12 m

extendible fiberglass linesman pole. Each mistletoe voucher specimen was collected with a winter twig or leafy

branchlet to substantiate the host tree species. Vouchers were mounted, labeled, and deposited in the Berea

College Herbarium (BEREA). A duplicate set of representative mistletoe specimens will be distributed to ILLS

(Appendix 1, 2), and other selected duplicate specimens will be sent to BRIT, EIU, MO, MU, and NCU. Acro¬

nyms for all herbaria follow Thiers (2014).

In 2012, to document Illinois distribution data of Phoradendron leucarpum ssp. leucarpum, all available

Illinois mistletoe herbarium specimens were annotated from BEREA, EIU, F, ILL, ILLS, IND, ISM, KY, MO,

MU, NCU, NY, OSU, and SIU. Descriptive data gathered from each American mistletoe specimen were Illinois

county, location, habitat, host tree identification (when listed), collector(s), collector number [or sino numero

(s.n.)], collection date, and herbarium depository.

During March 30-31 and April 1-2, 2013, a extensive mistletoe reconnaissance was made from 14 south¬

ern Illinois counties. The held search for mistletoe trees was focused in towns and cities and along terrain

bordered by major rivers. In previous Kentucky mistletoe surveys [e.g., Thompson & Noe (2003), Thompson

& Poindexter (2005), Thompson et al. (2008), Thompson & Evans (2010)], a high number of host trees were

typically associated with urban areas. Wooded terrain along Illinois rivers and streams tend to provide more

protected microclimates. Mohlenbrock (1990) reported American mistletoe as occasional from low woods in

the southern 19 Illinois counties (Fig. 3). He noted 15 counties were contiguous to the river borders of the Mis¬

sissippi, Ohio, Wabash, Cache, Saline, and other southern Illinois river tributaries.

The 2013 reconnaissance of 14 counties included the Mississippi River counties (Alexander, Union,

Jackson, Randolph) eastward to Williamson and Saline counties, the Wabash River counties (Wabash, White,

Gallatin), down the Ohio River counties (Gallatin, Hardin, Pope, Massac, Pulaski, Alexander), and the Cache

River counties (Johnson, Pulaski, Alexander). This collecting trip covered portions of the Coastal Plain Prov¬

ince, Salem Plateau Section (Ozark Plateau Province), Mt. Vernon Hill Country, the Till Plains Section (Central

Lowland Province), and Shawnee Hills Section (Interior Low Plateau Province) as delineated from the Illinois

physiographic provinces map of Leighton et al. (1993).

Thompson, Phoradendron in Cairo, Illinois, and incidence in Illinois

649

Southern Illinois

Fig. 3. Nineteen Southern Illinois counties ranked by total identified mistletoe host tree species plus unidentified hosts (Note: Table 2): 1) Alexander, 2)

Pulaski, 3) Hardin,4) Union, 5) Massac, 6) Pope, 7) Jackson, 8) Johnson, 9) Gallatin, 10) Randolph, 11) Wayne,12) Wabash, 13) St. Clair, 14) Crawford,

15) Williamson, 16) Saline, 17) White, 18) Lawrence, 19) Clark.

RESULTS AND DISCUSSION

Mistletoe Survey in the city of Cairo, Alexander County

Phoradendron leucarpum ssp. leucarpum was recorded in 547 hosts among 13 tree species (9 native, 4 intro¬

duced exotics) within Cairo (Table 1). Predominate mistletoe-infested tree species were silver maple (304

trees) followed by American elm (111 trees). Other significant host trees in order of abundance were sugar-

berry (29), red maple (22), paper mulberry (17), Callery pear and green ash (16 each), sugar maple (14), and

black walnut with 13 (Table 1).

650

Journal of the Botanical Research Institute of Texas 8(2)

From this Cairo inventory, nine host trees were documented for the first time in Illinois and are indicated

by an asterisk (*). Five of these new host trees, green ash, red maple, river birch, sugarberry, and sugar maple,

are native volunteers and/or planted. The remaining four hosts, Bradford pear, Himalayan white birch [Betula

utilis D. Don. ssp . jacquemontii (Spach.) WinklJ, Japanese pagoda tree [Styphnolobium japonicum (L.) Schott],

and paper mulberry, are planted and/or escaped exotics. Himalayan white birch and Japanese pagoda tree with

a double asterisk (**), are new host tree records for North America (Table 1).

Seven of the eight host trees listed by Overlease and Overlease (2005) were documented in the Cairo sur¬

vey (Table 1): American elm, black walnut, red maple, river birch, silver maple, sugarberry, and sugar maple.

Although not found in Cairo specifically, the remaining taxon, white ash, listed by Overlease and Overlease

(2005), was later documented in two counties. Silver maple and American elm were the highest two docu¬

mented hosts in the Cairo study as recorded in the Illinois survey by Overlease and Overlease (2005).

Mistletoe occurrence often varied from one to over 100 clusters among hosts of the same species, and the

MIS was highly correlated with number of overall host trees (Table 1). The host trees with MIS of only light in¬

festation (1-10 clusters) were honey locust, Himalayan white birch, Japanese pagoda tree, and river birch. Host

trees with moderate infestation (11-30 clusters) were black walnut, green ash, paper mulberry, red maple, and

sugarberry. Host trees with heavy infestation (31-100 clusters) included American elm and silver maple, and a

few older American elm and silver maple had extensive infestation (101+), which could lead to host mortality.

The majority of mistletoe-infested host trees were found in the Culturally Derived Community occupying

older populated areas interspersed among open street shoulder margins, paved and unpaved gravel roadsides

and medians, yards, city park lawns, church and school yards, and vacant city lots in Cairo proper. The 6.2 ha

lawn of St. Mary’s Park, established in 1872, had nine native or exotic planted tree species infested with mistle¬

toe. Some host trees were present in wet floodplain forest along the shoreline and levees of the Mississippi and

Ohio Rivers. Several other hosts inhabited small open-canopied groves or woodlot remnants of wet-mesic

floodplain and mesic floodplain forests throughout non-residential areas within Cairo. More concisely, Cairo

lacks any significant natural vegetated areas. Rather, the city is mainly comprised of land subjected to long¬

term anthropogenic disturbance or altered by bottomland flooding regimes.

As expected, few host trees with mistletoe were observed in the environs outside of the incorporated

Cairo city limits. The most common mistletoe-infested host trees were typically also the most numerous

canopy trees (i.e., silver maple and American elm). Those host trees in close proximity to each other generally

appeared to facilitate greater mistletoe seed dispersal, which likely contributed to the increased aggregated

(clumped) mistletoe distribution pattern observed within Cairo. Older and taller mature host trees character¬

istically formed open canopy crowns with better insolation for mistletoe success and provided greater infesta¬

tion opportunities from avian vectors over time.

The Mistletoe Occurrence Value (MOV) from 547 mistletoe-infested trees per 40.8 km (25.2 mi) of roads

was 13.41 in Cairo. This high density value is a result of the city focus, whereas countywide surveys include

much larger expanses of non-urbanized land; e.g., Thompson and Noe (2003) recorded 3502 host trees from

15 tree species in traversing 805 km for a MOV of 4.35 host trees/km in Rockcastle County, and in Garrard

County, Thompson and Poindexter (2005) documented 1740 host trees from 12 tree species in traveling 523

km for a MOV of 3.33 host trees/km.

Fifty-two specimens (26 vouchers and 26 duplicates) collected in the Cairo survey are presented with

complete label information (Appendix 1). The determination of Illinois state host tree records from Cairo was

based on the initial herbarium searches. Six incidental mistletoe specimens (American elm, an oak, black gum

(Nyssa sylvatica Marshall), silver maple, and two unidentified tree hosts) from other herbaria were examined

from Alexander County (Appendix 2). The additions of American elm, black gum, and silver maple specimens

added to the nine new tree species of the Cairo study (Table 1) accounted for 14 host trees among the 21 overall

hosts for Illinois (Table 2).

Previous Relevant Illinois Mistletoe Collections and Surveys

Schneck (1884a) collected seven Ulmus americana specimens in different phenological stages and wrote his

Thompson, Phoradendron in Cairo, Illinois, and incidence in Illinois

651

Table 1. Host specificity of Phoradendron leucarpummtWu) city limits of Cairo, Illinois.

Host Tree Species

No. Trees

Percent

Acer saccharinum L.

304

55.58

Ulmus omericono L.

111

20.29

*Celtis laevigata Willd.

5.30

*Acer rubrum L.

4.02

*Broussonetia papyrifera (L.) L'Her. ex Vent.

3.11

*Fraxinus pennsylvanica Marshall

2.93

*Pyrus calleryana Decne.

2.93

*Acersaccharum Marshall

2.56

Juglans nigra L.

2.38

*Betula nigra L.

0.36

Gleditsia triacanthos L.

0.18

**Betula utilis D. Don ssp.jacquemontii (Spach) Winkl.

0.18

**Styphnolobium japonicum (L.) Schott

0.18

Total Species: 13

547

100.00

(*) New Illinois host trees vouchered for American mistletoe.

(**) New United States host species vouchered for American mistletoe.

Table 2. Nineteen southern Illinois counties ranked according to identified mistletoe host tree species plus unknown hosts, collectors, specimens, and year(s) of

collection from 225 herbarium specimens (Note: Fig. 3, Appendix 1,2).

County

Host Species

+ Unknowns

Collectors

Number of

Specimens

Collection Years

1 Alexander

14 + 3

1939,1947,1970,1985,1992, 2011,2012, 2013

2 Pulaski

13 + 5

1860,1869,1919,1939,1949,1950,1985,1989, 2013

3 Hardin

7+1

1947,1949, 2013

4 Union

1948,1949,1955, 2013

5 Massac

1902,1948,1949, 2013

6 Pope

1931,1948,1949, 2013

7 Jackson

3 + 1

1948,1949,1956,1985, 2013

8 Johnson

3 + 1

1948,1949,1989, 2013

9 Gallatin

1948,1949, 2013

10 Randolph

2 + 2

1949,1955,1961,1977, 2013

11 Wayne

1949,1958

12 Wabash

1 + 1

1882,1883,1912,1949, 2013

13 St. Clair

1 + 1

1861,1955

14 Crawford

1949,1973

15 Williamson

1949

16 Saline

1949

17 White

1949

18 Lawrence

1949

19 Clark

1949

T = 225 specimens

descriptive notes on each label, which was a source for his article describing the life cycle of mistletoe. Schneck

(1884b) observed mistletoe on 28 honey locust trees, a pin oak, one red elm, 11 silver maples, and “Ulmus

americana, many thousand” over an 18 month period in 1882-1883 at Mt. Carmel, Illinois, in Wabash County.

Trelease (1916) excluded Champaign County, J. Perriam s.n. (ILL), no date, as “doubtless error of locality” and

no other Champaign County specimens were found. Thompson and Bennett (1938) reported [American] elm

and [silver] maple as the major trees infested by mistletoe at Horseshoe Lake in Alexander County near Cairo.

Tehon (1942) discovered that mistletoe prefers American elm and black gum in southern Illinois along the

Ohio and Wabash River bottomlands northward to Wabash County, and the Mississippi River bottomlands

652

Journal of the Botanical Research Institute of Texas 8(2)

north to Union County. Jones (1945) reported mistletoe as parasitic on American elm, black gum, oak, and

other deciduous trees in southern Illinois northward to Union and Lawrence counties. Jones (1963) later add¬

ed Randolph, Saline, and Crawford counties as containing mistletoe. Carter (1964) listed mistletoe occurrence

in 18 Illinois counties and hosts as mainly [American] elm and occasionally black gum, black walnut, honey

locust, oak, [silver] maple, and sycamore. Mohlenbrock (1990) listed American elm and sweetgum as host

trees. American elm was the predominant host named on herbarium specimens; sweetgum was not verified as

a host tree by any representative vouchers from either the herbaria search or this study.

Overlease and Overlease (2005) conducted a reconnaissance survey of American mistletoe (under

synonym, Phoradendron serotinum (Raf.) M.C. Johnston), with a focus on the distribution, host species, and

abundance across its geographical range in the eastern United States. Although they did not collect herbarium

vouchers, their identification of host trees in Illinois is accepted from their extensive study. Overlease and

Overlease (2005) listed eight host trees for Illinois with the two major hosts in tree numbers and clumps of

mistletoe as silver maple (93 trees/2334 mistletoe clusters) and American elm (39 trees/3339 mistletoe clus¬

ters). The remaining host trees listed were sugar maple (9/343), red maple (7/282), sugarberry (4/58), black

walnut (4/47), white ash [ Fraxinus americana L. (1/6)], and river birch (1/1). All eight taxa were documented

with representative voucher specimens during this study.

Kuijt (2003) cited 23 Illinois voucher specimens (counting six duplicates) from five host trees in 14 Il¬

linois counties from A, GH, ILL, K, MO, NY, US, and WIS. Host trees were a black gum, a black locust ( Robinia

pseudoacacia L.), an oak, a winged elm ( Ulmus alata Muhl.), 13 American elms, and three unidentified trees.

Eighteen vouchers cited by Kuijt (2003), which represented all 14 counties from ILL, MO, and NY, were exam¬

ined in this study. These 18 specimens are denoted by a dagger (t) in Appendix 2.

All 19 Illinois counties mapped for American mistletoe by Mohlenbrock (1990) were documented in the

herbarium searches. Ninety-seven voucher specimens were annotated from holdings at EIU (4), F (0), ILL (33),

ILLS (31), IND (0), ISM (16), KY (0), MO (2), MU (1), NCU (2), NY (3), OSU (0), and SIU (5). Twenty-six collec¬

tion years of these voucher specimens ranged from 1860 to 1992 (Table 3). These 97 specimens (84 vouchers

and 13 duplicates) are comprised of 77 identified hosts (10 duplicates) from eight host tree species, and 20

unidentified hosts (three duplicates) from 18 specimens without trees listed plus two unknown Quercus sp.

Mistletoe specimens with documentation of identified host tree on herbarium labels were counted as host tree

records. Similarly, specimens with a host tree identified on the labels while listing other mistletoe-infested

trees observed, were not counted without an actual mistletoe specimen documented.

The eight identified host tree species by abundance and number of collectors, counties, and herbarium

specimens (Appendix 2) are as follows: Ulmus americana (12 collectors/16 counties/54 specimens), Nyssa syl-

vatica (9/6/9), Gleditsia triacanthos (5/3/6), Acer saccharinum (2/2/2), Platanus occidentalis (2/1/2), Robinia pseu¬

doacacia (2/1/2), Juglans nigra (1/1/1), Ulmus alata (1/1/1), and no host identified including the two Quercus sp.

(10/9/20). In the Illinois herbaria search, the major host by far was American elm followed distantly by black

gum. These data were reported in previous botanical literature (Tehon 1942; Jones 1945,1963; Jones & Fuller

1955; Carter 1964).

A summary of the 97 voucher specimens is listed by decade followed by the 26 collectors, number of their

specimens, and collection year (Table 3). Fifty-five of the 97 specimens (56.70%) within 18/19 counties were

collected by three taxonomists, H.E. Ahles (7 counties/9 specimens), R.A. Evers (16 counties/23 specimens),

and G.S. Winterringer (12/23). Several of their representative specimens at ILL, ILLS, and ISM list the same

locality information and dates, denoting they often were held collecting companions (Appendix 2). The years,

1948-1949, were clearly the zenith of 20th century Illinois mistletoe collecting (Table 3). It is important to note

that 77/97 (79.38%) specimens were collected by 16/26 collectors during 1860-1949, or 154 to 65 years ago

(see: Tables 2,3).

Illinois Mistletoe Documented from 2013 Reconnaissance

The 2013 collecting trip to 14 southern Illinois counties on March 30-31 and April 1-2, 2013, provided 39

more mistletoe herbarium specimens with 37 ILLS duplicates, or 76 herbarium specimens on deposit at

Thompson, Phoradendron in Cairo, Illinois, and incidence in Illinois

653

Table 3. Summary of 97 Illinois Phoradendron leucarpum ssp. leucarpum specimens by decade, collector, number of specimens, and year (s).

Decade

Collector

Specimens

Collection Year(s)

1860s

Brendel

1860

Welsch

1861

Raymond

1869

1880s

Schneck

1882-1883

1900s

Gleason

1902

1910s

Trelease

1912

Palmer

1919

1930s

Schopf

1931

Boewe

1939

Evers

1939

1940s

Bailey

1947

Carter

1948

Evers

1948-1949

Winterringer

1948-1949

Ahles

1949

Sieve rt

1949

Walker

1949

1950s

Evers

1950

Buser

1955

Evers

1955

Neill

1955

Mohlenbrock

1956

Evers

1958

1960s

Neill

1961

1970s

Furray

1970

Huston

1970

Ebinger

1973

Shildneck

1977

1980s

Nickrent

1985

Winslip

1989

1990s

Phillippe

1992

Totals:

26 Collectors

97 Specimens

24 years represented

BEREA and ILLS (Appendix 2). These 76 Illinois specimens plus the 52 Cairo specimens contributed 128

mistletoe specimens toward this Illinois mistletoe study. Fourteen host tree species were documented during

this southern Illinois reconnaissance; eight were also present in the Cairo survey (Table 1). Four new host trees

documented for Illinois were Celtis occidentalis and Fraxinus americana (Hardin and Pulaski counties), and

Japanese maple (Acer palmatum Thunb.) and Quercus palustris (Pulaski County).

Aside from the Cairo survey (Appendix 1), 11 counties provided 76 mistletoe specimens from the 2013

reconnaissance (Appendix 2). County specimen collections were as follows: Alexander (American elm, silver

maple), Gallatin (silver maple), Hardin (American elm, black gum, black walnut, hackberry, red maple, sugar

maple, white ash), Jackson (black gum, silver maple), Johnson (American elm, silver maple), Massac (red ma¬

ple [2], silver maple), Pope (black gum, silver maple), Pulaski (American elm, black gum, black locust, green

ash, hackberry, honey locust, Japanese maple, pin oak, red maple, silver maple [3], sugarberry, white ash), Ran¬

dolph (black gum), Union (black gum, red maple, silver maple), and Wabash (American elm). Regardless of

extensive searches, mistletoe was not found in Saline, White, and Williamson counties, even though historical

collections are known.

Alexander County had the greatest host tree richness with 14 different hosts, Pulaski County provided

13 host tree species with 10 hosts from Mounds City and four from Mounds, and Hardin County accounted

for seven host tree species at Cave-in-Rock State Park (Table 2; Appendix 2). Excluding Cairo data (Table 1,

Appendix 1), the three leading hosts in the 2013 reconnaissance, were silver maple (8 counties/9 specimens),

black gum (6 counties/6 specimens), and American elm (5 counties/5 specimens). Six of the eight host tree spe-

654

Journal of the Botanical Research Institute of Texas 8(2)

cies previously annotated in the herbarium searches for Illinois specimens (American elm, black gum, black

locust, black walnut, honey locust, silver maple) were also collected (Appendix 2). Winged elm and American

sycamore were not redocumented as host trees during this study

The recent observations reveal silver maple to be the most abundant and prevalent mistletoe-infested tree

of southern Illinois counties (Table 1; Overlease & Overlease 2005; Appendix 1, 2). This feature is a contrast

to the incidence of mistletoe-infested American elm as evidenced from herbarium searches and botanical lit¬

erature (Appendix 2; Tehon 1942; Jones 1945; Carter 1964; Mohlenbrock 1990). Reasons are not clear for this

observation on the abundance of silver maple, although inferences may be made: historical collectors could

possibly have ignored or overlooked silver maple as a host tree as only 2/97 were documented with specimens

(Appendix 2). American elm certainly was the most collected mistletoe-infested host as evidenced with an

occurrence in 16/19 counties and 54 herbarium specimens prior to the 2013 survey The original Dutch Elm

Disease [ Ophiostoma ulmi (Buism.) Nannf.] and its more aggressive subspecies, the New World Dutch Elm

Disease ( Ophiostoma novo-ulmi Brasier spp. americana Brasier & S.A. Kirk) may have taken their toll upon

American elms (Brasier & Buck 2001). However, American elm and other host species are not as dominant in

2013 as it was in the 1940s-1960s and latter part of the 20th century. The intensive mistletoe survey of Cairo

was the one exception to the occurrence of mistletoe-infested American elm, even then, it was nearly a 3:1 ratio

of mistletoe-infested silver maple to American elm with mistletoe (Table 1).

The abundance of American mistletoe in the extreme southern counties and the scarcity as the geograph¬

ical range progresses upward to the northern and central counties was the paramount observation. Winter cli¬

mate was clearly the limiting factor in Illinois mistletoe distribution as evidenced by number of collections and

specimens through the historic herbarium searches, relevant literature (Schneck 1884a, b; Spooner 1983), and

the 2013 county reconnaissance. Phoradendron leucarpum ssp. leucarpum is near its northernmost distribution

in southern Illinois (e.g., its southern extraneous affinity) due to low and prevalent winter temperatures during

the past and at the present time. The American mistletoe distribution pattern has remained similar through

time: the occurrence of mistletoe-infested host trees and number of infested host tree species, continually de¬

creased as counties were inventoried northward along the Mississippi and Ohio Rivers and within the south-

central counties both in the past and the present (Table 1, 2,3; Appendix 1, 2).

Synopsis of Illinois American Mistletoe Study

1) A total of 225 herbarium specimens (97 from herbarium searches, 52 from Cairo, 76 from 2013 county

survey) were annotated from 19 southern Illinois counties (Appendix 1,2); counties ranked by number of

host species, collectors, specimens, and years of collection are located in the southernmost Illinois counties

(Table 2; Fig. 3).

2) American mistletoe was recorded from 21 different host tree species representative of 11 plant families (Ap¬

pendix 1, 2); three new exotic host trees for North America documented with Phoradendron leucarpum ssp.

leucarpum are Japanese maple (Acer palmatum Thunb.), Himalayan white birch [ Betula utilis D. Don ssp.

jacquemontii (Spach) WinklJ, and Japanese pagoda tree [Styphnolobiumjaponicum (L.) Schott].

3) The Cairo survey in Alexander County, documented 26 voucher specimens and 26 duplicates (52 speci¬

mens) from 13 hosts with 9 hosts new to Illinois; silver maple and American elm are the two dominant host

trees (Table 1; Appendix 1).

4) The herbarium searches for Illinois specimens generated 97 specimens (84 vouchers and 13 duplicates) and

provided eight identified host trees for Illinois; American elm and black gum are the two major host trees

(Appendix 2).

5) The 2013 reconnaissance survey of 14 southern Illinois counties yielded 39 vouchers and 37 duplicates (76

specimens) within 11 counties from 14 hosts; hackberry, pin oak, white ash, and Japanese maple are four

new Illinois hosts, and silver maple, black gum, and American elm are the three leading host trees by

county (Appendix 2).

6) American mistletoe was much more abundant and with greater host tree diversity near cities; e.g., Cairo,

Metropolis, Mounds, Mounds City, and the major rivers in the extreme eastern (Wabash), southern (Cache,

Thompson, Phoradendron in Cairo, Illinois, and incidence in Illinois

655

Ohio, Saline), and western (Mississippi) counties; mistletoe is very scarce in the northern and central coun¬

ties as evidenced in this study and the herbarium searches (Fig. 3; Table 2; Appendix 1, 2).

CONCLUSIONS

Cairo has the highest incidence of American mistletoe [Phoradendron leucarpum (Raf.) Rev. & M.C. Johnston

ssp. leucarpum] and the greatest number of host species for any town or city in Illinois. In Cairo, the Mistletoe

Occurrence Value (MOV) was 13.41 trees/km from 547 mistletoe-infested trees per 40.8 km of roads travelled.

The major parameter for mistletoe occurrence and distributional spread in southern Illinois is largely

related to climatic conditions (e.g., a continued mild winter climate and buffering effects by major rivers along

open lowland topography). Other important factors for the incidence of mistletoe include the availability of

mature, open-canopied host trees, culturally derived habitats of towns and cities, and influence of the Missis¬

sippi Flyway corridor for bird dispersal of fruits and seeds. The rather mild winter temperatures for the last

several years within the USDA Plant Hardiness 6b Zone has been conducive for evergreen mistletoe establish¬

ment, growth, proliferation, and migration in southern Illinois. The incidence of mistletoe in different hosts

species and in quantities correlates with its southern extraneous geographical affinity. Further American

mistletoe surveys, preferably with winter twigs or branchlets added to mistletoe vouchers specimens, would

enhance host tree identification and overall host species toward mistletoe distribution in southern Illinois

counties.

APPENDIX 1

ANNOTATED CAIRO MISTLETOE COLLECTIONS

ILLINOIS. Alexander Co.: Cairo: Cedar St. sidewalk jet. Cross St. adj. to abandoned Southern Medical Center Hospital, 37°00'9.50"N,

89°10'44.61"W, elev. 97 m, hemiparasitic on Acersocchorinum, 2 Mar 2011, R.L Thompson 11-02 (BEREA, ILLS); 25th St. & Sycamore St. in

yard behind Spirit's House Liquor, 37°00'32.49"N, 89°10'30.76"W, elev. 94 m, on Juglans nigra, 2 Mar 2011, R.L Thompson 11-05 (BEREA,

ILLS); 7th St. & Washington Ave. median, 36°59'59.09"N, 89°9'58.43"W, elev. 96 m, moderately-infested Acer saccharinum, 2 Mar 2011,

R.L Thompson 11-07 (BEREA, ILLS); DayStar Care Center, 2001 Cedar St. at end of Cross St., 37°00'39.52"N, 89°10'44.25"W, elev. 95 m, on

introduced Pyrus calleryana, 2 Mar 2011, R.L. Thompson 11-08 (BEREA, ILLS); St. Mary's Park, founded 1872, lawn bordering Magnolia

Drive, 37°00'38.18"N, 89°11'8.49"W, elev. 95 m, on Acerrubrum, 8 Mar 2011, R.L Thompson & R.J. Trites 11-14 (BEREA, ILLS); St. Mary's Park

lawn near park pavilion, 37°00'36.57"N, 89°11'9.24"W, elev. 95 m, on Betula nigra, 8 Mar 2011, R.L. Thompson & RJ. Trites 11-15 (BEREA,

ILLS); 36th St. & Washington Ave. median, 36°59'59.09"N, 89°9'58.43"W, elev. 96 m, on Acer saccharum, 8 Mar 2011, R.L Thompson & RJ.

Trites 11-21 (BEREA, ILLS); 19th St. jet. with Washington Ave. (US 51N) in yard behind Dairy Hut and Dairy Mart, 1808 Washington Ave.,

37°00'16.77"N, 89°10'29.33"W, elev. 96 m, heavily-infested on Ulmus americana, 8 Mar 2011, R.L. Thompson & R.J. Trites 11-24 (BEREA, ILLS);

St. Mary's Park lawn, 37°00'35.15"N, 89°11 '9.09"W, elev. 96 m, on lightly-infested ornamental Styphnolobium japonicum, 22 Apr 2011, R.L

Thompson & RJ. Trites 11-69 (BEREA, ILLS); St. Mary's Park lawn adj. to Magnolia Drive, 37°00'37.18"N, 89°11'8.14"W, elev. 95 m, on Celtis

laevigata, 22 Apr 2011, R.L. Thompson & R.J. Trites 11-70 (BEREA, ILLS); St. Mary's Park lawn adj. Park Place, 37°00'31.69"N, 89°11 '10.54"W,

elev. 96 m, lightly-infested Gleditsia triacanthos, 22 Apr 2011, R.L Thompson & R.J. Trites 11-71 (BEREA, ILLS); 418 Cross St. and M.L. King

Ave., 37°00'13.77"N, 89°10'41.47"W, elev. 95 m, on three infested Fraxinuspennsylvanica, 5 Jul 2011, R.L. Thompson 11-447 (BEREA, ILLS);

St. Mary's Park lawn, 37°00'35.15"N, 89°11'9.09"W, elev. 96 m, on exotic Styphnolobium japonicum with three clumps, 5 Jul 2011, R.L

Thompson 11-448 (BEREA, ILLS); St. Mary's Park lawn across from 2817 Park Place, 37°00'31.69"N, 89°11 '10.54"W, elev. 96 m, three clusters

on planted Gleditsia triacanthos, 13 Aug 2011, R.L Thompson 11-550 (BEREA, ILLS); St. Mary's Park yard near pavilion, 37°00'35.21"N,

89° 11'9.00"W, elev. 95 m, on Acer saccharum, 13 Aug 2011, R.L Thompson 11-551 (BEREA, ILLS); backyard behind 3111 Washington Ave.

residence adjacent to Magnolia Ave., 37°00'38.75"N, 89°11 '6.90"W, 96 m elev., on Betula nigra, 13 August 2011, R.L Thompson 11-556 (BEREA,

ILLS); Cairo High School, 37°1 '11.25"N, 89°11 '15.41 "W, elev. 96 m, on Acerrubrum, 13 Aug 2011, R.L Thompson 11-564 (BEREA, ILLS); Cairo

High School, moderately-infested ornamental Pyrus calleryana at front entrance, 37°1'9.57"N, 89°11'14.43"W, elev. 96 m, 13 Aug 2011,

R.L Thompson 11-565 (BEREA, ILLS); 25th St. & 2501 Holbrook Ave., 37°00'9.33"N, 89°10'45.36"W, elev. 95 m, a single clump on mature

Fraxinus pennsylvanica, 13 Aug 2011, R.L. Thompson 11-566 (BEREA, ILLS); 229 12th St. E of Poplar St., yard of Ronnie Garrett, 37°00'7.82"N,

89°10'9.95"W, elev. 95 m, on introduced ornamental Betula utilis ssp. jacquemontii with four clusters, 11 Sep 2011, R.L Thompson & K.

Rivers Thompson 11-633 (BEREA, ILLS); Johnson Terrace in yard at jet 38th St., on Juglans nigra, 9 Dec 2011, R.L Thompson 11-738 (BEREA,

ILLS); 229 12th St. east of Poplar St., yard of Ronnie Garrett, 37°00'7.82"N, 89°10'9.95"W, elev. 95 m, on lightly-infested planted Betula

utilis ssp. jacquemontii, 25 Jun 2012, R.L. Thompson &J.R. Abbott 12-827 (BEREA, ILLS); 424 16th St. jet. M.L. King Ave. (Walnut St.), yard of

William Little, 37°00'8.39"N, 89°10'26.41 "W, elev. 95 m, on planted Broussonettiapapyrifera with six clumps, 25 Jun 2012, R.L. Thompson &

J.R. Abbott 12-830 (BEREA, ILLS); abandoned Church of Christ yard at 26th St. & Sycamore St., 37°00'33.44"N, 89°10'48.40"W, elev. 94 m, on

Ulmus americana, 1 Apr 2013, R.L Thompson & W.W. Overbeck 13-63 (BEREA, ILLS); abandoned Church of Christ yard at 26th & Sycamore St.,

37°00'24.75"N, 89°10'42.72"W, elev. 94 m, on Celtis laevigata, 1 Apr 2013, R.L. Thompson & W.W. Overbeck 13-64 (BEREA, ILLS); abandoned

Church of Christ lawn at 26th St. & Sycamore, 37°00'34.87"N, 89°10'46.78"W, elev. 94 m, on escaped Broussonettia papyrifera, 1 Apr 2013,

R.L. Thompson & W.W. Overbeck 13-65 (BEREA, ILLS).

656

Journal of the Botanical Research Institute of Texas 8(2)

APPENDIX 2

ANNOTATED ILLINOIS MISTLETOE HERBARIUM COLLECTIONS

ILLINOIS. Alexander Co.: Miller City, tree in Mississippi River bottoms, 25 Nov 1939, R.A. Evers & F.W. Evers 93 (ILLt, ILLS); old floodplain

roadside between Horseshoe Lake and the Mississippi River, on Ulmus americana, 18 Oct 1947, W.M. Bailey &J.R. Swayne256 (EIU, NCU,

SIU); 1.0 mile S of Olive Branch, on Quercus, 10 Oct 1970, Furray 148 (EIU); Horseshoe Lake, wet woods, Sec.7-10,15-18, 20-22, T 16 S, R

2 W, wet bottomland woods, 5 Nov 1970,2.5. Huston 562 (SIU); a side road off Rt. 3/127 near pond and small homes in Cache, on Acer

saccharinum, 1 Nov 1985, D. Nickrent & K. Robertson 2076 (ILL); Ozark Hill Prairie Research Area, Shawnee National Forest, Mill Creek

7.5-minute topo, in Nyssa sylvatica on steep wooded slope, 8 Apr 1992, L.R. Phillippe&J. Olson 19792 (ILLS); Urbandale, jet. of US 51 & IL

37,37°2'23.14"N, 89°11 '10.37"W, elev. 94 m, heavily-infested Ulmus americana, 29 Mar 2013, R.L Thompson & W.W. Overbeck 13-04 (BEREA,

ILLS); Urbandale, jet. of US 51 & IL 37, behind residence, 37°2'20.90"N, 89°11'15.37"W, elev. 94 m, lightly-infested Acer saccharinum, 29

Mar 2013, R.L Thompson & W.W. Overbeck 13-05 (BEREA, ILLS). Clark Co.: E of West Union, on Ulmus americana, 3 Nov 1949, R.A. Evers 21997

(ILLS). Crawford Co.: SE of Palestine, on Ulmus americana, 26 Nov 1949, R.A. Evers 22043 (ILLS); Palestine, on Ulmus americana, 26 Dec

1949, H.EAhles 1892 (ILLt, ISM); Sec. 2, R 11 W,T 6 N., growing on Ulmus americana, 29 Dec 1973 ,J.E. Ebinger 14345 (EIU, ILLS). Gallatin

Co.: 2 mi. N of Lawler, on Ulmus, 13 Nov 1948, J.C. Carter s.n. (ILLS); 3.0 mi. N of Saline Mines, on Robinia pseudoacacia, 10 Dec 1949, H.E

Ahles 1894 (ILLt); 1.5 mi. N of Inman; on Ulmus americana, 10 Dec 1949, G.S. Winterringer2844 (ISM); 3.0 mi. N of Saline Mines, on Robinia

pseudoacac/a, 10 Dec 1949, G.S. Winterringer2851 (ISM); Cave-in-Rock Road (IL 1),yard ofGreg Mayne, Box71,37°36'03.75"N,88°12'27.04"W,

elev. 109 m, heavily-infested Acer saccharinum, 31 Mar 2013, R.L Thompson & W.W. Overbeck 13-35 (BEREA, ILLS). Hardin Co.: Cave-in-Rock

State Park, in the SE part of Hardin Co., T 12 S, R 9 or 10 E, bordering on the N shore of the Ohio River, tree growing on the sides of the

stony limestone bluffs, 11 Nov 1947, W.M. Bailey &J.R. Swayne292 (EIU, ILLS, NCU, SIU); 3.0 mi. from Cave-in-Rock, on Ulmus americana,

10 Dec 1949, H.EAhles 1897 (ILLt); S of Lamb, NE of Cave-in-Rock, on Ulmus americana, 10 Dec 1949, G.S. Winterringer 2849 (ISM); Cave-

in-Rock State Park, N of lodge, 37°28'10.65"N, 88°9'3.75"W, elev. 138 m, two small sprigs in Nyssa sylvatica, 31 Mar 2013, R.L Thompson

73-36 (BEREA); Cave-in-Rock State Park, S of lodge, 37°28'8.20"N, 88°9'7.28"W, elev. 141 m, 12-15 clumps on Acer rubrum, 31 Mar 2013,

R.L. Thompson 13-37 (BEREA, ILLS); Cave-in-Rock State Park, SE of lodge, 37°28'10.25"N, 88°9'2.45"W, elev. 130 m, moderately-infested

Juglans nigra, 31 Mar 2013, R.L Thompson 13-38 (BEREA, ILLS); Cave-in-Rock State Park, NE of lodge, 37°28'10.88"N, 88°9'2.78"W, elev. 134

m, lightly-infested Acersaccharum, 31 Mar 2013, R.L. Thompson 13-39 (BEREA, ILLS); Cave-in-Rock State Park, NW of lodge restaurant across

blacktop, 37°28'8.90"N, 88°9'9.79"W, elev. 146 m, in Fraxinus americana, 11-15 clumps, 31 Mar 2013, R.L. Thompson 13-40 (BEREA, ILLS);

Cave-in-Rock State Park, S of visitor center on blufftop, 37°28'10.72"N, 88°9'5.92"W, elev. 120 m, on Celtis occidentalis with 2-3 clusters, 31

Mar 2013, R.L. Thompson 13-41 (BEREA, ILLS); Cave-in-Rock State Park, W of lodge, 37°28'10.72"N, 88°9'5.92"W, elev. 144 m, in Ulmus

americana with 7-10 clumps, 31 Mar 2013, R.L. Thompson 13-42 (BEREA, ILLS). Jackson Co.: N of Grand Tower, on Ulmus americana, 12

Dec 1948, R.A. Evers 15674 (ILLS); N of Grand Tower, on Ulmus americana, 12 Dec 1948, G.5. Winterringer 1807 (ILLt, NYt); near Grand

Tower, on Ulmus americana, 2 Nov 1949, G.5. Winterringer2473 (ISM); N of jet. with Rt. 144 in tree along Rt. 3,12 Dec 1956, R.H. Mohlenbrock

8615 (SIU); 3.0 mi. NW of jet. with Rt. 149 along N side of Rt. 3 near a homestead along hillside creek, on a large Ulmus americana, 1 Nov

1985, D. Nickrent & K. Robertson 2075 (ILL); Grand tower, Redtown, S of IL 3 at Wills Rd., yard of Willie Smith at 1927 Brunkhorst Ave.,

37°38'28.36"N, 89°30'37.06"W, elev. 111m, moderately-infested Acer saccharinum, 30 Mar 2013, R.L. Thompson & W.W. Overbeck 13-30

(BEREA, ILLS); IL 3 N at Box 16178 in yard, 37°43'26,65"N, 89°17'14.95"W, elev. 110 m, seven heavily-infested Acer saccharinum, 30 Mar

2013, R.L. Thompson & W.W. Overbeck 13-31 (BEREA, ILLS); Murphysboro, S of IL 13 at 340 Niemann Lane near Walmart, 37°45'46.54"N,

89°17'14.95"W, elev. 118 m, moderately-infested Nyssa sylvatica, R.L. Thompson & W.W. Overbeck 13-32 (BEREA, ILLS). Johnson Co.: SW of

Belknap, on Ulmus americana, 11 Dec 1948, R.A. Evers 15670 (ILLS); 0.5 mi. S of Belknap, on Ulmus americana, 11 Dec 1948, G.5. Winter¬

ringer 1805 (ILLt, NYt); 4.0 mi. SW of Goreville below Draper Bluff, in rocky woods on Quercus, 2 Apr 1949, G.5. Winterringer 1910 (ILLt);

Lower Cache River Natural Area,T 14 S, R 3 E, Sect. 10, Karnak 7.5-minute topo map, tree near river, 30 Mar 1989, K. Winship 1007 (ILLS);

Barkhausen Cache River Wetlands Center, 200 m inside & S of county line on ditch floodplain, 37°18'30.36"N, 89°1 '5.59"W, elev. 100 m, a

single clump in young Ulmus americana, 31 Mar 2013, R.L. Thompson & W.W. Overbeck 13-48 (BEREA, ILLS); Cache River abandoned Railroad

bed adj. to S. Franklin St. between Belknap & Karnak, 37°18'38.15"N, 88°57'16.02"W, elev. 102 m, lightly-infested Acer saccharinum

specimens obtained with shotgun, 31 Mar 2013, R.L. Thompson & W.W. Overbeck 13-49 (BEREA, ILLS). Lawrence Co.: 4.0 mi. S, 0.5 mi. east

of Lawrenceville, woods, in Ulmus, 16 Mar 1949, J.P. Siverts.n. (ILL); 1.0 mi. NE of St. Francisville, on Ulmus americana, 11 Nov 1949, R.A.

Evers 22046 (ILLS); 1.0 mi. NE of St. Francisville, on Ulmus americana, 26 Dec 1949, H.EAhles 1891 (ILLt, ISM). Massac Co.: Metropolis, III.,

on Ulmus americana, 14 Aug 1902, H.A. Gleason s.n. (ILL); SE of Boaz in wet ground on Ulmus americana, 11 December 1948, R.A. Evers, J.

Hall, & G.5. Winterringer 15669 (ILLS, SIU); SE of Boaz, on Ulmus americana, 11 Dec 1948, G.5. Winterringer 1803 (ILLt, NYt); NE of Joppa, on

Nyssa sylvatica, 11 Dec 1949, G.5. Winterringer 2843 (ISM); Metropolis, US 45, Fort Massac State Park, lawn at Campground space 10 R,

37°8'47.64"N, 88°42'26.52"W, elev. 102 m, four clumps in Acer rubrum, 31 Mar 2013, R.L. Thompson 13-45 (BEREA, ILLS); Metropolis, Fort

Massac State Park in front of Visitor Center, 37°8'41.47"N, 88°43'18.95"W, elev. 102 m, a single large cluster in Acer rubrum, 31 Mar 2013,

R.L. Thompson 13-46 (BEREA, ILLS); Metropolis, in yard of Wilma Flemister, 719 A, East 5th St. (US 45) tree planted in 1956-57,37°8'44.45"N,

88°43'18.95"W, elev. 103 m, moderately-infested Acer saccharinum, 31 Mar 2013, R.L. Thompson & W.W. Overbeck 13-47 (BEREA, ILLS). Pope

Co.: Golconda Quad., R 8 E,T 12 S, on Lotton farm 1.0 mi. NE of town, on a Gleditsia triacanthos, 9 July 1931,2. Schopfs.n. (ILLS); Bay Creek

bottoms east of Rose Hill, on Ulmus americana, 11 Dec 1948, R.A. Evers, J. Hall, & G.S. Winterringer 15666 (ILLS); near Golconda, on Ulmus

americana, 11 Dec 1948, G.S. Winterringer 1806 (ILLt); N of Golconda, on Nyssa sylvatica, 10 Dec 1949, G.S. Winterringer 2842 (ISM); IL 1, S

of Zander's Lane, Rauch's Hill in hilly dry woods, 37°22'39.36"N, 88°29'31.45"W, elev. 138 m, on three Nyssa sylvatica, 31 Mar 2013, R.L.

Thompson & W.W. Overbeck 13-43 (BEREA, ILLS); Golcanda, IL 146, Patton St. off Bay City Dr., 37°21'29.20"N, 88°29'4.71"W, 147 m, in Acer

saccharinum with 15-30 clumps, 31 Mar 2013, R.L. Thompson & W.W. Overbeck 13-44 (BEREA, ILLS). Pulaski Co.: Phoradendron flavescens,

1860, F. Brendels.n. (ILL); Villa Ridge, tree with Phoradendron flavescens, 1869, Raymond s.n. (ILL); Mounds, III., in low ground, on Ulmus

alata, 1 Oct 1919, EJ. Palmer 16638 (MOt); 2.0 mi. NW of Olmstead along highway, on Ulmus americana, 11 April 1939, G.H. Boewes.n.

(ILLS); NW of Karnak in a swamp, 14 April 1949, R.A. Evers 15745 (ILLS); E of Karnak, on Ulmus americana, 11 Dec 1949, G.S. Winterringer

2847 (ISM); S of Mounds in swamp, on Acer saccharinum, 10 Jun 1950, R.A. Evers 23568 (ILLS); 1.0 mi. east of Rt. 3/127 on III. Rt. 3 in a field

Thompson, Phoradendron in Cairo, Illinois, and incidence in Illinois

657

adjacent to some homes, on Ulmus omericono, 1 Nov 1985, D. Nickrent&K. Robertson 2077 (ILL); Lower Cache River Natural Area, portion

of Porter Ditch Rd., 30 Mar 1989, K. Winship 1011 (ILLS); Hillerman, near 304Tick Ridge Rd. (Co. Rd. 2), Burnett family residence, 37°14'15.60"N,

88°56'27.98"W, elev. 140 m, one canker with sprigs in Fraxinus americana,31 Mar 2013, R.L Thompson & W.W. Overbeck 13-50 (BEREA, ILLS);

Cache River along abandoned railroad (So. Franklin Rd.), near Johnson Co. boundary, 37°18'3.04"N, 88°58'4.34"W, elev. 101 m, single

clump in small Nyssa sylvatica, 31 Mar 2013, R.L Thompson & W.W. Overbeck 13-51 (BEREA, ILLS); Olmstead, lawn of Olmstead Missionary

Baptist Church, IL 37S & jet. of Marigold St., 37°10'28.66"N, 89°5'53.21"W, elev. 105 m, lightly-infested Acer saccharinum 31 Mar 2013, R.L

Thompson & W.W. Overbeck 13-52 (BEREA, ILLS); Mounds City, off IL 37, N levee by welcome sign, 37°5'42.74"N, 89°9'16.24"W, elev. 100 m,

on Celtis occidentolis, 31 Mar 2013, R.L Thompson & W.W. Overbeck 13-53 (BEREA, ILLS); Mounds City, off IL 37, N levee by welcome sign,

37°5'42.74"N, 89°9'16.24"W, elev. 100 m, on Ulmus omericono, 31 Mar 2013, R.L. Thompson & W.W. Overbeck 13-54 (BEREA, ILLS); Mounds

City,off IL37, N levee adj. to welcome sign, 37°5'42.74"N,89°9'16.24"W, elev. 100 m,on Robinio pseudoacacia, 31 Mar2013,/?.L Thompson

& W.W. Overbeck 13-55 (BEREA, ILLS); Mounds City, Hopkin's Rd. (Commercial Ave.) adj. by IL 37 in lowland yard, 37°5'38.86"N, 89°9'14.04"W,

elev. 97 m, on Celtis laevigata, 31 Mar 2013, R.L. Thompson & W.W. Overbeck 13-56 (BEREA, ILLS); Mounds City, Hopkin's Rd. (Commercial

Ave.) adj. IL 37 in lowland yard, 37°5'38.86"N, 89°9'14.04"W, elev. 97 m, on Acer saccharinum, 31 Mar 2013, R.L. Thompson & W.W. Overbeck

13-57 (BEREA, ILLS); Mounds City, Hopkin's Rd. (Commercial Ave.) adj. to IL 37 in lowland yard, 37°5'38.86"N, 89°9'14.04"W, elev. 97 m,

seven clumps on Gleditsia triacanthos, 31 Mar 2013, R.L. Thompson & W.W. Overbeck 13-58 (BEREA, ILLS); Mounds City, Illinois Ave. & Pearl

St. by First Missionary Baptist Church near IL 37, 37°5'19.95"N, 89°9'44.11"W, elev. 97 m, on Froxinus pennsylvonico, 31 Mar 2013, R.L.

Thompson & W.W. Overbeck 73-59 (BEREA, ILLS); Mounds, off US 51N, yard of Zack Hannan, 124 N. Spencer St. jct.Thistletown St., 37°6'55.27"N,

89°11 '34.46"W, elev. 99 m, one large branch clump on Quercus polustris, 1 Apr 2013, R.L. Thompson & W.W. Overbeck 13-66 (BEREA, ILLS);

Mounds, off US 51N, yard of Zack Hannan, 124 N. Spencer St. jct.Thistletown St., 37°6'54.29"N, 89°11'33.93"W, elev. 99 m, on lightly-in¬

fested Acerrubrum, 1 Apr 2013, R.L. Thompson & W.W. Overbeck 13-67 (BEREA, ILLS); Mounds, off US 51N, yard of Zack Hannan, 124 N.

Spencer St. jet with Thistletown St., 37°6'54.70"N, 89°11 '34.46"W, elev. 99 m, small sprigs on two branches of Acer palmatum, 1 Apr 2013,

R. L. Thompson & W.W. Overbeck 13-68 (BEREA); Mounds, off US 51N, on Acer saccharinum, yard of Zack Hannan, 124 N. Spencer St. jet.

Thistletown St., 37°6'54.90"N, 89°11'34.68"W, elev. 99 m, 1 Apr 2013, R.L. Thompson & W.W. Overbeck 13-69 (BEREA, ILLS). Randolph Co.:

1.0 mi. NE of Fort Gage, on Nyssa sylvatica, 10 Dec 1949, R.A. Evers 22056 (ILLS); SW of Ellis Grove, 18 Aug 1955, J.O. Neill 7121 (ILLS); E of

Fort Gage in barnyard, on Nyssa sylvatica, 19 Aug 1955, R.A. Evers 49019 (ILLS); Chester, vicinity of cove off Rt. 3, Phoradendron in tree, 18

Nov 1961,3.0. Neill 15903 (ILLS); Dupo along III. Rt. 3, Ulmus americana, 26 Nov 1961 J.O. Neill 15904 (ILLS); 0.5 mi. west of Cora in cemetery

on slope by N shoulder of Rt. 3, on Nyssa sylvatica, 27 Mar 1977, P. Shildneck9231 (ISM); Chester, lawn at 8948 Edgemont Drive off IL 3

across from Mill St., 37°53'57.61"N, 89°48'37.23"W, elev. 196 m, two small clusters in a Nyssa sylvatica, 30 Mar 2013, R.L. Thompson & W.W.

Overbeck 13-34 (BEREA, ILLS). Saline Co.: NE part of Stonefort, on Ulmus americana, 14 Apr 1949, R.A. Evers &J. Flail 15731 (ILLS); Stonefort,

SW corner of Saline County, in Ulmus americana, 11 Dec 1949, FI.EAhles 1896 (ILLt); Stonefort, SW corner of Saline County, on Ulmus

americana, 11 Dec 1949, G.5. Winterringer2848 (ISM). St. Clair Co.: Mascoutah, III., Phoradendron flavescens, 1862, W. Welsch s.n. (ILL); SW

of Ellis Grove, in Nyssa sylvatica, 19 Aug 1 955,10. Neill 7121 (ISM).. Union Co.: Jonesboro, at the Ranger Station, on Nyssa sylvatica, 12 Dec

1948, R.A. Evers 15671 (ILLS); E of Wolf Lake, on Platanus occidentalis, 12 Dec 1948, R.A. Evers 15672 (ILLS); E of Wolf Lake, on Gleditsia tria¬

canthos, 12 Dec 1948, R.A. Evers 15673 (ILLS); Jonesboro, on Nyssa sylvatica, 12 Dec 1948, G.5. Winterringer 1804 (ILLt); on Gleditsia triac¬

anthos, MDec 1948, G.S. Winterringer 1810 (ILL); on Platanus occidentalis, 12 Dec 1948, G.5. Winterringer 1811 (ILL);foresters'headquarters,

on Nyssa sylvatica, 2 Nov 1949, G.5. Winterringer 2471 (ISM); 0.5 mi. E of town of Wolf Lake at S end of Pine Hills on low slope, Sec. 3,T. 11

S. , R. 3 W., on Juglans nigra, 10 Jan 1955, F.B. Buser5777 (ILL); Cavaness Road by 750 East State IL 3 near drainage ditch and railroad tracks,

37°28'26.96"N, 89°10'38.76"W, elev. 105 m, a single cluster in an Acer rubrum, 30 Mar 2013, R.L. Thompson & W.W. Overbeck 13-27 (BEREA,

ILLS); Wolf Lake, 4955 State IL 3, Murphy's family yard off Jacob St., 37°30'18.45"N, 89°26'20.14"W, elev. 107 m, a single clump on Acer

saccharinum, 30 Mar 2013, R.L. Thompson & W.W. Overbeck 13-28 (BEREA, ILLS); Jonesboro, off IL 146 & Main St. at USDA Shawnee Na¬

tional Forest, Ranger Station Picnic Shelter area, 37°27'25.51"N, 89°16'9.99"W, elev. 160 m, four clumps in a Nyssa sylvatica, 30 Mar 2013,

R.L. Thompson & W.W. Overbeck 13-29 (BEREA, ILLS). Wabash Co .: Mt. Carmel, Illinois, Phoradendron flavescens on Ulmus americana,

abundant fruits ripe, 25 Dec 1882,3. Schnecks.n. (ILL); Mt. Carmel, Illinois, Phoradendron flavescens fruits all gone from cold weather effects,

15 Jan 1883,3. Schnecks.n. (ILL,); Mt. Carmel, Illinois, Phoradendron flavescens inflorescences developing, 20 Apr 1883,3. Schnecks.n. (ILL,);

Mt. Carmel, Illinois, Phoradendron flavescens ovaries prominent, 26 Jun 1883,3. Schnecks.n. (ILL); Mt. Carmel, Illinois, Phoradendron flave¬

scens very small berries, 27 Jul 1883,3. Schnecks.n. (ILL); Mt. Carmel, Illinois, Phoradendron flavescens green fruits, staminate and pistillate

flowers open, 15 Sep 1883,3. Schnecks.n. (ILL); Mt. Carmel, Illinois, Phoradendron flavescens berries full grown, flowers full anthesis, 23

Oct 1883,3. Schnecks.n. (ILL); Mt. Carmel, Illinois, Phoradendron flavescens white berries ripe, 6 Dec 1883,3. Schnecks.n. (ILL); Mt. Carmel,

Illinois, Phoradendron flavescens on Ulmus americana, Dec 1912, Treleases.n. (MO); 2.0 mi. NE of Keensburg, on Ulmus americana, 26 Nov

1949, R.A. Evers 22047 (ILLS); 1.5 mi. NE of Allendale, on Ulmus americana, 9 Dec 1949, G.5. Winterringer 2845 (ISM); I-64E adj. to S. Water

St., 2.0 km S of Grayville on Wabash River floodplain, 38°13'42.21"N, 87°59'14.80"W, elev. 124 m, one clump on Ulmus americana, 2 Apr

2013, R.L. Thompson 13-70 (BEREA, ILLS). Wayne Co.: 5.0 mi. E of Fairfield, in Quercus, 20 Dec 1949, M. Walker s.n. (ILL); 5.0 mi. S of Fairfield

along roadside, in Gleditsia triacanthos, 20 Oct 1958, R.A. Evers 59540 (ILLS, MU). White Co.: NE of Crossville, on Ulmus americana, 10 Dec

1949, FI.EAhles 1898 (ILLt); NE of Crossville, III. Rt. 1, on Ulmus americana, 10 Dec 1949, G.5. Winterringer2850 (ISM); S of Herald in extreme

southeast corner of Williamson County, on Ulmus americana, 12 Dec 1949, R.A. Evers 22059 (ILLS). Williamson Co.: 1.5 mi. SW of Stonefort,

on Ulmus americana, 11 Dec 1949, FI.EAhles 1895 (ILLt); 1.5 mi. SW of Stonefort in extreme SE corner of Williamson County, on Ulmus

americana, 11 Dec 1949, G.5. Winterringer 2846 (ISM); village of Crab Orchard, on Ulmus americana, 12 Dec 1949, R.A. Evers 22058 (ILLS).

ACKNOWLEDGMENTS

I express my gratitude to three reviewers for their constructive comments of this manuscript as follows: J.

Richard Abbott, Missouri Botanical Garden; Thomas E. Hemmerly, Middle Tennessee State University; and

Barney L. Lipscomb, Botanical Research Institute of Texas. Special thanks are given to the herbarium curators

658

Journal of the Botanical Research Institute of Texas 8(2)

of EIU, F, ILL, ILLS, IND, ISM, MO, MU, NCU, NY, OSU, and SIU for access to mistletoe specimens and their

herbarium resources. I extend appreciation to Russell J. Trites, Murray State University, for his company in the

Cairo survey, and Will W. Overbeck, Eastern Kentucky University, for his keen sightings of mistletoe on the

southern Illinois counties reconnaissance, and Melanie G. Bentley, Eastern Kentucky University, for prepara¬

tion of the three figures.

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Agriculture and Natural Resources Conservation Service.

TETRAGONIA TETRAGONIOIDES (AIZOACEAE)

DISCOVERED IN LOUISIANA, U.S.A.

Sairah M. Javed

Lowell E. Urbatsch

Louisiana Natural Heritage Program

Louisiana Department of Wildlife and Fisheries

Baton Rouge, Louisiana 70898, U.S.A.

sjaved@wlf.la.gov

Louisiana State University

Department of Biological Sciences

Baton Rouge, Louisiana 70803, U.S.A.

ABSTRACT

Tetragonia tetragonioides (Pallas) Kuntze is reported for the first time as adventive to Touisiana. This largely southern hemisphere taxon has

become widespread around the World. Touisiana is added to the list of nine other states in the U.S. where this species is known to occur.

RESUMEN

Tetragonia tetragonioides (Pallas) Kuntze se registra por primera vez como especie adventicia en Louisiana. Este taxon, mayormente del

hemisferio oriental, se ha dispersado ampliamente alrededor del mundo. Lousiana se agrega a la lista de los otros nueve estados en los

EE.UU. donde se conoce la ocurrencia de esta especie.

The senior author discovered a single plant of Tetragonia tetragonioides (Pallas) Kuntze while performing a bo¬

tanical survey of Grand Terre Island in Jefferson Parish, Louisiana. This species was observed growing on dry

beach amongst coppice mounds between the dune ridge and the high tide line. Grand Terre is a barrier island

adjacent to the Gulf of Mexico and is situated just south of Barataria Bay and northwest of the Mississippi River

Delta. This represents the first report for the species in Louisiana. Because only one individual was observed,

the senior author collected a fragment for a voucher specimen. Associates of this species included Cakile Mill.,

Amaranthus gregii S. Watson, Sesuvium portulacastrum (L.) L., Spartina alterniflora Loisel., and Suaeda linearis

(Elliot) Moq.

The genus Tetragonia L. is comprised of 50-60 species, mostly of the southern hemisphere (Gathe and

Watson 2008). Tetragonia tetragonioides is the most widespread species and is native to New Zealand and Aus¬

tralia (Prescott 1984); however, it can be dispersed on ocean currents and consequently may have arrived

naturally to areas of South America (Fabris 1967; Taylor 1994). USDA, ARS, National Genetic Resources Pro¬

gram (2013) also includes China, Japan, and Taiwan within the native range of this taxon. In New Zealand, T.

tetragonioides inhabits coastal sand dunes, bluffs, and margins of coastal wetlands (Taylor 1994; Vivrette

2008). The seeds of T. tetragonioides can remain viable in salt water for over one month which allows them to

successfully disperse via ocean currents (Taylor 1994). Due to its apparent ease of dispersal and to anthropo¬

genic activities, this species can be found adventively or naturalized in many parts of the World (Taylor 1994).

In the U.S., T. tetragonioides has been introduced in California, Connecticut, Florida, Massachusetts, North

Carolina, Oregon, Pennsylvania, West Virginia, and Wisconsin (USDA, ARS, National Genetic Resources Pro¬

gram 2013). It is considered invasive only in California, where it is given an invasiveness rating of “limited,”

which indicates that environmental impacts are minor to moderate, or inadequate information is available to

designate a higher rating (California Invasive Plant Council 2006).

Tetragonia tetragonioides, commonly known as New Zealand spinach, is consumed by humans and culti¬

vated as a vegetable in various countries around the World, including the United States (USDA, ARS, National

Genetic Resources Program 2013; Roskruge 2011; Taylor 1994). Its flavor is similar to cultivated spinach Spina-

cia oleracea F., only milder (Roskruge 2011). Xing (2008) also reports that T. tetragonioides is used medicinally.

Voucher specimen: U.S.A.: LOUISIANA: Jefferson Parish: Grand Terre Island, growing on beach on south side of island near abandoned

buildings, only one plant observed, N 29.27574, W 89.93875, elev. near sea level, 23 May 2012, Javed 7 (LSU, #131803).

J. Bot. Res. Inst. Texas 8(2): 661 - 662.2014

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Journal of the Botanical Research Institute of Texas 8(2)

ACKNOWLEDGMENTS

The senior author thanks Ben Stultz with the Louisiana Department of Wildlife and Fisheries for boat trans¬

portation to Grand Terre Island and for being an excellent guide. We thank David Boufford, Harvard Univer¬

sity Herbaria, and Gustavo Heiden, Universidade de Sao Paulo as well as others in the Taxacom community for

their expertise in identifying this species. Our gratitude also extends to Anthony Gendall and Alison Kellow of

La Trobe University for helping obtain text from Flora of Australia. The senior author also extends a special

thank you to Chris Reid of the Louisiana Natural Heritage Program for his expertise and guidance. We greatly

appreciate Robert R. Haynes and Charlotte M. Taylor for their careful reviews of the manuscript.

REFERENCES

California Invasive Plant Council. 2006. California Invasive Plant Inventory Database, (http://www.cal-ipc.org/paf/). Ac¬

cessed 17 December 2013.

Fabris, H.A. 1967. Aizoaceae. In: A.L. Cabrera, ed. Flora de la Provincia de Buenos Aires, vol. 3. Instituto Nacional deTec-

nologia Agropecuaria, Buenos Aires, Argentina. Pp. 171-180.

Gathe, J & L. Watson. 2008. Tetragonia L. Western Australian Flerbarium (1998-). FloraBase—the Western Australian Flora.

Department of Parks and Wildlife, (http://florabase.dpaw.wa.gov.au/browse/profile/21407). Accessed 07 November

2013

Prescott, A. 1984. Tetragonia. In: A.S. George, ed. Flora of Australia, Vol. 4. Australian Government Publishing Service,

Canberra, Australia. Pp. 37-42.

Roskruge, N. 2011. The commercialization of kokihi or New Zealand spinach ( Tetragonia tetragonioides ) in New Zealand.

Agron. New Zealand 41:149-156.

Taylor, C.M. 1994. Revision of Tetragonia (Aizoaceae) in South America. Syst. Bot. 19:575-589.

USDA, ARS, National Genetic Resources Program. 2013. Germplasm Resources Information Network - (GRIN) [Online Data¬

base]. National Germplasm Resources Laboratory, Beltsville, Maryland, (http://www.ars-grin.gov/cgi-bin/npgs/html/

taxon.pl?36385). Accessed 07 November 2013.

Vivrette, NJ. 2008. Tetragonia. In: Flora of North America Editorial Committee. 1993-. Flora of North America north

of Mexico. 16+ vols. Oxford University Press, New York, New York, U.S.A. 4:77-78. (http://efloras.org/florataxon.

aspx?flora_id=1&taxon_id=132571). Accessed 8 July 2013.

Xing, F. 2008. Flora of China. eFIoras. (http://www.efloras.org, 9 January 2014). Missouri Botanical Garden, St. Louis, Mis¬

souri, and Harvard University Herbaria, Cambridge, Massachusetts, U.S.A.

AMSONIA LUDOVICIANA (APOCYNACEAE) NEW TO THE FLORA OF TEXAS (U.S. A.)

Chris Doffitt and Charles Allen*

Patricia and David Lewis

262 CR 3062

Newton, Texas 75966-7003, U.S.A.

Colorado State Univ., Fort Polk Station

1645 23rd St

Fort Polk, Louisiana 71459, U.S.A.

* Contact: charles.m.allen l.ctr@mail.mil

ABSTRACT

Amsonia ludoviciana Vail, Apocynaceae is reported new to Texas based on collections from Newton County and represents the western most

record of the species. The leaves are densely hairy on the abaxial surface, typical of this species.

ABSTRACT

Amsonia ludoviciana Vail, Apocynaceae se cita como nueva para Texas basada en colecciones de Newton County y representa la cita mas

occidental de la especie. Tas hojas son densamente pelosas en el enves, tipico de esta especie.

Our collections of Louisiana bluestar ( Amsonia ludoviciana Vail) from Newton County, Texas are apparently

the first report of this species for the state. It is not listed for Texas (Correll & Johnston 1970; Hatch et al 1990;

Jones et al 1997; Turner et al 2003). During fieldwork for the Flora of Newton County project, we have collected

specimens of the genus Amsonia. A sterile population (Site one) of Amsonia was discovered in west central

Newton County in November 2009 (Allen 21841) and at a different location (Site two), a fruiting specimen was

collected on June 25, 2010 (Allen 21985). Site two was revisited on April 7, 2011 and a collection of the popula¬

tion in flower, Allen 22212 was made. No flowering specimens have been collected at Site one. Site two is ap¬

proximately 2.5 km north of site one. The leaf blades from plants at both locations are densely hairy on the

abaxial surface and at site two, the flowers are hairy, both typical characteristics of A. ludoviciana. This species

is also reported from Georgia, Mississippi, and Louisiana (USDA NRCS 2014). In Louisiana, it is reported from

twelve parishes including three (Calcasieu, Sabine, and Vernon) adjacent to Texas. A. ludoviciana is ranked G3

and S2 in Georgia, S3 in Louisiana, and might be extirpated in Mississippi (Natureserve 2014). The counties in

Georgia are Rockdale, DeKalb, Gwinnett, and Walton. Associated herbaceous species include Dichanthelium

acuminatum (Sw.) Gould & C.A. Clark and Pteridium aquilinum (L.) Kuhn and associated woody species in¬

clude Cyrilla racemiflora L., Ilex opaca Ait., I. vomitoria Ait., Morelia cerifera (L.) Small, and Nyssa biflora Walt.

Our collection is the western most record of this species in the United States.

Voucher specimens for Amsonia ludoviciana: TEXAS. Newton Co.: site 1, forest, off County Road 3107 ca 0.1 mi W of Texas 87 and ca 7.5 mi

S of Bleakwood, 11 Nov 2011, Charles Allen and Brian Early 21841 (TSU); site two, forest, off Texas 82 at State Forest ca. 3 mi W of Texas 87

and 5 mi SW of Bleakwood, 25 Jun 2010, Charles Allen and Krisztian Megyeri 21985, (BRIT);.same location, 7 Apr 2011, Charles Allen 2212

(BRIT, TEX-TT).

ACKNOWTEDGMENTS

The authors would like to thank Sam Kieschnick and two anonymous reviewers for reviewing an earlier draft.

REFERENCES

Correll, D.S. & M.C. Johnston. 1970. Manual of the vascular plants of Texas. Texas Res. Found., Renner, Texas, U.S.A.

Hatch, S.L., K.N. Gandhi, & L.E. Brown. 1990. Checklist of the vascular plants of Texas. Texas Agric. Exp. Sta. Bull. MP 1655.

College Station, Texas, U.S.A.

Jones, S.D., J.K. Wipff, & P.M. Montgomery. 1997. Vascular plants of Texas; a comprehensive checklist including synonymy,

bibliography, and index. University of Texas Press, Austin, U.S.A.

NatureServe. 2014. NatureServe Explorer: An online encyclopedia of life [web application]. Version 7.1. NatureServe,

Arlington, Virginia. Available http://www.natureserve.org/explorer. (Accessed: Aug 19, 2014).

J. Bot. Res. Inst. Texas 8(2): 663 - 664.2014

664

Journal of the Botanical Research Institute of Texas 8(2)

Turner, B.L., H. Nichols, G. Denny, & 0. Doron. 2003. Atlas of the vascular plants of Texas, Vol. 1. Sida, Bot. Misc. 24:1-648.

USDA, NRCS. 2014. The PLANTS Database (http://plants.usda.gov,). National Plant Data Team, Greensboro, North

Carolina 27401-4901 USA.

THE VASCULAR FLORA AND PLANT COMMUNITIES OF CANDY ABSHIER

WILDLIFE MANAGEMENT AREA, CHAMBERS COUNTY, TEXAS, U.S.A.

Jason R. Singhurst

Wildlife Diversity Program

Texas Parks & Wildlife Department

4200 Smith School Road, Austin, Texas 78744 U.S.A.

jason.singhurst@tpwd.state.gov

David J. Rosen

Department of Biology

Lee College

Baytown, Texas 77522-0818, U.S.A.

Amos Cooper

District 7, Wildlife Division

10 Parks & Wildlife Drive

Port Arthur, Texas 77640, U.S.A.

Walter C. Holmes

Department of Biology

Baylor University

Waco, Texas 76798-7388 U.S.A.

ABSTRACT

Field studies at the Candy Abshier Wildlife Management Area, an area of approximately 83 ha (207 acres) of the Gulf Coast Prairies and

Marshes vegetation area, have resulted in a description of the vegetation associations and an annotated checklist of the vascular flora. The

following four associations occur on the property: (1) the newly described Upper Texas Coast Ingleside Sandy Wet Prairie; (2) Gulf Cord-

grass Herbaceous Vegetation; (3) Texas Gulf Coast Five Oak - Sugarberry Forest; and (4) natural depressional ponds. A total of 363 species

belonging to 220 genera and 84 families were recorded. A total of 24 exotic plant species were recorded. The families with the largest number

of species were Poaceae (63), Cyperaceae (48), Asteraceae (43), Fabaceae (14), and Juncaceae (12). Rhynchospora chapmanii and Euthamia

caroliniana were recorded as new to the Texas flora. Three Texas endemic plants were present: Thurovia triflora, Liatris bracteata , and Trad-

escantia subacaulis, the former two being tracked as rare plants. Other noteworthy species included Andropogon capillipes, Asdepias longifo-

liajuncus dliottii, Ludwigia hirtella, Pinguicula pumila, Rhynchospora latifolia, Rhynchospora plumosa, Sabatia gentianoides, and Xyris stricta.

RESUMEN

Investigaciones de campo en el Area de Manejo de Vida Silvestre, Candy Abshier Wildlife Management Area, con un area de aproximada-

mente 83 ha, resultaron en una descripcion de las asociaciones de comunidades de vegetacion y una lista anotada de la flora vascular. Son

cuatro las asociaciones que ocurren en la propiedad: (1) Pradera Mojada Arenosa de la Costa Superior de Texas, Ingleside - nuevamente de-

scrita; (2) Vegetacion Herbacea de Spartina del Golfo; (3) Bosque de Quercus-Celtis del Golfo de Texas; y (4) charcos en depresiones natura-

les. Se documentaron un total de 363 especies de plantas que pertenecen a 220 generos y 84 familias. Fas familias con las mayores represen-

taciones fueron Poaceae (63), Cyperaceae (48), Asteraceae (43), Fabaceae (14) y Juncaceae (12). Rhynchopsora chapmanii y Euthamia carolin¬

iana fueron registradas por primera vez en Texas. Se encontraron tres especies endemicas: Thurovia triflora, Liatris bracteata y Tradescantia

subacaulis, las primeras dos son plantas raras mantenidas en nuestra base de datos. Otras especies de interes son Andropogon capillipes, Asde¬

pias longifoliajuncus elliottii, Ludwigia hirtella, Pinguicula pumila, Rhynchospora latifolia, Rhynchospora plumosa, Sabatia gentianoides y Xyris

stricta.

INTRODUCTION

Currently, only 1.0% (26,304 ha) of the original 2,630,456 ha of coastal prairies still remain in or near its origi¬

nal condition in Texas (Smeins et al. 1991). The coastal prairies, known as the Gulf Coastal Prairie and Marsh¬

es vegetational area (Gould 1960), is a region of significant plant endemism. Carr (2009) mentions 104 of the

320 Texas endemic plants as occurring in at least one location in this region. The number of vascular plant

species (including introductions) in Texas varies from 4839 (Correll & Johnston 1979) to 5100 (Turner et al.

2003). MacRoberts and MacRoberts (2008) cite the number of species in the Gulf Coastal Prairies and Marshes

vegetational area as 2317, showing it to be the richest in species number of any of the vegetation areas of the

state. Yet, as of today, there are but few published floristic lists of portions of the area, thus indicating the area

has largely been neglected.

Topography of the Gulf Coastal Prairie and Marshes vegetational area is flat to very gently rolling from the

Texas-Louisiana border to Harris County (Houston). Included features are barrier islands, beaches, estuarine

J. Bot. Res. Inst. Texas 8(2): 665 - 675.2014

666

Journal of the Botanical Research Institute of Texas 8(2)

lagoons, saline and brackish marshes as well as inland prairies and woodlands of various sorts. This region of

the state is considered transitional and largely defined by the absence of pines, clearly distinguishing it from

the adjacent Pineywoods vegetational area. Elevations are mostly less than 46 m. Most of the region is under¬

lain by poorly consolidated clays, silts, and sands of Pleistocene or Holocene age. Nearest to the coast lies the

Beaumont Formation, which often appears as a featureless plain of black clay. However, sandy materials of

point bars, levees, and other depositional environments are also present. While scarcely perceptible topo¬

graphically, such sands have a major effect on the local diversity of vegetation. Also of note from the city of

Beaumont area is a series of former Pleistocene aged barrier islands that now lie on the current shoreline rather

than offshore. A strip of the Ingleside barrier island extends through the Candy Abshier Wildlife Management

Area (CAWMA) with the sands of this system supporting many species that do not occur on the Beaumont

clays to the west. Inland from the Beaumont Formation is the Fissie Formation, which typically has a loamier,

sandier surface and a more gently rolling topography. Both the Beaumont and Fissie formations routinely ex¬

hibit pimple mounds, several being up to 8 m in diameter and up to 1 m in height. These mounds occur

throughout the prairie portion of CAWMA.

STUDY SITE

Candy Abshier was established as a wildlife management area and selected to commemorate Catherine “Can¬

dy” Cain Abshier, a former Texas Parks and Wildlife employee who promoted wetland conservation recycling

and preservation of historic sites. The site was purchased in 1990. Candy Abshier Wildlife Management Area

(CAWMA) is recognized as one of the most popular points on the Great Texas Coastal Birding Trail (TPWD

2012). CAWMA (TPWD 2012) is located on Galveston Bay and Trinity Bay, near Smith Point in Chambers

County (Fig. 1). CAWMA ranges from a minimum of sea level to a maximum of 4 m in elevation. Climate of the

region is humid subtropical (Bomar 1995) with warm summers while relative humidity is almost always high.

Mean annual precipitation averages 140 cm in Chambers County. The average frost free period is 270 days in

Chambers County (Natural Fibers Information Center 1987). Hurricanes are relatively rare but over time have

exerted a considerable influence on regional vegetation.

The main soil type at CAWMA (Crout 1976) is the Stowell-Clodine association characterized by acid and

alkaline, sandy and loamy soils. This association is slightly above sea level and has little natural drainages. The

Stowell soils are on sandy ridges from 0.5 to 4 m above sea level. Clodine soils are in depressions 0.25 to 1 m

above sea level. Both soils have a surface layer of very dark grey sandy clay loam from 12.7 cm to 38.1 cm in

depth. A small portion of CAWMA near the western boundary consists of Veston soils, alkaline and saline soils

that support aquatic and salt tolerant grasses and sedges in loamy marine sediments that were deposited by

storm tides and wind action. CAWMA consists primarily of coastal prairie habitat with significant important

coastal woodlot (oak mottes) and freshwater ponds. Approximately 31 ha of the area are in this oak motte

habitat. The remaining area is coastal prairie (50 ha) vegetation and freshwater ponds (2 ha).

MATERIATS AND METHODS

The checklist is based upon specimens that were collected during eleven held trips that were made by two of

the authors (Rosen and Singhurst) between 2008 and 2013. Collections are deposited in the Baylor University

Herbarium (BAYFU) and University of Texas Herbarium (TEX-FF). The nomenclature follows Plants Database

(USDA, NRCS 2014). Plants considered non-native are those that are not part of the native to the Bora of Texas

and we followed Plants Database (USDA, NRCS 2014).

FLORISTIC RESULTS

Dominant vegetative over story on the CAWMA (Fig. 2) consists of slender bluestem ( Schizachyrium tenerum),

little bluestem ( Schizachyrium scoparium), brownseed paspalum ( Paspalum plicatulum), few flowered beaksedge

(Rhynchospora rariflora), fascicled beaksedge ( Rhynchospora fascicularis ), nutrushes ( Scleria spp.), gulf cord-

grass ( Spartina spartinae), coastal live oak ( Quercus virginiana), sugarberry ( Celtis laevigata), Carolina cherry

Singhurst et al., Flora of Candy Abshier Wildlife Management Area

667

Fig. 1. Candy Abshier Wildlife Management Area, Chambers County, Texas.

laurel ( Prunus caroliniana), and maidencane ( Panicum hemitomon). Forbs are mostly annual and perennials

with their presence being influenced by rainfall.

A total of 363 species representing 84 families and 220 genera were documented for CAWMA (Appendix

1). Of this total 33 (9.1%) are non-native species. Plant families with the largest number of species were Poaceae

with 63 (17.2 %), Cyperaceae with 48 (13.1%), and Asteraceae with 43 (11.7%). Several rare and regionally sig¬

nificant plant species were documented. These include threeflower broom-weed ( Thurovia triflora), bracted

blazing-star ( Liatris bracteata ), and stemless spider wort ( Tradescantia subacaulis).

The flora of CAWMA was compared with recently published floral checklists of three coastal prairies.

These include the flora of Nash Prairie (Rosen 2007) and Mowotony Prairie (Rosen 2010) in Brazoria County,

both owned by the Nature Conservancy of Texas and the Warren & Jack Road Prairies (Singhurst et al. 2014)

in Harris County, owned by the Katy Prairie Conservancy. The comparisons are presented in Table 1, which

shows comparable naturalness represented by a low percentage of non-natives (9% or less) for all four prairie

sites. This supports the importance for continued management of all four of these prairie preserves in a land¬

scape that is highly susceptible to much higher percentages of non-native plant invasions. In summary all four

of these prairies are extremely important to the conservation of the flora and associated biota of the upper coast

of Texas.

PLANT COMMUNITY RESULTS

Based on dominant species, landscape position, and soil water content, four (4) plant community associations

were documented at CAWMA. One association is a newly described plant community within Nature Serves

classification scheme (N VCS 2014). CAWMA is herbaceous rich with 90% of the species being herbaceous an¬

nuals and perennials while only 10% consists of trees, shrubs, and woody vines. For organizational purposes,

the plant community association descriptions are discussed by system categories (terrestrial and aquatic veg-

668

Journal of the Botanical Research Institute of Texas 8(2)

Fig. 2. Candy Abshier Wildlife Management Area Prairie Vegetation.

etation classes). Within each class, one or more associations may be present. The associations generally follow

the name, a brief description of each, with emphasis on major characteristic species.

Natural Terrestrial Associations

1. Upper Texas Coast Ingleside Sandy Wet Prairie (Schizachyrium tenerum - Rhynchospora rariflora - Rhyn-

chospora plumosa - Rhynchospora fascicularis - Scleria spp. Herbaceous Vegetation)

This community, which encompasses about 52 ha, is an upland coastal prairie of the West Gulf Coastal

Plain with extremely rich flora diversity. The area is dominated by Schizachyrium tenerum, Rhynchospora rari¬

flora, Rhynchospora plumosa, Rhynchospora fascicularis, and Scleria ciliata, S. georgiana, S. pauciflora, and S. re¬

ticularis. The community is developed on Stowell loamy fine sandy loam that is characterized by an abundance

of pimple mounds. Extreme variability in micro-topography is typical, particularly with sandy to sandy loam

soils within this prairie type. The most elevated mounds are very sandy and contain xeric sandhill flora found

at the summit. The inter-mound swales vary in depth, retaining water from days to weeks after rain events,

apparently caused by differential water holding capacity. In a few localized areas sandy ridges, in concert with

a series of mounds, act as dry upland features with seasonal seepage migrating from the bases of ridges and

mounds to lower areas that are similar to seepage bogs. These areas support such plants as Lycopodiella alope-

curoides and Schoenolirion croceum.

Other characteristic flora included Andropogon capillipes, Anthaenantia rufa, Arnoglossum ovatum, Dichan-

thelium acuminatum, D. scoparium, Drosera brevifolia, Eleocharis tuberculosa, Eupatorium rotundifolia, Gratiola

pilosa, Helianthus angustifolia, Hyptis alata, Hypericum crux-andreae, Eudwigia hirtella, Paspalum floridanum,

Singhurst et al., Flora of Candy Abshier Wildlife Management Area

669

Table 1. Floristic comparison of CAWMA flora to three upper coastal prairies in Texas.

Upper Coastal Prairie

County

Hectare

Native (%)

Non-Native (%)

Total

Candy Abshier WMA (TPWD)

Chambers

83 ha

330 (90.9%)

33 (9.1%)

363

Nash Prairie (TNC)

Brazoria

120 ha

289 (89.9%)

22 (7.1%)

311

Mowotony Prairie (TNC)

Brazoria

42 ha

195 (98%)

4 (2 %)

199

Warren & Jack Road Prairies (KPC)

Harris

321 ha

354 (93.1%)

26 (6.9%)

378

Pinguicula pumila, Polygala cruciata, Rhexia mariana, Sabatia gentianoides, Viola lanceolata, Xyris ambigua, and

X. stricta.

2. Gulf Cordgrass Herbaceous Vegetation ( Spartina spartinae Herbaceous Vegetation)

This community at CAWMA occurs on upland flats just above normal tidal reach and encompasses about

6 ha. Spartina spartinae is typically a monodominant, but Setariaparviflora is common and Spartina patens may

be locally codominant. Other characteristic species included Andropogon glomeratus, Cyperus spp., Lythrum

alatum, and Baccharis halimifolia. Several small patches of saline hardpans (slick spots) are embedded within

this plant community. These micro-habitats are sparsely vegetated with hyper-saline flora and included the

Texas endemics Liatris bracteata and Thurovia triflora (Poole et al. 2007). Additional saline flora includes Bor-

richiafrutescens, Limonium carolinianum, Lycium carolinianum, Monanthochloe littoralis, Salicornia depressa, and

Spergularia marina.

3. Texas Gulf Coast Live Oak - Sugarberry Forest (Quercus virginiana - Celtis laevigata/Prunus caroliniana

Forest)

This association (Diamond 1993) includes woodlands occurring along the upper Gulf Coast of Texas north of

Galveston Bay (Brazoria, Chambers, Galveston, and Jefferson counties, TX) on the Ingleside barrier-strand-

plain, a Pleistocene barrier ridge. This community at CAWMA encompasses about 24 ha. The canopy is domi¬

nated by Quercus virginiana. The patchy understory contains Ilex vomitoria and Prunus caroliniana, which may

also reach into the canopy. A few other oak species sporadically occur in this coastal forest and include Quercus

laurifolia, Q. nigra, and Q. phellos. Several Carex spp. and Allium canadense are prominent in the seasonally

damp shady understory during the spring and early summer.

Natural Aquatic Associations

Depressional Pond Types.—Pond flora at CAWMA encompasses about 2 ha and is variable due to differences

in substrate and water depth. Pond flora is mostly centric with the deeper center of ponds dominated by emer¬

gent aquatic plants and then migrating to seasonal flooded outer edges. The deeper portions of the ponds at

CAWMA included emergent aquatic flora such as Nymphaea odorata and Utricularia gibba. With water depth

of 15-61 cm (6-24 inches) Eleocharis quadrangulatajuncus effusus, Panicum hemitomon, Polygonum hydropiper-

oides, Proserpenica palustris, Sacciolepis striata, and Schoenoplectus tabernaemontana are prominent. A water

depth of less than 15 cm (6 inches) is dominated by Hydrolea ovatajuncus repens,Juncus roemeriana, Ludwigia

linearis, and Panicum rigidulum.

DISCUSSION

The overall goal of this research was to document the flora and plant communities of CAWMA. While some

data on the flora of upper coastal prairies have been collected on a few prairie sites, there is a great need to study

additional prairies. Thus this work will also serve as an invitation to conduct additional studies on the coastal

prairies. Southwestern Chambers County contains several small and one large (Middleton Prairie, 324 ha) in¬

tact prairies on the Ingleside geologic formation. Preliminary inventory of this prairie (Bridges et al. 2014) has

documented the presence of a number of species (~15) that seemingly maybe related to tract size and also blurs

the distinction between coastal prairie and the adjacent piney woods. Included among these species were indi¬

cators of tall grass prairie and seasonal seepage plants more typical of Longleaf Pine Wetland Savannahs of

670

Journal of the Botanical Research Institute of Texas 8(2)

southeast Texas. Included were such species as Marshallia graminifolia, Orbexilum simplex, Eriocaulon decangu-

lare, Helianthus mollis, Asclepias obovata, and Rhexia lutea. This circumstance supports a continued need for

additional study of the upper coastal prairies. In conclusion, CAWMA is an extremely important coastal prai¬

rie that contains a high diversity of flora, three Texas endemic plants, and a newly described plant community

association. We highly encourage further flora inventories of the coastal prairies of Texas.

APPENDIX 1. ANNOTATED CHECKLIST OF THE FLORA OF CANDY ABSHIER WILDLIFE MANAGEMENT AREA

The annotated checklist is divided into pteridophytes, gymnosperms, and angiosperms, which subdivided

into monocots and dicots. Families, genera, and species are arranged alphabetically beneath each heading.

Nomenclature follows USDA, NRCS (2014). Names of collectors are abbreviated as follows: DJR = David Rosen

andJRS = Jason Singhurst. An asterisk (*) denotes a non-native species (species not native to Texas). Voucher

specimens were verified at and deposited in the Baylor University Herbarium (BAYLU) and University of Texas

Herbarium (TEX).

The checklist consists of 84 families, 220 genera, and 363 species. The families with the most species are:

Poaceae (63), Asteraceae (43), Cyperaceae (48), Fabaceae (14), andjuncaceae (12).

PTERIDOPHYTES

Azollaceae Lycopodiaceae

Azolla microphylla Kaulf., JRS 20660 Lycopodiella alopecuroides (L.) Cranfill, JRS 15214

Dennstaedtiaceae Polypodiaceae

Pteridium oquilinum (L.) Kuhn var. psuedocoudotum, DJR 4020, Pleopeltis polypodioides [1.) Andrews & Windham ssp. michouxiono

JRS 15244 (Weath.) Andrews & Windham, JRS 20663

GYMNOSPERMS

Cupressaceae Pinaceae

Juniperus virginiana L., JRS 16042 Pinus toedo L., JRS 15163

Toxodium distichum (L.) Rich., JRS 14931

ANGIOSPERMS

MONOCOTYLEDONS

Alismataceae

Sagittariapapillosa Bunch, JRS 14971

Alliaceae

Allium conodense L., DJR 4006 & JRS 20629

Allium drummondii Regel, JRS 16053 & 20628

Nothoscordum bivalve (L.) Britt., JRS 20618

Amaryllidaceae

Cooperia drummondii Herbert, JRS 15260

Hypoxis hirsuta (L.) Coville, DJR 4026

Commelinaceae

Commelina erecta L., DJR 4346, JRS 15265

Tradescantia hirsutiflora Bush, DJR 4028, JRS 14944

Tradescantia subacaulis Bush, DJR 4025, JRS 15243

Cyperaceae

Carexcomplanata Torr. & Hook., DJR 4103& 4601

Carexfestucacea Schkuhr ex Willd., DJR 4084

Carexlongii Mack., DJR 4060

Carextribuloides Wahlenb., JRS 14981

Cyperus articulatus L., JRS 15231

Cyperus entrerianus* Boeckl., JRS 14976

Cyperus haspan L., DJR 4085, JRS 15226

Cyperus reflexus Vahl, DJR 4062

Cyperus strigosus L., JRS 15229

Cyperus virens Michx., JRS 14929

Eleocharis flavescens (Pour.) Urb., DJR 4814 & JRS 20601

Eleocharis macrostachya Britton, DJR 4112

Eleocharis montana (Kunth) Roemer. & Schultes, DJR 4063

Eleocharis montevidensis Kunth, DJR 4113 & JRS 20602

Eleocharispalustris (L.) Roem. & Schult., DJR 4112

Eleocharis quadrangulata (Michx.) Roemer & Schultes, JRS 20600

Eleocharis tuberculosa (Michx.) Roemer & Schultes, DJR 4073,4017,

1019, & 4088

Fimbristylis autumnalis (L.) Roem. & Schult., DJR 4661

Fimbristylis castanea (Michx.) Vahl, JRS 14963

Fimbristylis caroliniana (Lam.) Fernald, DJR 4341

Fuirena breviseta (Coville) Coville, DJR 4304 & 4654, JRS 15213

Fuirena squarrosa Michx., JRS 20603

Isolepis carinata Hook & Arn. ex Torr., JRS 14953

Kyllingaodorata\/ah\, DJR 4107, JRS 14954

Rhynchospora chapmannii M.A. Curtis, DJR 4660

Rhynchospora colorata (L.) H. Pfeiffer, JRS 14958

Rhynchospora corniculata (Lam.) A. Gray, JRS 20662

Rhynchospora debilis Gale, DJR 4094, JRS 15205

Rhynchospora divergens Chapm. ex M.A. Curtis, JRS 15205 & 16054

Rhynchospora fascicularis (Michx.) Vahl, DJR4291 &4293, JRS 15248

Rhynchospora filifolia A. Gray, DJR 4652

Rhynchospora globularis (Chapm.) Small var. globularis, DJR 4051,

4077, & 4083

Rhynchospora glomerata (L.) Vahl, JRS 15186

Rhynchospora gracilenta A. Gray, JRS 15250

Rhynchospora harveyi E. Boot., DJR 4653 & 4657

Rhynchospora inexpansa (Michx.) Vahl, DJR 4294, JRS s.n.

Rhynchospora latifolia (Baldw. ex Ell.) Thomas, JRS 15176

Rhynchospora macrostachya Torr. ex A. Gray, JRS 14972 & 15249

Rhynchospora plumosa Elliott, DJR 4068 & 4087, JRS 14959 & 15161

Rhynchosporarariflora (Michx.) Elliott, DJR 4092 &4052, JRS 15168

Singhurst et al., Flora of Candy Abshier Wildlife Management Area

Rhynchosporo recognita (Gale) Krai, DJR 4093

Sclerio cilioto Michx. var. globro (Chapm.) Fairey, DJR 4110

Scleria georgiana Core, DJR 4057, JRS 15253

Scleriopoucifloro Muhl. ex Willd. var. poucifloro, DJR 4067, JRS 15204

Sclerio reticularis Michx., DJR 4295,4342,& 4648, JRS 15273

Scleria triglomerata Michx., JRS 15180

Schoenoplectus californicus (C.A. Mey.) Palla, JRS 20659

Schoenopiectus tabernaemontani (K.C. Gmel.) Palla, JRS 20661

Iridaceae

Sisyrinchium atlanticum Bickn., DJR 4009, JRS 14943 & 16055

Sisyrinchium minus Engelm. & A. Gray, DJR 4080, JRS 14942

Juncaceae

Juncus brachycarpus Engelm., DJR 4003 & 4808

Juncus bufonius L., DJR 4676, JRS 16059

Juncus coriaceus Mackenzie, JRS 14980

Juncus dichotomus Elliott, DJR 4075, JRS 14973

Juncus elliottii Chapm., DJR 4082, JRS 14921

Juncus effusus L., DJR 4670

Juncus marginatus Rostk., DJR 4081, JRS 15192 & 15221

Juncus polycephalus Michx., JRS 15190

Juncus repens Michx., DJR4816

Juncus roemerianus Scheele, DJR 4066

Juncus tenuis Willd., JRS 15202

Juncus validus Coville var. validus, DJR 4297, JRS 14967 & 15295

Melanthiaceae

Schoenolirion croceum (Michx.) Wood, JRS 20658

Orchidaceae

Spiranthes cernua (L.) Rich., JRS s.n.

Spiranthes praecox (Walter) S. Watson, DJR 4058, JRS 14979

Poaceae

Agrostis hyemalis (Walter) Britton, Sterns & Poggenb., DJR 4004,

JRS 14957

Andropogon capillipes Nash (syn = Andropogon virginicus var.

giaucus), JRS 20567

Andropogon glomeratus (Walter) Britton, Sterns & Poggenb., JRS

20655

Andropogon ternarius DJR 4647 Michx., JRS 20656

Andropogon virginicus L. var. virginicus, DJR 4646

Anthaenantia villosa (Michx.) P. Beauv., JRS 20654

Aristida lanosa Muhl. ex Ell., JRS 20653

Aristida longespica Poir., JRS 20652

Aristida purpuroscens Poir., DJR 4649

Axonopus fissifolius (Raddi) Kuhlm., DJR 4022, JRS 15219

Axonopus furcotus (Fluegge) Hitchc., DJR 4659

Bothriochloa loguroides (DC.) Herter, JRS 20650

Briza minor* L.,JRS 14937

Bromus cothorticus* Vah\, DJR 4680

Cenchrus spinifex Cav., JRS 15281

Chloris goyono* Kunth, JRS 14955

Cynodon doctylon* (L.) Pers., JRS 14946

Dichonthelium ociculore (Desv. ex Poir.) Gould & C.A. Clark subsp.

aciculare, DJR 4050,4016,4053, & 4099, JRS 15252

Dichonthelium acuminatum (Sw.) Gould & C.A. Clark subsp. acumi¬

natum, DJR 4049,4055, & 4086

Dichonthelium acuminatum (Sw.) Gould & C.A. Clark subsp. lind-

heimeri (Nash) Freckmann & Lelong, DJR 4111, JRS 15247

Dichonthelium portoricense (Desv. ex Ham.) B.F. Hansen & Wun-

derlin, DJR 4054

Dichonthelium consonguineum (Kunth) Gould & C.A. Clark, DJR 4070

Dichonthelium oligosonthes (Schult.) Gould subsp. scribnerionum

(Nash) Freckmann & Lelong, DJR 4008

Dichonthelium scobriusculum (Ell.) Gould & C.A. Clark, JRS 20664

671

Dichonthelium scoporium (Lam.) Gould, DJR 4096, JRS 15175

Dichonthelium sphoerocorpon (Elliott) Gould, DJR 4102

Dichonthelium tenue (Muhl.) Freckmann & Lelong, DJR 4071

Distichlis spicata (L.) Greene, JRS 20649

Elymus virginicus L., JRS 20651

Eragrostis elliottii S. Watson, DJR 4655

Eragrostis lugens Nees., DJR 4658

Eragrostis refracta (Muhl.) Scribn., DJR 4662

Eragrostis spectabilis (Pursh) Steud., JRS 20648

Eragrostissecundiflora Presl ssp. oxylepis (Torr.) S.D. Koch., JRS 15188

Eustochys petroeo (Sw.) Desv., DJR 4106

Leersio hexondro Sw., DJR 4072

Monanthochloe littorolis Engelm., JRS 15242

Muhlenbergio copilloris (Lam.) Trin., JRS 20609

Ponicum onceps Michx., JRS 15194

Ponicum repens* L., DJR 4064

Ponicum rigidulum Nees, JRS 15234

Pospolum diiototum* Poir., JRS 15237

Pospolum floridonum Michx., DJR 4340, JRS 15187

Pospolum minus E. Fourn., DJR 4305

Pospolum monostachyum Vasey, DJR 4667 & JRS 20599

Paspalumplicatulum Michx., DJR 4098, JRS 15182

Paspalum setaceum Michx., JRS 4668

Paspalum urvillei* Steud., JRS 15238

Ponicum verrucosum Muhl., DJR 4662

Poo annua* L., DJR 4677

Polypogon monspeliensis* (L.) Desf., JRS 16063

Sacciolepis striata (L.) Nash, DJR 4651, JRS 15294

Schizochyrium tenerum Nees, DJR 4296, JRS 15288

Schizochyrium scoporium (Michx.) Nash, DJR 4666

Setorioporvifloro (Poir.) Kerguelen, JRS 15236

Sorghastrum nutons (L.) Nash, DJR 4665

Sportinopotens (Ait.) Muhl., JRS 15199

Sportino sportinoe (Trin.) Merr. ex Hitchc., JRS 15195 & 15169

Sporoboluspyromidotus (Lam.) Hitchc., JRS 20610

Sporobolus virginicus (L.) Kunth, JRS 15224

Stenotophrum secundotum* (Walt.) Kuntze, JRS 15191

Tridens strictus (Nutt.) Nash, JRS 20647

Vulpio octofloro (Walter) Rydb. var. octofloro, DJR 4076

Smilacaceae

Smilax bono-nox L., JRS 14910 & 15170

Smiloxglouco Walt., JRS 15290 & 14922

Smiloxsmollii Morong, JRS 15255

Typhaceae

Typho domingensis Pers., JRS 15228

Xyridaceae

Xyris ombiguo Kunth, JRS 15208

Xyris jupicoi Rich., DJR 4306

Xyris stricto Chapm., DJR, 4292, JRS 15203

DICOTYLEDONS

Acanthaceae

Dyschoriste linearis (Torr. & A. Gray) Kuntze, DJR 4747

Ruellio humilis Nutt., JRS 15013 & 15206

Altingiaceae

Liquidomborstyrocifluo L., JRS 16061

Apiaceae

Ammoselinum butleri (S. Watson) J.M. Coult. & Rose, DJR 4012

Choerophyllum tointurieri Hook. var. tointurieri, DJR 4674

Centello erecto (L.) Fern., JRS 15258 & 14917

Hydrocotyle bonoriensis Comm, ex Lam., JRS 15223

Hydrocotyle umbelloto L., DJR 4091

672

Journal of the Botanical Research Institute of Texas 8(2)

Limnosciadium pinnotum (Engelm. & A. Gray) Math. & Const., JRS

14960

Trepocarpus aethusae Nutt, ex DC., JRS 14938

Apocynaceae

Asclepios longifolio Michx., DJR 4823, JRS 15293

Asclepias vericillata L., JRS 14918

Cynanchum ongustifolium Pers., JRS 15266

Aquifoliaceae

Ilex vomitoria Walt., JRS 14903

Asteraceae

Ambrosia artemisiifolia L., JRS 15230

Aphanostephus skirrhobasis (DC.) Trel., JRS 15285

Amoglossum ovatum (Walt.) H.E. Robins., JRS 15178

Baccharis halimifolia L., JRS 15198

Bigelowia nuttallii L.C. Anders., JRS 15167

Boltonia diffusa Ell., DJR 4029 & JRS 14915 & 15173

Borrichia frutescens (L.) DC., JRS 15268 & 15012

Cirsium horridulum Michx., JRS 16060

Conoclinium coelestinum (L.) DC., JRS 20646

Coreopsis basalis (A. Dietr.) S.F. Blake, DJR 4105, JRS 14925 & 15261

Coreopsis tinctoria Nutt., JRS 14974 & 5280

Erigeron tenuis Torr. & A. Gray, DJR 4015

Eupatorium compositifolium Walt., JRS 15257

Eupatorium giaucescens Ell., DJR 4299 & JRS 15183

Eupatorium rotundifolium L., DJR 4298 & JRS 15254

Euthamia caroliniana (L.) Greene ex Porter & Britton, JRS 15467

Euthamia leptocephala (Torr. & A. Gray) Greene ex Porter & Britt.,

JRS 20646

Eurybia hemispherica (Alexander) Nesom, JRS 20645

Facelis retusa* (Lam.) Sch. Bip., JRS 20626

Gaillardiapulchella Foug., JRS 15216

Gamochaetapurpurea (L.) Cabrera, DJR 4104

Helianthus angustifolia L., DJR 4343 & JRS 15174

Heterotheca subaxillaris (Lam.) Britt. & Rusby, JRS 15179 & 16056

Iva angusti folia Nutt, ex DC., JRS 20644

Iva annua L., JRS 20643

Krigia dandelion (L.) Nutt., JRS 14948

Liatris acidota Engelm. & A. Gray, DJR 4349 & JRS 15184

Liatris bracteata Gaiser, JRS 15172

Palafoxia hookeriana Torr. & A. Gray, JRS 20667

Pityopsis graminifolia (Michx.) Nutt., JRS 20666

Pluchea rosea Godfrey, DJR 4300

Piuchea foetida (L.) DC., JRS 15165

Pyrrhopappus carolinianus (Walter) DC., JRS 16041

Rudbeckia hirta L., JRS 15165

Solidago sempervirens L., JRS 20604

Solidago odora Ait., JRS 20642

Solidago rugosa Mill., JRS 20641

Solidago tortifolia Ell., DJR 4023 & JRS 14965

Soliva sessilis* Ruiz & Pav., JRS 20627

Sonchus asper* (L.) Hill, JRS 14935

Symphyotrichum dumosum (L.) Nesom var. dumosum, JRS 20671

Verbesina virginica L., JRS 20614

Brassicaceae

Cardaminepennsylvanica Muhl. Ex. Willd., JRS 20607

Lepidium densiflorum Sch rad., JRS 16046

Lepidium virginicum L., JRSI4952 & 16042

Cabombaceae

Cabomba caroliniana A. Gray, JRS 20597

Campanulaceae

Lobelia appendiculata A. DC., JRS 20592

Lobeliapuberula Michx., DJR 4663, JRS 20633

Triodanisperfoliata (L.) Nieuw., JRS 14936

Cannabaceae

Celtis laevigata Willd., JRS 2063

Caprifoliaceae

Lonicera japonica* Thunb., JRS 14901

Sambucus nigra L., JRS 20617

Caryophyllaceae

Arenaria serpyllifolia* L., JRS 20620

Cerastium glomeratum* Thuill., DJR 4673 & JRS 20605

Silene antirrhina L., JRS 14907 & 16051

Spergularia salina J. Presl & C. Presl., DJR 4013

Stellaria media* (L.) Vill., DJR 4678

Chenopodiaceae

Salicornia depressa Standi., JRS 14977

Clusiaceae

Hypericum drummondii (Grev. & Hook.) Torr. & A. Gray, JRS 20640

Hypericum crux-andreae (L.) Crantz, DJR 4350, JRS 14902

Hypericum galioides Lam., DJR 4807

Hypericum gymnanthum Engelm. & A. Gray, DJR 4095

Hypericum hypericoides (L.) Crantz, DJR 4348, JRS 14919 & 15251

Hypericum mutilum L., JRS 20639

Hypericum punctatum Lam., DJR 4824

Convolvulaceae

Evolvulus sericeus Sw., DJR 4014

Ipomoea sagittata Poir., JRS 15189

Cuscutaceae

Cuscuta cuspidata Engelm., JRS 15262

Droseraceae

Drosera brevifolia Pursh, DJR 4032

Ebenaceae

Diospyros virginiana L., JRS 14909

Euphorbiaceae

Chamaesyce maculata (L.) Small, JRS 15232

Croton glandulosa L., JRS 15185

Triadicasebifera* (L.) Small, JRS 14905 &15220

Fabaceae

Baptisia bracteata Muhl. ex Elliott var. leucophaea (Nutt.) Kartesz &

Gandhi, DJR 4021, JRS 15200

Centrosema virginianum (L.) Benth., DJR 4347, JRS 15217

Chamaecrista fasciculata (Michx.) Greene, DJR 4344, JRS 15210

Crotalaria sagittalis L., DJR 4109

Desmanthus illinoensis (Michx.) McM. ex B.L. Rob. & Fern., JRS 15235

Medicago lupulina* L., DJR 4675

Medicagopolymorpha* L., JRS 16050

Melilotus officinalis * (L.) Lam, JRS 14961

Mimosa hystricina (Small ex Britt. & Rose) B.L. Turner, DJR 4822,

JRS15177

Neptuniapubescens Benth., JRS 15287

Sesbaniapunicea* (Cav.) Benth., DJR 4351

Tephrosia onobrychoides Nutt., JRS 20638

Vida ludoviciana Nutt., JRS 14939

Vida minutiflora Dietr., JRS 14949

Fumariaceae

Corydalis micrantha (Engelm. ex A. Gray) A. Gray, JRS 20606

Fagaceae

Quercus laurifolia Michx., JRS 14913

Quercus nigra L., JRS 14911

Quercus nuttallii Palmer, JRS 14930

Singhurst et al., Flora of Candy Abshier Wildlife Management Area

673

Quercus phellos L., JRS 14927

Quercus virginiono P. Mill., JRS 14912

Gentianaceae

Centaurium pulchellum* (Sw.) Druce, DJR 4010, JRS 14962

Eustoma exoltotum (L.) Salisb. ex G. Don ssp. exoltotum, JRS 15241

Sabatia campestris Nutt., JRS 14966

Sabatia gentionoides Ell., JRS 15211 & DJR 4813

Geraniaceae

Geranium carolinianum L., JRS 14945 & 16049

Haloragaceae

Proserpinacapalustris L., DJR 4815 & 4811, JRS 20668

Hydrophyllaceae

Hydro lea ovata Choisy., JRS 15222

Juglandaceae

Carya aquatica (Michx. f.) Nutt., JRS 15245

Lamiaceae

Hyptis alata (Raf.) Shinners, JRS 15279

Hedeoma hispida Pursh, JRS 20619

Monarda punctata L., JRS 15181

Physostegia intermedia (Nutt.) Engelm. & A. Gray, DJR 4089

Physostegiapulchella C. L. Lundell, JRS 14978

Salvia lyrata L., JRS 16064

Scutellaria integrifolia L., JRS 20611

Lauraceae

Cinnamomum camphora* (L.) J. Presl, JRS 14926

Persea borbonia (L.) Spreng., JRS 20608

Sassafras albidum (Nutt.) Nees, JRS 20634

Lentibulariaceae

Pinguiculapumila Michx., DJR 4027

Utricularia gibba L., DJR 4817, JRS 20637

Utricularia subulata L., DJR 4031, JRS 14934

Linaceae

Linum medium (Planch.) Britton var. texanum (Planch.) Fernald, DJR

4302,4078, JRS 15177 & 15263

Linum striatum Walt., JRS 15196

Linum sulcatum Riddell, JRS 16040

Loganiaceae

Gelsemium sempervirens (L.) J. St.-Hil., DJR 4672

Mitreola sessilifolia (J.F.Gmel) G. Don, JRS 15209

Lythraceae

Lythrum alatum Pursh, JRS 14975 & 15239

Lythrum lineare L., JRS 15272

Malvaceae

Hibiscus lasiocarpos Cav., JRS 1520

Malvaviscus arboreus Dill, ex Cav., JRS 20621

Sida rhombifolia L., JRS 14947 & 15215

Melostomataceae

Rhexia mariana L., JRS 15207

Myricaceae

Morelia caroliniensis (P. Mill.) Small, JRS 15246

Morelia cerifera (L.) Small, DJR 4018, JRS 14904

Nymphaeaceae

Nymphaea odorata Aiton, DJR 4818, JRS 20598

Nyssaceae

Nyssa sylvatica Marsh., JRS 15193

Onagraceae

Gaura lindheimeri Engelm. & A. Gray, JRS 15162

Ludwigia glandulosa Walter, JRS 4812

Ludwigia grandiflora* (M. Micheli) Greuter & Burdet ssp. grandiflora,

JRS15227

Ludwigia hirtella Raf., JRS 15256

Ludwigia linearis Walter, DJR 4352

Oenothera heterophylla Spach, JRS 15225

Oenothera laciniata Hill, DJR 4679

Oenothera linifolia Nutt., DJR 4005

Oenothera speciosa Nutt., JRS 14956,20613

Orobanchaceae

Agalinis fasciculata (Ell.) Raf., DJR 4097, JRS 15164 & 15283

Agalinis martitima (Raf.) Raf., JRS 20612

Agalinis purpurea (L.) Pennell, JRS 20669

Buchnera americana L., DJR 4079, JRS 14924 & 15286

Castilleja indivisa Engelm., JRS 20622

Oxalidaceae

Oxalis dillenii Jacq., JRS 16044

Oxalis violacea L., JRS 20670

Phytolaccaceae

Phytolacca americana L., JRS 15282

Plantaginaceae

Mecardonia acuminata (Walt.) Small, JRS 20636

Nuttallanthus canadensis (L.) Sutton, JRS 14940 & 20631

Plantago pusilla Nutt., JRS 16045

Plantago lanceolata* L., JRS 14950

Plantago virginica L., DJR 4011

Veronicaperegrina L., JRS 20623

Plumbaginaceae

Limonium carolinianum (Walt.) Britt., JRS 15267

Polygalaceae

Polygala cruciata L., DJR 4345, JRS 15292

Polygala leptocaulis Torr. & A. Gray, JRS 15259

Polygala mariana Mill., DJR 4108

Polygonaceae

Polygonum hydropiperoides Michx., JRS 14906

Rumexcrispus* L., JRS 16052

Rumexhastatulus Baldw., JRS 14941

Portulacaceae

Claytonia virginica L., JRS 20630

Phemeranthus parviflorus (Nutt.) Kiger,JRS 15269 & 14968

Primulaceae

Anagallis arvensis* L., JRS 15278

Anagallis minima (L.) Krause, R 4024, JRS 14982

Ranunculaceae

Anemone berlandieri Pritz., JRS 20625

Ranunculus hispidus Michx., JRS 20624

Rosaceae

Prunus caroliniana (Mill.) Ait., JRS 14933

Rosa bracteata* J.C. Wendl., JRS 14923

Rosa laevigata * Michx., DJR 4669

Rubus arguta Link, JRS 15166

Rubus r/ograndis Bailey, JRS 14916

Rubiaceae

Diodia teres Walt., JRS s.n.

Diodia virginiana L., JRS 15264

Galium aparine L., JRS 16048

Galium tinctorium L., JRS 14900

Houstonia pusilla Schoepf, JRS 20595

Houstonia micrantha (Shinners) Terrell, JRS 20593

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Journal of the Botanical Research Institute of Texas 8(2)

Houstonio roseo (Raf.) Terrell, JRS 20594

Oldenlandia uni flora L., DJR 4100 & JRS 15284

Sherardia arvensis* L., DJR 4681

Rutaceae

Zanthoxylum clava-herculis L., DJR 4819

Salicaceae

Salixnigra Marshall, DJR 4806

Sapotaceae

Sideroxylon lanuginosum Michx. JRS 20591

Saxifragaceae

Lepuropetalon spathulatum Elliott, JRS 20596

Scrophulariaceae

Gratiola brevifolia Raf., JRS 14920 & 15291

Gratiolapilosa Michx., DJR 4303, JRS 14914 & JRS 15212

Solanaceae

Lycium carolinianum Walt., JRS 14969 & 15271

Physalis angulata L., JRS 15276

Solanum ptycanthum Dunal, JRS 16057

Tetrachondraceae

Polypremum procumbens L., DJR 4301, JRS 15275

Ulmaceae

Ulmus crassifolia Nutt., JRS 20616

Valerianaceae

Valerianella radiata (L.) Dufr., JRS 16047

Verbenaceae

Lantana camara* L., JRS 14900

Phyla lanceolata (Michx.) Greene, JRS 14964 & 15240

Verbena brasiliensis* Veil., JRS 14951

Verbena halei Small, JRS 14928

Violaceae

Viola lanceolata L., DJR 4030, JRS 15289

Violasororia Willd., JRS 20632

Vitaceae

Ampelopsis arborea (L.) Koehne, JRS 15277 & 14932

Vitis rotundifolia Michx., DJR 4820

Vitis mustangensis Buckl., JRS 15233

ACKNOWLEDGMENTS

We are indebted to Michael Warriner, John Davis, Lynn Polasek, and David Forrester, all of Texas Parks and

Wildlife Department (TPWD, Austin) for suggesting and supporting this project. The careful reviews of Mike

Palmer and Larry E. Brown are greatly appreciated.

REFERENCES

Bomar, E.G. 1995. Texas weather. University of Texas Press, Austin, U.S.A.

Bridges, E.L., S.L. Orzell, K. Kindsher, & J.M. Poole. 2014. Middleton Prairie flora: Plant Resources Center Flora of Texas Data¬

base. University of Texas, Austin, U.S.A. http://orchid.biosci.utexas.edu/Texas_ll.html

Carr, W.R. 2009. No place but Texas: Annotated list of plant taxa endemic to the Lone Start State. Incomplete working

draft, The Nature Conservancy of Texas, San Antonio. U.S.A.

Correll, D.S. & M.C. Johnston 1979. Manual of the vascular plants of Texas. Texas Research Foundation, Renner, U.S.A.

Crout, J.D. 1976. Soil survey of Chambers County, Texas. United States Department of Agriculture, Soil Conservation

Service, U.S.A.

Diamond, D.D. 1993. Classification of the plant communities ofTexas (series level). Unpublished document. Texas Natural

Fleritage Program, Austin, U.S.A.

Gould, F.W., G.O. FIoffman, & C.A. Rechenthin. 1960. Vegetation areas ofTexas. Texas Agricultural Experimental Station,

Leaflet No. 492,Texas A&M Univ., College Station, U.S.A.

Jones, S.D., J.K. Wipff, & P.M. Montgomery. 1997. Vascular plants ofTexas. University ofTexas Press, Austin, U.S.A.

MacRoberts, M.H & B.R. MacRoberts. 2008. Species richness of vegetational areas ofTexas: A first approximation. J. Bot.

Res. Inst.Texas 2(2):1373-1379.

National Vegetation Classification System. 2014. Nature Serve Explorer, Ecological Classifications <http://www.natureserve.

org/explorer/classeco.htm>

Natural Fibers Information Center. 1978. The climates ofTexas counties. Austin Bureau of Business Research, University of

Texas, Austin, U.S.A.

Poole, J.M., W.R. Carr, D. Price, & J.R. Singhurst. 2007. A field guide to the rare plants ofTexas. Texas A&M Press, College

Station, U.S.A.

Rosen, DJ. 2007. The vascular flora of Nash Prairie a Coastal Prairie remnant in Brazoria County, Texas. J. Bot. Res. Inst.

Texas 1(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(1):489-495.

Singhurst, J.R., DJ. Rosen, & W.C. Holmes. 2009. Two new additions to the vascular flora ofTexas. Phytologia 91(1 ):69-72.

Singhurst, J.R., N. Shackelford, W. Newman, J.N. Mink, & W.C. Holmes. 2014. The ecology and abundance of Hymenoxystexana

(Asteraceae). Phytoneuron 2014-19:1-19.

Singhurst et al., Flora of Candy Abshier Wildlife Management Area

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Smeins, F.E., D.D. Diamond, & C.W. Hanselka. 1991. Coastal prairie. In: R.T. Coupland, ed. Ecosystems of the world: Natural

grasslands: Introduction and western hemisphere. Elsevier, New York, U.S.A. Pp. 269-290.

Texas Parks and Wildlife Department. 2012. http://www.tpwd.state.tx.us/huntwild/hunt/wma/find_a_wma/list/?id=36

Turner, B.L., H. Nichols, G. Denny, & 0. Doron. 2003. Atlas of the vascular plants ofTexas. Sida, Bot. Misc. 24. (2 vols.). Botani¬

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Journal of the Botanical Research Institute of Texas 8(2)

ANNOUNCEMENTS

2014 DELZIE DEMAREE TRAVEL AWARD RECIPIENT

The 26th Annual Delzie Demaree Travel Award was presented at the 61st Annual Systematics Symposium

(10-11 Oct 2014) at the Missouri Botanical Garden, St. Louis. One student was presented the Travel Award:

Keir Michael Wefferling, University of Wisconsin-Milwaukee.

The 2014 Travel Awards were underwritten by 1) Contributors to the Delzie Demaree Travel Award En¬

dowment, and 2) Members of the Delzie Demaree Travel Award Committee.

Anyone interested in making a contribution to Delzie Demaree Endowment Fund, which supports the

travel award, may make contributions by VISA or MasterCard or by a check, payable to Botanical Research

Institute of Texas, to Barney Lipscomb, 1700 University Drive, Fort Worth, TX 76107-3400, U.S.A. 1-817-332-

7432; Email: barney@brit.org. Thank you.

THE 2015 APPLICATIONS FOR THE DELZIE DEMAREE TRAVEL AWARD

Applications for the 2015 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 2015 are 9-10 October 2015 (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(2): 676.2014

IN MEMORIAM

LANDON E. McKINNEY

( 1949 - 2014 )

Ronald L. Jones Ralph L. Thompson

Eastern Kentucky University Herbarium Berea College Herbarium

Richmond, Kentucky 40475, US.A. Berea, Kentucky 40404, US.A.

Landon E. McKinney, a talented field botanist and naturalist, passed away on Thursday, June 5th, 2014, at the

Wade Park Veteran’s Medical Center in Cleveland, Ohio, at the age of 65. Landon Earl McKinney was born May

17, 1949 in Nashville, Tennessee, to the late Lawrence Vern and Constance Joy McKinney After graduating

from Donelson High School in Nashville in 1967, Landon joined the Navy and served as a Marine Navy Corps-

man from 1968-1970; he was a held combat corpsman in Vietnam during 1969. He earned a B.S. in Biology in

1973 from Middle Tennessee State University (MTSU) and subsequently his M.S. in Botany in 1977 from

MTSU. His master’s thesis was “ Preliminary Studies of the Acaulescent Blue Violets (Viola) with Special Reference

to Middle Tennessee,” which began his life-long fascination with the stemless blue violets. He was a Research

Associate in Plant Taxonomy at Vanderbilt University from 1985-1989, where he worked with Dr. Robert Krai.

After his family, Landon’s true passion was botany, which he focused upon throughout his life. He served

as a held botanist for the Tennessee Natural Heritage Program and the Kentucky State Nature Preserves Com¬

mission, and worked as an environmental consultant for several different companies in the eastern U.S.,

mostly in Kentucky, Ohio, and Tennessee. He conducted numerous status surveys for state and federal rare

plant species, worked on habitat analyses, assessed wetland communities, and made management recommen¬

dations on preserving natural habits and controlling invasive species. In recent years he established research

affiliations with Northern Kentucky University, University of Cincinnati, and Ohio State University.

Landon was active in the Tennessee Academy of Science and the Kentucky Academy of Science, serving

J. Bot. Res. Inst. Texas 8(2): 677 - 678.2014

678

Journal of the Botanical Research Institute of Texas 8(2)

terms as Chair and Secretary for both academies. In 2003-2005, he was a member of the Editorial Board of

Castanea, Journal of the Southern Appalachian Society, and served as Chair in 2005-2006. Landon was heavily

involved in the Kentucky Native Plant Society (KNPS) for many years; he regularly attended annual meetings

and frequently led held trips. He served as the KNPS President from 1992-1996 and from 2002-2006, and his

strong leadership was instrumental in maintaining and enhancing the activities of the society.

Landon was an avid plant collector throughout his career and collected thousands of specimens during

all his botanical activities and deposited them at regional herbaria such as EKY, TENN, and VDB. Landon ea¬

gerly collected difficult grasses, rushes, and sedges, and possessed a special skill in identifying these problem¬

atic species. Landon was especially skillful in Carex and Viola, and provided the keys to these two genera for

the 2005 book, Plant Life of Kentucky: An Illustrated Guide to the Vascular Flora. Landon’s primary passion

throughout his career was the genus Viola, particularly the stemless blue violets, where he was nationally

known for his expertise. The seminal publication of his career occurred in 1992 with a book on the stemless

blue violets—A Taxonomic Revision of the Acaulescent Blue Violets (Viola) of North America, published by the

Botanical Research Institute of Texas. This work is highly regarded and had a great influence on subsequent

studies of this group of violets, with other experts often citing Landon’s work and comparing it to their inter¬

pretations. Landon authored or co-authored approximately 50 scientific publications and professional presen¬

tations on Viola and on a variety of other topics. He was a frequent presenter at the Annual Meetings of the

Association of Southeastern Biologists, Kentucky Academy of Science, and Tennessee Academy of Science.

Landon continued his work and interest in Viola until the very end with the completion of his work with

R. John Little on the Violaceae for the Flora of North America which is now “in press.” This past April, Landon

sent an email to local botanists concerning the recent discovery of a new Kentucky county record for Viola

egglestonii, and noted,... “after 40+ years of working with violets, violets still excite me ... this population is as

classical cedar glade violet as you can get...”

Landon is survived by his devoted wife of 31 years, Lela; children Adam McKinney, Amanda (Troy) Chit¬

wood, and Eric (Amy) McKinney; grandchildren Landon, Lucas, Erica, Katelyn, and Whitney. Landon will be

sadly missed by his many extended family members, special friends, and botanical colleagues.

Reviewers

679

Reviewers

Volume 8 (2014)

120 reviewers: several individuals reviewed more than one manuscript.

We sincerely apologize if your name was accidently left out.

Thank you for supporting Journal of the Botanical Research Institute of Texas.

Abbott, J. Richard

Acedo, Carmen

Alexander, Jason A.

Alexander, Patrick

Almeda, Frank

Al-Shehbaz, Ihsan

Anderson, Loran C.

Baker, Brent Travis

Balslev, Henrik

Barker, Robyn M.

Barrie, Fred R.

Binder, Manfred

Boyd, Steve

Brown, Larry E.

Bryson, Charles T.

Budke, Jean C.

Cantino, Philip D.

Carlquist, Sherwin

Carmo E. Amaral, Maria do

Carr, Bill

Carter, J.R.

Chorghe, Alok

Colin, Rafael Torres

Cox, Pat

Darwin, Steven P.

Daschowdhury, Sangita

Dickinson, Timothy A.

Downie, Stephen R.

Duling, Darrin W.

Eliasson, Uno

Escobar Lechuga, Roberto

Estes, Dwayne

Estrada-Castillon, Eduardo

Fertig, Walter

Feuillet, Christian

Fleming, Gary

Franck, Alan

Freeman, Craig C.

Gandhi, Kanchi

Greuter, Werner

Hammel, Barry

Haynes, Robert R.

Hemmerly, Thomas E.

Henderson, Andrew

Henrickson, James

Herrera Arrieta,Y.

Hill, Steve

Hodgson, Wendy C.

Hogan, Tim

Holst, Bruce K.

Horn, Charles N.

Iganci, Joao Ricardo

Jaramillo, Patricia

Jarvis, C.E.

Ju, Yu-Ming

Kawasaki, Lucia

Keener, Brian R.

Keith, Eric L.

Kieschnick, Sam R.

Kirkbride, Jr., Joseph H.

Klitgaard, B.B.

Kriebel, Ricardo

Lammers, Thomas G.

Landrum, Leslie R.

Le Roux, M. Marianne

Leon de la Luz, Jose Luis

Lipscomb, Barney

Lorence, David H.

MacDougal, John

MacRoberts, Michael

Mansfield, Don

McDonald, Andrew

McMullen, Conley K.

Meyer, Harry

Miller, Joseph T.

Moore, Gerry

Morton, Cynthia M.

Nelson, Allan D.

O'Kane, Steve

Palmer, Mike

Pennington, T.D.

Pilz, George E.

Pipoly III, John

Poindexter, Derick B.

Powell, A. Michael

Rabeler, Richard K.

Rehman,Tiana

Reveal, James L.

Riefner, Rick

Rogers, Jack D.

Rondeau, Renee

Rosen, David J.

Rundel, Phil

Rye, Barbara L.

Saarela, Jeffery M.

Salywon, Andrew

Sanchez-Gonzalez, Arturo

Schlessman, Mark A.

Schwartzberg, Ezra

Serviss, Brett

Sirotnak, Joe

Skog, Laurence E.

Spaulding, Daniel Dieter

Steinmann, Victor

Suarez Suarez, Luz Stella

Swadek, Rebecca K.

Swenson, Ulf

Taylor, Charlotte M.

Tepe, Eric

Traverse, Alfred

Turner, Billie

Ulloa, Carmen

Utteridge, Timothy M.A.

Villasenor Rios, Jose L.

Vincent, Michael A.

Vovides, Andrew

We I dy, Troy

Wilson, Andy

Wunderlin, Richard P.

Zarucchi, James L.

680

Journal of the Botanical Research Institute of Texas 8(2)

Index to 65 Titles with 144 Authors

Volume 8 (2014)

We know you have a choice.

Thank you for choosing and supporting Journal of the Botanical Research Institute of Texas.

A floristic inventory of Dagny Johnson Key Largo Hammock Bo¬

tanical State Park and immediately adjacent lands (Monroe

County), Florida, U.S.A. by George J. Wilder, Susan V. Sprunt,

Janice A. Duquesnel, and Susan F. Kolterman— 8 ( 1):227

A gasteroid fungus, Palaeogaster micromorpha gen. & sp. nov.

(Boletales) in Cretaceous Myanmar amber by George O. Poinar,

Jr., Donis da Silva Alfredo, and luri Goulart Baseia— 8(1 ):139

A morphologically based taxonomic reevaluation of the genus Stip-

ulicida (Caryophyllaceae), with comments on rank by Derick B.

Poindexter, Kateland E. Bennett, and Alan S. Weakley— 8 ( 2):419

A new hedge-nettle (Stachys: Lamiaceae) from South Carolina,

U. S.A. by John B. Nelson and Douglas A. Rayner— 8 ( 2):25

A new species of Cremosperma (Gesneriaceae) from northeastern

Peru by Brian R. Keener and John L. Clark— 8(1 ):57

Anewspecies ofGarnotia (Poaceae) from Kerala, India by C.N.Sunil,

V. V. Naveen Kumar, and M.G. Sanilkumar— 8 ( 2):517

A new variety of Phanera glauca subsp. tenuiflora (Fabaceae:

Caesalpinioideae) from India by Rajib Gogoi and Subir Ban-

dyopadhyay— 8(1 ):71

A quantitative study of the vegetation surrounding populations of

Zigadenus densus (Melianthiaceae) at Fort Polk in west central

Louisiana, U.S.A. by Jacob Delahoussaye, Charles Allen, Stacy

Huskins, and Ariel Dauzart— 8(1 ):253

A quantitative study of the vegetation surrounding populations of

Uvulariasessilifolia (Colchicaceae) at Fort Polk in west central

Louisiana, U.S.A. by Ariel Dauzart, Charles Allen, Stacy Huskins,

and Jacob Delahoussaye— 8 ( 1):261

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

David J. Rosen— 8(1 ):381

Agave x madrensis (Asparagaceae), a putative hybrid from the

Sierra Madre Oriental, Mexico by Jose A. Villarreal Quintanilla,

Abraham Ramirez Gamez, Eduardo Estrada Castillon, Dino

U. Gonzalez Uribe, and Diana Jasso de Rodriguez— 8 ( 2):441

American mistletoe (Phoradendron leucarpum ssp. leucarpum,

Viscaceae) occurrence in host trees within the city of Cairo,

Alexander County, Illinois, and its incidence in Illinois, U.S.A.

by Ralph L.Thompson— 8 ( 2):641

Amsonia ludoviciana (Apocynaceae) new to the flora of Texas,

U.S.A. by Chris Doffitt, Charles Allen, Patricia Lewis, and David

Lewis— 8 ( 2):663

An annotated flora of Reed Plateau and adjacent areas, Brew¬

ster County, Texas, U.S.A. by Wendy Weckesser and Martin

Terry— 8(1 ):353

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

Daniel M. McNair and Mac H. Alford— 8(1 ):267

Browneopsis puyensis (Leguminosae: Caesalpinioideae: Detar-

ieae), a new species from Amazonian Ecuador by David A. Neill

and Mercedes Asanza— 8 ( 2 ): 511

Calathea cofaniorum and C. shishicoensis, new endemic species

of Marantaceae from Ecuador by Helen Kennedy— 8 ( 1):37

Calathea gordonii (Marantaceae), a new endemic Panamanian

species by Helen Kennedy— 8(1 ):31

Calathea rubribracteata, a new endemic species of Marantaceae

from Colombia by Helen Kennedy— 8 ( 2):493

Cyperusstewartii (Cyperaceae), a new species from Cocos Island,

Costa Rica by Gordon C. Tucker— 8 ( 1):25

Distigouania irregularis (Rhamnaceae) gen. et sp. nov. in Mid-

Tertiary amber from the Dominican Republic by Kenton L.

Chambers and George O. Poinar, Jr.— 8 ( 2):551

Dos nuevas especies de Myrcia (Myrtaceae), del Pacifico sur de

Costa Rica by Daniel Santamaria Aguilar, Armando Estrada Ch.

y Reinaldo Aguilar— 8 ( 2):449

Ecology and conservation of Acacia and Prosopis (Fabaceae)

woodlands of the Mojave Desert, U.S.A. by Scott R. Abella and

Kenneth L. Chittick— 8 ( 1):175

Errata: Four new annual species of Euphorbia section Tithyma-

lus (Euphorbiaceae) from North America by Mark H. May-

field— 8(1 ):85

Estructura de un zacatal de toboso (Hilaria mutica: Poaceae)

asociado a sustrato igneo en el noreste de Coahuila, Mexico

by Juan A. Encina-Dominguez, Jesus Valdes-Reyna y Jose A.

Villarreal-Quintanilla— 8 ( 2):583

Expanded distribution of Gratiola quartermaniae (Plantaginaceae)

in Texas, U.S.A. by Kimberly Norton Taylor and Robert J.

O'Kennon— 8(1 ):333

First valid place of publication of Duchesnea indica (Rosaceae:

Potentilleae) by James L. Reveal and Barbara Ertter— 8 ( 1):83

Five new Myrtaceae from southeastern Brazil by Marcos Sobral,

Marcelo da Costa Souza, Ludovic Jean Charles Kollmann,

Osny Tadeu Aguiar, Andre Luis Casarin Rochelle, and Kelly

Antunes— 8 ( 2):497

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

Mexico Jose Humberto Vega-Mares, Andres Eduardo Estrada-

Castillon, Jose Angel by Villarreal-Quintanilla, and Gustavo

Quintana Martinez— 8(1 ):151

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

de Cristo Mountains by Jill Larson, Brian Reif, B.E. Nelson, and

Ronald L. Hartman— 8 ( 1):271

Glossoloma velutinum (Gesneriaceae), a new species from the

Cordillera Central of the Colombian Andes by Larri A. Rodas

and John L. Clark— 8 ( 1):43

Hexasepalumteres(Rubiaceae), a new combination by Joseph H.

Kirkbride, Jr.— 8(1 ):17

Jarilla chocola (Caricaceae), un nuevo genero y especie para la

flora de El Salvador by Frank Sullyvan Cardoza Ruiz, Jose Ledis

Linares y Ana Eugenia Aguilar Grijalva— 8 ( 2):595

Landon E. McKinney (1949-2014) by Ronald L. Jones and Ralph L.

Thompson— 8 ( 2):677

Limitations to natural production of Lophophora williamsii (Cac-

taceae) III. Effects of repeated harvesting at two-year intervals

for six years in a SouthTexas (U.S.A.) population by Martin Terry,

Keeper Trout, Bennie Williams, Teodoso Herrera, and Norma

Fowler— 8 ( 2):541

Lomatium swingerae: a new species of Lomatium (Umbel-

liferae) from the Joseph Plains, Idaho, U.S.A. by Richard P.

McNeill— 8 ( 2):395

Low genetic diversity and poor dispersal, but not conservation sta¬

tus rank, are linked to climate change vulnerability by Cynthia

M. Morton and Matthew D. Schlesinger— 8 ( 1):305

New geographical and morphological data for Sideroxylon rec-

linatum subspecies austrofloridense (Sapotaceae), a taxon

endemic to southeastern peninsular Florida, U.S.A. by Paul T.

Corogin and Walter S. Judd— 8 ( 2):403

New species and combinations in Solanum section Androceras

(Solanaceae) by Stephen R. Stern, Lynn Bohs, and Jeffrey

Keeling— 8(1 ):1

Noteworthy vascular plant collections from the Red River of

Index

681

Arkansas and Louisiana, U.S.A. by Christopher Reid and M.

Jerome Lewis— 8(1 ):197

Passiflora soliana, una especie nueva de Passiflora (Passiflora-

ceae) del Pacifico Sur de Costa Rica by Armando Estrada Ch. &

Gerardo Rivera— 8(1 ):19

Pleurisanthes flava (Icacinaceae): A new record for Brazil by

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

Alves— 8(1 ):169

Primer registro de Astrocasia peltata (Euphorbiaceae) en Costa

Rica by Irene Calderon Sanou— 8(1 ): 165

Recent findings on the gypsum flora of the rim of the Guadalupe

Mountains, New Mexico, U.S.A.: A new species of Nerisyrenia

(Brassicaceae), a new state record, and an updated checklist

by Patrick J. Alexander, Norman A. Douglas, Helga Ochoterena,

Hilda Flores-Olvera, and Michael J. Moore— 8 ( 2):383

Redescubrimiento y nuevos registros de Cuscatlania vulcani-

cola (Nyctaginaceae) para El Salvador by Jose Ledis Linares,

Frank Sullyvan Cardoza Ruiz y Patricia Hernandez-Ledes-

ma— 8 ( 2):603

Sedum kiersteadiae (Crassulaceae), a newly described species

from the Klamath Region of California, U.S.A. by Barbara L.

Wilson, Richard E. Brainerd, and Nick Otting— 8(1 ):9

Synopsis of Galvezia (Plantaginaceae: Antirrhineae), including a

new cryptic species from southern Peru by Michael O. Dillon

and Victor Quipuscoa Silvestre— 8(1 ):47

Taxonomic status of Kobresia curvata and Kobresia fragilis (Cy-

peraceae) by Bikash Jana and R.C. Srivastava— 8 ( 1):77

Taxonomic studies in the Miconieae (Melastomataceae). XII. Revi¬

sion of Miconia sect. Miconiastrum, with emphasis on the

Miconia bicolor complex by Walter S. Judd, James Dan Skean,

Jr., Eldis R. Becquer, and Lucas C. Majure— 8 ( 2):457

Tetragonia tetragonioides (Aizoaceae) discovered in Louisiana,

U.S.A. by Sairah M. Javed and Lowell E. Urbatsch— 8 ( 2):661

The first naturalized occurrence of the Cannaceae family in the

Arkansas (U.S.A.) flora, with additional new and noteworthy

angiosperm records for the state by Brett E. Serviss, James H.

Peck, and Tiffany A. Graves— 8 ( 2):637

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

Junior Cezar Muneratto, Luiz Antonio de Souza, and Odair Jose

Garcia de Almeida— 8(1 ):127

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

phology by Harold W. Keller— 8 ( 1):87

The vascular flora and plant communities ofCandyAbshier Wildlife

Management Area, Chambers County, Texas, U.S.A. by Jason

R. Singhurst, Amos Cooper, David J. Rosen, and Walter C.

Holmes— 8 ( 2):665

The vascular flora of Fort Sumter and Fort Moultrie, South Carolina,

U.S.A. by Richard Stalter, Brent A. Berger, Eric E. Lamont, and

John Nelson— 8 ( 1):319

The vascular flora of Galveston Island State Park, Galveston County,

Texas, U.S.A. by David J. Rosen, Shiron K. Lawrence, and Andrew

Sipocz— 8 ( 1):339

Three new species of Senegalia (Fabaceae) from Brazil by David

S. Seigler, John E. Ebinger, Petala Gomes Ribeiro, and Luciano

Paganucci De Queiroz— 8(1 ):61

Ticodendron palaios sp. nov. (Ticodendraceae), a Mid-Tertiary

fossil flower in Dominican amber by Kenton L. Chambers and

George O. Poinar, Jr.— 8 ( 2):559

Tripogon malabarica (Poaceae: Chloridoideae: Chlorideae:

Tripogoninae), a new species from Kerala, India by Thoiba

Kottekkattu and A.K. Pradeep— 8 ( 2):523

Variability of vegetative flush colors in Zamia (Cycadales) by

Claudia Calonje, Lindy Knowles, Michael Calonje, and Chad

Husby— 8 ( 2):531

Vascular plant flora of the alpine zone in the Southern Rocky

Mountains, U.S.A. by James F. Fowler, B.E. Nelson, and Ronald

L. Hartman— 8 ( 2):611

Vegetation and vascular flora of tallgrass prairie and wetlands, Black

Squirrel Creek drainage, south-central Colorado: Perspectives

from the 1940s and 2011 by Sylvia Kelso, Leah Fugere, Miroslav

Kummel, and Sebastian Tsocanos— 8(1 ):203

Vegetation patterns in the Mediterranean-desert ecotone of Baja

California, Mexico by Sula Elizabeth Vanderplank, Jose Del¬

gadillo, Exequiel Ezcurra, and Lucinda A. McDade— 8 ( 2):565

Wandersong (Rubiaceae), a new genus from the Greater Antilles

by David W. Taylor— 8 ( 2):529

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

amber by George O. Poinar, Jr.— 8 ( 1):145

682

Journal of the Botanical Research Institute of Texas 8(2)

Index of 144 Authors

Volume 8 (2014)

Thank you for choosing Journal of the Botanical Research Institute of Texas.

Abella, Scott R.—8(1 ):175

Aguilar Grijalva, Ana Eugenia—8(2):595

Aguilar, Reinaldo—8(2):449

Alexander, Patrick J.—8(2):383

Alford, Mac H.—8(1 ):267

Allen, Charles—8(1):253, 261; 8(2):663

Alves, Marccus—8(1):169

Amorim, Bruno S8(1):169

Antonio de Souza, Luiz—8(1 ):127

Antunes, Kelly—8(2):497

Asanza, Mercedes—8(2):511

Bandyopadhyay, Subir—8(1 ):71

Becquer, Eldis R.—8(2):457

Bennett, Kateland E.—8(2):419

Berger, Brent A.—8(1 ):319

Bohs, Lynn—8(1 ):1

Calderon Sanou, Irene—8(1 ):165

Calonje, Claudia—8(2):531

Calonje, Michael—8(2):531

Cardoza Ruiz, Frank Sullyvan—8(2):595,

603

Casarin Rochelle, Andre Luis—8(2):497

Cezar Muneratto, Junior—8(1 ):127

Chambers, Kenton L.—8(2):551, 559

Chittick, Kenneth L.—8(1 ):175

Clark, John L.—8(1 ):43,57

Cooper, Amos—8(2):665

Corogin, Paul T.—8(2):403

Costa Souza, Marcelo da—8(2):497

Dauzart, Ariel—8(1 ):253, 261

Delahoussaye, Jacob—8(1):253, 261

Delgadillo, Jose—8(2):565

Dillon, Michael O.—8(1):47

Doffitt, Chris—8(2):663

Douglas, Norman A.—8(2):383

Duno de Stefano, Rodrigo—8(1):169

Duquesnel, Janice A.—8(1 ):227

Ebinger, John E.—8(1 ):61

Encina-Dominguez, Juan A.—8(2):583

Ertter, Barbara—8(1 ):83

Estrada Ch., Armando—8(1 ):19; 8(2):449

Estrada-Castillon, Andres Eduar¬

do—8(1):151; 8(2):441

Ezcurra, Exequiel—8(2):565

Flores-Olvera, Hilda—8(2):383

Fowler, James F.—8(2):611

Fowler, Norma—8(2):541

Fugere, Leah—8(1 ):203

Garcia de Almeida, Odair Jose—8(1):127

Gogoi, Rajib—8(1 ):71

Gomes Ribeiro, Petala—8(1 ):61

Gonzalez Uribe, Dino U.—8(2):441

Goulart Baseia, luri—8(1):139

Graves, Tiffany A.—8(2):637

Hartman, Ronald L.—8(1 ):271; 8(2):611

Hernandez-Ledesma, Patricia—8(2):603

Herrera, Teodoso—8(2):541

Holmes, Walter C—8(2):665

Husby, Chad—8(2):531

Huskins, Stacy—8(1 ):253, 261

Jana, Bikash—8(1):77

Jasso de Rodriguez, Diana—8(2):441

Javed, Sairah M.—8(2):661

Jones, Ronald L.—8(2):677

Judd, Walter S.—8(2):403, 457

Keeling, Jeffrey—8(1 ):1

Keener, Brian R.—8(1):57

Keller, Harold W.—8(1 ):87

Kelso, Sylvia—8(1 ):203

Kennedy, Helen—8(1 ):31, 37; 8(2):493

Kirkbride, Jr., Joseph H.—8(1 ):17

Knowles, Lindy—8(2):531

Kollmann, Ludovic Jean Charles—8(2):497

Kolterman, Susan F.—8(1 ):227

Kottekkattu, Thoiba—8(2):523

Kumar, V.V. Naveen—8(2):517

Kummel, Miroslav—8(1 ):203

Lamont, Eric E.—8(1):319

Larson, Jill—8(1 ):271

Lawrence,Shiron K—8(1):339

Lewis, David—8(2):663

Lewis, M. Jerome—8(1 ):197

Lewis, Patricia—8(2):663

Linares, Jose Ledis—8(2):595, 603

Majure, Lucas C.—8(2):457

Mayfield, Mark H.—8(1 ):85

McDade, Lucinda A.—8(2):565

McNair, Daniel M.—8(1):267

McNeill, Richard P—8(2):395

Moore, Michael J.—8(2):383

Morton, Cynthia M.—8(1 ):305

Neill, David A.—8(2):511

Nelson, B E — 8(1):271; 8(2):611

Nelson, John B.—8(1):319; 8(2):25

O'Kennon, Robert J.—8(1 ):333

Ochoterena, Helga—8(2):383

Otting, Nick—8(1 ):9

Paganucci De Queiroz, Luciano—8(1 ):61

Peck, James H.—8(2):637

Poinar, Jr., George O —8(1 ):139, 145;

8(2):551, 559

Poindexter, Derick B.—8(2):419

Pradeep, A.K.—8(2):523

Quintana Martinez, Gustavo—8(1 ):151

Quipuscoa Silvestre, Victor—8(1):47

Ramirez Gamez, Abraham—8(2):441

Rayner, Douglas A.—8(2):25

Reid, Christopher—8(1 ):197

Reif, Brian—8(1 ):271

Reveal, James L.—8(1 ):83

Richard E. Brainerd—8(1 ):9

Rivera, Gerardo—8(1 ):19

Rodas, Larri A.—8(1 ):43

Rosen, David J.—8(1):339, 381; 8(2):665

Sanilkumar, M.G.—8(2):517

Santamaria Aguilar, Daniel—8(2):449

Schlesinger, Matthew D.—8(1 ):305

Seigler, David S — 8(1):61

Serviss, Brett E.—8(2):637

Silva Alfredo, Donis da—8(1):139

Singhurst, Jason R.—8(2):665

Sipocz, Andrew—8(1 ):339

Skean, Jr., James Dan—8(2):457

Sobral, Marcos—8(2):497

Sprunt, Susan V.—8(1 ):227

Srivastava, R.C.— 8(1 ):77

Stalter, Richard—8(1):319

Stern, Stephen R.—8(1 ):1

Sunil, C.N.—8(2):517

Tadeu Aguiar, Osny—8(2):497

Taylor, David W.—8(2):529

Taylor, Kimberly Norton—8(1 ):333

Terry, Martin—8(1 ):353; 8(2):541

Thompson, Ralph L.—8(2):641, 677

Trout, Keeper—8(2):541

Tsocanos, Sebastian—8(1 ):203

Tucker, Gordon C.—8(1):25

Urbatsch, Lowell E.—8(2):661

Valdes-Reyna, Jesus—8(2):583

Vanderplank Sula Elizabeth—8(2):565

Vega-Mares, Jose Humberto—8(1 ):151

Villarreal Quintanilla, Jose A.—8(1 ):151;

8(2):441,583

Weakley, Alan S.—8(2):419

Weckesser, Wendy—8(1 ):353

Wilder, George J.—8(1 ):227

Williams, Bennie—8(2):541

Wilson, Barbara L.—8(1 ):9

Index

Botanical Names and Subject Index

Volume 8 (2014)

New names (52) in bold face

683

Acacia—8(1 ):175

Achnatherum nelsonii var. nelsonii (new

state record for New Mexico)—8(1 ):271

Agave—8(2):441

x madrensis—8(2):443

gentryi—8(2):441

lechuguilla—8(2):441

Aizoaceae—8(2):661

Alexander County (Illinois, U.S.A.)—8(2):641

Amazonian (Ecuador)—8(2):511

Amber

Dominican—8(1 ):145

Myanmar—8(1 ):139

American mistletoe—8(2):641

Andes (Colombian)—8(1):43

Androceras (sect.)—8(1 ):1

Antirrhineae—8(1 ):47

Apiaceae—8(2):383

Arizona (U.S.A.)—8(2):611

Arkansas (U.S.A.)—8(2):637

Artemisia borealis ssp. borealis (new state

record for New Mexico)—8(1 ):271

Ascomycota—8(1 ):145

Asparagaceae—8(2):441

Asteraceae—8(1 ):87

Astrocasia peltata (new record for Costa

Rica)—8(1 ):165

Bahamas—8(2):531

Baja California (Mexico)—8(2):565

Black Squirrel Creek Drainage (Colorado,

U.S.A.)—8(1 ):203

Blyxa aubertii (new state record for Missis¬

sippi)—8(1 ):267

Boletales—8(1 ):139

Brassicaceae—8(2):383

Brazil—8(1 ):61,169;8(2):497

Brewster County, Texas (U.S.A.)—8(1 ):359

Browneopsis

puyensis —8(2):51 1

Cactaceae—8(2):541

Caesalpinioideae—8(1 ):71; 8(2):511

Cairo (Illinois, U.S.A.)—8(2):641

Calathea

basiflora—8(1 ):31

cofaniorum —8(1 ):37

fredii—8(1 ):37; 8(2):493

gordonii—8(1 ):31

neillii—8(1 ):37

rubribracteata—8(2)493

shishicoensis — 8(1 ):41 ; 8(2):493

timothei—8(2):493

trianae—8(2):493

California—8(1 ):9

Calyptranthes

solitaria— 8(2):498

ubatubana —8(2):500

Candy Abshier Wildlife Management Area

(Texas, U.S.A.)—8(2):665

Canna indica (new record for Arkansas)

—8(2):637

Cannaceae—8(2):637

Carex

deweyana var. deweyana (new state

record for New Mexico)—8(1):271

gynocrates (new state record for New

Mexico)—8(1):271

rosea (new state record for New Mexi¬

co)—8(1 ):271

Caricaceae—8(2):595

Caryophyllaceae—8(2):419

Central America—8(1 ):25

Chambers County (Texas, U.S.A.)—8(2):665

Chione—8(2):529

exserta—8(2):529

seminervis—8(2):529

Chlorideae—8(2):523

Chloridoideae—8(2):523

Climate Change—8(1 ):305

Coahuila (Mexico)—8(2):583

Cocos Island (Costa Rica)—8(1):25

Colchicaceae—8(1 ):261

Colleteria—8(2):529

Colombia—8(1 ):43; 8(2):493

Colombian Andes—8(1 ):43

Colorado (U.S.A.)—8(1 ):203; 8(2):611

Conservation—8(1 ):305

Cordillera Central (Colombia)—8(1 ):43

Costa Rica—8(1 ):25,165; 8(2):449

PacificoSurde—8(1):19

Crassulaceae—8(1 ):9

Cremosperm

inversum —8(1 ):57

Crepis tectorum (new state record for New

Mexico)—8(1 ):271

Cretaceous Myanmar Amber—8(1 ):139

Cuscatlania vulcanicola (new record for El

Salvador)—8(2):603

Cycadales—8(2):531

Cymopterusspellenbergii (newstate record

for New Mexico)—8(1 ):271

Cyperaceae—8(1):25, 77

Cyperus

lentiginosus—8(1 ):28

panamensis—8(1 ):28

stewartii— 8(1 ):25

Dagny Johnson Key Largo Hammock

Botanical State Park (Florida, U.S.A.)

—8(1):227

Dalea lanata (new record for Louisiana)

—8(1):197

Dead-Man's fingers—8(1 ):145

Detarieae—8(2):511

Deutzia scabra—8(2):637

Diodella—8(1 ):17

Diodia teres—8(1 ):17

Distigouania —8(2):551,552

irregularis— 8(2):552

Dominican Amber—8(1 ):145; 8(2):559

Dominican Republic—8(2):551

Draba nemorosa var. nemorosa (new state

record for New Mexico)—8(1):271

Duchesnea indica—8(1 ):83

Echinacea—8(1 ):87

Ecology and Conservation (Mojave Desert,

U.S.A.)—8(1 ):175

Ecuador—8(2):511

Prov. Sucumbios—8(1):37

El Salvador—8(2):595,603

Encephalartos ferox—8(2):533

Erigeron

compositus (new state record for New

Mexico)—8(1):271

nivalis (new state record for New

Mexico)—8(1):271

Eugenia dipetala— 8(2):502

Euphorbia—8(1 ):85

ouachitana—8(1 ):85

Euphorbiaceae—8(1 ):85,165

Fabaceae—8(1):61,71,175; 8(2):511

Florida (U.S.A.)—8(1 ):227; 8(2):403

Flush Colors—8(2):531

Fort Moultrie (South Carolina, U.S.A.)—

8(1 ):319

Fort Polk (Louisiana, U.S.A.)—8(1 ):253,261

Fort Sumter (South Carolina, U.S.A.)—

8(1 ):319

Galveston Island State Park (Texas, U.S.A.)—

8(1 ):339

Galvezia

elisensii—8(1 ):49

fruticosa—8(1 ):49

grandiflora—8(1 ):49

leucantha—8(1):49

leucantha subsp. porphyrantha —

8(1):54

leucantha subsp. pubescens—8(1):54

speciosa—8(1):47

Garnotia—8(2):517

variyamensis —8(2):51 7

Gasteroid Fungus—8(1 ):139

Genetic Diversity—8(1 ):305

Gentiana aquatica (new state record for

New Mexico)—8(1 ):271

Gesneriaceae—8(1):43,57

Glossoloma ichthyoderma—8(1):45

Glossoloma velutinum—8(1 ):43

Grasslands (Halophytic)—8(1 ):151

Gratiola quartermaniae—8(1 ):333

Greater Antilles—8(2):529

Guadalupe Mountains (Texas, U.S.A.)—

8(2):383

Gypsum flora—8(2):383

Halophytic Grasslands—8(1 ):151

Heliotropium convolvulaceum (new record

for Louisiana)—8(1 ):197

Hexasepalum

angustifolium—8(1 ):17

teres— 8(1 ):17

Hilaria mutica—8(2):583

684

Journal of the Botanical Research Institute of Texas 8(2)

Hydrocharitaceae—8(1 ):267

Icacinaceae—8(1 ):169

India—8(1 ):71,77; 8(2):517, 523

Jarilla chocola (new record for El Salvador)

—8(2):595

Joseph Plains (Idaho, U.S.A.)—8(2):383

Kalamath Region (California)—8(1 ):9

Kerala (India)—8(2):517, 523

Kobresia

curvata—8(1 ):77

fragilis—8(1):77

La Sal Mountains (Utah, U.S.A.)—8(2):611

Lactuca biennis (new state record for New

Mexico)—8(1 ):271

Lamiaceae—8(2):431

Leguminosae—8(2):511

Lepidium densiflorum var. macrocarpum

(new state record for New Mexico)—

8(1 ):271

Lepidozamia peroffskyana—8(2):533

Loeflingia squarrosa (Arkansas)—8(1):197

Lomatium

bicolor var. leptocarpum—8(2):383

cous—8(2):383

swingerae— 8(2):395

Lophophora williamsii—8(2):541

Louisiana—8(1 ):253, 261

Malvaceae—8(1):127

Marantaceae—8(1):31,37; 8(2):493

McKinney, Landon—8(2):677

Mediterranean-Desert Ecotone (Baja Cali¬

fornia, Mexico)—8(2):565

Melastomataceae—8(2):457

Melianthiaceae—8(1 ):253

Mesoamericana, Flora—8(1 ):31

Mexico—8(2):441

Mexico (Chihuahua)—8(1 ):151

Miconia—8(2):457

barbata— 8(2):478

bicolor—8(2):457

bicolor var. patenti-setosa —8(2):474

cajalbanensis —8(2):483

cristalensis— 8(2)485

guajaibonensis —8(2):474

impressa —8(2):476

karsticola —8(2):463

maestrensis— 8(2):487

sect. Miconiastrum —8(2):461

Miconiastrum (sect, of Miconia)—8(2):457

Miconieae (Melastomataceae)—8(2):457

Mid-Tertiary Amber—8(2):551,559

Miscanthus

sacchariflorus—8(2):637

sinensis (new record for Arkansas)—

8(2):637

Mississippi (new state record)—8(1 ):267

Mistletoe (American)—8(2):641

Mojave Desert (U.S.A.)—8(1 ):175

Monroe County (Florida, U.S.A.)—8(1):227

Morphology

floral—8(1 ):127

floral, stem, petiole—8(1 ):87

Myanmar Amber—8(1 ):139

Myrcia

cacuminis— 8(2):504

paulii-jonesii— 8(2):449

riverae— 8(2):452

Myrtaceae—8(2):449,497

Nash Prairie (Texas, U.S.A.)—8(1 ):381

Nerisyrenia hypercorax— 8(2):384

New Country Records

Brazil (Pleurisanthes flava)—8(1 ):169

Costa Rica (Astrocasia peltata) —

8(1 ):165

El Salvador (Cuscatlania vulcanicola)

—8(2):603

El Salvador (Jarilla chocola)—8(2):595

New Mexico—8(1 ):271

New State Record for

Arkansas (Canna indica)—8(2):637

Arkansas (Miscanthus sinensis) —

8(2):637

Louisiana (Tetragonia tetragonioides)

—8(2):661

Louisiana (Dalea lanata)—8(1):197

Louisiana (Heliotropium convolvula-

ceum)—8(1):197

Mississippi (Blyxa aubertii)—8(1 ):267

New Mexico (Achnatherum nelsonii var.

nelsonii)—8(1 ):271

New Mexico (Achnatherum nelsonii var.

nelsonii)—8(1 ):271

New Mexico (Artemisia borealis ssp.

borealis)—8(1 ):271

New Mexico (Carex deweyana var.

deweyana)—8(1 ):271

New Mexico (Carex gynocrates)—

8(1 ):271

New Mexico (Carex rosea)—8(1 ):271

New Mexico (Crepis tectorum)—

8(1 ):271

New Mexico (Cymopterus spellenber-

gii)—8(1):271

New Mexico (Draba nemorosa var.

nemorosa)—8(1 ):271

New Mexico (Erigeron compositus)

—8(1):271

New Mexico (Erigeron nivalis)—8(1 ):271

New Mexico (Gentiana aquatica)—

8(1 ):271

New Mexico (Lactuca biennis)—8(1 ):271

New Mexico (Lepidium densiflorum var.

macrocarpum)—8(1 ):271

New Mexico (Paronychia wilkinsonii)

—8(2):383

New Mexico (Penstemon glaber var.

alpinus)—8(1 ):271

New Mexico (Polemonium occidentale

var. occidentale)—8(1 ):271

New Mexico (Potentilla fissa)—8(1 ):271

New Mexico (Ranunculus repens)—

8(1):271

New Mexico (Rudbeckia laciniata var.

laciniata)—8(1 ):271

New Mexico (Silene drummondii var.

striata)—8(1):271

New Mexico (Syringa vulgaris)—

8(1):271

New Mexico (Tripleurospermum inodo-

rum)—8(1):271

North America—8(1 ):85

North Central New Mexico—8(1 ):271

Nuevo Leon (Mexico)—8(2):441

Nyctaginaceae—8(2):603

Pacifico Sur de Costa Rica—8(1 ):19

Palaeogaster —8(1 ):140

micromorpha —8(1 ):140

Panama—8(1 ):31

Paronychia wilkinsonii (New State Record

for New Mexico)—8(2):383

Passiflora

brevifila—8(1):23

soliana—8(1 ):19

Peninsular Florida (U.S.A.)—8(2):403

Penstemon glaber var. alpinus (new state

record for New Mexico)—8(1 ):271

Peru—8(1):47,57

Phanera glauca subsp. tenuiflora var.

gandhiana—8(1 ):71

Phoradendron leucarpum ssp. leucar-

pum—8(2):641

Plantaginaceae—8(1):47, 333

Pleurisanthes flava (new record for Brazil)

—8(1):169

Plinia ambivalens—8(2):507

Poaceae—8(2):517, 523, 583

Polemonium occidentale var. occidentale

(new state record for New Mexico)

—8(1):271

Potentilla fissa (new state record for New

Mexico)—8(1 ):271

Potentilleae—8(1 ):83

Prosopis—8(1):175

Ranunculus repens (new state record for

New Mexico)—8(1 ):271

Red River (Arkansas, U.S.A.)—8(1 ):197

Reed Plateau (Texas, U.S.A.)—8(1 ):359

Rhamnaceae—8(2):551

Rocky Mountains (U.S.A.)—8(2):611

Rosaceae—8(1 ):83

Rubiaceae—8(1 ):17; 8(2):529

Rudbeckia laciniata var. laciniata (new state

record for New Mexico)—8(1):271

San Francisco Peaks (Arizona, U.S.A.)—

8(2):611

Sangre de Cristo Mountains (New Mexico)

—8(1):271

Sapotaceae—8(2):403

Sedum kiersteadiae—8(1 ):9

Senegalia

harleyi—8(1):64

hatschbachii—8(1 ):66

irwinii—8(1 ):62

Sida—8(1):127

regnellii—8(1 ):127

rhombifolia—8(1 ):127

urens—8(1 ):127

Sierra Madre Oriental (Mexico)—8(2):441

Silene

drummondii var. striata (new state

record for New Mexico)—8(1):271

hitchguirei (new state record for New

Mexico)—8(1):271

Index

685

Skkim (India)—8(1 ):77

tribulosum—8(1 ):6

Ticodendron—8(2):559

Solanaceae—8(1 ):1

South Carolina (U.S.A.)—8(1 ):319; 8(2):431

palaios—8(2):560

Solanum—8(1 ):1

South Texas (U.S.A.)—8(2):541

Tithymalus (sect.)—8(1 ):85

angustifolium—8(1 ):6

Southern Rocky Mountains (U.S.A.)—

Tripleurospermum inodorum (new s

angustifolium—8(1 ):7

8(2):611

record for New Mexico)—8(1):271

citrullifolium—8(1 ):6

Stachys

Tripogon

cordicitum —8(1 ):2

arenicola—8(2):431

malabarica—8(2):523

cordicitum—8(1 ):7

aspera—8(2):431

Tripogoninae—8(2):523

davisens—8(1 ):6

caroliniana—8(2):431

Umbelliferae—8(2):383

fructo-tecto—8(1 ):6

pilosa—8(2):431

Utah (U.S.A.)—8(2):611

grayi—8(1 ):7

Stipulicida—8(2):419

Uvularia sessilifolia—8(1 ):261

heterodoxum—8(1 ):7

lacerata — 8(2):426

Valle de Janos (Mexico)—8(1 ):151

johnstonii—8(1 ):6

Sucumbios (Prov. In Ecuador)—8(1 ):37

Viscaceae—8(2):641

knoblochii —8(1 ):5

Syringa vulgaris (new state record for New

Wandersong — 8(2):530

knoblochii—8(1 ):6

Mexico)—8(1 ):271

exserta—8(2):530

leucandrum—8(1 ):7

Tallgrass Prairie (Colorado, U.S.A.)—8(1 ):203

seminervis—8(2):530

lumholtzianum—8(1 ):7

Tertiary amber—8(1 ):145

Wetlands (Colorado, U.S.A.)—8(1 ):203

novomexicanum —8(1 ):6

Tetragonia tetragonioides (new record for

Xylaria antiqua — 8( 1 ):146

rostratum—8(1 ):6

Louisiana)—8(2):661

Xylariaceae—8(1 ):145

setigeroides —8(1 ):5

Texas (U.S.A.)—8(1 ):2, 333, 339, 359, 381;

Zacatal deToboso—8(2):583

setigeroides—8(1 ):6

8(2):665

Zamia—8(2):531

tenuipes—8(1 ):6

Ticodendraceae—8(2):559

Zigadenus densus—8(1 ):253

52 New names and New Combinations:

Volume 8 (2014)

Agave x madrensis Villarreal, A. Ramirez, & A.E. Estrada, hybrid

nov.— 8(2):443

Browneopsis puyensis D.A. Neill & Asanza, sp. nov.— 8(2):511

Calathea

cofaniorum H. Kenn., sp. nov.— 8(1 ):37

gordonii H. Kenn., sp. nov.— 8(1 ):31

rubribracteata H. Kenn., sp. nov— 8(2):493

shishicoensis H. Kenn., sp. nov.— 8(1 ):41

Calyptranthes

solitaria Sobral, Aguiar, & Antunes, sp. nov.— 8(2):498

ubatubana Sobral & Rochelle, sp. nov.— 8(2):500

Cremosperma inversum B.R. Keener & J.L. Clark, sp. nov.— 8(1 ):57

Cyperus stewartii G.C. Tucker, sp. nov.— 8(1):25

Distigouania K.L. Chambers & Poinar, gen. nov.— 8(2):552

irregularis K.L. Chambers & Poinar, sp. nov.— 8(2):552

Eugenia dipetala Sobral & Kollmann, sp. nov.— 8(2):502

Galvezia elisensii M.O. Dillon & Quipuscoa, sp. nov.— 8(1 ):49

Garnotia variyamensis Sunil, Naveen Kumar, & Sanilkumar, sp.

nov.— 8(2):517

Glossoloma velutinum J.L. Clark & L.A. Rodas, sp. nov.— 8(1 ):43

Hexasepalum teres (Walter) J.H. Kirkbr., comb. nov.— 8(1 ):17

Lomatium swingerae R.P. McNeill, sp. nov.— 8(2):395

Miconia

barbata (Borhidi) Judd, Becquer, & Majure, comb. nov.—

8(2):478

bicolor var. patenti-setosa (Borhidi) Judd, Becquer, & Majure,

comb. nov.— 8(2):474

cajalbanensis Judd, Becquer, & Majure, nom. nov.— 8(2):483

cristalensis (Borhidi) Judd, Becquer, & Majure, comb. nov.—

8(2):485

guajaibonensisJudd, Becquer, & Majure, nom. nov.— 8(2):474

impressa (Urb.) Judd, Becquer, & Majure, comb. nov.— 8(2):476

karsticola Judd, Becquer, Skean, & Majure, nom. nov.— 8(2):463

maestrensis Judd, Becquer, & Majure, nom. nov.— 8(2):487

Miconia sect. Miconiastrum (Bonpl. ex Naudin) Judd, Becquer, &

Majure, comb, nov.— 8(2):461

Myrcia

cacuminis Kollmann & Sobral, sp. nov.— 8(2):504

paulii-jonesii Aguilar, D. Santam., & A. Estrada, sp. nov.—

8(2):449

riverae A. Estrada, D. Santam., & Aguilar, sp. nov.— 8(2):452

Nerisyrenia hypercorax PJ. Alexander & MJ. Moore, sp. nov.—

8(2):384

Palaeogaster Poinar, Alfredo, & Baseia, gen. nov.— 8(1 ):140

micromorpha Poinar, Alfredo, & Baseia, sp. nov.— 8(1 ):140

Passiflora soliana A. Estrada &G. Rivera, sp. nov.— 8(1 ):19

Phanera glauca subsp. tenuiflora var. gandhiana Gogoi & Ban-

dyop., var. nov.— 8(1 ):71

Plinia ambivalens M. Souza & Sobral, sp. nov.— 8(2):507

Sedum kiersteadiae B.L. Wilson & R.E. Brainerd, sp. nov.— 8(1 ):9

Senegalia

harleyi Seigler, Ebinger, & P.G. Ribeiro, sp. nov.— 8(1):64

hatschbachii Seigler, Ebinger, & P.G. Ribeiro, sp. nov.— 8(1 ):66

irwinii Seigler, Ebinger, & P.G. Ribeiro, sp. nov.— 8(1 ):62

Solanum

cordicitum S. Stern, sp. nov.— 8(1 ):2

knoblochii (Whalen) S. Stern, comb. &stat. nov.— 8(1 ):5

novomexicanum (Bartlett) S. Stern, comb. & stat. nov.— 8(1 ):6

setigeroides (Whalen) S. Stern, comb. & stat. nov.— 8(1 ):5

Stachys caroliniana J.B. Nelson & D.A. Rayner, sp. nov.— 8(2):431

Stipulicida lacerata (C.W. James) D.B. Poind., K.E. Bennett, &

Weakley, comb, et stat. nov.— 8(2):426

Ticodendron palaios K.L. Chambers & Poinar, sp. nov.— 8(2):560

Tripogon malabarica Thoiba & Pradeep, sp. nov.— 8(2):523

Wandersong D.W. Taylor, gen. nov.— 8(2):530

exserta (DC.) D.W. Taylor, comb. nov.— 8(2):530

seminervis (Urb. & Ekman) D.W.Taylor, comb. nov.— 8(2):530

Xylaria antiqua Poinar, sp. nov.— 8(1 ):146

New Titles from BRIT Press

Manual of Montana Vascular Plants

August 2013

I 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-

I 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

I plants. It contains descriptions as well as habitat and distribution information

I 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,

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 |

tropical, conditions, just the opposite of much of Texas. This book explains I

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, I

and detailed maps. Be ready to be surprised by this special group of Texas I

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.

Press

: titles and others, plea